of the

Biological Society of Washington

VOLUME 108 1995

Vol. 108(1) published 24 March 1995 Vol. 108(3) published 19 September 1995

Vol. 108(2) published 22 June 1995 Vol. 108(4) published 28 December 1995


EDITOR C. Brian Robbins


Classical Languages Invertebrates

George C. Steyskal Jon L. Norenburg

Frank D. Ferrari Rafael Lemaitre

Plants Vertebrates

David B. Lellinger Thomas A. Munroe

Insects Wayne N. Mathis

All correspondence should be addressed to the

Biological Society of Washington, Smithsonian Institution

Washington, D.C. 20560

Allen Press Inc. Lawrence, Kansas 66044


of the


FOR 1994-1995


President JANET W. REID

President-Elect STEPHEN D. CAIRNS


Treasurer T. CHAD WALTER






Volume 108

Albert, James Sperling and Robert Rush Miller. Gymnotus maculosus, a new species of electric fish (Chordata: Teleostei: Gymnotoidei) from Middle America, with a key to species of Gymnotus

Bayer, Frederick M. A new species of the gorgonacean genus Narella (Anthozoa: Octocorallia) from Hawaiian waters

Brown, Rafe M., John W. Femer, and Luis A. Ruedas. A new species of lygosomine

lizard (Reptilia: Lacertilia: Scincidae: Sphenomorphus) from Mt. Isarog, Luzon Island,


Brown, Rafe M., John W. Ferner, and Rogelio V. Sison. Rediscovery and redescription

of Sphenomorphus beyeri Taylor (Reptilia: Lacertilia: Scincidae) from the Zambales

Mountains of Luzon, Philippines

Brown, Walter C. A new lizard of the genus Sphenomorphus (Reptilia: Scincidae) from

Mt. Kitanglad, Mindanao, Philippine Islands

Brown, Walter C. and Ely L. Alcala. A new species of Brachymeles (Reptilia: Scincidae)

from Catanduanes Island, Philippines

Bruce, Niel L. and J0rgen Olesen. Natatolana nukumbutho, a new species (Crustacea:

Isopoda: Cirolanidae) from deep water off Suva, Fiji

Cairns, Stephen D. New records of azooxanthellate stony corals (Cnidaria: Scleractinia

and Stylasteridae) from the Neogene of Panama and Costa Rica

Campos, Martha R. A new species of freshwater crab of the genus Strengeriana from

Colombia (Crustacea: Decapoda: Pseudothelphusidae)

Campos, Martha R. and Gilberto Rodriguez. Two new species of freshwater crabs of

the genus Hypolobocera from Colombia (Crustacea: Decapoda: Pseudothelphusidae)

Campos-da-Paz, Ricardo. Revision of the South American freshwater fish genus Stemarchorhamphus Eigenmann, 1905 (Ostariophysi: Gymnotiformes: Apteronotidae), with notes on its relationships

Carleton, Michael D. and Guy G. Musser. Systematic studies of oryzomyine rodents (Muridae: Sigmodontinae): definition and distribution of Oligoryzomys vegetus (Bangs, 1902)

Couri, Marcia Souto. Contribution to the knowledge of Reynoldsia Malloch (Diptera: Muscidae)-

Cuatrecasas, Jose. A new genus of the Compositae: Paramiflos (Espeletiinae) from Colombia

Cumberlidge, Neil. Remarks on the taxonomy of Sudanonautes chavanesii (A. Milne- Edwards, 1886) (Brachyura: Potamoidea: Potamonautidae) from Central Africa

Cumberlidge, Neil. Redescription of Sudanonautes faradjensis (Rathbun, 1921), a fresh- water crab from Central Africa (Brachyura: Potamoidea: Potamonautidae)

Dean, Harlan K. A new species of Raricirrus (Polychaeta: Ctenodrilidae) from wood collected in the Tongue of the Ocean, Virgin Islands

Erseus, Christer and Olav Giere. Otavius nicolae, a new gutless marine tubificid species (Oligochaeta) from Belize

Felder, Darryl L. and Raymond B. Manning. Neocallichirus cacahuate, a new species of ghost shrimp from the Atlantic coast of Florida, with reexamination of N. grandimana and N. lemaitrei (Crustacea: Decapoda: Callianassidae)

Ferrari, Frank D. Six copepodid stages of Ridgewayia klausruetzleri, a new species of copepod crustacean (Ridgewayiidae: Calanoida) from the barrier reef in Belize, with comments on appendage development

Goldman, David A. A juvenile of the scaled squid, Pholidoteuthis adami Voss, 1956 (Cephalopoda: Oegopsida), from the Florida Keys

Harold, Antony S. and Richard Winterbottom. Gobiodon acicularis, a new species of gobioid fish (Teleostei: Gobiidae) from Belau, Micronesia

662-678 147-152


6-17 388-391 392-394 212-219 533-550 98-101



338-369 281-291 748-750 238-246 629-636 169-179 491^95


180-200 136-146 687-694

Heyer, W. Ronald. South American rocky habitat Leptodactylus (Amphibia: Anura:

LeptodactyUdae) with description of two new species 695-716

Hobbs, Horton H., Jr. and H. H. Hobbs III. Procambarus (Ortmannicus) nueces

(Decapoda: Cambaridae), a new crayfish from the Nueces River Basin, Texas 54-60

Hobbs III, H. H. and Horton H. Hobbs, Jr. Macrobrachium catonium, a new troglobitic

shrimp from the Cayo District of Belize (Crustacea: Decapoda: Palaemonidae) 50-53

Hotchkiss, Frederick H. C. Loven's law and adult ray homologies in echinoids,

ophiuroids, edrioasteroids, and an ophiocistioid (Echinodermata: Eleutherozoa) 401^35

Hsieh, Hwey-Lian. Laonome albicingillum, a new fan worm species (Polychaeta:

Sabellidae: Sabellinae) from Taiwan 130-135

Jimenez, Maria, G. San Martin, and E. Lopez. Pionosyllis maxima Monro, 1930, P.

anops Hartman, 1953, and P. epipharynx Hartman, 1953, redescribed as Eusyllis

maxima (Monro, 1930), a new combination (Polychaeta: Syllidae: Eusyllinae) 496-501

Komai, Tomoyuki. A new species of the genus Discorsopagurus (Crustacea: Decapoda:

Paguridae) from Japan, previously known as D. schmitti (Stevens) 617-628

Komicker, Louis S. and Dale R. Calder. Hydroids colonizing the carapaces of the

ostracode Philomedes brenda from the Beaufort Sea, Arctic Ocean 125-129

Kritsky, Delane C. and Walter A. Boeger. Neotropical Monogenoidea. 26.

Annulotrematoides amazonicus, a new genus and species (Dactylogyridae:

Ancyrocephalinae), from the gills of Psectrogaster rutiloides (Kner) (Teleostei:

Characiformes: Curimatidae) from the Brazilian Amazon 528-532

Kropp, Roy K. Lithoscaptus pardalotus, a new species of coral-dwelling gall crab

(Crustacea: Brachyura: Cryptochiridae) from Belau 637-642

Lemaitre, Rafael. Charybdis hellerii (Milne Edwards, 1867), a nonindigenous portunid

crab (Crustacea: Decapoda: Brachyura) discovered in the Indian River lagoon system

of Florida 643-648

Lopez-Gonzalez, Pablo J. and Mercedes Conradi. Heteranthessius hoi, a new species

(Copepoda: Pseudanthessiidae) from a sea-anemone in the Straits of Gibralter with

remarks on the genus 107-1 16

Lotufo, Guilherme R. and John W. Fleeger. Description of Amphiascoides atopus, a

new species (Crustacea: Copepoda: Harpacticoida) from a mass culture system 117-124

Malabarba, Luiz R. and Andreas Kinder. A new species of the genus Bryconamericus

Eigenmann, 1907 from southern Brazil (Ostariophysi: Characidae) 679—686

Martin, Joel W. and Jennifer C. Christiansen. A new species of the shrimp genus

Chorocaris Martin & Hessler, 1990 (Crustacea: Decapoda: Bresiliidae) from

hydrothermal vent fields along the Mid-Atlantic Ridge 220-227

McClure, Matthew R. Alpheus angulatus, a new species of snapping shrimp from the

Gulf of Mexico and northwestern Atlantic, with a redescription of A. heterochaelis

Say, 1818 (Decapoda: Caridea: Alpheidae) 84-97

McLaughlin, Patsy A. and Janet Haig. A new species of Goreopagurus McLaughlin

(Decapoda: Anomura: Paguridae) from the Pacific, and a comparison with its Atlantic

counterpart 68-75

Messing, Charles G. Alloeocomatella, a new genus of reef-dwelling feather star from

the tropical Indo-West Pacific (Echinodermata: Crinoidea: Comasteridae) 436-450

Messing, Charles G. Redescription of a unique feather star (Echinodermata: Crinoidea:

Comatulida: Comasteridae) with the diagnosis of a new genus 656-661

Miura, Tomoyuki and Daniel Desbruyeres. Two new species of Opisthotrochopodus

(Polychaeta: Polynoidae: Branchinotogluminae) from the Lau and the North Fiji Bac-

arc Basins, southwestern Pacific Ocean 583-595

Ng, Peter K. L. and Takeharu Kosuge. On a new Somanniathelphusa Bott, 1968, from

Vietnam (Crustacea: Decapoda: Brachyura: Parathelphusidae 61-67

Norris, James N. and David L. Ballantine. Two new species of the red alga Chrysymenia

J. Agardh (Rhodymeniales: Rhodymeniaceae) from the tropical western Atlantic 153-165

Olson, Storrs L. and Helen F. James. Nomenclature of the Hawaiian Akialoas and

Nukupuus ( A ves : Drepanidini) 373-3 87

Ott, Jorg A., Monika Bauer-Nebelsick, and Veronica Novotny. The genus Laxus Cobb,

1894 (Stilbonematinae: Nematoda): Description of two new species with

ectosymbiotic chemoautotrophic bacteria 508-5 27

Palma, R. Eduardo. The karyotypes of two South American mouse opossums of the

genus Thylamys (Marsupialia: Didelphidae), from the Andes, and eastern Paraguay 1-5

Pamplona, Denise and Marcia Souto Couri. Morellia dendropanacis, a new species,

and other species with spotted wings: characterization and comparison (Diptera:

Muscidae : Muscinae) 45 1^57

Patton, James L. and Maria Nazareth F. da Silva. A review of the spiny mouse genus

Scolomys (Rodentia: Muridae: Sigmodontinae) with the description of a new species

from the western Amazon of Brazil 319-337

Pettibone, Marian H. New genera for two polychaetes of Lepidonotinae 577-582

Pohle, Gerhard and Fernando Marques. First zoa of Dissodactylus glasselli Rioja and

new range and host records for species of Dissodactylus (Brachyura: Pinnotheridae),

with a discussion of host-symbiont biogeography 247-253

Rambla, Juan Pablo Blanco, Ildefonso Liilero Arana, and Luis Cal Beltran Lares M. A

new calianassid (Decapoda: Thalassinidea) from the southern Caribbean Sea 102-106

Rausch, R. L. and V. R. Rausch. The taxonomic status of the shrew of St. Lawrence

Island, Bering Sea (Mammalia: Soricidae) 717-729

Roccatagliata, Daniel and Richard W. Heard. Two species of Oxyurostylis (Crustacea:

Cumacea: Diastylidae), O. smithi Caiman, 1912 and O. lecroyae, a new species from

the Gulf of Mexico . 596-6 1 2

Rodriguez, Gilberto and Austin B. Williams. Epilobocera wetherbeei, a new species of

freshwater crab (Decapoda: Brachyura: Pseudothelphusidae) from Hispaniola 76-83

Rodriguez-Almaraz, Gabino A. and Thomas A. Bowman. Sphaerolana karenae, a new

species of hypogean isopod crustacean from Nuevo Leon, Mexico 207-211

Ross, Charles A., Gregory C. Mayer, and Roger Bour. Designation of a lectotype for

Crocodilus siamensis Schneider, 1801 (Reptilia: Crocodylia) 298-301

Ruedas, Luis A. Description of a new large-bodied species of Apomys Meams, 1905

(Mammalia: Rodentia: Muridae) from Mindoro Island, Philippines 302-318

Russell, David E. Description of a new viviparous species of Dentatisyllis (Polychaeta:

Syllidae) from Belize with an assessment of growth and variation, and emendation of

the genus 568-576

Schultz, George A. Sinoniscus cavemicolus, a new genus and species of terrestrial

isopod crustacean from a cave in China (Styloniscidae: Oniscidea) 201-206

Simone, Luiz Ricardo L. Rissoella omata, a new species of Rissoellidae (Mollusca:

Gastropoda: Rissoelloidea) from the southeastern coast of Brazil 560-567

Stark, Bill P. A new species of Neoperla (Insecta: Plecoptera: Perlidae) from Mississippi 45-49

Tan, Cheryl G. S. Dittosa, a new genus of leucosiid (Crustacea: Decapoda: Brachjoira)

from southern Australia and New Zealand 465-476

Thomas, James Darwin and Manuel Ortiz. Leucothoe laurensi, a new species of

leucothoid amphipod from Cuban waters (Crustacea: Amphipoda: Leucothoidae) 613-616

Thompson, Fred G. A new freshwater snail from the Coosa River, Alabama

(Gastropoda: Prosobranchia: Hydrobiidae) 502-507

Todaro, M. Antonio. Paraturbanella solitaria, a new psammic species (Gastrotricha:

Macrodasyida: Turbanellidae), from the coast of Califomian 553-559

Turner, Richard L. and Robyn M. Heyman. Rediagnosis of the brittlestar genus

Ophiosyzygus and notes on its type species O. disacanthus (Echinodermata:

Ophiuroidea: Ophiomyxidae) based on the type specimens from Japanese waters and

new material from the Gulf of Mexico 292-297

Vazquez-Bader, Ana Rosa and Adolfo Gracia. A new crab species of the genus

Pseudorhombila H. Milne-Edwards, 1837 (Crustacea: Decapoda: Goneplacidae) 254-265

Wicksten, Mary K. Wlthin-species variation in Periclimenes yucatanicus (Ives), with

taxonomic remarks on P. pedersoni Chace (Crustacea: Decapoda: Caridea:

Palaemonidae) 458^64

Williams, Austin B. and Fred C. Dobbs. A new genus and species of caridean shrimp

(Crustacea: Decapoda: Bresiliidae) from hydrothermal vents on Loihi Seamount,

Hawaii 228-237

Zug, George R. and Ivan Ineich. A new skink (Emoia: Lacertilia: Reptilia) from the

forest of Fiji 395^00

Zunio, Gabriel E., Olga B. Vaccaro, Marcelo Canevari, and Alfred L. Gardner.

Taxonomy of the genus Lycalopex (Carnivora: Canidae) in Argentina 129-1 Al


Volume 108

(New taxa are indicated in italics; new combinations designated n.c.)


Antillocyathus gracilis 536

Asterosmilia irregularis 541

Narella omata 148

Paracy athus adetos 538

Oxy srm\\2i pliocenica 539

Septastracea altispina 534


Annulotrematoides ; 529

amazonicus 529


Paraturbanella solitaria 554


Laxus cosmopolitus 517

oneistus 509

MOLLUSCA Gastropoda

Opisthotrochopodus segonzaci 584

trifu reus 588

Pyrgulopsis hershleri 502

Rissoella (Rissoella) omata 56 1



Augenerilepidonotus 577

dictyolelpis n.c 578

Dentatisyllis mangalis 569

Eusyllis maxima n.c 496

Laonome albicingillum 131

Olgalepidonotus 580

kumari n.c. 580

Raricimis variabilis 169


Olavius nicolae 491



Alpheus angulatus 85

Amphiascoides atopus 118

Qhorocan^ fortunata 221

Discorsopagurus maclaughlinae 618

Dittosa 465

Epilobocera wetherbeei 77

Goreopagurus garthi 69

Heteranthessius hoi 108

Hypolobocera emberarum 652

kamsarum 649

Leucothoe laurensi 613

Lithoscaptus pardalotus 637

Macrobrachium catonium 50

Natatolana nukumbutho 212

Neocallichirus cacahuate 478

Opaepele 229

loihi 229

Oxyurostylis lecroyae 604

Procambarus (Ortmannicus) nueces 54

Pseudorhombila ometlanti 255

Ridgewayia klausruetzleri 182

Sergio guaiqueri 102

mericeae 267

Sinoniscus 201

cavemicolus 202

Somanniathelphusa /?ajc 62

Sphaerolana karenae 207

Strengehana florenciae 98


Morellia dendropanacis 451

Neoperla coxi 45


Alloeocomatella 437

pectinifera n.c 445

polycladia 438

Aphanocomaster 657


Bryconamericus lambari 680

Gobiodon acicularis 688

Gymnotus maculosus 666


Leptodactylus lithonaetes 708

mye rsi 712


Brachymeles minimus 392

Emoia mokosariniveikau 395

Sphenomorphus kitangladensis knollmanae

389 20

Akialoa .

ellisiana n.c

lanaiensis n.c. ...

obscura n.c

stejnegeri n.c. _- upupirostris n.c.



384 384 384 384 385 385

Apomys gracilirostris

Scolomys juruaense




Chrysymenia littleriana



glandulosus n.c.

154 159 748 749









ISSN 0006-324X


1994-1995 Officers

President: Janet W. Reid Secretary: Carole C. Baldwin

President-elect: Stephen D. Cairns Treasurer: T. Chad Walter

Elected Council

Robert J. Emry Susan L. Jewett

Richard C. Froeschner Lynne R. Parenti

Alfred L. Gardner F. Christian Thompson

Custodian of Publications: Austin B. Williams

PROCEEDINGS Editor: C. Brian Robbins

Associate Editors

Classical Languages: George C. Steyskal Invertebrates: Jon L. Norenburg

Frank D. Ferrari Plants: David B. Lellinger Rafael Lemaitre

Insects: Wayne N. Mathis Vertebrates: Thomas A. Munroe

Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $25.00 (for USA and non-USA addresses) include subscription to the Proceed- ings of the Biological Society of Washington. Library subscriptions to the Proceedings are: $40.00 for USA and non-USA addresses. Non-USA members or subscribers may pay an addi- tional $25.00 to receive the Proceedings by Air Mail.

The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued sporadically) are available. Correspondence dealing with membership and subscriptions should be sent to the Biological Society of Washington, P.O. Box 1897, Lawrence, Kansas 66044, U.S.A. Payment for membership is accepted in US dollars (cash or postal money order), checks on US banks, or MASTERCARD or VISA credit cards.

Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560.

Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044

Second class postage paid at Washington, D.C, and additional mailing office.

POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCI- ETY OF WASHINGTON, RO. Box 1897, Lawrence, Kansas 66044.

@ This paper meets the requirements of ANSI/NISO Z39.48- 1992 (Permanence of Paper).


108(l):l-5. 1995.

The karyotypes of two South American mouse opossums of the

genus Thylamys (Marsupialia: Didelphidae), from the

Andes, and eastern Paraguay

R. Eduardo Palma

Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, New Mexico 87131-1091, U.S.A.

Abstract. —The karyotypes of Thylamys macrura, from eastern Paraguay and adjacent Brazil, and T. pallidior, from the Andean Altiplano in Argentina and Bolivia, are reported for the first time. The karyotypes of somatic tissues from female T. macrura and female T. pallidior consist of 2N =14. However, the diploid number found in somatic cells in males of T. pallidior exhibits 2N = 1 3 because the Y chromosome is missing. The missing Y phenomenon might represent another case of chromosome mosaicism in American marsupials, although this time detected in didelphids.

Tate (1933) recognized five species groups in the genus Marmosa Gray, 1821 (sensu lato) based on phenetic characters. Subse- quent morphologic, chromosomic, and se- rologic studies have shown that these as- semblages approximate genera (Reig et al. 1985, 1987; Gardner & Creighton 1989). Among these, Thylamys Gray, 1843, has the southernmost distribution of mouse opossums in South America encompassing elevations as high as 3500 m, as well as lower temperate and subtropical habitats (Cabrera 1958). Thylamys differs from oth- er marmosines in having a characteristic tri- colored dorsal pattern, and the capacity to store fat in the tail (Tate 1933, Mann 1978) Recent revisions of the genus recognize five species (Gardner & Creighton 1989, Gard- ner 1993): Thylamys pusillus (Desmarest 1804), Thylamys macrura (Olfers 1818), Thylamys elegans (y^diXQrhousQ 1839), Thy- lamys velutinus (Wagner 1842), and Thy- lamys pallidior (Thomas 1902). Thylamys pallidior occurs on the rocky slopes of the Altiplano of Bolivia and Argentina (Tate 1933; fig. 1), whereas, the subtropical form T. macrura is found in the humid forests of eastern Paraguay and adjacent Brazil (Gard-

ner 1993; fig. 1). This constitutes the sixth published record of this species for Para- guay, since the original description by Olfers (1818) based on Azara's (1801) "Micoure a queue longue," or "Colilargo" (Azara 1845). Tate (1933) referred to this species as Mar- mosa marmota, and Cabrera (1 958) as Mar- mosa grisea. The name macrura (Olfers 1818) is available and T macrura is used by Gardner (1993). In this study, the kar- yotypes of T. macrura and T. pallidior are reported for the first time. This is the first study documenting the absence of one of the sex chromosomes of somatic cells in didelphid marsupials.


Chromosomal preparations were ob- tained directly from bone marrow using the standard velban technique described in An- derson et al. (1987). A minimum of 10 metaphase spreads were counted for each specimen. Nomenclature for chromosome morphology and fundamental number (FN) follows Patton (1967). Five specimens of T. pallidior from two localities in Bolivia were analyzed. The karyotype of T macrura was


Fig. 1 . Map showing the distribution of T. pallidior along the Andes of Argentina and BoHvia, and T. ma- crura in eastern Paraguay. The boxes and dot represent the collecting site given in the text.

obtained from a specimen from eastern Par- aguay. Voucher specimens, chromosome sHdes, and cell suspensions are deposited in the Museum of Southwestern Biology (MSB) and the American Museum of Natural His- tory (AMNH). Collection localities are (Fig. 1): Bolivia: Department of Chuquisaca, Ca- margo, 68 km (by road) N of Camargo, 3400 m, 20°09'S, 65°17'W (3 males MSB 57003, AMNH 262406, and AMNH 262407; 1 fe- male AMNH 262405); Department of Tar- ija, Serrania Sama, 3200 m, 21°27'S, 64°52'W (1 female, AMNH 263555). Par- aguay: Department of Concepcion, Escuela Agropecuaria, 7 km (by road) NE from Con- cepcion, 23°21'S, 57°23'W (1 female, MSB NK 27536).

Results and Discussion

The autosomes of T. macrura (2N =14, FN = 20; Fig. 2a) consist of three pairs of large submetacentrics (1-3), one pair of me-

dium-sized metacentrics (4), and two pairs of small acrocentrics (5-6). The X chro- mosome is a small acrocentric. The auto- somes of T. pallidior (Fig. 2b) are not dis- tinguishable from those of T. macrura, al- though the three males of the Andean spe- cies present 2N = 13, FN = 20; the Y chromosome was absent in all counted plates. The female T. pallidior exhibited the complete set of chromosomes, 2N = 14. A male of T. macrura was not available for karyotype.

The autosomic complement of the species of Thylamys reported here is similar to those documented previously for other species of the genus, such as T. elegans from Chile and Bolivia, which possess the identical three group pattern of autosomes and morphol- ogy (Reig et al. 1972, Palma & Yates 1995). Micoureus cinereus and M. constantiae have similar diploid and fundamental number (2N = 14, FN = 20; Palma & Yates 1995). Marmosa (sensu stricto), Marmosops, and Gracilinanus, have 2N =14, however the fundamental number in these taxa is FN = 24 (Reig 1968, Palma & Yates 1995).

The mouse opossum karyotypes present- ed in this paper reinforce the concept of chromosomic conservatism in marsupials, and support the fact that marsupial species that occur in remarkably different habitats share a common karyotype (Reig et al. 1977, Hayman 1990). The common 2N = 14 is shared by most of the marmosines in the Neotropics. Marmosa canescens (2N = 22), is the only known exception (Engstrom & Gardner 1988).

The conclusive evidence of the absence of the Y chromosome in T. pallidior is dif- ficult to determine under the methodology followed in this study. It is possible that the Y has been translocated to another chro- mosome, or this condition may be another example of chromosome mosaicism, i.e., a difference in sex-chromosome presence be- tween the germ line and cells of the somatic tissues (Hayman 1990). Similar patterns of absence of the Y chromosome for somatic



(f II II


H it mm


u n


H 6


Fig. 2, a, Standard karyotype of a female Thylamys macrura from Concepcion, Paraguay, 2N = 14, FN = 20. b, Standard karyotype of a male Thylamys pallidior from Camargo, Chuquisaca, Bolivia, 2N = 14,

FN = 20.

cells have been found for Chironectes min- imus (Palma & Yates 1995) and Dromiciops australis (Gallardo & Patterson 1987). Ad- ditional reports of chromosome mosaicism have been also made for Australian mar- supials of the family Peramelidae, where one of the X chromosomes is missing in somatic cells of females, and in Petauroides (Pe- tauridae), where the Y chromosome is miss- ing from the majority of cells obtained from bone marrow (Murray et al. 1979, Hay man 1990).

The missing Y chromosome found in Dromiciops australis (Gallardo & Patterson 1987), caused these authors to suggest that the microbiotheriid would be more related to Australasian marsupials than to Ameri- can metatherians, supporting Szalay's (1982) contention that Dromiciops and Australian marsupials constitute the cohort Australi- delphia. Data from this study and from Pal- ma & Yates (1995), prove that the Y chro- mosome is missing not only in Dromiciops and Australasian marsupials, but in Amer- ican marsupials as well. This scenario fits a typical case of parallelism or represents a plesiomorphic condition in the evolution of metatherian sexual chromosomes of both geographic regions. Hence, this character cannot be used as evidence for inferring

phylogeny between Australian and Ameri- can marsupial lineages.


Mike Bogan, Jennifer K. Frey, Jorge Sa- lazar, and Terry L. Yates provided valuable comments on a draft of the manuscript. Special thanks to Lucy Aquino from CITES- Paraguay, and Carl Shuster; to Raviano Colman and Luis Moran of the National Museum of Natural History of Paraguay, Maribe Robles, and to the National Parks Office of the Ministry of Agriculture and Livestock of Paraguay. Funding support was provided by a NSF Research Improvement Grant PDS- 105-774, the Latin American Institute, University of New Mexico, and Sigma Xi. Field work in Bolivia was sup- ported by NSF grants BSR-83- 16740 to the American Museum of Natural History (S. Anderson), and BSR-84-08923 to the Uni- versity of New Mexico (T. L. Yates). The map was prepared by Beth Dennis.

Literature Cited

Anderson, S., T. L. Yates, & J. A. Cook. 1987. Notes of Bolivian mammals 4: the genus Ctenomys (Rodentia, Ctenomyidae) in the eastern low- lands.—American Museum Novitates 2891:1- 20.


Azara, F., d'. 1 80 L Essais sur I'histoire naturelle des quadrupedes de la Province du Paraguay. Trad- uits sur le manuscript inedit de I'auteur, Pra. M. L. E. Moreau-Saint-Mery. Charies Pougens, Paris, 366 pp.

. 1845. Viajes por la America del Sur. Bibli-

oteca de Impresos Raros Americanos. Univer- sidad de la Republica, Facultad de Humani- dades y Ciencias, Institute de Investigaciones Historicas, Montevideo 1982, 318 pp.

Cabrera, A. 1958. Catalog© de los mamiferos de America del Sur.— Revista del Museo Argenti- ne de Ciencias Naturales "Bernardino Riva- davia", Ciencias Zoologicas 4:1-307.

Desmarest, A. G. 1804. Taubleau methodique des mammiferes. Pp. 5-38 in Nouveau dictionnaire d'histoire naturelle, appliquee aux arts, princi- palement a I'agriculture, a I'economie rurale et domestique: Par une societe de naturalistes et d'agriculteurs: Avec des figures tirees des trois regnes de la nature. Vol. 24. Paris, Deterville.

Engstrom, M. D., & A. L. Gardner. 1988. Karyotype of Marmosa canescens (Marsupialia: Didelphi- dae): a mouse opossum with 22 chromo- somes.—The Southwestern Naturalist 33:230- 233.

Gallardo, M. H., & B. D. Patterson. 1987. An ad- ditional 14-chromosome mosaicism in South American marsupials. Pp. 1 1 1-1 1 5 z>2 B. D. Pat- terson & R. M. Timm, eds., Studies in Neo- tropical mammalogy: essays in honor of Philip Hershkovitz.- Fieldiana: Zoology, new series 39:1-506.

Gardner, A. L. 1993. Order Didelphimorphia. Pp. 15-23 in D. E. Wilson & D. M. Reeder, eds., Mammal species of the world: a taxonomic and geographic reference. Second edition. Smith- sonian Institution Press, Washington, 1 206 pp.

, & G. K. Creighton. 1 989. A new generic name

for Tate's (1933) microtarsus group of South American mouse opossums (Marsupialia: Di- delphidae).— Proceedings of the Biological So- ciety of Washington 102:3-7.

Gray, J. E. 1821. On the natural arrangement of the vertebrose animals.— London Medical Reposi- tory 15: 296-310.

. 1843. List of the specimens of mammalia in

the collection of the British Museum of London. British Museum (Natural History), 2 1 6 pp.

Hayman, D. L. 1990. Marsupial cytogenetics.— Aus- tralian Journal of Zoology 37:331-349.

Mann, G. 1978. Los pequeiios mamiferos de Chile. Gay ana, Zoologia 40:1-342.

Murray, J. D., G. M. Mckay, & G. B. Sharman. 1979. Studies on metatherian sex chromosomes. IX. Sex chromosomes of the greater glider (Mar- supialia: Petauridae).— Australian Journal of Biological Sciences 32:375-386.

Olfers, I. 1818. Bemerkungen zu lUiger's Ueberblick der Saugthiere, nach ihrer Vertheilung iiber die Welttheile, riicksichtlich der Siidamerikanisch- en Arten (Species). Pp. 129-137 in Journal von Brazilien, oder vermischte Nachrichten aus Brasilien, auf wissenschaftlichen Reisen ges- ammelt, W. L. Eschwege, 129-137 in F. J. Ber- tuch, ed., Neue Bibliotek der wichtigsten Reise- beschreibungen zur Erweiterung der erd- und Volkerkunde; in Verbindung mit einigen an- deren Gelehrten gesammelt und herausgegeben. Weimar: Verlage des Landes-Industrie-Comp- toirs. Vol. 15, 304 pp.

Palma,R.E.,& T.L.Yates. 1995. The chromosomes of Bolivian didelphid marsupials.— Occasional Papers, The Museum, Texas Tech University, Lubbock (in press).

Patton, J. L. 1967. Chromosome studies of certain pocket mice, genus Perognathus (Rodentia, Het- eromyidae).— Journal of Mammalogy 48:27-37.

Reig, O. A. 1 968. The chromosomes of the didelphiid Marmosa robinsoni Bangs.— Experientia 24: 185-186.

, R. Fernandez, & A. Spotomo. 1972. Further

occurrence of a karyotype of 2n = 14 chromo- some in two species of Chilean didelphoid mar- supials.—Zeitschrift fur Saugetierkunde 37:37- 42.

, J. A. W. Kirsch, & L. Marshall. 1985. New

conclusions on the relationships of the opos- sum-like marsupials with an annotated classi- fication of the Didelphimorphia. Ameghi- niana 21:335-343.

, , & . 1987. Systematic relation- ships of the living and Neocenozoic American "opossum-like" marsupials (suborder Didel- phimorphia), with comments on the classifica- tion of these and of the Cretaceous and Paleo- gene New World and European metatherians. Pp. 1-89 in M. Archer, ed.. Possums and opos- sums: studies in evolution. Surrey Beatty and Sons and the Royal Society of New South Wales, Sydney, 1 and 2, 800 pp.

, A. L. Gardner, N. O. Bianchi, & J. L. Patton.

1977. The chromosomes of the Didelphidae (Marsupialia) and their evolutionary signifi- cance.—Biological Journal of the Linnean So- ciety 9:191-216.

Rofe, R., & D. Hayman. 1985. G-banding evidence for a conserved complement in the Marsupi- alia.—Cytogenetics and Cell Genetics 39:40-50.

Szalay, F. S. 1982. A new appraisal of marsupial phylogeny and classification. Pp. 62 1-640 in M. Archer, ed., Carnivorous marsupials. The Royal Biological Society of New South Wales, Sydney, Australia, 802 pp.

Tate, G. H. H. 1933. Systematic revision of the mar- supial genus Marmosa, with a discussion of the


adaptive radiation of the murine opossums (Marmosa).— Bulletin of the American Muse- um of Natural History 66:1-250.

Thomas, O. 1902. On Marmosa marmota and ele- gans with descriptions of new subspecies of the latter.— Annals and Magazine of Natural His- tory, series?, 9:408-410.

Wagner, J. A, 1842. Diagnosen neuer Arten brasi-

lischer Saugethiere. Archiv Naturgesch 8:356- 362. Waterhouse, G. R. 1839. Mammalia. Pp. 49-97 in C. Darwin, ed.. The zoology of the voyage of the H. M. S. Beagle under the command of Cap- tain Fitzroy, R. N., during the years 1832-1836. Smith, Elder and Company, London, 97 pp.


108(1):6-17. 1995.

Rediscovery and redescription of Sphenomorphus beyeri Taylor

(Reptilia: Lacertilia: Scincidae) from the Zambales

Mountains of Luzon, Philippines

Rafe M. Brown, John W. Femer, and Rogelio V. Sison

(RMB) Department of Zoology, Miami University, Oxford, Ohio 45056, U.S.A.;

(JWF) Department of Biology, Thomas More College, Crestview Hills, Kentucky 41017, U.S.A.;

(RMB, JWF) Department of Vertebrate Zoology, Cincinnati Museum of Natural History,

1720 Gilbert Avenue, Cincinnati, Ohio 45202-1401, U.S.A.;

(RVS) Zoology Division, National Museum of the Philippines, Executive House,

P. Burgos Street, Manila, Philippines

Abstract.— Sphenomorphus beyeri Taylor, 1922, long known from a single specimen, is redescribed on the basis of newly acquired material from the herpetologically unsurveyed area of the Zambales Mountains, west central Lu- zon Island, Republic of the Philippines. Phenotypically most similar to its distantly allopatric congener S. diwata, the holotype and our new series are distinguished from other Philippine Sphenomorphus by the combination of their small to moderate size (SVL = 46.6-67.1 mm), fused frontoparietals, 88- 96 paravertebrals, 38-42 scales at midbody, 19-21 subdigital fourth toe la- mellae, and unique coloration.

In his monograph on the lizards of the Philippine Islands, E. H. Taylor (1922a) recognized 16 Philippine species in the ge- nus Sphenomorphus (Lacertilia: Scincidae). He later supplemented this work on Phil- ippine lizards with descriptions of two more scincid species in a more general herpeto- logical contribution (Taylor 1922b). One of these was Sphenomorphus beyeri, a skink described on the basis of a single specimen found on Mt. Banahao in the Laguna Prov- ince of southern Luzon. Brown & Alcala (1980) later recognized 22 species of Phil- ippine Sphenomorphus. One of these was S. diwata Brown & Rabor (1967), which the authors hypothesized to be the closest rel- ative of S. beyeri. Brown & Alcala reported S. diwata from the Diwata mountains of northern Mindanao Island, and placed it and S. beyeri into the two-species non-phy- logenetic couplet "Group I Sphenomor- phus'" (1980: 1 54). As presently understood,

the genus Sphenomorphus contains over 1 20 species world-wide (Myers & Donnelly 1991) with 60 species in the Oriental and Australian zoogeographic regions alone (Brown&Alcala 1956, 1961a; Greer & Par- ker 1974; Greer 1979).

Sphenomorphus beyeri has, to date, only been known from the incomplete holotype, yet it invariably has been recognized as a valid species. In the course of a recent in- ventory of the fauna of the Philippines un- dertaken by the Cincinnati Museum of Nat- ural History (CMNH) and the National Mu- seum of the Philippines (PNM), we cap- tured 1 6 additional specimens of S. beyeri in the mossy cloud forests of the Zambales Mountains of west central Luzon. Collec- tion of this new material provided us with a unique opportunity to analyze intraspe- cific morphological variation of this endem- ic Philippine skink and to reconsider the validity of its specific rank.



We conducted field studies in the Zam- bales Mountain range from 1 7 February to 1 8 March 1 992. Drift fences and pitfall traps (25 m of 0.65 ml black plastic, stretched to 1 5 cm above ground, supported with wood- en stakes, with >50 cm deep plastic-lined pits spaced every 5 m for a total of six pit- falls per 25 m of drift fence at each 100 m interval on slope) and time-constrained searches were used to collect specimens. Specimens were photographed, then fixed in 10% buffered formalin; notes on color- ation, behavior, and habitat (including el- evation) were recorded at time of capture. Upon return to the U.S.A. (approximately one month later), specimens were trans- ferred to 70% ethanol.

Detailed examination of all material was conducted at the Cincinnati Museum of Natural History and at the National Mu- seum of Natural History. When possible (see character definitions below), we took mea- surements and scale counts following tech- niques detailed in Brown & Alcala (1980); illustrations of head scalation were made (by RMB) with a Wild microscope equipped with a camera lucida attachment.

Characters (measured to the nearest 0.1 mm) are defined as: snout-to-vent length (SVL), from tip of animal's snout to caudal margin of anal scale; tail length (TL), from caudal margin of anal scale to tail's tip (specimens with regenerated tails are not included in statistical analyses); axilla-groin distance (AGD), from caudalmost point where forelimb meets body to anteriormost point where hind limb meets body; hind leg length (HLL), from point where rear limb meets body to tip of longest (=4th) toe; head length (HL), from tip of snout to caudal edge of tympanum; head breadth (HE), width of head at its widest point (=ocular region) when viewed from above; snout length (SL), from anterior edge of bony orbit to tip of snout; eye diameter (ED), horizontal di- ameter across bony orbit; and tympanum

diameter (TD), horizontal distance across the tympanic annulus.

Lateral head scales (e.g., labial scales) were examined on both sides of the head and numbers for each side are given separately with a dashed line (— ) designating left from right respectively. Meristic and mensural data are given as means ± standard devi- ations (SD) and range.

Statistical analyses were carried out using the Statistical Analysis Software Program (SAS), version 6.03 (SAS Institute Inc. 1988a, 1988b), using UNIVARIATE pro- cedure for standard statistics. Significance of moment statistics (skewness, gi, and kur- tosis, g2) was calculated by hand (Sokal & Rohlf 1981:174-175).

Specimens are deposited in the California Academy of Science (CAS), Cincinnati Mu- seum of Natural History (CMNH), and Na- tional Museum of the Philippines (PNM). Material Examined includes: Holotype (Lu- zon Island, Laguna Province, Mt. Banahao) CAS 61 183, immature male, collected on a rock ledge on Mt. Banahao at 1500 m by E. H. Taylor (Taylor 1922b:285). Six fe- males (CMNH 3652, 3653, and 3658; PNM 2307, 2301, and 2304), nine males (CMNH 3655, 3657, and 3659; PNM 2300, 2302, 2303, 2305, 2306; USNM 337768) and one immature juvenile or hatchling (CMNH 3654), all collected by RMB and JWF. All were taken from Luzon Island, Zambales Province, Municipality of Masinloc, Bar- angay of Coto, Zambales Mountain range, Mt. High Peak.

Study Sites

Until the present study, (PNM/CMNH Philippine Biodiversity Inventory) the Zambales Mountains (Zambales Province, Municipality of Masinloc) were completely unsurveyed herpetologically due to a com- bination of major insurgency in this area and its close proximity to the Subic Bay Naval Base, 70 km south in the town of Olongapo. Following the eruption of Mt. Pinatubo in 199 1 and subsequent closing of


Fig. 1 . The Zambales Mountains. Shaded area rep- resents the major geologic formation constituting the Zambales Mountain range (see text). Darkened circle represents Mt. High Peak, and encompasses both lo- calities sampled.

the United States military facilities at Olon- gapo, an opportunity was made available to naturalists to survey this largely unknown region.

The Zambales Mountains (Fig. 1) are an isolated coastal mountain range encom- passing an area of approximately 6960 km^ on west central Luzon; the Zambales range extends from west of the Lingayen gulf to the western border of Manila Bay at an ori- entation of 20° West of due North. To the east, the Zambales Mountains are isolated from the nearest mountain range, the Cor- dillera Central, by the Plains of Tarlac (also known as the Pampanga Plains). To the west and north, these mountains end in the South China Sea; to the south they separate Subic Bay from Manila Bay. The Zambales Mountain range currently is contiguous with mainland Luzon, connected to the latter by the wide Plains of Tarlac, believed to have

been submerged below the Luzon sea during the Pliocene and Pleistocene (Rutland 1968; Hashimoto 1981a, 1981b; AufFenberg 1988). Geological evidence suggests that the Zam- bales Mountains were formerly isolated much farther west of their present position and were surrounded by water— not con- nected to mainland Luzon as they are today (Dickerson 1924, Auffenberg 1988). The Zambales currently represent a large "in- sular" massif, geologically isolated from the three other principal montane regions of Luzon (Bureau of Mines 1963; UNESCO/ ECAFE 1971; Hashimoto 1981a, 1981b; Auffenberg 1988).

Site 7.— Zambales Mountains, 1100 m, 15°35'N, 120°09'W. Sampled from 17 to 26 February, the site is a tropical moist decid- uous forest type (Whitmore 1984); virgin timber (predominantly Myrtaceae, Laura- ceae, and Tiliaceae) begins at an elevation of ca. 1030 m. Presence of Mt. Pinatubo ash deposits throughout the area combined with noticeable signs of rattan gathering efforts undertaken by local residents precludes des- ignation of the area as "undisturbed."

Site 2.— (Fig. 2) Zambales Mountains, 1500 m., 15°30'N, 120°08'W. Sampled be- tween 1 1 and 18 March 1992, this area lies within the broad category of tropical moist deciduous forest type but remains, none- theless, more typical of the upper montane (="mossy") rain forest (Whitmore 1984) due, presumably, to its somewhat higher el- evation. Virgin timber predominates (Fa- gaceae, Myrtaceae, Magnoliaceae, and Pin- aceae) and very little bamboo or other sec- ondary or disturbance indicator species were encountered.

A more extensive description of these study sites and a discussion of their geologic importance is provided by Ruedas et al. (1994).


Capture data.—K single specimen oi S. heyeri collected at Site 1 was a mature male


Fig. 2. Typical habitat of 5". beyeri between 1 500 and 1 600 m on Mt. High Peak, Zambales Mountains where most specimens of Sphenomorphus beyeri were found. The vegetation and forest floor in this area were coated with approximately 2-4 cm of volcanic ash from the eruption of Mt. Pinatubo. Photo from a color transparency by RMB, courtesy of CMNH.

(PNM 2300) collected under leaf litter on the forest floor at 1265 m. When disturbed, this lizard became alert, moved in a rapid serpentine manner and attempted to bur- row under debris and leaf litter. A pair of pitfall trapped specimens (CMNH 3652, a gravid female; PNM 2301, a mature female without eggs) from Site 2 were captured dur- ing the day, at 1510 m, and 1610 m, re- spectively. The rest of the new series was captured by splitting open rotten logs lying horizontally in contact with the forest floor. One specimen (male, PNM 2305) was dam- aged when the machete used to split the log struck it. No specimens were taken at night despite extensive search efforts.

Sphenomorphus beyeri Taylor, 1922:283 Fig. 3, 4

Diagnosis.— K small to moderate-sized Sphenomorphus, S. beyeri is readily distin-

guished from its congeners by the following combination of characters: (1) prefrontals moderate, usually separate; (2) frontopar- ietals fused except in immature specimens; (3) usually 6-7 labials; (4) four large supra- oculars (5) paravertebrals 88-96; (6) scales around midbody 38-42; (7) fourth toe la- mellae 1 9-2 1 ; (8) body proportion ratios as follows, SL/HL = 0.25-0.37; SL/HB = 0.38- 0.57; HB/HL = 0.60-0.69; HB/SVL = 0.13- 0.17; ED/SL = 0.30-0.43; (9) unique col- oration and color pattern.

Description (based on holotype and 1 6 re- ferred specimens). Details