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aD en nary

PROCEEDINGS

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

WASHINGTON PRINTED FOR THE SOCIETY

EDITOR

C. BRIAN ROBBINS

ASSOCIATE EDITORS

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

OFFICERS AND COUNCIL of the BIOLOGICAL SOCIETY OF WASHINGTON FOR 1994-1995

OFFICERS

President JANET W. REID

President-Elect STEPHEN D. CAIRNS

Secretary CAROLE C. BALDWIN

Treasurer T. CHAD WALTER

COUNCIL Elected Members ROBERT J. EMRY SUSAN L. JEWETT RICHARD C. FROESCHNER LYNNE R. PARENTI

ALFRED L. GARDNER F. CHRISTIAN THOMPSON

TABLE OF CONTENTS

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 22 ee Ee a ee ee Se ee eee

Bayer, Frederick M. A new species of the gorgonacean genus Narella (Anthozoa: Octocorallia) from: Haw aiwan< waters ss ed ee eee ent

Brown, Rafe M., John W. Ferner, and Luis A. Ruedas. A new species of lygosomine lizard (Reptilia: Lacertilia: Scincidae: Sphenomorphus) from Mt. Isarog, Luzon Island, Philippines ..22 bo a ee

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 [uzon; Philippines!2 = 2s ee ee

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’ Catanduanesylsl anes) P baal p pony cs eee ce eae ee ae eee ee

Bruce, Niel L. and Jgrgen 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 Sternarchorhamphus Eigenmann, 1905 (Ostariophysi: Gymnotiformes: Apteronotidae), with: notes on its relationships 222° 6) Sees eee a ie ee eee

Carleton, Michael D. and Guy G. Musser. Systematic studies of oryzomyine rodents (Muridae: Sigmodontinae): definition and distribution of Oligoryzomys vegetus RB earns 1D) an IS ST RIT AN le

Couri, Marcia Souto. Contribution to the knowledge of Reynoldsia Malloch (Diptera: VAS eae) 2 a 2 SoH Nd Tcl eh So a a each A eM ee LL

Cuatrecasas, Jose. A new genus of the Compositae: Paramiflos (Espeletiinae) from (Gro) (05101 5) b: Pane Ne ON en ee Oy Ae On Bey ure aA VE is ee ee he

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

Erséus, Christer and Olav Giere. Otavius nicolae, a new gutless marine tubificid species (Oligochaeta) ‘frome li Ze i ICN

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 onFappEendagerde velop mve mt ee ee tae ee

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

iV

662-678

147-152

18—28

6-17

388-391

392-394

212-219

533-550

98-101

649-655

29-44

338-369

281-291

748-750

238-246

629-636

169-179

491-495

477—490

180—200

136-146

687-694

Heyer, W. Ronald. South American rocky habitat Leptodactylus (Amphibia: Anura: Leptodactylidae) with description of two new species —____----________ Hobbs, Horton H., Jr. and H. H. Hobbs III. Procambarus (Ortmannicus) nueces (Decapoda: Cambaridae), a new crayfish from the Nueces River Basin, Texas ____ Hobbs III, H. H. and Horton H. Hobbs, Jr. Macrobrachium catonium, a new troglobitic shrimp from the Cayo District of Belize (Crustacea: Decapoda: Palaemonidae) ___. Hotchkiss, Frederick H. C. Lovén’s law and adult ray homologies in echinoids, ophiuroids, edrioasteroids, and an ophiocistioid (Echinodermata: Eleutherozoa) ____. Hsieh, Hwey-Lian. Laonome albicingillum, a new fan worm species (Polychaeta: Sabpellidae-sSaveliinae) trond dhabw an ore eatemntwa ee hs Slee Ee a Fe Jiménez, 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) ___ Komai, Tomoyuki. A new species of the genus Discorsopagurus (Crustacea: Decapoda: Paguridae) from Japan, previously known as D. schmitti (Stevens) —------------_-- Kornicker, Louis S. and Dale R. Calder. Hydroids colonizing the carapaces of the ostracode Philomedes brenda from the Beaufort Sea, Arctic Ocean _-_-_------ 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 —_- Kropp, Roy K. Lithoscaptus pardalotus, a new species of coral-dwelling gall crab (@iustacea> Brachyura; Cryptochiridac) from’ Belaw = Lemaitre, Rafael. Charybdis hellerii (Milne Edwards, 1867), a nonindigenous portunid crab (Crustacea: Decapoda: Brachyura) discovered in the Indian River lagoon system ORE LECTERNS ee i A ee RE TN. a Lépez-Gonzalez, Pablo J. and Mercedes Conradi. Heteranthessius hoi, a new species (Copepoda: Pseudanthessiidae) from a sea-anemone in the Straits of Gibralter with Be AAR SR OME CHS EC MU Ses waiter cae AL cat au el Lc ure ee Baa A a Bs Ve RE Lotufo, Guilherme R. and John W. Fleeger. Description of Amphiascoides atopus, a new species (Crustacea: Copepoda: Harpacticoida) from a mass culture system ___ Malabarba, Luiz R. and Andreas Kinder. A new species of the genus Bryconamericus Eigenmann, 1907 from southern Brazil (Ostariophysi: Characidae) —__--__-_-_-____- 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 —-_____ 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 Savemlolondoccapoda:, Candea: Alpheidac)) sa se McLaughlin, Patsy A. and Janet Haig. A new species of Goreopagurus McLaughlin (Decapoda: Anomura: Paguridae) from the Pacific, and a comparison with its Atlantic SOTTO as cP A MR A SR. RE a Re” Messing, Charles G. Alloeocomatella, a new genus of reef-dwelling feather star from the tropical Indo-West Pacific (Echinodermata: Crinoidea: Comasteridae) = Messing, Charles G. Redescription of a unique feather star (Echinodermata: Crinoidea: Comatulida: Comasteridae) with the diagnosis of a new genus __... Miura, Tomoyuki and Daniel Desbruyéres. Two new species of Opisthotrochopodus (Polychaeta: Polynoidae: Branchinotogluminae) from the Lau and the North Fiji Bac- Alcea aSinS WSOUthWeESterlybaCiiC, OCEAN yw ee ea Ng, Peter K. L. and Takeharu Kosuge. On a new Somanniathelphusa Bott, 1968, from Vietnam (Crustacea: Decapoda: Brachyura: Parathelphusidae _____»_»_____ Norris, James N. and David L. Ballantine. Two new species of the red alga Chrysymenia J. Agardh (Rhodymeniales: Rhodymeniaceae) from the tropical western Atlantic __- Olson, Storrs L. and Helen E James. Nomenclature of the Hawaiian Akialoas and INU PIS CAVeSss Dre pani Gini) == eset 2 cine eine ee erode ee ge a es ee Ott, Jorg A., Monika Bauer-Nebelsick, and Veronica Novotny. The genus Laxus Cobb, 1894 (Stilbonematinae: Nematoda): Description of two new species with cctosyimbiotic: Chemoautotrophic bacteria 228 eave Ty See 1) ee

695-716

54-60

50-53

401-435

130-135

496-501

617-628

125-129

528-532

637-642

643-648

107-116

117-124

679-686

220-227

84-97

68-75

436—450

656-661

583-595

61-67

153-165

373-387

508-527

Vi

Palma, R. Eduardo. The karyotypes of two South American mouse opossums of the genus Thylamys (Marsupialia: Didelphidae), from the Andes, and eastern Paraguay Pamplona, Denise and Marcia Souto Couri. Morellia dendropanacis, a new species,

and other species with spotted wings: characterization and comparison (Diptera: Miuscidae: Mii Sciniae)) settee 2 ios et cas Rd oe 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. 2.22: ee ee Pettibone, Marian H. New genera for two polychaetes of Lepidonotinae _________ 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 -_-----------------------_______________------ Rambla, Juan Pablo Blanco, Ildefonso Lifiero Arana, and Luis Cal Beltran Lares M. A new calianassid (Decapoda: Thalassinidea) from the southern Caribbean Sea ______. Rausch, R. L. and V. R. Rausch. The taxonomic status of the shrew of St. Lawrence Island Bering, Seax(ViammaliaS Sorncidac) Roccatagliata, Daniel and Richard W. Heard. Two species of Oxyurostylis (Crustacea: Cumacea: Diastylidae), O. smithi Calman, 1912 and O. lecroyae, a new species from the Gulf of Wile xtc ee a ae ea ee tone ee Rodriguez, Gilberto and Austin B. Williams. Epilobocera wetherbeei, a new species of freshwater crab (Decapoda: Brachyura: Pseudothelphusidae) from Hispaniola _______ Rédriguez-Almaraz, Gabino A. and Thomas A. Bowman. Sphaerolana karenae, a new species of hypogean isopod crustacean from Nuevo Leon, Mexico —_.________. Ross, Charles A., Gregory C. Mayer, and Roger Bour. Designation of a lectotype for Crocodilus siamensis Schneider, 1801 (Reptilia: Crocodylia) —----------------_------------ Ruedas, Luis A. Description of a new large-bodied species of Apomys Mearns, 1905 (Mammalia: Rodentia: Muridae) from Mindoro Island, Philippines ___-_-_-----_--_--____- 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 CEN fA UNS ae We es AN re el 8 BN La co ao Schultz, George A. Sinoniscus cavernicolus, a new genus and species of terrestrial isopod crustacean from a cave in China (Styloniscidae: Oniscidea) _____-____________ Simone, Luiz Ricardo L. Rissoella ornata, a new species of Rissoellidae (Mollusca: Gastropoda: Rissoelloidea) from the southeastern coast of Brazil —----______ Stark, Bill P. A new species of Neoperla (Insecta: Plecoptera: Perlidae) from Mississippi Tan, Cheryl G. S. Dittosa, a new genus of leucosiid (Crustacea: Decapoda: Brachyura) from southern Australia and New Zealand Thomas, James Darwin and Manuel Ortiz. Leucothoe laurensi, a new species of leucothoid amphipod from Cuban waters (Crustacea: Amphipoda: Leucothoidae) __ Thompson, Fred G. A new freshwater snail from the Coosa River, Alabama (Gastropoda: Prosobranchia: Hydrobiidae) Todaro, M. Antonio. Paraturbanella solitaria, a new psammic species (Gastrotricha: Macrodasyida: Turbanellidae), from the coast of Californian 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, materialstromithesGullfso fMex 1c @ yee ee en Vazquez-Bader, Ana Rosa and Adolfo Gracia. A new crab species of the genus Pseudorhombila H. Milne-Edwards, 1837 (Crustacea: Decapoda: Goneplacidae) __ Wicksten, Mary K. Wlthin-species variation in Periclimenes yucatanicus (Ives), with taxonomic remarks on P. pedersoni Chace (Crustacea: Decapoda: Caridea: Palaemiomidae) 25st ode oe le Re A ON Pe a i eS Williams, Austin B. and Fred C. Dobbs. A new genus and species of caridean shrimp (Crustacea: Decapoda: Bresiliidae) from hydrothermal vents on Loihi Seamount, Hawait __.22008 tts Secale she Er Rea Sd 8 Rr ee pe A, A PTE Ot Zug, George R. and Ivan Ineich. A new skink (Emoia: Lacertilia: Reptilia) from the forést of Fajitas. 3 ust, dante ie aor SUN ie lito Sean aE ee ads eo Zunio, Gabriel E., Olga B. Vaccaro, Marcelo Canevari, and Alfred L. Gardner. Taxonomy of the genus Lycalopex (Carnivora: Canidae) in Argentina —-_-

451-457

319-337

577-582

247-253

102-106

717-729

596-612

76-83

207-211

298-301

302-318

568-576

201-206

560-567 45—49

465-476

613-616

502-507

553-559

292-297

254-265

458-464

228-237

395—400

729-747

INDEX TO NEW TAXA

VOLUME 108

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

CNIDARIA Anthozoa PMA INGN SA CITY S102 7; C2 CL LES pees eee as es ea Seth Sag ee a J a 536 ETO SSTIMUT, WAR GIA CT AIS a ge ee re a 541 INE GN OIA LATD as ASE SV SIO ee 148 ESE AG AGN UES NCECL CLO Spot eine Pater ee ee IN oe OE la ee) 538 Sg STIMt ABH OC C111 CCl mca aaa Da a Ti sa a a aS eg ee Se 539 SEOUL, LILES TGS es a ee a a 534 PLATYHELMINTHES Trematoda PANPIRTEL OD LTCTIULLOUCLC Settee ree rasa Te ee a a ee 529 CLO LLY DI OVID. Sa BEI IIIA, APD LED ELLEN DILL ALL DE LOO ELE AD ALTE AEA, EN 529 GASTROTRICHA Reate ct GUNTUL ATV NAS LEE CA gd Co ete ies ea 554 NEMATODA LEASES CDM LY DOT TIA so 1 a I ef A ee SIL7/ OLCUS TLE S puerta re snk RARE Ea Si rel ronan rem se ada min ohare mc Svs a micas, PS PRESEN SL NY, Bae 509 MOLLUSCA Gastropoda O pisthomochOpoGuUsise ONL GCI oe ss ae ee eae a ae 584 AY YIU IS asa SN ts I ENE SES aera SAPS RR OS Ae SU A a i Taya rege Re 588 | PRETO] SINS BT LIAS I GI Ee RA Re el ec 502 BRASS Oe lam (ERAS SOe IA) 1511 CE Ca seers Is ee se ee 561 ANNELIDA Polychaeta PETE TEL CPLA OTL OLUES pete ea aE Nad a esd ee Si, (INSTEON DNA, Sj ha gen ST A a 578 Sra ALIS NNT S Wrst OCLLES pre meatier eee I ee ees ee Ee 569 | EUAN TORRES SUD T AY (1 Chg LN a eo OTe 496 EAC R LDL CLI OTL irr pmtietaa toes aeniemn rsa ele en eee eat Ee 131 ONDA DIL D YONI: a ET eB nae eB Ee 580 | UWP VET, TO Oy eS I I eS a a ve 580 j PEW EF CETURTAINSS LUA ZY UTI tas IE I os I Te aS a 169 Oligochaeta CON ER IES 1K CUL AAG goa 8a TEIN SIE RS IT EP ee vga den 491

Vill

ARTHROPODA

Crustacea

PN Tp SATU QTL ATUL S cp a te 6 ee AmmphiaSCO1deSs’ QLOp uss 222 ie ST i a a ee ee (COLO CATES Tf OMT GL Ge T E Discorsopaguius maclaughlinae —— ee aa ee ee DUI OS Goo oN a OE Rive ptt BEA EME NIE STE AE eae ek FAR Sy EAE AEE ENE OR OE OO RE Ra Sia nS ae Epilobocerawetherbeeiy Soret es er a GOLEOPASUNUS are I Ea Ie EE see F¥eteramthe sss (Oe 22s aa ee a eat nee ee ne Eby POlODOCE Ra CHD CIC TaN ts oe ae NE Ie eco EN Ea en ere ee ene

RATES GUAT SL a i an at SER CRA RAL Oe Sn ra Ree ae ea a ee IECUCOENOC QUT CHS a eh eee Ne see en eae ee FvehO@SCAptuS: Parada OL1US ee sae ee ae are tle ee TO gee le ee ee IMlaACE@ Ora Lanta CLONE LET ITS mise I ee ce ee ae nan ene Ce oer ee

Oxyurosty lis WecrOy ae oe eee ce ee Procambarus (Ortmannicus e7ueces si eee 0 ee eat Eseudornombilayoriet arti ee re oe a ee RIA 8 CWAyias KIQUSTUCTZVC TE 2-2 Sin LL NE NG aS ee ee a eed SEL SION SUOLGUCTE, Hk SO ss ad SE RN ee

VILCV UCC AC nN AR Ee ea Dl SETLOMLES CUS Ae Docc 8 Es eh SS TED AE AR Ee se se

Soman miatine lps apa xe eI ce Sphaerolanma karen <0 ea eae ge StKEMP ERAN A /OTEM Cia mL A EE Teg eae ae,

Morelliadendropanacising. 28 8 06Gs Dies Se eden RU rare 2 ON eae ee ee INe@o@perl aC niet iis ae (Ue Ik 9 eg SS ee SE a erie ee

VA LOC OCOniate Ta 2 te x 5 Beet eG eal al A ale il st TO alca A a PS CET UL Trea ot: Ce a I eee Sees Ne NE PCN oc VOWS CLG chp to tae NIE a lc ad oi ae SLO ZAP ILAN OC OTIGS ECT, = he A Sa ea SAC al S18 Tat Ut NO Nae, re RE Sea eo

CHORDATA

Pisces

Bryconamericus lambari Gobiodon acicularis

Keptodactylus*lithonaetes: wi sesarince. Sy Nineties eine ee ee eel ee LOLA AY Se Ra eRe ti eet a ee A ene) eee en eee og. fe eae ee AN ee en

Bach y ime les” 172trerrii ves mse ene eke ee ee LP CE RR TEE ERE elie OE ee EMOla INOKOSATINIVCIKGU ee ee a eee ee tet ere

SMe MOM Op SACL SLA CUS 1S ee we LB Ie PN UA OUST DLL LG ssc 20 Ss Nt ADS Nae RE nT ee eS

ANI OWID GY ca atl Bs EOI VE ESS NP eC SO PRIS AT eR ee ESS TRE TOO oS NS TE Pa aT a ed oe a dl A ON STO en Ung PUL | ENTS TNO al SE OR Ll ee RL gd COOLS CU Trea NF epee ee eerste ee ns BE Le a OE A ng a RS Ee © SG Tel IMs © spree oe ae ce a ee (Us BUNA SST Sy a Ie ee a eee

Mammalia

ENSOLEDSIS IRGC MITOSIS AE at eg he ed ee SCO MOMMY SW LEMLECLCIUS Cima sie wena IRE Lee a ene EB

CG intgy SW MTN TNA ELE C11 CTU Cas te eee eal a a ee a TU OO CLUL ONS (a me ne De SIA SAL PS SARI AE Ca ea I a I a de seo a i PECDIROLT TG DS, Na Tel WON OR A ae aL ete EE Ten ce STRING UNOS UI S UMC nypeeereto cers Dae nce PN a ee ee ee

384 385 385

1X

ee (oO bu, VOW a *

oy

, & 0 é &

Rae

VOLUME 108 NUMBER 1

24 MARCH 199

ISSN 0006-324X

THE BIOLOGICAL SOCIETY OF WASHINGTON

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.

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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.

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

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

108(1):1-5. 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 7. pallidior, from the Andean Altiplano in Argentina and Bolivia, are reported for the first time. The karyotypes of somatic tissues from female 7. macrura and female T. pallidior consist of 2N = 14. However, the diploid number found in somatic cells in males of 7. pallidior exhibits 2N = 13 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 mMarmosines 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 (Waterhouse 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 1s 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) ““Micouré a queue longue,” or ““Colilargo” (Azara 1845). Tate (1933) referred to this species as Mar- mosa marmota, and Cabrera (1958) as Mar- mosa grisea. The name macrura (Olfers 1818) is available and 7. macrura is used by Gardner (1993). In this study, the kar- yotypes of 7. 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.

Methods

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 7. pallidior from two localities in Bolivia were analyzed. The karyotype of 7. macrura was

2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1000 km

80 70 60 50

100 90

Fig. 1. Map showing the distribution of T. pallidior along the Andes of Argentina and Bolivia, and 7. 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 slides, and cell suspensions are deposited in the Museum of Southwestern Biology (MSB) and the American Museum of Natural His- tory (AMNB). 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- ya, 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 7. 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 7. 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 7. pallidior exhibited the complete set of chromosomes, 2N = 14. A male of 7. 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, 1.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

VOLUME 108, NUMBER 1

b)

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,

EN 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, Hayman 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.

Acknowledgments

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 Flaviano Colman and Luis Moran of the National Museum of Natural History of Paraguay, Maribé 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.

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

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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. Ferner, 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’’ (1980:154). As presently understood,

the genus Sphenomorphus contains over 120 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 16 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.

VOLUME 108, NUMBER 1 Methods

We conducted field studies in the Zam- bales Mountain range from 17 February to 18 March 1992. Drift fences and pitfall traps (25 m of 0.65 ml black plastic, stretched to 15 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