/ I r r H ' VOLUME 63 v L3 1976 '?7(* Cj 2l OF THE wm Published by the Missouri Botanical Garden Press, St. Louis, Missouri 63110 © Missouri Botanical Garden 1977 atimiCAC CONTENTS • Antonovics, Janis. The Nature of Limits to Natural Selection 224 Averett, John E. & Michael Powell. A New Gypsophilous Species of Gaillardia (Asteraceae) from Chihuahua, Mexico 375 Axelrod, Daniel I. Evolution of the Santa Lucia Fir (Abies bracteata) Ecosystem 24 Burch, Derek. A New Species of Chamaesyce ( Euphorbiaceae ) from the Bahamas 378 Carson, Hampton L. The Unit of Genetic Change in Adaptation and Speciation i 210 Correa, M. D. & A. S. Taylor B. Flora of Panama. Family 76A. Dros- eraceae 389 ■ Croat, Thomas B. Notes on Central and South American Cissus (Vita- ceae) 358 Croat, Thomas B. Flora of Panama. Family 105. Staphyleaceae 393 Croat, Thomas B. Flora of Panama. Family 108. Sapindaceae 419 Crosby, Marshall R. Rhynchostegiopsis carolae (Musci, Hookeriaceae ) : A New Species from Costa Rica 373 Curtis, William F. Chromosome Counts in Grielum and Cercis 379 Cruden, Robert William. Intraspecific Variation in Pollen-Ovule Ratios and Nectar Secretion — Preliminary Evidence of Ecotypic Adaptation _ 277 D'Arcy, W. G. New Names and Taxa in Solanaceae 363 D'Arcy, W. G. Chromosome Counts in Solarium 377 D'Arcy, W. G. Flora of Panama. Family 55. Bataceae 385 D'Arcy, W. G. Flora of Panama. Family 157. Symplocaceae 547 D'Arcy, W. G. Flora of Panama. Family 158. Oleaceae 553 Davidse, Gerrit. Evolution at the Population Level: The Twenty-second Systematics Symposium 209 Dietrich, Werner & Peter H. Raven. An Earlier Name for Oenothera stri- gosa ( Onagraceae ) 382 Eiten, Liene Teixeira. Inflorescence Units in the Cyperaceae 81 Eiten, Liene Teixeira. The Morphology of Some Critical Species of Cyp- eraceae 113 Furlow, John F. Nomenclatural Changes in Alnus (Betulaceae) 380 Gentry, Alwyx H. Studies in Bignoniaceae 18. Notes on S. Moore's Mato Grosso Bignoniaceae 42 Gentry, Alwyn H. Studies in Bignoniaceae 19. Generic Mergers and New Species of South American Bignoniaceae 46 Gentry, Alwyn H. Additional Panamanian Pa<:sifloraceae 341 Gentry, Alwyn H. A New Panamanian Sterculia with Taxonomic Notes on the Genus 370 1 200 Goldblatt, Peter. Evolution, Cytology and Subgeneric Classification in Moraea ( Iridaceae ) Goldblatt, Peter. Cytotaxonomic Studies in the Tribe Quillajeae (Rosa- ceae) v Goldblatt, Peter. Chromosome Number in Gomortega keule 207 Goldblatt, Peter. Bamardiella: A New Genus of the Iridaceae and its Relationship to Gynandtiris 309 Goldblatt, Peter. Chromosome Cytology of Hessea, Strumaria, and Car- polyza ( Amaryllidaceae ) 314 Goldblatt, Peter. The Genus Moraea in the Winter Rainfall Region of southern Africa 557 Goldblatt, Peter. New or Noteworthy Chromosome Records in the An- giosperms 889 Goldblatt, Peter & Richard C. Keating. Chromosome Cytology, Pollen Structure, and Relationship of Retzia capends 321 Graham, Alan. Studies in Neotropical Paleobotany. II. The Miocene Communities of Veracruz, Mexico 787 Howard, R. A. Flora of Panama. Family 106. Icacinaceae 399 Hunziker, Juan H. (see Wells, Philip V. & Juan H. Hunziker) 843 Johnson, George B. Enzyme Polymorphism and Adaptation in Alpine But- terflies 248 Keating, Richard C. (see Goldblatt, Peter & Richard C. Keating) 321 King R. M., D. W. Kyhos, A. M. Powell, P. H. Raven & H. Robinson. Chromosome Numbers in Compositae XIII. Eupatorieae 862 Kyhos, D. W. (see King, R. M. et al.) 862 Meyer, Frederick G. Flora of Panama. Family 181. Valerianaceae 581 Powell, A. M. (see King, R. M. et al.) 862 Powell, Michael, (see Averett, John E. & Michael Powell) 375 Prance, Ghillean T. Flora of Panama. Family 120. Caryocaraceae 541 Raven, Peter II. Generic and Sectional Delimitation in Onagraceae, Tribe Epi lobieae ... 32g Raven, Petkh H. Specific Status for CamiesotUa claviformis subsp. wig- ginsit (Onagraceae) ... 383 Raven, Peter H. (see Dietrich, Werner & Peter H. Raven) Raven, Peter H. (see King, R. M. et al.) _ Robinson, H. ( see King, R. M. et al.) 382 862 862 Sarukhan, Jose. On Selective Pressures and ^ergy Allocation in Popula- tions of Ranunculus repens L., R. bulbosus L. and R. acris **> Solbrig, Otto T. On the Relative Advantages of Cross- and Self-Pollina- ^ tion " Stearn, W. T. Union of Chionanthus and Linociera (Oleaceae) 355 Taylor, Peter. Flora of Panama. Family 176. Lentibulariaceae 565 Taylor B., A. S. (see Correa, M. D. & A. S. Taylor B.) 389 Wells Philip V. & Juan H. Hunziker. Origin of the Creosote Bush (Lar- tea) Deserts of Southwestern North America °* Wilbur, Robert L. Flora of Panama. Family 183. Campanulaceae 593 Wunderlin, Richard R. The Panamanian Species of Bauhinia (Legumi- ^ nosae) OF THE mm md VOLUME 63 1976 NUMBER 1 P ^976 CUMATRON. MISSOURI BOTANICAL GARDEN * GARWN LIBRARY CONTENTS Evolution, Cytology and Subgeneric Classification in Moraea (Iridaceae) ^ Peter Goldblatt Evolution of the Santa Lucia Fir {Abies bracteata) Ecosystem Daniel L ^ Axelrod Studies in Bignoniaceae 18: Notes on S. Moore's Mato Grosso Bignoniaceae ^ Alwyn H. Gentry " Studies in Bignoniaceae 19: Generic Mergers and New Species of South ^ American Bignoniaceae Alwyn H. Gentry Inflorescence Units in the Cyperaceae Liene Teixeira Eiten — The Morphology of Some Critical Brazilian Species of Cyperaceae Liene ^ Teixeira Eiten Cytotaxonomic Studies in the Tribe Quillajeae (Rosaceae) Peter Goldblatt 200 Chromosome Number in Gomortega keule Peter Goldblatt — - VOLUME 63 1976 NUMBER 1 OF THE The Annals contains papers, primarily in systematic botany, contributed from the Missouri Botanical Garden. Papers originating outside the Garden will also be accepted. Authors should write the editor for information concerning preparation of manuscripts and page charges. Editorial Committee * Gerrit Davidse, Editor-in-Chief Missouri Botanical Garden W. G. D'Arcy, Editor — Flora of Panama Missouri Botanical Garden John D. Dwyer Missouri Botanical Garden 6- St. Louis University Peter Goldblatt Missouri Botanical Garden Published St. Louis, Missouri 63110. ription information contact the Business Office of the Annals Box 368, 1041 New Hampshire, Lawrence, Kansas 66044. Subscription price is $40 per volume for 4 issues. Application to mail at second class rates is pending at Lawrence, Kansas 66044. © Missouri Botanical Garden 1976 OF THE M m VOLUME 63 1976 NUMBER 1 EVOLUTION, CYTOLOGY AND SUBGENERIC CLASSIFICATION IN MORAEA (IRIDACEAE) 1 Petkr Goldblatt 2 Abstract Moraea, an African genus widespread south of the Sahara, comprising ca. 92 species, is divided into five subgenera: Moraea, Monocephalae, Visciramosa, Grandiflora, and Vieusseuxia, and into several sections. Chromosome numbers of n = 10, 9, 8, 6, 20, and 12 are reported in the 52 species studied to date. A base number of x = 10 is postulated for Moraea, and Dietes is suggested as the closest living ancestor. Moraea is pictured as having evolved in the mid to late Tertiary in central southern Africa in response to the onset of a dry climatic regime border- ing the tropics. The great radiation of the genus in the southwestern Cape is seen as a more recent phenomenon resulting from the development of a Mediterranean climate in this i Several nomenclatural changes are made and four new species are described. area. Moraea omosome Iridaceae (Goldblatt, 1971a). In this work the karyotypes of several species were described and discussed with reference to the evolution of the genus, and al- though only a comparatively small number of species were studied, the cyto- logical evidence was found to be at odds in many instances with existing sub- generic treatments. Thus it was clear that further cytological investigation would provide invaluable information for taxonomic and evolutionary studies and that it would be especially useful at infrageneric levels. In preparation for a revision of the South African representatives of Moraea, which comprise the great majority of the species in the genus, I have undertaken a more extensive karyotypic study. Chromosomal data are now known for 52 species, approximately 70% of the genus, and this information has assisted con- tions with certain rphol itics of Moraea because of the strong correla- traits. The comparison of morphology and 1 This study was supported by a travel grant from the Council for Scientific and Industrial Research of South Africa, and by Grant BMS 74-18905 from the National Science Foundation of the U.S.A. 2 B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, 2315 Tower Grove Avenue, St. Louis. Missouri 63110. U.S.A. Ann. Missouri Bot. Gard. 63: 1-23. 1976. o ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 cytology has resulted in a complete rearrangement of the classification of Baker ( 1896 ) , the most recent and only detailed treatment of the genus, and has made interpret Afr The system presented here fall area (Goldblatt, 1973), but was necessarily incomplete because of the limited occurrence of Moraea in this area. Moreover, no formal ranks between genus and species were recognized in this revision so that the subgeneric treatment is essen- tially new in the present work. The cytological data are presented in the first section of the paper as these are central to the revised generic classification. The classification itself is then summarized, followed by an analysis of the karyology and by a phytogeographic and phylogenetic discussion. Finally, four new species are described and several necessary nomenclatural changes are explained. Chromosome Cytology method The paraffin section method described earlier (Goldblatt, 1971a), was aban- doned in favor of a rapid squash technique. Root tips from sprouting corms or germinating seeds were pretreated for three to four hours in 0.05% aqueous colchicine, fixed for two minutes in acetic ethanol 1:3, hydrolyzed in 10% HC1 for five minutes at 60° C and squashed in lacto-propionic orcein (Dyer, 1963). This method yielded excellent preparations in the minimum time, and although slides could not readily be made permanent, they kept well for two to three days allowing adequate time for study. OBSERVATIONS The outstanding feature that emerges from the cytological survey of Moraea (Table 1) is the predominance of species with chromosome numbers based on either x = 10 (2n = 20, 40) and x = 6 (2n = 12, 24, 36); the few exceptions to this so far discovered are M. papilionacea, 2ri = 18, M. jugax in which 2n = 20, 16 and 12 occur, and only one of the several populations of M. ciliata examined, also 2n = 18 (Table 1). Moraea papilionacea is clearly allied to the group of species with x = 10 and must be regarded as an aneuploid species and not, as might appear, a triploid based on x - 6. The existence of a 2n = 18 population in M. ciliata (predominantly 2n = 20), requires more detailed study as its signifi- cance is not yet known, but as M. ciliata is also related to other species with 2n 20, the 2n = 18 is clearly derived. The situation in M . fuoax, in which the three populations studied have quite different numbers, n = 10, 8 and 6, is more complex. The immediate impression is that this species, if in fact it is only one species, is the link between the two main chromosome groups, which are based on x = 10 and .t = 6. This interpreta- tion does not seem correct simply on morphological grounds, since M. fugax has a very distinctive morphology which suggests it is not a direct ancestor of the large and heterogeneous group with x = 6. 1976] GOLDBLATT — MORAEA 3 Table 1. Chromosome numbers in Moraea. The following abbreviations are used: C.P. Cape Province; Tvl. = Transvaal; S.W.A. West Species M. fergusoniae L. Bol. M. gawleri Spreng. M. lugubris (Salisb.) Goldbl. M. margaretae Goldbl. M. papilionacea (L.f. ) Ker Diploid Chromosome Number M. ramosissima ( L.f. ) 13 nice M. serj)entina Baker (or as M. framcsii L. Bol.) M. vegeta L. (as M. juncea) Collection Data or Reference to Previous Work 20 20 24 20 40 18 20 20 20 Subg. Moraea Sect. Moraea C.P.: Robinson Pass, Heirmtra s.n. (MO). Botrivier-Villiersdorp, Goldblatt 216 (BOL). C.P.: Cape Point Res., (Goldblatt, 1971a). C.P.: Van Rhyns Pass, Goldblatt 223 (J). C.P.: Southern Cape Peninsula, Goldblatt, no voucher. C.P. : Near Nababeep, Goldblatt 628 (BOL ) . Flats E of Nababeep, Goldblatt 3061 (MO). C.P.: Tulbagh Road, (Goldblatt, 1971a). Constantia Nek, (Goldblatt, 1971a). Kenil- worth flats, Goldblatt, no voucher. C.P.: Stellenbosch, (Goldblatt, 1971a) (Sakai, 1 952 ) . C.P.: Springbok, Goldblatt 626 (BOL ) . Van Rhyns Pass, (Goldblatt, 1971a). Slopes W of Springbok, Goldblatt 3043 (MO). 40 km S of Springbok, Goldblatt 3041 ( MO ) . C.P.: Groot Constantia, (Goldblatt, 1971a). (Femandes & Neves, 1961). M. bolusii Baker M. ciliata ( L.f. ) Ker M. falcifolia Klatt M. macromjx Lewis M. fugax (de la Roche) Jacq. (or as M. edulis) M. fugax var. gracilis Baker M. fdicaulis Baker (as M. diphylla Baker) Sect. Deserticola 20 C.P.: Between Steinkopf and Okiep, Gold Matt 2772 (MO). Sect. Acaules 20 20, 18 18 40 20 20 C.P.: Nieuwoudtville, (Goldblatt, 1971a). Verlate Kloof, Sutherland, Goldblatt 548 (BOL). Koedoes Mts., Goldblatt, no voucher. C.P.: Glenlyon, Nieuwoudtville, Strauss s.n. (NBG 90218). C.P.: Darling-Mamre road, Goldblatt, no voucher. C.P.: Between Queen Anne and Eseljacht, Goldblatt 2498 ( MO ) . G. P. : Nieuwoudtville, Goldblatt 564 ( BOL ) . 4 km W of Steinkopf, Goldblatt, no voucher. C.P.: Calvinia dist., (Goldblatt, 1971a). Near Avontuur, Goldblatt 2860 ( MO ) . Sect. Subracemosae 12 16 28 20 12 C.P: Hopefield, (Goldblatt, 1971a). C.P.: 10 km N of Malmesbury, Goldblatt 3025 (MO). (Sakai, 1952). C.P.: Flats S of Pikenierskloof Pass, Gold- blatt 3279 (MO). C.P.: Giftberg, Goldblatt 207 (BOL). 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Species A/, coopcri Baker M. angusta (Thunb. ) Ker M. cf. angusta M. anomala Lewis (as M. ncglccta) M. ncglccta Lewis M. bituminosa (Li.) Ker M. clsiac Goldbl. A/, odorata Lewis M. "viscaria" (Li.) Ker A/, carsonii Baker A/, clliotii Baker (or as A/, sctacea) M. crici-roscnii Fries M. ))olyanthos Li. i M. polystachya (Li.) Ker A/, striata Baker (as M. trita) M. aluocnsis Goldbl. A/, aristata (de la Roche) Ascli. & Graeb. M. barnardii L. Bol. M. bcllendcnii (Sweet) N.E.Br. A/, caeca Goldbl. A/, debilis Goldbl. A/, gigandra L. Bol. Af. insole ns Goldbl. A/, loubseri Goldbl., ined A/, lurida Ker Table 1. (continued) Diploid Chromosome Number Collection Data or Reference to Previous Work Sect. Tubi flora 20 C.P.: Caledon dist., (Goldblatt, 1971a) Subg. MONOCEPHALAE 20 20 20 20 C.P.: Malmesbury, (Goldblatt, 1971a). C.P.: Bainskloof, Goldblatt, no voucher. C.P.: Elim, (Goldblatt, 1971a). C.P.: Sandy flats E of Hermanus, Goldblatt 2999 (MO). Subg. VlSCIRAMOSA 20 20 20 20 C.P.: Hermanus, Goldblatt 3325 (MO). C.P.: Kenilworth Racecourse, Esterhuyscn 32358 (BOL). C.P.: Quoin Pt., (Goldblatt, 1971a). Ronde- bosch Common, Barnard s.n. (BOL). G.P.: Giftberg, Goldblatt 353 (BOL). Near Elim, Goldblatt, no voucher. Subg. VlEUSSEUXIA Sect. Polyanthes 12 12 24 12 12 12 24 36 Zambia: Kandalila Falls, Strid 2900 (C). ( Lewis, 1966). C.P.: 15 km S of Grahams- town, Goldblatt 2864 ( MO ) . C.P.: Fort Hare, Goldblatt 455 (BOL). (Lewis, 1966). C.P.: Ladismith, Goldblatt 661 (BOL). Oudtshoorn campsite, Goldblatt 2922 (MO). C.P.: Beaufort West, (Goldblatt, 1971a). Graaff Reinet, (Goldblatt, 1971a). 10 km W of Grahams town, Goldblatt, no voucher. S.W.A.: Huns Mts., Tolkcn 3985 (BOL). (Riley, 1962). Tvl.: Johannesburg, Goldblatt, no voucher. Tvl.: Haenertsburg, Goldblatt, no voucher. Sect. Vicusscuxia 12 12 12 12 12 12 12 12 12 12 C.P.: Wildehondekloof Pass, Goldblatt 2840 (MO). C.P.: Observatory grounds, Cape Town, Goldblatt 1299 (MO). C.P.: Shaws Pass, Strauss 35 (NBG). C.P.: Napier, (Goldblatt, 1971a). C.P.: Dasklip Pass, Goldblatt 678 (BOL). C.P.: Caledon dist., Goldblatt 673 (BOL). Ex hort., Goldblatt, no voucher. C.P.: Drayton, Caledon, (Goldblatt, 1971a). C.P.: Langebaan, Loubser 2228 (NBG). C.P.: Klein Hagelkraal, (Goldblatt, 1971a). Hermanus, Goldblatt 3311 (MO). Houw Hoek Pass, Goldblatt, no voucher. 1976] GOLDBLATT— MORAEA 5 Species Af. neopavonia Foster A/, tenuis Ker M. thomasiae Goldbl. M . tricuspidata ( L.f . ) Lewis A/, tripctala (L.f.) Ker Af. tulbaghensis L. Bol M. villosa Ker Af. alticola Goldbl. M. graminicola Oberm. subsp. graminicola M. huttonii (Baker) Oberm. M. macrantha Baker Af. moggii N.E.Br. subsp. moggii Af. schimperi (Hochst) Pichi-Serm A/, spathulata (L.f.) Klatt subsp. spathulata subsp. transvaalcnsis Goldbl subsp. unknown Table 1. (continued) Diploid Chromosome Number Collection Data or Reference to Previous Work 12 12 12 24 12 12 24 24 C.P.: Heuningberg, Goldblatt 655 (BOL). C.P.: Brandvlei, Goldblatt 587 (BOL). Steinkopf, Goldblatt 2777 (MO). Hills above Hermanus, Goldblatt 3010 (MO). C.P.: Burgher's Pass, Koo, Thomas s.n. (BOL). C.P.: Jeffreys Bay, Goldblatt 2893 (MO). C.P.: Kirstenbosch hill, Cape Peninsula, Goldblatt, no voucher. C.P.: Nieuwoudtville, (Goldblatt, 1971a). Ceres, Goldblatt 667 (BOL). Koedoes Mts., Goldblatt 547 (BOL). C.P.: Gouda Common, Goldblatt, no voucher. C.P.: Tulbagh, (Goldblatt, 1971a). Rie- beeck Kasteel, (Goldblatt, 1971a). Subg. Grandiflora 12 12 12 Natal: Drakensberg, Trauseld s.n. (BOL) Natal: Mooi River, (Goldblatt, 1971a). Natal: Nottingham, Moll 2666 (PRE). 12 12 12 12 12 12 Malawi: Pawck 6966 (K, MO). TvL: Lochicl, (Goldblatt, 1971a). Malawi: Nyika, (Goldblatt, 1971a). Living- stonia dist., Goldblatt 15 (J). C.P.: Knysna, (Goldblatt, 1971a). Ilarrismith, (Goldblatt, 1971a). TvL: Graskop, (Goldblatt, 1971a). (Riley, 1962). O.F.S.: Even with the three exceptional species in which base numbers other than x = 10 and x = 6 occur, the difference between the two main groups appears remarkable and it is unlikely with about 70% of the genus known cytologically that more species will be found linking these groups, especially as most of those unknown chromosomally are specialized and closely allied to species with x = 6. Species such as M. stricta (2n = 24, 36), M. villosa, and M. tulbaghensis (both 24) are clearly polyploid and not aneuploids derived from 2n = 20. This is 9 n clear both because the species closely related to these polyploids have diploid numbers of 2n = 12, and equally from considerations of chromosome length where the polyploid does in fact have approximately twice the amount of chromosome material compared with plants having 2/i = 12 or 2ri = 20 (Table 2). The occurrence of 2n = 20 in one population ( several individuals of M. gaw- leri), while a second (only 2 specimens) has 2n = 24 is curious. This species clearly belongs to the 2n = 20 group and the anomalous record of 2n = 24 may 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Table 2. Chromosome size in several species of Moraca. Species Diploid Number Haploid Chromosome 2n Length n Af. falcifolia Klatt 20 54 M. lugubris (Salisb.) Goldbl. 20 49.5 Af. cf. viscaria (Li.) Ker 20 68 Af. angusta (Li.) Ker 20 56.5 M. barnardii L. Bol. 12 57 Af. tenuis Ker Af. alticola Goldbl. 12 61.5 12 67.5 Af. thomaaiae Goldbl. 12 61 Af. tulhaghcmis L. Bol. 24 114 he due to the presence of B chromosomes. Unfortunately, due to difficulties in growing this species, no more information is availahle. While most species each has a characteristic karyotype, notable variations occur within forms of Moraea tripetala, particularly in the size, shape, and location of the satellite. In a few other species heteroploidy has been recorded. Variations in ploidy levels occur in M. ciliata, M. elliotii, and M. tricuspidata all of which have both diploid and tetraploid populations, while M. stricta evidently has both tetraploid and hexaploid populations. Aneuploid differences occur in Af. ciliata with most populations n = 10 but n = 9 in one population and in M. fugax where M. fugax var. fugax has n = 8 and 6 while var. gracilis has n = 10. All these examples of intraspecific variation in chromosome number require more detailed study and will undoubtedly have significance at the species level. For the pur- poses of interpreting subgeneric and interspecific relationships, heteroploidy is fugax which will discussed further. ^_ „ „ _romosomes are quite large in Moraea, ranging from about 4- 8 /i in the x = 6 group and from 2-5 fi in the x = 10 group with the method de- scribed here, considerable detail of the karyotype can be seen (Figs. 1-2). Rela- tive length of chromosomes and the position of the centromeres and satellites all contribute to greater understanding of species relationships. The size and position of the satellite are particularly useful features in indicating small natural groups. Details of karyotype relating to classification are discussed in the following pages. Subgeneric Classification SYNOPSIS OF TREATMENT Subg. 1. Moraea x= 10 Plants simple to many branched. Leaves (l-)2-many. Bracts and spathes usually acute. Corm tunics pale to black, never sticky. Outer tepals reflexed, inner either erect or spreading to reflexed. Seeds small and ± spherical-angled. Type species: Af. vegeta L. Species: ca. 21 Distribution: Cape Province, from Cape Town north to the Orange River and east to Kimberley and Graham stown. 1976] GOLDBLATT— MORAEA 7 Sect. 1. Moraea Plants few to many branched. Leaves (2-)3-many. Stem erect, produced above ground. Type species: A/, vegeta L. Species: ca. 10 Distribution: Cape Province, from Cape Town north to the Orange River and east to Kimberley and Humansdorp. Sect. 2. Acaules Baker Plants simple or branched. Leaves several. Stems entirely subterranean. Type species: M. ciliata (Li.) Ker Species: ca. 4 Distribution: Cape Province, from Cape Town north to Namaqualand and east to Graaf Rienett and Grahamstown. Sect. 3. Deserticola Goldbl. 3 Plants simple or few to many branched. Leaves solitary. Stems well developed. Type species: M. bolusii Baker Species: ca. 4 Distribution: Dry areas of the northwestern Cape, from the van Rhynsdorp district in the south to southern South West Africa. Sect. 4. Subracemosae Baker Plants few to many branched. Leaves 1 or 2, with leaf inserted well above ground at base of inflorescence. Stem well developed. Capsule strongly beaked. Type species: M. fugax (de la Roche) Jaeq. (lecti Distribution: Namaqualand to southwestern Cape. Sect. 5. Tubiflora Goldbl. 4 Species : 2 Plants much branched. Leaves 2-3. Flowers solitary in spathes, with a true perianth tube and inner tepals lacking. Type species: M. cooperi Baker Distribution: Southwestern Cape. Species: 1 T 10 Subg. 2. Monocephalae (Baker) Goldbl., stat. no\ Monocephalae Baker, Fl. Cap. 6: 10. 1896, basionym (sect.) Plants unbranched. Leaf solitary, terete. Stem well developed, nodes often viscous. Bracts usually obtuse. Seeds flattened, platelike. Type species: M. angusta (Thunb.) Ker (lectotype) Species: 3 Distribution: Southwestern and southern Cape as far east as George. x = 10 Subg. 3. Visciramosa Goldbl. 5 Plants several to many branched, conspicuously viscous below the nodes. Pro- duced leaves 2. Corm tunics pale to dark brown and oily on inner surfaces. Tepals spreading to reflexed. Seeds angled. Type species: M. bituminosa (Li.) Ker Species: ca. 4 3 Sect. Deserticola Goldbl., sect. nov. Planta simplex ad multiramosa. Folium solitarium. Caulis producta. Typus: M. bolusii Baker. * Sect. Tubiflora Goldbl., sect. nov. Planta multiramosa. Folia 2-3. Flores solitarii in spathae; tubus perianthii productus; tepala interiora absentia. Typus: M. cooperi Baker. 5 Subg. Visciramosa Goldbl., subg. nov. Planta multiramosa, viscosa subnodos. Folia 2. Tunicae conni pallidi brunneae, oleosae in pagina inferiora. Tepala effusa ad reflexa. Semina angulata. Typus: M. bituminosa (L.f. ) Ker. § ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Distribution: Cape Province, from Cape Town north to Namaqualand and east to Grahams town. Subg. 4. Vieusseuxia (de la Roche) Baker Plants simple to many branched. Leaves usually solitary or developed. Bracts and spathes acute. Seeds angled or with ± spongy testa. x = 6 3-5. Stems well Distribution: South Africa to Ethiopia. Sect. 6. Pohjanthes Goldbl. 6 lectotype) Species: ca. 35 Plants few to many branched. Leaves 1-5, usually solitary. Inner tepals entire, well developed. Seeds angled. Species: ca. 10 Type species: M. pohjstachya (Li.) Ker Distribution: Ethiopia to the southwestern Cape. Sect. 7. Vieusseuxia Plants simple or few branched. Leaf solitary (rarely 2). Flowers much modi- fied, inner tepals usually trifid or much reduced to absent. Seeds angled or with ± spongy testa. Type species: M. hellendenii (Sweet) N.E. Br. Species: ca. 25 Distribution: South Africa, Transvaal to southwestern Cape. x = 6 Subg. 5. Grandiflora Goldbl. 7 Plants unbranched, usually large. Produced leaf solitary. Flowers with inner tepals entire and erect. Seeds depressed, triangulate to discoid. Type species: M. spatJiuhita (Li.) Klatt Species: ca. 16 Distribution: Ethiopia to South Africa (excluding southwestern Cape). DISCUSSION Chromosome number has been a major factor in establishing natural species groups and, as a result, in influencing the subgeneric classification. Consideration of cytological data together with morphology has resulted in the placing of species with x = 10 (9) and including M. fugax with x = 10, 8 and 6 in three subgenera: Moraea, Monocephalae and Visciramosa; while species with x = 6 were segregated in two subgenera: Vieusseuxia and Grandiflora (Fig. 3). As thus subdivided, each subgenus is believed to constitute a natural assem- blage of species with a common ancestor. The base number of x = 10 is regarded as ancestral in the genus and the three subgenera with this number are probably best regarded as of great age and, while themselves natural groups, have diverged from one another to such a degree that they cannot be reasonably accommodated in a single subgenus without also including the subgenera with x = 6. The species with x = 6 are treated as comprising the two subgenera Vieusseuxia and Grandiflora. Since they are very distinct morphologically, they are, in my 6 Sect. Pohjanthes Goldbl., sect. nov. Flanta pauci- ad multiramosa. Folia 1-5. Tepala interiora integra, producta. Semina angulata. Typus: M. pohjstachya (L.f. ) Ker. 7 Subg. Grandiflora Goldbl., subg. nov. Planta simplices. Folium solitarium. Flores gener- ates; tepala interiora integra, erecta. Semina depressa, triangulata ad disco idea. Typus: A/. spathulata ( L.f. ) Klatt. 1976] GOLDBLATT— MORAEA 9 opinion, unlikely to have evolved in a single evolutionary line from the basic stock with x = 10. Thus the chromosome number of 2n = 12 is believed to have evolved independently, at least three times, in subg. Vieusseuxia, subg. Grancli- flora, and M. fugax (subg. Moraea). Subg. 1. Moraea. — This subgenus is morphologically heterogeneous and is here divided into five sections. Section Moraea comprises several quite unspc- cialized species having from 2 to many produced leaves, usually a well developed branching system (occasionally simple in depauperate specimens) and generalized flowers with entire, ± spreading inner tepals. This type of flower is also found in the three species of sect. Acaules but here the plant itself is greatly modified with a reduced aerial stem — only the upper part of the leaves and the inflorescence emerge above ground level. While the axis is still branched in M. falcifolia, the M are unbranched. The karyotypes of sect. Moraea and sect. Acaules are, with the exception of M. papiliotmcea (2n = 18), strikingly similar: there are usually 3-4 pairs (occa- sionally 5) of long chromosomes and a corresponding number of much smaller pairs. The satellites are usually large and located on a small chromosome pair (Figs. 1B-D). Moraea vegeta and M. papilionucea (Goldblatt, 1971a: 345-346) are exceptions with satellites on long chromosomes, and this may be indicative rpl In M. ciliata and M. macronyx there are a second pair of satellites on long chromosomes as well as the pair on short chromosomes. The affinities of sect. Deserticola, unusual in subg. Moraea in having a solitary leaf, are clearly with species such as M. serpentina of sect. Moraea and the karyo- type of M. bolusii (Fig. IE) is almost identical with that of M. serpentina (Fig. ID); the chromosome complement is distinctive in comprising three long, two medium long and five small chromosome pairs. The asymmetrical corm with its black, fibrous tunics of species of sect. Deserticola is very like those of M. ser- pentina and their relationship is very close; in fact, the presence of the single leaf was the only factor in deciding on sectional status for this small alliance. f 10, 8, and 6 have been recorded, as well as the dubious In = 28 by Sakai (1952), is placed together with the closely related M. filicaulis in sect. Subracemosae of subg. Moraea. The place- ment in this group may appear inconsistent since the base number in subg. Moraea is x = 10, while lower base numbers predominate in sect. Subracemosae. The decision was influenced primarily by the nature of the karyotype in M. fugax var. gracilis, which has n = 10; the karyotype matches exactly those found in sect. Moraea, and comprises six long acrocentric chromosomes and seven much shorter pairs ( Fig. IF). Moraea fugax var. gracilis appears, from the morphological point of view, ancestral to other forms of M . fugax, as well as M. filicaulis, as it is a large much branched plant in contrast with the smaller, fewer branched forms with lower chromosome numbers. In other respects it is similar to var. fugax and M . filicaulis, having in common with these the peculiar leaf insertion high up on the stem and the beaked ovary, unique to this group. A critical survey of the cytology of the heteroploid and also morphologically variable M. fugax seems 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 A B j 10 /* • C D i j 10 /* E G H I \' Figure 1. Karyotypes in Moraea subgenera Monica, Visciramosa, and Monoccphalac. — A. M. cooped. — B. M. lugubfis. — C. M. falcifolia. — D. M. serpentina. — E. M. bolusii. — F. M. fugax var. gracilis. — G. M. fugax var. /i/gfl*. — H. M. angusta (s.l.). — I. A/, eh/ae. [Upper scale applies to A-C, H-I; lower to D-G.] 1 976] GOLDBL ATT — MORAEA 11 called for and species limits will almost certainly be redefined in the light of further cytological and morphological study. Section Tubiflora, comprising only one species, M . cooperi, resembles sect. Moraea vegetatively in having a ramified axis. Its flowers are, however, unique in the genus, having a true perianth tube; the flowers are also unusual in being solitary in each inflorescence spathe and in lacking inner tepals. Unusual as M. cooperi is, the karyotype is like those found in sect. Moraea in having long pairs and large satellites on one of the seven short pairs of chromosomes (Fig. 1A). Subg. 2. Monocephalae. — Subgenus Monocephalae is small, comprising Moraea angusta and the two species M. neglecta and M. anomala segregated by Lewis ( 1949 ) . Lewis's treatment is followed here with reservation, as the species of the section require critical morphological study. Species in subg. Monocephalae have a very distinctive morphology with a simple stem and a solitary terete leaf, but the flowers are unspecialized, large and usually yellow. Without cytological data, the affinities of the species in this section would have remained misunder- stood, for the general morphological reduction, large generalized flowers, and the flattened seeds are strongly reminiscent of subg. Grandiflora and these two sub- genera were in fact regarded as a single section by Baker in his treatment. Other features, such as the somewhat obtuse bracts and spathes, and the sticky exudate below the nodes in some forms suggest affinity with subg. Visciramosa. While the chromosome number of 2n = 20 (Fig. 1H) places subg. Monocephalae close to subg. Moraea, the karyotypes are not particularly alike and size differences be- tween the chromosomes are not at all distinct. The similarities of subg. Mono- cephalae to subg. Grandiflora are believed to be independently derived, and M. angusta and its two allies probably evolved in parallel fashion with subg. Grandi- flora to achieve the reduction of branching and leaf number while retaining an ancestral chromosome number. Subg. 3. Visciramosa. — This remarkable group stands in a somewhat isolated position. The ± 4 species, which are characterized by a highly ramified in- florescence, curiously sticky patches on the stem below each node, a distinctive corm with brown tunics covered by an oily secretion, and flowers with broad, spreading to strongly reflexed segments and occasionally free stamens, give no particular indication of relationships to other groups within the genus. The chromosomes of the three species studied are similar and the karyotype of only one species, M. elsiae, is illustrated (Fig. II). There are no sharp size differences amongst the chromosomes, though 2 to 3 larger pairs occasionally stand out; a distinguishing feature of the karyotype is the characteristically small satellite found on a small chromosome pair. This satellite location is frequent though not invariable in the x = 10 group. Subg. 4. Vieusseuxia. — Although subg. Vieusseuxia is by far the largest sub- genus with ± 35 species, about 40% of the genus, it is morphologically more uni- form in vegetative habit than the x — 10 group, though the flower is much more varied. With the exception of M. polystachya, M. polyanthos, M. carsonii, its close allies, and a peculiar form of M. tripetala, all species have a single produced leaf and relatively few or no branches. In contrast to the vegetative uniformity, the flowers of many species have become specialized in remarkable ways. J9 ANNALS OK Tilt: MISSOURI BOTANICAL GARDEN [Vol. 63 Many natural species groups can be recognized in this subgenus but only two major groups are given recognition as sections. The least specialized species are placed in sect. Polyanthes with Moraea polystachya and its allies. They include forms with a much branched axis and 3-5 leaves, as well as more specialized forms with a solitary leaf and fewer branches. The flowers are generally unmodified and are blue. The karyotypes found in this section are perhaps the least special- ized of the species with a base number of x = 6, in that they are usually fairly symmetrical, an unspecialized karyotypic feature. Apart from similarities in the karyotypes of closely related species, the karyo- types found in sect. Polyanthes are varied and suggest that the section comprises several distantly related entities. A comparison of the karyotypes of M. polyanthos (Fig. 2F) with the previously published figure for M. stricta ( as M. trita) (Gold- blatt, 1971a: 348) illustrates this contention. Sect. Polyanthes is regarded as a link between subg. Moraea and sect. Vieusseuxia but chromosomal data place it firmly in the latter. In spite of its intermediate position it is not possible to give sect. Polyanthes higher rank as species like M. elliotii clearly link the group to sect. Vieusseuxia, exhibiting strong similarity to M. algoensis and M. tripetala, and the dividing line between the two sections is almost arbitrary. Sect. Vieusseuxia is characterized by modification and reduction of the inner tepals in most species, exceptions being Moraea insolens, and some forms of M. neopavonia and M. lurida. Typically the inner tepals are small and trifid but many may be quite reduced to short entire cusps or are absent as in M . banmrdii. The trend to reduction of the inner tepals is not unique and also occurs quite inde- pendently in sect. Moraea where some forms of M. fergusoniae have trifid tepals while in M. cooperi they are absent. The section contains several distinct group- ings each with characteristic morphology and karyotype. Moraea tripetala and its allies, such as M. algoemis, M. barnardii, M. debilis and M. tenuis (Figs. 2A, B, C), have similar karyotypes, usually with a small satellite on a long chromosome pair. Some variability occurs within species espe- cially in M. tenuis and M. tripetala where satellites differ in size and position. The so-called peacock moraeas, M. neopavonia, M. villosa, M. tulbaghenis, M. louhseri, and M. caeca (Figs. 2D, E), characterized by outer tepals with a broadly ovate to circular limb, usually with a bright color and with distinct markings, have a karyotype in which one of the longest chromosome pairs is submetacentric and bears small satellites. In contrast, Moraea bellendcnii, M. lurida, M. insolens, and Af. tricuspidata, which appear to be closely allied, do not share many karyotypic features. Large satellites are located on long metacentric chromosomes in M. bellendenii and M. insolens, and are variable in M. lurida, being either large and located on a long metacentric pair or on a smaller, acrocentric pair, while in the heteroploid M. tricuspidata, the satellites are small. The karyotype of Moraea thonwsiae with its acrocentric chromosome comple- ment (Fig. 2G) is rather different from other species in sect. Vieusseuxia. This tends to confirm its rather isolated position in the alliance. Several species in this section occur in the summer rainfall area of South Africa and await cytological examination. 1976] GOLDBLATT — MORjiEA 13 A B C D E F G H i J IO t Figure 2. Karyotypes in Moraea subgenera Viensseuxia and Grandiflora. — A. A/, bar nardii. — B. M. debilis. — C. M. tenuis. — D. M. caeca. — E. M. neopavonia. — F. M. polyanthos.— G. M ihomasiae. — H. M. alticola. Subg. 5. Grandiflora. — In marked contrast to the previous subgenus, subg. Grandiflora is florally uniform and all species have large, usually yellow flowers which have erect inner tepals and spreading outer tepals. Nevertheless the group is believed to be advanced and is very modified in vegetative character. The large produced leaf is always solitary and the branching is entirely reduced (only in the rarest cases are individuals of M. huttonii, or M. spathulata ever branched). Cytologically the section is very uniform and the karyotype of M. alticola (Fig. ]4 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 2H) is representative of the group; there are two pairs of submetacentric chromo- somes, the remaining four are acrocentric to almost telocentric, and a small satel- lite is present on a long, often apparently telocentric chromosome pair. All too few species of this widespread group are cytologically known and those from tropical Africa especially require investigation. Morphological uniformity how- ever suggests that few cytological surprises are to be expected. Classif Moraea much broader than the one accepted here, as two of his four subgenera, Dietes and Helixyra, are today regarded as distinct genera, the last mentioned now Gynandriris. Baker's remaining two subgenera, Vieusseuxia and Moraea ( as Eu- moraea), are retained in name only, for the composition and circumscription are entirely altered. Owing to the poorly known state of Moraea when Baker worked, his errors are easily understood. In some cases correction of his treatment was only possible with recourse to cytological data. As an example, Baker's Mono- cephahie can be quoted; in this section he included both M. angusta (2n = 20) and M. spathulata (2n = 12), as well as two other species not conforming even to his own definition of the section. Monocephalae, treated here as a subgenus, is retained in a narrow sense including only M. angusta and its two close allies. Wherever possible, Baker's sections are retained, though often redefined so as to conform to the revised system. Thus, when possible, the older treatment is fol- lowed or emended and the present system represents a continued refinement initiated by the author (Goldblatt, 1973) tudy Evolution The underlying assumption throughout this paper has been that the x = 10 (9) group of species, i.e., subgenera Moraea, Monocephalae, and Visciramosa, comprise the more primitive part of the genus, a hypothesis based primarily on morphological evidence. Most species in subgenera Moraea and Visciramosa have more than one leaf and numerous branches, a condition believed to be the an- cestral growth form in the genus. Though specialized species with a single leaf the 10, notably the species of subgenus Monocephalae, the great majority of the species with this reduced habit, including those with very specialized flowers, have low chromosome numbers, either 8 or predominantly x = 6. As indicated earlier, there is reason to believe that the low base number of x = 6 was derived independently in at least three lines of evolution (Fig. 3). The cytological data strongly support the basic assumption that x = 10 is the Moraea 10 have approximately the same amount of chromosome material as species with n = 6 and cannot therefore be derived by polyploidy and subsequent loss of chromo- somes from the latter group. Thus, x — 6 is seen as having been derived from an ancestral base number of x = 10 by decreasing aneuploidy, and the author's ear- lier tentative suggestion to this effect (Goldblatt, 1971a: 356) is therefore main- tained. One surprising feature, however, is the paucity of intermediates between x — 10 and x = 6; Moraea M n = 9 and M. fugax with n = 10, 8, and 6 being the only known exceptions. Sub- 1976] GOLDBLATT— MORAEA 15 ANCESTRAL STOCK X = 10 X = 10 Subg. MORAEA Sect. Moraea n= 10, 9, 20 Sect. Acaules n = 10, 9, 20 Sect. Tubi flora n = 10 Sect. Subracemosae n = 10, 8, 6 / / / / / X = 6 L Subg. VIEUSSEUXIA Sect. Polyanthos n = 6, 12, 18 T Sect. Vieusseuxia n = 6, 12 X = 10 Subg. VISCIRAMOSA X = 10 Subg. MONOCEPHALAE \ \ \ \ \ \ \ \ X = 6 Subg. GRANDI FLORA Figure genera and sections in Moraea. Diagrammatic representation of presumed phylogenetic relationships of sub- Moraea Grandiflora, as well as M. fugax and M. filicaulis of subg. 6, are probably not of particularly recent origin and their independent derivation (Fig. 3) suggests that this low chromosome number has ic. Once this was achieved, rapid evolution followed. consi difl The two subgenera with x by far the largest number of species with ± 75% of the species. Though both sub- genera may be of some age, a very recent spurt of evolution appears to have Med Moraea ranean region of South Africa. The hypothesis that x with the previously expressed view that the immediate ancestor of Moraea was the genus Dietes (x = 10). Dietes, a small genus of five African species with one 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 species on Lord Howe Island, Australia, is comprised of quite large, evergreen plants, usually with a branched inflorescence, a fan of many equitant leaves, and a simple Zm-like flower with subequal spreading tepals, petaloid styles, and distinct crests. Major primitive features are the evergreen habit, absence of a perianth tube (as in most Monicas), free stamens (joined in Moraea), and a large persistent rhizome. The flower of Moraea is similar, except in obviously highly derived forms, although the inner tepals are usually smaller than the outer and the stamens al- most always are joined in a column. The vegetative form of Moraea is, in contrast, much modified; the plant is deciduous, the rootstock is a conn, and the leaves are much reduced. The usual iridacaeous feature, the equitant leaf, is so reduced in Moraea that it is represented by the morphological equivalent of an extended leaf sheath, which is terminated by a small equitant apex (Arber, 1921; Lewis, 1954; Goldblatt, 1971a). All these features suggest that Moraea evolved in response to strong seasonal conditions possibly when either increasing cold or more arid climates prevailed. The underground corm would thus insure survival during unfavorable periods, while the reduction in size and branching and the nature of the leaf suggest an adaptation for rapid growth during a short growing period. Extreme cold such as accompanied by the severe glaciations in the Northern Hemisphere and a corresponding short growing season did not prevail in Africa either during the later Tertiary when Moraea was probably evolving or more recently. Seasonal aridity, however, does occur today and has been part of the African climatic regime since at least mid-Tertiary in areas where Moraea Ls found. As the climate deteriorated from the Miocene onward, areas bordering the tropics must have been subjected to increasing stress as a result of both summer drought and cooler winters. Such a setting is one in which Moraea is believed to have evolved. That the majority of species of Moraea including all those with a basic chromo- some number of x = 10 occur today in the southwestern Cape and Namaqualand under a winter rainfall, dry summer regime, might suggest this area as the place of origin of the genus. However, the origin of the genus under such a climate seems untenable in view of the current belief that true Mediterranean type cli- mates such as found in the southwestern Cape may not be much more than three million years old (Raven, 1973; Axelrod, 1973). Moraea is almost certainly con- siderably older and the primitive species of Moraea that now all occur in the Cape region may perhaps be relicts surviving only in this area owing to lack of competition. Alternatively, Moraea may have evolved entirely in this area under different climatic conditions. It is, however, evident that the extraordinary radiation of Moraea in the Cape winter rainfall area is a direct result of the violent climatic fluctuations during the Pleistocene (e.g., Schalke, 1973) and the development of an extreme Medi- terranean climate. Most of the present day Cape species are therefore probably of quite recent origin, probably less than 1-2 million years old, with the more localized species which occupy specialized habitats probably only a few tens of thousands of years old. The center of evolution of Moraea is perhaps somewhere to the north of the 1976] GOLDBLATT — MORAEA 17 Cape winter rainfall zone in the interior of central Southern Africa, between lati- tudes 10° and 30 °S which includes Namaqualand where several primitive species occur today. Parts of this region are arid semidesert while others are dry savanna, and this area is perhaps the most sensitive to climatic changes such as envisioned in the evolution of Moraea. Today, with the exception of the eastern mountain ranges and Namaqualand, this area is the habitat for few species of Moraea but contains a scattering of unrelated species. It is of interest, though, that the most primitive species of Ferraria* (base number x = 10), F. ghitinosa, the only freely branched species of this otherwise Cape genus, occurs throughout this zone. It is and perhaps evolved from Dietes in much the same way as allied to Moraea also once have inhabited this region. rn difl Grandifl also in West Africa, is probably a fairly recent development. These subgenera are undoubtedly the most specialized and thus probably of comparatively recent origin, although the montane habitat may be of considerable age. NOMENCLATURAL NOTES AND New SPECIES Although a detailed revision of the winter rainfall area species of Moraea is planned, and this is where most nomenclatural changes will be made, I have used what are now known to be correct specific names in this paper and several changes in current usage require explanation. The cytology of 4 undescribed species was discussed earlier in this paper and these new species are described bel ow. 1. Moraea gawleri Spreng., Syst. Veg. 5 (Index): 462. 1828, nom. nov. pro M. crispa (L.f. ) Ker. M. crispa (Li.) Ker, Curtis's Bot. Mag. tab. 754. 1804, horn, illeg., non Thunb., 1787. Iris crispa Li., Suppl. Pi. 98. 1781. Moraea nndulata Ker, Cen. Irid. 43. 1827, horn, illeg., non Thnnb., 1787. M. dccussata Klatt, Abh. Naturf. Ges. Halle 15: 367. 1882; Erganz. 33. 1882, syn. nov. Moraea gawleri, a new name proposed by Sprengel (1828) for the illegitimate homonym M. crispa (L.f.) Ker, is the eorreet name for the speeies variously known in herbaria as M. crispa (L.f.) Ker, M. undulata Ker, or M. decussata Klatt. The former two names are later homonyms while M. gawleri predates Klatt's A/, decus- sata by more than 50 years. 2. Moraea serpentina Baker, Handbook Irid. 52. 1892. Moraea framesii L. Bob, S. African Gard. 17: 418. 1927, syn. nov. Moraea serpentina Baker is regarded as a eonspeeific with Af . framesii, a com- mon N.W. Cape and Namaqualand species. The type specimens of these two species are very alike and fit within the author's concept of M. serpentina. 8 The chromosome count for this species, made by the writer, is as yet unpublished. ]Q ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 3. Moraea lugubris (Salisb.) Goldbl., comb. nov. Ferraria lugubris Salisb., Prod. Stirp. 42. 1796. type: Cape Town, base of Devils Peak, Thun- berg (UPS, lectotype). Moraea iriopetala Li., Suppl. PI. 98. 1781, nom. illeg., superfl. pro M. vegeta L. M. plumaria (Thunb.) Ker, Konig & Sims Ann. Bot. 1: 240. 1805, syn. nov. Iris plumaria Thunb., Diss. Irid. no. 16. 1782, nom. illeg., superfl. pro M . vegeta L. Moraea mira Klatt, Trans. S. African Philos. Soc. 3: 202. 1885, syn. nov. Moraea lugubris (Salisb.) Goldbl., better known as M. plumaria (Thunb.) Ker has had a complex nomenclatural history. This plant was first given the name M . iriopetala by the younger Linnaeus, this being known from the inscription on several specimens in the Linnaean herbarium. However, the way in which M. iriopetala was described makes the name illegitimate, and in fact nomenclaturally a superfluous synonym for M. vegeta L. The younger Linnaeus actually described two varieties under M. iriopetala, one (presumably the typical variety) listed with the synonym M. vegeta L., and a second with the name M. juncea L. cited. Subsequently, Thunberg (1782) described Iris plumaria, a species clearly match- ing the specimens annotated M . iriopetala in the Linnaean Herbarium. Unfor- tunately Thunberg cited as synonyms not only the illegitimate name M. iriopetala, but also M. vegeta and M. juncea, thus invalidating Iris plumaria. Before Ker (1804) transferred Iris plumaria to Moraea in which genus it can be treated as a legitimate new species, Salisbury (1796) provided a new name for it, Ferraria lugubris. Salisbury cited M. iriopetala and Iris plumaria as synonyms but both names, being illegitimate, were correctly not used. Salisbury's Ferraria lugubris is clearly a synonym of Thunberg's Iris plumaria and the younger Linnaeus's M . iriopetala (as this applies to material in the Linnaean collection), and Salisbury may even have seen the Linnaean specimens so annotated. Thunberg's specimens of this sDecies in the Thunbenj Herbarium at Uppsala are chosen as lectotype. 4. Moraea fergusoniae L. BoL, S. African Gard. 19: 294. 1929. Moraea fimbriata Klatt, Linnaea 34: 561. 1866, horn, illeg., non Loisel, 1822; syn. nov. Moraea fergusoniae and M. fimbriata are names both in current vise. Alt when described, M. fergusoniae was believed distinct from M . fimbriata, r name A/, fergusoniae must however be used as M. fim is a later homonym. 5. Moraea vegeta L. is the correct name of the species known for many years as M. juncea or as M. tristis. For an explanation of this change in usage see Barnard & Goldblatt (1975). 6. Moraea filicaulis Baker, Handbook Irid. 56. 1892. Moraea diphylla Baker, Bull. Misc. Inform. 1906:24. 1906, syn. nov. Moraea filicaulis is an older name for M. diphylla and the latter is reduced to snyonymy. 7. Moraea elsiae Goldbl., nom. nov. pro Homeria simulans Baker. Homeria simulans Baker, Fl. Cap. 6: 529. 1896. type: South Africa, Cape, Kenilworth, Cape Peninsula, H. Bolus 7931 (BOL, holotype; K, MO, isotypes). 1976] GOLDBLATT— MORAEA 19 Moraea elsiae is a new name proposed here for Homeria simulans Baker. In spite of the floral similarity of M. elsiae to Homeria, the whole appearance of the plant with its many branches, sticky internodes, short obtuse bracts, and oily corm tunics makes it clear that the plant is correctly placed in Moraea subg. Viscira- mosa. The original reason for placing it in Homeria was the reduced style crests but this character is now recognized as occurring in several species of Moraea also (Goldblatt, 1971b). The species is named in honor of Miss Elsie Esterhuysen, the indefatigable Cape botanist who is endeavoring to conserve its best known locality, Kenil worth Racecourse in Cape Town. A new name was necessary as M. simulans Baker (— Gijrandriris) blocks the transfer from Homeria. 8. For the explanation of the usage of Moraea striata Baker (for M. trita N.E. Br.), M. falcifolia Klatt (for M. fasciculate and M. galaxioides) and Af. elliotii (for M. mucra and M. stewartae) see Goldblatt (1973). NEW SPECIES Subg. Moraea sect. Moraea 1. Moraea margaretae Goldbl., sp. nov. type: South Africa, Cape, Namaqua- land, pipeline track, SW of Nababeep, Goldblatt 628 (BOL, holotype; K, MO, PRE, S, isotypes). Planta parva, ad 15 cm alta, ramosa. Folia producta 2-3, lineares, glabra, canaliculata an teres supra. Flores pallidi-lutei; tepala exteriora ad 3 cm longa, limbis 2 mm longis; tepala interiora erecta, lanceolata. Plants to 15 cm high, usually 1-2 branched. Corm 5-7 mm in diameter, the tunics brown, coarsely fibrous, the inner layers entire. Leaves 2-3, linear, canalicu- late, often terete and twisted near apex. Stem glabrous, branching usually from the base. Spathes herbaceous, with dark brown, acute or lacerated apices; inner spathe 3-4.5 cm long, outer 2-3 cm long. Flowers few, pale yellow; outer tepals 2-3 cm long with limb 1.5-2 cm, spreading to reflexed; inner tepals erect, spread- ing later, lanceolate, obtuse, to 2.0 cm. Filaments ± 5 mm long, joined for 4 mm; anthers 4-5 mm, red. Style branches ± 7 mm with lanceolate crests 6-10 mm long. Capsule and seeds unknown. Chromosome number In = 40. Flowering time: Late September and October. Distribution: Coarse sandy soils in Namaqualand; more common than the record suirtrests. This diminutive species is known from few collections but is nevertheless quite common in central Namaqualand, a very arid region of the Cape winter rainfall region. Its pale yellow, strongly veined flowers conform to the usual pattern for the genus, but its vegetative features, especially the characteristic branching from near the base and its 2 or 3 produced leaves, indicate a position in section Moraea and suggest particularly a close relationship with Af. papilionacea. The basic chromosome number of x — 10 confirms its sectional position. The diploid num- ber of 2n = 40 for the type population ( only two individuals examined ) suggests this species is tetraploid. Moraea margaretae is named after my wife whose com- pany and help on field trips has been invaluable. 2() ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 South Africa, cape: 29.17 (Springbok): 9 4 km W of Steinkopf (HA), GoldbJatt 2775 (MO). Koufontein, Steinkopf dist. (BC), Her re s.n. (STE 11835). Pipeline road S of Naba- beep (CA), Goldblatt 628 (BOL, K, MO, PRE, S). 8 km E of Nababeep (CB), Goldblatt 3061 (NBG, MO, PRE). 30.17 (Hondeklipbaai): 8 km N of Caries (BD), Leighton 1129 (BOL). Braekdam, hills, Schh'chter 11120 (BM). Without precise locality: Namaqualaiid minor, Scully 134 (BM). Subg. Vieusseuxia sect. Vieusseuxia 2. Moraea thomasiae Goldhl., sp. nov. type: South Africa, Cape, Koo (list., Burgher's Pass, Goldblatt 2422 (MO, holotype; K, NBG, PRE, S, isotypes). Planta parva, 15-20 cm alta, simplex. Cormus ad 1 cm diameter, tunicis atrobnmneis ad nigris. Folium productum solitarium, cananiculatum, glabrum, inflorescentiam excedentum. Flores lutei; tepala exteriora ad 3.5 cm longa, limbis ad 2 cm longis; tepala interiora lanceolata, ad 2 cm longa. Plants small, slender, 15-20 cm high. Conn about 1 cm in diameter, the tunics dark brown to black. Produced leaf solitary, ± basal, linear, about 3 mm wide, glabrous, exceeding the inflorescence. Scape simple. Spathes herbaceous with brown attenuate apices; inner spathe to 6 cm long, outer about half the length of the inner. Flowers yellow; outer perianth segments with a narrow, long, erect claw, 1.5 cm long, the limb reflexed, about 2 cm long; inner segments erect, nar- rowly lanceolate, up to 2 cm long. Filaments about 5 mm long, tree almost to the base; anthers about 6 mm long. Style branches about 1.5 cm long, bearing linear crests about 1 cm long. Capsules and seeds not known. Chromosome number 2/i 12. Flowering time: July to mid September. Distribution: Clay and shale slopes, usually on a south facing slope, in the Montanu and Worcester districts. Moraea thomasiae grows in the Worcester and Montagu districts of the south western Cape in fairly arid, semikaroid areas, where it grows characteristically on south facing shale slopes. It is a very distinct species with no close relatives. Its single leaf and slender, often unbranched stem suggests that it is best placed in sect. Vieusseuxia. Its simple flowers with entire inner tepals are not usual in the section, although these are known, for example, in M. incurva. The karyotype with 2u — 12, supports its subgeneric placement, but the predominantly acrocen- tric chromosomes do not suggest a close relationship with other species in sect. Vieusseuxia. Its somewhat superficial similarity to the A/, angusta complex is belied by its entirely canaliculate leaf, brown to black corm tunics, and above all, its very acute bract leaves and spathes. The difference in karyotype (A/, angusta has 2/i = 20) makes it quite clear that A/, thomasiae is not even remotely related to A/, an gust a. The species is named in honor of Margaret Thomas, an enthu- " Note on Citation of Specimens. — The arrangement of specimens examined, following the taxonomic treatment and discussion of each species, is based on the system currently gaining acceptance in South Africa (Edwards & Leistner, 1971). The system is based on a grid, and geographical degrees of latitude and longitude define each grid which is numbered accordingly. The one degree square grids are also designated by the name of a major town within it. Grids arc 4 divided into four quarters labelled from left to right A, B, C, or D and these quarter degree squares are again divided into four and labelled A, B, C, or D. Thus, all specimens are cited with data localizing them to one-sixteenth of a degree square. 1976] GOLDBLATT— MORAEA 21 ! siastic and untiring South African bulb grower and collector. The living material of the many species of Iridaceae she has provided me with is gratefully acknowl- edged. South Africa, cape: 33.19 (Worcester): Hex R. Pass, near summit (BO), Mauve ir Oliver (STE). Karoo Garden, Worcester (CD), Bayer 7 (NBG); Goklblatt 2422 (MO, NBC PRE, K). Onse Rug Farm, Worcester (list., Barker 9446 (NBG). 33.20 (Montagu): Oudeberg, Montagu dist. (GA), Acocks 20539 (NBG, PRE). Burger's Pass, Koo dist. (DB), Thomas s.n. (BOL); Mauve 6- Oliver 197 (STE). Vrolikheid, Mc- Gregor dist. (DD), Jooste 154, 181 (both STE). 3. Moraea debilis GoldbL, sp. nov. type: South Africa, Cape, clay slopes S\V of Caledon, Goklblatt 673 (BOL, holotype; K, MO, PRE, S, isotypes). Planta gracilis, ad 15-40 cm longa, ramosa. Cormus ad 1 cm diam., tunicis pallidis. Folium productum solitarium basale, canaliculatum, pubescentum an marginibus ciliatibus. Flores purpurei decolorentes ad malvini pallidi; tepala exteriora ad 2 cm longa; tepala interiora ad 1 cm longa, erecta, filiformia, plerumque tricuspidata. Plants slender, 15-40 cm high, usually branched. Corm ± 1 cm in diameter, the tunics pale, finely fibrous. Leaf solitary, basal, linear, pubescent on outer sur- face and/or ciliate on margins, exceeding the inflorescence. Stem laxly branched, rarely simple, the bract leaves dry. Spathes herbaceous, or dry above, the apex attenuate or lacerated; inner spathe 4-5.5 cm long, outer ± half the inner. Flowers purple, fading to a pale mauve and becoming lightly speckled; outer tepals spread- ing, ± 2 cm long with a slender bearded claw and a lanceolate limb ± 1 cm long; inner tepals ± 1 cm long, erect, filiform, usually tricuspidate with the central cusp much exceeding the laterals. Filaments ± 5 mm long, joined for about half the length; anthers ± 5 mm long. Style branches ± 8 mm long, the crests lanceolate, to 3 mm. Capsule narrowly ovoid-clavate. Seeds angled. Chromosome number 2/7 = 12. Flowering time: Late September and October. Distribution: Clay soils in the Caledon district, southwestern Cape. This slender, almost spindly plant with its small mauve flowers and reduced inner tepals is clearly allied to the well known and widespread Moraea tripetala. M. debilis is known from several collections, all from the Caledon district of the south western Cape, where it grows in clay soils amongst small shrubs and is typically late flowering. It is distinguished by its fine, pale corm tunics, pubescent leaf and small flowers with their threadlike, trifid to tricuspidate inner tepals. The karyotype, with a diploid number of 2n — 12, is very characteristic of A/. tripetala and its close allies. South Africa. Cape: 34.19 (Caledon): SW of Caledon (AC), Barnard s.n. (BOL 30694); Goklblatt 673 (BOL, K, MO, PRE, S). Between Bot River and Caledon, Mauve 4893 (PRE). Ca. 12 km E of Caledon (BA), Barker 10851 (NBG). Without precise locality: Caledon dist., Pappe s.n. (SAM 70709); Leipoldt 3562 (BOL); Schlechter 5527 (BOL, PRE). 4. Moraea caeca Barnard ex Goldbl., sp. nov. type: South Africa, Cape, top of Dasklip Pass near Porterville, Goklblatt 678 ( BOL, holotype; K, MO, PRE, S, isotypes ) . Planta 20—40 cm alta, raro ramosa. Folium productum, solitarium, canaliculatum, in- florescentiam excedentum. Flores malvescenti; tepala exteriora ad 3 cm long, ungue canalicu- 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 lato, pubescenti, limbis obovatis, ca. 2 cm latis; tepala interiora trieuspidata, cuspidc centrale, 5-8 mm longa, lineare, cuspidibus lateribus 2 mm iongis, obtusis. Plants slender, 20-40 cm high. Corm ± 1 cm in diameter, with light brown fibrous tunics. Leaf solitary, linear, glabrous, exceeding the inflorescence. Stem glabrous (occasionally pubescent), usually simple. Spathes herbaceous or dry above with brown attenuate apices; inner spathe ± 5 cm long, outer to ± 2.5 cm. Flowers mauve-lilac with a small yellow or black nectar guide; outer tepals spreading, ± 2.8 cm long with an erect, channelled, pubescent claw 8-12 mm long and a broadly ovate, spreading limb, to 1.8-2.2 cm wide; inner tepals tri- cuspidate, channelled, the central cusp 5-8 mm long, laterals to 2 mm. Filaments 2-3 mm long, united near the base only; anthers dark, ± 5 mm long. Style branches ± 5 mm long, the crests acute to obtuse, to 7 mm long. Capsule clavate, to 1.5 cm. Seeds angular. Chromosome number 2n — 12. Flowering time: Late September and October. Mou ville, and locally on the Cape Peninsula, southwestern Cape; confined to sandy soils. Moraea caeca is an attractive late-flowering species allied to the "peacock Moraeas," the usually large flowered, highly colored species with very broad, out- spread outer tepals with conspicuous nectar guides. This species is smaller than others of the group and instead of the bright nectar guide of yellow circled with contrasting colors, the guide is small and dark or absent. It grows in sandstone areas and is known from three somewhat isolated areas, the Piketberg and the Twenty Four Rivers Mountains on opposite sides of the Piketberg-Porterville valley and from a single collection from the Karbonkelberg on the Cape Penin- sula. The karyotype with In =12 is similar to that of M. neopavonia and A/, vil- losa (although the latter is tetraploid) in the number of metacentric and submeta- centric chromosomes and in the position of the satellite. South Africa, cape: 32.18 ( Clanwilliam ) : Kloof on SW side of the Piketberg (DC), Barnard s.n. (BOL). Moutons Valei, Marloth 11509 (PRE). Hills NW of Moutons Valei, Pillans 7488 (BOL). Piketberg (DD), Vfok s.n. (NBG 58948). Beyond Piketberg on Rede- linghuys road, Barnard s.n. (SAM 52394, BOL). 32.19 (Wuppertal): Dasklip (Cardouw) Pass above Porterville (CC), Barker 7599 (NBG, STE); Esterhuysen 16211 (BOL); Goldhlatt 678 (BOL, MO, K, PRE). Mountain above Porterville, Louhscr 856 (NBG). 33.19 (Worcester): Drieboscb, Groot Winterhoek (AA), Hatjnes 858 (STE). 34.18 ( Simonstown ) : Karbonkelberg, Cape Peninsula (AB), Salter 3288 (BM). Literature Cited Arber, A. 1921. The leaf structure of the Iridaceae considered in relation to the phyllode theory. Ann. Bot. (London) 35: 301-336. Axelrod, D. I. 1973. History of the Mediterranean ecosystem in California. Pp. 225-277, in F. di Castri & H. A. Mooney (editors), Mediterranean Type Ecosystems: Origin and Structure. Springer-Verlag, New York. Baker, J. G. 1896. Irideae. In W. T. Thiselton-Dyer (editor), Flora Capensis. Vol. 6: 7- 71. L. Reeve & Co., London. Barnard, T. T. & P. Goldblatt. 1975. A reappraisal of the application of specific epithets of the type species of Moraea and Dietes (Iridaceae). Taxon 24: 125-131. Dyer, A. F. 1963. The use of lacto-propionic orcein in rapid squash methods. Stain Technol. 38: 85-90. 19761 GOLDBL ATT— MORAE A 23 Edwards, D. & O. A. Leistner. 1971. A degree reference system for citing biological records in Southern Africa. Mitt. Bot. Staatssamml. Miinchen 10: 501-509. Fernandes, A. & J. B. Neves. 1961. Sur la caryologie de quelques Monocotyledones Afri- cains. Compt. Rend. IVe Reunion A.E.T.F.A.T.: 458-463. Goldblatt, P. 1971a. Cytological and morphological studies in the southern African Irida- ceae. J. S. African Bot. 37: 317-460. . 1971b. Moraea insolem. Fl. Pi. Africa 41: tab. 1639. . 1973. Contributions to the knowledge of Moraea (Iridaceae) in the summer rain- fall region of South Africa. Ann. Missouri Bot. Gard. 60: 204-259. Lewis, G. J. 1949. Moraea angnsta (Thunb.) Ker and allied species. J. S. African Bot. 15: 115-120. . 1954. Some aspects of the morphology, phylogeny and taxonomy of the South African Iridaceae. Ann. S. African Mils. 40: 15-113. Lewis, W. II. 1966. Chromosomes of two Moraea (Iridaceae) from Southern Africa. Sida 1: 381-382. Raven, P. H. 1973. The evolution of Mediterranean floras. Pp. 213-224, in F. di Castri & H. A. Mooney (editors), Mediterranean Type Ecosystems: Origin and Structure. Springer- Verlag, New York. Riley, H. P. 1962. Chromosome studies in some South African Monocotyledons. Canad. J. Genet. Cytol. 4: 50-55. Sakai, B. 1952. Zytologische Untersuchung bei Iridaceen Uber die Karyotypen verschiedener Arte der Unterfamilie Iridoideae. Cytologia 17: 104-111. Schalke, H. J. W. G. 1973. The Upper Quaternary of the Cape Flats Area (Cape Province, South Africa). Scripta Geol. 15. EVOLUTION OF THE SANTA LUCIA FIR (ABIES BRACTEATA) ECOSYSTEM 1 Daniel I. Axelrod 2 Abstract Abies scherrii Axelrod is described from tlie Miocene of western Nevada, and it resembles the living A. bract eat a of the Santa Lucia Mountains, coast-central California, a unique endemic representing the sole member of the subgenus Pscudotorrctja. Although the fossil species A. chancyi Mason and A. longirostris Knowlton have previously been considered allied to A. bracteata, those species are extinct, they are not closely allied to bracteata, and they are only distantly related to living Asian firs. A. scherrii occupied an ecotone between broadleaved ever- green sclerophyll forest and mixed conifer forest during the Miocene in western Nevada, and its descendant A. bracteata has a similar occurrence today. The Miocene communities were much richer than the living, including species whose nearest descendants occur only in regions with summer rainfall, or now occupy more restricted areas in California. The latter include Sequoia- dendron of the central and southern Sierra Nevada, species of Chamaecyparis and Picea that are confined to the Klamath-Siskiyou region of northwestern California, and species of Abies and Pinus that are largely subalpine in the Sierra Nevada. Abies scherrii and its associates shifted coastward as colder and drier climate developed over the interior. Abies bracteata has survived in a near-coastal climate where evaporation rate is not so high as in the interior or in areas to the south, and where it is largely removed from regular heavy winter snow and ice. Some of its associates were segregated into their present areas as the summer-dry mediterranean climate emerged in the Quaternary. Sequoiadendwn was confined to the relatively drier and sunnier central and southern Sierra Nevada where there was sufficient light in an open forest to enable it to reproduce. Chamaecy parts and Picea were restricted to the northwest sector where there is a longer precipitation season, some summer rain, and where evaporation rate is lower than in the Sierra. Present subalpine species of Abies, Pinus and Tsuga were eliminated from the mixed conifer forest in the Sierra Nevada as evaporation rate increased during summer and produced conditions inimical for seedling establishment, though they still persist in the upper mixed conifer forest in moister, more equable areas to the northwest A. bracteata may have had a wider distribution in the Coast Ranges during the moister phases of the Quaternary, together with the mixed conifer forest species. It was apparently restricted with them as drier, hotter climates spread during the later postglacial periods. Santa Lucia fir (Abies bracteata) is confined to the middle and upper slopes of the Santa Lucia Mountains, coast-central California (Figs. 1-2). Among the 60-odd species and varieties of fir it is the sole member of the subgenus Pseudo- torreya, the remainder representing the subgenus Abies (Liu, 1971). Its unique- ness is seen in the long-fusiform to ovoid-conical resinless winter buds, the cones with exserted awllike bracts 2 to 4 cm long, the sharply-pointed deep green needles like those of Torreya, the very thin bark (and hence the need for fire protection), and its tall spirelike habit that recalls that of Abies lusiocarpa or Picea engelmannii near timberline. The problem of its geologic history was raised by the recent discovery of a slab of shale on which are preserved three cone scales with long, exserted awllike bracts of a fir that indubitably represent a species similar to the living A. bracteata. This fossil, from the Late Miocene (13 m.y.) Purple Mountain flora of western Neveada, supplements a needle collected earlier at a nearby site that is similar to 1 This research project has been supported by the National Science Foundation and its support is gratefully acknowledged. The manuscript has benefited from critical reviews by Peter II. Raven and Steven N. Talley. 2 Department of Botany, University of California, Davis, California 95616. Ann. Missouri Bot. Card. 63: 24—41. 1976. 1976] AXELKOD— SANTA LUCIA FIR ECOSYSTEM 25 Figure 1. Present occurrence of Abies bracteata (see Griffin & Critchfield, 1972), and of the closely similar A. scherrii in the Miocene of Nevada. those produced by A. bracteata. In addition, new collections of the slightly older Middlegate flora ( Axelrod, 1956), situated about 80 miles southeast, have yielded a needle and winged seed that also represent fossil Santa Lucia fir. In each area the associated flora is composed of broadleaved sclerophylls (Arbutus, Castanopsis, Lithocarpus, Quercus) similar to species that live with Santa Lucia fir today. Both floras also have species of Abies, Picea, Pinus, Pseudo- tsuga, Chamaecyparis, Sec/uoiadendron (Figs. 4-20) and numerous dicots that now contribute to mixed conifer forest in the Sierra Nevada, Siskiyou-Klamath Mountains, and the high Coast Ranges. The Purple Mountain and Middlegate records thus provide a basis for comparing the Miocene ecosystem at two dif- ferent sites of slightly different age, and for outlining the post-Miocene history of the community. Geologic Occurrence PURPLE MOUNTAIN FLORA The site that yielded the Purple Mountain fossils allied to the living Santa Lucia fir is in the Truckee River canyon southwest of Wadsworth, Nevada ( Figs. 1, 3). Regional geologic reports (Rose, 1969; Bonham, 1969), coupled with my 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Figure 2. Santa Lucia fir on east slope of Cone Peak, altitude ~ 4,300 feet. Pinus lam- hertiana on left. Quercus chrysolepis in foreground is part of die evergreen sclerophyll forest that covers the nearby and distant slopes and includes Arbutus menziesii, Lithocarpus demi- florus, and Quercus wislizenii as codominants, all represented by fossils associated with Abies scherrii in the Nevada Miocene. own more detailed mapping in this loeal area, show that the plant-bearing beds are in the lower part of the Chloropagus Formation. It is composed chiefly of andesite flows but includes mudflow breccias and interbedded thin sections of limestone and organic shale that contain the remains of plants that lived on the borders of shallow ponds and small lakes. The Chloropagus is overlain by welded dacitic tuffs of the Kate Peak Formation, dated in the nearby region at 12-13 m.y. (Bonham, 1969). The Chloropagus rests unconformably on a 50-foot rhyo- lite tuff that has been correlated with the Old Gregory Formation in the hills west of Fallon. It lies unconformably on black flows of Alta Andesite, which in turn rests on the varicolored welded tuffs of the Hartford Hill Rhyolite (22 m.y.). The Hartford Hill covers a peneplaned basement of high grade metamorphic rocks intruded by granodiorite of Cretaceous age. 1976] AXELROD— SANTA LUCIA FIR ECOSYSTEM 27 ■ L . - - "'Vi yii ... . t T . . . '"". . :<> fr - : ;»:■ ::=:■&: : - . ■ \ ■ ■ >:. ■_■ ^*W^** i '. ■ . (fflttw ;. ■ /■■<•- ~ ♦ % «- V ':■ ■ - ■ ■ ' <& : - ■ ■ ■ -W.^--- "V - tfW* ■ • '"5 ■ '■■■-■■:: • .. ' s A ■ -H.,.. » *• -VW %» "^ -■*■ ■ ■ .f >; ■■■ ■■•■ -:, '^^ % ' ■ - . » ■ ■ & \ . - •■;■ ■ Figure 3. The Late Miocene Purple Mountain flora occurs in thin lake beds intercalated with andesites of the Chloropagus Formation on the low spur at the right, in the bank of the drainageway, and at the position of the observer. View is north, across Truckee River floodplain. Flows in the basal part of the Chloropagus Formation in Fort Defiance Can- yon 10 miles north of the fossil locality have been dated as 14 m.y., and a vitric tuff in the upper part of the sequence in Pierson Canyon 5 miles northwest is 13 m.y. The flora is therefore about 13.5 m.y. and is correlative approximately with the Late Miocene Fallon, Chloropagus, and Aldrich Station floras in the nearby region (Axelrod, 1956). MIDDLECATE FLORA This flora comes from a site on the north side of Middlegate basin, 5 miles northwest of Eastgate, Nevada (Axelrod, 1956). The plants are preserved in white to light gray, well-bedded opaline shales interbedded with fine vitric tuff now altered to bentonite. M gate Formation. They grade down into soft dark mudstone, very thin opaline shale, and with local conglomerates situated opposite the mouths of small streams that entered the lake. Middleg and flows of the Clan Alpine Volcanics directly north of the site. Since the flora was derived from volcanic slopes facing south, the warm dry exposure accounts for the rich representation of sclerophylls there. The Middlegate is overlain conformably by fine sandstone, mudstone, tuff, and conglomerate of the Monarch Mill Formation, the basal part of which has now yielded a rich (30+ taxa) mammal fauna of transitional Hemingfordian- Barstovian (~ 16 m.y.) age. 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Figures 4-20. Purple Mountain conifers associated with Abies scherrii. — 4. Abies scherrii Axelrod, no. 5489. — 5b. Abies scherrii Axelrod, no. 5491 from Middlegate; 5a, 5c. Abies brae- teata needles for comparison. — 6. Abies scherrii Axelrod, no. 5492 from Middletfate. — 7-9. Pseudotsuga sonomensis Dorf, nos. 5493-5495. — 10-11. Chamaecyparis sierrae Condit, nos. 5496-5497. — 12. Abies concoloroides Broun, no. 5498. — 13-14. Finns quinifolia Smith, nos. 5501-5502.— 15-16. Abies klamathcnsis Axelrod, nos. 5499, 5500.— 17-18. Picea sonomensis Axelrod, nos. 5503, 5504. — 19-20. Sequoiadendron ehaneyi Axelrod, nos. 5505, 5506. 1976] AXELHOD— SANTA LUCIA FIR ECOSYSTEM 29 Systematic Considerations The Nevada occurrences of a Miocene fir similar to the living A. bracteata is not the first report of a fossil that has been presumed related to it. Abies chaneyi Mason from Oregon (Mason, 1927; Chaney & Axelrod, 1959) and A. longirostris Knowlton from Colorado (Knowlton, 1923; MacGinitie, 1953) have also been considered allied to A. bracteata. However, comparisons now indicate they are not closely related to it, and some of these fossils certainly represent genera other than Abies. Thus, it is necessary first to clarify the status of the fossil records that have been presumed allied to A. bracteata, following which the history of Santa Lucia fir, and the community of which it forms a part, can be outlined in pro- visional manner. Abies scherrii Axelrod sp. nov. — Figs. 4, 5b, 6. Cone scales 9-12 mm broad, 6-8 mm long, broadly oblong to ovate, distal end truncate to broadly rounded, proximal part truncated, with thin, straight woody peg of attachment; bract exserted, over 2.3 cm long, awllike, 1 mm wide for most of length but widening to 2 mm in proximal 3 to 4 mm; lateral appendages not visible on exposed upper surface. Needles 2.0-2.8 cm long, broadest at middle, 2.5 mm; apex sharply acuminate, petiole somewhat curved, sharply truncated. Winged seed 12 mm long, wing terminal, 5 mm long, somewhat torn, about 5 mm wide distally; seed long-oval, 7 mm long, 3 mm broad. This species is represented in the Purple Mountain flora by a slab containing 3 cone scales with bracts and by a second specimen on which is a poorly pre- served needle. In the Middlegate flora a winged seed and a needle are referred to this species. The cone scales and attached bracts are similar to those of the living A. bracteata of the Santa Lucia Mountains, coast-central California. The lateral lobes of the bract are not visible, but this appears to be the result of curling so that they cannot be seen on the lower (inner) preserved surface. The needles and the single winged seed are also similar to those produced by the living Santa Lucia fir. This species is named for Annette K. Scherr, a student in my course in Forest History who, during a class field-trip to the Purple Mountain area, collected the slab of shale on which are preserved the diagnostic cone scales of fossil Santa Lucia fir, AI)ies scherrii. Occurrence: Nevada, Purple Mt.: LLC Mus. Pal. holotype no. 5489, hypotype no. 5490. Nevada, Middlegate: U.C. Mus. Pal. hypotype nos. 5491, 5492. Abies chaneyi Mason, Publ. Carnegie Inst. Wash. 347: 149, pi 4, figs. 1 and 7 (winged seeds only), fig. 2. 1927. Chaney & Axelrod, Publ. Carnegie Inst. Wash. 617: 137, pi 11, fig. 3. 1959. Axefrod, Univ. Calif. Publ. Geol. Sei. 51: 141 (winged seeds only). 1964. These eited specimens are winged seeds except for one that represents a cone scale with attached bract (Mason, pi. 4, fig. 2 only). The seeds have a slender, generally oblong to long-oval outline and the long narrow wing is attached high 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 up on the seed and is not appreciably widened or asymmetrical distally. The cone scale has a long exserted bract that is much wider at the distal edge of the scale than those of the living A. hracteata. These fossils are not closely related to any living fir. Although the winged seeds resemble those produced by the living A. hracteata, those of other species — notably A. chensiensis Van Tiegham of central and southern China and A. delavayi Franchetti of southern China — are also similar to them. The winged seeds of A. chaneyi resemble those of A. longirostris Knowlton from the Creede flora of Colorado (Knowlton, 1923), though the Creede specimens tend to average some- what smaller in size. However, the Creede fir has cone scales with exserted M In this regard, the Creede A. longirostris appears distantly related to A. delavayi of southern China, as judged from the winged seeds as well as the conspicuously exserted acuminate bract. Further affinity is also seen in the fact that typical fir foliage is well represented in the Creede flora. It is like that produced by many montane species, with the needles curved upward on the branchlets, a feature also exhibited by A. delavayi. Present evidence indicates that A. chaneyi is more nearly related to A. longi- rostris Knowlton than to any other fossil or living fir, and it may have been derived from the Creede species. Although both species seem distantly related to A. delavayi of southern China, there is no evidence of any close affinity between them and the living A. hracteata of the Santa Lucia Mountains. Occurrence: Oregon, Mascall: U.C. Mus. Pal., holotype no. 135, hypotype no. 5488, A, B (on same slabs as foliage specimens of Cephalotaxus, = "Abies chaneyi" branch let, nos. 134, and 130), 2826, 2828 (its counterpart), homeotype no. 2827. Oregon, Beulah: U.C. Mus. Pal. homeotypes nos. 779—780 (winged seeds only). Cephalotaxus bonseri (Knowlton) Chaney & Axelrod, Publ. Carnegie Inst. Wash. 617: 136, pi. 11, fig. 13. 1959 (see synonymy). Abies chaneyi Mason, Publ. Carnegie Inst. Wash. 346: 149, pi. 4, figs. J, 6 (counterparts of twig; not winged seed which remains A. chaneyi Mason). Chaney & Axelrod, Publ. Car- negie Inst. Wash. 617: 137, pi. 11, figs. 1—2 only (fig. 3 remains A. chaneyi Mason). 1959. Axelrod, Univ. Calif. Publ. Ceol. Sci. 51: 141. 1964 (in part). Torreya bonseri (Knowlton) LaMotte, Publ. Carnegie Inst. Wash. 455: 108, pi 3, fig. 9. 1936. The above-cited specimens of Ahies chaneyi are similar to the leafy twigs and needles of Cephalotaxus honseri, a species abundantly represented in the Mollala flora of western Oregon, and known also from the Spokane (Latah) and Neroly floras of Washington and California, respectively (Chaney & Axelrod, 1959: 136). The needles are not Ahies because they do not have the typical rounded bases, nor are the petioles twisted prominently as in A. hracteata. Furthermore, the leafy twigs do not reveal the rounded leaf scars that are diagnostic of Ahies. However, the longitudinal ridging on the fossil twigs is like that on the twigs of the living Cephalotaxus fortunei of central China, which bear long, sharply acuminate needles like the fossils, and they often are curved in a falcate manner much like the fossil foliage. Occurrence: Mascall, Ore. hypotype nos. 134 arid 136 (counter parts), homeotype no. 2829 (leafy branch); Stinking Water, Ore.: hypotypes nos. 2830, 2831, homeotypes nos. 2832, 1976] AXELROD— SANTA LUCIA FIR ECOSYSTEM 31 2833; Beulah, Ore.: homeotypes nos. 8573-8579 (needles only); 49-Camp, Nev.: hypotype no. 777. Summarizing, Abies scherrii from the Middle and Late Miocene of western Nevada is similar to the living A. bracteata. Reexamination of the fossil A. chaneyi Mason and A. longirostris Knowlton, previously considered related to A. bracteata, shows that they are not allied to it. They seem related to one another, and may be extinct members of an alliance of present Asian distribution, of which A. delavaiji of southern China is a surviving relict. Composition As now known, Abies scherrii occurs in the Purple Mountain and Middlegate floras of western Nevada. Among its associates that are common to both floras are the species listed in Table 1. As might be expected, each flora has species that are not now known from the other site. Species in the Purple Mountain flora that are not recorded at Middlegate are listed in Table 2. Table 1. Species associated with Abies scherrii in both the Purple Mountain and Middlegate floras. Fossil Similar Living Fossil Similar Living Species Species Species Species Abies concoloroides A. concolor Litliocarpus klamathensis L. demiflorus Abies klamathensis A. shastensis Quercus hannibalii Q. chrysolepis Picea sonomensis P. breweriana Mahonia simplex M. japonica; M. Picea magna P. polita lomariifolia Pinus quini folia P. monticola Malumia reticulata M. pinnata-insularis Pseudotsuga sonomensis P. menziesii Amelanchier alvordensis A. alnifolia Chamaecyparis sierrae C. lawsoniana Cercocarpus antiquus C. betuloides Sequoiadendron chaneyi S. giganteum Cercocarpus holmesii C. paucidentatus Populus eotremidoides P. trichocarjm Heteromeles sonomensis H. arbutifolia Pop id lis payettensis P. angustifolia Ltjonothamnus parvifolia L. extinct Populus pliotremuloides P. tremuloides Sorbus sp. nov. S. aucuparia Salix knowltonii S. lemmonii Acer columbianum A. glabrum Salix sp. nov. S. melanopsis Acer middlegateii A. saccharinum Salix wildcatensis S. lasiolepis Acer oregonianum A. macrophyllum Betula lacusiris B. papyrifera The taxa that have been found in the Middlegate but are not now known from Mount Table 2. Species associated with Abies scherrii in only the Purple Mountain flora Fossil Species Similar Living Species Salix sp. nov. S. nigra Castanopsis sonomensis C. chrysophylla H. glabrescens Holodiscus idahoensis Amorpha oklahomensis A. fruticosa Fossil Species Ceanothus leitchii Arbutus matthesii Leucothoe sp. nov. Similar Living Species C. velutinus Rtiamnus precalifornica R. californica A. menziesii L. davisiae 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Table 3. Species associated with Abies scherrii in only the Middlegate flora. Fossil Similar Living Fossil Similar Living Species Species Species Species Pinus florissanttt P. ponderosa Crataegus middlegateii C. chrysophyUa Salix owyheeana S. hookeriana Primus morganensis P. emarginata Salix hesperia S. lasiandra Acer minor A. negundo Abuts harneyana A. tent li folia Wins alvordensis R. glabra Betula vera B. lent a Ceanotlius precuneatus C. cuneatus Quercus ivislizenoides Q. wislizenii Styrax middle gat eii S. californica Mahonia sp. now M. nervosa Diospyros andersonae D. virginiana Hydrangea bcndii n> ii II. aspera A rbutus prexalapenMs A. arizonica Platanus paucidm it at a P. racemose Fraxinus millsiana F. anomala Plat anus dissect a P. extinct Fraxinus coulteri F. oregon a The differences in composition are attributable to two factors. First, the Mid- dlegate sample is larger (7,200 specimens) than that at Purple Mountain (1,200 specimens), so it would be expected to have more numerous species (50 vs. 35 taxa). Second, the somewhat greater age of the Middlegate (15-16 m.y. ) as com- pared with Purple Mountain flora (13 m.y.) accounts for the more numerous exotic taxa (Acer tyrrettii, Betula vera, Crataegus middle gat ei, Diospyros ander- sonae > Hydrangea bendirei, Platanus dissecta) in it. Nonetheless, it is noteworthy that the lists include many species which, in terms of their closest modern rela- tives, are associated frequently at the present time. Hence, the floristic differences are not as great as one might otherwise suppose. This becomes apparent if we compare the principal vegetation types that the modern species similar to the fossils represent, as well as other aspects of the paleoecology of these floras. Paleoecolocy VEGETATION Broadleaved Evergreen Forest. — Fossil Santa Lucia fir (Abies scherrii) occurs in floras in which broadleaved sclerophylls are commonly dominant. Quercus hannibalii, which is similar to the living Q. chrysolepis, makes up 85% of all specimens in the Middlegate flora, and it also dominates several of the 10 florules in the Purple Mountain area. Abies hracteata, which is similar to the fossil A. scherrii, is regularly associated with a dominant Quercus chrysolepis sclerophyll community in areas of its optimum development. Among its more common associates that make up broadleaved evergreen forest that interfingers with patches of conifer forest in the higher Santa Lucia Mountains are Arbutus men- ziesii, Lithocarpus densiflorus, and Quercus wislizenii, all with close equivalents in the western Nevada floras. Many of their associates have analogues in the Miocene floras that have fossil Santa Lucia fir, notably Acer negundo, 3 Acer macrophyllum* Ceanotlius cuneatus, Cercocarpus betuloides, Ileteromeles arbuti- folia, Platanus racemosa* Rhamnus californica, Salix lasiandra 3 and Salix mela- nopsis* Chiefly riparian. 1976] AXELROD— SANTA LUCIA FIR ECOSYSTEM 33 Other taxa in these floras have their nearest relatives in sclerophyll vegetation in areas of summer rainfall, from southern Utah and Arizona to western Texas and southward into Mexico, notably Acer grand identatum, Arbutus arizonica, Cercocarpus paucidentatus, Fraxinus anomala, Fraxinus velutina, 3 Populus an- gustifolia 3 (rare in So. Calif.), and Robinia neomexicana. Mixed Conifer Forest. — The sclerophyll-dominated slopes and flats near the Miocene basins of plant accumulation were bordered by a mixed conifer forest dominated by Sequoiadendron, with associates of Abies (concolor, magnified- shastensis) * Picea (breweriana, polita), Pinus (ponderosa), Pseudotsuga (rnen- ziesii), and Chamaecyparis (lawsoniana) among the common conifers. Their associates included fossil species of Acer, Alnus, Amelanchier, Crataegus, Fraxinus, Holodiscus, Mahonia, Platanus, Primus, Populus, Rosa, Salix, Sorbus, and others, as listed above under Composition. In addition, each flora also has a few forest taxa that indicate summer rain, notably Acer (grandidentatum, saccharinum) , Betula (lenta, papyrifera), Diospyros (virginiana) , Hydrangea (aspera) and Platanus (cf. occidentalis) . More numerous members of this alliance are in floras of similar age to the north in Oregon, or to the west in California. Their poorer representation in the Nevada floras is chiefly due to nearby terrain that produced local rainshadows over these basins. The Miocene occurrence of Abies scherrii with dominant evergreen sclero- phyllous vegetation, and with mixed conifer forest on bordering nearby slopes, Mount this chiefly in sites where it is protected from fire by rocky bluffs and cliffs. In such sites it is well removed from areas where it might contribute to a fossil record. The question may thus be posed: Is the rarity of A. scherrii in the fossil record to be attributed to its preference for well drained, rocky sites scattered in the ecotone between mixed conifer forest and broadleaved sclerophyll vegetation? There are important differences betwen the Miocene and the modern com- munities, the Mioc reconstruct climate Santa Lucia fir ranges through the canyon live oak-tan oak-madrone sclero- phyll vegetation, reaching up into the mixed conifer forest in the Santa Lucia Mountains at levels from 4,000 to 5,000 feet. This is shown by its occurrence with Pinus lambertiana, P. ponderosa, and P. coulteri at or near Cone Peak, with Calocedrus decurrens and Pinus lambertiana at or near Junipero Serra Peak, and with Calocedrus decurrens, Pinus ponderosa, and P. coulteri at South Ventana Cone. With certain qualifications noted below, thermal conditions in the ecotone from conifer forest to broadleaved sclerophyll forest may be considered to ap- proximate Miocene temperatures. Temperatures can be determined from records at stations in the Sierra Nevada where mixed conifer forest interfingers with Species in parenthesis are living plants that seem most nearly related to the fossils. 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN LVol. 63 70 10 20 C T T T T T PURPLE MTN. FLORA Estimated mean monthly temp 5 Aiue4> Iviacteata E. slope, Cone Pk Belknap Cr. & So. Fork Groves O^O^ Q5 5000 / Hastings Reservation 1750' I I l I I I l l I I i San Antonio Mission 1060' vie. Shasta Spr. Lakeshore Porterville Lemon Cove WM-t-CM 2 and A = WM-CM i i i i i i i i i i i i i 1 i i i i i i i i i 10 5 20 10 20 30 40 50 A = MEAN RANGE OF TEMPERATURE °F Figure 21. Estimate of thermal conditions under which fossil Santa Lucia fir lived during Late Miocene time. The data are based on present thermal conditions, modified by estimates of Miocene climate (see text). broadleaved sclerophyll vegetation, and also by calculating temperatures for specific sites from stations in the adjacent lowlands to the west. Temperatures in the area of Santa Lucia fir can be estimated from meterological records at stations in the nearby region to the east and north. iitur with Sequotadendron, Belknap Creek (4,800 ft.) and South Fork Grove (5,000 ft.) were calculated from temperatures at Lemon Cove (alt. 513 ft., T = 63.9°, A 35.7°F) and Porterville (alt. 393 ft, T = 63.2°, A = 34.6°F), using a lapse rate 1976] AXELROD— SANTA LUCIA FIR ECOSYSTEM 35 of 2.5 °F per 1,000 feet. 5 Figure 21 shows that they live under a mean temperature of ~ 53°F. A similar result is suggested by data used to estimate conditions where Santa Lucia fir overlaps members of the Sierra mixed conifer forest at altitudes between 4,000 and 5,000 feet. There the stations selected are Hastings Reserva- tion (alt. 1,750 feet, T = 59.1°, A = 21.5°F) and San Antonio Mission (alt. 1,060 ft, T = 60.2°, A = 28.4°F), utilizing a lapse rate of 2.5°F per 1,000 feet. The mean January and July temperatures estimated for the east slope of Cone Peak at 4,200 feet (see Fig. 21) are similar to those recorded there by Talley (1974) for those months in 1971. Significantly, this was a year in which temperatures at lowland stations in Salinas Valley to the east were close to the 30-year norm. Figure 21 also shows estimated temperature for the Shasta Springs area, where Chamaecyparis is in the ecotone between mixed conifer forest and broadleaved sclerophyll vegetation. The temperature was calculated from the records at Mt. Shasta City (alt 3,544 ft., T = 49.9°, A = 34.7°F) and Lakeshore (alt. 1,075 ft., T = 58.9°, A = 34.8°F). The range of temperature (A) in the Miocene was more nearly like that now in the forest-sclerophyll ecotone in the Coast Ranges than in the Sierra Nevada. This seems likely inasmuch as the fossil floras with Abies scherrii (cf. bracteata) all have a few taxa that live chiefly on the coastward slopes, notably Picea (cf. breweriana) , Chamaecyparis (cf. lawsoniana), and Castanopsis (cf. chrysophylla) in northwestern California, or are found in the coastal strip farther south, as exemplified by Cercocarpus (cf. blancheae), Lyonothamnus and Mahonia (cf. insularis). Furthermore, with some summer rainfall over the region — as compared with clear, cloudless summer skies today — the cloud deck would reduce the high summer temperatures. Also, Sequoia (sempervirens) was then living on the coastward slope of the Sierra Nevada (Condit, 1944) 90 miles west of the Purple Mountain flora. Its occurrence there together with Chamaecijparis, Lithocarpus, Persea, and Umbellularia clearly implies a low range of temperature. By infer- ence, comparable conditions must have extended inland since the northern Sierra was then only a low ridge without significant relief (Durrell, 1966: 192-195; Axelrod, 1956). Since the fossil floras with Abies (bracteata) represent an en- vironment like that now in the ecotone between broadleaved sclerophyll and mixed conifer forest, thermal conditions there were approximately as shown in Fig. 21: mean annual temperature, 53°F; mean range of temperature, 26°F; mean July temperature, 66° F; and mean January temperature 40°F. From these data it is estimated that the Miocene ecotone had an effective temperature (ET) or warmth (W) of 55.7°F, or 158 days with a mean temperature above 55.7°F. "'Temperature normally decreases as altitude increases. In air free from the surface, the standard lapse rate is -3.6°F/1,0()() ft. But close to the ground, heat is supplied to the overlying air during the day, so air temperatures measured in the instrument shelter record conditions warmer than those in free air. This "ground effect" thus reduced the lapse rate, and gives a "terrestrial' lapse rate that approximates 3.0°F/1,0()0 ft. under normal conditions (H. P. Bailey, written communication, Jan., 1975). The scrub form, var. minor, occurs to the south Coast Ranges. The tree form var. chryso- phylla has a relict occurrence in the Sierra Nevada near Pino Grande, Placer County, in the lower part of the mixed conifer forest where it is associated with Quercus chrysolepsis , Litho- carpiis densiflorus, and Arbutus menziesii. 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 The temperateness index was M 57, and about 4% of the hours of the year had frost (see Bailey, 1960, 1964). It is emphasized that Miocene thermal conditions differed from present ones in two important ways. First, winters must have been more mild because ice caps were not yet in existence and hence cold and freezing conditions (like the winter ice storm of 1974) were not yet present. Second, in view of warmer Miocene seas, summers were moist and warm, high evaporation such as exists today could not have been present, and hence summer temperatures were more moderate than those of today. The temperatures estimated above must therefore be extreme for the Late Miocene. Clearly, temperatures were more equable than those now in the ecotone from mixed conifer forest to broadleaved sclerophyll vegetation at .000-5.000 lives today. Precipitation in the mixed conifer-evergreen sclerophyll forest ecotone near 4,000 feet in the Santa Lucia Mountains is about 35-40 inches, distributed chiefly in the winter season as rain, and occasionally as light snow. As noted earlier, there was some summer rainfall over western Nevada during the Late Miocene, prob- ably amounting to several inches for the summer season. The minimum rainfall required to support the flora can be estimated also from the temperatures sug- gested above for the Miocene ecotone. Using a mean temperature (T) of 53°F and a range of temperature (A) of 26°F, the Water Need (N), which is based on the exponential relation between temperature and moisture (see Bailey, 1958), calls for a minimum precipitation of about 35 inches, including that of the summer season. ALTITUDE The general altitude of the lake basins in western Nevada that were situated in the ecotone between mixed conifer forest and broadleaved sclerophyll vegeta- tion was estimated earlier to be near 2,000 to 2,500 feet. This was based on the general relations of vegetation and climate, and on evidence that a major rain- shadow like that of the present had not yet developed (Axelrod, 1956). Altitude can also be estimated by comparing thermal conditions in the Mio- cene of Nevada with that at sea level to the west, and interpolating altitude from the difference in mean temperature (Axelrod, 1965, 1968). As estimated above, thermal conditions in the forest-sclerophyll ecotone were approximately T 53° F and A 26°F. During the Late Miocene the coastal strip had a mean temperature near 60° to 61 °F as judged from the slightly younger Neroly flora of western California that resembles vegetation in coastal Virginia (Condit, 1938). Tem- peratures were somewhat warmer during the Middle Miocene (~ 15 m.y. ) as judged from the Temblor flora near Coalinga (Rennie, 1972) which is situated 145 miles southeast of the Neroly flora. It shows relationship to vegetation in coastal North Carolina, where mean temperature is 62° to 63°F. Assuming a Late Miocene terrestrial lapse rate of 3.0°F per 1,000 feet (or 1° = 333 feet), a dif- ference in mean temperature of approximately 7°F-8°F between the Purple Moun- tain flora and the Miocene floras at sea level implies a minimum altitude of about 2,300-2,600 feet. Realizing that this is an estimate, it seems likely that the Piuple 1976] AXELROD— SANTA LUCIA FIR ECOSYSTEM 37 Mountain flora had an altitude near 2,500 feet. This agrees closely with earlier estimates for the Fallon and Chloropagus floras of the nearby region, estimates based on very different lines of reasoning (Axelrod, 1956). Post-Miocene Chances The mixed conifer forest that inhabited cooler, moister slopes and valleys bor- dering the sclerophyll-dominated basins was richer than the modern descendant vegetation. It included conifers related to those that are now restricted in area, notably Pieea breweriana and Chamaecyparis laivsoniana that are confined to northwestern California, Sequoiadendron giganteum which lives in the central and southern Sierra Nevada, as well as Abies magnified and Pinus montieola that now occur in subalpine sites in the Sierra Nevada well removed from Santa Lucia fir today. Similar relations are displayed by the broadleaved sclerophyll vegeta- tion, for the Miocene community included species similar to those now in coastal southern California, notably species of Cercocarpus, Lyonothamnus, and Mahonia. Furthermore, both vegetation zones had a few species related to those now in areas with summer rain. The emergence of modern communities of lower diversity and more restricted area is due to the gradual development of new moisture-thermal conditions during the Pliocene which culminated in the appearance of summer-dry mediterranean climate in the middle and late Quaternary. Increasing summer drought has resulted in progressively greater water stress during the critical period of seed- ling germination, growth, and establishment (Axelrod, 1976). Hence, taxa that were unable to adapt to these new conditions in the lower part of the mixed conifer forest near broadleaved evergreen sclerophyll vegetation were gradually confined to areas in which they could reproduce successfully. The regular occur- rence in the Miocene of such "subalpine" species as Abies (magnified), Pinus (montieola), Tsuga (mertensiana) and Popnlus (tremuloides) with mixed conifer forest taxa as Abies (coneolor), Pinus (ponderosa), Caloeedrus (deeurrens) , and Sequoiadendron (giganteum), and with broadleaved sclerophylls as Que reus (ehrysolepis, tcislizenii), Castanopsis (chrysophylla), Lithocarpus (densi floras) , Arbutus (menziesii), in the same fossil flora, is symptomatic of the nature of the post-Miocene changes in community composition that were due to changing cli- mate, and chiefly to increased water deficit in the summer season. Its effect can be summarized in terms of the modifications that appear to account for the dis- tributions of taxa whose Miocene relatives were associated with fossil Santa Lucia fir, and which now define the segregate communities of lower diversity that are confined to more local areas. Sierra redwood (Sequoiadendron), which is restricted to the central and southern Sierra Nevada, probably entered the range following the Late Miocene as rainfall decreased and as conditions became sunnier there. Sequoiadendron is now known from the Middle Pliocene Mount Reba flora (7 m.y.), Alpine County. The site is at wind timberline, above a subalpine forest of Abies magnified, Pinus montieola, and Tsuga mertensiarm. The Mt. Reba flora is dominated by broad- leaved sclerophyllous evergreens, as shown by the abundance of specimens of Que reus (ehrysolepis), Lithocarpus (densiflora), and Cupressus (cashmeriana) . 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Pseudotsuga (menziesii) is also common, and probably formed rich stands on nearby slopes much as it does today at levels near 2,000-2,500 feet in the lower foothills of the range. A few scraps of other conifers are present, notably Abies (concolor) and Sequoiadendron. The community shifted to lower levels as colder climates developed as the range was elevated later in the Pliocene and during the Quaternary. Reasons for the present absence of Sequoiadendron in the Sierra Nevada north of the American River drainage (Placer County) are not clear. However, the region receives much higher rainfall, and a number of taxa that are typical of forests in the Coast Ranges occur in the northern Sierra Nevada. This gives to the mixed conifer forest of that area a denser, richer understory of shrubs and small trees than in the Sierra farther south. As a result, relatively less light falls on the forest floor, and conditions are therefore unfavorable for taxa that are light demanding, especially in the seedling stage. That Sierra redwood seedlings thrive best under conditions of ample light is apparent from the heavy reproduc- tion that occurs about the margin of moist meadows in the southern Sierra today, as at Balch Park in Tulare County, east of Exeter. This agrees with the experi- mental studies by Stark (1968a) who noted "growth in full sunlight was superior to that in any shady forest" (p. 91) and that "healthy . . . seedlings grow best in full sunlight" p. 92). Stark (1968b: 276) also noted that the failure of trees to spread and expand their present range is the result of fire-suppression over the past decades, with the accumulation of a deeper-than-normal litter, as well as an increase in growth of understory shrubs and trees, and hence an increase also of shade within the forest. One other point is to be noted in terms of the need of Sequoiadendron seed- lings for ample light. Sierra redwood occurs at numerous sites in the Miocene of Nevada and also at a locality (Trapper Creek) in southeastern Idaho. How- ever, it has not been recorded in any of the Miocene floras that are scattered widely in Oregon, Washington, and western Idaho. The floras of that region are richer in taxa than in the area to the south chiefly because of the higher precipitation there. Mesic conifers (Cephalotaxus, Ginkgo, Ghjptostrobus, Meta- sequoia, Sequoia, Taxodium) are present in most of these floras, and deciduous hardwoods are especially abundant together with some associated broadleaved evergreens. But in the Miocene floras to the south, mesic conifers are rare to absent, deciduous hardwoods have a poorer representation, and sclerophyllous taxa increase in diversity and abundance. These relations imply climate was drier and sunnier to the south, which is consistent with the composition of the Miocene floras in the Mohave region where live oak woodland and thorn scrub are wholly dominant (Axelrod, 1958). The general distribution of Sequoiadendron during the Miocene is therefore consistent with conditions that it seems to require for best reproduction — ample light and sufficient moisture during the critical stage of seedling establishment. Taxa that are often considered "subalpine indicators," notably Abies magnifica, Pieea breweriana, Pinus monticola, Tsuga mertensiana, Popuhis tremuloides, and others, have close relatives in the Miocene floras of Nevada where they occur with sclerophyllous taxa and fossil Santa Lucia fir. Since geologic evidence shows 1976] AXELROD— SANTA LUCIA FIR ECOSYSTEM 39 that these plants lived in terrains of low relief, and since "subalpine" species are sufficiently abundant numerically to form codominants of some of the fossil floras, they must have lived near at hand (Axelrod, 1976). This means they were regular members of the mixed conifer forest. Such an occurrence is con- sistent with the presence of most of their living descendants in mixed conifer forest in areas to the north, where there is more rainfall in summer, a longer precipitation season, and a lower evaporation rate (Axelrod, 1976). The taxa that are now confined to subalpine forest in the Sierra evidently were restricted to that zone as dry summer, montane mediterranean-type climate spread during the later interglacial ages. Thus, the distinctness of the modern subalpine (or pure conifer) forest in the Sierra Nevada is due to the restriction of its taxa to this higher, cooler zone where the effect of high evaporation (= water deficit water stress) is less than in the mixed conifer forest at lower levels, a forest from which they are now largely excluded. The confinement of Abies bracteata to the Santa Lucia Mountains seems con- sistent with its mild climate and lower evaporation rate in summer as compared with the Sierra where many of its former associates occur. In this regard, the forested parts of the outer Coast Ranges have lower summer temperature and milder winters than comparable parts of the Sierra (Fig. 21). Thus, Santa Lucia fir may have been eliminated from the Sierra Nevada, where it probably occurred during the Pliocene, by the high evaporation rate in summer which was inimical to seedling establishment. A second factor was increasing winter cold with ac- companying snow and ice. That it would have had a disastrous effect on the trees at higher levels in the ecotone between mixed evergreen forest and mixed conifer forest may be inferred from the severe damage inflicted by the snow and ice storm on Santa Lucia fir during the winter of 1974, which resulted in numerous broken tops (with cones) and limbs (oral communication, S. Talley, 1974). Similar damage was inflicted on its broadleaved evergreen associates, notably Arbutus menziesii, Lithocarpus densiflorus, Quercus chrysolepis, and (). wislizenii. On this basis we may infer that conditions probably were too severe for A. bracteata in the Sierra Nevada during the glacial ages. In this regard, its present absence from the Coast Ranges farther north may reflect the more severe winters there, for the frequency of snow increases northward and the mean January temperature rapidly falls below 40°F, which appears to be near the minimum for the species today (Fig. 21). The present absence of Santa Lucia fir farther south in the Coast Ranges may reflect a recent restriction in range. It may possibly have been eliminated there by the xerothermic periods of the later Quaternary (Axelrod, 1966: 42-55). This seems consistent with the paucity of conifer forest taxa in the isolated areas where forest now occurs in the central Coast Ranges (Axelrod, 1976). Altitudes in these areas, whether in the Santa Cruz or Santa Lucia mountains or the interior Diablo and La Panza ranges, are relatively low and the forest has only one or two species. However, in the higher north Coast Ranges (north of Clear Lake), the Sierra Nevada, the Transverse Ranges and Peninsular Ranges of southern California, the forest has mixed stands of several conifers. It seems 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 likely that a warm dry xerothermic period would have eliminated many forest taxa from the central Coast Ranges simply because terrain was not sufficiently high to provide cool, moist sites for them, and hence they disappeared because drought stress militated against successful reproduction. That richer, more diverse forests were in the region earlier is shown by the occurrence of Calocedrus decurrens and Finns lambertiana in the Santa Cruz Mountains during the Plio-Pleistocene transition (Dorf, 1930: 18). This may also be inferred from the occurrence of a rich mixed conifer forest on the valley floor near San Jacinto in southern California ( Axelrod, 1966), a region now semi- desert. The occurrence of Sequoia sempervirens and Pseudotsuga menziesii along the Santa Barbara coast during the late Quaternary (Axelrod, 1967: 295-296), and the present relict stand of mixed evergreen forest (Arbutus menziesii, Quer- cus chrysolepsis, Lithocarpus densiflorus) in the nearby summit section of the Santa Ynez Range at San Marcos Pass also provide hints of the nature of the forests that probably inhabited the central Coast Ranges during the cooler, moister phases of the Quaternary. Literature Cited Axelrod, D. I. 1956. Mio-Pliocene floras from west-central Nevada. Univ. Calif. Publ. Geol. Sci. 33: 1-316. . 1958. Evolution of the Madro-Tertiary Geoflora. Bot. Rev. (Lancaster) 24: 433- 509. — . 1965. A method of determining the altitudes of Tertiary floras. Paleobotanist 14: 144-171. — . 1966. The Pleistocene Soboba flora of southern California. Univ. Calif. Publ. Geol. Sci. 60: 1-108. . 1967. Geologic history of the Californian insular flora. Pp. 267-315, in R. N. Phil- brick (editor), Proceedings of the Symposium on the Biology of the California Islands. Santa Barbara Bot. Garden, Santa Barbara, Calif. . 1968. Tertiary floras and topographic history of the Snake River Basin, Idaho. Bull. Geol. Soc. Amer. 79: 713-734. . 1976. History of the conifer forests, California and Nevada. Univ. Calif. Publ. Bot. 60: 1-62. Bailey, H. P. 1958. A simple moisture index based upon a primary law of evaporation. Geogr. Ann. Svenska Siillsk. Antropol. 40: 196-215. . 1960. A method of determining the warmth and temperateness of climate. Geogr. Ann. Svenska Sallsk. Antropol. 42: 1-16. . 1964. Toward a unified concept of the temperate climate. Geogr. Rev. (New York) 54: 516-545. Bonham, H. F. 1969. Geology and mineral deposits of Washoe and Storey Counties, Nevada. Bull. Nevada Bur. Mines 70: 1-140. Chaney, R. W. & D. I. Axelrod. 1959. Miocene floras of the Columbia Plateau. Publ. Car- negie Inst. Wash. 516: 1-237. Condit, C. 1938. The San Pablo flora of west central California. Publ. Carnegie Inst. Wash. 476: 217-268. . 1944. The Remington Hill flora (California). Publ. Carnegie Inst. Wash. 553:21-55. Dorf, E. 1930. Pliocene floras of California. Publ. Carnegie Inst. Wash. 412:1-108. Durrell, C. 1966. Tertiary and Quaternary geology of the northern Sierra Nevada. Bull. Calif. Div. Mines Geol. 190: 185-197. Griffin, J. R. & W. B. Critchfield. 1972. The distribution of forest trees in California. U.S.D.A. Forest Serv. Res. Pap. PSW 82. Knowlton, F. H. 1923. Fossil plants from the Tertiary lake-beds of south-central Colorado. Profess. Pap. U.S. Geol. Surv. 131: 183-197. Liu, Tang-Shui. 1971. A monograph of the genus Abies. Dept. Forest, Coll. Agric, Nat. Taiwan Univ., Taipei, Taiwan. 608 pp. 1976] AXELROD— SANTA LUCIA FIR ECOSYSTEM £\ MacGinitie, H. D. 1953. Fossil plants of the Florissant beds, Colorado. Publ. Carnegie Inst. Wash. 599: 1-188. Mason, II. L. 1927. Fossil records of some west American conifers. Publ. Carnegie Inst. Wash. 346: 139-158. Rennie, K. M. 1972. The Miocene Temblor flora of west central California. MS thesis, Univ. California, Davis. 106 pp. Rose, R. L. 1969. Geology of parts of the Wadsworth and Churchill Butte quadrangles, Nevada. Bull. Nevada Bur. Mines 71: 7-27. Stark, N. 1968a. The environmental tolerance of the seedling stage of Sequoiadendron gigantetim. Amer. Midi. Naturalist 80: 84-95. . 1968b. Seed ecology of Sequoiadendron giganteum. Madrono 19: 267-277. Talley, S. N. 1974. The ecology of Santa Lucia fir {Abies hracteata), a narrow endemic of California. PhD thesis, Duke Univ., North Carolina. 208 pp. STUDIES IN BIGNONIACEAE 18: NOTES ON S. MOORE'S MATO GROSSO BIGNONIACEAE' Alwyn II. Gentry 2 Abstract The 16 species of Bignoniaceae described by Moore from his own and Robert's Mato Grosso collections are identified. Three new combinations based on Moore's species are made. In preparation for the treatment of Bignoniaceae for Flora Neotropica I am attempting to identify the numerous, unaccounted-for species described in Big- nonia prior to 1900. Spencer Moore's Mato Grosso Bignoniaceae are among the most important of these. In 1895 Moore (1895) described 14 new species of Bignoniaceae collected during the Percy Sladen Mato Grosso Expedition of 1891- 1892. Later (Moore 1904, 1907) lie described two additional species of Big- noniaceae from the same area. Moore's species, described mostly in the genus Bignonia, have never been reinterpreted nor identified with known species of Bignoniaceae. The holotypes of Moore's collections are maintained in the her- barium of the British Museum of Natural History (BM); there are also partial sets at NY and MO. Through the kindness of the curator of the British Museum's Botany Department, I have been able to examine the holotypes of these plants. This paper identifies them to genus and species and proposes the three necessary new combinations based on Moore's names. Moore's other 13 species are reduced to synonymy. The nomenclatural significance of Moore's species is due to the fact that they were published just prior to Bureau & Schumann's (1896-1897) treatment of Bignoniaceae for Flora Brasiliensis. Published too late for inclusion in Flora Brasiliensis, these names have priority over those published in that work and subsequently. A few of Moore's plants were identified in the Flora Brasiliensis on the basis of the distributed specimens, however. The following 16 species of Bignoniaceae were described by Moore. 1. Bignonia rubescens S. Moore is Arrabidaea chica (II. & B.) Verl. (based on B. chica H.&B., PI. Aeq. 1: 107, tab. 31. 1808.) as noted by Bureau & Schumann (as B. erubescens) . 2. Bignonia tomentella S. Moore is Arrabidaea pubescens (L.) A. Gentry (based on B. pubescens L., Sp. PL, ed. 2, 2: 870. 1763.) and falls into the synon- ymy of that species. 3. Bignonia grewioides S. Moore is Arrabidaea fagoides (Cham.) Bur. (based on B. fagoides Cham., Linnaea 7: 680. 1832.) and becomes a synonym of that species which is itself uncomfortably close to A. platyphylla DC. 4. Bignonia melioides S. Moore is Pleonotoma brittonii Rusby (Bull. Torrey Bot. Club 27: 72. 1900.) and Moore's name is older. The new combination Pleono- 1 Supported by National Science Foundation grant GB-40103. 2 Missouri Botanical Garden, 2315 Tower Grove Avenue, St. Louis, Missouri 63110. Ann. Missouri Bot. Gard. 63: 42-45. 1976. 1976] GENTRY MOOR KS HIGNONIACEAE 43 toma melioides (S. Moore) A. Gentry (based on B. melioides S. Moore, Trans. Linn. Soc. London, Bot. 4: 414. 1895.) is necessary. 5. Bignonia caudigera S. Moore is Arrabidaea coleocalyx Bur. & K. Schum. (Fl. Bras. 8(2): 35. 1896.) and Moore's name is older. The new combination Arrabidaea caudigera (S. Moore) A. Gentry (based on B. caudigera S. Moore, Trans. Linn. Soc. London, Bot. 4: 415. 1895.) is necessary for this well-known species. 6. Bignonia modesta S. Moore is a species of Stizophyllum and Sandwith has annotated the type as S. riparium (H.B.K.) Sandw. sensu lato. Moore's plant differs from other specimens of S. riparium seen by me in its smaller serrulate leaflets. It differs from S. perforatum (Cham.) Miers in less noticeable pubes- cence, smaller serrulate leaflets, and especially the smaller uninflated calyx. The pink flower color agrees with S. perforatum or S. inaequilaterum Bur. & K. Schum. but not S. riparium sensu stricto. Species limits in Stizophyllum are hazy at best and only additional collections from Mato Grosso can tell whether B. modesta should be regarded as specifically distinct. For the present it may be tentatively included under S. riparium. 7. Macfadyena riparia S. Moore is a form of Phryganocydia corymbosa (Vent.) Bur. ex K. Schum. (based on Spathodea corymbosa Vent., Choix tab. 40. 1807.) and becomes a synonym of that species. The predominantly simple leaves of Moore's plant are insufficient grounds for species segregation. 8. Macfadyena bipinnata S. Moore is an otherwise undescribed species of mora Macfady guisl leaves with relatively large leaflets and lack of foliaceous pseudosti pules. A second collection of this species is Prance et al. 18906 from the Chapada dos Guimaraes, cerrado behind Colegio de Buriti, Mato Grosso. It differs from the type in simply pinnate leaves but is otherwise a good match. Memora axillaris Bur. & K. Schum. (a simply pinnate leaved species with foliaceous pseudostipules M M synonym should the two be united. 9. Macfadyena pubescens S. Moore is Macfadyena mollis (Sonder) Seem, (based on Spathodea mollis Sonder, Linnaea 22: 561. 1849.) which is itself synonymous with Macfadyena hispida (DC.) Seem, (based on Spathodea hispida DC, Prodr. 9: 205. 1845.). Seemann (1863) followed by Fabris (1965) separated M In either case Moore 10. Adenocalymma croceum S. Moore is Adenocalymma bracteolatum DC. (Prodr. 9: 200. 1845.) and becomes a synonym of that species. The only known fruiting collection of A. bracteolatum was collected by Roberts and identified by Moore with his A. croceum. This collection contains two immature capsules which 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 are oblong, obtuse at base and apex, 13-14 cm long, 2-2.2 cm wide, minutely lepidote, otherwise glabrous, drying dark with numerous pale lenticels, and some- what flattened in drying but probably subterete when fresh. The unwinged fruit of A. bracteolatum proves that the more widespread and closely related Adeno- calymma with winged fruits is not synonymous with this species and must be known as A. purpurascens Rushy (see Gentry, 1976). The ovary of Moore's col- lection agrees with that of the type of A. bracteolatum in being linear-oblong without lateral ridges, unlike the 4-ridged ovary of A. purpurascens. 11. Anemopaegma brevipes S. Moore is closely related to the earlier A. flavum Morong (Ann. New York Acad. Sci. 7: 188. 1893.) on account of its glabrous corolla tube and foliaceous pseudostipules. It appears to be distinct by its much denser pubescence, especially on the lower leaflet surface. I would tentatively assign a second collection to A. brevipes. This is Pirres & Leite 14692 (MO) from Roraima Territory. Its only noticeable difference is a more strongly bracteate less pubescent inflorescence, although its geographic disjunction is rather striking. While Anemopaegma chrysoleucum (H. B. K.) Sandw., A. flavum, and A. brevipes are clearly related and form a series subdivided mostly by increasing pubescence, they seem to be specifically distinct. The New York "isotype" of A. brevipes is actually A. flavum, however. Anemopaegma bifarium Bur. & K. Schum. (Fl. Bras. 8(2) : 124. 1896.) is based on Moore's collection, but Moore's name is older. 12. Anemopaegma decorum S. Moore is Clytostoma decorum Bur. & K. Schum. (Fl. Bras. 8(2): 1896.). Although Bureau & Schumann had not seen Moore's description, they saw a duplicate of his collection and redescribed the species in its correct genus. Moore's description is older than Bureau & Schumann's but C decorum Bur. & K. Schum. blocks a combination in Clytostoma based on A. decorum S. Moore. 13. Anemopaegma sylvestre S. Moore is Anemopaegma flavum Morong (Ann. New York Acad. Sci. 7: 188. 1893.) and becomes a synonym of the latter. 14. Tabebuia chapadensis S. Moore is Arrabidaea corallina (Jacq. ) Sandw. (based on Bignonia corallina Jacq., Fragm. Bot. 37, tab. 42, fig. 1. 1800-1809.) and has already been placed in the synonymy of that species (Gentry, 1973). 15. Cremastus sanctae-annae S. Moore (1904) is Arrabidaea sceptrum (Cham.) Sandw. (based on Bignonia sceptrum Cham., Linnaea 7: 710. 1832.). Sandwith (1968) suggested that C. sanctae-annae was probably a large-calyx form of A. pulchra (Cham.) Sandw., noting especially the agreement of its open pyramidal inflorescence with that of A. pulchra. I am hard pressed to distinguish A. pulchra from A. sceptrum; certainly the latter s inflorescence shows every kind of grada- tion from open to condensed. If A. pulchra is to be separated from A. sceptrum, its smaller calyx is the key character and the large calyx of C. sanctae-annae clearly allies it with the latter. 16. Jacaranda robertii S. Moore (1907) proves to be /. decurrens Cham, con- trary to my earlier (Gentry, 1974) interpretation. The BM holotype of /. robertii 5) is completely different from the MO "isotype" with the same col- 1976] GENTRY — MOORE'S BIGNONIACEAE 45 lection number. The holotype is clearly /. decurrens. The "isotype" is an un- described species mixed with corollas of /. decurrens. The numerous discrepancies I previously noted between the MO specimen of Roberts 675 and Moore's descrip- tion are thus explained: except for the corollas we were looking at two quite unrelated plants. The MO specimen of Roberts 675 has only vegetative parts, calyces, and very immature fruits of the new species, inadequate for its descrip- tion at the present time. Literature Cited Bureau, E. & K. Schumann. 1896-1897. Bignoniaceae. In C. F. P. Martius (editor), Flora Brasiliensis 8 (2): 1-452. Fabris, II. A. 1965. Bignoniaceae in Flora Argentina. Revista Mus. La Plata, Secc. Bot. 9: 273-419. Gentry, A. II. 1973. Bignoniaceae. In R. E. Woodson, Jr. & R. W. Schery. Flora of Pan- ama. Ann. Missouri Bot. Gard. 60: 781-977. 1974. Studies in Bignoniaceae 12: New or noteworthy species of South American Bignoniaceae. Ann. Missouri Bot. Gard. 61: 872-885. — . 1976. Studies in Bignoniaceae 19: Generic mergers and new species of South American Bignoniaceae. 46-80 Moore, S. 1895. The phanerogamic botany of the Mato Grosso Expedition 1891-92. Trans. Linn. Soe. London, Bot. 4: 265-516. . 1904. Mons. A. Robert's Mato Grosso Plants— I. J. Bot. 42: 106-107. . 1907. Note on some South American plants. J. Bot. 45: 404-406. Sandwith, N. Y. 1968. Notes on Bignoniaceae XXIX: Arrabidaea in Martius's Flora Brasili- ensis and subsequently. Kew Bull. 22: 403-420. Seeman, B. 1863. Revision of the natural order Bignoniaceae. Spathodea. J. Bot. 1: 225- 228. r STUDIES IN BIGNONIACEAE 19: GENERIC MERGERS AND NEW SPECIES OF SOUTH AMERICAN BIGNONIACEAE 1 Alwyn H. Gentry 2 Ahstract Preparation of the treatment of Bignoniaceae for Flora tie Venezuela and field work in Venezuela, Colombia, Ecuador, and Brazil have turned up several novelties and some taxonomic problems. In this paper the Atlenocalymma apurense, A. bract eolatum, and Amphilophium paniculatum complexes in Venezuela are reinterpreted necessitating one new variety, one new combination, and resurrection of two previously synonymized species. Anernopaegnia villosum, Arrabidaea prancei, Cuspidaria subincana, Ilaplolophium rodriguesii, Mansoa onohualcoides, Memora aspericarpa, M. tanacciicarpa, Tanaecium apiculatum, Tynnanthus villosus, Anenu)- paegma (datum, and A. patelliforme are described as new. Seven generic mergers necessitating a total of nine new combinations are proposed: Onohualcoa to Mansoa, Pseutlopaegma to Anernopaegnia, Sanhilaria to Paragonia, Nest or ia and Kuhlmannia to Pleonotoma, Roseodendron to Tabebuia, and Xerotecoma to Gotlmania. An eighth generic merger — Distictella with Dis- tictis — is rejected, but a new combination is needed for one species which is transferred to Distictis, Adenocalymma apurense (II.B.K.) Sandw. and relatives. As previously interpreted (Gentry, 1974a), A. apurense (including A. inun- daturn and A. calderonii) is a polymorphic widespread species ranging from Mexico to Para, Brazil. Additional collections of this complex indicate that it is even more polymorphic than previously supposed. The A. apurense complex con- sists of at least three distinct entities which deserve formal taxonomic recognition. Each of these three entities, virtually indistinguishable on the basis of vegetative and floral characteristics, has a distinctive fruit. A form with winged seeds and non-splitting capsule valves occurs from Mexico to northern Colombia and Venezuela and has been variously known as A. inundatum Mart, ex DC, A. hintonii Sandw. and A. calderonii (Standi.) Seib. The other two forms have wingless seeds and are restricted to South America. One, with non-splitting capsule valves and more coriaceous leaflets, has been known as A. inundatum var. surinamense. This form occurs from the upper Orinoco through the Guianas to Para, Brazil. I follow Sandwith in regarding it as not specifically distinct from the northern form with winged seeds. The third entity has wingless seeds and a shorter almost globose capsule with each valve splitting in half at maturity, i.e., effectively 4-valved. Although these three entities would not warrant any kind of taxonomic recognition on the basis of floral or vegetative characteristics, the 4-valved form can also be distinguished from the other two forms by its shorter (1-2 mm long) more square-ended ovary and is probably specifically distinct. The oldest name for any member of this complex is A. apurense, based on Bignonia apurense H. B. K. from the middle Orinoco region of Venezuela. I have previously (Gentry, 1974a) identified the type of A. apurense, which lacks fruit, 1 Support for this study was provided by NSF grant GB-40103. 2 Missouri Botanical Garden, 2315 Tower Grove Avenue, St. Louis, Missouri 03 110 Anx. Missouri But. Gaud. 63: 46-80. 1976] GENTRY— SOUTH AMERICAN BIGNONIACEAE 47 with the wing-seeded form of this complex, mainly because of its small subcoria- ceous leaflets. However, only specimens with wingless seeds have been subse- quently collected along the Orinoco so the A. apurense type should presumably be identified instead with one of the two wingless-seeded forms. The leaflets of the type collection are puberulous beneath, as are those of several collections of the form with subglobose capsules and splitting valves. Although this character is not constant [e.g., Gentry et ah 10694 (MO) has a 4-valved capsule and leaf- lets completely glabrous beneath], neither of the other two forms ever has leaf- lets puberulous below. Typical A. apurense is thus the small-capsuled, 4-valved form whose fruit has yet to be described. This capsule is subglobose-ellipsoid, rounded apically and basally, not at all compressed, 4-sulcate, splitting into 4 parts at maturity with each valve splitting down the midline, 2.4-6 cm long, each half-valve 1.3-2.5 cm wide, slightly lepidote when young, becoming densely papillate, drying tannish; seeds thick, wingless, 1.2-2 cm in diameter. In my opinion the difference between the fruit described above for A. apurense and that of the other two forms is adequate for specific segregation. The oldest available name for either (presumably) 2-valved form is A. inundatum. As long as winged versus wingless seeds are not considered adequate to justify specific segregation (see Gentry, 1973; Sandwith, 1955; Hunt, 1972), both the wing- seeded and wingless-seeded entities may be included in A. inundatum. The type of A. inundatum may represent the wingless-seeded form since no wing-seeded collections of this complex from Guiana or Amazonia have been seen. Adeno- calymma inundatum var. mrinamense , as defined by its wingless seeds, would thus be synonymous with typical A. inundatum, and a new infraspecific name for the northern entity with winged seeds would be necessary. Nevertheless, it seems advisable to retain the extant usage — A. inundatum var. mrinamense with wingless seeds, A. inundatum var. inundatum with winged seeds — until fruiting material from Amazonian Brazil is available. Adenocalymma purpurascens Rushy, Descr. S. Amer. PL 121. 1920. This species ranges from Bolivia to Venezuela and was treated by Sandwith (in the herbarium) as a synonym of A. bracetolatum DC. The fruit of neither A. purpurascens nor A. hracteolatum has been described. A recent collection [Gentry et al. 10672 (MO, VEN, duplicates to be distributed) from Bolivar, Venezuela] of flowering and fruiting material of Adenocalymma purpurascens taken from the same plant solves a longstanding puzzle and adds additional evidence of the inability of presence or absence of fruit wings to serve as an absolute generic criterion (cf. Gentry, 1973, 1974b, 1974c). The fruit of this species proves to be an oblong uncompressed capsule with the valves woody and prominently thick-winged along each margin. The fruit is thus tetragonal with raised angles in cross section. It is 10-19 cm long, 2.5-3.5 cm wide (including wings), 3-5 cm thick (including wings), drying light brown, the midrib not visible, glabrous, the surface slightly wrinkled-striate, the wrinkles making an acute angle with the axis of fruit. The seeds are bialate, 1.1-1.8 cm long, 3.3-6 cm wide, the wings more or less hyaline-membranaceous and indis- tinctly demarcated from the rather thick brown seed body. 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 This striking fruit is unique in Adenocalymma because of its conspicuous Cuspidaria-\ike wings. This prompted the late Dr. Sandwith to tentatively deter- mine a fruiting collection (Seibert 2012 from Madre de Dios, Pern) as Cuspidaria sp. nov. despite the collector s field notes that the plant was the same species as his number 2011, the latter an obvious Adenocalymma in flower. The identity of this interesting collection had otherwise remained an unsolved mystery. Actually, Seibert's flowering collection is not A. purpurascens but A. impressum (Rushy) Sandw. It is no wonder that the collector confused the two species: A. purpurascens and A. impressum are twins vegetatively. The main vegetative difference between the two species is a quantitative one — the pale-drying veins and veinlets of A. purpurascens dry a lighter color than those of A. impressum. In flower the smaller (6-9 by 5-7 mm) calyx with an irregularly toothed margin of A. purpurascens is quite distinct from the larger (11-14 by 7-12 mm) campanu- late calyx witli a truncate margin (remotely and minutely 5-denticulate) of A. impressum. The bracts and bracteoles of the latter species also tend to be slightly larger. In fruit these two plants are utterly distinctive, the subterete linear-oblong capsule of A. impressum showing no resemblance to the winged capsule of A. bracteolatum. Discovery of the winged fruit of A. purpurascens also creates a problem. The type of A. bracteolatum (d'Orbignij 758, P) from Mato Grosso lacks the 4-ridged tetragonal ovary otherwise associated with the wing-fruited species. Perhaps A. purpurascens is not the same as A. bracetolatum after all? This suspicion is con- firmed by two Mato Grosso collections at BM identified as A. croceum S. Moore. The flowering type of A. croceum proves to be the second collection of A. bracteo- latum (Gentry, 1976), likewise lacking a tetragonal ovary. The second collection of A. croceum is in fruit, comes from the same area as both of the flowering collec- tions, and agrees vegetatively with them. It is clearly A. bracteolatum. The fruit of A. bracteolatum is oblong and subterete with completely unwinged woody valves. Adenocalymma purpurascens with its strikingly winged fruits must be dis- tinct from A. bracteolatum. The only apparent characters, besides the ovary, for separating flowering A. bracteolatum from A. purpurascens are the tenuous ones of dark brown-drying (rather than black) inflorescence branches and a very slightly larger calyx. I cannot distinguish sterile material of these two species. The small leaves cited by Bureau & Schumann (1896-1897) as characteristic of A. l)racteolatum are present only in the inflorescence; even some sheets of the type collection include the larger leaves characteristic of both it and A. pur- purascens. Amphilophium paniculatum (L.) Il.B.K. in Venezuela. Three forms of this variable species, each characterized by its fruit, occur in Venezuela. The flowers of the three are indistinguishable. One of these forms occurs in the Sierra de Imataca region of Delta Amacuro and adjacent Bolivar State. These plants have very large (10-12 cm long, 4-5 cm wide, 1-3 cm thick), relatively flat capsules with muricate-rugose surfaces having the conspicuous projections widely scattered. Vegetatively these plants are distinguished by the yellowish, veiy finely and densely dendroid tomentose lower leaf surface. The 1976] GENTRY— SOUTH AMERICAN RIGNONIACEAE 4Q second form has a smooth granular-textured fruit surface, without prominent ridges or projections. This form has the leaves densely lepidote but otherwise glabrous beneath, except for tufts of long flexuous, mostly simple trichomes in the nerve axils and minute subappressed simple trichomes along the main nerves. It occurs along the dry inner base of the Cordillera de la Costa with fruiting collec- tions from Anzoategui, Miranda, Guarico, and Portuguesa States. This is the fruit type which occurs throughout the range of the species outside Venezuela. The third fruit form is similar to the preceding in size and convex valves, but the valves are sharply and closely reticulate-wrinkled, the wrinkles almost muricate. The leaflets associated with these fruits vary from quite densely dendroid pubes- cent beneath, especially along the main veins (Gentry et al. 11172), to densely lepidote but otherwise essentially glabrous (Agostini 110). Most trichomes of these plants are dendroid and stiff; tufts of lax simple trichomes in the nerve axils beneath are uniformly absent. These collections come from Portuguesa, Barinas, and Lara States and the Distrito Federal; I have also seen a collection [Davidse 5219 (COL, MO)] from Vichada Territory in the Colombian Llanos. A fourth form of A. paniculatum, characterized by large leaflets having a denser, uniformly dendroid, rather coarse indumentum, is also found in Venezuela; this has been called A. macrophyllum H.B.K. Its fruits are unknown. Outside Venezuela only the smooth surfaced fruit is known, and it is found associated with plants running the whole gamut from lepidote to densely den- droid pubescent leaves. I have previously followed Seibert (1940a) in treating the conspicuously pubescent-leaved forms (including A. macrophyllum) as A. paniculatum var. molle and the less pubescent forms as A. paniculatum var. paniculatum. In the absence of fruits, Pittier described two new species of Amphilophium from Venezuela based on differences in pubescence. A. xerophilum vegetatively matches the smooth-fruited Venezuelan collections and must be considered part of typical A. paniculatum. A. mollicomum is almost certainly a pubescent extreme (cf. Gentry et al. 11172, 11179) of the form with closely reticulate-ridged fruits which probably merits some kind of taxonomic recognition. I propose to treat it as a variety of A. paniculatum, A. paniculatum var. mollicomum (Pittier) A. Gentry, comb, et stat. nov. ( based on A. mollicomum Pittier, J. Wash. Acad. Sci. 18: 120. 1928.). Although var. paniculatum in Venezuela (and adjacent Colom- bia) can usually be vegetatively separated from var. mollicomum by the uniform leaf indumentum described above, the latter's great variation in leaf indumentum is exceeded by extra- Venezuelan smooth-fruited material, and determination of flowering collections is doubtful. In general, collections from the Cordillera de la Costa region are mollicomum, while collections outside this area are not. How- ever, the occurrence of mollicomum-\ike pubescent forms outside this area (in- cluding the fruiting Colombian collection of bona fide mollicomum) and occa- sional collections with tufted simple trichomes (Pittier 11566) inside it indicates some range overlap and/or breakdown of the vegetative characters. The fruits of macrophyllum are unknown and its possible identification with var. molli- comum remains moot. Only fruiting collections or specimens associated both geographically and by pubescence type with fruiting collections are assigned to 50 ANNALS OF THK MISSOURI BOTANICAL CARDEN [Vol. 63 var. mollicomum in the Flora de Venezuela. The uniformly densely pubescent- leaved forms usually referred to A. macrophyllum II.B.K. are still assumed to he A. paniculatum var. rnolle in the absence of fruiting material which could prove them to be var. mollicomum instead. Amphilophium paniculatum var. imatacense A. Gentry, var. nov. Distinguitur indumenta foliolorum subtili, dense, trichamatibus dendroideis; fructu magno, complanato, dissite muricato-rugoso. Type: Venezuela, bolivar-delta amacuho border: 61°44'W 8°4'N, pri- mary forest near Rio Grande o Toro, E of Upata, 300 m, liana, stem dull gray- brownish- green, tendrils gray-brownish-green, leaves papery, slightly glossy dark green above, dull, rough and paler green beneath, immature fruits dull pale green, 15 x 4.5 em, 8 Apr. 1967, de Bruijn 1662 (MO, holotype; isotypes WAG (3) ). In contrast to A. paniculatum and A. paniculatum var. mollicomum, the Sierra Imataea plant is vegetatively distinct and homogeneous. It clearly warrants taxo- nomic recognition. No collections of A. paniculatum from this area are known and it may well prove more than a variety. However, the variability inherent throughout this complex makes it prudent to accord it only varietal recognition at present. Additional collections examined: Venezuela, bolivar: El Palmar hacia Rio Crande, Sierra de Imataea, 300 m, hejuco grande, hojas bifolioladas, discoloras, calices verdes, sepalos ahiertos, simulando nn calicnlo, corolla en boton amarilla con apice morado, cnando ahierta (hilahiada) purpurea, luego blanca, frutos verrucosos, hasta 15 cm largos, pedasos, especie frecnente, 21 Feb. 1959, Bernardi 7192 (VEN, F). Tumeremo to Anacoco, N side of Cuyuni River, 19 km from Guyana frontier, 140-200 m, 18 Mar. 1974, Gentry et al. 10711 (MO, VEN). Sierra Imataea, rainforest between junction with Rio Re forma and 1 km below junction along river, between Rio la Reforma and Puerto Rico, X of El Palmar, 200-250 in, vine, leaves sub- coriaceous, dull green above, gray green below, calyx spreading, gray green, corolla cream) white, Steyermark SHI 17 (VEN). delta amacuro: Near the border (Rio (Irande o Toro) with Estado Bolivar, ca. 61°44'W 8°4'N, low primary forest, liana 20 m high, 5 cm in diameter, leaves papeiy dull, medium green above, paler beneath, fruits pale green, dull, Bidder 3833 (VEN, WAG). Bosque pluvial Este de Rio Grande, 37 km ENE de El Palmar, high-climbing liana, leaves tawny below, dull green above, corolla deep yellow, 10 Feb. 19o'4, Steyermark 93128 (VEN). Arrabidaea prancei A. Gentry, sp. nov. — Fig. 1A. Frutex scandens; ramuli subteretes, consociebus glandularum in nodis inter petioles; folia bifoliolata, interdum cirrhis simplicibus, foliolis ellipticis, acuminatis; inflorescentia floribus in paniculis parvis axillaribus dispositis; calyx cupulatus, truncatus, minute 5-denticulatus, lepi- dotus; corolla alba Lavandula suffusa, campanulata supra basem longam tubularem; stamina subexserta, thecis di\ aricatis; ovarium anguste oblongum, lepidotum; capsula linearis, ali(jnantum lepidota; semina bialata, bninneola. Liana; branchlets terete, striate, minutely lepidote or glabrous, when older drying reddish brown with numerous small, slightly raised almost circular lenti- eels, with 4 phloem arms in cross section; nodes with interpetiolar glandular fields; pseudostipules not evident. Leaves 2-foliolate, often with a simple tendril; leaflets elliptic, acuminate, truncate to broadly cuneate at base, subcoriaceous, 5-15 cm lone, 4.2-9 cm wide, minutely lepidote, otherwise glabrous, more or less 1976] GENTRY— SOUTH AMERICAN BIGXOXIACEAE 51 Figure 1. A. Arrabidaea prancei A. Gentry; photo of type, Gentry 12882; x Vvs- — B. Cuspidaria subincana A. Gentry; photo of type, Gentry 12825; X 1 /2- conspicuously 3- veined from base, with 3-4 secondary veins on each side; petio- lules 0.6-4.5 cm long; petiole 1-4 cm long, lepidote. Inflorescence a few-branched panicle, flattened and wider at and below each joint, lepidote throughout, puberu- lous only at the joints and on margins of the minute ( less than 1 mm long) bracts and bracteoles, the buds elongate, conical, tapering to an almost acuminate point. Flowers with calyx cupular, 7-10 mm long, 3-4 mm wide, more or less truncate, minutely and evenly 5-denticulate, occasionally slightly split on one side, densely lepidote, puberulous on margin and very sparsely in upper half, with often incon- spicuous plate-shaped glands in upper half; corolla white or greenish white with the tube tinted purple outside, tubular-infundibuliform above a very long tubular base, 4.5-6 cm long, the tube 3.5-4.5 cm long, 0.8-1.0 cm wide at top, the basal portion 2.0-2.6 cm long, 1.5-2 mm wide, the lobes 1-1.5 cm long, densely puberu- lous outside and on lobes inside, the upper part of tube glabrous inside, sparsely puberulous at and below level of stamen insertion, becoming densely glandular pubescent between 9 and 15 mm above base of tube and abruptly glabrous below this area; stamens didynamous, subexserted, the anther thecae divaricate, gla- brous, each 3 mm long, the longer filaments 1.6-1.8 cm long, the shorter filaments 1.2-1.5 cm long, the staminode 4-5 mm long, insertion 2.2-2.6 cm above base of corolla tube; pistil 4-4.5 cm long, the ovary linear oblong, tetragonal, the 4 angles raised (almost cross-shaped in cross section), 4 mm long, 1 mm wide, very densely lepidote, the ovules rather large, 2-seriate in each locule; disc 0.5 mm long, 1.25 mm wide. Capsule linear, compressed, blunt at apex and base, 25-36 cm long, 1.6-1.9 cm wide, the midline barely or not at all evident, the margins rounded, somewhat lepidote, drying brownish; seeds thin, bialate, the wings completely brown or with a thin hyaline marginal fringe, indistinctly demarcated from body of seed, 1.3-1.6 cm long, 2.6-4.8 cm wide. Type: Brazil, amazonas: Km 67 E of Manaus on Manaus-Itacoatiara Road, liana, flowers white, tube faintly lavender above without and sometimes below or within, fragrant, fruits green, 24 Nov. 1974, Gentry 12882 ( INPA, holotype; isotypes MO and to be distributed). 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Additional collections examined: Brazil. AMAZON AS: Basin of Rio Negro, road from Camanaus to Uaupes airport, roadside, vine, corolla white, exterior of tube purple, 30 Oct. 1971, Prance et al. 15863 (MO, NY). Rio Purus, Rio Ituxi, Lago Preto, 2 km N of Labrea, forest on terra firme, liana 10 cm diam. at widest point, calyx green-brown, corolla lavender- white outside but darker adaxially white inside, 26 June 1971, Prance et al. 13757 (MO, NY). Manaus, estrada Manaus-Itacoatiara, km 55, T. firme, argiloso, trepadeira, frutos maduros, 14 Oct. 1960, Rodriguez 6- Coelho 1839 (INPA). Estrada Castanho Tupana, entre o km 50-10; flores de color branca, tube violaceo, frutos verdes, 18 July 1972, Silva et al 847 (INPA, MO). : Cruzeiro do Sul, Rio Jurua & Rio Moa, Estrada Alemanha, disturbed secondary forest by road, corolla greenish white tinted purple on exterior, 14 Apr. 1971, Prance et al 11834 (NY). para: Rio Jari, Monte Dourado, flor lila clara, Oliveira 4479 (IAN). Rio Jari, Monte Dourado to Munguba, 13 May 1969, Silva 1955 (IAN). Rio Jari, Planalto A, flor branco aroxeado, 23 Apr. 1969, Silva 1955 (IAN). Rio Jari, Estrada de Munguba, km 4, 19 Apr. 1969, Silva 1876 (IAN). Rio Jari, Estrada entre Monte Dourado e Planalto A, km 3, 22 May 1969, Silva 203() (IAN). The affinity of this distinctive new species with Arrabidaea is not immediately obvious because of its elongate white corolla. In such features as its racemose- paniculate inflorescence, subexserted anthers, and elongate tubular white flowers it is reminiscent of Leucocalantlie. Nevertheless, it shares most of the important characters of Arrabidaea, including interpetiolar glandular fields, simple tendrils, stems with 4 phloem amis in cross section, corolla pubescent without, straight divaricate anther thecae, and a conspicuous nectariferous disc. In fact, were one to predict the characteristics to be expected in a species of Arrabidaea adapted for hawkmoth pollination, one would come close to a description of A. prancei. Its closest relative seems to be A. triplinervia which has very similar trinerved leaflets, reduced, almost racemose inflorescences, a similar corolla shape (except for the long narrow basal constriction), and sometimes white flowers. Cuspidaria subincana A. Gentry, sp. nov. — Fig. IB. Frutex seandens; ramuli subteretes, consociebus glandularum in nodis inter petioles; folia trifoliata vel bifoliata, interdum cirrhis simplicibus, foliolis ellipticis, obtusis vel breviter cuspi- datis, subtus canescentibus; inflorescentia floribus in panieula dispositis; calyx cupulatus, trun- catus, minute 5-dentieulatus, puberulus; corolla rubra, tubulo-infundibuliformis; antherarum lobi semicirculares, eonneetivo crasso; ovarium ovoideum, minute lepidotum; capsula linearis, subteres, pubemla, bisulcata, seminibus bialatis. Liana; branchlets terete, minutely striate, puberulous; interpetiolar glandular fields present; pseudostipules not noticeable; conspicuously lenticellate when older. Leaves 3-foliolate or 2-foliolate with a tendril or tendril scar; leaflets elliptic or obovate-elliptic to subrotund, the apex rounded or abruptly cuspidate, the base rounded to broadly cuneate, 3.5-14 cm long, 2.2-9 cm wide, chartaceous to sub- coriaceous, the main veins slightly raised beneath, densely canescent below except the darker-drying veins, glabrous above except on midvein, drying dark brown above, whitish below with the less pubescent main veins contrastingly darker; petiolules 1-3 cm long; petioles 2-5 cm long, puberulous. Inflorescence an axil- lary or terminal panicle, its branches puberulous. Flowers with the calyx cupular, 5-denticulate, 4-5 mm long, 2-3 mm wide, conspicuously puberulous, eglandular or with a few glands near margin, the marginal teeth often extending as raised lines on upper third of calyx; corolla magenta with a white throat, tubular- infundibuliform, 2.7-4 cm long, 0.7-1.3 cm wide at mouth of tube, the tube 2.3- 1976] GENTRY— SOUTH AMERICAN BIGNONIACEAE 53 3.4 cm long, the lobes 0.5-0.8 cm long, pubernlous outside and on lobes and at level of stamen insertion inside; stamens didynamous, the anther thecae bent and reflexed sharply forward near middle, 1-2 mm long, glabrous, the connective extremely thick, extended; pistil 1.7-2.4 cm long, the ovary ovoid, 1.5-1.8 mm long, 1-1.2 mm wide, with two longitudinal furrows on each side, densely mi- nutely lepidote, sometimes also scattered puberulous, the ovules 4-seriate in each locule; disc cupular-pulvinate, 1 mm long, 1.5 mm wide. Capsule linear, terete, 29 cm long, 1.3 cm wide, puberulous, drying grayish, with scattered small lenticel- like glands, each valve with a conspicuous median longitudinal furrow 2-3 mm wide and bordered by a slightly raised line on either side; seeds thin, flattened, bialate, 8-9 mm long, 3.7-4.6 cm wide, the hyaline membranaceous wings sharply demarcated from the seed body. Type: Brazil, amazonas: 2-5 km N of Manaus-Itacoatiara Road at km 79 near Rio Preto da Eva, 100-200 m, liana, flowers magenta with white throat, 24 Nov. 1974, Gentry 12825 (INPA, holotype; isotypes MO, MG, RB, to be dis- tributed ) . Additional collections examined: Venezuela, holivar: Bosques a lo largo de la frontera Venezolano-Brasilera, NE de la Serrania Pia-soi ( Pia-shauhy, pia-Savi), 3°53', 62°46', 650-800 m, high-climbing vine, leaves subcoriaceous, deep green above, gray-green below, calyx brown- ish, corolla deep rose, style pink-maroon at tip, whitish below, rachis of inflorescence tawny- green, 5-6 Jan. 1962, Steyermark 90694 (VEN). Brazil, amazonas: 2-5 km N of Manaus- Itacoatiara Road at km 79 near Rio Preto da Eva, 100-200 m, Gentry 12850 (INPA, MO). Manaus-Caracarai Road (BR 174), ca. km 100, roadside, vine, flowers magenta with white throat, 30 Nov. 1974, Gentry I- Ramos 12924 (INPA, MO). Estrado do Aleixo near Manaus, km 6-7 past INPA, second growth and forest edge, vine, fruits gray green, 2 Dec. 1974, Gentry 13042 (INPA, MO). Munic. Humayta, near Livramento, Rio Livramento, 12 Oct.-6 Nov. 1934, Krukoff 6878 ( K, MO, NY). Munic. Manaus, road to Aleixo, 12 Aug.-l Sep. 1936, Krukoff 8012 (K, MO, NY). Km 5.5-60 da Rodov. Manaus-Itacoatiara, cipo, sobre chao, flores lilas, 24 Oct. 1963, Oliveira 2770 (IAN, MO). Vic. of Labrea airport, Rio Purus, Rio Ituxi, fruit green, glaucous, 29 June 1971, Prance et al. 13974 (MO, INPA, NY). Mata da terra firme, solo argiloso, entre o rio Castanho e o Araca, trepadeira com flores violaceae, 12 July 1972, Silva et al. 549 (INPA, MO). 7 km N of Manaus on Estrada do Aleixo, vine, flowers light magenta, 21 Nov. 1974, Gentry 6 Prance 12787 (INPA, MO). Ca. km 80, Manaus-Caracarai Road (BR 174), road cut, vine, flowers magenta, 30 Nov. 1974, Gentry b Ramos 12927 (INPA, MO). Ca. km 70, Manaus-Caracarai Road (BR 174), road cut, vine, flowers magenta with white throat, reddish lines above each row of anthers, 1 Dec. 1974, Gentry 13006 (INPA, MO). Es- trada do Aleixo, Manaus, 15 Oct. 1947, Guedes 24 (IAN). Haplolophium rodriguesii A. Gentry, sp. nov. — Fig. 2. Frutex scandens; ramuli sexanguli sine consociebus glandularum in nodis inter petioles; folia bifoliolata, interdum cirrhis fissis, foliolis ellipticis, coriaceis, subtus puberulis; inflores- centia floribus in racemis axillaribus dispositis; calyx cupulatus, margine crispato, stellato- piloso; corolla rubra, tubulo-campanulata, ad medium flexa; stamina didynama, thecis divari- catis; ovarium ovoideum, dense tomentosum; discus patelliformis; capsula ignota. Liana; branchlets 6-sided, the angles ribbed, drying blackish, puberulous; interpetiolar glandular fields absent; pseudostipules not noticeable. Leaves 2- foliolate with a trifid tendril; leaflets elliptic, abruptly short-acuminate, the base rounded, 15-17 cm long, 9-12 cm wide, coriaceous, the veins and veinlets plane above, raised and intricately and conspicuously reticulate below, immersed-lepi- dote above, otherwise glabrous, simple-puberulous below, drying olive gray; 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Figure 2. Ilaplobphium wdriguesii A. Gentry; x Mj. [After Bodrigues 12951 (1NPA).] petiolules 2-4 cm long; petiole 6.5-7.5 cm long, puberulous, Inflorescence a fascicle of axillary racemes, puberulous with stellate trichomes, the flowers borne in pairs along rachis separated by 2-4 cm, each pair of pedicels subtended by two linear bracts 5-7 mm long and 1 mm wide, the pedicels 0.5-1 cm long, with a pair of linear 2-5 mm long bracteoles near middle. Flowers with the calyx hav- ing a cupular base and an expanded frilly margin, the base 6-8 mm long and 4-6 mm wide, the frilly margin ca. 5 mm wide, stellate-tomentose, with paired sub- linear glandular fields at top of base below the margin; corolla red, aromatic, relatively thick, tubular-campanulate, bent at 90° angle above base, 4-5 cm long, 0.8-1.2 cm wide, the tube ca. 1 cm to the bend, 2-2.5 cm above the bend, the lobes ca. 1 cm long, the throat with two longitudinal ridges, densely pubescent outside, pubescent inside on lobes and at level of stamen insertion, otherwise lepidote inside; stamens didynamous, inserted 7-8 mm from base of corolla tube, 1976] GENTRY SOUTH AMERICAN BIGNONIACEAE 55 the anther thecae divaricate, 3-4 nun long, the pollen grains 8-colpate; pistil 3.3- 3.5 cm long, the stigma elliptic, acuminate, 3-4 mm by 1.5 mm, the style stellate tomentose, the ovary ovoid, 2 mm long, 2 mm wide, densely tomentose, the ovules ca. 8-seriate in each locule; disc patelliform, 1-1.5 mm long, 4-5 mm wide. Cap- sule unknown. Type: Brazil, am azon as : Manaus, estrada do Igarape do Passarinho, terreno firme argiloso, mata, trepadeira com gavinhas, flores roxas, aromaticas, 22 May 1962, Rodrigues i? Chagas 4460 (INPA, holtotype; MO, NY, isotypes). This remarkable plant, only the second known species of Haplolophium, is very distinct from //. bracteatum Cham, in its much larger leaves with intricately reticulate and densely puberulous lower surfaces, in its much more open inflores- cence, and especially in its very inconspicuous linear inflorescence bracts and bracteoles. Nevertheless, it shares the fundamental characters of the genus in- cluding 6-angled branchlets, trifid non-disc-tipped tendrils, 8-colpate pollen, and, most notably, a calyx with a cupular truncate base and a frilly expanded margin. Haplolophiumis intermediate between Pilhecocteniiim and Amphilophium, agree- ing with the former in its tubular bent corollas, simple calyx, and echinate fruit and with the latter in its pollen, trifid tendrils, dendroid trichomes, and the pres- ence of a frilly calyx margin. The intricately reticulate and raised venation be- neath of //. rodriguesii is strikingly like that of Distictella monophijlla Sandw. Memora aspericarpa A. Gentry, sp. nov. — Fig. 3. Arbor parva vel frutex; ramuli subteretes, sine consociebus glandularum in nodis inter petioles; pseudostipulae crassae foliaceae; folia saepe 15-foliolata pinnis principal ibus tribus, foliolis lanceolatis vel anguste ellipticis, acuininatis, basi cuncatis vel rotimdatis, glabris; in- florescentiae racemosae terminales, bracteis mature deciduis, probabiliter minutis, bracteolis minutis; calyx coriaceous, campanulatus, bilabiatus, extus plcrumque tflaber intus lepidotns; corolla flava, tubulo-infundibuliformis, extus glabra; stamina thecis divaricatis; pistillum ovario glabre; discus pulvinatus; fructus oblongus, teres, exasperatus, seminibus exalatis, maxime crassis. Small tree or shrub, rarely subscandent, to 3 m tall; branchlets subterete, glabrous, lentieellate; nodes without interpetiolar glandular fields; pseudostipules thick-foliaceoiis, obovate, rounded. Leaves 2-3-compound, usually 15-foliolate with three 5-foliolate primary pinnae, each leaflet of lowermost leaflet pair of the ter- minal pinna sometimes replaced by a 3-foliolate secondary division, the leaf thus to 19-foliolate; tendrils not seen; leaflets lanceolate to narrowly elliptic, less com- monly elliptic or narrowly ovate, acuminate, cuneate to rounded at base, sub- coriaceous, 4-17 cm long, 1.2-8 cm wide, completely glabrous or with a few in- conspicuous simple trichomes along midvein above; petiolules and petiole very sparsely puberulous or glabrate. Inflorescence a contracted terminal raceme, drying dark, the pedicels 0.8-1.5 cm long, inconspicuously short-puberulous, the bracts early deciduous, probably minute, the bracteoles minute, subulate, 1-4 mm long, ca. 1 mm wide, attached at or below middle of pedicel 5-10 mm below calyx, lepidote inside, finely pubescent at tip and along margin. Flowers with the calyx coriaceous, campanulate, bilabiately split %-M» its length, 10-12 mm long, 8-10 mm wide, somewhat lepidote, with a few trichomes at apices of lobes, 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 N . Figure 3. Memora aspcricarpa A. Gentry. — A. Habit; X V>i- [After Steyermark 105375 (VEN).]— B. Fruit; X %. [After Ariste^uieta 6 Pannier 1842 (VEN).] otherwise glabrous outside, glandular lepidote inside, eglandular outside; corolla yellow, tubular-infundihuliform, 5.5-7.5 cm long, 1.2-2.4 cm wide at mouth of tube, the tube 4-5.8 cm long, lobes 1-1.5 cm long, mostly glabrous, the lobes sometimes with a few marginal trichomes, villous at and below level of stamen insertion; stamens didynamous, the anther thecae divaricate, 3-4 mm long, the longer filaments 3.7-3.8 cm long, the shorter filaments 2.7-2.8 cm long, the stami- 1976] GENTRY— SOUTH AMERICAN B1GNONIACEAE 57 node 10-11 mm long, insertion 16-18 mm from base of corolla tube; pistil 4.8-5 cm long, the ovary linear-oblong, 3 mm long, 1 mm wide, glabrous; disc patelli- form, 1-1.5 mm long, 2.5-3 mm wide. Capsule oblong, terete, 8-22 cm long, 3.3- 4.7 cm in diameter, the apex obtuse or acutish, the base more or less rounded, macroscopically conspicuously rough-surfaced with many minute rather flat tubercles somewhat resembling coarse sandpaper in appearance, these tubercles coalescing and not individually distinct under a lens; seeds very thick, wingless, angulate, 2.3-2.5 cm long, 2.0-3 cm wide, the hilum broad, convex, 7-12 mm wide, 17-21 mm long. Type: Venezuela, yaracuy: Selva nublada sobre piedras calcareas, entre el pueblo de Aroa y Altamira, 1050 m, small tree 3 m tall with slender trunk 1 cm diam., divergently branched above, corolla yellow, 22 Jan. 1972, Steyermark 105375 (MO, holotype; YEN, isotype). Additional collections examined: Venezuela, distrito federal: Caruao, (fruit), Pittier s.n. (VEX), yaracuy: Montana que queda enfrente del caserio de Quebrada Seca, trepadora de olor a ajos, 2 July 1953, (fruit), Atisteguieta isr Pannier 1842 (VEN). tachira: A la orilla de potreros, cerca a Jordan, bosque humedo tropical, arbolito, approx. 3 m alto, 26 Apr. 1964 (fruit), Yjjaz 351 (MY), lara: Terepaima, 1300-1600 m, arbusto trepador de flores amarillas (flowers, fruits), 24 Mar. 1959, Trujillo 4116 (MY). This species is restricted to upper elevations (mostly over 1,000 m) along the Venezuelan coastal cordillera east to the Distrito Federal. L M bilabiate (rather than spathaceous) calyx and thick-valved rough- surfaced fruit. It is perhaps most closely related to M. cladotricha Sandw. even though that species belongs to a different section of the arbitrarily subdivided genus. It resembles M. cladotricha in habit and fruit although lacking foliaceous inflores- cence bracts and bracteoles. The fruit of M. cladotricha, though similar to that of M. aspericarpa, has a macroscopically smoother surface with a less dull texture. The papillae covering the fruit of M. aspericarpa tend to coalesce and are indi- vidually obscure when seen through a lens; the papillae of M. cladotricha are individually distinct microscopically. Memora tanaeciicarpa A. Gentry, sp. nov. Frutex scandens; ramuli teretes, glabrati, sine consociebus glandularum in nodis inter petioles; folia 2-3-ternata, interdum cirrhosa, foliolis ellipticis vel ovatis, acuminatis; inflores- centia racemosa, axillaris, bracteis caducis; calyx campanulatus, minute 5-denticulatus; corolla tubulo-infundibulifonnis, tubo extus puberulo; stamina thecis divaricatis; ovarium oblongum, puberulum; discus pulvinatus. Liana; branchlets terete, glabrate, finely striate, elenticellate or with incon- spicuous elongate lenticels; nodes without interpetiolar glandular fields or pseudo- stipules. Leaves 2-3-ternate, the terminal 3 leaflets sometimes replaced by a simple tendril; leaflets elliptic to ovate, acuminate, the base cuneate to truncate, chartaceous, 4-15 cm long, 1.5-7 cm wide, very minutely and inconspicuously scattered lepidote or lepidote-papillose above and below, otherwise glabrous or very sparsely and minutely subpuberulous along midvein below, drying olive with contrasting yellowish or reddish main veins below, the main veins plane 58 ANNALS OK THE MISSOURI BOTANICAL GARDEN [Vol. 63 above, prominent below; petiolules and petiole glabrous or subpuberulous adaxially. Inflorescence an axillary raceme, puberulous, the bracts early decidu- ous, 2-3 mm long, 2-3 mm wide, yellowish puberulous and glandular, bracteoles absent. Flowers with the calyx campanulate, truncate, minutely 5-denticulate, 10-13 mm long, 6-9 mm wide, densely yellow-tomentose with branched trichomes, with scattered black-drying glands in upper half; corolla tubular-infundibuliform, ca. 4 cm long (mature corollas seen all shrivelled and partially destroyed), densely puberulous outside and on lobes inside, with plate-shaped glands at base of lobes outside, the tube mostly glabrous inside, densely villous at and be- low level of stamen insertion; stamens inserted 12-13 mm above base of corolla tube, the filaments 1.1-2.1 cm long, the anther thecae divaricate, 2.5-3 mm long; pistil 3.2-3.3 cm long, the style puberulous with branched trichomes, the ovary oblong, 3 mm long, 1 mm wide, densely puberulous; disc annular-pulvinate, 1 mm long, 2.5 mm wide. Type: Brazil, amazonas: Manaus, km 10 estrada Manaus-ltacoatiara, trepadeira com flores amarelas na mat a de t. firme solo argiloso, 16 May 1972, Loureiro, Fires 6- Athanagiklo s.n. (INPA 35794) (INPA, holotype; MO, isotype). Additional collections examined: Venezuela, amazonas: Dept. Apnres, Rio Orinoco, alrededores de Siquita entre la Isla Castillito y San Fernando de Atahapo, 100-140 in, Btnitin^ r£ al. 3631 (MY). Bhazil. amazonas: Duclce Forest Reserve, km 26 on Manans-Itacoatiara road, sterile vine, tendrils simple, stems rou^h, greenish, 23 Nov. 1974, Gentry 12S20 (MO, INPA). paha: Cabeceiras do Rio Uruara, flancos do Planalto Amazonico, mnnicip. de Prainlia, cipo ao lonRo do no, 11 May 1955, Froes 31 SSI, 31SS6 (both IAN). Santarem, Rio Maica, Serra de Taperinha, capoeira do pe da serra, cipo robnsto, fruto verde, 5 Feb. 1968, Silva 1371 (MC). This is quite unlike any other species of Memora in its large 4-valved capsule, its puberulous style and ovary, its yellowish-red puberulous 5-denticulate calyx, and its puberulous corolla tube. Except for the 2-3-ternate leaves it seems closer to Adenoeahjmma comosum and allies than to other Memora species, additional evidence of the artificiality of the separation of Memora and Adenoeahjmma. Tanaeciurn apiculatum A. Gentry, sp. nov. — Fig. 4. Frutex scandens; raimili teretes, sine consociebus glandularum in nodis inter petioles; folia bifoliolata, foliolis ellipticis, apiculatis, leviter lepidotis aliter glahris; inflorescentia florihus in racemo dispositis; calyx tubulosus, lepidotus, minute puherulous; corolla alba, elonj^ato-tubn- loso, puherula; stamina subexserta; ovarium lineare, dense lepidotum; fructus juvenis linearis, lepidotus. Liana; branchlets terete, striate, hollow, glabrous; interpetiolar glandular fields and pseudostipules absent. Leaves 2-foliolate with a (simple?) tendril; leaflets elliptic, sharply apiculate, rounded at the base, 16-18 cm long, 8.5-9 cm wide, membranaceous, lepidote above, inconspicuously scattered-lepidote below, other- wise glabrous, drying gray with a whitish cartilaginous margin, the apiculation with plate-shaped glands; petiolules 1.5-2.7 cm long; petioles 3.2-3.6 cm long, very slightly lepidote. Inflorescence a raceme, minutely lepidote and densely mealy puberulous, the rachis ca. 20 cm long, the pedicels ca. 0.5 cm long, the 3-4 Mow I X ers 6-/ mm 1976] GENTRY— SOUTH AMERICAN B1GNOMACEAE 59 Figure 4. Tanaecium apiculatum A. Gentry. — A. Inflorescence; X %> — B. Leaves; X ¥2 [After Smith 226 (US).] 60 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 wide, lepidote and minutely puberulous witli stellate trichomes, circumscissile and falling with corolla; corolla white, elongate-tubular, expanded apically, 15- 16 cm long, 14-16 mm wide at the mouth, the tube 13-14 cm long, the lobes 2-2.5 cm long, puberulous outside, the lobes glandular-lepidote inside, the tube gla- brous inside except for a few lepidote scales at base of filaments, with linear fields of plate-shaped glands near base of lobes outside; stamens didynamous, the anthers subexserted, the thecae 4(?) mm long, the filaments 10-12 cm long, insertion 9 cm above base of corolla tube, staminode 1 mm long, inserted 8.3 cm above base of tube; pistil 14-16 cm long, the ovary linear, the base somewhat widened, densely lepidote, 4-6 mm long, ca. 1 mm wide; disc annular-pulvinate, ca. 1 mm long, ca. 3 mm wide. Very young fruit linear, subterete, 8 cm long, 0.6 cm wide, scattered lepidote and with a few platter-shaped glands. Type: Venezuela, monacas: Caicara, important vine of the heavy woods, flowers white, wet woods, "caratero," 15 May 1952, Smith 226 (US 2121468, holo- type; US 2121469, isotype, MO fragments). This plant is utterly distinct from any other species of Bignoniaceae. Its elon- gate tubular white flowers are those characteristic of Tanaecium but the long tubular circumscissile calyx is unique in that genus. Cartilaginous leaflet margins are also unique in Tanaecium although found in several species of Adenocalymma; the striking terminal apiculations of the leaflets of this species are found in no other species of Bignoniaceae. Tynnanthus villosus A. Gentry, sp. nov. — Fig. 5. Fnitex scandens, in omnes partes villosus, sine consoeiebus glandularum in nodis inter petioles; pseudostipulae foliaceae; folia bifoliolata, foliolis oblongo-ellipticis; infloreseentia floribus in panicula axillari dispositis; calyx cupulatus, subtruncatus, puberulus; corolla cremea, bilabiata, extus pubernla; stamina didynama, thecis divaricatis, flexis; ovarium conicum puberu- lum; discus deficiens; capsula i^nota. Liana; branchlets terete, striate, villous with ca. 1 mm long reddish yellow trichomes; interpetiolar glandular fields absent; pseudostipules foliaceous, 0.9-1.5 cm by 0.8-1.5 cm. Leaves 2-foliolate with a tendril or tendril scar; leaflets more or less oblong-elliptic, acuminate, the base asymmetrically rounded thus usually subcordate on one side, 5-11 cm long, 3-7 cm wide, membranaceous, the main veins raised below, velutinous below with simple trichomes especially along the main veins, less conspicuously pubescent above, drying olive; petiolules 1-1.5 cm long; petioles 2-4 cm long, conspicuously villous. Inflorescence a racemose axil- lary panicle, villous, the rachis 4-12 cm long, the short, 2-5 mm long peduncles at right angles to it, each bearing 2-3 flowers and subtended by a narrow cadu- cous bract 2-3 mm long. Flowers with the calyx cupular, subtnmcate or minutely 5-denticulate, 1.5-2 mm long, ca. 2 mm wide, pubescent, eglandular; corolla cream ("amarillo blancuzco"), bilabiate, ca. 0.5 cm long, split about half its length, the 2 upper lobes almost fused, the 3 lower ones ca. 2 mm long, puberu- lous outside, inside puberulous on lower 3 lobes, floor of tube, at level of stamen insertion and on margins of upper 2 lobes; stamens didynamous, the anther thecae 1 mm long, divaricate and reflexed forward from a basal twist, the connec- 1976] GENTRY— SOUTH AMERICAN 1 BIGNONIACEAE 61 B Figure 5. Tynnanthus villosus A. ('.entry. — A. Habit; X M->. — B. Flower; X 5. [After Schunke6852 (MO).] tive extended 0.2 mm beyond anther attachment, the longer filaments ca. 2 mm long, the shorter filaments ca. 1.5 mm long, the staminode 1 mm long, insertion 0.5 mm from base of corolla tube; pistil 3 mm long, the stigma narrow, bilamellate, the style pubescent toward base, the ovary conical, ca. 1 mm long, 1 mm wide at base, densely pubescent, the ovules 3-4-seriate in each locule at bottom of ovary and 2-seriate at top; disc not apparent. Capsule not known. g2 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Type: Peru, san martin: Prov. Mariscal Caceres, Dtto. Tocache Nuevo, Que- brada de Canute, 400 m, liana de 8-9 m, flores amarillo blancuzco, sepalos brilliant yellow-green, en bosque secundario y con abundante luz solar, 18 Aug. 1973, Jose Schunke Vigo 6852 (MO, holotype; isotypes to be distributed). Additional collection examined: Peru, huanuco: Hills E of Tingo Maria, sterile liana, Croat 21121 (MO). This unmistakable plant has no obvious affinities with other species of Tyn- nanthus. It differs markedly from all other species of the genus in its shaggy indumentum, its narrow almost racemose inflorescence, and its persistent folia- ceous pseudostipules. Foliaceous pseudostipules are occasionally also found in the very different Tynnanthus myrianthus Bur. & K. Schum., but mostly on young bran chiefs. Tecoina peclicellata Bur. & K. Schum. in Mart. Fl. Bras. 8(2): 336. 1897. Handroanthus pcclicellatus (Bur. & K. Schum.) Mattos, Loefgrenia 50: 4. 1970. Bureau and Schumann noted the close relationship of their new species, actually a Tabebuia, with Tabebuia chrysotricha (Mart, ex DC.) Standi, but considered it distinct on account of its longer pedicels, smaller corolla and calyx, and different calyx indumentum. The species is known only from the type col- lection, Glaziou 1476 from Corcovado, Rio de Janeiro, Brazil. Presumably the holotype at Berlin has been destroyed. Isotypes at BR, K, and P have flowering branchlets resembling Tabebuia ochracea (Cham.) Standi, detached from branch- lets with leaves identical to those of Tabebuia chrysotrieha. Could the Claziou gathering be a mixture of these two species? A second isotype at Paris proves the suspicion well founded. This sheet has, in addition to two detached flowering branchlets resembling Tabebuia ochracea, a branchlet bearing both leaves and flowers. The attached flowers are those of Tabebuia chrysotricha, and T. pedicel- lata is indeed based on a mixed collection. I propose that Tecoma peclicellata be rejected under Article 70 of the Code of Nomenclature since the name was based on a type consisting of two or more discordant elements. OXOHUALCOA & MAXSOA Mansoa verrucifera (Schlecht.) A. Gentry, comb. nov. Bignonia verrucifera Schlecht., Linnaea 26: 655. 1853. type: Venezuela, Curucati, Wagener 307 (not seen). Onohualcoa verrucifera (Schlecht.) A. Gentry, Ann. Missouri Bot. Card. 60: 885. 1974 (1973). ( Includes complete synonymy. ) The monotypic genus Onohualcoa has been characterized (Sandwith, 1947) by its trifid tendril, rugose-warty calyx with ribs ending in subulate teeth, pink corolla, and tuberculate-echinate capsule. Dugand (1946) noted the differences between Onohualcoa (as Bayonia) and Adenocalymma, Pseudocalymma (now included in Pachyptera), Chodanthus (now included in Mansoa), and Petastonia ( now included in Arrabidaea). He separated his Bayonia from Chodanthus by the latter\s lack of prominent calyx ribs and tubercles on its capsules, although 197fi] GKXTRY— SOUTH AMERICAN BIGNOX1ACEAE 63 noting that the fruit is otherwise similar. The other genera mentioned by Dugand are clearly less closely related. Sandwith (1947) emphasized that Onohualcoa should be compared with Mansoa with a similar inflorescence but a smooth elongate-linear capsule. I have previously (Gentry, 1974a) aeeepted Sandwith's interpretation of Onohualcoa but united the two speeies recognized by him. On the other hand, Standley & Williams (1974) inexplicably sink Onohualcoa back into utterly un- related Adenocalymma. Recently I was able to see Mansoa difficilis (Cham.) Bur. & K. Sehum, in flower in the field and was surprised by its striking resemblance to Onohualcoa. A reexamination of the differences separating Onohualcoa and Mansoa shows that they are very closely related and should be united. In addition to the general similarity between the two genera (see Fig. 6) in such characteristics as subulate calyx teeth, the peculiar inflorescence, tendency to 3-foliolate leaves and 3- veined leaflets, corolla shape, color and pubescence, trifid tendrils, ovules 2- (-4-) seriate in each locule, and large nectariferous disc, may be added two more important similarities. The pollen of Onohualcoa has not been described. That of Mansoa is of an unusual and very characteristic camporeti dilate form with the alveolate exine broken into numerous discrete patches / (Fig. 6) proves to be the same as that of Mansoa and strongly supports the union of the two genera (Tombs & Gentry, in preparation). A second piece of new evidence comes from the capsules of the two genera. Onohualcoa \s chief claim to generic rank is its echinate-tuberculate capsule. The known capsules of Mansoa are relatively smooth-surfaced, although that of M. difficilis usually has raised ridges and may have a beaded almost subtuberculate surface texture. However, fruiting specimens of a new species of Mansoa from northeastern Brazil are now at hand which have the same type of echinate-tuber- culate surface as Onohualcoa (see below). If the fruit of Onohualcoa is not unique, then the genus cannot possibly be maintained as distinct. Even were the fruit distinct, merger with Mansoa would seem in order — the general trend now emerging in Bignoniaceae (Gentry, 1973, 1974b) is of genera defined by similar floral and vegetative characters with differences in fruits often characterizing & only species and varieties. Mansoa onohuaicoides A. Gentry, sp. nov. Fnitex scandens; ramulis teretibus, sine consociebus glandularum in nodis inter petioles; folia trifoliolata vel bifoliolata cum cirrho trifido, foliolis ovatis vel ellipticis, fere glabratis; inflorescentia anguste panieulata, axillaris; calyx cupulatus, 5-setaceus setis 1-2 nun longis, puberulus; corolla tubulo-intundibuliformis, tubo extus glabro; ovarium anguste oblongum, lepidotum vel lepidoto-tuberculatnm; discus pulvinatus; capsula anguste oblonga, teres, ver- rucoso-tuberculata, seminibus bialatis. Vine; branchlets terete, finely striate, with minute round whitish lenticels; nodes without interpetiolar glandular fields or pseudostipules. Leaves 2-3-folio- late, sometimes with a trifid tendril; leaflets narrowly ovate to elliptic, acute to acuminate, the base rounded to truncate, chartaceous, 2-11 cm long, 1-5 cm wide, 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 1976] GENTRY— SOUTH AMERICAN BIGXONIACEAE 65 distinctly scatterecl-lepidote above and below, otherwise glabrous or subpuberu- lous at base of midvein above and below, with clusters of plate-shaped glands in lower nerve axils below, drying olive to blackish olive, the main veins plane above, prominent below; petiolules 0.3-2.5 cm long; petioles 0.5-3 cm long, puberulous. Inflorescence veiy narrowly paniculate, the lateral branches each 2-3-flowered, puberulous, ebracteate. Flowers with the calyx cupular, 5-8 mm long (without teeth), 5-6 mm wide, basically truncate, 5-setate, the subulate teeth 1-2 mm long, very briefly extended as ridges near apex of calyx, puberulous; corolla tubular-infundibuliform, 4-4.5 cm long, the tube 3-3.5 cm long, the lobes ca. 1 cm long, the tube mostly glabrous outside, becoming sparsely puberulous toward top, glabrous inside except at level of stamen insertion, the lobes puberu- lous outside and inside; stamens didynamous, included, inserted 7-8 mm above base of corolla tube, the filaments ca. 2.5 cm long, the anther thecae divaricate, 4 mm long, the connective slightly extended; pistil 3 cm long, the ovary linear- oblong, 2.5-3 mm long, 1 mm wide, slightly narrowed toward base and apex, densely minutely lepidote or lepidote-tuberculate, the ovules 4-seriate in each locule; disc patelliform, 0.5-1 mm long, 1.5-2 mm wide. Capsule linear-oblong, subterete, verrucose-tuberculate, the enations ca. 1 mm long, ca. 15 cm long, 1.4 cm wide; seeds bialate, 0.7-0.9 cm long, 2.5-3.5 cm wide, the wings hyaline- membranaceous, clearly demarcated from rather thick brown seed body. Type: Brazil, ceara: Serra de Baturite, Quebradas occidentaes perto do sertao de Caninde, 24 Aug. 1908, Ducke s.n. (MG1594) (MG, holotype). Additional collections examined: Brazil, ceara: Guaramiranga, Matta, "cipo de cesto," 20 July 1908, Ducke s.n. (MG1356) (MG). Pernamhucxk Brejo de S. Jose, planta n. 5, 7 Sep. 1960, Lima 60-3525 (RB). Maranhao: Maracassiune River region, Campo do Casins, Froes 1857 (MO, NY). This species is somewhat intermediate between M. cliff icilis (Cham.) Bur. & K. Schnm., which ranges through most of eastern Brazil, and M . verrucifera, which ranges from Mexico to Guyana and south along the Andes to Acre, Brazil. The verrucose-tuberculate fruit is like that of M. verrucifera but narrower and with thinner, nonwoody capsule valves and smaller, more widely separated echinations; the thinness of the valves suggests M. difficilis, however. Immature fruits are more densely echinate and exactly like those of M. verrucifera. The flowers could easily be confused with either M. difficilis or M. verrucifera but the ab- breviated (or absent) calycine ridges, nonverrueose calyx surface, and almost glabrous corolla tube suggest the former. The nonribbed lenticellate branchlets URE 6. Anemopaegma and Mansoa. — A, D. Anemopaegma; B-C. Pseudopaegma. — A. orhiculatum (Jacq. ) DC; X r /is- — B. A. oligoneuron (Sprague & Sandw. ) A. Gentry; x 1 %7- C. A. longidens DC; X V.\- — D. A. chrysoleucum (H.B.K. ) Sandw.; X %s« — E, G. Mansoa; F, II. Onohualcoa. — E. M. difficilis (Cham.) Bur. & K. Schum.; x 1 %7- — F. M. verrucifera (Cham.) A. Gentry; X %i.— G. Pollen of M. difficilis; X 780.— H. Pollen of M. verrucifera; X 940. Note the similarity of flower shape in Anemopaegma and Pseudopaegma (A-D) and Mansoa and Onohualcoa (E-F), the subtruncate calyx like that of Anemopaegma in B, and the camporeticulate pollen of both Mansoa and Onohualcoa (G— II). gg ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 are unlike M. difficilis; branchlets of M. verrucifera are similar but less noticeably lenticellate. Pinnately veined leaflets with gland fields in the axils of several secondary nerves below are different from either species. Mansoa onohualcoides provides the heretofore missing link between Onohualcoa (i.e., M. verrucifera) and other species of Mansoa. In addition to the cited specimens from northeastern Brazil, a collection from southern Venezuela closely resembles M. onohualcoides and may be conspecific. The Venezuela collection [Lizot 90 from Mavacca, Territorio Amazonas (VEN)] has a shorter more noticeably branched inflorescence, lacks conspicuous lenticels, and has leaflets without glandular fields in the nerve axils below, not notably lepidote, and drying reddish black. PSEUDOPAEGMA & ANEMOPAEGMA Urban (1916) proposed Pseudopaegma as a segregate from Anemopaegma based mainly on the 5-6-colpate pollen of the former versus the ecolpate pollen of the latter. However, Gomes (1955) subsequently described the pollen of Anemopaegma (based on A. pachyphijllum and A. hilarianum, species not exam- ined by Urban) as 5-7-colpate and additional species of that genus also prove to have several-colpate pollen like that of Pseudopaegma ( Tombs & Gentry, in prep- aration). Clearly pollen does not support generic segregation. A second character used by Urban to distinguish Pseudopaegma from Anemo- paegma was the presence of subulate calyx teeth in the former but not the latter. However, Sprague & Sandwith (1932) and Sandwith (1955) later undermined this character by describing additional species of Pseudopaegma lacking subulate calyx teeth (cf. Fig. 6). Sandwith (1955) went so far as to note that "the length of subulate calyx teeth is suspect [even] as a specific character." The Venezuelan plant described here further proves this point. It usually has quite distinct calyx teeth 2-3 mm long but these are submarginal and sometimes hardly exceed the calyx margin. A third character separating Pseudopaegma from Anemopaegma was pro- posed by Sandwith (1955) who noted the presence of interpetiolar glandular fields only in the former. Interpetiolar glandular fields are a notoriously variable character, even from node to node on the same plant, and are rarely visible on species of Pseudopaegma with puberulous branchlets. Moreover, these glands are not present in the new species which is definitively linked to Pseudopaegma by its subulate calyx teeth. Apparently the fruits of none of the six species attributed to Pseudopaegma by Sandwith have been described. The fruits of three Pseudopaegma species (P. oligoneuron, P. longidens and the Venezuelan plant) are now at hand and all prove to have the same very characteristic form as those of Anemopaegma. In all other characteristics the two genera are also exactly the same (cf. Fig. 6). There seems to be no basis for continued segregation of Pseudopaegma and I propose that it be reduced to synonymy under Anemopaegma. Four new com- binations are necessary. 1976] GENTRY— SOUTH AMERICAN BIGNONIACEAE 67 Anemopaegma colombianum (Sandw.) A. Gentry, comb, no v. Pseudopaegmu colombianum Sandw., Kew Bull. 1953: 473. 1954. type: Colombia, Meta, Cuatrecasas 4605 (COL, K, US). Anemopaegma insculptum (Sandw.) A. Gentry, comb. nov. Pseuclopaegma insculptum Sandw., Kew Bull. 1954: 608. 1955. type: Colombia, Amazonas, Garcia-Barriga 14617 (COL, K, US). Anemopaegma oligoneuron ( Sprague & Sandw. ) A. Gentry, comb. nov. — Fig. 6. Pseuclopaegma oligoneuron Sprague & Sandw., Kew Bull. 1932: 88. 1932. type: British Guiana, Upper Demerara River, Jenman 4070 (K). Anemopaegma mirabile (Sandw.) A. Gentry, comb. nov. Pseuclopaegma mirabile Sandw., Kew Bull. 1953: 474. 1954. type: Brazil, Piauhy, Gardner 2679 (BM, K). Anemopaegma alatum A. Gentry, sp. nov. Frutex scandens, ramulis teretibus, sine consociebus glandularum in nodis inter petioles; folia trifoliolata vel bifoliolata cum cirrho trifido, foliolis ovatis, lepidoto-puncticulatis, leviter puberulis; inflorescentia racemosa, axillaris; calyx eampanulatus, 5-dentatus dentibus sub- marginalibus 2-3 mm longis, puberulus; corolla aurata, tubulo-campanulata, tubo glabro; ovarium ovoideum, basiliter contraction; discus pulvinatus; capsula elliptica, complanata, puberula, seminibus suborbiculatis. Liana; branchlets terete, finely striate, puberulous, elenticellate; interpetiolar glandular fields absent; pseudostipules minutely foliaceous, deciduous. Leaves 2-3-foliolate, the tendrils trifid; leaflets ovate, acute, rounded to shallowly cordate at base, 5-8.5 cm long, 3-6.5 cm wide, chartaceous, conspicuously lepidote-punc- tate above and below, puberulous along midvein above and main veins and very sparsely over surface below, the margin sometimes subciliate, drying olive to yellowish olive; petiolules 0.2-1.4 cm long; petioles 2.8-3.8 cm long, densely puberulous. Inflorescence an axillary raceme, the rachis and pedicels puberu- lous, the pedicels ca. 1 cm long. Flowers with the calyx campanulate, conspicu- ously 5-toothed ("5-winged" when fresh), the margin basically truncate, the subulate teeth submarginal, 2-3 mm long, extended as calycine ridges, 7-10 mm long (with teeth), 7-8 mm wide, puberulous and lepidote, also with plate-shaped glands below margin; corolla tubular-campanulate above a narrowly tubular basal portion, 5-6 cm long, 1.2-1.8 cm wide at mouth of tube, the tube 4.5-5 cm long, the lobes 0.7-0.9 cm long, the tube glabrous outside, the lobes ciliate, otherwise glabrous, the tube glabrous within except at and below level of stamen insertion, sometimes with plate-shaped glands below lobes outside; stamens didynamous, the thecae divaricate, 3-4 mm long, the longer filaments 1.9-2 cm long, the shorter filaments 1.3-1.5 cm long, the staminode 3 mm long, insertion 20-23 mm from base of corolla tube; pistil 4.3-4.4 cm long, the ovary ovoid, 2-3 mm long, 1.5 mm wide, papillose-lepidote, long-stipitate, the stipe 1 mm long and 0.7 mm wide, the ovules irregularly 4-seriate (appearing 2-seriate in cross section) in each locule; disc cylindric, 1.5 mm long, 2 mm wide. Capsule elliptic, thin-valved, flattened, acute at both ends, 5.5-7 cm long, 3-4 cm wide, the mid- line not at all raised, puberulous, also with scattered plate-shaped glands, drying tannish, the calyx persistent; seeds thin, suborbicular, the wing surrounding body gg ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 6:1 of seed, 2-2.6 cm long, 2.8-4 cm wide, outer margin of wing hyaline-membrana- ceous, this distinctly demarcated from the brownish basal portion wliich is poorly demarcated from the seed body. Type: Venezuela, sucre: Distrito Sucre, between La Sabana, Los Altos, and La Silleta, towards Zurita, 600 m, corolla yellow and white, 18 Aug. 1973, Steyermark et al. 107753 (MO, holotype; VEN, isotype). Additional collections examined: Venezuela, bolivar: Pequena meseta del Norte de Serrania Cararuban, SE de Canaima, 6° 15', 62°47', 300-600 m, trepadora, petalos amarillos, caliz verde, 5-alado, vista tambien en la sabana, 19 Feb. 1964, Agostini 386 (NY, VEN). Rio Carrao, Alto Caroni, alrededores de Salto Hacho, 6°15'N, 62°51'W, 350 in, Mar. 1954, Cardona 2868 (US). Rio Paragua, Salto de Anraima, 275 m, woody vine, corolla tube yellow, lobes white, common at edge of river in this part, 18 Apr. 1943, Killip 37544 (NY, US), monacjas: Selvas del Rio Sabaneta, cerca del pueblo de Quiriquire, trepadora de flores blanco-eremoso, crece al margen de la selva, 9 Aug. 1955, Lasscr ir Varesclii 4089 (VEN). This plant keys to A. jucundum in Sandwith's key but is most closely related to A. longidens on the basis of its conspicuous rather thick calyx teeth, glabrous corolla tube, and only slightly pubescent leaves. It differs from A. longidens in lacking glandular fields at the nodes, having subfoliaceous pseudostipules, leaf- lets densely punctate-lepidote and puberulous along main veins (and slightly over surface) beneath, slightly shorter calyx teeth (2.5-5 mm long in A. longi- dens), and smaller fruit. In addition its leaves dry olive instead of green and the base of the corolla tube is cylindrical rather than flared out around the ovary as in A. longidens. Anemopaegma alatum is also close to A. magnirei Sandw. of Surinam, the type of which has a shorter (4-6 mm long) calyx similarly pilose pubescent but with a sinuate, scarcely denticulate, margin and lepidote-punctate leaves. That species also differs from A. alatum in the corolla tube densely lepi- dote outside, shorter petiolules (2-4 mm long), and lack of subfoliaceous pseudo- stipules. Treatment of the Venezuelan plant as conspecific with A. magnirei is certainly not possible even though a collection [Cowan 39224 (NY) from British Guiana, determined as A. magnirei by Sandwith] somewhat intermediate be- tween A. magnirei and A. alatum exists. The Cowan collection has a larger ( 10 mm by 8-9 mm) calyx than A. magnirei with minute but definite submarginal denticulations, and the corolla tube is scattered-lepidote outside. It is unlike both A. maguirei and A. alatum, especially in having its leaflets only very slightly (not densely) punctate lepidote beneath, although its short (2-4 mm) petiolules match those of the former; its glabrous (except for a few trichomes near margin) calyx is also unlike both of the other plants. This may represent yet a third spe- cies whose relationship with A. parkeri, another species with glandular-lepidote corolla tube, must be considered. The Venezuelan plant is now represented by 5 different collections, all of which are quite homogeneous (one lacks foliaceous pseudostipules). Several of these were variously determined (with queries) by Sandwith as A. carrerense ( A. karstenii Bur. & K. Schum., Cardona 2868), A. parkeri (Agostini 386), and Pseudopaegma sp. cf. P. oligoneuron (Killip 37544) but the relationships seem instead with A. longidens and A. maguirei. 1976] GENTRY— SOUTH AMERICAN B1GNONIACEAE gQ Anemopaegma patelliforme A. Gentry, sp. nov. Frutex scandens; ranmlis teretibus, sine consociebus glandularum in nodis inter petioles; folia trifoliolata vel bifoliolata cum cirrho, foliolis ovatis, plerumque glabratis, petiolulis lateralibus 0.8-1.2 cm longis, petiolis 4-10 cm longis; inflorescentia racemosa, axillaris brevis; calyx patelliformis, truncatus, pubenilus; corolla aurata, tubulo-campanulata, tul)o lepidoto; ovarium oblongo-ellipticum, lepidotum, basaliter contractum; discus pulvinatus; capsula ignota. Liana; branchlets terete, finely striate, glabrate; interpetiolar glandular fields absent; pseudostipules spathulate-foliaceous, to 5 mm long, 2 mm wide, caducous. Leaves 3-foliolate or 2-foliolate with a simple (?) tendril; leaflets ovate, acute to acuminate, the base rounded to very broadly cuneate, 10-15 cm long, 5-7.5 cm wide, membranaceous to chartaceous, the main veins slightly raised above, promi- nent below, scattered impressed-lepidote, otherwise mostly glabrous, usually with a few minute inconspicuous trichomes scattered along main vein above and be- low, with conspicuous clusters of plate-shaped glands in axils of lateral nerves below, drying olive; lateral petiolules 0.8-1.2 cm long, the terminal petiolule to 3.5 cm long; petioles 4.0-10 cm long, puberulous. Inflorescence a short 4-7- flowered axillary raceme, the pedicels and rachis puberulous. Flowers with the calyx expanded-campanulate, truncate, ca. 3 mm long, 6-8 mm wide, minutely puberulous, with plate-shaped glands in upper half; corolla yellow, tubular- campanulate, 4.3-5.2 cm long, 1.4-1.7 cm wide at mouth of tube, the tube 3.5-4 cm long, the lobes 0.7-1.1 cm long, glandular lepidote outside, the lobes mostly glabrous, with a few trichomes along the subciliate margins, the tube inside mostly glabrous, with some glandular-lepidote scales, densely pubescent at level of stamen insertion; stamens didynamous, the anther thecae divaricate, 3 mm long, the longer filaments 1.8-1.9 cm long, the shorter filaments 1.4-1.6 cm long, the staminode 4-5 mm long, insertion 9-10 mm from base of corolla tube; pistil ca. 2.5 cm long, the ovary oblong-elliptic, 2 mm long, 1 mm wide, contracted slightly at base, densely glandular-lepidote, the ovules 4-seriate in each locule in cross section; disc pulvinate, tapering to base of ovary, ca. 1 mm long, ca. 2 mm wide. Capsule unknown. Type: Venezuela, amazoxas: Mavaca, Alto Orinoco, Indios Guaicas (Yano- mano), las flores son utilizadas para adorno en las orejas perforadas de las mujeres, trepadora de flores amarillas, Jan. 1970, Aristeguieta dr Lizot 7374 (VEN, holotype; MO, fragments). Vegetatively this plant is similar to A. longipetiolatum Sprague of Paraguay because of its long petioles. That species is very different in puberulous leaves, much larger narrowly campanulate calyx, and longer inflorescence. The calyx llif f ( Schlecht. ) Sandw. and matched in Anemopaegma only by very different pubes- cent-leaved A. oligoneuron (Sprague & Sandw.) A. Gentry and by A. insculptum (Sandw.) A. Gentry. patellif lacks lepidote scales on the outside of the corolla as well as having shorter (3-5 cm long) petioles and longer (1.5-2 cm long) lateral petiolules. Anemopaegma insculptum also differs in having interpetiolar glandular fields and lacking glandular fields in the axils of the lateral nerves below. 7Q ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Anemopaegma villosum A. Gentry, sp. nov. Frutex scandens; raniuli teretes, villosi, sine consociebus glandularum in nodis inter petioles; pseudostipulis foliaceis; folia 2-foliolata, foliolis ellipticis, acutis, subtus villosis; in- florescentia floribus 1-3 in axillis foliorum dispositis; calyx cupulatus, fere membranceous, trun- catus; corolla tubulo-campanulata, tiibo extus lepidoto; stamina thecis divaricatis; ovarium ellipsoideum, lepidotum; discus pulvinatus; capsula ignota. Vine; branchlets terete, finely striate, villous; interpetiolar glandular fields absent; pseudostipules thinly foliaceous, elliptic, acute, 5-15 mm long, 2-8 mm wide, noticeably veined. Leaves 2-foliolate, often with a minutely trifid tendril; leaflets ovate, acute to short-acuminate, rounded to truncate at base but often abruptly attenuate at top of petiole, 6.5-9 cm long, 3.5-5.5 cm wide, membrana- ceous, pubescent with scattered trichomes above, villous beneath, especially along main veins, the trichomes to almost 1 mm long, drying olive; petiolules 1.5 cm long; petioles 2 cm long, villous. Inflorescence 1-3 axillary or terminal flowers, the pedicels pilose, 4-7 mm long. Flowers with the calyx cupular, almost mem- branaceous, truncate, 5-6 mm long, 5 mm wide, appressed-pilose, eglandular; corolla "y e " ow with white," tubular-campanulate, 4.5 cm long, 1.4-2 cm wide at mouth of tube, the tube 4 cm long, the lobes ca. 0.7 cm long, conspicuously glandular-lepidote outside and on lobes and upper part of tube inside, sparsely pubescent at level of stamen insertion; stamens didynamous, inserted 6-7 mm above base of corolla tube, the filaments 1.5-2 cm long, the anther thecae divari- cate, 3 mm long; ovary ellipsoid, slightly narrowed at base, 2 mm long, 1 mm wide, densely minutely lepidote; disc conical pulvinate, 1 mm long, 1.5 mm wide. Capsule not known. Type: Venezuela, portuguesa: Selva tropofila sobre penasco calcareo, 5 km este-noreste de Agua Blanca 22 km noreste de Acarigua, 190 m, twining around shrubs on bluffs, corolla yellow with white, leaves membranaceous, gray green below, 24 Aug. 1966, Steyermark 6 Rabe 96455 (VEN, holotype; NY, US, iso- types ) . This species is most closely related to A. rugosum (Schlecht. ) Spr. which dif- fers in conspicuously bullate leaflets, obtuse pseudostipules, a more coriaceous calyx, and especially a pilose indumentum of stiff erect trichomes on leaves and branchlets. Anemopaegma rugosum is known only from altitudes of 600-1,300 m in Distrito Federal and Aragua State. Sanhilaria & Paragonia Paragonia brasiliensis (H. Baill.) A. Gentry, comb. nov. SanJiilaria brasiliensis H. Baill., Hist. Pi. 10: 27. 1888 (1891). Examination of the type specimen of Baillon's monotypic genus indicates that it is very similar to Paragonia pyramidata (L. Rich.) Bur., the only species of Paragonia. There can be no doubt that Sanhilaria and Paragonia are synonymous. The question which must be resolved is whether Sanhilaria brasiliensis is even specifically distinct from P. pyramidata. 1976] GENTRY— SOUTH AMKHICAN BIGNONIACEAE ^\ The type collection of S. brasiliensis — St. Hilaire 745 (P) from Minas Gerais, Brazil — has all essential characters of Paragonia, including the characteristic subulate pseudostipules. It differs from P. pyramidata in its softly puberulous short-petioled leaves, narrower inflorescence, and especially in the conspicuously costate almost winged calyx; no corollas are extant. A second sheet of the same collection has an immature fruit which is more compressed than in P. pyramidata and lacks the characteristic sandpaperlike surface of that species. The fruit of Sanhilaria is densely lepidote. Another collection of Sanhilaria is now at hand (Heringer 10277, Jequie, Bahia, Brazil (NY) ) and has corollas exactly like those of P. pyramidata except for the acute corolla lobes. This collection has glabrous leaflets but the leaves are otherwise similar to those of the type in their small size, narrowly elliptic shape, and very short petioles and petiolules. A third col- lection of Sanhilaria, also from Bahia (Serra da Agua de Rega, 28 km N of Seabra, 1000 m, Irwin et al. 31159 (MO) ), was originally misidentified by me as Adenocalymma sp. This collection is in young fruit and the fruits are densely lepidote, smooth-surfaced, and strongly compressed. The characteristic costate calyx is persistent on one of them. The leaflets of the Irwin et al. collection are glabrous (except for lepidote scales) as in the Heringer collection and have the characteristic narrowly elliptic shape and blunt apex of Sanhilaria. As in the other two Sanhilaria collections, the petioles and petiolules are extremely short to essentially nonexistent. Patagonia pyramidata also occurs in Brazil ranging southward to Rio Grande do Sul. However, I have seen no collection of P. pyramidata which approaches Sanhilaria in leaf form, the species differing constantly in having much longer petioles and petiolules and a wider ovate or elliptic-ovate leaflet. Paragonia pyramidata is known from Goias and the Distrito Federal where it typically has its leaves softly puberulous below as in the type of Sanhilaria. I regard such variation in pubescence as taxonomically unimportant. Collections of P. pyra- midata from Brazil all have the convex sandpaper-surfaced capsule valves, broadly paniculate inflorescences, and ecostate calyces characteristic of the species. The evidence supports recognition of Sanhilaria brasiliensis as a distinct spe- cies and the new combination Paragonia brasiliensis (Baill. ) A. Gentry is necessary. two Table 1. Nestoria. Kuiilmannia & Pleonotoma Pleonotoma albiflora (Salzm. ex DC.) A. Gentry, comb. nov. Bignonia albiflora Salzm. ex DC, Prodr. 9: 167. 1845. type: Brazil, Bahia, Salzmann 346 (G-DC, P). Memora albiflora (Salzm. ex DC.) Miers, Proc. Roy. Ilort. Soc. London 3: 185. 1863. M. obtusifoliolata Bur. & K. Schum. in Mart., Fl. Bras. 8(2): 261. 1896. type: Brazil, Bahia, Riedel 750 (K, P). Nestoria obtusifoliolata (Bur. & K. Schum.) Urb., Ber. Deutsch. Bot. Ges. 34: 752. 1916. N. albiflora (Salzm. ex DC.) Sandw., Candollea 7: 248. 1936. Kuhlmannia colatinensis J. C. Gomes, Arq. Serv. Florest. 10: 201, fig. 1. 1956; Notul. Syst. (Paris) 15: 224, fig. 2. 1956. type: Brazil, Espirito Santo, Kuhlmann 6567 (RB, MO). 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Table 1. Contrasting features of Paramimia pyramidata and P. brasiliensis. P. pyramidata Tendril tip minutely bifid (rarely trifid) Petioles and petiolules well developed Leaflets elliptic or ovate-elliptic, the apex obtuse to acuminate Calyx ecostate Inflorescence broadly paniculate Capsule subterete, sandpaper-surfaced Corolla lobes rounded Mexico to southern Brazil P. brasiliensis Tendril tip minutely trifid Petioles and petiolules reduced, subobsolescent Leaflets narrowly elliptic to oblanceolate, the apex obtuse Calyx conspicuously ridged (cf. Fridericia) Inflorescence racemose-paniculate Capsule strongly compressed, smooth- surfaced, densely lepidote (immature) Corolla lobes acute Eastern Brazil (Bahia and Minas Gerais) The monotypic genus Nestoria was segregated from Memora by Urban ( 1916) on the basis of its trifid tendril and tricolpate pollen. Bureau & Schumann (1896- 1897) had earlier suggested that its single species seemed out of place in Memora because of its Anemopaegma-\ike flower and open inflorescence. Sandwith (1936) pointed out that the pollen and tendril characters used by Urban to segregate Nestoria were just those of Pleonotoma which likewise has biternate leaves and similar white or cream corollas. However, he retained Nestoria as rather doubt- fully distinct from Pleonotoma on the basis of its many-ribbed rather than tetrag- onal branchlets and its "remarkable inflorescence." These characters are insuffi- cient for generic segregation in my opinion. The Salzmann type collection of Nestoria albiflora does have conspicuously tetragonal branchlets with the four angles ribbed and contrastingly lighter colored than the rest of the branchlet. This tetragonal branchlet is readily visible even in the Field Museum photograph (negative 7674) of the type and is just that of Pleonotoma. The Salzmann collection is obviously very close to Pleonotoma. On the other hand the Riedel type collection of Memora obtusifoliolata has mostly very inconspicuously, or not at all tetragonal, 6-8-ribbed branchlets. Only above the nodes are the branchlets of this collection clearly tetragonal. The calyces of this collection are also somewhat shorter than those of the Salzmann collection and the leaflets are round-tipped. Nevertheless, the two collections are so similar as to leave little doubt that they are conspecific. Now we must remember that neither Schumann (collaboration with Bureau being purely nominal) nor Urban had seen the Salzmann collection. Memora obtusifoliolata was described from the Riedel collection as having terete branch- lets, thus seeming to accord with Memora in this important aspect. When Urban surveyed pollen and tendrils of the Bignoniaceae, he discovered that M. obtusi- foliolata differed from other members of Memora as noted previously and con- sequently erected the monotypic Nestoria for it. We may presume that had these 1976] GENTRY— SOUTH AMERICAN B1GNONIACEAE 73 authors seen the more tetragonal-stemmed Salzmann collection they might have recognized its affinity with Pleonotoma, especially as that genus has both the trifid tendrils and 3-colpate pollen which led to Urban's rejection of the species from Memora. Meanwhile Miers, who had seen the Salzmann collection, had transferred Bignonia albiflora to Memora rather than to Pleonotoma despite the tetragonal branchlets. This is hardly surprising when one remembers that Memora and Pleonotoma were defined very differently by Miers than by subsequent authors with foliaceous pseudostipules as the major diagnostic character. Species with terete stems and simple tendrils were thus lumped with unrelated species having tetragonal branchlets and trifid tendrils. We have, then, a species known from two collections, one with quite obviously tetragonal branchlets, the other with 6-8-ribbed subtetragonal or subterete branchlets. We also know that several accepted species of Pleonotoma may have inconspicuously tetragonal and even multi-ribbed (though with four ribs more conspicuous) older branchlets. How fundamental is the difference between tetragonal and multi-ribbed branchlets? Evidence from other genera indicates that in several species of Bignoniaceae normally having tetragonal, 4-ribbed branchlets [e.g., Mussatia hyacinthina (Standi.) Sandw., Cydisia diversifolia (II.B.K.) Miers], occasional plants or branches of plants may be perfectly 6- angled and 6-ribbed. The change from 4-angled to 6-angled may be reversed or repeated even on a single branch. Whereas ribbed and angled branchlets may differ fundamentally from unribbed terete ones, the number of ribs is less im- portant, even in species delimitation. The ribbed branchlets of Nestoria ally it with Pleonotoma. The inflorescence of Nestoria is a very lax open raceme while the typical in- florescence of Pleonotoma is a relatively few-flowered and contracted raceme with a different facies. However, such species of Pleonotoma as P. clematis (II.B.K.) Miers, P. melioides (S. Moore) A. Gentry, or P. jasminifolia (H.B.K.) Miers may have inflorescences ranging in size from a few flowers to an elongate 18-or-more-flowered raceme with successive flower pairs separated by 3 cm and pedicels at least 2.5 cm long. The inflorescences of these species have exactly the same appearance as that of Nestoria. Additional evidence also supports the affinity of Nestoria with Pleonotoma. While the tetragonal branchlets of Pleonotoma have received all the taxonomic attention, the petioles and petiolules of Pleonotoma are also characteristically tetragonal with raised angles. The petiolules (and sometimes the petioles) of Nestoria, even in specimens with subterete branchlets, are sharply tetragonal with raised angles just as in Pleonotoma. Another kind of evidence comes from an unexpected source: examination of type material of Kuhlmannia colatinensis proves that monotypic genus synony- mous with N. albiflora. The Kuhlmannia type includes the first known fruit of Nestoria and this fruit is also similar to that of some species of Pleonotoma. All available evidence (trifid tendril, 3-colpate microreticulate pollen, biter- nate leaves, inflorescence, flowers, ribbed subtetragonal branchlets, fruit) sup- ports merger of Nestoria (and Kuhlmannia) with Pleonotoma. The single new 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 combination Pleonotoma albiflora (Salzm. ex DC.) A. Gentry reduces two un- necessary and monotypic genera to synonymy. This rare species has been assumed endemic to eastern Brazil (Bahia and Espirito Santo) so the discovery of a disjunct population in lowland Guyana is entirely unexpected. Nevertheless, I am unable to distinguish from P. albiflora a suite of specimens from Venezuela, Surinam, French Guiana, and Amapa, Brazil. Amazingly, the collection from French Guiana was made by Aublet long before the first Brazilian collections. Guyana area collections I have examined are: Venezuela, delta amacuro: Rio Grande cerca de los limites del Estado Bolivar, Blanco 498 (VEN). Surinam: Mapanegebied, Kamp 8, Vreden 11320 (WAG). French cuiana: without locality, Aublet s.n. (BM). Brazil. Rio Jari (border of Amapa and Para), Agua Branca, Silva 2459 (IAN). A sterile collection from Amazonas, Brazil [Gentry ir Ramos 13333, km 120, Manaus-Itacoatiara Road (MO, INPA)] may be a juvenile form of this species. Two of these collections are in fruit, permitting amplification of Gomes's fruit description. The fruit of P. albiflora is linear-oblong, flattened but with rather woody valves, glabrous, 35-51 cm long, 2.1-3 cm wide, acute at base and apex, drying light green. The seeds are very thin, bialate, ca. 1.8 cm long and 7.5 cm wide, drying uniformly brownish except for very narrow hyaline wing tips. The seeds are uniseriate as in Memora but unlike other species of Pleonotoma. This difference is inadequate for generic segregation of Nestoria. XEROTECOMA & GODMANIA Godmania dardanoi (J. C. Gomes) A. Gentry, comb. nov. Xcrotccoma dardanoi J. C. Gomes, Rev. Bras. Biol. 24: 405. 1965. type: Brazil, Pernanihueo, Lima 61-3598 (RB). Gomes compared his monotypic new genus with Tabebuia (Tecoma to him) because of its arboreous habit, 3-colporate reticulate pollen grains, and palmately compound leaves. He separated it from Tabebuia because of its striate, spirally twisted capsule, campanulate-gibbous corolla, barbate anthers, and patelliform calyx. Unfortunately, Gomes overlooked the fact that these are the same charac- ters by which Godmania is distinguished from Tabebuia. Gomes's illustration of X. dardanoi also shows such GodmaniaAike features as attenuate leaflets, curved ery short anther thecae, and a seed with an indetermin wing Through the courtesy of the Jardim Botanico of Rio de Janeiro, I was able to examine the type of Gomes's plant during a recent visit to Rio de Janeiro. It proves to match Godmania in all essentials. In fact, it is so close to G. aesculifolia (H.B.K.) Standi., the only recognized species of the genus, that its specific separation needs to be justified. Godinania aesculifolia ranges from Mexico to Venezuela and Bolivia and even into extreme northern Brazil (Roraima, Para) but has not been collected south of the Amazon in Brazil. It is thus disjunct from G. dardanoi which occurs in the caatinga of eastern Brazil. The corolla of G. aescidifolia is considerably smaller (1.0-1.6 cm long) than that of the caatinga plant (2.5-3.0 cm long) 1976] GENTRY— SOUTH AMERICAN BIGNONTACEAE 75 though of the same peculiar shortly and widely campanulate form and with the same unusual short triangular lobes. The 15-38 cm long fruit of the caatinga plant is shorter than the 45-100 cm long fruit of G. aesculifolia and the pubes- cence is of a different consistency. These differences are adequate to maintain the plant of eastern Brazil as specifically distinct and the new combination God- mania dardanoi (J. C. Gomes) A. Gentry is in order. I have seen two additional specimens of G. dardanoi besides the type and the two other collections from Pernambuco cited by Gomes. One of these was col- lected at Santa Elena in Bahia State (Zehntner 378, RB 6383); the second (P. de Campo Porto s.n., RB 29630) is without data. Interestingly enough these two col- lections have been annotated by Kuhlmann with an unpublished manuscript name as a new species of Godmania. ROSEODENDRON & TABEBUIA Tabebuia millsii (Miranda) A. Gentry, comb. nov. Cybistax millsii Miranda, Bol. Soc. Bot. Mexico 26: 129. 1961. type: Mexico, Chiapas, Gomez- Pompa 312 (US). Roseodendron millsii (Miranda) Miranda, Bol. Soc. Bot. Mexico 29: 43. 1965. A reconsideration of the relationships of Roseodendron with Tabeuia makes the former's reduction to Tabebuia seem advisable, contrary to my former opinion (Gentry, 1970). The two species of this group known to Seibert were placed by him in Cybistax (Seibert 1940a, 1940b). Both had been described in Tabebuia. They were distinguished from Tabebuia by the striate ovary, costate fruit, thin membranaceous calyx, and capitate inflorescence trichomes. These characters were thought to be closer to those of Cybistax, a monotypic genus of Central Brazil and adjacent Peru. Miranda subsequently described a third species of the alliance in Cybistax as C. millsii. Later (Miranda, 1965) he correctly noted that the relationship of these species with Cybistax is only superficial. He erected for them the new genus Roseodendron, separated from Cybistax by the more deli- cately membranaceous, narrower, ecostate calyx, narrowly oblong, estipitate ovary, and indumentum of branched trichomes and lepidote scales with raised borders. He might also have mentioned Roseodendron \s much narrower linear fruit with relatively inconspicuous ribs, very different from the oblong fruit with narrow, raised, 1-5 mm thick ribs of Cybistax. Clearly Roseodendron is much closer to Tabebuia than to Cybistax. Indeed all of the characters (except calyx texture) noted by Miranda as distinguishing Roseodendron from Cybistax are also characteristic of Tabebuia. Discovery of the fruits of Tabebuia guayacan (Seem.) Hemsl. and T. capitata (Bur. & K. Schum.) Sandw. [especially a form of the latter from Peru, Castillo 37-MCS (WIS)] lessens the importance of the capsule ribs of Roseodendron as a generic character. Tabebuia capitata has varyingly striate fruits as does T. heptaphylla (Veil.) Toledo (see Gentry, 1975), while T. guayacan has irregularly muricate-reticulate or interruptedly costate fruits. Indeed, these fruits can be more costate than that of Roseodendron chrysea (i.e., Tabebuia chrysea Blake). Capitate trichomes are a notoriously variable characteristic in Bignoniaceae 76 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 and many natural genera (e.g., Arrabidaea, Lundia, Mansoa, Piriadacus) have species both with and without capitate trichomes. Their presence in Roseoden- dron does not merit its generic segregation. The thinner calyx of Roseodendron is less different from that of Tabebuia in fresh material than in the herbarium. In any case it too differs only in degree from the relatively large thin calyces of such Tabebuia species as T. elliptica (DC.) Sandw. and wet-forest forms of T. chrysantha (Jacq.) Nichols. The well-developed central axis of the inflorescence of Roseodendron is, in my opinion, a more significant difference from Tabebuia than any of those mentioned by Seibert (1940a) but still insufficient for generic segregation. In the field Roseodendron has exactly the appearance of Tabebuia in flowering, fruiting, or vegetative condition and is routinely recognized by col- lectors as Tabebuia. Even the colloquial names for species of Roseodendron are generically the same as those of Tabebuia, e.g., in Mexico "macuelis de cerro" for R. mills ii and "macuelis de bajo" for T. rosea (Bertol.) DC. (fide Miranda, 1961). I now consider Roseodendron best treated under Tabebuia. Tabebuia millsii is the only new combination needed. Tabebuia millsii presents another problem. The species has been supposed to be restricted to the Caribbean side of southern Mexico from Veracruz to Chiapas. However, recent collections from central Bolivar State in Venezuela seem indis- tinguishable from T. millsii. The Venezuelan plant agrees with T. millsii, rather than T. donnell-smithii of Pacific Mexico to El Salvador, in (mostly) 5-foliolate leaves, entire leaflet margins, dense simple pubescence of leaflet undersides, somewhat flattened reticulately veined as well as longitudinally costate fruit, and dimensions of the mature leaf (leaflets to 17 X 7 cm and terminal petiolule 3 cm long; the same dimensions for T. millsii are 13 X 6.5 cm and 3 cm, for T. donnell- smithii 25 X 14 cm and 7 cm). In some respects the Venezuelan population is intermediate between T. millsii and T. donnell-smithii. Its fruit has 8-12 major ribs (compared to 8 in T. millsii and 10-12 in T. donnell-smithii) . Juvenile leaves of the Venezuelan plant (Mar- cano-Berti 2560 (VEN), Gentry it Berry 15060 (MO, VEN)) tend to be 7-f olio- late and the leaflets are much larger than mature leaflets and usually somewhat serrate; these juvenile leaves thus approximate the mature leaves of T. donnell- smithii (juvenile leaves of T. millsii from Mexico are not known). The wood I have seen [Gentrti ir Berrtt 15071 (MO)] is yellow and thus similar to the clear yellow wood of T. donnell-smithii rather than to the heavy tough dark wood described by Miranda (possibly in confusion with T. chrysantha or T. guayacan?) for T. millsii in Mexico. However, lumbermen around La Paragua at the eastern edge of the range of the Venezuelan plant claim that trees from further west have a much darker wood. Despite its long disfunction, I cannot separate the Venezuelan population from the Mexican one. The only consistent difference between the Venezuelan and Mexican populations of T. millsii is in their fruits (based on a single fruiting col- lection of each). The Venezuelan fruits are only 1.4-2.1 cm wide while those of Mexican T. millsii are 2.1-3 cm wide and those of T. donnell-smithii are (fide Seibert, 1940a) 2-3 cm wide. The Venezuelan fruits have mostly simple eglandu- lar trichomes while those of T. millsii have gland-tipped and dendroid trichomes. 1976] GENTRY— SOUTH AMERICAN BIGNONIACEAE 77 However, the Venezuelan fruits do have occasional branched and gland-tipped trichomes and the difference in capsule width does not warrant a taxonomic dis- tinction. Long range disjunctions are not uncommon in Bignoniaceae (although often collecting artifacts) but in no other case known to me does a disjunction involve such limited areas of distribution at both ends. Tabebuia donnell-srnithii is introduced in Ecuador and one wonders if a similar introduction might have taken place in Venezuela. However, the Venezuelan tree, known locally as "eacho de venado," is certainly native and is in fact one of the three major timber trees of the La Paragua area (along with "cedro" (Cedrella) and "laurel" (Cordia alliodora ) ) . It seems probable that the morphologically somewhat intermediate Venezue- lan population represents the ancestral form for both Mexican plants with dif- ferentiation of T. donnell-smithii in western Mexico and minor changes in T. millsii in eastern Mexico occurring subsequent to disruption of a once more continuously distributed ancestral stock. DiSTICTIS & DlSTICTELLA Distictis pulverulenta (Sandw.) A. Gentry, comb. nov. Distictella pulverulenta Sandw., Brittonia 3: 91. 1938. type: Brazil, Amazonas, Krukoff 8685 (BM, BR, GH, F, K, MO, NY, U). My recent treatment of Distictis (Gentry, 1974d) concluded that several closely related genera (Phaedranthus, Anomoctenium, W unschmannia) should not be segregated from it. One other genus also needs to be considered in relation to Distictis. This is Distictella, a segregate which agrees with Distictis in large- meshed (alveolate) acolpate pollen, trifid tendrils, phloem 4-anned in stem cross section, a more or less racemose, usually bracteate inflorescence with large tubu- lar flowers, campanulate, truncate cupular calyx, conspicuous disc, and flattened ovoid woody fruit. Bureau ( 1864 ) was the first to propose the separation of Distictis ( as Macro- discus) from Distictella (as Distictis). Distictella (his Distictis) was charac- terized by the ovaiy containing a number of series of ovules in each locule and surmounting a cylindrical disc, an oblong, tomentose, flattened capsule with two woody curved valves (one concave, the other convex), the tip of the replum at- tached to the septum which bears numerous series of linear seed scars along each border, and pubescent seeds. Distictis (his Macrodiscus) was said to differ from the preceding by the glabrous uncurved capsule with a furrow in place of a rib on each valve, free replum ends extended beyond the tip of the septum, glabrous seeds, and a punctiform hilum; all these characters are those of the fruit. Urban (1916) later pointed out that Bureau had misapplied Distictis and that Macro- discus was actually Distictis while Distictis sensu Bureau must be known as Distictella. Schumann (1894) introduced new characters in separating Distictis (as Macrodiscus) from Distictella (as Distictis) on account of its thinner corolla without a conspicuous bend in it. He also noted that Distictella differs from Pithecoctenium in having terete twigs lacking the detachable ribs of the latter' s 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 63 Table 2. Real and supposed differences between Distictis and Distictella. Distictis Capsule glabrous Capsule not curved Capsule midline impressed Ends of replum free and extended beyond end of septum Seed scars punetifonn Seeds glabrous, with relatively short: wings Corolla thinner in texture Corolla straight or slightly curved Branchlets 6-angled and ribbed Distictella Capsule pubescent Capsule curved Capsule midline raised (implied) Ends of replum attached to end of septum Seed scars linear Seeds pubescent with longer wings Corolla thicker in texture Corolla strongly bent Branchlets terete, not ribbed 6-angled twigs. Melchior (1927) did not consider Distictis in his synopsis of subtribe Pithecoctineae but considered this character of taxonomic importance, separating Distictella from Neves-Armondia and Pithecoctenium on the basis of round rather than 6-angled branchlets. Sprague & Sandwith (1932) further emphasized the character of absence of detachable ribs in Distictella. Although considerations of terete versus angled and ribbed branchlets were intended to apply to Distictella as compared with Pithecoctenium, they are equally valid to separate it from Distictis. The differences that have been proposed to separate Distictis (sensu stricto) and Distictella are listed in Table 2. A number of new species have been discovered in both genera since these generic differences were outlined by Bureau (1864) and Schumann (1894) and the fruit differences prove not to be correlated with the floral and vegetative characters. For example Distictis gnaphalantha (A. Rich.) Urb. has pubescent fruits but is certainly congeneric with Distictis lactiflora (Vahl) DC, the type species of Distictis. Distictella obovata Sandw. and D. monophylla Sandw. have the flowers and twigs of Distictella but the fruit of Distictis (fruits witl arcely or not at all evident midrib, acuminate, the valves not curved, seeds glabrous). My (Gentry, 1974d) reduction of Anomoctenium — which agrees with Distictis in uncurved fruit, 6-angled branchlets, and nearly straight corolla — to Distictis has expanded that genus to include species with pubescent seeds. In general the characters of the fruit elucidated by Bureau lack taxonomic significance at the generic level. Seed scars vary considerably in shape even within a single species and the "punctiform" seed scars of D. lactiflora are not fundamentally different from the "linear" seed scars of Distictella; indeed Bureau's own illustration of Distictis (as Macrodiscus) showed elongate rather than punc- tate seed scars. The ends of the replum are extended beyond the end of the septum in species of both genera which have acuminate fruits but not in those species with apically blunt fruits. A tendency to seed pubescence is found in 1976] GENTRY— SOUTH AMERICAN BIGNONIACEAE 79 several species of Distictis while some species of Distictella have glabrous seeds. The capsule midline may be raised or not and the capsule curved or not in Distictella; the capsule is usually pubescent in Distictis as well as Distictella. We are left with only vegetative and floral characters separating the two genera. With the exception of a single species these characters appear to be constant and correlated. The flowers of Distictella (with one exception) match the syndrome of xylocopid (or Pithecoctenium-type) flowers (Gentry, 1974b) in having a thickened corolla which is strongly bent near the middle. With a single exception, these flowers are white, usually with a yellow throat. The flowers of Distictis are some- what thinner in texture and more or less straight Most are of the " Anemopaegma- type" and are adapted for generalized bee pollination. These species have corollas ranging in color from cream to lavender or magenta. Two species of Distictis have the corolla deeper red, more elongate, and stamens exserted or subexserted; these are adapted to hummingbird pollination. All species of Distictis have 6- angled branchlets with the angles ribbed; all species of Distictella (with one exception) have terete unribbed branchlets. Otherwise there appears to be no vegetative difference. There are several exceptions to the characters noted above for Distictella. Happily enough a single species is responsible for all the exceptions: Distictella pulverulenta Sandw. has ribbed 6-sided branchlets, a straight corolla, and flowers described as rich purple. Distictella pulvertilenta is actually a Distictis and the new combination Distictis pulverulenta (Sandw.) A. Gentry is necessary. Distictis pulverulenta becomes the tenth species of Distictis (see Gentry, 1974d). Should the rest of the species of Distictella remain segregated from Distictis: The removal of D. pulverulenta leaves Distictella as a homogeneous and well- circumscribed group. While its nearest relationships are clearly with Distictis, it is convenient, at least provisionally, to accept the now traditional separation of Distictella while noting that the fruit characters on which this segregation was originally based are of minimal value. r> Literature Cued Bureau, E. 1864. Monographic des Bignoniacees. Bailliere et fils, Paris. & K. Schumann. 1896-1897. Bignoniaceae. In C. F. P. Martins (editor), Flora Brasiliensis 8(2): 1-452. Duc;and, A. 1946. Noticias Botanicas Colombianas VI. Caldasia 4: 51-65. Gentry, A. H. 1970. 264. America. Brittonia 242. 1973. Generic delimitations of Central American Bignoniaceae. Brittonia 25: 226- 1974a (1973). Bignoniaceae. In R. E. W