t > ' t \ V sr/-ri5i:^ ^ n VOLUME 56 C ^ 1969 ANNALS OF THE Missouri Botanical Garden Published by the Missouri Botanical Garden Press St. Louis, Missouri 63110 CONTENTS FLORA OF PANAMA, PART VI Family 88. Zygophyllaceae Duncan M. Porter FLORA OF PANAMA, PART VIII r Family 16 L Menyanthaceae Thomas S. Elias Family 167. Boraginaceae Joan W. Noxvicke Family 169. Labiatae loan W. Nowicke £r Car Viable Soil Algae from the Herbarium of the Missouri Botanical Garden Bruce C. Parker, Noel Schanen & Richard Renner Scanning Electron Microscopy as an Aid to Pollen Taxonomy John E. Ridgway & John J. Skvarla On Tropical Tree Seedlings. I. Seeds, Seedlings, Systems, and Systematics James A. Duke Petalostemon Sally A. Walker NOTES Supplementary Notes on the Phytolaccaceae, I. Joan W. Nowicke 1 Family 96. Polygalaceae Walter H. Lewis & Oha Herrera-MacBryde 9 29 33 71 113 121 125 CHI) Viktor Muehlenhach 163 Notes on the Galapagos Euphorbieae (Euphorbiaceae) Derek Burch . 173 The Genus Cienfuegosia Cav. (Malvaceae) Paul A. Fryxell ... 179 The Initiation of Vascular Cambium and Production of Secondary Xylem in Flower Bud Pedicels of Asclepias curassavica L. F. M. Safwat . . 251 Cytological and Chromatographic Evidence of Interspecific Hybridization in 261 The Genus Hoffmannia (Rubiaceae) in Panama John D. Dwyer . . 269 The Skunk Cabbage in Missouri Erna R. Eisendrath .... 287 288 Kallstroemia (Zygophyllaceae) in Missouri Duncan M. Porter 290 Tropical Island Biogeography: The Missouri Botanical Garden's Sixteenth Annual Systematics S>'mposium Duncan M. Porter 293 Seasonal Flowering Behavior in Central Panama Thomas B. Croat ... 295 Studies in Neotropical Paleobotany. I. The Oligocene Communities of Puer- to Rico Alan Graham & David M. Jarzen 308 Wood Anatomy of Goodcniaceae and the Problem of Insular Woodiness Sherwin Carlqidst The Cahfornia Islands Robert F, Thome NOTES 358 391 417 419 Distribution Patterns of West Indian Mosses Marshall R. Crosby ... 409 Drosophilidae of Hawaii Hampton L. Carson Vegetation of a Galapagos Island before and after an Ice Age ?aul A. Colinvaiix Pollen Morphology of Saprophytic Taxa in the Gentianaceae Siwert Nils- son &■ John Skx'arla Cytotaxonomic Notes on Some Neotropical Gentianaceae Richard E. Weaver, ]r Spores of the Heterophyllous Selaginellae of Mexico and Central America Robert Lee Hellwig 420 439 444 The Ozark Flora— Some Collections of Note W. G. D'Arcy 465 468 Additional Type Material of Bouvardia (Rubiaceae) Will H. Black- well, Jr. Gr Marshall C. Johnston A New Species of Scaer^ola (Goodcniaceae) from Tahiti Sheruin Carl- quist Chromosome Numbers of Phanerogams. 3 Some new combinations in Protium (Burseraceae) Duncan M. Porter 475 469 471 ANNALS Missouri VOLUME 56 1969 NUMBER 1 Garden CONTENTS FLORA OF PANAMA, PART VI If Family 88. Zygopbyllaceae by Duncan M. Porter ^"^ Family 96. Polygal by Walter H. Lewis & Olga Herrera-MacBryde 9-28 FLORA OF PANAMA, PART VIII Family 161. A^enyonffiaceoe by Thomas S. Ellas .... . . . • • • • * ^^"^^ J r Family 167. Borag/naceoe by Joan W. Nowlcke ^^'^^ Family 169. LabiataB by Joan W. Nowicke & Carl C. Epiing 71-111 ■A N. VOLUME 56 1969 NUMBER 1 ANNALS OF TH ■^ - - ' _ L " ■ ^ n , ^ Missouri Botanical Garden - ■ ' ._-"■'-"" A journal containing scientific contributions from the Missouri Botanical Garden and tlie Department of Botany of Washington University in affiliation with the Missouri Botanical Garden, Outside contributions in systematic botany and allied fields will also be con- sidered. These papers are subject to a charge of $2 5 per printed page. ■_ h J— f -.^ Editorial Committee ' - Derek BuRCH, EJffor, pro few. Missouri Botanical Garden & Washington University Sheri Davis, Assistant to the Editor W - X - Marshall Crosby, Missouri Botanical Garden John D, Dwyer, Missouri Botanical Garden & St. Louis University , < ■■ ' Joan Nowicke, Missouri Botanical Garden & Washington University Duncan Porter, Missouri Botanical Garden £r Washington University -*, * -■ v ^ . -r Information The Annals of the Missouri Botanical Garden appears times during the calendar year. Three numbers totaling about 400 page stitute a volume • ■■ ^ ... - , - Beginning with Volume 53, 1966: Subscription Price ...... ... . $15.00 per volume three Single Numbers 5.50 each Contents of previous issues of the Annals of the Missouri Botan- ical Garden are listed in the Agricultural Index, published by the H. W. Wilson Company. r y^ Beginning June 1, 1962 the Stcchert-Hafner Service Agency, Inc., 31 East 10th St., New York 3, N.Y., becanie sole agent for the Annals of the Missouri Botanical Garden. The Agency handles all subscriptions, all claims beginning with volume 50 (1963) and all requests for back issues. Out-of-print numbers will be reprinted as may be required. Under the arrangement with the Stechcrt-Hafner Service Agency, the Agency handles also all purchases of any portion of the Annals subseries known as the "Flora of Panama.'* All matters regarding exchange continue to be handled by the Missouri Botanical Garden, 2315 Tower Grove Ave., St. Louis, Missouri 63110, as in the past. '■ h ■ ■■■'■.--.- ..■-.■.. The Annals of the Missouri Botanical Garden are listed in Cur- rent Contents Agricultural, Food and Veterinary Sciences. < ^ L H -I n_ < , ■- - - Printed by Bardgctt Printing and Publishing Company, St. Louis, Mo. 63103. VOLUME 56 1969 NUMBER 1 ANNALS OF TH Missouri Botanical Garden FLORA OF PANAMA BY Robert E. Woodson, Jr. and Robert W. Schery AND Collaborators Part VI Family 88. ZYGOPHYLLACEAE' BY Duncan M. Porter Missouri Botanical Garden Annual to perennial, often suffrutescent, herbs, shrubs, or occasionally trees; branches usually divaricate, with angled or swollen nodes, growth sympodial. Leaves opposite or occasionally alternate, usually even-pinnate, occasionally simple or 2-foiiolate, rarely 3-7-folioIate, often fleshy to coriaceous, petiolate to subsessile, persistent; leaflets usually entire, inequilateral, petioluhite to subsessile; stipules paired, free, foliaceous, fleshy, or spinescent, persistent or rarely deciduous. Flowers (4-)5(-6)-merous, bisexual, hypogynous, regular or occasionally slightly irregular; peduncles terminal or pseudaxillary, 1 -flowered, solitary or occasionally few to many; sepals (4-) 5 (-6), free or rarely slightly connate basally, imbricate, persistent or occasionally deciduous; petals (4-)5(-6), free or rarely connate basallv, often clawed, sometimes twisted, imbricate or convolute, deciduous, rarely marcescent; extrastaminal and/or intrastaminal disc usually present; stamens m (l-)2(-3) whorls of 5 each, outer whorl usually opposite petals, often alternately unequal in length or sterile, the filaments free, subulate to filiform or rarely winged, frequently glandular or appendaged basally, occasionally outer whorl basally adnate to petals, inserted on or below disc, the anthers 2-loculed, sub- (20 C2-)5(-12) ( l-)2-manv placcntation axilc or rarely basal, the style terminal, usually simple, the stigma minutely and obscurely lobed to distincll} ridged. Fruit a (2-)5-lobed loculicidal or septicidal capsule or a schi/oearp splitting lengthwise into 5 or 10(-12) hard tuberclcd to spiny or winged mericarps, rarely a drupe or a drupaceous berry; seeds iC-manv) per loeule, endosperm present or absent. Assisted by National Science Foundation Grant No. GB'5674. Ann. Missouri Bot. Gard. 56(1): 1-7, 1969. 2 ANNALS OF THE MISSOURI [Vol. 56 BOTANICAL GAKDEN A family of about 2 7 genera and 2 50 species, widely distributed, but mainly in the warmer, drier regions of the world. Two genera, Kalhtroemia Scop, and Tribulus L., have been collected in Panama. Guaiacum sanctum L., from the northern Caribbean, is culti\ated in Balboa, while G. officinale L., from the south- ern Caribbean, is cultivated in Panama City and the Canal Zone. The latter also has been reported from Bocas del Toro Province by Allen (Ceiba 10: 35-36, Although specimens of Guaiacum collected in the wild have not been ) fj / O "J — ^.- occur along the Caribbean coast. Both Guaiacum officinale {guayacdn, iron uood, lignum vitae) and G. sanctum (guayacdn, lignum vitae) are widely cultivated in the tropics for their globose growth habits and myriads of blue or purple flowers. Their wood is very hard, tough, and resinous, and G. officinale, in particular, has been used to a con- siderable extent in the manufacture of small turned objects and stern bearings for ships. a. Herbs; nowcrs yellow, stamens iinainuTKl;iged; fruit a schizocarp separating at maturity into 5-10 mericarps. b. Fruit tuberclcd, at maturity separating into 10 mericarps, beak persisting- intrastaminal glands absent 1. Kalhtroemia l)b. Iruit spmy, at maturity separating into 5 mericarps, beak falling witb meri- carps; intrastaminal glands present 2. Trihiilus aa. Trees or shrubs; Howers blue or purple, stamens appendaged; fruit a 2-5-lobed loeulicidal capsule (-; itaiacHm 1. KALISlROIuMIA Kallslroemia Scop., Introd. 212, 1777. EhrcHbcrgia Mart., Nov. Gen. Sp. Brasil. 2: 72, 1826, non Spreng., Neue Entdeek. 2: 129, 1821. Hetcmzygis Bunge, Mem. Acad. Imp. Sci. St.-Pctersbourg Divers Savans 2: 82, 183 5. Amiual or occasionall) perennial herbs; stems herbaceous to suffrutescent, difl'usely branched, prostrate to decumbent or ascending, terete, somewhat suc- culent, becoming striate on ilr\ing, densely pubescent to glabrate, spreading radially from a central lap root to 1 m or more long. I.eaves opposite, elliptical to broadly obovate, abruptly even-jMnnate, one of each pair alternatelv small sometimes abortive; leaflets 2-10 pairs, opposite, entire, subscssile, elliptical to broadly oblong or obo\ate, somewhat uneciua! in si/.e, those on one side of rachis slightly smaller, lowest pair markedly unec|ual, terminal pair pointed forward and more falcate, pubescent to glabrate; stipules foliaceous. Fhuers solitary, pseudaxil- lary, regidar; peduncles emerging from axils of alternately smaller leaves; sepals er or 5(-6) ) orange, the base same or green to red, free, spreading heiiu'sphericallv, fugaceous, usually marcescent, convolute; disc flesh\, annular, ohscurelv lo'C-1 2>]()bed'; stamens lOC-12), 5C-6) opposite petals exterior, somewhat longer, and adnate basally to petals, 5 (-6) opposite sepals subtended to extcrinr h\ a small bilobed land, the hlaments filiform, subiflate, or rarely basally winged, unappendaged, inserted on disc, the anthers globose or ovoid to linear-obiong or rarelv linear, those opposite sepals rarely aborting; ovary sessile, 10(-1 2)-lobed anddoculed, cr]„bose or 1969] FLORA OF PANAMA (Family 88. Zygophyllaceae^ 3 ovoid or occasionally conical or pyramidal, glabrous to pubescent, the ovules 1 per locule, pendulous, placentation axile, sometimes 1 or more aborting, the style sim- ple, cylindrical to broadly conical, more or less 10(-12)-ridged, persisting to form a beak on fruit, the stigma capitate, oblong, or clavate, 10(-12)-ridged or -lobcd, papillose or rarely coarsely canescent, terminal or rarely extending down almost to base of style. Fruit 10(-12)-lobed, ovoid, occasionally conical, or rarely pyra- midal, glabrous or pubescent, at maturity dividing septicidally and separating from a persistent styliferous axis into 10('12) or occasionally fewer mericarps; mcricarps 1-loculed, 1-seeded, obliquely triangular, wedge-shaped, more or less tuberculate or rugose abaxially; seeds oblong-ovoid, obhquely pendulous, the testa membranaceous, the embyro straight, endosperm absent. The largest New World genus of the family, with 17 species occurring in warm arid and tropical areas from the southern half of the United States to central Argentina. Also introduced into WY'st Africa and India. The two species found in Panama apparently are indigenous. a. Ovary and fruit ovoid, g]a])rous or occasionally strigose basally; sepals ovate, hirsute with appressed to spreading trichomes, in fruit clasping but not entirely covering mature mericarps, only scarious margins becoming involute 1. K. maxima aa. Ovary pyramidal, fruit ovoid, both densely appressed short-pilose; sepals lan- ceolate, hispidulous with fine trichomes of two lengths, spreading from base of mature fruit and appearing hnear lanceolate through margins becoming sharply involute 2. K. pubescens 1. Kallslroemia maxima (L.) Hook. & Arn., Bot. Beechey 282, 1838. Trihiihis maximiis L., Sp. PI. 386, 1753. T. trijiigatiis Nutt., Gen. 1: 277, 1818. Kallstrocmia trihidus Meisn., PI. Vase. Gen. 2: 43, 1837, nom. superfl. Trihidus decolor Macfad., Fl. Jamaica 186, 1837, nom. superfl. T. dimidiatus Raf., Autikon Bot. 176, 1840, nom. superfl. T. trijiigus Steudel, Nom. Bot. ed. 2, 2: 699, 1841, nom. superfl. T. tiiherculaiiis Sesse & Moc, Fl. Mex. ed. 2, 109, 1894. Kallstrocmia cancsccns Rydb. in Vail & Rydb., N. Amer. Fl. 25: 113, 1910. Annual herb; stems prostrate to decumbent, to 1 m long, sericeous and spar- ingly hirsute with apically-directed white trichomes. J eaves obovatc, 1-5.5 cm long, 1-3.5 cm wide; leaflets 3-4 pairs, broadly oblong to elliptical, appressed- hirsute to glabrate, margins and veins sericeous, 5^18 mm long, 3-11 mm wide, terminal pair usually largest. Flowers ca 1 cm in diam; peduncles to 28 mm long; sepals ovate, 4-5 mm long, 2 mm wide, as long as or little shorter than petals, hirsute with appressed to spreading trichomes, in flower ca as long as style, in fruit clasphig but not entirely covering mature mericarps and shorter than beak, onlv scarious margins becoming involute, persistent; petals yellowy fading orange, obovate, 4-6 mm long, to 6 mm wide, marcescent; ovary ovoid, ca 1 nun in diam, usually glabrous but occasionally strigose basally, the style 2 mm long, cylindrical, base slightly conical, the stigma capitate, ca 1 mm long, obscurely lOdobed, papillose. Fruit ovoid, 4-5 mm in diam, usually glabrous but occasionally strigose basally, the beak 3-6 mm long, glabrous, cylindrical, the base widely conical; mericarps 4 mm high, 1,5 mm wide, abaxially tubercled, cross-ridged, and slightly keeled, sides pitted, adaxial edge angled. [Vol. 56 4 ANNALS Ob THE MISSOURI BOTANICAL GARDEN South Carolina, Georgia, Tlorida, and Texas, Sinaloa and Tamaulipas, Mex- ico, to Central America, the West Indies, and northern South America. Called verdohiga throughout much of Latin America, as it closely resembles Portulaca oleracca L. {PortiiUwacfae), the true rcrdooa. The latter is much used b} the populations as a potlierb. lii Paiuuna Kalhtrocuiia niaxhua is known as hicrha de poUo (hde Pro. Hcrihcrto 3 5). CANAL zone: Catun, Cowcll 3/4 t.i\Y), Hio. Hcrihcrto 35 (LS); Farfan Beach, Duycr 6798 (MO); Ancon, Grccnniau & iWccnmau 50/9 (MO); Frijolcs, railroad track! liilliV 3445 in part (US); vie of Ancon, Piper 58/5 (US); bctw Panama 8c Corozal, alt 20-30 m, Pitticr 4443 (F, (ill, NY, US); Balboa, roadside, Staudlcy 26/03 (US); C'hiva Chiva Trail nr Miraflorcs Lake, disturbed area, Tyson /385 (ATTC); Mirallores Locks area, roadside, Tyson 1609 (ATTC, MO); Pedro Miguel garbage dump, Tyson & Bluw 4017 (ATTC). ciiiRiyui: David airport, disturbed grassland & edge of marsh, alt 2 5 ft, Lcuis ct al. 737 (MO), los santos: 5-9 mi from C:hitre on rd to Las Tablas, roadside, Biirch ct al. 1216 (F, MO); 10 km S of Chitre, Corrcu A. 85 (ATTC). Panama: Panama, roadsides, Hayes 728 (BlM); \ era C:ru/, beach & sandy areas behind, Lcuis ct al. 3015 (MO); sabanas N of Panama, Bra. Paid 526 (US), phovtncf unknown: Bahia Soldado, C.oicaU 227 (NY); s. loc, Sccmauu s.n. (BM). 2. Fvallsiroemia piihesrens (G. Don) Dandy in Keav, Kew Bull. 10: 138, 1955. — Fig. 1. Trihulus imhcsccns G. Don, Ccn. Syst. 1: 769, 183 1. KaUstroemia minor Hook. f. in Hook., Niger. Fl. 269, 1849, nom. superfl. Trihulus maxiwus var. minor (Hook. f. in Hook.) Oliver, Fl. Trop. Afr. 1: 284, 1868. KaUstroemia carihaca Pydb. in Vail & Pydb., N. Amer, Fl. 23: 111, 1910. K. fihihrata Rydb. in Vail ik P\db., loc. cit. 1 12. K. longipes R\db. in \ail & liydb., loc. cit. I 14. Annual Jicrh; stems prostrate to decimibent, to 1 m long, hirsute and sericeous with apically-directed fine white trichomes. Leaves obovate, to 2 cm long and 1 cm \vidc; leaflets (2-) 3 pairs, elliptical to obovate, appressed-hirsute to glahrate, \eins and margins more or less sericeous, to 1 1 nnn long and 5 nmi wide, terminal pair largest. Vhnvers ea 1 cm in diam; peduncles 5-15 mm long; sepals lanceolate, 3.5-4 nun long, 1.5 mm wide, in flower ca as long as st)le, hispidulous with line white trichomes of two lengths, spreachng from base of mature fruit and appearing linear-lanceolate through margins becoming sharply involute, persistent; petals yellow, obovate, ca 6 mm long and 5 mm wide, marcescent; ovar\ pvra- midal, 4 mm long including style, densely appressed short-pilose with hne while trichomes, the style stout, conical, the stigma capit1(1 fare type refer to descriptions; numbers in ronian type refer to synonyms; numbers with dagger (t) refer to niimes incidentally mentioned. rhrenbergia 2 Cuaiacnm 2t officinale 2 sanctimi 2t Heterozygis 2 Kallstroeniia 2t, 2 canescens 3 caribaea 4 glabrata 4 longpipes 4 maxima .'J, 4t minor 4 pubescens 4 tribulus 3 Portulacaceae 4t Portidaca oleracea 4'*' Tril)ulus 2t, 5 cistoides 6t decolor 3 dimidiatus 3 maxim us 3 — var. minor 4 pubescens 4 terrestris 6t, 6 trijugatus 3 trijugus 3 tuberculatus 3 FLORA OF PANAMA BY Robert E. Woodson, ]r. and Robkht \V. Schkry AND COLLABOI^VTORS Part VI FAMILY 96. POLVGALACEAE^ BY Walter II. Lewis and Olc;a Herhlla-MacBryde Missouri Botanical Garden, St. Louis Herbs, erect or scandeiit shrubs, or rarely small trees. Leaves alternate, sonie- tiiiies opposite or vertieillate, simple; stipules absent. Fluvers 5 , ± /ygomorphic, usually racemose, bracteate; sepals 5, unequal (slightly unequal in \]()HtahecO, the 2 inner large and often petaloid, wing-like; petals 3 or sometimes 5, declinatc, if 3 the lower usually keel-shaped, often appendaged, the 2 upper ligulate or ovate, usually adnate to the lower petal and connate to the staminal sheath, if 5 connate above the middle into a tube and adnate with the sepals, subequal; sta- mens 8, rarely fewer, the hlamcnts connate for most of their length to form a c ) (1-) r 1 per loculc, pendu- lous. Fruit a capsule, drupe, berry, or samara, deliisccnt or indehisccnt; seeds 1 per loeule, rarely 2, often pilose, usually arillatc, llie endosperm straight. Eighteen genera with over 1000 species; temperate and tropical regions. Four ( ) a. Calyx free, the sepals iine(|ual with the 2 inner sepals (win^s) larger, petaloid; petals 3; ovary 1-2'locnlar. b. Ovary and fruit 2-locular, the fruit a dehiseent capsule; mostly delicate herbs 1- Polygala bb. Ovary and fruit l-loeular, the fruit indehiscent; mostly somewhat woody. c. Scandcnt shrubs; keel clawed with a fimbriate crest, plicate; fruit a samara with a large wing; lowlands, occasionally cloud forests. . . . 2. Securidaca cc. Erect herbs, shrubs, or rarely trees; keel neither plicate, clawed, nor ap- pendaged; fruit drupaceous, not at all or only verv obscurely winged; cloud forests onlv 3. MoitJiina aa. Calyx adnate to the corolla, the sepals ± e(]u.il, not petaloid; petals 5; ovary (2')4-5-locidar 4. Moutahca 1. POLYGALA Polygala L., Sp. Pi. 701, 1753. Shrubs, suhshrubs, or commonly herbs in Panama. Leaves alternate, some- times opposite or vertieillate, simple, short petioled or sessile. Racemes terminal or lateral, sometimes contracted into heads. Floucrs often showy; sepals 5, un- equal, the outer 3 herbaceous, free or sometimes 2 connate basally, the inner 2 ^Assisted bv National Science Foundation Grant No. GB-8438. Ann. Missouri Box. Card. 56(1): 9-28, 1969. 10 ANNALS OI- THE MISSOURI BOTANICAL GARDEN [Vol. 56 large, petaloid, wing-like; petals 3, connate basally, the lower keel-shaped, crested, lobed, or occasionally unappendaged, the 2 upper hgulate to ovate, connate to the keel or adnate to the staniinal sheath or both; stamens 8, the hlaments connate nearly to the apex into a sheath spht above and adnate to the petals below; ovary 2-locular, the ovules solitary, the style incurved, the stigma dilated, entire to 2- lobed, often tufted. Capsules usually membranous, compressed, obovate to orbi- cular, generally wingless, loculicidally dehiscent; seeds usually 2, often pilose, mostly arillate, with or without endosperm. A genus of perhaps 500 species, ± cosmopolitan; 13 are recogni/ed in Panama. n. Leaves ahernate (those of the basal '/4 of plant usually vcrticillate or sul)vcrti- eillatc). 1). Leaves linear-lanceolate to elliptic or ovate, 2-^5 mm wide; keel without ap^ pcndages. e. Annual; leaves linear to lineardanceolate, sometimes elliptic, 1-4. S cm long, 2-9 mm wide; outer sepals oblong ovate, obtuse, usually glandular-ciliatc on the margin, the larger one 1.8-2.2 mm lon^, the smaller 2 almost en- tirely connate, 1.5-1.9 mm long; capsule oblong-ellipsoid, 2.9-3.5 mm long, glabrous; seeds 1.9-2.5 mm long; aril corneous, crest-like, with 2 short appendages, 0.6-0.8 mm long, glabrous 1. P. hrizoiile$ cc. Suflruticulose; leaves elliptic to ovate, 2.5-8 cm long, l--^.5 cm wide; outer sepals lanceolate, acute or acuminate, spreading pih)se, usually all 3 ca the same si/e, 3-5 mm long; capsule ovoid to orbicular, 7.5-11 mm long, spreading pilose; seeds 3.5-5 mm long; aril thin, umbraculiform, without appendages, 2.5-4 nun long, somewhat pilose 2. P. inwamcnsis bf). Leaves narrowly linear or linear-Ianceolatc, 0.2-3.5 mm \\ ide; keel with appendages. d. llacemes elongate, loosely n(j\\ered, to 8 cm long. e. Flowers white; seeds ca 1 mm long, with a few weakly hook-tipped hairs; aril obsolete; rare 3. P. fcndlcri var. hcterothrix ee. Flowers rose to pur|)lish, rarely white; seeds 1-1.8 nun long, with all hairs straight; aril present. f. Stems glabrous; seeds 1-1.3 mm long; aril barely 2-lobed, ca 1/10 as long as the seed; occasional 4. P. Icptocaitlis H. Stems tlensely pedicillate-glandular; seeds 1.4-1.8 nun long; aril strongl) 2-lobed, '/4-I/2 as long as the seed; \ery conunon dd. luaccmcs compact, densely flowered, to 2.6 cm long. 5. P. ]Hiniciilata g. Keel not gland dotted; capsule suborbicular; seeds ellipsoid, sparsely short-pubescent, the 2 lobes of the aril 0.9 1.6 mm long. . . . 6. P. Jn-j^rophila gg. Keel gland dotted apicalh and basally; capsule ovoid (jl)long; seeds conical, sericeous, comose basally, the 2 lobes of the aril 0.5 mm long to absent. h. Wings strongly cuspidate apically; seeds 1.8-2.5 mm long (in- cluding coma); aril minute or absent; frequent 7. P. lon^icaiilis hh. Wings not or rarely bluntly mucronate apically; seeds 2.5-2.7 mm long C including coma); aril 2-lobed, 0.5 mm long; J"'»r^' 8. P. variahilis aa. Leaves all verticillate (rarely alternate in apical V4 of P. iiujouiou'). i. Mostly delicate herbs to 0.3 m tall; leaves 4-21 mm long, 2.7-8 mm wide; seeds 1-1.8 mm long, lobes of the aril narrowly oblong, 0.9 1.6 mm long. j. Racemes compact, very densely flowered, 8-11 mm witle; sepals acumin- ate apically, 1.5-2.4 mm long; wing much larger than the keel, mucro- nate at the rounded ape\, 3-4.6 mm long, 1.8-3.2 mm wide 9. /'. timoutoH 1969] FLORA OF PANAMA (Family 96. Polygalaceae^ 11 jj. Racemes loose or somewhat densely flowered, 4-6 mm wide; sepals ob- tuse apieally (the lower 2 often somewhat acute in P. aparinoides^^ 0.7-2 mm long; wings ea the same length as the keel, not mucronate at the rounded apex, 1.6-2 mm long, 1-1.2 mm wide. k. Margin of leaves denticulate; seeds often equaled by the 2 narrowly o])long lobes of the aril 10. P. aparinoides kk. Margin of leaves entire or rarely remotely crenate; lobes of the aril ca half the length of the seed 11. P. asperidoides ii. Sufl'rutescent perennials to 2 m tall; leaves 4-7.5 cm long, 1.8-3 cm wide; seeds 4 mm long, lobes of the aril wide, 2.5-3 mm long. I. Leaves apieally acute, sparingl}- punctate; sepals without glands; aril lobes broadly obovate 12. P. wurdackiana II. Leaves apieally acuminate, densely punctate; smaller pair of sepals each with 2 large glands; aril lobes oblong 13. P. jefensis 1. Polygala hrizoides St.-Hil., Fl. Bras. Mcrid. 2: 30, t. 88, 1829. P. anoustifoUu H.B.K., Nov. Gen. Sp. PI. 5: 405, t. 511, 1823, non Gilib. (FL Lituan. 2: 11-3, 1781). Erect or ± ascending, simple or branched annnah, 6-39 cm high; stems ter- ete, ± densely incurved- or loosely-spreading incurved-pubescent. Leaves alter- nate, linear to lineardanceolate, sometimes elliptic, acute, acuminate or mucro- nate, rarely obtuse apieally, acute or acuminate basally, the lateral nerves 3-6 pairs or obsolete (then 1 -nerved), sparsely incurved-pubescent or glabrescent, 1-4.5 cm long, 2-9 mm wide; petioles 0.7-2 mm long. Racemes loosely flowered, includ- ing the axis to 8 cm long; bracts deciduous, ovate-lanceolate, acute, the margin ciliate, 1-1.8 mm long. Flowers purple, greenish-white, or greenish-lavender, on slender pendulous pedicels, 1.2-1.5 mm long; outer sepals oblong-ovate, obtuse, usually glandular-ciliate on the margin, rarely eglandular, the largest 1.8-2.2 mm long, the smaller 2 almost entirely connate, 1.5-1.9 mm long; wings unequal, broadly obovate, retuse apieally, cuneatc basally, the margin somewhat undulate, obscurely flabellinerved, glabrous, 3.2-4 nun long, 2.5-3.4 mm wide; upper petals angustate in the middle, suborbicular-obovate, 3-3.2 mm long, retuse apieally, the margin often undulate, the basal half ± hairy inside; keel 3.5-4 mm long without appendages; staminal tube sparsely pid^escent basally or glabres- cent, the fdaments 1.4 1.6 mm long; ovary oblong, 0.9-1.5 mm long, 0.8-1.1 nmi wide, the style curved, 2.4-2.7 mm long, the stigma lateral, concave, villous- ciliate. Capsules oblongdlipsoid, 2.9-3.5 mm long, 2.1-2.8 mm wide, glabrous; seeds (Fig- Ic) oblong, 1.9-2.5 mm long, densely pilose; aril corneous, crest-like at the top of the seed \\ith 2 short appendages, 0.6-0.8 nmi long, glabrous. Mexico to Parayuav and the West Indies, CANAL zone: Pipeline Rd, at end of rd, nr Gatiin Lake, Croat 4728 (MO); Ft Kobbc nr BOQ, Duke 3934 (MO); Sosa Hill, Duke 4649 (MO); rd to Coro/al, Gcrvais 144 (US); Balboa Heisbts, dry hillsides, Killip 3062 (US); W slope of Aneon Hill, vie of Balboa, alt 20-7 5 ni, Seihert 382 (GH, MO); Ballwa, moist thieket, scarce, Standky 25467 (US), weedy field, Standley 2566? (US), moist thicket, Stattdlcy 28323 (US). CHiRiQUi: Boquete, savannas, alt 4000 ft, Duvichon 754 (MO, US); David airport, dis- turbed grassland & edge of marsh, alt 25 ft, I^enis et al 770 (MO), 773 (MO); Sabana dc la Tortuga, bctw El Boquetc & Caldera, alt 300-700 m, Vittier 3337 (MO); vie of Boquetc, llanos Francia, alt 3300 ft, Stern ct til 1171 (MO), cocle: vie of Xata, alt 50 m, AUett 834 (GH, MO, US); Bio Hato Airstrip, Blum C- Duycr 2464 (MO); Fl Vallc, ca 3.6 km from Fl Greco Hotel along rd to InterAmerican Hwy, Currca & Dressier 1- ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 1012 (MO); Aguadulce, savannas, nr sea level, Pitticr 4867 (US); betw Las Margaritas & El Valle, Wooihou ct al. 1272 (GH, MO, NY), panama: savanna nr Rio Pacora & Chepo Hvvy, Duke 5935 (MO); along rd toward top of Cerro Campana, bald savanna-like areas, Duke 5952 (MO); nr Chepo, savannas, Duke 6052 (MO); Ccrro Campana, grass- land, Ehinger 944 (MO); San Jose I, Hast Harbor, roeky coastal bluffs, Erianson 441 (GH, MO), grassy bluffs, Erianson 583 (GH, MO); Tumba-Muerto, Bro. Ucrihcrto 187 (US); N of Panama City, savannas, Bro. raid 521 (US); Sabana de Juan Corso, nr Chepo, alt 60-80 m, Pitticr 4506 (US), Pitticr 4743a (US); Laguna de Portalla, nr Chepo, alt 50 m, Pitticr 463 5 (US); vie of Juan Franco Race Track, savanna, scarce, Standley 27728 (US); E of Pacora, wet savanna, alt ca 25 m, Woodson ct al. 753 (MO); nr Arraijan, thickets & forests, alt ca 15 m, Woodson ct al. 7373 (Gil, MO, NY, US). Considered a variety of P. munticola II.B.K. by Stcvermark (Fieldiana: Bot. 28: 300, 1952). 2. Polygala panamensis Chodat, Mem. Sue. Phys. Geneve 31(2): 35, 1893. /f 15-68 em long, pilose, leaves, alter- nate, elliptie to ovate, aeute or aeuminate apieally, cuneate basally, (the lower sometimes oval and obtuse), with ea 5 pairs of lateral nerves, somewhat retieulate, pilose above and beneath, usually more densely so along the veins beneath, 2.5-8 em long, 1-3.5 em wide; petioles 1-2 mm long, pilose. Racemes loosely flowered, 2-10 em long (ineluding the short axis), densely pilose; bracts deciduous, linear to linear-lanceolate, pilose, 1.8-3 mm long. Flowers blue or purple, the pedicels 1.5-3.3 nun long; outer sepals lanceolate, aeute or acuminate, spreading pilose, all 3 ca equal, 3-5 mm long (occasionally 1 to 0.5 mm longer); wings oblong, rounded apieally, cuneate basally, pilose along the eosta, 7-10 mm long, 3.8-5 mm wide; upper petals narrowly obovate-spatulate, ciliate below or glabrescent, equaling the keel; keel 6.5-8 mm long, without appendages; staminal tube slightly split to ca halfway into 2 bundles each of 4 partly free filaments, the filaments 1.5-2.2 mm long; ovary oblong, 1.5-3.5 mm long, 0.8-1.1 mm wide, pubescent, the style curved, 3.2-4.5 mm long, the stigmas 2, slightly spaced, the outer sub- apical on a small stalk, the inner sessile. Capsules ovoid to orbicular, 7.5-11 mm long, 6.5-10 mm wide, spreading-pilose; seeds (Fig. li) flattened, 3.5-5 mm long, pilose; aril membranaceous, umbraculiform, 2.5-4 mm long, pilose, the mar- gin irregularly toothed. Panama. cuiKiyui: Vic of El Boquetc, alt 1000-1500 m, Cornman 2054 (US); Boquete, sa- vannas, alt 4000 ft, Davidson 749 (MO); El Boquete, roadside, alt 1300 m, Killip 3501 (US); vie of Boquete to 3 mi N, alt 3300-4200 ft, second growth, cultivated areas & roadsides, Leu is ct al. 375 (MO); El Bocjuete, pastures, alt 1000-1300 m, Pitticr 2891 (NY, US); betw Cerro Vaca & Hato del Loro, 850-1100 ft, Pitticr 5394 (F, GH, US); Rio Chiriqui Viejo Valley, nr El Volcan, White 232 (GH, MO, US); valley of' upper Rio Chiriqui Viejo, White & White 102 (MO), cocle: vie of El Valle, alt 600-1000 m, Allen 1153 (MO); vie of Ola, alt 100-350 m, Pitticr 5077 (US). Closely related to if not eonspecific \\ ith P. custaricensis Chodat and P. diir- andii Chodat of Guatemala and Costa I'ica. 3. Polygala fendleri Chodat var. heterothrix Blake, Bull. Torrcv Bot. Club, 51 : 85, 1924. Frect, very slender, usually branched annual; 6-15 cm high; stems ribbed, glabrous. Leaves whorled in 3-4's at the base of the stem, lancc-obovate, 3-5 nun 1969] FLORA OF PANAMA (Family 96. Polygalaceae^ H Jong, 1-1.8 mm wide, the upper alternate, linear or linear-lanceolate, 4-10 mm long, 0.4-1.5 mm wide, acute to cuspidate, 1-nerved, the margins slightly revo^ lute, glahrous, subsessile. llacemes loosely flowered, cylindric, to 7 cm long, 5.5-6.5 mm wide, the axis to 3 cm long; bracts deciduous, linear-lanceolate, acute, glabrous, 0.8-1.1 mm long. Flowers white, the pedicels to 0.8 mm long; largest sepal broadly ovate to suborbicular, rounded, 0.9-1.1 mm long, the smaller 2 oblong, obtuse, 0.8 1 mm long; wings elliptic or subobovate, rounded apically, short-cuneate basally, 2.2-3 mm long, 1.2-2.1 mm wide; upper petals oblong, rounded, equaling the keel; keel 2-2.6 mm long, 4-lobed, the lobes 0.6-0.8 mm long; staminal tube glabrous inside, anthers subsessile; ovary subor- bicular, 0.5-0.7 mm long, 0.4-0.6 mm wide, the style ascendent, 0.5-0.7 mm long, developed distally into a navicular cavity, the inner end of which has a sessile, globose stigma. Capsules narrowly elhpsoid, 1.8-2 mm long, 1 mm wide; seeds (Fig. Id) ca 1 mm long, the hairs mostly straight, a few weakly hook- tipped; aril obsolete. Panama. CHiRTQiiT: Boqucte, Llanos IVancia, 4 mi from Boqiietc toward Dolcga, llanos border- ing creek, alt ca 4500 ft, Dwycr & Hayden 761 HA (MO). coci.E: vie of El Valle de Anton, alt ca 600 m, Allen 1933 (MO, US); betvv las Margaritas & El Valle, open llano, Woodson i't al 1261 (MO, US). 4. Polygala leplocaiilis Forrey & Gray, Fl. N, Amer. 1: 130, 1838. Erect, simple and usually slender mniiials; 12-40 cm high; stems subangu- late, glabrous. Leaves alternate, linear, acute to acuminate, 1 -nerved, the mar- gins slightly revolute, glabrous, 4-2 5 mm long, 0.2-1 mm wide, subsessile. Racemes cylindric, loosely flowered, 1-8 cm long, 5-5.5 mm wide, the axis to 10 cm long; bracts deciduous, lanceolate, acute, glabrous, 0.7-1.2 mm long. Floirers deep rose, the pedicels 2 mm long; outer sepals 0.8-1 mm long, the largest broadly ovate, the smaller 2 ovate to oblong; wings obo\ate or elliptic, rounded apically, cuneate basally, 3-nerved, 1.8-2 mm long, 0.7.0-9 mm wide; upper petals oblong or ovate, usually emarginate at the somewhat narrowed apex, 1.7- 1.9 mm long; keel 1.3-1.7 mm long, the crest 5-10-lobed, the lobes 0.2-0.7 mm long, usually single, sometimes forked apically; staminal tube glabrous inside, the filaments very short; ovar) oblong or o\oid, 0.3-0.6 mm long, 0.2-0.3 mm wide, the style minute, developed distally into a navicular cavity, the apical end of which has a tuft of hairs, the inner end a globose, sessile stigma. Capstdes oblong, with a row of glands on each side of the septum, 1.5-1.8 mm long, 0.7-0.9 mm wide; seeds (Fig. Ig) oblong, 1-1.3 mm long, sparsely appressed-pubescent, aril minute, 2-lobcd, appressed, ca 1/10 as long as the seed, glabrous. Louisiana and throughout Central America and the West Indies to northern Argentina. Frequently confused with P. pajiiculata L. CANAL ZONE: Chiva-Chiva, dry fields, Killip 37 28 (MO, US), chiriqui: Llanos Francia, vie of Boquete, alt 3300 ft. Stern et al 1158 (GH, MO, US); vie of Boqucte, rooted in water, alt 1200-1500 m, Woodson & Schcry 781 (MO), cocle: vie of El Valle, alt 800-1000 m, Allen 99 (MO); El Valle de Anton, along Rio Anton, lower portion of valley & marshes, alt ca 500 m, Uuntcr & Allen 376 (MO); Pieacho de Ola, alt 350-600 m, Pittier 5070 (US); El Valle de Anton & vie, on tussocks in bogs, alt 500-700 m, Seihert [Vol. 56 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN 488 (GH, MO, NY, US), panama: along rd bctw Panama & Chcpo, in savannas, Dodge ct al. 16691 (MO); nr Rio Pacora & Chcpo Hwy, savanna, Diikc 5936 (MO), 5942 (MO); Tocumen, field, Dwyer 2569 (MO); Tocunien airport, Dwycr 2583a (GH); Pa- cora, Mendez 107 (MO); E of Pacora, wet savanna, alt ca 2 5 m, Wuudsoti ct ul. 731 (GH, MO, US). 5. Polygala paniculata L., Syst. Nat. ed. 10, 1154, 17 59. Erect, mostly much-branched annuah, 10-50 cm high; stems terete, densely pedicellate-glandular. Leaves 4-5-\vhorled or pseudowhorled at the base of the stem, otherwise alternate, lanceolate to linear-lanceolate, acute apically, cuneate basally, 1-nerved, 5-32 mm long, 0.6-3.5 mm wide, short petioled. Racemes cylindric, loosely flowered, 3-7 cm long, 5-7 mm wide, the peduncle to 14 cm long; bracts deciduous, lanceolate, acute, glabrous, 0.8-1.1 mm long. Fkmers rose or purplish, rarely white, the pedicels 0.7-1.2 mm long; outer sepals bearing 2 glands basally, the largest sepal broadl) ovate, obtuse, 1-1.2 mm long, the smaller 2 oblong-ovate, obtuse, 0.9-1.1 mm long; wings obovate, spatulate-obovate or oblong-obovate, rounded apically, cuneate basally, 3-nerved, 2-2.8 mm long, 0.7-1 mm wide; upper petals narrowly ovate-lanceolate, attenuate to the apex, 2-2.5 mm long; keel 1.9-2.7 mm long, the crest 6-10-lobed, the lobes 0.4-0.9 mm long, rarely partly forked apically; staminal tube glabrous inside, the filaments very short; ovary obovoid to almost orbicular, 0.6-0.8 mm long, 0.4-0.6 mm wide, the style straight or ascending, 0.4-0.8 mm long, developed distally into a navicular cavity, the apical end of whicli has a tuft of hairs, the inner end a globose, sessile stigma. Capsules ellipsoid to oblong, 1.8-2.6 mm long, 0.9-1.5 mm wide; seeds (Fig. Ij) oblong, 1.4-1.8 mm long, shortly appressed-pubescent, the 2 lobes of the aril narrowly oblong, appressed, 0.4-0.8 nun long. A pantropical weed; in the Americas extending from Texas to southern Brazil. The species is very common in Panama from sea level to at least 2 300 in. BOCAs DEL TORO: s. loc, Ciirlctou 126 (GH, XY, US); Changiiinola to 5 mi S at jet of Rios Changuinola & Tcrebc, edge of river & railway, & adjacent rain forest, alt 100- 200 ft, Leuis ct al. 798 (MO), canal zonk: Barro Colorado T, Axiles 70 (F, MO), 943 (F); Ft San Lorenzo, Burch ct al. 1026 (MO); Ancon, Cclcsthic 59 (US); nr BOQ, Ft Kobbe, Duke 3925 (MO); vie of Hill C-6, Ft Sherman, shaded areas, Duke 4384 (MO); (Rio) Chagres, Fcndlcr 55 (MO, US); Ancon Hill, Greenman & Grccnman 5106 (MOy, Orchid I off Barro Colorado I, clearing at French Lock site, Kc7iovcr 406 (US); Ancon Hill, dry hillsides, alt 100 m, Killip 3019 (MO, US); betw Frijoles & Monte T irio, along railroad, alt 30 m, Killip 12122 (US); top of bluff above gulf, McDauicl 5177 (MO); Balboa, Mell 5 (MO); Ancon, Bro. Paul 159 (US); nr Old Ft San Lwenzo, mouth of Rio Chagres, Piper 5933 (US); vie of Ancon, Piper 6001 (US); W slope of Ancon Hill, vie of Balboa, Scihcrt 110 QG]], MO, NY); Barro Colorado I, Gross Pt., Shuttuck 943 (MO); Balboa, moist thicket, Standley 25567 (US), wet open bank, Staiidley 25590 (US), weed in garden, Standley 26110 (US), in cornfield, Sta7id!cy 29305 (US); hills \V of Canal, nr Gatun, Standley 27300 (US); Gamboa, brushy slope, Standley 28372 (US); vie of Summit, brushy slope, Standley 30079 (US); vie of Ft Sherman, Standley 30923 (US); Daricn Station, open bank, Standley 31564 (US); vie of Rio Cocoli, Rd K-9, along road- side, Su-rn ct al. 40 (MO, US), chiriqui: Boquctc, alt 3800 ft, Davidson 594 (F, GH, MO, US); Fl Bocjuete, roadside, alt 2300 m, Killip 3503 (US); vie Bw]uete, from Bo- quete to 3 miles N, second growth, cultivated areas & roadsides, alt 3300-4200 ft, Lcuis et al. 330 (MO), 344 (MO); vie of Fl Boquete, alt 1000-1300 m, Maxun 4929 (US), brushy pasture, Maxon 5114 (US); pastures around El Boquete, alt 1000-1300 m, Pittler 2892 (US); vie of Boquete, alt 1200-1500 m, Woodson & Schcry 720 (GH, MO). cocle: El Valle, ca 3.1 km from El Greco Hotel, along rd, Correa £r Dressier 1010 1969] FLORA OF PANAMA (Family 96. Polygalaceae^ 15 (MO); 12 mi NE of Penonome, granitic river gorge, alt 1200 ft, Lewis et al 1508 (MO); hills NE of El Valle de Anton, disturbed forest edges, alt 2000 ft, Lcxvis ct al 1808 (MO); El Valle de Anton, edge of cloud forest & roadside, 1000 2000 ft, Lcuis ct al 2545 (MO); at foot of Cerro Pilon, cloud forest, alt ca 2000 ft, Dwycr & Correa 7986 (MO); above Penonome, Williams 277 (NY, US); betw Las Margaritas & El Valle, Wood- son ct al 1240 (GH, MO, NY, US), 1747 (GH, MO), colon: Maria Chiquita, along shore above beach, Ebi7igcr 444 (MO); 2.4 mi SW of Pina, rain forest by beach & rd, Lewis ct al 1846 (MO); betw^ Rio Piedras & Puerto Pilon, roadside, Lewis et al 3214 (MO); vie of Palenque, nr sea level, Pittier 4131 (US), darien: village of Mannene, Kirkbride & Bristan J 596 (MO), herrera: 10 mi S of Ocii on Las Minas Rd, Graham 246 (GH). PANAMvv: vie of provincial boundary, along rd between Las Uvas, Province of Panama, and El Valle, Code Province, Bartlett & Lasser 16664a (MO), 5-6 mi E of Chepo on PanAmerican Hwy, Duke 4022 (GH, MO); Rio Chagres, 1 mi above Madden Lake, Duke 4478 (MO); PanAmerican Hwy at Rio Mamoni, ca 4 mi beyond Chepo, Duke 5565 (GH, MO); Cerro Azul, Duke 9366 (MO); Alhajuela, river bank, Bro. Heriberto 1 (US); 2-3 mi S of Goofy Lake, rd to Cerro Jefe, lower elev of cloud forest, alt 2000-2200 ft, Lewis et al 239 (MO); tributary of Rio Chagres, 5 mi SW of Cerro Brewster, sandy & rocky river banks, alt ca 1000 ft, Lewis et al 3506 (MO); Cerro Campana, common along roadside nr moist woods, alt 2900 ft, McDaniel 6873 (MO); banks of Mamoni River, above Chepo, alt 20-25 m, Pittier 4727 (US), san blas: airport at Irandi, Duke 6505 (GH, MO); mainland opposite Ailigandi, from mouth of Rio Ailigandi to 2.5 mi inland, marshy rain forest, often native plantations, Lewis et al 64 (MO), province un- know^n: s. loc, Moore 42 (F); western Panama, Stork s.n. July-Aug 1923 (US). 6. Polygala hygrophila H.B.K., Nov. Gen. Sp. PI. 5: 395, 1823. Slender, erect annuals, simple or slightly branched, 7-50 cm high; stems wing-angled, glabrous. Leaves subverticillate at the base of the stem, the others alternate, linear to linear-lanceolate, mucronate or acute apically, cuneate basally, 1-nerved, the margins shghtly revolute, glabrous, 6-12 mm long, 0.5-1.2 mm wide, sessile. Racemes very dense, cylindric or cylindric-ovoid, slightly comose apically, 1-2.6 cm long, 6.5-12 mm wide, the peduncle to 5 cm long; bracts linear-lanceolate, acuminate-subulate, usually carinate, ciliolate basally, glabrescent toward the apex, 2-3 mm long. Flowers white or greenish-white, tinted pink or lavender, subsessile; sepals elliptic-oblong to ovate, subacute, occasionally with a ridge along the costa on the dorsal side, then usually appearing carinate, 1.7-2.2 mm long; wings oval, sometimes suborbicular, rounded apically, cuneate-rounded basally, sometimes with a ridge along the costa on the dorsal side, then usually appearing concave, 5-7-nerved, 4,3-5 mm long, 3-3.8 mm wide; upper petals oval or obovate-oblong, rounded apically, ca equaling the keel; keel 2.5-3 mm long, the crest irregularly trilobate on each side; staminal tube glabrous inside, the hiaments 0.4-0.6 mm long; ovary ovoid to orbicular, 0.5-1.3 mm long, 0.4- 0.7 mm wide, the style ascendent, 0.5-1 mm long, developed distally into a navicular cavity, the apical end of which has a tuft of hairs, the inner end a globose, sessile stigma. Capsules suborbicular, 1.5-1.9 mm long, 1.4-1.9 mm wide; seeds (Fig. le) ellipsoid, 1.7-1.9 mm long, sparsely short-pubescent; aril corneous above, the 2 lobes narrowly and obliquely oblong, apprcssed, 0.9-1.6 mm long, glabrous. British Honduras, Costa Rica, Panama and Santo Domingo; northern South America. CANAL ZONE: Rio Azotc Caballo, savannas along drowned river, alt 66-70 m, Dodge et al 16854 (MO); Sabana de Marcelito, nr El Vigia, Pittier 2384 (US), chiriqui: nr [Vol. 56 16 ANNALS OF THE MISSOLIU BOLAINICAL GARDEN El Boqiictc, pastures, alt 1000-1300 m, Pitticr 2^93 (US); Sahana dc la Tortuga, bctw El Boqucte & CaUlcra, after fire, alt .300-700 m, Pitticr 33J7 (US); Ccrro Vaca, alt 900- 1136 m, Pitticr 5347 (F, US); Jarcmillo, Ro<|ucte, alt 4S00 ft, Terry 1274 (F, GH, MO, US); vie of Boqiiete, alt 1200-1500 m, Woodson & Schcry 780 (MO), cocll: Penon- onie, level grcjiintl outside town, Ehingcr 1013 (MO); Aguadulee, savannas nr sea level, Pitticr 4876 (GH, NY, US); Pieaeho de Ola, alt 350-600 m, Pi7//er 5058 (US). Pan- ama: Casa Larga, dry savannas, alt 65 m. Alien 2971 (GH, MO, US); hills NE of Haeienda La Joya, alt 50-300 ni, Dod^e ct id, 16877 (MO); nr Rio Paeora & Chepo hwy, savanna, Dnkc 5914 (MO); Juan Diaz, dry fields, alt 50 m, KUlip 3267 (MO, US); Sli slope of Cerro Campana, roadside & grasslantl, Leiris ct ah 3ii5 (MO); Cerro Campana, eoninion in hog in grassland, alt 2400 ft, McDanicI 6921 (MO); Sahana dc Dormisolo, nr C:hepo, alt 60-80 m, Pitticr 4651 (US). 7. Polygala lonj^iraulis 11. U.K., Nov. Gen. Sp. PL 5: 396, 1823. Slender erect autmals, simple or sparsely branched, 6 60 cm high; stems glabrous. I cares alternate, linear or linear-lanceolate, somelimes with a whorl of 3 reduced narrow obo\ate leaves at base of stem, acute or cuspidate apically, acute or attenuate basally, 1 -nerved, the margin slighth rexolute, glabrous, punc- tate, 4-28 mm long, 0.5-3 mm wide, subsessile. Racemes depresscd-subglobose or globose, densely llowered, 6-12 mm long, 6-18 mm wide, the axis to 3.5 cm Jong, the peduncle to 12 cm long; bracts lance-ovate, subulate, the margins glandular-ciliate, usually gland-dotted on back, 0.8-1.2 mm long. Floivers vary- ing from white to deep purple, the pedicels 1.5-3 mm long; largest sepal ovate or elliptic, cuspidate, 5-nerved, 1.7-2.2 nun long, the smaller 2 ovate, occasion- ally elliptic, ctispidate, gland-dotted basally, 1.6-1.9 mm long; wings elliptic, obong or OAate-oblong, cuspidate apically, usually cuneate-rounded basally, 3- ner\ed, bearing a few glands on back, 4.7-5.8 mm long, 1.5-2 mm wide; upper petals suboblong to nearly obovate, obtuse, 3.6-5.3 mm long; keel 3-4.3 mm long, gland-dotted apicall\ and basally, the crest lobed, the lobes 0.6 1 mm long, usually 6, rarely 8, single or irregularly forked; staminal tube glabrous inside, the filaments very short; o\ary oblong, 1-1.4 mm long, 0.7-^0.9 nun wide, the style slender, straight, 1.8-2.2 mm long, developed distally into a navicular cavity, the apical end of which has a tuft of hairs, the inner end a globose, papillose stigma supported on a minute stalk. Capsules ovoid oblong, bearing a double line of glands along the septum, 2.8-4 mm long, 1.4-2.2 mm wide; seeds (Fig. Ih) conic, sericeous, comose basally, 1.8-2.5 nun long (including the coma); aril minute or absent. Mexico through Central America and the West Indies to Argentina; frequent in Panama. CANAL /onf: nr BOQ, Ft Kohhe, Didw 3912 (MO); summit of Ancon Hill, dry hill- sides, Killip 3021 (GH, NY, US); open grassy slope, common, Staudlcy 25217 (US). CHTRiyui: W of Gualaea, level ground, eommon, alt 500 ft, AUcu 5052 (MO); hi Valle, ca .^.6 m from II Creeo Hotel along rd to InterAmeriean Hwy, Corrca & Dressier 1008 (MO); David airport, disturhed grassland, edge of marsh, alt 2 5 ft, Lewis ct al 768 (MO), cocll: Nata, alt ca 50 m, Allen 825 (GH, MO, NY, US); Llano Bonito, .\ of Las Margaritas, alt 400^500 m, Scihert 522 (GH, MO, NY); hetvv has Margaritas & El Valle, Woodson ct id. 7266 (GH, MO, NY, US), 1274 (GH, MO). Panama: vie of provincial houndary, along rd betw has Uvas (Panama) and El Valle (Coele), Bartlett & Lasscr /666S (MO); Tahoga I, alt 900 ft, Collciistie 475 (US); nr beach at Nueva Gorgona, Didic 4543 (MO); rd toward top of Cerro Campana, hald savanna-like areas, Didie 5951 (MO); nr Chepo, savannas, Duke 6038 (MO); Tocumen airport, Duyer 1969] FLORA OF PANAMA (Family 96. Pulygalaceae) 17 I a b d g ■ V ' \j^^r .■-'■^^^ -^ .f -'i ' V* V I "x f I *f^to.-i.^..x\ e^ '^ f - j k ? rl ^V-'.'^V.'flu m Fig. 1. Seeds of Polygala species. A, V. aparinoidcs Hook. & Arn.; B, V. a^peruloidcs H.B.K.; C, P. brizoides St.-HiL; D, P. {cndleri var. hetcrothrix Blake; E, P. hygrophila H.B.K. F, P, jefensis W. H. Lewis; G, P. leptocaulis Torrey & Gray; H, P. longicaitU^ H.B.K. I, P. panamensis Chodat; J, P. panicidatu L.; K, P. timontou Aiiblet; L, P. variahdis H.B.K. M, P. ivtirdackiana W. H. Lewis, All X 10. [Vol. 56 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2575 (MO); nr llioniar, open pasture land, Ebi)i^cr 505 (MO); Ccrro Cainpana, grass- land, Ehingcr 930 (MO); San Jose I, East Harbor, roeky eoastal l)luffs, frhmson 445 (GH, US), vie of South Beaeh, rocky coastal bluffs, Erlaiisoii 52 7 (GH, US); San Jose I, East Harbor, on bald spots reach inj; greater height than surrounding grasses, ilarluw 22 (GH, US); I'umba Muerto, Bra. Hcriherto 170 (US); San b)se I, East Bav, rocky open place at head of sea-cliffs, ]nhusU)n /93 (GH, MO, US); laboga I, dry fields, 200 m, KiUip 3189 (MO, US); (Eas) Sabanas, Bro. Paul 15 (US); C:hepo, alt 60 m, Pilticr s.n., Oct 1911 (US); Eaguna de Portala, nr Uhepo, alt 50 m, Pitticr 4572 (US); Eas Sabanas, open grassy slope, common, Staiidlcy 259 72 (US); nr Punta Paitilla, savanna, common, StuucUcy 26294 (US); E of Paeora, wet savanna, alt ca 2 5 m, Wuochon ct (il. 754 (MO). VEK.VGUAS: 2 mi W of Santiago on PanAmerican Huv, common along banks of buy, l)u}crcf ill. 7549 (MO). 8. Polyf,'aIa variabilis II.B.K., Nov. Gen. Sp. Pi. 5: 397, 1823. Slender erect aiiiiiuils, simple or sparsely branched, 14-20 cm high; stems ribbed, ghibruus. Lanes all alternate, linear, acute or acuminate, 1 -nerved, glab- rous, usually punctate, 3-9 mm long, 0.3-0.8 mm wide, subsessile. Racemes somewhat dense, conic, to 1 cm long and to 7 mm wide, tbe axis to 3 cm long. l-luwers purple, the pedicels 0.5-0.8 mm long; largest sepal ovate, obtuse or blunt, 1.5-1.9 mm long, sometimes with a few basal glands; the smaller 2 ovate to sub- oblong, obtuse, 1-1.4 nun long, gland-dotted along the costa; wings elliptic to oblong, rounded or rarely bluntly mucronate apicalh, cuneate basally, 3-ner\ed, bearing a few glands basally and rarely alojig the costa, 2.6-3.2 nun long, 2.5-3.4 e; upper petals o\ ate-lanceolate, obtuse, ca equaling the keel; keel 2.5-3.4 mm long, gland-dotted apically and basally, the crest lobed, the lobes 6 or 8, 0.5-0.8 mm long; slaminal tube glabrous inside, the filaments very short; ovary oblong, 0.5-0.8 mm long, 0.1-0.2 mm wide, the style straigbt, 0.3-0.4 jnm long, developed distally into a na\icular cavity, the apical end of which has a tuft of hairs, the inner end a globose stigma supported on a minute stalk. Capsules ovoid- oblong, 3 mm long, 1.2-1.6 mm wide; seeds (big. 11) conical, sericeous, comose basally, 2.5-2.7 nnn long (including the coma), the 2 lobes of the aril narrowly oblong, appressed, 0.5 nun long, glabrous. rare in Patiama. C Amazon basin) c:oci.r,: Aguadulce, savannas nr sea level, Pilticr 4875 (US). This species is closely related to P. lo)iojcauUs II.B.K. 9. Fohfrala tinioutou Aublet, Hist. PI. Gui. Fr. 737, 1775. Slender erect a}i}itiah, sijnple or slighth branched, 8-18 cm high; stems wing- angled, glabrous. J eaves nearly all in whorls of 3, occasionally the uppermost, opposite or alternate, oval to obovate, mucronate at the acutish apex, cuneate basall), 1-nerved, glabrous, 4-16 mm long, 3-5 mm wide, sessile or subsessile. Racemes eylindric, \'ery dense, slightly comose apically, 1-4 cm long, 8-11 mm wide, the peduncle very short, to 12 mm long; bracts linear-lanceolate, subulate, 2-3 nnn long. Flouers pink, stibsessile; sepals lanceolate, acuminate, 1.5-2.4 mm long; wings o\al to suborbicular, mucronate at the rounded apex, cuneate-rounded basally, ca 3-nerved, 3-4.6 mm long, 1.8-3.2 mm wide; upper petals oblong to oboxate-oblong, rounded apicallv, ca ec|ualing the keel; keel 1.8-2 nnn loner, '^the 1969] FLORA OF PANAMA (Family 96. Polygalaceae') 19 crest 6-9-lobed, the lobes 0.3-0.6 mm long; staminal tube glabrous inside, the filaments 0.4-0.8 mm long; ovary ovoid to almost orbicular, 0.5-0.9 mm long, 0.4-0.6 mm wide, the style ascendent, 0.4-0.6 mm long, developed distally into a navicular cavity, the apical end of which has a tuft of hairs, the inner end a globose, sessile stigma. Capsules orbicular to orbicular-ovoid, 1.2-1.4 mm long, 1-1.2 mm wide; seeds (Fig. Ik) ellipsoid, 1.2-1.3 mm long, sparsely short- pubescent; aril corneous above, the 2 lobes narrowly oblong, appressed, 0.6-0.9 mm long. British Honduras, Costa Rica and Panama; Trinidad and northern South America to the Amazon delta. CANAL zone: along drowned Rio Azote Caballo, alt 66 70 m, Dodge et al J 6855 (MO); Sabana de Marcclito, nr Fl Vif^ia, Fittier 2385 (US), panama: Casa Larga, dry savannas, alt 65 m, Allen 2972 (MO). 10. Polygala ai)ariiu)i(les Hook. & Arn., Bot. Beech. Voy. 227, 1836. Erect or spreading, simple or branched slender perennials, 5-27 cm high; stems winged, glabrous. Leaves verticillate, in whorls of 3-5, obovate to elliptic- obovate, sometimes the uppermost lanceolate to elliptic-lanceolate, 7-21 mm long, 4-8 mm wide, cuspidate apically, attenuate basally, 1 -nerved, the margin denticu- late, glabrous, glandular-punctate; petioles ca 1 mm long. Racemes cylindric, acuminate, rather dense, 4-20 mm long, 4-6 mm wide, the axis to 3.5 cm long; bracts deciduous, lanceolate, acuminate, glabrous, 0.6-0.8 mm long. Tluwers purple or greenish-purple, the pedicels 0.4-0.7 mm long, outer sepals often mar- inally ciliate from the apex to the middle, glabrous toward the base, usually gland-dotted, the largest ovate, obtuse, 1-1.2 mm long, the smaller 2 oblong to oblong-ovate, often somewhat acute, otherwise obtuse, 0.7-1 mm long; wings obo- vate to elliptic-obovate, rounded apically, short-clawed basally, glabrous, usually gland-dotted, 1.6-2 mm long, 1.1-1.2 mm wide; upper petals 1.9-2.2 mm long, ovate to rhombic-ovate, obtuse or rounded; keel 1.4-1.6 mm long, the crest 4-6- lobed, the lobes 0.3-0.7 mm long; staminal tube often split ca halfway into 2 bundles, each of 4 short filaments; ovary oblong to ovoid, 0.4-0.6 mm long, 0.3-0.5 mm wide, the style ascending, 0.2-0.3 mm long, developed distally into a navicular cavity, the apical end of which has a tuft of hairs, the inner end a globose, sessile stigma. Capsules broadly ellipsoid, 1.7-2.2 mm long, 1.4-1.9 mm wide; seeds (Fig. la) oblong, 1.4-1.8 mm long, appressed-pilose, often equaled by the 2 narrowly oblong, appressed lobes of the aril. Mexico (Jalisco), Guatemala, British Honduras and Panama. CHiRiyui: Boqucte, alt 3800 ft, swamp in potrcro, Davidson 574 (MO, US); El Boquctc, alt 1300 m, Killip 3500 (US); 12.4 mi N of David, cdgf of steep river bank, Lewis et al. 721 (MO); vie of Boqucte, alt 1200-1500 m, Woodson & Schery 779 (MO), 785 (MO). Closely allied to P. asperuloides H.B.K. 11. Polygala asperuloides H.B.K., Xov. Gen. Sp. PI. 5: 403, 1823. Erect or spreading, simple or branched slender annuals, 5-13 cm high; stems winged, glabrous. Leaves verticillate, often in whorls of 4-5, the uppermost some- -0 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 times in 2's or 3's, lanceolate or lance-elliptic, 7-9 mm long, 2.7-7 mm wide, the lowermost usually ovate or orbicular-ohovate, 7-9 mm long, 4.5-5 mm wide, cuspidate apically, attenuate basally, 1-nerved, the margins entire or rarely re- motely crenate, glabrous, glandular-punctate; petioles less than 1 mm long. Ra- cemes cylindric, loosely flowered, 4-22 mm long, 4-6 nnn wide, the axis to 3.5 cm long; bracts deciduous, lanceolate, subulate, glabrous, 0.5-0.7 mm long. i' loners purple, the pedicels 0.4-0.7 mm long; outer sepals sometimes ciholate, with 2 large glands on the back, the largest sepal o\ate to orbicular-ovate, obtuse, 1-2 mm long, the smaller 2 oblong, obtuse, 0.7-0.9 mm long; wings elliptic to obovatc, rounded apically, cuneate basally, glabrous, gland-dotted, 1.6-1.8 mm long, 1 mm wide; upper petals 1.8-2 nun long, triangular-ovate to rhombic-ovate, obtuse; keel 1.4-1.6 mm long \\ith 2 large glands apically, the crest 6-lobed, the lobes 0.4-0.7 mm long; staminal tube glabrous inside, the hlaments 0.1-0.3 mm long; ovary oblong or ovoid, 0.3-0.5 mm long, 0.3-0.4 mm wide, gland-dotted, the style ascending, 0.2-0.3 mm long, developed distallv into a navicular ca\itv the apical end of which has a tuft of hairs, the inner end a dobose, sessile stigma. Capsules broadly ellipsoid, 1.2-1.4 mm long, 1-1.2 mm wide, gland-dotted; seeds (Fig. lb) oblong, 1-1.2 mm long, appressed-pilosc; aril with the 2 lobes oblong, appressed, 0.5-0.7 mm long. Guatemala, British Honduras and Panama; also in Colombia and Venezuela. cocle: Vic of LI Vallc, ah 800-1000 m, Allen 773 (MO, US); vie of Fl Vallo dc \nt()n, alt 800 m, Allen 1932 (iMO, US); Llano Bonito, N of Las Margaritas, open llanos, ah 400-500 m, Seihert 523 (MO, US). Panama: along rd toward top of (Vrro Cam- pana, bald savannadike areas, Duke 5950 (MO); Cerro Campana, grassland, Ehhii-cr 920 (MO). / 12. Polygala wunlackiana W. 11. Lew is, Ann. Missouri Bot. Card. 55: 365 fig. L 1969. Suffrutescent perennials to 1.3 m, branched above, often with 2-4 branchlets at a node, glabrous, angular, green. Leaves (2-) 3-5 in whorls; petioles 4-10 mm long, glabrous; blades elliptic to less commonly ovate, basally somewhat attenu- ated, apically acute, the margins remotely crenate, glabrous, sparingly punctate, 4-7.5 cm long and 1.8-3 cm wide decreasing in si/e apicalh. Racemes glabrous, the axis elongating to ca 9 cm, the flowers loose; bracts o\ate, glabrous, deciduous, 0.8-1 mm long. LUmers white, the pedicels to 3 mm long; outer sepals margin- ally ciliate, the largest ovate, concave, 2.3-2.5 mm long, the smaller 2 ovate* to oblong and usually short-connate basally, sometimes free, 1-1.5 nnn long; wings ol)ovate to oblong-elliptic, apically rounded, basallv short-clawed, maroinallv cih- ate, persistent, 3.2-3.8 mm long; petals with upper pair oJKjvate, rounded, equal- ing the keel; keel 2.5-2.8 mm long with a crest 8-22-lobed (often irregularly forked and varying in si/e). Capsules widely oblong, glabrous, 4-5 mm long, 3.2-4 mm wide; seeds ovate, 4 mm long, densely pubescent with hairs copper colored at maturity; arils 2.5-3 mm long, the 2 lobes broadly obovate, appressed. Apparently endemic to the cloud forest above 2 500 ft in the vicinity of El Valle de Anton, Panama. 1969] FLORA OF PANAMA (Family 96. Polygahceae} 21 cocle: Ccrro Pajita, hills N of El Vallc de Anton, 1000-1200 m, common, Alien £r Allen 4170 (MO); Cerro Caracoral (vie of El Valle de Anton), elfin forest^ alt ca 1000 m, Duke & Buyer 15101 (MO, NY); mountains N of El Valle dc Anton, cloud forest, alt 2500-3000 ft, Lewis et al. 7723 (holotype MO; isotypes DUKE, K, UC, US). 13. Polygala jefensis W'.H. Lewis, Ann. Missouri Bot. Gard. 33: 367, 1969. Herb to shrub 2 m high CDuke 943 1), branched above with 2-3 branehlets per node, glabrous, angular. Leaves 4-5 in whorls; petioles usually 4-7 mm long, glabrous; blades elliptic, somewhat attenuated basally, acuminate apically, the margins remotely crenatc, glabrous, densely punctate, the more mature 4-5.5 cm long and 2-2.5 cm wide. Pujcemes terminal, glabrous, the axis to 7 cm long, the HowTrs ± loose; bracts ovatedanceolate, ciliate, deciduous, 0.8-1 mm long. Flowers greenish-white, the pedicels to 2 mm long; outer sepals marginally ciliate, the largest ovate, concave, 2 mm long, the smaller 2 ovate to oblong, 1 mm long, usually shortly connate basally, each with 2 large glands; wings broadly obovate, apically rounded, basally short-clawed, marginally ciliate, persistent, 3.5 mm long, 2.5 mm wide; petals with upper pair obovate, rounded, 3 mm long, glab- rous; keel 3 mm long with a crest 6-20-lobed (mostly 6). Capsules widely oblong to subglobose, glabrous, 4 mm long, 3-3.5 mm wide; seeds ovate, curved, 4 mm long, pubescent with hairs copper colored at maturity; arils 2.8-3 mm long, the 2 lobes oblong, appressed. Endemic to Cerro Jefe, Province of Panama; closely allied to P. inirdackiana. PANAMA: Cerro Jefe, alt 3400 ft, Bouche s.n. (holotype MO), Duke 9431 (MO), alt 2700-3000 ft, Tyson ct al 4397 (MO). 2. SECURIDACA Securidaca L., Syst. Nat. ed. 10, 1155, 1759, nom. cons., non L. (Sp. PL cd. 1, 707, 1753). Elsota Adanson, Fam. 2: 3 58, 1763, Woody vines, usually scandent shrubs, or trees. Leaves alternate, entire, short- petioled, generally with corneous peziziform stipular glands. Racemes terminal or axillary, often paniculate. Fhmers usually pink or purple; sepals 5, free, the outer 3 herbaceous, the inner 2 (wings) much larger, petal-like; petals 3, the lowest (keel) boat-shaped, clawed, with a plicate, fimbriate crest, the 2 upper adnate to the base of the staminal tube but distinct from the keel; stamens 8, their filaments connate nearly to the apex into a sheath split on the upper side, adnate to the keel and upper petals toward the base; anthers confluently 1 -celled, open- ing by a large introrse-apical pore; ovary unilocular through the abortion of the second loculc, with an appendage on one side which develops into the wing of the fruit, the style sickle-shaped, excavated apically, the stigma 2-lobed, approxi- mate, the ovule solitary, pendulous; disc a low^ fleshy ring at base of the ovary. Fruit a samara, the wing large, 1 -seeded; seed glabrous, cxarillate, endosperm wanting, the cotyledons thick-fleshy, oily. Perhaps 50 species in the tropics and subtropics excluding Australia; two species occur in Panama. [Vol. 56 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN a. Leaves acute or obtuse apically, rounded or cuncatc basally, puberulent, stri- ^ose, or glabrescent beneath; ilowers small; largest sepal from the outer whorl 3-4 mm long» the smaller two ca 2 mm long; wings glabrous on the outside; upper petals 5-7 mm long; keel with a wide many-plicate crest, 1-1.8 mm long 1 . S. diversifolia aa. Leaves obtuse to roiuided or sometimes emarginate apically, broadly rounded or truncate basally, softly and densely velvetypilose beneath; flowers large; largest sepal from the outer whorl 3.5-5 mm long, the smaller two 2.6-4 mm long; wings pubescent on the outside; petals 8-11 mm long; keel with a narrow few- plicate crest less than I mm long, sometimes obsolete 2. S. tcniiifolia 1. Securidaca diversifolia (L.) Blake in Standloy, Contr. U, S. Nat. Herb. 23^ 594, 1923. Polygala diversifolia L., Sp. Pi. 703, 1753. Scandent shrubs or woody vines; branches terete, puberulous, strigose or glabrescent. Leaves elliptic-oblong or oblong-o\ate to broadh o\ate or o\al, acute or obtuse apically, rounded or cuneate basally, 3,5-8.5 cm long, 1.5-5 cm wide (lea\'es of the flowering branches much smaller, usually oval or suborbicular), pergamaccous or chartaceous \\ith veins densely reticulate and prominulous on both sides, somewhat shining, puberulent or glabrous above, puberulent, strigose or glabrescent beneath, usualh more densely so along the veins; petioles strigose or puberulous, 2-7 mm long. Racemes loose or rather dense, terminal or axillary, sometimes branched, 1.5-8 cm long, 1.8-2.5 cm wide; peduncles 2 cm or less, the axis to 6 cm long, densely strigose or puberulous; bracts lanceolate to ovate, acuminate, pubescent, sometimes persistent, 1.5-3 mm long. Floirers pink or pur- ple, the pedicels 3-7 mm long, pubescent; outer sepals oval, rounded, puberulent except near the ciliate margin, the largest usually concave, 3-4 mm long, the smaller 2 ca 2 mm long; wings suborbicular to oval or o\cite, broadly rounded apically, usually emarginate, short-clawed basally, usual!) unequal, the margin sometimes ciliate, the outside glabrous (including the claw), 7-10 mm long, 6-8 mm wide; ui")pcr petals subspatulate, truncate apically, usually sparsely pub- erulent basally inside, rarely ciliate on the margin, 5-7 mm long; keel 7.5-10 mm long, as folded together 1.9-2.8 mm wide, the middle portion of the apex form- ing a wide, many-plicate, irregularly toothed or fimbriate crest, 1-1.8 mm long, the keel frequentK ciHate on the margin, sometimes the basal half sparsely puberulous on the outside or glabrous, usually puberulous inside on the saccate portion; basal half of staminal sheath pubescent, the free part of the filaments 2.5-3.2 mm long; o\'ar\- compressed-ellipsoidal, glabrous, 0.8-1 mm long, 0.6- 0.8 mm wide, slightly margined on one side, the appendage on the other side ovale, acute, densely puberulent, extending beyond the ovary, 7-9 nun long, the style 9-11 mm long, tlie stigma 2-lobed, approximate. Sajiiara (not seen at maturity) greenish or straw-colored, puberulous, 4-6 cm long, fruiting locule plump, elexated-reticidate, 5-8 mm long, margined on upper side, the margin 1-2.5 mm wide, prolonged bexond the fruiting locule and connate with tlie wing, usually with a triangular free tooth apically; wing obovate, narrower basally, 3,2-5 cm long. 3-4 mm wide (excluding the connate upper wing-margin of fruiting locule), 11-17 mm near middle (after Blake, N. Amer. Fl. 25: 373, 1924). 1969] FLORA OF PANAMA (Family 96. Polygalaceae') 23 Mexico to Peru and the Lesser Antilles. HOC AS DEL TORO: Changuinolii Valley, DiDiIap 577 (F, US), canal zoive: Barro Colorado T, shoreline SW of rairchild Point, Crout 4865 (MO), 48 73 (MO); vie of former town of Empire, Culebra Cut & vie, alt 80 m, Hunter & Allen 772 (GH, MO); Gamboa ]\i], Junes 311 (US), 3J2 (US); s. loe. JusepJi 462 (MO); Ancon, hospital grounds, alt 20-80 m, Mason 4 (F, GH, NY, US); Red lank to Pueblo Nuevo, Chiva-Chiva Trail, Piper 5126 (F, GH, NY, US); Barro Colorado I, Shattuck 798 (F, MO), 834 (MO); Anton Hill, moist thicket, Standley 26327 (US); nr Ft Randolph, Standley 28670 (US); Balboa, Standley 29266 (US); betu France Field & Catival, Standley 30i87 (US); Obispo, Standley 31747 (US); rd from Ft Sherman to Ft San Forenzo, Tyson & Blum 3757 (MO), 3758 (MO); nr Miraflorcs, White 77 (G!?, MO); Barro Colorado I, Wooduard & Vestal 335 (A, F, GH, MO), 453 (A, F, GH, MO), 556 (F, GH, MO), 6i8 (A, F, GH, MO); Ancon, hospital, Zetek Z-2526 (F). cocle: vie of Fl Valle, alt 600-1000 m, Allen 1634 (GH, MO, NY, US); mountains bevond Fa Pintada, alt 400-600 m. Hunter & Allen 543 (MO); Penonome & vie, Williams 110 (NY), 138 (NY), 411 (NY), 412 (NY), los SANTOS: Bahia Honda, Elmore 1117 (F). Panama: "La Victoria," \ic of La Jo\a, alt 200 m, Allen 4317 (MO); San Jose \, Camphell 9 (GH); 2 mi S of Cerro Azul, alt ca 700 ft, Dw\er 3531 (MO); summit of Cerro Campana, rain forest. Buyer et al. 4831 (MO); San Jose I, Erhwson 33 (GH, NY, US), Johnston 527 (GH), 572 (GH, MO, US), 903 (GH), 1067 (GH); Taboga I, forested slopes, Machride 2815 (US); Camino del Boti- cario, nr Chepo, alt 30-50 m, Pittier 4542 (F); vie of Juan Franco race track, nr Panama, moist thicket, Standley 27725 (US). ( 1879) from Panama (Lion-Hill railway stati cUvcrsifolia, a misidcntilication which Oort ( ) 678, ) notes as having boon frequent. Seeiiiann's S. puhesceJis ( 80, 1852), collected near Panama (City), will probably prove to be S. pub- esceus ft ovata DC. (Prodr. 1: 341, 1824), and if so then it should also be in- cluded under S. dirersifoUa (see Oort, loc. cit. 680). Finally, S. tomcntosa of Ilemsley (loc. cit.), but not S. tanientosa St.-Hil. (Fl. Bras. Mcrid. 2: 68, t. 96, 1829), collected near Panama City, may be included here though this is not certain. 2. Seouridaca tenuifolia Chodat, Bull. Herb. Boissier 3: 54 5, 1895. Ehotu chrrsotricha Rilcv, Kcw Rull. 1927: 1 19, 1927. E. tenuifolia (Chodat) Sandvvith, Kcw Bull. 1929: 79, 1929. Scandent sJiriihs or woody vines; brandies terete, softly pilose. Leaves oval, ()l)long-ovate or ovate, obtuse to rounded, sometimes emarginate apically, broadly rounded or truncate basally, 3.5-9.5 cm long, 2-3.5('4) cm wide, chartaceous, veins densely reticulate and prominulous on both sides, somewhat shining, sparse- ly pilose (usually more densely on margin) or glabrescent above, softly and densely vclvcty-pilose beneath, especially along the veins; petioles pilose, 2-4 mm long. Race)}ies loose or somewhat dense, terminal; peduncles 1 cm or less long, the axis 2.2-5 cm long, pilose; bracts lanceolate, acute to acuminate, pilose, decid- uous, 1.8-2 mm long. Fhmers pink; pedicels short^pilose, 5-8 mm long; outer sepals o\al, rarely suborbicular, rounded, densely pilose on the outside except near llie ciliate margin, the largest 3.5-5 mm long, the smaller two 2.6-4 mm long; wings suborbicular, broadly rounded apically, usually emarginate, short- clawed basally, the margin ciliate, tlie outside pubescent (including the claw), 9-12 mm long, 6-9 nmi wide; upper petals obovate or obovate-spatulate, rounded, 24 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 usually undulate apically, attenuate and gibbous toward the base, glabrous, 8-11 mm long; keel 9.5-13 mm long, as folded together 5-6 mm wide, the apex form- ing a narrow, few-plicate, minutely toothed crest less than 1 mm long, some- times obsolete, the keel ciliate on the margin, sometimes sparsely puberulent on the outside, puberulent inside on ca the basal half of the saccate portion; staminal sheath pubescent basally, the free portion of the lllaments 2-2.8 mm long; ovary compressed-ellipsoidal, densely pubescent, 1.2-1.8 mm long, 0.8-1 mm wide, the style 7.5-11 mm long, the stigma 2-lobed, approximate. Samara densely short- pilose, 4.5-7 cm long, plump, elevated-reticulate, 7-11 nmi long; wing obovate, narrower basally, 4-6 cm long, 14-22 mm wide near middle. Panama and Trinidad. (ANAL zone: Ciirundu, Thuntlor Hill, McDmiicl S186 (MO); Las Sabanas, Pitticr 6642 (US); betw I't Clayton & Coro/al, aloriK old Las Cruccs Trail, moist thickft' Stand- ley 29/09 (US); vie of T.io Cocoli, Hd K-9, Stem ei al. 301 (MO); Corozal, Stevens 114 (US); Barro Colorado I, Wooduan! & Vestal SOS (A, MO), colon: Santa Rita lUdgv, loHKing rd 19 km from Transisthmian Hwy, Dwyer 8576 (MO), darien: nr Piinta Garachinc, thorn iurest, Duke 1074 (US); vie of Id Real, Rio Tuira, ed^e of rd on Don Pablo Othon's farm. Stem et al. 800 (GH, MO, US). Panama: vie of El Llano, Dnhe 55 7 2 (GH, MO); nr Rio Tapia Juan Diaz region, roadside thieket, Maxon t llaney 6749 (GH, NY, US); Camino del Hotieario, nr Chepo, alt 30-50 m, rUtier 4542 (US); Old Panama, Riley 14S (t>iie of Llsota ehrysutrieha; MO, US); Ixfw Matias Hernandez & Juan Diaz, Staiidley 32034 (US), vi raguas: alon^ buy nr Rio Tabasara, Wundsnn et at. 466 (MO), i-kovlnce cnknown: s. loe., Cyfisehaeh s.)i. in 1857 (MO). Our specimens of S. teiiiiifolia had been misidentified as S. coriacea Bonpl. 3. IMONNINA Moiinina lluiz & Pavon, Syst. Veg. Fl. Peru. Chile. 169, 1798. Hcrhs, shrubs, or small trees. ]. eaves alternate, entire, short-petiolate, stipulate or with stipular glands. Flouers pm-ple or blue; sepals 5, the outer 3 herbaceous, free, the inner 2 (wings) lower (keel) concave; petals 3, the adnate below to the staminal sheath; stamens 8, \ery rarely 6, their filaments con- nate nearly to apex into a sheath split on the upper side, the anthers confiuently 1 -eel led, opening by a large introrse apical pore; ovary 1-locular, the style genicu- late, the stigma with 2 dissimilar lobes \Aith ihe outer (lower) ± acute and the inner (upper) tubercled, the ovule 1, pendidous; disc reduced to a gland at base of the ovary. Dntpe with a fleshy coat, the surface rugose, marginate, 1- seeded; seeds glabrous, exarillate, the cotyledons thick. About 150 species from the southern U.S. and Mexico to Argentina and Chile; predominantly northwestern South American. A comprehensive treatment of the genus is needed and will idtimately be provided by Ferreyra (for Peru see Jour. Arnold Arb. 27: 123-167, 1946; for Fcuador see Moydia 16: 193-226, 1953; for Colombia see Smithsonian Misc. Coll. 121: 1-59, 1953; for Vene- zuela see Hrittonia 9: 9-18, 1957). Two species are recognized for Panama. a. Racemes slightly eomose toward the apex; braets lance-subulate, 4.5-5 mm long; leaves elliptic to oval, cuneate-attenuate or obtuse basally, 6-13 em long, 2.5-7 cm wide; lower sepals of the outer whorl without a lateral lobe 1. M. srlratica 1969] FLORA OF PANAMA (Family 96. Polygalaceae') 2 5 aa. Racemes not at all comose; bracts triangular-ovate, acute or obtuse, 1.4-2.2 mm long; leaves oblanceolate to obovate elliptic, acute to acuminate basally, 3-10 cm long, 1-3.5 cm wide; lower sepals of the outer whorl usually with a small lateral lobe 2. M. xalapensis 1. Monnina sylvalica Schlecht. ik Cham., Linnaea 3: 231, 1830. — Fig. 2. M. crhputu Blake, N. Amer. Fl. 25: 377, 1924. Shrubs or herbs to 3 m high; branches striate, incurved-puberulous or gla- brous. Leaves elHptic, elhptic-ovate, or oval, acute or acuminate apically, cuneate to euneate-attenuate or obtuse basally, 6-13 em long, 2.5-7 cm wide, usually glabrous, sometimes pubescent on both sides, the eosta prominulous beneath with 4-6 pairs of conspicuous lateral veins; petioles 3-7 mm long, glabrous or pubes- cent. Racemes cylindric, somewhat dense, obtuse, slightly comose toward the apex, to 9 cm long, 8-18 mm wide; peduncles to 5,5 cm long, several or numer- ous, simple or branched, terminal or axillary, the axis to 16 cm long, pubescent; bracts lance-subulate, attenuate, ciliate, deciduous, 4.5-5 mm long. Flowers purple or blue; pedicels 1-2 mm long, pubescent; outer sepals ovate to oblong-ovate, obtuse, rarely acute, ciliate on the margins, sometimes short-pubescent or glabres- cent on the outside, the largest 2.5-3 mm long, sometimes concave, the smaller two 1.5-2.5 mm long; wings suborbicular to suborbicular-obovate, hooded, broadly rounded apically, rounded or rounded-cuncate basally, rarely ciliate on the margin, occasionally sparsely pubescent within, 3.5-5 mm long, 3-4.2 mm wide; keel orbicular, usually plicate laterally, emarginate apically, 4-7 mm long, 5-7.8 mm wide, rarely ciliate on the lower part of the margin, sparsely pubescent within or glabrous; upper petals yellow, spatulale, pubescent within, sparsely pubescent or glabrous on the outside, 3.8-5.4 mm long; staminal sheath pilose apically, the free part of the filaments 0.7-1.4 mm long; ovary ellipsoidal, 1.1-1.8 mm long, 0.9-1.1 mm wide, usually glabrous, rarely pubescent, the style usually geniculate, 2.3-3 mm long, the stigma 2-lobed, the outer acute, the inner tubercled. Drupe ovoid, obtuse or emarginate, conspicuously rugose-reticulate, glabrous, 5-8 mm long, 3.5-4.5 mm wide. Mexico to Panama at higher elevations. CHiRiQui: llanos on slopes of Volcan de Chiriqui Viejo & along Rio Chiriqui Viejo, alt 1200 m, Allen 984 (MO); trail from Paso Ancho to Monte Lirio, upper valley of Rio Chiriqui Viejo, alt 1500 2000 m, AUcn 1576 (MO); vie of Boquete, lumber rd into hills E of Rio Caldera, Alien 4662 (MO); Bajo Mono-Rohalo Trail, W slopes of Cerro Hor- queta, alt 5000-7000 ft, Alien 4791 (MO); Cerro Horqueta, alt ca 1500 m, DuJze 13653 (MO); Fred Collins Finca, edge of coffee plantation, alt 6000 ft, Ebinger 661 (MO); vie of Boquete, from Boquete to 3 miles N, second growth, cultivated areas, roadsides, alt 3300-4200 ft, Leivis ct al 412 (MO), 473 (MO); vie of Cerro Punta, cloud forest & disturbed edge, alt 6800 ft, Rid^way & Solis 2408 (F, MO); Cerro Horqueta, cloud for- est, alt 6500 ft, von Hagcn Gr von Hagen 2074 (MO); betw Concepcion & El Volcan, White 311 (MO); vie of Bajo Mona & Quebrada Chiquero, alt 1500 m, Woodson & Schery 540 (MO), cocle: betw Cerro Pilon & Fl Valle de Anton, cloud forest, alt 700- 900 m, Duke & Dwyer 13900 (GH, MO); El Valle de Anton at foot of Cerro Pilon, cloud forest, alt ca 2000 ft, Divycr Gr Correa 7959 (GH, MO); summit of Cerro Pilon, above El Valle de Anton, rain forest, alt ca 2700 ft, Dwyer et al. 4500 (MO); mountains beyond La Pintada, alt 400-600 m, Hunter & Allen 545 (MO); mountains N of El Valle de Anton, cloud forest, alt 2500-3000 ft, Leivis et al 1735 (COL, MO, UC, US); El Valle de Anton, edge of cloud forest & roadside, alt 1000-2000 ft, Lewis et al 2539 (DUKE, 26 I VNNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 I A ^ ^^^ # G F C D E Fig. 2. Monnina syliatica Schlccht. & C:ham.; A, habit (X '/O; B, inflori-sccncc (XI); C, sepals (X 31/2); D, kc'cl (X 3'/2); E, stamens and upper petals (X iVz'); F, pistil (X 31/2); C, fruit (X 31/2). After Leu is ct al 2S39 (MO). K, MO, NV); Cerro Pilon, alt 2000 ft, /'o;7er ct al 4603 (MO), los santos: Cerro Pilon, rain forest, alt 2 700 ft, Dn ycr & Lulhithin 861 2 (MO), 2. Monnina xalapen^is II.B.K., Nov. Gen. Sp, PI. o: 414, 1823. Al. lathcpala Blake, N. Anier. Fl. 23: 3 76, 1924. Herbs, shrubs or even small trees to 6 m high; branches striate, shortstrigose or becoming ± glabrescent. Leaves oblanceolate, elhptic, or obovate-elliptic, acute to acuminate, sometimes cuspidate or falcate apically, acute to acuminate basally, 3-10 cm long, 1-3.5 cm \vide, membranaceous to chartaceous, rather sparsely pubescent on both sides, usually more densely so along the costa and yeins b "^^ c ncath, sonictiincs glabrescent, the costa prominulous beneath with 5-7 pairs of lateral veins; petioles 3-12 nini long, pubescent. Racc'))U's not at all comose, to 1969] FLOKA OF PANAMA (Family 96, Polygahceae^ 27 3.5 cm long and 1.5 cm wide, terminal or axillary; peduncles to 1.5 cm long, the axis 10 cm or less, conspicuously pubescent; bracts triangular-ovate, acute or obtuse, ciliate and pubescent, deciduous, 1.4-2.2 mm long. Flowers purple or blue; pedicels 1-2.4 mm long, pubescent; outer sepals free, triangular- or oblong- to suborbicular-ovate, acute to obtuse, sometimes mucronate at the obtuse apex, margin ciliate, short-pubescent or glabrescent outside, 2.5-3.2 mm long, usually all 3 ca equal, sometimes the upper sepal slightly longer and concave, usually the lower 2 with a small lateral lobe; wings suborbicular, hooded, broadly rounded apically, cuneate basally, usually cihate on the margin, sparsely pubescent within, 5-6.2 mm long, 4.5-6 mm wide; keel orbicular, plicate laterally, emarginate apically, 5.5-6.5 mm long, 5-7.8 mm wide, rarely ciliate on the lower part of the margin, glabrous within; upper petals yellow, spatulate, pubescent within, sparsely pubescent or glabrous on the outside, 3.4-4.8 mm long; staminal sheath pilose apically, the free part of the hlaments 0.8-1.2 mm long; ovary ellipsoidal to ovoid, 1.3-2 mm long, 0.8-1 mm wide, pubescent or glabrous, the style geniculate, 2.5- 3.5 mm long, the stigma 2-lobed, the outer acute, the inner tubercled. Drupe ellipsoid-ovoid, obtuse, rugose-reticulate, glabrous, 6-7.5 mm long, 3.5-4 mm wide. Mexico to Panama at higher elevations. CHiRiyui: trail from Bambito to Cerro Piinta, 1400-2300 m, Allen 318 (MO) (as M. pittieri Chodat); summit & S\V face of Ccrro Copctc, alt 9000 ft, Allen 4899 (MO); Volcan dc Chiriqui, alt 7000 ft, Daiidson 889 (A, MO); Alto dc Cuesta, Pitticr 3119 (holotypc of M. latiscpalu US; isotypc NY); valley of upper Rio Chiriqui Viejo, vie of Monte Lirio, alt 1300-1900 m, Seihert 306 (MO); Rio Chiriqui Viejo valley, White 49 (MO); Chiriqui, valley of upper Rio Chiriqui Viejo, in open sunlight, White & White 26 (A, MO, NY) (as M. pittieri Chodat?); Finea Lerida to Pena Blanca, alt 1750-2000 m, Woodson Sr Schery 304 (MO); Casita Alta to Ccrro Copete, ah 2300-3300 m, Woodson & Sehery 338 (GH, MO); Potrero Muleto to summit, Volcan dc Chiriqui, 3500-4000 m, Woodson & Sehery 441 (GH, MO); vie of Casita Alta, Volcan de Chiriqui, alt 1500-2000 m, Woodson et al 802 (A, MO), province unknown: Sierra Madre, Seemann s.n. (GH). We suspect that A/, xalapoisis and AL sylvatica hybridize; one collection from Cerro Horqueta at 5000-5800 ft [Dwyer et al 43S (MO, UC, US)] is inter- mediate between these species and is considered a putative hybrid. 4. MOUTABI'A Moutabea Aublet, Hist. PI. Gui. Fr. 679, t. 247, 1775. Erect or seandent shrubs or small trees. Leaves alternate, petiolate, the blades thick and coriaceous, usually glabrous. Racemes short, axillary. Flouers white or yellow; calyx adnate with the corolla into a tube, the sepals 5, ± equal, the petals 5, subequal with the lower shgluly boat-shaped; stamens 8, joined into 2 groups of 4 on the margin of an obhque sheatli which is adnate to the corolla-tube, the anthers sessile; ovary 4- or rarely 2- to 5-locular, surrounded by a disc, the ovules 1, pendulous, the style hliform, slightly dilated apically. Berry (drupaceous?) glo- bose, indchiscent, 2-5-seedcd, edible; seeds shining in the fleshy pericarp, without aril or endosperm. Perhaps 10 species of tropical South America and Panama; only one in Panama, a hrst record for the genus in North America. [Vol. 56 28 VNNALS OF THE MISSOURI BOTANICAL GARDEN 1. Moutabea art". longifoHa Pocppig & Fndl., Nov. Gen. Sp. IM. 2: 62, t. 186, 1838. SJinihs 2.1 111 tall, the branches with dark, straight hairs. I. eaves with short, thick petioles 1 cm long; blades oblong to obovate, glabrous, prominently reticulate- veined above and below, ca 17 cm long and ca 7 cm at widest point 12 cm from the petioles. Racemes terminal and axillary, several per axis, the peduncles to 2.5 cm long, sparsely puberulent with numerous puberulent o\ate bracts 2 mm long. Flojvers white (drying almost black), tubular; tube 1 cm long; sepals 6-8 mm long, pubescent within; petals 8-10 mm long, linear-spatulate, pubescent within; 8 sessile anthers grouped 4 on each margin of the sheath, the margin 2-3 mm long. Priiit unknown. cocu:: Bisniark ahovr Pcnonome, Williams 565 (US). Allied to and perhaps conspecific with M. longifolia of South America which, however, has more lanceolate leaves. A second specimen collected by W illiams f966J in Darien (Cana & vie, US) may also prove to be a species of Montahea. The material is sterile but the leaves closely approximate those of the original plate of M. Ion 21 folia. Index of Latin Names NunibcTs in bold fuco t>pc reefer to tk'scriptions; numbers in roman iyyc refer to synonyms; numbers witb dagger (f) refer to names incidentally mentioned. Flsota 21 chrysotrica 2??, 24 1 tcnuifolia 23 Monnina 24 crispata 2 5 latisepala 26, 27 1 pittieri 27t sylvatiea 25, 27t xalapensis 26, 27t Moutabea 9t, 27, 28t aff. lonRifolia 28 lon^ifolia 28t Polygala 9t, 9 anyustifolia I I aparinoides 19 asix'ruloitles 19t, 19 brizoides 11 costaricensis 12t divcrsifolia 22 durandii 12t fendleri var, hctcr- othrix 12 hygrophila 15 jefensis 21 leptoeaulis 13 longicaulis 16, 18t monticola 12t panamensis 12 panieulata 13t, 14 timoutou 18 variabilis 18 wurdackiana 20, 21 f Seeuridaca 21 coriacea 24 1 diversifolia 22, 23t pubescens 23t — /? ovata 23t tenuifolia 23, 24t tomentosa 23 f volubilis 23 1 FLORA OF PANAMA BY Robert E. Woodson, Jr. and Robert W. Schery AND Collaborators Part VIII Family 161. MENYANTHACEAE' I5Y Thomas S. Elias- St. Louis L'nivcrsit)-, St. Louis, Missouri Herbs, aquatic or semiaqiiatic. Leaves alternate, entire, simple or trifoliolatc, often orbicular and peltate, petiole sheathing at tlie base, exstipulatc. Inflores- cences racemose, solitary, paired, or fasciculate, or in many-flowered cymes, panicles, or involucrate heads. I'hniers bisexual, regular; calyx 5-merous, free or fused basally; corolla 5-merous, fused basally to form a short tube, the lobes valvate or induplicate-valvate, the margins and/or interior often fimbriate or barbate; stamens 5, alternate with the petals and inserted near the base of the corolla tube, the anthers 2-locular, sagittate, \ersatile; hypogynous nectaries usually present; pistil 1, the ovary superior, 1-locular, with 2 parietal placentas, ovules CO, the style simple, the stigma bifid. Fruit a capsule, 2- or 4-valved, rarely Heshy and indehiscent; seeds few to numerous, sometimes winged, endosperm copious, embryo small. — x = 9, 17. A small family of five genera and ca 30-3 5 species occurring in temperate and tropical regions of both hemispheres. The genera have been included by earlier workers (Bentham & Hooker, Gen. PI. 2: 819, 1876; Gilg in Engler & PrantI, Nat. Pllan/enfam. 4(2): 62, 1876) as a tribe or subfamily of the Gentianaceae. later workers (Hutchinson, Fam. Fl. PI. 1: 451, ed. 2, 1959; Cronquist, Fvol. & Class. Fl. PI. 288-289, 1968) have maintained them as a distinct family, separated from the Gentianaceae by the alternate lea\es and the valvate or induplicate-valvate aestivation. Onlv the genus NxnipJioides is found in Panama. 1. NYMPHOIDFS Nymphoides Scguier, PL Veron. 3: 121, 1754. Lirnnanthcmitm S. G. Gmcl., Novi Comment. Acad. Sci. Imp. Pctrop. 14(1): 5 25, 1770. Herhs, aquatic, perennial; stems petiole-like. Leaves simple, usually broadly ovate or orbicular, peltate, cordate at tbc base, floating. Flowers wbite or yellow; calyx basally fused; corolla broadly campanulate or almost rotate, the lobes fimbri- ate; stamens included, the anthers ovate to linear; ovary ellipsoid, the style short, ^Assisted by Xational Science Foundation Grant No. GB-5674. -Present address: Arnold Arboretum of Harvard Univ., 22 Divinity Ave., Cambridge, Mass. 02138. Ann. Missouri Box. Card. 56(1): 29-32, 1969. 30 i \NNALS ()!• THE MISSOURI BOTANICAL GARDEN [Vol. 56 Fig. L N) Dipljoides liiiJiiholdtiiuiu))! (II.R.K.) O. Kiintzc: A, habit (X V2); B, lloucr (X 2'/^); C, two corolhi lobes and a stamen (X 5); 1), fruit \\ iib niarccsccnt cahx and style CX 5). After Croat H2n (MO). 1969] FLORA OF PANAMA (Family 161. Menyanthaceae^ 31 the stigmas usually broad. Capsules indchisccnt or eventually rupturing irreg- ularly, firm-walled; seeds many, unwinged, the seed coat hard. — x = 9. A small genus of ca 20 species chiefly of tropical and subtropical areas in the New and Old W^orlds. Only one species is known from Panama, the common N. hiimholdtianjim. Nyinphoides huniboldlianum (ILBK.) O. Kunt/e, Rev. Gen. PL 2: 429, 1891. (^^s Nymphodes^.^Vi^. 1. Villarsia humhohltianuni H.B.K., Nov. Gen. Sp, PI, 3: 187, 1819. Limnanthemum humholdiianum (H.B.K.) Griscl)., Gen. Sp. Gent. 347, 1839. Herbs aquatic, from submerged rhi/omes; stems 8-150 cm long, to 6 mm in diam, ascending, green, glandular (?) dotted near the apex, succulent, weak, vertically septate with several continuous air spaces, the stems with a single terminal leaf or usually branching near the apex to gi\e rise to an inflorescence and additional leaves. Leaves orbicular to subovate, cordate at the base, often deeply so, obtuse at the apex, often becoming emarginate on the larger leaves, 2-18 cm in diam, succulent, green above, dark reddish-violet beneath, coriaceous. hiflorescejices of man)' fascicled flowers, apju^aring umbellate, axillary. Flowers white, 1-8 cm long, pecHcellate, tlie pedicels (mature) 3-7 cm long, succulent, glandular (?) dotted, erect to suberect; calyx-lobes lanceolate, acute to short apiculate at the apex, the margins scarious, 4.5-8 mm long; corolla 1-1.8 cm long, the lobes o\'ate to ovate-oblong, acute at the apex, very membranous, ± transparent when dried, 8-12 mm long, conspicuously fimbriate, the hairs 1-3 mm long, the lobes fused near the base; stamens introrse, 5-7 mm long, the filaments ca 3-5 mm long, ± flattened, the anthers 2 nun long, sagittate and spreading at the base; pistil 9-12 nmi long, the o\ary ellipsoid, sessile, 3-5 mm long, the style cylindrical, the stigma lobes to 1 mm long, flattened. Capsules indchisccnt and eventually rupturing irreguarly, ellipsoid, 4-6 mm long, ca 10-1 8-seeded the persistent style rostrate; seeds orbicular, slightly flattened dorsiventrally, 1-1.2 nun long, shiny. CANAL zone: Barro Colorado I, shore line, (juict water, Croaf 8215 (MO); Rio Chagres, Fairchild 2045 (MO); Madden Lake, Woodsou & Schcry 958 (MO), cocle: El Vallc de Anton & vie, in ])oss, Scihcrt 440 (MO); hetw Las Margaritas & El Vallc de Anton, in bog, \Voods()}i ct al. 7 753 (MO), panama: along rd bctw Panama 8c Chepo, in pools in savannas, Dodge ct ah 16711 (MO); Sabanas, nr Chepo, in shallow pond, Hunter & Allen 66 (MO), Hunter & Steyermark s.n. (MO); wet savanna E of Pacora, in lake, depth 1 m, Woodson et ah 752 (MO). A common aquatic herb in the West Indies and Central and South America, N. hiiiiiholdtiainuii appears to be restricted to quiet and non-running waters. When growing in shallow bogs or marshes, the stems are considerably shortened. The length of the stem is apparently determined by the depth of the water in which it is groA\ing. In marsh habitats in Panama the stems are ca 8-15 cm long (Seihert 440, Woodsoji et ah 7 753 j, while those growing in lakes ha\'e been ob- served to be as lono as 1.5 m (Croat 821 S). In addition to sliorter stems, the marsh or bog plants of this species ha\'e smaller leaxes than those plants which [Vol. 56 32 / VNNALS or THE MISSOLHI BOTANICAL GARDEN arc found in lakes. The small-lea\ed populations were gi\en varietal status (var. panifolini}!^ by Crisebacli (Gen. Sp. Gent. 347, 1839). Ornduff, in a treatment of the Meso-Anieriean and West Indian species of Nyniphoide^, suo;ocsts that N. hiDJiholdtiana is conspccific with the OldW'orld N. iudica (L.) O. Kunt/c. He makes his case \ery strongly, but I prefer to hold to the later name until the taxonomy of the N. hidica complex is completely worked out. Index of Latin Names Numbers in bold face type refer to descriptions; numbers in roniiui lype refer to synonyms; iiuin]>ers with d^i^^er (t) refer to niunes incidentally mentioned. Gentianaceac 29t Limnanthcmiim 29 humboldtianum ^ I Nymphoidcs 29t, 29 humboldtianum 3 If, 31 — var. parvifohum 32t indiea 321 Villarsia humboldtianum 31 FLORA OF PANAMA BY Robert E. Woodson, Jr. and Robert W. Schery AND Collaborators Part IX FAMILY 167. BORAGINACEAE^ BY Joan \V. Nowicke Missouri Botanical Garden and Department of Botany, Washington University, St. Louis, Missouri Herbs, shrubs or trees, rarely lianas, frequently scabrous, the hairs simple, uniseriate, or stellate, often with a hardened base, or sometimes glabrous. Leaves alternate or sometimes opposite, simple, generally petiolate, the blade pinnately- nerved, entire or rarely dentate. Inflorescences cymes, generally helicoid or scorpioid, paniculate and open, spike-like, or glomerate, or rarely the flowers sol- itary, the bracts seemingly absent. Flouers perfect or functionally unisexual, acti- morphic or rarely zygomorpbic; bractcoles generally absent; sepals 5, rarely 4, free or connate basally, sometimes irregular; petals 5, united, salverform, funnel- form, or campanulate, the lobes distinct or obscure, the tube sometimes with folds or appendages in the throat; stamens 5, functional or not, epipetalous, alternate with the corolla lobes, the filaments simple or with dorsal appendages, glabrous or fimbriolate basally, the anthers linear, introrse, dehiscing longitudinally; gynoecium syncarpous, 2-carpellate, ovary superior, bilocular and becoming falsely 4-locular, placentation axile, ovules 4, or fewer by abortion, anatropous, style one, gynobasic or terminal, simple, cleft, or twice cleft, stigmas 1, 2, or 4. Fruit 4 nutlets, a 1-4-secded nut, or a drupe; seeds generally without endosperm. x = 4, 10, 13. A widely-distributed family of about 100 genera and 2000 species. Seven genera are found in Panama. The Boraginaccae are of little economic importance; a few are cultivated as ornamentals, and some species of Cordia, whose wood varies in color and weight, arc utilized as a source of lumber. Useful references: Johnston, I. M. Studies in the Horaginaceae, XV. Notes on some Mexican and Central American species of Cordia. Jour. Arnold Arb. 21: 336-355, 1940. — . Studies in the Boraginaceae, XVH. A. Cordia section Varronia in Mexico and Central America, loc. cit. 30: 85-104, 1949a. — . Studies in the Boraginaceae, XVI II. Boraginaceae of the south- ern West Indies, loc. cit. 30: 11M38, 1949b. . Botany of San Jose Island. Sargentia 7: 1-306, 1949c. Key to Genera a. Herbs. b. Nutlets with hooked spines; style gynobasic. ^ Assisted by National Science Foundation Grant No. GB-5674. Ann. Missouri Bot. Card. 56(1): 33-69, 1969. [\'()L. 56 34 ANNALS OI- THE MISSOL'RI BOTANICAL C;APJ)LN c. Fruit distinctly 4 l()t)ccl, the nutlets attached at the apices; \eins tending to form a network in the leaves 1. Cyno'^Iossuvi cc. Fruit pyramidal, the nutlets attached medially; \eins not forming a net- work in the leaves 2. Huckclia bb. Nutlets strigose or glabrous but without hooked spines; style terminal. ^. Ucliotropiiim aa. Shrubs, trees, or sometimes vines. d. Styles and stigmas simple 4. Tour uc for da dd. Styles once- or twice-di\ itied, the stigmas 2 or 4. e. Stigmas 2, capitate or peltate. f. Calyx closed in bud; leaves entire 5. Bourrcriu ff . Calyx open in bud; leaves serrate 6. Ehretia ee. Stigmas 4, slender or clavate 7. Cordia scor- 1. CYNOGLOSSLM Cynoglossum L., Sp. PL 134, 1753. Herbs, mostly perennial, sonietinies annual or biennial, slender to very robust, glabrous to eoarsely pubescent. Leaves alternate, simple, pinnately-nerved, entire, the basal ones long-petiolate. Inflorescences racemes or irregular panicles, pioid, the bracts mostly not apparent. Vlmvers perfect, actinomorphic; calyx of 5 sepals, ± connate basally, enlarging somewhat in fruit; corolla salverform, funnclform, or ± campanulate, blue, violet, or reddish, 5-lobed, the lobes ± spreading, 5 faucal appendages generally A\elLdevcloped; stamens 5, included or barely cxserted at the corolla tliroat, subsessile or the filaments very short, the anthers oblong; ovary 4-lobcd, distinctly so, the gynobasc disc-like, the style gynobasic, the stigma 1 and ± capitate. fn 4 cm wide; inflorescences > 10 cm long 1 . //. indictiin aa. Decumbent herbs; leaves > 1.5 cm wide; inflorescences < 9 cm long. b. Plants glabrous; leaves somewhat succulent, usually glaucous. ... 2. U. curassaiicum bb. Plants pubescent, the hairs white; leaves not succulent nor glaucous. 3. //. procumhcns 1. Heliotro]uuni inclicuni L., Sp. PI. 130, 17 53. Heliophytum iruUcum (L.) DC, Prodr. 9: 5 56, 184 5. Herbs, annual, coarse and weedy, to 1.5 m, the stems with short and longer hairs. Leaves alternate, ovate, or sometimes ± deltoid, acute, finely to coarsely undulate, the bases obtuse to rounded and long-attenuate, to 1 1 cm long and 6.5 cm wide, sparsely pubescent on both surfaces, the veins prominent beneath; petioles ± winged or indistinct, to 8 cm long. Inflorescences spicate, scorpioid, to ca 28 cm long, mostly terminal, the bracts absent. Flowers sessile; calyx of 5 free sepals, subulate, ca 1.5-2 mm long; corolla salverform, blue, violet, or rarely white, the tube ca 3-4 mm long, each lobe ca 1 mm long; stamens ± sessile, the anthers ca 0.6-0.8 mm long; ovary weakly 4-lobed^ the style somewhat stout, the stigma conical. Fruit deeply 4-lobed, the lobes horizontally divergent, sep- arating into 4 nutlets at maturity, the nutlets angulate, glabrous to sparsely strigose, 2-3 mm long. A roadside weed, widely distributed in Panama and in the American Tropics. BOCAs DEL TORO: vic of Chiriqui Lagoon, Old Bank I, von Wedel 1968 (MO), canal ZONE: s. loc., Eppehhei777er s.n. (F). cocle: 12 mi NE of Penonome, Lewis ct al. 1524 (MO), darien: vic of El Real, Rio Tuira, Stern et al. 452 (MO), los santos: Rio To- nosi, vic of Tonosi, Lewis et al. 155H (MO), panama: roadside betw Chepo & wharf, Dodge s.n. (F); San Miguel I, Duke 10937 (MO); Tahoga I, MacBriile 2792 (F, MO). VERAGUAS: 2 mi S of Canazas, Tyson 3 725 (MO). A medicinal tea is made by boiling the leaves. [Vol. 56 38 ANNALS OF THE MISSOUIU BOTANICAL GARDEN 2. Heliotropiuni curassavicuni L., Sp, PL 130, 1753. Herbs, prostrate, the stems 0.5 m long or longer, glabrous, suceulent, some- times glaucous on the younger parts. Leaves alternate or somewhat fasciculate, oblanceolate, lanceolate, or spatulate, obtuse, entire, the bases attenuate, to 4 cm long and 1 cm wide, glabrous, succulent, sonu^what glaucous; petioles indistinct. Inflorescences ± spikes, unilateral, -mostly terminal, ca 2-5 cm long, the bracts absent. Fhnvers sessile, ± actinomorphic; calyx of 5 sepals, slightly connate basally, elongate-deltoid, ca 1.2-1.5 mm long; corolla salverform, w^hite, the tube somewhat saccate, ca 1.2 mm long, each lobe ca 0.8 mm long; stamens 5, ± sessile, the anthers ca 0.6-0.7 mm long; ovary 4-lobed, the disk obscure, the style absent, the stigma conical, annular-pubescent at the widened base, fruit 4-lobcd, separating into 4 nutlets at maturity, the nutlets glabrous, WTdge-shaped, ca 1.6- 1 .7 mm long. W idely distributed in the warmer regions of America, particularly along coasts and inland on saline soils. LOS SANTOS: Monagrc Beach, Dnyer 4177 (MO), Tyson ci ul. 3023 (MO). 3. Heliotropiuni procunihens Mill., Card. Diet, ed 8, no. 10, 1768. — Fig. 2. //. inundatiim Sw., Prodr. 40, 1788; non L., Sp. PI. 130, 1753. Herbs, prostrate, somewhat wiry, the stems 15-20 cm long, pubescent, the hairs white. Leaves alternate, ± elliptic to ovate, bluntly acute, entire, the bases obtuse, to 1.5 cm long and 0.8 cm wide, pubescent on both surfaces, the hairs Willi enlarged bases; petioles ca 3-5 nun long. Inflorescences spike-like, unilateral, to 7 cm long, the bracts absent. Flouers ± sessile; calyx of 5 sepals, shghtly connate basally, the lobes lanceolate, subequal, ca 1 mm long, pubescent; corolla weakly salverform, white, the tube ca 0.8 nun long, sparsely pubescent, the lobes ± unequal, erect, and ca 0.2-0.4 mm long; stamens 5, ± sessile, the anthers ca 0.2 mm long; ovary 4-l()bed, the style absent, the stigma conic. Fruit 4'lobed, separating into 4 nutlets at maturity, the nutlets wedge-shaped, strigose, ca 0.4- 0.5 mm long. Widely distributed in tlie warmer parts of America. LOS SANTOS: Iaio ToHosl, vic of Tonosi, Lewis ct ul 1575 (MO). 4. TOURNl lOlVriA Tournefortia L., Sp. PI. 140, 1753, Shriibs, sometimes decumbent, vines, or small trees, glabrous to densely pubes- cent. Leaves alternate, sometimes opposite, simple, pinnately nerved; sessile or petiolatc. Inflorescences cymose, scorpioid, sojnctimes borne in irregular panicles, the bracts seemingly absent. Flowers perfect, ± actinomorphic, sessile or shortly pedicellate; calyx of 5 sepals, free or weakly connate basally, lanceolate to ovate, ± glabrous to densely strigose; corolla \\hite, yellow, or ± green, salverform or funnelform, glabrous or strigose, the tube cylindrical, the limb usually 5-lobed and spreading, the lobes frequendy keeled; stamens (4-) 5, rarely more, borne on the corolla throat, included, the fdaments very short, the anthers elongate, free, 1969] FLORA OF PANAMA (Family 167. Boraginaceae~) 39 Fig. 2. Heliotropium procuinhcns Mill.: A, habit (caX%); B, flower (caXlS). After Lewis et at. 1575 (MO). 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 or ± connate apically; ovary 2-carpellate, each carpel with two ovules, disc ± distinct, sometimes cup-shaped, style long and slender or ± absent, the stigma peltate or conic. Fruit Heshy, ± globose and unlobed to distinctly 4-lobed, sep- arating into 2-4 nutlets at maturity. A large genus of about 120 poorly-dehned species in the warmer regions of the world, but centering in the neotropics. Eight species are found in Panama. a. Fruits distinctly 2-4-lohcd; anthers connate apically. b. Leaves with dense, curled pubescence; corolla tube < 2.5 mm long; style ca 1 mm long i j vohihilis bb. Leaves pubescent, but the hairs not dense nor curled; corolla tube ca > 3 mm long; style elongate and delicate, ca 2-3 mm long 2. T. maciilata aa. Fruits pyramidal or globose, not 2-4-lobed; anthers free apically. c. Flowers with a long corolla tube, 10-12 mm, and short sepals, ca < L5 "^m 1""K 3. T. anguitiflura cc. Flowers without the above combination of a long corolla tube and short sepals. d. Plants essentially glabrous on the stems. e. Branches of the inflorescence elongate; stigma on a short style; leaves me ■mbranaceous 4. T. glabra ee. Branches of the inflorescence short and tightly scorpioid; stigma ± sessile; leaves coriaceous 5. T. hicoJor dd. Plants conspicuously pubescent on stems, leaves, and sometimes fruits. f. Inflorescences densely-flowered, markedly scorpioid; sepals mostly 2-6 mm long. g. Stem hairs 4-6 mm long; leaves acuminate apically 6. T. cuspidata gg. Stem hairs 1-2 mm long; leaves acute apically 7. T. hirsutisshna ff. Inflorescences once or twice dichotomously branch.^d, the branches elongate; sepals very long, ca 8-10 mm 8. T. johnstouii 1. Tournefortia volubilis L., Sp. PI. 140, 1753.— Fig. 3. Vhies, woody, the stems with curled, appressed hairs. Leaves alternate, ± ovate, acute, entire, the bases obtuse or slightly rounded, ca 4-6 cm long and 1.5-2.5 cm wide, pubescent on both surfaces, the hairs curled and very dense beneath; petioles ca 0.5-1 cm long. lufloresceiices cymose, arranged in open panicles, the branches 3-4 cm long. Flowers sessile to pedicels ca 1 mm long in fruit; calyx of 5 sepals, slightly connate basally, lanceolate, ca 1 mm long, pubes- cent; corolla white, pubescent, the tube ca 2 mm long; the limb 5-lobed, the lobes subulate and ca 1.5-1.8 mm long; stamens 5, ± sessile, the anthers ca 0.6-0.7 mm long, connate apically; ovary ovoid, the style about equal to the sepals, the stigma conical. Fruit distinctly ( l-)2-4-lobcd, ca 3-4(?) mm in diam. Florida, Mexico, Central America, the West Indies, and South America. LOS SANTOS: Monagre Beach, Divyer 4182 (MO). An extremely variable species closely related to T. maciilata, Tournefortia voluhilis may be distinguished by its dense, curled pubescence, shorter corolla tube, and more delicate appearance of the inflorescence. 2. Tournefortia niaculala Jacq., Enum. 14, 1760. — Fig. 4. T. svrhigacfoUa Vahl, Symb. 3: 23, 1794. T. peruviana Poir., Fncyc. Suppl. 4? 42 5, 1816. SJiruhs, decumbent, or small trees to 5 m, the younger parts pubescent. Leaves alternate, ± elliptic to ovate, entire, acute to acuminate, the bases obtuse or rarely 1969] FLORA OF PANAMA (Family 167. Boragifiaceae^ 41 Fig. 3. Tournefortia volithilis L.: A, habit (ca X %); B, under surface of leaf (ca X 1.5). After Dwyer 4182 (MO). oblique, to 14 cm long and 5 cm wide, ± glabrous to moderately pubescent on both surfaces; petioles to ca 2 cm long, hiflorescences cymose, arranged in irreg- ular panicles, the branches to 5 cm long, Fhnrers sessile or with pedicels to 2 mm long in fruit; calyx of 5 free sepals, deltoid or ovate-elliptic, ca 1.5-2 mm long, [Vol. 56 42 ANNALS OF THE MISSOURI BOTANICAL GARDEN ^ V^t *rt4. ^ ^ /"/<- ; Fig. 4. Tourncfortia maculata Jacq.: A, fruiting branch (X Vs); B, flowering branch CX 3/8). A, after Tyson ct al. 4284 (MO); B, after Davidson 853 (MO). sparsely scurfy to lightly strigosc; corolla white to pale yellow-green, the tube ca 4-5 long, the lobes subulate, each < 1 mm long, ± scurfy; stamens 5, borne ca halfway up corolla tube, sessile, the anthers ca 1 mm long, connate apically; ovary ovoid, the style delicate, ca 2-3 mm long, the stigma conical. Fruit orange to yellow, distinctly (2-) 4-lobed, ca 4-5 mm wide, separating into (2-)4 nutlets at maturity. Mexico, Central America, the \\ est Indies, and northwestern South America. CANAL zone: Barro Colorado I, Builey & Bailey 237 (F); Ft San Lorenzo, Maxori 7011 (F). CHiRT^^ui: Boquctc, Davidson 853 (MO), cocle: bctvv Las Margaritas & El Valle, Woodson ct al. 1279 (MO), los santos: Loma Pricta, Duke 11879 (MO). Pan- ama: Cerro Aziil, Goofy Lake, Dwyer 2412 (MO); San Jose I, Frlanson 114 (GH), 501 (GH); rd from Chepo to i:i Llano, Tyson & Sn7ith 4119 (MO); Taboga I, Woodson ct a!. 1471 (MO). VERAGCAS: Coiba I, Du'\er 2iiO (MO); 5 mi E of Santiago, Trson ct al 4284 (MO). 1969] FLORA OF PANAMA (Family 167. Boragiuaceae') 43 3. Tournefortia angustiflora Ruiz & Pav., Fl. Peru. 2: 25, pi 151, 1799. T. billbergiaua Beurl., Kongl, Vetcnsk. Acad. Handl. Stock. 139, 18 54. SJiriih to 3 m, or vine-like, sparsely pubescent. Leaves alternate, elliptic, lan- ceolate-elliptic, or rarely ovate-elliptic, entire, acute, the bases obtuse, to 1 5 cm long and 8 cm wide; petioles to ca 1.5 cm long. Inflorescences spikes, scorpioid, to 6 cm long. Flmcers sessile; calyx of 5 sepals, slightly connate basally, broadly ovate, ca 1.2-1.5 mm long, ± glabrous to sparsely strigose; corolla white to yellowy glabrous to sparsely strigose, the tube ca 10-12 mm long, the lobes keeled, each ca 2 mm long; stamens 5, borne halfway up the corolla tube, ± sessile, the anthers ca 2 mm long; ovary ovoid, the style absent, the stigma closely appressed on the ovary. Fruit pyramidal, ca 4-5 mm in diam, the exocarp fleshy, separating into 4 nutlets at maturity. From Honduras to Peru, common in Panama. BOCAS DEL TORO: 10-15 mi from mouth of Rio Changuinola, Lewis ct ah 876 (MO); Chiriqui Lagoon, Water Valley, von Wcdd 1600 (MO), 1785 (MO), 1802 (MO), 1832 (MO), Old Bank I, von Wedel 2004 (MO); vie of Chiriqui Lagoon, Colon I, von Wedel 2867 (MO), 2960 (MO), canal zone: W end Gatun Lake Dam, Blum & Tyson 2000 (MO); Barro Colorado I, Wooduorth & Vested 378 (A), cocle: vie of El Valle de Anton, Allen 2059 (MO), darifn: El Real, Quebrada Trapiehe, Duke & Bristun 37 5 (MO); Cocalito, Dwyer 4465 (MO), herrera: 10 mi S Oeii, Tyson ct al. 2863 (MO). PANAMA: nr Chepo, Hunter & Allen 94 (MO), san elas: mouth of Rio Ailigandi to 2.5 mi inland, Lewis ct al. 167 (MO). The combination of long corolla tube and short sepals makes this species easily recognizable. 4. Tournefortia glabra L., Sp. Pi. 141, 1753. — Fig. 5. Shriih, or small tree to 8m, ± glabrous. Leaves alternate, ± crowded, lan- ceolate-elliptic to ovate-elliptic, entire or very finely undulate, acute or acuminate, the bases obtuse, to 23 cm long and 10 cm wide, sparsely strigose on upper sur- face; petioles to ca 4 cm long. Inflorescences cymose, the branches elongate, ca 12(-30) cm long at maturity. Flowers sessile; calyx of 5 free sepals, subulate to lanceolate, sparsely strigose, ca 1.2-2(-3) mm long; corolla yellow to green, sparsely strigose, the tube ca 4-5 mm long, the lobes ca 1 mm long; stamens 5, borne ca halfway up the corolla tube, sessile, the anthers ca 1 mm long; ovary globose, the style ca 1 mm long, the stigma conical. Fruit globose, the exocarp ± fleshy, ca 4-5 mm in diameter, separating into 4 nutlets at maturity. Mexico, Central America, and parts of the West Indies. }!OCAS DEL TORO: Almifantc, Daytonia Farm, Cooper 376 (MO); Chiriqui Lagoon, Water Valley, von Wedel J573 (MO), 1746 (MO). CHiRigui; Boqucte, nr Chiqucro, Davidson 552 (MO); Cerro Horqucta, Cloud Forest Litter Study, Duke et al 13637 (MO); Boquete, Finca Collins, Dw\cr & Hay den 7644 (MO); Rio Chiriqui Vicjo Valley, nr Bambita, White 46 (MO); Finca Lcrida to Pcna Blanca, Woodson & Schery 285 (MO); vie of Casita Alta, Volean de Chiriqui, Woodson et al 986 (MO), darien: bctw Quebrada Venado & Peje Swamp, Bristan 1006 (MO); vie of Paya, Rio Paya, Stern et al 396 (MO). PANAMA: Rio Bavano nr PanAm Hwy, DuJie 3974 (MO), 3998 (MO). [Vol. 56 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 5. Toiirnefortia glabra L.: A, fruiting branch (XVh); B, flowering branch CX %). After Woodson & Schery 285 (MO). 1969] FLORA OF PANAMA (Family 167. Boragijiaceae') 4 5 5. Tournefortia bicolor Sw., Prodr. 40, 1788. T. laevigata Lam., Tab. Encyc. 1: 416, 1791. Shrubs, scandcnt, or woody \incs, the stems ± glabrous, or rarely sparsely pubescent, tlie hairs appressed. Leaves alternate, ± elliptic, acute or rarelv acute-acuminate, entire, the bases obtuse, to 14 cm long and 7.5 cm wide, ± glabrous or rarely with appressed hairs, somewhat coriaceous, falcate, tending to dry brown-black; petioles ca 0.5-2 cm long. Inflorescences cymose, scorpioid, densely-flowered. Floivers sessile; calyx of 5 sepals, slightly connate basally, sub- a white (yellow on drying), ± glabrous (-6) C-2.5) the way up the corolla tube, almost sessile, the anthers ca 1.5 mm long; ovary ovoid, the style absent, the stigma thick and conical. Fruit globose, white, ca 5-6 mm in diam, the exocarp fleshy, containing (1)4 nutlets. Widely distributed in the American Tropics, and well represented in Panama. BOCAS DtL TORo: Water Valley, von Wedel 986 (MO), 1838 (MO); vie of Chiriqui Lagoon, Old Bank I, vou Wedel 1952 (MO), 1992 (MO), Fish Creek Hills, von Wedel 2427 (MO), Cocoa Cay, von Wedel 2877 (MO), canal zone: Miraflores locks, Victoria Fill, Allen 1713 (MO); Ft Clayton, Duycr 3932 (MO); Barro Colorado I, Shattuck s.n. (MO), 807 (MO); nr Gorgas Memorial Lab, White 83 (MO), darien: trail bctw Cana and Boca de Cupe, Stern et al 619 (MO); vie of Cana, Stern et ah 692 (MO); Tucute, Terry & Terry 1397 (MO), los santos: vie of Tonosi, Stern et al. 1867 (MO). Pan- ama: San Jose I, Johnston 758 (GH), 1355 (GH). 6. Tournefortia cuspidata II.13.K., Nov. Gen. Sp. PL 3: 83, 1818. — Fig. 6. T. ohscura DC, Prodr. 9: 517, 1845. Shrubs, decumbent and liana-like or ± erect to ca 3.5 m, the stems pubes- cent, the hairs 4-6 mm long, brown. Leaves alternate, elliptic, ovate, or rarely obovate or ovate-lanceolate, entire, acuminate, sometimes narrowly so, the bases obtuse, to 18 cm long and 8 cm wide, pubescent on both surfaces, especially on the veins beneath; petioles to ca 2 cm long. Inflorescences cymose, densely- flowered, scorpioid, mostly terminal. Flowers sessile; calyx of 5 sepals, lanceolate to subulate, densely strigose, ca 5-6 (-9) mm long; corolla white to yellow, strigose, the tube ca 6-7 mm long, the lobes ca 2-2.5 mm long, ± keeled; stamens 5, borne halfway up the corolla tube, ± sessile, the anthers ca 1 mm long; ovary ovoid, the style very short or absent, the stigma conical, a swollen ring basally. Fruit fleshy, somewhat lobed basally, ca 3-4 mm long, separating into C2-)4 nutlets at maturity. Central America, the West Indies, and northern South America. BOCAS DEL TORo: Changuinola Valley, Dujilap 411 (F); junction of Rios Changuin- ola & Tcrebe, Lewis et al 951 (MO); Almirante, nr Dos Nulla, McDaniel 5138 (MO); Chiriqui Lagoon, Water Valley, von Wedel 770 (MO), 1282 (MO), 1674 (MO), Old Bank I, von Wedel 1959 (MO), Little Bocas, von Wedel 2510 (MO), 2559 (MO). CANAL ZONE: Barro Colorado T, Avilcs 957 (F), Broun 90 (F), Woodivorth & Vestal 328 (F), 523 (F). darien: Boca Grande I, Duke 8843 (MO), panama: E of Pacora, Woodson et al 721 (MO), 722 (MO). 46 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 /Mi B Fig. 6. Tourncfnrtia cns-pidatu [[.B.K.: A, habit (ca X V2); B, upper surface of kaf (ca X 4). After Aviles 957 (F). 7. TournefortJa liirMili.shima L., Sp. PI. 140, 1753. Fig. 7. SJiruhs, vines, or rarely trees, llie stems pubescent, the hairs 1-2 mm lon^. Leaves alternate, elliptic to ovate, acute, sometimes slightly acuminate, entire, the bases weakly rounded to obtuse, to 1 7 em lon^,^ and 8 em wide, pubescent on both surfaces, frequently scabrous on the upper; petioles ca 0.5-2,5 cm long. Inflorescences cymose, seorpioid, axillary or terminal, densely-flowered. Floners sessile; calyx of 5 sepals, ± free, subulate to lanceolate, ca 2-4 mm long, pubes- cent on both surfaces and sometimes noticeably so on the margins; corolla white, densely strigosc above and glabrous near the base, the tube 4.5-5.5 mm long, the lobes ca 2 mm long; stamens 5, sessile, the anthers ca 1 mm long; ovary globose, 1969] FLORA OF PANAMA (Family 167. Boragiuaceae^ 47 B Fig. 7. Tournefortia hirsutissima L.: A, habit (ca X Vi); B, upper surface of leaf (ca X4.5); C, fruit (ca X 4.5); D, fruit, very pubescent (X 4.5). A-C, after Alien 4654 (MO); D, after Tyson 1572 (MO). the Style absent, the stigma stout and eonical. Fruit pyramidal, yellow-green ± pubescent, often densely so, ca 5-6 mm high, containing 4 nutlets. Widely distributed in Central America. [Vol. 56 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN BOCAS DFiL TORO: s. loc, voji Wi'dcl 4 52 (MO). t:ANAT, /ONE: Madden Dam, nr Rio Chagrcs, Scihert 560 (MO); Barro Colorado I, Gross Point, Shattuck 970 (MO); Ft San Lorenzo, Tyson J 572 (MO), Tyson & Blum 3698 (MO). CHiRiyui: Llanos on slopes of Volcan de Chiricjui Viejo, Allen 968 (MO); vie of Boquete. Allen 4654 (MO). Pan- ama: Toeumen, Duyer 4056 (MO). 8. Tournefortia johnstonii Stanclley, Publ. Field Mus. Nat. Hist., Bot. Scr. 18: 991, 1938. Shrubs, to ca 3 m, stems densely pubescent, the hairs short, brown. Leaves opposite to siiboppositc, ovate to ovate-elliptic, finely undulate to irregularly finely dentate, acute or acute-acuminate, the bases ± obtuse, to 17 cm long and 8 cm wide, densely to moderately short-hairy, more so on the veins beneath, cystolilh- dotted; petioles to ca 3 cm long. Inflorescences cymose, once or twice dich- otomousl}' branched, the branches to ca 12 cm long, axillary or terminal. Flowers ± sessile or with pedicels to ca 1 mm long; calyx of 5 free sepals, narrow- lanceolate, 7-10 mm long, strigose; corolla green to yellow, strigose, the tube ca 5-7 mm long, the lobes keeled, eacli ca 2 mm long; stamens 5, borne near the apex at the corolla tube, sessile, the anthers ca 1.8 mm long; ovary ovoid, the style ca 1-2 mm long, the stigma conical. Fruit pyramidal, unlobed, fleshy, ca 5-7 mm in diameter, separating into 1-4 nutlets at maturity. Apparently occurring only in Costa Rica and Panama. CHiRiyci: Volean de Chiriqui, Davidson 976 (F), vie of Casita Alta, Woodson ct ah 888 (MO, NY). Standley, in his Flora of Costa Rica (Publ. Field Mus. Nat. Hist. Bot. Ser. 18: 991, 992, 1938), described two new species, T. johnstonii and T . ramonensis. The Panamanian collections cited above have the longer sepals and corolla tubes which Standley attributed to T. ]ohn$tonii. However, when these specimens are compared with a plioto of the type of T. ramonensis (/^r6';/^s 6118, photo GH from F), I ha\e serious doubts about tlie validity of their separation. 1 have not seen sufficient material to make this reduction with assurance. Because of insufficient material, one collection of Tournefortia remains un- named: Woodson & Schery 609 (MO) (Chiriqui: vie of Bajo Chorro): shrub to 3 m, the branches densely brown-pubescent; leaves ± elliptic, ca 11-14 cm long and 4-5 cm wide, pubescent and cystolith-dotted on both surfaces; sepals lanceolate, ca 5-7 mm long in fruit; style ca 2 mm long, the stigma conical; fruit with a dry exocarp (?), unlobed. This collection may be referable to either T. johnstonii or T. ramonensis, but the fruit lacks the fleshy exocarp of these species. I am unable to determine if this is only an immature condition in the Woodson & Schery collection. See also the discussion of T. johnstonii. 5, BOURRERIA Bourreria P. Br., Hist. Jamaica 168, 1756. Trees or shrubs. Leaves alternate, rarely almost whorled, simple, mostly lanceo- late or elliptic; petiolate. Inflorescences cymes, open and many-floAvered or occasion- ally few-flo\\ered. Flowers perfect, ± actinomorphic, sessile or pedicellate; calyx 1969] FLORA or PANAMA (Fnmilv 167. BoragUiaceae^ 49 campanulatC; closed in bud, 2-5-lohcd; corolla funnclforni to salvcrforni, 5-lobcd, the lobes frequently rounded; stamens 5, included or exserted, borne on the corolla throat, subsessile or the filaments only shortly free, the anthers narrow to broadly oblong; ovary ovoid, unlobed, 2- or incompletely 4docular, the style bifid near the apex, the stigmas 2 and capitate or peltate. ¥niit drupaceous, ± spherical, the pericarp thin, enclosing 4 nutlets. About 30 species, mostly in the W^^st Indies, but a few occur in Venezuela, Central America, and Florida. a. Corolla tubes ca 17-2 3 mm lon^, the lobes ca 9 mm in diam; leaves to 2 5 cm long 1. /^- panawensis aa. Corolla tubes < 6 mm long, the lobes ca 4-5 mm in diam; leaves to 16 cm long 2. B. oxrvhxUa 1. Bourreria pananiensis I. M. Johnston, Jour. Arnold Arb. 29: 229, 1948. R. supcrha var. glabra Schcry, Ann. Missouri Bot. Card. 29: ^^66, 1942. Trees, to 17 m, glabrous. Leaves ovate to obovatc, acute, entire, the bases obtuse, to 25 cm long and 10 cm wide, somewhat coriaceous, glabrous; petioles ca 1-2.5 cm long, bifl few -flowered, mostly terminal. Floirers with pedicels ca 0.5-1.5 cm long; calyx campanulate, coriaceous, glabrous, the tube ca 6-9 mm long, 5-lobed, the lobes deltoid, subccjual, and ca 3.5-5.5 mm long; corolla funnelform, wiiite, densely papillose-glandular on both surfaces, the tube ca 17-23 nmi long, 5-lobed, the lobes semi-orbicular, ca 8-9 mm deep and 9-10 wide; stamcjis 5, included, borne on the corolla throat, sub- sessile, the hlaments glabrous, the anthers ca 2.5-3 mm long and 2-2.2 mm wide; ovary subovoid, minute at anthesis, ca 1-2 mm high, the style stout, ca 9-11 mm long, bifid near the tip, the stigmas 2 and peltate. Fruit unknown. Known only from the Chiriqui Lagoon region of Panama. liocAS DEL TORo: Cliirlqiii Lagoon, Colon I, von Wcdcl 2472 (MO), 2949 (GH, MO), 2974 (GH). 2. Bourreria oxyphylla Standi., Trop. Woods 16: 40, 1928.— Fig. 8. Trees, small, the younger branches with sparse appressed pubescence. Leaves ± elliptic, acute, entire, the bases obtuse, to 16 cm long and 8 cm wide, ± glabrous on both surfaces; petioles ca 1-1.5 cm long. Inflorescences cymose, paniculate, axillary or terminal, the branches softly pubescent, riuners subsessile to peduncles ca 3 mm long; calyx weakly campanulate, softly pubescent, the tube ca 4-5 mm long, 4-5-lobed, the lobes deltoid, subequal, ca 1-1.8 mm long; corolla funnelform, white, softly glandular-pubescent, the tube ca 4.5-5.5 mm long, 5-lobed, the lobes semi-orbicular to obovate, ca 4 mm deep and 5 nmi wide; stamens 5, exserted, borne on the corolla throat, the hlaments ca 4 mm long, the anthers ca 1.7-1.9 mm long; ovary oblong-ovoid, the style ca 4 mm long, the stigmas 2 and peltate. Fruit drupaceous, pyramidal, ca 5-7 mm high, the peri- carp hardened. Apparently known only from British Honduras, Honduras, and Panama. colon: beach betw Fato & Playa dc Damas, Pittier 3941 (GH). uariEN: Chusomo- catro & Rio Arcti, Dtike 8712 (MO), san blas: Mulatuppu, Duke 8 533 (MO); Soska- tupu I, Duke 8940 (MO). 50 ANNALS or THE MISSOURI BOTANICAL GARDEN [Vol. 56 ■ V Jl t^/ 1 cw-^^t-tVl^- '~1 A I'iK- 8. Boiirrcria oxypJnlla Standlc^y: A, habit (X ^/h); B, Hower (X 2). After Duke 8940 (MO). 6. Kill! I 11 A Elirelia P. Br., Hist. Jamaica 168, 1756. Trees or shrubs. Leaves alternate, simple, pinnately-nerved, entire or dentate; mostly petiolate. hifloiesceiiees i)anieulate or cymose, mostK terminal, riouers aetinomorphie, subsessile to pedicellate; calyx of 5 sepals, open in buti. perfect. free or somewhat connate basallv; corolla fimnelform, small, white, 5-lobed, tlie lobes spreading; stamens 5, mosth exserted, the anthers linear or broadly oblong; ovary 2- or 4-loeLdar, the style bilid near the tip, the stigmas 2, and capitate or claxate. Fruit drupaceous, mosth spherical, with one 4doculed and 4-seeded stone, with two 2-loculed, 2-seeded stones or with 4 separate stones. 1969] FLORA OF PANAMA (Family 167. Horag'uiaceae 51 A i'ig. 9. Ehrctia austiii-smilhii Standlcy: A, liabit (Xy6); B, fruit (ca X 2). After White 220 (F). / Vbout 40 species, mostly in the Old World tropics, with a few in the West Indies and Central America. The wood is of economic value in some of the tree species. 1. Ehretia auslin-Miiithii Standi., Publ. Field Mus. Nat. Hist., Bot. Ser. 18 984, 1938. Fig. 9. Trees to 2 5 m, the vounger branches pubescent. Leaves ± ovate, acute, serrate, the bases rounded, or shallowly truncate, to 6 cm long and 5 cm wide, ± scabrous abo\e and sparsely pubescent beneath; petioles to ca 1 cm long. Inflorescences cymose, arranged in an open panicle, mostly terminal, the branches scurf\. Fhuers subsessile to pedicels ca 1-2 mm long; calyx of 5 sepals, connate Vol. 56 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN basally, open in bud, ca 1.6-1.8 inni long o\cr-all, the lobes ovate, the margins finibriolate; eorolla ± eampanulate, white, the tube ca 1.8-2 mm long, 5-Jobed, the lobes ovate, ea 1.2-1.4 mm long; stamens 5, exserted, borne low in the corolla throat, the lilaments ca 1.6-1.8 nnn long, glabrous, the anthers ea 1 mm long; ovary ovoid, the style stout, ea 1 mm long, verv shorth bifid, the stigmas 2 and recurved. Vrait drupaceous, white to green, ± oxoid, ca 10-12 mm in diam. ApparentI) occurring only in Costa Puca and Panama. CHiRiyii: nio Chiriqui \ kjo Valley nr I3ambito Woods, W'hWc 220 (F). The above collection is in fruit only; the floral description was completed from Costa Hican material [A//,s7/;/ S;;///// /i528 (MO, isotype)]. 7. CORD I A CorcHa L., Sp. PI. 190, 1753. Slirnhs, sometimes scandent, or trees, sometimes functionally dioecious, stri- gose to softly pubescent, the hairs simple, uniseriate, or stellate. Leaves mostly alternate, rarely subopposite, simple, mostly homomorphic, sometimes dimorphic, petiolate. hifloreseences cymes, very irregular, paniculate and open, spike-like, or rarely densely glomerate, axillary or terminal, the bracts generally absent. FUniers perfect and homomorphic or functionally unisexual and slightly heteromorphic, actinomorphic, ± sessile; calyx of 5 connate sepals, tubular or ± eampanulate, sometimes costate, densely strigose to softly pubescent, sometimes with resinous dots when young, 5-Iobed or splitting irregularly and 2-4-lobed; corolla of 5(12) connate petals, funnelform to saherform, marcescent or falling soon after anthesis, sparsely strigose to glabr(Kis, white to cream, 5(-12)-lobed, the lobes oblong and conspicuous, almost as broad as long, or shallow and obscure; stamens as many as the corolla lobes, functional, or reduced and abortive, epipetalous, borne on the corolla throat, alternate, the filaments sometimes finibriolate and /or hooked near the base, the anlliers dehiscing longitudinally, introrse; ovary 4-loculed, functional or reduced and abortixe, unlobed, the ovules 1-4, usually only one developing, style slender, elongate, twice bifid, the stigmas 4 and capitate or cla\ate. Fruit usually drupaceous, unlobed, the exocarp dry or mucilaginous; seed usually 1. A genus of about 200 species, distributed in tlie warmer regions of tlie world. Se\enteen species are found in Panama. ;i. Inflorescences open paniculate cymes, repeated!} branched; corolla lobes readily delimited. b. Flowers functionally dioecious, heteromorphic, sometimes only slightly so; stamens reduced in 9 ilowers, the style reduced in S flowers. c. Calyx costate; leaves homomorpbic 1. C. divcrsifolia cc. Calyx not conspicuously costate; leaves large and beteromorpbic, some ovate, the others much smaller and ± orbicuhir 2. C. pufunNcnsis bb. Flowers perfect, homomorphic; stamens and style fully de\eloped. d. Hairs stellate ^i. C. aUiodora dd. Hairs simple. e. Leaves conspicuously bullate 4. C. durcri __ w ee. Leaves not bullate. 1969] FLORA OF PANAMA (Family 167. Eoragiuaccae^ 53 f. Corolla-lohcs broader than lon^; leaves ovate to obovate and ± toothed on the distal portion 5. C dcntata If. C'orolla-lohes not broader than long; if leaves ovate or obovate, then not ± toothed on the distal portion. g. Calyx persistent, enveloping the maturing fruit; corolla ver- niillion; leaves large and coarseh scabrous 6. C. schcstoia gg. Calyx persistent, but not enveloping the maturing fruit; corolla white or cream; leaves not conspicuoush large and scabrous. h. Branching noticeably dichotomous; veins and vc inlets of under surface of leaves covered by appressed slender hairs which cover and converge over the vein let areoles 7. C. hicolor hh. Branching not regularly dichotomous; undersurface of leaves glabrous or pubescent but not vvith veins and veinlets of undersurface of leaves covered by appressed slender hairs which cover and converge over the veinlet- areoles. i. Leaves ± obovate, conspicuously and abruptly long acuminate; branches of the inflorescence essentially glabrous 8. C. lasiocalyx ii. Leaves ovate to elliptic-lanceolate, acuminate but not al)ruptly so; branches of the inflorescence softly pu])e scent. j. Leaves rigid, noticeably falcate; calyx ca > 7 mm long 9. C. protracta jj. Leaves not conspicuously rigid or falcate; calyx ; petioles ca 1-4 mm long 13. C. lineata oo. Inflorescences internodal or terminal; petioles ca 5-10 mm long 14. C. inermis nn. Calyx-lobes caudate, ca 2-4 mm long 15. C. glohosa mm. Inflorescences spike-like. p. Bases of petioles decurrent on the axillary peduncles for 5-10 mm; inflorescences axillary 16. C. spinesccns pp. Bases of petioles not decurrent on axillary peduncles; inflorescences mostly terminal or sometimes internodal 17. C. ciirassavica 1. Cordia diversifolia Pavon ex DC, Prodr. 9: 474, 1845. C. johustOiiii Cufodontis, Arch. Bot. Sist. 10: 41, 1934. C. pctensis Lundcll, Wrightia 4: 49, 1968. Shriih or tree, to 10 m, functionally dioecious, branchlets densely soft-pubes- cent. Leaves alternate, lanceolate-elliptic, entire or sometimes sparsely dentate near tlie apices, acute to acute-mucronate, the bases obtuse to slightly oblique, to [Vol. 56 54 ANiXALS OF THE MISSOURI BOTANICAL CAKDEiN 20 cm long and 7 cm wide, roughl} pubescent, especially on the upper surface. hiflorcscoices cymes, ± paniculate, the tips scorpioid, terminal, ca 15 cm long, sot'tly-pubescent. rioirers functionally unisexual, ± sessile. Stamiuate f hirers with tlie calyx tubular, ca 4-5 mm long, ca 5'dentate, conspicuously ca 10-ribbcd, pubescent; corolla white, the tube ca 4-5 nnn long; the lobes oblong, ca 2-3 mm long and 1-1.2 mm wide; stamens 5, exscrted, the filaments sparsely pubescent near the base, ca 4 mm long, the anthers ca 1.6-1.8 nun long; ovary reduced and abortive, the style absent. Pistillate flowers with the calyx tubular, ca 4-5 mm long, 4-5-dentate, weakly ribbed, pubescent; corolla white, the tube ca 4 mm long, the lobes oblong, ca 2 mm long and < 1 mm wide; stamens ca 5, reduced and abortive, weakly exscrted, the filaments ca 1 mm long, glabrous, the anthers ca 0.5 nun long; ovary ovoid, the st\]e well-developed and exscrted, the branches elongate, ca 4 nun long in late flowering. Fruit ellipsoidal, fleshy, ca 7-9 mm long, the calyx persistent and saucer-shaped. Central America. BOCAS DFX TORO: Almlrantc, rd to the "Bomba", Blum 1314 C^IO); Changuinola, to S mi S of junction of Ilios Changuinola & Tcrc1u\ Leu is ct al. 795 (MO), 9i7 (MO), 946 (MO); ('hiriquicito to 5 mi S along Ilio Guarunio, Ix'uls ct al, 1994 (MO); \ ic of Chiriqiii lagoon, vou Wcdcl 1070 (MO), Old Bank I, von Wcdel 1894 (GH), 205/ (MO), Pumpkin 1^., von Wcdcl 2580 (MO), 2595 (MO), Water Vallcv, ion Wcdcl 2745 (MO); vie of Nieveeita, Woodsou ct id. IH24 (F, MO). 2. Cordia paiiamensis Riley, Kew Bull. 1927: 135, 1927. Shrubs or trees to 15 m, functionally dioecious, strigose. Leaves alternate or suhopposite, dimorphic, the larger o\ate to ovate-elliptic, acute, ± entire to rarely finely dentate, the bases rounded, ± ohlicpie or rarely obtuse, to 30 cm long and 15 cm wide, the smaller fewer in numlx'r, subopposite the larger, ovate-orbicular, acute, entire, the bases rounded and < half the si/e of the larger, scabrous or strigose on the upper surface, tomentose or strigose beneath; petioles, ca 5-15 mm long, strigose. \n\lorcsccnces cymes, paniculate, mostly terminal or borne in the dichotomous axils of the branches, to ca 15 cm lon<^ and wide. Flouers function- all\ unisexual, ± sessile. Stamiiiate flowers subglobose in bud; calyx campanulate, ca 4 nun long, 5 dentate, fainth ribbed, pubescent on both surfaces; corolla campanulate, while, the tube ca 3 mm long, the lobes oblong, ca 2 nun long and ca 1.2 nnn wide; stamens conspicuously exscrted, the lilaments ca 3-4 mm long, pubescent basally, the anthers ca 1 mm long; o\ary reduced and abortive, the style somewliat developed, included, branched, each branch bihd near the apex. Pistillate flouers clavate-eylindric in bud; calyx tubular-campanulate, ca 4 mm long, 5-dentate, 5-ribbed, pubesccJit on I)oth surfaces, becoming saucer-shaped in fruit; corolla saberform, white, the tube ca 2.5-3 min long, the lobes oblong, ca 1 mm long and 1 mm wide; stamens 4-5, reduced and abortive, subsessile to hla- ments ca 1 nun long, glabrous, the anthers ca 0.4-0.5 mm long; ovary ovoid- oblong, the style well developed, exscrted, branched, each branch bifid near the apex, the lobes clavate and recurved. Fruit a drupe, ovoid, white to green, black to dark brown on drying, glabrous, ca 6-9 nun long, coriaceous. 1969] FLORA oi PANAMA (Family 167. BoragiJwceacO 55 Central y\nicrica, from El Sahador lo Panama, and probably in Colombia and Ecuador. CzVNAL zone: KJl tracks along rd to Tropic Test Center, Corrca 261 (MO); Farfan Beach area, Dnycr 6766 (MO), Tyson JH23 (MO); Barro Colorado I, Miller Point, Shattuck 1155 (MO); Albrook A.F.B., Tyson 1114 (MO); Curundu, nr Survival School, Tyson 4174 (MO); vie of Miraflores, White 134 (MO), cocle: Hills S of El Valle de Anton, Allan 24H1 (MO); Rio Hato airstrip, Blum £r Dwycr 2471 (MO), colon: Nuevo Chagres, beach and adj rdside, Lewis el al i858 (MO). CHIRIQUI: 1: of Gualaca, Allen 5032 (MO); 2 mi SW Guabala, Tyson el al, 4247 (MO), darien: 2 mi E of Santa Fe, Tyson ct al 4676 (MO), panama: San Jose I, Andersson s.n. (MO), Erlanson 226 (GH), 2 76 (NY); Cerro Campana, Dnhe 8667 (MO); Pedro Gon/ales I, Dwycr 1705 (MO); E of Pacora, Woodson ct al. 757 (MO); Taboga I, Woodson ct al 1544 (MO), 1547 (MO). SAN HLAS: Mulatuppu, Duke H540 (MO); Ailigantli, Dwycr 6827 (MO). VERAGUAs: Coiba 1, Dwycr 1574 (MO), 2343 (MO); Santiago, 4 mi from Transisthmian Ihvy, Divyer & Kirkhridc 7412 (MO). Cordia pananiensis is one of the more perplexing species in this difficult genus. Other species QCordia macwphylla L., C. mlcaia DC.) also have large di- morphic leaves and indistinguishable (lowers and fruits, Johnston (Jour. Arnold Arb. 30: 126, 1949) maintained their separation on the basis of geographical ranoe and vewtative characters, but admitted that C. nuKrophylla, C. pajiamcnsis, and C. sulcata were closely related and were not "strong" species. From his key, and from the fact that their geographical ranges do tend to overlap, as well as from an examination of large numbers of Panamanian collections (whose vi hibit continuous variation), it appears that this group may well represent a polymorphic species centering in the West Indies and Central America. To add to the above confusion, fruiting specimens of C. paiiameusis look suspiciously similar to those of C. stcllifera I. M. Johnston, the species name of which refers to the star-like appearance of the calyx in fruit. This latter species is perfect-flowered and, although it does not appear to he in Panama (I have not seen large-leaved, perfect-flowered specimens of Cordia from that country), the question can be raised concerning the identihcation of fruiting collections of "C. stellifera" from other localities: are these in fact female specimens of C. pan- amensis, since both have a star-like calyx in fruit? 3. Cordia alliodora (Ruiz & Pav.) Cham., T innaea 8: 121, 1833. Ccrdona AUknhni Ruiz & Pav., Fl. Peru. 2: 47, />/. 2 84, 1799. Trees, to 20 m, stems and branches with stellate hairs, nodes swollen and inhabited by ants. Leaves alternate, oxate-elhptic, occasionally ovate, entire, acute or acute-mucronate, the bases obtuse or oi)lic|ue, to 18 cm long and 8 cm wide. sparsely to densely covered with stellate hairs, especially on the undersurface; petioles to ca 3.5 cm long. In florescences irregular cymes, spreading and loose, or sometimes compacted, the primary panicles to 18-20 cm long. Flouers perfect, ± sessile or with pedicels to ca 1 nun long; calyx tubular, ca 4-5 nmi long, densely soft-pubescent, ca 10-ribbed and with 5 small teeth; corolla-tube cylindrical, ca 3 nun long, lobes 5, spreading, each ca 5-6 nun long, white to cream, somewhat hirsute at the throat, ± persistent; stamens 5 (rarely more), borne at the corolla throat, the hlaments ca 5-6 mm long at maturity, each with a hooked fimbriolate [Vol. S6 56 ANNALS OI THE MISSOURI BOTANICAL f;ARl)KN base, tlic aiuluM-s ca 1.5-1.8 nun long; o\ary cyliiulrical, ca 1 mm long, ribbed or wrinkled, the style elongate, 3-4 mm long, the stigmas 4, as a result of 2 divisions, eaeh < 1 nun long. Fruit ejlindrical, not driipaeeous, the periearp fibrous and ehartaeeous, ea 5-6 mm lon^. W idel) distributed in tropieal Ameriea. nocAS DEL toko: vie of Chiri(iui Lagoon, Oltl Hank I, von Wcdcl 1H77 (MO). CANAL zont: vie of Gorgas Hospital, Blum 2210 (MO); Tropic Test Center, Duycr & Rohytis 56 (MO); Government Forest alon^ Las Cruees Trail, Hunter & Allen 449 (MO, NY); Chiva-Chiva Trail, Red Tank to Pueblo Nueve, Piper 5755 (NY), coton: Juan Mina, Rio Chagres, Allen 4198 (MO), hlhhlra: vie of Oei'i, hill ahove eantera of Sr. Joaquan Carri/o, Stern et til. 172 1 (MO), los santos: vie of Tonosi, Guanieo, Stern et al 1856 (MO). 4. Cordia dwyeri Nowieke, Ph\tologia (in press). — Fig. 10. Shrubs, to 3.5 ni, the stems densely pubeseent, tiie hairs simple, brown. Leaves ovate, sometimes ± obo\'ate, aeuminate, revolute, the bases ± roimded, to 31 em long and 13 em wide, sparsely pubeseent abo\e, densely pubeseent be- neath, coriaceous and eonspieuously bullate; petioles 1-2 cm long, stout, pubeseent. Inflorescences cymose, paniculate, densely flowered, the branehes pubescent. FloU' ers perfect (?), ± sessile; ealyx cupulate, densely strigose, the tube ca 4-5 nun long, 3-4-lobed, tlie lobes deltoid and ea 1.5-2 mm long; corolla salverform, color unknown, the tube 4-5 mm long, 5dobed, the lobes oblong, 2.5-3 mm long and 2-2,5 mm wide; stamens 5, exserted, the hlaments ca 7-8 mm long, pubescent basally, the anthers ca 1.5 mm long; ovary ovoid, the style ca 1,2 mm long, the stiigmas clavatc. Fruit unknown. Apparently only from Panama. colon: Santa Rita Ridge, 19 km from Transisthmian Ihvv, Dwyer H857 (holotvpe MO; isotypcs F, GH). The bullate condition of the leaves reacUIy separates C. duyeri from all other Panamanian species of Cordia, 5. Cordia dentata Poir., Ijicvc. 7: 48, 1806. Shnihs, or trees to 2 5 m, coarsely pubescent. leaves alternate, ovate to obo\ate, sparsely nuieronate-clentate, blunt-acute (slightly mucronate), the bases obtuse to rounded, to 12 cm long and 7 cm wide, coarsely pubescent, the hairs with multicellular bases; petioles to 2 cm long, hiflurescciices irregular panicles, the apices scorpioid, ca 10-13 cm long and wide, terminal, riowers perfect, ± sessile; calyx tubular, ca 10-ribbed, 3-4 mm long, 3(-5)-lobed, sparsely strigose on the ribs; corolla white to cream, t'unnelform, the tube ca 3-4 mm long with 5 widely spreading lobes 6-8 mm long, glabrous; stamens 5, borne on the corolla throat, the free hlament 3-4 mm long, sparsely strigose at the point of attach- ment, the anthers ca 1.7-1.9 mm long; ovary bottle-shaped, the style ca 2-3 mm long, dixided and the branches redixided near the apices, the stigmas 4. Fruit ovoid, with fleshy exocarp and bony endocarp, white, to 1.5 cm long. Mexico, Central America, the West Indies, and northern South America. 1969] FLORA OF PANAMA (Family 167. Boraginaceae') 57 Fig. 10. Cordia dwyeri Novvickc: A, habit CcaXV2); B, llower (ca X 2). After Dwyer 8857 (MO). [Vol. 56 58 AiNWALS or THE MISSOURI BOTANICAL GARDEN Hi-KKLKA: vie of Pcsc, Alloi 805 (MO, NY), los santos: vie of Las Tahlas, Diner 1 167 C.i\10); Monagre Beach, Du'\cr 4165 (MO). vera(;uas : 5 mi E Santiago, T\sofi et ill 4289 (MO). This is ihc "Cordia alba" of most aulhors. 6. Cordia sehestena L., Sp. PI. 190, 1753. Shrubs or trees, the younger stems pubescent. Leaves ovate, sometimes broadly so, acute, entire, the bases ± rounded, to 21 cm lung and 15 cm wide, scabrous above, pubescent on the veins beneath; petioles ca 3-4 cm long. Jiifloresceiices eymose, paniculate, loosely-flowered. Houers perfect, pedicels ca 0.5-1.5 cm long, pubescent; calyx cylindrical, the tube ca 10 12 mm long at anthesis, 3(-4)- lobed, the lobes deltoid and ca 12 mm long, greatly enlarging in fruit; corolla funnelform, vermillion, the tube ca 20-2 5 mm long, 5-^6dobed, the lobes ca 10 nun long and 8-9 mm wide; stamens 5(^6), included, the filaments free from the corolla tul)e for ca 2-3 n)m, ± glabrous at tlie point of attachment, the anthers ca 2.2-2.4 nun long; ovary ovoid, the style ca 20-2 5 nun long, the lobes clavate and flattened. Fruit drupaceous, o\oid, hard, ca 2-2.5 cm in diam, the calyx tightly adhering and becoming white and fleshy. Native to the West Indies and ranging from the Bahamas to nortbern Vcne/.uela. SAN ULAS: Soskatupu I, Duke 8963 (MO), 15476 (MO), Elhis 1698 (MO). This species, which is widely cultivated in the tropics, is very distinctive among the Panamanian members of CAndla because of the showy flowers and the large fruit with its adherent calyx. Tlie collections cited above, which are in bud or late-flower-early-fruiting stage, were not adequate for a complete description, so \AVst Indian material [\r/7.so// 7972 (MO); Stimsou 4200 (MO)] was also utilized. 7. Cordia hicolor DC, Prodr. 9: 4S5, 1845. C. trichostylu Pittier, Contr. U.S. Nat. Herl). 18: 252, 1917. Trees, 10-20 m, the younger branches and stems appressed-pubescent. I.eaies elliptic to ovate, acuminate or sometimes acute, entire, the bases obtuse, to 18 cm long and 8 cm wide, darker with appressed pubescence abo\e, paler with slender appressed hairs below that are borne on the veins and \einlets and conxer^e and cover the veinlets-areoles; petioles ca 0.5 cm long. Jiifloresceuce^s cvmes, paiu'cu- late and ± open. Flouers perfect; cal\\ tubular-campanulate, ca 4 nnn long. 5'dentate, the teeth ± unecjual, unribl)cd, pubescent; corofla saberform, uhite, the tube ca 3-4 mm long, the lobes oblong, ca 1.5-1.7 nun long and ca 1.2 nun wide; stamens exserted, the filaments ca 3 mm long, linibriolate basaily. the anthers ca 1.5-1.6 mm long; o\ary ovoid, the style well-developed, the lobes exserted and clavate. Fruit a drupe, ovoiil, ca 1 cm long, dull green, inibescent with minute grey hairs. Sporadically distributed in Central America, the southernmost West Indies, and northeastern South America. 1969] FLORA OF PANAMA (Family 167. Boraginaceae') 59 CANAL ZONE: Barro Colorado I, Oppcnhcimer 252 (MO). Panama: San Jose I, Johnston 620 (GH), 621 (GH), 963 (GH); Alios del Kio Pacora, edge of river and cloud forest, Leivis ct al 2267 (MO). Described at various times by Johnston (Jour. Arnold Arb, 21: 3 50, 1940; Sargcntia 7: 255, 1949) as being perfcct-flowered and functionally unisexual, the plants examined from Panama appear to be perfect-flowered: normal sized stamens and enlarging ovaries are found in the same flower. 8. Cordia lasioealyx Pittier, Contr. U.S. Nat. Herb. 18: 251, 1917. Fig. IIA. Trees, small or medium sized, glabrous. Leaves elliptic to obovate, conspicu- ously and abruptly long-acuminate ( 1.5-3 cm long), entire, the bases obtuse, to 14 cm long and 5 cm wide, glabrous on both surfaces, chartaceous; petioles ca 0.5-1 cm long, hiflorescences cymes, paniculate, axillary or terminal, A ' £f}t4^ ^ UX/^1 c Fig. 11. Cordia lasiocalyx Piuier: A, leaf (ca X %), after Shattiick 813 (F). C. hici- dula I. M. Johnston: B, leaf (ca X Vh), after Dunlap 322 (F). C. protraciu I. M. Johnston: C, leaf (ca X %), after Cooper 244 (GH). the branches glabrous. Flowers perfect, sessile; calyx cupulate, glabrous, ca 2.5-3 mm long, 3-lobed, the lobes deltoid and ca 2 mm long; corolla funnelform, white, the tube ca 4.5-5.5 mm long, 5-lobed, the lobes oblong, ca 4 mm long and ca 2.2 mm wide; stamens 5, exserted, the filaments ca 3-3.5 mm long, slightly pubescent basally, the anthers ca 1.3-1.5 mm long; ovary ovoid, the style slender, the stigmas clavate. Fruit irregularly ovoid, the stylar scar laterally placed, ca 5-6 [Vol. 56 60 ANNALS OF THE MISSOURI BOTANICAL GARDEN mm high and ca 9-10 mm wide, color unknown. Known only from Panama. CANAL zone: Barro Colorado I, Dock area, Sluitiuck 813 (F, MO), Gross Point, Shattuck 853 (F, MO), nr Termite House on shore, Wilson 69 (F). darien: around Garachiiie, Vittier 5694 (holotype US); forests around Finogana, Pitticr 6560 (LS). The collections cited above have been identified as Cordia nitida Vahl, which Jolinstop (1940) considers as restricted to the West Indies. I am inclined to agree with him after an examination of the W'est Indian material at the Missouri Botanical Garden. Among the I\mamanian species of Cordia it is very distinctive because of the leaves: relatively uniform in size and shape with the apices con- spicuously acuminate. I have seen only one fruiting specimen, but the drupe appears to mature asymmetrically, with the stylar scar becoming laterally placed. See also the discussion of C. protracta. 9. Cordia protracia 1. M. Johnston, Jour. Arnold Arb. 21: 349, 1940. Fig. lie. Shrubs, erect, the young stems softly pubsecent, the mature stems sparsely so. Leaves oblong-ovate, acuminate, entire, the bases rounded to ± obtuse, to 22 cm long and ca 10 cm wide, sparsely pubescent on the veins of both surfaces, coriaceous, noticeably falcate: petioles ca 0.5-1 cm long, lujlorescences cymes, paniculate, axillary or terminal, the branches softly pubescent. Flowers perfect, ± sessile, the buds clavate, ca 6-7 mm long; calyx ± tubular at anthesis, ca 6.5-7.5 nun long, sparsely strigose within and without, 5-lobed, the lobes deltoid and ca 1 mm long; corolla fuimclform, white, the tube ca 7-8 mm long, 5-lobed, the lobes oblong, ca 3-4 mm long and 1.6-2 mm wide; stamens 5, exserted, the iilaments ca 3-4 nun long, pubescent at the bases, the anthers ca 2 nmi long; o\ary elongate-ovoid, the style ca 7 mm long, the lobes slender. Yruli a drupe, ± ovoid, ca 1 cm long, white. Panama. SAN HLAS: Perme, Cooper 235 (US), 244 (holotype C\\\ isotype US). Several Panamanian species of Cordia have acuminately-tipped leaves as one of the more conspicuous characters of identihcation. They are also closely allied in floral characters, but, after an examination of type material, 1 think that they are all distinct species. Cordia lasiocalyx has the most ob\iously acuminate condition, but its regular and relatively small and thin leaves make it easily separable; C. protracta, with its large, rigid and noticeably falcate leaves, provides a striking contrast to C. lasiocalyx; the leaves of C. lucidida are admittedlv intermediate between the above-mentioned conditions, but this species is still easily distinguish- able, especially when the pubescence of the inflorescence and floral si/e are also considered. 10. Cordia lucidula I. M. Johnston, Jour. Arnold Arb. 21: 352, 1940.— Fig. IIB. Trees, small, to ca 8 m, mostly glabrous, the young stems softly-pubescent. Leaves ovate, sometimes ovate-elliptic or elliptic-lanceolate, acuminate, entire, the 1969] FLORA OF PANAMA (Family 167. Boraginaceae) 61 bases obtuse, to 16 cm long and 8 cm wide, ± glabrous on both surfaces, or very sparsely pubescent on the veins beneath, somewhat falcate; petioles ca 0.5-1 cm long. Inflorescences cymes, paniculate, open, mostly terminal, the branches softly- pubescent. Fhners perfect, ± sessile; calyx cupulate, very sparsely pubescent within and without, ca 2.5 mm long, 3-4'lobed, the lobes deltoid and ca 1 mm long; corolla campanulate, white, the tube ca 4 mm long, 5-lobed, the lobes oblong, ca 2 mm long and ca 1.2 mm wide; stamens 5, cxserted, the filaments ca 3.5-4 nnn long, pubescent basally, the anthers ca 1.2 mm long; ovary ovoid, the stigmas slenderly clavate. Fruit unknown. Panama and Costa Rica. BOCAs DEL TORo: Daytonia Farm, Cooper 372 (F); Research Lagoon, Cooper 406 (F); Changuinola Valley, Potrero I, Dunlap 284 (US; holotype, F; isotype), 322 (F); ChangLiinola Valley, Diinhip 356 (F); Chiriqui Lagoon, Fish Creek Hills, von Wedc4 2425 (MO). See discussion of C. protracta. 11. Cordia poreata Nowicke, Phytologia (in press). — Fig. 12. Shriihs^ to ca 3 m, the younger branches softly-pubescent. Feaves alternate, elliptic, acuminate, entire, the bases obtuse, to 24 cm long and 9 cm wide, ± rigid, slightly falcate, ± glabrous and grey-green above, sparsely pubescent be- neath; petioles ca 2-10 mm long. Inflorescences cymes, loosely-flowered, the peduncles softly-pubescent. Flowers perfect, ± sessile; buds clavate; calyx cupu- late, the tube ca 4 mm long, glabrous, 3-4-lobed, the lobes deltoid, ca 1.5-2 mm long; corolla salverform, white, the tube ca 4-4.5 mm long, 5-lobed, the lobes oblong, ca 5 mm long and ca 1.8 mm wide; stamens 5, exserted, the filaments ca 3-4 mm long, pubescent basally, the anthers ca 2 mm long; ovary ovoid, the style ca 4 mm long at anthesis. Fruit obliquely ellipsoidal, 10-12 mm high, apex acute. Known only from Panama. colon: Santa Rita Ridge lumber rd, Correa & Dressier 742 (MO), 1076 (holotype MO); Santa Rita Ridge, 19 km from Transisthmian Hwy, Dnyer 8581 (MO), hvvy to 8 mi E, Dwyer et al. 9027 (MO), los santos: Cerro Pilon, Divyer & Lallathin 8586 (MO). 12. Cordia bifurcata Roem. & Schult., Syst. Veg. 4: 466, 1819. — Fig. 13. Shrubs, small, the stems strigose. Leaves ovate to elliptic, ± acuminate, en- tire, the bases obtuse, to 10 cm long and 4 cm wide, sparsely pubescent on the upper surface, more so on the lower surface; petioles ca 3-10 mm long. Inf lores- cences cymes, the branches very reduced, sometimes appearing as a capitulum in bud, terminal or internodal. Flowers perfect, sessile; calyx cupulate, strigose, ca 1.8-2 mm long (4-)5-lobed, the lobes shallowly deltoid; corolla weakly cam- panulate, white, ca 3 mm long, obscurely 5-lobed, the lobes very small; stamens 5, weakly exserted if at all, the filaments ca 1 mm long, pubescent basally, the anthers ca 0.5 mm long; ovary ovoid, the style ± included. Fruit drupaceous, ± ovoid, enclosed within the calyx, ca 2-3 mm long. Costa Rica, Panama, and southward along the Andes. 62 ANNA! S OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 Fig. 12. Coniia porcata Nowickc: A, habit (X V2}; B, flower (ca X 3); C, fruit (ca X 2). After Corrcu & Dressier 1076 (MO). nociAS DFL roRO: Almirante, llesearch Grounds, Cooper 8 3 (F, GH, NY); Changuin- ola Valley, DunJap 90 (GH). darii \: vie of Paya, ilio Paya, Stern et al. 268 (GH). 13. Coniia lineala (L.) Rocm. & Schult., Syst. Vcg. 4: 464, 1819. Varro)iiu lineatu L,, Syst. Nat. ed. 10, 916, 17 59. Fig. 14. SJniihs, sciindent, or small trees, softly-pubescent. Leaves alternate, elongate- acuminate, the bases ± sessile elliptic to elongate ovate, coarsely dentate, acute to ± rounded to obtuse, to 12 cm long and 4 em wide coarsel\ pubescent; to petioles 3-4 nmi long, hiflorcsceiices cymes, densely glomerate, mostly axillary, 1969] FLORA OF PANAMA (Family 167. Boraguiaceae^ 63 Fig. 13. Cordia hifurcata Rocm. & Schiilt.: habit (ca X 5/6 ). After Stern ct al 268 (GH). [Vol. S6 64 ANNALS or THE MISSOL'RI BOTAMCAL CAHDEN Fig. 14. Cordia liiiccita (L.) Roem. & Schiilt.: habit (caXVn). After Duke 5580 (MO). 1969] FLORA OF PANAMA (Fnmilv 167. Vyoragiuaceae') 65 the peduncles to 4.5 cm long. Flowers perfect, sessile; calyx cupular, 2-3 mm long, densely strigose, 5 small teetli; corolla tubular, wliite, ca 3-4 mm long, with 5 inconspicuous lobes; stamens 5, borne on the corolla throat, the filaments ca < 1 mm long, the anthers ca 0.50.7 mm long; ovary globose, the style slender, ca 1.5-2 mm long, forked, the branches rediAided near the apices, the stigmas 4. Fruit a drupe, red, enveloped by the cal\\, ca 3-4 mm long. Mexico, Central America, the northern West Indies, and possibly northern South America. dahikn: Chipi^ana, Duke & liristti}i 264 (MO). Panama: PanAnu'rican II\\y at Rio Mamonica, Duke 55HO (MO); forests bordering grasslands on Cerro Campana, Duke 867 7 (MO); bctw Pacura & Cliepo, hoggv grasslands & marginal thickets, Woodsoji ef al. 1671 (MO). SAN BLAS: MiilatLipo, Rio Ihcdi, Cooper 276 (NV), Duke H4H7 (MO); Rio Mulatupo headwaters, riius 1742 (MO); mouth of Rio Ailigandi to 2.5 mi inland, I.eivis et a}, 204 (MO). 14. Cordia inermis (Mill.) I. i\l. Johnston, Jour. Arnold Arb. 30: 95, 1949.— Fig. 15. ] autana inerma Mill., C7ard. Diet. 1768. CAndia cana Mart. & Gal., Bull. Aead. P^ov. Soe. Bruxelles IK2): 331, 1844. Shrubs, to 2 ni, the younger stems seabrous, the mature stems less so. Feaves ovate, aeute-aeuminate, serrate or doLd)ly serrate, tlie bases entire, ± rounded and slightly attenuate, to 9 em long and ea 3-4 em wide, sparsely seabrous on both surfaces; petioles ea 0.5-1 em long. Jiifloresccuccs eymose, appearing as dense capitula, 5-7 mm wide, terminal or internodal, the peduncles slender, pubescent, < 3 cm long. Fhnrcrs perfect, sessile; calyx cupulate, strigose, ca 1.8 mm long, 5-lobed, the lobes deltoid; corolla weakly campanulate, white, the tube ca 2.5-3.5 mm long, obscurely 5dobed, the lobes ca 0.4 long and 0.6 mm wide; stamens 5, slightly exserted, the fUaments ca 1 mm long, pubescent basally, the anthers ca 0.5 mm long; ovary semi-spherical, style not exserted. Fruit unknown. Mexico, Central America, and the north coast of Colombia. PANAMA: Tabo^a 1, Woodsou et ui 147H (F, GH;. Johnston (Jour. Arnold Arb. 30: 95, 1949) also reports this species from Taboga Island. The specimen cited [Staudley 27039 (US)] has not been seen. 15. Cordia glohosa (Jacq.) ll.Bd<., Nov. Gen. Sp. PI. 3: 76: 1818. Varronia glohosa Jacc]., Enum. 14, 1760, Shrnhs, erect, 1-2 m, the stems scabrous. Leaver ovate, sometimes broadly so, acute, serrate, the bases rouncled to obtuse and ± oblique, to 6 cm long and 5 cm wide, scabrous on both surfaces; petioles ca 0,5-1.5 cm long, liiflorescoices cymosc, appearing as dense capitula, ca 1 cm in diam, the peduncles slender, scabrous, ca 1-2 cm long. Flowers perfect, sessile; calyx cupulate, ca 2-3 mm long, strigose, the lobes caudate, ca 2-3 mm long; corolla funnelform, white, ca 6-7 mm long, the lobes shallow; stamens 5, the fdaments ca 2-2.2 mm long, the anthers ca 0.8-0.9 mm long; ovary ovoid, the style slender, the stigmas clavate. Fruit drupaceous, enclosed by the calyx, ellipsoidal, ca 4-5 mm long. Mexico, Central America, the West Indies, Vene/uela, and eastern Bra/il. [Voi . 56 66 V.NMALS OF THE MISSOURI BOTAMCAL GAllDEN ^^ ; ;:; $ H ^ K A Fig. 15. Cordia iucrwis (Mill.) I. M, Johnston: A, hahit (X 5 7); B, infl (ca X ?5). After Wooihou ct al I47H CC}\~). orc'sccncc 1969] FLOPwV OF PANAMA (Family 167. Boragiuaceae^ 67 PANAMA: Punta Paitilla, Siandlcy 26268 (GH). Johnston (Jour. Arnold Arb. 30: 98, 1949) described C. glohosa as com- prising two geographic varieties. The typical variety was limited to the Lesser Antilles, Venezuela, and 13ra/il (with larger, more pointed, and sharply serrate leaves and larger flower-heads), and var. hnmilis to Florida, Mexico, Central America, and the Greater Antilles. The collection cited above is in fruit only. The floral description was taken from a Mexican specimen [Hinton 13209 (MO)] which Johnston (loc. cit.) c ited. 16. Cordia spinescens L., Mant. 2, 206, 1771. Varro}iia fcrntgiuca Lam., Tab. Fnc>'c. 1: 418, 1791. Cordia fcrrii^inea (Lam.) Rocm. & Schult., Syst. Veg. 4: 458, 1819. Shmhs or trees, to 5 m. Leaves alternate, ovate to ovate-elliptic, coarsely dentate to rarely ± entire, acute to acuminate, the bases rounded to slightly obtuse, to 17 cm long and 9 cm wide, sparsely strigose aboxe and softly-pubescent beneath, sometimes densely so; petioles ca 1-2.5 cm long. Juflorescences spikes, scorpioid, to 21 cm long, axillary, the peduncles decurrent with the leaf petioles. Flowers perfect, sessile; calyx cupular, ea 2-3 mm long, strigose, 5-toothed, faintly 10-lined or -ribbed; corolla tubular, white to yellow, ca 3-4 mm long, weakly Sdobed, fimbriolatc near the throat; stamens 5, borne on the corolla throat, the filaments ca 1 mm long, the anthers ca 0.6-0.7 mm long; ovary globose, the style slender, ca 3 mm long, twice forked, the stigmas 4. Fruit drupaceous, red, ca 3-5 mm long, ovoid; seeds 1-4. Mexico, Central America, Colombia, and southward along the Andes to Bolivia. IJOCAS DEL TORO: Chan^uinola to 5 mi S at junction of Rios C^hanguinola & Tercbc, Lexvis ct al 845 (MO); Water Valk-y, von Wedel 1633 (MO), 1672 (MO), 1801 (MO); Cliiri(|ui Tagoon, Old Bank I, vou Wcdcl 2169 (MO); vie Chiriqui LiiKoon, Quebra Nigua, von Wcdcl 2741 (MO), C-oUni I, vou Wcdcl 2839 (MO), canal zonh: Frijolcs, fhiu^^cr 307 (MO); Corozal, Crccumau & Crccnman 5207 (MO); Sahanas, YMlcy 117 (MO); Barro Colorado I, nr F. Miller #11, Shuttuck 480 (MO), Gross Trail #8, Shuttiick 1073 (F, MO); Miraflores Locks area, Tyson 1134 (MO); nr Ft San Lorenzo, Tyson & Blui}i 3676 (MO); X side of Canal hevond bridge, P. White 91 (F, MO); vie Miraflores Lake, P. WJiite 166 (MO). 24 5 (MO), chihkjli: Boquete, \V slopes of Cerro Horqucta, Allen 4781 (MO), Finea Collins, lilnui & Buyer 2525.4 (MO), fJjinger 705 (MO); Chiquero, Davidson 557 (F, MO); Cerro llorqueta, Duke ct al 13628 (MO), Duyer & Haydvu 7692 (.MO), Duyer ct al 53:^ (MO); Palto Alto, F of Boquete, Stern ct al 1088 (MO); linca Lerida to Pena Blanea, Woodson & Schcry 314 (MO); vie of Bajo Chorro, Woodso}i CT ScJicry 613 (MO); \'olcan de Chiricjui vie of Casita Alta, Woodson ct al 912 (MO). cocit:: El Vallc de Anton, trails nr Finca Tomas Arias, Allen 4232 (F, MO), dariln: Piio Balsa, betw N Quebrada C busomoeatre & Rio Areti, Duke 8706 (MO); Isla Casaya, Duke 10385 (MO). Panama: vie of junction of Rio Paeora 8c Pdo Corso, Duke 12009 (MO); tliiekets & forest nr Arraijan, Woodson ct al 1357 (MO), san blas: Veraguas, bills \V of Sona. Allen 1044 (MO); Mulatuppu (Rio Ibedi), Duke 8479 (MO); Sasardi, Duke 10144 (MO); mainland opposite Ailigandi, from moutb of Rio Ailigandi to 2.5 mi inland. Leu is ct al 165 (MO). [Vol. 56 68 ANNALS or THE MISSOURI BOTANICAL GAIUJEN 17. Cordia furassavica (Jacq.) Piocm. & Schult., Syst. Vcg. 4: 460, 1819. Vario)iui curassmica facq., I'.mim. 14, 1760, V. niacrostachya Jacq., loc. cit. Cordia macrostachia (Jac(|.) Room. & Schult., Syst. Vvg. 4: 461, 1819. C. chcpcnsis Pitticr, Conti. U.S. Nat. Herb. 18: 253, 1917. Shrnhs or small trees, to 3 in. Leaves alternate, lanceolate-elliptie to ovate- elliptic, coarsely to finely tlentate or subentire, acute, the bases obtuse or sometimes attenuate, to 13 cm long and 5 cm wide, sometimes ± coriaceous, the upper surface with small incrustations, each usually bearing a hair, the lower surface softly and densely pubescent; subsessile to petioles ca 5-6 inm long, hi florescences spikes, scorpioid, to 13 cm long, terminal. Floners perfect, sessile; calyx cupular, ca 2.5-3 mm long, 5-loothed, softly-pubescent and covered with resinous dots when young; corolla tubular, white, ca 3.5-4.5 mm long, weakly 5-lobed; stamens 5, borne on the corolla throat, the filaments ca 1.6-1.8 mm long, fimbriolate near the base, the anthers < 1 mm long; o\ary ovoid, the style slender, ca 2 mm long, twice divided near the apex, the stigmas 4. Fruit drupaceous, ovoid, ca 4-6 mm long and 3.5-5 mm wide, pink to red; seed 1, the cotyledons wrinkled and folded. Mexico, Central America, the West Indies, and northern South America. nocAs DEL TORO; Chiri(]iii Lasoon, Colon I, ro„ Wcdcl 2923 (MO), canal zonk- nr beach at Ft Kobbc, Diikc 4196 (MO); Toro Point nr Ft Slicrman, McDuuicl 4996 (MO), cocle: 10 km W of Agua Diilce on Intcranu-rican Hwy, Corrca 87 (MO); El Vallc de Anton, Scihert 4i9 (MO); bctw /VKiiadulcc and Anton, Woodson ct al. 1207 (MO); bctw Las Margaritas & HI Valle, Woodson et al J 293 (MO), colon: Nucvo Chagrcs, beach and adj rdsidc, Lvuh et al. 7 8 56 (MO), herreka: bctw Las Minas & Pese, Duke 123JH (MO); 12.5 mi S of Oci'i, Lewis et al. 1633 (MO), los santos: 1-2 mi W of Candelaria, Duke 12435 (MO). Panama: nr beach at Nucva Gorgona, Duke 4514 (MO); Ccrro Campana, Duke 10721 (MO), san blas: Soskatupu, Duke' 10191 (MO). VERACCAS: 5 mi N Santiago, vie of Rio Santa Maria, lihim & Tyson 624 (MO); 2-4 mi F of Santiago, Duke 12366 (MO); 2 mi W of Santiago on Transisthmian Hwy, Duyer et al 7550 (MO). Because of insufficient material, a few collections of Cordia remain un- named; (1) Cooper 429 (F, Gil) (Bocas del Toro: Daytonia Farm) and Cooper & Slater 307 (F, US) (Chiriqui: Progreso): trees 6-20 m; leaves obovate, ^, brown-pubescent beneath; fruits yellow to rcddish-vellow. These mav be referable to C. collococca F., which is conspicuousl) absent from Panama, but a note by Johnston on the Cooper 429 (GH) sheet refers both collections to C. invennsta, an undescribed species. I have not been able to find a valid publication of the above and do not wish to describe a new species from fruiting material alone. (2) \eivis et al 1806 (MO) (Code: hills NF of Fl Valle de Antcm): tree to 10 m; leaves ovate-elliptic, sparsely pubescent to ± glabrous on both surfaces; calyx 3-5-lobed; fruits immature. This may be referable to C. eriostigma Pitticr, but the absence of flowers and the damaged leaves make positive identification difficult. 1969] iLORA OF PANAMA (Family 167. Boraginaceae 69 Indjx of Lamn Nam is Numbers in InAd face t\p€ refer to descriptions; niniihers in roman type refer to s\non\ms; numbers uitb dagger (t) refer to Ucunes incidentally mentioned. Boraginaceae 33t Bourrcria 48 oxyphylla 49 panamcnsis 49 superba var. glabra 49 Cerdona alliodora 5 5 Cordia 331, 52, 55t, 56t, 581, 60t, 68t alba 58t alliodora 55 bicolor 58 bifurcata 61 cana 65 chepensis 68 colloeocca 68t curassavica 68 dcntata 56 diversifolia 53 dvvyeri 56, 56t eriostigma 68t ferruginea 67 globosa 65, 67t — var. humilis 67 inermis 65 invenusta 68t johnstonii 53 lasiocalyx 59, 60t lineata 62 lucidula 60t, 60 macropbylla 5 5 niacrostachia 68 nitida 60t panamensis 54, 5 5t pctcnsis 53 porcata 61 protracta 60t, 60 sebestena 58 spinescens 67 stellifera 5 51 t J, I sulcata 5 5't tricbostvla 58 Cynoglossum 34 mexicanum 3 5 officinale 34 Ehretia 50 austin-smithii 51 Hackelia 35 costaricensis 3 5 mexicana 35 Ileliophytum indicum 37 Heliotropium 37 curassavicum 38 indicum 37 inundatimi 38 lagoense 37t prcx'uml)ens 38 Lantana inerma 65 Lappula costaricensis 3 5 Tournefortia 38, 48t angustiflora 43 bicolor 45 billbergiana 43 cuspidata 45 glabra 43 hirsutissima 46 jobnstonii 48, 48t laevigata 45 maculata 4 Of, 40 obscura 4 5 peruviana 40 ramonensis 48t svringaefolia 40 volubilis 40, 40t Varronia 3 31 curassavica 68 ferruginea 67 globosa 65 lineata 62 macrostachva 68 FLORA OF PANAMA BY Robert E, Woodson, Jr. and Robert W. Schery AND Collaborators Part IX Family 169. LABIATAE^^^ BY Joan \V. Nowicke and Carl C, Epling ^ Missouri Botanical Garden and Department of Botany, Washington University, St. Louis, Missouri, and Department of Botan)-, Lniversity of California at Los Angeles, I OS Angeles, California Herbs, annual or perennial, half shrubs, or shrubs, rarely lianas or trees, usually nith aromatic oils, the stems and branches quadrangular, glabrous or pubescent, the hairs frec|uently uniseriate or glandular. Leaves opposite or some- times whorled, simple or rarely compound, petiolate or sessile, estipulate, the blade usually pinnately-nerved, entire to frequentl) dentate, glabrous to pubescent, sometimes with resinous' dots, hifloresceiwes usually cymes, 2-cc flowered, some- times reduced to a single flower,'' frequently sessile, each cyme then termed a verticillaster, generally opposite, each pair of verticillasters then forming a verticil, arranged as a spike-like raceme, or the verticils congested and forming a head or eapitulum, each verticillaster, capitulum, or flower subtended by a bractCs), persistent or deciduous. Flowers generally perfect, /ygomorphic or rarely actino- morphic; bracteolate or not; sessile to pedicellate; sepals (4-) 5, united, bilabiate or regular, persistent, 5-, 10-, 13 , or 15-ribbed; petals 5, sometimes appearing as 4 through reduction or fusion, united, typically bilabiate, the tube straight, arcuate or more sharply bent; stamens 4 and paired, or 2, staminodes rarely present, epipetalous, included or exscrted, distinct or rarely monadelphous, the hlaments straight or witli a xariously de\'e]oped connective, the anthers with 2 functional thecae or sometimes one, dehiscing longitudinally, the thecae parallel Or divergent; ovary superior, syncarpous, 2-carpellate, 4-lobcd, 2-locular or seem- ingly 4docular by a secondary di\ision, placentation basal, gynobase usually present, variously de\xdoped as a disc, gynophore, or nectary, the style 1, com- pletely gynobasic or terminal, bifid near the tip, the branches frequently unequal, 'Assisted by National Scienee Foundation Crant No. GBo674 (Principal hnestigator, Walter H. Lewis). '-^This paper is based on an extensive revision of a manuscript submitted by Epling in December, 1960 - January, 196L Almost all reductions to synonomy are those found in the original manuscript, and may in some cases be appearing for the first time. Also, as the phites date from 1961 some have no collection cited, and it was impossible to determine the specimens from which they were drawn. ■^Deceased, November, 1968. * "resinous*' is used here in a very general sense, and designates a yellow-green exudate. ■'Most of the Labiatae have vcrticillate inllorescences, and it is my opinion that the sol- itary flower may well represent the ultimate in reduction of the sessile cyme, i.e., only the central flower remains. I have, therefore, described all the inflorescences as being variations of a cyme. Ann. Missouri Bot. Card. 56(1): 7L111, 1969. [Vox . 56 72 ANNALS OF THK MISSOURI BOTANICAL GARDEN iJic stii^iiias minute at the tips, riiiit a collection of 4 nutlets, verv rarelv fleshv, attached at the base or laterally to a columnar extension of the gynobase; seed 1, the testa tliin, the endosperm absent or scanty, x = 6-11, 13, 17-20. A large family of about 200 o;cnera and 3200 species, cosmopolitan, but with a center of distribution in the Mediterranean region. iAbout 10 genera are found in Panama, but only Scdria and Hxpt'is are rep- resented in any di\'ersitv. ^ > Useful references : Fpling, C. C. Synopsis of the South American J aJ?iatac. Ilepert. Spec. Nov. Beih. «3C1): 1-96, 1935; 85(2): 97-192, 1936; 85(3): 193-288, 1936; 85(4): 289-341, 1937. Ki:y to Centra a. Stamens 2. 1). Only one fertile thcea in each anther, calyx bilabiate 1. Suhia bb. Roth thecae fertile in each anlber; calyx ± regular with 5 equal teeth ... 2. Ciiuila aa. Stamens 4. c. Stamen filaments united into a short tube above the attachment to the corolla; leaves frequently variegated; eoroUa tube reHexed near base ?>,Colcus cc. Stamen filaments free; leaves not \ariegated; corolla tube not sharpK reflexed. d. Middle lobe of lower eoroUa lip clearly dipper shaped; calyx ± regular. e. Mature nutlets attached at the base; cahx mostiv tul)ular with \er- tical vcination 4. Uypth ee. Mature nutlets attached lateralh to a columnar extension of the g\ no- base; calyx canipanulate and w ith horizontal \eins connecting the vertical 5. Marsyp'unithes dd. Middle lobe of lower corolla lip plane and spreading; eal\ \ weakly or conspicuously /ygomorphic. f. Lips of the cahx entire, the upper bearing an erect protuberance flat- tened contrary to tlie floral axis 6. ScuteUaria ff. Lips of the calyx variously toothed and without lateral projection of the upper lip. g. Calyx conspicuously bi'abiate, the upper lip with margins partially or completely deciuTent on the tube 7. Ocimum gg. Calyx weakly bilabiate, the upper lip lacking decurrent margins. h. Verticillasters reduced to 1 flower; calyx becoming saccate at maturitv 8. Tcucrhim hh. Verticillasters ca ,^41owered; calyx enlarging at maturit)' but not becoming saccate. i. Leaves elongate-tleltoid, the upper surfaces rugose-areo- late 9. Ltpechiuia ii. Iea\cs t)\ate, the upper surfaces flat 10. StacJiys 1. SALMA Salvia L., Sp. PI. 23, 1753. Hcrhs, annual or perennial, hall' sliruhs or shrubs, j^lahrous to pubescent. Leaves simple or rarely pinnatcly conipoimd, opposite, pctiolatc, pinnatcly-ncrvcd, entire or generally dentate, ghibrous or pubescent, the liairs frequently imiseriate. Inflorescences verticils, cacli with 4 or more flowers, or rarely the Ihjwers soHtary and opposite, the bracts deciduous or persistent, sometimes sliowy. Flouers sub- sessile to shortly pedicellate, the bracteoles generally not apparent; calyx tubular or canipanulate, bilabiate, rarely the teeth subequal, the upper lip generally entire or sometimes 3-lobed, 3- or 5-9-\eincd, the lower lip 2-lobed and generally 6'l0-\eined, sometimes witli glandular capitate hairs; corolla tubidar, /ygomorphic, 1969] FLORA OF PANAMA (Family 169. iMhiatae^ 73 *_ of 5 united petals, blue, white, or red, naked within or with 2-4 papillae or a ring of hairs above the ovary, bilabiate, the upper lip generally entire and plane, the lower B-Iobed; stamens 2, staminodia sometimes present, mounted on the eorolla throat, included in the upper lip or exserted, the hlaments distinct, the connecti\e elongate, the lower portion straight, retrorsely toothed or geniculate, the anthers generally with one functional theca; ovary 4-lobed, provided with a dis- tinct gynobase generally forming a nectarium, the st)lc arising from the gynobase, long and slender, bilid near the tip, the branches ± equal, or the posterior branch longer. ISutlets (104, attached at tlie base, oblong-ovate, brown. Salvia is a large genus of 500-900 species with an almost cosmopolitain distribution, being well represented in the tropics, tlie subtropics of the Mediter- ranean region and Mexico, and temperate areas. The genus is easily distinguished from the remaining Jahiatae found in Panama h\ its stamen number, the presence of only one functional thcca in each anther, and the distinct connective. Ten species of Salvia are described from I^mama. In the original manuscript of Epling, S. tiliaefolia Vahl was treated also, being represented by a single collection, Allen 217. Howe\er, after comparing the abo\e specimen with non- Panamanian material of S. tiliaefolia (all determined by Tpling) as well as a photo (MO) of the type [Herllardi sji, (C)], I have concluded that Allen 2i7 is a dc pauperatc specimen of S. lov^imarghiata Briq. and that S. tiliaefolia is absent from Panama or at least as of this date has not been collected there. a. Corolla generally bliiL", somctiincs rose or uliite, the tube 2.5-15 mm long; the bracts inconspicuous; the eal}.\ green, or rarely red. b. At least some upper calyx lips 5-9-veinecl, styles glabrous or liglitly pubescent. c. Corolla tube 2.5-6 mm long, naked within; styles glabrous. d. Verticils 6-10-Howered, the bracts ± lanceolate, ca 2-3 mm long and to ca 2 nun wide, e. Mature calyces to 4 mm long; stamen connective geniculate below^ the mitldle and bearing an assurgent arm 1. S. occidcu talis ce. Mature calyces ca 6 mm long; stamen connecti\e straight, bearing a retrorse tooth near the midtlle 2. .S\ micrantha i\i\. Innorescences den sel\ -flowered, globose in appearance, the bracts broadly reniform, ca 10 mm wide 3. S. hyptouk's cc. Corolla tube ca 7-10 mm long, ])caring 2-4 welbdefined papillae within above ovary; st>le pubescent near the tip 4. S. ptcroiira bb. Upper calyx lip 3-veined; styles pubescent. f. Corolla tid>e bearing 4 papillae within above the ovary. g. Corolla tube ca 11-15 mm long; verticils ca 0.5-2 cm distant; at least some pedicels ca 4-6 nun long 5. S. mcnihranacea gg. Corolla lube ca 6-7 mm long; verticils crowded; pedicels ca 0.5-2.5 mm long 6. S. polystachya ff. Corolla tube naked within. h. Leaf bhules conspicuously attenuate along the petiole; bracts ca 4 mm long, mostly deciduous; corolla !> 10 mm long, i. Delicate herbs; leaves small, ca 2-3 cm long, the bases mostly rounded-truncate, the petioles slender 8. S. trichopcs ii. Coarse herbs; leaves larger, at least some 6 cm long, the bases obtuse 9. S. alvajaca aa. Corolla crimson, the tube 18-34 mm long; the bracts show^y; calyx crimson 10. S. u'ivj^Tieriana [Vol. 56 74 ANNALS or THE MISSOURI BOTANICAL GARDEN 1. Salvia occidenlalis Swart/, Prodr. 14, 17S8. Herbs, anniuils, prostrate, the brandies ea 30-40 em long, finely pubeseent with small ± eiirxed Iiairs, or glabrous. J eaves ovate to ± deltoid, aeute, erenate- serrate along the distal margins, the bases obtuse, some truneate or rarely attenuate, to 6 em long and 3 em wide, glabrous, or the lower surfaee pubeseent Willi short hairs; subsessile to petioles ca 1 em long on lower leaves, liiflorcs' eeiiees vertieils, 6-10'ilowered, pubescent with glandular-capitate hairs, the bracts broadly ovate and long-acuminate, or rhomboid, ca 2-3 mm long, pubescent. riouers with pedicels ca 1-2.5 cm long; calyx tubular, ca 2-4 mm long at matLuity, with glandular-capitate hairs, bilabiate, the upper lip mostly 5-9-\eined; corolla tubular, ca 2.5 imn long, blue with white streaks, naked within, the upper lip ca 1,2-1.8 nun long, the lower lip weakly 3d()bed, ca 1.5-2 mm long; stamens mounted on the corolla throat, the anthers slender, ca 1 mm long; style slender, the stiuma 2-lobed, the lobes flattened. Nutlets smooth, ca 1.5 mm long, usuallv onl\' 1 maturing. A weedy species of the American tropics. iiiHRKRA: Ocii, 100 111, AUcn 4089 (MO); vie of Ocii, Stcyu et al 1705 (MO), los SANTOS: rd betw Macaracas & C-hitrc Hwy, Duycr 3155 (MO); rd l)Ltw Tonosi & Macara- cas, Oliver et at, -1S33 (MO), panama: Tahoga T, AUcu HI (MO), I 350 in, Allen 1284 (MO); hills bctw C^opira & rotrcro, 80-130 m, Dod-e & Hunter 8611 (MO). 8629 (MO); forest alon^ telcphoiu* trail l)Lit as a "weed" on trail f)t't\\ l^io Indio Ihdro^raphic Sta. & Natural Bridge of Ilio PiR'nti\ Doiliie & Allen 17480 (MO); \'cra Cru/ bcaclics, I.euis et al. 3017 (MO). 2. Salvia niirran!ha \ ahl, Fnuin. 1: 235, 1805. S. orhieuhiris Iknth., Hot Voy. Suliiluir 151, 1884. Herbs, annuals, the stems with appressed hairs, to 40 cm. I. eaves ± deUoid, acute, erenate, the bases ± truncate or sli^^htK cordate, to 3 cm long and 2 cm A\'ide, ± glabrous abo\e and sparsely pubescent on the \eins beneath; petioles to 8 mm long. Jiiflorescenees \'erticils, ca 6-flowered, the bracts lanceolate, ca 2 nun long, earl\' decichiotis. Flouers pcLhcellate, ea 2-4 mm long; bracteoles single, narrowK-lanceolate, ca 11, 5 mm lon^; calw tid)ular, 5-6 mm long at maturit\', pubescent w itii gland idar capitate hairs, the upper lobe 2-toothed and mostly 5-9-\eined, the lower 3-toothed; corolla tul)ular, ca 4-5 mm long, blue, naked A\ithin, the lo^ver lip much larger; stamens 2, the connective straight or with a slight hook at the tip; st\le long and slender, bifid at the tip, one branch much longer. Nutlets (3-)4, oblong, ca 2 mm long, pale brown. W ideh distributed in Florichi and the West Indies; knoA\ n in Panama onlv from Man/anillo Island. PANAMA: Man/anillo I, Ihiyes 716 (NY). This species difFers from S, oeeideutalis, to which it is undoLd)tedly closely related, b\ its more robust aj'^pcarance, the structiu'c of its stamen connecti\e, and its slender and tmetiual stiuma lobes. 1969] FLORA OF PANAMA (Family 169. Lahiatae') 75 3. Salvia hyptoides Mart. & Gal., Bull. Acad. Roy. Sci. Bruxcllcs 11(2): 74, 1844. Fig. 1 . Herbs, annual, erect to 60 ein. Leaves deltoid-ovate, acuminate, serrate, the bases truncate to slightly obtuse, to 5 cm long and 3.5 cm wide, both surfaces sparingly pubescent, more so on the veins beneath; petioles ca 1-2 cm long. Iniloresceuces verticils, densely-tlowercd, globose, 12-15 mm broad, the verticils Fly. 1. Sail ill hyptoidcs Mart. & Gal.: A, habit (ca X V2); B, flower (ca X 8); C, sta- mens (X 10); D, nutlets (ca X 10). Specimen unknown. 76 ANxNALS OF THE MISSOURI B0TAMC:AL GAHUKN [Vol. 56 ilistaiu below and crowded above, llie bracts reniforin and acuminate, often membranous, reticulately \eined, ca 8-10 mm wide, i hirers ± sessile or witb pedicels to 1.5 mm long; calyx ± tubular, 3.5-4 mm long, tbe upper lip with single tooth and ca 5-9-veined, the lower with 2 teeth, pubescent near the apex; corolla tubular, 2.5-3.5 mm long, blue, naked within, the upper lip ca 1 mm long and with glanduhu-capitate hairs, the lower lip ca 1.5 mm long and spread- ing; stamens mounted on tbe corolla throat, tbe connective retrorsely toothed below the middle, the anthers ca 0.3-0.4 mm long; style glabrous, the stigma \vith 2 branches acute and rounded. Nutlets 3-4, ovate and ± flattened, ca 0.5-0.7 mm long, the testa smooth and pale brown. Central America, Colombia, and Vene/uela. CANAL zone: around Tl Paraiso, Pittier 2575 (F). cuiiuyci: upper vallc\- of Rio Chiriqui Vicjo, White & White 105 (MO), dakiln: Rota de Ciipc, 40 m, /\//f» 904 (MO). mlrrlha: Olu, 100 m, Allot 4045 (MO). Los santos: rd Ixtw Tonosi & Macaracas, Oliver ft al. 3564 (MO), panama: Sabanas nr Chcpo, ^0 m. Hunter & Allen 23 (F, MO). SAN luAs: vie of Miilatuppu, Duke 8488 (MO). 4. Siilvia pleroura Bricj., Ann. Conserv. Jard. Hot. Geneve, 2: 139, 1898. Herbs or subshrubs, the branches ± glabrous, to 2 m. Leaves ovate to ellip- tic, acute, serrulate, tbe bases obtuse to rounded, sometimes extended along the petiole, to 1 1 cm long and 5 cm wide, ± glabrous, resin-dotted, petioles ca 1.5-3.5 cm long. Inflorescetices verticils, 6-1 2-n()wered, arranged in a spike-like raceme, ca 7-10 long, the bracts broadly awl-shaped, ± saccate at tbe base, long- acuminate, ca 3-5 nun long, deciduous, b'hnrers with pedicels 2-4 mm long; calyx tubular, ca 5.5-8 mm long, membranaceous, sometimes slighth pubescent, tbe upper lip entire and ± 5-ribbed, the lower lip di\ided and ca 5-7-ribbed; corolla tubular, 7-10 mm long, blue, with 2-4 papillae near base, the upper lip extended and ca 5-7 nmi long, the lower ca 4-5 nun long, pubescent near die distal portion; filaments with a retrorse tooth below the middle, the anthers slender, ca 1.5-1.8 mm long; style long, slender, pubescent near the tip. Nutlets 2-4, ovoid, ca 1.5 long, pale brown. Costa Pdca and Panama. CHnu^)ui: Ccrro Vaca, Pittier 5312 (F). c:ocle: vie of Fl \'alle de Anton, Allen 93 (MO), 600 m, 2764 (MO), White & White 68 (MO); la Pintada, 400-600 m, Hunter & Allen 53 7 (MO). 5. Salvia iiiembranarea Benth., Lab. Gen. et Sp. 259, 1833. S. inenihrn)uicea var. villostila Benth., loe. eit. 720, 183 5. S. eompueta var. iniziiensis O. Kt/e., Rev. Gen. PI. 2: 530, 189 1. S. irctziiensis lern., Proc. Amer. Acad. 35: 540, 1900. llerhs, somewhat suflrutescent, tlie upper stems and branches with pale brown uniseriate hairs, erect to 2 m. Leaves ovate, acute, crenate-serratc, tbe bases rounded, to 10 cm long and 5 cm wide, sparsely pubescent on botb surfaces more so on the veins, petioles to 7 cm long, hifloresceiiees verticils, 8-I2-flowered, 5-25 nun distant, the bracts rounded, caducous. Flowers with pedicels 2-6 nun long, the bracteole single, inconspicuous, caducous, ca 0.3-0.5 mm long; calyx tubular, ca 5-7 mm long, the upper lip entire and 3-veined, the 1969] FLORA OF PANAMA (Family 169. iMhiatae^ 77 lower lip 2-toothcd and 5-7-vcincd, slightly rcsin-dotted; corolla tubular, ca 15-20 mm long, rose-purple, with 4 papillae abo\e the ovary, the lips ca equal in length, ca 4-5 mm long, the lower more spreading, pubescent at the tip; con- nectives straight, the anthers ca 1.5 mm long; style long and slender, bifid and pubescent near the tip, the branches unequal. Nutlets unknown. Mexico and Central America. CHiRiyui: trail from Ccrro Punta to headwaters of Hio Caldera, Allen 1442 (GH,i\Y); Bajo Chorro, Duiidsou 81 (F, MO); Volcan dc Chiriqui, Davidson 926 (A, F), Woodson et ill 1037 (GH). 6. Salvia poljsiachya Ort., llort. Matrit. Dec. 5 5, 1797 S. compacta O. Ktzc, Rev. Gen. PI. 2: S30, 1891. Herbs, perennial, stems and branches with appressed hairs, rarely glabrous, to 3 m. Leaves o\ate, acute^acuminate, serruhite, the bases ± rounded, to 6 cm long and 4.5 cm wide, sparsely pubescent on both surfaces, more so on the veins beneath; petioles ca 1-3 cm long, liiflurescejues verticils, closely appressed to one another, 1 2-1 8-Howered, the bracts ovate and long-acuminate, ca 3 mm long, early caducous. Flowers with pedicels 0.5-2 mm long; bracteoles not apparent; calyx tubular, ca 4 mm long, pubescent with uniseriate hairs, resin-dotted, the upper lip entire and 3-veined, the lower 2-toothed and 6-7 \eined; corolla tubular, ca 6-7 mm long, blue-violet, with 4 papillae above the ovary, the upper lip ca 4 mm long and pubescent near the tip, the lower Hp ca 3 mm long; connectives straight and with a small tooth near the point of attachment, the anthers ca 1 mm long; style long and slender, bihd and pubescent near the tip, the branches unequal. Nutlets 4, oblong, ca 1.5 mm long, pale brown. Central America. CHiRiQei: Uanos on slopes of Volcan de Chiriqui, Allen 988 (MO); upper valley of Rio Chiruiiii Viejo, Allen 1590 (GH, MO), White & White 83 (MO), White 208 (GH); vie of Boquete, Lewis et al 396 (MO). This species is sometimes difficult to separate from S. nieuihranacea. The compactness of the verticils in S. polystachya is the most distinct difference be- tween them. 7. Salvia loiifiimargiiiata Bric]., Ann. Conserv. Jard. Bot. Geneve 2: 127, 1898. Herbs, perennial, somewhat suifruteseent at the base, to 1-2 m. Leaves ob- ovate to rhombate, acute-acuminate, serrate, the bases long-attenuate, to 8 em long and 5 cm wide, ± glabrous and slightly resin-dotted; petioles indistinct, to ca 5 cm long, winged. J }i florescences verticils, each ca 6-flowered, the bracts ca 1.5-2 mm long, ± awl-shaped, seemingly persistent. Flowers pedicellate, ca 1.5- 3.5 mm long, lightly pubescent; calyx tubular, ca 4-5 mm long, the upper lip entire and 3-veined, the lower lip 2-toothed and ca 7-9-veined; corolla tubular, ca 5-6 mm long, white to blue, naked within, the upper lip ca 1.2-1.3 mm long, the lower lip ca 1.7-1.8 mm long; connectives with a retrorse tooth below the middle, the anthers ca 0.8 mm long; styles long and slender, bifid, the branches conspicuously unequal. Nutlets ca 4, oblong-ovate, pale brown, ca 1.5 mm long. [Vol. 56 78 ANNALS OF TlIK MISSOURI BOTANICAL GARDEN Central Anicrica, Colombia, and Venc/ucla. CANAL ZONE: foicst along telephone trail hetw Ilio Indio Ilydrographic Sta & Natural Bridge of Rio Puente, Dodfic & Allen 17479 (MO). CHiuiyLJ: hetw Hato del Joho & Cerro Voca, Pitticr 5408 (GU); Bismarck, Williams 255 (NY), cocll: vie of El Vallc, Allen 217 (GH, MO). 8. Salvia trichopes Epling, Bull. Torrcy Bot. Club 68: 564, 1941. Herbs, or balf sbrubs, ± glabrous. Leaves o\atc to ± deltoid, acute, serrate, tlie bases rounded, ± truncate, or oecasionall) oblique, to 3.5 cm long and 2 cm wide, glabrous to rarely sparsely pubescent, sligbtly resin-dotted; petioles slender, ca 1-4 long. Inflorescences \crticils, each ca 4-6-no\\cred, llie bracts ca 5-6 nmi long, lojig-acuminate and deciduous, rioners willi pedicels ca 1-4 nnn lont' pubescent; calyx tubular, ca 6-7 mm long, bilabiate, tbe upper lobe entire and 3-veined, tbe lower lobe 2-lootbed and ca 7-veined, sparsely pubescent; corolla tubular, rose, ca 13-15 mm long, naked \\itbin, tbe lips about equal in lengtb, ca 2-2.2 mm long, ligbtly pubescent near tbe tip in bud; connectives uitb a retrorse lootli below tbe middle, tbe antbers ca 1 mm lony; styles lon^^ and slender, bifid, (lie branches unequal. Nutlets ca 4, oblong^ovate, pale brown, ea 1 .5 mm louLi. CHuutjLi: vie of Rajo C:horro, Woodson G- Seheiy 693 (isotype MO). Known onl\ from ibc t\pe collection, tbe delicate leaves and slender petioles of S. trichopes, as well as its corolla lengtb, indicate its distinctness from otber Stilvia species. 9. Salvia alvajaca Oersted, Vidensk. Meddcl. Dansk Naturbist. Foren. Kj0ben- bavn 38, 185 3. — Fie. 2 S. ourophylla Briq., Ann. Conserv. Jard. Bot. Geneve 2: 126, 1898. S. pcruiixta Briq., loe. eit. 129. 5. psilophyUa hpling. Bull. Torrey But. Chih 67: 5 2 7, 1940. SiihsJiriihs, brandies birtellous, to 2 m. Leaves o\ate to obovate, acute- acuminate, serrate or weakly serrulate, tbe bases obtuse, sometimes attenuate or rarely rounded, to 12 cm long and 8 cm wide, sparsely pubescent above and ± glabrous beneatb; petioles ca 2-9 cm long. Inflorescences verticils, 3-6 llowcred, 1-2 cm distant, tbe bracts awl-sbaped and keeled at tbe base, ca 4-6 cm lon<' deciduous. Floners with pedicels 2-6 mm long; calyx tubular, conspicuously 2-lipped, tbe lower 2 lobcd, the lobes lanceolate, tbe upper lip entire and 3-veined, tbe tube ca 4-5 mm long, tbe lobes ca 3-4 nnn long, membranaceous, ca 10-ribbed; corolla tubular, white and tinged with pink or blue, tbe tube ca 8-10 nnn long, naked within, the lips ca equal, ca 3-4 nmi long, pubescent near tbe tips; stamens mounted on tbe corolla throat, tbe anthers elongate, ca 1.5-2 mm long; style lonn and slender, ca 10-12 nnn long, bifid, tbe branches unequal. Nutlets (2-) o\()id, pale brown, ca 1.5 mm long. Apparently only in Costa P.ica and Panama. cniuiyci: Bajo Chorro, Davidson 60 (paratvpe of S. psiloplnlla, MO); lil Boqiute, Maxon 4964 (F); S slope Volean dc Ghiri(|ui, Tern U04 (F); F.'l Volc;in, Chiriqui Viejo Valley, White 99 (GH, MO), 167 (GH); V\ Volean, Potrero Muleto to summit, Woodson & Schery 452 (MO); Fl \ olean, vie of Callejon Seeo, Woodson & Sehery 4H5 (MO) 1969] FLORA OF PANAMA (Family 169. Ijihiatae^ 79 Fig. 2. Salvia alvajaca Oersted: A, habit (ca X Vz^; B, flower (ca X 5); C, stanuns (ca X 6); D, nutlets (ea X 10). Specimen unknown. Vol. 56 80 ANNAl S OF THli MISSOURI BOTANICAL GARDEN 10. Salvia wajiiieriana Polak., Linnaca n.s. 7: 591, 1877. S. toniluzii Briq., Ann. Conserv. J;)rd. Bot. Ccncvc 2: 157, 1898. Fis- 3. S. aJhopileata I'pling in Woodson & Schcr\ , Ann. Missouri Bot. Gard. 27: 337, 1940. Hcrhs,, somewhat suffrutcsccnt at llic base, perennial, the younger steins with uniseriate hairs, ereet to 2 m. heaven ovate, aeiniiinate, serrulate, the bases obtuse Fis- 3. Salvia xiaoucriana Polak.: A, habit (ca X 1/2); R, flowcT (X 2.5); C, stamuns (ca X 3.5); D, nutlets (ta X 3). Prohably after White & White Ml (MO). 1969] FLORA OF PANAMA (Family 169. Lahiatae^ 81 or rounded, to 1 3 cm long and 7 cm wide, ± glabrous and rcsin-dottcd; petioles ca 2-5 cm long, hifloresceiices verticils, paired, ca 1-1.5 cm distant, each with 5-7 flowers, bracts broadly rounded, 6-8 mm long. Flouers with pedicels ca 2-10 mm long, covered with uniseriate hairs; bracteoles not apparent; calyx tubular, ca 12-15 mm long, rose, bilabiate, the upper lip entire and ca 7-ribbed, the lower lip 2-toothed and ca 7-ribbed, pubescent with uniseriate and glandular capitate hairs; corolla tubular, ca 18-34 mm long, crimson, with 2 papillae above the ovary, the upper lip ca 3 mm long, the lower lip ca 2 mm long, both pubes- cent; stamens mounted near the top of the corolla throat, the connective with a long arm, the anUiers ca 2-2.2 mm long; styles long and slender, pubescent, the tips bifid, the branches unequal. Nutlets (2-)4, oblong, ca 2-2.5 mm long, mot- tled dark and pale brown. Apparently only from Costa Piica and Panama. CHiRiQUi: Volcan de Chiriqui, Terry /35] (F, Gil); upper Rio Chiriqui, White & White 321 (holotypc of S. alhopikata, i\l()). This is perhaps the most shoA\\ of the Salvias found in Panama. 2. CUNILA Cunila L., Sp. PI. ed. 2, 30, 1762. Half'shnihs or shrubs, scandent or erect. Leaves opposite, simple, subsessile to petiolate, pinnately-nerved, generally small, hifloresceuces verticils and densely- flowered, or loose umbel-like cymes, or capitula and few- to densely-flowered; the bracts generally leaf-like, hhnvers sessile or pedicellate; bracteoles usually present; calyx tubular, ± actinomorphie or bilabiate, 10 15-veined, pubescent on the inner throat; corolla tubular, white or purple, the tube enclosed or somewhat exsertcd, bilabiate, sometimes weakly so, the upper lip erect, almost flat, emargi- nate, the lower lip 3dobed, the middle lobe widest; stamens 2, exsertcd, the filaments distinct, the connective not developed, the anthers with 2 functional thecae; ovary 4-lobed, gynobase present, ± cylindrical, the style gynobasic, bifid near the tip, the branches subulate or the anterior somewhat longer and flattened. Nutlets (l-)4, attached at the base, ovoid, the pericarp smooth. A genus of ca 15 species in the New World. 1. Cunila polyanlha Benth., Lab. Gen. et Sp. 362, 1834. ^Fig. 4. Shnihs, scandent or ± erect, to 1.5 m, softly pubescent on the younger branches or glabrous. Leaves ovate to ovate-lanceolate, acute-acuminate, weakly serrulate to subcntire, the bases obtuse to ± rounded, to 6 cm long and 3 cm wide, slightly pubescent, more so on the veins beneath; petioles ca 2-5 mm long. Inflorescences verticils, rarely cymose, few and crowded, each ca 5-7-flowcred, bracts leaf-fike, ca 4-5 mm long, deciduous. Flouers with pedicels 1-3 mm long; bracteoles present, ± linear, ca 1.5-2 mm long; calyx tubular, ca 3.5-4 mm long, 5-toothed, the teeth deltoid and ± equal, ca 12-1 5-veined, pubescent, especially on the inner surface of the teeth; corolla tubular, white, exsertcd, ca 3-4 mm long, weakly bilabiate, the lips ca 1-1.5 mm long, the upper lip cleft, the lower hp 3-lobed, the middle larger, pubescent near the tip; stamens exsertcd, mounted [Vol. 56 82 ANNAT.S OF THE MISSOURI BOTANICAL GARDEN Fig. 4. Cunila polyantha Bcnth.: A, habit (ca X 1/2); B, floucr (ca X 10). Probably after Davidson 898 (MO). 1969] FLORA OF PANAMA (Family 169. Lahiatae') 83 low in the corolla throat, the anthers small, ca 0.5 mm long; gynobasc straight, the style slender, exserted, bifid near the tip, the branches unequal. Nutlets ovoid, smooth, mature size unknown. Mexico and Central America. CHiRiQui: Volcan de Chinqui,Davichon 543 (F), 898 (A, MO). Examination of non-Panamanian collections of C. polyantha reveals a notice- able variation in the development of the cymes and their arrangement. In some Mexican material the cymes are numerous, large and conspicuously pedunculate, m contrast to Panamanian specimens where the cymes are few, small and sessile, thus forming verticillasters. A photograph (MO) (G)] 7 not identical to either of the above mentioned specimens, shows \erticils with the more extensive cymose development. In the absence of any comprehensive treatment of the genus, I am not sure of the signihcance of this variation and hesitate to describe a new species. 3. COLFUS Coleus Lour., Fl. Cochinch. 2: 3 72, 1790. Herbs, half shrubs, or shrubs. Leaves opposite, simple, petiolate, pinnately- nervcd. Inflorescences \'erticils, ca 6-flowered, or loose cymes arranged in panicles, racemes, or spikes, the bracts reflexed or deciduous. Floirers ± pedicellate; calyx campanulate, bilabiate, the upper lip 3-lobed, the middle lobe largest, the lateral lobes pointed or rounded, the lower lip 2-toothcd or dobed; corolla tubular, re- flexed, bilabiate, the upper lip 3-4-lobed, the lower lip boat-shaped or concave and entire; stamens 4, curved into the lower lip, monadelphous, sometimes weakly so, the short tube free from the corolla, the anthers with 2 functional thecae; ovary 4-lobed, the gynobasc variously developed, sometimes as a nectary, the style gynobasic, bifid near the tip, the branches equal and acute. Nutlets ca 4, attached at the base, ovoid to ovoid-spherical, smooth. Variously described as having 100-200 species, mostly in warmer regions of the old world. A single introduced species in Panama. 1. Coleus blumei Benth., Lab. Gen. et Sp. 56, 1832. Herbs, somewhat succulent, sprawling, the stems to 70 cm long, the younger branches pubescent. Leaves ovate, ± acute, crenate, the bases truncate, to 4 cm long and 3.5 cm wide, sparsely pubescent above and on the veins beneath, resin-dotted, sometimes variegated; petioles ca 1-4 cm long. Inflorescences cymes, loosely-flowered, the lateral branches developed, the bracts deciduous. F/oicers with pedicels 1-2 mm long; bracteoles not apparent; calyx ca 2 mm long, pubes- cent, resin-dotted, the upper lip 3-lobed, the middle lobe much larger, the 2 laterals rounded, the lower lip much smaller and 2-toothed; corolla ca 1 cm long, purple, exserted, the tube sharply recurved, the upper lip 2-lobed, the lower lip boat-shaped, entire and becoming enlarged; filaments glabrous, united near their bases, the anthers ca 0.6-0.7 mm long; ovary 4-lobed, the gynobasc asymmetrical and becoming enlarged in fruit, the style bifid near the tip, the branches ± equal. 84 [Vol. 56 ANNALS OF THE MISSOUIU BOTANICAL CARDLN Nutlets 4, somewhat compressed laterally, ca 1 mm in diam, brown. Nati\c of the Kast Indies, but \\k\d\ introduced elsewhere because of its ornamental properties. cocLE: r\ \'allc df Anton, Lcitis et al. 25/4 (MO), 4. IIVPTIS Hyj)tis Jaeq., C:oll. 1: 101, 1787. nom. cons. Mcsosphacriini l\ Br., Hist. Jamaic. 2S7, pi. IH. f. 3, 17S6. Hi'ihs, coarse, half-shrubs, shrubs, or rarely trees. Leaves opposite, simple, mostly petiolate, pinnalely-ner\ed. hifloreseeiiees \erticils, arranged as a spike or raceme, or congested into dense capitula, ± sessile or pedunculate; bracts linear to ovate. FUmers sessile or pedicellate; bracteoles present or not apparent; calyx tubidar or campanulate, ± actinomorphic, rarely bilabiate, often enlarging in fruit, generally membranaceous and ca lO-xeined, the tu!)e straight, beiU, or cur\ed, the teeth deltoid, linear or spinulose; corolla tubular, white, blue, or rose-purple, glabrous or pubescent, bilabiate, the upper lip 2-lobed, the lower lip usually longer and 3-lobed, the nuddle lobe saccate; stamens 4, paired, mounted on the corolla throat, curved into the lower lip or exserted over it, the fdaments distinct, the connective not developed, the anthers with 2 functional th frequently red to purple, mostly divergent; ovary 4-lobed, gvnobasc lobed and ecae, somewhat elongated, the style gynobasic, long and slender, branched near th tip, the lobes small, elavate and ± e{|ual. Nutlets (l-)4, attaehed at the base, ovoid-cylindrieal or ± flattened, sometimes with 2 slight depressions on the lobes of the gynobase adhering, brown. About 300 species in the warmer regions of the New World, especiallv Brazil, and a few representatives in the Old World tropics. Useful reference: Epiing, C. C. Revision del genero Hvptis il.ahiatae). P.evista Mus. La Plata Bot. 7: 153-497, 1949. a. Verticils fcvv-flovvcrcd, distant. b. Caly.x teeth deltoid 1. f^. rcrtkiUata DO. C.ahx teeth linear or spinulose. c. Robust herbs; infloreseences open, peduneles ca 0.5-1 em lon^; calyx- tubes 5-7 mm lon^; bracts ± deeitUious 2. H. siancolcns cc. Weak herbs; intlorescenees more compacted, sessile, or with peduncles < 0.5 cm long; calyx tulses < 3 mm lon^; bracts persistent. d. Bracteoles linear; inflorescences ± scorpioid; conspicuous white tuft o f hair between calyx teeth 3. H. pectin peciinuta dd. Bracteoles ovate; inflorescences semi^lobose; no conspicuous white tuft of hair between calyx teeth 4. 11. miitahilis aa. Verticils congested into dense capitula, or if doubtful, then plant with branched hairs. e. Inflorescences sessile or sliortly pedunculate; the cleft between the upper calyx teeth cut more deepK 5^ j/_ pulcoioidcs cc. Inflorescences pedunculate; calyx teeth cut ± evenly. f. Outer bracts hliform to linear; cab x tube at maturity recurved above the middle 6, //. recunuta ff. Outer bracts lanceolate, spatulate, or ovate; calyx tube ± straight at ma- turitv. 1969] FLORA OF PANAMA (Family 169. Lahiatae^ 85 g. Stems and under surfaces of leaves with branched hairs . . 7. H. ohlo}ii;ifoUa gg. Hairs simple. h. Leaves definitelv sessile, the bases truncate and clasping; inflorcs- cences conspicuously whitish-pubescent 8. /i. cUlatata hh. Leaves petiolate, or if subsessile, then with narrowed bases; inflo- rescences not conspicuously whitish-pubescent. i. Calyx tube naked within, sometimes pubescent near orifice but not pilose-annulate above the nutlets. j. Peduncles ^ mm long, the teeth spinulose . ... 10. H. hreiipcs ]}. At least some peduncles 2 em long or longer. I. l^iobust, erect herbs; peduncles stout; at least some leaves > S cm long 1 1. H. brachiatu II. Slender, prostrate herl>s; peduncles slender; leaves mostly 2.5 3.5 cm long 12. f/. atroruhcns ii. Calyx tube distinctly pilose-annulate within above the nutlets. m. Calyx tube conspicuously longer than the teeth. . 13. H. capitafa mm. Cal> X tube and teeth about ecjual in length. n. Slender herbs; leaves ± ovate, ca 3 4 long; bracts ovate and crcnate 14. //. Umtanacfoha nn. Piobust, coarse herbs; leaves lanceolate, or if rarely ovate-lanceolate then > 5 cm long; bracts ± awl- shaped and entire, o. Bracts glabrous; mature capitula ± globos 15. H. savannarum oo. Bracts with soft pale brown pubescence; mature capitula ± hemispheric 16. H. conferta 1. Hyptis verlicillata Jacq., Coll. 1: 101, 1787. Mcsosphaerum vcrticillatum (Jaeq.) O. Ktze., Pev. Gen. PI. 2: 525, 1891. Herh or half shrub, perennial, 1-3 m tall, the branches apprcsscd-puberulent and cTcnerallv bearing spreading longer bairs along the angles. J eaves elliptical r\^ > > • • or oblong-lanccolatc, acute, serrulate or subentire, the bases obtuse, to 7 cm long and 2 cm wide, both surfaces lii^htly pubescent and often bearing longer hairs along the principal veins, resin-dotted; petioles ca 5-10 mm long, the upper leaves sometimes subsessile. Inflorescences verticils, 12-1 5-flowered, ± moniHform, arranged in panicles in the axils of the upper reduced leaves; bracts linear, ca 1-2 mm long, setaceous. Ylowers with pedicels ca 0.5-1 mm long, tufts of hairs at the bases; calyx tubular, ca 2 mm long at maturity, teeth ± lanceolate-deltoid, ca 1 mm long, the tube distended at maturity' b\ the nutlets, the teeth somewhat connivent; corolla ca 2-3 mm long, blue and white, exserted, the lobes ca 1 mm lona + pubescent; stamens exserted, the fdaments lightly pubescent, the anthers ca 0.1-0.2 mm lon^; crvnobase lobed, the style ca 2 mm long. Niitlets 2-4, ovoid-oblong, ca 1.5-1.7 mm long, the lobes of the gynobase sometimes adhering, the pericarp somewbat corky. A widespread weed of the American tropics. BOCAS DEL TORo: Changuinola to 5 mi S at junct of Rios Changuinola & Terebe, Lewis et ah 920 (MO); Chiriquieito to S mi S along Rio Guarumo, Lewis et al J 983 (MO); s. loc, von Weilel 326 (MO); vie of Chiriqui Lagoon, Cocoa Cay, von Wcdel 2878 (MO). CANAL zone: Rio Chagrcs, Fendlcr 221 (MO); Ft San Lorenzo, Tyson 2240 (MO); vie of Miraflores, G. White 123 (F, MO); s. loc, s. coll. (MO 1119508), darien: Rio Balso betw Qucbrada Chusomocatre & Rio Areti, Duke 8274 (MO); Tucute, Terry & Terry 1401 (F, MO). PANAMA: Taboga I, AUcn 1302 (MO); Rio Pita, 1-3 above confluence ^vith [Vol. 56 86 AiNNALS OF THE MISSOURI IJOTAINICAL (JARUEN Puu Macstra, Ditke 4732 (MO); liio Cliagrcs tributary, 5 mi S\V Cx'rro Brew .stcr, Ixui is ct a/, 3466 CMO); Chcpo, Tv,so;2 7427 (MO); Canita, 18 m V. of Chcpo, Tyson & Smith 4/38 (MO). SAN BLAS: mainlaiul opposite Achitiippu to 1.5 mi inland, Lewis ct al. 96 (MO). PROVINCE unknown: Audcrssoii s. n. (MO). 2. Hyptis suaveoleiis (L.) Foil., Ann. Miis. Hist. Nat. Paris 7: 472, pL 29, f. 2, 1806. Bullota siicivcolcus L., Syst. Nat., ctl. 10, 1100, 1759. Mcsosphucnim siturculois (L.) O. Ktze., J\v\ . Gun. Pi. 2: 525, 1891. Uerhs, coarse, sliglilly sufl'rtitesccnt at tlic base, to 3 m, ligluly to densely pubescent, tbe liairs iiniseriate and to 3 nun long, some glandular-capitate. Leaves deltoid, ovate, or rhoinbate, acute, serrate to doubly serrate, the bases cordate to rounded, to 8(-12) cm long and 6(-8) cm wide, pubescent on both sin-faces, densely so on the veins beneath; petioles to 7 cm long, hi florescences cymes, 3-6 flowered, the peduncles ca 0.5-2 cm long, the brads leal'like, caducous. Llouers subsessile; calyx narrowly campanulate, ca 4.5-6C-8) mm long at maturity, teeth spinuluse, ca 2 (-3. 5) mm long, ca 10-ribbed, membranaceous, pubescent; corolla purple, the tube ca 4-5 nnn long, exserted, the lips ca 1.5-2 mm long, pubescent; lilaments pubescent, the anthers ca 0.3 mm lonu; stvlc slender, ca 4 mm long, weakly 2-lol)ecl. Mittlets lC-2), reetangular, ea 3 nnn long, the pericarp corky and dark brown. Widely distributed in the tropics of North and South America, and also found in the Fast Indies to Hong Kong and lormosa, Australia, and Africa. I30CAS DEL lOHO: C:()Kimhus I., 6 mi inland, ion Wcdcl 34 (MO); vie of Chiricini Lagoon, Careening Cny, van Wcdcl 2H18 (MO), canal zonf: sal)anas alon^' drowned Ilio Azote Caballo, Dodi^c ct cd. ]6H^4 (MO), coclk: vie Illo li-ta & Pan Am Hwv, Blum & Tyson 1S92 (MO), uarii-n: vie of F.I Llano, Duke 5795 (MO), pana.ma: betw Capira & Potrero, Dodoe & Hunter 8640 (MO); S 6 mi L of Chcpo on Pan Am Hwy, Duke 4049 (MO); nr beach at Nueva Goryona, Duke 4490 (MO). 3. Hyplis peclinata (L.) Poit., Ann. Mus. Hist. Nat. Paris 7: 474, /;/. 30, 1806. Nepcta pecthiatu L., Sp. PI. 799, 1753. MesospJiaeruni pectinatum (L.) O. Kt/e., Hev. Gen. PL 2: S2 5, 1891. Her])s, perennial, to 3 m, stems pubescent and izenerallv scabrous alonu the angles. J eaves ovate to ± deltoid, acute, double serrate, the bases truncate to slightly obtuse, to 3.5C-8) cm long 2(-4.5) cm wide, pubescent on both surfaces, densely so beneath, sparsely resin-dotted; petioles to 5 cm long on mature leaves. Inflorescetices cymes, each with 2-3 semi-scorpioid liranches, the peduncles ca 1-2 nnn long, the bract leaf-like, 3-4 mm long, ilouers subsessile; bracteole linear, ca 3-4 mm long, pubescent; cah.x tubular, ca 2 nnn long at maturity, 10 ribbed, pubescent, the teeth spiiudose, ca 1-2 nnn long, a conspicuous tuft of white hairs between the teeth; corolla tube ca 1.5 mm long, exserted, rose-pinple abo\e, ± cream below, pubescent, small and inconspicuous, the lips ca 0.5-0.6 mm long; stamens ± sessile, the anthers ca 0.1 nnn long; g\nobase somewhat lobed, style seemingly enclosed, hhttlets 3-4, oblong, ca 0.7-0.9 mm long. A widespread but not abundant weed of the American tropics; to be expected at lower elevations. 1969] FLORA OF PANAMA (Family 169. iMbiatae^ 87 darien: Ccrro Pirrc, Dnke & Ulias 13676 (MO), los santos: 17.8 mi S of Macara- cas, Lewis et al. 1614 (MO); 25 mi S\V of Tonosi, headwaters of Rio Pedrcgal, Lewis et ah 2963 (MO). PANAMA: Ccrro Jefc, Duke 9389 (MO), Dwyer et al. 7267 (MO); Isla del Rcy, Dulic 9544 (MO). 4. Hyptis miilahilis (A. Rich.) Biiq., Bull. Herb. Boissicr 4: 788, 1896. Nepeta mutahilis A. I'ich., Actcs Soc. Hist. Nat. Paris 110, 1792. Mcsosiyhacriim nmtahile (A. Rich.) O. Ktzc, Rev. Gen. PI, 2: 525. 1891. Herbs, to 3,5 m. Leaves ovate or rhomboid, acute, serrate to doubly serrate, the bases truncate or obtuse, to 6 cm long and 3.5 cm wide, ± hirsute on the upper surfaces, ± pubescent below, particularly along the veins, subsessile to petioles ca 2-3 cm long, liifluresecnces verticils, 3-10 Howered, the peduncles ca 0.5-1 cm long at maturity, arranged as a panicle 5-15 cm long, the bracts leaf- like. Floivers sessile; bracteoles ovale or ovate-lanceolate, ca 4-6 mm long, pubes- cent and ciliate along the margins; caKx tube ca 1.5-2 mm long, elongating to ca 5-6(-7) nmi long at maturity, membranaceous, pubescent, 10-ribbed, the teeth spinulose, ca 1 mm long; corolla lube ca 4-5 mm long, purple to lavender, the lips <1 mm long, pubescent; stamens exserted, the filaments pubescent, the an- thers ca 0.2 mm long; style ca 4.5-5.5 mm long. Nutlets (2-)3-4, ovoid, ca 2 mm long, dark brown. A common weed of roadsides and second growth areas, widely distributed in the American tropics. CANAL zone: hetw Gorgona & Mamci, Pittier 2240 (F). chiriqui: vie of Boquetc, Davidson 586 (F, MO); Valley of Rio Chiriqui Viejo, N of Vokan City, Duke 9030 (MO); from Boquetc to 3 mi N, Lewis et al. 345 (MO), 349 (F, MO), cocle: vie El Valle, Allen 1179 (MO); El Valle de Anton, nr Einca Tomas Arias, Allen 4224 (MO); El Valle de Ant6n, Lewis et al. 2520 (MO); Rio Hata Military Res alon^ Pan Am Hwy, Tyson & Blimi 2556 (MO),DARnN: Ridge NW of Yavi/a, Duke 6532 (MO); 8 mi N of Santa Fe, Agua Eria, Duke 10117 (MO); Cerro Piria(|ue, Tyson et al. 3822 (MO), herrera: vie of Chitre, Allen 1107 (MO), los santos: Punta Mala, Tyson 2710 (MO). 5, Hyptis pulegioules Pohl ex Benlh., Lab. Gen. el Sp. 128, 1833. Mesosphaenini puleg.!odes (Pohl e.\ Henth.) O. Ktze,, Rev. Gen. PI. 2: 527, 1891. Herbs, erect to 60 cm, pubescent with uniseriate appressed hairs. Leaves elliptic-lanceolate, acute, entire or mostly dentate, to ca 3.8 cm long and ca 1-2.5 cm wide, pubescent on both surfaces, more so on the veins beneath. Inflorescences capitula, ca 1 cm wide, axillary, sessile to peduncles ca 1 cm long, the bracts iiliform, ca 3-4 mm long. Lloucrs crowded, very pubescent at their bases; calyx lube ca 2-3 mm long, membranaceous, liie teeth lanceolate, ca 1 nmi long, the cleft between the upper teeth cut more deeply; corolla ca 1-1.5 mm long, the tube naked within, the lips ca 0.2-0.3 mm long, sparingly pubescent; stamens weakly exserted, the anthers 0.1-0.2 nun long; style shortly exserted. Nutlets (3-)4, o\ate, ca 0.5 nun long, pale brown. Infreciuent in Panama; scattered throughout Central America and northern South America. chiriqui: Cerro Vaea, in sa\annas, Vittier 5343 (F)- rANA.\L\: sabanas nr Chepo, Hunter & Allen 55 (MO, in part). ^^ AiNNAI S Ol THE MISSOUHI HOTAiMCAL GARDEN [Vol. 56 6. Hyplis ret'urvata Poit.. Ann. Mus. Hist. N;it. Paris 7: 467, pi. 28, f. 1, 1806. — Fig. 5. Mcsosphticnini wcitrratiini (Poit.) O. Kt/c, Hcv. Ccn. IM. 2: S27, 1891. Ilcrha, annuals, slender, liulul\ pubeseent, to 1 ni. Letues ovate to elliptie, aeute, erenate, the bases rounded to ± obtuse, to 3 cm long and 2 cm wide, lightly pubeseent on both surfaces; petioles ca 0.3 (-4) cm long, hifluresccuccs capitula, ca 1 cm diam, the peduncles slender, ca 1-3 cm long, the bracts filiform to linear, ca 2.5-3 mm long, fimbriolate. Flovcrs subsessile; calyx tubular, ca 3 mm long, becoming saccate at the base and constricted and distinctly recurved above the middle, pubescent near the orifice, the teeth spinulosc, ca 1-1.2 mm long; corolla tube ca 2 mm long, white, the lips ca 0.7-1 mm long, sparsely pubescent; stamens exserted, the filaments glabrous, the anthers 0.2 mm long; style exserted, the branches minute. Nutlets 4, oblong-ovoid, ca 0.8-1 mm long, pale broAvn. Parts of Central America and northern South America. liocAs DEL TORO: Santa Catalina, Hlackucll ct al 2737 (MO), canal zone: Rarro Colorado I, Croat 4737 (MO); s. loc, rppleslicinier s.n. (F). CHiRiyui: llanos W of (uia- laca, Allen 5045 (MO); 12.4 mi N of Davitl. Lenis ct al. 720 (MO); vie of San Icli.x, rittier 5433 (F). Panama: Pacora, 35 mi, .Allen lOOi (MO); San Jose I, disturbed area! Johnston 1294 (MO); Panama, Bro. Paul 527 (MO). 7. Hyi»tis oblongifolia Benth. in DC, Prodr. 12: 12 5, 184S. U.vulcanica Seem., Bot. Voy. Herald, 188, 1854. Mesosphaemm ohlongiioliiini (Renth.) O. Kiz., Rev. Gen. PI. 2: 526, 1891 M.vulcanicuin (Seem.) O. Kt/e., loc. eit., 527. Shnihs, sometimes small trees, to ca 2 m, the branches ± lanate with branched hairs. Leaves o\ate-elliptic, acute, finely serrate, the bases rounded to obtuse, to 8 cm long and 4 cm wide, pubescent on both surfaces, the hairs generally simple and uniseriate above, branched and extremely dense beneath; petioles ca 1-3 cm long, densely pubescent. Inflorescences ± capitulate, axillary, loosely-flowered, ca 0.5-1 cm diam, the peduncles ca 0.5-1 cm long, the capitula sometimes arranged in a cymose pattern, the bracts linear, masked by the dense pubescence, llmvers subsessile; calyx campanulate, ca 2.5-3 mm long, naked within, densely pubescent without, obscurely 10-veined, the teeth spinulosc, ca 2-3 mm long; corolla tube ca 4-5 nun long, laxender, the lower hp ca 1.5 mm long, both lips pubescent; stamens exserted, the filaments pubescent, the anthers ca 0.4 mm long; style long exserted, ca 4-5 mm long, the lobes rounded, small and equal. Nutlets unknown. Common in Mexico and sparsely distributed in Guatamala, Costa Rica, and Panama. CHiRiyui: Roquete, M. E. Terry 1272 (A, F); Terry & Terry 1639 (F). panama: swamp F of Rio Tncumen, Standley 26623 (GH). Although the branched hairs are unique, the density of the pubescence tends to mask this character. 1969] FLORA OF PANAMA (Faiiiil) 169. fahiatae^ 89 ^.9:n ^ Fig. 5. Hyptis recurvata Poit.: A, habit (ca X 1); B, flower (ca X 8); C, mature calyx (ca X 20). A-B, specimen unknown. C, after Croat 4737 (MO). [Vol. 56 90 ANNALS OI THE MISSOURI BOTANICAL GARDLN 8. Hyptis dilalata Bcnth. in DC, Prodr. 12: 103, 1848. — Fiu. 6. //. crcnata var. augustifoUa Iknth. ex Scorn., Bot. Voy. Herald, 187, 1854. Mcsosphacnim dilatatiim (Benth.) O. Ktze., Ilev. Gen. PL 2: 526, 1891. Herb's or subslirubs, pubescent with uniseriate and glandular hairs, to ca 1 ni. Leaves oblong-laneeolate, acute, crenate, tlie bases truncate, to 5 cm long and 1.5 cm wide, pubescent on both surfaces, densely so beneath; sessile. Ju flores- cences, capitula, ca 1-1.5 cm wide, the peduncles 1.5-3.5 cm long, the bracts linear or subulate, ca 7-10 mm long, densely pubescent. Flouers sessile, tufts of uniseriate hairs at the base, each 2-3 mm long; calyx ± campanulate, ca 2.5-3 ri^. 6. Hrptis dilutata Berith.: A, habit (X 1.2 ); B, flower (ca ^ 8); C, stamens (ca X 10); D, nutlets (ca X 8). After Mloi 1015 (MO). 1969] FLORA OF PANAMA (Family 169. iMhiatae^ 91 mm long at maturity, membranaceous, ± 10-ribbed, resin-dotted, the teeth ca 1-1.8 mm long and conspicuously pubescent; corolla tube ca 3-3.5 mm long, blue (?), the lips ca 1 mm long, slightly pubescent; stamens exserted, the fda- ments ± glabrous, the anthers ca 0.3-4 mm long; style long exserted. Nutlets (2-) 4, ovoid, ca 1.5 mm long, pale browji. Panama and northern South America. cocle: betw Paso del Arado & Ola, Pitlicr 5015 (F). herrera: 5 mi S of Ocii, Stern ct ah 1687 (MO), veraguas: vie of Santiago, Alien 1015 (MO). 9. Hyptis oblusiflora Presl ex Benth., Lab. Gen. et Sp. 107, 1833.— Fig. 7. Mesosphaerum ohtiisifloruui (Presl ex Benth.) O. Ktze., Rev. Gen. PI. 2: 525, 1891. Herbs, ± suffrutescent at the base, to 1.5 m. Leaves o\ale, the upper ± elliptic, acute, serrate, tlie bases obtuse to attenuate, to 8 cm long and 3.5 cm wide, sparsely pubescent on both surfaces, more so on the veins beneath, resin- dotted; subsessile to petioles 0.5-1 cm long, hifloresceuces capitula, dense, axil- lary, sessile to peduncles ca 1 cm long; bracts ovate-lanceolate, ca 3-4 mm long. Flowers sessile; calyx short tubular, ca 1-1.5 mm long, ± naked within, teeth ± deltoid, ca 0.7-1 mm long, saccate at maturity and distended by the nutlets; corolla tube 2-3 mm long, white, shghtly pubescent, the lips ca 1 mm long; stamens slightly exserted, the filaments pubescent, the anthers 0.3 mm long; style long exserted. Nutlets (3-)^, ovate, ca 0.6-0.7 mm long, dark brown, pubescent at the distal portion. Costa Rica, Panama, and northwestern South America. BOCAs del toro: Santa Catalina, Blaehuell et al. 2729 (MO); Chiricjuicito to 5 mi S along Rio Guarumo, Lewis et al 2070 (MO), cocle: El Valle de Anton, Lewis et al. 2513 (MO), colon: along Rio Igiianitas, EUas & Kirkhride 1631 (MO). 10. Hyptis brevipes Poit., Ann. iMus. Hist. i\at. Paris 7: 465, 1806. Mesosphaenun hrevipes O, Kt/c, Rev. Gen. PI. 2: 525, 1891. Herbs, annual, erect or sometimes sprawhng, to 60 cm, pubescent, the hairs long and uniseriate. Leaves lanceolate-cHiptic, rarely rhomboid-ovate, acute, doubly or irregularly serrate, the bases attenuate, to 5 cm long and 2 cm wide, pubescent on both surfaces, more so on the veins beneath, resin-dotted; sessile to petioles ca 1 cm long, hifloresceuces capitula, ca 1 cm in diam, the peduncles ca 0.5-1 cm long, the bracts awl-shaped and ciliate, ca 4-6 nmi long. Floivers sessile, tufts of hairs at the bases; calyx ± campanulate, 3-3.5 mm long at maturity, lightly pubescent within near the orihce, obscurely 10-nerved, the teeth spinulose, ca 1.5-2 mm long; corolla ca 3 nmi long, white, the lower lip ca 0.6 0.8 mm long, pubescent, the hairs uniseriate; stamens weakly exserted, the hlaments ± pubescent, the anthers ca 0.2 mm long; styles ca 3 mm long, the lobes small. Nutlets 4, oblong-ovoid, ca 0.6 mm long, the pericarps slightly roughened, dark brown. A widely distributed weed in tropical America, from Mexico to Argentina, but probably native to Brazil (Epling, 1949). CANAL zone: Barro Colorado I, Miller Trail, Shattitek 479 (F). colon: along Rio Sirri, Trinidad Basin, Pittier 4023 (F). 92 ANNALS or THE MISSOURI BOTANICAL GARDEN [Vol. 56 Fig, 7. Hyptis ubtusiflora Prcsl.: A, habit (X 1/2); B, flower (ca X 18); C, stamens (ca X 15); D, nutlets (ca X 30). After Woodson ct ul. 1906 (location unknown). 1969] FLORA OF PANAMA (Family 169. Lahiatae) 93 11. Hyptis braehiata Briq., Ann. Conscr\ . Jard. Bot. Geneve 2: 215, 1898. Mesosphaerum hrachiatum Britj,, loc. cit., pro s\n. Herbs or subshrubs, robust, ± densely soft-pubeseent, to 2 m. Leaves ovate or sometimes ± elliptic, acute, crenateserrate, the bases rounded to obtuse, to 8.5 cm long and 4.5 cm wide, pubescent on both surfaces, densely so beneath; petioles ca 0.5-1 cm long. Inflorescences capitula, axillary, the peduncles ca 1-2 cm long, pubescent, the bracts mostly ovate, ca 6-9 mm long, pubescent. Vlowers sessile; calyx tubular, ca 2 nun long, ± naked within, pubescent and resin-dotted, membranaceous, the teeth spinulose, ca 2.5-3 mm long, cihate be- tween; corolla tube ca 4-5 mm long, the lower lip longer, ca 2 mm long, slightly pubescent and resin-dotted; stamens exserted, the filaments glabrous, the anthers ca 0.2-0.3 mm long. Nutlets ca (3-)4, flattened ovoid, 2 depressions near the base, 1.4-1.5 mm long, the pericarp verrucose. Costa Rica, Panama, Colombia, and Venezuela. CANAL zone: hill opposite Paraiso Development, Dxvyer 7152 (MO), cocle: hills S of EI Valle de Anton, Allen 28 i 6 (F). Panama: alon^ nl nr top of Cerro C^ampana, Duke 5963 (MO). 12. Hyplijs alrorubens Poit., Ann. Mus. Hist. Nat. Paris 7: 466 pi 27, /. 3, 1806. — Fig. 8. Mesosphaerum atroruhcns (Poit.) O. Ktzc., Rev. Gen. PI. 2: 525, 1891. Herbs, slender, prostrate, rooting at the nodes, stems to 3 m long, lightly pubescent. Leaves ovate, acute, crenate-serrate, the bases rounded and attenuate, 2.5-3.5 cm long and 1-1.5 cm wide, sparsely pubescent, resin-dotted; petioles ca 0.5-1.5 cm long, narrowly margined. Inflorescences capitula, axillary, hemis- pheric, 8-10 mm in diam, the peduncles ca 0.5-2.5 cm long, the bracts obovate, 4-6 mm long, ciliate. Fknvers sessile, tufts of hair at the bases; calyx tubular, ca 3.5-4 mm long at maturity, naked within, the teeth acicular to setaceous, ca 1-1.2 mm long, pubescent between; corolla ca 4-5 mm long, white, the lower lip ca 1 mm long, pubescent; stamens shortly exserted, the filaments lightly pubescent, the anthers ca 0.2 mm long; gynobase weakly lobed, the style ea 4-5 mm long, minutely bifid near the tip. Nutlets 4, ovoid-oblong, ca 0.6 0.8 mm long, the pericarp verrucose. Mexico, Guatemala, Panama, the West Indies, and northern South America. BOCAS DEL TORO: Colon I, SW of Bocas del Toro at Macaw Hills, von Wedel 526 (MO). Known only from a single collection in Panama, Hyptis atrorubens has been characterized as a widespread tropical weed (Epling, 1937) — its paucity in this area may be due to poor collecting. Superficially, it resembles H, lantanaefolia, but lacks the pilose-annulate condition of the calyx tube associated with the latter species. 13. Hyptis capitata Jacq., Coll. 1: 102, 1787. Mesosphaerum eapitatum (Jacq.) O. Ktze., Rev. Gen. PL 2: 525, 1891. Hyptis capitata var. pilosa Briq., Ann. Conscrv. Jard. Bot. Geneve 2: 224, 1898. 94 [Vol. 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 8. Hyptis atroruhens Poit.: A, habit (ca X 1); B, flower (ca X 6); C, stamens (ca X 10); D, nutlets (ca X 10). Probably after i oh Wcdel 526 (MO). 1969] FLORA OF PANAMA (Family 169, Lahiatae^ 95 Herbs, perennial, sparsely pubescent, to 2 ni. Leaves ovate to ± rhomboid, acute, irregularly serrate, the bases obtuse to ± attenuate, to 12 cm long and 6 cm wide, glabrous to sparsely pubescent; subsessile to petioles ca 2-3 cm long. Inflorescences capitula, densely-flowered, axillary, 2-3 cm in diam, the peduncles ca 2-7 cm long; the bracts ovate-lanceolate, ca 5-7 mm long, sparsely pubescent, frequently reflexed and hidden at maturity. Floirers sessile; calyx tubular, ca 6-8 mm long at maturity, membranaceous, pilose-annulate within above the nutlets, sometimes only sparsely so, teeth spinulose, ca 1.5-2 mm long; corolla tube ca 3-3,5 mm long, white, the lower lip larger, ca 1 mm long, pubescent; stamens exserted, the filaments ± glabrous, the anthers ca 0.2 mm long. Nutlets (3-)4, ovoid, the lobes of the gynobase adhering, ca 1-1.2 mm long, pale to dark brown, the pericarp ± smooth. A common weed of tlie American tropics, from Mexico to Colombia and X'cne/uela and west to Peru and Ecuador. BOCAs DEL toko: rcgion of Almirantc, Cooper 82 (F). canal zone : Barro Col- orado I, Fairchild Pt, Ehinger 284 (MO); Wetmore & Ahhe 59 (F); Barro Colorado I, nr Laboratory, Wctniore & Abbe 172 (F). CiiiRiyui: vie of Boquctc, Woodson £r Schcry 798 (MO), cocle: s. loc, Blum & Tyson 1888 (MO); El Vallc de Anton, Lewis et ah 2527 (MO), darien: vie of Santa Fc, Duke 9489 (MO); vie of Manrunc, Kirkhridc & Bristan 1585 (MO); vie of Tueuti, Terry & Terry 1402 (MO); Piinta Mala, Tyson 2716 (MO); Tonosi, on river bank, Tyson et ah 2998 (MO), los santos: vie of headwaters of Rio Pedrogal, Lewis et al 2970 (MO), panama: Isla del Rev, Duke 9522 (MO), 9567 (MO); E slope of Cerro Jefe, Tyson 3414 (MO), san blas: mainland opposite Ailigandi, Lewis et al 69 (MO). 14. Hyptis lanlanaefolia Poit,, Ann. Mus. Hist. Nat. Paris 7: 468, i. 29, /. 1, 1806. Mesosphaerinn laiitanifolium (Poit.) O. Ktze., Rev. Gen. PI. 2: 525, 1891. Herbs, perennial, procumbent to ± erect, to 1.5 m, incanoiis in the upper parts. Leaves ovate-elliptic, acute, crenate-serrate, the bases attenuate, sparsely pubescent above, the hairs appressed on veins beneath; sessile to petioles ca 0.5 cm long. Inflorescences capitula, densely-flowered, axillary, the peduncles to 4 cm long, the bracts ovate, sometimes broadly so, 5-7 mm long, the margins crenate. Flowers sessile, tufts of hair at the bases; calyx funnelform-tubular, the tube ca 1.5-1.8 mm long, pilose-annulate within above the nutlets, the teeth spinulose, ca 2.5-3.0 mm long; corolla tube ca 4-5 mm long, white or sometimes pale pink, the lower lip ca 2 mm long, pubescent; stamens ± mcluded in the lower lip, the hlaments pubescent near the base, the anthers ca 0.3 mm long; style exserted or included in the lower lip. Nutlets (3-)4, ovoid, pale brown, ca 1 mm long. A common herb of grassland areas, and widely distributed from Mexico south to Sao Paulo, Brazil, west to Bolivia, and also in the West Indies. CHiRiQui: vie of Boquctc, Woodson & Schcry 795 (MO), cocle: El Vallc. Dwyer 1823 (F, MO). PANAMA: Cerro Campana, Duke 8660 (MO), Ehiu^er 915 (MO), McDaniel 6816 (MO); San Jose I, Bald Hill, Johnston 46 (MO); nr Arraijan, Woodson et al 1405 (MO). [Vol. 56 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN 15. Hyplis savannaruin Briq., Ann. Conserv. Jard. Bot. Geneve 2: 216, 1898. Mcsosphaennn savannarum T?riq., loc. cit., pro syn. Herbs, perennial, coarse, to 3 m, ± glabrous. Leaves lanceolate-ovate, the upper ± lanceolate, acute, serrate, the bases obtuse or attenuate, to 13 cm long and 4 cm wide, sparsely pubescent on the veins, resin-dotted; ± sessile. Inf lores- cences capitula, dense, axillary, ± globose at maturity, the peduncles 2.5-5 cm long, the bracts lanceolate or awl-shaped, ca 1 cm long, ± glabrous. Flowers sessile; calyx tubular-campanulate, ca 3-3.5 mm long at maturity, pilose- annulate within above the nutlets, the teeth acicular, ca 3-3.5 mm long, pu- bescent between; corolla tube ca 6-7 mm long, white and purple, the lower lip ca 1.5-2 mm long, hghtly pubescent; stamens exserted, the filaments lightly pubescent, the anthers ca 0.4 mm long; ovary 4-lobed, the gynobase slightly elongated, becoming lobcd, the style minutely bifid near the tip. Miitlets 4, oblong-flattened, the lobes of the gynobase sometimes adhering, ca 1.5-1.7 mm long, brown. Guatemala, El Salvador, Costa Rica, Panama, Colombia, and Peru. PANAMA: vie of Pacora, Allen 1005 (MO); Cerro Campana, SE slope, Lcxiis ct ah 31i4 (MO). Hyptis savannarum is closely related to H. couferta and can be confused with it, especially with the narrow -lca\ed forms of the latter. However, the larger llowers and more spherical capitula, as well as the glabrous bracts of H. savan-^ narum proxide characters that e / 16. Hyptis conferta Pohl ex Bcnth., Lab. Gen. et Sp. 112, 1833. Mesosphacrum coufcrtiim (Pohl ex Bcnth.) O. Ktzc, Rev. Gen. PL 2: 526, 1891. Hyinh comtricta Briq., Ann. Conserv. Jard. Bot. Geneve, 2: 217, 1898. Herln, perennial, coarse, to 1.5 ni, appressed-pubescent on the branches. Leaves lanceolate or narrowly elliptical (in Panama), acute, serrate, the bases attenuate, to 18 cm long and 2.5 em wide, glabrous or densely soft-pubescent above, glabrous or pubescent on the veins beneath, the midrib and secondary veins pronounced; sessile to petioles ca 0.5 cm long. Inflorescencei, capitula, dense, axillary, ± hemispherical, the peduncles ca 1-4 cm long, the bracts lan- ceolate or awl-shaped, ca 8-9 nun long, densely pubescent, rioners sessile, tufts of hairs at the bases; calyx tubular-campanulate, ca 3-4 mm long at maturity, pilose-annulate within above the nutlets, the teeth acicular to setaceous, ca 3-4 mm long; corolla tube ca 4.5-5 mm long, white and rose, the lower lip ca 1.5- 1.8 mm long, the lips and tube lightly pubescent; stamens exserted, the filaments sparsely pubescent near the base, the anthers ca 0.4 mm long; gynobase some- what elongated, becoming lobcd, the style minutely bifid near the tip. Nutlets 4, oblong-flattened, the lobes of the gynobase adhering, ca 1.5-1.6 mm long, brown. A connnon species in the American tropics, found mostly at lower elevations, from Mexico to southern Brazil, and west to Bolivia. CHiRiyui: Boquete, Pitticr 2862 (GH). Panama: rd betw Chepo and Panama, Dodge ct ah 16648a (MO); savanna nr Rio Pacora and Chepo Hwy, DuJu; 5911 (GH). 1969] FLORA OF PANAMA (Family 169. Labiatae^ 97 5. IMARSYPIANTHES Marsypianthe 1833. Herbs prostrate or erect, often somewhat viscid. Leaves opposite, simple, petiolate, pinnatcly-nerved, membranaceous or rugose, generally dentate. Inflores- cences capitula, loosely-flowered and pedunculate, or rarely flowers solitary, the bracts hnear-lanceolate to ± ovate or awl-shaped. Flowers sessile or shortly pedi- cellate; bracteoles not apparent; calyx campanulate or funnelform, ± actinomor- phic, with 5 ± equal lobes, lanceolate to deltoid; corolla tubular, bilabiate, the upper lip 2-lobed, the lower lip prominently 3-lobed, the middle lobe saccate; stamens 4, paired, mounted near the top of the corolla throat, decurved into the lower saccate lip, the filaments short, distinct, the connective not developed, the anthers with 2 functional thecae; ovary 4-lobed, the gynobase unlobed, the style bifid near the tip, the branches ± even. Nutlets 4, attached laterally to a columnar extension of the gynobase and ± open along this line at maturity. Three species in tropical and subtropical America: M. chamacdrys in the West Indies and Mexico, south to Brazil and Peru; and two closely related species, M. montana Benth. (including M. foUolosa Bcnth.) from Brazil and M. hassleri Briq. from Paraguay and Brazil. Useful reference: Ephng, C. C. Supplementary notes on American Labiatae. VII. Brittonia 12: 140-150, 1960. 1. Marsypianthes chamaedrys (Vahl) O. Ktze., Rev. Gen. PI. 2: 524, 1891.— Fig. 9. Clinopodium chamacdrys Vahl, Symb. 3: 77, 1794. Herbs, annual, prostrate or semi-erect, to 0.7 m, glandular-viscid and often with attached grains of sand or dirt. Leaves ovate, acute, crenate-serrate, the bases obtuse, subtruncate or cordate, to 7 cm long and 2 cm wide, both surfaces sparingly villous but often viscid; petioles ca 1-2.5 cm long. Jnfl itula, ± loosely flowered, peduncles ca 1-2.5 cm long, the bracts lanceolate, sometimes lanceolate-ovate, ca 8-10 mm long, fimbriolate. Floorers sessile or with pedicels to ca 2 mm long; calyx campanulate, ca 6-8 mm long at maturity, per- sistent, 10-ribbed, pubescent, membranaceous, the teeth deltoid; corolla tubular, purple, ca 7-9 mm long, bilabiate, the upper lip 2-lobed, the lower 3-lobed, the lips weakly pubescent; filaments pubescent, the anthers ca 0.5 mm long; style slender, ca 2-4 mm long, the stigma 2-lobed, the one acute. Nutlets (3-)4, ca 2 mm long, pale brown. Mexico, Central America, the West Indies, south to Brazil and Peru. CANAL ZONE: Coco Solo, US Army Trop Test Center, Mine Emplacement Center, Divycr & Duke 7909 (MO), chiricjui: Boquete, navhhon 603 (MO); llanos betvv Boquete & David, Ebinger 746 (MO), panama: Cerro Campana, Duke 5995 (MO); Altos del Rio Pacora, Lewis ct ah 2354 (MO). 6. SCUTELLARIA Scutellaria L., Sp. PL 598, 175 3. Herbs, mostly perennials, or half shrubs, erect or procumbent, rarely climbers, glabrous or pubescent. Leaves simple, opposite, petiolate or subsessile, pinnately- [Vol. 56 98 ANNALS or THE MISSOURI BOTANICAL GARDEN Fig. 9. Marsypianthes chmnacdrys (Vahl) O. Ktzc; A, habit (X Vz); B, flower (ca X 5); C, nutlets (ca X 12); D, stamens (ca X 8). Specimen unknown. nerved. // vcrticillaslcrs, or flowers solitary and opposite, or irreg- ularly placed in the axils of bracts, the bracts leaf-like and persistent or deciduous. Fluwers generally pedicellate; bracteoles usually apparent; calyx ± saccate, mem- branaceous, bilabiate, the lips entire, rounded and becoming concave, the upper 1969] FLORA OF PANAMA (Family 169. Lahiatae^ 99 lip deciduous and usually with a transverse squama becoming an erect scutellum at maturity, the lower lip persistent; corolla tubular, bilabiate, generally long- exserted, red, orange, yellow, blue, violet, or white and one of the previous colors, the tube cylindrical and ± sigmoid, arcuate, or more sharply bent, the upper lip cup-shaped or galeate and ± joined obliquely to the lateral lobes of the lower lip, glabrous or pubescent within or without; stamens 4, paired, included in the upper lip, the pairs attached at different levels on the corolla tube, the filaments distinct, the connective not developed, the lower anthers with 1 functional theca; ovary 4-lobed, gynobase generally conspicuous and asynnnetrical, variously de- scribed or functioning as a gynophore, the style long and slender, bifid near the tip, the upper style branch small or not apparent. JSiitlets ca 3-4, attached at the base, ± ovoid or spherical, the pericarp variously tubercular or verrucose, some- times provided with a band near the equator. A large genus of ca 180 species found in most parts of the world. In the New World it ranges from the Arctic Circle to Tierra del Fuego and is represented by about 110 species. The number of species found in Panama is uncertain; Epling in the original manuscript cited only S. glabra Leonard, but there arc at least two, and possibly four, additional species. Scutellaria is probably the most distinctive genus in the Labiatae, chiefly be- cause of the pccuhar structure of the calyx, the squama or scutellum, (well- developed in all but 4 or 5 species) seen at anthesis. (1942) larly homogeneous for its si/e and distribution, lie bases the sections largely upon characters more often associated with specific differentiation, which then becomes correspondingly tenuous. Specimens must have mature flowers, since corolla length is a constant character in Fpling's ke}s both to sections and species. For this reason two collections cannot be identified: Stern et ai 1031 (MO), from Cliiriqui, has only two immature flowers and ambiguous vegetative characters; Leu is et ah 3522 (MO), from the Province of Panama, lacks corollas and although similar to S. purpurascens Swart/ does not appear to be conspecific with it. Useful references: Epling, C. C. The American species of Scutellaria, Univ. Calif. Publ. Bot. 20: 1-146, 1942. Leonard, F. C. The North American species of Scutellaria. Contr. U.S. Natl. Herb. 22: 703-748, 1927. a, Subshrubs, robust; at least some leaves > 12 cm long 1- S. glabra aa. Herbs or shrubs; leaves < 8 cm long. b. Herbs; leaves deltoid-ovate; corolla 12-15 mm lon^^ 2. S. piirpurasccns bb. Shrubs; leaves elliptic; corolla 20-25 mm lon^ 3. S. Icnisiana 1. Scutellaria glabra Leonard, Contr. U.S. Natl. Herb. 22: 715, 1927. Subshrubs, erect, to 1.5 m, mostly glabrous. Leaves ovate or elliptic-ovate, subentire or undulate, the bases shallowly cordate, to 18 cm long and 8 cm wide, glabrous; petioles ca 3-5 cm long, hifloresceuces verticillastcrs, l-2'flo\vered, arranged in a false raceme, ca 10-2 5 cm long, the bracts lanceolate, ca 3-5 mm 100 [Vol. 56 ANNALS OF THE MISSOURI BOTANICAL CAHDEN Fig. 10. Scutellaria glabra Leonard: A, habit (X '/2); B, iloucr (ca X 2.5); C, sta meiTi (ca X 2.5); D, nutlets (X 5). After White 106 (MO). 1969] FLORA OF PANAMA (Family 169. iMhiatae') 101 long, early deciduous. Flowers with pedicels ea 2-6 mm long; bracteoles 2, minute, < 0.5 mm long; eal)'x 2.5-3 mm long in flower, enlarging to ea 5-6 mm in fruit, the squama of the upper lip extending to ca 4-5 mm in fruit; corolla red or blue, ca 2-2.5 cm long, the tube constricted abo\'e the cal\x, then weakly expanded, the upper lip arched, the lower lip flat, naked within; lower stamen pair attached 13-15 nun from tiic base, the anthers ca 0.6 nnn long; gynobase conspicuous, asymmetrical and gynophore-like, the style long and slender, the branches unecjual. Nutlets (3')4, semi-spherical, black, evenh' tuberculate, ea 2 mm in diameter. CHiKi^L I: upper Cluritjui Vicjo \ allcv, trail from l^aso Ancho to Monte Lirio, AUoi 1478 (MO); Chiriqui Vicjo Valley, White 106 (MO). 2. Scutellaria purj>iirascens Swart/, Prodr. 89, 1788. Herbs, perennial, erect or somewhat decumbent, to 40 cm, short-pubescent throughout, l.eax^es deltoid-o\ate, acute, ± erenate, the bases rounded or shal- loA\ly cordate, to 7 cm long and 4 cm wide, sparsely pubescent to glabrous above, pubescent on the veins beneath; petioles to 3 cm long, hiflorescences \erticillas- ters, mostly IHowcred, the bracts lanceolate and ciliate, 2 4 mm long. Flouers with pedicels to 6 mm long at maturity; bracteoles 2, minute, ca < 0.5 nnn long; calyx 2 mm long in flower, ca 4-^5 mm long in fruit, the squama then 4-5 mm long, sparsely and minutely pubescent; corolla blue, 12-15 nnn long, naked within or very sparsely pubescent; lower stamen pair attached 6-8 mm from the base; gyno- base conspicuous and asvnnnetrical, the style apparentK unlobed. Nutlets (2-)4, ± spherical, brown, ca 1.5-1.7 mm in diameter. The Lesser Antilles, Costa Hica, Panama, Colombia, Venezuela, and Brazil. BOCAs DKL TORO: Sixola Vallc}', Sihubi Falls, Bowlec & Rowlce 378 (NY), canai zone: Garden at Gatiin Station, Jlnycs 59 (NY), darikn: 0-4 mi up Puo Sabana from Santa. Fe, Duke 4135 (MO); Cana & vie, WilUanis 764 (NY), 949 (NY). 3. Scutellaria lewisiana Nowicke, Phytolo^ia (in press). ^ — Fig. 11. Shrubs, tbe upper stems maroon and with curled pubescence. Leaves elliptic, acute, serrate, the bases obtuse, to 6 cm long and 2 em wide, glabrous, resin- dotted, dark-green above, pale green beneath; petioles ea 1-2 cm long. Inflores- cences verticils, 2'flo\vered, ca 1 cm distant, arranged in a false raceme, the bracts elliptic, ca 3-4 mm long. Flowers with pedicels ca 4-6 mm long; bracteoles 2, 0.5 mm long; calyx ca 4 mm long in flower, softly pubescent; corolla ca 23-2 5 mm long, lavender, the lower lip white, ± straight, hghtly pubescent; lower stamen pair attached on the upper half of the corolla tube, the anthers ca 0.6 mm long; gynobase conspicuous, asymmetrical, the style unlobed. Nutlets unknown. CHiRigui: Caldera-Chiriquicito trail, betw Qucbrada Hondo and Divide, Kirkhr'ule & Duhe 920 (holotype MO). 102 ANNALS Ob THE MISSOURI BOTANICAL GARDEN [Vol. 56 Fig. 11. Scutellaria hwidana Nowickc: A, habit (ca X 5/7); B, Howcr (ca X 2) After Kirkhride &r Duke 920 (MO). 7. OCIMUM Ocinium L., Sp. PI. 597, 1753. Herbs, annual or perennial, or small shrubs, the stems glabrous or pubescent. Leaves opposite, simple, pinnateh-nerved, mostly petiolate. ]}iflorescences verti- cils, ca 6-flo\\ered or sojnetimes lO-llowered, appearing overall as a spike or a raceme, the bracts leaf-like. Flowers mostly pedicellate; bracteoles not apparent; calyx ± campanulate, frequently deflexed in fruit, bilabiate, the upper lip entire, rounded, the margins deciuTent partially or completely to tlie base of the calyx tube, the lower lip 4't()othed, the teeth aristate and unequal; corolla tubular, enclosed or exserted, bilabiate, the upper lip 4-lobed, the lower entire and flat or concave; stamens 4, paired, e(|ual to the corolla tube or exserted, the fdaments distinct, the upper pair naked, the lower pair glabrous or pubescent, with or 1969] FLORA OF PANAMA (Family 169. Lahiatae^ 103 without an appendage at the base, the anthers with 2 functional thecae; ovary 4-lobed, the gynobase lobed or unlobcd, the lobes sometimes as high as the nutlets at anthesis, the style slender, bifid near the tip, the branches equal or unequal. Nutlets ca 4, attached at the base, ± spherical, the pericarp smooth or warty. Commonly called Basil, the genus consists of about 50 to 60 species in the warmer regions of the world. An unusual representation in Panama with the presence of O. gratissiuiiim L. as well as the widespread weeds O. nncrajithuui W'illd. and O. americaniim L., and the apparent absence of O. hasiliciiui L., the most frequently cultivated species. a. Lower stamen pair with naked filaments; mature calyx 7-9 mm long, the de- current margins of the upper calyx lobe extending to the base 1. O. micranihum aa. Lower stamen pair w ith pubescent or appendaged filaments, calyx ca < 6 mm long at maturity, the deeurrent margins of the upper calyx lobe not extending completely to the base. b. Herbs < 0.5 m; lea\es < 2 cm wide, sul)entire; calyx pubescent at the base with stiff hairs ca 2 mm long, the deeurrent margins of the upper lip extend- ing almost to the base 2. O. amehcanmn bb. Subshrubs, to 2 m; leaves > 3 cm wide, ± crenate; calyx softly pubescent at the base, the deeurrent margins of the upper lip ca Vs to V2 as long as the calvx 3.0. gratissimum 1. Ocinium micrantlium W'illd., Fnum. Ilort. Berol. 630, 1809. — Fig. 12. Herbs, annuals, erect to 1 m, the branches cymose-divaricate, glabrous. Leaves rhomboid, obovate or ovate, acute, serrate, the bases attenuate, to 7.5 cm long and 3.5( 5 in the type) cm wide, resin-dotted, ± glabrous to lightly pubescent on the veins; petioles to 4 cm long. ]}ijloresceuces verticils, each ca 6'flowercd, 0.5-1.5 cm distant, the false raceme to 15 cm long, bract leaf-like, ca 4-6 mm long. Flowers with pedicels ca 4-5 nun long, pubescent; calyx 7-9 mm long at maturity, membranaceous, pubescent on the veins, the upper lobed rounded, 3-veined, margins deeurrent on the calyx tube to its base, the lower ca 6-8-veined, teeth upwardly curved; corolla tubular, ca 3-4 mm long, the upper 4-lobed, the lowXT entire; filaments naked, the anthers ca 0.5 mm long; ovary 4-lobed, gyno- base ± unlobed, the style slender, ca 3-4 mm long, the branches unequal. Nutlets (3-)4, ca 1.7-2.0 mm long, the pericarp dark brown, minutely warty. W^idely distributed in the American tropics. BOCAS di:l toro: Old Bank I, von Wcdel 1865 (MO); Colon I, von Wcclel 2838 (MO). CANAL ZONE: MiiallorL's Lake, White 199 (MO), dakien: Yaviza, alons Qucbrada Uvital off Hio Chucunaquc, Duke 4999 (MO); Canipamcnto Bucna Vista, Stern ct al 814 (MO), hkrrera: Pcsc, Allen 803 (MO); Ocu, Allen 4090 (MO). LOS santos: 3 mi S of Carrcta, Biireh et al 1249 (MO), panama: sabanas nr Chcpo, Hunter £r Allen 68 (MO). SAN HLAs: Mulatuppu, Duke 8481 (MO), 8522 (MO); Pino 1, Elias 1719 (MO); mainland opposite Ailigandi, Leivis et al. 74 (MO). 2. Ocimum amerioanuiii L,, Amoen. Acad. 4: 276, 1759. O. canum Sims, Bot. Mag. 51: ^ 2452, 1824. O. stamineiun Sims, loc. cit. Herbs, annuals, to 40 cm, pubescent with uniseriate curled hairs. Leaves elliptic, acute, subentire, the bases obtuse, to ca 3 cm long and 1 cm wide, sparingly pubescent above, more so on the veins beneath, resin-dotted; petioles ca [Vol. 56 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 12. Ocimiim micranihum Willtl.: A, habit (ca X V2); B, flower (ca X 10); C, sta- mens C^^^ X 8); D, nutlets (ca X 15). Specimen unknown. 1969] FLORA OF PANAMA (Family 169. Lahiatae^ 105 0.5-1 cm long. Influrescences verticils, ca 6 flowered, 1-1.5 cm distant, the false raceme to 17 cm long, the bracts elliptic, ca 3.5-4.5 mm long, the margins pubescent. Fluuers with pedicels ca 2-2.5 mm long; bracteoles not apparent; calyx 4-5 long at maturity, pubescent within and without at the base, the hairs ca 2 mm long, the upper lip rounded and acutely tipped, 3-veined, the decurrent margins extending almost to the base, the lower lip 5-7-veined, the lateral teeth wider; corolla tubular, white to yellow -green, the tube ca 2-3 mm long, the lips ca 1-1.5 mm long, the lower lip ± concave; stamens exserted, the lower pair with a small appendage at the base of the hlament, the anthers ca 0.6 mm long; ovary 4-lobed, the gynobase lobed, the style bifid near the tip, the branches ± equal. Nutlets C3-)4, oblong, ca 1.2 mm long, black, the pericarp ± warty. Frequent in the American tropics. darien: headwaters of Rio Chico, specimen cultivated by Choco Indians, Allen 4627 (MO). 3. Ocimum gralissimum L., Sp. Pi. 1197, 1753. Subshrubs, erect, to 2 m, glabrous to softly pubescent on the younger branches. Leaves ovate or elliptic, acute, crenate to subentire, the bases rounded and weakly attenuate or ± obtuse, to 7 cm long and 4.5 cm wide, resin-dotted, glabrous above and sparsely pubescent on the veins beneath; petioles to ca 4.5 cm long, pubescent. Inflorescences verticils, each ca 6-flowered, 0.5-1 cm distant, the false racemes 10-15 cm long, the bracts ± deltoid, 2-3 mm long, pubescent. Flouers with pedicels ca 1-2 mm long; bracteoles not apparent; calyx 3-4 mm long, membranaceous, resin-dotted, pubescent on the veins and at the base, naked within, the upper lip entire, 3-veined, lateral margins decurrent only on upper Vi of tube, the lower lip 5-7-veincd; corolla tubular, white and purple, ca 2.5 mm long overall; stamens exserted, lower filament pair with conspicuously pubescent appendages at the base, the anthers ca 0,6 mm long; ovary 4-lobed, gynobase weakly lobed, the style bifid near the tip, the branches ± equal. Nutlets (3-)4, ± spherical or slightly elongated, ca 1.2 mm long, the pericarp warty. The tropics of America and Africa, and Ceylon, Java, and the East Indies. VERAGUAS: Canazas, Tysoti 3651 (MO). 8. TEUCRIUM Teucrium L., Sp. PI. 562, 1753. Herbs, perennial, rarely annual herbs or half-shrubs, branching from the base or in the inflorescences. lobed parted, pctiolate or subsessile, pinnately-nerved. Inflorescences verticillasters, 1-00 flowered, solitary and axillary, or arranged in spikes, the bracts leaf-Hke. Flowers subsessile or with slender pedicels; bracteoles not apparent; calyx tubular or campanulate, sometimes becoming saccate at maturity, actinomorphic or weakly zygomorphic, 10-veined, the teeth ± equal, or the lower tooth larger; corolla tubular and short, or funnelform and slightly exserted, scarcely bilabiate, seem- ingly 1 -lipped and 5-lobed, in actuality the upper hp 2-lobed, the lobes laterally [Vol. 56 106 ANNALS OF THE MISSOURI BOTANICAL GARDEN placed, the lower lip 3-lobed, the middle lobe largest, the 2 lateral lobes about equal to the lobes of the upper lip; stamens 4, paired, exsertcd, the lower pair longer, tlie filaments distinct, the connective not developed, the anthers with 2 functional thecae; ovary 4-lobed, the style terminal, long and slender, bifid near the tip. Nutlets ca 4, temporarily united, ovoid to wedge-shaped, glabrous or pubescent, the pericarp sculptured or smooth. A ± cosmopolitan genus of 100 species with a center of distribution in the Mediterranean area; one species has been collected in Panama. Useful reference: McChntock, E., and C. C. Epling. A revision of Teiicrinm in the New World, with observations on its variation, geographical distribution, and history. Brittonia 5: 491-510, 1946. 1. Teucrium vesicariuni Mill., Gard. Diet. ed. 8, 1768. — Fig. 13. Herbs, perennials, to 1 m, rootstocks creeping, pubescent with curved hairs or with spreading ± glandular hairs. Leaves opposite, ovate or oblong-ovate, acute, crenate-scrrate, rounded or subtruncate at the base, to 7 cm long and 3.5 cm wide, the upper surfaces pubescent or glabrous, the lower paler with soft curved hairs, or ± glabrous, sometimes with longer straiglil hairs on the veins; petioles ca 2-2.5 cm long, hifloresceuces verticillasters, 1 -flowered, arranged as a terminal spike, to ca 1 7 cm long, the bract leaflike, lanceolate, ca 4-6 mm long, pubescent beneath. Floners with pedicels ca 1 mm long; calyx weakly bilabiate, 5-7 mm long, the upper lip 2-toothed, the lower lip 3-toothed, the teeth deltoid, pubescent, inflated and ± saccate at maturity; corolla tubular, exsertcd or equal to the calyx, the upper small and inconspicuous, the lower ca 4 mm long; stamens 4, exsertcd, the anthers 0.5 mm long. Nutlets wedge-shaped, glabrous, wrinkled. 2-2.5 mm long. A variable species in tropical America. BOCAS DEL TORo: Water Valley, Chiriqiii Lagoon, von Wcdcl 2668 (MO); Nit-vccita 50 m, Woodson ct al. 1857 (MO). 9. LEPi;CHlNlA Lepechinia W'illd., Hort. Berol. 1: 21, 1803. Sphacelc Benth. in Edwards, Bot. Reg. 1.^: pi. 7 28 9, 1829. Herbs, half-shrubs, or shrubs. Leaves opposite, simple, generally petiolate, pinnately-nerved. hifloresceuces verticils, loosely or densely flowered, arranged in open or crowded spikes, or sometimes reduced to a single flower, the bracts leaf-like. Floivers sessile or pedicellate; bracteoles generally not apparent; calyx campanulate, actinomorphic to weakly bilabiate, ca 10-veined, often membranaceous, enlarg- ing at fruiting time, the teeth sometimes conivent; corolla tubular to campanulate, enclosed or shortly exsertcd, white, blue, or purple, naked within or a ring of hair-like appendages near the base, weakly bilabiate, the upper lip weakly 2-lobed, the lower 3-lobed, the lobes ± flat; stamens 4, paired, the filaments distinct, the connective not developed, the anthers with two functional thecae, the thecae 1969] FLORA OF PANAMA (Family 169. Lahiatae) 107 Fig. 13. Tcucrmm vcsicarium Mill.: A, habit (X V2); B, flower (ca X 5); C, section of flower with stamens and nutlets (ca X 5). After ro)i Wcdcl 1668 (MO). diverging or parallel; ovary 4-lobcd, the gynobase generally uniform and thick, the style gynobasie, bifid near the tip, the branches ± flattened or awl-shaped. Nutlets 4, attached at the base, ovoid, black or brown, smooth or puberulcnt. [Vol. 56 108 ANNALS OF THE MISSOURI BOTANICAL GARDEN Levcchinia is a predominantly iNcw World genus of about 38 species, ranging from California to Chile. Epling (1935) reduced SpJuwele Benth, to Lepechiuia and removed Dckiuia Mart. & Gal., which Bricjuet O cihiatae, Naturl. PHan/en- fam. IV 3: 183-375, 1897) had included in his treatment of the genus. Usaful reference: Epling, C. C. A S)nopsis of the tribe I.epechiJiieae (Lahiatae). Brittonia 6: 352-364, 1948. 1. Lepechinia st-hiedeana (Schlecht.) Vatke, Verh. Bot. Vercins Prov. Bran- denburg 17: Sitzungsber. 36, 1875.— Fig. 14. Stachys schiedcana Schlecht., Linnaea 7: 398, 18 32. Herbs, procumbent, the stems ca 20-40 cm long, pubescent throughout, the hairs uniseriate. Leaves elongate-deltoid, acute, the bases truncate-cordate, to 5 cm long and 2 cm wide, pubescent on both surfaces, densel> so beneath, the upper surface rugose-areolate; subsessile to petioles to 6 cm long on basal leaves. Inflores- cences verticils, each 6-flowered, crowded at anthesis, becoming remote at fruiting, the bracts leaf-like, enlarging with maturation. Fhmers with pedicels 2-3 mm long; bracteoles not apparent; calyx campanulate, 4-6 mm long in flower, enlarg- ing to 10-11 mm in fruit, reticulate between the veins, membranaceous, pubescent at anthesis, weakly bilabiate, the upper lip 3-toothed, the lower 2-toothed, the teeth ca equal in length or the lower teeth slightly shorter; corolla ca 4-5 mm long, blue or purple, a ring of hair-like appendages near the base; stamens slightly exserted, the thecae ± diverging; gynobase present, the style lobes somewhat flat- tened. Niitlets (3-)4, ovoid, ca 1.8-2 mm long, the pericarp reticulate. Central America. CHiRioui: Ccrro Copctc, summit and SW slope, Allen 4HH7 (MO); Volcan de Chiriqui, Potrero Mulcto to summit, Woodson & Schcry 428 (MO). 10. STACHYS Stachys L., Sp. PI. 580, 1753. Herbs, annual or perennial, rarely half shrubs or shrubs, usually pubescent. Leaves simple, opposite, pinnately-nerved, petiolate or subsessile. Inflorescences verticils, 4-oo flowered, appearing as spikes, the bracts leaf-like or absent. Flou perfeet, subsessile or shortly pedicellate; bracteoles generally apparent; calv: ers X weakly zygomorphic, the teeth subequal and frequently aristate or spinulose-tipped; corolla tubular, exserted or included, purple to pink, the tube frequently con- stricted above the nutlets, sometimes witli a spur on the lower side, bilabiate, the upper lip subgaleate and 2-lobed, the lower lip 3-lobed, the central lobe largest, sometimes with a pilose annular ring above the nutlets; stamens 4, paired, slightly exserted or inclosed in the upper lip, the fdaments distinct, glabrous or pubescent, the connective not developed, the anthers with 2 functional thecae, the thecae strongly divergent or parallel; ovary 4-lobed, the gynobase unlobed or forming a nectarium, the style bihd near the tip, the branches equal and acute. Nutlets ca 4, attached at the base, ± obovate. 1969] FLORA OF PANAMA (Family 169. iMhiatae^ 109 Fig. 14. Lcpechinia schiedcana (Schlecht.) Vatkc: A, habit (ca X V2); B, flower (X 7); C\ anther top view (ca X 15), C-, anther side view (ca X 15); D, nutlets (ca X 10). After Woodson 1036 (location unknown). Up to 200 species in the North Temperate Zone and extending to South America and South Africa. Useful reference: Epling, C. C. Preliminary revision of American Stachys. Repert. Spec. Nov. Beih. 80: 1-75, 1934, 1. Stachys coMtaricensis Briq., Bull. Soc. Rov. Bot. Belgique 30: 240, 1891. — Fig. 15. Herbs, prostrate, rooting at the nodes, the younger branches sparsely pubes- cent, the hairs uniseriate and ± retrorse. Leaves ovate to elongate-deltoid, acute, crenate-serrate, the bases ± ± truncate, ca 3-5 cm long and ca 2-3.5 cm wide, sparsely pubescent on both surfaces and on the margins; petioles ca 1.5-3 cm long, sparsely pubescent. Inflorescences verticils, ca 6-flo\vered, appearing as spike- like racemes, ca 5-10 cm long, the bracts lanceolate, ca 6-10 mm long, reflexed, or the lowermost verticil in the axils of reduced leaves. Flowers subsessile; bracteoles 110 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 Fig. 15. Stachys costuriccnsis Briq.: A, habit (ca X 1); B, flower (ca X 8); C, stamens (ca X 10); D, nutlets (ca X 5). After Woodson & Scficry SSI (MO). 1969] FLORA OF PANAMA (Family 169. Lahiatae') 111 present, awl-shaped, ca 1-2 mm long, pubescent; calyx campanulate, weakly zygo- morphic, the tube ca 2-3 mm long at maturity, the teeth deltoid-lanceolate, ca 2 mm long, pubescent, obscurely nerved but each tooth apex with a spine like tip, the uppermost largest; corolla tubular, 5-6 mm long, ± pink, constricted ca halfway up and with a small spur on the lower side, the upper lip 2-lobcd and concave, ca 2-3 mm long, the lower lip 3-lobed and spreading, ca 3-4 mm long, pubescent within and without, annular-pubescent within near the constriction; stamens ex- serted, the hlaments pubescent, the anthers ca 0.5 mm long; ovary 4-lobed, the equal. gynobase unlobed, the st}le glabrous, bifid near the tip, the branches Nutlets ca 4, ovoid, ca 1-2 mm long, dark brown. Mexico and Central America. CHiHiQui: nr Boqucte, Pitticr 2966 (location unknown, ace. EpHng, 1934); vie of Bajo Mono & Quebrada Chiqucro, 1500 m, Woodson & Schery 552 (MO). Indi X oi Latin Names Numbers in bold fare type refer to descriptions; numbers in ronian type refer to synonyms; numbers with dagger (t) refer to names incidentally mentioned. Ballota suaveolens 86 Clinopodium chamaedrys 97 Coleus 83 blumei 83 Cunila 81 polyantha 81, 8^t Dekinia lOSf Hyptis 72t, 84, 84t atrorubens 93, 93t brachiata 93 brevipes 91 capitata 93 var. pilosa 93 conferta 96t, 96 constricta 96 crenata var. augustifolia 90 dilatata 90 lantanaefolia 93t, 93 mutabilis 87 oblongifolia 88 obtusiflora 91 peetinata 86 pulegioides 87 reeurvata 88 savannarum 95, 96t suaveolens 86 vertieillata 85 vulcanica 88 Labiatac 72t, 73t, 84t, 99t, 108t Lepechinia 104, lOSf schiedeana 106 Lepechineae 108t Marsypianthes 96 chamaedrys 97t, 97 foliolosa 97t hassleri 97t montana 971 Mesosphaerum 84 atrorubens 9 3 brachiatum 93 brevipes 91 capitatum 93 eonfertum 96 dilatatum 90 lantanifolium 95 mutabilc 87 oblongifolium 88 obtusiflorum 91 pectinatum 86 pulegiodes 87 recurvatum 88 savannarum 9 5 suaveolens 86 vertieillatum 8 5 vulcanicum 88 Nepeta mutabilis 87 peetinata 86 Ocimum 102 americanum 103t, 103 basilicum 103t canum 103 gratissimum 103t, lOS micranthum 103t, 103 stamineum 103 Salvia 72t> 72, 73t, 78t, 81t albopileata 80, 8 If alvajaca 78 compacta 77 — var. irazuensis 76 hyptoidcs 75 irazuensis 76 longimarginata 73t, 77 membranacea 76, 77t — var. villosula 76 micrantha 74 occidentalis 74, 74t orbicularis 74 ourophylla 78 I>ermixta 78 polystaehya 77, 77t psilophylla 78, 78t ptcroura 76 tiliaefolia 73t tonduzii 80 trichopes 78, 78t wagneriana 80 Scutellaria 97, 99t glabra 99t, 99 lewisiana 101 purpurascens 99t, 101 Sphacele 106, lOSf Stachys 108, 109t costaricensis 109 schideana 108 Teucrium 105, 106t vcsicarium 106 The previous issue of the ANNALS OF THE MISSOURI BOTANICAL GARDEN, Vol. 55, No. 3 pp. 171-402, was published on April 30, 1969. Published by the Missouri Botanical Garden Press St. Louis, Missouri 63110 f ) I ^ ■- ANNALS VOLUME 56 1 969 NUMBER 2 ^ f Missouri r y CONTENTS '..■■"■■ ■ ■■ ■ ■ ■ ^ - . - m ^ h - r '■- r Viable Soil Algae from the Herbarimn of the Missouri Botanical Garden Bruce C. Parker, Noel Schanen h Richard Rentier . . ... - £ ' - -r * Scanning Electron Microscopy as an Aid to Pollen Taxonomy John E. Ridgxvay &■ John J. Skrarla . . . .... •' ■. _ I" On Tropical Tree Seedhngs. I. Seeds, Seedhngs, Systems, and Systematics James A. Duke . . . ^^^ ^1 ^^P ^^L ^^M ^^F ^^h ^^h ^L ^^ ^^ ^^ ^^^ ^^r ^^B ^^H Adventive Plants New to the Missouri Flora (III) Viktor Muehlenhach AT- ^ Notes on the Galapagos Euphorbieae (Euphorbiaceae) Derek Burch . The Genus Cienfuegosia Cav. (Malvaceae) Paul A. Fryxell . . . The Initiation of Vascular Cambium and Production of Secondary Xylem in Flower Bud Pedicels of Asclepias curassavica L. F. M. Safwat Cytological and Chromatographic Evidence of Interspecific Hybridization in Petalostemon Sally A. Walker The Genus Ho^wa«w/« (Rubiaceae) in Panama John D. Dwyer . . NOTES The Skunk Cabbage in Missouri Erna R. Eisendrath . . . . . Supplementary Notes on the Phytolaccaceae, I. Joan W. Nowicke . ^Ihtroemia (Zygophyllaceae) in Missouri Duncan M. Porter . . 113 121 125 163 173 179 251 261 269 287 288 ¥' 290 VOLUME 56 1 969 NUMBER 2 ANNALS OF THE -.J. - - i Missouri Botanical Garden > - 1 The Annals contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden and the Department of Biology of Washington University. Papers originating outside the r ■ Garden or University will also be accepted. These papers are subject to a charge of $25 per printed page. Editorial Committee . ^ " ■ - ■-. _ -_ * ■— X r r Derek Burgh, Editor, pro tem, Missouri Botanical Garden &■ Washington University - - - -- . - - - - - ^ - ' - Sheri Davis, Assistant to the Editor Marshall Crosby, Missouri Botanical Garden John D. Dwyer, Missouri Botanical Garden £r St, Louis University Joan Nowicke, Missouri Botanical Garden & Washington Universe Duncan Porter, Missouri Botanical Garden & Washington Unive\ Information The Annals of the Missouri Botanical Garden appears three times during the calendar year. Three numbers totaling about 400 pages con- stitute a volume. Beginning with Volume 53, 1966: Subscription Price $15.00 per volume Single Numbers 5.50 each Contents of previous issues of the Annals of the Missouri Botan- ical Garden are listed in the Agricultural Index, published by the H. W. Wilson Comoanv. ■ ' J L Beginning June 1, 1962 the Stechert-Hafner Service Agency, Inc., 31 East lOth St., New York 3, N.Y., became sole agent for the Annals of the Missouri Botanical Garden. The Agency handles all subscriptions, all claims beginning with volume 50 (1963) and all requests for back issues. Out-of-print numbers will be reprinted as may be required. Under the arrangement with the Stechert-Hafner Service Agency, the Agency handles also all purchases of any portion of the Annals subseries known as the "Flora of Panama." -. ^ All matters regarding exchange continue to be handled by the Missouri Botanical Garden, 2315 Tower Grove Ave., St. Louis, Missouri 63110, as in the past. J- The Annals of the Missouri Botanical Garden arc listed in Cur- rent Contents Agricultural, Food and Veterinary Sciences. t Printed by Bardgett Printing and Publishing Company, St. Louis, Mo. 63103. VOLUME 56 1969 NUMBER 2 ANNALS OF TH Missouri Botanical Garden VIABLE SOIL ALGAE FROM THE HERBARIUM OF THE MISSOURI BOTANICAL GARDEN BY Bruce C. Parker/ Noel Schanen, and Richard Renner- Department of Botany, Washington University St. Louis Abstract Soil particles from herbarium sheets of aquatic, bog and terrestrial plants were used as innoculum on Bold's modified Bristol's medium. Forty-six of the 124 innoculations were suc- cessful, and algae were obtained from specimens up to 60 years old. Green algae were the most frequent survivors from plants collected within the last 30 years, and blue-greens from the period before that. Introduction Although many have recognized the importance of desiccation resistence to survival of microorganisms (Evans, 1959; Hortobagyi, 1960; Lund, 1962), few investigators have approached this question through attempts to culture algae from soil samples kept dry in excess of a few years. Bristol (1919) identified 11 genera (7 blue-green, 3 green, 1 diatom) which grew from 16 samples of partially air-dried English soils sealed in lead-capped bottles for 23-70 years. Algae and fungi developed even in the 70-year-dry soil when moistened with Bristol's inorganic salt solution and placed in light for some months. Lipman (1941) attempted to grow algae from fragments of numerous her- barium sheets of green and blue-green algae. The only culture which developed was a specimen of Nostoc commune collected in 1853. Lipman repeated this ex- periment twice, estabhshing a record viability of 86 and 87 years for this species. Becquerel (1942) took dry soil from 10 non-disinfected herbarium specimens of various mosses and fern allies spanning 56-98 years in age. When placed in sterile inorganic salts medium (pH 5) of unreported composition in window light, 14 different genera ultimately developed (4 blue-green, 9 green, 1 diatom). All 1 Present address: Department of Biology, Virginia Polytechnic Institute, Blacksburg, Virginia 24060. 2 Present address: Department of Biology, Macon Junior College, Macon, Georgia 31206. Ann. Missouri Bot. Gard. 56(2): 113-119, 1969. [Vol. 56 114 ANNALS OF THE MISSOURI BOTANICAL GARDEN soils yielded viable algae including the 98-year-old sample from which the diatom Nitzschia palea grew. In contrast to these reports, Parker (unpublished) failed to obtain viable algae from numerous herbarium specimens of lichens 35 to more than 100 years old following mild surface sterilization of the plant fragments. We know of no efforts besides these to treat directly the longevity of algae or population viability under drought conditions. Therefore, we set out during 1967-'69 to examine more extensively the questions treated by Bristol (1919), Lipman (1941), and Becquerel (1942). Methods Using the herbarium of the Missouri Botanical Garden (MO), we obtained, after much searching, 50 specimens of bog or aquatic vascular plants dating from 1831 to 1966 on which small crumbs of dry soil remained. A second series comprising 74 specimens of Gramineae (mostly terrestrial) dating from 1885 to 1966. Unhke the herbaria used by Lipman (1941) and Becquerel (1942), all our source material except the 1966 Gramineae had received some kind of dis- infection. At least back to 1955, all plants at MO were fumigated with carbon tetrachloride, naphthalene and paradichlorobenzene before mounting. Before this time formaldehyde or mercuric chloride were used. After fihng, the specimens are continuously subject to the vapors of the same chemicals used for the initial fumigation. We picked dry soil crumbs no more than a few millimeters in diameter from roots with ethanol-flamed forceps and placed them ascptically into 50-ml cotton- plugged Erlenmeyer flasks or Pyrex screw-cap test tubes, both approximately half full of sterile Bold's (1949) modified Bristol's medium. Cultures were developed in approximately 3000 lux of constant, cool-white fluorescent hght at 22-23° C, then placed in subdued light. After approximately two months, containers with visible growth were examined microscopically to determine the taxonomic affiha- tion of the viable algae, ^\'e made no attempts to identify all algae to species or genus level, a task now requiring elaborate sub-culturing and specialized skills for many groups (Bold & Parker, 1962; Chantanachat & Bold, 1962; Bischoff & Bold, 1963; Brown & Bold, 1964; Cox & Bold, 1966; Koster, 1966; Smith & Bold, 1966; Pearson & Kingsbury, 1966; Allen & Stanier, 1968). As new cultures developed, they too were observed, allowing in most cases, nearly t^vo years for the appearance of visible growth. Results Although visible growth appeared as late as one year in a few cultures, most algae grew to maximum proportions within a few months. In all, 46 of the 124 soil crumbs yielded viable algae contrasting strikingly with results of Bristol (1919) and Becquerel (1942) who obtained viable algae from all their soil samples. Fungi developed in a greater number of cultures than did algae. Table 1 predominantly terrestrial Gramineae. th 1969] PARKER, SCHANEN & RENNER — SOIL ALGAE 115 Date Collected 12-18-66 5-27-62 5- 5-55 3-11-51 4-24-4 3 9-28-62 9- 2-62 9- 2-61 9-17-60 6-17-59 6-24-57 6- 6-57 6- 5-57 8-10-55 4-28-53 8- 6-51 5-29-48 8-19-46 7-13-46 7-11-45 10- 3-43 8- 7-42 9-12-41 Table 1. Chronological List of Herbarium Source Material Yielding Viable Algae. Aquatic Plants (first series) AND GrAMINEAE (sECOND SERIES) Source Plant Algae Idefitified 12-18-66 LiinnocJjaris flava Oscillatoria, chlorococcaccan, Vlothrix (Uronema?) Limnocharis flava Alisma gramineum Isoetes melanopoda Eqtiisetum hyemale 9-26-46 Triglochin sp. Equisetum arvense 9-27-40 Eqidseturn hyemale 5-26-38 Equisetum ferrissii 7-23-3 5 Typha latifolia 4-14-3 5 Equisetum hyemale 7- 9-31 1910 Saggittaria umeata Triglochin maritima Oscillatoria, chloroccacean, Navicula Oscillatoria, Aulosira 2 chlorococcaccan spp. chlorococcacean sp. Phorjnidium? 2 chlorococcacean spp. 2 chlorococcaccan spp. chlorococcaccan sp. Nostoc Nostoc chlorococcacean sp. Nostoc 12-24-66 Unidentified, unsterilizcd Lyngbya, Gleocystis (or palmelloid chlorococcalean) 12-19-66 Unidentified, unsterilizcd chlorococcacean, Protosiphon 12-14-66 Unidentified, unsterilizcd chlorococcalean, Protosiphon and Botrydium? 12-14-66 Unidentified, unsterilizcd Oscillatoria^ chlorococcacean, Sccnedesmus 12-13-66 Unidentified, unsterilizcd chlorococcalean, chaetophoracean 12-12-66 Unidentified, unsterilizcd chlorosphaeralean 12- 7-66 Unidentified, unsterilizcd chlorosphaeralean, Cylindrocapsa 12- 7-66 Unidentified, unsterilizcd Anahaena, Lynghya Stigonema, chlorococcalean, chlorosphaeralean Protosiphon Uniola laxa 9- 4-62 Panicum agrostoides Manisaria rugosa 7-13-62 Paspalum pubescens Protosiphon NostoCj Gloeocystis (or palmelloid chlorococcalean), Sccnedesmus? 9-30-61 Echinochloa frunentacea chlorococcacean Aristida wrightii Trioda sp. Avena fatua Stipa comata Bouteloua curtipendula Aristida wrightii Panicum agrostoides chlorococcacean 2 chlorococcaccan spp. chlorococcalean Anabaena, chlorococcacean Anahaena chlorococcacean chlorococcalean 8-22-53 Echinochloa frumentacea chlorococcalean Sple7iophalis sp. Calamogrostis epigejos Trehouxia, chlorococcalean, Gloeocytis chlorosphaeralean 8-23-49 Calamogrostis canadensis chlorococcalean Panicum polyanthes Stipa Columbiana Andropogon virginicus Stipa comata Bouteloua curtipendula Paspalum pubescens Muhlenbergia setifolia 6-27-40 Panicum agrostoides 9- 3-33 Paspalum ciliatifolium Hapalosiphon, chlorococcalean chlorococcacean Nostoc chlorococcacean Chroococcus? , chlorococcaccan Anahaena chlorococcacean chlorococcacean, chlorosphaeralean Anahaena [Vol. 56 116 ANNALS OF THE MISSOURI BOTANICAL GARDEN These data illustrate that no alga survived more than 60 years. Green algae dominated in frequency of appearance during the most recent 30 years, while blue-green algae dominated the preceding 30-ycar period. Only one diatom grew and this from the most recent material. Our results show further that similar viabilities occurred for the aquatic plants as for the predominantly terrestrial ones. Although we did not identify species, it was clear that recent collections (1966) (P J ) older collections, especially the non-preserved Gramiueae. Figure 1 illustrates the steady loss in viability of algal populations with increasing age of dried soil sample. 120 110 100 90 80 7.0 60 50 40 30 20 10 l-{7^ 7.5 1 i 1 1 1956-66 1946-56 1936-46 1926-36 1910-26 1881-1910 Fig. 1. Graph showing loss in algal viability, expressed in two ways: (1) as % of cul- tures developing algae; (2) as (Minimum No. Species Populations/No. Cultures) X 100. For convenience, calculations were based on approximately 10-year periods as shown on abcissa. 1969] PARKER, SCHANEN & RENNER SOIL ALGAE 117 Noteworthy among the genera not reported by earher workers but found in our cultures were a chaetoplioracean alga, Cyliudrocapsa, Scenedcsmus, JJlothrix, and Navicula, all from recent specimens, and the genus Protosiphon in some- what older source material. Among the blue-green algae, filamentous forms dominated in our cultures as with those of Bristol (1919) and Becquerel (1942), the Nostocaceae appearing form inated in all these studies. Diatoms were of rare occurrence. Discussion (1919) (1942) that a variety of soil algae, mainly belonging to the Chlorophycota and Cyanophy- cota, remain viable in dry soil for several decades in sufficient numbers to re- generate a visible population in response to favorable growing conditions. Table 2 .^H _H — ■_ ^H ^H ^ ^1 Wl that of Lipman (1941) which suggested that blue-green algae, especi and Anahaena, possess greater survival capacities than other algae. (1919) (1942) Table 2. pREguENCY of Occurrence of Viable Algae (by genus) in Dry Soil Reported by Bristol (1919), Becquerel (1942) and Us. Genus and Major Group Cyanophycota: Anahaena Aulosira Chroococcus? 7 Cy lindrospermitm 2 4 1 11 Uapalosiphon Lyngbya Nodularia Nostoc* OsciUatoria Phormidiuin Plectonema Siphononcma Stigonema Chlorophycota: Botrydina 1 4 chaetophoracean sp. chlorococcaccan spp. chlorococcalcan spp. Chlorococcum 11 Parker Bristol Becquerel et al. 1 1 1 2 1 1 5 1 1 1 2 5 4 1? 1 1 23 9 Cjcnus and Major Group chlorosphacralean spp. Cocconiyxa Cylindrocapsa Dactylothece Dictyococcus Glococrstis Microcystis? ¥ * (a green) Oocystis Protosiphon Sccnedesmus Stichococcus Trehouxia (= Cystocuccus) Truchiscia 4 4 Ulothrix (Uroncma?) Chrysophycota; Botrydium? Navicula Nitzschia 1 Parker Bristol Becquerel et al. 1 1 1 1 1 3 4 1 5 1 3 4 2 1 1 1 1 » » See also Lipman (1941) for record longevity. Becquerel's reference to this blue-green algal genus as a green alga renders its identity questionable. 118 [Vol. 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN urn. Xan- (1919) medium nor Bold's (1949) modified Bristol's medium favoi and Becquerel (1942) did not describe the composition of his med thophycean algae, however, grow satisfactorily in these media, so their absence suggests a low viability for this group in air-dried soil. The pattern of reduced viability with increasing age of soil samples terminat- ing at 1910 in our study lends credence to our assumption that our experimental technique was entirely sterile and that algal contamination of soil crumbs in this herbarium is infrequent. Bristol (1919) also took precautions to avoid con- (1942) to maintain sterile conditions. If the algae which developed in Becquerel's soil samples were of the same age as the source material (65-98 years), then we might conclude that some factor(s) brought about reduced maximum longevity of algae in our soil samples. For example, treatment of our herbaria with preserva- tives or differences in soil moisture contents might explain our apparent higher mortality. Indeed, Bristol's soil samples possessed moisture contents somewhat higher than most fully air-dried soils, and neither her soils nor those used by Becquerel were pretreated with carbon tetrachloride or continuous vapors from paradichlorobenzenc and naphthalene. Another possible explanation for the older soils of Bristol (1919) and Becquerel (1942) yielding viable algae is that they used larger amounts of soil in their inocula. Our soil crumb inocula, which were unusually small (1-3 mm) as a result of meticulous cleaning of herbarium mounted plants, may have con- tained significantly fewer viable algal cells initially. This feature would tend to produce a more steeply sloping mortahty curve. Bristol (1919) encountered blue-green algae more frequently than green algae, while we and Becquerel (1942) obtained the opposite result. This dif- ference between our data and that of Bristol could have resulted from differences in media or soils. The salts in Bristol's medium are 4X the concentration of Bold's (1949) modified Bristol's medium, while the ratios of salts (e.g., sodium nitrate/potassium phosphate) are the same. One might postulate that, over a long enough period of time, moist soil will experience some dentrification via the mixed microbial community present, and the N/P ratio will drop. This ratio should drop more in Bristol's solution than in Bold's modified medium, because the phosphate level will remain essentially constant. Such a lower N/P ratio may have favored the development of nitrogen-fixing blue-green algae which dominated Bristol's cultures, especially following the appearance of green algae. Our data demonstrates the remarkable ability of soil algal populations to sur- vive for many years in an air-dry condition. We cannot conclude that individual algal cells live this long, because cell division might have occurred during the long holding period. At least, given sufficient moisture, some soil algae can reproduce (Parker, 1961) occ a population may be considerable with but a few cells surviving many years. The importance of this feature to mechanisms of algal dispersal and survival over od 1969] PARKER, SCHANEN & RENNER — SOIL ALGAE 119 Literature Cited Allen, M. M. and R. Y. Stanier. 1968. Growth and division of some unicellular blue- green algae. Jour. Gen. Microbiol. 51: 199-202, Becquerel, p. 1942. Reviviscence et longevite de certaines algues en vie latente dans les terres dessechees des plantcs des vieux hcrbiers. Compt. Rend. Acad. Sci. 214: 986-988. BiscHOFF, 11. W. and H. C. Bold. 1963. Phycological Studies IV. Some soil algae from Enchanted Rock and related algal species. Univ. Texas Publ. 6318: 1-95. Bold, H. C. 1949. The morphology of Chlamydomonas chlamydogama, sp. nov. Bull. Torrey Bot. Club 76: 10M08. Bold, H. C. and B. C. Parker. 1962. Some supplementary attributes in the classification of Chlorococcum species. Arch f. Mikrobiol. 42: 267-288. Bristol, B. M. 1919. On the retention of vitality by algae from old stored soils. New Phytol. 18: 92-107. Brown, R. M. Jr. and H. C. Bold. 1964. Phycological Studies V. Comparative studies of the algal genera Tetracystis and Chlorococcum. Univ. Texas Publ. 6417: 1-213. Chantanachat, S. and H. C. Bold. 1962. Phycological Studies 11. Some algae from arid soils. Univ. Texas Publ. 6218: 1-74. Cox, E. R. and H. C. Bold. 1966. Phycological Studies Vll. Taxonomic investigations of Stigcoclonhim. Univ. Texas Publ. 6618: 1-167. Evans, J. H. 1959. The survival of freshwater algae during dry periods. Part II. Drying experiments. Jour. Ecol. 47: 5 5-81. pon Hortoragyt, T. 1960. The several month long lite ot algae trom a driea up laboratory conditions. Acta Biol. Acad. Sci. Hungar. 10(%): 299-300. KosTER, J. T. 1966. Some remarks on the taxonomy of the Cyanophyceae. Acta Bot. Neerl. 15: 57-62. Lipman, C. B. 1941. The successful revival of Nostoc commune from a herbarium spec- imen eighty-seven years old. Bull. Torrey Bot. Club 68: 664-666. Lund, J. W. G. 1962. Soil algae. IN: Lewin (ed.). Physiology and biochemistry of algae. Academic Press, N.Y., 759-770. Parker, B. C. 1961. Facultative heterotrophy in certain soil algae from the ecological viewpoint. Ecology 42: 381-386. Pearson, J. E. and J. M. Kingsbury. 1966. Culturally induced variation in four mor- phologically diverse bluc-grecn algae. Am, Jour. Bot. 53: 192-200. Smith, R. L. and H. C. Bold. 1966. Phycological Studies VI. Investigations of the algal genera Eremosphaera and Oocystis. Univ. Texas Publ. 6612: 1-121. SCANNING ELECTRON MICROSCOPY AS AN AID TO POLLEN TAXONOMY'^ BY John E. IUdgway and John J. Skvarla Department of Botany, Washington University, St. Louis and Department of Botany and Microbiology, Samuel Roberts Noble Laboratory of Electron Microscopy, University of Oklahoma, Norman Abstract Information obtained from scanning electron microscopy of whole pollen grains of 7xa and \va indicates direct correlation with that gathered from light microscopy and electron microscopy of thin sections. This information helps clarify and support past analyses. Introduction The portrayal of pollen surfaces has been greatly improved by the scanning electron microscope. Although still a relatively new instrument, there are already several publications describing its utility in pollen and spore investigations (see Echlin, 1968, for an extensive bibliography). For a number of years one of us (J.J.S.) has been concerned primarily with the morphology and taxonomy of grass and composite pollen. This has involved studies with the light microscope and the transmission electron microscope. Since the scanning electron microscope offers a rapid means of observing pollen wall surfaces, as well as providing greater resolution than the light microscope, the importance of this instrument to our work is obvious. Although preparative techniques for the scanning electron microscope are relatively simple and have been described (Echhn, 1968), comparatively Httle information is available on the two major processing techniques employed in taxonomic studies of pollen (viz. fresh and acetolyzed). Moreover, the aperture regions, so critical in taxonomic diagnoses, have been neglected at the expense of the unaperturate portions of the pollen wall. This report illustrates the effectiveness of scanning electron microscopy in studies of pollen morphology. Materi.vls and Methods Fresh pollen of Zea mays (Gramineae) and species of ha (Conipositae) were used. The pollen was examined in three stages: (1) fresh, unstained, (2) fresh, stained with OsO,, and (3) acetolyzed. After OsO, staining or acetolysis, the pollen was dehydrated through graded alcohols and allowed to dry after final rinses in either propylene oxide or reagent grade acetone. iThe authors wish to ac]** w ■ *^ -It ^ i H A ^4.= I ■r^'. — JW ■->>^;-TYvft-^r---- --AA^ ***H .' ' -s^.. 0. S ^ **^ I ^fi^^m.^ ^ 3? ■1^^ !^^ ii ©■ .'i^ -x ■"1^ ^^- m % % Fig. 1-9. Scanning electron micrographs of pollen. Fig. 1. Intact pore of fresh, un- stained pollen of Zea maySy X 2900. Fig. 2. Annulus and pore of fresh, unstained pollen of Zea mays, X 5800. Fig. 3. Pore of fresh, OsO^ stained pollen of Zea maySy X 5800. Arrows indicate endexine beneath the operculum. Fig. 4. Pore of acctohzed pollen of 7xa mays, X 3100. Fig, 5. Colpus and pore of fresh, unstained pollen of Iva xanthifoUa, X 6500. Fig. 6. Same as previous figure, after contraction of colpus. Fig. 7. Fresh, unstained pollen of Iva ciliata showing infolding of exine at colpus, X 3300. Fig. 8. Expanded colpus of acetolyzcd pollen of Iva ciliata, X 5400. Fig. 9. Colpus and pore of acetolyzed pollen of Iva xanthifoUa, X 7100. In areas away from colpi the morphology of tlie exine could be directly related to published work on transmission electron microscopy of thin sections (Skvarla and Larson, 1966), In such areas untreated, OsO^-treated, and acetolyzed pollen gave similar results. 124 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 Discussion Information obtained from scanning electron microscopy of Zea and Iva pollen has indicated direct correlations with that gathered from light microscopy of whole mounts and thin sections, and electron microscopy of thin sections. These results did not in any way put a new interpretation on past analyses, but did, however, help clarify, put into a better perspective, and support these data. Although most scanning studies have tended to emphasize' the ornamentation of the pollen wall \ier se, we feel that the information presented here on germinal apertures is also of great value. Dimensions of colpi and pores can be observed easily and with greater accuracy than with light microscopy. The utilization of these parameters in taxonomic problems is obvious. While the scanning electron microscope is designed primarily to give surface data, it is interesting to note that we have also been able to observe pollen wall stratification. It is likely that techniques will be developed which will provide more such data. That this is already nearing reality is indicated by scanning photographs of pollen which has been sectioned (F.chiin, 1968). The present study emphasizes the necessity for preparing pollen by different methods in order to gain maximum information. We have found that while fresh, untreated pollen is suitable for study, it is necessary to use fixation (OsOJ and acetolysis for supplemcntalion. Use of OsO, to stabilize labile algal and microbial specimens has provided satisfactory results (Fchlin, 1968). Since procedures of OsO^ fixation are standard in pollen studies using the transmission electron micro- scope, by using OsO, preparations in our scanning electron microscope studies, we can make direct correlative comparisons which are otherwise not possible. References EcHLiN, P. 1968. The use of the scanning reflection electron microscope in the study of plant and microbial material. J. R. iMicr. Soc, 88: 407-418. Skvakla, J. J., and D. A. Larson. 1965. An electron microscopic study of pollen mor- phology in the Compositae with special reference to the Ambrosiinae. Grana Palyn. 6: 210-269. ON TROPICAL TREE SEEDLINGS^ I. SEEDS, SEEDLINGS, SYSTEMS, AND SYSTEMATICS BY James A. Duke Battcllc Memorial Institute, Columbus Laboratories Columbus, Ohio Illustrated by Peggy K. Duke I \bstract The scattered information on characters useful in seed and seedling identification is brought together and terms defined. The problem of information retrieval in identifying dlings is discussed and an example given of the use of a polyclave for naming an un- known. Seedling characters are given for a number of tropical families. Introduction For the past three years, as contractor to the Atomic Energy Commission, Battelle Memorial Institute, Coknnbus Laboratories, has been involved in a study of the biological feasibility of nuclear exeavation of a sea-level canal in Panama or Colombia. Fiscal and temporal limitations forced emphasis on anthropological feasibility. My role, as botanist for the survey, has been to determine how man utihzes plants in the study area. Many times I have been faced with an isolated seed or fruit used by the natives. Other scientists, studying herbivore diets, may have only a seed from a fecal sample. Some of these can be identified, but usually only with difficulty. Internal morphology of the seeds provides valuable clues, Manv of the seeds are still viable. Some exhibit improved germinability after passing through animals. Characters of the resulting seedlings ofFer more valuable clues. As seed coats often remain intimately associated with seedlings, it seems rather narrow-minded to divorce seed from seedling studies or viee versa. This introduction to the study of tropical tree seedlings presents some of the important terms and concepts. Seeds and Systematics The importance of seeds to systematics is readily apparent in Gray's Manual (Fernald, 1950) where the second major division is termed *'Speniiatophyta'' meaning seed-plants, but there are cultivated spermatophytes which bear no seeds. Subdivisions also bear titles reflecting seed characters, Gymnospermae and Angio- spermac, depending on whether the carpels are open or closed, but there are exceptional angiosperms which have open carpels {Anchietea, Butomus, Decorsella, Firmiatia, Mitrasacme, Moringa, Reseda, Trillium) (Melville, 1962). The classes, Dicotvledones and Monocotvledones, are based on the number of cotvlcdons but many dicots (especially in Caryophyllales and Ranalcs) have only one cotyledon normally developed. Some monocots, e.g., Arisaema, Arum, Commelina, Dioscorea, Paris, Rajajiia, Sagittaria, Tannic, Tinantia, Trichopus, and Trillium, regularly ^Studies supported in part by the U.S. Atomic F.ncrgy Commission, Nevada Operations Office, Contract No. AT(26-1)-171. Ann. Missouri Box. Card. S6(2): 125-161, 1969. [Vol. 56 126 ANNALS OF THE MISSOURI BOTANICAL GARDEN or aberrantly reveal traces of a second cotyledon. Some taxa arc stenospermous with little seed variability, while others arc curyspermous with much variability. ( ) have dif- ferent types of seeds in the "ray" and "dise" flowers of the same eapitulum. Species of Stellaria have capsules containing seeds with two types of sculpture, the per- centages of each type being en\ironmentally determined. In Axyris amaranthuides (1946) (1) (2) exalate, lenticular seeds with satiny dark gray body. Thus, seeds are important to systematics for the definitions of taxa, from the division to the subspecific categories. Study of seeds will make one more sympathetic to segregation of the classic water lily assemblage into several families. Seeds of CeratophyUum exhibit a rare combination of characteristics: (1) radicle not apparent, (2) plumule foliate, (3) testa spiniferous, (4) embryo investing, and (5) endosperm absent. The water lotus, Nelnmho, shares several fundamental characteristics with Ceratophyl- lum, such as (1) radicle not apparent, (2) plumule foliate, (3) embryo investing and endosperm absent [a correlated combination of characters rendering Martin's (1946) classification of the embryo as "broad" untenable]. An amateur readily could separate the seeds of Nelumho and CeratophyUum by their external mor- phology. Nymphaea, classically included in the same family with Nelumho, has a radically diff'erent seed, which like its pollen, anatomy, etc., suggests mono- cotyledonous afl'inities. Definitive characters are (1) broad embryo, (2) puber- ulent fines, (3) white aril, and (4) copious amyloidiferous endosperm. Seed characters suggest that Nelumho is closer to CeratophyUum than to Nymphaea. Seeds and Retkieval Systems Murley (1951) voiced the still-unfulfilled challenge "identification of isolated seeds is important in paleobotany, archaeology, and in the practical work of seed laboratories, for the seed is usually the only part of the plant available for iden- tification. . . . Seed keys should be great timesavers compared to the former trial and error process." A novice confronted for the first time with water lily seeds would probably need to spend hours or even days to find out to which family the seed belonged. Toward the solution of this novice's problems, Duke (1964a, b; 1965b, c, d, e) started the polyclave, an information-retrieval system designed primarily to facilitate identification of unknowns. The polychne is a coordinated index to around 800 characters, some rarely employed by seed technicians, e.g., latex, leaf type, germination type, indument, etc. Seed characters alone overcrowded the standard 5x8 edgepunch cards. The polyclave system recognizes 480 families arranged roughly after the Dalla Torre and Harms sequence. Abbreviations of these 480 families are shown in the Family Polyclave Underlay (Plate 1). Plate 1 also contains underlays for Missouri, Panama, and Viet Nam, with abbreviations of the families reported in their respec- tive floras. These underlays, or positives, are printed in color, with plus signs in each corner. Plate 2 shows various character overlay cards or negatives updated 1969] DUKE — SEEDLING CHARACTERS 127 since publication of the polyclave. Negatives are printed in black and white or black and transparent and bear a minus ( — ) sign in each corner. Blanks in the over- lays represent families recorded to have the characters. Acetate transparencies of these cards can be superimposed over the corresponding underlay or positive. Then only the abbreviations which show through in color correspond to families pos- sessing the characters. Seeds of the water lily allies afford good examples of the polyclave approach, but some poisonous fabaceous seeds afford a more colorful demonstration. In Plate 2, abbreviations of families reported to have red seeds have been erased. Superpose the corresponding transparency over the family underlay and you read in color abbreviations of those families reported to have red seeds. Similarly, superpose the transparency for black seeds over an underlay and you read in color only those families reported to have black seeds. Superpose both transparencies over the underlay and you read in color only those families reported to have both characters. Add to these the transparency for poisonous seeds, and you see in color only the few families reported to have the three characters. Refer to Figure 23 and note that the seedlings of Ormosia have stipels, a rare character reported only in those families whose abbreviations have been removed from the transparency for stipel- late eophylls. Superpose all four transparencies and you will read in color "FAB," for Fabaceae, to which Ormosia belongs. With four transparencies, most families have been eliminated from consideration. In the polyclave (Duke 1969b) there are 770 character negatives scored for various taxonomic characters, about half of which are useful in determining the family of unknown seeds and seedlings. Superposition of any combination of these eliminates families not reported to have the characters. Theoreticallv, if an unknown has nine uncorrelated charac- ters, each of w^hich occurs in only half of the 480 families, superposition of the nine corresponding character transparencies would eliminate all but one family. Because of parallel evolution and flaws in existing classifications, however, several families may turn up, although perhaps few or no species in each may have the combination of nine characters. A retrieval system at the generic level would be more efficient, but it would be a sizeable chore to score some 20,000 genera, many with over a hundred species each, for a thousand characters. Beginnings have been made for the genera of grasses (Duke, Gunn, and Terrell, unpub- hshed), legumes (Duke, 1965e, 1969), and palms (Duke, 1965d). Seeds OVULES Seeds, like other plant entities, defy rigid definition, but a common definition is "fertilized ovule." The angiosperm ovule consists of a central body, enclosed more or less completely by one or two integuments and supported on a basal stalk, the fiinicle. The central body consists of the nncellus, a distal part, in which sporogenous tissue is borne, and the chalaza, a basal part where funicle, integu- ments, and nucellus merge. In sessile ovules, the funicle is absent, but in some the funicle may be elongate and adnate to the ovule body forming a ridge, the raphe. The scar on mature seeds known as the hilum represents die point of attachment of the funicle (or carpel wall in sessile ovules). Generally there are one or two integu- [Vol. 56 128 ANNALS OF THE MISSOURI BOTANICAL GARDEN o VI i < r ■ ■ u ^ ^ Ci w C5 MH tr^ 1— < b o X UJ r o CL, > en X c < c/3 o H L^ > UJ < o •< < 5 ^ < S z ti ^ 7" n 1— 1 rt :d!: < -H J-H < ce X "A < :£; < UJ u ^ O ^ ^ ■Ti 5 H UJ < 1— 1 b -fc X r H o. < ^. 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X X UJ — -< X -1: X -J UJ E k— 1 L l-H 1 u: X — UJ X c X z X X hJ UJ < X X X < < CI X UJ X A X 1-H i— ' X X L X X a: >- c Q < X -1 L Art nJ ^ X E UJ 1 — 1 u: c 3 1— UJ X I— CJ X - G ■< X Q UJ U, r_5 ^ Ltf u E Z X 3- X X H H > :■ X < '^ F*- tJ^ pU F ■ M ■*n ;^ ^ E H H :> X X L^ V j_ :» X B X A 3 i X -hH_b E Z* ^1* T** r** ft A nA < < X < < z C H-* r T H UJ n l-H < ^4h t/3 y4 X H n 1 ■Art 2 UJ A u: UJ X < "A E > < < ^ ^ z E < X UJ X X U U :t si < y. 3 b < ui < X X X UJ 2: X < X c u. X X X L E UJ h ■ 1 < X EIj X UJ H X X' IH > X UJ H-i 7: 3 u: a. X u: 3 rA u: X UJ t— 1 ^ X X UJ rA 5 31 F X X X ■-J X X f- X UJ X UJ UJ a. > hH < X UJ < a. < F-_> 0. X X X < 3 E z UJ X 1 r ^ -J UJ u UJ H 'yj X Ir- ;1 X 'J5 UJ X X UJ b_ c >- u X Z X UJ F X 3 UJ X l-H Z E -J X 1— 1 P- 3 ("^ CM c X X X X < A- < X ? ^ UJ X < X X H-l X 1— X < X -< X *-H X u UJ ft 4-H X H c X oJ F-i X X -T CO u: h-l X '-J X c_ « »— 4 E >^ X u: l-H u! X E < UJ ^ - -H E Q UJ E r^ X UJ X z. 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X A ■■^4 < c r^ UJ X T, < X X 5 r 'A < LJ_L c E X X c UJ X b^^ ^ UJ X 3 3 F^ X X E X' 21 ft a C X X X UJ —— c UJ < I- U- < 3= X c 0- X X Hr- -*— E h-iM *-- r < T 'H > >- X Z >■ r 1 A -Jh z UJ E ij^ h- 1 y < r*-; -H X oJ ry] X UJ X < »-H X y; X < >: c X X X MM < 4 z :2 X X l-H Z UJ -J X UJ X X ^ 5 N J u >- 5 1— 1 X c T < ^ z X X X X X X < T -J 1-1-1 X X L ^^ f-H z r " DC E Q HH X a »^ X UJ H X J Q < UJ X r C3 X >■ X ^ > > X >■ LJ X X X z u: X ^ 3 < X X Q X -M X -J- X SJ X UJ X X E 1— 1 < a n :^ ::: UJ E F- - UJ < UJ F^ft l-k. aH < X a UJ u. r" X 4— ri -^ u: -1 E Z X D- X X i— 1 ■ > :x -< 130 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 ments, more or less fused to each other and to the central body, but in specialized ategmic types the integuments are absent, e.g., some Balanophorales and Santalales. The opening between the tips of the integuments where the nucellus is exposed is termed the tnicropyle. Arils, loosely defined as investing outgrowths from the chalazal region, have been questionably interpreted as third integuments and as primitive. Freedom of the nucellus from the integuments as in Amphipterygmm, Cananga, Casnarina, Cleome, Juglans, and Myrica seems rare. Freedom of the two integuments occurs in some Amentiferae, Capparaceae, Fabaceae, Ranuncu- laceae, and Rosaceae. Where there are two, the outer usually is more massive and longer than the inner. Occasionally in the Annonaceae, Cactaceae, Proteaceae, and Trapaceae, the inner projects beyond the outer. Bitcgmic anatropous ovules usually have only one integument obvious on the side adjacent to the funicle. Number of integuments is not a constant indicator of relationships. Most species of Popidus have two integuments, but two species and all Salix species known, have only one (Fames, 1961). Peperomia has one integument, while other Piperaceae have two. Both integuments are absent in some Olacaceae and Opilia- ceae. As a rule, gamopetalous dicots are unitcgmic but Primulaceae and Cucur- hitaceae are bitegmic. OVULAR TYPES Ovulcs are termed crass inn cellar when the nucellus is mas- sive and the megaspore mother cells arc deep in the distal tissue and tenuinucellar when the nucellus is small and delicate and the spore mother cells (usually one) are directly below the epidermis. The two types intergrade. Crassinucellar is more primitive, usually accompanied by two integuments. The ovule is orthotropous (loosely synonymous with atropous^ when straight and upright on the placental surface, with the micropyle distal and the funicle short or absent. It is anatropous when bent upon itself and more or less adnate to the funicle, with the micropyle facing the placenta. Cronquist (1968) states that the anatropous condition is eventually derived from the orthotropous. It is campylutropous when the ovule is reniform, attached near the middle, and the micropyle faces the placenta. It is amphitropous when the funicle is adnate to the ovule for about half its length and the micropyle faces laterally. Anatropy is more common and characterizes many primitive taxa (e.g., in Ranales, Helobiae, Nymphaeaceae'). Orthotropy occurs in more specialized groups with solitary ovules and basal placentation — Juglandaceae, Najadaceae, Piperaceae, Polygonaceae, and Restionaceae. Campy- lotropy characterizes Apocynaceae, Capparaceae, Caryophyllaceae, Geraniaceae, and Verbenaceae. Orthotropy, though superficially more simple, seems to be derived as in the Urticales. Ulmaceae and Moraceae have suspended anatropous ovules, Laportea is transitional and Urtica and Boehmeria have basal orthotropous ovules. Circinotropous or coiled ovulcs are rare but occur in Cactaceae. STOMATAL TYPES and in some cases, stomatal types may serve as indicators: Actinocytic: Stoma encircled by a rosette of subsidiary cells. (Fames, 1961) ("cruciferous") : Stoma surrounded by three cells of which one is distinctly smaller than the other two. 1969] DUKE — SEEDLING CHARACTERS 131 Anomocytic ("ranunculaceous") : Stoma surrounded by a limited number of cells that are indistinguishable in size, shape, or form from those of the remainder of the epidermis. (Type 4; Stebbins and Khush, 1961.) Diacytic C'caryophyllaceous") : Stoma enclosed by a pair of subsidiary cells whose common wall is at right angles to the guard cells. Didymocytic C'arecaceous") : Stoma enclosed by four subsidiary cells, the pairs dissimilar. (Type 2; Stebbins and Khush, 1961.) "Gramineous": Guard cells with the middle portions much narrower than the ends giving a dumbbell appearance in surface view. Stomata tend to disappear as the seed matures, but they are reported in Carina indica, Carpolyza spiralis^ and Nerine howdeni (Boyd, 1932). Paracytic C'rubiaceous") : Stoma accompanied on either side by one or more subsidiary cells parallel to the long axis of the pore and guard cells. (Types 1 and 3; Stebbins and Khush, 1961.) Triacytic ("gordoniaceous") : Guard cells with three subsidiaries similar to each other, but differentiated from the epidermis. ENDOSPERM The so-callcd double fertilization results in two chains of events, the one leading to embryo formation, the other to endosperm formation. In all monocots and dicots so far investigated (Swamy and Ganapathy, 1957) endo- sperm formation follows one of three sequences in development. Nuclear: Initial division and usually subsequent ones are not followed by wall deposition. Nuclei may remain free or later be separated by walls. Cellular: First and several subsequent divisions accompanied by wall for- mation. Helohial: A transverse wall is laid down following the initial division, dividing the embryo sac into a micropylar and a chalazal chamber. Subsequent divisions are generally free-nuclear and may take place in both chambers, but invariably the main bulk of the endosperm is formed by the micropylar chamber. These developmental types will be of little use in identification of unknowns, but may be correlated with other characteristics. Sporne (1954), working on external morphological correlates concluded that the nuclear type was primitive, while Swamy and Ganapathy (195 7), correlating with wood anatomy, concluded that the cellular type was less advanced. Cronquist (1968) notes that in the Gentiauaceae, characterized by nuclear endosperm, two advanced mycotrophs, Voyria and Yoryriella, have cellular endosperm. Presence or absence of endosperm is a useful character. Ruminations in the endosperm are frequently diagnostic, and may arise (1) through expansion of the developing endosperm into furrows in the integument or (2) throu The latter may explain the peculiar tendency to cryptocotylar germination in some Annonaceae, The chemical and physical nature of the endosperm is occasionally diagnostic. Dissection in water of many seeds characterized by oily endosperm will yield an [Vol. 56 132 ANNALS OF THE MISSOURI BOTANICAL GARDEN obvious oil-slick. The IKI starch test can be used to detect amyloidiferous endo- sperm in embryos. Sivnnondsia chinensh is unique among plants in that its seed oil is not a fat but a hquid wax. (albuminous seeds). Cronquist ( (exalbuminous seeds) tion may take place rather late in ontogeny, as in many of the Sapotaceae, in rm rm the ground with or without the testa. In members of the Myrtales and many oco cotyledons to form the type of embryo termed macrojjodial. Perisperm, nutritive tissue derived from the nucellus or integuments, may be present in addition to endosperm, and is associated with the "peripheral" embryos of Martin (1946). Cronquist (1968) states that perisperm is clearly an advanced condition. During germination, the food is absorbed from the perisperm by the endosperm and thence is passed on to the embryo. SEED SIZE AND SHAPE As Eamcs (1961) notes, both large and small seeds have been considered primitive. Large seed size has quite naturally been correlated with the tree habit, and therefore deemed primitive. For examples of variation in the famihes, witness the palms, with huge seeds hke Cocos and small ones like Prestoea, and the legumes, with small seeds like TrifoUum and large seeds like Mora. Nonetheless, the novice faced with an unknown could eliminate much work if he knew what groups had the same size seeds as the unknown. The same is true of seed shapes. Who knows which shapes are most fre- quent — ellipsoid, reniform, lenticular, globose, ovoid, oblong, or Among the myriad of other shapes possible, some are quite rare, and the knowl- edge of which families exhibit which types can save an immense amount of random searching. segmen TESTA As the seed develops from the fertihzcd ovule, the integuments de- velop the seed coat or testa. Both integuments may still be recognizable in some Euphorbiaceae, Rosaceae, and Rutaceae, but more commonly the inner is lost or drastically reduced. In others, such as Opiliaceae and Pelliceriaceae, there is no testa. Appendages are quite often useful clues to affinities. Most important are wings, arils [fleshy transformations of the outer layers of the outer integument ( = sarcotestae), of the chala/a or of the distal part of the funicle], arilloids (those developing from the micropylar rim of the integuments), strophioles (fleshy seed) ), caruncles ( seed Bombacaceae, Malvaceae, Convolvulaceae, and Caryophyllaceae or concentrated in comas at one end of the seed, as in some Onagraceae, Apocynaceae, and Asclepiadaceae. Specialized appendages, much resembling modifications of the caruncle, are cap-Hke operculi (e.g., in Commelinaceae') which seem to facilitate 1969] DUKE — SEEDLING CHARACTERS 133 ( ) above characteristics, and several of the above characteristics intergrade. Corner (1954) suggests that the truly arillate seed is primitive in angiosperms, but arils and sarcotestae are generally regarded as adaptations for seed dispersal. Alatc and comosc seeds are clearly derived adaptations when compared to seeds with no special dispersal mechanisms. Less prominent markings of the testa have been treated in detail by Murley (1951) Brown probably is the most frequent seed color, with other colors occurring in less than half the famihes. Particular types of seeds, often seen in necklaces, are ocellate and halonate seeds, which are frequent in legumes. Ocellate seeds have a large spot of color, like an eveball such as the black-eved red seeds of AhmSy Ormosia, etc. Halonate seeds have a circle of Hghter or darker color prominent in the seed as in Enterolohiuni cyclocarpum (Duke, 1965a). CHEMISTRY Earlc and Jones (1962) published results of chemical analyses of seed samples from 1 1 3 plant families and their research continues to expand the number of families, genera, and species investigated. Valuable biochemical data presented at a specific level are percent ash, protein, and oil; weight per 1000 seeds; fraction of alcohol-soluble nitrogen; fraction of trichloracetic-acid-soluble nitrogen, reactions of starch test, alkaloid test, tannin test, etc. UTILITY AND GEOGRAPHY Most botauists at scvcral points in their career will have received seeds for identification with no other information than: "This seed is used by the natives (1) to poison fish, (2) to eat raw, (3) to make neck- laces, (4) to poison animals, (5) as an anthelmintic, (6) as an oil source, (7) pulverized as an insecticide, (8) as a starch source, (9) as a candle, etc." Or the letter may read, 'The enclosed seed was found: (1) germinating in sea drift, (2) among the belongings of a jaz/ musician, (3) to constitute 90% of the tufted titmouse's food intake, (4) clinging to the clothing of an exile Cuban, (5) in a Navajo campsite, (6) mixed in with pollen 1000 feet above New York City, (7) to cause dermatitis among three prisoners, (8) in the ejecta of a regur- gitating child, (9) as a frequent fossil in a Pleistocene deposit, etc." Unsolicited letters arrive by the hundreds in botanical gardens all over the world with statements like the above followed by the question "What is it?" Responsible taxonomists faced with sugh queries have spent days combing the Hterature to find out which seeds are barbascos, poisonous, edible, common in sea drift, airborne, allergenic, ornamental, etc. It is sad that most of their hand- scribbled notes could not be transcribed to an information retrieval system avail- able to subsequent botanists. Knowledge of the origin of the seed can be equally useful if entered into the system. There are many genera with brilhant red seeds, but how many occur in your backyard, in your state, in your country, in your continent, in your hemisphere? 134 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 Piperad type Embryo PROEMBRYO The following key serves to distinguish the main types of pro- embryo : I. Division of zygote longitudinal II. Division of zygote transverse: A. Division in terminal cell transverse: 1. Basal cell plays essential part in development of the embryo proper Chenopodiad type 2. Basal cell plays no essential part in development of the embryo proper: (a) Basal cell divides no further and becomes a large suspensor cell (suspensors of a few cells, but derived from the terminal cell, may be present) Caryophyllad type (b) Basal cell usually forms a suspensor of two or more cells Solanad type B. Division in terminal cell longitudinal: 1. Both basal and terminal cells take part in fur- ther development of embryo Asterad type 2. Basal cell plays httle or no part in development of the embryo Onagrad type ("crucifer type") Further information on embryology will be found in Davis (1966). EMBRYO Martin (1946) recognized 12 embryo types among seed plants. Some categories overlap with others, but this in no way destroys their taxonomic subdivided A. Basal (Embryo at One Pole of Seed) 1. Rudimentary: Small, nonperipheral embryo in medium to large seed, more Martin types scheme based on embryo types. It roughly groups the rmae onocot small embryo and copious endosperm are rather characteristic of both the woody and herbaceous Ranales and are frequent in the Liliaceae- Amaryllidaceae alliance as well as in palms and sedges. Intergrading with the rudimentary are two small categories, the Broad and the Canitate. 2, Broad: Martir rm Martin, should be sified as investing. Broad embryos occur infrequently in aquatic or palustrine, more or less primitive dicots and monocots, and in a few parasites. 1969] DUKE SEEDLING CHARACTERS 135 3. Capitate: A capitate or turbinate, more or less basal, embryo in copious occurs 4. in about ten families. Capitate and broad embryos seem to be correlated with operculate seeds. Lateral: Although characteristic of more genera than the preceding, the lateral embryo is characteristic of fewer families, one of them being the grasses. Reeder (1957) illustrates the usefulness of the embryo in grass systematics. Embryos of Cynomoriaceae, Bromeliaceae, and Centrolepi- daceae could be included according to the definition "embryo basal-lateral or lateral and evident from the exterior, usually with copious starchy endosperm." The lateral subdivision, found only in monocots, except for apparent occurrence in Cynomoriaceae, indicates a transition to the Peripheral division. B. Peripheral Embryo ordinarily elongate and large, quarter to dominant, contiguous in part, at least to the testa; perisperm starchy, central or rarely lateral; cotyledons narrow or expanded; dicots with one cotyledon occasionally abortive. This roughly coincides with the Centrospermae but other taxa approach the peripheral condition, e.g., members of Cactaceae, Cuscu- taceae, Frankeniaceae, Ulmaceae, etc. (To further suggest the resem- blance of Centrospermae to monocots, consider also the farinose starchy endosperm, characteristic of the Farinosae of monocots, and the tendency toward geniculate, ensheathed nodes.) In the peripheral division are straight, arcuate, hippocrepiform, annular, and spirolobal embryos. Many dicot peripheral embryos have the second cotyledon reduced (anisocotylar, as in Wittia, Fig. 60) or absent. Peculiarities shared by many families with peripheral embryos are possession of bcta-cyanin and polyporate pollen grains. Polyporate pollen grains also occur in Alisniataceae where, e.g., Sagittaria has rudiments of a second cotyledon. C . Axile 1. Linear Subdivision: (a) Linear: Embryo axial, generally several times longer than broad, straight, curved or coiled; cotyledons not expanded; seeds normally not minute; endosperm not starchy (rarely so in monocots). Martin notes a distinct group, Cannaceae, Marantaceae, Pontederiaceae, Sparganiaceae, and losteraceae, which are unique among linear seeds in having starchy endosperm and in having the base of the embryo extend into a depression at one end of the seed, as in some Commelinaceae, and Arecaceae, in Caulophyilum thalictroides (Po- dophyll monocots, and dicots. Included are straight, arcuate, hippocrepiform, annular, and spirolobal em- rm their nerinheral counterparts only by the excentric endosperm 136 [Vol. 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN supposedly would not give a starch test when present. Many aquatic monocots absorb all the reserves into a "foot." Macropodial embryos occur, e.g., in Alismataceae, Climaceae, Hydrocharitaceae, Lecythi- daceae, WelwUschiaceae, Zosteraceae, etc. (b) Dnarf: Embryo \ariable in relative size, small to total, generally stocky, cotyledons often poorly developed; seeds exclusive of testa mostly 0.3-2.0 mm long, often nearly as broad as long. This type occurs in many sympetalous families, e.g., Ericaceae, Geiitiaiiaceae, Luganiaceae, Scrophulariaceae, and Sulatiaceae. Parasitic derivatives of these are more likely to ha\e the following: (e) Micro: Seeds usually minute, less than 0.2 mm long, exclusive of (50 to 150) 2. Foliate Subdivision: (a) Spatulate: Embryo erect; cotyledons variable, thin to thick and slightly expanded to broad. The spatulate type occurs in more fam- ilies than any other, closely rivaled by the hnear (Martin, 1946). Since these are most common, they are least valuable for purposes of identifying unknowns. It would be practical to divide the spatu- late into subclasses based on cotyledon shape and relative lengths of hypocotyl and cotyledon (the H:C ratio in the polyclave). (b) Bent: \\\\h. expanded cotyledons in an axile position but with the cotyledons bent upon the hypocotyl in a jackknife fashion; cotyle- dons are generally thick or planoconvex but not necessarily so. In the commonest type, the bend is in the plane of the cotyledons. (o=) (0 ) mbent (o //). The term oblique is unfortunate since certain "linear** embryos are oriented obliquely to the long axis of the seed, e.g., Clintonia. In a fourth type (orthoplocal), the cotyledons in addition to being in- cumbent, are conduplicate (o>>). Such an embryo occurs in Avicennia. Martin would probably have classified this as a "folded" embryo. Spirolobal embryos have incumbent cotyledons, once folded (o j I II) while diplecolobal embryos have two or more folds (o | || 1 1)- I I I I I ^ All these "bent" embryos occur within the Brassicaceae and sporad- ically in other families. Rare convolute cotyledons occur in Ter- m in alia. (c) Folded: "Embryo with cotyledons usually thin, extensively expanded and folded in various ways." Many of the embryos diagrammed by Martin could be called incumbent or diplecolobal. Weirdly folded or contortuplicate cotyledons are frequent in the Sapindales and Malvales. The Convolvidaceae, also characterized by folded cotyle- dons, frequently share with the Malvaceae and Myrtaceae the char- acteristic of punctate cotyledons. 1969] DUKE — SEEDLING CHARACTERS 137 (d) Investing: The last of Martin's categories is defined under his AXILE division: ''Embryo erect and with thick cotyledons overlap- ping and encasing the somewhat dwarfed stalk for at least half its length; endosperm wanting or limited." This would merge at the halfway point with both the folded and the spatulate types. Species having both foliate plumules and straight embryos would usually fit this category. Investing embryos are not common, but frequently are associated with cryptocotylar germination. POLYEMBRYONY Polyembryony, the presence of more than one embryo in an ovule or seed, is not uncommon, and is characteristic of some groups. Secondary embryos may be adventive, arising from mother sporophytic tissue, or may derive from cleavage of the daughter sporophyte (embryo), or they may be haploid and derived from gametophytic nuclei other than the egg. Adventive embryos of nucellar origin are more common than those of integumentary origin, and have been said to exist in 19 percent of Trillium undulatum ovules. Citrus, Eugenia, and Mangifera are tropical genera notorious for polyembryony. Dormancy and Vivipary Some seed technologists avoid the use of internal morphology and the seedling. Many small seeds germinate rapidly without scarification and the seedling yields additional characters for determination of an unknown. Faced with seeds brought to me for identification in Panama, I planted the seeds and ten days later had seedlings of Cannabis (Fig. 10), thereby confirming my suspicions. Some seeds are germinating before they are shed. In Cycadaceae and Ginkgoaceae, there may be no embryo when the "seed" is shed, but in most if not all angiosperms, the embryo is present in some stage of development, frequently continuing its devel- opment after shedding. Some seem to have no dormant period, e.g., Annona, Durio, Myristica, and Thalassia, Fames (1961) suggests that dormancy represents an advanced stage in the evolution of the seed. After-ripening— morphological and physiological — may delay readiness to germinate until long after shedding. Fames states that size of seed seems less important, from the standpoint of primitiveness, than time of inception of dormancy and stage of specialization of the embryo. Since primitive gymnosperms and a few angiosperms lack dormancy, and since there is a long-continuing, after-ripening process in some primitive families, absence of dormancy seems primitive. Duke (in press) finds larger seeds and quicker germhiation in species of the rain forest than in drier forest types. Examples reported by Fames (1961) of time necessary to complete growth from seed-shedding stage to germinating stage are Vnmaria, 8 days; Caltha, 10 days; Clematis, 17 days; Actaea, Hepatica, and Thalictrtim, 2 months; Cocos, Fraxintis, and Paris, 4 months; Crocus, 6 months; Corydalis, 10 months; Trih lium, 12 months. Data presented by Marrero (1949) for tropical tree species, suggest that cryptocotylar species (22) average 41 days between sowing and germination w^hile phancrocotylar species (35) average 23 days. [Vol. 56 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN Vivipary, an obvious example of lack of dormancy, has been attributed to some tropical tree species, especially mangroves. Seeds not truly viviparous, e.g., Capparis, Hiira, and higa (Fig. 17), have been reported as germinating before falling from the tree, but such cases are probably due to unusual meteorological conditions rather than inherent characteristics of the species. Often, seedlings will be found intimately associated with fruits on the forest floor. On account of such an association, the author was able to identify the seedlings of Apeiba, Clusia, and Enallagma, all with many-seeded fruits, sometimes important in mammal diets. The flattened subspherical tuberculate fruits of Apeiba, the 5-locular superior fruits of Clusia, and the large globose fruits of Enallagma ofFer combinations of characters occurring in few families. The combination of operculate fruits with palmilobed cotyledons that characterizes Cariniana (Fig. 66) apparently occurs only in the Lecythidaceae. Coupling fruit characters with those of the seeds and seedlings renders determinations more certain. A second dormancy occurs in many tropical seedlings, in the dipterocarps in Asia and, e.g., in Connarus (Fig, 16) and Ormosia (Fig. 23) in Panama. Such species go into a state of dormancy after the cotyledons and first eophylls are raised and fully expanded, awaiting some ecological trigger to further their development. Seedlings GERMINATION There are two types of germination, phanerocotylar, in which the cotyledons emerge from the seed, and cryptocotylar, in which the cotyledons do not emerge from the seed (Duke, 1965a). Phanerocotyly is more common in dicotyledons, cryptocotyly in monocots. The peanut is a transitional type, while many genera have species with both types, e.g., Acer, Bauhinia, Caesalpinia, Clematis, Couratari, Lecythis, Ormosia, Passiflora, Phaseolus, Pithecellobium, Prunus, Pterocarpus, Qnerciis, Rhamniis, Riibia, Sapindns, Sophora, Stercnlia, Termifialia, Theobroyfia, and Trichosanthes. Eames (1961) states that the hypogeal (cryptocotylar) method is clearly advanced. However, in Phaseolus, cryptocotyly is a MendcHan dominant to phanerocotyly (Compton, 1912). Among cryptocotylar dicots, two types seem rather rare and definitive: those in which scale leaves (cataphylls) preceding the foliage leaves are opposite (Bignoniaceae, Clusiaceae, Melastomaceae). The common type has small spiral or alternate cata- phylls preceding the foliage leaves as in Giistavia (Fig. 65). Annonaceae, Hura, and several Rubiaceae, e.g., Pentagonia (Fig. 77) and Tocoyena (Fig. 79) exhibit a peculiar transitional condition. The cotyledons tardily emerge from the seeds, or may break off in the seed, unless man or accident breaks the seed coat. Usually the radicle is the first structure to emerge from the seed, but rarely the plumule emerges first. The hypocotyl is the portion of the axis where the stem-root transition occurs. Occasionally there is an external demarcation, the collet, between hypocotyl and root. The cryptocotylar forms are often described as being devoid of hypocotyl, but the transition must occur. Compton (1912) has differentiated three types of transition in the legumes. The epicotyi, the embryonic axis above the cotyledonary 1969] DUKE — SEEDLING CHARACTERS 139 node, is often not obvious in the seed until after germination, but it tends to be better developed in cryptoeotylar species before germination. SEEDLING AXIS After germination, the distinction between hypocotyl and epicotyl is more easily made (cf. Virola, Fig. 12). Frequently the hypocotyl will be swollen, pegged, or annulate near the soil. In many woody species such as Tamarindus and Virola (Fig. 12) the epicotyl and hypocotyl have different indu- ments and textures. In species destined to become stilted or buttressed, such as Cecroina jieltata and Sloanea herteriana, adventitious roots are often obvious at the seedling stage (Duke, 1965a). Swollen hypocotyls are not uncommon in species destined to have swollen trunks. In Puerto Rican tree seedlings studies (Duke, 1965a), all laticiferous species except Pterocarpiis had obvious latex by the first eophyll stage. In the seedling stage, Biirsera and Dacryodes already exhibit the characteristic aroma of turpen- tine; seedlings of many Lauraceae and Myrtaceae, e.g., Pimenta, may be dis- tinguished by their odors; seedlings of Cedrela odorata and Cordia alliodora are said to possess the characteristic alliaceous odor. A notebook containing a seedling of Qiiararibea smells about as pleasant as a herbarium case containing dried specimens. Spines may be evident quite early (second eophyll stage in Acacia farnesiana') (Duke, 1965a) and tardily develop in the axils of the cotyledons of Pereskia bleo. Bailey's survey (1956) of 99 dicot famihes showed that 77 percent of the seedlings have an even number of strands at the cotyledonary node, and 60 percent of the cotyledons have two independent traces related to a single gap. The double-trace, unilacunar node is not confined to a few genera, but is of common occurrence in many orders of dicotyledons. This contrasts markedly with the foliar nodes, where the majority of dicots have an odd number of traces. The double- trace unilacunar node has been reported for foliar nodes only in certain Am- borellaceae, Austrohaileyaceae, Calycanthaceae, Chloranthaceae, Hernandiaceae, Lamiaceae, Lactoridaceae, Lauraceae, Solanaceae, Trwieniaceae, and Verhenaceae. Bailey observed no cotyledons with multilacunar attachments, but pentalacunar attachments are reported for Fitchia speciosa (Asteraceae) and multilacunar for Gustavia (Lecyihidaceae) (Carlquist, 1961). If the trilacunar condition is primitive, and ontogeny recapitulatory, one would expect ontogenetic reduction of trilacunar to unilacunar nodes. No seedling studied by Bailey displayed such reduction; on the contrary, first eophylls of a consider- able number examined were unilacunar succeeded by tri- or even multilacunar nodes. Unilacunar nodes are usually associated with exstipulate leaves, w^hile tri- lacunar and multilacunar are associated with stipules (Carlquist, 1961). Two is the common number of traces in the lower taxa of both dicots and monocots, but four or three by fusion of the middle pair are frequent. Large numbers seem uncommon but occur in some Araceae and Cannaceae. In many monocots there are two strong lateral traces supplying the sides of the sheathing leaf base. (1961) roots [Vol. 56 140 ANNALS OF THE MISSOURI BOTANICAL GARDEN is frequent in monocotyledons. Tetrarchy has been considered more primitive than diarchy because of association with woody types, as for example in the Fabaceae. In Ranales, the Ranmicidaceae have diarch w^hile woody famihes have tetrarch primary roots. In Asteraceae both types occur, with neither obviously associated with primitive types. In dicots there are few \'ariations from tet- rarchy and diarcliy, but in monocots, polyarchy with 4, 6, 8, or 12 poles is not uncommon. COTYLEDON NUMBER It has bccu suggcstcd by Eames (1961) that dicotyly is the rule among angiosperms and gymnosperms, and that such features as polycotyly, schi/ocotyly, anisocotyly, gamocotyly, monocotyly, and cryptocotyly are derived. Pseudomonocotylar embryos may result from the conferruminate (gamocotylar) condition as in Eugenia, Ncliunho, RhizopJiora, etc., or from reduction of one of the cotyledons, as in species of Abroiiia, Caruni, Claytonia, Corydalis^ Cyclamen, Dicentra, Erauthis, Erigoiia, Mamviillaria, Peperomia, Vin- guicida, Ranunculus,, Scaligera, Trapa, etc. Cronquist (1968) notes in Peperoniia a gradual differentiation of two cotyledons for different functions: P. pellucida has tw^o equal cotyledons w^hich are withdraw'n to become the first functional leaves of the seedling; P. peruviana also has two, but only one is withdrawn from the seed, the other remains to function as an absorbing organ; P. parrifolia has tw^o initially unequal cotyledons — the larger remains within the seed coat as an absorbing organ, the smaller becoming the first leaf of the seedling, a situation hardly to be distinguished from that in many monocots. In some Gesneriacae, the cotyledons are the only leaves the plant develops. Among monocots, some Alismataceae, Araceae, Conunelinaceae, and Dioscoreaceae show rudiments of a second cotyledon. Anisocotyly is well exemplified by such embryos as Petiveria. Tricotyly is a frequent aberration, occurring in such diverse plants as coffee, maple, mesquite, and walnut. Polycotyly has been reported for 2 to 4 percent of the "British flora*' (Eames, 1961), and is frequent in parasitic dicots, e.g., Eoran- thus, ISuytsia, and Persoonia. In woody Ranalean taxa, the percentage of tricotylar embryos may reach as high as 87 percent, witli 13 percent tetracotylar and none dicotylar (Eames, 1961). In some cases, e.g., conifers and crucifers, the two cotyledons may be so deeply lobed as to appear tetracotylar. In Podocarpus coria- ceiis (Duke, 1965a), the first pair of leaves is attached so close to the cotyledonary node as to give the seedling a tetracotylar appearance. Superficially the cotyledons and eophylls appear similar but the cotyledons have two veins prominent while the eophylls have only one. Lobing of the cotyledons appears to be constant in some taxa such as Bursera, Dacryodes, Eepidium, and Tilia where the lobing is palmate and in Amdnckia, Eschscholzia, Eucalyptus, Haematoxylum, Ipomoea, Pittosporum, Pterocarya, and Schizopetalon, where the cotyledons are deepl} bifid. Retuse cotyledons are more or less characteristic of Bignoniaceae and Convolvidaceae. Lobed cotyledons are not necessarily correlated with lobed leaves. Peculiar combinations such as trifoHolate cotyledons and entire eophyll 1 (Bursera), entire cotyledons, and tri- foholate eoph)ll 1 (Ceiha), bilobate cotyledons and paripinnate eophyll 1 (Haema' 1969] DUKE — SEEDLING CHARACTERS 141 toxyhim), rctuse cotyledons and imparipinnate eophyll 1 (]acaranda) can be very diagnostic. Plicate, crcnate cotyledons seem to be characteristic of the arboreal segregates of the Boraginaceae, Glandular margins occur on the cotyledons of several seed- lings whose adult leaves also have glandular margins CRosaceae, Rutaceae, etc.)- Cotyledon SHAPE AND SIZE Among entire cotyledons, there is a great variety of shapes and sizes which may be useful in determinations. Linear cotyledons are exceptional in some families, e.g., Dodonaea in Sapindaceae, and Spondias in Anacardiaceac, while they are the rule in others, especially in monocots and several dicots possessing "linear" embryos. Reniform cotyledons prevail in some famihes, especially those with "folded" and "spatulate" embryos (e.g., Bignonia' ceae, Comhretaceae, and Malvaceae^, Many genera have species with narrow and species with broad cotyledons, e.g., Coreopsis and Galiian. Broad cotyledons are not necessarily followed by broad leaves. Coccoloha, Vagus, Gomidesia, Hakea, and Myrcia have species in which the cotyledons are broader than long while the eophylls arc longer than broad. Other species have very narrow cotyledons followed by broad eophylls, e.g., Acer, Cissampelos, Cochlospermum, and Menispermiim. Absolute size of the expanded cotyledons is probably less important than relative size. Auriculate cotyledons occur (e.g., many Fahaceae and Meliaceae^ while inte- grading cordate cotyledons seem less frequent (e.g., some Raniincnlaceae'). Peltate cotyledons occur but rarely (e.g., PeperoiuiiO^ peltate eophylls being more frequent (e.g., Hernandiaceaey Menispermaceae, Piperaceae, and Tropaeolaceae^. COTYLEDON VENATION Bailey (1956) notes than many cotyledons are characterized by having a pseudo-palmate or palmate-parallel venation. Such cotyledons, scored plinerved in the polyclave, commonly have three or more con- spicuous primary veins that extend outward from a locus at the base of the cotyledons, the laterals diverging and often reconverging parallel to the margins. Many such cotyledons show a transition from an even number (2) of vascular strands at the nodal level to an odd number (3 or 5) of primary veins in the lamina. There are numerous deviations from this common type, even among seed- lings of the same genus or species. Cotyledons with different forms and venations may have similar basal (nodal) vasculatures and conversely cotyledons of similar form may have different nodal patterns. Pinnate venation is less frequent in cotyledons than in foliage leaves, but there are more penninerved cotyledons among those attached at a single-trace node. Parallel venation is the common type in monocotyledons. In cryptocotylar forms, whose cotyledon tips are modified for suctorial purposes (scutellum), there is usually only one trace which divides to form a median and two strong lateral bundles, but only the median vein continues to the scutellum. In more primitive forms, in which there is little or no downbending of the distal part, all the vascular bundles, including those of the sides of the sheath, usually continue into the scutellum. [Vol. 56 142 ANNALS OF THE MISSOURI BOTANICAL GARDEN Carlquist (1961) notes that seedling anatomy is adduced as evidence of the origin of monocots from dicots but adds that vascular anatomy of cotyledons is perhaps of little signihcancc because the duality of traces in each dicot cotyledon is as frequent as the well-known duality of traces in monocot cotyledons. The con- servative nature of seedling anatomy promises useful evolutionary studies. Cotyledons with a pair of midveins, as are frequent in gymnosperms like Podocarpus coriaceiis, are only occasional in angiosperms (some Austr oh alley aceae, Chloranthaceae, Moniniiaceae, and Poaceae'), Examples of cotyledons and eophylls exhibiting similar venation patterns are rare. In the polyclave the term seed leaf is used for the cotyledons after ger- mination. VERNATION Vcmation (prefohation) is the disposition of the leaves in the bud. The terminology in existence may be apphed also to the relation of the cotyledons to each other in the seed. The types of vernation are: Coudiiplicate induplicate: The lamina is folded lengthwise along the veins so that the halves of the upper surface lie together. This type is prevalent in leaves, but rare in cotyledons. The cotyledons of Avicennia are conduplicately folded, one so that its lower surfaces are opposed (reduplicate), the other induplieately embracing it (Duke, 1965a). Monniera, Raphauus, and other genera with ortho- plocal embryos are similar to Avicennia. Conduplicate reduplicate: The lamina is folded lengthwise along the veins so that the halves of the lower surface lie together as in some palm eophylls. Plicate (pleated): The lamina is folded along the veins, like a closed fan, as in cotyledons of Cordia (Duke, 1965a), and in leaves of Acer. Convolute: The lamina is rolled lengthwise forming a coil as in the leaves of certain Rosaceae and the cotyledons of many Combretaceae as in Terminalia (Duke, 1965a). According to Lubbock (1891) leaves of Drimys, Kadsiira, and Schisandra are convolute. Among Asian mangroves, Rhizophora and Bruguiera have convolute vernation of the leaves, while Carallia, Ceriops, Gynotroches, and Pellacalyx have involute vernation. Involute: Both margins of the lamina are inrolled lengthwise on the upper surface, as in certain Violaceae and Rhizophoraceae. Revolute: Both margins of the lamina are inrolled lengthwise on the lower surface, as in many Ericaceae. Reclinate (inflexed): The upper part of the blade is bent on the lower, as in leaves of Liriodendron. One of the cotyledons of Petiveria in addition to being involute, is doubled back on itself. The second broader cotyledon is convolute or involute about the first. It seems that the scutellum could in many cases rep- resent the end of a reclinate cotyledon. Circinate: The upper part of the blade is rolled back on the lower, so that the tip is in the center of the coil, as in Cycadaceae and Droseraceae. Certain legume eophylls, e.g., Parkia (Duke, 1965a), approach this type of vernation, while the cotyledons of Cariniana (Fig. 66) have a complex circinate vernation. 1969] DUKE — SEEDLING CHARACTERS 143 Strict: The blade is straight in the bud and expands by growth alone as in many cotyledons and adult leaves. Vernation of the cotyledons frequently differs from that of eophylls. Major mangrove genera of Panama may be distinguished by virtue of vernation and eophyll arrangement alone. The distinguishing characters are: Cotyledons strict; leaves involute Pelliceria Cotyledons not strict; leaves not involute: Cotyledons conduplicate Avicennia Cotyledons convolute or conferruminate: Cotyledons conferruminate; leaves convolute Rhizophora Cotyledons convolute; leaves not convolute: Eophylls opposite Laguncularia Eophylls alternate Conocarpus PETIOLES AND STIPULES Scssilc and long-petiolate cotyledons may occur on seedlings of the same genus or family, but there is rarely much variability at the specific level. In Bryoftia dioica the cotyledons are long-petiolate while in B. laciniosa they are subsessile. Lubbock (1892) speculates that cotyledons tend to be sessile when they are supported by a long hypocotyl, but long-petiolate when borne close to the ground, citing Vitis hypoglauca with short hypocotyls and long cotyledonary petioles and V. cehennensis with long hypocotyl and short petioles. Petioles are often connate at the base as in Lupinus, Podophyllum, and Sangui- sorba. On eophylls, the geniculate or incrassate petiole often foreshadows a com- pound leaf. Stipules offer good diagnostic characters. Even the cotyledons of certain seedlings, e.g., in Caesalpiniaceae, Pahaceae, Mimosaceae, and Rubiaceae, have obvious stipules. The cotyledons of Prosopis jiiUflora, in addition to being stipu- late, exhibit the nyctinastic tendencies of the foliage leaves. Among some crypto- cotylar legumes, e.g., Erythrma spp., (Duke, 1965a), the cataphylls tend to have paired stipules almost equal to those of the eophylls. Stipules of the second eophyll of Acacia farnesiana are spinescent (Duke, 1965a). In stipulate plants, the stipules are usually apparent no later than the second eophyll stage. Stipules are usually apparent on the seedlings but absent from mature branches of species with fugaceous stipules. Stipels are obvious on the first eophylls of some Mimosaceae and Moringaceae, and may be expected in other families whose foliage leaves are characterized by stipels. INDUMENT Presence or absence of indument is not highly definitive, but certain types of indument can be very useful in defining seedhngs. Indument provides a main clue for distinguishing three weed-tree species' seedlings in Panama. Eophylls of Cecropia have a white arachnoid indument on the lower surface, while those of Didymopanax have a diffuse, more or less colorless indu- ment. Ochroma eophylls have a close felted grayish indument, rusty at the base of the petioles (Duke, 1965a). [Vol. 56 144 ANNALS OF THE MISSOURI BOTANICAL GARDEN Punctate cotyledons arc characteristic of several families (Elaeagimceae, Mal- vaceae, Myrtaceae, Vrimiilaceae, Riitaceae). Beneath a tree of Thesiyesia popiihiea near Guanica, Puerto Rico, where seedlings similar to those of Ipomoea hispida, as figured by Lubbock (1892). Examination with a lens showed the characteristic pellucid punctation of Thespesias cotyledons, showing that this was merely an aberrant type with bifid cotyledons such as occur normally in Ipomoea spp. This is apparently the first such aberration reported for the Malvaceae. Montezuma, another malvaceous genus in Puerto Rico, often coexistent with Thespesia, may be distinguished on basis of its stellate hairs. MICROSCOPY When critical determinations are needed, microscopic exam- ination of the seedling will yield further clues. It has been generally agreed that the stomata on various plant parts are of the same type, but Paliwal and Bhandari (1962) point out that "The stomata on the leaves of Michelia and Magnolia are syndetocheilic while those occurring on the outer integument of ovules . . . are of the haplocheilic type/' Stomatal types in cotyledons and in adult leaves could also differ; the point deserves investigation. It would be worthwhile to see if distribution of stomata is the same in cotyledons, eophylls, and adult foliage. Hydathodes or glandular mucrons are frequent at the tips of the cotyledons as in Rhus typhina. EOPHYLL SEQUENCE Tomlinsou (1960) proposed the term "eophyll" for the first few leaves with green, expanded laminae developed by the seedlings, as opposed to the brown rudimentary scale leaves of fixed number which precede them in most cryptocotylar species. In palms, he noted that the shape and size of the first eophyll is constant and the distribution of its armature and indument is diagnostic. Each palm has its characteristic series of transitional leaves between the eophylls and adult leaves. Similarly characteristic eophyll sequences occur in Anacardiaceae, Bignoniaceae, Caesalpiniaceae, Fahaceae, Meliaceae, Mimosaceae, Sapindaceae, etc., probably in most seedlings w^hose adults have compound leaves. The eophyll sequence may be abrupt, as in many Mimosaceae with the first eophyll pinnate, the second bipinnate; or delayed, as in many woody Fahaceae, Meliaceae, Sapiiidaceae, etc., where several eophylls are simple and succeeding ones gradually add leaflets. Often the tumid petiole of an eophyll foreshadows a compound leaf. In some species, there is no sequence, e.g., in Giiaiacum, where paripinnate eophylls immediately follow the cotyledons (Duke, 1965a). There are few reverse sequences in which eophylls are compound and adult leaves simple (Acacia, Ulex). I suspect this would pro\e to be true of Suartzia sp. (Fig. 22). In Biirsera simaruha, the cotyledons are trifoliolate, eophylls simple, and adult leaves compound. In others, the cotyledons are lobate and the eophylls and adult leaves (mctaphylls) simple (Tilia). Having cotyledons and eophylls pre- cisely similar, as reported in Dowingia by Lubbock (1892) seems to be quite exceptional. There are few phanerocotylar species with cotyledons broader than the first 12 eophylls CAsclepiadaceae, Cactaceae, Casuarinaceae, Myrtaceae, Fro- teaceae, e.g.). Several species are known where the eophylls arc more conspicu- ously dentate or lacerate than the adult leaves QCasearia sp., Cordia spp., Ficus, Gustavia (Fig. 65), Querciis, certain members of Quiinaceae, and Rapanea sp.). 1969] DUKE SEEDLING CHARACTERS 145 / (1892) remarkable amount of variation in Ulex europaeus. In some seedlings, the leaves are all simple, in others the primary leaves are simple, subsequent ones trifolio- late; in still others all leaves are trifoliolate. They may be alternate or opposite indiscriminately. All ultimately become modified into simple spines or may be altogether aborted, while the branches form compound or branching spines. Al- though there are exceptions, the eophyll sequence seems to be characteristic of the species, as do most other characters used to distinguish seedlings. Blastogeny may be added to the other exciting and relatively unexplored fields of investiga- tion, where both variations and constancy may be used in evaluating taxonomic and systematic concepts. Even variability, e.g., heterophylly, as in Morus, Sas- safras, and, in Panama, Ruiqmla, seems to be a constant attribute of some taxa. Preliminauy Systematic Survey^ The following descriptions are based on the limited information available, mostly from the author's researches except where otherwise noted. Regrettably, the family description is often based on a single species. To amplify these studies, seeds arc requested from any tropical American woody species accompanied by a voucher specimen. Especially desired are representatives of families not included in this paper, then genera, then species. When possible, the seeds will be ger- minated, illustrated, preserved in the seedling herbarium, with duplicate seedlings returned to the collector who will be duly acknowledged. Herbarium specimens from Latin America are available in exchange for those seeds accompanied by voucher specimens. podocarpaceae: Germination phanerocotylar, the two cotyledons linear, bincrvatc. Eophylls pseudocotyledonary, like the cotyledons, but uninervate. gnetaceae: Seeds broadly oblongoid, the outer coat pinkish-red, not striped, the inner coat olive-brown, longitudinally striped. Germination cryptocotylar, with alternate cataphylls (Fig. 1). arecaceae (paemae): Germination cryptocotylar. Cataphylls alternate. Eophylls simple, bifoliolate, or paripinnate; usually possessing the armature and indmnent of the adult. Vernation induplicate or reduplicate. The species illus- trated (Figs. 2, 3, 4) all have bifoliolate eophylls. Astrocaryum eophylls are already armed and glaucous like the metaphylls (Fig. 4). The specimen illustrated had been grazed by some herbivore. araceae: Germination cryptocotylar, the cataphylls alternate, linear- lanceolate, the first eophyll long-petiolate, hastate, spirally convolute CMontrichar- dia, Fig. 5). smilacaceae: Germination cryptocotylar. Cataphylls alternate; eophylls al- ternate, with venation like the metaphylls (Fig. 6). 1 Presented at the Second Symposium on Amazonian Biota in Florencia, Colombia, Janu- ary 23, 1969, and submitted for publication in Spanish in the proceedings of the symposium. 146 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 casuarinaceae: Germination phancrocotylar, the cotyledons narrowly ob- ovate; coph)lls vcrticillatc, apprcsscd. piperaceae: Germination phancrocotylar to subcryptocotylar, monocotylar, anisocotylar, dicotylar, or tricotylar; the cotyledons sometimes retuse. Eophylls usually with aroma, indument, and venation of the mctapiiylls. moraceae: Germination cryptocotylar (usually in large-seeded species, Figs. 7 and 8) with alternate cataphylls to phanerocot>lar (usually in small-seeded species like Finis, Fig. 9) without cataphylls, often anisocotylar. Latex, stipules, and indument often characteristic. Eophylls in cryptocotylar species alternate, stipulate, often more conspicuously dentate than the metaphylls, as in Broswnun hernadettae (Fig. 8) [but entire and witli caducous stipules in B. utile (Fig. 7)]. Cecropia and Vuuroiima suggest transitions to Urticaceae. cannabaceae: Germination phancrocotylar; the cotyledons sessile, entire, uniner\ed, or triplinerved. Eophylls near the cotyledons, subsessile, penninerved, with an undulate, rugulose margin, sometimes basally dentate (Fig. 10). urticaceae: Germination phancrocotylar, often anisocotylar. Cotyledons long-petiolate, often plinerved and rcniform. Indument, venation, and arrange- ment of the eophylls often diagnostic. olacaceae: Germination cryptocotylar, the stalked cotyledons sccund. Hypo- cotyl glabrous, somewhat swollen. Cataph)lls alternate, the upper subtended by spines, grading into alternate, penninerved, glabrous, entire, mucronate eophylls, subtended by spines (Ximeiiia). polygonaceae: Germination phancrocotylar, the two cotyledons rcniform, subplinerved. Eophylls supracotyledonary, alternate, subconvolute, penninerved, ochreate (Coccoloha). My efforts to germinate Triplaris have failed. Peculiar that a weed-tree should possess such low vitality! nyctaginaceae: Germination phancrocotylar, at first anisocotylar, the cotyle- dons becoming subcqual, subsessile, rcniform, plane, entire, penninerved. Eophylls supracotyledonary, alternate, entire, short-petiolate, penninerved (Ncca; Fig. 11). annonaceae: Germination phanerocot)lar, often tardily so, probably crypto- cotylar in large-seeded species. Cotyledons lanceolate, subpenninerved. Eophylls alternate, conduplicate. Aroma, indument, and \enation often diagnostic. myristicaceae: Germination cryptocotylar, the sccund cotyledons borne in the corrugated seed well above the ground. Eophylls supracotyledonary, alternate, conduphcatc, exstipulate, penninerved, entire (Virola, Fig. 12). Latex, indument! and aroma often diagnostic. monimiaceae: Germination phancrocotylar, the hypocotyl basally swollen, the cotyledons subsessile, broadly o\ate, subplinerved and bluntly apiculate in MolUncdia (Fig. 13), uninerved and emarginatc in Siparuna (Fig. 14). lauraceae: Germination cryptocotylar, the epicotyl usually emerging from the cotyledons, i.e., cotyledons not sccund. Cataphylls alternate. Eophylls alter- nate, entire, exstipulate, penninerved. Aroma and indument often diagnostic. 1969] DUKE SEEDLING CHARACTERS 147 Fig. 1-16. Fig. 1. Gnetum leyholdii (X 0.4). Fig. 2. Genoma binervia (X 0.8). Fig. 3. Welfia georgit QDuke 15030; X 0.6). Fig. 4. Astrocarynm standleyanum (X 0.2). Fig. 5. Montrichardia arhorescens (X 0.4). Fig. 6. Smilax sp. QDuke 15077; X 0,4), Fig. 7. Brosium utile (X 0.2). Fig. 8. Brosium bernadettcae (X 0.4). Fig. 9. Ficus insipida QDuke 15195; X 0.8), Fig. 10. Cannabis sativa (X 0.4). Fig. 11. Neea sp. (Duke 15332; X 0.4). Fig. 12. Virola sp. (X 0.4). Fig. 13. MoUinedia CDuke 15234; X 0.4). Fig. 14. Siparuna QDuke 15128; X 0.8). Fig. 15. Capparis pittieri (X 0.4). Fig. 16. Connarus panamensis (X 0.4). [Vol. 56 148 ANNALS OF THE MISSOURI BOTANICAL GARDEN hernandiaceae: Germination cryplocot\Iar, the stalked eotyledons seeund. Cataphylls alternate, grading into alternate, peltate, subplinerved, entire eophylls {ller}ia}idia). papavekaceae: CJermination phanerocot\]ar, tlie cotyledons subsessile, elliptic, entire, plinerved. Eophylls cotyledonary, quercoal, with orange latex (Bocconia). capparaceae: Germination phaneroeot}'lar, the coivledons plane or slightly planoconvex, subconvolute. liophylls alternate, entire, penninerved, conduplicate. Capparis pittieri (Fig. 15) has the largest cotyledons of any Capparaceae studied, characteristically raising the gray pubescent testa with them. Several nerves arise at the base of the cotyledon and the hypocotyl is somewhat swollen. morinc;a(:eae: Germination cryptocotylar, the short-stalked cotyledons re- maining in the trialatc seed. First eophylls supracotyledonary, opposite, decom- pound, stipellate. As is often true in cryptocotylar species, the plumule is foliate in the ungcrminated seed. chkysobalanaceae: Germination cryptocot}lar, the cotyledons seeund. Cataphylls alternate, sometimes stipulate, grading into alternate, stipulate, entire, lanceolate to ovate, penninerved eophylls. connaraceae: Germination cryptocotylar, cataphylls absent. First eophylls supracotyledonary, opposite, often incrassipetiolatc, ovate, subcordate, entire. Coiniarns panamemh (Fig. 16), like Rcmrea, resembles the seedhngs of many Fahaceae, e.g., Orwosia (Fig. 23), and is often dormant at the stage depicted. The absence of stipules on the eophylls separates the Connaraceae from most Vahaceae. MiMOSACEAE: Germination phanerocotylar to cryptocotylar, with or without cataphylls; eophylls simple to decompound, the sequence gradual to abrupt. The species of Inga investigated, like J. spectahilis (Fig. 17), have nonphoto- synthetic cotyledons, rarely seeund, usually auriculate, which are cryptocotylar but escape from the testa shortly after germination. These seeds frequently germinate while still in the pod. Cataphylls are sometimes present, and stipulate. All first eophylls studied are bifoliolate. Pithecellohium longifoliuui (Fig. 18) has seedhngs comparable to J)iga, sub- cryptocotylar with nonphotosynthetic cotyledons and bifoliolate eophylls. All species of Pithecellohiuuf studied generate the odor of a mercaptan in germinating. Cotyledons of Pentaclethra (Fig. 19) are somewhat similar, but arc green and probably photos) nthetic, usually gaping apart at the surface of the soil in the swamps they frequent, thus not exactly hypogeal nor epigeal, nor cryptocotylar nor phanerocotylar. The first eophylls are decompound. caesalpiniaceae: Germination phanerocot\lar to cryptocotylar, with or without cataphylls, cryptocotylar cataph\lliferous germination usually associated with the larger seeds. Probably the most affirmative demonstration of tlie impor- tance of seedlings to systematics is Leonard's (195 7) excellent work on African Caesalpiniaceae, in which he proposes the following hypotheses: 1969] DUKE SEEDLING CHARACTERS 149 Fig. 17-27. Fig. 17. Jnga spcctahiUs (X 0.4). Fig. 18. Pithecellohiiim hn^ifolhim (iDuke 8212; X 0.8). Fig. 19. Pentaclcthra macrohha (X 0.1). Fig. 20. Vrioria copaifera (X 0.1). Fig. 21. Mora okifera (X 0.04). Fig. 22. Swartzia simplex (X 0.4). Fig. 23. Or- mosia sp. (X 0.4). Fig. 24. Oleiocarpon panamensis, (X 0.2). Fig. 25. Maiighania sp. (/J/J^' 15448; X 0.4), Fig. 26. Myroxylon hahamum (X 0.4). Fig. 27. Ptcrocarpus hayesii (X 0.4). 150 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 (a) The establishment of synonymy between genera aeeording to morpholog- ical data should be provable by the similarity of their seedlings. (b) Morphologically related genera, which have the same seedlings, may not be generically distinct. (c) The partition of a heterogeneous genus into several genera according to their morphological characters, should be provable by the existence of a particular seedhng type for each of them. (d) The existence of several seedling types within one genus may be an in- dication of a generic heterogeneity that must be checked by other morphological data. Priuria copaifera (Fig. 20), a swamp forest species, is cryptocotylar with alternate cataphylls succeeded by bifoliolate eophylls which gradually give way to paripinnate metaphylls. Mora oleifera (Fig. 21), a brackish swamp forest species, with the largest dicot seeds known, is also cryptocotylar with alternate cataphylls succeeded by bifoliolate eophylls succeeded by paripinnate metaphylls. fabaceae: Germination phancrocotylar to cryptocotylar, with or without cataphylls (in the same genus occasionally). Cataphylls often stipulate. Simple incrassipetiolate eophylls often foreshadow compound metaphylls. Latex, indu- ment, stipules, stipellae, leaf margin, venation, and arrangement often diagnostic. Cryptocotylar Swartzia (Fig. 22) exhibits the rare reversed eophyll sequence, the imparipinnate eophylls gradually giving way to unifoliolate metaphylls. Ormosia (Fig. 23) is barely phancrocotylar, the nonphotosynthetic cotyledons often embraced by the nigrescent testa. Oleiucarpon panamensis (Fig. 24) exhibits an abrupt sequence, the phan- crocotylar cotyledons being followed by eophylls similar in structure to the metaphylls. Manghayiia, an introduction becoming a regular constituent of the savanna flora, is cryptocotylar, apparently without cataphylls, the eophylls unifoliolate (Fig. 25). Myroxlon (Fig. 26) with the first im- paripinnate eophylls opposite. This combination of characters shows up in several families with alternate leaves and cryptocotylar seedlings: (1) Species which have cataphylls have the first eophylls alternate while (2) species without cata- phylls have the first two eophylls opposite, with subsequent eophylls alternate. The seedling of Pterocarpus hayesi (Fig. 27) resembles that of P. indica more than that of P. officinalis. It differs from both in having the first eophylls opposite. It is transitional from cryptocotyly to phanerocotyly. Some cotyledons, probably photosynthetie, emerge from the seed but remain secund, others appar- ently never escape from the seed. erythroxylaceae: Germination phancrocotylar, the cotyledons oblong, uninerved. Eophylls alternate, supracotyledonary, entire, penninerved, convolute, usually stipulate. 1969] DUKE — SEEDLING CHARACTERS 151 ZYGOPHYLLACEAE : Germination phancrocotylar, the cotyledons subsessile, subtriplinerved, entire, elliptic, glabrous. First eophylls supracotyledonary, op- posite, paripinnate (Giiaiacuni). rutaceae: Germination cryptocotylar to phancrocotylar, often anisocotylar. Cotyledons and eophylls often aromatic and punctate, a marginal row of glands frequent. Eophyll sequence gradual in compound-leaved species. Poly- embryony frequent. Citrus limomim (Fig. 28) is cryptocotylar, anisocotylar, the cotyledons se- cund; the eophylls are alternate, punctate, dentate, pennincrved, with alate, armed petioles. Miirraya (Fig. 29) is cryptocotylar with alternate cataphylls and dentate, aromatic eophylls. simarubaceae: I have not yet observed germination in Quassia (Fig. 30) but suspect it is cryptocotylar. The eophylls suggest certain Mimosaceae, Caesal- piniaceae, and Sapindaceae. burseraceae: Cotyledons contortuplicate and subcryptocotylar (Dacryode^ to simple and planoconvex (Tctragastris, Vrotium) or trifoliolate (Bitrsera) and phancrocotylar. First eophylls simple and opposite (Protium, Tetragastris) or alter- ( to trifoholate and opposite (Dacryodes). Aroma often diagnostic. Tetragastris luinamensis (Fig. 31) f with auriculate apiculate cotyledons, the eophylls opposite, supracotyledonary, pennincrved, entire, or basally auriculate. meliaceae: Germination cryptocotylar with alternate cataphylls and eophylls iSwietenia) or with no cataphylls and opposite, entire to dentate eophylls (Gtiarea, Trichilia) to phancrocotylar (McUa, Cedrda). Carapa (Fig. 32) conforms to the Guarea'Trichilia type. Cedrela (Fig. 33) has subsessilc, entire, plane, narrowdy ovoid cotyledons, with alternate, supracotyledonary, trifohohite, often dentate, eophylls. Melia (Fig. 34) has pctiolate, obovoid, triplinervcd, entire cotyledons with the aceroid first eophyll apparently borne at the cotyledonary node. Trichilia cipo (Fig. 35) differs from other species in that the cotyledons sep- arate but are not strictly opposite or secund. MALPiGHiACEAE: Germination cryptocotylar (especially in alate seeds) with- out cataphylls to phancrocotylar, often tardy (1 year in Byrsouima^. Eophylls alternate to opposite, exstipulate, entire, pennincrved, the indument often di- agnostic. Although germination in Bunchosia (Fig. 36) is phancrocotylar, the greenish- white, planoconvex, subsessile cotyledons are probably not photosynthetic. Al- though I found hundreds of these seedlings beneath a parent tree on San Jose Island and all had aerial cotyledons ("epigeal") I believe that, if the seeds w^re planted beneath the surface of the soil, the cotyledons would not emerge ("hypo- geal"). The eophylls are supracotyledonary, opposite, decussate, entire, penni- ncrved, pubescent. [Vol. 56 152 ANNALS OF THE MISSOURI BOTANICAL GARDEN polygalaceae: Germination cryptocotylar, the subsessile cotyledons secund. Cataphylls alternate, grading into the alternate, entire, narrowly ovate eophylls (Securidaca). euphorbiaceae: Germination mostly phanerocotylar with long-petiolate cotyledons, subcryptocotylar in Ilura and cryptocotylar in Hevca, both, however with long-petiolate, secund cotyledons. F.ophyll type, arrangement, indument, venation, glands, and latex often diagnostic. Hippomane already has poisonous latex at the stage illustrated in Fig. 37. The cotyledons, often cyanotic or bron/e-colored, are entire, triplinerved, the petioles with a copious white latex. The First eophylls are supracotyledonary, op- posite, dentate, penninerved, subsequent eophylls alternate and stipulate. anacardiaceae: Germination cryptocotylar (Coiuocladia) to phanerocotylar with secund [AHatardinm excehmn (Fig. 38) and A, occidcntalc] or opposed cotyledons. Fophylls alternate (Comocladia), opposite (Spuudicn) or pscudoverti- cillate {Anacardinm, Mangijcra). Alternate cataphylls usually present in crypto- cotylar species. Foph}l] sequence gradual in compound-lea\ed species. staphyleaceae: Germination phanerocotylar, the cotyledons subsessile, ovate, entire to retuse, subplinerved. First eophylls supracotyledonary, opposite, trifoliolate, stipulate, and stipellate, the leaflets denticulate. Fophyjl sequence gradual (Turpiiiiii). sapindaceae: Germination cryptocotylar with aUernate cataphylls and eophylls (Alluphylns, Mdiococcm, Supiudus) or without cataphylls and with the first eophylls opposite {Cupama, Maytayha, VauUinia, Serjmiia, Thoniiiia); rarely phanerocotylar {Dodovaea). Talisia nervosa (Fig. 39) suggests Mdiococcus, with cryptocotylar germina- tion and alternate cataphylls and bifoliolate alternate eophylls. sabiaceae: Germination phanerocotylar, the cotyledons foliar, plane, entire to undulate, subsessile, penniner\ed. Hypocot>l swollen, glabrous. Eophylls supra- cotyledonary, ahernate, dentate, penninerved (Mcliusma, Fig. 40). rhamnaceae: Germination phanerocotylar, the cotyledons equal, tripli- nerved, long-petiolate. First eophylls supracotyledonary, opposite, penninerved (Coluhriua) or plincrxed (Zizyphus), subsequent eophvlls alternate. tiliaceae: Germination phanerocotylar, often associated with the rotting fruit, the cotyledons petiolate, broadly obovatc, basally subtruncate, triplinerved, entire QApeiha aspera, Fig. 41). MALVACEAE: Germination phanerocotylar, the cotyledons often convolute, reniform triplinerved to multiplinerved, long-petiolate, often punctate. First eophylls supracotyledonary, alternate, stipulate. Indument and venation often diagnostic. Hihisnis codeusis (Fig. 42) illustrates the similarity between seedlings of Tiliaceae and Malvaceae. 1969] DUKE SEEDLING CHARACTERS 153 Fig. 28-40. Fig. 28. Citrus limonum (X 0.4). Fig. 29. Murraya exotica (X 0.6). Fig. 30. Quassia amara (X 0,4). Fig. 31. Tetragastris panamcnsis (X 0,4). Fig. 32. Carapa sp. CDuke 14988; X 0.4). Fig. 33. Cedrcla sp. (X 0.4). Fig. 34. Melia azcderach (X 0.8). Fig. 35. Trichilia cipo (X 0.4). Fig. 36. Biinchosia cor77ifoIia QDnke 1S034; X 0.4). Fig. 37. Hippomane mancinella QDukc 11732; X 0.4). Fig. 38. Anacardium excehum QDuke 8383; X 0.3). Fig. 39. Talisia nervosa (X0.4). Fig. 40. Meliosma punamensis (X 0.4). [Vol. 56 154 ANNALS Ol THE MISSOURI BOTANICAL GARDEN bombacaceae: Germination cryptocotylar to phancrocotylar, the cotyledons ovate to reniform, subsessile to long-pctiolatc, subplinerved, the eophylls supra- cotyledonary, alternate, stipulate, the transition to compound leaves gradual. Hypocotyl often swollen, especially in those species destined to be ''bottle trees." Hampea (Fig, 43) illustrates the convolution of the emerging cotyledons. Pachira is cryptocotylar with both the cataphylls and eophvlls alternate (Fig. 44). All Quararihea examined are cryptocotylar with alternate cataphylls but with the first pair of eophylls opposite or verticillate (Fig. 45). Dried seedlings have the odor characteristic of the adults. (Fig. 46) merocot} sterculiaceae: Germination cryptocotylar to phancrocotylar, often in the same genus (Sterculia, Theohroma). Cotyledons convolute in phancrocotylar, se- cund in cryptocotylar, species of Stercnlia, as e.g., S. costaricense (Fig. 47) and S. apetala, dilleniaceae : Germination phancrocotylar, the cotyledons conduplicate, penninerved Cl^oliocarpiis, Fig. 48). ochnaceae: Germination phancrocotylar (often tricotylar), the cotyledons sessile, enervate, planoconvex, acuminate, often cyanotic, probably not photo- synthetic. Eophylls supracotyledonary, alternate but appearing subverticillate, narrowly ovate, exstipulate, penninerved, more conspicuously dentate than the metaphylls QOuratea, Fig. 49). marcgraviaceae: Germination phancrocotylar, with little or no hypocotyl, the cotyledons subsessile, uninerved (Fig. 50). Although many seeds germinated, none showed development of the epicotyl. Developmental studies of the family should prove interesting, pelliceriaceae: Germination subcryptocotylar, the exocarp falling away and exposing the reddish, sessile, planoconvex, nonphotosynthetic cotyledons. Eophylls supracotyledonary, alternate, involute, exstipulate, weakly nerved (Pe//i- ceria, Fig. 51). clusiaceae: Germination cryptocotylar, the cotyledons secund, cataphylls supracotyledonary, opposite (in all genera studied except Chisia with phancro- cotylar, submultiplinerved, equal cotyledons). Latex present in the youngest seedling studied. Rheedia (Fig. 52) is characteristic of the cryptocotylar species, with opposite, deltoid cataphylls, followed by opposite, entire penninerved eophylls. The illus- trated specimen took a year to germinate. bixaceae: Germination phancrocotylar, the entire, ovate cotyledons weakly triplinerved, punctate-lineate. Eophylls supracotyledonary, alternate, ovate, sub- cordate, subplinerved, punctate-lineate (Bixa, Fig. 53). 1969] DUKE SEEDLING CHARACTERS 155 Fig. 41-54. Fig. 41. Apeiba aspcra (X 0.8). Fig. 42. Hishiscus cocleamis QDuke 13934; X 0.8). Fig. 43. llampea sp. (X 0.4). Fig. 44. Pachira aquatica (X 0.1). Fig. 45. Quarari- hea asterolepis Qlirhtan ]335; X 0.4). Fig. 46, Matisia cordata (X 0.2). Fig. 47. Stcrculia costaricana (X 0.3). Fig. 48. DoUocarpus sp. (X 0.4). Fig. 49. Ouratea lucens QDuke 8298; X 0.4). Fig. 50. Marc^raviaceae (X 3.2). Fig. 51. PcUiceria rhizophorae (X 0.4). Fig. 52. Rheedia sp. QDukc 11980; X 0.4). Fig. 53. Bixa orellana (X 0.6). Fig. 54. Cochlospcrmum vitifolium (X 0.4). [Vol. 56 156 ANNALS OF THE MISSOURI BOTANICAL GAKDEN coCHLOS PERM ACE al; Gcmiination (after 1 year) phaneroeotoylar, the cotyledons short-petiolate, linear-lanceolate, entire, nninerved. Fopliylls supra- cotyledonary, alternate, aceroid. Latex orange QCocJilosperninm, Fig. 54). violaceae: Germination phanerocotylar, the cotyledons subsessile, entire, tripliner\ed, the eophylls supracotyledonary, alternate, penninerved, coarsely den- tate [more so than metaphylls, stipulate (Duke No. 15005)]; or germination cryptocotylar, the cotyledons sessile, not secund, with subopposite cataphylls grading into subopposite eophjlls QGloeospcrmum, Fig. 55). flacourtiaceae: Germination phanerocotylar, the eophylls supracotyledon- ary, alternate, stipulate, penniner\ed, often pellucidpunctate or lineate, usually more conspicuously dentate than the metaphylls. Mayna (Fig. 56) is a typical flaeourtiaceous seedHng. PASSU loraceae: Germination phanerocotylar, the cotyledons long-petiolate, elliptic, entire, plinerved, the eophylls supracotyledonary (Fig. 5 7), alternate, variable in shape and venation; less commonly cryptocotylar (Fig. 58). cactaceae: Germination phanerocotylar, anisocotylar, the cotyledons sub- sessile, plane, broadly ellipsoid, entire, penninerved, ultimately becoming almost equal, with axillary spines; eo]:)hylls supracotyledonary, subopposite, tardily spines- cent, rather like the cotyledons QPereskia, Fig. 59). The anisocotylar, planoconvex, linear cotyledons of Wittia (Fig. 60) are the only leaves on this epiphyte. lecythidaceak: Apparently there are two germination types in Panama's Coiiratari (Fig. 61 and Fig. 62). Correa and Dressier No. 793 is phanerocotylar, the hypocotyl shghlly tetragonal, the cotyledons ovate, rugulose but plane, penni- nerved. In C. pauamoisis, however, the cotyledons remain in the winged seed, followed by supracot}ledonary eophylls, suggestive of the cotyledons in the phanerocotylar species. Catapliylls are absent. The seedlings of EscJnreilera (Fig. 63) and I.ecythis (Fig. 64) conform to the Custavia type (Fig. 65), cryptocotylar with alternate cataphylls and eophylls. Cnstaria snperha (Fig. 65) is cryptocotylar, the epicotyl arising between the corrugated, planoconvex cotyledons, with alternate cataphylls, grading into alter- nate, exstipulate, lanceolate to oblanceolate eophylls, proportionately more dentate than the metaphylls. Carwiana pyriformis (Fig. 66) has very distinctive phanerocotylar gcmiina- tion, the aceroid lobate cotyledons subcircinately embraced in the testa which is raised with the cotyledons. The drip-tips of the cotyledons are very conspicuous as they unfold. The eophylls are supracotyledonary, alternate, penninerved, and more conspicuously dentate than the metaphylls. rhizophoraceae: Germination cryptocotylar, without cataphylls, the eophylls supracotyledonary, opposite, exstipulate, penninerved, entire, glabrous (Rhizo^ phora). 1969] DUKE SEEDLING CHARACTERS 157 Fig. 55-68. Fig. 5 5. Gloespcrmum sp. CDukc 15268; X 0.4). Fig. 56. Mayna sp. QDulie 74963; X 0.4). Fig. 57. Fassiflora sp. CDuJic 13270; X 0.4). Fig. 58. Pas^siflora sp. CDukc 15287; X 0.4). Fig. 59. Pcreskia hico (A X 1.2; B & C X 0.4). Fig. 60. Wittia sp. (X 0.4). Fig. 61. Couratari panamcnsis (X 0.4). Fig. 62. Couratari sp. QCorrea & Dressier 793; X 0.4). Fig. 63. Eschwcilera QBristan 22; X 0.4). Fig. 64. Lecythis sp. QBlume s.n. "mon- key pot"; X 0.4). Fig. 65. Gustavia supcrha (XO.l). Fig. 66. Carinimia pyriformis (X 0.4). Fig. 67. Syzygium sp. CDukc 15302; X 0.8). Fig. 68. Eugenia umlaccense (X 0.4). 158 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 combretaceae: Germination phanerocotylar, the cotyledons reniforni, con- volute, the eophylls supracotyledonary, alternate or opposite, exstipulate. mvrtaceae: Seedlings, like the embryos, diagnostic for certain generic oco 7) ( ruminate planoconvex cotyledons ^Eugenia, Fig. 68; Syzygium, Fig. ( phylls present or absent); phanerocotylar with broadly ovate subsessile cotyledons {Gomideda, Myrcia), or with lanceolate, uninerved cotyledons (Psidium). Punc- tations, indument, and aroma are often diagnostic. melastomaceae: Commonly the germination is phanerocotylar, the eophylls decussate with the cotyledons. Muuriri parvifoUa (Fig. 69), howe\'er, is cryptocotylar, the cotyledons secund in the testa, which is raised above the soil. Supracotyledonary, opposite cataphylls and eophylls are mostly borne in the saine plane. araliaceae: Germination phanerocotylar, the cotyledons broadly ovate, pli^ nerved to penninerved, the eophylls simple, often telescoped at the cotyledonary node; the eophyll sequence gradual. myrsinaceae: Germination phanerocotylar, sometimes operculate, the cotyle- dons equal or anisocotylar, the margins entire or undulate, often punctate or lineate and cyanotic; eophylls usually supracotyledonary, alternate, conduplicatc exstipulate (Ardisia spp., Figs. 70 and 71). sapotaceae: Germination cryptocotylar, without cataphylls but with secund cotyledons, to phanerocotylar, the cotyledons dark green and coriaceous. Eophylls usually supracotyledonary and alternate. Latex usually obvious at an early stage. oleaceae: Germination phanerocotylar, the cotyledons subsessile; eophylls supracotyledonary, opposite. apocynaceae: Germination usually phanerocotylar, the cotyledons ovate, cordate to auriculate, triplinerved. Eophylls supracotyledonary, alternate, lanceo- late to oblanceolate, entire, penninerved. Latex copious (Plumeria). Lacmellea edtilis (Fig. 72) is cryptocotylar, the stalked cotyledons secund; cataphylls opposite, eophylls lanceolate, opposite, weakly nerved. Latex copious, sticky, white. Stemniadenia (Fig. 73) has phanerocotylar, ovate, entire to undulate, gla- brous, penninerved, subsessile cotyledons with copious latex, the eophylls supra- cotyledonary, entire, penninerved, decussate with the cotyledons. asclepiadaceae: Germination phanerocotylar, the ellii^tic cotyledons long-petiolate, penninerved, with white latex. First eophylls supracotyledonary, opposite (Calotropis). cordiaceae: Germination phanerocotylar, the cotyledons deltoid, subsessile to long-petiolate, plicate, undulate to crenulate. Eophylls supracotyledonary, alternate, penninerved, usually more prominently dentate than the metaphylls. Cystoliths and indument often diagnostic. 1969] DUKE SEEDLING CHARACTERS 159 Fig. 69-82. Mouriri parx'ifolia (X 0.4). Fig. 70. Ardisia sp. QDuke 14989; X 0.4). Fig. 71. Ardisia sp. (Dwfee 15132; X 0.4). Fig. 72. Lacmellca cdidis (X0.4). Fig. 73. Stemmadcnia sp. QDuke 13161; X 0.4). Fig. 74. Enallagyna latifolia (X 0.4). Fig. 75. Crcs- centia cujete QBlume 3656; X 0.4). Fig. 76. Coussarca ccrrojcfcnsis (X 0.4). Fig. 77. Pentagonia sp. (X 0.4). Fig. 78. Ccphaelis corrcac QD'ike 15010; X 0.8). Fig. 79. Toco- yena pittieri (X 0.4). Fig. 80. Alibertia edulis QlUume 3734; X 0.6). Fig. 81. Borojoa sp. CBristan 140; X 0.4). Fig. 82. Faramca occidentalis (X 0.4). \(:)L. 56 160 ANNA].S Ol' THE MISSOUIU BOTANICAL GAUDliN vehhenaclal: Ccrniijiation pliancrocotylar, the cotyledons ovate, entire, sub- tripliner\ed, short-petiolecl. Fopliylls supraeotjlcdonar)', opposite, decussate with the cot}Iedons, often more dentate than the metaphylls. Induinent often diagnostic, e.g., the punctate ghinds in Tectoaa impart a red-spotted outhne of {he seedHng to the newspaper in which it is pressed. avtcenniaceae: Germination phanerocot}Iar, one cot\]edon conduplieate about the otlier, slightly unequal, broadly reniforni, subcordate, sometimes emargi- iiate. lioph\]ls supracot)ledonary, opposite, decussate with tlie cotyledons, entire, hmceolatc to o\ate, weakh' penninerxed. solanaceae: Cicrniination plianerocotylar, the cot\ledons ovoid, pennincrved, entire, long-petiolate. noplnlls supracotylcdonary, alternate, exstipulate (Cestrnni). BiGNOMACEAE: Germination usuallv phanerocotylar, the cotyledons narrowly to broadly reniform, ener\ate to pliner\ed, deeply emarginate to bilobate, cor- date to auriculate. Fophylls supracot\ledonary, usually opposite and decussate with the cotyledons, simple (in species with simple or digitate metaph)lls, e.g., Crescentia, Fig. 7 5), dentate (in species with pinnate metaphylls) or compound (in species with decompound inetaplnlls). Enallaguia (Fig. 74) is the only cryptocotylar species studied. Note the opposed cataplnlls Avhich follow the retuse cotyledons. Although the cotvledons may escape from the testa as it putrifies, they are brownish-black and non- photos\nthelic. RL'BiACT-AE: Germination usually phanerocotylar, the cotyledons narrowly to broad!) ovoid, subpliner\ed, entire, subsessile to long petiolate, usually stipulate. l-oph\Ils supracotxledonary, opposite, decussate with the cot\le.dons, stipulate. Tocoyeua pitticri (Fig. 79) illustrates a fairly conunon deviation in which the testa (and/or endosperm) is raised characteristicallv with the cotvledons (cf. also Fig. 76 and 77) whicli tardily escape to become phanerocotylar. This state is but a step away from true cryptocotyly, as in Vavauica occideutalh (Fig. 82). Bllil KX.HAPHV Hailey, T. W. 19 56. Nodal anatonn and \asculatiirc of seedlings. Jour. Arnold Arb. H7: 269-287. Boyd, L. 1932. Monoc()t\lons seedlings: Morphological studies in the postseminal de\el- opment of the embryo. Trans. Proc. Hot. Soe. idinhurgh li I : 5 224. Cahli)uist, S. 1961. C-omparative plant anatomy. Holt, llinehart 8; Wins;on, New York. 146 pp. CoMPTo.N, R. H. 1912. An investigation of the seedling structure in the Leguminosae. Jour. Linn, Soc. London, Butanv 11: 1 122. CoKNi:n, E, J. H. 19S4. The durian theor} extended. II. The arilhitc fruit and the com- pound leaf. Phytoniorph. 1: 152165. CRONyLisT, A. 1968. The evolution and classification of Houering plants. Houghton Mif- flin Co., Boston. 396 pp. Davis, G. L. 1966. Systematic embr\()logy of the angiosperms. Jolin Wiley & Sons, New York. 528 pp. Duke, J. A. 1964a. Prelude to the polyclave. L Embr>o. Roneo, Durham, North Carolina. 30 pp. + iv. . 1964b. Prelude II: Seed, lioneo, Durham, Nortli Carolina. 19 pp. 1969] DUKE SEEDLING CHARACTERS 1 6 1 — . 1965a. Kcvs for the identification of seedlings of some pruniinL-nt wood} species in eight forest types in Puerto llico. Ann. Missouri Bot. Card. 52: ^i4 .^50. . 1965h. Prelude III; Sterile specimens, lloneo, Durliani, North Carolina. 39 pp. ^ — . 1965c. Prelude IV: Flower, lloneo, Diuham, North Carolina. 28 pp. . 1965d. Prelude to a palm polyehne. Ronco, IJurham, North Carolina. 14 pp. , 1965e. Prelude to a legume polyclave. I. Hahit. Ptoneo, Durham, North Car- olina. 8 pp. . 1969. Legume polyclave. 288 eharacler cards for legume genera. 36 pp. Xerox transparency. Columbus, Ohio. Jan. 13. . 1969b. Family polyclave. 770 character cards for spermatophyte families. Battelle Memorial Institute. 77 pp. . (in press). Woody seedlings. In: II. T. Odum (ed.), Radiation Ecology and a Rain Forest. ., C. R. GuNN, and E. E. Terrfi k (unpublished). Prelude to a grass polyclave. 3 30 character cards for grass genera. Eamks, a. J. 1961. Morphology of the Angiosperms. McGraw Hill Book Co., Inc., New York. 5 18 pp. Eahif, F. l\. and Q. Jones. 1962. Analyses of seed samples from 113 plant families. Fcon. Bot. 16; 221-250. Feknai D. M. L. 1950. Graves manual of botany. 8th cd. American Book Co., New York. 1632 pp. Leonard, J. 1957. Genera des Cynomctreae et des Amherstieae africaines (Leguminosae — Caesalpinioideae); Essai de blastogenie applii|uec a la systemati(]ue. Aeademie royale de Belgique, Classe des sciences, Memoires, Collection in-8 \ 30(1677), Fas. 2: 1-314, 2 3 pho:os. FuHisocK, J. 1891. On stipules, their form and function. Pt. I. Jour. Linn. Soc, London, Botany 28: 217-243. . 1892. A Contribution to Our Knowledge of Seedlings. 2 vols. London. 608, 646 pp. Marrero, J. 1949. Tree seed data from Puerto Pdco. Caribbean Forester 10: 11-26. Martin, A. C. 1946. The comparative internal morphology of seeds. Amer. Midland Nat. 36: 513-660. Melville, 1L 1962. A new theory of the angiosperm flower L Kew Bull. 16: loO. MuRLEY, M. 1{. 1951. Seeds of the Cruciferae of northeastern North America. Amer. Mid- land Nat. 46: 1-81. Paliwal, G. S. and N. M. Bhandari. 1962. Stomatal development in some Magnoliaccae. Phvtomorph, 12: 409 412. Reeder T. R. 1957. The embryo in grass systematics. Amer. Jour. Bot. 44: 756-768. Sporne, K. p. 1954. A note on nuclear endosperm as a primitive character among dicotyle- dons. Phytomorph. 4: 27 5-278. Sterbtns, G. L. and G. S. Khush. 1961. Variations in the organization of the stomatal complex in the leaf epidermis of monocotyledons and its hearing on their phylogeny. Amer. Jour. Bot. 48: 51-59. Swamy, B. G. L. and P. M. Ganapathy. 19 57. On endosperm in dicotyledons. Bot. Gaz. 119: 47-50. ToMLiNSON, P. B. 1960. Seedling leaves in palms and their morphological significance. Jour. Arnold Arb. 41: 414-428. ADVENTIVE PLANTS NEW TO THE MISSOURI FLORA (III) BY Viktor Muehlknhach Missouri Hotanical Garden, St. Louis / \bstrac:t Twenty-four ta\a (mostly species) are proposed as additions to the adventivc flora of Missouri. Of these, 11 are nati\e to tlie U.S. mostly originating in the South or West, four are European but are already known from the U.S., two are Asiatic, one is from Tropical America and six are escapes from cultivation. In 1957 and 1960, the author published lists of adventivc plants which were collected on the railroads in the City of St. Louis and were new^ to the Missouri flora. The hrst Hst contained 16 plants, and the second 33 (Muehlenbach, 1957, 1960), the great majority of which were species, but in a few cases varieties. J. A. Steyermark (1963) included a further 25 unpublished plants in his Flora of Missouri — seven in the text, and another 18 in the supplement. Since that time the writer has found a few more new plants. Omittint^ all not surely deter- mined plants, the total is 24. Six are escapes from cultivation. Four originate in Europe, but are already known from the U.S., two are Asiatic species, one is con- sidered to come from Tropical America. The remaining 1 1 plants are native to some area of the U.S., most (at least 8) originating in the South or in the West, and one possibly coming from the Atlantic coast. All plants were sent to speciahsts for determination or at least for corroboration, as mentioned in the description of each plant. The writer acknowledges with gratitude the help so received. Vouchers for these records (with one exception — Ver1?e}ia halei^ have been deposited in the herbarium of the Missouri Botanical Garden. The initials and numbers in parentheses below are those of the writer in field collections. Eragrostis oxylepis (Torr.) Torr. One small colony along the fence of the Produce Market at North Market Street, St. Louis Avenue freight vard of the Norfolk and WY-stcrn Railwav (at that time Wabash R.R), October 29, 1961 (V,M. 1931). Determined by G. B. Van Schaack. According to Hitchcock (1951) it is a characteristic southern species (in older hteraturc fre(|iiently called E. sccundiflora Presl.). It is dis- tributed through almost all states from Fla, to Calif. Even though it is found in four neighboring states (Tenn., Ark., Okla., Kans.) one would licsitate to consider this iirass a native to Missouri. Too manv t\pical adventives were found in its vicinity, and it also disappeared the following year, although no signs of any threat to the vegetation could be detected. All this suggests that E. oxylcpis is an adventivc plant in this case, since so many of these plants do disappear after one season ^^'ithout anv obvious reason. Ann. Missouri Bot. Card. 56(2): 163-171, 1969. [Vol. 56 164 ANNALS Ob THE MISSOURI BOTANICAL GARDEN Jhuchs gerardi Loisel. Two almost confluent colonics in tlic southern (now abandoned) part of the Ranken yard of the Ternnnal R.R. Association, east of Conipton Ave., June 14, 1964; June 27, July 24, August 29, 1965 (V.M. 23S6, 2476, 2505, 2522). Also observed in later years, altliougli a Hre licavily damaged the whole vegetation in this locality in the autumn of 1966. Determined by F. J. Hermann. A com- mon plant along the entire Atlantic coast from Me. to Fla., also in Wash. (Jones, 1936), but quite a few times introduced inland. The nearest place is Ind. (Deam, 1940 — along R.P,.!). Sjiiall (1933) adds Great Lake Lowland, 111. to Mich., but later Jones and Fuller (1955) stated that there is no evidence for the occurrence of jniicits geraidi in III. Finally Fcrnald (1950) mentions Minn., Tidestrom (1925) Utah (in Salt Lake Citv in salt marshes!) and Stevens (1961) \. Dak. Polygonuhi pe)is}ivaiiiciiiii T . var. rosacflonim J. B. S. Norton. One specimen on the levee of the Mississipjii River, between the Fads and Veterans Memorial Bridges, in the vicinity of a Terminal R.H. Association track, September 15, 1963 {V.M. 2222): five specimens on the right of-way of the Missouri-Pacific R.R., between Davis St. Junction and Tesson St., October 12, 1963 (V.M. 2275); one huge specimen in the Baden freight yard of the Missouri- Kansas-Texas R.R., November 4, 1967 (V.M. 2914). Determined by II. A. Wahl (as were all the other Polygouniu forms). Gleason and Cronquist (1963) do not mention this variety of the common plant, but Fcrnald (1950) cites this variety, which he considers a nati\c, in Del., Md. and Va. The status of this variety in Missouri is uncertain. In Jiumerous floras of states between the Atlantic coast and Mo. it is not mentioned, so it could well be an introduced plant for us. Pulygonum Lipathifuliiini L. var. salicijoliuui Sibth. Vcw specimens in the now abandoned \\estern part of the Pumken yard of the Terminal R.R. Association, September 7, 1964 (V.M. 243Ha); one small colony in the North St. Louis freight yard of the Burhngton R.R., August 21, 1966 (V.M. 2709). Gleason and Cronquist (1963) do not mention it at all, but Fcrnald (1950) cites a number of states, inostly northern, throughout the United States. Generally it is considered a native plant, but St. John (1963) calls it a European weed. However, Hegi (1911, 1957) does not mention it in either edition of his flora. Polygonum lapalJiifoIiiiDi L. var. prostration Wimm. One specimen on the right-of-way of the Manufacturers Railway, south of Victor St., July 27, 1963 (V.M. 2150). Fcrnald (1950) gives this description: "shores, railroad-yards and waste places, local, N.S. to Alta. and Wash., s. to N. J., e. Pa. and Calif. (Nat/d. from Fu.)" This plant has apparently recei\ed little attention, because no other report could be found in the literature consulted. 1969] MUEIILENBACH MISSOURI ADVEMIVES 165 Polygo)in))i scahnini Mocnch. Three speeiniens in the Inbound freight yard of the lloek Ishind, Friseo and Chicago and llastern IHinois \\.l\., OTallon St., October 23, 1966 fV.AI. 2766). According to Fernald (1950) tlie distribution is ". . . New Fngl., N.Y., Mich., (i\atzd. from Fur.) )5 5) and Colo. 111. (1 ) cited under other names. Fhe adventi\e nature of P. i^cahrnm is generally accepted, althouoli Cleason and Cronquist (1963) think that this plant is probably botli nati\e and introduced lure. Volygouiim persicaria F. forma alhiflora Millsp. One colony in Tyler St. ireiglit \ard of the Terminal R.IF Association, Au- gust 2, 1966 (V.M. 2695); one colony on tlie riglit-of-way of the same railroad between Destrehan St. and Mckinley Bridge, September 18, 1966 (V.M. 2731). This form of tlie common Volygonuni persicaria, naturalized from Europe, seems to be quite frequent too, l)ut in most floras it is mentioned only in the descripti\e (1963) f (1950) mentions tlie rare forma albifl not gi\c any more precise distribution. It is cited in only two of the floras con- 1 and Core (1953) and Minn, by Fakela (1965). 'cnv test (his No. P875, coll. July 27, 1967) on V.AF 2 73]. The growth of this test specimen is bigger than the original, and the color of the flowers is pure white. Beta vulgaris F. One specimen on die big a acant place adjoining the right-of-way of the Frisco IFR., east of Kingshighway (now completely built over), June 2, 1962 (V.AF 2 963). Determined by 11. A. W ahl. There are only a few records in the United States for Beta vulgaris. As frecjuendy happens with other escaped plants, it has been for the most part o\erlooked or simply neglected. Only a few floras have apparently tried to register all of them. Fspecially careful registration of escaped plants has been done in New Fngland. Beta vulgaris is cited from four states — Me., N.IF, Mass., Conn. (Bean, Flill, and Faton, 1958). From the other coast, Calif, is mentioned and Munz and Keck (1963) write: "Fscaping from gardens and sometimes natur. in low damp places." The root of the St. Fouis plant is remarkably thin, so it is not probable that it is a direct garden escape. Cheiu)pocliiiiii uatsoiii A. Nels. One specimen in the Carrie Avenue freight yard of the Hock Island R.R., along the car cleaning track, Jul\ 1, 1961 fV.AF 1795). Determined by H. A. Wahl. According to him (1954) N.M., Ariz, and Kans. In connection with the examination of this collection he wrote "An item of particular interest was the collection of C. watsoni, V.AF 7 795, which is essentially a species of dry areas in the Rocky Mountain region and this iv; flip ^1r^f roll(>(tion as an adventivc in a weedy habitat" (in litt.). 166 AiNNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 J Atriplex truncata (Torrcy) Gray One small colony in the North St. Louis freight yard of the Burlington R.R., uie 30, 1956 {VM. 974); one colony on the right-of-way of the Manufacturers Railway, south of Arsenal St., June 7, 1959 {VM. 1521); possibly on the same place one colony (June 23, July 27, September 15, 1963 {V.M. 2106, 2147, 2210). H. A. W'ahl expressed his opinion that some of these collections could be A. truncata. C. Frankton from the Canada Department of Agriculture in Ottawa confirmed them. He also wrote, "This annual occurs as a natixe west of 105" Long, in the Northwestern States and Canada. I have not encountered it (in litt.) western states. AtripL'x hetewi^peniia Bunge. One colony on the right-of-way of the Lerminal ll.R. Association east of Geraldine Ave., opposite A.C.L. Haase Co. Food Products plant, July 22, 1956 (V.M. 1026). Frankton discovered this species amidst a collection of A. hastata L. In a paper with Bassctt (1968) he now one of the least known ad\cnti\es in North America. According to them it has spread rapidly in western Canada, especially B.C. in this decade. The first collection was made in 193 7. The herbarium of the Canada Department of Agri- culture in Ottawa, Ont. also has collections from the U.S.: N. Dak. (Fargo, twiVe, including what is apparently the oldest collection in the U.S. made by Stevens in 1954) and Wash. (Oro\illc). Weber (1966) is the only U.S. writer who mentions this plant (Colo., Larimer Co., and Mont., Powell Co., found in both places by G. N. Jones). According to the Canadian paper, Atriplcx Jwtewspernia is considered to be native to Furnpcan and Asiatic Russia. It apparently began to spread in the third decade of this century, at first in Europe, then in North America as well, but because of confusion with other species of this genus (espe- cially A. hastata L. and A. iilteiis Schkuhr), it was onlv tardilv recocmi/ed Salsola colUna Pall. (now abandoned) ii the Terminal R.R. Association, east of Compton A\e., October 4, 1964 fV.AL 2451h); several specimens in T}lcr St. freight vard of the Terminal ]\.\\. Associa- tion, July 17, 1965 (V.M. 2503). In both places growing with Salsola kali L. V.M. 2503 was tentatively determined b\ H. A. W'ahl as Salsola colllua. C. Frank- ton confirmed this finding, as well as an earfier collection, V.M. 2451h. He adds. We have a sheet from Semipalatinsk \\hkh matches >our collection \ery well" (in litt.). This adxentive species is (]uite scarce in the U.S. Pohl and Gillespie (1959) mention Minn, (first discovered 1937 by J. W. Moore), Colo, and Li. Later this plant was found in N. Dak. (Stevens, 1961). It is a widely distributed Asiatic species, reaching into the southeastern corner of European Russia. Vhiladelphns curunariiis L. At least six shrubs over 2 m high on the right-of-way of the Missouri-Pacific R.R. at the crossing with Pennsylvania A\e., along a high stone wall, May 10, 1969] MUEHLKNBACH MISSOURI ADVENTIVES 167 1964, May 28, 1966 (V.M. 23?>(), 2614). Several more shrubs are growing on the exeeedingly steep southern embankment of the same railroad east of Broadway but were not eoUeeted sinee the ground here was verj' unstable. Determined by D. B. Dunn. Philadelphus cuwnariiis is widely eultivated in the U.S. and often escapes. New England reports arc available from five states (Me., Vt., Mass., R.I., Conn., Seymour, 1969). Jones and Fuller (1955) and Deam (1940) have en- countered it in 111. and Ind., which is in keeping with the careful search for escapes that has been jnade in tliese two states. In addition this shrub has been reported from N.Y. (House, 1924) and la. and S. Dak. (Rydberg, 1932). Fruinis armeniaca L. One small shrub without fruits or flowers. The St. Louis A\enue freight yard of the Norfolk and Western Railway, September 22, 1968 (VM. 2987). Before the next visit it had been cut down, but in June 1969 some suckers could be ( ) (Gleason and Cronquist, 1963 Pi^um sativum L. var. sativum (P. sativum L. s. str.). One small colony with well-developed white flowers in the Bulwer yard of the Terminal R.R. Association, south of the station building of the Carrie Avenue freight yard of the Rock Island R.R., May 8, 1966 (V.M. 2597). Determined by 11. Culler. Like many other garden escapes, peas are very rarely reported. Mun/ and Keck (1963) mention it as a white flowering waif from cultivation in Cahf. More frequently, the varieties of Pisuin sativum are not distinguished. Crutuu li)iilhei))ieriaiius Scheele. One small colonv on a siding of the Missouri-Pacific R.IL, along the National Lead Co. plant west' of Macklind Ave., June 2, 1962 fV.AL i965j. Determined by L. H. Shinners. It is a western and southwestern plant. Kearney and Peebles (1960) mention Ariz, (introduced?), Kans. to Tex.; it is reported for Okla. (Stemen and Myers, 1937) and for N.C. (Radford, Ahlcs, and Bell, 1968). Partheiiocissus tricuspidata (Sieb. & Zuce.) Planch. One specimen in the now abandoned western part of the Ranken yard of the Terminal R.P.. Association, west of Compton Ave., creeping on the ground, Au- gust 2, 1964 (V.M. 2411). Determined by D. B. Dunn. One small specimen climbing on the wall of the Frigidaire Warehouse along a siding of the Frisco R.R., east of Kingshighway, May 29, 1965 (V.M. 2466); one specimen climbing on the wall of Ahrens & McCarron building along the Christy Lead of the Missouri- Pacific R.R., north of Beck Ave., May 19, 1968 (V.M. 2927). Determined by \V. H. Lewis. In these two localities PartJicnocissus tricuspidata could be a garden escape, because it was observed on other buildings in the vicinity. In the first localitv, however, there are no gardens or houses nearby, and an introduction from outside is possible. Also, it disappeared shortly after the discovery. As with so many other cultivated plants, the reports from other parts of the country are [Vol. 56 168 ANNALS OF THE MISSOURI BOTANICAL GAP.DEN o scanty, rcrnald (1950) gives the distribution Mass. to O., and it is included as an escape in III. (Jones and Fuller, 1955) and Okla. (Waterfall, 1950). Gaura sinuata Nutt. ex Ser. One small but dense colony on the most western track of the Baden freight yard of tlie IMissouri-Kansas-Texas R.R., October 12, 1966 (V.Af. 2760). Deter- mined by P. Raven. The area of origin, according to liim (in litt.) Okla., and it is introduced sporadically in the Southeast. Further introductions are reported from Calif. (Mun/ and Keck, 1963) and from N. Y. City (House, 1924). Finally, it is found also in Kans. and Ark. (Rydberg, 1932) and S.C. (Radford, Allies, and Bell, 1968), but with no ju( states. There is no doubt that Gaura sinuata is an ad\enti\e in St. Louis, llie track where it was found was used intensivelv in previous \ears for cleanina the freight cars. Numerous other adventives had grown there, as is often the case in similar places. A httle surprising was the fact that the cleaning of the cars was discontinued some years before, and e\en the track from tlie frcidit vard itself disconnected so tliat no more traffic wliatsoever couki occur. On tlie same day another novcit) for Mo. — Ph) sails acutifcAia^wds discovered nearby. The place was visited every summer several times, and it is thfficult to understand that two such conspicuous plants could be constantly o\erIookcd for several years. Between this track and the freight \ard is a \ery wide strip of vacant land richlv covered with vegetation, and it is not reasonable to assume that both plants represent a recent introduction from the site of the freight yard. During the study of plant introductions, one runs from time to time into pu/zles like this which su^nrcst that much more study is needed on distribution mechanisms and factors affecting seed dormancy in wild plants. Torilis arvoisis (Huds.) Link Right-of-way of MissourLRacihc R.IL, north of Flwood St., June 13, 1954 (V.M. 151); one colony along the northern sidings of the grain elevator of the Missouri-Pacific R.R., south of Steins St., September 2, 1956 (V.M. 1060); one colony in Lindenwood freight yard of the Frisco ILR., east of die R.R. bridge over the Des Peres sewer, July 21, 1962 (V.M. 1987); abuudant along a siding of the Manufacturers ILiilway between its shops and Potomac St., Julv 15, 1967 (V.M. 283 6). Tuiilis arreusis is in all probability the most interesting and per- plexing find. On two separate occasions the plants were sent to M. F. Mathias and were identihed as T. arveiisis. This unexpected result suggested a close examination of the Torilis material of the Missouri Botanical Garden. The 23 sheets determined as Torilis japoiiica (Iloutt.) DC. from Mo., were all T.arreusisl Of 16 sheets from other states, 13 (from nine states) were also found to be T. arreiisis. Onlv two sheets (from two states) represent T. jajwiiica. In one case no decision was possible. Mathias and Constance (1944) give the characters of the two species: T. japouica has an involucre of several bracts, one to each ray, while T. arvciisis either lacks an involucre, or has a single bract. They also differ in fruit characters. Meanwhile M. F. Mathias informed me (in litt.) that J. T. Howell (1962) found this same confusion to be general throut'hout studies ^1 — ^"- .....wu^ 1969] MUEHLKNBACH — MISSOUHI ADVENTIVES 169 of these taxa in the U.S. He cites "frequent misapplication of the name T. japonica {T. aiithriscus) in North America. Whereas most floras of the central, southern and eastern United States refer tlie loosel) branching hedge-parslc) to T. japonica (or T. aiithnsciis} most of the plants examined (CAS, UCO) belong to I . ar- roisis. Torilis japonica (Iloutt.) DC. appears to be a rare plant in the United States . . .". Until then T. arveusis was listed only from Calif, and Ore. Tlie Cal- ifornian herbarium material increased the number of states to 1 1 , leaving only three states with proven T. japonica. The herbarium material of the Missouri Botanical Garden increases the Torilis anensis states by three (Ala. and \'a. be- sides Mo.), and the Torilis japonica states by one, Ind. G. Hegi underlines that both species, which have quite distinct geographical distributions, were frequently confused even in Furope. Apparently this paper of Howell has not receixed the attention it deserves. In some newer floras no reference to this controxersv is given. The only exception seems to be l^adford, Ahles, and Bell (1968) in their flora of the Carolinas, but in their preliminary guide (1964) even they considered T. arveusis and T, japonica as synonyms. Now that T. arveusis is known to be quite frequent on the R.R. in St. Louis, the question remains whether T, japoiiica will also be found. AscJepias suhverticillata (Gra\') \^ail One big colony spread on several sidings of the Burlington R.R. south of Destrehan St., July 28, 1962, July 28, 1963 (V,M, 2000, 2152). Still present in the summer of 1969. Determined b\ the late R. E. W^oodson, Jr. In his mono- graph (Woodson, 1954) he gi\es the range of this western species as Ariz., Colo., Ida., N.M., Tex. and Utah. Kearney and Peebles (1960) added Kans. Kearney and Peebles emphasi/c that this plant is very poisonous to livestock and "should be eradicated from pastures whenever practicable.*' At its locality in St, Louis it can not create any danger. One thing was striking, namely the extreme viability. The tracks were frequently treated with weed killers, and once the whole dried vegetation was burned down to the bare ground, but the colony flourished again, as though nothing had happened. Most probably adventivc in Mo. Ipoinoea purpurea (L.) Roth \ar. diversifolia (Lindl.) O'Donell Right-of-way of the Missouri Pacific R.R. between Reilly Ave. and the signal box at the Davis St. Junction, September 25, 1955 (V.M. 811), October 14, 1961 (V.M. 1912), probably both collections are from one locality. Determined by ^\^ H. Lewis, It seems to be a very rare plant. In the floras consulted Ipo}uoea purpurea with 3-lobed leaves is mentioned only bv Gleason and Cronciuist (1963) (1940) (1953) has studied Ipomoea purpurea var. diversifolia in material from N.M., Tex. and Ariz. Verbena halei Small. One specimen in the Carrie Avenue freight yard of the Rock Island R.R., July 4, 1957 fV.M. 1256); two specimens in the St. Louis Avenue freight yard [Vol. 56 170 ANiNALS Ol THE MISSOURI BOTANICAL GAKDliN of the Norfolk and Western Railway, June 28, 1958 (V.M. 1423); at the same plaee four plants (only two colleeted) May 22, 1960 (V.M. 1607). Determined by n. N. Moldenke. Aeeording to him (1963) this speeies is wide-spread in the southern states from N.C. to Ari/. Colleeted also in two neiizhborins states— o ""' o Ark. and Okla. In the first mentioned state the nearest point to St. Louis is Stone County, but Moldenke's opinion (in lit.) is that these findings represent aceidental introduetions. Physalis aaiiifoha (Miers) Sandw. (P. xvrightii Cray) One speeimen in the same locality in the Baden freight yard of the Missouri- Kansas-Texas R.R. as Gaiira siniiata, and discussed there, October 12, 1966 (1958) (1967) he used an older name, P. aciitifolia. Two states could be added: N.M. (Wooton and Standley, 1915) and Colo. (Mun/ and Keck, 1963). Ciicumis sativHs L. One specimen in the Carrie Avenue freight yard of the Rock Island R.R., on the edge of the roadway separating it from tlie Bulwer yard of the Terminal R.R. Association, September 9, 1961 (V.M. 1847). Determined by H. Cutler. As with other garden escapes, few records are known from other states. Gleason (1952) and Fernald (1950) speak in general terms: "frequently spontaneous in waste ground" or "incline to appear spontaneously." Only 111. (Jones and Fuller, 195 5) has a concrete report. In spite of the dangerous locality, the plant liad escaped injuries and even formed edible cucumbers. LlTERATLKE Cl iJEAN, R. C, A. F. Hii.1., & H. J. Eaton, 1958. Twelfth report of the committee on plant distrihution. Rhodora 60: 297-305. Deam, (;. C. 1940. Flora of Indiana. Binford Printing; Press Co., Tntlianapolis. Fernald, M. L. 1950. Gray's Manual of Hotany, ed. 8. American Book Co., New York. Frankton, C. & I. J. Bassett. 1968. The genus Airiplcx (Chcnopodiaccae) in Canada. 1. Three introduced species: A. hetcrospcrwa. A. nblougifolia, antl A. hortcnsis. Canad. Jour. Dot. 46: H09-1313. Gleason, H. A. 19 52. The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada. N.Y. Bot. Card., New York. & A- CHONyuiST. 1963. Manual of Vascular Plants of Nortlieastern United States and Adjacent Canada. Van Nostrand, Princeton, N.J. Harrington, H. D. 1964. Manual of the Plants of Colorado. Sage Books, Denver. Hegi, G. 1911. Illustricrte Flora von Mittel-Furopa. Band 111. Dicotyledones (I. Tcil). J. F. Lchmann's Verlag, Munchen. • 1957. Illustricrte Flora von Mittcl Fiiropa. Band III 1. Teil. Dicotyledones, ed 2. Carl Hanser Verlag, Munchen. Hitchcock, A. S. 1951. Manual of the Grasses of the United States, cd. 2. rev hy A. Chase. U.S.D.A., Misc. Publ. 200. House, II. D. 1924. Annotated list of the ferns and flowering plants of New York State N.Y. State Mus. Bidl. 254: 5 7-59. Howell, J. T. 1962. Five days to Reno: A botanical motnrlogue. Leafl. West Bot 9- 233-242. Jones, G. N. 1936. A botanical survev of the Olympic Peninsula, Washington. Univ. Wash. Puhl. Biol. 5: 1-286. 1969] MUKHLKNBACII — MISSOURI ADVENTIVES 171 & G. D. Fuller. 1955. V^ascular plants of Illinois. Univ. of Illinois Press, Ur- bana, and Illinois State Miis., Springfield. Kearxey, T. II, & R. H. Peebles. 1960. Arizona Flora, ed. 2. Univ. of California Press, Berkeley & Los Angeles. Lakela, O. 1965. A Flora of Northeastern Minnesota. Univ. of Minnesota Press, Min- neapolis. Mathias, M. F. & L. Constance. 1944. Umhelliferae. N. Amer. Fl. 28B: 45-295. MoLDENKE, H. N. 1963. Materials toward a monograph of the genus Verbena. Phytologia 9: 160-177. MuEHLENBACH, V. 1957. Adventitious and escaped plants new to Missouri. Rhodora 59: 27-31. . 1960. Adventive plants new to the Missouri Flora. Amer. Midi. Nat. 64: 161-168. MuNZ, P. A. & D. D. Keck. 1963. A California Flora. Univ, of California Press, Berkeley & Los Angeles. O'Donell, C. A. 1953. Convolvulaeeas nue\as o criticas. IV. 1 illoa 26: 353-400, PoHL, R. W. & J. P. Gillespie. 19 59. Distributional and cytological notes on Sahola collUia. Pihodora 61: 265-267. Radford, A. E., H. E. Ahles, & C. W, Bri \ . 1964. Guide to the Vascular Flora of the Carolinas. Ihe Book Exchange, Univ. of North Carolina, (hapel Hill. — . 1968. Manual of the Vascular Flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill. Hydblhg, p. a. 1932. Flora of the Prairies and Plains of Central North America. N.Y. Bot. Gard., New York. St, John, H. 1963. Flora of Southeastern Washington and of Adjacent Idaho, ed. 3. Out- door Pictures, Escondido, C>alif. SEYMOLfR, F. C. 1969. The Flora of New En^^hind. Tuttle, Putland, Vt. Small, J. K. 1933. Manual of the Southeastern Mora. Univ. of North Carolina Press, Chapel Hill. Stemen, T. R. & W. S. Myers. 1937. Oklahoma Flora. Harlow Publ. Corp., Oklahoma City. Stevens, O. A. 1961. New records for North Dakota, llhodora 63: 39-46. Steyekmark, J. A. 1963. Flora of Missouri. Iowa State Univ. Press, Ames. Straushaugh, p. D. & E. L. Core. 1953. Flora of West Virginia (Part 11). W. Va. Univ. Bull. Ser. 53, 12(1): 271-570. TiDESTROM, I. 1925. Flora of Utah and Nevada. Contr. U.S. Nat. Herb. 25: 1-663. Wahl, H. A. 19 54. Preliminary study of the s<^'nus Cheno-podiiim in North America. Bar- tonia 27: 1-46. Waterfall, U. T. 19 50. Some additions to the Oklahoma flora. Rhodora 52: 3 5-41. . 19 58. A taxonomic study of the genus Physalis m North America north of Mexico. Rhodora 60: 152-173. . 1967. Physalis in Mexico, Central America and the West Indies, llhodora 69: 319-329. Weber. W. Wo Phys. Sci. 23: 1-24. )DsoN, R. E., Jr. 1 Bot. Gard. 41: 1-211. Ann. Missouri WooTON, E. O. & P. C. Standley. 1915. Flora of New Mexico. Contr. U.S. Nat Herb. 19: 1-794. NOTES ON THE GALAPAGOS EUPHORBIEAE (EUPHORBIACEAE) BY Derek Burch^ Missouri Botanical Garden and Department of Botany, Washington University, St. Louis Abstract The tribe is represented by 1 1 species in two genera, nine of which (eight species of Chamaesycc and the only Euphorbia^ are endemic. A novelty, Chamaesyce abdita, is de- scribed, and a key furnished to the species of this genus. The morphological diversity found in the Linnaean genus Eiqyhorhia has sug- gested to a number of workers that the 1500 or more species might well be divided into several, more homogeneous, groups. The element segregated as the genus Chamaesyce S. F. Gray appears to be natural and relatively well defined, and has been accepted as distinct in describing this tribe for the Flora of the Galapagos Islands (Wiggins & Porter, Stanford University Press, in press). This necessitates several new^ combinations, and recent collections have included one that must be described as a new species. Eight of the ten species of Chamaesyce found are endemic. They share a number of characteristics, and for the most part are rather distinct from other New W^orld species. Several of them are shrubs and even the herbs have a strong tendency to woodiness at the base or are perennial by their rootstock. In many cases the stems are somewhat swollen at the nodes and pseudoarticulate. Most have a somew^hat coriaceous texture, short-petiolate leaves, a reduced number of staminate flowers in each cyathium, and cyathial glands which are dark-colored (at least after drying). Tlie amount of infra-specific variation is high (as seems to be the case throughout the genus), but there is not much overlap in general appearance of the species. (Figures 1 and 2). The other two species which occur are Chamaesyce ophthalmica (Pers,) Burch, a common but rather patchily distributed weed of the American tropics, and C. hirta (L.) Millsp., a pan-tropical weed. The two species often grow in mixed stands elsewhere in the New World, with C. hirta by far the more aggres- sive of the two, but this is not the case in the Galapagos where C. hirta is known from only one collection. Key to Species of Chamaesyce 1. Capsule glabrous. 2. Leaves linear, mature stems leafless with leaves clustered on short spurs. 1. C. viminalis 2. Leaves linear-lanceolate or broader, leaves mostly on laterals when stems ma- ture but internodes not condensed to form spurs. 3. Base of leaf blades deeply cordate enveloping stem, blades orbicular; seed plump, obscurely angled, ca 1.2 mm long, faces smooth .... 2. C. amplcxicaidis ^Work on the genus Chamaesyce supported by National Science Foundation Grant No. GB- 5778 (Principal Investigator Derek Burch). Ann. Missouri Box. Card. 56(2): 173-178, 1969. [Vol. 56 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. Base of blades at most cordate and scarcely extending over stem, blades elliptic or narrower; seed strongly angled, faces flat, variously sculptured or, if smooth, seed < 1 mm long. 4. Leaves linear-lanceolate; faces of seed irregular with broken, ± trans- verse ridges 3. C. puuctulata 4. Leaves elliptic; faces of seed smooth or with unbroken transverse ridges. 5. Stipules ca 1 mm long, bifid or with lacerate margins; cyathial gland appendage merely a light rim to gland; seed > 1.2 mm long, faces with broad transverse ridges 4. C. rcciirvu 5. Stipules ea 0.5 mm long, lacerate to base; gland appendage prominent, to twice width of gland; seed < 1 mm long, faces smooth 5. C. hindhcHsis 1. Capsule pubescent. 6. Cyathia rather few, not grouped in dense glomerules. 7. Leaves narrowly elliptic to linear, 12-2 5 mm long; plants tomcntose throughout; faces of seed with fine reticulate markings 6. C. galapagcia 7. Leaves ovoid to suborbicular or, if elliptic, < 10 mm long; plants hirsute or strigose throughout; faces of seed essentially smooth. 8, Seeds plump, obscurely angled; leaf blades broadly ovoid to sub- orbicular with a cordate base; plants hirsute throughout. . 7. C. nummidaria 8. Seeds strongly angled, faces flat; leaf blades elliptic with base oblique; plants strigose throughout 8. C ahdita 6. Cyathia many, collected into dense glomerules. 9. Glomerules terminal on main axes (often in pairs) or on leafy laterals. 9. C. opIitJudnnca 9, Glomerules axillary on short leafless peduncles or terminal 10. C. hirta 1. Chawaesyce viniinea (Hook, f.) Burch, conib. nov., based on Euphorbia vhninea Hook, f., Trans. Linn. Soc, London 20: 184, 1847. Type: Albc- marie Island, Macrae s.n. (K). — Fig. la. E.viyntnca f. alhcmarlcnsh Eobinson & Greenm., Amer. Jour. Sci. 3rd ser. 50: 138, 1895. Syntypes: Albemarle Island, Macrae s.n. (K); Baur 251 (GH), 252 (GH). E. vitninca f. jacohcnsis Hobinson & Greenm., loc. cit. Tvpe: James Island, nr Orchilla Bay, Baur 249 (GH). E. viminca f. casteUana Robinson & Greenm., loc. cit. Type: Tower Island, Baur 247 (GH). E. viminca f. chathatnoisis Robinson & Greenm., loc. cit. Tvpe: Chatham Island, lower re- gion, SW end, Baur 245 (GH). E. viniinea f. carolcnsis Robinson & Greenm., op. cit. 139. Tvpe: Charles Island, Andcrsson 27 (GH). E. viminca f. barringtoncnsis Robinson 8; Greenm., loc. cit. Tvpe: Barrington Island, Baur 244 (GH). E, viminca f. jcrvcnsis Robinson & Greenm., loc. cit. Type: Jervis Island, Baur 250 (GH), E, viminca var. ahingdoncnsis Robinson & Greenm., loc. cit. Type: Abingdon Island, Baur 246 (GH). The variety and forms proposed by Robinson & Greenman ean not be dis- tinguished when all the collections now available are assembled. The leaf size, shape, and arrangement, on which their separations were based, are extremely variable even in one collection, and no consistent pattern of variation could be found. In mature plants tlie linear leaves are carried on spurs. The collar-like stipules persist after the leaves have fallen. The capsules are glabrous when young. No mature fruits have been seen, and this might bear some relation to the separation of the sexes that occurs in many 1969] BURCH — GALAPAGOS EUPHORBIEAE 175 cyathia. Some collections have cyathia with well developed flowers of each sex, but in most cases those with normal-looking female flowers have only tiny males present while those with normal males have only tiny traces of a female. In at least one case both "male" and "female" cyathia occur on the same plant. Chamaesyce vagiunlata (Griseb.) Millsp. of the southern Bahamas is similar in appearance to this species. 2. Chamaesyce aniplexicaulis (Hook, f.) Burch, comb, nov., based on Euphor- hia amplexicaulis Hook, f., Trans. Linn. Soc, London 20: 183, 1847. Type: Chatham Island, Danvin s.ji., end of Sept. 1835 (K). — Fig. lb. In his original description Hooker noted a resemblance between this species and one from the Bahamas. He was probably referring to C. huxifolia (Lam.) Small which has similar though larger capsules and smooth plump seeds but does not show the strongly clasping leaf bases, lacerate stipules and black cyathial glands of C. amplexicaulis. 3. Chamaesyce punrlulata (Anderss.) Burch, comb, nov., based on EupJiorhia piuictulata Anderss., Stockh. Vet. Akad. Handl. 1853: 235, 1854. Type: locis sicissimis insulae, Albemarle, Audersson s.n. (S — not seen). — Fig. 2a. Euphorhia diffusa Hook, f., Trans. Linn. Soc., London, 20: 184, 1847. Type: Albemarle Island, Macrae & Cuming s.n, (K). Non Jac(|uin, Misc. 2: ^11, 1781, ncc Dufour, Bull. Soc. Bot. Fr. 7: 445, 1860. E. ar tic III at a Anderss., Stockh. Vet. Akad. Handl. 1853: 236, 1854. Type: locis siccis, lapi- dosis regionis inferioris et mediae insularum Chatham et Indefatigable, Andersson s.n. (S — not seen; ? isotypc: K). Non Aublet, Hist. PL Gui. Fr. 480, 1775. E. andcrssonii Millsp., rubl. Field Mus. Nat. Hist. Bot. Ser. 2: 63, 1900, based on E. articu- lata Anderss. The linear-lanceolate leaves, together with the 4-angled irregularly-ridged seed in glabrous capsules, distinguish this species. The name E. diffusa Hook. f. has been widely used for this taxon but is a homonym and illegitimate. Since Andcrsson's epithet piuictulata is available it seems better to transfer this rather than tr\ing to justify the retention of the well-known epithet. 4. Chamaesyce recurva (Hook, f.) Burch, comb, nov., based on Euphorhia recurva Hook, f., Trans. Linn. Soc, London 20: 182, 1847. Type: Chatham Island, Danrin s.n., end of Sept. 1835 (K). — Fig. Ic. E. apiculata Anderss., Stoekh. Vet. Akad. Handl. 1853: 2:^4, 1854. Type: supra lapides juxta litora insulae Chatham, Andersson s.n. (S — not seen). E. flahcUaris Anderss. ex Boiss. in DC, Prodr. 15(2): 17, 1862. Type: Insula Indefatigable, Andersson s.n. (S — not seen). E.nesiotica Robinson, Proc. Amer. Acad. 38: 167, 1902. Type: Seymour Island, South, Snodgrass & Heller 589 (GH). The overall appearance, habit and leaf characters are extremely variable in this species. Extremes are well-marked and are the basis for most of taxa here placed in synonymy. There are many intermediates, however, and since such characters as stipule and seed shape (which arc usually conservative) are rather constant it seems wiser to treat them as part of a single variable species. [Vol. 56 176 ANNALS OF THE MISSOURI BOTANICAL GARDEN ^ , / f'^ X ?>';«.n«:'^ J Fig. 1. Chamaesyce of the Galapagos. A, C. viminea (Hook, f.) Burch, Habit (X 1); B, C. amplcxicaulis (Hook, f.) Burch. Habit (X 1); C, C. rectirva (Hook, f.) Burch. Habit CX 1); D, C. ^alapagcia (Robinson & Grecnman) Burch. Habit (X 1). A after Wi^^^ius & Porter 340 (MO); B after Wiggins & Porter 308 (CAS); C after Snodgrass & Ilellcr 383 (GH); D after Stcivart 18H9 (DS). 5. Chamaesyce bindloensis (Stewart) Burch, comb, ct stat, no\' based on Euphorbia articulata Andcrss. var. hhuUoensis Stewart, Proc. Cal. Acad. Sci. 4th ser. 1: 91, 1911. Type: Bindloe Island, low shrubs in tufaceous soil in the vicinity of the shore, Stewart 1868 (GH).— Fig. 2c. 1969] BURCH^GALAPAGOS EUPHORBIEAE 177 6. Chamaesyce galapageia (Robinson & Grecnman) Burch, comb, nov., based on Euphorbia galapageia Robinson & Grecnman, Amer. Jour. Sci. 3rd scr. SO: 144, 1895. Type: Charles Island, June 1891, Baiir 261 (GH).— Fig. Id. Euphorbia stevensii Stewart, Proc. Cal. Acad. Sci. 4lh scr. 1: 92, 1911. Type: Albemarle Island, Iguana Cove, Stewart 1890 (GH, photo DS). Tliis and the preceding two species have elliptic leaves and are much more similar in overall appearance to other New World species than are most of the endemic species. In particular C galapageia suggests the group, including C. nutans (Lag.) Small and C. hyssopifolia (L.) Small, with cyathia in more or less dense clusters made up of dichotomising laterals. None of the species in this group show the extreme reduction from this situation that is found in C. galapageia and all three of the endemic species differ in seed, cyathia and pubescence characters. 7. Chamaesyce nummularia (Hook, f.) Burch, comb, nov., based on Euphorbia nummiilaria Hook, f., Trans. Linn. Soc, London 20: 183, 1847. Type: Chatham Island, Darwin s,n. (K). — Fig. 2b. A distinctive species with small, hirsute, suborbicular leaves, and seeds with face smooth and so plump as to obscure the angles. Robinson & Greenman described Euphorbia nummularia var. glabra based on QBaur 3S6, GH) (A ) distinctive, 2-3-lobed leaves with a deep sinus between the unequal lobes. It is much more similar to Chamaesyce numnudaria than to any other New World species, but I should prefer to see more collections from the island before making a final judgment on its status. 8. Chamaesyce abdila Burch, sp. nov. — Fig. 2d-g. Herha prostrata perennis ex caudice ut videtur; omnes partes minutae, stri- gosae sparsim. Folia elliptica, Integra; stipulae junctae vel distinctae, intcgra. Cyathia soHtaria; glandes stipitatae parum, atropurpurae, appendices obsoletae; flores masculini 3-7 per cyathium. Semen anguste ovoideum, 4-angulare, super- ficiebus plus minusve planis, laeve. Herh, somewhat woody at base, apparently perennial from the rootstock; stems numerous to 1 mm diam. branching freely from base, the nodes somewhat en- larged, prostrate, to ca 1 cm, sparsely strigose. Leaves membranous; stipules joined or distinct, entire, to ca 0.3 mm long; blade elliptic, 1.5-3.5 x 1-1.5 mm, the base oblique, the margin entire, the apex obtuse, sparsely strigose, particularly on lower surface. Cythia solitary; glands small, somewhat stipitate, deep purple, appendages obsolete; male flowers 3-7 per cyathium; styles joined at base, bifid and somewhat clavate for Vi length. Capsule broadly ovoid, strigose, to 1.2 mm long, 1.5 mm wide; seed narrowly ovoid, 4-angled, to 0.7 mm long, 0.5 mm wide, the faces flat, smooth. Type: Galapagos Islands: Santa Fe: grassy slopes among Opuntia and Bursera, plateau about 2 km from landing beach at NE of island, alt ca 40-50 m, Wig- i>ins & Porter 584 (holotvpe: MO; isotypes: CAS, USF). [Vol. 56 178 ANNALS OF THE MISSOURI BOTANICAL GARDKlVf J^ i^y-L^*-^*^ , ^ 97^^M,.*^ «A , C B M ihAjl/K- Fig. 2. Chamaesyce of the Galapagos. A, C. piinctuhna (Andcrss.) Burch. Hal)it (X 1); B, C. nummularia (Hook, f.) Burch. Habit (X 1); C, C. bhuUocnsis (Stewart) Burch. Habit (X 1); D-G, C. ahdita Burch. D, habit (X 1); E, leaf bases & stipules (X 2.5); F, cyathium & capsule (X 7); G, seed (X 12). A after VVig^ms & Porter 730 (MO); B after Suoir 231 (DS); C after Wifigins & Porter 300 (CAS); D-G after VVj<'«r«s & Porter 584 (MO). This tiny species is known only from the type collection, and has perhaps been overlooked by other collectors because of its si/e. It has some resemblance to Chamaesyce micromera ( ) Wooton & Standley, a North American species also reported from Peru, but differs in pubescence, style length, seed size and smooUiness of seed coat. The characters distinguishing it from C. nummularia are noted in the key. Only one EupJiorhia is kno\\n, E. equisetifurmis Stewart, and this from a single collection. From the rather sparse material available it is difficult to place it in relation to the rest of the genus. The articulate stems and whorled scales or leaves suggest lioissier's section Alectoroctoumn but it docs not appear to be close to other species of this group. The representation of the remainder of the tribe is extremely poor, and even the weedy species of Poinsettia are apparently absent. Their failure to invade the areas disturbed by man is as remarkable as is the lack of success in the colonisa- tion by Chamaesyce hirta. THE GENUS CIENFUEGOSIA CAV. (MALVACEAE) BY Paul A. Fryxell Research Geneticist, Crops Research Division, Agricultural Research Service, U.S. Department of Agriculture, Texas A & M University, College Station, Texas Abstract One novelty, Cienfucgosia rosei, is included in the 24 species recognized in this taxonomic revision of the genus. A new subgeneric classification is presented, with two subgenera and seven sections. Introduction and Historical Account Cieufitegosia is a member of the tribe Cossypieae of the Malvaceae. Some authors have placed these plants (tribe Hihisceac sens, lat.) in the Bombacaceae, but the evidence now suggests tliat they are better retained in the Malvaceae (Fryxell, 1968). The genera included in the Gossypieae, in addition to Cienfuegosia, are Cos- sypinm L., Thespesia Corr., Gossypioides Skov. ex Hutch., Hanipea Schlccht., Kokia Lewt., Cephalohibiscus Ulbr. and Lehronnecia Fosb. The principal unifying features that characterize the members of this tribe and which distinguish them from other genera (especially from the Hihisceac sens, str.) are (a) the con- duplicate embryos, and (b) the presence of pigment glands distributed to various parts of the plant, together with the presence of certain sesquiterpene-like pigments associated with these glands. These pigments (principally gossypol but also several other chemically related compounds) are apparently unique to tlie tribe Gossypleae (Lukefahr & Fryxell, 1967; Fryxell, 1968). Various botanists have dealt with species of Cienfiiegosia since Cavanilles established the genus in 1787. Garcke (1860) was the first of the few who have dealt comprehensively with the entire genus. Fully two-thirds of the species now known were undcscribed in Garcke's time, Hochreutiner (1902) dealt inclusively with the group, but he simply listed the species known at that time, and indicated synonymy and effected nomcncla- tural changes. He did not attempt to deal with the classification, distribution, or phylogeny of the group. He accepted a circumscription of the genus that was much broader than is now considered valid. Ulbrich (1914) also simply listed the species and accepted a similarly broad conception of the genus. Hutchinson (1947) provided a delimitation of the genus with which I am in closer agreement, although his treatment of the individual species is not entirely convincing. Giirke (in Martius, 1892) considered only the species that occurred in or near Brazil; Robinson (1895) treated only the North American species; Rodrigo (1941, 1948) dealt only with the Argentine species; Hutchinson and Dalziel Ann. Missouri Box. Card. 56(2): 179-250, 1969, [Vol. 56 180 ANNALS OF THE MISSOURI BOTANICAL CAKDEN (1928) and Excll and Wild (1961) included only the species from specilic regions of Africa. The present study intends to deal in a comprehensive manner with the descrip- tion and classification of the genus Ciciifuegoua. Cavanilles (1787) hrst described this genus and named it Cienfuegosia in honor of the 16th century Spanish botanist Bernardo Cienfuegos, author of "Historia de las plantas." Two years later, A. L. dc Jussieu (1789) shortened the generic name to Fiigosia, the name under which these plants were most commonly known during the ensuing se\enty years. During these sevent) years, three other : published. Linnaeus (posthumously, under 1 ) published the variant "Cieiifiigiosd." Anotl fiiegia, of W'illdenow (1800). Cieiifnoosia v ler published by DeCandolle (1824), an error that was perpetuated recently by (1947) / and not as a distinct concept, its orthography has been so greatly altered that combinations widi the one name cannot automaticalh be transferred to the other, but must be handled as distinct combinations. Gareke (1860) reviewed this history and noted the priority of Cavanilles' name. Consequently he transferred several species to CieJifnegusia that had been published under the invalid name Fiigo^ia. Ventenat (1800) published the name Redutea for a plant that must now be included in Cieiijnegosia. Ventenat chose this name in honor of the botanical artist, P. J. Hedoute, but intentionally altered the orthography in naming the plant. \'entenat's spelling was used by Persoon (1806), but almost all other subsequent authors have cited the name as Rcdoiitea Vent., -^vhich is the spelling that Kunth used in publishing lledoutea Vent., which is the spelling that Kunth used in publishing Rcdoittca tripartita ll.B.K. This error may have been perpe- tuated partly because Index Kewensis lists the name in this way, as does \\ illis (1966), who moreover attributes Rediitca to Persoon. (1862) ( Elias Durand) for a plant that must be held indistinguishable from that which Gray (1852) had previously described as a species of Fugosia. The name FJidii- randia tcxaiia Buckl. has never achieved eurrenc\ bexond the original publication ) because Gray (1863) (AlthoLi^li Sliinners ( ; correct in ) Relationships oi the Genus Recently Fryxell (1968) presented a key, distinguishing Cienfuegosia from other genera of the Cossypieae. There is sufficient variability among the species / / (for which ;; = 10 or 11) clearly from the other "enera (for which ;/ = 12 or 1969] FRYXELL — CIJ:M UEGOSIA 181 13); but counts arc not yet known for llic two monotypic genera Cephalohibiscus and Lehroiuiecia. Plant habit also serves to distinguish Cieufuegosia from other members of the tribe. Cieufuegosia varies from the relati\cly robust shrub, C. affiuis, to pro- cumbent herbs such as C. argentina. Arboreal forms never occur in Cieufuegosia. 13y contrast TJiespesia, Ha^tpea, CepJudohibisciis, Lehroiuiecia, and Kokia are typically arborescent (and occasionally shrubby); Gossypium and Gossypioides arc typically shrubby (or occasionally arborescent in the former genus). None of the genera of Gossypicae other than Cieufuegosia includes herbaceous plants. A few additional characters merit brief comment. Lew ton (1925) discussed the unsatisfactory nature of the number of cells in the capsule as a character for distinguishing genera. Cell number is constant for some genera (three in Hampea, Kokia, and Lehrouuccia; five in CephaloJiihiscus^, but varies from three to five among the species of Cieufuegosia, Cossypiu}u, Gossypioides, and Thespesia, It is often useful in delimiting lower taxa, hoAve\er. The presence of an involucel is characteristic of most plants of the Gossypieae, The number of parts of the involucel is of value in classifying these plants at both the generic and specific levels. The genera Gossypium, Gossypioides, Hampea, CcpJudoJiihiscus, J.ehrouuecia, and Kokia uniformly have involucels of three bract- lets. The involucral bractlets of Thespesia and Cieufuegosia arc more variable, (in Cieufuegosia sect. Friesia') to as many as 20 ( ;s.); but within these genera the number and d ispositio bractlets are of taxonomic significance. Attempts to use the form of the bracts as a diagnostic character have not succeeded (Lewton, 1925). The bracts are often caducous in lliespesia and in two species of Gossypium, but are persistent in the remainder of the tribe. Involucral nectaries are found in threes at the summit of the pedicel directly below the point of insertion of the involucral bracts. (An additional set of three may also occur, alternate with and above the bracts, in the cultivated species of Gossypium.^ These nectaries are characteristic of the tribe Gossypieae, although they do not occur in all of the species. They are absent in the Hihisceae sens, str., although species of lUhiscus sect. Furcaria have nectaries located directly on the calyx (cf. Janda, 1937). The presence or absence of involucral nectaries is useful in delimitimr lower taxa. The gametic chromosome number in Thespesia, Hauipea, and Gossypium is u 13, with allotctraploidy occurring in Gossypiu})i. In Gossypioides and Kokia the number is u- 12. The numbers for Jehrouuecia and Cephalohibiscus are not known. In Cieufuegosia numbers of ;? 10 and 11 have been reported (cf. Table 2). In addition to the morphological similarities (Fryxell, 1968) that imply phylogenetic relationships, other evidence further substantiates such relationships, Skovsted (1944) presented evidence from experimental studies with C, hil- dehraiidtii that suggests a distant relationship with Hibiscus. Hibiscus rubidus Skov. and H. hifurcatus Cav. set fruit with empty seeds when pollinated by C. hildebraudtii. (According to Men/el and W^ilson (1961) these two species are 1^2 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 correctly identified as H. acetosella W'elw. & Hiern. and H. radiatus Cav. respec- tively.) In addition, C. hildehrandtii grew vigorously and flowered when onto HdmcHs escuJeutus L. (= Ahelmuschiis csculentiis (L.) Moench). An affinity of Cicufucgosia with Gussypiuin is indicated by indirect evidence. One line of evidence concerns the ability of the cotton rust QVuccinia stakmanii Presl.) to infect C. argcntina to a fimited extent; this rust fails to infect other species outside of Gassy pinm (cf. Savile, 1954). The cotton rust is an oWigate hetcroecious rust that alternates between Gossypium spp. and nouteloiia spp. (Presley and King, 1943; Blank and Leathers, 1963). It occurs in northern Mexico and adjacent parts of Arizona, New Mexico, and Texas. Its primary grass hosts are Bouteloua aristidoides (H.B.K.) Griseb. and B. harhata Lag., both annuals, though not closel) allied. The aecial stage will develop on most species of Gossypium (Presley and King, 1943; Blank, personal communication), and can constitute an economic hazard in commercial plantings of cotton in years favorable to rust development. The rust, however, is not known to infect malvaccous plants other than Gossypiniii spp. and Gieufitegosia argentina. On C. argentina pycnial lesions will form, but these quickly regress and do not develop to the aecial stage. Attempts to infect other species of Cienfnegasia with Pnccinia stakmanii have given negative results with C. affinis, C. drummondii, C. suJfiirea, C. hetcrophylla, C. yncatancnsis, C. hildehrandtii, and C, hearnii (Blank, unpublished). It was therefore of interest to discover aecial rust lesions occurring naturally on Gienjiicgosia in Argentina. Two specimens in the Geneva herbarium QRodri- guez 12, of C. argentina; and Monetti 14711, of C. hispida^ each show several well-developed aecial lesions. Both were collected in Tucuman in 1913, which may have been a favorable year for the rust in that area. tacion Fxperimental Ayronecuaria, P. R. A request to Ing. M. Gutierrez (E tive response. ) for Held observations of rust on Cienfnego f' bearing aeeial lesions. Schulz observed that 10% of the plants in a natural pop- ulation in eastern Chaco were infected. Plants of C. drummondii and C. sidfurea growing in the same area were not infected. One can conclude, therefore, that a rust endemic to northern Argentina occurs naturally on species of Gicnfuegosia sect. Friesia, and is probably co species. Moreover, the rust is probably a taxon distinct from Pnccinia stakmanii. (b) (a) crop (Gutierrez, personal comiiumication), even though both Bouteloua aristi- doides and B. harhata occur in Argentina. Observations of uredial or telial rust stages on these grasses arc at present lacking. Another line of evidence is given by the host relations of certain insect species that have become pests of cotton and hence of economic importance, particularly those whose host range is relati\ely narrow. The evidence given by Szumkowski (1952, 1953), W'hitcomb & Britton (1953), and Lukefahr & Martin (1962) jfinis witb Gossypium, in that Alahama argillacea ( 1969] niYXELL — CIENFUEGOSIA 183 ) ff The red bolhvorni of cotton, Diparopsis castanea, also occurs on C. hilde- hraudtii which is probably its original host (iMarshall et al., 1937; Pearson, 1954). Dipawpsis watersi and D. tephragranuua occur on C. digitata Cav. and Gossypium (Pearson, 1954) (Emsley, 195 7) ffiiiis (see p. 217) translated roughly as "wild cotton," and of the Arabian Cienfiiegosia ( ) Ecology / indicates an adaptation to relati\ely dr\ or otherwise unfavorable habitats. A few species occur in the severest of xeric environments, notably C. tripartita, C. chia- rugii, C. hearnii, C. rosci, and C. wehhii. A few of the more shrubby species are grassland plants; C. heteroclada from Nigeria and Ghana is especially adapted to its dry grassland environment because it flowers and fruits directly from the perennial root within a relati\ely short season (Fmsley, 1957), and C. integrifoUa from Paraguay may be similarly adapted. Many of the South American species occur in chaparral association on sandy soils. Others favor heavy soils. The species of sect, briesia often occur in saline environments. Cienfuegoda yucatanensis pro- vides an interesting example in that in Yucatan it occurs on arid, stony soils, while in Florida, Cuba, and Bahama it is found in more mesic environments. Distribution Subgenus Articuhta is African (Fig. 1) and subg. Cienfiiegosia is American (Figs. 2-5), with the exception of C. digitata, which occurs in Africa. Subgenus Articidata is herein divided into three sections: Articulata; Dioica; and Carckea. Section Articidata has a single species, C. gerrardii, with a relatively narrow distribution in Natal and the Transvaal. Section Dioica also has a single species, C. heteroclada, with a restricted distribution from northern Nigeria to northern Ghana. Section Carckea has five species, four of which occur around the borders of the Gulf of Aden, the fifth occurring primarily in southeast Africa. None of these species, with the possible exception of C. hildehrandtii, is particu- larly widespread or abundant. The somewhat greater range of C. hildehrandtii may result from its probably greater potential for seed dispersal. The distributional pattern of these species does not suggest a precise localization of a "center of origin" for the subgenus. Subgenus Cieujitego: f Paraguay ana; and Vriesia. Section Cieufuegosia (Fig. 2) includes six widely scat- tered species (cf. Fryxell, 1967b), one of which has a disjunct distribution in southern and in northwestern Africa. The other five have discrete and restricted distributions in the following places: Paraguay, Ecuador-Peru, Venezuela, Mexico, and around the Gulf of Mexico (parts of Yucatan, Florida, Bahama, and Cuba). 184 [Vol. 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 1. Distribution of Cienfucgosia subg. Articulata. iree species, two of wliich nrc primarily ^'■ne/uela and Paraguay; the third is \e Section Rohusta (Fig. 3) inchides t) Brazilian in distribution, extending to Mexican. Section Paraguayana (Fig. 4) includes four species that occur in Para- guay and northern Argentina, although one of them, C. drummondii, also occurs in southern Texas. Section Vriesia (Fig. 5) includes four species found primarily in northern Argentina and Paraguay. 1969] FRYXELL CIENFUEGOSIA 185 atan^niis C. hetaropliylla *t C. subrftrnata C. intarmvdio C. digjtafa 2 3 Fig. 2. Distribution of Cienfucgosia sect. Cicnfucgosia, Fig. 3. Distribution of Cienfue- gosia sect. Robusta. Fig. 4. Distribution of Cicnfucgosia sect. Varaguayana, Fig. 5. Distrib- ution of Cicnfucgosia sect. Fricsia. The distribution patterns point to Paraguay as the center of dispersal of the group. Nine species occur within Paraguay, primarily in the southeastern half of the country; three of these are endemic to Paraguay. No other area has com- parable diversity. The general pattern of disjunction previously discussed for C. drummondii has been noted for many elements of the American flora; C. druuimojuUi was cited as an example in this connection by Bray (1900). This entire question was reviewed by Raven (1963), w^ho concluded that such disjunction of xero- phytes probobaly occurred by dispersal from South America to North America in relatively recent time. The distribution of C. drummondii accords with this inter- pretation because: (a) the center of dispersal (or "source") of this group is South American, so that it is more reasonable to suppose that propagulcs were dissemi- nated from south to north rather than in the opposite direction; and (b) the plants of this species from the two areas are scarcely distinguishable, a factor that suggests [Vol. 56 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN that the disjunction is of relatively recent origin. The only apparent distinction is that the petal spot may be present or absent in the South American specimens but is uniformly absent in the Texas specimens, a variabiUty pattern which further supports the idea of a southern origin. The distribution of C. digitata provides a second example of disjunction in Cienfuegosia, The species occurs from southern Angola to the Transvaal in South Africa and from Mauritania to northern Nigeria in northwestern Africa, flutch- inson (1947) stated that the area of the greatest variability of C. digitata is in the southern part of its range, and that this is therefore its primary distribution; the more northerly distribution is the result of secondary colonization. I find no such difference in variabiUty. On the basis of the distributional data, implying an American origin and a transatlantic crossing, I prefer the opposite conclusion: that the West African distribution is primary and the South African distribution secondary. The distributional disjunction of C. digitata is somewhat parallel with the disjunct distribution of Gossypiunt anomalnm in roughly the same areas of Africa, as noted by Pearson (1954) and discussed by Fryxell (1967b). Perhaps those more famihar with the flora of Africa might add additional examples and proffer a phytogeographic interpretation. The lack of morphological divergence between the disjunct populations, both in C. digitata and G. anomalnm, suggests that the disjunction is of relatively recent origin. The parallel with G. anomalnm is noteworthy in another way. The other taxa closely allied to C. digitata occur in America. Thus, the distribution of the entire group spans the Atlantic. Similarly, taxa closely allied to G. anomalnm have recently been described from the Cape Verde Islands; this indicates a dis- tributional extension into (if not across) the Atlantic. These distributional data are particularly significant because this species group or its near relatives have been suggested as possible ancestors of the amphidiploid species of Gossypinm (see Phillips, 1963). The strong distributional parallel that exists between the taxa allied to C. digitata and those that are allied to G. anomalnm may suggest additional clues to the phylogenetic history of Gossypinm. The most primitive (as well as the least known) of the six species in sect. Cienfuegosia appears to be C. snbternata, which is endemic to Paraguay. The Fcuadorian-Peruvian C. tripartita is apparently derived directly from it. An expan- sion or migration north w^ard along the Pacific coast of South America is indicated. The Mexican C. rosei may be a speciali/ed derivative of this line. The African C. digitata and C. heterophylla from Colombia, Venezuela, and Brazil, appear to constitute a separate phylogenetic line; this suggests a migration northward along the Atlantic side of South America, and, indeed, across the Atlantic. The insular C. yncatanensis is a specialization of C. heterophylla. The overall picture indicates an expansion from a center in or near Paraguay northward as far as Yucatan and southern Florida and eastw^ird across the Atlantic. The pattern of distribution of C. yncatanensis suggests that it wvas achieved via salt-water dissemination to coastal and insular sites. Perhaps C. digitata was disseminated across the Atlantic by the same means. The transatlantic migration presumably occurred earlier than the African migration that gave rise to the present-day disjunction in the distri- bution of this taxon. 1969] FRYXELL — CIENFUEGOSIA 187 Reproductive Biology Cienf f gosia but is known elsewhere in the Gossypieae in Hampea, and elsewhere in the Malvaceae in Kydia and Napaea. In subg. Articulata self-pollination occurs easily and presumably regularly (except perhaps in C. gerrardW) because the decurrent stigma is in close proximity to the androecium. The showy corolla is clearly an adaptation for attracting insects. That cross-pollination also occurs regularly, there- fore, seems likely. In subg. Cienfuegosia the basic (primitive) pattern show^s a capitate, greatly exserted stigma; thus self-pollination is probably infrequent. This basic pattern has evolved in two directions, both of them involving a reduction in relative style length. The first departure from the basic pattern is found in some species of sect / subg. Articulata and in Gossypium. The clavate style with decurrent stigmatic lobes that only partially exceed the androecium (expressed in most extreme form in C. heter- ophylla^ is clearly an adaptation for increased self-pollination. An overall decrease in flower size accompanies this adaptation. The second departure from the basic pattern occurs principally in C, siilfurea and C. argentina, of sect. Varagiiayana and sect. Vriesia, respectively. In these species, the upper portions of the style branches are free and partially recurved, so that the stigmatic lobes are brought nearer to the androecium, thus enhancing the possibihty of self-pollination. In these same species the petal spot is less prominent, and facultative cleistogamy occurs. fnegosia ( ) base of each petal in all taxa, except that the spot is small in C. arge^itina, vestigial in C. siilfurea, absent in C. yucatauensis, and sometimes absent in C. drummondii. Flowering is diurnal in Cienfuegosia. The flower generally opens in early morning and withers later in the same day. In some species the flowers close by noon, in others not until late in the afternoon. The corolla, with the androecium attached, falls the next day. Three species, C. somaliana, C. uelshii and C. hearnii (and possibly also C. chiarugii^ are exceptional in that their flowers do not open until midday, then close later in the afternoon. Facultative cleistogamy occurs in C. drummondii, C. sidfurea, and C. argoitina, but has not been observed in the other species that are in culture. I tentatively suggest that the phenomenon is confined to the sections Friesia and Paraguayaua. Cleistogamic flowers (which readily produce fruits and seeds) apparently develop in response to environmental factors such as shading. Such flowers are greatly reduced in size (and sometimes number) of parts and have been illustrated by (1963) (Fryxell, 1963 and unpublished; F. D. Wilson, ) The flowers of Cienfuegosia are generally campanulate, narrow^ly so in C. hearnii. However, in C. yucatanensis the petals flare widely to a fully rotate form. [Vol. 56 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN Subgenus Cienfuegosia ^^ \ I / Phylogenetic diagram of the genus Cwnfv^gosla Cav. (time axis perpendicular.) Subgenus Arfrcufafa I Fig. 6. Phylogenetic diagram of Cienfucgosia. Phylogeny The postulated phylogeny of tlie genus is illustrated diagrammatieally in Fig. 6. Each of the two subgenera of Cicufnegosia is considered to be mono- phyletic, but whether the tw^o groups had a common origin is not clear. The premises upon which the discussion of specific phylogenetic hncs is based appear in Tabid. Table 1. Primitive and advanced traits in Cicufnegosia Cav. Trait Primitive Advanced Growth ha])it Pollination adaptations Petal spot Stigma Flowering branches Capsule Pigment glands Number of seeds per locule Large and shrubby Outbreeding Present Exserted from androecium Complex Pubescent Extended to all parts Many Small and herbaceous Inbreeding Absent Included in androecium Reduced Glabrous Restricted to certain parts Few 1969] FRYXELL — CIENFUEGOSIA 189 111 subg. Articulata, C. gerrardii from South Africa is the most primitive rep- resentative, as indicated by: its large size and shrubby growth habit; its many- jointed fruiting branches; its more widely distributed pigment glands; and its flower structure that favors outbreeding. Hutchinson (1959) drew attention to the phylogenetic position of this species. (Although it has many primitive traits, C. gerrardii exhibits an advanced character in its reduced seed number.) The more complex fruiting branches of C. gerrardii have become reduced in two distinct evolutionary lines, while remaining articulated and sometimc^s several- Howered. One line is represented by the highly specialized C. heteroclmla, occur- ing in northern Nigeria and Ghana, which dcxclops many-flowered racemes. The other line includes the shrubby C. hildehrandtii from eastern and southeastern Africa and the subshrubs C. chiarugii, C. somaliana, C. ivelshti, and C. hearnii from Somalia, Ethiopia, and Aden. In most of these species the flowers arc borne singly on articulated peduncles in the axils of the leaves (sometimes more than one in C. ivehhii and C. hildehrajidtiiX This pattern of two evolutionary lines suggests an expansion from south to north. The amount of divergence among the species of this subgenus, including the development of dioecy in one species, in- dicates that the group is ancient. In subg, Cienfttegosia, sect. Robiista is the most primitive group and C. affinis is the most primitive species. It is a large, woody shrub with peduncles that are sometime articulated and even (rarely) several-flowered. The flowers are large and clearly adapted to outbreeding, with long styles (nearly equaling the petals) and capitate stigmas. Moreover, C. affinis shows the greatest similarity to species of sect. Articiilata, notably in peduncle articulation and in the more widespread dis- tribution of pigment glands, especially in the corolla. Cienfiiegosia gerrardii and C. affinis, the two species interpreted as the most primitive of their respective subgenera, both have hairy fruits. Cienfiiegosia ciiyahensis and C. intermedia are intermediate between C. affinis and species of sect. Cienfiiegosia and thus represent transitional forms to or offshoots of this evolutionary line. Cienfiiegosia suhternata is the next step in this series as the most primitive of the six taxa included in sect. Cienfiiegosia. Th distributional and phylogenetic relationships among the six species in this section (noted on p. 183), involve one development that leads to C. tripartita and C. rosei and another to C. heterophylla, C. yiicatanensis, and C. digitata. Hutchinson (1947) referred to a cline in leaf shape in these species but failed to account for the geographical relations among these plants. The flower form of C affinis (including such features as the long style, the large corolla, and the prominent petal spot with its characteristic yellow radii) can be traced to at least one element in each of the other sections of subg. Cien- fiiegosia. These examples are: (1) C suhternata of sect. Cienfiiegosia, as already noted; (2) C. idmifolia of sect. Friesia; and (3) C. subprostrata of sect. Para- guayana. The last named species also shares wath C. affinis the sericeous vcstiture of the fruit that is not found elsewhere in the subgenus. Because of this relation- ship, these species are considered to be the most primitive representatives of their respective sections. Phylogenetic relationships of sections Friesia and Paraguayana, hoW'Cver, cannot be clearly inferred. [Vol. 56 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN The four species of sect. Friesia are closely allied morphologically and do not exhibit any noteworthy phylogenetic series. Cioifuegosia ulmifolia is probably the most primitive representative; C. hasslerana is closely related to it; C. hispida and C. argentina are apparently more highly speciah/ed. The four species of sect. Paraguayana are more divergent among themselves than those of sect. Friesia; C, suhprostrata and C. integrifolia are closely allied and relatively more primitive, while C. driimmondii and C. sidfiirea are more specialized. In both of these sections the more primitive species are distributed in or near the proposed center of origin of the subgenus, Paraguay, while the derived species have distributions that extend beyond this center to the south and west. As in subg. Articidata, the great divergence among the species of subg. Cien- fiiegosia indicates that the group is ancient. Chromosome Counts Previously published information on the chromosome numbers of Cieiifuegosia is meager, Youngman (1931, p. 54) published a somatic (?) chromosome count of 26 for C. hildehraudtii, which is identical to the numbers reported for Cos- sypium, Thespesia, and Hampea. However, Skovsted (1935, p. 287) later pub- lished a count of 2;/ = 22 for the same species, and Palacios and Tiranti (1966) and F. D. Wilson (unpublished) confirmed tlie latter number. Longley (1933, p. 224) reported 2n ~ 20 for C. heterophylla. He described the plant as ''C. heterophylla (Vent.) Garcke, a species native to the southern part of the United States, . . ." Presumably, his material came from the Florida Keys and in fact represents C. yucataneusis Millsp. Palacios and Tiranti (1966) published values of n = 10 for three Argentine species, C. sidfnrea, C. driimmondii, and C. ulmifolia. Living material is available for study of twelve taxa, and additional counts have been made as given in Table 2. Table 2. Chromosome numbers in Cienfiicgosia. Gametic chromosome Species n u m I yer Authority C. hililchrandtii Garcke C. wchhii (T. Anders.) Garcke C. hearnii Fryx. C somaliana Fryx. C. di^itata Cav. C. heterophylla (Vent.) Garcke C. yucatanensis Millsp. C. tripartita (H.B.K,) Garcke C. rosei Fryx. C.affinis (H.B.K.) Hochr. C. sulfurea (Jiiss. in St.-Hil.) Garcke C. drummondii (A. Gray) Lewt. C. argentina Gtirke C. tdmifolia Fryx. 11 11 II 11 10 10 10 10 10 10 10 10 10 10 Skovsted (1935); Palacios & Tiranti (1966) Wilson (Unpubl.) Wilson (Unpubl.) Wilson (Unpubl.) Wilson (1968) Wilson (1968) Longley (1933) Wilson (1968) Wilson (Unpubl.) Wilson (Unpubl.) Palacios & Tiranti (1966) Palacios & Tiranti (1966) Wilson (Unpubl.) Palacios & Tiranti (1966) ^ The unpublished data of F. D. Wilson cited in this table were graciously provided by Dr. Wilson from a manuscript in preparation dealing with the karyology of Cienfuegosia, 1969] FRYXELL — CIENFUEGOSIA 191 Hybridization Twelve of the twenty-four taxa included in Cienfuegosia are currently avail- able in the living state for experimental study. Many interspecific crosses have been attempted in this group in order to assess genetic relationships. The following crosses have been successful: C. welshii X C hearnii C. digitata X C. heterophyUa C. yucatanensis X C. heterophyUa C. yucatanensis X C. digitata C, yucatanensis X C. tripartita C. heterophyUa X C. tripartita C. argentina X C. affinis C, sulfurea X C. drumviondii (lethal as seedlings^ — Palacios and Tiranti, 1966) Additional crosses are being made. The results of these experimental studies will be reported elsewhere. Systematic Treatment Cienfuegosia Cav., Diss. 3: 174., t. 72., f, 2, 1787. Fugosia Juss., Gen. PL 274, 1789, pro syn. Cicnf ligiosa Gis., Prael. (Linn.) 471, 1792. Cicnfuegia Willd. in L., Sp. PL cd. 4. 3: 723, 1800. Rcdiitea Vent., Hort. Ccls t. U, 1800. Rcdoutea H.B.K., Nov. Gen. Sp. PL 5: 293, 1822. Cienfiigosia DC., Prodr. 1: 457, 1824. Elidurandia BuckL, Proc. Acad. Sci. Philadelphia 1861: 450, 1862. Perennial herbs or subshrubs (sometimes shrubs) with herbaeeous to woody stems arising from a woody rootstoek. Stems procumbent (but never repent), lax and ascending, or upright; glabrous to pubescent, commonly angled, usually punctate. Leaves entire or coarsely serrate to variously divided, sometimes deeply and secondarily so; linear, elliptic, cuneiform, reniform, or divided digitately; pubescent to glabrous; sometimes punctate; foliar nectaries present (subg. Articu- lata^ or absent (subg. Cienfuegosia'). Phyllotaxy variable, including 2-, 3- and 5-ranked arrangements. Petioles often canaliculate, glabrous to densely pubescent, often with pubescence confined to distal end. Stipules large and foliaceous (some- times clasping) to minute and subulate; glabrous to pubescent; caducous or persistent. Peduncles axillary; usually unifloratc although sometimes multiflorate, with or without articulation, trigonal, incrassate above, surmounted by three involucral nectaries or these lacking. Involucel sometimes wanting, but usually present, persistent, and 9'12-foliolatc (except reduced to 3-foliolate in C.heter- oclada and partially so reduced in C. somaUana'); bractlets minute, subulate (and in 3 groups in subg. Articulata') or prominent and linear to spatulate, equaling the calyx. Calyx gamosepaluus, 5-lobed, usually punctate, glabrous to densely pubescent, sometimes costulate; lobes long-acuminate to rotund-apiculate. Petals 5, distinct, sometimes punctate, stellate-pubescent without, usually showy, yellow (sometimes nearly white), pink, or purple, with dark basal spot (sometimes lack- [Vol. 56 192 ANNALS OF THE MLSSOLRI l^OTANICAL GARDEN ing); flowers withering in a day, sometimes eleistogamie. Anther mass globose, oblate, elliptie, or obo\ate; staminal column yellowish or dark red, sometimes punctate; stamens sometimes S-ranked; anthers unilocular, dark red, orange, or yellow; pollen globose, eehinate. Style single, sometimes divided apically, usually elongate so that the stigma exceeds the androecium, sometimes punctate; stigmas 3-5-lobed (or if style divided, stigmas distinct), usually exceeding androecium, decurrent or capitate, dark red or yellowish, sometimes pubescent. Fruit a capsule, usually Sdoculed (sometimes 4-5-loculed), o\oid, globose, sometimes obovoid, or notably elongated (in C. somaliana and C. welshiO, glabrous or weakly to densely ascending-sericeous, usually punctate, sometimes with hairs on inner margin of suture. Seeds 3-8 mm, turbinate, minutely puberulent (in C. afjinis^ to densely comose; hairs tightly appressed (C. driimuiondii), loosely appressed, or patent, hght tan to rufous-red, up to 10 mm long. Embryos conduplicatc. Cotyledons cuncifonn to sub-reniform, obtuse to emarginate, sometimes punctate. The indumentum is variable; some species are densely pubescent while others are essentially glabrous. Pubescence is generally greater on the under surface of the leaves. Plant hairs (except for those on the seed coat) are usually stellate and vary in size from minute puberulence to hairs 1-2 mm in length. Pubescence is characteristically red-brown in C, gerrardii. Where longer hairs occur, they are often reduced from stellate clusters to single, semi scabrous structures. Pigment glands are found in all above-ground parts of the plant in a few^ species and are variably distributed ill others. The varying distributional patterns have taxonomic significance, especially in the extension of these glands to the floral parts. The glands are always present on the calyx (except sometimes in C. hearui'O, although they are often very obscure in species of sect. Friesia. Glands of the calyx are especially prominent in sect. Cienfiiegosia. Hutchinson (1947) described these calyx glands as "conlined to a double row along each nerve" and used this as a diagnostic trait to distinguish Cieufitcgosia from its allied genera, in which he described the calyx glands as "scattered, sometimes absent." Although the calyx glands in Cieufuegosia are often oriented in the regular fashion de- scribed by Hutchinson (e.g., C. tripartita, C. heterochnhO, they are also quite irregularly placed in certain species (e.g., C. drnmmondii, C. hildehrandtii)\ there- fore, it is necessary to reject this trait as diagnostic for the genus. The comparative morphology of the embryos of Cieufuegosia spp. (Fryxell, 1968) shows them to bear a marked resemblance to those of Cossypinm^ except that most of the species of Cieufuegosia have embryos lacking the dark pigment glands, while only a few of the species of CossypiKm arc lacking them. Type species: Cieufuegosia ligitata Cav. Artificial Key to the Speciis of Cienfuegosia, Based on Vegetative Characters a. Lea\t^s deeply dissected or compound. b. Plants procumbent herbs. c. Leaf segments rounded to obovate, crenate 21. C. argcniina cc. Leaf segments linear, dentate 24. C. hassk ran a bb. Plants ascending herbs or subshrubs. d. Foliage odoriferous 4, C. somahana dd. Foliage non-odoriferous. 1969] FRYXELL — CIENFUEGOSIA 193 e. Leaves trifoliolate 12. C. subternata cc. Leaves not compound. f. Leaves large, deeply 3-partrd (at times trifoliolate) . . 13. C. tripartita ff. Leaves moderate, digitately divided. g. Leaf lobes acute 8. C. digitata gg. Leaf lobes obtuse 16. C. intertnedia aa. Leaves simple or moderately lobed. h. Leaves simple, entire. i. Plant hirsute 14. C. affinis ii. Plant glabrate. j. Foliar nectaries present, apical 7, C hcicroclada jj. Foliar nectaries lacking. k. Stipules often equaling petioles 19. C. intcgrifolia kk. Stipules inconspicuous. L Leaves orbicular 20. C. subprostrata 11. Leaves otherwise. m. Leaves 1-2 times as long as broad. n. Leaves cuneiform to obovate, obtuse 5. C. chiarugii nn. Leaves ovate, apiculate 10. C. rosei mm. Leaves much longer than broad. o. Leaves 4-lO times as long as broad 9. C. yucatanensis oo. Leaves 2-4 times as long as broad. p. Leaves elliptic, pcnninerved 15, C. cuyahensis pp. Leaves ovate, palminerved 1 1. C. hcterophylla hh. Leaves more or less lobed or, if simple, then dentate. q. Stipules foliaceous. r. Stipules symmetrical; leaves cordate 1. C. gcrrardii rr. Stipules auriculate-clasping; leaves truncate. s. Plant procumbent herb 23. C. idmifoUa ss. Plant upright subshrub 2. C. hildcbrandtii qq. Stipules inconspicuous. t. Foliage odoriferous. u. Leaves ciliate-margined 4. C. somaliana uu. Leaves glabrous 6. C. hearnii tt. Foliage non-odoriferous. V. Plants hirsute. w. Stipules subulate, symmetrical 17. C. sulfurea WW, Stipules auriculatc, asymmetrical 22. C. hispida vv. Plants glabrate or puberulent. X. Plants procumbent 2 1 . C. argentina XX. Plants ascending. y. Foliar nectaries present. z. Petioles shorter than lamina 7. C. heteroclada zz. Petioles longer than lamina 3. C. wehhii yy. Foliar nectaries lacking. A. Leaves simple, dentate 18. C. drummondii AA. Leaves trilobed; lobes entire. B. Leaf lobes obovate 16. C. intermedia BB. Leaf lobes lanceolate to ovate. C. Leaves large, deeply & variably lobed. . 13. C. tripartita CC. Leaves moderately sized and lobed. D. Leaves mostly acute to obtuse. . 11. C. hcterophylla DD. Leaves* apiculate 10. C. rosei No previous subgencric classifications have been proposed in Cienfuegosia^ although Garcke (1860) separated an American and an African group, and Hutchinson (1947) suggested certain species alHances. Neither author, however, gave taxonomic status to these groups or provided them with a nomcnclatural basis. The present treatment divides the genus into two subgenera and seven sections. 194 [Vol. 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN Key to the Suiu;enera 194) a. Peduncles articulate (sometimes obscurely so at base); flowers solitary or on several-tlowered sympodial branches; petals punctate; stigmas decurrent; foliar nectaries usually present; bractlets of involucel in three groups I. ARTICVLATA (p. aa. Peduncles non-articulate; flowers solitary in the leaf axils; petals epunctate (or if peduncles articulate or petals punctate, then stigma capitate); stigmas capitate or decurrent; foliar nectaries lacking; bractlets of involucel not grouped, sometimes '^^^i"8 II. CIENFVEGOSIA (p. 204) I. Surg. Ariiculaia Fryxcll, subgcn. nov. Folia punctata, plcTumcjuc cum ticctariis infra. Pcdunculi articulati. Involu- cmin breve, sacpissime 9-ph)llQm; foliolata iinolucrorum tribus in turmis dis- posita. Petala punctata. Stigmata decurrentia. Leaves of variable form, punctate, with 0-3 dorsal foliar nectaries. Flowers solitary or on several-flowered sympodial branches. Pedtmcles articulate, bracteate at articulation, surmounted by 3 prominent involucral nectaries. Involucel of 9 bractlets arranged in 3 groups (or these reduced to 3), each group surmounting an involucral nectary (or nectaries absent in sect. D/o/a/); bractlets inconspicuous, triangular or subulate, persistent. Calyx punctate (punctae sometimes restricted in distribution, or lacking); lobes acute or rotund-apiculate. Petal punctate, vellow, purple, or cream-colored with dark spot on claw. Stigmas decurrent, 3-4-lobed. Fruits glabrous or minutely pilose, usually punctate. Type species: Cicufnegosia gcrrardii (llarv. ex Harv. & Sond.) Hochr. The name of this subgenus is chosen in reference to the articulated peduncles that characterize it. Key to Sections of Subgenus Articulata a. Shrubs; leaves markedly cordate, palmately trilobed, lol^es rounded, entire; stipules large, foliose, oblong, sessile; pedicels subtended by foliar leaf. . . 1. Articulata (p. 194) (contains single species: \. C. gcrrardii') aa. Subslirubs or perennial herbs; leaves cuneate, entire or dentate, sometimes shal- lowly lobed or deeply dissected; stipules triangular, subulate or auriculate; pedicels subtended by small bracts. b. Leaves cuneiform to flabelliform or reniform, entire, dentate, or dissected; pedimcles in axils of leaves, l-2-flo\vcred; stipides triangular, auriculate or subulate; bractlets of involucel 9 2. Garckca (p. 196) bb. Leaves obovate, entire (sometimes apically tridentate); flowers dimorphic, in short racemes appearing before leaves; stipules subulate; bractlets of involucel 3, irregular 3. Djoica (p. 203) (contains single species: 7. C. hetcrochula') 1. Sect. Articulata Cienfiu'gosia subg. Articulata Fryx. sect. Articulata Leaves profoundly cordate. Stipules prominent, obtuse, foliaceous. Flowers borne on sympodial fruiting branches. Peduncle articulated, bearing foliar leaf at articulation. Type species: The type of sect. Articulata is the type of subg. Articulata, C. gcr- rardii. 1. Cienfuegosia gcrrardii (Harv. ex Harv. & Sond.) Hochr., Ann. Cons. & Jard. Bot., Geneve 6: 56, 1902. 1969] FRYXELL — CIENFUEGOSIA 195 Fiigosia Gcrrardi Harv. ex Ilarv. & Sond., Fl. Cap. 2: 588, 1862 [Type: Ladysmith, Natal: Gerrard 632 (TCD-n.v.)]- ThespesiaRehmannii, Szyszl., Polypet. Thai. Rehmann. 44, 1888 (=^Ro7pr. Spraw. Posiedzen 17: 136, 1888) [Type: Ladysmith, Tichman 7128 (n.v.)]. Illustrations: Fig. 17, d-c; also Wood (1911) t. 563; Janda (1937) Figs. 18, 28, 34; Fryxell (1968) Fig. 3,a. Branched open shrub; stems upright or lax, weakly ridged to terete, punctate, moderately covered with a minute stellate tomcntum having a characteristic red- brown color. Leaves cordate, acute, sub-entire to moderately 3-5-lobed, sparingly pubescent to glabrate, ciliate-mnrgined, 2-3 cm long, usually broader; lobes rotund; foliar nectary single, basal, elongate (1-2 mm). Petioles pubescent, terete, ob- scurely punctate, about V2 lengtli of lamina. Stipules fohaceous, entire, persistent, elliptic to oblong, obtuse, 1 -nerved, cuncatc-scssilc, pubescent to glabrate, punctate, 1-2 cm long (equaling or exceeding petioles), 3-8 mm broad, symmetrical. Fruit- ing branches sympodial, several-flowered, articulate and foliate at base of each pedicel. Pedicels Vz-lVz cm, puberulent to glabrate, surmounted by 3 large bordered nectaries. Involucel of 9 bractlcts in 3 groups; bractlets irregular, subulate (1-3 mm) or occasionally larger (up to 8 nun) and then often spatulate and somewhat foliose, punctate, l-ner\ed, glabrate. Calyx 11 Vz cm, costulate, puberulent to glabrate; lobes 3- (rarely 5-) ribbed, triangular, acute (sometimes rounded, sometimes acuminate, rarely 1 -toothed), 4-7 mm long. Petals yellow, 2-3 !/2 cm with small dark-red spot on claw (spot sometimes lacking). Anther mass globose to obovoid; filaments 2-4 mm, reddish [in sicco]; androecial column and occasionally the filaments sparingly punctate; anthers red; pollen light-colored. Style markedly punctate, exceeding androecium; stiguia 3-lobed. Fruit 3-loculed, globose to obovoid, 1-11/2 cm, rostrate, short-pilose without, glabrous within. Seeds one per loculc, 8 mm long, 5 mm wide, oval in outline, soft-pubescent; seed hairs dense, brown, V3 mm. Embryo punctate. Distribution (Fig. 7): Transvaal: in the region of Lydenburg and Middle- burg; Swaziland; Natal: nearly to the coast, and as far south as the Umzimkulu River; Mozambique: west of Laurenyo Marcjues (Excell and Wild, 1961). Marshall, Parson, and Hutchinson (1937) note that the ecological distribu- tion of this species in South Africa and Swaziland is confined to the middle veld district, which is defined roughly as the area included within 2000 to 4000 feet elevation. The plant occurs ''in colonies growing in scanty soil in the hilly ground above 1400 ft." South Africa: natal: Zululand: Gerrard 257 (BM); Magul, Ngotsche, Parsons s.n., Dec 1925 (PRE); Port Natal, Gerrard & MeKen, s.n. (S- this specimen is incorrectly de- noted as the type); Camperdown, 2000 ft, Wooj^239 (LE), 11722 (PRE); Haygarth s.n.. 1883 (BM). Griqualand East: In cHvis pr. TTydcsdale" ad flum. Umzimkulu, 2500 ft, Tyson 1408 (G, Z). Transvaal: Lydcnhurg, Wilms 83 (BM, G, L, M, P, PII), 4800 ft, Schlechter 3950 (G, M, UC, Z); outside village on Farm Rovidraai, open scrub in riverbank 4800 ft, Liebcnhcrg 3492 (PRE); procumbent on riverbank, 4700 ft, Galpin 12193 (PRE), Rogers 14561 (Z); Kruger National Park, Pretoriashap, 2000 ft, v. d. Schijff 2076 83b (G, L), Kassner 121 (BR); bci Brankhorst-Spruit, Wilms 83a (BM, Z). Wilm Swaziland: Bremersdorp, Hutchinson 5 (PRE). [Vol. 56 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2. Sect. Garckea FryxcII, sect. nov. Folia cuneiformia vcl rcniformia, simplicia vcl dissccta. Stipulac magnae ctiam auriculatae-acuminatac, aut parvac et subulatae. Pcdunculi plcrumque axillarcs, articulati, squamis parvulis ad articulos. Subshrubs. Stems glabrous to pubescent, weakly angular, punctate, often ob- scurely so. Leaves cuneiform to reniform, entire (except sometimes dissected in C. somaliana^ or sometimes 3-lobed, dentate- or serrate-margined on upper por- tion, punctate (sometimes translucently so), glabrous to pubescent. Foliar nectaries 0-3. Petioles glabrous, puberulent, or scabrous, punctate, usually canaliculate. Stipules large and clasping or inconspicuous and subulate. Flowers 1 or 2 (rarely more) on axillary peduncles or short sympodial branches. Peduncles articulate, with much-reduced bract at articulation, sumiounted by 3 involucral nectaries. Involucel of 9 bractlets in 3 groups, each group usually surmounting a nectary; bractlets subulate, sometimes laterally connate. Calyx costulate (except in C. chi- ariigii'), splitting and pseudo-bilabiate in flower and fruit; lobes rotund-apiculate, 3-5-ribbed. Petals yellow, purple, or cream-colored, with dark claw. Androecium purple or orange; pollen cream-colored or orange. Fruit prominently or obscurely punctate, glabrous to minutely pilose, ovoid to slightly obovoid or greatly elongated. Type species: Cieufnegosia hildehrandtii Garcke. Chiovenda (1929) published C. chiarugii and established the section Syno- dontos to include that species, but did not provide a diagnosis for the name. Moreover, he based tlie section on (and derived its name from) a single trait, a gamophyllods involucel, which C. chiarugii does not possess. This name cannot be accepted as valid because the descriptio genericO'Specifica pertains only to monotypic genera (Article 42). Section Synodoutos is not monotypic and was not considered to be by Chiovenda. It is possible to validate Chiovenda's name by providing the necessary diagnosis, but since the name was descriptively ill-chosen, it is considered preferable to publish a new name not encumbered with a misconception. Such a course is not in violation of Article 62 since the name Synodontos was not validly pubhshed. The name Garckea is chosen to honor A. Garcke, who made the first signifi- cant evaluation of Cicnfuegosia (1860) and who later described tlie type species of this section. Key to Species of Section Garckea a. Stipules prominent, auriculate-clasping; foliar nectaries ?^, basal; petals yellow; androecium orange 2. C. hildehrandtii aa. Stipules inconspicuous, subulate; foliar nectaries 1-3, medially positioned, some- times absent; petals yellow, cream, or purple; androecium purple. b. Petioles as long as lamina or longer, puberulent or scabrous, especially at distal end; plant puberulent or pulx:scx?nt (at least on petioles); petals yellow or cream. c. Leaves reniform, denticulate, sparsely scabrous below and on petiole, non- odoriferous, S-ranked; foliar nectaries 3; peduncle exceeding leaves, articu- lated near center; petals cream 3. C. wehhii cc. Leaves flabelliform, slightly 3-lobed to deeply dissected, dentate, minutely puberulent on margin and petiole, odoriferous, 3-ranked; foliar nectary 1 (-3 or rarely 0); peduncle shorter than leaves, articulated basally; petals yellow 4. C. somaliana 1969] FRYXELL — CIENFUEGOSIA 197 bb. Petioles shorter than the lamina, glabratc; plant essentially glabrous; petals purple. d. Leaves obovate; calyx ecostiilatc, prominently punctate; foliar necta- ries lacking 5. C. chiarugii dd. Leaves cuneiform, odoriferous; calyx costulate, punctate or epunctatc; foliar nectaries 1-3 6. C. hcarnii 2. Cienfuegosia hildebrcwdtii Garckc, Kichl. Bot. Jahrb. Bot. Gart, Berl. 2: 337, 1883 [Holotype: Txamtei in Durunia. Im Gras auf salziocm Bodon, Hilde' hrandt 2325 (probably Bt). Neotypc: Tanzania: Lushoto District: 3 miles NW of Mombo, 29 Apr. 1953, Drnmmond & Hemsley 2285 (BR, isonco- types: B, FI)]. Illustrations: Fig. 17, a-c; also Marshall, Parson, and Hutchinson (1937) Figs. 1-3; Exell& Wild (1961) t. 85; Fryxell ( 1968) Fig. 3,b. Small, branched, upright shrub. Stems usually minutely pubescent, sometimes glabrous, terete. Leaves broader than long, cuneate, flabelliform, unlobed to weakly trilobcd, irregularly dentate, 3'nerved, glabrate above, sometimes minutely pubescent below, ciHate-margined. Fohar nectaries 3 (rarely lacking), basal. Petioles about Vs length of lamina, densely pubescent throughout to moder- ately pubescent on upper surface only. Stipules auriculatedasping, asymmetrical (rarely lanceolate, symmetrical), acuminate, entire, pubescent or glabrate, cihate- margined, 6-10 mm long (V^2-% length of petiole) rarely longer, half as broad. Peduncles solitary (rarely multiple), equaling or barely exceeding petiole, articu- late and bracteatc near center, punctate, often pubescent, nectaries prominent. Involucel bractlets 1-3 mm, central bractlet of each group often exceeding lateral ( ) brous or pubescent; lobes shallow, 3-ribbed, rotund-apiculate. Petals 3-4 cm (rarely smaller), sparingly punctate, yellow with large dark-red spot on claw. Androecial column pallid, punctate; filaments dark-red; anther mass globose; an- thers unranked, orange; pollen orange. Style epunctatc, exceeding androecium; stigma dark-red, pubescent. Fruit o\'oid to obo\oid, obscurely punctate, glabrous to weakly pilose, 3-4-loculed, 10-14 mm long. Seeds turbinate, 5-6 mm long, densely comose; hairs 8-10 mm, straight, soft but appearing bristly, red-brown to dirty-tan color. Embryo epunctatc, but with reddish punctae developing on meso- cotyl and cotylcdonary margins upon germination. Chromosome number: w ^= 11. The variation in vestiture that is characteristic of this species can be noted even on individual collections. For example, both glabrous and pubescent branches occur among the duplicates, even on a single sheet, of Schlechter 11770 (Z). Hochreutiner separated two branches of the collection of Braga 82 (Z), dif- fering in pubescence, as distinct varieties in manuscript names. I do not believe that a varietal distinction based on pubescence is warranted because: (a) there is no correlated variation in other traits, and (b) there seems to be no geographical basis for the distinction. Therefore, the varietal epithets suggested by Hochreutiner shall herein remain unrecognized. [Vol. 56 198 ANNALS OF THE MISSOURI BOTANICAL GARDEN Distribution (Fig. 8): Natal, Swaziland, and the Transvaal in the south northward along the eastern coast of Africa to central Mozambique. It also occurs in Tanzania, but according to Pearson (1954) is absent between there and central Mozambique. Specimens were cited by Gon^alves (1963) from southern and central Mozambique; Marshall, Parson, and Hutchinson (1937) cited its occur- rence in Southern Rhodesia, northern Mozambique, and southern Tanzania, indicating a relatively continuous distribution. These localities are included in the distribution map since there appears to be no question of the identity of the plants concerned. In the southern part of its range, where it overlaps the range of C. gerrardii, it is separated ecologically from this species, occurring at lower alti- tudes. Ulbrich (1914) asserted that C. hildebrandtii occurs in Madagascar, but no specimens from there are known to me. Marshall, Parson, and Hutchinson (1937) gathered important ecological data on the distribution of C. hildehrandtii, at least in the soutliern portion of its range, which Pearson (1954) indicated to be its principal distribution. They found it to occur in close association with Acacia harhertuuensis Schweick. at altitudes below 1300 feet, to be absent from areas with "thin, stony soils," and to be present in regions of "better and deeper soils." They divided the habitats in which C. hilde- hrandtii occurs into three types: "(a) the crests and sloping sides of low elevations, (b) along water channels, and (c) pans (vleis)." (The last named are depressions often characterized by impermeable (saline?) soil, vegetated only on the margins.) They noted that the occurrence of the plant on the water channels and pans is "consequent on proximity to watercourses" and drew particular attention to the dissemination of seeds by water, since their bristly vestiture renders them buoyant. However, C. hildehrandtii occurs primarily on the crests of low hills having good soil cover. Water-borne dissemination does not account for this occurrence. The seeds of this species have bristly hairs that stand straight out, and thus appear to be well-adapted to wind-transport (as a tumbleweed). The seeds drop from the plant very quickly upon maturation, and will germinate with the first adequate rainfall, since they apparently have no dormancy. Plants have an extensive root system, which can reach layers of permanent moisture (Marshall, Parson, and Hutchinson, 1937). These characteristics seem to account for the primary hilltop distribution. The capacity of this species for wind dissemination may also account for its greater range, as compared to the other species in subg. Articulata. Gos- sypium australe F. Muell., native to arid areas of Australia, may have been dis- seminated in a similar manner (Fryxell, 1965). South Africa: natal: Zululand: Ntambanana, Taylor s.n. (PRE); 9 mi from Hluhluwe, Reserve gate, Hlabisa rd, Wells 2048 (PRE); Umfolosi Game Reserve, 600 ft alt, Ward 3240 (PRE), Edwards 3298 (PRE); Ingvvauuma, red sand nr vlcy, Strey 4757 (PRE). Transvaal: Barberton Distr.: Komati Poort, ah 1000 ft, Schlcchter 11770 (BM, BR, G, L, LE, MO, P, S, US, Z), Rogers 12624 (Z), 22389 (Z); Lebombo Elats, Strey 4028 (PRE); Kruger National Parl<, Nelspruit, SVi mi N of Malelanc, 1000 ft, Codd 5229 (PRE); Skukuza, on brackish land, 1000 ft alt, v. d. Schijff 3409 (MO), 3938 (PRE). Swaziland: Stegi to eastern foot of Lubombo, Burtt-Davy 10572 (PRE); Bremersdorp, Wood s.n., 25 Apr 1927 (PRE); Sipofaneni, Murdoch 123 (TAES). Mozambique: sul do save: Sabie, Moambo, Pcdrogdo 27 (LIE), 225 (LIE); Mo- amba, Harland s.n., 1930 (C); Umbeluzi, Boric 257 (BM), 55 J (PRE), Lcmos 127 (LIE). Marracuene, Pedro & Pedrogdo 643A (LIL-p.p.), 663 (LIE); dc Carvalho 219 (BM); 1969] FRYXELL — CIENFUEGOSIA 199 ^TXS3^ F Fig. 7. Distribution of C. gerrardii • and C. heteroclada O. Fig. 8. Distribution of C. hil- dehrandtii •, C. chiarugii ®, C. somaliana X, C. welshii O, and C. hcarnii A. Boane, Braga 82 (Z); Guija, Pciro & PeJrog^o 2ii4 (LIL); Goba, Wood 6828 (PRE); bci Pisini zwischen Lebombo und Dclgao Bai, Schenck 825 (Z); Myre £r Bahinhas J 663 (BM). Tanzania: lushoto distr.: 3 mi NW of Mombo, on waterlogged soil, alluvial plain, 425 m, Drummond & Hemsley 2285 (B, BR, FI). IN cultivation: Fryxell 14001 (ARIZ, TAES); Fryxell 14007 (provenance: Sipofa- neni, Swaziland) (TAES). 3. Cienfuegosia welshii (T. Anders.) Garcke, Eichl. Bot. Jahrb. Bot. Gard. Berl. 2: 337, 1883 (as C. Welshii^. Hibiscus Welshii T. Anders., Jour. Linn. Soc., London 5 (Suppl. 1): 8, 1860 [Type: Aden, rarissime: Hook, f. & T. (K-n.v.)]. Fugosia Wclschii Hochr., Ann. Cons. & Jard. Bot., Geneve 4: 174, 1900. Illustrations: Fig. 18, d-g; also Anderson (loc. cit.) t. 1; Janda (1937) Figs. 27, 33; Fryxell (1968) Fig. 3,c. Upright subshrub from woody rootstock. Stems woody, glabrous, punctate. ( s), abou (below) medially to sub-apically positioned. Petioles as long as or longer than lamina canaliculate, espec upper side (h LIS (1-4 mm), 1/2-1 ) or sometimes ( ) flowered, articulate and with small bract near center, glabrous, [Vol. 56 200 ANNALS OF THE MISSOURI BOTANICAL GARDEN punctate, often exceeding leaves (1-7 cm), nectaries large. Involucel bractlets 1-2 mm, glabrous, punctate, central bractlct of each group often exceeding lateral bractlets. Calyx V2-I cm, minutely pilose to glabrate, costulatc, prominently punc- tate (punctae V3 mm, distributed in rows along veins throughout calyx); lobes shallow, 3'ribbcd, apically pubescent within. Petals 2 3 cm, pale yellow or cream-colored with dark-maroon basal spot covering V3 of petal, sparingly punctate. Androecium epunctate, dark maroon, anther mass globose; pollen yellowish. Style epunctate, exceeding androecium; stigma dark purple, pubescent. Fruit 3'4-loculed, prominently punctate (punctiie V3 mm diam.), minutely and sparsely pilose without, glabrous within, elongate (7-10 mm broad, 12 30 mm long). Seeds 4-5 mm, copiously hairy; hairs fulvous, loosely apprcsscd to nearly straight, to 8 mm. Embryo epunctate, but with pinkish punctae developing on mesocotyl and translu- cent punctae developing as a single marginal row on the cotyledons upon germ- ination. Chromosome number: n ^^ 11. The vernacular name "Blamhar is recorded on Gooding's specimen from Somaha, and Hearn (1968) notes that in southern Yemen the term "atah baladi' (cotton of the country) is applied to Cieufuegosia spp. as well as Gossyphan spp. Distribution (Fig. 8): This species occurs under very arid circumstances in Aden, as well as in Somalia below 700 m. Brettell (1966) states that C. wehhii "is to be found in some profusion" in the Jebel Sirah range of mountains just to the east of Dirgag. Hearn (1968) cites additional locations and indicates its altitudinal range to be up to several hundred meters above sea level. Hutchinson (1947) noted a greater range of variability, at least in leaf form, in material from Somalia, which he retained in C. nelshii. Examination of the specimens, however, indicates that specific differences exist in this material that merit taxo- nomic recognition (Fryxell, 1967a). The separation of the following species (C. somaliivia) provides this distinction. Southern Yemen (Aden): Ad fances mantis Scham-Scham: Deflers s.n., 3 Apr 1886 (Z), Deflers 60 (K, P); Goldmore Valley, Schwcinjnrth 75 (BM, C, Z); 2 m alt, Ilihle' hrandt 780a (BM, LE); Beecari s,n,, Mar 1870 (FI); Waring 19 (K). Somalia: Los Anod, 2300 ft alt, Uetnming 2J87 (K); Mordalc, alt 740 m, Gooilmg 207 (4) (K). IN cultivation: Fryxell 14006 (TAES). 4. Cienfiiegosia somaliana Fryx., Britt. 19: 33, 1967. [Type: Gan Libali, Urgo/Jilbadaig, 11 July 1945, Glover & Gilliland 1206, (Holotype: K)]. Illustrations: Fig. 18, a-c; also Fryxell (1967a) Fig. l,g, (1968) Fig. 3,d. Subshrub with odoriferous foliage. Stems angled, punetate, minutely puberu- lent (hairs ca. 0.1 mm, often simple) beeoming glabrate. Leaves 3-ranked; 3-5-veined, 8-25 mm long, somewhat broader, 3-lobed to deeply disseeted, having charaeteristie musky odor when handled; leaf segments obovate to linear, secon- darily divided, dentate, punctate (especially on margins), margins minutely ciliate, otherwise puberulent to glabrate; foliar nectaries 1-3 (or rarely lacking), medial, inconspicuous. Petiole minutely puberulent (most densely so near juncture with lamina) to glabrate, about equahng lamina. Stipules minute (2-6 mm), triangular, puberulent to glabrate, caducous. Peduncles Vi-lVz cm, minutely 1969] FRYXELL — CIENFUEGOSIA 201 puberulcnt to glabrate, punctate, articulate and bracteate near base. Involucel of ca 9 unequal bractlets, imperfectly arranged in 3 groups; bractlets triangular, 2-5 mm, punctate, puberulcnt to glabrate, central one of each group often longest. Calyx 8-10 mm, costulate, punctate (punctae translucent); lobes shallow, 3- ribbed (intercostal areas scarious), minutely puberulcnt to glabrate without, spar- ingly short-pubescent at apices within. Petals 15-25 mm, prominently punctate (punctae dark red), bright yellow^ with large maroon spot covering basal half. Androecium maroon; anther masses globose; pollen yellow. Styles slender, exceed- ing androecium and nearly equaling petals, red, epunctate; stigmas decurrent. maroon, glabrous. Fruit ascending-pilose when young, becoming minutely and sparsely pilose at maturity, prominently punctate (punctae Vs mm diam,)> 3- loculed, about twice as long as broad. Seeds 5-7 mm, copiously hairy, the hairs nearly straight, tan or brown, 6-8 mm. Embryo light-punctate along cotyledonary margin, punctae darken upon germination, those of the mesocotyl becoming bright red. Chromosome number: n ^^ 11. Distribution (Fig. 8): Somalia: in mountains along northern coast; Ethiopia: in vicinity of Dire Dawa. Cioifuegosia somahana occurs at elevations from 900 m to 1700 m, while C. wehhii occurs from sea level up to 700 m. The vernacular name 'l)iei* is recorded on llally 9674. As previously noted (p. 189), these plants were included in C, ivelshii by Hutchinson (1947) who referred to a range of variability in leaf form. However, the two taxa that are here distinguished are differentiated by more than variation in leaf form, as is shown in the following tabulation ( 1967a, Table 1). from Fryxell, C. u'chhii C. somaliana Foliage : Leaf form: Non-odoriferous Odoriferous Broader than long, reniform, denti- Slighdy broader than long, suborbi- culate cular to dissected, dentate Vestiture : Scabrous (on stipules, petioles, and Minutely pubcrulent to glabrate underside of leaf} Greater than lamina Petiole length: Peduncle length: 3-7 cm, medially articulate Petal color: Equaling lamina V2-2V2 cm, basally articulate Pale yellow with small spot cover- Bright yellow with large spot cover- Stigmas: Seed length: ing low^r third of petal, sparingly punctate Pubescent 4-5 mm ing lower half of petal, promi- nently punctate Glabrous 5-7 mm Somalia: gan libah: Urgo/Jilbadaig, Glover & Gilliland 1206 (K), Golis Range, Glover & Gilliland 1216 (K); on first terrace facing N, 5100 ft, aromatic leaves. Bally 11910 (K). hargeisa: sandy ground by stream bed, aromatic, 4 300 ft, Gillett 4239 (FI, Fodjor Esc. W W / — on crest of spur, leaves fragrant when crushed, Newhould 716 (K). Ethiopia: Generally dry area, 2 km SE of Dire Dawa, 3000 ft alt, Getahun s.n,, 11 Nov 1965 (TAES). IN cultivation: Vryxell s.n. (provenance: Dire Dawa» Ethiopia) (BM, CTES, TAES). [Vol. 56 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN 5. Cienfuegosia chiarugii Chiov., Fl. Som. 101, 1929 [Type: Somalia, Costa del Migiurdni: Altipiano calcareo presso Hafun, 23 May 1924, Piiccioni £r Stefa- nini 617 (680) (Holotype: FI)]. Illustrations: Chiovcnda (1929) PI. Ill, fig. 3 (not PI. Ill, fig. 1 as stated by Chiovcnda, nor PI. 1, fig. 3 as cited in Index Londonensis.) Low woody subshrub. Stems glabrous, punctate. Leaves cuneiform-obovate, entire, obtuse to weakly and irregularly undulate apically, glabrous, punctate, almost as broad as long; foliar nectary lacking. Petioles 3-4 mm (ca Vb length of lamina), slightly scabrous near juncture with lamina. Stipules minute (1 mm), subulate. Peduncles 3-4 mm, prominently punctate, glabrate, basally articulate with small scale at articulation (articulation obscured by shortness of struc- tures). Involucel bractlets 1-1 V2 mm, prominently punctate, glabrate. Calyx ) (punctae denser toward apices of lobes) ower, glabrate without, pilose within (01 In addition Chiovenda's description adds: Androecium purple; filaments 6 mm. Style simple; stigma clavate, 3-lobed. Petals violet. The fruits and seeds are unknown. Chiovenda's complete and generally accurate diagnosis of tliis species included the description of a gamophyllous involucel and, indeed, emphasized it as a dis- tinctive feature. Moreover, he used this trait as the sole diagnostic feature for sect. Synodontos, which he erected to include this species (cf. p. 196). Examina- tion of the type indicates that the involucel is not gamophyllous but is similar to that typical of the genus with bractlets wholly distinct. Chiovenda's description must therefore be amended. Distribution (Fig. 8): Known only from the type locahty, in northeastern Somaha, the only region of Africa beyond 50° east longitude. Somalia: Costa dci Migiurtini: Altipiano calcareo presso Hafun, Puccioni & Stefanini 617 (680) (FI). 6. Cienfuegosia hearnii Fryx., Britt. 19: 33, 1967 [Type: In cultivation, Tempe, Arizona, Fryxell 14005 (Holotype: ARIZ, isotypes: LE, SD, TAES, US)]. Illustrations: Fig. 18, h-k; also Fryxell (1967a) Fig. 1, e-f, (1968) Fig. 3,e. Low perennial subshrub, branches erect, fohage odoriferous. Stems glabrate. Leaves 5-ranked, leathery, rhomboid to subrcniform, sometimes weakly 3-lobed, coarsely serrate, undulate to crisped, cuneate to truncate, glabrous, 3-5 nerved, 15-20 mm long, 20-2 5 mm broad (or smaller under stressed conditions), with characteristic musky odor when handled; foliar nectaries 1-3, median. Petioles 5-15 mm long, scabrous near juncture with lamina, sparingly so elsewhere. Stip- ules minute (1 mm), sessile, subulate, glabrous, caducous. Peduncle axillary, 5-20 mm, articulate near center or below with small bract at articulation, punctate, glabrous, nectaries large. Involucel bractlets triangular, 2 mm long, punctate. Calyx 7-8 mm, costulate, scarious in intercostal areas, densely to sparingly punc- tate (sometimes epunctate); lobes shallow, ovate, 3-5-ribbed. Petals 2 cm. 1969] FRYXELL — CIENFUEGOSIA 203 sparingly punctate, dark purple to roseate (rarely pallid) within, greenish margin without (where exposed in bud), very dark spot covering lower third of petal within. Anther mass ellipsoid; staminal column dark purple, punctate; anthers numerous, dark purple; pollen cream-colored to orange. Style yellowish, punctate, roecmm elongated, 1-1 Vz cm long, 3-4 loculed, rostrate, weakly strigose, punctate. Seeds ( ;ht) on mcsocotyl and translucent punctac as single marginal row on cotyledons upon germination. Chromosome number: w^ 11. The original description of this species, which was based on cultivated material from a single accession, may now be broadened to account for the variation found in a wider range of material. Although the collections cited exhibit an appreciable variability in several traits, they appear to represent but a single taxon. Four vernacular names are on record for C. hearnii. Hearn (1968) refers to the name "atah haladi" ("cotton of the country") used for both Cienfuegosia spp. and Gossyinum spp. in Southern Yemen. In addition, collectors' notes record "Lawa" (Thesiger s.n.), "Tamrat el Ghanam" ("sheep browse"), and "Shejra el Ham" ("plant of distress"), the latter two from Maxwell-Darling 94. Distribution (Fig. 8): Hearn (1968) states that this species is known from three localities on the southern margin of the Arabian peninsula groudng in the coastal httoral and to an altitude of 1000 m. Additional specimens (Thesiger s.n. & Popov et al. 4236) extend this distribution more than 100 miles inland. This is apparently an environment of extreme aridity. Southern Arabia: hadhramaut: Mcifa Hajar, on pebbly plain, alt 50 ft, Maxwell- Darling 94 (BM), Giiichard 122 (BM); Riyan, Guichard 122A (BM); Radum, alt 100 ft, __^ .— _^Av ___^^ - p1. /> l"1 1 T^ _. _j _T A ^ ^ ^ Hearn 4 CBM); W Halaya Well, Thesiger s.n., 16 Feb 1946 (BM). Pap IN cultivation: Tempe, Arizona, P. Fryxell 14005 (ARIZ, LE, SD, TAES, US). 3. Sect, Dioica Fryxell, sect. nov. Folia cuneata, simplicia vel exigua lobata. Flores dimorphi, in racemis brevibus antequam folia portata sunt. Foliola involucrorum tria. Plant glabrous. Stems angular, punctate. Leaves simple to weakly lobed, cuncate, entire, with medial to subapical foHar nectaries. Petioles short. Stipules hnear, minute. Flowers arising in short racemes from ground level before leaves. Peduncles elongate. Involucral nectaries lacking. Involucel irregular, typically of (punctae ) epunctate. Androccium palhd, narrow. Fruit obovoid, glabrous, punctate. Seeds densely comose; hairs straight, rufous-red. Type species: Cienfuegosia heterodada Sprague. The name of sect. Dioica is chosen in reference to the dioecious condition found in its only species. 204 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 7. Cienfuegosia heteroclada Sprague, Kew Bull. 1907: 48, 1907 [Type: North- ern Nigeria, Kontagora, in the bush after burning, Dalziel 122. (Holotype: K-n.v., isotype: G)J. Fugosia heteroclada Sprague, op. cit., 49, pro syn. Illustrations: Fig. 17, f-g; also Hutehinson & Dal/iel (1928) Fig. 112. Mueh-rcduced subshrub from woody rootstoek. Species dioecious. Stems ridged; inflorescence a short raceme, exceeded by vegetative stems. Leaves elliptic to obovate, or sometimes apieally 2'3-lobed or 2-5-dentate, glabrous above and below, 3-5-nerved. Fohar nectaries 1-3, the central one near apex of leaf, the laterals near middle. Petioles canaliculate, glabrous, punctate. Stipules 2-5 mm, glabrous. Rachis of inflorescence punctate, glabrous, angular, pallid, pedicels arising from nodes of rachis, bracteate at nodes, 1-3 cm, glabrous. Involucral bracts irregular in form and number; typically 3, each triangular (or sometimes apieally laciniate or 2-3-parted, suggesting a fusion of parts), 4-5 mm long, 2 mm broad, glabrous. Calyx infundibular to campanulate, 9-15 mm, glabrous, punctae sometimes con- lined to tips of lobes, green, sometimes torn in flower; lobes sometimes sufi^used with purple. Petals 2-3 cm. Androecium 6-12 mm, pallid; filaments 1-3 mm. (Pistillate flowers not seen.) Fruit 18 mm long, ca. 1 cm broad. Seeds ca. 8 mm, hairs ca. 1 cm long. Emsley (1957) notes the existence of floral dimorphism in this species and states that about 90% of the plants are staminate, 10% pistillate. The illustra- tion presented by Hutchinson and Dal/iel (1928) shows a flower with a greatly reduced gynoecium that apparently represents the staminate form. The occurrence of dioecism is unknown elsewhere in Cienfuegcnia, and occurs elsewhere in the tribe Gossypieae only in the genus Hampea. Distribution (Fig. 7): Nigeria; vicinity of Kontagora (and Naraguta?); Ghana. It has a limited distribution, although described as "locally abundant" (Emsley, 1957). Plants of C. heteroclada are nearly acaulescent, flower directly from the perennial rootstoek, and fruit in a relatively short time. The species is thus well-adapted to the periodically burned-over savannahs in which it occurs. Nigeria: Kontagora, Northern Nigeria, Dalziel 122 (G). Ghana: Tongo Hills, E side of Zuarungu, Morton A 1286 (K); Tumu, nr Rest House, Ahiekpor s.n., 10 June 1966 (GC) Adains Gr Akpabla 4364 (GC); roadside ca 2 mi S of Han, Adams 4013 (GC); footpath from Pwalgu Bridge to Gbeigo village, Enti & Agyakwa s.n., 20 Jan 1966 (GG). II SuBG. Cienfuegosia Cienfuegosia Cav., Diss. 3: 174, 1787 (as genus). Leaves of variable form, generally punctate, but often obscurely so; fohar nectaries lacking. Flowers axillary, solitary, on unjointed peduncles (or peduncles sometimes articulated in C.affinis^. Involucral nectaries 3 or none. Involucel of 8-12 bractlets (or involucel lacking in section Friesia'); bractlets ungrouped, minute and subulate to prominent and lanceolate or spatulate. Calyx punctate (punctae sometimes obscure); lobes acute to acuminate. Petals punctate or epunc- tatc, yellow (or nearly white in C. tripartita') with dark spot on claw often having yeflow radii, or without spot. Stigmas decurrent or capitate, 3-5-lobed; styles 1969] FRYXELL — CIENFUEGOSIA 205 sometimes divided apically. Fruits glabrous or densely villous, punctate or epunc- C. digitata Cav. suture line (in sect. Cienfuegosia) . fiicgosia is the type of the genus Cienf Key to Sections of Subgenus Cienfuegosia a. Stigmas decurrent (or at most subcapitatc); fruits glabrous, cpunctate; bractlets of involucel ca. 9, usually minute, linear-subulate; inner suture hairs present on capsule; involucral nectaries usually present 4. Cienfuegosia (p. 205) aa. Stigmas capitate; fruits glabrous or pubescent, usually punctate; bractlets of in- volucel 8-10, lanceolate to spatulate, subequal to calyx, or lacking; suture hairs lacking; involucral nectaries lacking: b. Upright shrub or perennial herb; stipules filiform; involucel subequal to calyx; petals punctate 5. Rohusta (p. 216) bb. Perennial herb, decumbent or ascending; stipules subulate or auriculate; in- volucel subequal to calyx or lacking; petals epunctate (or at most obscurely punctate). c. Involucel subequal to calyx : stipules subulate, symmetrical; plants pro- cumbent or ascending 6. Paragiiayana (p. 220) cc. Involucel lacking; stipules auriculate, asymmetrical; plants procumbent 7. Vriesia (p. 228) 4. Sect. Cienfuegosia Cienfuegosia Cav., Diss, 3: 174, 1787 (as genus). Herbs or subshrubs arising from perennial woody rootstock. Stems ascending, green, 5-angled, minutely scurfy-puberulent to glabrous, obscurely punctate. Leaves entire, tripartite, or digitately divided, minutely puberulent to glabrous, punctate (but punctae obscure and usually visible only with magnification and careful illumination). Petioles canaliculate, minutely punctate, glabrous or mi- nutely puberulent especially near juncture with lamina. Stipules minute. Peduncles glabrous to scurfy-puberulent, incrassate above, minutely punctate, surmounted by 3 involucral nectaries (sometimes lacking). Involucel of ca. 9 elements; bractlets distinct, minute, filiform, persistent. Calyx 15-veined, prominently punctate, pu- berulent to glabrate; lobes acuminate or sometimes acute. Petals epunctate, yellow (sometimes white) with or without dark spot on claw. Stigmas decurrent (or sub- capitate in C, subternata^, 3-lobed; styles undivided. Fruits epunctate, 3-valved, glabrous without, with hairs on inner suture line. Lewton (1925) drew attention to the capsulary fringe of hairs that char- acterizes this section, as it is found in C. heterophyUa (Vent.) Garcke and C. yiicatanensis Millsp. One of the most striking features of this section is the variability of leaf form within plants of all of the species. \\ hen speaking of leaf form in this group, therefore, it is necessary to use the concept of "leaf spectrum" (Melville, 1953), Type species: The type of sect. Cienfuegosia is the type of subg. Cienfuegosia, C. digitata Cav. Key to Species of Section Cienfuegosia a. Leaves deeply digitately divided, often secondarily so; anthers 5-ranked; petal spot (when present) and staminal column carmine; distribution African 8. C. digitata aa. Leaves entire, lobed, rarely with secondary divisions as above or trifoliolate; an- thers usually unranked; petal spot and staminal column dark-maroon (or spot [Vol. 56 206 ANNALS OF THE MISSOURI BOTANICAL GARDEN absent and staminal column light); distribution American. b. Leaves narrowly lanceolate to oblong, entire (lower leaves rarely trilobed); petiole glabrous; involucral nectaries usually lacking; corolla rotate; petals yellow, small (1-2 cm), lacking basal spot; pollen yellow; distribution Yuca- tan, Florida, Bahamas, Cuba 9. C. yucatanensis bb. Leaves broad and entire or variously divided; petiole puberulent at distal end; involucral nectaries present; corolla campanulate; petals yellow or white with basal spot; pollen orange or reddish; distribution Middle or South American. c. Leaves broadly ovate to moderately 3-lobed; distribution northern South America and Middle America. d. Style and stigma exceeding androecium; involucral bractlets 3-10 mm; leaves apiculate 10. C. rosei dd. Style and stigma largely included within androecium; involucral bract- lets 1-3 mm; leaves usually acute to obtuse 11. C. heterophylla cc. Leaves large, entire to deeply divided or tripartite. e. Leaves trifoliolate; leaflets narrow; petals yellow, with large basal spot having yellow radii; distribution Paraguay 12. C. suhternata ee. Leaves entire, cuneiform to variously divided (sometimes secon- darily so); petals white or pale yellow, with small basal spot usually lacking radii; distribution Peru and Ecuador ... 13. C. tripartita 8. Cienfiiegosia digitata Cav., Diss. 3: 174, 1787 [Type: Senegal: Adanson MSA (Holotype: P, isotypes: MA, P)]. Fugosia digitata Pers., Syn. 2: 240, 1806. Hibiscus Cavanillesii Kuntze, Rev. Gen. PI. 1: 69, 1891 (based on Cienfugosia digitata Cav.) non H. cavanillesianus H.B.K. C. digitata var. lineariloba Hochr., Ann. Cons. Jard. Bot., Geneve 6: 56, 1902 [Type: S.W. Africa: Grootfontein, Hereroland, 21 May 1899, Dinter 669 (Z)]. C. junciformis A, Chev., Rev, Bot. Appliq. 30: 267, 1950 [Type: Niger: Chevalier s.n. 16 Feb 1950 (PAT ? -n.v.)]. C. junciformis var. ruyssenii A. Chev., loc. cit. 268, (as ruysseni^ [Type: Niger: Ruyssen s,n. (PAT ? -n.v.)]. Illustrations: Fig. 20, a-d; also Cavanillcs (1787) PI. 72, fig. 2, f-m; Lamarck (1819) iii, t. 577 (n.v.); Chevalier (loc. cit.) PI. XII; Fryxell (1968) Fig. 3,h. Plant minutely puberulent on meristems, becoming essentially glabratc through- out. Stems minutely and obscurely punctate. Leaves 5-ranked, digitately 3-5-parted; lobes often secondarily divided apieally, sometimes simple, lanceolate to oblan- ceolatc, acute. Petioles punctate, ca. V2 length of lamina or less, minutely puberu- lent near juncture with lamina. Stipules 1-3 mm, linear, pubescent (sometimes glabrate). Peduncles axillary, solitary, unjointed, costulate, obscurely punctate, minutely puberulent to glabrate, 1-4 cm long QVz-Wz times length of leaf), usually surmounted by 3 prominent involucral nectaries. Involucel bractlets 1-4 mm, usually glabrate. Calyx 1-1 V2 cm long, minutely puberulent to glabrate, costulate (intercostal areas whitish), punctae V4 mm diam., deeply lobed; lobes 3-ribbed, acuminate (sometimes acute). Petals yellow with carmine spot having yellow radii on claw (spot rarely lacking), 1^/2-2^2 cm (rarely larger), with undulate margin. Androecium carmine or pale; anther mass globose to obovate; anthers numerous, arranged in 5 ranks opposite the petals (or ranks divided to appear as 10); pollen orange. Style pallid, exceeding androecium; stigma decur- rent, pubescent, whitish. Fruit ovoid, 8-12 mm, copiously hairy (hairs 2 mm) on internal suture margins. Seeds 3-4 mm, densely comosc; hairs appressed, tan, 6-8 mm long. Embryo epunctate. Chromosome number: n^ 10. 1969] FRYXELL CIENFUEGOSIA 207 ^^i%^i -■ ^. 10 Fig. 9. Distribution of C. digitata. Fig. 10. Distribution of C. yucatanensis. Fig. 11. Dis- tribution of C. heterophylla A , C. affinis •, and C. cuyahensis O. Cienfuegosta digitata is relatively uniform morphologically, although leaf form exhibits considerable developmental plasticity. Herbarium specimens and cultivated plants both show this Such phenotypic variation is well within the response potential of an individual plant, as indeed can be noted on the holotype of the species. Hochreutiner's var. lineariloha was based on such variation and has therefore been reduced to synonymy. One aberrant specimen from the Transvaal (Schlieben 9248) produced multiple calyx lobes, a tendency not noted elsewhere. Most of the calyces on this specimen have 6 or 7 lobes rather than 5. This plant is, in all other respects, representative of C. digitata. The aberration therefore is considered not to have taxonomic significance. Another exceptional plant is Davey 224, which lacks a petal spot. [Vol. 56 208 ANNALS OF THE MISSOURI BOTANICAL GARDEN Distribution (Fig. 9): As previously noted (p. 186 and Fryxell, 1967b), this species has a disjunct distribution, occurring in western Africa and in southern Africa, but without any concomitant morphological differentiation. The northern segment of the distribution of C. digitata extends from Senegambia and southern Mauritania through Mali and northern Ghana to extreme western Niger and northern Nigeria. Reported occurrences as far east as the Sudan have not been substantiated and possibly refer to specimens of Gossypium somalense (Giirke) Hutch., as noted by Knight (1949). The southern S( of C. digitata extends from the southern parts of Angola and Zambia and north- eastern SAV. Africa through Botswana to western Rhodesia and the Transvaal. (1905, p. 225) (CN) verify its identity, and no other records from there are known. Mauritania: Tagant Plateau bctw Madjcria & Tidjikdjc. silty flood plain, G. 62 (BM). Senegambia: Perrottct 32 (G), 66 (BM, G, P, S,); Bords du Senegal Q\V Ao^Pcrrottct s,n., 1831 (G, L); Plaines de Richard, Perrottct s,n., 21 Sep 1824 (G, LE); Sichcr 58 (G, LE, M?, MO?, OXF?, S, UC); Dagana, Lepricur 1 (G, L, P); Adanson I45-A (MA, P); Roger 13 (LE); Berhaut 1344 (BR, Z); Ueiuielot s.n., 1835 (LE), s.n., 1839 (EI, OXE), Mali: Ansongo, Olufsen 385a (C), Hagerup 385a (BR); Goumal (Dioura): Daxey 224 (K), 233 (B); Env. de Tombouctou, Legagneiix s.n,, Jim 1920 (L); Goiirma : de Fada a Koupcla, Chevalier 24535 (BR). Nigeria: kano: Honohl 20 (Z). Angola: Huila bctw Humbe and Ruacana, Morros de Cualeque, Dungoc^na, c 1150 m, Exell £r Mendon^a 2872 (BM); Ruacana, R Cunene, c 1000 m, Exell & MendoiK^-a 2694 (BM), 2789 (MO). South-west africa: grootfontein distr.: Tsumeb, Dinter 7459 (B, M), Meyer 21 (B); Earm Onguma, Gelbhol/dline, Walter 429 (M, PRE); Farm Eockshof, Volk 125 (M); Amboland, Schinz 212 (Z), 2]3 (Z), Raittancn 19 (LE, Z), Dinter 669 (Z). outjo distr.: Farm Franken, Walter 2/187 (M); Otiizemba, MerxmiUler & Ceis,^ 1355 (M, PRE); Farm Kaross, Merxmiiller Sr Geiss 1549 (BM, M); Farm Ha/eldene, Karos Block, deW inter £r Lcistner 5110 (M). simkue distr.: Story 6509 (PRE); Klein Nanuitoni, Breyer 20592 (PRE); Onknanjumi, Widfhorst 21 (Z); Ondonga, Raiitanen 401 (Z); Vleyrand (Kalk) bei Naruchas, Dinter 7215 (B); Gautscha Pan, E of Karakuwise, Maguire 2183 (PRE). Zambia: Mazabuka, 3400 ft, Behhington 1693 (K), 3300 ft, Vet Officer 249 (PRE). Botswana; Northern Bechuanaland betw Mumpswe and Sigara pans, ca 26 mi NNW of mouth of Nata River, Mopane Woodland, 900 m, Drummond & Scagrief 5215 (BM); Mochudi, Harbor 6481 (K); Crocodile River, KUngherg s.h., Sep 1876 (S); nr Derdcpoort, 3000 ft, Codd 8873 (PRE). South Africa: transvaal ; N of Soutpansberg, betw Vivo and Waterpoort rd to Mopane, Schliehen 9248 (G, M); Soutpansberg, Liliput, Strcy 3493 (PRE), 5 mi S of Grass Valley, Potgietersrust, Meeusc 9581 (B, PRE); Klippan, Boshveld, Rehtnann 5223 (Z); Bestersput im Distrikt Blomfontein, Welti 17 (Z), Schlechter 4664 (Z). in cultivation: Fryxell 14008 (provenance: Outjo, SW Africa) (TAES). 9. Cienfuegosia yiicatauensis Millsp., Publ. Field Mus. Nat. Hist. Bot. Ser. 2: 74, 1900 [Tyiie: Yucatan: arid stony soil about 6 km S of Progreso, 5 Mar 1899, Millspaiigh 1693 (Holotypc: F-61693)]. Illustrations: Fig. 20, eg; also Small (1933) p. 860; Fryxell (1968) Fig. 3,1. Plant glabrous throughout. Stems minutely and obscurely punctate. Leaves entire, oblong to lanceolate, acute, 4-10 (rarely 20) times as long as broad; lower leaves sometimes broader and trilobed. Petioles punctate, V2 length of lamina or less, glabrous. Stipules 1-2 mm, linear, glabrous. Peduncles axillary, solitary, 1969] FRYXELL — CIENFUEGOSIA 209 unjointcd, costulate, obscurely punctate, glabrous, 2-5 cm long QVi-lVz times length of leaf) ; involucral nectaries usually lacking (rarely present, small) . Involuccl of 6-9 ungroupecl bractlets; bractlets V2-2 mm, glabrous. Calyx glabrous, costulate (intercostal areas light green), punctae Ve mm diam, 8-12 mm, equal- ing or exceeding fruit, deeply lobed; lobes 3-ribbed, acuminate. Petals yellow throughout, 1-2 cm, opening flat to form a rotate corolla. Androecium pallid, epunctatc; anthers few (10-30), unranked; pollen yellow. Style pallid, sparingly punctate, exceeding androecium; stigma decurrent, pubescent, whitish. Fruit 3- (occasionally 4-) valved, 6-8 nun, ovoid, copiously hairy on internal suture mar- gins. Seeds 2-3 mm, densely comose; hairs appressed, greenish-brown, 4-5 mm long. Embryo epunctate. Chromosome number: w^ 10. This species appears to be a specialized derivative of C. heterophylla (Vent.) Garcke. Its most notable specializations are in leaf form and in flower structure. The flower is small, opens to a fully rotate form, lacks the dark spot at the center characteristic of related species, and has significantly fcnver anthers and produces smaller fruits and seeds. \\ hether or not these floral differences are related to a specific adaptation to local poUinators, is not known. Smafl (1933) distir Indies'' and that (of C. heterophyUa') from South America but did not associate the former with C. yucataneush from Yucatan. He gave no nomenclatural recog- nition to the distinction. Two accessions of C. yucatanensis are available in cultivation, one from Yuca- tan, the other from Great Inagua, Bahamas. The examination of these living plants shows that the two accessions are clearly conspecific, yet that they are demon- strably different in leaf spectrum and in certain other characteristics. The study of additional accessions of this species from different segments of its distribution may reveal distinctions requiring taxonomic recognition, but if so, at no higher rank than that of forma. For the present, no such distinctions are made. Distribution (Fig. 10): Yucatan: in vicinity of Progreso and Merida; Florida: Grassy Key, Long Key, Lower Matccumbe Key, Lignum Vitae Key, and Key West; Cuba: Prov. Camaguey: Cayo Romano, Cayo Sabinal, and Cayo Guajaba; Ba- hamas: Great Inagua Island. Such a distributional pattern, around the southeastern shores of the Gulf of Mexico, suggests salt-water dispersal of seed. Mexico: yucatan: Progreso, moist open places, Sieere 3019 (F, MEXU, MICH); arid stony soil ca 6 km S of Progreso, Milhpaugh 7 693 (F); km 31 Merida Rd, Lundell & Lun- dell 8017 (F, GH, MEXU, MICH, TEX, US); Lukcfahr s.n., 26 Jul 1964 (GH, LE, SD, TAES); Chichankanab, Gamrjer 1269 (F); rd betw Sisal and Merida, Schott 693 (F), United States: Florida: Lowlt Matccumbe Kev, rocky hammocks, Small ct ah 11599 (GH, MICH, MO, S, US), Small 8392 (C, M, S, UC), in dry sandy soil, Moldenke 619 (ILL, MO, S, US), Busivell 1387 (ARIZ); Lignum Vitae Key, Blodgctt s.n. (GH), Chap- man 747 (US), Garber s.n., Aug 1877 (GH, PH, US); Grassy Key, coral soil, Curtiss 398 (F, GH, LE, M, MO, PH, US); Long Key, Simpson 469 (F, GH, S, US); Key West, Chap- man s,n. (US); S. Florida, Chapman s.n. (F, MO). Cuba: camaguey: Cayo Guajaba. Shafcr 634 (NY), 686 (F, GH, NY, US); Cayo Romano, Lomo dc Loro, Shafer 2639 (F, GH, NY, US); Cayo Sabinal, in a kind of sweet water meadow, Ekman 15487 (F, G, GH, MO, NY, S, US). Bahamas: Great Inagua, Dtmhar 146 (GH). [Vol. 56 210 ANNALS OF THE MISSOURI BOTANICAL GARDEN IN cultivation: Fryxell 15013 (provenance: Yucatan) (ARIZ, TAES); Fryxcll s.n. (provenance: Great Inagua, Bahamas) (C, CTES, GH, MICH, TAES). 10, Cienfuegosia ro?*ei Fryxcll, sp. nov. [Type: MEXICO: Oaxaca: between San Geronimo and La Venta, alt 200 ft, 13 July 1895, Nelson 2779, (Holo- type: US, isotype: GH)]. Illustration: Fig. 20, o-p. Frutex humilis, e radicc pcrcnni exoriens, puberula dense et minute in meri- stematibus (pili stellati, ea. 0.2 mm), plus minusve glabrescens omnino, Caules virides, 5-angulati, ascendentes, punctati minute; internodia brevia. Folia ovata et Integra vel trilobata moderate, 3-8 cm longa, apiculata, obscure punctata, circa latiora quam longa vel aliquantum angustiora. Petioli canaliculati, punctati, quam laminae V^-Vz longiores, puberuli minute praesertim prope juncturas laminarum. Stipulae 3-7 mm longae, lineares, punctatae, puberulae. Pedunculi axillares, soli- tarii, inarticulati, incrassati supra, punctati minute, puberuli minute vel glabrati, 2- 8V^ cm longi, circa longitudines foliorum subtentiorum aequantes vel parum exce- dentes, 3 nectariis involucellis terminati. Involucella ca. 9 bracteolis inaggregatis formata; bracteolae involucelliorum distinctae, 3-12 mm longae, filiformes vel subspathulatae, interdum punctatae, puberulae vel glabratae, persistentes. Calyces puberuli vel glabrati, costati leniter, punctati (glandibus ad basem tubi calycis seria- tis dispositis), 12-2 5 mm longi, profundc lobati; lobis calycibus 3-costatis, acutis vel acuminatis. Petala epunctata, IVz-'iVz cm longa, flava pallida; unguis saepe macula marronina radios flavos habente. Androecia marronina; antheris numer- osis (plus quam 30), imperfecte 5-fariam; pollenibus fusco-incarnatis; cumulis antherarum sphericis vel obovatis. Styli unici, punctati, androecia 6-8 mm ex- cedentes. Stigmata decurrentia, marronina, trilobata, pubescentia, petala fere aequantia ad apicem lata. Fructus 3-loculati, epunctati, 10-13 mm, ovoidei, glabri extus, ciliati sparse in margine suturae. Semina 4-5 mm, dense comosa; pili crispi- patentes, brunnei pallide, 5-6 mm longi. Chromosomatum numerus: w= 10. Cienfuegosia rosei is in some respects morphologically similar to C. hetero- phylla but may be distinguished from the latter species by: its longer involucral bractlets (and stipules); its apiculate leaves; its larger flowers with the st}de notably exceeding the androecium rather than largely included within it; and the less prominently punctate calyx but more prominendy punctate fohage. The plant appears also to achieve a smaller stature than C. heterophylla, with shorter internodes. As indicated in the discussion on phylogeny (p. 189), C. rosei is interpreted as having its closest affinity with C. tripartita. Its somewhat closer superficial resemblance to C. hcterophylla is considered to be an example of con- vergent evolution. Biosystematic studies will permit verification or qualification of this hypothesis. Rose recognized this plant as a new species in a manuscript name written on the isotype kept at the Gray Herbarium, but the name was never published. Rose's epithet, "humilis," is not taken up in publishing this species because of the possibility of confusion with the name published by Giirke, C. phloniidifolia var. humilis, even though Gurke*s epithet is illegitimate and of different rank. The name Cienfuegosia rosei is chosen instead to honor J. N. Rose. 1969] FRYXELL — CIENFUEGOSIA 211 Distribution: Mexico: Oaxaca: in the lowland coastal plain around Tehuan- tepec. The plant is described by King (1356) as "abundant [in] flat grazed areas [among] cacti and thorny leguminous shrubs", as I can verify from observation of the same site. Mexico: oaxaca: betw San Geronimo and La Vcnta, Nelson 2779 (GH, US); 4 km NNE of Tchuantepec, King 1356 (MICH, TEX), Fryxcll 749 (BH, CTES, F, MEXU, MO, NA, TAES, U); 12 km NNE of Tehuantcpec, King 394 (MICH); ccrca Juchitan, Miranda 8551 (MEXU); 7 mi W of Nilotcpec, Fryxell & Bates 906 (BH, TAES). 11. Cienfuegosia heterophylla (Vent.) Garcke, Bonpl. 8: 150, 1860. Rediitea heterophylla Vent., Hort, Cels, t. 11, 1800 [Type: Ventenat s.n. (Holotype: G, isotypes: B (photo in GH, MO), FI, MO?, P)]. Fiigosia heterophylla Spach, Hist. Veg. Phan. 3: 397, 1834. Hibiscus Redoutei Kuntze, Rev. Gen, PL 1: 69, 1891 (based on Redoutca heterophylla Vent.)- Fiigosia punctata Turcz. Bull. Soc. Nat. Mosc. 31: 196, 18 58 [Type; Brazil: Serra de Jacobina, Bahia, Blanchet 2702 (Isotypes: BM, G, NY, OXF, P)] (non Fugosia punc- tata Cunn. ex Benth. 1863). Illustrations: Fig. 20, h-k; also Ventenat (1800) t. 11; Redoute (1827) t. 118 (n.v.); Spach (1846) t. 4218; Martins (1892) PL 112, fig. 2; Sampaio (1926) PI. 1, fig. 2 (illustr. of a single seed of doubtful identity, labelled "Flacourtiaceae- Cienfuegosia heterophylla."); Fryxell (1968) Fig. 3,g. Plant minutely scurfy-puberulcnt (appearing lepidote) throughout, or glabrate on older foliage. Stems minutely and obscurely punctate. Leaves 5-ranked, entire, broadly ovate (sometimes rhomboid or obovate, rarely rotund) to moderately 3-lobed, acute to obtuse, rarely acuminate or apiculate, 2-3 times as long as broad (or broader, if lobed), ciliate-margined. Petioles punctate, V<4'V2 length of lamina, minutely puberulcnt especially near juncture with lamina. Stipules 2-5 mm, linear, pubescent. Peduncles axillary, solitary, unjointed, costulate, obscurely punc- tate, minutely puberulcnt to glabrate, 2-5 cm (rarely longer), approximately equaling or shghtly exceeding (rarely greatly exceeding) length of leaf, sur- mounted by 3 involucral nectaries. Involucel of 9 ungroupcd bractlets; bractlets 1-3 mm, glabrate. Calyx minutely puberulcnt to glabrate, costulate (intercostal areas light green), punctae Vb mm diam., 8-15 mm, deeply 5-lobed; lobes 3'ribbed, acute to acuminate. Petals l-lVz cm, yellow with maroon spot having yellow radii on claw. Androccium maroon; anther mass globose; anthers numerous (> 30); pollen orange. Style pallid, barely exceeding androccium; stigma decurrent, red- dish, pubescent. Fruit ovoid, 8-11 mm, copiously hairy (hairs 2 mm) on internal suture margins. Seeds 4 per locule, 3-4 mm, densely comose; hairs appressed, brown or tan, 6-8 mm long. Embryo epunctate. Chromosome number: m= 10. The Geneva specimen from the Herbier dc Ventenat is evidently the holotype since it matches Ventenat's plate (executed by Redoute) quite closely, whereas the Paris and Berlin plants do not match it. Distribution (Fig. 11): Northern Colombia and Venezuela; certain of the Lesser Antilles (Aruba, Margarita, Trinidad, and St. Thomas); Brazil: provinces of Bahia, Piaui, and Ceara (?). Specimens from Oaxaca, Mexico that were ten- [Vol. 56 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN tatively referred to C. heterophylla (Fryxell, 1967a), are here segregated as a distinct species, C. rosei. This taxon occurs primarily around the borders of the Caribbean Sea, but it also occurs in northeastern Brazil in what may constitute a distributional dis- junction. The Brazilian material is scanty, but differs from the typical form in being more densely puberulent throughout, and in having broadly acute (rather than acuminate) calyx lobes that are less prominently punctate (punctae ca. 1/10 mm diam.)- These differences are too shght to merit recognition at any level higlier than that of forma. Should the existence of such a taxon be con- firmed by future studies, it would, of course, be best established on Turczaninow's description of Fugosia juoictata. Colombia: magdalena: in dry soil, La Paz, alt 200 m, Haujit 4352 (US); Hwy, 7 km S of Riohacha, alt 20 m, Uaught 4473 (F, NY, US); Santa Marta,.alt 150 ft, Smith 779 (F, G, GH, L, MICH, MO, NY, P, PH, S, UC, US); Wagcner 76 (FI, UC). Venezuela: Maracaybo, Plee s.n, (P). sucre: Peninsula dc Paria, Stcycrmark & Agos- tint 91287 (US); Cumana, Moritz 529 (BM). bolivar: along water in ditch betw Rio Caroni and Ciudad Bolivar, alt 200 m, Steyermark 57634 (F, NY), Moritz 7 038 (BM, NY, P, US); Orinoco, Hunihohlt £r Bonpland s.n. (P). Antilles: aruba island: Bohlingh 6383 (L, NY, P). Margarita island: El Valle, Miller £r Johnston 66 (F, GH, MO, NY, P, US); Porlamar, Stephens 1 (GH). trinidad: Chacachacare Island, Britton 521 (GH, NY, US), Richardson 899 (IJ, US). ST. thomas: Krum Bay, von Eggers 350 (G, L, LE, M, Z), rocky hillside, Paradise Bay, Britton & Britton 211 (NY, US), HjaJmarson s.n., 24 Mar 1851 (S), Rose s.n., 1-5 Mar 1913 (NY, US), Krebs s.n., 23 Jan 1898 (C), Raiinkiacr s.n., 28 Mar 1906 (C), Bhmmcr 9 (G). Brazil: piaui: Gardner 2399 (Fl, NY but labelled "Prov. Ceara", OXF); baiiia: Serra de Jacobina, Blanchet 2702 (BM, G, NY, OXF, P). IN cultivation: Hurt Gels (from St. Thomas), Ventenat s.n. (B (photo in GH, MO), FI, G, MO?, P); (provenance: Isla Margarita, Venezuela) Fryxell 15004 (ARIZ, TAES); (provenance: Maracay, Venezuela) Fryxell 15010 (TAES). 12. Cienfuegosia subteniata (Hassl.) Fryx., Taxon 16: 321, 1967. C. heterophylla (Vent.) Garcke subsp. siihternata Hassl., Repcrt. Spec. Nov. 7: 380, 1909 [Type: Peguaho, Sierra de Amambay, Paraguay, Hassler 10899 (Lectotype: G, specifi- cally the sheet bearing Hassler's complete collection data; isotypes: G-2 additional sheets)]. Illustrations: Fig. 20, 1-n. Plant minutely puberulent on young foliage, becoming glabrate. Stems very obscurely punctate. Leaves trifoliolate, often unifoliolate above, rarely trilobed below; leaflets usually distinct, entire, lanceolate, acute, 7 cm long or less, ca. 1/10 as broad as long. Petioles V4 length of lamina or less (rarely longer on lower leaves), minutely puberulent especially near juncture with lamina. Stipules 2-3 mm, linear, puberulent. Peduncles axillary, solitary, unjointed, costulate, obscurely punctate, minutely puberulent to glabrate, 3-5 cm long (approximately equaling the leaf), surmounted by 3 involucral nectaries. Involucel of ca. 9 ungrouped bractlets; bractlets 1-3 mm, puberulent to glabrate. Calyx 1-1 Vi cm, glabrate, costulate (intercostal areas whitish), punctae < Vi mm diam, deeply lobed; lobes 3'ribbed, acuminate. Petals 2-3 cm, yellow with large (1 cm) maroon spot on claw having yellow radii. Androecium maroon; antlier mass globose; anthers numerous (> 40); pollen yellow^ (?). Style pallid, punctate, glabrous, greatly exceeding androecium (nearly equaling petals); stigma sub-capitate (i.e., 1969] FRYXELL — CIENFUEGOSIA 213 intermediate between deeurrent and capitate forms), dark red. Fruit ca. 1 em, eopiously hairy (hairs 2 mm) on internal suture margins. Seeds (immature) ca. 3 mm, angular, brownish-pubescent. This species is a narrow^ endemic, clearly alhed to the other species of this section, most obviously so in the nature of certain of its fruiting structures: involucel, calyx, and capsule. It is distinctive in its trifoliolate leaves and its large flowers with sub-capitate stigmas. Distribution (Fig, 12): restricted to the valley of the Rio Apa in northern Paraguay. Paraguay: in campo humido argilloso, Pc^iiaho, Sierra dc Amaml)ay, Hassler 10899, (G); Nordl. Paraguay (22-23° lat.) zwischcn Rio Apa iind Rio Aquidahan, Centurion, Nie- dcrcr Camp, Sumpfrand an grasfrcie Stcllu, Tkhrig 4022, (0-4 sheets (photo of 1 in MO), CH, L, M; there are 2 additional sheets at Geneve labelled "K. Yichrig 5230" that may be duplicates of 4022). \?>.Cienfiiegosia tripartita (H.B.KO Giirke in Mart., Fl. Bras. 12 (3): 578, 1892. Rcdoutca tripartita II.B.K. Nov. Gen. Sp. PI. 5: 294, 1821 [Type: Peru: Jaen de Braea- moros: Humboldt & Bonpland s.n. (Ilolotype apparently lost; photographs arc at MO and NY); Neot>pe: Peru: Dept. Cajamarea, Prov. Jaen, near the confluence of the Chinchipc and Maranon, 600-800 m, Wchcrhaticr 6214, Mav 1912 (US, isoneotvpe: F, GH)]. Fiigosia tripartita Steud. Nom, ed. II, 1: 649, 1840. Hibiscus tripartita Kunt/e, Rev. Gen. PI. 1 : 69, 1891. Fiigosia cuneata Benth., Bot. Voy. Sulph. 68, 1844 [Type: Ecuador: Guayaquil: Sinclair s.n, (K-n.v., US-photo)]. Hibiscus cuneata Kuntze, Rev. Gen. PI. 1: 69, 1891. Cienfuegosia Iwtcrophylla var. cuneata Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 13: 477, 1956. Illustrations: Fig. 19. Plant minutely scurfy-pubcrulent becoming glabrate. Stems punctate, some- times obscurely so. Leaves of variable form : sometimes ovate-entire or ovate- erenate, more eommonly 3- (rarely 5-) lobed in various degrees to completely trifoliolate, 2-7 cm long; leaflets lanceolate to obovatc, entire or rarely secondarily divided, sometimes single and narrowly oblong. Petioles usually less than Vz length of lamina, punctate, minutely puberulent near juncture with lamina (rarely puberulent tliroughout). Stipules linear, 1-3 mm, caducous. Peduncles axillary, solitary, unjointed, costulate, minutely punctate, glabrate, 2-9 cm long (approximately equaling or slightly exceeding length of subtending leaf), usually surmounted by 3 involucral nectaries. Involucel of 9 ungroupcd bractlets; bract- lets 1-3 mm, rarely longer, glabrous. Calyx glabrate, costulate, prominently punc- tate (punctae Vt> mm diam.), \-\V2 cm, deeply lobed; lobes 3-ribbed, acuminate. Petals epunctate (or rarely punctate), pale yellow to white with small maroon spot on claw (usually lacking radii), 3-4 cm (rarely smaller). Androccium maroon (sometimes light); anther mass elliptic; pollen orange. Style pallid, usually greatly exceeding androccium (nearly equaling petals), epunctate, (rarely punctate); stigma deeurrent or rarely sub-capitate, pinkish; stigmatic lobes sometimes par- tially free. Fruit ovoid, l-Wz cm, copiously hairy (hairs 2 mm) on internal 214 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 \'- 12A Fig. 12. Distribution of C. suhternata O and C. drummondii in South America. Fig. 12a. Distribution of C. drummondii in Texas. Fig. 13. Distribution of C. tripartita. Fig. 14. Distribution of C sulfurea C. intc^rifolia O, and C. suhprostrata ®. 1969J I RYXELL CIEM LEGOSIA 215 suture margins. Seeds 5-9 per locule, 2-3 mm, comose; hairs appressed, tan, ca. 5 mm. Chromosome number; ;/ = 10. The holot)pe of Redoiilea tripartita was in the Berhn herbarium and is now apparently lost. Photographs of this plant are in the NY and MO herbaria with the designation \V. 12836 from the W'illdenow colleetion. The number 3601 also appears on this specimen, although it was not numbered by Humboldt and Bonpland. Apparently there were no isotypes, and no syntypes or paratypes WTre cited in the original description b)' Kunth. The situation, therefore, requires the selection of a neotype. One might under the Piules select as neotype either the pliotograph of the lost holotype or a suitably chosen specimen. Since the holotype w^as a poor specimen and accord ijig to Kunth was wormy, so that he had to rely on Bonpland*s notes to some extent in preparing his description, it seems wiser to select a specimen as neotype. The specimen chosen is Weberhaiier 6214 (US- #1496233), which was collected near the type locality and which, in a rather variable species, conforms well to the holotype. Duplicates of this specimen are in F (628149) and GH. Previous authors (Garcke, 1860; Hassler, Repert. Spec. Nov. 7: 380, 1909; Svenson, 1946; and Macbride, loc. cit.) noted an alliance between C. tripartita and C. heterophylla, but only Macbride recognized the alliance nomenclaturally. His choice of Bentham's epithet, cnneata, has priority in varietal rank. The characteristically high degree of variabiHty of leaf form found in sect. Cicjifuegosia is especially well expressed in C. tripartita. The extremes of ex- pression can often be found among the leaves of a single specimen of this species. The extent of this variability is aptly characteri/ed in a collector's note, [Haught 40 (US)], which says, "Leaves extremely variable in form — if found fossil, how many genera might be founded on those of a single specimen!" The taxon is also quite variable in certain other respects, wdiich may be best noted in Hutchison & Wright 5422, which show^s the following atypical charac- teristics. In this specimen the herbage is more pubescent, the petioles shorter, the black glands more prominent and more widely distributed (even to the petals), and the stigma is of a subcapitate form. Tbese arc all deviations in the direction of C. intermedia. Further material may reveal a form that deserves taxonomic recognition. Distribution (Fig. 13): Ecuador: from the Santa Elena peninsula and the vicinity of Guayaquil southward, including the Island of Puna; Peru: provinces of Tumbes and Piura, inland to Cajamarca and Ama/onas. This taxon is appar- ently }iot contiguous with C. heteropJiylla in northern Colombia as has sometimes been implied. Ecuador: guayas: Safios, Asphoul 5661 (US); Giia)'ai]iiil, Sinclair s.n. (K n.v., US-photo). ANCON DiSTR.: Santa Elena Peninsula, Slicppard s.n. (K); Salinas, nr sea level, laiidcman 46 (K); Chanduy, in litore Maris Pacifiei, Spruce 6393 (BAI); Puna, E^^crs 14767 (M), Andersson 135 (S). Peru: piura: Talara, Ihuight 40 (NY, US); near the Amotapcs where rain falls nearly every year, Uau^ht s.n., 1928 (F); Cerro Prieto, 20 mi E of Cape Parinas, 1500 ft alt, Haught 191 (US), tumhks: cerca de Zarumilla, alt 30 m, Fcrreyra 5962 (F, US). CAJAMARCA: Prov. Celendin, westl. Talwand des Maranon iiher Balsas, 2000 m alt, Wchcr- hauer 4262 (G); Prov. Jaen, nr eonfliience of Chinchipe and Maranon, 600-800 m, [Vol. 56 216 ANNALS OF THE MISSOURI BOTANICAL GARDEN Wchcrhaiicr 6214 (F, GH, US), ama/onas: I'rov. Chach.iiioyas, Balsd-s on lliu Maranon and upstream ca 4 km, alt 800 m, II iitchisuu t \Vi-i'j.ht 5422 (L'C, US); VidulSeuege s.n., 1876-77 (P). IN cultivation: (provenance: Playas, Ecuador) F/yxc'// s.ii. (BM, TAES); (prove- nance: Chilete, Peru) Vryxcll s.n. (TAI'S). 5. Sect. Rohusia Fryxcll, sect. nuv. Folia clliptica vcl trilobata. Stipulac nniuitao, lilit'onncs. Involucrum 8-10 phyllum; foliolia iinolucroruni calyccni subac(|uantia. Ncctaria involucrorum ct lolioruiii nulla. Stigmata capitala, rubra. Pctala punctata. Capsulac glabrae intus. Plants upright or decunibent, shrubby or herbaceous, densely pubescent to axillary, solitary (oeeasioucilK' obtuse) (or peduncles some- times articulate). Tinolucral nectaries lacking. Tnvolucel of 8-10 bractlets; bractlets not grouped, prominent, sube(|ual to calyx, cHstuiet, linear-lanceolate to ) Cp e> to acuminate. Petals moderate to (sometimes obscureh so) yellow radii. Stigmas capitate, red. Stxle greatly exceeding androecium, undivided. Fruits glabrous or densely villous without, glabrous within. Type species: The type of sect, liubusta is Cieiifiie^osia afjiiiis (ll.B.K.) Hochr. The name of tlie section is chosen in reference to the larger, shrubbier urowth habit found in the type species of this section. Key to Specjfs oi Sic i ion I^ohlsia a. Leaves ovate to elliptic, entire, short pctiokd (ca '/s-l/lO lamina length). b. Plants densely pubescent except on upper surface of leaf (rarely glabrate); fruit ascending-villous; seed suh-ghihrous to minutely puherulent 14. C. affiuis bb. Plant glabrous; fruit glabrous; seeds hairy 15. C. ciiyahemh aa. Leaves trilobed; petiole > V2 length of lamina 16. C. intermedia 14. Cienfiiegosia affiuis (H.B.K.) Ilochr., Ann. Cons. & Jard. Bot., Geneve 6: 54, 1902. Hibiscus affiuis II.B.K., Nov. Gen. Sp. PI. 5: 289, 1821 [Type: inter Angostura et Trapiehe de D. F. Farreras (Orinoco): lliimholdt & Boupland 1076 (P)]. (This location is near the present Ciudad Bolivar; see Sandwith 1925.) //. sulphurcus H.B.K., loc. cit. [Type: Quebrada de Coteeita, Caracas: Uiimholdt & Boiipland 593 (P)] non Wall ex Voigt, Tlort. Suburb. Calc. 120, 1845. Cieufucgosia sulphurea Hassl., Ostenia 343, 1933 (illegitimate, Article 64) non Garcke. //. sulphurcus var. aeutifolius DC, Prodr. 1 : 45], 1824 [Type: ?]. Fugosia Jauccolatu Juss. in St. Hil., IT Rras. Mer. 1: 253, 1825 [Type: Brazil: St.Uilaire s.n. (P)]. F. affinis Juss. in St.IIil., loc. eit. [Type: Brazil: St.llilairc s.n. (Holotvpe: P, isotvpe: P)]. Hibiscus hilairci Kuntze, llev. Gen. PI. 1 : 69, 1891 (as //. lUlairii). Fugosia phlomidijoUa Juss. in St. Ilil., Fl. Bras. Mer. 1: 253, 1825 [lype: Brazil, in campis propc vicum Chapada in parte pro\i]iciae Minas Geraes dicta Minas Novas: St.'Uilaire s.n. (Holotype: P, isotypes: F, P)]. Cicnfuegosia phlomidifolia Garcke, Bonpl. 8: 150, 1860. Hibiscus phlomidifolia Kuntze, Rev. Gen. PI. I : 69, 1891. Fugosia campcstris Benth. ex Hook., Jour. Bot. 4: 120, 1842 [Type: British Guiana: dry savannahs, Bio Branco: Schomhurgk S38, July 1840 (1839?) (Isotvpes: BM F FT G GH, L-2 sheets, NY, OXF, P (photoMO), US)]. 1969] FRYXELL — CIENFUEGOSIA 217 Hibiscus campestris Kimt7.e, Rev. Gen. PL 1: 69, 1891. Cienfuegosia affinis var. campestris Hochr., Ann. Cons. & Jard. Bot. Geneve 6: 54, 1902. Fugosia guianensis Klotzsch e\ Schomb., Reise Brit. Guiana 3: 1171, 1848 [Type: Auf der Savanne an der Miindung des Pirara, Schomhiirgk s.n. (K-?)]. F,retusa Tiircz., Bull. Soc. Nat. Mosc. 31: 197, 1858 [Type: Venezuela: prov. Cumancnsis, prope Guanaguana, alt 2000 ped, Mar. 1846, Vunck & Schlim 700 Ql^typcs: G, LD, LE, W as photo MO)]. Cienfuegosia phlomidifolia var. humilis Giirkc in Mart., Fl. Bras., 12(3): 575, 1892 [Type: Fiuicli & Schlim 700] (illegitimate, Artiele 63). Cienfuegosia affinis var. humilis Hochr., Ann. Cons. Jard. Bot., Geneve 6: 54, 1902 (illegiti- mate, Article 63). C.riedcUi Giirke in Mart., Fl. Bras. 12(3): 576, 1892 [Type: in campis siccis prope Rio Pardo, Sept 1826, medel 543 (Type: LE)]. Hibiscus rectiflorus Rusby, Mem. N.Y. Bot. Gard. 7: 300, 1927 [Type: Bolivia; near Reyes, 24 Oct 1921, White J 538 (Holotype: NY, isotype US)]. Illustrations: Fig. 21, a-c; also St.TIilairc (op. cit.) t. 50; Schumann (1890) fig. 20 L; Martins (1892) PL 112, fig. 1, PI. 113, fig. 1; Szumkowski (1952) Fig. 1-6; (1953) Fig. 2-6; Fryxcll (1968) Fig. 3,f. Upright shrub, 1 to 6 or more feet high. Stems woody, densely pubescent (though pubescence sometimes absent on lush regrowth and often lost on old woody branches), terete, punctate. Leaves ovate to elliptic, entire, penninerved (veins prominent below), acute, sometimes obtuse, 3-12 cm long, about V2 as broad (rarely narrow^er), rarely ghibrate, generally weakly pubescent to glabrous above and densely to moderately pubescent below (young growth often yellowish- canescent), punctate. Petioles 1-15 nun (approximately 1/10 length of lamina or less, the leaves sometimes subsessile), punctate, usually pubescent. Stipules filiform, pubescent, 3-10 mm, early-caducous. Peduncle axillary, solitary or some- times multiple, unjointed or sometimes articulate, punctate, pubescent, 1-8 cm. Involucel of 8-10 bractlets; bractlets linear lanceolate to sub-spatulate, pubescent, punctate (punctae often obscured by pubescence), 8-22 mm (nearly equaling calyx, persistent. Calyx I-21/2 cm, punctae 1/10 mm diam., often obscured by pubescence, densely to moderately pubescent; lobes 3-ribbed. Petals 2^2-5 cm, yellow^ with maroon spot on chuv ha\ing yellow radii. Androecium maroon TV- throughout, punctate; anther mass obovate. Style pallid, punctate, greatly ex ceeding androecium (sometimes equahng petals); stigma 3-4-lobed, dark-red, pubescent. Fruit 3-4-valved, punctate, 1-1 '/2 cm, o\oid, densely ascending-villous without (hairs 1-2 mm). Seeds turbinate to sub-globose, 2-3 mm, sub-glabrous (covered with very short, sparse, rusty hairs). Embryo cpunctate. Chromosome number: n ^ 10. Vernacular names; In Venezuela C. afji)ns is known as ''algodon de sahana* (cot- ton of the savanna). In Brazil the names "algoddo do ca}}ipo" (field cotton) and ''algodclo hravo" (wild cotton) have been recorded. This species occupies a key position to our understanding of the subg. Cien- fuegosia, It is herein interpreted as the most primitive species on the basis of its morphology. At the same time a remarkably clear transitional series connects this species directly with the more specialized representatives of sect. Cienfuegosia. This plant is remarkably plastic in phenotype when grown in culture. Com- parison of field-grown and greenhouse plants (the latter receiving partial shade and few temperature extremes) indicates that leaf shape, leaf size, petiole length, 218 AiNNALS or THE MISSOURI BOTANICAL GARDEN [Vol. 56 amount of pubescence, and articulation of the peduncle are among the traits affected. Seasonal changes (temperature and/or photoperiod) have similar effects. Much of the considerable \ariation encountered in herbarium collections of this species is probably caused by ecologic (or seasonal) differences and does not rep- resent genetic or taxonomic diversity, although ecot)'pic variation may exist as well. 1 therefore feel justified in presenting so rich a synonymy for a single variable taxon. For example, the long-petioled glabrate phenot\pe that characterizes the holo- 'eolatij (and also Woroiioir 7S28) is similar to the phenotvpe fiic'gusia aff ) ( season, develops the short petioles and dense pubescence that arc typical of C. affuiis. Thus, the type of F. lauceolata can conhdently be assigned to C. affinis on the basis of its sericeous capsule and subglabrous seeds. The type of C. riedelii is an exceptionally tomentose specimen, but appears to differ in no other respect. Perhaps more than one taxon exists within this group. If so, only the com- parative culture of a wide range of material could demonstrate their existence. The flowering response of C. affliiis is under strong photoperiodic control when plants grow under conditions of fluctuating daylength (i.e., at higher latitude). # ''^ N latitude to about 22 S latitude. 0\'er the greater part of this area daylength does not vary significantly through the course of the year. Collation of collection dates of flowering specimens shows that this plant flowers in the field more or less throughout the year. When culti- vated at higher latitudes, however, it flowers only under short-day conditions. At Tempc, /\rizona (33^^' N latitude), for example, it flowers only from November through April and is completely xegetative during the summer months. Distribution (Fig. 11): Venezuela and adjoining parts of Colombia; Brazil: Rio Branco ) Liay and BoHvia). Szumkowski (1953) discussed the ecological and geographical distribution of C. affinis in the Venezuelan part of its range. His map (loc. cit., p. 6) does not indicate the com- plete distribution of the species in Venezuela, but only the locations where he made routine field observations on a seasonal basis. Szumkowski's field observa- tions showed that the plant is widespread and abundant in the chaparral zone of Venezuela, and occurs primarily along the eastern slopes of the Cordillera de Merida (at altitudes up to 1100 m) and northeast to the coast in the vicinity of Caracas and Cumana. It is also abundant in the plains that border the chaparral zone to the southeast (at an altitude of 100-200 m) in the states of Barinas and Portuguesa. The species apparently occurs under sinu'lar (chaparral and savanna) ecological conditions throughout its range. Colombia: Moritz 98 (G, LE); Wa^^oier 350 (UCy Venezuela: Cuman;i, S. Antonio, Flinch IHl (HM, C, LE, P); Cocollar, Fmick & Schlim 700 [G, LD, LE, MO (as photo of W)]. miranda: S slope of La Silla' above Los Palos Grandes (Caracas), ca 1100 m, Dctniis 2278 (K); tliickcts on roadsides, butw Tur- mcro and Ocumare del Tiiy, alt 1000 m, WilliuDis LS5 5 cm); calyx and involucel nearly glabrous; fruit glabrous; petioles 1/10^4 length of lamina 19. C. integrifoUa cc. Leaves orbicular to ovate, 5-nerved; stipules caducous, inconspicuous; pedicels short (1-4 cm); calyx and involucel scabrous; fruit ascending-sericeous; petioles about V2 length of lamina 20. C. siihprostrata 17. Cienfuegosia sulfiirea (Juss.) Garcke, Bonpl 8: 150, 1860 (as C. sulphurea') non (H.B.K.) HassL, Ostcnia 343 (1933 Fugosia sulfiirea Juss. in St.-Hil., Fl. Bras. Mer. 1: 252, 1825 [Type: Uruguay: paturagc seca pres le village de S. Salvador, province Casplatina, St.'Uilairc 241 (Holotype: P, isotype: P)]. Hibiscus Jussieui Kuntze, Rev. Gen. PI. 1: 69, 1891 (based on THgosia sulfiirea Juss.). Cienfuegosia stilphurea var. gcnuina Gijrke in Mart., Fl. Bras. 12(3): 577, 1892 (illegiti- mate, Article 26). C. driimmondii (A. Gray) Lewt. emend. HassL, Ostenia 342, 1933 pro parte (illegitimate, Article 67). C. drummondii var. puhescens HassL, loc. cit. 343. [Tvpes; prope Frav Bentos: Arechavaleta 1524 (MVM? -n.v.), and Sellow 105i (?) syntypes]. Even though the combination C. driimmondii (A. Gray) Lewt. emend. HassL is in contravention of Article 67, in that llassler did not adopt the earlier epithet snlfurca from Jussieu [as Robinson (1895) had done earlier in combining these taxa], nevertheless the varietal epithet puhescens must be accepted as valid under the provisions of Article 68. Illustrations: Fig. 22, e-h; also St.-Hilaire (op. cit.) PI. 49; Rodrigo (1941) Fig. 2; Fryxcll (1968) Fig. 3,k. [Vol. 56 222 AiNNALS OF THE MISSOURI BOTANICAL GARDEN Procumbent perennial herb arising from woody root-stock, canescent on young growth. Stems terete to weakly ridged, densely hirsute (becoming glabrate in age), minutely punctate. Leaves orbicular to reniform, sometimes weakly 3-lobed, dentate to erenate, cordate to truncate, obtuse (or sometimes acute), B-S-nerved, as broad as or broader than long, hirsute below, sparingly hirsute to glabrate above (hairs Vil mm), densely cihate-margined. Petioles pubescent (especially near jiuicture with lamina), punctate, Vz length of lamina or less. Stipules nar- rowly subulate, acute to acuminate, punctate, pubescent below, ghibrous above, ciliate on margin, 1-4 mm. Peduncles pubescent, 1-5 cm, punctate. Tnvolucel of ca 9 bractlets; bractlcts hirsute (especially on margin), lanceolate to spatulate, acute, 5-10 mm long, 1-2 mm broad, 1-nerved, punctate. Calyx 10-15 mm, exceeding involucel, hirsute without, glabrous within, sometimes costulate; lobes broadly ovate, acute, 3-vcined, densely ciliate (hairs Vzl mm) on margins. Petals 1-21/2 cm, yellow (sometimes turning bluc^grecn in sicco), lacking basal petal spot or soinctimes with vestigial spot. Androecium wliite (sometimes reddish in sicco), epunctate; anther mass obovatc; stamciis 5-rauked; anthers yellow. Style greenish-white, sometimes punctate, often slightly 3 4-parted apically; stig- matic lobes 3 to 4, usually free, hght red. Fruit glabrous within and without, minutely punctate, 3-4-\alved, 5-10 mm, globose. Seeds ca 4 mm, turbinate, comosc; seed hairs tan to red-brown, apprcssed, 5-7 mm long. Embryo epunctate. Chromosome number: ii ^^ 10. The original spelling of the specific epithet, salfiirea, was altered by Garcke to sulphitrea, and his orthographic change has been perpetuated by most sub- sequent authors. The original spelling is here restored. imically, (cf. Fryxell, 1963). Distribution (Fig. 14): Argentina: northwestern Cordoba, Santa Fe, Entre Rios, Corrientes, and eastern Chaco; Uruguay: extreme western portion; Para- guay; in Rio Paraguay valley. The trailing stems of this species form carpets over the ground in dry scrub and in moist meadows. It occurs both on heavy clay soils and on sand. In Entre Pios and Chaco it is sympatric with C. drnijiniondii and apparently occurs in nn'xcd populations with it, as is attested to by several mixed collections (Loroitz 142H, Aguilar 117], Krapovichas 7977, SchiiH 169, and Ifiiio & CAos s.ii.) that have been made. In spite of such confusion, however, the two species can be easily distinguished (see accompanying key, p. 221, and that given by Rodrigo, 1941), do not intergrade in nature, and h\bridi/e in culture onlv with difficidtv. The hybrids are lethal at an carlv aye (Palacios & Tiranti, 1966). Another mixed collection from Corrientes (ScJinarz 9949) indi- cates that C. sulfiirea also occurs together with C nhiiifoUa. Paraguay: in arenosis pr. Conccpcion, Ilassler 7315 C^M, G, K, MO, \Y, P, S, UC); San Salvador, Rojns 2592 (M, G); Prov. Pilar, Curuparit/, Schiilz 7978, (TTI ). \rgentina: CHACO: Dpto. Rcsistcncia: C'olonia Bcnitcz, Scliith 769 (BAB-p.p.), 627 (S); Margarita Bclcn, Aiiiiilar 640 (LIL), 7037 (LiL), N7J (LlL-p.p.), Meyer 2653 (LIL); Campo cic Tajamar, Schrocter 37 (BAB). coRRn:x\ ris : Dpto. C^iiruzu Ciiatia : Sjicgazzini s.w. (149?), 2-4 Apr 1940 (BAB), park-like country on licavy clay, Riita 14, ca Km 62S, Pcdcrscu 4698 (BR, C, US), Pcrugorria, Cahnya 10572 (LIL, M), Iharrola 2S56 (LIL), 2263 (BAB, LIL); al norte dc Sauce, Custclhnws 34437 (BA). Dpto. I I 1969] FRYXELL — CIENFUKGOSIA 223 Mbiiriiciiya: Estancia "Santa Maria/' Pcdcrscn 5283 (C, P, S,). Dpto, San Cosmc: Paso de la Patria, Costa Toledo, 5 5 m, Meyer 8792 (LIL). Dpto. Capital: Santa Catalina, Iharrola 944 (LIL), Maharez 1473 (LIL), Ilicken s.n., 26 Mar 1902 (SI), dOrhigny H6 (P), Iharrola 2668 (LIL), Dupre s.n. (P), Corn. Fscuela del Ceiitcnario 78 (BA). Dpto. Mer- cedes: Spegazzini s.n. (69?), 23-31 Mar 1940 (BAB), Justino Solari cercanias, Iharrola 2607 (SI), Camino de la Fcria, Pvodrigo 606 (NY). Dpto. Lnipedrado: Schivarz 9949 (LIL-p.p.), EI Sombrero, Scliuarz 10084 (LIL). Dpto. Monte Caseros: Mocoreta cercanias, Iharrola 2423 (LIL). Dpto. San Martin: en boscjue de Quebracho Blanco, Nandubay 5 km N de Pellegrini, Krapovickas 7956 (LIL), entrc Mercedes y Pellegrini, en monte de Nandubay y espinillos, Krapovickas 7961 (LIL); Bella Vista, Rojas Acosta s.n., Oct 1904 (BAB), santa FE: Dpto. San Cristobal: Estacion Ilcrsilia, Mauicz s.n., 25 Jan 1907 (BAB), BaJcgno 612 (LIL). ENTRE Rios: Dpto. Concordia: Federal, Molfino & Clos s.n., 31 Mar 1927 (BAB-p.p.) L, H. s.n., 15 Feb 1917 (BA), Fedcracion, Crovctto & Piccinini 4701 (BAB), Crovctto & Legiiizamon 4913 (BAB, TAFS). Dpto. La Paz: Lorentz 1428 (UC-p.p.), Sauce de Luna, Krapovickas 7971 (LIL, SI-p.p.); Colon, Meyer 10S95 (LIL), Castellanos 1168 (BA); Gualeguaychu, Jurado 15 his (SI), Garcia s.n., 17 Jan 1943 (LIL); Pehuajo Norte, Serie & Mijoxa 665 (BA); Concepcion del Lruguav, Lorentz 883 (BM, Fl, G, GH, L, LE, M, OXF, P, LC, Z), 57J (UC), 253 (UC), s.n., April 1880 (P), s.n., 3 May 1880 (CORD), Schulz 305 (LIE), Baez 75 (BA); Picada, Mahonlet 164 (BA); Villaguay, Meyer 11273 (LIL), Miisch 995 (GH). cordoha: Bio Tcrcero, Estancia, Stnckert 9679 (CORD, G, S), 10042 (G, S); Rio Primero, Estancia "San I eodoro," Stnckert 11558 (CUP.D, G), 21440 (CORD, G, TAES); San Justo, Sacanta, Stnckert 7081 (G), 9940 (G, S). Uruguay: Entre Belen y Bella Lnion, Hagonere & Piccinini 6554 (BAB, TAES); Paturage scca pres le village de S. Salvador, Provincia Cisplantina, St.-Uilaire 241 (P); Manatial cerca de Mercedes, CastcUanos s.n., 23 Jan 1947 (LIL); ccrca de Bella Union, Castellanos s.n., 28 Mar 1948 (LIE); Nera (Soriano), licrro 1466 (G); Salto, Ostcn 5407 (US); Fray Bentos, Fruchard s.n., 29 Jan 1879 (1899?) (P). IN cuETivATioN : (provenance : Colonia Benite/., Chaco, Argentina) FryxcU 1 5007 (ARIZ, TAES). 18. Cienfiiegosia dnutujiondii (A. Gray) Lc\\t., Rull. Torr. Bot. Club 37: 473, 1910. Fiigosia drummondii A. Gray, PI. Wright. 1: 23, 1852 [Type: Gonzales, Texas: Drnmmond 42, 1834 (Holotype: K,'isotypes: BM, GH, LF, OXF, P)]. Hihiscns drinnmojidii Kuntzc, Rev. Gen. PL 1: 69, 1891 non Turcz. Bull. Soc. Nat. Mose. 31: 195, 1858. Cienfiiegosia sulphnrca Garcke emend. Robins, in Gra) & Robins., Syn. Fl. N.A. 1(1): 337, 1895 pro parte. C. sulphnrca var. drummondii Hoclir., Ann. Cons. Jard. Bot., Gene\e 4: 173, 1900. Elidnrandta tcxana Buckl., Proc. Acad. Sci. Philad. 1861: 450, 1862 [Type: Texas: on the banks of the Colorado River above Austin, Buckley s.n., May-June 1861 (PH)]. Hihiscns pnlveridentus Griseb., Abb. Koniglich. Ges. Wiss. Gott. 24: 49, 18 79 [Type: Argentina: Salta: Lorentz & Uieronwius 1082, i\ov 1873 (IIolot>pe: GOET, iso- type: UC)]. Fugosia pidvcrulenta Hochr., Ann. Cons. Jard. Bot. Geneve 4: 173, 1900. Cienfiiegosia sidpliurea var. glahra Giirke in Mart., Fl. Bras. 12(3): 577, 1892 [Type: Lorentz &■ Hieronymus J 082]. C. sulphurea var. glahra f. intermedia Cbod. 8c Hassl., Bidl. Herb. Boiss., ser. 2, 5: 302, 1905 [Type: Paraguay: in arenosis prope Concepcion, Sep 1901, Hassler 7315h (G)]. C. sidphurea var. major Hassl., Repert. Spec. Nov. 7: 379, 1909 [Types: Paraguay: Gran Chaco, ad ripam occidcntalum flum. Paraguay, latit. S. 22° 2 1' - 23° 30', Oct 1903, Hassler 2416 (Lcctotype: G- Herb. Delessert, isotypes: G, GH, NY, P, S, UC); Rojas 100 (Syntype: G)]. C. drummondii var. genuina Hassl., Ostenia 343, 1933 (illegitimate, Article 26). Fiigosia sidphurea var. trifida Griseb. ex Rodrigo, Darwiniana 5: 220, 1941 nomen nudum. Although the type of Hibiscus piilvendentus Grisb. was cited by Giirke in publishing Cienfiiegosia sulphurea \'ar. glahra Giirke, the latter epitliet cannot be rejected under Article 63 as nomenclaturally superfluous, but must be retained as [Vol. 56 224 ANNALS OF THE MISSOURI BOTANICAL GARDEN legitimate since it is of different rank. The two names are, of course, synonymous, since they are based on the same material. Illustrations: Fig. 22, a-d; also Smith (1895) PL 49; Seckt (1929-1930) PI. 13, Fig. 370; Rodrigo (1941) Fig. 3, C & 1); Fryxell (1963) Fig. 1, a & b, (1968) Fig. 3,j. Perennial herb or subshrub, arising from woody rootstock. Stems semi- decmiibent to ascending, angled, sparsely puberulent to glabrate, minutely punc- tate. Leaves coriaceous, ovate to elliptic, sometimes weakly 3-lobed, coarsely serrate to denticulate (rarely subentire), glabrate (margin not or scarcely ciliate), mi- nutely punctate, about IV2 times as long as broad, truncate, acute. Petioles V2 length of lamina or less, minutely puberulent near juncture with lamina or glabrate throughout, triquetrous, punctate. Stipules sul)ulate or lanceolate, 1-8 mm long, Vzl mm broad, punctate, caducous, glabrate to ciliate-margincd, acute to acuminate. Peduncles sparingly puberulent to glabrate, 2-11 cm, punctate. Involucel of 8-10 bractlets; bractlcts lanceolate to spatulate, 6-14 mm long, 1-nerved, acute to acuminate, punctate, minutely puberulent or glabrate (rarely setose), 1-2 mm broad. Calyx 10-20 mm, exceeding in\olucel, inconspicuously punctate, minutely puberulent or glabrate (rarely setose), sometimes weakly costulate; lobes lanceolate, acute to acuminate, 3-ner\ed, rarely sparingly ciliate- margined, sometimes laciniately dixided into 2 or 3 teeth. Petals 1^2-3 V2 cm, yellow (sometimes turning blue-green in sicco), with solid dark-red basal petal spot, or with petal spot lacking. Androecium cpunctate, w^hitish (or dark-red if petal spot is present); anther mass obovate; stamens 5-ranked; pollen yellow. Style grecnish-w^hite, punctate; stigma 4-5-lobed, dark red. Fruit glabrous within and without, minutely (and sometimes obscurely) punctate, 4-5-valved, obovoid- obtuse (sometimes ovoid-rostrate), 8-14 mm. Seeds angularly turbinate, tan, 3-4 mm; seed hairs tightly appressed (seeds appearing hairless). Embryo cpunctate. Chromosome number: w^ 10. Hochrcutincr (loc. cit.) noted that Grisebach's Hihiscus pnJvendoitns be- longs in Fugosia (i.e., Cicufiiegosia') but did not consider the problem beyond making the new combination. Kearney (1953) independently made the same observation, noting the existence of the UC isotype, but he identified the plant as C. siilphiirea (St.-Hil.) Garcke on the basis of the keys of Rodrigo (1941) and Hutchinson (1947). I have examined this specimen, as well as the holotypc (GOET), and it unquestionably belongs in C. drunnnoidii, not in C. sulphitrea. These two species are not distinguished by Hutchinson, but arc clearly dis- tinguished by Rodrigo. Three vernacular names and three medicinal uses for this species are on record. Seckt (1929-30) records the name "iiicrcnrio" for what is apparently (on the basis of his illustration) this species, although he identifies it as C. sidpJinrea (St.-Hil.) Garcke. On Escobar 13 (13AB) from the province of Chaco the same name is recorded, together with the information that "la medicina lugarena la utiliza como purgante." On a specimen from Colon ia Benitez, Chaco, without indication of collector or date (BAB), the name "yerha del indio" is recorded together with the statement "para llagas gargarismo." On Rojas 13589 and Pier- 1969J FRYXELL — CIENFUEGOSIA 225 rotti 1 the name ^'guaicurii cad" is given together with the indieation that the plant is used as an astringent. Tliis name is evidently in the guarani language (Cadogan, 1957) and may best be translated as 'plant of the Guaieurii Indians," a name comparable to the Spanish "yerha del ifidio/' In none of these cases is the part of the plant used for these medicinal purposes given. This species is polymorphic for petal spot and pigmentation of the androecial column, these two traits simply being two expressions of the same condition. This variation is assumed to have a simple genetic basis and httle or no taxonomic signihcancc. Mixed populations apparently occur, judging from mixed collections that I have seen, e.g., iJudwan A2235 (S). Distinctive features of this species include the tendency to develop divided calyx lobes, which are notable in Ventnri 551 (SI) for example, and the 4-5- lobed stigma and 4-5-valved capsule, that are in contrast to the basically trimerous condition found generally in the remainder of the genus. Distribution (Fig. 12 and 12a): Argentina: Salta, Tucuman, Santiago del Estero, eastern Chaco and Formosa, Corrientes, Santa Fe, Entre Rios, and Cor- doba; central Paraguay; Brazil: southernmost Mato Grosso in the vicinity of the Rio Apa; United States: Texas, in a narrow belt from Brownsville north to Austin, being fairly abundant in the vicinity of Corpus Christi. There appears to be no basis for SmalFs (1933) suggestion that this species may occur on Key W^cst, Florida. In Argentina C. driiunuoiulU grows to some extent in mixed populations with C. snlfurea and also with C. iihiiifoUa. It is found in low, moist areas with heavy soils that are often saline; its salt-tolerance appears to be high. Subsequent to the original description of Fngosia drummondii and Elidurandia texana by Gray and Buckley, respecti\'ely, the history of the Texas occurrence of this plant has received considerable attention (Coulter, 1891; Heller, 1895; Smith, 1895; Lewton, 1910; Hanson, 1921; Hutchinson, 1947), and its notably disjunct distribution has been discussed by Bray (1900), Hutchinson (1947), and Fryxell (1967b). The essential facts may be summarized as follows: Collector Locality Date Collector Locality Date Drummond Gonzales 18 34 Han son Gregory 1919 Buckley near Austin ca. 1861 Schulz Brownsville 1924 Heller Corpus Christi 1894 Eifrig Bishop 1925 Nealley Corpus Christi 1894 Tharp Woodsboro 192 5 Lewton Taft 1910 Lewton S. of Dallas 1925 More recent collections [including several seed collections (Lukcfahr and Martin, 1962) made available to the author by Dr. M. J. Lukcfahr] indicate that the plant occurs commonly, not rarely as reported earlier, in the areas of Browns- ville and Corpus Christi. The extension of its range to the north of Corpus Christi (including both type locations) is more problematical, however, since it has not been collected in this area for over 100 years, if one ignores the unsatis- factory designation of locality on Lewton's 1925 collection. The distributional disjunction shown by C. drummondii was j^robably achieved relatively recently by dispersal from the southern hemisphere to the northern [Vol. 56 226 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Fryxcll, 1967b). The time or means of this dispersal are not immediately evident. IIo\ve\er, eircumstanlial evidence suggests that bird dispersal in recent times is a probable explanation, Crudcn (1966) and Proctor (1968) discuss the plausibility of birds as plant dispersal agents and note that members of the Charadr a formes are likely agents of longdistance dispersal. Proctor emphasizes the capacity of tliese birds to retain seeds in their crops for long periods, especially seeds in the size range shown by C. driimmoiidii, and subsequently to disgorge them intact and thus to disperse them in migration over great distances. Crudcn emphasizes the dispersal potential of seeds adhering in mud to the feet and feathers of migrating birds. This explanation is also plausible for C. dmmmondii, since it occurs in moist habitats, although such a means of dispersal lacks force as a general explanation of disjunction since many of the disjunct plant species occur in dry, upland habitats, which the birds do not frequent and where the conditions are not favorable for the adherence of the seeds to the birds. In any case, the latitudinal similiarity of the two portions of the distribution of C. driun- mondii supports the idea that migrating birds are the responsible dispersal agents. United States: Texas : Gonzales, Brummond 42 (BM, GH, K, LE, OXF, P); banks of the Colorado above Austin, Buckley s.n. (Holotypc of Elidurandia tcxana Buckl.: PII); S of Dallas, Lcwton s.u., 25 Aug 1925 (S). Calhoun County: Port Alto, DrushcII 114 (P). Refugio County: Drushdl 176 (P); Woodsboro, Baysidc, Tharp 3606 (TEX, US). San Patricio County: weed in cotton liclds, Gregory, Hanson s.n., 7 Jun 1919 (GH, MO); Welder Wildlife Refuge near Sinton, Tharp s.n., 9 Oct 19 58 (TEX); roadside, 4 mi NE of Sinton, Tryxell 701 (TAES); Bishop, Eifrii>, 21 (F); slightly saline soil on open bottom, near Sinton, Wolcott £r Barkley 16T416 (F, TEX); 8 mi SW of Taft in swale, Jones 413 (SMU). Nueces County: 1 mi N of Robstown, Chapman 3014 (TEX); nr Gardendalc, S of Corpus Christi, Johnston 53175.14 (TEX). Kleberg County: Agua Camp, Johnston 541181 (SMU, TEX). Corpus Christi: Heller 1808 (GH, PH, S, UC), Nealley s.n., 1894 (F, LIL, MO). Cameron County: "Victoria" soils area, Johnston s.n., Jiuie 1952 (TEX); Brownsville, Schidz 2969 (F); Laguna Atascosa Wildlife Refuge, Fryxell 436 (CTES, TAES), Van Fleet 6095 (TEX); Olmito Rd, near Arroyo Colorado, Clover 1259 (MICH). Willacy County: among mcsquitc and cactus, Shiller M5 (TAES); Sauz Ranch, Johnston 53175.15 (TEX). Brazil: mato grosso: Porto Murtinho, Mahne s.n., 2 Jan 1903 (S), Robert 887 (BM); Porto Pachico, Moore 1056 (BM). Paraguay: Chaco, Puerto Casado, Rojas 1935 (G, M, SI), Greoory et al, 10601 (LIE), Lindman A.2235 (S); Puerto Guarani, campo palmar, Rojas 13589 (G, LIL, MO, TEX); ad ripam occidentalum flum, Paraguay, Hassler 2416 (G, GH, K, MO, NY, P, S, UC), Pride s.n. (K), Ficbrio 1233 (G, K, M, S, Z); en face dc I'Assomplion, Balansas 1875 (P). concepcion: Hassler 73l5h (G), 7317 (BM, G); Pilcomayo River, Moron^ 929 (BM, GH, MO, NY, PH, US), Rojas 100 (G); in regionc locus Yparacav, Cordillera de Altos, Hassler 12390 (C, G, GH, K, L, MO, NY, US, Z); ad ripam rivi Igneri, Hassler 1477 (BM, K, G); von nassem Weisen, Euerto Olympo, Ansits 2163 (S); poor saline soil, foot of Cordillera de Villarica, Jorgensen 3814 (BA, F, GH, LIE, NY, PH, S, SI); Puerte Yatayba, Daguerie 967 (BA), Kuntze s.n., Nov 1892 (NY). Argentina: Formosa: Dpto. Pilcomavo: Ruta 11 Cda. a Eaguna Blanca al km 7, Morel 2192 (LIE, UC), al km 2, Morel 2141 (GH, LIE); Ruta 86 al S. O. a 36 km de Puente Gobernacia, Morel 2756 (EIL, MO); al E dc Puerto Portena a 2 km, Morel 2388 (LIE); Huta 11 Clorinda a Formosa km 4, Morel 1684 (LIE); eamino de Clorinda a Paraiso, Rojas 8975 (SI). Dpto. Pirane: Pilaga, Morel 734 (LIE, US), 33 (GH, LIE), Pierotti 4226 (EIL), 4268 (LIL); Casco Cue, Morel 896 (LIE), NY); Guayacan (20 km al noroeste de Pirane), Pierotti 6 (LIE, SI); Estero Grande, Rcales 63 (LIL); Estancia Lacore, Pierotti 1 (LIL); Laguna de los Rios, Reales 73 (LIL); Monteagudo, Rcales 172 (LIL). Dpto. Clorinda: Curema, Reales 312 (LIL). Dpto. Patino: Las Lomitas FCE, Krapovichas 1293 (LIL), J305 (LIL). Dpto. Formosa: 10 km dc la ciudad, sobrc ruta 11 al N, Krapo- vichas 992 (LIL), Jorgensen 3054 (BA, MO), Kerincs 282 (BAB, TAES). chaco: Dpto. 1969] FRYXELL — CIENFUEGOSIA 22 7 Resistencia: Colonia Bcnitez, Schulz 169 (BAB-p.p.); Margarita Belcn, Aguilar J 263 (LIL), 1171 (LIL-p.p.). Dpto. Florcncia: La Sabana, Basaldua s.n., Oct 1904 (BAB). Dpto. Nopalpi: Las Cuchillas (Lote F), Buratovich 338 (LIL); La Clotildc de P. R. Saenz Pcna, Buratovich 284 (LIL), Baez 12 (BAB); Rio Bcrmejo, Rosario del Dorado, Floss- dorf s.it., 27 Aug 1906 (BAB); Seccic'm 17, Flossdorf s.n,, 28 Nov 1906 (BAB, TAES); Fontana, en suclos arcillo-arenosos, Meyer 2074 (GH, LIL); en campos humedos, Meyer 684 (LIL). ENRIQUE urien: Meyer 4685 (LIL), Schulz 3780 (LIL); Las Palmas, Jorgensen 2316 (BA, GH, LIL, SI, US); Sannihi, Escobar 13 (BAB, TAES); Chaco Central, Asp 89 (SI); Laguna Blanca, Aguilar 63 (LIL). corrientes: Dpto. Monte Caseros: Libcrtad, costa arroya Curuza, Ibarrola 2511 (LIL); Goya, Boelke 1519 (SI), entre rios: Dpto. Feliciano: arroyo Puerto, Casteltanos 1170 (BA); Feliciano, Crovetto & Cherini 6109 (BAB). Dpto. Concordia: Federal, Molfino & Clos s.n., 31 Mar 1927 (BAB-p.p.), L. H. s.n., 15 Feb 1917 (BA). Dpto. La Paz: Alcazar, Krapovickas 7972 (LIL), Saphis 239 (SI); Sauce de Luna, Krapovickas 7971 (SI-p.p.); Al^orrobenwald bei La Paz, Lorentz 1428 (UC-p.p.); Guale- guaychu, Jurado 15 (SI), s.n.. May 1920 (BA); Villaguay, Est. Santa Martha, Mihch 36 (SI); Colonic Hernandarias, Lorentz 1372 (UC); Colonia "Avigdor" Bovril FCE, Krapo- vickas 261 (LIL). santa fe: Dpto. San Cristobal: Arrufo, Balegno 729 (LIL), "La Gua- suncha" cntrc E. Rams y Ilercitia, Krapovickas 743 (LIL). Dpto. Gral. Obligado: Villa Ana, Hayward 01464 (LIL), 0J465 (LIL). Dpto. San Justo: La CrioUa, CasteUanos 19560 (BA); 14 km S de Vera y Pintado, Krapovickas & Cristobal 14325 (CTFS, TAES). Dpto. Vera: Caraquatay, borde de camino, isleta dc sosque, Krapovickas 7818 (LIL). Chaco Santa- fecino: Mocovi, Venturi 190 (SI), Lynch s.n., Nov 1903 (BA); Indio Muerto, Comczaria 2727 (LIL); Videla, Bagonese 2089 (BA-p.p.); Malabrigo FCSF, en el terraplen del FC, Krapovickas 348 (LIL). cordoba: Dpto. Union: Escuela de Bell-Ville, s.n., 24 Apr 1907 (BAB). Dpto. San Justo: Balncario, Balegno 1097 (LIL); Jeanmarie, en las inmediaciones de la Estacion, Hunziker 11367 (CORD, TAES); San Francisco, Borsini 1254 (LIL). San- tiago DEL estero: Dpto. Pellegrini: orilla del salitral, 500 m, Venturi 5678 (GH, LIL, SI, US), tucuman: Dpto. Leales: Chafiar Poso, hab. tcrrcnos salitrosos, 300 m, Venturi 551 (BA, G, LIL, SI, US); Diaz 63 (BAB), salta: Lorentz & Hieronyinus J 082 (GOET, UC). Dpto. Capital: Tiro Federal, Corrca 29 (LIL). IN CULTIVATION : (provcnancc : Argentina) Fryxell 15012 (TAES) ; (provenance : Texas) Fryxell 15006 (ARIZ, TAES). 19. Cienfiiegosia integrifolia (Chod. & Hassl.) Fryxell, comb et stat. nov. C.sulphurea var. integrifolia Chod. & Hassl., Bull. Herb. Boiss. 2 ser., 5: 302, 1905 [Type: in campis pr. Igatimi, Paraguay, 4 Nov 1898, llassler 5444 (Holotype: G-Herb, Boissier, isotypes: BM, G, NY, P)]. C. subprostrata var. vera Hassl., Repert. Spec. Nov. 7: 379, 1909 (illegitimate, Article 63). The same specimen is cited as type of both names cited above. Therefore, the name C. siihprostrata var. vera Hassl. is nomenclaturally superfluous and must be rejected under Article 63. Illustrations: none. Low ascending herb arising from perennial rootstock. Stems glabrous, ob- scurely punctate, weakly angled. Leaves glaucous, minutely punctate, entire, 3(-5)-nerved, orbicular to obovate, truncate to cuneate, obtuse to apiculate. Petiole glabrate, canaliculate, obscurely punctate, 1-5 mm, rarely longer. Stipules narrowly linear, acuminate, often equaling or exceeding the petioles, 1 -nerved, punctate, glabrate, persistent. Peduncles 3 10 cm (rarely to 18 cm) (exceedir leaves), glabrous, minutely punctate, incrassate above. Involucel of ca. 9 bractlets; bractlets often unequal, punctate, glabrous, 1-nerved, linear-lanceolate to weakly spatulate, Vz , 1-2 mm broad (adjacent bractlets sometimes con- ). Calyx, 12-18 mm, equaling or exceeding involu- )rous to sparingly scabrous (on veins); lobes broadly [Vol. 56 228 ANxNALS OF THE MISSOURI BOTANICAL GARDEN ovate, 5-8 mm broad, acute to acuminate, 3-5-ncrvcd. Petals 3Vi to 5 cm long, yellow with dark maroon spot ha\ing yellow radii. Androecium dark maroon throughout; anther mass obovate; anthers 5-ranked (?). Style greatly exceeding androecium ("t.^arly equaling petals), punctate, reddish (?); stigma 3dobed, dark red. Fruit (from immature example) ovoid, epunctate, rostrate, glabrous. Seeds unknown. Distribution (Fig. 14): Paraguay: along the northeastern border in the Sierra de Maracayu and the Sierra de Amambai and southward as far as Caaguazii (at ca. 26° S. Lat.). This species is known ecologically only from the few brief comments made by collectors: *'in campis siccis,'' 'campo colina," and ''in campis combustis." The last comment suggests a similarity in its adaptation to that of C. heteroclada of Nigeria, a similarity that extends in part at least to its vegetati\e morphology. Paraguay: In campis in regionc vicinc Igatimi, Sierra dc Maracavii, Hassler 5444 (BM, G, NY, P); prope Caaguazii, in campis siccis, Hassler 9376 (G, K/NY, UC); Sierra de Amambay, in campis combustis Estrclla, Uassler 10221 (G); campo colina, Rojas 6207 (G). 20. Cieufuegosia stihprostrata Hochr., Ann, Cons. & Jard. Bot. Geneve 6: 57, 1902. [Type: In regione vicine San Estanislao, Sierra de Maracayu, Jan 1899, Hassler 6011 (Holotype: G, isotypes: B (photo in MO), BM, GH, K, MO, NY, P, S, UC)]. Illustrations: Fig. 21, f-g. Stems weak (traihng?), glabrous, ridged, obscurely punctate. Leaves entire, orbicular to ovate-elliptic, obtuse to acute, S-ncrved, obscurely punctate, reticulate, glabrous. Petioles canahculate, punctate, V4 length of lamina or longer (6-12 mm), glabrous or slightly hirsute near juncture with lamina. Stipules incon- spicuous (1-2 mm), filiform, glabrate, caducous. Peduncle 2-41/2 cm, costulate, obscurely punctate, glabrate to sparingly scabrous above. Involucel of 9-10 bractlets; bractlets lanceolate to spatulate, acute, 1-3-nervcd, punctate, sparingly scabrous on margin and midrib (hairs 1 mm), 7-14 mm long, 1-3 mm broad. Calyx 12-17 mm (exceeding involucel), obscurely punctate, costulate, scabrous on veins and margin (hairs 1 mm); lobes 3-nerved, acute, ovate-lanceolate. Petals 3-4 cm, yellow with large maroon spot (with yellow radii). Androecium maroon throughout; anther mass obovate; anthers 5-ranked. Style greatly exceeding an- droecium (nearly equaling petals), punctate, whitish; stigma red, 3-lobed. Fruit 3-celIed, ovoid, rostrate, 10-12 mm, ascending-sericeous. Seeds 4 mm, turbinate, comose; seeds hairs appressed, tan. Distribution (Fig. 14): known only from central Paraguay in the vicinity of San Estanislao. Paraguay: san pedro: In arcnosis, in regionc vicinis San Estanislao, Sierra de Mara- cayii, Hassler 6011 (B, BM, G, GH, K, MO, NY, P. S, UC); Rio Tapiracuay, 8 km dc San Estanislao, camino a Rosario, campo cerca del rio, Krapoxickas et al J 3945 (CTES, TAl'S). 7. Sect. Friesia Fryxell, sect. nov. Ilerbae prostratae. Folia simplicia vcl profunde clissecta. Stipulac auriculatae. Involucrum nullum, Petala epunctata. Capsulac glabrae intus. 1969J FRYXELL — CIENFUEGOSIA 229 Plants prostrate, herbaceous, arising from perennial woody rootstocks, glabrate (but punctae very obscure) ( ) ascending in flower, recurved in fruit. Involucral nectaries lacking. Involucel lack- c ) with or without dark spot on claw; spot when present sometimes having yellow radii. Style single or slightly divided apically. Stigmas capitate. Fruits 3-valved, glabrous within and without. Seeds appressed-hairy. Type species: The type of sect. Friesia is Cienfiiegosia argentina Giirke. The section is named in honor of R. E. Fries who, in describing C. hispida R. E. Fries, noted the alliance of this group of species. Key to Species of Section Friesia a. Leaves weakly lobed or variously parted; leaf margins crenate; tips of calyx lobes often hirsute. b. Leaves nearly simple to moderately parted (sometimes deeply and secondarily so), glabrate; style divided apically, stigmas li<;ht-colorcd 21. C. argentina bb. Leaves weakly tri-lobed, hispid below and on petioles; style nndivided; stig- mas dark-red 22. C. hispida aa. Leaves simple or deeply dissected; leaf margins serrate; tips of calyx lobes gla- brous. c. Leaves simple; stipules prominent (3-11 mm long, 2-6 mm broad); petioles V2 length of lamina or less 23. C. nhnifolia cc. Leaves deeply dissected (or if simple, much reduced); stipules small (2-7 mm long, 1-3 mm broad); petioles % length of lamina or more 24. C. hasslcrana 21. Cienfiiegosia argentina Giirke in Mart., FI. Bras. 12(3): 579, 1892 [Type: Argentina: Salta: La Florida, Rio de Tala, Feb. 1873, Lorentz & Hieronymus 384 (Lectotype: UC; isoleetotypes: MO (photo), BR, G, NY, US)]. Hibiscus argentiniis Kuntzc, Rev. Gen. PI. 3(2): 19, 1898 non Speg. (1901). Fugosia argentina Hochr., Ann. Cons. & Jard. Bot. Geneve 4: 172, 1900. Giirke eitcs Lorentz & Hieronymus s.n, and Lorentz & Hierony^mis 1006, in addition to the leetotype. These speeimens therefore beeome paratypes. Illustrations: Fig. 23, a^e; also Martins (1892) PI. 113, fig. 2; Rodrigo (1948), Fig. 3 (labeled C. argentina var. Hasslerana /. escholtzioides'); Fryxell (1963) Fig. 1, c & d (labeled C. argentina var. hasslerana^; (1968) Fig. 3,1. ( ) c times minutely ciliate-margined, rarely hirsute), truncate, obtuse to acute, 5- nerved, punctate (but punctae detectable only on youngest leaves); lobes rounded, sometimes secondarily divided apically. Petioles canaliculate, i/4-V^ (rarely to %) length of lamina, more or less pubescent near juncture with lamina, otherwise glab- rate (rarely pubescent throughout), minutely punctate. Stipules sessile, auriculate (sometimes falcate), ciliate-margined (hairs ^^ mm), glabrous ventrally, pubes- cent to glabrate dorsally, 2-6 mm long, Vs mm broad, punctate, 1- (sometimes 3-) 230 [Vol. 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN nerved, persistent. Peduncles (especially above) glabratc, minutely punctate, slightly costulatc. Involucel lacking. Calyx 15 ribbed, 10-18 mm, equaling or exceeding fruit, minutely pubescent throughout or glabrate (but usually pubescent apically), scarious intercostally, minutely and sparingly Ml/2 (but ); lobes 3'ribbed, ciliate-margined, generally pubescent at apex (hairs ascending). Flowers sometimes cleistogamic. Petals with undulate margin, 2-3 cm, yellow with small dark-red spot on claw\ Anther mass globose; filaments from pale to dark red; anthers yellow-orange. Style apically 3-parted, greenish, punctate, exceeding androecium; stigmatic lobes white to somewhat red- dish in sicco, pubescent. Fruit epunctate (?), 3-loculcd, 6-12 mm, ovoid. Seeds 3-4 mm, turbinate, comose; seed hairs appressed, brown, 3-5 mm. Embryo epunc- tate. Chromosome number: n -= 10. Distribution (Fig. 15): Argentina: abundant in the provinces of Tucuman and Salta, occurring in adjacent parts of Santiago del Estero and Catamarca, and extending as far southward as Cordoba. It extends northward into Bolivia and northeastward to central Paraguay. Cienfuegosia argentina is a trailing herb that occurs primarily in open fields, apparently preferring sandy soils. Its principal distribution occurs at altitudes of 700-1 500 m, but it probably also occurs at lower altitudes in Paraguay and at least as high as 1 700 m in Bohvia. Bolivia: Gran Chaco, Tatarenda, in campo aprico procumbcns, Fries 1446 (S); Quiru- sillas-Vallcgrande, 1700 m. Troll 929 (B, M). Paraguay: Chaco Paraguayo, [sla Poi, Rojas 7019a (G); Carandayty, Eojas 7179 (G). Fig. 15. Distribution of C. ar^eutina C. hispida O and C. hasslcrana and C. ulnii folia O. Fig. 16. Distribution of 1969] FRYXKLL — CIENFUEGOSIA 231 Argentina: salta: Dpto. Anta: P Seccion, El Dorado, Luna 1037 (LIL); 3' Seccion, Luna s.n., 13 Feb 1948 (LIL); 10 km NE of Lumbrcras, alt 710 m, West 8396 (GH, MO, UC). Dpto. Candelaria: Ruiz de las Llanos, O'Doncll 4715 (LIL), Schrciter 9439 (BA, LIL); en los campos, 1000 m, R. S[chreiter] s.n., 26 Mar 1929 (LIL), Venturi 8761 (GH, US); Qucbrada Sta. Barbara, 1000 m, Schrciter 5808 (F, LIL), O'Donell 4718 (LIL); entre Ruiz dc los Llanos y R. de la Frontcra, O'Donell 5460 (LIL). Dpto. Metan: camino de Horcones a El Porvenir, Carenzo 322 (LIL); Lumbrera, Ruiz 494 (LIL); Metan a Salta, O'Donell 5482 (LIL); entre Metan y R. de los Llanos, Legname & Cuezzo 3035 (LIL). Rosario de la Frontera: Schrciter 1549 (B, BAB, F, LIL, TAES, US), LiUo 4402 (F, G-3 sheets, one of which is labelled 'Trancas, Prov. Tucuman,'* LIL, NY), de la Sofa 86 (LIL). Rio dc Tala: La Florida, Lorcntz & Hieronymus 384 (B, BR, G, MO - as photo, NY - labelled "prov. Catamarca" rather than 'prov. Salta," UC, US); cerca de La Caiiada, 2540 (LIL), Meyer 21185 (LIL); Cormel Moldcs, a orilla del Rio Chanapampa, Hunzikcr 1159 (LIL, NY); camino de Salta a Tucuman, Villalon s.n. (LIL); Joaquim V. Gonzales, Aguilar 269 (LIL); Giiemes, Meyer 8368 (LIL). tucuman: Dpto. Trancas: Rio Tala, Venturi 7370 (US), Rodriguez 12 (G, LIL, US); zwischen Alduralde und Trancas, Lorentz & Hieronymus 1006 (UC); a San Pedro de Colalao, Schrciter 7414 (LIL); Burrucayu, Stuckert 12821 (G), 9J28 (G), J6969 (G, LIL, S), Wall 37 (S), Diaz 3 (BAB, TAES). CORDOBA: Los Mogotes, Sierra Chica de Cordoba, Holuiherg s,n., 6 Feb 187 7 (BA). cata- marca: Dpto. Ancasti; Las Tunas, Spcgazzini s.n,, 14 Feb 1910 (BAB), s,n„ 20 Feb 1910 (BAB). SANTIAGO DEL ESTERO: Dpto. Roblcs: Furena, Mahlonado 241 (NY, S), Arganards 144 (SI); Garza, Krapovickas 10043 (LIL); entre Bcltan y El Simbolar, Krapovickas 10042 (LIL); El Simbolar, Krapovickas 10041 (LIL). IN cultivation: (provenance: Colonia Benitez, Chaco, Argentina) Fryxell 15003 (ARIZ, LE, TAES). 22 . Cienfuegosia hispida R. E. Fries, Kungl. Sv. Vet. Handl. 24: 33, 1947 [Type: Argentina: Salta: orilla de un salitral, Cerro Negro, Antillas, 700 m, 19 Mar 1930, Vefitiiri 10311 (Holotype: S, isotypes: GH, MO, NY, US)]. Illustrations: Fig. 23, Mi; also Fries (op. cit.) PI. 7, fig. 2; Rodrigo (1948) Fig. 1, A-E. Stems procumbent, hirsute (hairs 1 mm) becoming glabratc, apparently epunctatc. Leaves ovate to weakly 3-lobcd, dentate, truncate, obtuse to acute, spar- ingly hirsute becoming glabrate above, hispid below (primarily on veins), cihate- margined, nearly as broad as long, 3'5-nerved. Petioles densely hispid throughout, canaliculate, punctate, V4-V3 (rarely V2 ) length of lamina. Stipules subulate to auriculatc, sessile, hirsute (on vein) to 1-3 mm broad (sometimes larger), per c m ( ), sparsely hispid, obscurely punctate. Involucel lacking ( let is discernible on the holotype). Calyx 15 ribbed, 12-20 mm, densely hispid in bud, moderately so in flower and fruit (especially on veins and margin), apparently epunctate; lobes lanceolate, acuminate, 3-ribbed. Petals 2-3 cm, yellow with large dark-red spot. Androccium dark red; anthers orange (?). Style exceed- ing androecium, whitish; stigma red, pubescent. Fruit 8-10 mm, glabrous (?). Seeds comose; seed hairs tan. Although C. hispida is closely allied to the other species of sect. Friesia and is known only from a relatively few specimens, it appears to be a distinct taxon, distinguished by its vestiture, its large seeds ( ) spot, and its leaf fo Jtion (Fig. 16): Arj (1948) cites and illustrates a specimen ( [Vol. 56 232 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 17. A-C, Cicnfuegosia hildebrandtii Garckc: A, flower [Frvxcll 14007 (TAES)]; B, mature fruit [FryxcU 14001 (TAES)]; C, leaf [Fryxcll 14007 (TAES)]. D E, Cicnfue- gosia gcrrardii (Harv. ex Harv. & Sond.) liochr.: D, bud [Llehenhcrg 3492 (PRE)]; E, leaf [Wood 11722 (P1\E)]. F-G, Cicufucgosia luicroclada Spraguc: F, tip of vegetative shoot [Ahickpor s.n. (GC)]; G, flowering shoot, arising from base of preeeding year's stem [Futi Gr Agyakna s.n. (GC)]. All X 1. 1969] FRYXELL CIKNFUEGOSIA 233 a f Fig. 18. A-C, Cicnfuegosia somaVuma Fryx. [Fryxetl s.n. (TABS)]: A, leaf; B, mature D-G, Cienfuciiosia wdshii (Anders.) Garckc [Fryxell 14006 (TAFS)]: fruit; C, flower. D, leaf; E, mature fruit; F, bud; G, flower. H-K, Cienfucgosia hcarjtii Fryx. [Fryxell 14005 (TAES)]: H, leaf; I, mature fruit; J, seedling with cotyledons shown; K, flower. All X 1. [Vol. 56 234 ANNALS OF THE MISSOUKI BOTANICAL GARDEN Escalante 417, LP -n.v.) from Corrientes. Tlic type is from a saline environment. Argkntina: salta: Antillas, Cerro Negro 700 m, orilla dc un salitral, Ventiiri 10311 (GH, MO, NY, S, US), tucuman: Dpto. Lcales, La Florida, hab. bosquc, 300 m, Monctti 14711 (G, LIL). SANTIAGO DEL ESTERO: Dpto. Lcalcs, Los Puestos, Meyer 12798 (LIL). fuegusia iilmifuU (Ilolotype: BAB)] Illustrations: Fig. 23, i-k; also Rodrigo (1941) Fig. 1, Fig. 3A (both labelled argcntma'); Fryxell (1967a) ) ( to elliptie, serrate, truneate, acute, glabrate (or sparingly hirsute below), some- times sparingly cihate, 5-veined; veins reddishbrown below. Petioles 14-V2 length of lamina, obscurely punctate, glabrate or sparingly puberulent near juncture with lamina, canaliculate, reddish-brown. Stipules sessile, auriculate (rarely falcate), acuminate (rarely toodied), glabrate or weakly ciliate, persistent, 3-11 mm long, 2-6 mm broad, 1 -nerved, sometimes exceeding petioles. Peduncles 1-6 cm, weakly ridged, sparsely pubescent to glabrate, obscurely punctate. Involucel lacking. Calyx 1 5-ribbed, 10-20 mm (exceeding fruit), obscurely punctate, glabrous (or some- times with a few scabrous hairs toward base of calyx tube, but none on apices of calyx lobes); calyx lobes 3-ribbed, lanceolate, acuminate to acute, glabrous. Petals 2-4 cm, yellow with large maroon spot on claw Inning yellow radii. Androecium maroon throughout; anther mass obovoid. Style sparsely punctate, exceeding an- droecium, undivided (or sometimes divided apically in sicco); st pubescent, 3-lobed. Fruit 3-locuIed, epunctate, 8-10 mm, globose to ovoid. Seeds 4-5 mm long, hairy; hairs appressed, tan. Embryo epunctate. Chromosome num- ber: }i = 10. Distribution (Fig. 15): Paraguay: in the valley of the Rio Paraguay from its confluence with the Rio Apa south to the confluence with the Rio Parana; Aroen- tina: Chaco, Corrientes, and Santa Fe in the vallev of the Rio Parana. A sinele specimen from Tucuman extends this distribution to the west. Tliis species occurs on sandy soils, on heavy clay, and frequently in saline situations. Paraguay: boqueron: Puerto Casadn, Estancia "Maria Casilda," Vcdcrscn 4099 (BR, C, G, GH, US), Grciiory ct ah 10604 (LIL); Chaco Paraguayo, OrihiicLa, campo palmas, Rojas 5208 (G); Isla Poi Orihucla, campo ticrra arcnosa, Rojas 7019 (G); San Salvador, dans les argiles impermcablcs, Bahinsu 1875a (P-p.pO; Itapucumi, Rojus 2599 (G, M), Argentina: chaco: Dpto. Pcsistcncia, Colonia Iknitez, Schidz 170 (BAB), 622 (S), 10998 (LIL); Fontana, Meyer 2075 (GH, LIL). corrientes: Dpto. Mburucuya: Estancia Santa Teresa, Pedersen 477 (BR, C, P, S, US), Schuarz 8700 (LIL); Empcdrado, Schuarz 9949 (LIL-p.p.). Dpto. San Luis del Palmar: 25 km E de S. Luis del Palmar, Krapovickas & Crhtohal 11805 (TAES). santa fe: Mocovi, Ocampo, Venturi 190 (BA, BAB, LIL, TAES), Veniuri 190 bis (SI); Videla, Rai!,onese (BA-p.p,); Malabrigo, Castellanos 19563 (BA). tucuman: Diaz 56 (BAB). Cienf 5: 302, 1905 [Type: Para^^uay: in arenosis salsis in rcgione cursus supcrioris (Lectotype: G, isotvpes: B ( MO), BM, GH, MICH, NY, P, S, UC)1 1969] FRYXELL — CIENFUEGOSIA 235 Fig. 19. Cienfucgosia tripartita (H.B.K.) Giirke [Fryxcll s.n. (TAES)]: A-E, leaves, showing some of the variability encountered; F, flower; G, mature fruit. All X 1. 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 1969] FRYXELL — CIENFUEGOSIA 237 J C. argentina var. hassJcrana Hassl., Rcpcrt. Spec. Nov, 7: 381, 1909. C. escholtzioiiies Hochr. ex Chod. & Hassl., Bull. Herb. Boiss., scr. 2, 5: 302, 1905 [Type: Paraguay : in arcnosis salsis in regionc cursus superioris fhiminis Apa, Nov 1901, Hassler 7686a (Lectotype: G, isotypcs: BM, NY, S, UC)]. C. argentina var. hasslcraua L escholtzioides Hassl., Repert. Spec. Nov. 7: 381, 1909. None of the duplicates of Hassler 7686 can be definitely identified as the holotype of C. hassleraua. Only the Geneva and British Museum specimens bear the name C. hassleraua Hochr., and in both cases the handwriting is Ilassler's rather than Hochreutiner's, Unless the holotype is subsequently discovered, the specimen kept at Geneva is here designated as lectotype. The problem of identif)^- ing the holotype of C. escholtzioides is similar^ and the Gene\a specimen of Hassler 7686a is also designated as lectotxpe: Illustrations: Fig. 23, l-o. Stems slender, weakly ridged, obscurely punctate, glabrate. Leaves glabrate, pinnatisect to moderately parted (or sometimes simple, cuneiform, and apicall} dentate on reduced lower leaves). Petiole canaliculate, glabrous or slightly hirsute near juncture with lamina, % length of to equaling lamina, obscurely punctate. Stipules auriculate (sometimes subulate), sessile, glabrous to ciliate, 1 -nerved, minutely punctate, 2-7 mm long, 1-3 nun broad. Peduncles glabrate (or with a few bristles at summit), obscurely punctate, 2-4 Vi em, ridged. Involueel lacking (vestigial bractlets discernible on Balaiisa 1875a and Krapovickas et ah 14249^. Calyx 10-18 mm, 15-ribbed, glabrate or somewhat bristly basally but glabrous apically, scarious intercostal!}, obscurely punctate (sometimes appearing epunc- tate); lobes 3-ribbed, glabrous, narrowly lanceolate, acuminate, sometimes acute). Petals 2-3 Vi cm, yellow with large maroon spot having poorly developed yellow radii on claw. Androccium maroon throughout; anther mass obovoid. Style greatly exceeding androccium (nearly equaling petals), punctate, pallid, undivided (or sometimes sHghtly divided); stigma pubescent, pink to red, 3-lobcd. Fruit epunctate, elliptic, 8-10 mm, 3-loculed. Seeds 3-4 mm, comose; seed hairs ap- pressed, tan. The leaf of C hassleraua is characteristically deeply dissected, but plants pro- duce occasional reduced, undissected leaves. This variability reaches its extreme expression in an isotype of C. hassleraua (^Hassler 7686, UC). Other sheets of this same plant, notably the lectotype (G), show^ the deeply dissected leaf typical of the species. Both types of leaves are found on the same branch in many specimens, such as another isotype (S). Distribution (Fig. 16): Paraguay and southward into the Argentine province of Corrienties. Fig. 20. A-D, Cienfnegomx digitata Cav. [fryxcU J4008 (TAES)]: A, two leaves; B, bud; C, mature fruit; D, flower. E-G, Cicnfiicgosia yucatancmis Millsp. : E, mature fruit [Fryxell s.n, (TAES)]; F, day-old flower [Dunbar 146 (GTI)]; G, three leaves [Dioihar 146 (GH)]. H-K, Cienfuegosia heterophylla (Vent.) Garcke [Miller &r ]ohnston 66 (GH)]: H, bud; EJ^ leaves; K, flower. L-N, Cicnfiiegosia suhteniata (Hassl.) Eryx. [Fiehrig 4022 (GH)]: L, two reduced lower leaves; M, two climax leaves; N, bud. OP, Cie)ifiiegosia rosei Fryx. [Nelson 2779 (GH)]: O, three leaves; P, flower. All X E 238 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 Fig. 21. A-E, Cienfuegosia afjinis (H.B.K.) Hochr. [fryxell 15011 (TAES)]: A. leaf; B, llovvcr; C, mature fruit; D, seedling with cotyledons shown; E, bud. F G, Cienfuegosia suhprostruta Hochr. [Hassler 6011 (GH)]: F, two leaves; G, flower. All X 1. 1969] FRYXELL — CIENFUEGOSIA 239 f Fig. 22. A-D, Cienfuegosia drunimondii (A. Gray) Lewt. : A, flower [Fryxell 15005 (TAES)]; B, mature fruit [Fryxell 436 (TAES)]; C^D, leaves [Fryxell 15005 (TAES)]. E-H, Cienfuegosia sulfurea (St.-Hil.) Garcke [Fryxell 15007 (TAES), except F, Miisch 995 (GH)]: E-F, leaves; G, flower; H, immature fruit. All X 1. The ecological adaptation of this species is apparently similar to that of the other species of this section. A notation on the type, "in arenosis salsis," indicates an affinity for sandy soils and a saline environment. Paraguay: In arenosis salsis in regione cursus superioris fluminis Apa, Hassler 7686 (B, BM, G, GH, MICH, MO, NY, P, S, UC), 7686^/ (BM, G, NY, S, UC); zwischen Rio Apa und Rio Aquidaban, Fiehrig 5000 (G, K, L); alto Paraguay, San Salvador, Rojas 2587 240 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 1969] FRYXELL CIENFUEGOSIA 241 (G, M), Balansa 1875a (P-p.p.); Belen (20 km SE de Concepcion) en campo, Krapovickas et al 14249 (CTES, TAES). Argentina: corrientes: Arroyo Mocoreta, Ccntellanos 34435 (BA, M); Dpto. Mburucuya, Santa Maria, Pedersen 5042 (C, P, S); Paso dc los Librcs, 10 km al NW, Cristobal et al 86 (LIL). Rejected Names and Excluded Species Rejected names: Cienfuegosia affinis var. humilis (Giirke) Hochr. — illegitimate, Article 63. C. Benthamii Hochr. — illegitimate, Article 67. C. Drummondii var. gcniiina Hassl. — illegitimate, Article 26. C. phlomidifolia var. humilis Giirke — illegitimate, Article 63. C. punctata (Cunn. ex Benth.) Domin — illegitimate Articles 64 and 67. C. subprostrata var. vera Hassl. illegitimate, Article 63. C. sulphurea (H.B.K.) Hassl. — illegitimate, Article 64. All combinations with Fugosia Juss. must also be rejected, as discussed on p. 180, because of the priority of Cienfuegosia Cav. Excluded species (including combinations both in Cienfuegosia and in Fugosia'): C. althaeoides Chiov. C. anornahim (Wawr.) Giirke C. australis (F. Muell.) K. Schum. F. australis (F. Muell.) Benth. C. australis var. niicrocarpa Domin. C. Benthamii Hochr. [basionym: F. punctata Cunn. ex Benth.] C. Bricchctti Ulbr. C. cuneiformis (DC.) Hochr. ^ F. cuneiformis (DC.) Benth. C. EUenheckii Giirke C. flaviflora (F. Muell.) Hochr. ^ F. flaviflora F. Muell. (^ Hibiscus ludwigii Eckl. & Zeyh.) (^ Gossypium anomalum Wawr. ex Wawr. & Peyr.) (^ Gossypium australe F. Muell.) (^ Gossypium australe F. Muell.?) (^ Gossypium cunninghamii Tod.) (^ Gossypium somalense (Giirke) Hutch.) (^ Alyogync cuneiformis (DC.) Alef.) (^ Gossypium somalense (Giirke) Hutch.) C. gossypioides (R. Br.) Hochr. C. hakeacfoUa (Giord.) Hochr. F, hakeaefolia (Giord.) Hook. C. hakeaefolia xar, lilacina (Lindl.) Hochr. (^ Thespesia lampas (Cav.) Dalz. ex Dalz. & Gibs. var. thespesioides (R. Br. ex Benth.) Fryx.) (= Gossypium sturtianum J. H. Willis) (^ Alyogyne hakeaefolia (Giord.) Alef.) F. lilacinus (Lindl.) G. Don ex Loud. (= Alyogync hakeaefolia (Giord.) Alef.) C. incana Schwartz C. latifolia (Benth.) Hochr. = F. latifolia Benth. C. Palmeri Rose C. pedata (Bail.) Domin. F. pedata Bail. C. pentaphylla K. Schum. (^ Gossypium incanum (Schwartz) Hillc.) (^ Gossypium costulatum Tod.) (= Gossypium aridum (Rose & Standi.) Skov.) (= Gossypium bickii Prokh.) (^ Gossypium anomalum Wawr, ex Wawr. & Peyr.) Fig. 23. A-E, Cienfuegosia argentina Giirke [Fryxell 15003 (TAES)]: A, seedling with cotyledons shown; B, immature fruit; C, mature fruit; D, flower; E, leaf. F-H, Cienfuegosia hispida R. E. Fries [Venturi 10311 (GH)]: F, flower; G, three leaves; H, mature fruit. I-K, Cienfuegosia ulmifolia Fryx.: I, leaf [Krapovickas & Cristobal 11805 (TAES)]; J, im- mature fruit [Pedersen 4099 (GH)]; K, flower [Krapovickas & Cristobal 11805 (TAES)]. L-O, Cienfuegosia hasslerana Hochr. ex Chod. & Hassl. [Hassler 7686 (GH)]: L, two climax leaves; M, two reduced, lower leaves; N, flower; O, immature fruit. All X L [Vol. 56 242 ANNALS OF THE MISSOURI BOTANICAL GARDEN C.popuUfolia (Benth.) Hochr. F. populifolia Benth. C. pulchella C. A. Gardn. F. pulchella C. A. Gardn. C. punctata (Cunn. ex Benth.) Domin. = F. punctata Cunn. ex Benth. C. robinsonii (F. Muell.) Hochr. F. robinsonii F. Muell. C. so7nalcnsis Giirke C. thcspcsioides (R. Br. ex Benth.) Hochr = F. thespesioides R. Br. ex Benth. C. triphyUa (Harv. ex Harv. & Sond.) K. Scluim. — F. triphyUa Harv. ex Harv. & Sond F. arcysiana Dcflcrs F. Grantii (Mast, ex Oliv.) Hochr. F. hakcaefolia var. coronopifoUa (Miq.) Benth. ( — Gossypium p op uli folium (Benth.) F, MuelL ex Tod.) (^ Gossypiiun pulchellmn (C. A. Gardn.) Fryx.) (=^ Gossypium cunningharnii Tod.) (= Gossypium robinsonii F. Muell.) (^ Gossypium somalense (Giirke) Hutch.) (^^=^ Thespesia lampas (Cav.) Dalz. ex Calz & Gibs. var. thespesioides (R. Br. ex Benth.) Fryx.) ( — Gossypium triphyllum (Harv, ex Harv. & Sond.) Hochr.) (= Gossypium areysianum Deflers) (= Kostelctzkya Grantii (Mast, ev Oliv.) Garcke) (= Alyogync hakcaefolia (Giord.) Alef.) In addition to the preceding rejections and exclusions, a few additional names have appeared in the literature that have apparently never been validly published. These names have no standing under the Rules and must be regarded as nomina nudae. They are: C. Dinteri Ulbr. ex Skov nom. nud. (— Gossypium hcrhaccum var. dinteri Ulbr.?) F. hildebrandti ex Youngman C. cuneata (Benth.) Hutch, nom. nud. F. sulphurea var. trifida Griseb. ex Rodrigo nom. nud. Literature Cited Blank, L. M. & C. R. Leathers. 1963. 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Chromosomes and crossing behavior of Hibiscus cannahinus, H. acetosella, and H. radiatus. Amer. J. Bot. 48: 651-657. Palacios, R. a. & I. N. TiRANTi. 1966. Numero chromosomico de especies argentinas de Cienfuegosia Cav. (Malvaceas). Inst. Nac. Tccn. Agropec. (Argentina) Bol. 40. Pearson, E. O. 1954. The host plants of Diparopsis and Sacadodes. Emp. Cotton Gr. Rev, 31: 283-284. Persoon, C. H. 1806. Synopsis Plantarum. Tiibingen. (Vol. 2). [Vol. 56 244 ANNALS OF THE MISSOURI BOTANICAL GARDEN Phillips, L. L. 1963. The cytogenetics of Gossypium and the origin of New World cottons. Evolution 17: 460-469. Presley, J. T. & C. J. King. 1943. A description of the fungus causing cotton rust, and a preliminary survey of its hosts. Phytopathology 33: 382-389. Proctor, V. W. 1968. Long-distance dispersal of seeds by retention in digestive tract of birds. Science 160: 321-322. Raven, P. H. 1963. Amphitropical relationships in the floras of North and South America. Quart. Rev. Biol. 38: 151-177. Redoute, P. J. 1827. Choix des Plus Belles Fleurs . . . Paris, n.v. Robinson, B. L. 1895. Cienfuegosia. hi: Gray, A., & Robinson, B. L, Synoptical Flora of North America. The Polypetalae. pp. 3 37-3 38. RoDRiGO, A. P. 1941. Las especies argentinas del genero Cienfuegosia, Darvviniana 5: 215-223. . 1948. Addenda a "las especies argentinas del genero Cienfuegosia^ Not. Mus. La Plata 13: 25-33. Sampaio, G. 1926. Exemplos de arilo na flora Brasileira. Bob Mus. Nac. Rio de Janeiro 2(6): 5-34. Sandwith, N. Y. 1925. Humboldt and Bonpland's itinerary in V^enezuela. Kew Bull. 1925: 295-310. SavilEj D. B. O. 1954. The fungi as aids in the taxonomy of flowering plants. Science 120: 583-585. Schumann, K. 1890. Malvaceae. In: Englcr, A. and PrantI, K. Die natiirlichen Pflanzen- familien 3(6): 30-53. Seckt, Hans. 1929-30. Flora Cordobensis. Cordoba, Argentina. Shinners, Lloyd H. 1962. Evolution of the Gray's and Small's manual ranges. Sida 1: 1-31. Skovsted, a. 1935. Chromosome numbers in the Malvaceae. L Jour. Gen. 31: 263-296. . 1944. Some hybridization experiments in the tribe Hibisceae. Comp. Rend. Trav. Lab. Carlsberg, Ser. Physiol. 24: 1-30. Small, J. K. 1933. Manual of the Southeastern Flora. Univ. of N. Carolina Press. Smith, J. G. 189 5. Notes and observations on new or little known species. Missouri Bot. Gard., Ann. Rcpt. 6: 113-120. Spach, E. 1846. Fugosia heterophylla. Hook. Bot. Mag. 72: ^ 42] 8. SvENSON, H. K. 1946. Vegetation of the coast of Ecuador and Peru and its relation to that of the Galagapos Islands. 11. Catalogue of plants. Amer. Jour. Bot. 33: 427-498. SzuMKOwsKi, W. 1952. El "algodon de sabana,'! Cienfuegosia affinis, huesped del "picudo del algodon," Anthonomus grandis in Venezuela. Agron. Trop. 1 : 279-286. . 1953. Ciertfuegosia affinis, malvacea silvestre, hospedera de insectos dafiinos del algodonero en Venezuela. Agron. Trop. 3; 3-12. Ulbrich, E. 1914. tJber einige Malvaceen Gattungen aus der Vervvandtscbaft von Gossy- pium L. Bot. Jahrb. 50(suppl.): 357-362. Ventlnat, E. p. 1800. Description des plantes nouvelles ... J. M. Cels. Whitcomb, W. & L. A. Britton. 1953. The control of cotton boll weevil Anthonomus grandis Boh. in Venezuela. Enip. Cotton Grow. Rev. 30: 177-181, Willdenow, C. L. 1800. Species Plantarum. I.d. 4. Vol. III. Willis, J. C. (revised by H. K. Airy Shaw). 1966. A Dictionary of the Flowering Plants and Ferns. 7th ed. Cambridge Univ. Press. Wilson, F. D. 1968. lOPB Chromosome Number Counts XIX. Taxonl7:577. Wood, J. Medley. 1911. Natal Plants. Bennet & Davis: Durban. Vol. 6, part 3. Youngman, W. 1931. Studies in the cytology of the Hibisceae. II. Ann. Bot. 45: 49-72. Acknowledgments This study would have been impossible without the cooperation of many herbarium curators who lent specimens for study. Grateful thanks for their cooperation is extended to the curators and other personnel of all the herbaria whose specimens are cited herein. Herbarium citations follow the abbreviations given in Index Herbariorum (Regnum Vegetabile, vol. 31, 1964). Thanks is also ow^d several individuals who have supplied viable seed samples of several species of Cienfuegosia, The opportunity to observe living plants is almost a necessity in monographic studies, and the availability of living material enables biosystematic work to be 1969] FRYXELL — CIENFUEGOSIA 245 undertaken. I am therefore especially grateful to Miss M. Gerber, Mr. A. Getahun, Dr. F. Gillfiam, Ing. M. Gutierrez, Mr. A. B. Ilearn, Dr. A. Krapovickas, Dr. C. F. Lewis, Dr. M. J. Lukefahr, Dr. L. L. Phillips, Mr. J. H. Saunders, and Mr. F. Yepez for making seeds avail- able from various parts of the world. Finally, I wish to express a special word of appreciation to Dr. F. D. Wilson, both for a critical commentary on the manuscript and for providing some of the cytological data reported in Table 2. Indkx of Collectors (Type specimens in boldface) Adams 4013 C. hetcroclada & Akpabla 4364 C. hetcroclada Adanson 145A Aguilar 269 640, 1037 63, 1263 1171 Ahiekpor s.n. Andersson 135 Ansits 2163 Arechavaleta 1524 Argaiiaras 144 Asp 89 Asplund 5661 Baez 12 75 Balansa 1875 1875a Balegno 612 — 729, 1097 Bally 9674, 11910 Basaldua s.n. Bebbington 1693 Beccari s.n. Berhaut 1344 Berro 1466 Berti & Escalante 417 Blanchet 2702 Blaumer 9 Blodgett s.n. Boelcke 1519 Boldingh 6383 Boric 257, 551 Borsini 1254 Brade 13440 Braga 82 Breyer 20592 Britton 521 & Britton 211 Buckley s.n. Buratovich 284, 338 Burchell 5620 BurU Davy 10572 Buswell 1387 C, digitata C. argentina C. sulfurca C. drummondii C. sulfurea & C. drummondii (mixed) C. hetcroclada C. tripartita C. drummondii C. sulfurca C, argentina C. drummondii C. tripartita C. drummondii C. sulfurca C. drummondii C. ulmifolia & C. hasslerana (mixed) C. sulfurea C. drummondii C, somaliana C. drummondii C. digitata C. welshii C. digitata C. sulfurea C. hispida C. heteroph} Ila C. heterophylla C. yucatanensis C. drummondii C. heterophylla C. hildebrandtii C. drummondii C. affinis C, hildebrandtii C. digitata C. heterophylla C. heterophylla C. drummondii C. drummondii C. affinis C. hildebrandtii C. yucatanensis Cabrera 10572 Carenzo 322 Castellanos 3443 5 C. sulfurea C. argentina C. hasslerana s.n., 1168, 34437 C. sulfurea C. ulmifolia 19563 1170, 19560 Chapman s.n. 3041 C. drummondii C. yucatanensis C. drummondii C. digitata Chevalier s.n., 24 535 Claussen 374, 549, 590 C. affinis Clover 12 59 Codd 5229 8873 Comezaria 2727 Correa 29 Cristobal, Ahumada & de la Sota 86 C. drummondii C. hildebrandtii C. digitata C. drummondii C. drummondii C. hasslerana Crovetto & Cherini 6109 C. drummondii & Leguizamon 4913 C. sulfurea &Piccinini 4701 C. sulfurea Curran & Haman 1072 C. affinis Curtiss 398 DaCosta 27A Dagueri 967 Dalziel 122 Davey 224, 233 DeCarvalho 219 Deflers s.n., 60 Dennis 2278 DcWinter & Leistner 51 10 C. digitata C. yucatanensis C. affinis C. drummondii C. hetcroclada C. digitata C. hildebrandtii C. welshii C. affinis Diaz 3 56 63 C. argentina C. ulmifolia C. drummondii Dintcr 669, 7215, 74 59 C. digitata drummondii Drummond, T. 42 Drummond, Pi. B. & Hemslcy 2285 C. C. hildebrandtii & Seagrief 5215 C. digitata drummondii Drushell 114, 176 Ducke 2068 Dunbar 146 Dupre s.n. Edwards 3298 Eggers 3 50 13560 14767 Eifrig 21 C. C. affinis C. yucatanensis C. sulfurea C. hildebrandtii C. heterophylla C. affinis C. tripartita C. drummondii [Vol, 56 246 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ekman 15487 Elias 183, 223, 398 Enti & Agyakwa s.n. Escobar 13 Excll& Mendon^^a2694, 2789, 2872 Fcndlcr 105, 106 Fcrrcyra 5962 Ficbrig 1233 4022, 5230 C. yucatancnsis C. affinis C. hetenxlada C. drummondii 5444, 9376, 5000 Flossdorf s.n. Fries 1446 Fruchard s.n. Fryxcll s.n. s.n. C. digitata C. affinis C. tripartita C. drummondii C. subternata C. hasslerana C. drummondii C. argentina C. suifurea C. somaliana C. tripartita 436, 701, 15006, 15012 749 14001, 14007 14005 14006 14008 15003 15004, 15010 C. hctcrophylla C. drummondii C. rosei C. hilticbrandtii C. hcarnii C. wclshii C. digitata C. argentina 15007 15011, 15016 s.n., 15013 & Bates 906 Funck 181 & Schlim 700 Galpin 12193 Garber s.n. Garcia s.n. Gardner 1459 2399 Gaudichaud 44 Gaumer 1269 Gerrard 257, 632 & McKen. s.n. Gillett 4239 Glaziou 18882, 20714 Glover &Gilliland 1206, 1216 Gooding 207 Graham 248 Gregory, Krapovickas & Pietrarelli 10601 10604 C. sull'urea C. affinis C. yucatancnsis C. rosci C. affinis C. affinis C. gerrardii C. yucatancnsis C. suifurea C. affinis C. hcterophylla C. affinis C, yucatancnsis C. gerrardii C. gerrardii C. somaliana C. affinis C. somaliana C, c. vvclshii aflmis 9908 Grosourdy s.n. Guichard'l22, 122A Hageriip 38 5a Hanson s.n. Harbor 6481 Harland s.n. Hasslcr 10899 — — — 8219, 10706 7315 1477, 2416, drummondii tdmifolia affinis affinis * * C. C, c. c. C. hearnn C. digitata C. drummondii C. digitata C. hildebrandtii C. subternata C. affinis C. suifurea 10221 6011 7686, 7686a Haught 4352, 4473 s.n., 40, 191 Haygarth s.n. C. integrifolia C. subprostrata C. hasslcrana C. hcterophylla C. tripartita C. gerrardii Hayward 01464, 01465 C. drummondii Hearn 4 Heller 1808 Hemming 2187 Heudelot s.n. Hicken s.n. Hildebrandt 780a 2325 C. c. c. hcarnii drummondii Hjalmarson s.n. Hochnc 171, 198, 4615 C. affinis vvclshii C. digitata C. suifurea C. welshii C. hildebrandtii C. hcterophylla Holmberg s.n. Honold 20 Hooker & T. s.n. C. argentina C. digitata C. welshii s.n. Humboldt & Bonpland s.n. C. hcterophylla C. tripartita C. affinis C, argentina C. drummondii 593, 1076 Hunziker 1159 11367 Hutchinson 5 C. gerrardii Hutchison & Wright 5422 C. tripartita Ibarrola 944, 2263, 2423, C. suifurea 2556, 2607, 2668 2511 Jeullet 727 Jobert 1037 Johnston s.n., 53175.14, 53175.15, 541181 Jones 413 Jor^enscn 2316, 3054, 3814 Jurado 15 bis s.n., 15 Kassncr 121 Kermes 282 King 394, 1356 Klingberg s.n. Krapovickas 10041, 10042, 10043 C. drummondii C. affinis C. affinis C. drummondii C. drummondii C. drummondii C. suifurea C. drummondii C. gerrardii C. drummondii C. rosci C. digitata C. argentina 261, 348, 743, 992, 1293, 1305, 7818, 7972 7971 C. d rummon dii 7956, 7961 C. suifurea & C. drummondii (mixed) C. suifurea & Cristobal 1I805C. ulmifolia 14325 C. drummondii & Ahumada I 394 5 C. subprostrata 14249 C. hasslcrana Krebs s.n. Kimtzc s.n. Landeman 46 2094 7315b, 7317, 12390 C. drummondii Legagneux s.n. C. hcterophylla C. drummondii C. tripartita C. affinis C. digitata 1969] FRYXELL — CIENFUEGOSIA 247 Legname & Cuczzo 3035 C. argentina Lcmos 127 Leprieur 1 Lewton s.n. Licbenberg 3492 Lillu 4402 Linden 14 56 Lindman A 303 7 A2235 C. hildebrandtii C. digitata C. drummondii C. gcrrardii C. argentina C. affinis C. affinis C. drummondii Lorentz s.n., 253, 571, 883 C. sulfurca 1428 1372 & Hieronymus C. sulfurca & C. drummondii (mixed) C. drummondii 1082 384, 1006 C. drummondii C. argentina C. yucatanensis C. argentina Lundell & Lundcll 8017 C. yucatanensis Lukefahr s.n. Luna s.n., 1037 Lynch s.n. Macedo 372, 4197 Maguirc 2183 Maldonado 241 Malmc s.n. 1666 s.n. Malvarez 1473 C. drummondii C. affinis C. digitata C. argentina C. affinis C. cuyabensis C. drummondii C. sulfurca Martins 1 397, 2680, 2681 C. affinis C. hcarnii Maxwell-Darling 94 Mecuse 9581 Merxmtiller & Geiss 1355, 1549 Meyer 21 Meyer, T. 2653, 8792, C. digitata C. digitata C. digitata C. sulfurca 10595, 11273 684, 2074, 4685 C. drummondii C. ulmifolia 2075 12798 ^ 8368, 21185 Miller & Johnston 66 Mdlspaugh 1693 Miranda 8551 Mocquerys 960 Moldenke 619 Molfmo & Clos s.n. Monetti 14711 Moore 1056 Morel 33, 734, 896, 1684, C. hispida C. argentina C. heterophylla C. yucatanensis C. rosci C. affinis C. yucatanensis C. sulfurca & C. drummondii (mixed) C. hispida C. drummondii 2141, 2192, 2388, 2756C. drummondii Moritz 529, 1038 98 Morong 929 Morton A 1286 Muni s.n. Murdoch 123 Musch 36 995 Myre & Balsinas 1663 C. heterophylla C. affinis C. drummondii C. heteroclada C. hildebrandtii C. sulfurca C. drummondii C. sulfurea C. hildebrandtii Naboulet 164 Neallcy s.n. Nelson 2779 Ncwbould 716 O'Donell 4715, 4718, 5460, 5482 Olufsen 385a d'Orbigny 86 Osten 5407 Parsons s.n, Pedersen 4698, 5283 477, 4099 C. sulfurca C. drummondii C. rosei C. somaliana 5042 C. argentina C. digitata C. sulfurea C. sulfurea C. gerrardii C. sulfurea C. ulmifolia C. hasslerana Pedro & Pedrogao 643-A C. hildebrandtii & Gossypium sp. (mLxed) C. hildebrandtii C. hildebrandtii C. digitata — 663, 2114 Pedrogao 27, 225 Perrottet s.n., 32, 66 Pierotti 1, 6, 4226, 4268 C. drummondii Pilger 289 Pittier 8143 Plee s.n. Pohl 223 C. cuyabensis C. affinis C. heterophylla C. affinis Popov, Tilling & Gilliland 4236, 4277 Pride s.n. Puccioni & Stefanini 617 Rabcn 886 Pagonese 2089 C. hearnii C. drummondii C. chiarugii C, affinis C. ulmifolia & C. drummondii ( d) & Piccinini 6554 C. sulfurea Raunkaicr s.n. Pautanen 19, 401 Reales 63, 73, 172, 289, 312 Rehmann 5223 7128 Richardson 899 Riedel s.n., 543, 2024 Robert 887, 891 Rodrigo 606 Rodriguez 12 Roger 13 Rogers 14 561 12624, 22389 C. heterophylla C. digitata Rojas 2592 C. drummondii C. digitata C. gerrardii C. heterophylla C. affinis C. drummondii C. sulfurea C. argentina C. digitata C. gerrardii C. hildebrandtii C. sulfurca 100, 1935, 8975, 13589 6207 7019a, 7179 C. drummondii C. intcgrifolia C. argentina 2599, 5208, 7019C. ulmifolia 2587 C. hasslerana Rojas Acosta s.n. Rose s.n, Ruiz 494 Ruyssen s.n. Saphis 239 Schenck 82 5 V. d. Schijff 2076 C. sulfurca C. heterophylla C. argentina C. digitata C. drummondii C. hildebrandtii C. gerrardii [Vol. 56 248 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3409, 3938 Schinz 212, 213 Schlechkr 3950 11770 — 4664 Schliebcn 9248 C. hildcbrandtii C. digitata- C. gerrardii C. hildcbrandtii C. digitata C. digitata Schomburkgs.n.,45 5,838 C. affinis Schott 693 Schreitcr s.n., 1549, 5808, 7414, 9439 Schroctcr 37 Schulz, A. 62 7, 3780 169 C. yucatancnsis 7978 C. argcnlina C. sulfurea C. sulfurea C. drummondii C. sulfurea and C. drummondii (mixed) 170, 622, 10998 C. ulmifolia Schulz, C. 305 Schub, E. 2969 Schwarz 9949 8700 10084 Schvveinfurth 75 Sellow 1053 Serie & Mijoya 665 Shafer 634, 686, 2639 C. yucatanensis C. sulfurea C. drummondii C, ulmifolia & C. sulfurea (mixed) C. ulmifolia C. sulfurea C. vvelshii C. sulfurea C. sulfurea Sheppard s.n. ShilJer i\l5 Sieber 58 Silva 144 Simpson 469 Sinclair s.n. Small, Bailey & Matthaus 11599 G. D. Smith 253 H, H. Smith 779 de la Sota 86 Spegazzini 69, 149 C. tripartita C. drummondii C. digitata C. affinis C. yucatanensis C. tripartita s.n. Spruce 6393 Steere 3019 Stephens 1 Stcyermarlv 5 7634 55189 C. yucatanensis C. affinis C. heterophylla C. argcntina C. sulfurea C. argcntina C. tripartita C, yucatanensis C. heterophylla C. heterophylla C. alTinis & Agostini 91287 C. heterophylla St.-Hilaire s.n., 1021 241 Story 6509 C. affinis C. sulfurea C. digitata Strey 4028, 4757 ^ 3493 Stuckert 7081, 9679, 9940, 10042, 11558, 21440 ^ 9128, 12821, 16969 Tatavinow s.n. Tate 99 Taylor s.n. Tharp s.n., 3606 Thesiger s.n. Troll 929 Tyson 1408 Ule 7979 Van Fleet 609 5 Van Landsbcrge 249 Velasco 7 Ventenat s.n. Venturi 5 51, 5678 7370, 8761 10311 190 190 bis Vidal'Senege s.n. Villalon s.n. Volk 125 Wagener 76 _ 350 Wall 3 7 Walter 2/187, 429 Ward 3240 Waring 19 Warming s.n., 442 C, hildcbrandtii C. digitata C sulfurea earnii C. argcntina C. affinis C. affinis C. Jiildebrandtii C. drummondii C. h C. argcntina C. gerrardii C. affinis C. drununondii C. affinis C. affinis C. heterophylla C. drummondii C. argcntina C. hispida C. ulmifolia (see also C. drummondii) C. ulmifolia C. tripartita C. argcntina C. digitata C. heterophylla C. affinis C. argcntina C. digitata C. hildcljrandtii C. welshii C. affinis Wcberbaucr 4262, 6214 C. tripartita Weddell 2641 Wells 2048 Welti 17 West 8396 White 1538 Williams 13585 C. affinis C. hildcbrandtii C. digitata C. argcntina C. affinis 13646 C. affinis Wilms 83, 83a, 83b C. gerrardii Wolcott & Barklcy 16T416 C. drummondii J. M. Wood 239, 11722 C. gerrardii C. hildcbrandtii R. C.Wood s.n., 6828 Woronow 7528 Wulfhorst 21 C. affinis C. digitata 1969] FRYXELL CIENFUEGOSIA 249 Index to Latin Names New taxa arc in boldface, all other taxa are in roman type; numbers in boldface refer to descriptions, numbers in roman refer to synonyms, and numbers with dagger (f) refer to names mentioned incidentally. Abelmoschus esculentus 182t Acacia barbertoncnsis 198t 4. t Alabama argillacca 182 Alyogync cuneiformis 24 If; hakcaefolia '241-2421 Anthonomus grandis ISSf Bombacaceae 179t Bouteloua 182t; aristidoides 182t; barbata 182t Cephalohibiscus 179t, 18 If Cienfuegia 1801, 191 Cienfuegosia 191; subg. Articulata 183t, 187t, 189-191t, 194, 198t; sect. Articu- lata 183t, 187t, 189t, 194; sect. Dioica 183t, 194t, 203, 203t; sect. Garkea 183t, 196; sect. Synodontos {nom. niid.) 196t, 202t; subg. Cienfuegosia 183t, 187t, 189-191t, 204-205, 205t, 217t; sect. Cienfuegosia 183t, 186-187t, 189t, 192t, 20S, 2051, 217t; sect. Friesi 181-184t, 187t, 189-190t, 192t, 204t, 228-229, 2291; sect. Paraguayana 183- 184t, 187t, 189-1901, 220-221, 22It; sect. Robusta 183-184t, 189t, 216, 216t; affinis 181-183t, 189t, 191-192t, 204t, 216-217, 216-218t, var. cam- pestris 217, var humilis 217, 24 If; althaeoides 241; anomalum 241; argen- tina 181-1821, 1871, 190-1911, 229- 230, 229-230t, 234t, var. hasslerana 229t, 237, f. escholtzioides 2291, 237; australis 241, var. microcarpa 241; ben- thamii 241, 24 If; bricchettii 241; chi- arugii 1831, 1871, 189t, 196t, 202; cuneata {nom. nud.) 2421; cuneiformis 241; cuyabcnsis 189t, 219-220, 2201; digitata 183t, 186t, 189-192t, 2051, 206, 206-208t, var. lineariloba 206, 207t; dinteri {nom. nud.) 242t; drum- mondii 182t, 184-1851, 187t, 190- 192t, 221-2221, 223-224, 224-226t, var. genuina 223, 24 If, var. pubescens 221; ellenbeckii 241; escholtzioides 237, 2371; flaviflora 241; gcrrardii 183t, 187t, 189t, 192t, 194-195, 194t, 198t; gossypioides 241; hakcaefolia 241, var. lilacina 241; hasslerana 190t, 234, 237, 237t; hearnii 182-1831, 187t, 189t, 191-192t, 202-203, 203t; heter- oclada 1831, 187t, 189t, 191-192t, 203t, 204, 204t, 228t; heterophylla 182t, 186-187t, 189-1911, 205t, 209- 210t, 211, 212t, 215t, ssp. subternata 212, var. cuneata 213, 215t; hilde- brandtii 181-1831, 189-190t, 192t, 196t, 197, 198t; hispida 182t, 190t, 2291, 231, 231t; incana 241; integri- folia 183t, 190t, 227-228; intermedia 1891, 215t, 220; junciformis 206, var. ruyssenii 206; latifolia 241; palmcri 241; pedata 241; pentaphylla 241; phlomidi- folia 216, var. humilis 21 Of, 217, 24 If; populifolia 242; pulchella 242; punctata 241t, 242; riedelii 217, 218t; robinsonii 242; rosei 1831, 186-1871, 189t, 210, 21 Of, 212t; somalensis 242; somaliana 187t, 189t, 191-1921, 1961, 200-201, 200-201t; subprostrata 189-1901, 228, var. vera 227, 227t, 24 If; subternata 186t, 189t, 205t, 212-213; "sulfurea (Juss.) Garcke 182t, 187t, 190-191t, 216, 221-222, 221-222t, 224-225t, var. drummondii 223, var. genuina 221, var. glabra 22 3, 22 3t, f. intermedia 223, var. integrifolia 227, var. major 223; sulphurea Garcke emend. Robins. 223; sulphurea Hassl. 216, 24 If; thespe- sioides 242; tripartita 1831, 186-187t, 189t, 19M92t, 204t, 2101, 213, 215, 215t; triphylla 242; ulmifolia 1821, 189-190t, 222t, 225t, 234; welshii 183t, 187t, 189t, 19M92t, 199-200, 200-201t; yucatanensis 182-183t, 186- 187t, 189-1911, 205t, 208-209, 209t Cienfugiosa 180t, 191 Cienfugosia 180t, 191 Diparopsis castanea 183t; telephragramma 183t; waters! 183t Elidurandia ISOf, 191; texana ISOf, 223, 225t Fugosia 180t, 191, 224t, 241t; affinis 216; areysiana 242; argcntina 229; aus- tralis 242; campestris 216; cuneata 213; cuneiformis 241; digitata 206, 2081; drummondii 223, 225t; flaviflora 241; gerrardi 195; grantii 242; guianensis 217; hakcaefolia 241, var. coronopifolia 242; heteroclada 204; heterophylla 211; hildebrandti {nom. nud.) 242t; lanceolata 216, 2181"; latifolia 241; lilacinus 241; pedata 241; phlomidifolia 216; populi- folia 242; pulchella 242; pulverulenta 223; punctata Turcz. 211, 212t, 241- 242; punctata Cunn. ex Benth. 21 If; retusa 217; robinsonii 242; *sulfurea 221, var. trifida {nom. nud.) 223, 242t; thespesioides 242; tripartita 213; tri- phylla 242; welschii 199 Gossypieae 179-1811, 187t, 204t Gossypioides 1791, 181 f 250 [Vol. 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN Gossypium 1791, 18M83t, 186-187t, 190t, 192t, 200t, 203t; anomalum 1831, 186t, 241t; 242t; aridii m 241t; arcysianiim australe 198t, 24 If; bickii 24 If; costulatnm 24 If; cunning- hamii 241-242t; hcrbaccnm var. dinteri 242t; incanum 24 If; popiilifolium 242t; pulchellum 242t; robinsonii 242t; so- malensc 208t, 24 l-242t; sturtianum 241t; triphyllum 242t Hampea 179t, ISlf, 187t, 190t, 204t Hibisccae 179t, 18 If Hibiscus IBlf, 187; sect Fiircaria 181t; afFinis 216; acetosclla 182t; argcntinus 229; bifurcatus ISlf; campcstris 217; cavanillesianus 206t; cavanillcsii 206; cuncata 213; drummondii 223; csculcn- tLis 182t; hihiirei 216; jussicul 221; ludwigii 241; phlomidifolia 216; pulvor- ulentiis 223, 223-224t; radiatus 182t; rectiflorus 217; redoutci 211; rubidus 1 8 1 1 ; sulphurous 216, var. acutifolius 216; tripartita 213; welshii 199 Kokia 1791, 181t Kostclctzkya grantii 24 2 f Kydia 18 7t Lebronnccia 1791, 181 f Malvaceae 179t, 187t Napaca 18 7t Puccinia staknianii 182t Rcdoutea 18 Of, 191; hcterophylla 21 If; tripartita ISOf, 213, 2151 Rcdutca 180t, 191; helerophylla 211 Sacadodcs pyralis 183t Thcspesia 179t, 18 If, 190t; lampas var. thcspcsioidcs 241-2421; multibractcata 18 If; rchniannii 195 *The alternate spellings of Jussieu^i epithet, sulfurea and sulphurea, are not separated in this index. THE INITIATION OF VASCULAR CAMBIUM AND PRODUCTION OF SECONDARY XYLEM IN FLOWER BUD PEDICELS OF ASCLEPIAS CURASSAVICA L. IN CULTURE^ BY F. M. Safwat Department of Biology, University of Massaehusetts at Boston I \bstrac:t Flower buds of Asclepias curasaavka were cultured to determine the effects of indole- 3-acetic acid (lAA) and kinetin (6-furfuryIaminopurine) on the secondary growth of the pedicels. Initiation and activity of the cambium were achieved on media containing both of these substances as well as on media containing 1 mg/1 auxin and 10 mg/1 autoclaved deoxyribonucleic acid (DNA). The highest camhial activity and secondary xylem production occurred in pedicels grown on the latter medium. This study suggests that the presence of lAA and kinetin together in the medium enhances the mobility of both, and that there is an interaction between auxin and kinetin in initiating vascular cambium and influenc- ing its subsequent activity. Such an interaction may also take place in the pedicels of fruit of AscJepias curassavica undergoing secondary growth. Introduction The flower pedicels of many plants are herbaceous and consist entirely of primary growth. However, following pollination, fertilization, and the subsequent development of fruit, they very often undergo considerable thickening and become quite woody. Such a change in the habit of the pedicels of some plants has been attributed to the effects of auxins (Nitsch, 1952, 1965). Auxins are known to occur in relatively large quantities in the developing fruits of a number of plants (Nitsch, 1950; Wright, 1956; and others), and it is conceivable that they play a role in the secondary growth of pedicels. This hypothesis is further supported by the fact that auxins induce cambial initiation and xylem differentiation, both in tissue cultures (Camus, 1949; Wetmore and Sorokin, 1955; and others), (Jacobs and Morrow, 1957) (1957) ingly apparent that the processes of organ and tissue differentiation in plants are influeneed by interactions between various growth substances. There is experi- mental evidence, for instance, that kinins interact with auxins in initiating cambial activity in radish roots grown in culture (Loomis and Torrey, 1964; Torrey and Loomis, 1967) and in pea epicotyl segments (Sorokin et al., 1962). Cytokinins are known to occur in relatively large quantities in the developing fruits of a number of plants investigated (Steward and Simmonds, 1954; Goldacre and Bottomley, 1959; Letham and Bollard, 1961; Letham, 1967; Nitsch and Nitsch, 1961), but their effects on the secondary growth of pedicels have not been previously reported. 1 Supported by an institutional grant, C52-67 (1), from the Graduate School of the Uni- versity of Massachusetts at Amherst. Ann. Missouri Box. Card. 56(2): 251-260, 1969. 252 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 I first became interested in this problem while culturing flower buds of Asdepias curassavica. I noticed that the flower bud pedicels showed signs of lateral growth after growing for several weeks on White's basic medium plus 1 mg/1 indole-3-acetic acid (lAA) and 10 mg/1 autoclaved deoxyribonucleic acid (DNA), but flower bud pedicels showed little or no secondary growth when grown on W'liite's medium to which only 1 mg/1 lAA had been added. This ob- servation led me to explore the i>ossible interaction of kinetin C6-furfurylaminopu- rine) and lAA in the initiation of vascular cambin and the secondary growth of the pedicels, since autoclaved DNA, in combination with lAA, is known to produce effects similar to that of kinetin in inducing cell division in tissue cul- tures (Miller et al., 195 5). Materials and Methods Excised young inflorescences of A. curassavica were sterilized for five minutes in a mixture of equal volumes of 70% ethyl alcohol and a saturated aqueous solution of calcium hypochlorite. (By using the mixture instead of the two solu- tions separately, we were able to reduce contamination of the cultures to about 2%.) The inflorescences were then rinsed four times in sterile glass-distilled water, and buds that were three to four nun long with pedicels about one-third of this length were excised. The entire sequence of sterilization and rinsing was performed under a hood and the buds were kept in petri plates placed in an ice bath until they were aseptically transferred onto the appropriate media. The media used in these experiments consisted of the following: L White's basic medium without the added growth substances (White, 1963). 2. White's basic medium and lAA in concentrations of 0.5, 1, and 2 mg/1. 3. 4. tions of 0.5, 1, and 5 mg/1 ( 6-furfurvlaminopurine) lAA at the various concentrations of 0.5, 1, and 2 mg/1 :/l 5. White's basic medium plus 10 mg/1 of DNA and 1 mg/1 lAA. White's basic medium and DNA were autoclaved for 20 minutes under 15 pounds of pressure. Kinetin and lAA were sterilized by Milliporc filtering and then added aspcctically to the media. 1 CI Difco agar was added to all media, and the pi I was adjusted to 5.5. The media were poured into 30 ml Falcon plastic tissue culture bottles. Fight buds were transferred aspcctically onto each of the media, and four replicas were made of each experiment. The bottles were then Fig. 1-5. Fig. 1. A flower bud of A<,clcpias cumssavicn before culturing (X 22). Fig. 2. Wh the swelling of the basal portion of the pedicel (X 22). Fig. 3. A flower bud after growing for four weeks on White's basic medium plus 1.0 mg/1 TAA and 0.5 mg/1 kinetin. Notice the uniform growth of the pedicel. Fig. 4. Transverse section of the pedicel of a young bud before culturing; notice the amphiphloic siphonostele in the center and the poor differentia- tion of the xylem elements (X 150). Fig. 5. Transverse section of a callus showing nests of vascular tissues and the origin of a root (X 140). 1969] SAFWAT — ASCLEPIAS CURASSAVICA PEDICEL CULTURE 253 \ ■rt V*- itf- ■^ ST^ftp:; >^ '-■, *^ 5: I e « .^' « .3 > * - / • ^ *■. =(^- ^ r^ #1^ *4^ w '? Yv^ i 1 *. ■'V *** ► %- Xf *- V- ■^- * * ^ ft-! ^. s> ^ > - -^•^^ <* ^ V ^- ■'■ .*^**». ^-H*^^ ■fifr. *ri-^«t -I* iJiM 3* *' .I-' » ^V .* ,^ 4 ■■*, fr / "i^^ «!* ■^ ■'^. "*?**' **♦'* * / ■ 'rt <'■ Y 4 '"'^j^ ^. I* % A *f ''"4 •- ^ '** * * V t 4. -1& -*»> ^\ »-^ . t ^ < ■ ! 1- >^ . 'J f ^ V u*.' ^ *-■* #'* .■**(' ' '^ ■>. > * ^^***. .a / ^; $ ■■^ W ^^.'^ ^ ^^ m ■jf v^ V ■ ■ J IW^^WBi-^jji- »*^ ^ .■^■. *!■ ^ ?. * > r • -Q -K. %- f .■^ .-^ *- I^^ r^ .^ ^ -of- [Vol. 56 2 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN in a growth PC. and a night temperature of 20±1°C.; the light intensity was set at 500 foot candles for 12 hours daily. After heing kept in culture for periods from two to six weeks, the buds were measured, killed and fixed in formalin-acetic acid-alcohol (FAA) solution. The preserved material was embedded in paraffin, either by the standard alcohol'xylene method, or by the tertiary butyletliyl alcohol procedure. Microtome sections were cut at 10-20/t. Aqueous crystal violet and erythrosin in clove oil produced satisfactory staining that rendered vascular tissues quite distinct. Fast green in 95% ethyl alcohol and 1% aqueous safranin were also used. Observations and Results 1. Anatomy of the Pedicel The pedicels of a young bud of Asclcpias ciirassavica is terete and has an (S ) and corresponds to th (Fia. 4) The vascular cylinder appears dissected and not continuous because certain provascular cells have not been differentiated into xylem elements. The pedicel wood consists primarily of protoxylem elements having relatively poor development of the metaxylem. No vessels are apparent, and the secondary walls of the tra- cheids are either spiral or annular. During the development of the bud, the pedicel elongates; the cessation of this elongation marks the completion of the metaxylem development. Unlike the young stem, the pedicel contains no fibers, 2. Experiiucntal. Buds placed on the White's basic medium did not grow satisfactorily and showed signs of yellowing in approximately two weeks. There was some elongation of the pedicels, but they did not grow^ laterally, nor did diey form noticeable calluses. The cross sections of these pedicels resembled those of intact buds. Pedicels of buds grown in the auxin medium elongated to almost double their initial length. The cut ends of these pedicels began to form calluses within t\vo weeks, and a conspicuous swelling occurred during this period (Fig. 2). The buds were sustained in culture for about four weeks, after which time most began to yellow and some died in culture. Both serial cross and serial longitudinal sections revealed vascular differentiation in the callus. Near the base of the callus there was a random distribution of vascular tissues, and, at a slightly higher level, sev- Fig. 6-9. Portions of transverse sections of pedicels grown on Whitens basic medium plus 0.5 mg/1 kinetin and 1.0 mg/1 lAA, showing the ontogeny of cambial initiation and activity. Fig. 6. The lignification and further differentiation of primary xylem elements (X 130). Fig. 7. The initiation of the vascular cambium near the primary xylem (X 130). Fig. 8. The further development of the vascular cambium and the production of secondary wood (X 400). Fig. 9. The origins of the two vascular cambia as observed in a few speci- (X 150). 1969] SAFWAT 1 VSCLEPIAS CURASSAVICA PEDICEL CULTURE 255 ^-'^'^- ^ ^ w^ ^ ^ ;"**b T ^ ■^. N^ j% '«**, / ^ ^ i. -y. _j" ■1-. r- -v "■v^ ■"¥ ■^^ ■»- j>p «4it . jr.- - V «/ ^ -fr- 3I< ,^' *■ * -^' ^ > ^^,,-^k^*^' J. ^ *T K . 4. V y ^ #*" v^r 'A *- y ■1r. # ?^ i^ ^# V /T>^, #9 i ^ > '^■ 4^ ■< .- j^ ti to « »#-^ ^' ■/ -J> 1 X y^ *^ ^^ -> > s. i» '- -- 'S r ^ •«*^ ;# Jr ■^' r* '«:f ^ ■ ? ^ » **. ' -i'^ # ^ r%r " ^ * [Vol. 56 256 A.\i\ALS OF THE MISSOURI BOTAMC Al. (;AK1)EN eral iicsts of vascular tissues were observed, each nest surrounded by its own meristem. In some pedicels, roots originated from tliese meristems (Fig. 5). y\t a still higher level, in the pedicel proper, cambial initiation was noticed in a very few specimens, but the amount of \ylem produced was very limited. The cambial activity was confined to the lower one third of the pedicel in all the specimens examined. The increase of the concentration of lAA from 0.5 mg/T to 2 mg/1 cHd not change the caml)ial acri\it\ appreciabl). Tlie growth of l)iids in llie kinetin medium was generally poor; slight elonga- tion of the pedicels was noticed, but with no signs of cambial initiation or activity. The buds were maintained on this medium for about three weeks. The auxin-kinctin medium supported the growth of sepals and petals (the gynostegium failed to grow on any of the media), and the buds were maintained in culture for a maximum of six weeks. The pedicels elongated and calluses were produced at their cut ends (Fig. -i). The anatomy of the callus was very similar to that of the callus produced on the auxin medium. Nests of vascular tissues were observed; root development occurred in only a few specimens. 3. Initiutiou and activity of tJw vascular cauihiuni in pedicels groivii on the auxin- kin etin media. Tlie initiation of cambial activity takes place at a level above the callus within the pedicel proper, and gradually progresses toward the opposite end. In none of the buds examined did cambial activity extend over the entire length of the pedicel. The first sign of dilfercMitiation begins with the development of the metaxylem and the strong hgnillcation of the xylem elements (Fig. 6). Then a vascular cambium originates from parenchyma cells between the primary xylem and the phloem. Cambial formation begins outside tlie xylem elements, gradually extend- ing tangcntially so that a complete layer of cambium is finall)' produced (Figs. 7 and 8). From this cambium, secondary tissues are derived in a manner similar to their production in stems. In the periphery of the secondary phloem, patches of hbcrs are seen. The cork cambium originates from a subepidermal layer and is, therefore, considered to be superficial in origin. Lenticels were also observed in a number of specimens. In a few^ pedicels, I encountered a different pattern of cambial activity in w^hich two distinct cambia arose. The first of these originated as described above while the second was deri\'ed frorii the activity of the inner phloem region and produced xylem centrifugally and phloem centripelally (Fig. 9). When two cam- bia arose, only the outer one continued its activity, whereas the inner cambium ceased functioning after an initial production of secondary xylem. In stems liaving Fig. 10-13. Fig. 10. Iransvorse section of a pedicel after growing for five weeks on White's basic niediuni plus 1.0 nig/'l lAA and 10 mg/1 aiitoclaved l)NA; notice the propor- tion of the secondary xylem to the rest of the section (X 110). Fig. 1 1. A portion of l-ig. 10 at a higher magnification ("^ 400). Fig. 12. Longitudinal section of a pedicel showing vessels and tracheids (X 400). Fig. 13. Transverse section of a young stem of Asclepias curassavica: notice the relative amount of secondary xylem compared with tlie rest of the section (X 100). 1969] SAFWAT — ASCTFPIAS CL'RASSAVICA PFDICEI CULTURE 257 V \ ■K --t.! ^ ? "%. >^ Jx-^ •SBtS" ^?, ^^"^"^^^-^m^ -■*^^*v .j'« 1ft C - ^^. |9» ^' -Vr- :%■: '^""v^ 1^ «» I). A ■"^ ^ >. i rj , ■'*^. '--*.. H ^- ■^>^-.». '- >y ■r:ifr>^ i -> ^1P*W. Oil^ r,-. -M a^^PWi ■-x- -ii .ji ^™*^^ V If [Vol. 56 258 ANNALS OF THE MISSOURI BOTANICAL GARDEN amphiphloic steles — such as, tobacco, potatoes, and tomatoes — one cambium normally originates and from the outer phloem alone (Fsau, 1960). In all specimens the secondary wood consists of tracheids and vessels, the latter having simple perforations and measuring 150-200/c in diameter (Fig. 12). Fittings on the secondary walls of the vessels were alternate. In general, cells of tissues in pedicels are larger lluin their counterparts of intact plants. In none of the experiments was the secondary xylcm production as spectacular as in pedicels grown on a medium containing 10 mg/1 autoclavcd DNA and 1 mg/1 lAA. On this medium the pattern of initiation and activity of the cam- bium was similar to that obser\ed in pedicels grown on the auxin-kinetin media, but the amoiuit of secondary wood was most remarkable. In 1 igure 10 notice the proportion of the xylcm to the rest of the pedicels. When this is compared witli a cross section of a young stem of die same plant measuring about 3 mm in diam- eter (Fig. 13), the extent of the eambial activity in the pedicel is quite apparent. Discussion The effects of auxins on the stimulation of eambial activity and xylem dif- ferentiation have been reported by a number of investigators (C ) it seems that auxin alone is not sufficient for the initiation and subsequent activity of the vascular cambium, at least not in bud pedicels grown in vitro. Thus, in this series of experiments, the differentiation of tlie vascular cambium of buds grow^n on media containing only lAA either did not occur at all, or, when present, the secondary xylem formation w^as very limited. This is in sharp contrast to eambial activity in pedicels grown in the kinetin-auxin or the DNA-auxin media. It seems that in buds grown on media containing auxin alone, the auxin tended to be localized in the callus as is indicated by the swelHngs of this region. This is similar to the situation of callus swellings on plants following the appli- cation of auxin (Leopold, 1964). The presence of kinetin or DNA in addition to the auxin seems to cause a more uniform growth of the pedicel. It is ])ossible that kinetin is involved in increasing the mobility of the auxin. Kinetin by itself is poorly translocated in plant tissues (Miller, 1961). But a number of investigators have shown that kinins enhance the mobility of auxins in plants (McCready et al., 1965), and that auxin may, in turn, increase the mobihty of kinetin (Seth et al., 1966). Therefore, it seems probable that auxins and kinens increase the translocation of each other (McCready, 1965). Such an increase in the auxin and kinetin mobility may also be operational in the pedicel. One evidence for this conekision is the swelling of the callus in the absence of kinetin. Further experiments are now^ underwa\ to determine the possible interdependence of tlie translocation of these growth substances in the pedicel. The interaction between kinins and auxins in differentiation has been stressed by Skoog and Miller (1957); and by Torrey and Loomis (1964, 1967) who showed experimentally that kinetin is very important in the initiation and activity 1969] SAFWAT^ASCLEPIAS CURASSAVICA PEDICEL CULTURE 259 of the vascular cambium of excised roots of radish grown in culture. Such an interaction is also reported by Sorokin et al. (1962) in the pea epicotyl segments. There is a strong possibility that kinetin and auxin are required for the initia- tion and activity of the vascular cambium in the pedicels. Kinetin alone did not support the growth of pedicels and lAA without kinetin produced very Hmited cambial activity and in only a few of the buds. Such activation may be a result of the presence of some endogenous kinins in the buds when they were excised. It may be concluded that, in addition to the probable increase in the mobility of the auxin and kinetin, there seems to be an interaction beween the tvvo growth substances leading to cambial initiation and wood production in the pedicel. Al- though 0.5 mg/1 of kinetin and 1 mg/1 of lAA showed "good" cambial activity, no optimum concentrations of these substances have been established for cambial activity in bud pedicels. However, preliminary observations indicate that increas- ing the concentrations of kinetin and lAA to 1 mg/I and 2 mg/1, respectively, seems to increase the cambial activity in bud pedicels. The fact that flower pedicels of intact plants do not show signs of lateral growth is probably because insufficient quantities of these substances are pro- duced. On the other hand, both auxins and cytokinins are known to occur in relatively large quantities in developing fruits of a number of plants. This may also be true in the development of Asdepias fruits. Further experiments will be conducted to explore the possibility of the interaction of kinins and auxins in the secondary growth of fruit pedicels in various species of Asdepias. LlTERATLiRE CiTED Camus, G. 1949. Rccherches sur le role des bourgeons dans les phcnomencs de morphogenesc. Rev. CytoL Biol. Vcgetalcs 11: M99. Davies, C. R., a. K. Seth and P. F. Wareing. 1966. Auxin and kinetin interaction in apical dominance. Science 151: 468. Esau, K. 1960. Anatomy of Seed Plants. Wiley, New York. Goldacre, p. L., and W. Bottomley. 1959. A kinin in apple fruitlets. Nature 184: 555-556. Jacobs, W. P. and I. B. Morrow. 1957. A quantitative study of xylem differentiation in the vegetative shoot apex of Colcus. Amer. Jour. Bot. 44: 823-842. Leopold, A. C. 1964, Plant Growth and Development. McGraw-Hill, New^ York. Letham, D. S. and E. G. Boleard. 1961. Stimulators of cell division in developing fruits. Nature 191: 1119-1120. Letham, D. S, 1967. Chemistry and physiology of kinetin-like compounds. Ann. Rev. Plant Physiol. 18: 349-364. LooMis, R. S. and J. G. Torrey. 1964. Chemical control of vascular cambium initiation. Nat. Acad. Sci. U.S.A. 52: 3-11. Miller, C. O., F. Skoog, M. II. von Saltza, and F. M. Strong. 1955. Kinetin, a cell division factor from deoxyribonucleic acid. Jour. Amer. Chem. Soc. 77: 1392. Miller, CO. 1961. Kinetin and related compounds in plant growth, Ann. Rev. Plant Physiol 12: 395-408. McCready, C. C, D. J. Osborne, and M. K. Black. 1965. Promotion by kinetin of the polar transport of two auxins. Nature 208: 1065-7. NiTSCH, J. P. 1950. Plant and morphogenesis of the strawberry as related to auxin. Amer. Jour. Botany 37: 211-215. NiTSCH, J. P. 1952, Plant hormones in the development of fruits. Quart. Rev. Biol. 27: 33-57. NiTSCH, J. P., and C. Nitsch. 1961. Growth factors in the tomato fruit, pp. 687-705. In R. M. Klein (ed.), Plant Growth Regulation. Iowa State University Press, Ames, Iowa. [Vol. 56 260 ANNALS OF THE MISSOURI BOTANICAL GARDEN Nttsc:h, J. P. 1965. Physiology of flower and fruit development. In "Eneyelopedia of Plant Physiology'^ (W. Ruhland, ed.), Vol. XV/1, pp. 15.^7-1647. Springer," Berlin. Safwat, F. M. 1962. The floral morpliology of Sccuukuic and the evolution of tlie pollina- tion apparatus in Asclep'uulaccac. Ann. Missouri Hot. Card. 19; 95-129. Seth, a. K., CJ, p.. Davies, and P. F. W'arking. 1966. Auxin l^rt'ects on tlie Mohility of Kinetin in the Plant. Science I.">1: 587-588. v Skoog, F., and C. O. iMillek. 1957. Chemical regidaiion of growth and organ formation in plant tissues cultured in vitro. Symp. Soc. F.xptl. Hio. 11: 118 1.^1. SoROKiN, H. P., S. N. Matiiur, and K. V. Ihimann. 1962. The efleets of auxins and kinetin on \yleni differentiation in the pea epicotyl. Anier. Jour. Bot. 49: 444-4 54. Steward, F. C, and N. \V. Simxtomjs. 1954. Growth promoting suhstances in ovary and immature fruit of banana. Nature 173: 1083. Torre Y, J. C, and R. S. Loom is. 1967. Auxin-c\ tokinin control of secondary \ascular tissue formation in isolated roots of Raphinms. Amer. Jour. Bot. 54: 1098 1106. Wetmore, H. H., and S. SoR()KI^. 1955. On difl'erentiation of xylem. Jour. Arnold Aib. 36: 305-317. White, P. I». 1963. The Cultii atio)i of Animal and Vhnit Cells. Second edition. The Ronald Press, New York. Wright, S. T, C. 1956. Studies of fruit development in relation to plant hormones. Illinois Jour. Hort. Sci. 31: 196-211. CYTOLOGICAL AND CHROMATOGRAPHIC EVIDENCE FOR INTERSPECIFIC HYBRIDIZATION IN PETALOSTEMON^ BY Sally A. Walker^ Department of General Biology, Vanderbilt University, Nashville, Tennessee Abstract Chromatographic profiles of fla\onoid extracts of P. purpurcum and P. gattingcri are quite distinct, P. piirpureiun showing 12 species-speciiic compounds, and V. gattingcri show- ing 6 specific compounds. Chromatographic profiles of all putative hybrids except one exhibit compounds specific to each parental species. Three types of chromosomal abnormalities exist in putative hybrids which are not present in either parental species, but completely normal meiosis is reported in several hybrids. In a revision of the genus Petalostemoii Michx., Wemple (1965) reported that interspecific hybridization is possible within taxononiic sections of this genus but not between species of different sections. He created fertile, artfieial hybrids between Petalostemon gattingcri (Heller) Heller and P. purpiiretini (Vent.) Rydb., members of sect. Purpiirei. In nature, these species are usually ecolog- ically isolated. Petalostemon gattingeri is endemic to hmestone glades in Tennessee and Alabama, and P. piirptireton is a prairie species. In the central basin of middle Tennessee, Petalostenioji gattingeri is wide- spread on limestone glades, while only a few individuals of P. purpureiim have been found in the area. Naturally occurring populations of putative hybrids of (1968) County along a roadside and on patches of open limestone. Petalostemon pnrpiireiim was found in adjacent areas which Brecden considered to be prairie relicts. He measured fourteen morphological characters of the two parental species and of putative hybrids and found that the hybrids were generally intermediate between the tw^o parental species or within the range of variation of one of them. The present study was undertaken to supplement morphological data with chromotog- raphic and cytological data. Materials and Methods The following populations of tlie parental species and hybrids WTre studied P. gattingeri Davidson County, Tennessee: Mount View. Walker 195 Davidson County, Tennessee: Couchville Pike. Walker 196. ^ Based on a dissertation submitted to the Graduate School of Vanderbilt University in partial fulfilhnent of the requirements for the degree of Doctor of Philosophy. This work was supported by a National Defense Education Act Predoctoral Fellowship. Dr. William M. Clement, Jr. is gratefully acknowledged for his supervision of this research. 2 Currently Research Associate, Center for the Biology of Natural Systems, Washington University, and Missouri Botanical Gatdcn, St. Louis, Missouri. Ann. Missouri Bot. Card. 56(2): 261-267, 1969. [Vol. 56 262 ANNALS OF THE MISSOURI BOTANICAL GARDEN P. purpureiim Sevier County, Arkansas: Ben Lomond. Demaree 53998. P. pnrpiireiini X gdttingeri Wilson County, Tennessee: Cedar Forest Road. Walker 1 76. Wilson County, Tennessee: Hurrieane Creek Road. Walker 190. Voucher specimens from these populations are deposited in the Vanderbilt University Herbarium (\^DB). Chromatography . Chromatograplne patterns of flavonoid compounds in the two parental species were established from plants collected from pure populations in which there was no evidence of hybridization. Ten plants of each parental species and ten plants from each of the two putative hybrid populations were analyzed. Flavonoid compounds were extracted from stems and leaves in 99% methanol : 1% normal IICl in a ratio of one gram dry weight of plant ma- terial : ten ml. extracting solvent. Twenty drops of extract were spotted on a sheet of Whatman #3 MM chromatographic paper \\]uch was developed in the first direction in a solution of n-butanol: acetic acid: water (6:1:2) for seven hours and in the second dimension in 15% acetic acid. The chromatograms were observed under ultraviolet light both before and after the application of ammonia fumes. The chromatograms were then dipped into a solution of one part 1% potassium ferricyanide and one part 2% ferric chloride, in dilute HCl, and finally in water. The spots were identified by means of die Rf values in two dimensions and color reactions with the treatments described above. Cytology. Inflorescences were fixed in absolute ethanol: glacial acetic acid: chloroform (9:3:1). After fixation, the material was transferred to 70% ethanol and stored at — 15"" C. Pollen mother cells were stained with iron-propionic car- mine. The pachytene stage, as well as later stages of meiosis, was examined in putative hybrids and in the parental species. Results Chromatography. A fist of Rf values and color reactions of compounds found in Petalostemon gattiugeri and P. purpnreum is given in Table 1. Twelve compounds are specific to P. purpnreum and six compounds specific to P. gat- tbigeri. The chromatographic profile of P. purpnreum is shown in Figure lA and that of P. gattiugeri in Figure IB. Willi the exception of one plant, every putative hybrid was found to have some compounds specific to each of the two species. However, no single plant exhibited all of the compounds specific to each species. Tabic 2 shows the number of plants in which each species-specific compound occurred in each population. Cytology. poll of the parental species. The pachytene stage of meiosis is particularly useful for karotype analysis in this genus. Figures 2 and 3 show pachytene cells of Petaloste- mon gattiugeri and P. purpnreum, respectively. The nucleolus and nucleolus organizer are prominent, as well as chromatic and achromatic regions of the seven pairs of chromosomes. No significant differences were found in the karyotypes of the two species (Walker, unpublished). 1969] WALKER INTERSPECIFIC HYBRIDIZATION IN PETALOSTEMON 263 ♦ ^ L CO - # |i %, '■*. -^ « fv ^ \ -V >■ * jl|s^*«R^ « \ * ** % f # V^ #^ ^Mi^ y . ^ii^. f ^♦^ V \ •%.* I ^ s ^ 1^ :^ ::#. rf. ^^ ■-i^JSW ■3* -i' ^ .41 ^k ;« -.*?" t d* * -' R- 5S ■3^ \ *f^ .. * ■*■' .f* # iJ. - ^ X^/ # * ^ ^ ■^ « f r jf^^j^ V »^» F' ^r. ^ At ^ '-- ■'OJ^ ■^^ ^ Fig. 2-5. Pachytene stage of meiosis in pollen mother cells. Fig. 2. Petalostcmon gattiyi- geri,X 868. ¥ig. 3 /p. ptirpiireum, X 1073. Fig. 4. Hybrid, X 1 195. Fig. 5. Hybrid, X 1200. some hybrid dcri^ ativcs this suppression does not oceur because of the particular gene combination present. The third type of abnormahty, sticky hetcrochroniatin, may also be the result of gcnic imbalance. Nielsen (1961) has found this condition in mciotic prophase derivatives of Agroelymns tnrneri, a putative hybrid of Agropyron dasystachyum and Elymus innovatiis. This condition, described by Nielsen as ^^accumulation of pycnotic materials/' produced lethality in the microsporocytes. He concluded that this abnormahty may be the result of an unbalanced en/yme and nucleo- protein condition . Since no structural diifcrenccs in the chromosomes of Petalostemon gattingeri and P. jmrpureiiui were revealed by karyotype analysis, and since meiosis in the F^ hybrids is regular, it is concluded that abnormalities in hybrid derivatives of 266 i VNNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 Spot # 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 Rf, 1 0.74 0.59 0.48 0.39 0.36 0.19 0.31 0.26 0.16 0.21 0.32 0.14 0.02 0.33 0.57 0.83 0.66 0.45 0.11 0.46 0.5 3 0.57 0.46 0.42 0.74 0.41 Table 1 Compounds Found in Parental Species Rf, 2 0.55 0.73 0.41 0.55 0.69 0.07 0.18 0.66 0.79 0.90 0.72 0.65 0.27 0.85 0.88 0.10 0.05 0.02 0.91 0.30 0.39 0.47 0.55 0.74 0.85 0.11 uv white light bhie purple purple purple white yellow purple white white white pink pink bright blue pink light blue white ^vh ite purple light blue purple light blue yellow to pink UV-f MI.. yellow yellow vcllow yellow vcllow Feci, + K,Fe(CN),, P + + + + + + + + + + + + + + + + + + + + + + + G + + + + + + + + + + + P, compound found in PctaJostcmon piirpiircum; G, compoinul found in P. gattingcri. Table 2 Occurrence of Species-Specific Compounds in Putative Hybrid Populations 1. 2. 3. 4. 5. 6. 14. 15. 16. 18. 19. 20. 21. 22. 23. 24. 25. 26. Compound Specific to ]'. imrpurciim + + + + + + + Specific to P. gattingcri + No. of plants in No. of plants in population # 1 with population # 2 with compoimd compound (total: 10 plants) (total: 10 plants) 6 4 8 5 3 8 3 1 6 6 7 2 5 2 6 5 3 4 4 1 8 8 7 10 2 9 10 6 4 6 4 6 4 1 2 1969] WALKER — IiNTERSPECIFIC HYBRIDIZATION IN PETALOSTEMON 267 these two species are the result of genie disharmony and not the result of struc- tural chromosomal differences. The chromatographic data provide evidence for natural hybridization between these two species and support \\ cmple's conclusion that barriers to hybridization arc ecological and not genetic. Anderson (1949) has pointed out that the evolutionary role played by intro- gressive hybridization is the enrichment of variation in the participating species. In considering the significance of hybridization in the two populations studied here, one must take into account that PetalosteiiioJi gattingeri is much more abundant in these areas than P. piirpiireiim since the limestone glade habitat of P. gattingeri is more prevalent in the area than is the typical prairie habitat of P, jnirpureum. Therefore it appears that the consequences of introgression will be an increase in the variability of the P. gattingeri and a gradual disappearance of P, purpureiun individuals from the population. Literature Cited Anderson, E, 1949. Introgressive Hybridization. 109 pp. Jolin Wiley and Sons, New York. Breeden, E. 1968. Ecological tolerance in the seed and seedling stages of two species of Petalostemon (Lcgnminosae). Unpub. Ph.D. Thesis, 198 pp. Vanderbilt University Li- brary, Nashville, Tennessee. Navaschin, M. 1934. Chromosomal alterations caused by hybridization and their bearing upon certain general genetic problems. Cytologia 5: 169-203. Nielsen, E. 1961, Cytology and fertility of derivatives of A^^wclymus tumeric Cytologia 26: 137-154. Wemple, D. K. 1965. Revision of the genus Petalostemon (Leguminosae). UnpubL Ph.D. Thesis, 320 pp. Iowa State University of Science and Technology, Ames, Iowa. i THE GENUS IIOFFMANNIA (RUBIACEAE) IN PANAMA BY John D. D\\ yer St. Louis University and Missouri Botanical Garden, St. Louis, Missouri Abstract Eleven new species of IloffDiuiniia (^Ruhuiccae^ and diagnoses of nine previous!) de- scribed species from Panama are presented. A key to the species is included. The tropical genera Hoffiiunniia S\v., Hamelia Jacq., and Bertiera Aubl. con- stitute the tribe Hamelieae of the Ilnhiaceae, with the first two genera restricted to the tropics of the New World. Paul Standley, in liis treatment of Hoffuiannia in the North American Flora QPvithiales — Riihiaceae (pars) 32(3): 190-208, 1934), recomized 50 species extending from Mexico to Panama, with two species, Sw. and H. tuhiflora Grise (in Willis, A Dictionary of Cambridge Univ. Press, 1966) i7 (loc. cit,, pp. 19M93) Tlora of Costa Rica (Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 131M313, 1938) furnishes the only keys to species of Hoffmannia published in relatively recent times; these are accompanied by specific diagnoses. Hoffmannia is a genus of the rain forest wdth the majority of species occurring at higher altitudes. In Panama, for example, Hoffmannia is generally found at altitudes of 3000 to 6000 ft, although a few species may occur at sea level or at low elevations. Panama seems to represent a good collecting ground for Hoffmannia and is rivaled or surpassed in number of species only by Costa Rica and Guatemala. The fact that eleven new species from Panama are described in this paper suggests the need for an intensification in the collecting of the genus, at least in Central America. Prior to 1959 onlv about 50 collections from Mexico and Central Amer- ica were to be found in the herbarium of the Missouri Botanical Garden, with the majority of these from the Province of Chiriqui in Panama. Not a solitary collection from the Panamanian Pro\'incc of Darien was numbered among these. In the last decade about 50 new Panamanian collections have been added to the herbarium. Particularlv noteworthv are the 1968 collections of Dr. Tames Duke and Mr. Joseph Kirkbride, Jr. from the provinces of Bocas del Toro, Chiriqui, and Darien. Currently, Hoffmannia is known from seven of the nine provinces of the Republic; thus far the genus has not been collected in the provinces of Herrera and Veraguas. The genus Hamelia seems to be the lowland counterpart of Hoffmannia, In Panama Hamelia is rarely seen at high elevations, although the author has seen H. patens Jacq. in the region of Boquete, Prov. Chiriqui, at approximately 5000 ft elevation. Several characters give evidence of the natural relationship of the two genera: imbricate petals, numerous ovules borne on axile placentas, reticulate Ann. Missouri Bot. Card. 56(2): 269-286, 1969. [Vol. 56 270 ANNALS OF THE MISSOURI BOTANICAL GARDEN ( ). Hoff Hamelia: A number of species of Hoffmannia are herbaceous, while Hamelia is consistently suffrutesccnt or arborescent. Such additional features as interspcx:ific variability in floral color in Hoffmannia, or the development of tumescences on the petioles of two species (H. vescicuUfera Standley and H. kirkhridei Dwyer) suggest the greater adaptability of Hoffmannia. Vegetative, Floral and Fruit Characters From approximately 1916 to 1953 Paul Standley pubHshed extensively on the systematics of Hoffmannia, While he exhibited remarkable ability to recognize novelties in the genus, he all too frequently failed to describe the species in pre- cise fashion. His keys to species are not fundamentally sound, as firstly, he utilized few morphological characters in his couplets, and secondly, some characters (e.g. the length of the calycine teeth or the relative length of the corolla lobes to the corolla tube) used in the leading couplets of the keys are extraordinarily difficult to assess in herbarium material. The fact that he rarely utilizes androecial and gynoccial characters in his keys seems to confirm this criticism. a. Vegetative Characters The fact that the stipules are rarely observed in Hoffmannia even in the field, despite the presence of obvious stipular scars, deprives the taxonomist of a pre- sumably important diagnostic character. Leaf shape and petiole length are the most important foliage characters in Hoffmainiia. The leaves of the vast majority of species tend to be sparsely hairy; those which appear to be glabrous usually show under magnification a few hairs on the proximal portion of the costa on the lower side of the blade. In texture the leaves range from membranaceous to sub- coriaceous and do not tend to be succulent except in H. davidsoniae Standley and H. eliasii Dwyer. Frequently coarse hairs (?) matted into a crystalline crust are visible under high magnification in H. kirJd?ridei. The occurrence of a few stellate hairs in this same species is noteworthy in view of the fact that Solereder C^ys- tematische der Dicotyledonen 501, 1899) and Metcalfe and Chalk C^natomy of the Dicotyledons 1: 445. Oxford Univ. Press, 1950) consider the Rnbiaceac to be characterized by unicellular trichomcs. Raphides and cystoliths are abundant in the foliage and should be studied in this genus as well as in Hamelia, The pig- mentation of the leaves of H. ruhripigmenta Dwyer is noteworthy. b. IS! on-vegetative Characters Hoffmannia is characterized by axillary inflorescences, and no exceptions have been noted. Peduncle length, pedicel length, and the relative density of the flowTrs may be important diagnostic features, although at least one well-collected Panamanian species (H. pittieri Standley) shows a disconcerting amount of intra- specific variation. The multiflow^rcd conglomerate inflorescences of H. areolata Standley, H, aeruginosa Standley, and H. vescicuUfera, as well as H. longicalycina Dw^er simulate those of Cephaelis, although those of Hoffmannia are simpler as they lack the complex of bracts and bracteoles subtending tlie flowers of Cephaelis. 1969] DWYER — PANAMANIAN HOFFMANNIA 271 Bud morphology may be a useful taxonomic character; e.g. the buds of H. lew- isiana Dwyer are remarkably clavate at the apex, while those of H. ruhripignienta # specie (H. psychotriaefolia, H. pittieri, and H. woodsonii^ indicate that the colors are constant intraspecifically. Hoffmannia davidsojiiae, on the other hand, seems to have a corolla which varies from yellow to a mixture of yellow and red. In view of Standley*s reluctance to emphasize floral characters in his specific diagnoses, particularly of the androecium and gynoecium, it may be profitable to examine these in detail, at least for the Panamanian representatives. In the case hypanth and abundance of raphides are difficult to assess taxonomically. Intraspecific variation in the length of the calycine lobes of H. lougicalycina Dwyer is obviously diagnostic. No glands were observed alternating with the calycine lobes [as are found e.g. in the Mex- ican H. discolor (Lemaire) Hemsley], except in part of one flower of a collection of H. pittieri Standley {von Hagen & von Uagen 207 S). The ratio of corolla lobe length to tube length is particularly unsatisfactory as a diagnostic character, the lobes tending to reach full size before the tube com- pletes its elongtaion. The corolla tube is usually glabrous within; externally it may be glabrous or pubescent. The external surface should be examined carefully under magnification as the hairs arc frequently very minute. Tlie stamens offer more important characters than perhaps appreciated by previous workers. The connective of the anthers is produced apically in H. trood- souii and basally in H. fimhrianthera Dwyer, While the stamens usually number four in Hoffmannia, H. eliasii usually has three, and some species occasionally have five stamens, e.g. H. leuisiana. The pistil is usually bicarpellate, although the ovary of H. p sy ch at riae folia may have three or very rarely four locules. The pulvinate ovarian disc is less prominent than in Hamelia and is presumably not an important diagnostic fea- ture. The style is elongate, regularly slender, and glabrous. The terminal stigmas are usually connate, may separate naturally at anthcsis (e.g. in H. psychotriae- folia^y or may be teased apart with dissecting needles. In the Panamanian Hoff- mannia at least three basic shapes characterize the stigmas: club-shaped or clavate connate stigmas (most species), linear or narrowly oblong (e.g. H, psychotriaefolia, H. areolata'), and lance-shaped (H. lancistigma Dwyer). Predictably, stigma mor- phology will play a major role in establishing sections within the genus when it is monographed. The fruits of Hoffmannia at the terminal stage of maturity are fleshy, although this stage is rarely encountered in collections, possibly because the fruits are soon deciduous in the field. The most widely distributed species, H. psychotriaefolia, is usually collected in an immature stage of fruit and can be identified immediately by the fact that the fruit is narrowly oblong and is twice as long as wide. The importance of ribbing in the fruits of Hoffmannia is diff'icult to assess as so few fruiting collections are available for study. The seeds usually show very little variation interspecifically, a circumstance also noted in Hamelia. [Vol. 56 272 ANNALS OF THK MISSOURI BOTANICAL GARDEN KlY to Tin; PANA^L\NL\N HoFrNL\NNL\ a. Petioles vcsciculiferous. b. Leaf-blades to 2 5 cm long; petioles to 4 cm long; fruits red .... 19. U. vcscicidifcra bb. Leaf-blades to 40 cm long; petioles to 20 cm long; fruits wbite . . . 10. //. kirkhridei aa. Petioles not \esciculiferoiis, or petioles absent. b. Leaves petiolate. c. Hairs of hypantliiimi to 1.7 mm long; inflorescences usually epedunculate and densely capitulate. d. Small trees; petioles 0.5-0.8 cm long; stigmas clavate 1. //. aeruginosa dd. Herbs; petioles 1-4 cm long; stigmas linear-oblong 3. H. areolata cc. Hairs of hypathium absent or less than 1 mm long; inflorescences usually pedunculate or patulous. c. Fruits (immediately before becoming flesb> ) at least 2X longer than wide; stigmas linear-oblong, spreading, ca Vi the length of the style 16. //. psychotriae folia ee. Fruits less than 2X longer than wide; stigmas if linear-oblong, very short and not patulous. f. Calycine lobes, at least in fruit, to 12 mm long; seeds visible through the wall of the fruit 13, //. longicalycina ff. Calycine lobes 13 mm long; seeds not visible through the wall of the fruit, g. Inflorescences rhizomatous; stamens 3 to 4. h. Leaves oblong-rotund; flowers yellow to yellow-brown or yellow-red; corolla lobes ca 3 X the length of the corolla tube; anthers 4; stigmas clavate-rotund 6. //. davidsoniae hh. Leaves oblong or obovate-elliptic; flowers red; corolla lobes less than 2 X the length of the corolla tube; anthers usually 3; stigmas narrowly oblong 7. H. eliasii gg. Inflorescences not rhizomatous, but axillary in the erect parts of the stem; stamens 4(-5), rarely 6 (often 3 in H. ostaiirca'). i. Anthers with the connective not produced apically or basally. j. Flowers at maturity ca 2 cm long; calycine lobes 4-5(-6); stamens 4-5; stigmas 3-4; ovaries 3-4-lo- culate 12. H. leuisiana jj. Flowers at maturity to 1.5 cm long; calycine lobes reg- ularly 4; stamens 4; stigmas 2-3; ovaries 2-3-loculate. k. Lamina of leaf membranaceous; stamens 3(4) 14. H. ostaurca kk. Lamina of leaf thin-chartaceous to subcoriaceous; stamens 4(-5). I. Stigmas lanceolate 1 1. JL Jancistigma II. Stigmas capitate-rotund to digitiform. m. Leaf blades rubescent, the pigment ex- tractable on boiling; flowers subsessile 17. //. ruhripigmcnta mm. Leaf blades not rubescent, not yielding pigment on boding; flowers obviously pedicellate. n. Leaves oblong-rotund or widely ob- long, exceeding 5 cm in width; caly- cine teeth to 1.2 mm long, o. Leaves chartaceous to subcoriaceous, not membranaceous and translu- cent on drying. p. Leaves widely oblong to ovate- subrotund, to 1 1 cm long, sub- coriaceous, the lateral veins 7-9; inflorescences compressed, the flowers few. ... 5. //. cercidifoUa 1969] DWYEn PANAMANIAN HOFFMANNIA 273 pp. Leaves elliptic or obovate-ellip- tic, 9-3 5 cm long, chartaccous, the lateral veins (11) 15-20; inflorescences usually patulous, the flowers numerous. . 15. //. pittieri 00. Leaves membranaceous and trans- lucent on drying. q. Leaves to 25 cm long; pedicels 0.5 mm wide; calycine lolx^s to 1.1 mm long; corolla tube pubescent; stamens 3(-4) . . . 14. H. ostaurea qq. Leaves 10-16 cm long; pedi- cels ca 0.2 5 mm wide; calycine lobes 0.4-0.8 mm long; corolla tulx' glabrous; stamens 4 . . . 4. H. capillacea ««***ttl •*«««*» nn. Leaves narrowly oblong, usually ca 5 cm wide; calycine teeth 1-2 mm long 2, //. an^ustifolia ii. Anthers with the connective produced apically or basally. r. Stems cane-like; leaves to 9.5 cm wide; anthers with the connective produced basally; flowers purple . . . . 8. H, finibrianthera rr. Stems not cane-like; leaves usually 3 cm wide; anthers with the connective produced apically; flowers yellow 20. //. woodsonii bb. Leaves sessile. s. Leaves rounded toward the apex; calycine lobes 3-4 mm long 9. H. haydenii ss. Leaves acute or acuminate toward the apex; calycine lobes less than 1 mm long 18. H. suhanriculata 1. Hoffmannia aeruginosa Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1313, 1938. Shrubs to 4 m, the branchlets slender, terete, the bark pale-brown, ferru- gineous-villosulosc, the hairs deciduous, leaving a minutely tuberculate surface. Leaves with petioles 5-8 mm long, often slightly curved or reflexcd, densely ferrugineous-villosulose, to 3 mm wide; blades oblong, shortly acuminate at the apex, widely to narrowly cuneate at the base, to 20 cm long and 8 cm wide, stiffly chartaccous, scarcely discolorous, drying brown or green, minutely foveolate, glandular-shiny, the raphides more conspicuous beneath, moderately abundant, white, divergent, to 0.5 mm long, the costa subplane and slender above, prominu- lous beneath, minutely ferrugineous to glabrescent, the secondary veins ca 15, arcuate, prominulous, the intervenal areas often appearing lineolate under mag- nification. Inflorescences disposed in uppermost axils, the cymes with 3-10 flowers on peduncles which are reduced to a thickening (or absent), fcrrugineous- pubescent or purple-villose, the pedicels absent or the flowers subsessilc. Flowers with the hypanthium with elongate soft ferrugineous hairs to 1.7 mm long, the calycine teeth subulate, to 1.5 mm long; corolla densely villose on the outside, the lobes acute; stamens with the anthers ca 3 mm long, not produced at apex and base, acute at the apex; ovary 2-loculate, the stigmas ball-shaped. Fruit not seen. [Vol. 56 274 ANNALS OF THE MISSOURI BOTANICAL GARDEN Costa Rica: Zarccro, 1,500 in, Smith A 86 (F, holotype); Guadalupe de Zarccro, 1,525 m, Smith H 659 (F, MO). Panama: bocas del toro: Fish Creek Mts, vie Chiriqui Lagoon, ion Wcdcl 2289 (i\10). 2, Hoffmannia angustifolia Standley, Contr. U.S. Nat. Herb. 20: 206, 1919. Distribution: Known to occur from Mexico to Panama, Panama: cHiRiyui: Qucbrada Vela, Woodson cr Schery 261 (F). The solitary collection cited for Panama appears to belong to this species. As the narrow-leaved species of Hoffmannia are so poorly known taxonomically I am hesitant about preparing a detailed diagnosis at this time, although numerous collections of the species exist, especially from Costa Rica. The leaves of Woodson & Schery 261 are elongate and very narrow, and are attenuate-acuminate at the apex. The linear-oblong buds have calycine lobes which are deltoid and 1-2 mm long; the hypanthium is deeply ribbed; the ovary is 2-loculate; and the stigmas are connate, forming a clavatc mass. 3. HofJnuuDiia areolata Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 52, 1940. Herbs to 2 m tall; stems smooth, glabrous or villose. Leaves with petioles 1-4 cm long; blade elliptic, attenuate-acuminate at the apex, the acumen to 3 cm long, cuneate at the base, to 25 (?) cm long and 8.5 cm wide, chartaceous, concolorous, the costa immersed above, piano-compressed below, apparently glabrescent above, appressed villosulose on the costa below, distinctly areolate, the raphides numerous, scattered, white, hnear, to 0.3 mm long, the secondary veins to 2 5, arcuate. Inflorescences in the uppermost axils of the leaves, sessile, conglomerate, to 2 cm long or longer, often wider than long. Flowers with the liypanthium oblong- rotund, covered with elongate ferrugineous hairs to 1.7 mm long, the calycine lobes 4, ca 3.5 mm long, obtuse; corolla ca 4.5 mm long, the tube cylindrical, ca 1 mm wide, the lobes shorter than tube; stamens 4, ovate, ca 1.5 mm long, the connective not produced at apex and at base; ovary 2-loculate, style ca 2.5 mm long; stigmas 2, digitiform, spreading, ca 1.5 mm long. Fruits scarlet (fide Davidson 40^, Panama: chiriqui: Boqucte, vie Bajo Chore, 6000 ft, Davidson 40 (F, holotype; GH, MO isotypes); Woodson & Schery 661 (F, GH, MO). The flowers are described as yellow^ or greenish. This species is marked by having leaves with numerous secondary veins, densely conglomerate inflorescences, and hypanthia with elongate ferrugineous hairs. 4. Hoffman}iia capillat*ea Dwyer, spec. nov. Suffnitices, caulibus hgnosis lacvibus glabris, cicatricibus petiolorum modice conspicuis, nodis hie ca 1 cm distantibus. Folia petiohs ad 5 cm longis, ca 0.2 cm latis, Icviter alatis, strictis; lamina oblonga, apice subacuta acuminataque, acumine ad 1 cm longo, basi acuta et vix symmetrica, 10-16 cm longa, 5-7 cm lata, membranacea, discolor, supra opaco-viridior, glabra, costa utrinque subcom- planata infra leviter alata, venis lateralibus ca 10, vix prominulis, venis tertiariis 1969] DWYER — PANAMANIAN HOFFMANNIA 275 prominulis et patulis. Influrescentiae in axillis superioribus dispositae, glabrae, pcdimculo 1-15 mm longo, ca 0.3 mm lato, capillacco, floribus 2-4 in ramulis patulis sacpe subradiate dispositis, pcdicellis 6-8 mm longis, capillaceis, ad 0.4 mm longis, ad 0.4 mm latis, glabris. Floras (hie in gemmis solum vidi) hypanthio ca 3 mm longo, squarroso, glabro, carnoso, lobis 4, vix conspicuis, acutis, ca 0.4C-1?) mm longis, pubcrulis; corolla lobis 4, acutis, glabris; stamina 4, antheris ca 2.5-3 mm longis, conncctivis apicc basique non productis, filamentis brevibus, ca 0.7 mm latis; ovarium 2-loculare, pariete carnoso, stigmatibus oblongo- rotundis, ca 1.5 mm longis, quam stylo latioribus. Fructus non visi. Panama: cocle: Mts N El Valle, 2500-3000 ft, Lewis, MacBrydc, Oliver & Ridguay 1783 (MO, holotypc). This new species is readily distinguished by its membranaceous leaves and dcHcately capillaccous peduncle and pedicels. The flowers are described as white. 5. Hoffmannia cercidifolia Dwyer, spec. nov. Frutices, ramulis gracihbus laevibus glabris. Folia petiolis gracilibus, ad 4.5 cm longis, ca 0.15 cm latis; lamina ovato-subrotunda vel lato-oblonga, apice cuneata, acumine falcato vel recto, ad 1 cm longo, ultime subacute, basi obtusa vel truncata, 7-11 cm longa, 5.5-8 cm lata, in sicco discolor, supra opaco-rubro-fusca, omnino glabrcscens (fortasse in costa vcnisque subtus pilosa) sub lente minute nitido- foveolata, venis laterahbus 7-9, arcuatis, subtus prominuhs. hiflorescentiae non rhizanthae, ad 1 cm longae, floribus paucis, nutantibus, pcdunculo paene nullos, ptdicellis brevibus. Flares (hie gemmas solum vidi) hypanthio brevi campanulato, costato, glabro, cupula calycis brevi inconspicuaque, dentibus 4, triangularibus, ca 1 mm longis, carnosis glabris; corolla tubo brevi, ca 2.5 mm longo, subcarnoso intus glabro cxtus pubescenti, lobis 4, oblongis, ca 4 mm longis; stamina 4, an- theris ca 3.5 mm longis, ca 0.8 mm diam, conncctivis apice basique non productis, fdamentis complanatis; ovarium 2-loculare, stylo ca 5.5 mm longo, stigmatibus 2, in capitulum clavato-rotundum dispositis, ovulis multis. Fructus non visi. Panama: DAKitw: Cloud forest, Ccrro Campamcnto (S of Cerro Pirre), Duke 15597 CMO, holotype). This new species is readily distinguished by its ovatc-subrotund leaves with very few lateral veins. The general appearance of the leaves is suggestive of the redbud, Cercis canadensis L., so common in much of the United States. 6. Hoff 53, 1946. Herbs to 1.5 m; stems terete, rigid-carnose, smooth, glabrous, usually drying tan, the internodes to 9 cm apart. Leaves with petioles 2-7 cm long, slender and somewhat wiry, 1-2 mm wide, puberulent; lamina oblong-rotund, rounded and acuminate at the apex, the acumen 5 18 mm long, rounded or tapering widely, then abruptly cuncate at the base, 8-17 cm long, 4-12 cm wide, thinly and stiffly succulent, smooth, lustrous, brown, minutely white-puberulent to glabrous above, minutely puberulent on veins beneath, the costa prominulous to subplane above, 276 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 prominent below, the principal veins 12-15, widely arcuate, prominulous, 6-12 mm apart. Inflorescences mostly rhizomatous, the peduncles elongate, ascending, 5-10 cm long, 1-2 mm wide, rubescent, glabrous, terminated by 3-4 usually radiately disposed patulous branches, the cymules ascending, each branch 7-17 flowered, the pedicels stiff, glabrous, to 5 mm long, ca 0.3 mm wide. Flowers with the buds obviously acute at the apex, the hypanthium strongly 6-8-ribbed, emerald-green then red, to 3 mm long, the calycine lobes flaring, deltoid, 0.3-0.5 mm long; corolla to 6 mm long, the lobes 4, much longer than tube, glabrous, the anthers 4, oblong, to 3.8 mm long, the connective not produced at apex or at base, the filaments to 0.6 mm long; ovary 2-loculate, the style to 4 mm long, the stigmas 2, clavate-oblong, ca 1.1 mm wide, each stigma somewhat uncinate. Fruit not seen. Panama: chiriqui: Cerro Horqueta, ca 6000 ft, Blum & Dwyer 2623 (MO), Dwyer & Lallathin 8756 (MO), 8789 (MO); Bajo Chorro, ca 6000 ft, Davidson 216 (F, MO, type collection); Bajo Mono & Quebrada Chiquero, 1500 m, Woodson & Schery 595 (F, MO). Like Hoffmannia eliasii, this species is succulent and has rhizomatous inflores- cences. The corolla is yellow to yellow-brown or yellow above and red at the base. The corolla lobes appear to be almost 3 X the length of the very short corolla tube and in this species represent a rehable diagnostic character (cf. remarks in introduction, p. 5). When the 2 stigmas are pulled apart with dissecting needles they have a claw-like or cheliped appearance. 7. Hoffmannia eliasii Dwyer, spec. nov. Herbae ad 1 dm altae. succulenfae. < simplicibus hgnosis, hie 15-20 cm longis, ad 1.2 cm latis, radicibus fibrosis hgnosis. Folia subsessiha aut plerumque petiolata petiolis ad 12 cm longis, plerumque leviter alatis; lamina oblonga vel obovato-elliptica, apice cuneata ad subobtusa, vix acuminata, basi cuneata ad contractocuneata, ad 25 cm longa, ad 9.5 cm lata, rigido-chartacea, vix discolor, glabrescens, ciliis in costa venisque paucis, costa supra prominula, subtus subprominenti ad subplana saepe leviter alata, venis lat- eralibus principahbus 12-16, leviter arcuatis prominulis plerumque minute alatis. locata. Infl intermediis irregularibus et mox ramosis ilo-pinnatiformibus, vena marginali con peduncuhs for marcescentibus, canaliculatisque pilosis cihis debilibus diffusisque, pedicelUs fere cam nullis vel ad 4 mm longis. Flores hypanthio longo, glabro, in x-sect costato, dentibus calycinis 4, vix inaequalibus erectis subu- latis 1-1.8 mm longis, minute-puberulis; corolla omino glabra, tubo cylindrico, ad 2.5 mm longo, lobis 4, lanceolatis quam tubo longioribus, ad 4.5 mm longis; stamina 3, antheris 3-4 mm longis, subsessilibus, filamentis ca 0.5 mm longis, stylo lineari-cylindrico, connectivis apice basique non productis, stigmatibus 2, inaequali-digitiformibus, ad 0.7 mm longis quam stylo vix latioribus; ovarium 2-loculare. Fructus (hie immaturi) pedicellati, pedicellis fortiter adscendentibus. 1969] DWYER — PANAMANIAN HOFFMANNIA 277 diffuso-pilosis, ad 4 mm longis, oblongi, ad 0.5 mm longi, ad 2.5 mm lati, calyce persistent! coronati, lobis 1-2.5 mm longis, adscendentibus, in sicco rubri, modice costati, minuto puberuli. Panama: darien: Cana-Cuasi Trail, Chepigna, 4000 ft, Terry & Terry 1508 (F, GH, MO); Rio Cuasi, mainstream, 0-2.5 mi S Tres Bocas, Kirkbride & Duke 1129 (MO, holo- type). SAN BLAs: headwaters Rio Mulatupo, Elias 1779 (MO). Colombia: choco: Rio Nercua, Quebrada Barrial & Quebrada Ambrosio nr Camp Curiche, Duke 11438 (MO), Hoffmannia eliasii is named in honor of Dr. Thomas EHas, one of the collec- tors of the species. It is readily distinguished from its obvious ally H. davidsoniae, from the Province of Chiriqui, by its oblong rather than subrotund leaves and much more strict inflorescence. The stamens of the new species are usually three, and the stigmas are narrowly oblong. A Costa Rican collection, Austin Smith 90S (NY), resembles the new species in leaf shape, but its inflorescences are subsessile. 8. Hoffmannia fimbrianthera Dwyer, spec. nov. Suffrutices caulibus laevibus arundinoideis glabris teretibus, internodis ad 15 cm distantibus. Folia petioHs gracilibus laxis, ad 7 cm longis, ad 1.2 mm latis, minute arachnoideo-puberulis; lamina oblongo-rotundata, saepe subfalcata, ad apicem cuneata, acumine ad 1,5 cm longo, ad basim acuta vel angusto-cuneata, ad 19 cm longa, ad 9.5 cm lata, papyracea, vix discolor, in sicco grisea, sub lente conspicue foveolata, supra glabrescens, infra in venis solum granuloso-puberula, costa supra plana, subtus vix prominenti, venis lateralibus ca 15, divergenti- arcuatis, subtus conspicue prominulis, venis intermediis patulo-pinnati-formibus, vena marginal! undulata a margine ca 8 mm distanti, Inflorescentiae sessiles, flori- bus paucis fortasse (3-)5(-10) subsessilibus vel ad 1 mm loneis. Flores hypanthio ineo- camosa extus ferrugineo-puberula intus minuto-farinosa, lobis fortasse 4; stamina 4, antheris oblongis, ad 3.2 mm longis, minute apiculatis, connectivo basi fimbria patula petaloidea ad 3 mm longo ornato; ovarium 2-loculare, ovulis multis, stylo ad 3.5 mm longo, ad 0.2 mm lato, ad apicem latiore, stigmatibus 2 clavato- connatis, ca 0.35 mm latis, Fructus non visi. Panama: bocas del toro: Rio Teribe just below Puerto Palenque, 350 m, Kirkbride Gr Duke 556 (MO, holotype). rs as a "cane-like, coppiced shrub; This ramiflorous, tetramerous. Durnlish." The and tures of the species. The same species 9. Hoffmannia haydenii Dwyer, spec. nov. Herbae caulibus in vivo fortasse teretibus, in sicco 4-angularibus, glabrae. Folia sessilia; lamina obovato-elhptica vel oblonga, versus apicem rotunda, apice acuminata, acumine ad 1.5 cm longo vel fortasse longiore, versus basim contracto- [Vol. 56 278 ANNALS OF THE MISSOURI BOTANICAL GAKDEN cuneata. tunc ukimc auricula.a. auriculis ad 1 cm latis. ad 35 cm longa ad 14 cm lata, tcnui-coriacea. discolor, supra in sicco Ijiunnca. supra glab c en . subtu in osta vcnisque pubcrula, costa supra subplana ct kv.tcr cana he h subU^ vix prominula. ad 3 nnn lata, vcnis latcralibus - 2\(f°"f.%P';f ,"J,^;^ "^1 complanatis subtus prominulis. Inflore.centiae r uzanth..c, 1.5-7 ™ '™8a ^cUs dunculis 1.2-6 en, longis. capillaccis, ca 0.3 mm lat.s vcl ^-^ ™" '^ -;^^^'™^ nubcrulis in vivo rubris, ad 2.5 mm longis, pubcruhs. Ftor.s gen mis apicc u':.ato-aeutis. hypan.hio pubcrulo vix costato, lobis calyeinis 4. -ect. aneeola^i 3-4 mm longis, in sicco purpurco-rubris pubcruhs; corolla lob.s 4, '"-. <^» ^ mm lonois cxtus pubcrulis quam tubo longioribus; stamma 4, anthcris ca 3-8-4t-''- j mm Lgis, c nncctivis apicc hasiquc non productis. fdamcntis brcv.bus; cnanum 2leulari, stigmatibus in capi.ulum clavatum rotundumque aggrega.is. VruCu. non visi. Panama. ch,r,bci: Finca Collins, ca 5000 ft, Uayd,,, 117 (MO, holotype). This new species is obvously related to Standley's H. s„ha,mcuk,a known only from fruiting material collected at Cartago Costa Rica, which also has Icaf- hlades that are dilated and clasping. H. haydem. however, has much th.cUr III j-4 illT 111 LA l_ kA 1- V- ^-* -* -* *'• 1, ^*- V- » *-• -* — ^-* — - - - ^^ *- leaves, longer pedicels, and much more elongate calycine lobes. ffrutices ad 2 m alti, caulibus teredbus (?) alabris. folia petiohs ad 13 cm longis, in med.o ca 3 mm lat.s. infra basnn an mac conspicue turgidis, inflationibus oblongis, 6-10 cm longis. m sicco ea ' ™ '"^^ lamina elliptico-cblonga, apicc subrotunda ad lato-cuneata acummc ad longo. ad 40 em longa, ad 20 em lata, rigido-chartacca, in s.cco '^'^™'«' ^» " supra prominula canaliculataque subtus subprominenti versus ''asm, d.st nc c siibercso-alata. vcms latcralibus 15-25. ad 3 em distantibus. »[™»"^ "P" ™ prominulis. subtus conspicuo-prominulis. vcnis intcrmediis plus^divcrgcnt.bus mox evanescenlibus, venis tcrtiariis patulis irregulari-pinnatiformibus, supra tor a se glabra, subtus in costa veniscjuc interdum tcgeticulis ciiorum simpl'J'"™ » ", sorum crystalhnorum ornata. ciliis paueis stellatis. h.flore^c-enUae P'^^q-fJ* per axillaL. pcduneulis rcflcxis, M.5 em longis, ea 1.3 mm '^"-..'^'g^f"; "'"^ is ciliorum ervstallinorum ct raphidium instructis, ciliis panes, ad 0.7 mm longis, rare stellatis, floribus subsessilibus. F/«rc.s (hie in frucfhus Juven hbu ) ) calycis brevi subturgida villosa. dcntibus 4, crasso-carnosis (m med.o solu. i cihatis, ad 4.5 mm longis. ad 3.5 ram latis. vcnis paueis adscendent.bus; ovarium 2-loculari. paricte hypanlhii crasso. ovulis muhis. Fructm matur. non v.si. nibus crasso-reticulatis rho..iboideis. ca 6 mm dia.Ti. PAN.VMA: BOCAS OB. xo.,o: ^..^na Vis.a, Aln.iran.. C„o,„.j^225 (F^ rapid streams through evergreen seasonal forest, ca 300 400 tt, Kirklniac c holotype). ff Standley, known from the Provinces of Chiriqui and Code, the only other species of the uenus with veseicnliferous petioles. Hoffmannia kirkhrulei, named after 1969] DWYER — PANAMANIAN HOFFMANNIA 279 Mr. Joseph Kirkbride, Jr., is readily dis perhaps the largest of the genus, whicl (Piuhiaceae^ If presence of occasional stellate hairs on the leaves of the new species is noteworthy, as stellate hairs have not been reported in Hoffniauuia. 11. Hojfmannia lanoistignia Dwyer, spec. nov. Friitices cauhbus arundinoideis teretibus et laevibus, ciliis minutis crebris fcrrugineo-moniliformibus deciduis. Folia valde adsccndentia, bene distantia, 2.5- 3.5 em distantia, petiolis ad 7.5 em longis, ferrugineis; lamina oblonga, apiee obtusa, vix acuminata, ultime obtusa, basi cuneata, plerumque asymmetriea, in sicco discolor, papyracea vel rigido-chartacea, sub lente minute granulosa foveolataque, ciliis subtus paucis praeter eostam denso pubescentem, costa supra prominula, sub- tus prominula, ad 1.5 mm lata, venis secundariis ca 15, arcuatis, subtus prominulis, venis tertiariis patulo-pinnatiformibus. hiflorescentiae saepe 1 per axillam, saepe a partibus caulis aphyllis orientes, pedunculo adscendenti, ad 10 cm longo, modice tcnui, ca 2 mm lato, substricto vel arcuato, glabrescenti vel sub lente ciliis paucis crystallinis ornato. Flores fortasse omnino glabri, hypanthio oblongo, ad 3.8 mm longo, costato, dcntibus calycinis 4, erectis triangularibus, ca 1 mm longis, corolla tubo brevi tenui-carnoso, lobis 4(-6), ca 4.5 mm longis, carnosis; stamina 4, antheris ca 3.8 mm longis, ad 1.2 mm latis, connectivis apicc basique non pro- ductis, filamentis ca 1 mm longis, complanatis, ad 1 mm latis; ovarium 2-locularc, stylo ca 4 mm longo, stigmatibus 2, lanceolatis, ca 2.8 mm longis, ad 0.8 mm latis, crassis quam stylo latioribus in medio constrictis. Panama: chiriqui: Cerro Horqueta, ca 6000 ft, Dwycr 7735 (MO, holotypc). As the specific name suggests, this new species is distinguished by its lance- shaped stigmas. A preliminary study of more than 20 Panamanian species of Hoffmannia suggests that stigma morphology may well represent one of the most important diagnostic characters in the genus; these fall into three kinds: 1) ball-shaped, resulting from 2 or 3 stigmas being subconnate, 2) linear, and 3) lanceolate. 12. Hoffmannia lewisiana Dwyer, spec, nov, Friitex, ramuhs laevibus glabris canaliculatis. Folia petiolis ad 4.5 cm longis, glabris; lamina oblonga, apiee subcontracto-acuta acuminataquc, acumine ad 1.5 cm longo, basi acuta et fere symmetrica, concolor, supra glabra, subtus solum in costa venisquc puberula, costa supra subplana, canaliculata, subtus prominula, ad 0.15 cm lata, venis lateralibus ± 12, supra planis, subtus prominulis, arcuatis, postremo manifeste patulo-reticulata; stipulae non visae. Jiiflorescoitiae ad 6.5 cm longae, ad 4 mm latae, patulae, pedunculo brevi, ram is pluribus, 2-4 cm longis, 0.8-1.8 mm latis, cymis paucis alternatim vel saepe subradiate dispositis, pedicelhs 1-7 mm longis, ca 0.4 mm latis, dense albido-puberulis. Flores gemmis (lob ) ad 7 mm longo, carnoso, puberulo et conspicue costato, dcntibus 4-6, erectis, [Vol. 56 280 ANNALS OF THE MISSOUKI BOTANICAL GARDEN cuncatis vel triangularibus, inacqualihus (uno saepc conspicuc rcducto), ad 2.5 mm longis, puberulis; corolla tube ad 11 mm longo, carnoso, lobis 5, ad 5.5 mm longis, cxtus puberulis; stamina 4-5, antbcris ca 3.5 mm longis, ca 1.3 mm latis, connectivis apicc basique non productis, filamcntis complanatis, ca 1 mm latis; ovarium (2-)3-4-locularc, stylo 12-14 mm longo, stigmalibus (2-) 3-4, in capitu- lum rotundum dispositis. Fructns non visi. Panama: c:HiRiyui: vie Ccrro Punta, 6800 ft, cloud forest and disturbed lcI^c, Ruli!,uay & Solis 2413 (MO, holotypc). This new species, named in honor of Dr. Walter Lewis of the Missouri Bo- tanical Garden, is readily distinguished by its extraortlinarily large flowers, whose perianth parts and stamens are usualh in fives, and whose gynoecium is occasion- ally quadriloculate. As the diagnosis indicates the perianth parts may occasionally b e ui sixes. 1 cm distantibus. Folia petiolis 13. IIuff))ia)i)iia longicalycina Dwj'cr, spec. nov. Suffnitices ramulis lae\ ibus ad 10 cm longis, angulari ad plano-compressis, ad 2 mm latis, ciliis albis simplici- bus, crasso-subulatis tegctes crustaceas formantibus; lamina oblonga, ad apicem cuneata, acumine brevi, ad 1 ciri longo, ultime acuto, basi attenuatocuneata, costa subtus prominula porcataque, ad 2 mm lata, rigido-chartacea, vix discolor, in sicco viridis, supra glabrescens, subtus in costa venisque appresso-ciliata, ciliis interdum rigidis appresso-falcatis albidis, venis lateralibus ca 16, patulo-arcualtis prominulis. liiflorcscentiae ad 3.5 cm longae, in capitula axillaria floribunda conglomerata dispositae, pedicellis ad 3 mm longis, puberulis. Flores non visi. Fmctus oblongi, in x-sect terctes, hie ad 12 mm longi, ca 7 mm lati, extus levitcr costati, pubescentes ciliis aliquibus ad 2 mm longis ferrugineis debihbusque, lobis calycinis pcrsistentibus 5, erectis lineari-lanccolatis, ad 12 mm longis, ca 2 mm latis, petaloideis pubescentibusque, 2-loculares, seminibus multis crasso-reticulatis, ca 0.5 mm diam, per parietem tenuem visibilibus. PaxNama: darien: Cana-Cuasi Trail bctw Ccrro Campiaincnto & La Fscalcra to "paramo" E of Trcs Rocas, KirJihridc & Dtikc 1278 (MO, holotypc). This new species is distinguished from all other species of Hoffmajiiiia by the length of the calycinc lobes. The elongate calyx suggests the genus Amj?hidaysa Standley, with three species in the New World tropics. Hofjmainiia lougicalyciua, however, has a complete septum, at least in the basal part of the fruit, and two locules, and has the reticulate seeds characteristic of Hoffniaunia. The seeds arc visible through the thin wall of the dried fruit, a character which I have not hitherto observed in the manv Iloffiiiaufiia fruits examined. 14. Huff)iiaii)!ia ostaurea Dwyer, spec. nov. Suifrittices succulenti, ramuhs in sicco canalieulatis, ^ areabus ciliorum aggregato-crustaceorum ornatis, ramulis juvenihbus crcbris raphi- dibus notatis, internodis 1.5-3 cm distantibus. Folia petiohs arcuato-ascendentibus, ad 8 cm longis, in sicco 2-15 cm latis, glabris, raphidibus crcbris; lamina elhptica, apicc cuneata brevi-acuminata acumine ad 1 cm longo, basi subcontracto vel lato- 1969] DWYER PANAMANIAN HOFFMANNIA 281 cuneata, ad 2 5 cm longa, ad 7,5 cm lata, tcnui-mcmbranacea, concolor, glabra sed raphidibus vcrmiformibus ad 0.3 mm loiigis ornatis, costa utrinquc complanata, vcnis latcralibus 12-15, patulo-arcuatis gracillissimis, vcnis tcrtiariis irrcgulari- pinnatiformibus, areabus rcticulatis conspicuc patulis. Flares (hie gemmas solum vidi) glabri, hypanthio oblongo-rotundo, ca 2 mm longo, non costato, lobis calycinis 4 inacqualibus subulatis, ad 1.2 mm longis, raphidibus crcbris albidis; corolla subcarnosa lobis fortasso 4; stamina 3 (4), anthcris ad 3 mm longis, connectivis apice basique non productis, filamcntis brevibus; ovarium 2-loculare, stigmati- bus 2 in capitulum rotundum ca 1 mm longum dispositis. Fructus rotundi, ad 5 mm diam, in vivo rubri, in sicco rubro-purpurei, pcdicellis gracilibus, ad 5 mm longis. Panama: bocas del toro: Rio Tcriblc below Oucbrada Huron & Qucbrada Schlunjik, Kirkbride & Duke 501 (MO, holotypc). The specific latin name ostaiirea means *'mouth of the bull" and suggests the provincial name "Bocas del Toro." This new species is marked by extraordinarily thin leaves; in the dried state these are so translucent that one can read newsprint through them. The axillary inflorescences are very reduced and few-flowered, the latter borne on very slender ligneous pedicels. The fruit at maturity is a striking purple-red color. 15. Hoffmanniapittieri Standley, Conlr. U.S. Nat. Herb. 18: 140, 1916. Hoffrnamiia panaynensis Standley, Contr. U.S. Nat. Herb. 20: 204, 1919. Shriihs to 8 m tall, the thicker branches smooth, drying tan, terete, glabrous, hollow^ in the center. Leaves with petioles 5.5-7 cm long, 1.5-3 mm wide, the lamina often slightly decurrent, elliptic or obovate-elliptic, acute to cuneate, or rarely subrotund at the apex, acuminate, the acumen to 2 cm long, acute to attenuate-cuneate at the base, 9-3 5 cm long, 8-12 cm wide, the larger blades 2.5-3 X longer than wide, chartaceous, usually shghtly discolorous, usually drying tan or brown, glabrous, the costa prominulous above, prominulous to prominent beneath, often ligneous, to 2.5 mm wide, the lateral veins (IT) 15-20, widely arcuate, 1(2.2) cm apart, with T2 conspicuous irregular veins between each pair of laterals. Inflorescefices in the uppermost axils, patulous, 3-9 cm long, the primary peduncle compressed as a nodal pad, or elongate, the secondary peduncles 1-4 glabrous, the flowers usually patulous, the pedicels to 1 5 mm long, stiff, glabrous. Flowers wath the hypanthium carnose, ribbed, sparsely puberulent to gla- brous, the calycine teeth 4, deltoid, 0.6-2 mm long; corolla to 13.5 mm long, subcarnose, glabrous to sparsely puberulent, the lobes 4(-5), 2.3-4.3 mm long, 1.7-3 mm wide; stamens 4, the anthers 2.3-4.5 mm long, the connectives not produced at apex and at base, the filaments to 1.8 mm long; ovary 2-4-loculate, the style to 1 1 mm long, the stigmas usually ball-shaped, 1.2-1.8 mm long, wider than style, each stigma ovate-oblong to cordate. Fruits subrotund, tlie calycine lobes persistent, to 6.5 mm long, glabrous, drjing red-brown, scarcely ribbed, the seeds brown, foveolate. Distribution: Known only from the higher elevations of Chiriqui and neigh- boring Bocas del Toro. [Vol. 56 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN Panama: bocas del toro: l^ol)aldo Trail, N slopes Cerro Horqucta, Allen 4962 (MO). CHiRigui: Cerro Punta, 2000 m, Allen J533 (MO); Boquete, Finca Collins, ca 5000 it, Buyer & Ilaydcn 7654 (CTT, K, MO, UC, US), 7665 (F, GH, K, MO, NY, UC, US), Stern et ah 1127 (MO); El IJoquetc, Bajo Chorro, Davidson 47 (F, MO), 51 (F, MO); Boquete, Cerro Horqueta, Blum & Dxvyer 2599 (MO), 2643 (MO), 2647 (MO), Duize et al. 13651 (MO), Dwyer 7654 (MO), Dwyer & nayJ4jn 7745 (GH, MO), Buyer & Lallathin 8756 (MO), 8783 (MO), 8790 (MO), Kirkhride 147 (MO), von Hagen & von llagen 2076 (F, NY), 2078 (NY); betw Pinola & Quebrada Hondo to summit on Chiriqui Trail, Kirkhride & Duke 894 (MO); Upper Valley Rio Chiriqui Viejo, vie Monte Lirio, 1300-1900 m, Scihert 165 (F, NY, MO); Rio Chiriqui Viejo, White & White 2 (F, NY, MO), 74 (F, GH, MO); vie Casita Alta, Volcan de Chiriqui, 1500-2000 m, Woodson et al 839 (F, MO, NY). Virtually all of the collections cited above were made at or very close to Cerro Horqueta in the Province of Chiriqui, the type locahty of the species. H. pittieri is the tallest of tlie Panamanian Hoffnianjiia species, reaching up to 25 ft in height. It is distinguished from the other species of the Republic by its hollow stem (older branches), large leaves with numerous lateral veins, and large flowers usually born on elongate pedicels. The inflorescence is very variable as to the structure of the peduncles. Noteworthy is the fact that the pedicel length in the fruits does not appear to be correlated with the length of the floral pedicels, the latter being longer in the material examined. The flowers are orange-yellow (tliose of Davidson 51 are described as 'whitish green"), and the fruit is red at maturitv. le.Uoffniannia psychoUiaefoUa (Benth.) Griseb., Fl. Brit. \V. Ind. 321, 1861. Higginsia psychotriac folia Bcnth. in (3erst., Vidcnsk. Meddcl. Dansk Naturhist. Foren. Kjoben- havn 1852: 50, 1852. Shrubs to 2.5 m tall, the stems ultimately slender, terete, glabrous, rubescent. Leaves with petioles to 9 cm long and 2 mm wide; lamina oblong, acute and finally attenuate-acuminate at the apex, the acumen to 2.5 cm long, acute and inequilateral at the base, to 20 cm long and 8 cm wide, usually 3-4 X longer than wide, stiffly chartaceous, concolorous or discolorous, glabrescent, usually with minute, scattered, white granulations on both surfaces, the lateral veins 10-12, widely arcuate. Inflorescences in the axils of the uppermost leaves, the peduncles reduced to a pad or to 8 mm long, the flowers radiately disposed, to 7 per inflorescence, the pedicels slender, to 4 mm long. Flowers to 12 mm long, the hypanthium 2-4 mm long, scarcely ribbed, often with abundant red raphides, the calycine teeth 4, cuiieatc-dcltoid, 0.8-2 mm long, acute; corolla 8-11 mm long, the lobes 4, thickly petaloid, often longer than the tube at maturity; stamens 4, the anthers 2-2. 8 (-4) mm long, the connective not produced at apex and base, the filaments ca 0.5-1 mm long; ovary 2-3-loculate, the style to 5.5 mm long; stigmas 2, linear, equal, spreading at maturity, 23. 3 mm long. Fruits with pedicels radiately disposed, slender, shorter than the pericarp, oblong to ovate- oblong, obtuse or truncate at apex, to 8 mm long, at least 2 X longer than wide, at first delicately ribbed, smooth, glabrous, often drying yellowy finally fleshy, china-red, pink, or purple. Distribution: Guatemala to Panama. 1969] DWYER — PANAMANIAN HOFFMANNIA 283 Panama: ciiiriqui: vie "New Switzerland", central valley, Rio Chiriqui Viejo, Alhn 1329 (F, MO); Boquete, Ebinger 671 (MO); Bajo Chorro, 6000 ft, Davidson 198 (MO); Finca Collins, Dwyer Or Hay den 7666 (MO), 7681 (GH, MO, US), Hayden 162 (MO), Kirkhride 120 (MO); Cerro Horqueta, Divycr et ah 518 (GH, K, MO, NY, UC, US), Dwyer & Hayden 7653 (GH, US), 7664 (GH, K, MO, UC, US), von Hagen & von Hagen 2054 (F, NY), 2174 (NY); Palo Alto, E Boquete, 5000 ft, Stern et al 1015 (MO); Quebrada Velo, 1800 m, Woodson &■ Schery 258 (F, MO); Bajo Mono & Quebrada Chiqucro, Wood- son & Schery 564 (F, MO); Casita Alta, Volcan de Chiriqui, Woodson et al 884 (MO, NY), cocle: betw Cerro Pilon & El Valle dc Anton, 700-900 m, Duke & Divyer 13972 (MO). Hoffmannia psychotriaefolia is readily distinguished in the fruiting stage, at least prior to a terminal period of fleshiness, as the fruits are at least twice as long as wide, a condition not observed in Panamanian species or in many extra-Panama sheets examined in various herbaria. In the flowering stage it is often confused with H. woodsonii, but it is distinguished from the latter by its much larger leaf blades with more lateral veins. The flowers are yellow, and the fruits vary from pink to red, presumably being a china-red as they approach maturity. 17. Hoffmannia rubripigmenta Dw^yer, spec. nov. Herbae lignosae, humiles, ramulis glabris raphidibus crebris minutis oblongis ornatis. Folia pctioHs ad 1 cm longis, leviter alatis; lamina obovato-elliptica, apice late cuneata, ultime obtusa, versus basim contracto-cuneata et vix decurrens, ad 23 cm longa, ad 9.5 cm lata, chartacea, in sicco opaco-rubra, fortasse glabra, costa utrimque prominula, venis lateralibus ca 8, arcuatis, venis intcrmediis plus divergentibus, tunc ramosis et evanescentibus; stipulae non visae. Inflorescentiae petiolo quam pauUum longiores, pedunculo gracili, erecto, glabro, ad 2 cm longo, ca 5 mm lato, in sicco rubro, floribus ca 8, subsessilibus. Flares (hie gemmas solum vidi) gemmis angusto-oblongis, ca 1 mm latis, hypanthio oblongo, ad 2.5 mm longo, carnoso, glabro, ecostato, dcntibus calycinis erectis cuneatis acutis, ca L5 mm longis, vix connatis, corolla carnosa, tubo fortasse brevi, lobis 4, glabris; stamina 4, antheris ca 4 mm longis, connectivis apice basique non productis; ovarium 2-loculare, pariete hypanthii crasso ad 3.5 mm lato, stylo cylindrico, stigmatibus erectis digitiformibus, ca 1.3 mm longis, vix quam stylo crassioribus, Fructiis non visi. Panama: darien: upper Rio Membrillo & Camp 7 on Construction Rd to San Bias, 100-800 m, Duke 10870 (MO, holotype). On boiling the flowers and the leaves one extracts a red pigment. I have not encountered this in any other species of the genus studied. The buds are very narrow and the digitiform stigmas suggest a possible relationship with H. psycho- triaefolia. The flowers are described as pinkish. 18. Hoffmannia siihauriculata Standley, Jour. Wash. Acad. Sci. 18: 179, 1928. Herbs, short, w^oody, the roots fibrous, the stems strongly quadrangular to terete, smooth, glabrous. Ixaves sessile, the blade elhptic to obovate-elliptic, or obovate-spatulate, acute and short-acuminate at the apex, cuneate and often widely so at the base, to 2.5 cm wide at the base, ultimately obtuse or vaguely auriculate, 9-30 cm long, 5-10 cm wide, stiff'ly chartaceous, glabrous with oo [Vol. 56 284 ANNALS OF THE MISSOURI BOTANICAL GARDEN linear raphidcs, the costa porcate above, prominent beneath, the lateral veins 10-25, widely arcuate, prominent beneath, usually 1.5-2.5 em apart, with usually a conspicuous intermediate vein between a pair of principal veins. Inflorescences with the peduncle arising at or near the ground line, the peduncles wiry but not stiffly erect, here to 4 cm long, the cymules few-flowered. Fhmers (only buds seen) with the hypanlhium glabrous, the calycine teeth 4, cuneate-subulate, acute, puberulent, less than 1 mm long; stamens and petals not seen; ovary 2- locular. Fruits oblong, here to ca 6 mm long, 4-ribbed, the seeds ca 1 mm long, coarsely foveolate. Distribution: Known from one collection at Cartago, Costa Rica, and from Bocas del Toro and Darien, Panama. Panama: bocas del toro: s. loc, von Wcdd 361 (MO), dakien: Ccrro Pirrc, Brh- tan 627 (MO). Standley, in his description of the type, states that the leaves are membra- naceous; those of the Panamanian collections are stiffly chartaceous. The flowers of von Wedel 361 are described as white, and those of llristan 62 7 as lilac. 19. Uofimannia vesciciilifera Standley, Publ. Field Columbian Mus., Bot. Ser. 4: 285, 1929. Shrubs, presumably densely ferrugineous and arachnoid flocculose. Leaves with petioles to 4.5 cm long, conspicuousl) inflated for most of tlieir length, the inflations to 1 cm wide, located nearer the blade than the node, densely fcrrugineous-flocculose; leaf-blades oblong-rotund to obovate-oblong, briefly acumi- nate at the apex, cuneate to rounded at the base, to 2 5 cm long and 15 cm wide, chartaceous, discolorous, drying brown, usually foveolate above, the hairs diffuse, to 5 mm long, often calcareous at base, confined to veins and venules, the costa immersed above, prominulous to piano-compressed beneath, the principal secon- dary veins 12-25, widely arcuate to substrict, the margin often weakly ciliolatc. Inflorescences to 5 cm long, the peduncles to 2 cm long, the flowers numerous in densely bracteate axillary capitula, these rounded or compressed-rotund, the pedicels to 4 mm long, glabrous; bracts hnear-lanceolate, to 1.5 cm long, 0.3-1.5 mm wide, with 1-2 irregular ascending submarginal veins, the margins glandular- pectinate-ciliate, the hairs to 4 mm long. Flowers with the hypanthium oblong, truncate apically, glabrous, delicately 4-ribbed, the calycine lobes 4, unequal in length and width, lanceolate, tapering narrowly above middle, 7.5-10 mm long, 1.5-2 mm wide, the hairs ferrugineous, marginal, to 2 mm long, usually densely aggregated above middle; corolla with lobes shorter than tube; stamens 4, the anthers with connective not produced at apex or at base; ovary 2-locular, the style slender, the stigmas clavate, spreading at maturity, wider than and ca V4 length of style. Fruits oblong-rotund, here to 5 mm long and 4.5 mm wide, dehcately- ribbed, the ribs ca 10, the pericarp thin-walled, the raphides abundant, the hairs deciduous, the calycine lobes 4, persistent, linear-lanceolate, to 10 mm long, the hairs deciduous in patches, the seeds mostly rectangular to cuneate, coarsely reticulate. 1969] DWYER — PANAMANIAN HOFFMANNIA 285 Panama: bocas del toro: Chiriqui-Caldcras Trail bctw Uigucron & Gutiorrca, Kirk- hride & Duke 761 (MO); E slope La Zorra to divide, mossy forest, cloud forest, Kirkhride & Duke 829 (MO); Water Valley, vie Chiriqui Lagoon, von WcM 1514 (F, MO); Fish Creek Mts, vie Chiriqui Lagoon, von Wedel 2284 (MO), cocle: Ccrro Caracoral, elfin forest, Kirkhride 1117 (MO). This is perhaps the easiest of the Panamanian Hoffmannia to recognize because of the elongate hairs of the inflorescence and the inflated bases of the petioles which resemble those of Tococa Aubl. CMelastomaceae), Kirkhride & Duke 761 note the absence of ants on the swollen petioles. They also describe the stipules as *'horny/' The inflorescence of H, vesciciilifera is red, the hypanthiuni pink, and the corollas yellow. The mature fruits are described as red. 20. Hoffmannia woodsonii Standlcy, Ann. Missouri Bot. Gard. 28: 471, 1941. Shrubs 2-3 m tall, the stems glabrous, smooth, the raphides dot-like, white. Leaves with petioles 2-6 cm long, to 0.8 mm wide; lamina oblong, attenuate- acuminate at the apex, the acumen to 1 cm long, attenuate-cuneate or acute at the base, 5-12 cm long, l-5(-6) cm wide, membranaceous or thin-chartaccous, the raphides scattered, white, linear on lower surface, subgranular above, to 0.4 mm long, the costa piano-compressed beneath, the lateral veins 4-8, prominulous (especially beneath), glabrous or occasionally puberulent. Inflorescences 1-2 per axil, to 1.5 cm long, compressed, the peduncles reduced to a pad or to 6 mm long, ca 0.5 mm wide, glabrous to ferrugineous-puberulent, the flowers usually 2-6 per cymule, the pedicels 5-7 mm long, often radiately disposed or deflexed- candelabraform, capillaceous, glabrous to ferrugineous-puberulent. Flowers with the hypanthium squarrose in x-sect, subcarnose, glabrous or puberulent, the caly- cine teeth 4(-5), cuneate-subulatc, 0.5-1.3 mm long; corolla 7-8 mm long, the lobes 4, ca 4 mm long, delicately carnose, diffusely puberulent on the outside, often delicately-ribbed; stamens 4, the anthers partly exserted at an thesis, 3-4.7 mm long, the connective produced at the apex, then usually ovate-cuneate, to 1 mm long; ovary 2-locellate, the style 4-5.5 mm long, the stigmas 2, oblong- capitate, 1-1.2 mm long, wider than the style. Fruits (here not seen in fleshy mature stage) oblong-subrotund, truncate, to 4.5 mm long, the calycine lobes persistent, glabrous, dehcately ribbed, the ribs well-spaced, varying in prominence. Known only from Panama. Panama: bocas del toro: betw Gullierrez Quebrada & E slope La Zorra, headwaters Rio Mali, Chiriqui Trail, Kirkhride & Duke 715 (MO), canal zone: Barro Colorado I, Croat 5118 (MO), chiriqui: Cerro Horqueta, Blum Gr Dwyer 2619 (MO), Duke et al 13634 (MO); betw Pinola & Quebrada Hondo toward summit Chiriqui Trail, Kirkhride & Duke 915 (MO); Boquete, Finea Collins, Stern et al 1149 (MO); upper Rio Chiriqui Viejo, 1300-1900 m, White & White 6 (MO), 26 (MO); Quebrada Velo, 1800 m, Woodson & Schery 260 (F); vie Callejon Seco, Volcan de Chiriqui, 1700 m, Woodson & Sehery 503 (holotype, F; isotype, MO), cocle: Cerro Pilon & El Valle de Anton, 700-900 m, Duke £r Dwyer 13972 (MO); Cerro Gaital Caracoral, 2700-3000 ft, Divyer £r Correa 8910 (MO); Cerro Pilon nr La Mesa, ca 2700 ft, Dwyer £r Lallathin 8593 (MO), 8627 (MO), 8659 (MO), 868J (MO), Lallathin 5090 (MO); mts N El Valle de Anton, 2500-3000 ft, Lexris et al 1732 (G, MO, US), Stinison 5033 (MO), los santos: Loma Prieta, 800-900 m, Duke 11849 (MO); Loma Prieta, Cerro Grande, 2400-2800 ft, Lewis et al 2212 (GH, MO, US), 2242 (GH, MO, US), darien: s. loc, Bristan 103 (MO). [Vol. 56 286 ANNALS OF THE MISSOURI BOTANICAL GARDEN This species is often difficult to distinguish from H, pittieri and H. psycho- triaefolia. Its leaf blades are usually much smaller; Croat 5118, probably the sole collection of the species made near sea level and the only collection of the genus from the Canal Zone, has larger and wider leaves. The consistently produced apex of connective of the anthers, visible under magnification, readily distinguishes H. ivoodsonii from the two aforementioned species. Its clavate-rotund stigmas readily separate it from H. psych otriaef alia, and its smaller leaves with few lateral veins serve to distinguish it from H. pittieri. Excluded Species 1. Hoffmannia discolor (Lemairc) Hemsley (Biol. Centr. Am. 2: 36. 1881) probably docs not occur in Panama. One collection, Terry Gr Terry 1508 (MO) from Boquetc, Prov, Chiriqui, Panama, has been erroneously assigned to this species. It docs not have the minute calycine glands and linear anthers which Standley in the North Am. Flora (loc. cit., p. !i2) includes in his floral description. 2. Three collections of von Wcdel [2363 (MO), 717 (F), and 963 (F)] from Chiriqui Lagoon, Prov. Bocas del Toro, probably represent a new species. As these collections belong to the complex of Hoffmannia marked by narrow leaf-blades with attenuate-acuminate apices and elongate petioles, a complex in need of critical study on a New World basis, I have elected not to prepare a diagnosis of it. NOTES THE SKUNK CABBAGE IN MISSOURI A single specimen of Symplocarpiis foetidus (L.) Nutt. has flowered and vegetatively flourished in Jefferson County, Missouri during the spring seasons of 1967-9; whether it has borne fruit during the summers of 1967 and 1968, I do not know. However, the appearance of Skunk Cabbage in Missouri is, so far as I have been able to determine, new; its range is described by Fernald in Gray's Manual (8th ed.) as "Que. to se. Man., s. to w. N.S., N.E., L.I., Va., upland to Ga. and Tenn., W. Va., O., Ind., centr. 111. and la." It is not mentioned in Stcyermarks' Flora of Missouri, Jones and Fuller, in Vascular Plants of Illinois, report its most westerly appearance in that state a good deal to the north of us (Henderson County), its closest approach in our approximate latitude in Jasper County. Specimens in the Missouri Botanical Garden Herbarium, dated 1903 and 1907, with notations that they were collected at the Garden by John H. Kellogg, can be presumed to be material from plants cultivated on the Garden premises by Mr. Kellogg. This leaves us, then, with two anomalies: the first, did the single plant ob- served to be growing so happily in Missouri arrive there by natural means? It is growing in what is, admittedly, an unnatural — or artificially prepared — habitat: a bog created by walling in the bank of a stream just below a flowing spring. This is on the estate of Jay G, Rice, approximately 8 miles south of Antonia, Missouri, where Charles M, Rice built the wall and did introduce a number of plants before his death. However, Mr. Charles Rice died in 1950 and, to the best of my knowledge, the Skunk Cabbage did not appear for 17 years therafter; I am also entirely sure that no plants have been introduced since 1950. The second anomaly is the question of why the plant has not spread, even within the boggy area in which the single specimen is growing. The flowers are bisexual, and although proterogynous, Trelease reported (Amer. Nat., Sept 1879, p, 580) that, after the stigmas of the lower flowers have matured, **somc can evidently be fertilized by the mere falling of pollen from the upper flowers without any extraneous aid, for pollen is shed in such quantities that it covers the bottom of the spathe." Trelease's classic work on the pollination of this plant was carried on at Cornell, long before he came to Missouri — if the Skunk Cab- bage is now following in his footsteps, its pace is a very slow one. — Erna R. Eisen- drath, Missouri Botanical Garden, St. Louis. Ann. Missouri Bot. Card. 56(2): 287-291, 1969. [Vol. 56 288 ANNALS OF THE MISSOURI BOTANICAL GARDEN SUPPLEMENTARY NOTES ON THE PIIYTOLACCACEAE, I. A study of material at the British Museum and Kew Gardens has yielded a new species of LopJiiocarpus and the recognition of Vhytolacca polyandra Batalin. Lophiocarpiis latifolia Nowicke, sp. nov. Herha annua vel perennis, glabra. Folia lanceolata-elliptiea, ad 10 cm longa et 1 cm lata. — Fig. 1. Herbs, sparsely branched, slightly suffrutescent at the base, glabrous, to 1 m. Leaves lanceolate-elliptic, acute, entire, to 4 cm long and 1 cm wide, calcium oxalate crystals present; sessile or with petioles 2-3 mm long, lujlorescences spikes, up to 20 cm long, the larger terminal, the smaller axillary, flowers in clusters of 2-3. hlouers perfect, ± actinomorphic, yellow-green; the central flower subtended by a single bract, ca 1.8-2 mm long, weakly 3-lobed, the central lobe much elongate, the 2 lateral flowers subtended by a single bract ca 1.2 mm long and 2 bracteoles ca 0.8 mm long; sepals 5, ± equal, oblong and rounded, ca 2-2.5 mm long and 1 1.6 mm wide, thin, membranaceous; stamens 4, 3 alternate and one opposite, exserted, the filaments ca 2.2-2.5 mm long, the anthers at least 3x longer than broad, ca 0.8 mm long; ovary spherical, 1 -carpel- late (?), style absent, stigmas 4, thready, ca 1 mm long. Fruit an achene, ± globose, ca 1.8 mm long, weakly 8 ridged and ± muricate; seed one, the testa shiny black, the embryo large. Ilolotypc : Mozambique : louhenco marques : . Bcla Vista, betw Tcnonganinc and Floresta de Licuati, 12 Sept 1961, Lc7nos'£sr Balsinhas 263 (BM). Pollen grains single, subprolate, ca 2 3-24/i. (E) X ^^ 28-29/x (P) 3-colpate, colpi ca 17/x long, exine ca 2-2. 5/t in thickness, ca 3/t at the poles, sexine ± equal to the nexine and finely reticulated. Pollen examined: Lemos & BahiiiJias 263 (BM). Known only from the type collection, L. latifolia can be readily distinguished by its leaves which have a conspicuous blade in marked contrast to the remaining species of the genus, all of which have filiform or linear leaxes. PJiytolacca polyaudra Batalin, Acta llorti Petropol. 13: 99, 1893. (Syntypes: Potafiin s.n,, Beresoivski s.n. both LE) P. clavigera W. W. Smith, Bot. Mag. 149: pi 8 9 78, 192 3. Herbs, coarse and very robust, to 2 m. f eaves ovate to elliptic, acute to mucro- nate, entire, the bases obtuse to attenuate, to 32 cm lony and 13 cm wide. ± glabrous, the veins shglitly scabrous; sessile or with petioles ca 3 cm long. Inflorescences racemes, stout, up to 30 cm long, densely flowered, mostly axillary. Flowers with pedicels ca 5-11 mm long, ± winged at the base; bract single, ca 2 mm long, lanceolate; bracteoles two, ca 1 mm long, lanceolate; sepals 5, ovate-elliptic, ca 3-3.5 mm long and 2-2.5 nmi wide; stamens 12-16, arranged in 2 irregular whorls, the filaments ca 2.5 mm long, awl-shaped, the anthers ca 1 mm; ovary of 7-9 carpels, ± completely united or with apices slightly free, 1969] NOTES 289 f > . - 1 ^ .' 1 ^ *^^ t- - ■■'■ ■ "L--*^ V - It V : ^.■' ; ^" Ij^^ CiN'^^o 1 -- 'i^t fr t i^**S*fc^-^TV^ a ^a i^ >^ 4 . ry> t ^]'* n-iv. I Ij /^^m«tA ^^ i«*5":»tl4 .: "■ "«*^ fltf!Viftra«,*d** * : A, ^;trt- .^ r,-;"-!^.^^a= « *s C^l. . Fig. 1. Photograph of holotype of Lopliiocarpus latifoUa Nowicke. [Vol. 56 290 ANNALS OF THE MISSOURI BOTANICAL GARDEN the Styles as many as the carpels, ca 1 mm long, the stigma on the upper surface. Fruit a berry, ca 7-9 ribbed, ca 7-10 mm in diam, seeds 7-9, each ca 4 mm long, the testa shiny black. Specimens examined. China: Kansu, Beresowski s.n. (LE); Potanm s.n. (LE); SW Szechuan, Mts bctw Litang and Shou-Chu Rivers, bctw Wa-Erh-Dje and Gam, Rock 16738 (GH). Cultivated specimens from Kew Gardens (type of Phytolacca clavigcra W. W. Smith). Grown from seeds sent by Forrest from Yunnan, China (K). In an earlier paper on the Phytolaccaceae (Nowicke, Ann. Missouri Bot. Gard. 55: 294-364, 1968) 1 omitted treatment of this species because of in- sufficient material to establish its vahdity. Even at a first consideration, its dis- tinguishing characters, coarse, robust growth form and geographical location of China, appear as unsound bases for specific recognition, the more so in a genus which is noted for its plasticity. However, no other taxon in PJiytohcca even approximates the growtli form and dense inflorescences of P. polyandra except P. achiosa Roxb. which is easily distinguished by its separate carpels. — Joan W. Noiricke, Missouri Botanical Garden, and Department of Botany, Washington University, St, Louis, KALLSTROEMIA (ZYGOPHYLLACEAE) IN MISSOURI In his monumental Flora of Missouri (Iowa St. Univ. Press, Ames, 1963, p. 966), Stcyermark lists the Missouri taxon of Kallstroemia Scop, as K. hirsutis- sima Vail. Kallstroemia intermedia Rydb. is given as a synonym. Stcyermark discusses these tw^o concepts, indicating that some authors maintain them as separate species. During my studies on Kallstroemia [The genus Kallstroemia (Zygophyl- laceae). Contr. Gray Herb., in press], 1 have found that, indeed, there is a single taxon of the genus in the state, but that the correct name to be applied to it is Kallstroemia parviflora Norton, Kallstroemia intermedia is a synonym of K. parvi- flora, while K. hirsutissima is a species of the Chihuahuan Desert of the south- WTstcrn United States and adjacent Mexico. Kallstroemia parviflora is a species of the semiarid grasslands of the Southwest, introduced into Missouri along railroad rights-of-way. The following collections have been seen from the state: JACKSON CO.: Courtney, rare along railroad, Bush 767 (MO); Kansas Cit>', waste ground, Bush 8168 (GH, MO, NY, US), 8168A (CAS); Sheffield, Bush 7281 (GH, MO, NY). ST. LOUIS CO.: Allenton, Lettcrman s.n,, 1897 (MO); Meramec Highlands, Drushcl 5273 (MO). ST. LOUIS city: on railroads, Egocrt s.n., 1882 (NY), railroads, banks, Eg- gert s.n., 1883 (NY). Steyermark also maps Kallstroemia as occurring in Jasper, Jefferson, and Law- rence counties. — Duncan M, Porter, Missouri Botanical Garden, St, Louis, 1969] NOTES 291 H. L. Chakravarty, who published Lagenaria hicorniita as a new species in this journal (Ann. Missouri Bot. Gard. 55: 69. [Sept.] 1968.), had previously pubhshed Lagenaria hicornata [sic] (Bull. Inst. Fondam. Afrique Noire, Ser. A, Sci. Nat. 30: 412. [April] 1968.) based on the same type. ^Editor. Erratum: Ann, Missouri Bot. Gard. 55: 398, line 22. For "ca 1.5 longis" read "ca 1.5 mm longis/' The 1969 Jesse M. Grcenman Award was presented to Clifford M. Wetmore for his "Lichens of the Black Hills of South Dakota and Wyoming" (Publ. Mus. Michigan State Univ., Biol. Ser. 3(4): 209-464. 1967 [1968].). The Grcenman Award is given by the Alumni Association of the Missouri Botanical Garden for the best Ph.D. dissertation in plant systematics pubhshed during the previous year. The award honors Jesse M, Grcenman, Curator of the Herbarium from 1919 to 1943. Papers published in 1969 are now being con- sidered and may be submitted until 1 May 1970. Papers may be sent to David M. Gates, Director, Missouri Botanical Garden, 2315 Tower Grove Ave., St. Louis, Missouri 63110. — Editor. Edgar Anderson, who was associated with the Garden for over 45 years, died suddenly of a heart attack on 18 June 1969. — Editor. The previous issue of the annals of the Missouri botanical garden, Vol. 56, No. 1, pp. M12, was pubhshed on October 13, 1969, Published by the Missouri Botanical Garden Press St, Louis, Missouri 631 1 ANNALS VOLUME 56 1969 NUMBER 3 CONTENTS Tropical Island Biogeography: The Missouri Botanical Garden's Sixteenth Annual Systematics Symposium Duncan M, Porter . . . . . . Seasonal Flowering Behavior in Central Panama Thomas B. Croat . . . Studies in Neotropical Paleobotany. I. The Oligocene Communities of Puer- to Rico Alan Graham & David M. Jarzen Wood Anatomy of Goodeniaceae and the Problem of Insular Woodiness Shenvin Carlquist The Cahfornia Islands Robert F. Thome . . . Distribution Patterns of ^^^cst Indian Mosses Marshall R. Crosby . . . Drosophilidae of Hawaii Hampton L. Carson . . Vegetation of a Galapagos Island before and after an Ice Age Vaiil A. Colinvaux Pollen Morphology of Saprophytic Taxa in the Gentianaceae Siwert Nils- son & John Skvarla Cytotaxonomic Notes on Some Neotropical Gentianaceae Richard E. Weaver, Jr Spores of the Heterophyllous Selaginellae of Mexico and Central America Robert Lee Hellivig The O/ark Flora— Some Collections of Note W, G. D'Arcy NOTES ' ■ ' * Additional Type Material of Bouvardia (Rubiaceae) Will H. Black- well, Jr. & Marshall C. Johnston . . . . . A New Species of Scaevola (Goodeniaceae) from Tahiti Sheruin Carl- Chromosome Numbers of Phanerogams. 3. . ] . . ' ' Some new combinations in Protium (Burseraceae) " Duncan m' Porter 293 295 308 358 391 409 417 419 420 439 444 465 468 469 471 475 VOLUME 56 1969 NUMBER 3 ANNALS OF THE Missouri Botanical Garden The Annals contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden and the Department of Biology of Washington University. Papers originating outside the Garden or University will also be accepted. For information on prep- aration of manuscripts, see the inside back cover. The Annals appears three times a year, and three numbers, totaling about 400 pages, constitute a volume. For information con- cerning subscrijitions, see the back cover. Matters regarding exchange of publications are handled by the Library, Missouri Botanical Gar- rlpfi. 2^1 S Tnwer Grove Avenue. St. Louis, Missouri 631 10. Editorial Committee Marshall R. Crosby, Editor Missouri Botanical Garden Sheri G. Davis, Assistant to the Editor Missouri Botanical Garden Duncan M. Porter, Editor, Flora of Panama Missouri Botanical Garden 6- Washington University John D. Dwi:er Missouri Botanical Garden Qr St, Louis University Joan W. Nowicke Missouri Botanical Garden &- Washington University ( Publication of the Annals is aided by National Science Foundation grant GN-810 to the Missouri Botanical Garden, Published by the Missouri Botanical Garden Press, St. Louis, Missouri 63110. Printed by Bardgett Printing and Publishing Company, St. Louis, Mo. 63103. VOLUME 56 1969 NUMBER 3 ANNALS OF TH Missouri Botanical Garden TROPICAL ISLAND BIOGEOGRAPHY: THE MISSOURI BOTANICAL GARDEN'S SIXTEENTH ANNUAL SYSTEMATICS SYMPOSIUM^ Duncan M. Porter^ MacArthur and Wilson (1967: 3) have stated, "The study of insular bio- geography has contributed a major part of evolutionary theory and much of its clearest documentation. An island is certainly an intrinsically appeahng study object. It is simpler than a continent or an ocean, a visibly discrete object that can be labelled with a name and its resident populations identified thereby. In the science of biogeography, the island is the first unit that the mind can pick out and begin to comprehend. By studying clusters of islands, biologists view a simpler microcosm of the seemingly infinite complexity of a continental and oceanic biogeography. Islands offer an additional advantage in being more numer- ous than continents and oceans. By their very multiplicity, and variation in shape, size, degree of isolation, and ecology, islands provide the necessary replications in natural 'experiments' by which evolutionary hypotheses can be tested." Tropical islands were chosen as the subject of the Sixteenth Annual Systematics Symposium in part because of the above reasons. In addition man, as he continues to interfere with his environment, is rapidly decimating many tropical organisms (Porter, 1970), especially those on islands, where native species are less able to compete with disturbed conditions than are introduced weedy species. The Symposium was held in the Museum building of the Missouri Botanical Garden on Friday and Saturday, 17-18 October 1969. About 215 botanical ai zoological systematists, representing 62 institutions in the United States and Ca ada, attended. d mposium being directed by the Symposium moderator, Dr. Marston Bates, University of Michigan. Speakers and their topics were Dr. Hampton L. Carson, Washington University, '^Drosophilidae of Hawaii"; Dr. Paul A. Cofinvaux, Ohio State Univer- sity, "Vegetation of a Galapagos Island before and after an ice age"; Dr. Thomas B. Croat, Missouri Botanical Garden, "Seasonality in the flora of Barro Colorado ^The Symposium was supported by NSF Grant GB-14722. 2 Missouri Botanical Garden, 2315 Tower Grove Avenue, St. Louis, Missouri 63110. Ann. Missouri Bot. Gard. 56: 293-294. 1969. [Vol. 56 294 ANNALS OF THE MISSOURI BOTANICAL GARDEN Island, Panama"; Dr. Marshall R. Crosby, Missouri Botanical Garden, "Distribu- tion patterns of West Indian Cryptograms"; Dr. Alan Graham, Kent State Uni- versity, ^'Tertiary phytogeography of Puerto Rico"; and Dr. Terrell 11. Hamilton, University of Texas, "Adaptive radiation and the control of avian endemism in the Galapagos Archipelago." In addition Dr. Sherwin Carlquist, Rancho Santa Ana Botanic Garden, presented an e\ening talk on "Evolution in the Hawaiian Islands*' at Rebstock Hall, Washington University. The papers given by Dr. Croat, Dr. Crosby, and Dr. Graham, plus the ab- stracts of the papers by Dr. Carson and Dr. Colinvaux follow in this issue of the Annals of the Missouri Botanical Garden, The papers of Dr. Carlquist and Dr. Thorne are the result of discussions which took place during the Symposium, The Symposium is structured so that a maximum amount of time is available for discussion following presentation of the papers. The discussion periods were particularly rewarding, as they not only provided an opportunity for members of the audience to question the speakers, but also for them to review pertinent aspects of their own research. This situation encouraged a lively exchange of ideas and data, as investigators and their graduate students actively participated in the formal discussions. Both these discussions and the informal conversations that took place outside the formal meetings proved valuable to the participants. The stimulating papers presented by the speakers, the direction of the dis- cussions by Dr. Bates, and Dr. Carlquist's colorful evening talk all combined to produce a successful Symposium. Literature Cited MacArthur, R. H. & E. O. Wilson. 1967. The Theory of Island Biogeography. Princeton. Porter, D. M. 1970. The rape of Panama. Missouri Bot. Card. Bull. 53(1): 11-17. SEASONAL FLOWERING BEHAVIOR IN CENTRAL PANAi\L\ Thomas B. Croat ^ occurrin Abstract Based upon data obtained from study of herbarium material as well as from field observa- tions on Barro Colorado Island, species are enumerated according to seasonal flowering and fruiting times. Nearly the same number of species flower or fruit during the dry season months as in the wet season. The total number of species in flower or fruit in any one month varies considerably with flowering peaks occurring in February, July, and December. The December peak in flowering is caused largely by trees and vines and to some extent by shrubs and small trees. The July peak is caused by the flowering of both shrubs and small trees as well as herbs. The beginning of the rainy season is a period of maximum vegetative competition, and few species use this period for initiation of flowering. An understanding of the seasonal fluctuations of plants is essential to our study of ecology and evolution in the tropics. In an attempt to better understand the seasonal pattern of flowering and to establish the sequence of flowering for J on Barro Colorado Island, I have made a tabulation of seasonal records based on herbarium specimens. The data was obtained from studies made at the Field Museum, Chicago, and the Missouri Botanical Garden and is based on collections made in central Panama with heavy emphasis on Barro Colorado Island. This study of herbarium material follows eight months of observations on the island. Though this report is a preliminary one, the picture of seasonal patterns of flowering represented in central Panama is believed to be reasonably accurate. Barro Colorado Island, with an area of six square miles, is about three miles in greatest diameter. The island was formed by the damming of the Chagres River which resulted in the formation of Gatun Lake, a principal part of the Panama Canal route. It is centrally located in the Canal Zone about midway between the Pacific and the Caribbean. The topography of Barro Colorado Island consists of a mass of hills, the largest of which is 450 feet above the level of the lake or about 537 feet above sea level. The steep undulating terrain is traversed by over 20 miles of trails which make nearly all of the island accessible. However, since the trees are in general quite tall and the canopy closed for much of the year, collecting plants in the forest can be rather fruitless without the aid of climbing equipment. The island's approximately 25 miles of shorehnc is very irregular and has undergone drastic changes since its fonnation. The northern and eastern sides of the island undergo considerable erosion on account of wakes produced by ships and strengthened by trade winds. Many of the coves have silted in so that during the dry season, when the level of Gatiin Lake drops several feet, the deposits become tongues of land which arc invaded by a number of weedy species. This is particularly true of the more protected coves on the southern side of the island. It is here also, in the quiet waters of the southern and western edges of the 1 Missouri Botanical Garden, 2315 Tower Grove Avenue, St. Louis, Missouri 63110 Ann. Missouri Bot. Card. 56: 295-307. 1969. 296 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 island, that several aquatic associations of plants occur most frequently. These associations form the early stages of a hydrarch succession important in building the shoreline. Perhaps the most important of these is the Acrostic iim-Ann'ona glabra association. Although no major disturbances of the island's flora or fauna have taken place since 1923, when the island was set aside as a biological preserve, a major portion of the island was cutover 60-70 years ago. This cutting included much of the eastern third of the island, major promontories of the northern side of the island (including Orchid Island), and small portions of the western periphery. Most of the rest of the island is considered by some authorities to be covered with primeval forest. While I doubt this, it is nonetheless very old. The vegetation is thus primarily advanced secondary to nearly primary forest. It could perhaps be best referred to as semi-evergreen moist tropical forest. The island is rich in plant species — I have recorded 103 species of vascular crypto- gams and 1181 species of phanerogams. Dennis Knight, University of Wyoming, who has been making successional studies on the flora for the past several years, reports (personal communication) that even in what appears to be the most homogeneous areas of the island, the species-area curve for trees with greater than I inch dbh does not level off in as much as 1.5 hectares (about 3.7 acres). He reports that there are in such areas about 50-60 species of trees per acre. The island receives about 107 inches of rain in an average year, most of this falling between May and December (Fig. 1). Thus the climate is seasonal with a pronounced dry season beginning sometime in December and extending through January to April. Rains gradually build up to reach a peak in November. With the exception of wind and rainfall, other facets of the physical environment are remarkably invariable particularly at the forest floor. For example AUee (1926) reports that the total variation of monthly mean temperature during the year is only 1.5° F. The decrease in rainfall during the dry season is reflected in a marked in- crease in leaf fall. During the mid to late dry season, before leaves are re-initiated, the forest, with leaf litter piling up to a considerable depth on the floor, takes on a much difi^erent appearance than during the rainy season. This openness, coupled with the higher wind intensities of the dry season, have a beneficial effect on those species whose fruits are adapted for ^^ind dispersal. This will be discussed later. Despite the many objections which may be raised, the use of the herbarium for studying seasonality of flowering and fruiting will remain an important tool in our understanding of phenolog}'. This is particularh true for species which are either so rare or so obscure that firsthand observation of their seasonality is very limited. Lowland tropical rain forests ha\e many such species. It is only after the accumulation of many specimens that a composite picture of the seasonality of such species may emerge. Even tliis picture would be tentatixe, and only on the scene observation of many representatives of a species over a long period would present a definitive picture. 1969] CROAT FLOWERING IN CENTRAL PANAMA 297 17.9 11.7 11.0 10.9 3.4 2.2 ■■ 1 1.3 ^m J F M A M J J A S O N D Figure 1. — Monthly rainfall, in inches, on Barro Colorado Island. Based on 39 year data from the Meteorological and Hydrographic Branch, Panama Canal Company. One of the foremost requirements in using herbarium specimens for coming to conclusions concerning phenology is an adequate knowledge of the species in- volved. Identification of herbarium material is often clearly not to be trusted. It is also helpful to know the species from field experience. Knowledge of the appearance of the fruit at maturity, how the seeds are distributed, and how long a fruit of mature size persists on the plant are all important in evaluating herbarium specimens. There are many species wdiich have fruits collected all year but none of which are mature. Mature fruits are frequently not represented on herbarium specimens because they have fallen or been eaten or because of their size or soft nature. Even observation of some species in the field can be confusing. Species of Apeiba (Tiliaceae), for example, often have fruits on the tree for a full year, yet fruits with viable seeds are quickly removed by monkeys. Old fruits left on the trees by the monkeys probably contain no viable seeds. In the case of Liiehea (Tiliaceae) and a number of other species the valves of the fruit persist for some time after the seeds have been dispersed with fittle change in appearance. A chief disadvantage of using herbarium collections for seasonality studies is that the range of flow^ering time may be exaggerated, because a species may flower 298 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 03 o^ O m o OS oi fH . r^ ro I Ih if ;3 Ih 3 Ih * r^ fH o I 0^ Ih % O I I ! I I I I I 3 IH 3 IH • ^ I Ih O 0^ I Ih Ih o Ih o Ih 3 Ih IH o |i^ t-M m tin U. PM Ih Ih 03 o Ih 3 Ih * #v I Ih iH O^ 00 ro O^ CT. 00 ro Tf n] ro -^ in ^ Ti- o t I I I I I I I I I I iH I > IH T3 >> M 3 Ih 3 ^ Ih Ih 3 iH nj CO in in 00 rJ ON rj m ro CM •— t CN I I I I I I I I I O o 0; Ih o t -? >H OJ O [M m a; Ih O Ih o OJ O Jh CD o o 3 Ih c qj O ^ 00 o PLh [Jh PLh P-) 03 C o o: en in so r~ 00 0^ rMroT:tinvDi>ooa^ o 00 o in •-1 < H O H n3 Cfl Ih 03 c > 3 cd ^ i-i cd Ih C a; Ih Cd s 1969] CROAT FLOWERING IN CENTRAL PANAMA 299 abnormally early in one year and abnormally late in another year. On the other hand, the peak flowering period can be accurately determined in the herbarium, provided enough collections arc available. In originally attempting to categorize the seasons of flowering on Barro Colorado Island no more than a half dozen patterns were envisioned. It was found that the patterns are much more numerous and quite complex. In order to avoid competition, plants have altered their flowering periods to fill every possible niche. Species can be grouped into what I feel are 18 meaningful groups of biolog- ical importance. Table 1 depicts the seasonality types. The solid line indicates flowering period and the intermittent hne indicates fruiting period. The lines indicate the general spans of the various seasonality types and are not meant to depict any particular species. Thus, for example while all the species in season type 1 both flower and fruit within the dry season, they need not all have flowers and fruits simul- taneously. In many season groups several kinds of flowering patterns may be found. Individuals of some species ha\e a progressive development of their flowers and have small fruits which develop quickly. Thus they have flowers and fruits simultaneously throughout much of their flowering period. Good examples of such flowering behavior are found among members of the Gramineae, Cyperaceae, Musaceae, Marantaceae, Polygonaceae, Amarantaceae, Euphorbiaceae, Labiatae, Acanthaceae, and Compositae. More commonly individuals have distinct flowering and fruiting periods. Matu- ration of fruit for most species (even those with small fruits) usually occurs after most flowering has ceased. Species with large fruits usually require considerable time for fruit maturation. Consequently the time of flowering and the time of fruit maturation of any individual plant may be separated by a long period of time. Though perhaps most true of species having small or medium fruits, the stag- gered onset of flowering of difFerent individuals insures that both flowers and fruits are present on a population of plants for a considerable part of their flower- ing season. As a result the flowering season of a species is usually much longer than the flowering season of any individual of that species. Many species are found in flower or fruit over a relatively long period of time. There are 480 such species, about 41% of ah phanerogams, which are in flower for more than nine months of the year. These species generally have a definite peak of flowering and fruiting at some time during the year. It is quite possible that the large number of species flowering for nine months or more is merely a reflection of the total range of variation in flowering time as exhibited by spec- imens collected over a period of many years. Some of these species may be found to flower less than nine months in any given year. Being unable to decide which phenomenon was of more importance, the sea- sonal peak or the duration of flowering, those species in flower for greater than nine months were grouped into difl"ercnt classes as seen in the second column of figures of Table peak of a species season was the optimum period of flowering for that species and thus that this peak should be used to classify the species. In many cases the peaks were quite 300 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 pronounced with an insignificant number of individuals flowering or fruiting at other times of the year. Thus for this purpose, most of the original set of 18 flowering and fruiting patterns were duplicated for those species flowering more than during nine months. To the right is the total of both of these columns. From this data I had hoped to come to some conclusion as to which season was more important from the standpoint of species reproduction. The largest single category, disregarding the 161 species for which too little is known to establish seasonality, is the "non-seasonal" group — those species which flower and pea son (85 six'cies) (54 species). Tf species (139 versus 133) There are 49 species which are either strongly bimodal or which I suspect to be bimodal in their flowering pattern. Of these 2 7 flower more than nine months. While it is by no means easy to categorize the bimodal species, most of them flower once during the dry season (mostly mid dry season), then flower again sometime in the wet season (usually mid to late wet season). Another tendency for many of the bimodal species is for one burst of flowering to be smaller than the other. Chrysophyllum panamense (Sapotaccae) is a good ex- ample. Although verifying field work is required, Chrysophyllum panamense, on the basis of herbarium material, appears to have a few flowers from January July July fruits six months later during January through March. The period from April through June shows neg- ligible herbarium material with none seen for October through December. In the case of Hirtella triandra and particularly H. raceniosa there may be tri- modal flowering. Although this might be difficult to detect in the field, herbarium material shows three strong equidistant clusters of fruiting specimens of H. race- niosa. Hirtella triandra shows similar tendencies though not so strong. These species have been included with the bimodal species. Much more important than I would have originally thought are the "transi- tion" species— those which utilize more than one season for either flowering or fruiting. There are a total of 375 species which flower through such transitional periods, either dry to wet season or wet to dry season. There are 368 species which fruit in a similar manner. These figures are higher than for those species whose flowers or fruits are restricted to either the dry or wet season or for those which peak in either the dry or wet. In other words, there are more transition species than there are restricted species. Perhaps the transition species take ad- vantage of the benefits of both seasons— the openness of the dry season and moisture of the wet season. While no attempt has been made to define the actual triggering mecbanisms for flowering, there can be little doubt that the onset of the dry season acts in some manner to synchronize flowering of a large percentage of the island's flora. the Except for a small peak of flowering activity by some species during J 1969] CROAT FLOWERING IN CENTRAL PANAMA 301 well within the eonfines of the wet season, all major components of the vegetation have their major flowering activity sometime during the dry season. In all prob- ability flowering activity of species making up these peaks is due to a reaction to conditions of drought. In an attempt to establish a composite picture of seasonality a graph (Fig. 2 A) has been prepared of the total number of species collected in flower and in fruit for every month of the year. The fact that the fruiting specimens are less fre- quently collected than flowering specimens is reflected in their almost consistently lower values as depicted on the graphs. A species need have been collected only once to be reported for a particular month. The assumption is made that most species have been collected at least once during most of the months they are in flower or fruit. Although irregular collecting opens the possibility that more initial collections (i.e. species collected for the first time for a particular month) will be made at a particular period of time, it does not seriously affect the data. If one were attempting to establish the peak of flowering of a particular species based upon the total number of specimens, the effect would be a more serious one. Three major peaks of both flowering and fruiting activity, namely July, De- cember and February, are seen in the graph for all species collected in flower per month (Fig. 2A). There is no significant difference in the peaks of flowering and fruiting except when fruiting drops off in November and flowering increases. If the species flowering per month are separated according to type of habit (Fig. 3), the three points of maximum flowering or fruiting as seen on the composite graph are seen to be caused by different components of the vegetation. Since differences in seasonal flowering of monocotyledonous and dicotyledon- ous herbs appear to be insignificant (Fig. 2B) all herbaceous plants are included together for purposes of comparison with other types of habit. Though most of the palms have been excluded from the more typical herbaceous plants, the herbs remain a diverse group of plants. They include annual, perennial, and suffruticose species which may be erect, sprawling, vinelike, epiphytic, hemi-epiphytic, para- sitic, or aquatic. Included also are the palmettos and giant monocotyledonous herbs 2 A. TOTAL SPECIES 700 600 500 400 F LOWERING aoo 2B. F LOWE RING HERBS M O N OC O T I 150 t> I C O T I 100 N 7 Coal layer /9 eyzft Dar/c ^rej^ llynltic clcu/ Samples 14-/8 Grej/^ fine -grained llgnltlc ^and^lone (hits of curbofiized p(ani remains) \ t>tt Figure 4. — Stratigraphy, sequence of lithologies, and position of samples from Locality B, San Sebastian Formation, Puerto Rico. 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 315 Systematics A total of 165 morphological types of pollen and spores were recovered from the San Sebastian Formation. From this assemblage 44 (ca. 25%) have been identified to genus. In the following discussion the palynomorphs are cited accord- ing to locality (A, B, or C) and sample number (1—50). The photomicrographs are cited according to locality, sample number, and slide number (1-15), and coordinates of the England Slide Finder are also given. A summary of relative abundance is given in Table 1 and the species composition of the San Sebastian microflora in Table 2. Detailed descriptions are deferred to a later publication where the microfossils will be given formal taxonomie status. LycopodlH}}i (Fig. 5—6) The genus I.ycopoditim consists of approximately 100 species widely dis- tributed throughout tropical and temperate regions of the w^orld. The sjwres of many temperate forms and those most frequently reported from Ceno/oic deposits in the Northern Hemisphere are characterized by a distinct reticulum on the distal surface. Certain tropical species difl^er, however, in having slightly protruding, broadly blunt apices and/or a rugulate to less distinct reticulate sculpture pattern. The spores of Lycopodiiim Type A are similar to those of L. carolinianum. They were recovered from samples C-43 and C-50, with an abundance of 0.5%. The spores of Lycoyodiam Type B are most similar to the modern L. serratuui. Although we did not see collections of the plant from Puerto Rico, it does occur on Cuba and Haiti. Single specimens were recovered from samples C-48 and C-50. Selagiiiella (Fig. 7) Members of the genus Selaginella number about 700 species and are chiefly tropical in distribution, with a few ranging into temperate regions. At least three species are recorded from Puerto Rico, S. laxifolia, S. krugii, and S. suhstipitata. The spores are rare in the San Sebastian microflora and occur at Locaflties B and C. Cyathea (Fig. 8) The genus Cyathea is a tropical to subtropical assemblage of approximately 800 species. It is an arborescent fern of the wet tropics especially in the moun- tainous regions from Mexico to Chile. In the Antilles it occurs on all the islands and is represented by about 35 species including C. arhorea, C. horinquena, and C.puhescens. Cyathea arhorea is the most widespread and is abundant on Puerto Rico. The plants are common secondary invaders of impoverished soils in damp climates, especially at lower altitudes (Richards, 1966: 398). The fossil spores range in size from 35 to 55//., suggesting more than one species may be represented. They are present in Locality B, samples 15, 19, 25, and 33 with an abundance of 0.5-1.5%, and in Locality C, samples 41, 43, and 50 with an abundance of 0.5—1.0%. [Vol. 56 316 ANNALS OF THE MISSOURI BOTANICAL GARDEN Table 1. Per cent composition of identified taxa from San Sebastian Formation (Oligo- cene), Puerto Rico (based on 200 grains counted). P =^ present. Locality A Locality B Locality C Taxon 3 4 8 10 12 13 1 15 19 25 33 40 41 43 45 48 50 Lycopodium A .5 .5 Lycopodium B P P Sela^inella P P P Cyathca 1 .5 1.5 .5 .5 1 .5 Hemitelia 1 (Cnemidaria) P Jamcsonia (Eriosorus) Pteris (A,B,C) 5 .5 .5 P Podocarpus P P Ahutilon Acacia .5 .5 P Aetanthus Alchornea .5 1 1 1 .5 1 1 .5 .5 .5 .5 BcrnouUia .5 .5 5 2.5 1 Bomhax 1 Brunellia P 1 1.5 Bursera 1 i Casearia Catostemma .5 P 2.5 1 P 1 Chrysophyllum .5 P Corynostylis .5 .5 .5 Dcndropanax P En^clhardia 1.5 1 2 2.5 15 3 2.5 .5 .5 44.5 31 54 1 41.5 37 Eugenia .5 P .5 Fagiis I Faramea 1.5 P Guarea .5 P P .5 Hauya .5 P Ilex Jacaranda Liqtiidamhar 1.5 .5 2 .5 Marcgravia Merremia .5 Myrica 1 Norantea 1 P Nyssa .5 .5 1.5 4.5 .5 Oxalls PcUiciera 2 1 1 1 P P 1 P Pleodcndron .5 .5 .5 Rauivolfia Bhizophora 89 95 96 95.5 47.5 36 1 77 61.5 71 .5 .5 1.5 Salix 1 .5 Tecoma Tetrorchidium .5 1.5 2 .5 P 3 1 1 ,5 Tournefortia P Zanthoxyhim j .5 1.5 P Monolete Fern 1 Spores (Poly- podiaceae) 1 2 6.5 9 5 5 30 5 12 18 10 8 5 Trilete Fern 1 1 Spores 2 9 16 10 5 4 10 10 8 12 13 5 7 Palmae 1 2 3.5 3.5 8 5 7 50 2 5 15 9 12 12 Unidentified 4.5 1 2 3.5 32.5 34.5 48 1 15.5 8.5 11 24.5 8 6 19.5 31.5 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 317 Table 2. Systematic listing of identified genera occurring in the San Sebastian Forma- tion, Puerto Rico. Family : Lycopodiaceae Genus : Lycopodiutn Family : Selaginellaceae Genus : Selaginella Family : Cyatheaceae Genus: Cyathea Genus: Hemitelia (Cnemi- daria) Genus: Liquidambar Family : Oxalidaceae Genus: Oxalis Family : Rutaceae Genus : Zanthoxylum Family : Burseraceae Genus: Bursera Family: Meliaceae Family : Gymnogrammaceae Genus : Guarea Genus: Jamesonia (Eriosorus) Family: Euphorbiaceae Family : Polypodiaceae Genus: Pteris Family : Podocarpaceae Genus : Fodocarpus Family : Salicaceae Genus: Salix Family : Myricaceae Genus: Myrica Family : Juglandaceae Genus : Engelhardia Family: Fagaceae Genus: Fagus Family : Loranthaceae Genus: Aetanthus Family : Brunclliaceae Genus: BrunelUa Family : Hamamelidaceae Genus: Alchornea Genus : Tetrorchidium Family : Aquifoliaceae Genus: Ilex Family : Malvaceae Genus: Abutilon Family : Bombacaceae Genus: Bomhax Genus: Catostemma Genus: Bernoullia Family : Marcgraviaceae Genus : Marcgravia Genus : Norantea Family : Theaceae Genus: Pelliciera Family : Canellaceae Genus : Vleodendron Family Genus Family Genus Family Genus Family Genus Family Genus Family Genus Family Genus Family Genus Family Genus Family Genus Family Genus Family Genus Genus Family Genus Violaceae Corynostylis Flacourtiaceae Casearia Rhizophoraceae Rhizophora Myrtaceae Eugenia Onagraceae Hauya Araliaceae Dendropanax Nyssaceae Nyssa Sapotaceae Chrysophyllum Apocynaceae Rauwolfia Convolvulaceae Merremia Boraginaceae Tournefortia Bignoniaceae Jacaranda Tecoma Rubiaceae Faramea Previous reports of fossil spores of Cyathea include Hammen and Gonzalez (1960) from the Pleistocene of Colombia and Hammen (1963) from the Quater- Megafossils have been reported by Berry ( :, Hollick and Berry (1924) from the Plk ) (1968) from the Chuckanut Formation (Paleocene?) of northwest ern Washington. Th Hemitelia QCnemidaria; Fig. 9) of the tree ferns is currently being revised, and there are nomenclatural differences between the older and newer systems. According to some treatments the arborescent ferns belong to the genus Cyathea, while others recognize a number of genera. In a forthcoming revision ferns previously rec- ognized as Hemitelia will be called Cnemidaria (R. Tryon, personal communica- common to moist tropical mountains, and three ). The plants are species, including H. horrida, are native to Puerto Rico. They are occasional to & Wadsworth, ) and about 40 (Fig. 69) ports (1960) Miocene [Vol. 56 318 ANNALS OF THE MISSOURI BOTANICAL GARDEN ..*™''-^ ■>--v^-- 'v. V m^ 5 ( ' -^5i 6 • fj.^l 7 ■*flW^" ^.-. ^ C > ./ *M \ *«^ }^' %. ■ * Z*^ .- - ,.^ -^ 8 ^ % 9 10 >^ ■t 11 12 ■^«?. ■^ ^. 13 «k i ftft *' ■j^ 4 "Jl-yli. 14 4 8" "« # ft ^> « ■^ft-. «^ 15 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 319 Solo Formation of Veracruz, Mexico. Wolfe (personal communication, 1969) has megafossils, probably related to Asiatic forms, from Tertiary material in Alaska. (1924) Brazil, as H. hranneri. lifolia w (1928) of H. hranneri from the Collazo Shales of Puerto Rico. Jamesonia (Fig. 10) J (A, Try on, personal communication, 1969) J J about 19 species distributed from southern Mexico to central Bolivia and Brazil. It is a high altitude plant (1,500-5,000 m) of the paramos or other cool, wet highlands (Tryon, 1962); it characteristically grows at elevations higher than Eriosoriis. The genus Eriosoriis consists of about 30 species occurring from Andean South America, Brazil, Bolivia, and on the sandstone massifs of Venezuela north to Guerrero, Mexico. Eriosoriis hispidulus grows on the isolated Atlantic island of Tristan da Cunha (Tryon, 1966) and on El Yunque, Puerto Rico. It is a member of the wet cloud forest vegetation growing most commonly at elevations above 2,000 m. J was recovered from sample C-49. J (Hammen & Gonzalez, 1960), anc Tristan da Cunha (Hafsten, 1960) Pteris (Fig. 11-13) The spores of Pteris are common in the San Sebastian Formation. Several species are probably represented. Three relatively distinct types can be recognized, based on differences in sculptural elements on the distal surface. In Type A the sculpturing elements are a series of irregularly distributed sharp ridges; in Type B, low rounded ridges; and in Type C, verrucae. The genus consists of about 250 species. At least some grow under relatively xcrophytic conditions and may be primary invaders in dry barren regions (Rich- ards, 1966: 271). Spores of Pteris are common at Locahties B and C, being most abundant in samples C-43, 4 5, and 50, where they represent as much as 5% Figures 5-15. — San Sebastian (Oligocc 5. Lycopodium Type A, 40/i (C-50, 1; ESF Y-35). — 6. Lycopodium Type B, 40/i CC-50, 15; hbt U-21,3J. 7. Selaginella, 28fi (C-48,1; ESF N-28). — 8. Cyathea, 33/z (C-43, 15; ESF B-45)^— 9. Hemitelia, 42fi (C-48,1; ESF 0-31,1.-10. lamesonia (Eriosorus), 44^ (C-49,1; ESF K-25,2). — 11. Pteris Type A, 54/. (C-42,1; ESF S-38,4). — 12. Pteris Type B, 43/i (C- 44,5; ESF L-26). — 13. Pteris Type C, 45^ (C-46,2; ESF K-28,4). — 14. Acacia, 72/i (C-45,10; ESF J-38,3). — 15. Bernoullia, 59/i (C-48,10; ESF G-30). 320 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 . -^ 16 17 ^' ■Vi, x..^ - "-"v^ / 19 i t 20 ;^ 21 22 S& 23 24 27 -■^L _.^-' ..--l*^" 33 ,^«-»--^- ■ ^3^>fr i .- ''/ r / 34 \ 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 321 of the total assemblage. Spores similar to Pteris have been reported by Hammen and Gonzalez (1960) from the Pleistocene of Colombia as Triletes "tipo J." the Podocarpus (Fig. 16) The genus Podocarpus consists of about 100 si in (Fig. 70) and is the only native conifer. The plants grow in high-mountain, tropical forests, Cook, 1926: 115). It is a medium-size tree (ca. 10 m tall) nearly throughout the year (Little & VVadsworth, 1964). (Gl Podocarpus pollen is rare, in the San Sebastian Formation, with only 3 or 4 grains recovered (samples C-41, 42, 45). Previous reports arc those of Hammen and Gonzalez (1960) from the Pleistoccne-Holocene of Colombia; Tsukada (1966) from core samples of Lake Petenxil, Guatemala, absolute date (C^O 4,000 years before present; and Heusser (1966) from Late Pleistocene deposits in the Province of Llanquihue, southern Chile. The genus is also present in our samples from the Miocene Paraje Solo Formation of Veracruz, Mexico. Specimens of fossil wood and leaves of Podocarpus have been reported from the lowermost Miocene of Argentina (Berry, 1938), the Late Tertiary of Bolivia (Berry, 1939: 33), and the Eocene of western Tennessee (Dilcher, 1969). An account of the distribution of Podocarpaceae in Southern Hemisphere Cretaceous and Tertiary (1953) Abutilon (Fig. 20) (Malvaceae) 100 species of herbs and shrubs, widely distributed around the world. About 20 species occur in the Antilles with 4-5 in Puerto Rico. Species of Abutilon are adapted to a wide range of ecological conditions, including dry rocky slopes, stream- sides, open plains, and more tropical shaded forest conditions. A single grain of Abutilon was recovered from sample A-1. The fossil pollen is very similar to that of the modern A. incanum. This species is not found in Puerto Rico today, but grows in western Arizona, Texas, and southward into Baja California, Nuevo Leon, San Luis Potosi, and Sinaloa, Mexico, on dry rocky slopes (Shreve & Wiggins, 1964: 880). This distribution Figures 16-36.— San Sebastian (Oligocene) microfossils, Puerto Rico. — 16. Podocar- pus, 40 X 68/1 (C-41, 7; ESF U-37, 1-2). — 17. Actanthus 32/x (C-46,1; ESF H-41,3). — 18, 35-36. Alchornea, 22fi (C-45,10; ESF G-46,4). — 19. Brunellia, 24/* (C-42,15; ESF U-31,1). — 20. Ahutilon, 45/t (A-1,1; ESF H-36,1-2). — 21. Faramca, 32/: (C-46,6; ESF T-44,3). — 22. Bomhax, 43/: (B-14; ESF 0-17,2). — 23. Bursera, 29/i (B-17,5; ESF V-16,2. — 24-25. Catostemma, 33/x (C-43,1; ESF K-39,2). — 24. Outline.— 25. Surface.— 26. Casearia, 18X23/: (C-42,15; ESF S-36,4). — 27. ChrysophyUum, 13.5X18.9/: (C-41,1; ESF K-33,4). — 28-29. Corynostytis, 25/. (C-47,1; ESF P-22,2-4).— 28. Out- line.— 29. Surface. — 30-31. Dcndropanax, 25/: (C-46,15; ESF J-20,4). — 30. Outline.— 31. Surface. — 32. Guarea, 36/: (C-43,2; ESF U-42,1). — 33-34. Fagus, 39/: CA-1,1; ESF D-32,1). [Vol. 56 322 ANNALS or THE MISSOURI BOTANICAL GARDEN _xy^ ^1 .^*#%^ ^ «> ^ ■^ 37 « -IV ,j#" 38 39 i;. A^ ^ 41 J *.• • - 45 ■*\ f / 46 V , •1' 49 \ # \ ■'-^ \V± I; *' i z" 50 w^-j jtf ( »p \ XL. V ■i- 52 53 1969] GRAHAM & JARZEN — OLIGOCEiNE OF PUERTO RICO 323 pattern is somewhat different from otlier living eounterparts of tlie microfossils, but there is some megafossil evidenee that A. incanum may have had a wider dis- tribution in the past. Rollick (1928) described fossil fruits from the Collazo Shales of Puerto Rico as Malvocarpon clarum. He states that ''. . . generic relation- ship with Ahutilon is strongly indicated by the shape and arrangement of the carpels" (p. 214) and compares the specimens to fruits of the modern A. texense, a synonym of A. incanum Cfide Shrevc & Wiggins, 1964: 880). Other reports of megafossils of Ahutilon are Hollick (1936) from the Paleocene or Eocene of Alaska, and Berry (1925:217), w^ho described Malvacarpus tertiarins from Miocene deposits in Argentina and compared it to several modern genera includ- ing Ahutilon. Acacia (Fig. 14) Acacia includes some 700 species distributed in warm-temperate regions; the largest concentrations are in Africa and Australia. The number of native species in regions with wdiich the fossil assemblage has affinities is relatively fcnv (e.g., 1 species in Cuba, 17 species in northern Mexico; fide Shreve & Wiggins, 1964). The pollen of 60 species, including most of those from Latin America, were compared with the microfossils. The fossil is characterized by its exceptional size (72/^t) and scabrate cxine. These features are not common in modern species of Acacia, but are characteristic of A. farnesiana. This plant is a spiny shrub or small tree (3-4 m tall) of thickets and forests in the dry coastal limestone regions of Puerto Rico. Its geographic range is throughout I atin America and into south- em Texas, California, and Arizona, and through cultivation along the Gulf States to Florida. It is considered to be introduced in the Antilles, although there is some debate about its natural occurrence in Cuba. 1 he presence of A, farnesiana pollen in the San Sebastian microflora suggests it may be native to Cuba and other islands of the West Indies. Tsukada (1966) found Acacia polyads in Recent deposits from Guatemala, and Ileusser (1966) reported Acacia from Late Tleistocene deposits in southern Chile. Megafossils identihcd as Acacia also occur in the Pliocene floras of Potosi, Bolivia (Berry, 1939: 45). Aetanthus (Fig. 17) The genus Aetanthus (Loranthaceae) is presently restricted to the northern Andes. There are about 15 species which grow in temperate forests at elevations of about 2,000 m. A single specimen was recovered from sample C-46. The genus has not previously been reported from the fossil record. Figures 37-53. — San Sebastian (Oligoccnc) microfossils, Puerto Rico. — 37, Eauya, 85/i (C-42,9; ESF P-23,2).^38. ]acaranda, 50/z (C-46,8; ESF J-37,2). — 39-40. Marc- gravia, 19/i (C-42,15; ESF R-29,4). — 39. Outline.— 40. Surface. — 41. Eugenia, 27^ (C-45,3; ESFL-19),— 42. Ilex, 25^ (C-47,15; ESF H-19). — 43 44. Nyssa, 50/. CC-41,1; ESF D-37,4.— 43. Outline.— 44. Surface. — 45. Oxalis, 32^ (A-12,1; ESF P-32,1). — 46. Rauivolfia, 35/x (B-39,1; ESF C-45,4). — 47-48. Zanthoxylum, 13 X 18^ (C-50,15; ESF N-26).— 47. Outline.- 48. Surface. — 49. Pleodendron, 23/x (C-41,6; ESF E-24,4). 50. Engelhardia, 27/i (C-44,1; ESF U-24,3). — 51 52. Liquidaynhar, 35/t (C-43,1; ESF V-37,3). — 51. Outline. — 52. Surface. — 53. Norantca, 33/x (C-50,15; ESF N-30,4). [Vol. 56 324 ANNALS OF THE MISSOURI BOTANICAL GARDKN Alchornea (Fig. 18, 35-36) Members of the genus Alchornea (Eupliorbiaceae) are pantropical. There are about 70 widely distributed species (Fig. 71), represented in the New World tropics by a well-circumscribed section, Alchornea. Forty species occur in South America where they are common members of hydrosere successions along river banks and swamps. Species from Mexico and Central America are fewer in num- ber and commonly grow^ in rain forests, wet lowlands, and rocky montane areas. A single species, A. latifolia, growls in Puerto Rico today (Little & Wadsworth, 1964: 262). It is a medium-sized, rapid-growing evergreen tree attaining heights of 10—20 m. The trees are most common in the montane rain forests (e.g., Cerro de Punta), but they also occur in moist hmestone coastal regions, growing mainly in openings and along roadsides. This species also occurs on Cuba, Jamaica, Haiti (A. haitiensis also present), and Tortola; and from Veracruz, Mexico, south to western Panama (G. L. Webster, personal communication, 1969). Microfossils of Alchornea are common in the San Sebastian Formation, occur- ring in nearly every sample (0.5-1.0% ). They also occur in Pleistocene deposits from El Salvador and Guatemala (Tsukada & Deevey, 1967), Upper Cretaceous- Tertiary of Guyana (Hammen & Wijmstra, 1964, as Psilatricolporites opercula- tus^y in the Lower and Middle Eocene of Tibu, Colombia (Gonzalez-Guzman, 1967), and in our Miocene samples from the Paraje Solo Formation, Veracruz, Mexico. Bernoullia (Fig. 15) There are two species of Bernoullia (Bombacaceac) distributed from southern Mexico through Central America into the tropical regions of northern South Amer- (Fig. 72) (30 m tall) and are deciduous during the latter part of the dry season. Microfossils are common at Locality C with an abundance as high as 2.5%. There are no other reports of the genus in the fossil record. Bomhax (Fig. 22) Members of Bomhax are tropical and include about eight species in Mexico, Central and South America. The plants arc common in damp tropical forests. Microfossils of Bomhax are rare in the Puerto Rican samples, with a single spec- imen recovered from Locality B-14 and B-24. Previous reports include Hammen (1954, as Bomhacacidites annae') from the Lower Tertiary of Colombia, Norem (1955) from the Eocene of Venezuela, and Hammen (1963) from Quaternary deposits in Guyana. Tsukada (1966) also reports Bomhax from Recent core ma- terial of Lake Petenxil, Guatemala. Brunellia (Fig. 19) The genus Brunellia (Brunelliaceae) consist of 4 5 species distributed in the Neotropics from Mexico through Central America to Peru. It is represented in Puerto Rico by B. coniocladifolia, a medium-size evergreen tree (5-10 m tall) of the montane forests to elevations of 1,500 m or higher (Little & Wadsworth, 1969] GRAHAM & JARZEN — OLICiOCENK OF PUERTO RICO 325 1964). The range of B. comodadifulia is throughout the Greater Antilles, Guade- loupe, and Venezuela and Colombia. Microfossils were recovered from samples C-43 and C-48 (1.0-1.5%). The fossils are similar to several modern species of Brunellia, including B. comocladi- folia. The genus has not previously been reported from the fossil record. Eursera (Fig. 23) The genus Bursera (Burseraceae) is warm-temperate to tropical in distribu- tion, consists of about 100 species, and occurs throughout Mexico, Central Amer- ica, the Antilles, and northern South America (Brizicky, 1962fl). The western Mexican species have recently been monographed by McVaugh and Rzedowski (1965). The most widespread species is probably B. simaruha which grows through the entire range of the genus (Little & Wadsworth, 1964). In Puerto Rico this species is a medium-size deciduous tree 20-40 feet tall and is common in deciduous forests with soils derived from limestone. A single pollen grain was recovered from sample B-17. It is similar to a number of modern species including B. simaruha. Pollen of Bursera is also known from Recent core sediments from Lake Petenxil in northern Guatemala, Tsukada, ) (MacGinitie, 1953; cf 1965: 318) Casearia (Fig. 26) (Flacourtiaceae) tion. Some 40 species occur in Mexico and Central America; 90-100 grow in South America and about 25 in the West Indies (Fig. 73). At least seven species are native to Puerto Rico; C. arhorea, C. aculeata, C. hicolor, C. decandra, C. guianensis, C. ramiflora, and C. sylvestris. In Puerto Rico the genus is widely distributed along roadsides, thickets, and in the lower montane moist hmestone (10 m tall) the understory of moist tropical forests. Pollen of Casearia occurs only in samples C-42 and C-47, with an abundance of less than 0.5%. (1964) ( 106-107) reported leaf material from the Lower Miocene of Rio Pichileufu, Argentina, as similar to the modern C. arguta. In the same paper Berry states that the only previous report was that of Engelhardt from the Lower Miocene of Chile. Catostemma (Fig. 24-25) The genus Catostemma (Bombacaceae) is a New World group of six species ditributed in Guyana, Venezuela, and Brazil. The trees are most abundant in the tropical rainforests of Guyana where the average annual rainfall is over 250 cm. Here they reach heights of 30 m and are codominants in the canopy layer along with Mora and Eschweilera (Richards, 1966: 238-239). Pollen of the six species of Catostemma can be separated into two groups. One type (four species) is 4-6 stephanocolpatc with a densely and evenly per- forated tectum. The second type is 4-6 stephanoporate and echinate; C. praecox 326 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 and C. micranthum are inclucled lierc. The microfossils are of the latter type. Presently C. jnaecox and C. micranthum are restricted to the rain forests of Brazil. The abundance of Catostcmnia pollen in the San Sebastian Formation is 0.5% in sample A- 12, and it is common at Locality C. Pollen has also been reported from the Upper Eocene to earliest Oligocene of Guyana and is cited ) ( in tlie Oligocene and Lower Miocene. Other reports of fossil pollen of Catostemma are those of Hammen (1963) from the Quaternary of Guyana and probably that of Norem (1955) from the Upper Eocene of Venezuela, listed as "tetracolpate pollen." The presence of CatustcDima pollen in the Puerto Piican samples con- siderably extends the kno\\n geographic range of the genus during Tertiary times. ChrysofhyUum (Fig. 27) This large genus of Sapotaceae is distributed throughout the tropical regions of the world. It is especially common in the American tropics (Fig. 74). Species in the New World are found in the moist wooded and montane forests of Mexico, Central America, and throughout the West Indies; one species, C. oUriforme occurs in southern Florida (Wood & Channell, I960). Three species arc common to Puerto Rico and adjacent ii,hmds—C.argenteum, C. cainito, and C. oliviforme. These are evergreen shrubs or trees reaching heights up to 15 m (Little & Wadsworth, 1964). The trees are common in the moist limestone and lower montane forests, primarily in northern and western Puer- to Rico. Microfossils of Chrysophyllum are present in samples B-25 (0.5% ) and C-41 and C-47 as single specimens. The pollen of ChrysuphyUum is similar to some members of the Burseraceae, although pollen of the Burseraceae is usually about 50% larger. Hollick (1928) described two species and one variety of megafossils of Chrysophylhim from the Colla/o River area. He compared this material to C. argenteiim and C. cainito. Other reports of megafossils are Berry (1939: 154) from the Phocene of Vene/uela, and Berry (1938: 45) from the Eocene of Venezuela. Corynostylis (Fig. 28-29) (Violaceae) from Veracruz, Mexico, into South America (Fig. 75). The most wide-ranging species, C. arhorea, is a woody vine reaching tree girth. It is common along river banks in tropical forests. This species grows throughout the range of the genus, while the others are restricted to Brazil, Colombia, and Peru. Microfossils of Corynostylis occur in samples B-19, C-43, C-47, and C-50 with an abundance of 0.5%. There arc no other reports of Corynostylis in the fossil record. Dendropanax (Fig. 30-31) The genus Dendropanax (Araliaceae) occurs in tropical and subtropical re- gions throughout the world and includ(>s about 75 species. The American species (about 45) occur in Mexico as far north as the state of Tamauhpas, throughout 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 327 Central America, and into South America (Fig. 76). Deiulroyanax is common in montane forests and mainly consists of trees with a few shrubs. A medium-size evergreen tree reaching heights of 1 5 m or more, D. arhoreus is common in Puerto Rico. It grows in the lower montane and moist limestone regions of Puerto Rico and is common in the coffee plantations of the Central and Western Cordilleras. Another species, D. laurifolia, also grows in Puerto Rico but is not as common. The fossil pollen is rare, being recovered only from sample C-48 (0.5%). The only other fossil record of Dendrupatiax is by Dilcher and Dolph (1970), who reported leaves in Eocene deposits of western Tennessee. Engelhardia (Fig. 50) At present there are four species recognized in Engelhardia — E. pterocarya from Costa Rica; E. mexicana from Mexico, Guatemala, and Costa Rica; E. nicara- guensis from Nicaragua; and E. guatemalciisis from Guatemala. According to D. E. Stone these can be reduced to two, E. pterocarya and E. mexicana, the latter with three subspecies (cf. also Manning, 19 59). The genus has a discontinuous distribution along the cordillera from Orizaba, Mexico, to Boquete, Panama, and one collection is known from Colombia. EngeUiardia is not known from the West Indies. These medium-size trees commonly grow at elevations of 1,000-1,500 m in wet mountains. Fossil pollen of Engelhardia occurs at all locahties, and at Locality C it con- -50% of the samnles. Other reports include those of Traverse (1955) stitutes 30 (?) Brandon Lignite of Vermont; Wodehouse (1933) the Eocene Green River Formation of Colorado and Utah; and Langenheim et al. (1968) from Oligo-Miocene deposits in Chiapas, Mexico. It is common in the Paraje Solo (Miocene) Formation of Veracruz, Mexico. Pollen of the genus is also recorded in several presently unpublished theses and manuscripts on Mississippi Embayment and British Columbia deposits. If all these reports are valid, Engel- hardia formerly had an extensive geographic range and probably broader ecological tolerances. Reported megafossils are common in Tertiary deposits of the western United States (LaMotte, 1952). Engetiia (Fig. 41) goiia (Myrtaceae) contains se\ (1960). McVaugh (1963) stal 500 ac- have been referred to Eugenia. It is widely distributed throughout tropical and subtropical regions. About 150 species occur in Mexico and Central America, and at least 135 are found in South America, concentrated in Brazil. Eighty or more (F ) (10-20 m tall) found in the montane forests at elevations up to about 1,000 m. (0.5%) (1960) (1962) and Tsukada (1966) from Recent core samples of Lake Petenxil, Guatemala. [Vol. 56 328 ANNALS OF THE MISSOURI BOTANICAL GARDEN Megafossils of Eugenia have been reported by Rollick (1928) from the Oh of Puerto Rico, Berry (1929) from the Loja Basin (Late Miocene) of Ec Berry (1937: 59) from the Uppermost Miocene of Trinidad, and Berry ( ) Fagiis (Fig. 34-35) Vagus, with about ten species, is mostly north temperate in distribution with species tion of 1,800 m. An account of the distribution of Vagus in Mexico is given by Fox and Sharp (1954). Fossils are present in sample A-3 with an abundance of 1.0%. Pollen of six species was compared to the microfossil, and it is most similar to F. americana. Faramea (Fig. 21) (Rubiaceae) (Fig. 78) (3-10 m tall) and is common in the understory of moist limestone and lower mon- tane regions. The pollen of Faramea is dimorphic (Willis, 1966: 444), with 2- and 3- porate grains. The pollen of F. occidentalis is typically 3-porate, as is the micro- fossil. The pollen is present in samples B-25 (1.5%) and C-41 (less than ) Gnarea (Fig. 32) ( ) and some 20 species in Africa. In Puerto Rico they are shrubs (ca. 2-3 m tall) to large evergreen trees (15-25 m tall) ecies // // n fi 7 (Gleason & Cook, 1926) seedlings of this Microfossils were recovered from Localities B and C (0.5% ). Hollick (1928) reported G. opinahilis from the Ohgocene of Puerto Rico, and Hollick and Berry (1924) described leaves of the genus from the Late Tertiary of Brazil and re- ferred them to the modern G. trichilioides. Graham (19621?) transferred fruits described as Ficus ceratops from Upper Cretaceous beds of Wyoming to Guarea. Hauya (Fig. 37) (Chiap mala), H. microcerata (Chiapas south to Honduras and El Salvador), and E. elegans (Hildago and San Luis Potosi, Mexico, south to Costa Rica). Hauya glauca is considered a subspecies of H. microcerata (Raven, personal communication, 1969). The pollen of other genera closely related to Hauya (Calylophus, Clarkia, and Gaura') was examined and found to be distinct from Haura. 1969] GRAHAM & JARZEN^ — OLIGOCENE OF PUERTO RICO 329 Ecologically Hauya occurs with oaks and other lush middle elevation vegeta- tion. It does not range upward into the pine belt or downward into the seasonally drier deciduous vegetation. It is always rare, and phylogenetically is one of the most primitive genera of Onagraceae (Raven, personal communication, 1969). During the Oligocene Hauya had a distribution considerably different than at present, and it grew in the West Indies where it no longer occurs. Microfossils were recovered from Locahty C (less than 0.5% ); there are no other reports of Hauya in the fossil record. Ilex (Fig. 42) The genus Ilex is widespread and includes about 400 species. At least 60 grow in Mexico, Central America, and the West Indies; and some 180 species occur in South America. These trees and shrubs occupy a wide variety of habitats from wet, tropical rain forests, pine forests, marshy thickets and swamps to moun- tain slopes. Four species are known from mountain slopes and wooded valleys of Puerto Rico — I. niacfadyenii, J. nitida, J. ohcordata, and T. sintenisii. A single specimen w^as recovered from sample C-47. Previous reports from Latin America are those by Hammen and Gonzalez (1960) from Upper Pleisto- cene and Holocene deposits near Bogota, Colombia, Hammen (1962) from Recent sediments (5,000 yrs.) of Laguna dc los Bobos, Colombia, Hammen (1963) from the Quaternary of Guyana, and Tsukada (1966) from core ma- terial of Lake Petenxil, Guatemala. Jacaranda (Fig. 38) There arc about 50 species of lacaranda in Latin America icaranda in Latin America (Fig. 80). J. mimosifolia, is introduced from Argentina ( ) w^ooded hillsides. The pollen of Jacaranda is similar to Tecoma Qcf, Fig. 38, 58), but the for- mer is considerably larger. The single microfossil is present in sample C-46. There are no other reports of the genus in the fossil record. Liquidamhar (Fig. 51-52) Eastern Asia (Ernst, 1963). The species from the eastern United States, L. styracijlua, is common in deciduous forests along river flood plains and other mesic temperate habitats. It ranges into Mexico and Honduras (as L. macrophyUa or L. styracijlua and grows in the montane forests at elevations of 1,000-2,000 m. ) Mic 0.5-2.0%. Tsukada (1966) poll (1960) (LaMotte, 1952) MarcQravia (Fig. 39-40) The genus Marcgravia (Marcgraviaceae) occurs in Central and South America and the West Indies (Fig. 81). There are about 55 species of these climbing, 330 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 ^- 55 '^- ' ■i'- 64 54 'K ^^ ;^1 .^lp> 57 ■*'>X*»^ ^ 67 -'^^ 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 331 epiphytic shrubs, including M. sintenisii and M. rectiflora. The latter is common in the rain forests of Puerto Rico and under favorable conditions the adult plant, 2-3 m in diameter, becomes a dominant feature of the forest. occur rence is consistent with the small amount of pollen produced by the cleistogamous flowers of some species. The genus has not previously been reported from the fossil record. Merremia (Fig. 55) Merremia (Convolvulaceae) grows in warm temperate regions throughout the world (Willis, 1966). There are about 80 species, and 40 of these are found in the neotropics (Fig. 82). The plant is a herbaceous liana, commonly an early invader in secondary successions (Richards, 1966: 393). Eight species grow in the West Indies occupying open places, bluffs, and dry lowland habitats (M. aegyptia, M. data, M. cissoides, M. dissecta, M. qmnqiiefolia, M. tridentata, M. tuherosa, and M. umhellata'). The single microfossil specimen from sample C-42 was compared to pollen of species presently growing in the Antilles and is most similar to M. dissecta. Fossil pollen has also been reported by Gonzalez-Guzman (1967) from the Lower and Middle Eocene of the Tibu region in Colombia. Myrica (Fig. 62) The genus Myrica is almost cosmopolitan in distribution with 35 known species. There are about 17 species of these trees and shrubs widely distributed throughout Mexico, Central America, and the Antilles. An additional 1 5 or more are known from South America. Two connnon members of the Puerto Rican flora defl all). These are medium-size, evergreen trees ( nearly all the islands of the Antilles. A single microfossil was recovered from sample C-42. Previous reports of fossil pollen are those of Hammen and Gonzalez (1960) from the Upper Pleistocene and Holocene of Colombia, Hammen (1962) from Recent sediments (5,000 yrs.) from Laguna de Los Bobos, Colombia, and Tsukada (1966) from Lake Petenxil, Guatemala. Norantea (Fig. 53) The genus Norantea (Marcgraviaceae) includes about 3 5 species of the American tropics, being most abundant in South America (Fig. 83). They are hanas which superficially resemble Vhilodendron (Willis, 1966: 777). "Figures 54-68.— San Sebastian (Oligocene) microfossils, Puerto Rico. — 54, 66-68. Pelliciera CC-48,1; ESF D-38,2).— 54. 72/£.— 66. 55/i.— 67. Outline, 54/;.— 68. Surface, 54^. _ 55. Merremia, 63/i (C-42,9; ESF P-41,3). — 56-57. T etror chidium , 32^, (B-14,10; ESF M-22,1).— 56. Low focus.— 57. High focus. — 58. Tecoma, 27/* (€-48,1; ESF U-20,4). 59. Tournefortia, 28m (A-12,10; ESF U-34,4). — 60-61. Salix, 21 ii (C-46,15; ESF L-30,3).— 60. Outline.— 61. Surface. — 62. Myrica, 20/i (C-42,4; ESF N-43). — 63-65. Rhizoph 63. 14/i. — 64. 24/1.-65. 28/i. [Vol. 56 332 ANNALS OF THE MISSOURI BOTANICAL GARDEN A single specimen was recovered from sample C-50. There are no previous reports of Norantea from the fossil record. Nyssa (Fig. 43-44) (Nyssaceae) ) (Eyd Maine, the Great Lakes, and southern Canada south to the Brazos River in Texas and Lake Okeechobee in Florida. The latter also occurs in the highlands of Hidalgo, Puebia, and Chiapas, Mexico, at elevations of 1,000-1,700 m. It is frequently associated with Ijquidanihar, Qiiercns, and Pinns strohiis. Miranda (1945) has discussed the presence of f^yssa in Mexico. Microfossils from the San Sebastian Formation are present at Localities A and C and are most common in sample C-48 (4.5% ). The fossil record of Nyssa is extensive (Eyde, 1963; Eyde & Barghoorn, 1963); however, we know of no other records from northern Latin America. Oxalis (Fig. 45) The genus Oxalis is widely distributed, consisting of about 800 species. It is particularly abundant in Central and South America. At least 20 species grow in the Antilles as small herbs of diverse habitats. Microfossils of Oxalis are rare occurring only in sample A-12 (less than 0.5%). There are no previous reports of the genus in Tertiary deposits of Latin America; Brown (1938) has reported Oxalis from the Pleistocene of Louisiana, the only other record know^n to us. Pelliciera (Fig. 54, 66-68) The genus Pelliciera (Theaceae; Pellicieriaceae fide Willis, 1966) is repre- sented in the modern flora by a single species, P. rhizophurae. It is a small tree, 8-10 m tall, with buttress roots and a mangrove habit. It occurs along sandy beaches, swamps, and mud flats, frequently associated with other mangrove ith through (Fig. 84) (Colombia, Ecuador) The present range of Pelliciera is considerably restricted when compared to its former occurrence. Wijmstra (1968) has reported pollen from Oligocene to Miocene beds of the Guiana Basin of northern South America. Wijmtsra also cites references to other occurrences in the Marago Basin of Brazil (Miocene) and from a Miocene pollen flora in Venezuela. Langenheim et al. (1967) have identified Pelliciera pollen from Oligo-Miocene beds in Chiapas, Mexico. It is also present in our samples of Middle Tertiary age in the Canal Zone, Panama. The fossil record of Pelliciera is from the Middle Oligocene to the Present, and these sites are distributed as far north as Chiapas, Mexico and Puerto Rico. The pollen recovered shows considerable diversity in size and morphology. The grains range in size from 50-90/^ and include coarsely scabrate to verrucate forms. The same variation is evident in pollen of the modern P. rhizophorae. 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 333 Figures 69-70. — Present Latin American distribution of genera identified from the San Sebastian (Oligocene) Formation, Puerto Rico. — 69. Hemitelia.— 70. Podocarpus. [Vol. 56 334 ANNALS OF THE MISSOURI BOTANICAL GARDEN Vleodendron (Fig. 49) The genus Pleodendron (Caiicllaceae) with two species is restricted to the West Indies. Vleodendron ekmanii is known only from wooded hillsides in Haiti at elevations of 800 ni, and P. micranthum, a tall tree of virgin forests, occurs in the Sierra de Luquillo, Puerto Rico, at elevations of about 1,200 m. Microfossils of Pleodendron are present in samples C-41, 43, and 50 (0.5% ). There are no other fossil records of the genus. Rauwolfia (Fig. 46) Rauwolfia (AixK-ynaceae) consists of about 100 species distributed through- out tropical regions of the world. There are 15-20 species in Mexico and Central America, about 40 species in South America, and 10-15 from the Antilles. In Puerto Rico the genus is represented by R. nitida, a small tree or shrub (7-14 m tall), and R. viridius, a shrub about 1-4 m tall (Fig. 85). The plants grow in tlie coastal and limestone forests and thickets, usually in open habitats. A single specimen was recovered from sample B-39. There are no other fossil records. Rhizophora (Fig. 63-65) The genus Rhizophora contains 6-9 species distributed throughout the world along tropical coasts. The plants are gregarious trees or shrubs inhabiting mud flats of coastal tidal marshes, brackish streams and lagoons (S. Graham, 1964). Two species grow in areas with which the microflora has affinities— R. raceniosa, infrequent in the Americas but more common along the African shore, and R. mangle which ranges from southern Florida to northern South America. The pollen recovered includes as extremes a small, psilate type with distinct colpi transversales and a larger form with a scabrate surface pattern and less distinct colpi transversales. According to Langenheim et al. (1967) these varia- tions are included within the pollen of 7^. mangle. Fossil pollen of Rhizophora is found at all localities witli an abundance of 96% at Locality A, 77%, at Locality B, and 1-2% at Locality C. Latin Amer- ican records of the genus are common and include those of Hammen (1963) from the Quaternary of Guayana and Langenheim et al. (1967) from the Ohgo- Miocene of Chiapas, Mexico. It is also present in our samples from the Miocene of Veracruz, Mexico, and the Middle Tertiary (Oligo-Mioccne to Middle Mio- cene) of Panama. Hollick (1928) records megafossils from the Oligocene of Puerto Rico. Salix (Fig. 60-61) The genus Salix includes about 500 species, chiefly north temperate in dis- tribution, but with at least 1 5 species occurring in Mexico, Central America, and the Antilles. An additional ten species are known from South America. Species of Salix range from trees to shrubs and subshrubs, and although most common in moist, mesic environments, they occupy a wide range of habitats. Microfossils were recovered from samples C-43 (1%), C-46 (0.5%), and C-48 (0.5%). We know of no other reports of the genus from Latin American deposits. 1969] GRAHAM & JARZEN^ — OLIGOCENE OF PUERTO RICO 335 Figures 71-72.— Present Latin American distribution of genera identified from the San Sebastian (Oligocene) Formation, Puerto Rico. — 71. Alchornea.— 72, Bernoullia. [Vol. 56 336 ANNALS OF THE MISSOURI BOTANICAL GARDEN Figures 73-74.— Present Latin American distribution of genera identified from the San Sebastian (Oligocene) Formation, Puerto Rico. — 73. Casearia. — 74. Chrysophyllum. 1969] GRAHAM 8r JARZEN — OLIGOCENE OF PUERTO RICO 337 CORYNOSTYLIS Figures 75-76.— Present Latin American distribution of genera identified from the San Sebastian (Oligocene) Formation, Puerto Rico. — 75. Corynostylis. — 76. Dendropanax. 338 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 \ \ \ l^ -^ I I ) c:r — "^ ^■> ^ / V / \ < l-p \ ^ > /- \ /' yA \ i Figures 77-78.— Present Latin American distribution of genera identified from the San Sebastian (Oligocene) Formation, Puerto Rico. — 77. Eugenia. — 78. Faramea. 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 339 Figures 79-80. — Present Latin America distribution of genera identified from the San Sebastian (Oligocene) Formation, Puerto Rico.— 79. Guarea.— 80. Jacaranda. 340 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 MARCGRAVIA Throughout Mexico & C. Am. MERREMIA Figures 81-82.— Present Latin American distribution of genera identified from the San (Olig 81. Marcgravia. — 82. Merremia. 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 341 PELLICIERA Figures 83-84. — Present Latin American distribution of genera identified from the San Sebastian (Oligocene) Formation, Puerto Rico. — 83. Norantea. — 84. Pelliciera. 342 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 Figures 85-86.— Present Latin American distribution of genera identified from the San Sebastian (Oligocene) Formation, Puerto Rico. — 85. Rauwolfui. — 86. Tctrorchidiiun. 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 343 Tecoma (Fig. 58) Tecoma (Bignoniaceae) ranges from Florida to Mexico, Argentina, and the West Indies. There are about 16 species and one, T. stans, is native to Puerto Rico* This species is a small evergreen tree (ca. 8 m tall) and has an extensive geographic range. Pollen of Tecoma is rare, occurring only at Locality C (0.5% )• There are no other fossil records. Tetrorchidium (Fig. 56—57) The genus Tetrorchidium (Euphorbiaceae) consists of 16 species distributed in Central and South America, the West Indies (but not Puerto Rico), and tropical west Africa (Fig. 86). Gomcz-Pompa has recently found the genus in the San Andres Tuxtla area of Veracruz, Mexico (W^ebster, personal communica- tion, 1969). The plants commonly grow at the upper limits of the montane rain local (1-3%) Toiirnefortia (Fig. 59) This genus of Boraginaceac is a tropical to subtropical assemblage of about 150 species. At least 130 are known from the New World, and a number occur on the islands of the West Indies (Fig. 87). These small trees and shrubs are common along moist coastal regions where they are often an important feature of primar)^ succession (Richards, 1966). Microfossils were present only in sample A-12 and were rare. There are no other fossil records. Zanthoxylum (Fig. 47—48) The genus Zanthoxylum (Rutaceae) is a widely distributed assemblage of 20—30 species in temperate and subtropical regions (Fig. 88). At least seven species grow in Puerto Rico — Z. flarrum, Z. carihaeum, Z. martinicense , and Z. monophyUtim being common. These medium-si/e deciduous trees (7— 17 m tall) were common on Puerto Rico and adjacent islands, but because of their highly valued timber they are now scarce and restricted to protected forests (Little & Wadsworth, 1964). In its original habitat Zanthoxylum was widely distributed in the coastal limestone and lower montane forests, chiefly of western Puerto Rico. Gleason and Cook (1926) also reported the trees as common in mesophytic coastal forests. Pollen of Zanthoxylum is present at Localities B and C with an abundance of 0.5-1.5%. Berry (1923, 1925, 1939) described Fagara wadii from the Miocene of Oaxaca, Mexico, and F. miocenica from Trinidad and Cuba. *'The occurrence of s|>ecies apparently transitional in the character of the perianth between Zanthoxylum and Fagara is ample reason to regard both as components of a single genus*' (Brizicky, 1962^; 7). The genus Fagara is also listed as a synonym of Zanthoxylum in LaMotte's (1952) catalogue. Other Microfossils: In addition to these genera, 120 other types wxre re- covered. Many are referable to family, while others can only be placed in an artificial morphological category. Some of the forms represented are numerous fern 344 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 TOURNEFORTIA Figures 87-88. — Present Latin American distribution of genera identified from the San Sebastian (Oligocene) Formation, Puerto Rico, — 87. Tournefortia. — 88. Zanthoxylum. 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 345 spores similar to many genera of Polypodiaceae, about 1 5 types of palm pollen, and representatives of Bombacaceae, Burseraceae, Caryophyllaceae, Ericaceae, Eu- phorbiaceae, Gramineae, Magnoliaceae, Malpighiaceae, Nyctaginaceae, Oleaceae, Polygonaceae, Proteaceae, Rubiaceae, Theaceae, and Tiliaceae. Fungal spores were also common, but in the absence of adequate reference collections, and with little ecological data available for tropical fungi, identification and environmental inter- pretations were not possible (Graham, 1962fl). Discussion Initial studies of vegetational history in neotropical regions are likely to yield a miscellaneous assemblage of data difficult to integrate into a documented sum- mary of paleoenvironments. Stratigraphic control and environmental trends estab- lished through studies of sequential fossil floras provide the necessary context for interpreting paleontological data, and this context is not available for northern Latin America. The San Sebastian assemblage is the only fossil flora known from the Antilles, and it is the only Middle Oligocene flora from Latin America for which recent published information is available. Until other studies are completed only qualified interpretations may be made of geologic events and vegetational response. New Fossil Records, — One category of data from the Puerto Rican studies is reports of genera with no previous known fossil record. These include Aetanthus (Loranthaceae), Bernoullia (Bombacaceae), Brunellia (Brunelliaceae), Cory- nostylis (Violaceae), Dendropanax (Arahaceae), Faramea (Rubiaceae), Hauya (Onagraceae), ]acaranda and Teconia (Bignoniaceae), Marcgravia and Norantea (Marcgraviaceae), Fleodendron (Canellaceae), Rauwolfia (Apocynaceae), Tet- rorchidiiim (Euphorbiaceae), and Tournefortia (Boraginaceae). These 15 genera constitute 35% of the identified microflora. Oligocene Communities of Puerto Rico. — Thirteen genera (ca. 30%) of the San Sebastian microflora no longer grow in Puerto Rico and represent extinctions after Middle Oligocene times. Three of these ^Norantea, Tetrorchidium, Bomhax^ presently grow in the West Indies but not on Puerto Rico; two QAetanthus, Catos- temma) are South American; two are Mexican and Central American QHauya, Engelhardia^^\ and three occur in temperate regions of the eastern United States and eastern Mexico (Fagus, Liquidambar, Nyssa). The remaining three genera (Bernoullia, Cory nostylis , Pelliciera) are widespread in Latin America except the Antilles. Changes in the composition of geofloras through time is common, particu- larly when ecologically comparable taxa are involved. With the exception of the cool-temperate element, these difEerences in composition between the San Sebas- tian microflora and the present vegetation of the West Indies do not imply sig- nificant climatic changes. ^A single collection is known from Colombia (D. E. Stone, personal communication, 1969), [Vol. 56 346 ANNALS OF THE MISSOURI BOTANICAL GARDEN The arrangement of the OHgoeene genera into eommunities is tentative because of meager ecological data on the modern forms. An estimate of the associations and their composition is presented in Table 3. At least three communities were present: 1) a lowland, coastal brackish-water community of Rhizophora and Pelliciera; 2) an upland warm-temperate to subtropical community, with several members common along river banks and lake margins and others which are still part of the present elfin or mossy forest of Puerto Rico; and 3) a highland tem- perate to cool-temperate community of Vagus, Liquidamhar, and Nyssa, Several genera w^re probably common to more than one association (e.g., Ilex, Tecorna, ]acaranda, Bursera, and Bernoullia^. Since approximately 75% of the genera identified presently grow on Puerto Rico or in the West Indies, the aspect and composition of the Oligocene commu- nities were similar to those of the present. There were, however, three principal differences. There was the group of seven genera that no longer grow in the West Indies, but which are found in ecologically similar habitats elsewhere in Latin America QAetanthus, Catostemma, Hauya, Engelhardia, Bernoullia, Corynostylis, Pelliciera^. A second difference w^as that of the above group, Engelhardia was particularly common. At Locality C pollen of this genus constitutes 31-54% of the microflora (Table 1), indicating it was an abundant and conspicuous element Table 3. Suggested 01igoa?ne communities of Puerto Rico. Asterisk C*) designates plants presently growing in the elfin forest of Pico del Oeste, elevation 1050 m (cited by Howard, 1968). 1. Coastal brackish-water community. Rhizophora Pelliciera 2, Upland warm-temperate to subtropical community. '^Cyathea Hemetelia (=^Cnemidaria) Jamesonia Cor Eriosorus) * Lycopodium * Sclaginella Podocarpus Alchornea * Bernoullia Botnbax Brunellia Bur sera Casearia Catostemma Chrysophyllum Corynostylis Dendropanax EngeJhardia * Eugenia Far ante a Guarea Hauya "Ilex Jacaranda Marcgravia Merremia Norantea Pleodendron Tecoma Tetrorchidium Tournefortia Zanthoxylum Csi>ecies also common along river banks and lake margins) Alchornea Casearia Corynostylis Rauwolfia Salix Tournefortia Zanthoxylum 3. Highland temperate to cool-temperate community. Eagus Liquidamhar Nyssa "Lycopodium '^Selaginella (genera possibly ranging into the higher altitudes) ""Ilex Myrica 1969] GRAHAM & JAR2EN — OLIGOCENE OF PUERTO RICO 347 that has been eliminated from the flora since OHgoccne times. The other six genera were less common (0.5—2.5%). The third and climatically the most sig- nificant difference was the presence of a temperate to cool-temperate element CFagus, Liquidambar, Nyssa'). The presence of these plants is important in con- siderations of paleophysiography and will be discussed in a later section. The gymnosperm element present today on extreme highlands in the Greater Antilles is absent from the microfossil assemblage. In the Dominican Republic the elfin or mossy forest is replaced at the highest elevations by a forest of Pinus occidentalis. In view^ of the prolific pollen production of pine, it is likely that pollen from such a community would be incorporated into the accumulating sedi- ments. The only gymnosperm pollen recovered was a few grains of Podocarpus (Table 1). Hollick (1928) reports only Zamia from the megafossil flora of Puerto Rico. These data do not reveal the time of introduction of the high-altitude gym- nosperm element in the Antillean flora, but this apparently took place in post- Oligocene times. In addition to the microfossils, Ilollick (1928) reported 56 genera of mega- fossils from the Collazo Shales (Tabic 4). Several were tentative identifications, and others need verification. Consequently the present megafossil record is of limited use in environmental reconstructions. Pollen of six megafossil genera was recovered (^Hemitelia, Chrysophyllum, Eugenia, Guarea, Malvocarpon [cf. Abu- tilon], and Rhizophora), and others C^actris, Palmacites, Palmocarpon, Cassia, Inga, Sophora^ may be among microfossils presently identified only to family (Palmae, Leguminosae). However, the reliability of the megafossil record cannot be established nor comparisons made between the mega- and microflora until the megafossil flora has been revised; at present only 10% of the megafossil genera was found as microfossils. Table 4. Genera of megafi ossils from the Collazo Shal es, Puerto Rico CHolIick, 192! 8). Asterisk (*) indicates ^cnus also represented by pollen. Algae Copaiva Musophyllum Chondrites Cynomctra Myrcia Ferns and Allies Dipholis Myrsine *Hemitelia Echites Oreodaphne Isoetes Echitoniutn Palmacites '^ Eugenia Palmocarpon Gymnosperms Ficus Palmophyllum Zamia *Guarca Pithecellobium Guettarda Plumiera Angiosperms Hancornia Protorhipis Acrodiclidhim Hufclandia Psidium Aniba Icacorca Pamulus Annona Inga * Rhizophora Apocynophyllum Iriartea Sapindus Aspidosperma Juglans Sapota Bactris Lonchocarpus Sideroxylon Cassia "^Malvocarpon Sophora '^ Chrysophyllum Manicaria Stylogyne Coccoloha Melicocca Trichilia Comhretutn Misanteca Zizypkus [Vol. 56 348 ANNALS OF THE MISSOURI BOTANICAL GARDEN Oligocene Climates and Paleophysiography. — In the preceding section it was noted that about 75% of the identified OHgocene flora presently grows in the West Indies. Of the 13 genera no longer present, 10 grow in habitats similar to those available on Puerto Rico. Consequently the Oligocene climates and general environmental conditions must have been comparable to those of the present. These conditions include an insular environment, although considerably greater in extent than the present landmass of Puerto Rico; a central highland with greater relief than at present; trade winds from the northeast during the winter months, shifting to the east and southeast by July; a rainfall near 180 inches on the windward slopes with a rainshadow efFect reducing annual precipitation locally to near the present 10 inches to the lee; and a mean annual temperature of about 78^ F on the south coast to several degrees less than the present 70^^ F in the mountains. The most significant difference between the Oligocene communities and those of the present was the temperate element in the fossil flora. If Ohgocene climates in the Antilles were comparable to those of today, accounting for the distinct similarities between past and present vegetation, physiographic relief must have been greater. The higher altitudes provided the temperate environments required by Fagus, Liqiiidambar, and Nyssa, without altering conditions necessary for the growth of more tropical species. An estimate of physiographic relief in the Greater Antilles during the OHgo- cene may be made from two kinds of evidence. One is on the basis of the vegeta- tion on present highlands compared to those required by the temperate fossil communities. The highest present elevation on Puerto Rico is Cerro de Punta, 1,338 m, where the vegetation is an elfin or mossy forest (Howard, 1968). In Cuba the highest elevation is Pico Turquino, about 2,000 m. The vegetation there is similar to that on Cerro de Punta. The highest altitude in the Greater Antilles is Pico Duarte, 3,175 m, in the Dominican Republic. The vegetation of Pico Duarte (Holdridge, 1945; G. I. Gastony, personal communication, 1969) is a rocky, windswept, open community of Pinus occidentalis with festooning lichens reminiscent of the elfin forest at lower elevations. Other plants at the summit in- clude Pellaea ternifolia, Plagiogyra semicordata, Lobelia ekmanii, Scrophidaria den- sifijlia, Hypericum pycnophyllum, Gnaphalium eggersii, Dendropemon pycnophyi- lus (parasitic on pines) and Lyonia heptamera. The vegetation on other Antillean highlands has been described by Asprey and Robbins (1953; Jamaica), Howard (1968; Puerto Rico), Seifri/, (1943; Cuba) and Urban (1909-1910). Several widespread temperate genera are mentioned : Apium, Aster, Bidens, Clematis, Clethra, Drosera, Eupatoriiim, Hypericum, Ilex, Myrica, Oxalis, Potentilla, Ranun- culus, Ruhiis, Smilax, Yaccinium, Viburnum, and others. Absent from this assem- blage of herbs, shrubs, and small trees are the larger arboreal forms found in the fossil deposits. If it is assumed that altitudes greater than at present provided habitats for the growth of a cool-temperate element in an insular environment at 18^ N latitude, the mountains of the Greater Antilles during the Lower and Middle Tertiary ex- ceeded 3,175 m (10,300 feet). Erosion and diff'erential subsidence reduced these 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 349 heights with concomitant ehmination of the cool-temperate habitats. The low per- centages (0.5-4,5% ) of the arboreal temperate element in the fossil deposits are consistent with the hypothesis that this element grew in a highland habitat re- moved from the basin of deposition. In evaluating the possibility of Oligocene highlands that were of significantly greater altitude than those of today, it must be noted that Fagus, Liquidamhar, and Nyssa all presently grow in east-coastal Mexico, most commonly at elevations of about 1,800 m (6,000 feet). This is a continental rather than insular en- vironment, however, which is not as affected by the Guiana-Caribbean-Florida Gulf Stream current. Further, although these floristic data may only suggest the possibility of Oligocene highlands greater than 3,175 m, this estimate of paleo- physiographic conditions in the Greater Antilles is supported by recently available geologic data. The following information has been provided by R. P. Briggs, U.S. Geologic Survey, San Juan. There is no good evidence for the existence, prior to the middle or late part of the Late Cretaceous, of any sizable islands or landmasses in this part of the crust of the world. From that time probably through the middle Eocene some reasonably good-sized islands may have been present, but nothing approaching the size of Puerto Rico today, rather perhaps more of the order of some of the Virgin Islands. After the middle Eocene there was great crustal uplift, forming a landmass considerably greater than exists today, something perhaps approaching 100 miles north to south and surely connecting with the Virgin Islands on the east and prob- ably with Hispaniola on the west. It is quite possible that elevations on this land- mass approached 15,000 feet. This estimate is based as follows. An oil test well on the north coast of Puerto Rico revealed the middle Tertiary San Sebastian Formation resting on an eroded volcanic basement at a depth presently about 5,500 feet below sea level. The present highest point on Puerto Rico is 4,390 feet. This gives a present elevation of Cerro de Punta, above the eroded basement, of a little less than 10,000 feet. Our evidence though shows that there probably was differential subsidence, with the coastal area subsiding somewhat more than the present interior area. But, considering later erosion and the fact that the unconformity probably goes to somewhat greater depth northward beneath the Atlantic Ocean, I would not be surprised if there were 10,000 feet of relief at the time of deposition of the Middle Tertiary rocks in the test well. Perhaps there was a considerably greater relief at the maximum time of uplift, probably in the late Eocene. You estimate the maximum altitude of Puerto Rico during the middle Oligocene to have been between 6,500 and 7,000 feet above the sites sampled. If we average the present elevations of your sites to 1,000 feet and the average highlands of Puerto Rico to about 4,000 feet, it would appear you are thinking in terms of 3,500 to 4,000 feet of post-middle Oligocene erosion. Going back to my oil test well versus present relief estimate, this certainly would seem to make your estimate and my estimate similar. That is 1,000 feet plus 6,500 feet plus 5,500 feet equals a minimum altitude of 13,000 feet. Of course, much of what I said above is grossly oversimplified, and there is evidence for differential subsidence. Despite this, I think it is safe to say that the relief of ancestral Puerto Rico from Late Eocene to Middle Oligocene time was ap- preciably greater than exists today. It is also fair to say it was being degraded at a very fast rate, A comparison of Table 1, Localities A and B with Fig. 4 reveals other details of the vegetation, physiography, and some facies control of the microfossils. At Locahty A (second column, Table 1) all identified genera are represented at 2.5% or less except Rhizophora which constitutes up to 96% of the microflora. This site marks the approximate position of the shoreline during the Middle [Vol. 56 350 ANNALS OF THE MISSOURI BOTANICAL GARDEN Oligocenc bordered by a mangrove swamp of Rhizophora mangle and Pelliciera rhizophorae (the highest percentages of the latter were also found at Locahty A). Samples 19, 25, and 40 from Locahty B (Fig. 4) arc coalified lignites, and Rhizophora pollen makes up 61-77% of the samples. Samples 15 and 33 are clays (high mineral content), Rhizophora pollen constitutes 1% of sample 15 and was absent from sample 33, The alternate layering of organic-rich versus mineral strata indicates the shoreline was vacillating, probably in response to gradual short-term pyrogenic (rather than eustatic) changes. During the times of subsidence mangrove vegetation dominated at Locahty B and the developing soils incorporated a large amount of organic material. With slight and gradual short-term emergence the site was drained, Rhizophora was ehminatcd at its inland boundary, and more upland vegetation became established on soils of greater mineral content. One of the plants commonly approaching sites vacated by the mangroves w^as Engelhardia. At each locahty it increased considerably more than other species as Rhizophora decreased (e.g., from a maximum of 3% where Rhizophora is abundant to 54% where it is scarce). The reciprocal relationship in percentages of Rhizophora-Pelliciera and Engelhardia pollen reflects the struc- tural instability of the Oligocenc shorehne and records the response of these communities to the orogenic changes characterizing the Caribbean Basin during Middle Tertiary times. Pathiiays of Migration, — Equatorial climatic belts are known to have ex- tended considerably beyond their present hmits during the Early and Middle Ter- tiary. The Eocene Goshen flora of w^estern Oregon, near latitude 44 degrees north contains a significant tropical element (Chancy & Sanborn, 1933), and even the revised Tertiary floras of Alaska are yielding some species of tropical affinities (Wolfe, personal communication, 1969). Geological evidence from the Antilles reveals the presence of larger and more continuous landmasscs (lessening the physical, oceanic barrier to migration), and paleobotanical data reveal climatic conditions favorable to the interchange of northern and southern elements. That many southern species actually did extend into the southeastern United States is evident both from paleobotanical studies of Mississippi Embayment floras and floristic affinities of the modern communities (Graham, 1965/0- The question is whether the Antillean Arc or the Isthmian-coastal Mexico region was the prin- cipal migration route between the North and South American biotas during Early Tertiary times. Data from Puerto Rico make possible some preliminary specula- tions, although results from our studies in Panama and Veracruz, Mexico, may require revision of these. If ancestral Puerto Rico was a landmass some 100 miles wide and included at least the Virgin Islands, Puerto Rico, and Hispaniola, the sea barrier between the West Indies and the southeastern United States may seem significantly re- duced. However, the Tertiary seas extended northw^ard across at least part of peninsula Florida, and Oligocenc strata are exposed in southern South Carohna, Georgia, Alabama, Mississippi, and as far as 120 miles inland along the Texas coast (Fig. 89). In eastern Mexico and the Yucatan Peninsula, the closest point between Mexico and the Antilles (Cuba), the inland extent of Ohgocene seas was 1969] GRAHAM & JARZEN OLIGOCENE OF PUERTO RICO 351 3000 - 2000 - 1000 - Figure 89.— Paleophysiography and Caribbean migration routes during the Middle Oligocene. considerably less (Salas, 1968). The maximum dimensions of ancestral Puerto Rico, its east-west connections (viz., with Cuba, Yucatan, and the Lesser Antilles), and the history of other islands in the Caribbean are not well known, but available evidence suggests the physical barrier between the southeastern United States and the Greater Antilles during the Middle Oligocene was probably more extensive than the Greater Antilles and Mexico greater than at present and possibly less. A second approach to evaluating the Antilles as a Tertiary landbridge is by an West M (commonly eastern Mexican) with QFagus, Liquidamhar, Nyssa') also grow in Latin America (e.g., eastern Mexico). The same would hold if Rollick's (1928) identification of Juglans were included. Thus the 44 identified genera of the San Sebastian Ohgocene flora all have afEini- northern Latin America, and none have exclusive affinities with the ties with northern southeastern United States. I On the basis of these geological and botanical considerations a prehminary model may be made of Tertiary migrations along the Caribbean Basin (Fig. 89). The time of maximum extent of tropical chmates (Early Tertiary) corresponds with the period of extensive uplift and maximum availabihty of land surface in 352 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 the Antilles. In the absence of both physical and climatic barriers, two migration routes were available for extension of tropical and subtropical species into north- ern regions — the Isthmian-coastal Mexico route and the Antillean Arc. Those species migrating along the former were able to extend into southern United States (cf. genera hstcd by Ball, 1931, 1939; and Berry, 1916 from Eocene deposits). After Middle Oligocene times these tropical elements either became extinct as a result of climatic trends toward cool-temperate conditions, evolved new ecotypes capable of growing under temperate environments (e.g., Dirca, Thymelaeaceae; Asimina, Annonaceae; Diospyros, Ebenaceae) or survived in subtropical outhers such as the southward expanding Florida peninsula (e.g., Rhizophora, Laguncu- laria, Tillandda, Avicennia, Annuna, Diphulis; Graham, 1965^). The interchange of species along the Antillean Arc between South America and east coastal Mexico (Yucatan) was facilitated by greater landmasses and physiographic diversity which provided an array of habitats. Further northward migration was hindered, however, by an oceanic barrier at least as extensive as at present. Regarding the Bahama Islands which presently form the nearest com- plete connection between the Greater Antilles and southeastern United States, Butterlin (1956: 173) states that from the Lower Cretaceous "until the Quater- nary, shallow calcareous sedimentation consisting of reef or inter-reef formations were established in the marine subsidence basin. The area remained out of the scope of the orogenic forces and these series are not folded. During the Quaternary and particularly the Pleistocene, relative movements of the marine level and sub- merged shelf have enabled the latter to emerge." Reconstruction of the geologic history of the Florida peninsula is complicated by Phocene and younger strata overlying the Lower and Middle Tertiary deposits. Much of the pertinent data must come from studies of subsurface geology based on available well cores. According to the correlation charts of Cooke, Gardner, and Woodring (1943) the Ocala, Byram, and Suwannee Limestones are marine units deposited during Upper Eocene through Middle Oligocene times. Further, the Suwannee Limestone is encountered in subsurface cores as far north as Fort Pierce, and isopach maps (Toulmin, 195 5) show that it thickens to the south. These data mean that at least the lower one-third of the Florida peninsula was submerged during much of Eocene and Oligocene times. In the vicinity of Ocala, Florida, and in the northern Florida-Georgia region exposed Eocene strata are surrounded by younger rocks. Geologically this could mean these areas were emergent during Middle and Late Tertiary times and no younger overlying rocks were deposited. According to this interpretation these areas would be islands, and their presence would reduce the oceanic barrier be- tween the Antilles and the southeastern United States. Another possibility is that the Eocene units were not emergent during post-Eocene times but have been exposed through erosion of the overlying beds. The Eocene rocks in question form the cap of the Ocala Uplift. The association of the beds with a geomorphic feature ) (personal turc and not due to continuous post-Eocene emergence. In summary, the gelogical 1969] GRAHAM & JARZEN — OLIGOCENE OF PUERTO RICO 353 evidence indicates that much of peninsula Florida was submerged and no inter- vening islands were present during the time the floristic composition of the San Sebastian flora was developing. In contrast, the Lesser Antilles, forming the present connection between the Greater Antilles and northern South America, were probably significant land- masses during much of the Tertiary. According to Butterlin (1956) "the Dutch islands must have remained above w^ater (from Cretaceous) until the Upper Eocene. During that last period the sea flooded the region depositing hmestones and marls. . . . The islands recmerged again and the sea did not flood the area until the Miocene.'' As a consequence of the difference in physiographic history between the north- ern and southern connections of ancestral Puerto Rico, the Oligocene communities include numerous elements common to the present plant associations of eastern Mexico, Central and northern South America, and arc floristically distinct from the vegetation of the southeastern United States. The herbaceous and shrubby temperate elements in the modern Antillean flora, including many species with broad ecological tolerances, constitute cither holdovers from older Tertiary times (the distinctly cool-temperate arboreal forms being eliminated through erosion of the highland habitats) or modern introductions through natural, long-distant dis- persal and anthropogenic factors. Summary In the preceding section a model has been presented for early Tertiary migra- tion patterns around the Caribbean Basin (Fig. 89). The model is consistent with paleobotanical and geological data as presently understood, but it can also be evaluated on the basis of natural Qi.e., non-anthropogenic) distribution patterns of modern organisms. We accept as a generalization that animal, particularly land vertebrate, distribution is influenced to a considerable extent by the range of the plants serving as a source of food and protection. If this is true, and if the model is essentially correct, certain plants and animals should have at least vestiges of a distribution pattern consistent with Fig. 89. The organisms exhibiting this pattern should be 1) those whose origins date back at least to Early or Middle Tertiary times, and 2) those with conservative distribution potentials and without excep- tionally broad ecological tolerances Ce.g., non-weedy species). Changes in the range of such organisms can be expected to commonly reflect changes in the physical and/or climatic barriers to migration. Based on the model, plant and animal species of the southeastern United States having affinities with Latin America should show^ strongest affinities with eastern Mexico and/or Central America and/or northern South America, rather than with the flora and fauna of the Antilles even though the latter is geograph- ically closer. Although the literature of biogeography can be surveyed for examples pertinent to evaluating the model, critical tests can only be made by specialists in various groups who can differentiate between biologically meaningful patterns and those created by poor taxonomy. One range reflecting the expected pattern is that of the plant eenus Cuphea (S. Graham, personal communication, 1969). The spe- [Vol. 56 354 ANNALS OF THE MISSOURI BOTANICAL GARDEN cies occurring in the southeastern United States are generally more closely related to those of Mexico, Central and even northern South America than to those pres- ently growing in the Antilles. Similarly several genera of trees in the eastern deciduous forest of the United States occur disjunct in eastern Mexico and regions to the south and are absent in the Antilles. Martin and Harrell (1957) cite similar patterns among the vertebrates. A final consideration relates to the history of the isthmian region during lower Tertiary times. Studies on the vegetational history of Panama are just beginning, but according to Woodring (personal communication, 1969) study of the geology and fossil faunas reveals that the area was submerged during at least the major part of the Tertiary (from Paleocenc to Early Pleistocene). 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Ser., Prelim. Map 8 5. WOOD ANATOMY OF GOODENIACEAE AND THE PROBLEM OF INSULAR WOODINESS^ Shehwin Carlquist- Abstract Goodcniaccac, a basically herbaceous family with a clear center of origin and diversifica- tion in Australia, has become woody to a limited extent in several genera and genuinely arborescent in some species of Scaevolu. The most woody of these arc montane species of Pacific islands. The probable evolutionary patterns of these and correlations between wood anatomy and ecological conditions are reviewed. Quantitative and qualitative data arc pre- sented in tabular form for 78 collections of 43 taxa of the family. Xermorphic Goodcniaccac tend to have short, narrow vessel elements with helical sculpturing, whereas the reverse is true in mesic species. Absence of axial parench)ma and abundance of crystals also charac- terizes xeric species. The predominance of erect ray cells or raylessness, as well as occurrence in a few species of scalariform or elliptical pits on lateral walls of vessels are probably in- dicators of juvenilism (paedomorphosis). Goodcniaccac do not have libriform fibers, but rather tracheids or fiber-tracheids, with a tendency toward the latter in arborescent species. Multiscriate rays are tall. Correlations between wood anatomy and habit are summarized under the headings: Australian short-lived perennials and short-lived shrubs; montane Pacific species of Scaevola; and maritime species of Scacvola. IS an Introduction Wood anatomy of Goodcniaccac has been little studied. Tliis no doubt derives from the fact that no species in the family pro\ides any economically used wood. Although a few species of Scaevola become small trees, most of the family can be described as herbs. Ho\ve\'er, a predominantly herbaceous family that becomes woody during its evolution provides interesting patterns of wood anatotiiy and ought to be an especially valuable subject for that reason. Goodeniaccae outstanding example of this mode of evolution. The center of origin and divcrsihcation of the family is obviously Australian. If we take into account Bninonia and the recently named Neogoodeuia (Gardner & George, 1963) and Coupcnwokia (Carolin, 1967), ten of the 16 genera are endemic to Australia. Tn four genera, only one species extends beyond Australia, and that species occurs in Australia also: Selliera radicans (southern and south- eastern Australia, Tasmania, New Zealand, Chile); Velleia spathulata (eastern Australia, southeastern New Guinea); Calugyne pilosa (northern Australia, Mo- luccas, southern New Guinea, southern China); and Ixscheriaultia filiformis (northern Australia, Moluccas, New Guinea). The remainder of these genera arc exclusively Australian species. Goodeuia kouingsJ?crgii occurs only in Indo- nesia and southeastern Asia, but this is a single species of a large genus. That none of these genera (except Goodeuia^ has preponderantly primitive features for NTcn^^oo'onf '^^n^f.™" supported by grants from the National Science Foundation, NSFG 23396, GB 4977X, and GB 14092. Preliminary comments on this topic were pre- sented at the 10th Pacific Science Congress. _ 2Claremont Graduate School, Rancho Santa Ana Botanic Garden, Clarcmont Califor- nia 91711, Ann. Missouri Rot. Card. 56: 358 390. 1969. 1969] CARLQUIST — GOOUENIACEAE 359 the family suggests strongly that the origin and diversificaion of Goodeniaccae has taken place within Austraha. The only other genus with non-Austrahan species is Scacvola. This genus, too, has been regarded as specialized within the family (Krause, 1912; Carolin, 1959, 1966). It is the only genus of the family which becomes markedly shrubby or arborescent. The most arborescent species are S. gau- dichandiana and S. glahra of the Hawaiian Islands; these can form trees five or six meters tall. On the basis of circumstantial evidence alone, the family has an herbaceous origin and has become woodier on islands. Floral morphology and geographical distribution seem to support this view. Of the genera other than Scaevola, few show any appreciable woodincss. This seems to be related to adaptation to the dry temperate Australian environment. Many Australian Goodeniaceae are annuals or short-lived perennials. The latter are often branched from the base, so that little wood accumulates, and a wood sample, if one can be collected at all, has to be taken from the short rootstock. Wood in this structure is probably not really comparable to wood of a true shrub, and the grain is often so contorted that sections are difficult to prepare and of hmited value. Growth forms such as these are related to Mediterranean climates. In the present study, Dampiera hrounii and D. glahrescens represent caudex- bearing perennials, branched from near the base. Short-lived shrubs, usually less than a meter high, are formed by Goodenia scayigera, G. ovata, Coopernookia polygalacea, C. strophiulata, and Verreauxia reinuardtii. Of these, only Goodenia ovata grows in a mesic situation, the cucalypt forests in which tree ferns may also be found. The other shrubby species listed grow in relatively dry sandy areas of Western Australia. Within the genus Scaevola some of the Australian species correspond to the above growth forms, but some are more genuinely woody. Scaevola sericophyUa and S. fasciculata are short-lived shrubs of Western Australian scrub heath, whereas S. crassifolia, S. nitida, and S. porocarya are larger shrubs (to one meter or more), probably of relatively short duration (10 years or less). These three species occupy sites near the coast: S. crassifolia is typically a dune plant. Scaevola spinescens is a shrub, usually less than a meter in height, that grows near salt lakes in Western Autralia and other states of Australia. Scaevola tomentosa, al- though a close relative of S. spinescens, is a prostrate shrub of coastal limestones in Western Australia. Scaevola holosericea and S. paludosa are Australian species, little more than perennial herbs, in which only a few older stems have sufFicient wood for study. None of the Australian scacvolas could be called clearly mesic in habitat except S. oppositifolia, a scandent species of disturbed places in wet forests of Queens- land. In localities north of Australia, including islands as far north as the Philip- pines, S. oppositifolia has been given a multiplicity of segregate specific names. These have been justifiably designated as synonyms by Lecnhouts (1957a). With respect to taxonomy, Krause (1912) has placed all of the AustraHan species studied here in section Xerocarpaea except S. spinescens, S. tomentosa, S. oppositifolia, and the wide-ranging S. taccada. Scaevola spinescens and S. tomentosa 360 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 rf — 1-4 Wood sections of Coopernookia. — -1-2. Coopernookia polygalacca (Carl quist 3475). — 1. Transection. Note narrowness of vessels. — 2. Tangential section. Rays are numerous, narrow. — 3-4. Coopernookia strophiolata (Carlquist 339^). — 3. Transection. Note growth rings.— 4. Tangential section. Rays are few, wide; light portions arc the wider vessels from early wood. — Scale to the left of Fig. 1 is a stage micrometer photographed at the same scale as Fie. 1-41. Leneth shown is 1.H mm* rlivi^inn*; 10;/ n^rh 1969] CARLQUIST — GOODENIACEAE 361 are members of sect. Crossotoma; S, oppositifoUa belongs to sect, Euantiophyllum, and S. taccada belongs to sect. Scaevoh. Species of section Scaevola (formerly sect. Sarcocarpaea'), sometimes termed the "fleshy-fruited scaevolas/* bear purple or white fruits with a fleshy exocarp attractive to birds and a woody endocarp in which seeds are embedded. Because of the excellence of this dispersal mechanism for long-distance dispersal (Carl- quist, 1967), the fleshy-fruited scaevolas have spread around the world. This is undoubtedly a relatively recent, although pre-human, pattern. The widespread pan-Pacific beach Scaevola, S. taccada, occurs in Australia and might have orig- inated there. Scaevola phunieri does not occur in Austraha, but its distribution might be said to represent a radiation from the Australian region. The species of section Scaevola are otherwise non-Australian. However, perhaps the closely- related fleshy-fruited sections, Crossotoma and Enantiophyllum represent remnants of this stock on the Australian continent. Scaevola calendulacea, a beach species of eastern Australia, has fleshy white fruits, tinged purple, that float in seawater and remind one of fruits of S, taccada. It supposedly belongs to section Xerocarpaea, but if so (and that disposition needs to be reinvestigated), it shows rapid evolution of a beach species with fleshy fruits both attractive to birds and dispersiblc by flotation in seawater. Either interpretation of S. caleudnlacea as a derivative of dry-fruited Scaevola species or hypothesizing it as a relict of sect. Scaevola stresses the same result — origin of the fleshy-fruited stock termed section Scaevola in the Australian region. Two species of section Scaevola are exclusively maritime and distributed by oceanic drift of the fruits: S. taccada koeni2ii, etc.) and S. plumieri frutescens lia'). Guppy (1917) stratcd that S. taccada fruits can float for a year or more in seawater, while fruits of S. phunieri can witlistand only four or five months of flotation. Scaevola taccada ranges across the Pacific, reaching afl tropical Pacific coasts (except the west coasts of Mexico, Central and South America) and coasts of the Indian Ocean, including Madagascar, the Comoro Islands, the Seychelles, the Andamans, and India (Krause, 1912). Scaevola phunieri has a more restricted distribution: Flor- ida and the Gulf Coast, the West Indies and south to Brazil; India, Ceylon, east (Stern & Bri/ieky, 1958; Bri/icky, 1966) icific Coast of North America (Clarion Isla fornia) evola plumieri may have a distribution more limited than that of S. taccada because of its lesser abihty to float in seawater. However, the fleshy purple fruits of S. 2'?m- mieri would lend themselves to transport by birds, especially shore birds (Carl- ) (St John, 1962) pos Fruits of S. taccada are distributed by seawater, but also have been observed ( ) (G 362 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 Figures 5-8. — Wood sections of Goodcnia — 5-6. Goodcnia ovata {Carlqiiht 831), 5. Transection. Portion of pith below, showing one of the primary bundles in which sec- ondary xylem docs not form; wood is rayless at first. — 6. Tangential section from periphery of larger stem. Tall multiseriate rays are visible. — 7-8. Goodcnia decurrcns (Carlquist 1308). 7. Transection.^ — 8. Tangential section. Wide multiseriate rays and many uniseriate rays are evident. — ■ Scale of magnification shown beside Fig. 1. 1969] CARLQUIST GOODENIACEAE 363 1906) and Hawaii (St. John, 1952). However, these plants are within the mari- time influence and probably are not isolated genetically from beach populations, nor can they become isolated due to the excellent dispersal mechanism of this species. When a littoral insular species cannot become isolated for long periods of time, jwpulations of minor distinction can arise (^.g., S. taccada var. tuamotensis St. John, 1960), but speciation beyond such minor variants is impossible (Carl- quist, 1966/0- O^ the basis of these considerations alone, one would not expect S. taccada to evolve into montane species. In fact, comparative gross morphology of the fleshy-fruited scaevolas shows that the montane species of Pacific islands form a phyletic series separate from S. taccada (or S. plnniierO. The reverse hy- pothesis, that the various purple-fruited montane species have each originated independently from S. taccada is considerably the more difficult hyix)thesis and not borne out by the evidence. However, this interpretation was advanced by Brown (1935), who claimed the Marqucsan S. siihcapitata to be a derivative of S. taccada, and evidently also by Lauterbach (1908), who regarded the Samoan S. nuhigena as closest to S. taccada. Brown's conclusion is amusing, for though he claims S. siihcapitata as close to S. taccada, he regards a second species, S. marque- sensis, as close to the Hawaiian S. procera (a Kauai population of S. gaiidichau- diana'). However, S. siihcapitata and S. marquesensis seem barely separable, and probably should be united. Scae^'ola nuhigena is worlliy of specific recognition; it resembles S. florihiinda of Fiji most closely, and Lauterbach does acknowledge the similarity in leaf shape between these species. The montane fleshy-fruited scaevolas form an interesting trans-Pacific series. They probably represent a sort of island-hopping, but perhaps not in a perfect sequence (^.g., Samoan derived from Fijian, Fijian from New Caledonia, etc.) from west to east. These insular species, together with S. plumieri and S. taccada, a cohesive group. The montane insular species probably originated from a form (1966) species certainly not clear at present, and Carolin's phylogeny of fruit types is open to ques- tion. For example, he envisions fruits of S. mollis (*Typc 3") and S. chamissoniana ("Type 2") as independent derivatives from the type represented by an Australian species, S. porocarya, and not known outside of S. porocarya. However, S. mollis and S. chamissoniana are very closely related Hawaiian species which very likely derive from a single Hawaiian ancestor, an ancestor tliat probably resembled the Fijian S. florihiinda but certainly not S. porocarya. The species of sect. Crossotoma, with their one-flowered cymes, do resemble the species of sect. Scaevola in a num- ber of respects, certainly. A recently-named species of New Guinea, S. pauciflora (Leenhouts, 19571?), shows similiarity to both sections. The New Caledonian species of sect. Scaevola seem to represent at least t^vo introductions. Scaevola hechii, a lowland species of dry serpentine areas, is quite unlike the upland species in habit, inflorescence type, and various other technical features. The upland scaevolas of New Calendonia are, however, closely related to species of islands to the east. Scaei'ola montana in particular is similar to S. flori- hiinda, S. marquesensis, etc. Hybrids between S. montana and S. indigofera may [Vol. 56 364 ANNALS OF THE MISSOURI BOTANICAL GARDEN Figures 9 12. — Wood sections of Leschcnauhia. 9-10. Ix'schenaiiltia splendens (Carlquist 3437). — 9. Transection, pith below. — 10. Tangential section. Note raylessness of wood. — 11-12. Leschcnauhia hiloba {Carlquist 911). — 11. Transection. Lower portion of this photograph is rayless w^ood. — 12. Tangential section, taken from periphery of older stem. Rays have developed at this stage. — Scale of magnification shown beside Fig. 1. 1969] CARLQUIST — GOODENIACEAE 365 be found in various places on New Caledonia. Scaei'ola montana typically is a lower-elevation montane species, but it comes into contact with S. indigofera in some places. Hybrids in New Caledonia scaevolas have very likely been named as species, and study of various taxa in the field in New Caledonia is very much needed. Scaevola cylindrica might be such a hybrid. The species of Scaevola from Fiji to the Marquesas form a single phyletic series. Of these, S. floribunda of Fiji and S. niihigena of Samoa are most closely related to each other. Scaevola floribunda can become a tree to 6 meters, but S. niihigena and S. marquesensis are shrubs. A distinctive new species from Tahiti, S. tahitensis, is described in an accompanying paper (Carlquist, 1969); this species represents the first montane Scaex'ola to be reported between Savaii, Samoa, and the Marquesas Islands. Most of the Hawaiian species of Scaevola are shrubs, but S. glabra, S. gaudi- chaudiana, and S. "procera" can be small trees, to about 5 meters. Scaevola glabra probably represents an early introduction of sect. Scaevola to the Hawaiian Islands. It exemplifies loss of dispersibility (Carlquist, 1966a) and adaptation to orni- thophily (Carlquist, 1970a). However, there are no basic features in S. glabra alien to the fleshy-fruited species of Scaevola — curved flowers borne singly in an axillary fashion can be found in S. indigofera and Australian scaevolas; yellow flower color occurs in S. gaiidichaiidii (Gillett, 1969) and S. tomentosa. If S. gla- bra were recognized as a separate genus, Camyhiisia de Vriese, the genus Scaevola could with justification be broken into a number of genera. Scaevola kauaiensis is regarded here as a synonym of S. glabra. The Hawaiian species other than S, glabra and S. taccada appear to represent a single phyletic series closely related to the series beginning with S. montana de- scribed above, although one or two further exceptions might be entertained. At any rate, S. gaiidichaiidiana and its close allies (S. chamissoniana, S. mollis) rep- resent adaptations to wet forest. Scaevola kilaiieae and S. gaudichaudii favor drier habitats. These two species tend not to be true upright shrubs, but are typically branched from near ground level. Scaevola coriacea is a sprawling species adapted to dry locahties near the coast. It might be a derivative of the S. gaudichaudiana complex, but the possibility of a relationship to S. plumieri, which it resembles in a number of ways, ought to be investigated. Both Scaevola gaudichaudiana and S. chaniissoniana can range from relatively wet to relatively dry forest; S. gaudi- chaudiana and S. mollis, a wet-forest species characteristically, hybridize. Such folia by Skottsberg (1927) (1888) of hybridization between S. gaudichaudiana and S. mollis has been analyzed excel- (1966) ri ability (called ) with S. gaudichaudiana. Scaevola gaudichaudii, described well in habitat and morphology recently by Gillett (1969), has one-flowered cymes like the species of section Crossotoma. The nature of its relationships, whether to the S. gaudi- [Vol. 56 366 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig ■s 13 16. — 13 14. Dampicra hrownii (Carlquist /376).— 13. Transection, pith below. Note occlusion of vessels by unidentified substances.— 14. Tangential section. Rays are wide, distortions in course of axial elements arc evident — 15 16. Scaevola spincscens (Carl- quht 914). — 15. Transection. Note narrowness of vessels, which are, however, numerous. 16. Tangential section. Rays are very wide and rather short. — Scale of magnification shown beside Fig. 1. 1969] CARLQUIST — GOODENIACEAE 367 chaiidiana complex or to species of Crossotoma, needs to be investigated. For illus- trations and descriptions of ecological adaptations of the various Hawaiian species of Scaevola, the reader is referred to Carlquist (1970a). Evolution of Scaevola on Pacific islands has featured both arborescence and entry into wet forest. The Hawaiian species represent not only these tendencies, but adaptation to drier areas as well. The succulent leaves of S. coriacea and S. kilaueae are external indicators of xeromorphy. On the other hand, the leathery, glabrous leaves of S, glabra represent a definite adaptation in leaf characteristics to rain forest — -areas occupied by S. glabra receive more than 100 inches of rain per year. Succulence characterizes Scaei'ola sphiescens, S. plumieri, and S. taccada leaves and can be called a response to halophytic ecology. The wide variety of growth forms and ecological adaptations in the genus Scaevola make this genus, together with the other genera of Goodeniaceae, an in- teresting group for study of wood anatomy in relation to ecology. Are changes in ecological adaptation followed by (or accompanied by) changes in wood anatomy, and if so, what are these changes in wood anatomy? Can longer as well as shorter vessel elements evolve in such a group? Is tlie theory of paedomorphosis (Carl- quist, 1962) relevant to wood evolution in Goodeniaceae? What is the nature of wood evolution in a primarily herbaceous group, and how does it differ from patterns in a truly woody group of dicotyledons? Questions like these seem best answ^ered by study of groups such as Goodeniaceae. Data on w^ood anatomy of Goodeniaceae is extremely scanty. The description of w^ood for this family by Metcalfe and Chalk (1950) is derived almost exclu- sively from S. taccada, as those authors acknowledge. The data on wood anatomy of Goodeniaceae offered by Krause (1912) are insignificant. Kanehira (1921) described wood anatomy only for S\ taccada. Data on vegetative anatomy other than wood were offered by Colozza (1908), Vesque (1876), Kienholz (1921), and Mullan (1931, 1933). Solercder (1885) summarized w^ood anatomy for the family, but his description is based on three species which are not identified with respect to particular features in his text, so his generalizations are not very useful. The only significant comparative wood study to date is that of Stern and Brizicky (1958), w^ho analyzed wood of S. phimieri and compared it to wood of S. taccada, S. gaiidichaudiana, and S. floribiinda. Materials, Methods, and Acknowledgments Because woods of Goodeniaceae are virtually absent from institutional wood collections, most of the woods studied here had to be collected in the field. Some samples from the U.S. National Museum (USw) and the Arnold Arboretum (Aw) w^ood collections w^re available. Many individuals have assisted my field work, and I wish to extend thanks particularly to Dr. William L. Stern and Dr. George W. Gillett. Others who deserve thanks for aid in my field work or in microtechnical tasks include Mrs. Jane Benjamin Baker, Mr. Norman Carlson, Mr. Alex George, the late Mr. Fred W. Humphreys and Mrs. Humphreys, Dr. Robert Jancey, Mr. Tim O. Magee, Mr. John Parham, Dr. Harold St. John, Prof. A. C. Smith, Dr. Robert F. Thome, Miss Linda Thorne, and Mrs. Ann Piatt Walker. Wood samples w^re dried in the areas where they w^ere collected. Wood samples of Goodeniaceae dry readily without molding, especially if bark is partly removed. Sections and macerations of the various samples w^ere prepared using the usual techniques. The vessels, tracheids, and fiber-tracheids of many Goodeniaceae have what could be called a gelatinous 368 ANNALS OF THE MISSOLHU BOTANICAL GARDEN [Vol. 56 Figures 17-20.— Wood 17-18. Scaevola fasciculata (Carlqnist 897). — 17. Transection. Several growth rings are evident. — 18. Tangen- tial section. Rays are narrow and inconspicuous. — 19-20. Sccwiola nitida (Cuilijiiist 1047). 19. Transection. — 20. Tangential section. Rays are wider and more conspicuous; early- wood vessels are shown. Scale of magnification shown heside Fig. 1. 1969] CARLQUIST — GOODENIACEAE 369 nature. Although this is not pronounced in many species, it often causes difficulties in sectioning. Features considered to show significant differences or similarities among the taxa are summarized in Table 1, and further explanation of symbols is given below. For quantitative data, 50 or more measurements per feature of each collection were taken. The diameter of fiber-tracheids and trachcids was obtained from measurement of these cells at their widest diameter (usually from a maceration) and averaging such measurements for each collection. The number of species and collections studied here is probably sufficient to give reliable results, at least in many characters of wood anatomy. Anatomical Descriptions The quantitative data in the table arc self-explanatory. For qualitative data, symbols have been used. ap = axial parenehyma more abundant at end of growth ring, or present only at end of growth ring. bo ^ axial parenchyma present only in bands whieh repre- sent the margin of a growth ring, da = diffuse, relatively abundant, ds ^ diffuse, relatively scanty. ft = fiber-traeheids present, no tracheids. mv = more numerous vessels at margin of growth ring. t = traeheids present, no fiber-tracheids. tft = both tracheids and fiber-tracheids present, vs = vasicentric scanty. wv^ vessels wider at beginning of growth ring. -|- = feature present. — feature absent. With respect to ray histology, the letters indicate presence of cell types as seen in radial section: U = upright (erect); S - square; P^= procumbent. Upper case indicates abundance of a cell type, lower ease indicates paucity, and absence of the letter indicates no cells of a category could be found. Thus USP indicates abundance of upright, square, and procumbent cells; Us indicates upright cells abundant, with a few square cells and no procumbent cells. VESSELS Dimensions, — In Asteraceae, length and diameter of vessels in secondary xylem bears a very close relationship to degree of mesomorphy or xeromorphy (Carlquist, 1966r). Does this apply to Goodeniaceae as well? If we examine the data in Table 1 and arbitrarily designate average vesschclement length of more than 400/x as long and less than 200//, as short, we find a marked correlation with ecology. The rain forest species of Scaevola ha\'e long vessel elements almost with- out exception. Such long vessels are shown in Fig. 24, 26, 28, 34, and 36. Of the Goodeniaceae in which vessel-element length averages between 200/i and 300/t w^e find that all arc xerophytes: (Fig. 2) (Fig. 4), Darnpiera glabrescens, Leschenanltia hiloha (Fig. 12), L. ft y)lendens (Fig. 10), Scaevola coriacea (Fig. 30), S. dielsii^ S. holos leae, S. pahidosa, S. spinescens (Fig. 16), and S,tonientosa, To be 370 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 o •X3 03 O QJ o 03 QJ OJ :3 QJ :3 qj a IT) Of o 03 c o s o OJ C O O M 4 ^ /-N n » < ^-^ H ^ (/) • ^H ;-( a> C/5 W ^sodAX IPD luaojiaqn 3uiqsin3ui;siQ s[pisXj3 ^t'lUAq.iuojHj I'M'^V ^i^aojojsiij Xcy S[[03 'oSnjOAy 4fd ^^ rf 'oStuoAV ^spioqoeix JOtlU JO spioqiujx J'^l'TiuniQ ri 'oSejaAy 'sprnq-^uax -10(11 [ JO sp[.^i{.:)FJx qi^ii.i^ T/ 'SJTJ [OSSOA -1>^}'">'-LI'']Q dnojQ joj s|.issoy\^ Ti '[ossoA IsopjAV r/ 'o3t?j.iAy o 03 H + ir^ ^ ^ 0> in ro (N 03 u QJ > QJ o '^ in m in ro O O 03 C/^ -H ^ »n IN. a a a a CO 03 03 "i > > > >" ^ ^ & & o a OJ^ o o o o o o o o o c in vO ^ in in ^ (N ^ in 1- ^ CTi -t SO in ^ m \D rr, cn (N q ':^ ^0 in in in in in O in in ro ro m ro Ti- ro in ro in ro O 00 ^ ro 00 ro a> ^ ro GO i K ^ s ^ =: ^ ^ ;^ "^ ^ ■ ^C o Q O pc; 00 o ro ■ E ro ^' 00 CQ 00 ^ > > + + s^ ^ ro Tf ^ O rJ ro ro 1> in in CO ro ro ro ro OJ \0 00 cji CO in ^ 'v". /'^ ^'"N < < c/^ cr p; CC V-/ * v^/ f-K c 00 lil- V a; 00 w = « = 4 t^^ ^^ ^ Q (^, O o X < IN ro 00 u QJ c o o -0 03 T3 .0 > > n3 3 3 a D a a a a PLH CD 3 P P m 00 4 in 4 q in in vO J ^K ro ^ '^ in ^ r- vX) ro CM ■ oJ (^ * in in 4 GO O; (N 00 03 fH CO 6 oi ^H V • « * -^ 4 ro IN ro 00 0^ CO in a^ in I:^ (N 370 0^ ro \0 00 ro CO nJ rsi ^.0 0^ in in ■0 CO ro in tN 0^ 0^ CO in '-0 p: X in ro 4 i- o3 03 o p: 2 ^ ■^. .2 c/^ ^ ^ g ►-J nJ 5^ V^i u c/^ 1969] CARLQUIST — GOODENIACEAE 371 o o o o o Ch CO CQ > o o o o o o o o o o o o o o o o o o ++ + o o o o o o o o ++ o o o o o yo <>r» :i^ T3 ^3 ^ ^3 ^ o o rJ2 (/) ui u^ t/; VI ■^ ^ ^ Tj "T^ 03 cd cd cd ct3 ■^ T3 ^ '^ '^ Ph Cm DP Ph P PP Ph Ph PP P p PP Ph Ph Ph CLh fL, ^ (^D v: c/^ c/i P P PP P ^ Ph PP CI, &, ;:^ c«.^ tr t/) c/) c/) L/J D D p P p « ro r-- 00 tN o O vC o (N CM CO in ^ ro o 00 t^ fH O 00 O in tJ- in ro in r>J \0 ^ ro r^ ^ in m I— 1 in (N !>■ GO iri '* ^ A A O rf- vTi O (Ti ro O ^ A * A B rj ^ c- ro ro ro ro r^J n3 On! in oa o O^ OJ O I-- t-- -H ro O O rt --^ CO --^ 00 I> ^ CO X 00 •i- -t ^ u^ 00 O O^ nJ ^ ^ -H 00 O O ^ O t^ in ^ ro in ^ rf ro \0 in in i> in ^ in in ro ^ t^ r^ (Ti ^ o^ rj- O ro m r- r- ra o ro ni O O ro ro in oi ri O Tf in in rf CO ^ B p « # * « ^ * « * » * « m # -^ » ^ m * m * * * O ^ 135 103 160 CO CO in 00 125 135 o 103 o o 111 in o 103 CO 00 143 107 CO 103 r- 00 Oi in o ■^ ^ in ^ ^ 00 ro ro in o in •^ ^ 00 m in ro o -f GO 00 J>- 00 in ro in vo 538 406 381 429 561 433 246 840 585 667 585 690 527 620 504 456 772 632 918 616 889 374 00 ^ 388 233 260 302 377 296 422 507 426 411 393 500 448 466 303 315 403 480 533 359 454 268 Q s ■^ St "3 < < p= p: in CO nj ni (^ ^ /'^ P= O O c-r ^ ^ V4 "13 f^ 5 QJ < c^ p:; Zl^lD t: »n 1^ s t^- 2 O "- >:= *r: P^ P= . Ln r:h ^ ^ ^ to t^ N CO P^ CO CM o '^ ^ l±\ '^ CP < CO p:: < C3 — O ? K ^ o^ E a: CO 00 in 00 in to CO ro in S ^ ^ ^ I > ^3 rO rO ro in I > J U O O P P o c<3 O ^ - 00 CQ i^j IN or c^ &.T3 ^ P^ 03 M CO sP p: C. o r- J p; P^ In "n IN. CO ■^ OJ !5^ to v; U ~ •= •= ■= ct) r^ V. U. k. V^ 2 ^ ^ ^ « ^ u u u u ^ ^ > i>j M fvi r^t r . "^ « ^ u < IN Vi ^ 5 CO t/:3 uo v:> ^ t/5 C/5 C/^ ty^ C/D ^ 372 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 o o c CO u o a; C/5 o 03 03 U C Vi cd 3 .6 o 03 > c^ s a nato s o rt ^4h ^ ^ Q OJ o 3 ^ .s C o * U ta * )j ^^ cQ » < V^ H ^ c/) • rH ^ s vi [aiqx rf 'oSejOAy SpioqoHix t/ *o3rjoAy *spioq.">BJX jDqij JO spir>qoBJX qj3uo^ ri 's;ij [.7SS3A jojouibiq dnoj^ joj s[nss.^^ t/ *pss3A J^-^PIA\ rl *o3rj,iAy I^SS3A 1S33U01 ri *33bioav O X H ^_l c^H !-f- o o o o o o O (N u^ r- t^ un o o ro r- i> r-- CO o r- ro (N ro 00 \C t^ t^ v^ in r4 iri f^ Ln T^ ^ o o > o o o > > > j^ o ^ ^ o o o o + o o o o o o o o o r^ (N (M O (N CM O no Tt vO vO O IT) \sC o ^ ra CTi O^ O vO m CO (M rj m in ^ O^ CO O ro 00 r- ^ ro CO t:J- m v£> vO (ji ^ O O in Tj- o^ i> r^ vO r^ o> (N in ro 00 ^ in ro O in 00 in nJ O in \0 m in 00 1-H O in in Tj- CJ\ ^ ^ 00 in 00 Tj- O ro 1- 'I- -1- Tj- ^ 'i- c^ i> ^ (N rj in o o^ ^ rr) \0 m ^ Tt 3 /~N /'^ ^~^ < < < ^ t/:) c/D p:; p: c:; *^^ ^^ >^^v o CD ro oo O r^ (N- 00 X < < ■^ ro ro O IV 00 <^ <^ ^ 3 s s (5^ Q^ t^l "3 ::: :: v; '^ !:u 'S^ ^ C3 ^3 r> V*. *«- r^ < on 1:1? v_/ o in OJ (N &£ OJ 1^ Q s ^ ^ awefl Car/ ■< c/: c/^ •-G (=^ ct; •^ o t^ N. ^ VO in 1^4 4** +*A r/i v: cxi v:i *'** i-^ **^ * •^ • 2 s >s 3^ ^ ^ t:^ 01 CA) Crt C/!) LO C/3 C/^ p:; cc tf p; tf K v_/\_^ ^y v_^ V_y ^_/ o Tj- in ^ t^ a\ ^ ^ r^ r^ ^ rf t^ r^ r^ IN r-- IN > (M CO in in ro m < P= O 4- + W t/l C/l -Q "-a ^ C/3 tft ^3 XJ > > T3 T3 03 ffi o3 oj rt o 00 00 1- O < en N> X 00 p? V, »^ *^ >^ -2 Q Q t3 a u u ^ s Q VI VI v: c/^ t-ri t/*, -^_ v; v^ ' *^ *f^ *v^ * *^ *^^ CU 'v^ 2 a s s 2 a -Nt V; ^ c/:j ^•5 c/: s .P ^ Si. o 1969] CARLQUIST — GOODKMACEAE 373 CO o o o o n 5 n 5 000000000 o > B CO o o + + o o + o o + + 000000000 o + o o o o o o c/j v: ui K_ tn rt t/) ^ T3 "^ C/1 u^ nzi > P-- W Wl ■^ y^ > > > V5 (« TD O O Oh a a DP P Oh 3 PPPPPPPPP a P Ph V) 3 CI. 0- PP 06 o 00 m ID 00 \DOoO'-Hr-iOra'-;fN| ir^Ot^t^i^^DrO^O in IT) m o 00 in 00 r^ 00 o o 00 o (N '^cXJO^LnOmiN-rC so CO O ro in in -rf \D m ^ ^ \£J inooT3--^^Tfin^^ in O in -O rJ 00 o ro ro CM O rj ro ro in ro rsi cjt^^'1"COinO^^'^o^ f^J^oror^irororo^orO o CM ro (N O ^ ro fM in CO in O000^ln^0^'-^0 oor^-^^roinro^oji— f ^-1-^i^^in^Oin ^ r- in ro in O ^ in in '^ Tt u-i ■^ in in ■^ ro i^t^OvsDsor-r-mi>- \D ^ \0 in O ro 00 O (N "^ O l> u^^^]0000^•-;ra ro in rJ ro 00 ^ ro -rj- 00 in ro 00 00 rO00i-HTi-rO'i-rja> ■^ra-^'-H'^a^^ioiro o^ o so ro in r^ in in ro O O ro O CO 00 ■^ r- ^ a^soroo^so■^so^^o^ CXJCJiOCOO^roOOOO ro ro ■'i- ro O in 00 vO o in in ro 00 in \0 00 Tf in co 00 00 ro oorJO^r^(Ni>coo sC^£}r\io^CMsOvOroin Tj-inOT^ininvOu^i^ ro in ro I> sD rJ ro in in OJ ^ ro n PC . ^ ^^ 1^ <3^ ^ ^ ^ ^ ^ nj O r^l h-._ Q f^ ^ Q "3 Q UOU U U U P o ro ro in p p: p CT 03 in 03 o K ^ < in t^ C5- v: t-T) C/5 X PQ > Q < 00 CO P^ 00 *^ s s 374 [Vol. 56 ANNALS OV THE MISSOURI BOTANICAL GARDEN Figures 21-24.— Wood sections of New Caledonian species of Sawvohi. 21-22. Scac- volahccldi (Carlquist 705J. ^2 1. Transection. Note narron vessels and thick-walled tracheids. 22. Tangential section. Tiavs are tall. 23 24. Scaciola nio)itinia (Carlquist 714). 23. Transection. Note wide vessels.^24. Tangential section. Rays lar^e-eelled and conspicu- ous. Scale of magnification shown heside Fig. 1. 1969] CARLQLIST — GOODENIACEAE 375 finds some species in which vessel elements do not, at first, seem to agree witJi tliis distinction between xeromorphy and mesomorpliy. Goodeiiia deciirrens has relatively long vessel elements, but this is a short-lived perennial of moderately wet forest in which growdi takes place during spring months w^hen water is avail- able. This might also be true of Scaevola platyphyUa, S. sericophylla, and possibly S. beckii (Fig. 22). Relatively long vessel elements occur in most collections of S. taccada (Fig. 40) and S\ ylumieri (Fig. 38) as well as in S.porocarya. Scaevola taccada seems paradoxical in this respect, for we would expect a halophyte to be a xeromorph at first glance. However, S\ taccada might qualify as a succulent, and succulent species appear to exemplify paedomorphosis or perhaps actual mesomorphy. With respect to vessel diameter, trends of xeromorphy and mesomorphy are even clearer than they are with vessel-element length. Thus, narrow vessel diameter characterizes the species of dry habitats: Coopernookia polygalacea (Fig. 1^,C. stropliiolata (Fig. 3), Leschenaultia /?/7o/?a(Fig. 11^, L,florihuuda, L. splen- dens (Fig. 9), Scaevola beckii (Fig. 21), S. fasciculata (Fig. 17), S. nitida (Fig. 19), S. paliidosa, S. platyphylla, S. porocarya, S. sericophyUa, S. spiiiescens (Fig. 15), and S, tomeutosa. The montane insular species of Scaevola have relatively wade vessels (Fig. 23, 25, 27, 33, 35). The only Australian Goodeniaceae to have relati\'ely wide vessels are from wet forest: Goodenia ovata (Fig. 5) and Scaevola oppositifolia (Fig. 41). Wide vessels in S. op posit i folia probably relate not only to this, but to the scandent habit of that species. This factor very likely applies also to S. coriacea (Fig. 29), which could be described as a prostrate non-twining vine. On the basis of vessel diameter, S. plumieri (Fig. 37) and S. taccada (Fig. 39) ought to qualify as mcsic. Perhaps these succulent or semi-succulent seaside shrubs, like comparably succulent species of Euphorbia (Carlquist, 1970/?), can be said to have a mcsic internal environment in their stems. Abundance. — Metcalfe and Chalk (1950) claim absence of vessels from (1958) (1912) (1876) ( ) In general, Goodeniaceae show vessel abundance in inverse proportion to vessel diameter. Because I selected the latter measure, data on vessel abundance w^as not presented here. Perforation Plates. — Goodeniaceae have basically simple perforation plates. Scalariform perforation plates, mixed with simple ones, were reported for Scaevola plumieri, S. taccada, and S. gaudichaiidiana by Stern and Bri/icky (1958), Such a condition also occurs in Goodenia ovata, Scaevola mollis, S.montana, and S. "pro- cera" More species could undoubtedly be added to this fist by more intensive ob- servation. Tlie occurrence of such plates has been interpreted as a vestige of a primitive condition by Stern and Brizicky (1958). However, in the species where scalariform perforation plates occur, most plates are simple and the scalariform plates (which generallv have 10-15 bars) arc definitely in the minority. Many of 376 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 56 Figures 25-28. — Wood sections uf Fijian and Tahitian species of Scacvola, — 25-26. Scacvola florihunda (Carlquist 700). — 25. Transection. Note wide vessels. — 26. Tangential section. Rays arc wide and tall. 27-28. Scaevohi taliitoisis (Carlquist 654). — 27. Transec- tion. Note thick-w^allcd fiher-tracheids. — 28. Tangential section. Ray cells are mostly up- right. Scale of magnification shown l)cside Fig. 1. 1969] CAULQUIST — GOODENIACEAE 377 the scalariform plates, where present, have aberrant structure with anastomosing and variously-oriented bars. If scalariform plates were present as a vestige, one w^ould expect them to be in the majority, and if simple plates were also present, all grades of transition from many to few bars ought to occur. This does not appear to characterize any Goodeniaceae. Occurrence of scalariform perforation plates does not appear to be an indicator of primitiveness in Cichorieae (Carlquist, 1960(7). In that group these plates are not common and tend to occur only in mesomorphic species. This appears to be true in Goodeniaceae also. Lateral-Wall Pitting. — The presence of scalariform lateral-wall pitting in a group with the characteristics of Goodeniaceae seems more an indicator of juven- ihsm than of primitiveness. ScaevoJa ghhra (Fig. 45) exemplifies this pitting condition more clearly than any other Goodeniaceae and was cited in this regard (Carlquist, 1962) c ) in areas of intervascular pitting. Such a scheme of lateral-wall pitting was also observed in S. floribuuda, S. gaiidichaudiana, S. kilaneae, S, mollis, S. montana, and S. tahitensis. Stern and Bri/icky (1958) listed "alternate and transitional" pitting for intervascular areas of vessels in S. gandichaudiana. This list of species suggests that wet-forest species are most likely to have scalariform or transitional lateral-w^all pitting. This also proved to be the case in another genus of dicotyledons that has adapted to WTt forest in the Hawaiian Islands, Euphorbia (Carlquist, 19701;). Scalariform or transitional intervascular pitting also occurs in S. taccada and, to a hmited extent, S. pluniieri, as noted by Stern and Bri/icky (1958). If the longer, wider vessels in these two species are indications of mesomorphy or suc- culence or pacdomorphosis (concepts closely interrelated), then this pitting type in these species is understandable. This appeared to be true in insular species of Euphorbia (Carlquist, 1970/;). Vegetative portions of Scaevola taccada and S. plumicri are certainly succulent, as shown by Kienholz (1921) and Mullan (1931, 1933). Xeromorphy appears to be basic to other vessel-wall modifications. The species of Goodeniaceae not listed above may be assumed to have alternate pitting exclu- sively or nearly so. However, a few^ of these show, in addition, occurrence of grooves or bands. Vessel pits in which the pit apertures arc elongate, forming short grooves to either side of a pit, w^ere observed in Da}}ipiera glahrescens, Scaevola nitida, and S, oppositifolia. More marked grooves, which tend to interconnect sev- eral apertures in a helix on the \essel w^-^ll, were observed in S. beckii, S, coriacea, and S. spinescens (Fig. 43). Scaevola spijicsceus has the most conspicuous grooves in this series. A species closely allied to S. spinescens, S. tomentosa, lacks grooves, however. Scaevola tomentosa is a less xeromorphic species than S. spinescens. The two species of Coopernookia exhibit a more marked form of xeromorphy — presence of fine bands, the so-called "tertiary helical thickenings" of some authors. This condition is illustrated for C. polygalacea (Fig. 42). Coopernookia wood qual- ifies as the most xeromorphic of Goodeniaceae on the basis of vessel dimensions. [Vol. 56 378 VNNALS OF THE MISSOURI BOTAMCAL GARDEN Figures 29 32.— W 29 30. Scaevola coriacca (Carlquist 2125). ^29. Transection. Note the rather wide vessels; these arc prohal)ly related to the sprawling habit of the plant. ^30. Tangential section. Note small cell size. — 31-32. Scaevola gaiaUchaudii {Carhiuht 2380). — 31. Transection. Growth rings are evident. — 32. Tangential section. — Scale of magnification shown beside Fig. 1. 1969] CARLQUIST — GOODENIACEAE 379 Grooves and bands have been suj^gested as indieators of xeromorphy in Asleraccac (Carlquist, 1966c') and other families (Webber, 1936). Diameter of pits in Goodeniaeeae ranges from 3 to 7/x (to 9/-i in S. glabra because elhptieal pits arc extremely common). This ranee is not great, and no correlations are proposed. Howe\er, pits in vessels of Australian Goodeniaeeae seem generally smaller than those in the insular species of Scaevola. Groti])ing. — In studies of woods of Astereae (Carlquist, 1960Z?) and other tribes of Asteraeeae, degree of vessel grouping proved to be an interesting indicator of xeromorphy or phyletie speciali/ation, or perhaps both. Goodeniaeeae as a whole have a much lower degree of vessel grouping than do Astereae as a whole, and tliis probably reflects lower degree of specialization in w^oods of Goodeniaeeae. The high figure for the family is in Scaevola heckii, the distinctive serpentine species of New Caledonia. Other species are low^ in degree of vessel grouping, and minor differences probably are not significant. TRACHEIDS AND riBER-TRACIIEIDS Borders are clearly present on pits of all imperforate tracheary elements in Goodeniaeeae, and no libriform fibers can be said to be present. Designating the imperforate elements as tracheids or liber-tracheids is a rather arbitrary matter. For convenience, I have used a criterion employed elsewhere (Carlquist, 1961). If the ellipse of the pit aperture extends beyond the outline of the pit cavity, the element is designated as a fiber-tracheid ; if the aperture is shorter than the diameter of the pit cavity, the element is said to be a tracheid. The distinction in the case of Goodeniaeeae is not a ver)' great one, for most elements have pits in which the aperture is only slightly longer than, or slightly shorter than, the diameter of the pit cavity. Consequently, stress should not be placed upon this feature, summarized in the last column of Table I. However, there seems to he a correlation between increasing woodiness (or arborescence) and the tendency toward production of fiber-tracheids rather than tracheids. The Hawaiian species of Scaevola in particular might be cited in this regard. Fiber-tracheids and tracheids in Goodeniaeeae range in wall thickness, but exceptionally thick walls eharacteri/e Scaevola glabra (Fig. 35, 36), S. tahitensis (Fig. 27, 28), S. beckii (Fig. 21, 22), and S. spinesceus (Fig. 15, 16). In S. spinesceus, fiber-tracheids and tracheids are relatixeh narrow, so that with thick w^alls, the lumina of the cells are virtually occluded, a fact noted by Vesque (1876). Tracheids and fiber-tracheids of S. gaudichaiidiaua (Fig. 33), S.pliimieri (Fig. 37), S. taccada (Fig. 39), and S. oppositifolia (Fig. 41), look thin-walled, but these cells are relatively wide, so tliat the wall thickness leaves room for a lumen much wider than that of S. sp'mescens. Gelatinous \\alls characten/e tracheids, fiber-tracheids, and vessel elements of many Goodeniaeeae, judging from staining reactions and shrinkage patterns Avithin walls. These were seen most strikingly in Scaevola glabra, S. platyphylla, and S. spinesceus. [Vol. 56 380 ANNALS OF THE MISSOURI BOTANICAL GARDEN ^35 Figures 3 3-36. — Wood sections of upland Hawaiian sprcics of Scacvola, — 3 3-- 34. Scac- vola gandichaUiVuwa (USw-J 5333). — 33. Transection. Vessels arc rather wide and isolated. — 34. Tangential section. Multiseriatc rays are rather short and narrow. — 3 5-36. Scacvola glabra (Carlquist 1975). — 3 5. Transection. Note deposits in \esscls; axial parenchyma is abundant. — 36. Tangential section. Rays arc very wide and hi^h; vessel elements are rather long. — Scale of magnification sho\\n beside Fig. 1. 1969] CARLQUIST — GOODENIACEAE 381 VASCULAR RAYS Dimensions. — Species with exceptionally tall rays (averaging more than 3 mm) for Goodeniaccae include Scaevola chamissoniana var. cylindrocarpa, S. glabra (Fig. 36), S. oppositifoUa, S. palndosa, S. platyphylla, S. porocarya, and S. taccada (Fig. 40). However, rays of Goodeniaccae generally are unusually tall. For ex- ample, the average ray height in Asteraceae at large is only 1.44 mm. However, tall rays characterize dicotyledons of various types, and ray height shows no clear correlation with habit. The correlation between ray height and vessel-element length is not perfect in Goodeniaccae, although it is more often true than not. Scaevola pahidosa has tall rays, but short vessel elements. Short rays occur in the following Goodeniaccae: Coopernookia strophiolata (Fig. 4), Goodenia decurrens (Fig. 8), Leschenaiiltia biloba (Fig. 12), Scaevola crassifolia, S, diehii, S/'pro- cera^' S. scricophylla, and Verreauxia reinwardtii. These species all have relatively short vessel elements, although again the correlation of short rays w'ith short vessel elements is imperfect. Cell Shape, — With respect to ray histology, Goodeniaccae show some interest- ing tendencies. Predominance of erect ray cells, a criterion cited earlier (Carlquist, 1962) as an indication of paedomorphosis, characterizes many Goodeniaccae: Danipiera brmoiii, Goodenia decurrens, G, ovata, G. scapigera, Leschenaultia bi- loba, Scaevola coriacea, S. floribunda, S.glabra, S, indigofera, S. montana, S. nitida, S. platyphylla, S. phiniieri, S. sericophylla, S. taccada, and S. tahitensis. There is a correlation between erectncss of ray cells and longer vessel elements. Both may be related to mcsomorphy, as seems the case in Asteraceae (Carlquist, 1966c). Predominance of procumbent cells characterizes only a few Goodeniaccae: Cooper- nookia polygalacea, C. strophiolata, Scaevola holosericea, S, kilaueae, S. porocarya, S. spinescens, and S. tomentosa. All but one of these is from a rather dry habitat in Australia; S. kilaueae is a species of an area which is effectively dry (new^ lava of Kilauea) in the Hawaiian Islands. Rayless woods are, in effect, those in which ray cells are exclusively erect, and thus equivalent to libriform fibers (or axial parenchyma). Raylessness charac- terizes all of the species of Leschenaultia, At an early stage, Leschenaultia woods are definitely rayless, as shown for L, splendens (Fig. 9—10). Later in ontogeny, rays developed, as shown for L. biloha. The same is true of Goodenia ovata (Fig. 5—6), This relation between ontogeny and raylessness was noted by Barghoorn (1941). Rayless woods tend to occur in herbaceous groups which show juvenilism (Carl- quist, 1962). In particular, herbaceous groups which show^ a limited increase in w^oodiness tend to include rayless species. This appears to be true in Goodeniaceae. Goodenia ovata suggests this clearly, for the secondary xylem originates not from the vascular bundles of the primary cylinder, but outside them (Fig. 5) as mentioned and illustrated by both Vesque (1876) and Krause (1912). This condition can be said to represent an anomalous secondary xylem condition and possibly a return to woodiness in a phvlad that has lost the abilitv to form secondarv xvlem in bun- dies of the primary cylinder. If so, the relationship between herbaceous groups and ravlessncss would certainly be clear. 382 ANNALS OF THE MISSOUEI BOTANICAL GARDEN [Vol. 56 Figures 37-40.— Wood sections of littoral species of Scacvola. — 37-38. Scaevola phimicri fVV. G. EarJe s.n.). — 37. Transection, Note wide vessels. — 38. TanRential section. Vessel elements are relatively long. — 39-40. Scaevola taccada (Carlquist 701), — 39, Tran- section. Vessels arc relatively sparse and isolated. — 40. Tangential section. Rays are wide and tall. Scale of magnification shown beside Fig. 1. 1969] CARLQUIST — GOODENIACEAE 383 Ray Cell Walls. — Relatively thin-walled ray cells characterize most Goode- niaceae. These cells are lignified in nearly all species. A few cells of larger rays in Goodcnia decurrens (Fig. 8) are not hgnified and tend to collapse. Notably thick ray cell walls were observed in Scaerola heckii (Fig. 22), S. diehii, S, glabra (Fig. 36), S, plumieri (Fig. 38), and S, spiuescens (Fig. 16). Conspicuous pits char- acterize ray cells of Cooperiiookia polygalaceae (Fig. 46) and Dampiera hrownii. Notably large ray cells occur in Scaevola glabra (Fig. 36), S. plumieri (Fig. 38), and S. taccada (Fig. 40). Uniseriate Rays, — Uniseriate rays are present in all Goodeniaccae studied (except the rayless stems of Lescbenaultia~). Uniseriate rays are usually composed of erect ray cells. On superficial examination of wood, a uniseriate ray can be confused with axial parenchyma strands. The latter occur as shorter series, roughly the length of vessel elements, that form smooth fusiform outhnes as seen in a tangential section. The cells of a uniseriate rav are more rounded and undulate in outline, and ordinarily they are taller tlian a strand of axial parenchyma. AXIAL PARENCHYMA The range in axial parenchyma in Goodeniaccae is rather marked, as can be seen in Table 1. Axial parenchyma is absent or nearly so in Leschenaultia. Axial parenchyma is absent except for a very few^ cells at the end of a growth ring in Coopernookia polygalacea, C. strophiolata, Daiiipiera browuii, Goodenia decurrens, Scaevola diehii , S. f S. sericophylla. Interestingly, these arc all relatively shortdivcd perennials from Australia. Con- sistent formation of parenchyma seems characteristic of longer-lived species in Goodeniaccae. Very sparse axial parenchyma arranged in a diffuse fashion characterizes most species of Goodeniaccae. Where a sparse axial parenchyma in a diffuse distribution occurs, one cannot really distinguish between apotracheal and paratrachcal (vasi- centric) satisfactorily, because a few cells will be adjacent to vessels although many will not be. In a few Goodeniaccae, however, only \ asicentric axial paren- chyma cells wxre seen: Scaerola beckii, S, plumieri, and S. taccada. This w^oukl seem an interesting species character, for it emphasizes the difference between these three species and the purple-fruited montane insular species of Scaevola, which have either abundant or sparse diffuse parenchyma. Relatively abundant diffuse parenchyma characterizes a few Goodeniaccae: Goodenia ovata, Scaevola coriacea, S. crassifolia, S. florihunda, S. glabra, S. nion- tana, S, oppositifolia, S. spiuescens, and one collection of S. taccada. These are all woody rain forest plants or succulent plants. Correlation with mcsomorphy is sug- gested by the fact that abundance of axial parenchyma is greatest in the species adapted to the wettest forest: S. glabra of the Hawaiian Islands. GROWTH RINGS I Growth rings are present in many Goodeniaccae, particularly those of tem- perate Australia, but few are strongly pronounced. Probably none could be called truly ring-porous. Growth rings have been illustrated here for Coopernookia poly- 384 ANNALS or THE MISSOURI BOTANICAL GARDEN [Vol. 56 1 II! Figures 41-45.— 41. Scaevola oppositifolia (Carlqiiist 7386). Transection. The wide vessels correspond to the scandent habit of this species. — Scale of magnification shown above. 42-45. Portions of vessel walls to show types of lateral-wall pitting.— 42. CoopcDiookia poly- galacca (Carlqiiist 3475). Fine bands are evident.— 4.^. Scaevola spirwsccns (Carlqiiist 914). Grooves interconnect pits adjacent in a helix.— 44. Scaevola chamissoniana var. chamis- soniana (Carlqiiist 2241). Scalariform pitting at left, alternate pitting at right in this vessel.— 4 5. Scaevola glabra (Carlqiiist 1975). While some vessels show a near-perfect scalariform pattern, this vessel shows a transitional or near-scalariform pattern. — Scale of magnification for Fig. 42-45 is shown to right of Fig. 45. Divisions 10^ each. 1969] CARLQUIST — ^GOODENIACEAE 385 galacea (Fig. 1, top), C. strophiulata (Fig. 3), Goodenia decurreus (Fig. 7, mid- dle), Leschenaultia hiloha (Fig. 11, top), Scnevola aimiescens (Fig. 15), S. fasci- culata (Fig. 17), S. tahitensis (Fig. 27), and S. gandichaudii (Fig. 31). Rain forest species such as Scaevola glabra (Fig. 35) show very subtle or virtually no growth ring phenomena, whereas the most marked growtli rings are seen in the shrubby perennials of southwestern Australia. INSOLUBLE DEPOSITS Deposits insoluble in water, alcohol, and xylene are present in woods of many Goodeniaceae. These appear resin-like, but even that vague term might be mis- leading. Chemical identihcation of these prominent deposits is very much needed. Massi\ __„^...a cells, were observed in the following species: Damjiicra hroivnii (Fig ), Goodenia scapigera, Leschenaultia splendens (a few vessels near pith, Fig. 9) Scaevola heckii (Fig. 21-22) ( 29-30), S.crassifolia, S. gandichaudiana(¥\o. 33-34), S. gaudichaiidiKVig. 31), S. glabra (Fig. 35-36), S. holosericea, S. indigojcra, S. kilaueae, S. oppositifolia (Fig. 41), S. spinescens (Fig. 15-16), and S". tahitensis (Fig. 27-28). Scantier accumulations of these materials, present only as droplets in parenchyma and other cells, were observed in woods of Coopernookia puly galacea (Fig. 1-2), Goodenia decurrens (Fig. 7-8), Scaevola fasciculata (Fig. 17-18), S. mollis, S.montana (Fig. 23), S. "procera" and S. taccada (Fig. 39-40). Goodeniaceae in which few or no deposits were observed in woods include Coopernookia strophiolata (Fig. 3- 4), Dampiera glahrescens, Goodenia ovata (Fig. 5-6), Leschenaultia hiloha (Fig. 11-12), Scaevola dielsii, S. florihunda (Fig. 25-26), S. uitida (Fig. 19-20), S. pahidosa, S. platyphylla, S. plumieri (Fig. 37-38), S. porocarya, S. sericophylla, S. tomentosa, and Verreauxia reinwardtii. CRYSTALS Crystals, presumably composed of calcium oxalate, occur in ray cells of many Goodeniaceae, as shown in Table 1 and Fig. 46-49. These crystals show birefrin- gence in polarized light, often exhibiting irridescent colorations (Fig. 49). Crystals of Goodeniaceae could all be described as rhombic. A few are 4-sided rhombic as seen in face view; in the majority, additional faces arc added, so that a hexagonal appearance is achieved (Fig. 49). In a few, the ends develop flattened faces so that an octagonal shape occurs. Large hexagonal crystals are abundant in rays of the following species: Cooper- nookia strophiolata (Fig. 47), Scaevola crassifolia, S. diehii, S. holoscricea, S. nitida, S. oppositifolia, S. porocarya (Fig. 48), S. sericophylla, S. spinescens (Fig. 49), and S. tomentosa. Somewhat smaller crystals, more elongate in shape, were observed in Coopernookia polygalacca (Fig. 46), Scaevola coriacea, S. gandichaudii, S. kilaueae, S. pahidosa, and S. platyphylla. Scaevola heckii has distinctive crystals: octagonal in face view, with a rather rounded outline, and pronounced fractures and gaps in the central portions. 386 [Vol. 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN FiGUKLS 46-49. — Portions of rays from radial sections of wood of Goodcniaccac show- ing crystals. — 46. Coopeniookia polygalacca (Carlquist 34 7SJ. Crystals are small and some- what obscured by prominent pitting in ray cells. — 47. Coopcriinokia strophiolata (Carlquist 3393). Crystals are medium to large and mostly hexagonal in face view. ^ 48. Scaevola porocarya [Carlquist 2957), Crystals are medium-si/ed and hexagonal Crectangular as seen in side view). — 49. Scaevola spinesceus (Carlquist 914). Crystals iihotograpbed with polarized light; these would Ije relatively large in Goodeniaeeae as a whole. Scale of magnification for Fig. 46 49 is shown above Fig. 46. Divisions lO/i each. 1969] CARLQUIST — GOODENIACEAE 387 There appears to be a very elear correlation between xeromorphy and presence of crystals. None of the upland montane scaevolas of Pacific islands have crystals. The only exception is S. oppositifoUa, wliicli does not belong to sect. Scaevola in any case. In addition to rhombic crystals in rays, druses occur in pith and cortex of Goodeniaceae (Colozza, 1908; Krause, 1912; Metcalfe & Chalk, 1950). In the present study, such druses were observed in pith of Scaevola taccada and S. ta- hitensis. They probably occur in more species, but pith and cortex were not investigated. Discussion and Conclusions Systematic application of data from A\ood anatomy in Goodeniaceae is obvious enough. Contrasts and similarities arc self-evident in the table, illustrations, and text of this paper. However, as the reader will have noticed, adapation of Goode- niaceae to diverse habitats has carried with it marked shifts in modes of anatomical structure. These modes may be summarized as follows: Australia)! Short-Lived Perejniials aiul Short-Lived Shrubs. — This group in- cludes Cooperjiookia, Coodenia, Danipiera, some species of Scaevola. It possesses relatively short, narrow vessels, which are numerous per area of transection. Per- foration plates are almost invariably simple; lateral wall pits are relatively small and alternate. Grooves along vessel walls or even bands (Cooperuookia) may be present. Tracheids and fiber-tracheids are present. Rays of some species contain a preponderance of procumbent cells, w^hile others QLescheiiaultia spp.; Goodeiiia ovata^ show preponderance of erect ray cells and even raylessness earlier in the ontogeny of stems. Some wide, short rays are present (Goodenia deciirrens and Scaevola spinescens), but most species in this group have relati\'ely narrow^ rays of medium height. Axial parenchyma is absent or very sparse and is sometimes present only as a few cells in growth rings. In no species is it abundant. Growth rings are prominent, and most species have large rhomboidal crystals in the rays. Goodenia ovata is an exception to the above pattern, but it occupies a mesic locality, so that its relatively long, wide vessels, relatively abundant axial paren- chyma, lack of growtli rings, and lack of crystals — all of these characteristics of the group beknv — are understandable. Scaevola oppositifolia is also exceptional in virtually the same respects C<^'>^ccpt that it has crystals), and again, it occupies mesic sites in Queensland and the Indo-Malaysian islands. Montane Pacific Species of Scaevola.— These species are characterized by long, relatively wide vessel elements, which are rather few per unit area of transection. There is a strong tendency toward transitional or even scalariform lateral-wall pitting on vessels; grooves or bands are absent. Perforation plates are basically simple, but occasional scalariform plates (often with aberrant bars) are present. Both tracheids and fiber-tracheids are present, but there is a tendency toward the latter. Corresponding to greater length of vessel elements, erect ray cells are rel- atively more common than procumbent cells. Rays are mostly relatively tall, of varying width (some very wide). Axial parenchyma is diffuse, ranging from sparse to very abundant. Growth rings are few^ and inconspicuous. Crystals are lacking. [Vol. 56 388 ANNALS OF THE MISSOURI BOTANICAL GARDEN Exceptions to the above pattern are found in lowland insular species which represent less or no adaptation to wet-forest conditions. Thus vessels are narrow in S. beckii and S. gaiidichaiidii, short in S. heckii, S. coriacea, S. gaudichaudii, and S. kilaueae. Procumbent ray cells predominate in S. kilaucae. Crystals are present in rays of S. heddi, S. coriacea, S. gaudichaudii, and S. kilaueae. In these respects, S. beckii, S. coriacea, S. gaudichaudii, and S. kilaucae show greater xeromorphy and more resemblance to the Australian Goodeniaceae. Maritime Scaevulas. — Surprisingly, the maritime scaevolas resemble the wet- forest species much more than they do the Austrahan species. Growth in areas of high salinity is not accompanied by xeromorphy. E\idently the succulent nature of these species has as a consequence a mesomorphic pattern, instead. This appears Scaevola (Carlquist, 1970 /;) parallels between wood anatomy in Euphorbia and that of Scaevola are quite marked, in fact. Increased arborescence and adaptation to mesic, relatively uniform conditions by an herbaceous group are well shown by montane insular species of Scaevola. These features, detailed above, are placed in relief by the contrasting patterns of the lowland insular species of Scaevola and the Australian Goodeniaceae. While each group of dicotyledons will sho\\ somewhat different tendencies in evolution (and continents) ;■ seem to follow this course and have been studied in this regard seem to show phenomena referable to paedomorphosis: Asteraceae, Lobelioideae, Euphorbia. The fact that juvenilism (or neoteny) is a mechanism that permits change in mature wood pat- terns seems clear by now. Woods of Goodeniaceae showing these tendencies have been described as having a primitive aspect (Stern & Bri/lcky, 1958). The ex- planation for this appearance seems to involve paedomorphosis rather than the simple retention of primitive features. Because primary xylcm is a reservoir of relatively primitive features, woods in which primary xylcm patterns are protracted will tend to show a more primitive aspect than expected. How'ever, Goodeniaceae do seem to show some retained primitive features re- gardless of juvenilism. These include occurrence of tracheids and fiber-tracheid; rather than libriform hbers. Libriform fibers are the imperforate tracheary cell ;y) ( The relative abundance of uniscriate rays also seems validly a primitive feature- again, Asteraceae, Solanaeeae, and Campanulaceac have a paucity of uniscriate rays. The low degree of grouping in vessels of Goodeniaceae might also be a primiti\'e feature. LiTERATURi: Cited Bargiioorn, E. S. 1941. The ontogenetic development and phylogenetic specialization of rays in the xylcm of dicotyledons. III. The elimination of rays. Bull. Torrey Bot Club 68: 317-?525. Brizicky, G. K. 1966. The Goodeniaceae in the southeastern United States. Jour Arnold Arbor. 47: 293 300. 1969] CARLQUIST — GOODENIACEAE 389 Brown, F. B. II. 1935. Flora of southeastern Polynesia. III. Dicotyledons. Bernice P. Bishop Mus. Bull. 130: 1-386. Carlquist, S. 1960a. Wood anatomy of Cichorieae (Composite). Trop. Woods 112: 65-91. 196017. Wood anatomy of Astereae (Compositae). Trop. Woods 113: 54-84. 1961. Comparative Plant Anatomy. New York. 1962. A theory of paedomorphosis in dicoyledonous woods. Phytomorphology 12: 30-45. — . 1966a. The biota of long-distance dispersal. III. Loss of dispersibility in the Ha- waiian flora. Brittonia 18: 310-335. — . 1966J;. The biota of long-distance dispersal. IV. Genetic systems in the floras of oceanic islands. Evolution 20: 4 3 3-4 5 5. — . 1966c. Wood anatomy of Compositae: a summary, \.'ith comments on factors controlling wood evaluation. Aliso 6(2): 25-44. — . 1967. The biota of long-distance dispersal. V. Plant dispersal to Pacific Islands. Bull. Torrcy Bot. Club 94: 129-162. — . 1969. A new species of Scaevola (Goodeniaceae) from Tahiti. Ann. Missouri Bot. Card. 56: . , 1970a. Hawaiian Natural History. New York. . 19701?. Wood anatomy of Hawaiian, Macaronesian, and other species of Euphor- bia. Biol. Jour. Linn. Soc. (in press). Carolin, R. C. 19 59. Floral structure and anatomy in the family Goodeniaceae Dumort. Proc. Linn. Soc. New South Wales 84: 242-255. ■ -. 1966. Seeds and fruit of the Goodeniaceae. Proc. Linn. Soc. New South Wales 91: 58-83. , 1967. Coopernoohia: A new genus of Goodeniaceae. Proc. Linn. Soc. New South Wales 92: 209-216. CoLozzA, A. 1908. Studio anatomico sulle Goodeniaceae. Nuovo Giorn. Bot. Ital. 15: 5-92. Gardner, C. & A. George. 1963. Eight new plants from Western Australia. Jour. Roy. Soc. Western Austral. 46: 129-136. GiLLETT, G. W. 1966. Hybridization and its taxonomic implications in the Scaevola gaiidichaiidiaiia complex of the Hawaiian Islands. Evolution 20: 506-516. 1969. The nomenclatural and taxonomic status of the Hawaiian shrub Scaevola gaudichaiidii H. & A. Pacific Sci. 23: 12 5-128. GupPY, H. B. 1906. Observations of a Naturalist in the Pacific between 1896 and 1899. 2. Plant Dispersal. London. Plants, seeds, and currents in the West Indies and Azores. London. 1917. Hillebrand, W. 1888. Flora of the Hawaiian Islands. Heidelberg. Kanehira, R. 1921. Anatomical Characters and Identification of Formosan Woods. Bu- reau of Productive Industries, Taihoku, Taiwan. KiENHOLZ, R. 1926. An ecological anatomical study of beach vegetation in the Philip- pines. Proc. Amer. Philos. Soc. 65 (suppl.): 58 100. Krause, K. 1912. Goodeniaceae. In A. Engler CEditor)/*Das Pflanzcnreich." 54: 1-207. Lauterbach, C. 1908. Beitrage zur Flora der SamoaTnseln. Bot. Jahrb. 41: 215-238. Leenhouts, p. W. 1957a. Goodeniaceae, In C. G. G. J. van Steenis (Editor), "Flora Malesiana." 5: 335-344. 1957fc. Miscellaneous notes on New Guinea plants. IV. Nova Guinea n.s. 8: 175-177. Metcalfe, C. R. & L. Chalk. 1950. Anatomy of the Dicotyledons. 2. London. Mullan, D. p. 1931. Observations on the water-storing devices in the leaves of some In- dian halophytes. Jour, Indian Bot. Soc. 10: 126-133, 193 3. Observations on the biology and physiological anatomy of some Indian halophytes. Part 2. Psammophilous halophytes. Jour. Indian Bot. Soc. 12: 165-182. St. John, H. 1952. Notes on Hawaiian species of Scaevola (Goodeniaceae). Hawaiian Plant Studies 19. Pacific Sci. 6: 30-34. . 1960. The name of the Indo-Pacific strand Scaevola. Taxon 9: 200-208. . 1962, A new Scaevola (Goodeniaceae) from Socotra Island. Webbia 17: 45-58. Skottsberg, C. 1927. Artemisia, Scaevola, Santaliim, and Vaccinium of Hawaii. Bernice P. Bishop Mus. Bull. 43: 1-89. [Vol. 56 390 ANNALS OF THE MISSOURI BOTANICAL GARDEN SoLEREDER, H. 1885. Ubcr den systcmatischen Wert der Ilolzstruktur bei den Dicot\]e- donen. Miinchen. Stern, W. L. & G. K. Brizicky. 1958. The woods and flora of the Florida Keys. Good- eniaceac. Trop. Woods 109: 38-44. Ves^)ue, J. 1876. Note sur Tanatomie dii Goodenia ovata. Ann. Sci. Nat. Bot., Ser. 6. 3: 312-326. Webber, Irma E. 1936. The woods of sclerophyllous and desert shrubs and desert plants of California. Amur. Jour. Bot. 23: 181 188. THE CALIFORNIA ISLANDS^ Robert F. Tiiorne^ Abstract The California Islands off the Pacific coast of southern and Baja California display almost diagrammatically the insular phenomena that characterize fringing archipelagos. To a lesser extent they also illustrate some of the insular features associated with oceanic islands. Several of these islands have been much studied and are relatively accessible. The location, size, elevation, and geological history of the islands arc described. To deter- mine the degree of disharmony of the island floras, they are analyzed in comparison with selected California mainland floras. The degree of richness of the island floras is discussed in relation to size of the islands, proximity to the mainland, and other variables. Such phenomena of island life as the immigrant pattern of vertebrate distribution, relict versus autochthonous endemism of plants and animals, insular gigantism, continuous flower- ing, high degree of hybridization, vulnerabflity, and loss of dispersibflity in island species are considered in reference to the California Islands. Some suggestions as to available facflities on the islands and persons or institutions to contact are given in an epflogue. Off the Pacific coast of southern Cahfornia and Baja CaHfornia, Mexico, lie the islands variously called the Channel Islands or CaHfornia Offshore Islands and the Baja California Islands or, collectively, the California Islands. Although they are in no sense tropical, they are worthy of mention in this symposium be- cause they are a good example of a fringing archipelago and because they illustrate many of the insular phenomena tliat one might otherwise have to travel far to tropical areas to study. Some of them are quite accessible from the heavily pop- ulated areas of southern California, and some of them have been rather thoroughly studied by biologists. Although they are, thus, rather well known to Cahfornia biologists, they are surprisingly little known to non-Californians. It is hoped that this short discussion of them and of some of their insular characteristics will create more interest in them among those biologists concerned with evolutionary and environmental phenomena. The Southern California and Baja California Islands consist of 16 major islands or island groups spread over some 500 miles between Point Conception, Cahfornia, and Punta Eugenia, Baja California, Mexico (Fig. 1). From east to west the Northern Channel Islands are Anacapa, Santa Cruz, Santa Rosa, and San Miguel. The Southern Channel Islands are San Nicolas, Santa Barbara, Santa Catalina, and San Clemente Islands. The Baja Cahfornia Islands include Los Coronados just south of San Diego, Todos Santos off Ensenada, San Martin off San Quintin, San Geronimo south of El Rosario, the San Benitos, Cedros, and Natividad off Punta Eugenia, and Guadalupe 157 miles off the Baja Cahfornia peninsula. In order of size, with area in square miles, the islands are: Cedros (134), Guadalupe (98), Santa Cruz (96), Santa Rosa (84), Santa Catahna (75), San II am grateful to Dr. Reid Moran for critically reviewing the manuscript, for mal