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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
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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
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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.
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’. 1801. Essais sur l’histoire naturelle des
quadrupédes de la Province du Paraguay. Trad-
uits sur le manuscript inédit de l’auteur, Pra. M.
L. E. Moreau-Saint-Meéry. Charles Pougens,
Paris, 366 pp.
1845. Viajes por la América del Sur. Bibli- oteca de Impresos Raros Americanos. Univer- sidad de la Republica, Facultad de Humani- dades y Ciencias, Instituto de Investigaciones Historicas, Montevideo 1982, 318 pp. Cabrera, A. 1958. Catalogo de los mamiferos de
América del Sur.— Revista del Museo Argenti- no de Ciencias Naturales ‘““Bernardino Riva- davia”, Ciencias Zooldgicas 4:1-307.
Desmarest, A. G. 1804. Taubleau méthodique des mammiféres. Pp. 5-38 in Nouveau dictionnaire d@histoire naturelle, appliquée aux arts, princi- palement a l’agriculture, a l’@conomie rurale et domestique: Par une société de naturalistes et d’agriculteurs: Avec des figures tirées 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. 111-115 in B. D. Pat- terson & R. M. Timm, eds., Studies in Neo- tropical mammalogy: essays in honor of Philip Hershkovitz.—Fieldiana: Zoology, new series 39:1-5S06.
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, 1206 pp.
——., &G.K. Creighton. 1989. 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), 216 pp.
Hayman, D.L. 1990. Marsupial cytogenetics. — Aus- tralian Journal of Zoology 37:331-349.
Mann, G. 1978. Los pequenos mamiferos de Chile. — Gayana, 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.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Olfers, I. 1818. Bemerkungen zu Illiger’s Ueberblick der Sdugthiere, nach ihrer Vertheilung tuber die Welttheile, riicksichtlich der Sudamerikanisch- 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 chromosome 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. 1968. The chromosomes of the didelphiid Marmosa robinsoni Bangs.—Experientia 24: 185-186.
— , R. Fernandez, & A. Spotorno. 1972. Further occurrence of a karyotype of 2n = 14 chromo- some in two species of Chilean didelphoid mar- supials.— Zeitschrift fir 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. 621-640 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
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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 7, 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.
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 (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 (HB), 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
7
diameter (ITD), 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, g,, and kur- tosis, g,) 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 61183, 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 1991 and subsequent closing of
16° N—
SOUTH CHINA
SEA
IS? IN =
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
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
been submerged below the Luzon sea during the Pliocene and Pleistocene (Rutland 1968; Hashimoto 1981la, 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 198la, 1981b; Auffenberg 1988).
Site 1.—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 11 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).
Results
Capture data.—A single specimen of S. beyeri collected at Site 1 was a mature male
VOLUME 108, NUMBER 1
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.—A 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 19-21; (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 16 re- ferred specimens). — Details of the head sca- lation of an adult male (PNM 2302; cap- tured by RMB on 16 March 1992 in mon- tane cloud forest at 1460 m, between 1400 and 1630 hr) are shown in Fig. 3 from dorsal (A), ventral (B) and lateral (C) perspectives.
Head scalation.— Rostral 1.2-2.9 (X = 2.2 + 0.4 SD, n = 14) mm wide (holotype
10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Dorsal (A), ventral (B), and lateral (C) per- spectives of head scalation of Sphenomorphus beyeri (PNM male 2302; HL = 12.0 mm). Arrow indicates the presence of the azygous interprefrontal scale on snout.
= 1.8). Frontonasals 1.7-2.4 (X = 2.2 + 0.3 SD, n= 14) mm wide and 1.1-1.6 (¥ = 1.3 + 0.2 SD; n = 14) mm long (holotype = 1.9 x 1.1). Ten of 17 specimens (including holotype) with separate prefrontals, four specimens with prefrontals in narrow to moderate contact, and three with prefron- tals separated by an azygous interprefrontal (Fig. 3). Interprefrontals wider anteriorly; their caudal tip projecting slightly beyond prefrontal/frontal suture.
Frontal triangular, longer than wide, nar- rowed to a point caudally; in contact with two anterior supraoculars; length 2.8-3.7 mm (X = 3.3 + 0.4 SD, n= 14) width 1.7- YS mmm OF = 20) ae 0.4 SD, a= 1) io lotype = 3.1 x 1.9). Frontoparietals fused; D3 .¢\ won OF = D& +2 O.4 SID, m= 1.2) long and 2.5-4.3 mm (X = 3.5 + 0.4 SD, n = 14) wide (holotype = 2.7 x 3.2). Pa- rietals behind interparietal in contact for a distance shorter than the parietal itself. In- terparietals 1.5-2.8 mm (X = 1.9 + 0.5 SD, n = 14) long and 1.2-1.6 mm (X¥ = 1.3 + 0.2 SD, n = 14) wide (holotype = 1.7 x 1.3).
Nasal large and single with nostril at cen-
ter, bordered posteriorly by two pairs of overlapping (superimposed) loreals; most dorsal larger than ventral. Holotype with only 10 supraciliaries; supraciliaries varying considerably among and between our series of specimens: 1 1 — 12 (two specimens), 12— GD ISO GY), WAS Oy, WAH M3 (@), 13—12 (2), 13—13 (3), 13—14 (1), 14—13 ()yand 14= 141() x66 OnO = 0.9 SD; range = 11-14—11-14; n = 16— 16. All specimens with four large supraocu- lars; anterior most strongly triangular; the second widest. Caudalmost supraocular fol- lowed by three or four curved rows of very small scales, each with two or three scales. Tympanum exposed, not strongly de- pressed or sunken. Holotype with 6—7 up- per labials and 6 —6 lower labials. Our series varies considerably in both upper and lower labial scale counts: Upper labials: 6—5 (one specimen), 6—6 (10), 7—7 (4) or 7—6 (1); X—=(63— 6.6 = 05 11S srances—s6= 7—S-7; n = 16—16. Lower labials: 5—5 (1), 5—6 (1), 6—6 (12), 6—8 (1), 7—7 (1), or 8—7 (1); X = 6.3—6.4 + 1.0—1.5 SD; range = 5-8 —5—-8; n = 16—16. Mental chin scale 2.3-3.4 mm (X = 2.8 + 0.3 SD, n= 14) wide and 1.5-2.4 mm (X = 1.9 + 0.3 SD, n = 14) long (holotype = 2.6 x 1.6); followed by one postmental, the latter in contact with two lower labials.
Dorsal scalation.—Nuchal scales undif- ferentiated. Scales (transversely) around midbody = 38 (two specimens), 39 (6), 40 (S, including holotype), 41 (1), and 42 (2); X = 39.9 + 1.17 SD, n= 16. Paravertebrals = 88 (one specimen), 89 (2), 90 (2), 91 (2), 92 (6), 93 (1), 94 (holotype), and 96 (2); X = Wiles) am AS esvOh Tp = 10,
Subdigital lamellae and ventral scala- tion. — All specimens with long digits, as re- flected in lamellae scale counts. Both the holotype and our specimens display a_lon- gest to shortest toes rank of 4, 3, 2, 5, 1. Holotype has 19—19 fourth toe lamellae, whereas our series with 19—19 (two spec- imens), 19—20 (1), 20—20 (5), or 21—21 (8); X¥ = 20.3—20.4 + 0.8—0.7 SD; range
VOLUME 108, NUMBER 1
11
Table 1.— Measurements (in mm) from all known specimens of Sphenomorphus beyeri (character abbreviations in text). Standard univariate statistics, presented below, are abbreviated as follows: X, mean; SD, standard deviation; W, Shapiro-Wilk test for normality (N = normal); g, = skewness statistic; g, = kurtosis statistic. None of the skewness or kurtosis statistics were found to be significant (P > 0.05). CMNH 3654 and the holotype were excluded from the analysis as they were not sexually mature. Tail length (TL) also was excluded from the analysis as several individuals showed scars indicative of caudal autotomy and regeneration.
Character
Specimen # Sex SVL TL AGD HLL HL HB SL ED TD CMNH 3652 f 59.4 76.5 31.4 29.5 12.3 7.8 31) 2.3 1.3 CMNH 3653 f 67.1 Udo 35.0 21.9 9.2 12.9 8.9 2.8 tod CMNH 3654 v 358) 44.1 14.8 123 8.1 5.3 Dog 1.8 le CMNH 3655 m 64.9 90.8! 31.3 24.3 14.4 9.8 4.7 3.1 2.0 USNM 337768 m_ 63.4 56.4! 31.8 24.3 1350 8.7 4.0 Srl 15 CMNH 3657 m 66.7 100.0 34.6 25.0 14.1 8.9 4.6 B52 1.7 CMNH 3658 f Sl 33.5! DIS 20.8 Well 7.4 BoP) Dp N72 CMNH 2359 m 62.8 Vol 32.9 24.1 13.4 9.3 4.3 25 1.5 PNM 2300 m 56.6 70.8 DAES 22.9 12.0 7.6 4.6 od 1ES PNM 2301 ff 46.6 66.4 Wie) 16.9 OD 6.9 3.3 1.6 lee PNM 2302 oy. | (a7) 91.6 33.3 D3%5 13.1 9.1 4.6 3.1 1S PNM 2303 m 48.0 49.1! 21.8 20.3 10.8 6.5 Sod) 2 1.1 PNM 2304 fy 55.8 69.8 28.6 20.3 11.8 7.4 4.1 Dee) 1.4 PNM 2305 m 56.5 80.4 29.8 DDS) IDES 8.2 4.5 2.8 1.5 PNM 2306 ol Oa 61.8! 30.3 22.4 W2e7/ 8.2 4.1 2.6 1.3 PNM 2307 fe 50.0 62.4 26.3 18.1 2 13. 3.8 2.4 NZ CAS 61183* m 43.0 _ 20.9 17.1 Wo3) 6.5 3.9 325 2.4 X 58.9 ~ 30.2 22.3 1225 8.2 4.2 Def 1.5 SD 6.1 — 3.6 2.0 1.1 0.9 0.4 0.3 0.2 Range 19.2 — Sot To 3.6 3)3} 1S 0.9 0.9 n 14 — 14 14 14 14 14 14 14 WwW N — N N N N N N N £1 =0.39) — —0.83 —0.60 =0),05 =){05 —0.67 i209 —0.66 Zo —=(0),2)1 — —0.86 —0.02 —0.66 =(0)i331 O07 =I —0.42
1 Tail recently autotomized or showing scars of caudal autotomy and subsequent regeneration.
* Holotype.
= 19-21—19-21; n = 16—16. Holotype with 5—5 first finger lamellae and ours have 5—5 (five specimens), 6—6 (7) or 7—7 (4); Nee) 2 323) SID. range — 6— 7—5-7; n= 16—16. Preanal scales strongly enlarged.
Body size.—Table 1 contains the mor- phological measurements taken from all known specimens of S. beyeri. Body size proportions of holotype and our new series are discussed below.
Coloration. —Color plates (Fig. 4) contain lateral views of male PNM 2302 in life. Dorsal surfaces dark umber to lavender brown. A mid-vertebral line of very dark
brown to black spots variably obvious; ver- tebral spots (or line) at midbody usually en- compassing one scale only, but occasionally two or three scale rows; spot series (or line) fading caudally on some specimens. Very dark brown to black series of irregular blotches laterally, usually forming a solid band, strongest at the pre-nuchal region, next strongest above hind limbs at pectoral gir- dle; band extending anteriorly through tym- panum to rostral scale and caudally to thick- est portions of tail base (‘4 total tail length). Dorsally, lateral band bordered by a thin golden yellow edge; ventrally, golden yellow edge becoming solid yellow on caudal por-
12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Sphenomorphus beyeri in life before preservation (PNM male 2302; SVL = 62.7 mm). Note presence of grey volcanic ash on substrate. Photographs by RMB, courtesy of CMNH.
VOLUME 108, NUMBER 1
tions of body. Laterally, axilla-groin region bright orange with round yellow markings arranged in spots. Distinctive black tip on tail of all complete specimens. Undersides of arms and legs bright golden yellow. Pos- terior regions of hind limbs with distinct yellow spots surrounded with brown bor- ders fading into a lavender brown back- ground. Ventral side of body from nuchal region to tail golden yellow with pinkish tan, light grey, or very light sky blue flecks on nuchal region, ventral side of head, torso, and tail base. Lavender brown series of spots arranged into irregular, disjunct stripes of four to eight scales in length and one scale in width on ventral side of neck. Chin pink- ish tan or translucent. Labial scales lavender to purplish, each with white spot in center or on ventral border of scale.
In alcohol: coloration generally faded throughout; ventral yellow and lateral or- ange fading to dull pinkish tan; ventral nu- chal region devoid of blue tones; purple on labials fading to lavender or brown.
Comparisons. — Table 2 compares S. be- yeri with closely related, sympatric, and/or morphologically similar species of Sphen- omorphus.
Discrepancies with Taylor’s original de- scription. —Taylor’s (1922b) description of S. beyeri stated that the holotype exhibited separated prefrontals, as do many of ours (10 of 16). However, three of our series have prefrontal scales in narrow to moderate con- tact and three others show the presence of an azygous interprefrontal (Fig. 3). In ad- dition, Taylor (1922b) counted 40 scales at midbody in the holotype but did not record paravertebrals. Brown & Alcala (1980) and Alcala (1986) confirmed his midbody scale counts, and reported also that there were 94 paravertebral scales in the holotype. Our examination of the holotype confirms both of these reports, which fall within the range of variation for the new series (88—96; Table 2). Taylor (1922b) counted 17 lamellae un- der the holotype’s longest (=fourth, rear) toe, but Brown & Alcala (1980) and Alcala
13
(1986) reported that the holotype had 19 fourth toe lamellae. Our examination of the holotype confirms that the specimen has 19—19 subdigital lamellae under the fourth toe. Taylor (1922b) reported a SL/HL ratio for the holotype of 0.34; calculations for our series are remarkably close (¥ = 0.32, range = 0.25-0.37; n = 16). Taylor (1922b) re- ported that the SL/HB ratio was 0.60 for the holotype; our series averaged 0.49 (range = 0.38-0.57; n = 16). Taylor’s specimen had a HB/HL ratio of 0.57, and ours average 0.66 (range = 0.60-0.69; mn = 16). Taylor reported a HB/SVL ratio of 0.15; our spec- imens average 0.14 (range = 0.13-0.17; n = 16). The holotype also had an ED/SL ratio of 0.50, but the eyes on our specimens seemed somewhat smaller on average (X = 0.37; range 0.30-0.43; nm = 16). Disparities in these calculated ratios may reflect the small sample size (n = 1) available to Tay- lor; the distinctive black tip on the tails of all complete specimens undoubtedly would not have been missed by Taylor had he been able to recover the portion of the holotype’s tail which was autotomized and lost in cap- ture.
Remarks.—Two of the Zambales speci- mens were gravid females (CMNH 3652 and 3653), each containing two yellowish, thick- ly-shelled, oviductal eggs, the texture of which suggest oviparity. One of our speci- mens appears to be recently hatched (CMNH 3654). It is well within the range of mor- phological variation described here for S. beyeri except that it has unfused frontopar- ietals. As stated above (site descriptions), both areas surveyed in this study contained ash deposits from the June 1991 eruption of Mt. Pinatubo. This rendered the forest floor, vegetation, and much of the remain- ing habitat very dry (Fig. 2). Where water might be expected to collect (i.e., in Pan- danus axils or other depressions in rocks or vegetation), the ash rapidly absorbed water deposited by rain or dew. Taylor (1922b) mentioned that he found the holotype on a rock ledge on Mt. Banahao; none of our
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
14
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VOLUME 108, NUMBER 1
specimens were collected anywhere near rocks or outcroppings in the Zambales but, instead, from inside moist rotten logs. Only two specimens were caught in our extensive pitfall trap system, suggesting either low lev- els of activity or semi-fossorial existence. However, due to the disturbed nature of the forest on Mt. High Peak, it is impossible to establish the true microhabitat of S. beyeri. The apparent semi-fossorial nature of this species in the Zambales could have been due to animals retreating from the aridity of the immediate area which was blanketed by Mt. Pinatubo’s ash.
At present, S. beyeri is known to exhibit an allopatric insular montane distribution. However, little biogeographical informa- tion can be inferred from this fact alone due to lack of reliable survey data for southern Luzon, specifically the areas between the two localities discussed here. While the two mountain ranges included in this species’ locality records are located on separate geo- logic components of Luzon, adequate hab- itat connecting these insular regions prob- ably existed during the last glaciation (Rut- land 1968; Hashimoto 1981la, 1981b; Auf- fenberg 1988). Mountains like Mt. Makiling and Mt. Arayat may support undocumented populations of S. beyeri as suggested by their intermediate placement between the two known localities (Mt. High Peak and Mt. Banahao). Studies of poorly known high el- evation montane environments (e.g., Brown & Alcala 1961b, Custudio 1986, Auffenberg & Auffenberg 1988) are needed in such areas in and between these two disjunct localities to ascertain the true distribution of S. beyeri as well as other relict species. While ele- vational species succession of scincid liz- ards (Custudio 1986) and small mammals (McCoy & Connor 1980, Rapoport 1982, Rickart et al. 1991, Ruedas et al. 1994) has
—_—
15
recently been documented in some areas of the Philippines, little is known about the potential effects of elevational gradients on lizard species stratification, diversity, and endemism in the Philippine archipelago. Our unpublished survey data suggest that spe- cies diversity 1s inversely proportional to elevation, but that endemism is positively correlated with elevation at least on the is- lands of Luzon, Panay, Mindanao, and Mindoro. The latter topic is subject of an- other work in progress (Ferner, pers. comm.).
Acknowledgments
Collecting permits were facilitated by the Protected Areas and Wildlife Bureau of the Philippines Department of the Environ- ment and Natural Resources, especially by A. Alcala, A. Ballesfin, J. Caleda, and C. Catibog-Sinha. J. F. Barcelona, J. R. Dem- boski, R. E. Fernandez, and F. T. Wacdisen assisted in the collection effort and capture of several specimens. We thank R. I. Crom- bie for his help in obtaining the S. beyeri type specimen from the California Acade- my of Science, for facilitating our work at USNM, and for confirming the identity of our S. beyeri specimens along with W. C. Brown. Statistical analyses were carried out by L. A. Ruedas who also provided the map used in the manuscript. This study was funded by a grant from the John T. and Catherine C. MacArthur Foundation (to P. C. Gonzales and R. S. Kennedy) and by the following grants from the Miami University community (Oxford, Ohio) to RMB: The Zoology Student Enrichment Fund Grant, the Roschman Student Enrichment Fund Grant, the Undergraduate Research Fund Grant, and the Hefner Museum of Zoology Independent Studies Grant. JWF acknowl- edges the continued support of Thomas
4 Differs from S. a. abdictus by the presence of four (as opposed to five) supraoculars. > Differs from S. c. coxi by the presence of four (as opposed to five) supraoculars and coloration.
© New species (Brown et al. 1995, this issue).
16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
More College. Equipment was made avail- able to us by H. Eshbaugh and G. Kritsky; outdoor apparel and other gear was provid- ed by Outdoor Adventures in Cincinnati, Ohio. Comments on preliminary drafts of the manuscript were provided by R. I. Crombie, R. F. Inger, S. M. Moody, and L. A. Ruedas. This paper is Contribution No. 3 to the Results of the PNM/CMNH Bio- diversity Inventory of the Philippines.
Literature Cited
Alcala, A. C. 1986. Guide to Philippine flora and fauna. Vol. X, Amphibians and reptiles. Natural Resource Management Center Ministry of Nat- ural Resources and University of the Philip- pines, xv + 195 pp.
Auffenberg, W.G. 1988. Gray’s monitor lizard. Uni- versity of Florida Press, Gainesville, iv + 419 pp.
— .,&T. Auffenberg. 1988. Resource partitioning in a community of Philippine skinks (Suaria: Scincidae).— Bulletin of the Florida State Mu- seum, Biological Sciences 32(2):150—219.
Brown, R. M., J. W. Ferner, & Luis A. Ruedas. 1995. A new species of lygosomine lizard (Reptilia: Lacertilia: Scincidae; Sphenomorphus) from Mt. Isarog, Luzon Island, Philippines. — Proceedings of the Biological Society of Washington 108:18— Dy.
Brown, W.C., & A.C. Alcala. 1956. A review of the Philippine lizards of the genus Lygosoma (Leiol- opisma).—Occasional Papers of the Natural History Museum, Stanford University 3:1-10.
—,,& . 1961a. A new sphenomorphid liz- ard from Palawan Island, Philippines. —Occa- sional Papers of the California Academy of Sci- ence 32:1-4.
——.,,& . 1961b. Populations of amphibians and reptiles in submontane and montane forests of Cuernos de Negros, Philippine Islands. — Ecology 42(4):628-636.
—,, & 1970. The zoogeography of the herpetofauna of the Philippine islands, a fring- ing archipelago. — Proceedings of the California Academy of Science, fourth series 4(38):105- 130.
——.,& . 1980. Philippine lizards of the fam- ily Scincidae. — Silliman University Natural Sci- ence Monograph Series 2. 264 pp.
—,&D.S.Rabor. 1967. A new sphenomorphid lizard (Scincidae) from the Philippine islands. — Proceedings of the Biological Society of Wash- ington 80:69-72.
Bureau of Mines, Philippines, in coordination with the Board of Technical Surveys and Maps. 1963. Geological map of the Philippines, edition 1. Single sheet map.
Custudio, C. C. 1986. Altitudinal distribution of liz- ards of the Scincidae in Mt. Makiling, Laguna. — Sylvatropical Philippine Forest Research Jour- nal 11(3, 4):181-202.
Dickerson, R. E. 1924. Tertiary paleogeography of the Philippines. — Philippine Journal of Science 25(1):10-55.
Greer, A. E. 1979. A phylogenetic subdivision of Australian skinks.—Records of the Australian Museum 32:339-371.
—, & F. Parker. 1974. The faciatus species group of Sphenomorphus (Lacertilia: Scincidae): notes on eight previously described species and de- scriptions of three new species. — Proceedings of the Papua New Guinea Science Society 25:31- 64.
Hashimoto, W. 198la. Geologic development of the
Philippines. Pp. 83-170 in T. Kobiyashi, R. To-
riyama, & W. Hashimoto, eds., Geology and
paleontology of Southeast Asia, CCXVII, Vol.
DOP
1981b. Supplementary notes on the geologic development of the Philippines. Pp. 171-190 in
T. Kobiyashi, R. Toriyama, & W. Hashimoto,
eds., Geology and paleontology of Southeast
Asia, CCXVIII, Vol. 22.
McCoy, E. D., & E. F. Connor. 1980. Latitudinal gradients in the species diversity of North American mammals.—Evolution 34:193-203.
Myers, W. C., & M. A. Donnelly. 1991. The lizard genus Sphenomorphus (Scincidae) in Panama, with a description of a new species. — American Museum Novitates 3027:1-12.
Rapoport, E. 1982. Areography: geographical strat- egies of species. Pergamon Press, New York, xvi + 269 pp.
Rickart E. A., L. R. Heaney, & R.C. Utzurrum. 1991. Distribution and ecology of small mammals along an elevation transect in Southeast Luzon, Philippines. —Journal of Mammalogy 72:458- 469.
Ruedas, L. A., J. R. Demboski, & R. V. Sison. 1994. Morphological and ecological variation in Otop- teropus cartilagonodus Kock, 1969 (Mammalia: Chiroptera: Pteropodidae) from Luzon, Phil- ippines. — Proceedings of the Biological Society of Washington 107:1-16. ;
Rutland, R. W. 1968. A tectonic study of part of the Philippine Fault Zone. — Quarterly Journal of the Geological Society of London 123(4):293-325.
SAS Institute Inc. 1988a. SAS/STAT user’s guide, release 6.03 edition. SAS Institute Inc., Cary, North Carolina, 1028 pp.
VOLUME 108, NUMBER 1 7
1988b. SAS procedures guide, release 6.03
. 1922b. Additions to the herpetological fauna
edition. SAS Institute Inc. Cary, North Caroli- of the Philippine Islands, II.— Philippine Jour- na, 441 pp. nal of Science 21(3):253-303. Sokal, R. R., & F. J. Rohlf. 1981. Biometry, second UNESCO/ECAFE. 1971. Geologic map of Southeast edition. W. H. Freeman and Co., New York, Asia. United Nations Publication, No. 69-30632. Xvlil + 859 pp. Whitmore, T. C. 1984. Tropical rain forests of the Taylor, E. H. 1922a. The lizards of the Philippine Far East. Clarendon Press, Oxford, England. xvi islands.— Monograph of the Bureau of Science, + 718 pp.
Manila, 17:1-115.
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108(1):18-28. 1995.
A new species of lygosomine lizard (Reptilia: Lacertilia: Scincidae; Sphenomorphus) from Mt. Isarog, Luzon Island, Philippines
Rafe M. Brown, John W. Ferner, and Luis A. Ruedas
(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.; (LAR, RMB, JWF) Department of Vertebrate Zoology, Cincinnati Museum of Natural History, 1720 Gilbert Avenue, Cincinnati, Ohio 45202-1401, U.S.A.; (LAR) Department of Biology, Cayey University College, Cayey, Puerto Rico 00736
Abstract. —Sphenomorphus knollmanae, a new species, is described on the basis of recently collected material from Mt. Isarog, Bicol Peninsula, south- eastern Luzon, Philippines. The small series (7 = 5) differs from its congeners by the combination of its fused frontoparietals, relatively low number of par- avertebrals (73-83) and midbody scales (34-39), the presence of 17—20 sub- digital fourth toe lamellae, distinctive patterns of coloration, and a host of measurements related to its small body size (SVL = 47.5-51.0 mm). To better distinguish between the new species and two closely related congeners, uni- variate and multivariate analyses were performed on a suite of morphological characters. The three species were found to be well differentiated morpholog-
ically.
Worldwide, the genus Sphenomorphus contains over 120 species and is a “taxo- nomically residual’? plesiomorphic taxon that “remains a convenient repository for
. species, pending further phylogenetic analysis’”> (Myers & Donnelly 1991:2). Brown & Alicala (1961b) reported that Ori- ental and Australian zoogeographic regions contain over 60 scincid species in Spheno- morphus. In their key to Philippine Scin- cidae, Brown & Alcala (1980) recognized 22 species of Sphenomorphus, subdividing these into five groups based on external morphology. The Group I species of Phil- ippine Sphenomorphus are S. beyeri (Taylor 1922) and S. diwata (Brown & Rabor 1967, see Brown & Alcala 1980, for review). Until recently, S. beyeri was known only from the holotype, collected by E. H. Taylor on Mt. Banahao, Laguna province, southern Luzon Island (Taylor 1922). During a recent bio- diversity inventory of the Philippines con-
ducted by the National Museum of the Phil- ippines (PNM) and the Cincinnati Museum of Natural History (CMNH), we rediscov- ered and redescribed Sphenomorphus beyeri from specimens taken on Mt. High Peak, Zambales Mountains, west central Luzon Island (Brown et al. 1995). Sphenomorphus diwata also is currently known only from a small number of specimens collected in the Diwata Mountains, Surigao del Sur Prov- ince, northern Mindanao Island (Brown & Rabor 1967, Brown & Alcala 1980). While examining material in the United States National Museum of Natural History (USNM), R. I. Crombie brought to our at- tention a small series of Sphenomorphus skinks that appeared very similar to our specimens of S. beyeri from the Zambales. At the time, we were not confident in the assignment of these specimens to our con- cept of S. beyeri (from the type locality or from Mt. High Peak) as several inconsis-
VOLUME 108, NUMBER 1
tencies immediately were apparent. Follow- ing detailed examination of these specimens and a host of univariate and multivariate statistical analyses we concluded that dif- ferences between this series and its most closely-related congeners were sufficient to warrant its recognition as a distinct species.
Methods
Morphological characters and scale counts used here follow definitions and abbrevia- tions in Brown & Alcala (1980) and Brown et al. (1995). Measurements were taken to the nearest 0.1 mm with digital calipers. All measurements are based on specimens pre- served in 70% ethanol. In cases where scales of interest are found on both sides of the head (e.g., labials), scale numbers are given in pairs, separated with a long dash (—), designating left from right respectively. Mensural and meristic character abbrevia- tions (defined in Brown et al. 1995) include: snout-to-vent length (SVL), tail length (TL), axilla-groin distance (AGD), hind leg length (HLL), head length (HL), head breadth (HB), snout length (SL), eye diameter (ED), tym- panum diameter (TD), paravertebrals (PVS), midbody scales (MBS), supralabials (SUL), and infralabials (IFL). Specimens examined are deposited in the California Academy of Science (CAS) the Cincinnati Museum of Natural History (CMNH), the National Museum of the Philippines (PNM), and the United States National Museum of Natural History (USNM).
Statistical analyses were performed using the Statistical Analysis System software, version 6.03 (SAS Institute Inc., 1988a, 1988b). Sexually immature specimens (S. beyeri, PNM 2303 and CMNH 3654, CAS 61183; S. diwata, CAS 133514; Spheno- morphus sp., USNM 318343) were exclud- ed from univariate and multivariate anal- yses. A Student-Newman-Keuls multiple range test was performed on both raw and log (base 10) transformed data to determine patterns of significant character variation. Two principal component analyses were
19
performed, both on the correlation matrix of the variables. The first included only raw (untransformed) data; the second was car- ried out on the log (base 10) transformed data, in order to minimize the effects of size differences among the different populations examined herein; in the case of the log (base 10) transformed analysis, the size compo- nent of the variation is restricted to prin- cipal component axis one. In both in- stances, the first and second and the first and third principal component scores were then plotted in order to ascertain morpho- logical differentiation among groups.
Results
Sphenomorphus beyeri, S. diwata, and S. n. sp., distinctly segregated into discrete groups in the principal component analysis (Fig. 1). In the PC analysis based on raw data (Fig. la, b), principal component one differentiates between S. n. sp. and S. diwata and between the new species and S. beyeri. This component loads heavily on HL, SVL, HB, HLL, AGD, and SL. Principal com- ponent two distinguishes between S. diwata and remaining Group I Sphenomorphus. This component loads heavily on fourth toe lamellae and MBS as well as TD, ED, and PVS. The third principal component differ- entiates between S. beyeri and remaining Group I Sphenomorphus. This component loads primarily on fourth toe lamellae, PVS, and TD. Together, the first three principal components account for 85.7% of the vari- ation (PC I, 55.0%; PC II, 16.7%; PC III, 14%).
In the PC analysis based on the log trans- formed data (Fig. lc, d), principal compo- nent one and two discriminate between S. n. sp. and S. diwata, while principal com- ponents two and three discriminate between S. diwata and S. beyeri. Principal compo- nent three also discriminates between S. be- yeri and the new species. The first four prin- cipal components account for 91.6% of the variation (56.4, 16.8, 13.2, and 5.2, respec- tively). Factor loadings along the first prin-
20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
pe)
—s
-4
4 2 0 -2 0 PC III PC II
Fig. 1. Plots of principal component scores for two species of Group I Sphenomorphus and the new species examined in this study. Component I versus compo- nent II (a); component I versus component III (b); component I versus component III for log (base 10) transformed data (c); component I versus component III for log (base 10) transformed data (d). Symbols are: squares = S. beyeri; triangles = S. diwata; circles = S. knollmanae, new species.
cipal component are relatively homoge- neous, indicating that the size variation gen- erally has been isolated to this component, while shape is more important among re- maining principal components. High load- ings 1n principal component two are shown in discrete characters (scale row counts). An additional PC analysis carried out on the correlation matrix of the log (base 10) trans- formed variables but using only measure- ments (not shown), indicates that tympa- num and eye diameters were the heaviest contributors to the variation in PC axes two and three.
An important point to make regarding these analyses concerns the orientation of the principal component axes of each pu- tative species group’s dispersion in multi- variate space. Orthogonal orientation of the axes 1n multivariate space has been inter- preted as indicative of differing allometric growth patterns (Voss et al. 1990, Voss & Marcus 1992); a corollary of the foregoing is that different orientations of the principal component axes are thereby good evidence of distinct specific status. In the case of the
analyses of the populations of Sphenomor- phus examined herein, it is quite clear that the orientation of the axes of dispersion are quite distinct both in analyses based on raw data as well as log (base 10) transformed data. This particular distinction 1s especially severe between S. beyeri and remaining Sphenomorphus examined and is clearly ob- served in the raw data plots, but more sig- nificantly in the log (base 10) transformed data, which minimizes the contribution of size to principal component axis one.
In view of the quantum separation in al- lometric growth patterns, as well as mor- phology between the Mt. Isarog Spheno- morphus, S. beyeri, and S. diwata in both multivariate (PCA) and univariate analyses (ANOVA) of discrete and continuously varying characters, we describe the series from Mt. Isarog as:
Sphenomorphus knollmanae, new species Figs. 2, 3
Holotype. —PNM 2311 (formerly USNM 318342), adult male, collected by L. R. Hea- ney on | May 1988, in loose leaf litter along- side a fallen, partially decomposed log on the forest floor in primary mid montane forest at 1125 m on Mt. Isarog (Philippines, S. Luzon, Bicol Peninsula, Camarines Sur Prov.), 4.5 km N, 20.5 km E Naga City, 13°40'N, 123°22’E (map: fig. 1 in Goodman & Gonzales 1990).
Paratypes.—(4) USNM 318341 (female) and 318343 Guvenile), same data as above except as follows: USNM 318341, collected by L. R. Heaney on 29 Apr 1988; USNM 318343, collected by S. M. Goodman, 22 Mar 1988. USNM 318344 and CAS 191800 (formerly USNM 318345) collected by S. M. Goodman, 19 Mar 1988, and by R. C. B. Utzurrum on 20 Mar 1988 respectively, at 4 km N, 21 km E Naga City, 13°40’N, 123°22’E, at 1350 m in primary upper mon- tane forest. All specimens collected in loose leaf litter and loose topsoil on forest floor; USNM 318344 and CAS 191800 associated
VOLUME 108, NUMBER 1
with fallen, partially decomposed logs on forest floor.
Etymology. —Named in honor of the late Margy Knollman, friend and teacher, who guided the senior author through his first scientific experiment at age seven and con- tinued to encourage his herpetological pur- suits until the time of her death in Novem- ber 1989.
Diagnosis. —A small to moderate species of Sphenomorphus (SVL, 47.5—51.0 mm) differing from its congeners by a combina- tion of the following characteristics: fron- toparietals fused; prefontals separate, in contact, or with azygous interprefrontal; 73 to 83 paravertebrals; 34 to 39 scales around midbody; 17 to 20 subdigital fourth toe la- mellae; unique coloration (see below).
Description of holotype.—(PNM 2311) Total Length, 118.7 mm; SVL, 48.7 mm; nie 0.0mm, 115) mime? SL, 3:9 mm: HB, 7.1 mm; ED, 3.1 mm; lower eyelid scaly with translucent window, oval in shape, ar- ranged horizontally; ear opening and tym- panum exposed, not deeply sunken, verti- cally oval, 1.2 mm in width; ear opening without spines or lobules; limbs pentadac- tyl, well developed; HLL, 18.3 mm; AGD, 22.7 mm; head (viewed from above) ta- pered, snout rounded dorsally and laterally; dorsal, lateral, and ventral scales smooth, unstriated; rostral large, visible from above, broader (1.8 mm) than long (0.7 mm), form- ing a curved suture with frontonasal; latter wider (1.5 mm) than long (1.1 mm); pre- frontals in broad contact; frontoparietals fused (2.8 mm wide, 1.8 mm long) frontal moderate, rhomboidal, pointed caudally, 1.9 mm wide, 3.0 mm long, in contact with two supraoculars; interparietal moderate, point- ed, 1.3 mm wide, 1.9 mm long; parietals in contact behind interparietal; nasals large and single with round nostril at centers, widely separated by frontonasal and nasal bordered caudally by two pairs of overlapping loreals, dorsal pair slightly larger than ventral; 3— 3 large preoculars, most ventral contacts su- ture between third and fourth labial; 4—4
21 oo LK 4 heheh “Sih eG
Fig. 2. Sphenomorphus knollmanae holotype (PNM 2311)—Lateral (A), and dorsal (B) head scalation and subdigital lamellae, right hind foot (C).
large supraoculars, anteriormost triangular, second widest; last supraocular followed by 3 rows of small scales clustered in postoc- ular region, each row containing 2-3 scales; 3—3 temporals, dorsalmost wraps one- fourth of way around posterior edge of pa- rietal scales; nuchals undifferentiated, ex- cept for most lateral pair which is very slightly enlarged; 2—2 rows of small scales between eye and labials; 12—12 supracili- aries; 15—16 lower ciliaries; 6—6 suprala- bials; 6—6 infralabials; 73 paravertebrals; 34 midbody scales; 20 subdigital fourth toe lamellae; 5 first finger lamellae; toe length (shortest to longest) 4, 3, 2, 5, 1; two strongly enlarged preanal scales apparent; mental chin scale followed by single postmental bordered caudally by two pairs of chin shields; subcaudals only slightly larger than ventrals. The holotype had a live weight of 3.0 g. Body size proportions and coloration discussed below.
Coloration. — Field notes recorded by R. C. B. Utzurrum (courtesy L. R. Heaney, Field Museum of Natural History) state that in life CAS 191800 had a “golden venter, dark brown dorsum, mottled on sides.” In alcohol, dorsal surfaces very dark brown with black spots and a darkly pigmented (=2-3 scale rows) black mid-vertebral line. Mid-vertebral line darkest on CAS 191800,
22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
22 S) sé en NS ez l2 Oe 6! i i é\ ol g| 71 a el il Ol 6 8 i SS) S 7 £ ty Naliidtedlndt hon usdvnbselsusotsnlastunebvittisshsntiscticsteuabsaticatintisalinatscalioutiesdinetinstes sve tiaptavsrlnetiey
Bee cemOCe Gc Vcc ccmlcer: Ocgirol ieaeee| 91 i] 7\ gi 2! WW Ol 6 8 Z 9 Ss 7 € mint iyntunluntonhuntantoutwubralidiutonfostysbotusluebuolotonbuntunbaertidstouelvvtuliutte valent aytirdyutaatonlntunlutadaduntystuonaty
Fig. 3. Dorsal (top) and ventral (bottom) views of the type series of Sphenomorphus knollmanae placed underneath five specimens of S. beyeri for comparison.
VOLUME 108, NUMBER 1
23
Table 1.— Morphological measurements and scale counts taken from all known specimens of Sphenomorphus knollmanae (see text for abbreviations of characters). Standard univariate statistics are presented below as means + one standard deviation (sexually mature specimens only).
Character
Specimen # Sex SVL TL AGD HLL HL HB SL ED TD PVS’' MBS_ SUL _ IFL 4th Toe MISINIMESISS842 7 -f 49°78 71:5 24-97 18:5. 1010 6.2 3.6 2:5 1.2 76 39 7-7 6-6 19 PNM 23112 ma AS VOW)» 22 MSS lle Wel Be SSE wales 7/8} 34 6-6 6-6 20 USNM 318343 ? 47.5 Sirol 4 Selo) NOLO O.00 395.25) 1E2, 83 39 7-7 6-6 19 WSNM 3183445 of S10 41.4 26.5 17.2 10:0. 6.4 3.5 2.7 1.4 76 37 7-7 6-6 17 CAS 191800 f 50.4 SOD SH Sse eR) 6:4 32 27, ICS 76 35 6-6 7-7 18
Mean 49.1 — DSS 4 Ost OsOm Seige.) eles) OPN 36-5) 36:5) 62317 18-5 SD 1.36) = Did a OMtte ORG, OFS SOSH HOLD 083) PA2he 92845. 0:6 (0:5 1.3 @ Holotype.
> Tail autotomized and partially regenerated.
but also heavy on PNM 231 and USNM 318341, somewhat lighter (1-2 scale rows) on USNM 318343 and 318344. Mid-ver- tebral line ending abruptly at pectoral girdle where dorsal mottling coalesces into trans- verse bars that fade caudally (this pattern not apparent in USNM 318343). Laterally, with a heavy series of black blotches, form- ing a solid stripe in canthal region, and ex- tending posteriorly from nostril, through eye and typanum, to groin. Lateral black stripe anteriorly bordered ventrally by a distinct white line intersecting the tympanum at one- half its height and extending from caudal edge of eye to region dorsal to forearm. Lat- eral white line on midsection breaking up into series of white spots that continue cau- dally through anterior one-third length of tail. Ventral surfaces in complete specimens pale yellow from chin to tip of tail (USNM 318344 has a regenerated tail that is com- pletely black; USNM 318343 and CAS 191800, with autotomized tails). Throat pale yellow with dark umber flecks (darkest on CAS 191800 and almost invisible on ho- lotype and USNM 318341). In specimens with heavy speckling, pattern wraps around onto lateral portions of neck, extending to approximately one-half the height of the tympanum.
Variation. —Our sample includes one male, three females and a sexually undeter-
mined juvenile. USNM 31834 and CAS 191800 both were gravid at the time of pres- ervation, each containing two thinly-shelled eggs. USNM 318341 may have been gravid when preserved (remnants of what appear to be eggs remain), but some breakdown of the ovaries has occurred. Table 1 contains morphological measurements of the five specimens of S. knollmanae and characters and measurements differing from holotype description follow below.
Variation in head scalation is as follows: FOstraleles= 22) (Xe— 1S = 0.408): 0 — 4) mm wide; frontonasal 1.5-1.8 (1.7 + 0.1; n = 4) mm wide and 1.0-1.1 (1.1 + 0.1; = 4) mm long; frontoparietal 2.5—2.9 (2.7 + 0.2; n = 4) mm wide and 1.7-2.1 (1.9 + 0.1; n = 4) long; frontal 0.9-1.9 (1.5 + 0.4; n = 4) mm wide and 1.5-3.0 (2.4 + 0.67; n= 4) mm long; interparietal 1.1 to 1.4 (1.2 + 0.1; n = 4) mm wide and 1.7 to 1.9 (1.8 + 0.1) mm long. Prefrontals are in broad contact (three specimens) or separated by an azygous interprefrontal (one specimen), the latter somewhat wider anteriorly where contacts frontonasal; increasingly narrow caudally where its most caudal border ex- tends slightly beyond margin delineated by contact between prefrontal and frontal. Su- praciliaries 12-16 (¥ = 13.4 + 1.5 SD; n= 5) on right and 12-15 (2.6 + 1.3; n= 5) on left; lower ciliaries, 5-18 (16.2 + 1.l; n=
24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 2.—Comparisons of selected measurements and scale counts for Sphenomorphus knollmanae and two closely related congeners. Presented below are standard univariate statistics (means + one standard deviation, sexually mature specimens only) and results of Student-Newman-Keuls multiple range tests. Superscript letter by means indicates group assignment (means with same letter are not statistically different at the P > 0.05 level).
See text for definitions of abbreviations used in this table.
Character
4th
SVL TL AGD HLL HL HB SL ED TD PVS-' MBS_~ SUL _ IFL_ Toe S anova X AGIE = O52 IIBZE NOs bce Sse ZIP 1.3? T/0P B6.5° 65 63 18.5% n=4 SD VAS) We 25 0.1 OG Ws Os OLA Os 47 2:4 » (0:6 055 ay es S. beyeri 2X SBOB = ZO22 WEP MDS Bye ae Bye 5° OBE BOS 6.5 GA 20.3" n= 14 SD S621 7 = 316 2.0 hei O9) OA Osss - O27 1 1.2 O18 Ld S088 S. diwata 26 SAP = 5G DAD Ngj.se OP Ase Bie Die See alse 55.5 5.5 13.5 n= SD 47 — 5.4 8.6 0.4 06 06 06 0.8 1.4 Aes] OL O55) OL7/
5) on right and 14-17 (16.0 1.27 7 — 5) on left; supralabials 6-7 (6.6 + 0.5; n = 5); infralabials 6-7 (6.2 + 0.4; n = 5) on right and 5-6 (5.8 + 0.4; n = 5) on left.
In specimens with regenerated tails (USNM 318343 and CAS 191800), normal subcaudals replaced by a series of narrow scales that cover entire ventral surface of regenerated tail. Besides holotype, only CAS 191800 was weighed before preservation (3.4 g).
Ratios of morphological measurements for the series (holotype in parentheses) are as follows: SL/HL, ¥ = 0.34 + 0.03, range = 0.29-0.36 (0.35); SL/HB, ¥ = 0.55 + 0.03, range = 0.50-0.58 (0.55); HB/HL, X¥ = 0.61 + 0.03, range = 0.60-0.64 (0.64); BV OMIL, 26S 0.13 = O01, memes = 0,12 0.15 (0.15); HL/SVL, X¥ = 0.21 + 0.01, range = 0)20=0123(0. 23); BD/SID X0— 077) =: 0.06, range = 0.71-0.84 (0.77); ED/HB, X = 0.42 + 0.01, range = 0.41-0.44 (0.41); AGD/SVL, X = 0.52 + 0.04, range = 0.47- 0.57 (0.47); HLL/SVL, X = 0.36 + 0.02, range = 0.34—-0.38 (0.37).
Comparisons. —Table 2 compares S. knollmanae with Group I Sphenomorphus species. Excepting its low number of par- avertebrals, S. knollmanae adheres to the gestalt of Group I Sphenomorphus members (Brown & Alcala 1980); accordingly, the new species appears closely related to S. beyeri and S. diwata. Besides size and characters
He I+
listed in Table 2 of this study, S. knoll- manae differs from the former by coloration and disposition of color pattern (Brown et al. 1995). It differs from the latter by char- acters in Table 2 and also in that it invari- ably has only 4 supraoculars (vs. 5-6 in S. diwata) and fused frontoparietals (vs. 2 in S. diwata; Brown & Rabor 1967, Brown & Alcala 1980). As in S. beyeri (Brown et al. 1995) contact, or lack thereof, between the prefrontal scales is not fixed in this species as itis in S. diwata (Brown & Alcala 1980). The azygous interprefrontal scale exhibited by USNM 318344 also is apparent in some specimens of S. beyeri (Brown et al. 1995), but not in any known specimens of S. di- wata (Brown & Rabor 1967, Brown & Al- cala 1980).
The low number of paravertebrals in S. knollmanae assigns this species to Group III of Brown & Alcala’s (1980) key; accord- ingly, comparisons with S. /eucospilos and S. laterimaculatus, as well as S. decipiens, are warranted. Differences between S. knollmanae. S. leucospilos, and S. decipiens are as follows: Sphenomorphus knollmanae distinct by its 73-83 paravertebrals (vs. 63- 68 in S. leucospilos and 57-66 in S. deci- piens), 34-39 scales around midbody (vs. 32 in S. leucospilos and 32-38 in S. deci- piens), and 17-20 fourth toe lamellae (vs. 14-18 in S. decipiens). Sphenomorphus de- cipiens also has a smaller overall body size
VOLUME 108, NUMBER 1
(SVL = 31-45 mm; Brown & Alcala 1980) than S. knollmanae. Differences in color pattern and body proportions between these three species also are apparent (Brown & Alcala 1980).
Body measurements and scale counts of the single known S. /aterimaculatus speci- men are very close to the range of variation of both S. leucospilos and S. knollmanae (see Brown et al. 1995). While the range of paravertebrals and midbody scales in S. knollmanae do not overlap with S. /ateri- maculatus (73-83 vs. 72 and 34-39 vs. 40 respectively), the small number of known S. laterimaculatus specimens (n = 1) pre- cludes classification based solely on these characters. However, others are apparent: Brown & Alcala (1980) describe the fronto- parietal of S. laterimaculatus as “long and pointed, almost as long as frontoparietals and interparietal together” (1980:178), a description which does not accord with the relative size of these scales in S. knollman- ae, especially since the frontal of S. /ateri- maculatus touches three supraoculars, whereas the frontals of S. knol/manae only contact two supraoculars. In addition, the holotype of S. /aterimaculatus has eight in- fralabials and all specimens of S. knoll- manae have six or seven. There are six or seven first finger subdigital lamellae in S. laterimaculatus and five to six in S. knoll- manae. Coloration and body proportion dif- ferences between these species are also ap- parent (see Brown & Alcala 1980).
Discussion
At the present time, the new species is only known from the type locality on Mt. Isarog (fig. 1 in Goodman & Gonzales 1990). Detailed habitat descriptions (see Brown 1919, and Whitmore 1984 for review of for- est classifications), habitat photographs, and a map of the type locality for S. knollmanae are included in Goodman & Gonzales (1990).
Very little is known about the habitat and
pe)
ecology of S. Anollmanae and its closely- related congeners, S. beyeri, S. diwata, S. laterimaculatus, S. decipiens, and S. leu- cospilos. Excepting S. decipiens and S. di- wata, all are known only from the Luzon faunal region; excepting S. decipiens, all are known only from small series. Taylor (1922) reported that the S. beyeri holotype from Mt. Banahao was collected on a rock ledge at 1500 m, but we collected most of our specimens from the Zambales Mountains by splitting open rotten logs, 1n pitfall traps, or under leaf litter—at high elevations (1265-1610 m; Brown et al. 1995). Brown & Alcala (1980) reported that S. diwata were found under leaf litter between 1600 and 1700 m on Mt. Hilong-hilong, northern Mindanao (see Brown & Rabor 1967). Sphenomorphus decipiens is also semi-fos- sorial at low to medium elevations (100- 1200 m; Brown & Alcala 1980). The known specimens of S. knollmanae were taken from similar semi-fossorial environments on the forest floor on Mt. Isarog. All were captured by L. H. Heaney, A. Alcala, and coworkers (under leaf litter, in loose topsoil, occasion- ally beside rotten logs), while digging for worms to be used as bait for mammal traps. No habitat data are available for S. /ateri- maculatus and S. leucospilos (Brown & A\l- cala 1980).
Studies of high elevation scincids and their habitats have been sorely lacking, with the exception of a few instances (Brown & Al- cala 1961la, Custudio 1986). The effects of altitudinal gradients on species richness, abundance, diversity, and distributional patterns have been addressed to a greater extent in birds (Goodman & Gonzales 1990) and mammals (Heaney et al. 1989, Rickart et al. 1991). Efforts to provide a preliminary report of altitudinal effects on scincid lizard distribution in the Philippines currently are under way. While mountain tops have been neglected by many collectors and surveyors in the past, a recent renewal of interest in their unique flora and fauna has produced discoveries (e.g., Gonzales & Kennedy 1990,
26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Lazell 1992, Ross & Gonzales 1992) and rediscoveries (e.g., Ross & Lazell 1990; Brown et al. 1995; Crombie, pers. comm.) of many taxa endemic to the Philippines.
The works of Goodman & Gonzales (1990) and Oliver et al. (1992) have both stressed the importance of continued study of montane regions in order to fuel conser- vation efforts aimed at preserving these fragile centers of endemism and diversity (see Balate et al. 1992, for a bibliography of conservation in the Philippines). We sup- port their invocations to public awareness with respect to this central issue of Philip- pine conservation given that we repeatedly have witnessed and participated in discov- eries of endemic animals new to science which inhabit extremely limited distribu- tions at high elevations in disappearing frag- ments of pristine habitat. The loss of such habitat can and often does have effects det- rimental to populations of amphibians and reptiles restricted to the immediate area (pers. obs.).
Finally, while we do not wish to engage in speculation (sensu Lazell 1992) of exactly what species may await biologists in similar environments on Philippine mountains, we do agree with Ross & Gonzales (1992) that the northern Philippines (especially the Lu- zon faunal zone) is zoogeographically com- plex and contains more centers of endem- ism than previously thought. Our recent studies suggest that the higher volcanic peaks of southern Luzon (Mt.’s Bulusan, Mayon, Labo, Banahao, Isarog, Samat, Natib, Cuadrado, Angilo and Maquiling) all war- rant intensive, long-term survey efforts of the kind that have produced (and continue to produce) many new discoveries on their neighbors.
Comparative material examined. — Sphenomorphus beyeri holotype, CAS 61183; S. beyeri, PNM 2300-2307, CMNH 3652-3655, 3657-3659, USNM 337768. S. diwata holotype (CAS 2478), S. diwata (CAS 133514 and 133515).
Acknowledgments
Collecting permits were facilitated by the Protected Areas and Wildlife Bureau of the Philippine Department of the Environment and Natural Resources, especially by A. Al- cala, and C. Catibog-Sinha. The Philippine Bureau of Forestry Development and the Bicol University College of Fisheries, Ta- baco, Albay assisted during the collection of these and other specimens during L. R. Heaney’s group’s field work on Mt. Isarog. R. I. Crombie of the United States National Museum (USNM) facilitated loans and pro- vided many helpful comments and sugges- tions throughout this and related research. W.C. Brown and J. Vindum facilitated loans of specimens in the California Academy of Science (CAS) collections and A. Alcala and L. R. Heaney generously provided access to the specimens they collected on Mt. Isarog with R. de Leon, S. M. Goodman, E. A. Rickart and R. C. B. Utzurrum. RMB thanks the Miami University Zoology Department for its encouragement and JWF acknowl- edges the continued support of Thomas More College. Comments on preliminary drafts of the manuscript were provided by W. C. Brown, R. I. Crombie, R. F. Inger, S. M. Moody, S. Simon, and one anony- mous reviewer. We owe a debt of gratitude to Pedro C. Gonzales (PNM) and Robert S. Kennedy (CMNH) for their continued sup- port of our work with Philippine herpeto- fauna.
Literature Cited
Alcala, A. C. 1986. Guide to Philippine flora and fauna. Vol. X, Amphibians and reptiles. Natural Resource Management Center Ministry of Nat- ural Resources and University of the Philip- pines, 195 pp.
Auffenberg, W. G. 1988. Gray’s Monitor Lizard. University of Florida Press, Gainesville, 419 pp.
Balate, D. S., H. C. Miranda, L. R. Heaney, & J. F. Rieger. 1992. Diversity and conservation of Philippine land vertebrates: an annotated bib- liography.— Silliman Journal 36(1):129-149.
Brown, R. M., J. W. Ferner, & R. V. Sison. 1995. Rediscovery and redescription of Sphenomor-
VOLUME 108, NUMBER 1
phus beyeri Taylor (Reptilia: Lacertilia: Scinci- dae) from the Zambales Mountains of Luzon, Philippines. — Proceedings of the Biological So- ciety of Washington 108:6—17.
Brown, W. C., & A. C. Alcala. 1961la. Populations of amphibians and reptiles in submontane and montane forests of Cuernos de Negros, Philip- pine Islands.— Ecology 42(4):628—636.
,& . 1961b. A new sphenomorphid liz-
ard from Palawan Island, Philippines. —Occa-
sional Papers of the California Academy of Sci-
ence 32:1-4.
=, 2 1970. The zoogeography of the herpetofauna of the Philippine islands, a fring- ing archipelago. — Proceedings of the California Academy of Science, fourth series 38(6):105-— 130.
——.,& . 1980. Philippine lizards of the fam- ily Scincidae. Silliman University Natural Sci- ence Monograph Series No. 2, 264 pp.
—., &D.S. Rabor. 1967. A new sphenomorphid lizard (Scincidae) from the Philippine islands. — Proceedings of the Biological Society of Wash- ington 80:69-72.
Brown, W. H. 1919. Vegetation of the Philippine mountains. Bureau of Printing, Manila, 434 pp.
Bureau of Mines, Philippines, in coordination with the Board of Technical Surveys and Maps. 1963. Geological map of the Philippines, edition No. 1.
Custudio, C. C. 1986. Altitudinal distribution of liz- ards of the Scincidae in Mt. Makiling, Laguna. — Sylvatropical Philippine Forest Research Jour- nal 11(3, 4):181-202.
Dickerson, R. E. 1924. Tertiary paleogeography of the Philippines. — Philippine Journal of Science 25(1):10-55.
Gonzales, P. C., & R.S. Kennedy. 1990. A new spe- cies of Stachyris babbler (Aves: Timaliidae) from the island of Panay, Philippines.— Wilson Bul- letin 102:367-379.
Goodman, S. M., & P. C. Gonzales. 1990. The birds of Mt. Isarog National Park, Southern Luzon, Philippines, with particular reference to altitu- dinal distribution. Fieldiana 60:1-39.
Hashimoto, W. 1981la. Geologic development of the Philippines. Pp. 83-170 in T. Kobiyashi, R. To- riyama, & W. Hashimoto, eds., Geology and Paleontology of Southeast Asia, CCXVII, Vol. UD
. 1981b. Supplementary notes on the geologic
development of the Philippines. Pp. 171-190 in
T. Kobiyashi, R. Toriyama, & W. Hashimoto,
eds., Geology and paleontology of Southeast
Asia, CCXVIII, Vol. 22.
Heaney, L. R. 1986. Biogeography of mammals in SE Asia: estimates of rates of colonization, ex-
Di,
tinction and speciation. — Biological Journal of the Linnean Society 28:127-165.
, P. D. Heideman, E. A. Rickart, R. B. Utzur- rum, & I. S. H. Klompen. 1989. Elevational zonation of mammals in the central Philip- pines. — Journal of Tropical Ecology 5:259-280.
Lazell, J. 1992. New flying lizards and predictive biogeography of two Asian archipelagos. — Bul- letin of the Museum of Comparative Zoology 152(9):475—505.
McCoy, E. D., & E. F. Connor. 1980. Latitudinal gradients in the species diversity of North American mammals.—Evolution 34:193-203.
Myers, W. C., & M. A. Donnelly. 1991. The lizard genus Sphenomorphus (Scincidae) in Panama, with a description of a new species. American Museum Novitates 3027:1-12.
Oliver, W. L. R., C. R. Cox, P. C. Gonzales, & L. R. Heaney. 1992. Cloud rats in the Philippines— preliminary report on distribution and status. — Oryx 27(1):41-48.
Rapoport, E. 1982. Areography: geographical strat- egies of species. Pergamon Press, New York, 269 pp.
Rickart, E. A., L.R. Heaney, & R.C. Utzurrum. 1991. Distribution and ecology of small mammals along an elevation transect in Southeast Luzon, Philippines.—Journal of Mammalogy 72:458- 469.
Ross, C. A., & P. C. Gonzales. 1992. Amphibians and reptiles of Catanduanes Island, Philip- pines. — National Museum Papers (Manila) 2(2): 50-76.
—., & J. D. Lazell, Jr. 1990. Amphibians and reptiles of Dinagat and Siargao Islands, Phil- ippines.—The Philippine Journal of Science 119(3):257-286.
Ruedas, L. A., J. R. Demboski, & R. V. Sison. 1994. Morphological and ecological variation in Otop- teropus cartilagonodus Kock, 1969 (Mammalia: Chiroptera: Pteropodidae) from Luzon, Phil- ippines. — Proceedings of the Biological Society of Washington 107:1-6.
Rutland, R. W. 1968. A tectonic study of part of the Philippine Fault Zone. — Quarterly Journal of the Geological Society of London 123(4):293-325.
Taylor, E.H. 1922. Additions to the herpetological fauna of the Philippine Islands, II.— Philippine Journal of Science 21(3):253-303.
SAS Institute Inc. 1988a. SAS/STAT user’s guide, release 6.03 edition. SAS Institute Inc., Cary, North Carolina, 1028 pp.
1988b. SAS procedures guide, release 6.03 edition. SAS Institute Inc., Cary, North Caro- lina, 441 pp.
Sokal, R. R., & F. J. Rohlf.
1981. Biometry, second
28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
edition. W. H. Freeman and Co., New York, ——, , & P. Escalante P. 1990. Morpholog- 859 pp. ical evolution in muroid rodents I, Conservative
UNESCO/ECAFE. 1971. Geologic map of Southeast patterns of craniometric covariance and their Asia. United Nations Publication, No. 69-30632. ontogenetic basis in the Neotropical rodent ge-
Voss, R. S., & L. F. Marcus. 1992. Morphological nus Zygodontomys. —Evolution 44:1568—1587. evolution in muroid rodents II. Craniometric Whitmore, T. C. 1984. Tropical rain forests of the factor divergence in seven Neotropical genera, Far East. Clarendon Press, Oxford, England, 718 with experimental results from Zygodonto- pp.
mys. —Evolution 46:1918—-1934.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
108(1):29-44. 1995.
Revision of the South American freshwater fish genus Sternarchorhamphus Eigenmann, 1905 (Ostariophysi: Gymnotiformes: Apteronotidae), with notes on its relationships
Ricardo Campos-da-Paz
Secao de Peixes, Museu de Zoologia da Universidade de Sao Paulo, Avenida Nazaré 481, Ipiranga—Sao Paulo—SP, CEP 04263-000, Brasil; Laboratorio de Ictiologia Geral e Aplicada, Departamentos de Zoologia e Biologia Marinha, Instituto de Biologia/CCS, Universidade Federal do Rio de Janeiro, Caixa Postal 68049,
Ilha do Fundao— Rio de Janeiro— RJ, CEP 21944-970, Brazil
Abstract. —A revision of the apteronotid genus Sternarchorhamphus Eigen- mann is presented. Sternarchorhamphus muelleri (Steindachner) from the Amazonas and Orinoco river systems, is redescribed and a lectotype is des- ignated. Sternarchorhamphus hahni Meinken, a nominal species previously assigned to this genus, is transferred to another gymnotiform genus on the basis of evidence presented herein. As a consequence, Sternarchorhamphus is now considered monotypic. A brief discussion of Mago-Leccia’s apteronotid sub- family Sternarchorhynchinae is provided and its monophyly is tentatively ac- cepted. Relationships of Sternarchorhamphus muelleri within the Sternar- chorhynchinae remain obscure, due to an absence of information on the phyletic history of closely related species, such as Orthosternarchus tamandua (Boulen- ger) and Ubidia magdalenensis Miles. A discussion on the taxonomic status of nominal species previously assigned to Sternarchorhamphus is provided.
Resumo. —Uma revisao do género de Gymnotiformes Sternarchorhamphus Eigenmann (familia Apteronotidae) € apresentada. Sternarchorhamphus muel- leri (Steindachner), das bacias dos rios Amazonas e Orinoco, é redescrita, e lectotipo é designado para esta espécie. Uma espécie nominal previamente referida a Sternarchorhamphus, S. hahni Meinken, é removida deste género; como consequéncia, Sternarchorhamphus é agora considerado género mono- tipico. Uma breve discussao sobre a subfamilia de Apteronotidae proposta por Mago-Leccia, Sternarchorhynchinae, é¢ apresentada, e sua monofilia é tentati- vamente aceita. A posicao filogenética de Sternarchorhamphus dentro da sub- familia permanece obscura, 0 que se deve, em parte, a auséncia de informacoes sobre espécies relacionadas aS. muelleri, tais como Orthosternarchus tamandua (Boulenger) e Ubidia magdalenensis Miles. E feita uma discussao sobre o status taxon6mico de espécies nominais previamente incluidas em Sternarchorham- phus.
The Neotropical gymnotiform genus Sternarchorhamphus Eigenmann (in Eigen- mann & Ward 1905) was established to in- clude three nominal species with elongate snouts, originally described in Sternarchus
Bloch & Schneider (currently a junior syn- onym of Apteronotus Lacépéde; e.g., Fowler 1951), namely: Sternarchus (Rhamphoster- narchus) macrostomus Gunther, 1870 (from Jeberos [=Xeberos], Rio Maranon basin,
30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Peru), Sternarchus (Rhamphosternarchus) muelleri Steindachner, 1881 (type species of Sternarchorhamphus, from the Rio Ama- zonas, at Para State, Brazil), and Sternar- chus tamandua Boulenger, 1898 (from the Rio Jurua, a tributary of the Rio Amazonas in Brazil). Eigenmann (in Eigenmann & Ward 1905:166), however, noted that this latter species was probably a member of “‘a genus distinct from Sternarchorhamphus as here understood” and, in fact, it was sub- sequently used by Ellis (1913) to establish the monotypic apteronotid genus Ortho- sternarchus.
The first appearance of the name Stern- archorhamphus was in a key to genera of Gymnotiformes (““Gymnotidae”’ of earlier authors) presented by Eigenmann & Ward (1905). An unequivocal citation present in that publication (p. 165) makes Eigenmann the sole author of the genus, according to Article 50a of the International Code of Zoological Nomenclature (1985). In that key, the new taxon was first included in the ““Sternarchinae”’ (=Apteronotidae), and distinguished from Sternarchorhynchus Castelnau on the basis of its “snout straight, the gape moderate,” rather than “snout strongly decurved, mouth minute.”’ Addi- tionally, it was stated that Sternarchorham- phus seemed to be “intermediate between Sternarchus and Sternarchorhynchus, hav- ing the long snout of the latter and mouth size approaching the former’ (Eigenmann & Ward 1905:165), a position also held by Ihering (1907).
Some years later, Meinken (1937) de- scribed Sternarchorhamphus hahni on the basis of a single specimen from the Rio Pa- rana drainage in Argentina. That was the first record of the genus outside of the Am- azon basin. As discussed below, however, the examination of the holotype of S. hahni showed that it does not belong to the Ap- teronotidae.
Mago-Leccia (1976) first noted the pres- ence of Sternarchorhamphus in the Rio Ori- noco system. He also proposed a phyloge-
netic scheme for Venezuelan gymnotiform genera where Sternarchorhamphus and Sternarchorhynchus were considered sister- groups within the Apteronotidae (Mago- Leccia 1976, fig. 99). Along with Orthoster- narchus Ellis and Ubidia Miles, those two genera were assigned to the Sternarcho- rhynchinae (not of Hoedeman 1962), a pro- posed apteronotid subfamily whose mem- bers were recognized by their “very elongate snout and reduced mouth gape” (Mago- Leccia 1976:206; see also Mago-Leccia 1978:14 and Mago-Leccia 1994:13).
Campos-da-Paz (1992) had recently ex- pressed some doubts about the monophyly of the Sternarchorhamphus plus Stern- archorhynchus clade, but tentatively rec- ognized Mago-Leccia’s subfamily Stern- archorhynchinae.
In a recent investigation, Triques (1993) did not find characters supporting a close relationship between Sternarchorhamphus and Sternarchorhynchus. Instead, he pro- posed Sternarchorhamphus as the sister- group to a subunit of the Apteronotidae in- cluding the short-snouted genera Adonto- sternarchus Ellis, plus Sternarchella Eigen- mann and Porotergus Ellis (Triques 1993: 123, fig. 24).
Mago-Leecia (1994) recently included two species in Sternarchorhamphus: S. muelleri and S. hahni. He stated in a footnote (p. 36), however, that this latter species could belong to a different gymnotiform genus, but without additional discussion. Sternar- chorhamphus macrostomus was used by him to establish a new monotypic apteronotid genus, Platyurosternarchus Mago-Leccia, whose relationships are currently unknown. The Sternarchorhynchinae appear only as part of a classificatory system proposed by Mago-Leccia (1978) (Mago-Leccia 1994:13).
The present study is a revision of Stern- archorhamphus Eigenmann. A detailed morphological description and osteological analysis of the type species, S. muelleri, is presented, along with data on geographic distribution based on recent collections from
VOLUME 108, NUMBER 1
the Amazon and Orinoco drainages. Re- examination of the holotype of S. hahni re- sulted in a transfer of this taxon to another gymnotiform genus.
Methods. —Most measurements were taken according to Mago-Leccia (1978) and were made point-to-point with dial calipers and ruler on the left side of specimens when- ever possible. Measurements that require further explanation are the following: LEA (Mago-Leccia et al. 1985) is the distance from the tip of snout to posterior end of anal-fin base; interorbital width is the space between the dorsal margins of the eyes; head depth was taken at the occiput; and tail depth was measured at posterior end of the anal fin. ““HL” denotes head length.
In the counts of pectoral-fin rays, the an- terior unbranched rays are indicated by low- er-case roman numerals, and branched rays are indicated by arabic numerals. The term “branched rays”’ refers to all rays posterior to the anterior unbranched rays, even if the posterior terminal ray is unbranched at its base or distal segment. All specimens with indications of regeneration and/or injuries at the posterior region of body were exclud- ed from measurements of total length (TL) and tail length (CL), and counts of caudal- fin rays. Morphometric and meristic data for the paralectotype of Sternarchorham- phus muelleri are given between brackets.
Osteological illustrations were prepared with the aid of a camera lucida, from a spec- imen cleared and counterstained for bone and cartilage with Alizarin red and Alcian blue respectively, according to the proce- dure of Taylor & Van Dyke (1985). Verte- bral counts were made on this specimen and from radiographs of additional specimens, and follow Mago-Leccia et al. (1985). Bone terminology follows Mago-Leccia et al. (1985), except that the terms mesethmoid (e.g., Fink & Fink 1981) and anguloarticular (see Nelson 1973) were used instead of eth- moid and angular, respectively. In the list of material examined, “‘c&s’’ indicates a cleared and stained specimen.
31
Specimens examined for the present study are deposited in the following institutions: American Museum of Natural History, New York (AMNH), Academy of Natural Sci- ences of Philadelphia, Philadelphia (ANSP), Instituto Nacional de Pesquisas da Ama- zonia, Manaus (INPA); Museu Nacional, Rio de Janeiro (MNRJ), Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo (MZUSP), Naturhistorisches Museum, Wien (NMW), National Museum of Natu- ral History, Smithsonian Institution, Wash- ington D.C. (USNM), and Zoologisches Museum, Berlin (ZMB). In the ‘‘Material Examined”’ section, names of collectors are cited when known, and the term ““EPA”’ re- fers to the ““Expedicao Permanente da Ama- zonia,” a Brazilian field expedition under the direction of P. Vanzolini (MZUSP) fi- nanced by the Fundacao de Amparo a Pes- quisa do Estado de Sao Paulo (FAPESP), Sao Paulo State Government.
Sternarchorhamphus Eigenmann, 1905
Sternarchorhamphus Eigenmann in Eigen- mann & Ward 1905:160, 165-166, pl. VIII, fig. 7, partim, type by original des- ignation Sternarchus (Rhamphosternar- chus) muelleri Steindachner, 1881.— Ihering 1907, in key to ““Gymnotidae,”’ key to species of Brazil.—Eigenmann 1910, list of species.—Regan 1911, list- ed.—Ellis 1912, in key to “‘Sternarchin- ae.” —Ellis 1913, in key to “‘Sternarchin- ae,’ described, food items, mutilation and regeneration of caudal peduncle. — Jordan 1920, listed, Eigenmann & Ward consid- ered as authors.—Eigenmann & Allen 1942, in key to “Apteronotinae,”’ briefly described, list of species.— Mules 1945, compared to Ubidia. —Miles 1947, com- pared to Ubidia.—Fowler 1951, Brazil, list of species.—Travassos 1960, synon- ymy, inclusion of S. hahni.—Gery & Vu- Tan-Tué 1964, briefly compared to Gym- norhamphichthys Ellis, present in key to Apteronotidae, upper Amazon.—Britski
32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1. of apteronotid fishes. a. Widespread condition in the Apteronotidae (modified from Meunier & Kirsch- baum, 1984:140, fig. 5a). b. Caudal-fin skeleton of Ster- narchorhamphus muelleri.
Diagrammatic view of caudal-fin skeleton
1972, reported from Rio Parana basin. — Mago-Leccia 1976, described from Ven- ezuela, osteology, first record in the Rio Orinoco basin, sister group of Stern- archorhynchus, assigned to the “Stern- archorhynchinae.’’— Mago-Leccia 1978, listed in the “‘Sternarchorhynchinae.” — Bullock et al. 1979, listed. —Kramer 1990, discussion on electric organ discharge patterns.—Campos-da-Paz 1992, com- pared to Sternarchorhynchus. —Triques 1993, relationships, Rio Orinoco. — Mago- Leccia, 1994, diagnosis, brief description, notes on osteology, species.
Diagnosis. — According to recent studies on interrelationships of gymnotiform gen- era (Mago-Leccia 1976, Triques 1993; see also ““Note on Relationships,’ below), the following characters are derived for Stern- archorhamphus, among closely related ap- teronotids: caudal-fin rays reduced in num- ber (two to five rays; Fig. 1); presence up to five irregular rows of diminutive dentary teeth (most easily observed on the anterior portion of this bone in adults); presence of a mesocoracoid; and snout slightly turned dorsally.
The following combination of characters,
either primitive or of uncertain polarity, distinguishes Sternarchorhamphus from all other known apteronotids: snout elongated and laterally compressed; upper jaw pro- duced and pointed, lower jaw somewhat in- cluded; teeth minute, present on both jaws at all ages; mouth small to moderate, 11.2- 16.5% [16.5%] in HL; eyes small, 2.5—4.4% [3.6%] in HL; lateral ethmoids present; scales rare or absent on antero-dorsal, and dorsal regions of body; dorsal fleshy filament orig- inating on anterior third of total length and extending to vertical through posterior end of anal fin.
Etymology. —From Sternarchus, an ear- lier nominal genus of the Apteronotidae and currently junior synonym of Apteronotus, and the Greek rhamphos, for beak. Gender masculine.
Distribution. —Most examined speci- mens of Sternarchorhamphus were collect- ed at localities near the main channels of the Rio Amazonas and Rio Orinoco, and their large tributaries. Additional speci- mens came from the vicinities of Belém (Para State, Brazil), Amapa (Amapa State, Brazil), and the Rio Ucayali basin (Peru; Fig. 2). Mago-Leccia (1976) was the first to record Sternarchorhamphus from Venezue- la, studying specimens from Rio Portuguesa and Rio Apurito. Recent collections from the Rio Orinoco originated in the deep river channel (examined specimens from the Ori- noco Delta and middle Orinoco, collected by the crew of R/V Eastward, 1978-1979, deposited at AMNH, ANSP, and USNM). Ellis (1913) and Santos et al. (1984) ex- amined specimens from the Rio Tocantins basin.
Note on relationships. —Published phy- logenetic hypotheses of the Apteronotidae do not include more than six of the ten cur- rently accepted genera (e.g., Mago-Leccia 1976, Triques 1993) and, therefore, a com- prehensive cladogram for the family is still unavailable (efforts on this subject are in progress elsewhere; J. Albert, and M. Tri- ques, pers. comm.). It is beyond the scope
VOLUME 108, NUMBER 1
NN
oA x SWE oY
33
:
Fig. 2. Geographic distribution of Sternarchorhamphus muelleri. Some symbols represent more than one
collection locality and/or lot of specimens.
of this study to present a detailed discussion on the intrafamilial relationships of the Ap- teronotidae. Furthermore, specimens of the apteronotids Orthosternarchus tamandua (Boulenger, 1898) and Ubidia magdalenen- sis Miles (1945) are extremely rare in col- lections, making it impossible to examine their internal anatomy and precluding a rig- orous test of Sternarchorhynchinae mono- phyly (O. tamandua MZUSP 2647 [1 ex.] and U. magdalenensis USNM 123795 [1 e€X.; paratype] were examined for external features).
The characters used by Mago-Leccia to define the Sternarchorhynchinae (1976), “‘reduced mouth gape’’ and “elongate snout,” and to include Sternarchorhampus in it, are rather vague; their status as sec- ondary homologues (=synapomorphies; see Pinna 1991) depends not much from their overall similarity, but rather on a well-cor- roborated hypothesis of relationships of closely related groups. An attempt to give
these characters a more objective approach is presented below.
Sternarchorhynchines all have the mouth gape length less than 38% of the snout length. In remaining apteronotids, most gymnoti- forms (except rhamphichthyids), and most closely related non-gymnotiform ostario- physans (Characiformes and Siluriformes; see Fink & Fink 1981), this value is con- sistently greater than 55%. When mouth gape length is compared to postorbital length, the value is always less than 31% in sternarchorhynchines, and greater than 40% in the proposed non-rhamphichthyid out- groups.
In most apteronotids, remaining gym- notiforms (but, again, not rhamphi- chthyids), and most closely related non- gymnotiform ostariophysans, snout length represents less than 40% in head length. In Orthosternarchus, Sternarchorhamphus and Sternarchorhynchus this value is always greater than 50%; Ubidia, however, is a
34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
unique case. Although it also has a con- spicuously elongate snout, the eye in this genus is located well forward in the head (an autapomorphic feature; see Mago-Lec- cia 1994:159, fig. 56B), resulting in the snout length/head length ratio decreasing to around 40%. Comparing snout length to postorbital length results in values of more than 90% for sternarchorhynchines (except Ubidia which, because of eye position, has the snout 70% of postorbital length) and less than 80% in remaining non-rhamphi- chthyid groups cited.
Currently available evidence indicates that the Rhamphichthyidae is distantly re- lated to the Sternarchorhynchinae (Mago- Leccia 1976, 1978; Triques 1993), and the “reduced mouth gape” and an “elongate snout” can be hypothesized as being inde- pendently acquired in these taxa.
Some authors have called attention (e.g., Schaefer 1987) and discussed (e.g., Pimentel & Riggins 1987) the questionable general utility, in phylogenetic analyses, of propor- tional differences observed between mor- phological characters. It should be noted, however, that the above discussion is based on previous phylogenetic hypotheses con- cerning external (Fink & Fink 1981) and internal (Mago-Leccia 1976, 1978; Triques 1993) gymnotiform relationships. Further- more, the utilization of morphometry in the present case is an attempt to make the terms “‘reduced”’ (the mouth) and “elongate” (the snout) as objective as possible (this kind of strategy is commonly observed in literature; e.g., Weitzman & Fink 1985 [fig. 79, char- acters 18, 32, and 40]). Additional justifi- cation for utilization of morphometry was found in Chappill (1989:231), who stated that “‘[q]uantitative characters should gen- erally only be used [i.e., in phylogenetic analyses] when the choice is between ex- amining them or abandoning the analysis entirely for want of sufficient characters.” In some cases, however, there is evidence that morphometry can explicitly be used for phylogenetic purposes (e.g., Pinna 1989:24
[character 2], Costa 1990 [fig. 32, characters 13, 34, 39, 46, 80 and 89], Schaefer 1991 (fig. 15, character 13], and Vari 1991 [fig. 11, characters 17 and 42]).
In conclusion, the monophyly of the Sternarchorhynchinae, including Sternar- chorhamphus, is tentatively accepted. The position of this genus within the subfamily, however, remains uncertain until additional material of related genera is available, which will allow a more detailed analysis of this question.
Sternarchorhamphus muelleri (Steindachner, 1881) Figs. 1-5, Table 1
Sternarchus (Rhamphosternarchus) Mulleri Steindachner 1881:99, original descrip- tion, Para State, Brazil.—Steindachner 1882:15, pl. V, fig. 4, described, no exact locality, Rio Amazonas at Para State, Brazil.
Sternarchorhynchus mulleri (not S. mulleri Castelnau 1855).—Eigenmann & Eigen- mann 1891:62, listed, Para. —Eigenmann 1894:625, listed, Para.
Sternarchorhamphus mulleri. —Eigenmann & Ward 1905:165-—166, pl. VIII, fig. 7, as- signed as type species of Sternarchorham- phus, original designation, Para. — Eigen- mann 1910:449, listed, Para to Peru.— Ellis 1913:142-143, 174, 182-183, figs. 10 and 23-25, described, food habits, re- generation of caudal peduncle, Alcobaca (=‘*‘Alcoboca’’), Rio Tocantins, Para.— Meinken 1937:79, compared to Sternar- chorhamphus hahni.—Fowler 1939:276, Contamana, Peru.—Eigenmann & Allen 1942:320, listed, lower Amazon to the Ucayali.— Fowler 1945:180, fig. 65, Peru (Contamana) and Amazonas.— Miles 1947:184, compared to Ubidia magda- lenensis.
Sternarchorhamphus muelleri. —\hering 1907:277, Amazonas, Para.— Mago-Lec- cia 1976:244—249, fig. 78, described from Rio Orinoco, Venezuela. — Ortega & Vari
VOLUME 108, NUMBER 1
1986:12, Peru.—Campos-da-Paz 1992: 24, 134, briefly compared to Sternar- chorhynchus. —Mago-Leccia, 1994:35, 36, 71, 107, 156, 203, fig. 52, brief descrip- tion, notes on osteology, listed.
Sternarchorhamphus mulleri.—Starks 1913: 23, described, variation of anus position, Para.—Santos et al. 1984:18, 78, figure, in list of species from lower Rio Tocan- tins, common names, Brazil.—Triques 1993:91, listed, discussion of relation- ships to other gymnotiforms.
Sternarchus mulleri. —Jordan 1920:512, cited as type species of Sternarchorham- phus.
Sternachorhamphus [misspelling] muel- leri. —Magalhaes 1931:178, cited.
Sternarchus mulleri.—Travassos 1960:24, cited as type species of Sternarchorham- phus.
Sternarchoramphus [misspelling] mul- leri. —Begossi & Braga 1992:107, tb.2, and 11, fig. 5, common name in Rio Tocan- tins, listed as fish avoided as food.
Material examined. —Brazil: NMW 65328:1, lectotype (photograph; new des- ignation), 260.5 mm LEA, no exact collec- tion site, Rio Amazonas at Para State, F. Steindachner donation, accession file num- ber “1874.1.299 & 299a”; NMW 65328:2, paralectotype, 249.0 mm LEA, same data of lectotype; INPA 4850, 7 specimens, Ca- maleao, Ilha da Marchantaria, Rio Soli- moes, Amazonas State, 3 Feb 1982, G. M. Soares; INPA 4852, 5 specimens, Cama- leao, Ilha da Marchantaria, Rio Solimoes, Amazonas State, 1 Jun 1981, G. M. Soares; MNRJ 1221, 1 specimen, no collection data; MNRJ 1222, 2 specimens, no collection data; MNRJ 9022, 2 specimens, ‘Ver-o- Peso’ market, Belem, Para State, 10 Feb 1958, L. Travassos & F. Pires leg.; MNRJ 12182, 1 specimen, Amapa, Amapa State, Apr 1981, G. W. Nunan et al.; MZUSP 6983, 7 specimens, Rio Madeira, 25 km below Nova Olinda, Amazonas State, 27 Sep 1967, EPA; MZUSP 9580, 1 specimen,
35
fish market at Manaus, Amazonas State, 17— 19 Sep 1968, EPA leg.; MZUSP 23321, 1 specimen, mouth of Parana do Catito, be- low mouth of Rio Jurua, Rio Solimdées, Amazonas State, 4 Oct 1968, EPA; MZUSP 24675, 1 specimen, Santa Luzia, Rio Purus, Amazonas State, 11 Jan 1975, P. E. Van- zolini; USNM 52547, 2 specimens, Para to Manaus, Rio Amazonas, 1901, J. B. Steere.
Peru: ANSP 95834, 2 specimens, Rio Ucayali at Cantamana, Jul-Aug 1937, W. C. Morrow; ANSP 120348, 2 specimens, Pucallpa, on Rio Ucayali, 18-19 Jun 1969, E. J. Huggins.
Venezuela: ANSP 149460, 3 specimens, shallow river, N side of river across from Isla Tres Canos, Delta Amacuro, 13 Nov 1979, R/V Eastward (H. Lopez, M. Cor- coran); ANSP 149457, 1 specimen, Rio Ori- noco, below Barrancas, ca. km 140, depth 26 m, Delta Amacuro, 17 Feb 1978, R/V Eastward (Lundberg & Baskin); ANSP 160250, 6 specimens, Rio Guariquito at confluence of Rio Orinoco, Estado Bolivar, 25 Nov 1985, B. Chernoff et al.; ANSP 162297, 2 specimens, Rio Orinoco, near mouth of Rio Caura, Estado Bolivar, 22 Nov 1985, G. J. Lundberg et al.; ANSP 166792, 1 specimen, Caicara, L. Bartolico, Estado Bolivar, Rio Orinoco basin, 20 May 1987, M. Rodriguez & R. Richardson; ANSP 166793, 1 specimen, Caicara, Cas- tillero, Estado Bolivar, Rio Orinoco basin, 19 Apr 1988, M. A. Rodriguez & A. Mar- tinez; USNM 226339, 1 specimen, Rio Ori- noco, deep river channel, Brazo Imataca, 82 n.m. upstream from sea buoy, Territorio Federal Delta Amacuro, 22 Feb 1978, D. Taphorn; USNM 226491, 2 specimens, Rio Orinoco, deep river channel, north side of Isla Tortola, 123 n. m. from sea buoy, Delta Amacuro Fed. Territory, 19 Feb 1978, J. N. Baskin; USNM 226495, 3 specimens, Rio Orinoco, Brazo Imataca, south from Isla Remolino, ca. km 82.9 from sea buoy, Delta Amacuro Fed. Territory, 22 Feb 1978, Baskin/Lundberg; USNM 228805, 6 spec- imens (1 C&S), across from Palua, ca. 182
36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.—Morphometric data for the paralectotype (NMW 65328:2) and additional non-type material of Sternarchorhamphus muelleri. TL, LEA, HL, and CL are expressed in mm; measurement | is expressed as % of TL; measurements 2 to 7 are expressed as % of LEA; 8 to 21 as % of HL; 22 as % of CL. SD, standard
deviation; and n, number of examined specimens.
Paralectotype Range X SD n TL 292.0 261.0-446.0 Ss 15 LEA 249.0 188.0—375.0 - — 17 HL 30.0 20.5-63.3 - — 28 CL 43.0 35.0-77.0 - — tS) 1) Standard length 85.2 81.4-88.2 83.9 + 1.7 15 2) Anal-fin length 90.4 89.1-93.0 Oiie sale 15 3) Body depth 10.1 8.7-12.2 10.2 + 0.8 18 4) Head length 12.1 11.4-14.5 12.4 + 0.9 18 5) Pre-anal distance 9.3 8.0-10.8 Wh se 07 18 6) Pre-pectoral distance Doi 11.3-15.4 WoW ae Ilo 18 7) Tail length 72 16.7—22.7 HOLS =e9 14 8) Snout length 49.5 45.9-52.9 49.3 + 2.0 28 9) Eye diameter 3.6 2.54.4 3.3 + 0.4 28 10) Mouth width 16.5 11.2-16.5 13.3 + 1.4 25 11) Interorbital width 11.9 8.0-13.0 10.6 + 1.4 28 12) Snout to occiput 90.1 83.5-91.7 87.4 + 2.3 28 13) Postorbital distance 53.4 46.9-53.8 50.5 + 1.9 28 14) Pectoral-fin length 45.2 43.5-59.0 SOLO) as S77 28 15) Pre-anal-fin length 715.9 68.8-79.8 VAS) se ZO 26 16) Snout to anus 58.7 48 .5-60.2 54.8 + 3.5 Di) 17) Anterior naris—posterior naris 5.6 3.3-5.6 4.3 + 0.6 26 18) Posterior naris—eye MoT 22.0-30.3 26.8 + 2.1 26 19) Head depth 62.0 52.1-63.9 S75 se Sats} 26 20) Head width 24.1 22.2-29.9 2p eal 28 21) Branchial opening 19.4 15.6-19.4 1 2 IED 28 22) Tail depth 9.7 6.3-10.6 Sie ales 14
n. m. from sea buoy, 9 Nov 1975, H. Lopez & O. Riviero; USNM 228806, 6 specimens, shallow river, downstream from sea buoy 82, near mouth of a small cano, Delta Ama- curo Fed. Territory, 21 Nov 1979, H. Lopez et al.; USNM 228808, 1 specimen, shallow river, north shore, 49 n. m. from sea buoy, Delta Amacuro Fed. Territory, 20 Nov 1979, Lopez et al.
Diagnosis. —As for the genus.
Description. —Morphometric data for the paralectotype (NMW 65328:2) and addi- tional specimens of S. muelleri are given in Table 1. The lectotype (NMW 65328:1) is shown in Fig. 3.
Body strongly compressed laterally, es- pecially posterior to abdominal cavity, deepest in this region; dorsal profile from
nearly straight to gently convex. Lateral line complete, extending to caudal peduncle, but not onto caudal fin in some specimens.
Head laterally compressed, more mark- edly so anteriorly, deepest at occiput and widest in opercular area; snout pointed and conical, slightly turned dorsally in most specimens; eyes reduced in size, completely covered with skin, located dorso-laterally on head; small (paired) sensory pore pres- ent, dorsal to eye and usually near vertical through anterior border of eyes.
Mouth small, sub-terminal (inferior in some specimens), its gape usually not reach- ing vertical through anterior border of an- terior nare; upper jaw projected, lower jaw somewhat included. Single patch of numer- ous diminutive conical teeth present on pre-
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38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
maxilla (25-40, ¥ = 34.1, nm = 7 [approx. 35 in paralectotype]). Usually four of five irregular rows of similar teeth on dentary (25-45, X = 36.4, n = 7 [approx. 25 in par- alectotype]). Maxilla and mesopterygoid edentulous. Small rounded papillae cover- ing part of the roof and floor of the oral cavity. Anterior nares tubular; posterior ones rounded, without a tube and remote from eyes.
Branchial opening reduced to a short slit immediately anterior to pectoral-fin origin; branchial membranes joined or partially joined to isthmus. Anterior chamber of gas bladder small, posterior elongate. Anus and urogenital papilla adjacent and located ven- tral to opercular region, shifting anteriorly with age.
Pectoral fins moderate, elongate and slightly pointed, with 11 + 12-14 rays [ii + 13]. Anal fin with 230-262 rays [231], its origin near vertical through nape; first an- terior anal-fin rays less developed (some un- divided) and smaller than posterior ones.
Scales small, cycloid, absent or greatly re- duced in number on anterior region of body and dorsally; lateral line perforated scales usually larger than those immediately dor- sal and ventral to it. Scales above lateral line four to nine. Small scales, sometimes in single series, present on caudal fin in some specimens.
Dorsal filament (=“‘dorsal thong”; a mod- ified muscle according to Mago-Leccia 1994) originating on anterior third of total length, its tip reaching vertical through posterior end of anal fin. Tail (region from base of last anal-fin ray to tip of caudal fin) elongate, laterally compressed, ending in reduced caudal fin with two to five rays [four]; con- dition unique among Gymnotiformes. Cau- dal peduncle sometimes with discrete con- striction at caudal-fin base.
Osteological features of Sternarchorham- phus muelleri, relevant at the generic level, are as follows: premaxilla broadest anteri- orly, and extending posteriorly to near the maxilla; maxilla elongate and curved pos- teriorly, its anterior portion well-developed
(Figs. 4 and 5); dentary large, extending pos- teriorly and covering anterior portion of re- troarticular; Meckel’s cartilage well-devel- oped and elongate, partially associated with anguloarticular and dentary; coronomeck- elian bone reduced in size, compared to ad- jacent bones; retroarticular well-developed, with pointed antero-ventral process; lateral ethmoids present; vomer elongate, its an- terior portion “arrow-shaped” and contact- ing parasphenoid through small cartilagi- nous bridge; posterior end of vomer point- ed; palatine cartilage present, well-devel- oped anteriorly and contacting anterior portion of maxilla; mesopterygoid broad, edentulous, with poorly-developed ascend- ing process; parasphenoid elongate, bifur- cate anteriorly and posteriorly; infraorbital series represented only by canal-bearing portions of bones; mesethmoid elongate, rounded and reduced at its anterior portion; two cranial fontanels present and well-de- veloped (interfrontal larger than interpari- etal); supraoccipital small, with reduced crest; posttemporal fossae absent; opercle ornamented with numerous small trabecu- lae; mesocoracoid present; scapular fora- men absent; coracoid with well-developed postero-ventral process, but not reaching cleithral symphysis; posttemporal fused to supracleithrum; extrascapular present; two postcleithra; four pectoral radials; four branchiostegal rays, first and second almost filamentous, others large and laminar; uro- hyal broad and expanded posteriorly with reduced head, and approaching in size the basihyal and first ceratobranchial; gill rakers not ossified; four infrapharyngobranchials, fourth one cartilaginous; five epibranchials, fifth one cartilaginous; upper pharyngeal tooth plate with seven or eight teeth, con- nected to third epibranchial through a lig- ament; lower pharyngeal tooth plate with 12-13 teeth; Weberian apparatus without claustrum; 16-17 precaudal vertebrae (We- berian complex included); two “‘rib-like bones”’ (modified ribs?) present in posterior wall of abdominal cavity, anterior one en- larged, laterally compressed and turned an-
VOLUME 108, NUMBER 1
39
de
—
Fig. 4. Jaws of S. muelleri, USNM 228805, 188.8 mm LEA. Lateral view of left side. Scale bar = 1 mm. aa, anguloarticular; de, dentary; ma, maxilla; pm, premaxilla; ra, retroarticular.
teriorly; proximal pterygiophores of anal fin with expanded, pointed, symmetric projec- tions anteriorly, directed dorsally on the an- terior third of pterygiophore, well-devel- oped posterior to abdominal region; 91—99 vertebrae to base of last anal-fin ray (We- berian complex included); well-developed intermuscular bones, especially conspicu- ous dorsally and ventrally to vertebral col- umn, and posterior to abdominal region; caudal-fin skeleton consolidated into a sin- gle element of reduced size.
Color in alcohol.—Body light yellowish brown, covered with diminutive irregular dark spots (chromatophores), most on dor- sal portion of head and snout, and on back.
The overall coloration can vary from pallid
to melanistic. Pectoral fins hyaline at base, -and usually dark distally (chromatophores
on fin membranes); anal fin of some spec- imens with continuous black margin along entire length, with melanophores also pres- ent over fin membranes; specimens from the Rio Orinoco, however, can be nearly completely pallid (J. Lundberg, pers. comm.). Caudal fin hyaline.
Food habits. —Stomach-contents of Sternarchorhamphus showed fragments of
partially digested insect larvae (tentatively
identified as Diptera), along with some un- identified Annelida. These findings agree with those by Ellis (1913:174), who had also
ra Fig. 5.
de
Lower jaw of S. muelleri, USNM 228805, 188.8 mm LEA. Medial view of left side. Scale bar = 1
mm. aa, anguloarticular; cb, coronomeckelian bone; de, dentary; mc, Meckel’s cartilage; ra, retroarticular.
40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
noted a single unidentified Entomostraca specimen and additional insect larvae, other than Diptera.
Remarks. —Santos et al. (1984) listed the names “itui” (a common name for most apteronotid fishes in Brazil) and “‘tuvira”’ (also used for other Gymnotiformes, except the Electrophoridae) for S. muelleri in the lower Rio Tocantins region. The same au- thors also observed that this species is the only apteronotid with some commercial value as a food fish in that area. Begossi & Braga (1992), curiously, listed S. muelleri as a fish avoided as food by local fishermen at Rio Tocantins (in States of Maranhao and Tocantins), and recorded the common name ‘“‘lampréia”’ to this species (that name is also assigned for other non-apteronotid gym- notiform fishes occurring in that area).
Langner & Scheich (1978) and Kramer (1990) discussed the electric organ discharg- es (EODs) of some gymnotiforms, including specimens they identified as Sternarchor- hamphus, which showed a fundamental fre- quency of EOD between 1300 and 1700 Hz. A figure presented by Langner & Scheich (1978:236, fig. 1), and later reproduced by Kramer (1990:198, fig. 4.57), however, does not show Sternarchorhamphus, but Stern- archorhynchus.
Etymology. —Steindachner proposed the name Mulleri (the original spelling in his 1881 paper) for the new species in honor of Johannes Muller, a famous German ichthy- ologist of the 19th century. Several different spelling forms for this species are currently widespread in literature (see synonymy, above). According to the International Code of Zoological Nomenclature (1985; Article 32d), however, the correct form is muelleri.
Taxonomic status of Sternarchus (Rhamphosternarchus) macrostomus Gunther, Sternarchus tamandua Boulenger, and Sternarchorhamphus hahni Meinken
Gunther (1870) described Sternarchus (Rhamphosternarchus) macrostomus (“‘ma-
crostoma’’ in the original spelling; not Ster- narchus macrostomus Fowler [1943]) on the basis of a single specimen from Jeberos (=Xeberos), Rio Maranon drainage, Peru. This nominal species was subsequently as- signed to Sternarchorhynchus by Eigen- mann & Eigenmann (1891; “Upper Ama- zon’’). Eigenmann (in Eigenmann & Ward 1905) included S. macrostomus in Sternar- chorhamphus, without any detailed expla- nation. Ellis (1913) did not examine spec- imens of S. macrostomus and simply quot- ed Gunther’s original description, following Eigenmann’s placement of the species. This view remained unalterated since then, until the recent work of Mago-Leccia (1994), who used the species to establish the monotypic apteronotid genus Platyurosternarchus. Platyurosternarchus first appeared in a key to apteronotid genera (Mago-Leccia 1994: 26). Later, in the same study, that author stated that the genus was “proposed in order to locate properly the species Sternarchus macrostomus,’”’ that the new genus is “‘clear- ly different from Sternarchorhamphus”’ (p. 37), and provided a photograph of the head of a specimen from the Orinoco basin (p. 160, fig. 57A). A list of distinctive features (including uniquely derived characters, such as the overall morphology of caudal fin) was also given by Mago-Leccia, who definitely demonstrated that P. macrostomus and S. muelleri are different species (but note also characters already pointed out by Gunther 1870). Relationships of Platyurosternar- chus, however, were not discussed in that work, and the closest relatives of that genus remain uncertain. A preliminary view of this problem suggests that Platyurosternarchus and Sternarchorhamphus belong to differ- ent subsets within the family Apteronotidae and are not sister groups. As in S. muelleri, the snout in P. macrostomus is elongate (around 50% in head length), but the latter species does not have the reduced mouth gape length characteristic of sternarcho- rhynchines (approximately 65% in snout length, and 75% in postorbital length vs. less
VOLUME 108, NUMBER |
than 35% in snout length, and less than 31% in postorbital length, respectively). The pos- sibility of Platyurosternarchus macrosto- mus constitutes the sister-group of the Sternarchorhynchinae cannot be dismissed at this time, but further detailed investiga- tion is needed (see discussion on the Stern- archorhynchinae, above).
Sternarchus tamandua was described by Boulenger (1898) on the basis of a single specimen, with an injured tail, from Rio Jurua (Brazil). The species was transferred to Sternarchorhamphus by Eigenmann (in Eigenmann & Ward 1905), who was fol- lowed by Ihering (1907) and Eigenmann (1910). Later, the species was used by Ellis (1913) to establish the monotypic apter- onotid genus Orthosternarchus. The name of this genus, however, appeared first in key to genera to ““Gymnotidae” (=Gymnoti- formes) from British Guiana provided by Ellis (1912; type-species’ name was not mentioned), and that should be considered its original description. Curiously, not a sin- gle specimen of Orthosternarchus was re- corded from Guiana until the present date. Little has been published on this poorly known species since then, mainly because of its scarcity in collections. Detailed infor- mation on its internal morphology and os- teology is unavailable. The overall form of the snout is an autapomorphic feature, clearly suggesting that O. tamandua is not conspecific with S. muelleri or any known apteronotid (see, for example, Ellis 1913: 144, fig. 11, Mago-Leccia 1994:147, fig. 41). Since this genus is currently assigned to the Sternarchorhynchinae, the possibility of Orthosternarchus and Sternarchorhamphus are its sister-groups must be left open until enough material is available to allow this kind of investigation. If a sister-group re- lationship between these two genera can be demonstrated, then an interesting taxonom- ic question will arise, since their included species were once referred to a single genus, Sternarchorhamphus. Current available ev- idence is scarce and cannot definitively show
41
a close relationship between O. tamandua and S. muelleri, so these species are here kept in their own separate genera.
A study on the single known specimen of Sternarchorhamphus hahni Meinken (ho- lotype, ZMB 31367), collected near Corri- entes, Rio Parana basin in Argentina, re- vealed that it lacks the diagnostic features of the Apteronotidae (e.g., caudal fin inter- nally supported by a single bony element; a dorsal fleshy filament) and should rather be referred to Rhamphichthys Miller & Tros- chel, 1846 (Rhamphichthyidae). In addi- tion to several features observed only in rhamphichthyids and closely related groups among gymnotiforms (Hypopomidae; see Mago-Leccia 1978, and Triques 1993; e.g., no teeth on both jaws; anterior nares not tubular and located close to upper lip), S. hahni has 323 anal-fin rays (330 recorded by Meiken 1937; vs. a maximum of 290 anal-fin rays in related groups), a condition apparently uniquely derived for Rhamphi- chthys (or a subset of it) among closely re- lated taxa. Mago-Leccia (1976, 1994) pre- viously posed some doubts on the taxonom- ic status of S. hahni but, because of lack of access to type material, did not discuss the question in greater detail (a more detailed approach on this subject is currently being published elsewhere; Campos-da-Paz & Paepke 1994).
In conclusion, it seems reasonable not to include the nominal species S. hahni Mei- ken and S. macrostomus (Gunther) in Stern- archorhamphus which, otherwise, would make it non-monophyletic. Sternarchus ta- mandu Boulenger remains in Orthosternar- chus Ellis until a detailed phylogenetic study on the Sternarchorhynchinae reveals its po- sition regarding that genus. The solution presented herein is to consider Sternar- chorhamphus a monotypic subunit of the Apteronotidae, comprising only S. muelleri (Steindachner). As discussed above, how- ever, its phylogenetic relationships to other sternarchorhynchine apteronotid genera re- mains uncertain, depending on more com-
42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
plete and conclusive investigations than those presented so far.
Acknowledgments
For conversations and discussions on sys- tematics and taxonomy of gymnotiform fishes, I am grateful to J. Albert, H. A. Brit- ski, W. J. E. M. Costa, C. Cox-Fernandes, J. G. Lundberg, N. A. Menezes, H.-J. Paepke, M. C. C. de Pinna, J. P. Sullivan, S. Toledo and M. L. Triques. I am deeply indebted to the following individuals and institutions for their hospitality, access to collections and/or to specimens under their care, logistic and equipment support, loan of specimens and all kinds of additional as- sistance: N. Feinberg, G. J. Nelson and M. S. T. Piza (AMNH); E. B. Bohlke, P. A. Buckup, W. G. Saul, and S. A. Schaefer (ANSP); M. C. C. de Pinna (Field Museum of Natural History, Chicago); M. Jégu (INPA); C. A. Bizerril, U. Caramaschi, D. F. Moraes, Jr., I. B. Moreira, G. W. Nunan and J. C. de Oliveira (MNRJ); F. A. Bock- mann, H. A. Britski, J. L. Figueiredo, N. A. Menezes, O. T. Oyakawa, L. M. S. Porto, and M. L. Triques (MZUSP); B. Herzig (NMW); J. G. Lundberg (University of Ar- izona, Tucson); E. P. Caramaschi, W. J. E. M. Costa, and R. Sachsse (Universidade Federal do Rio de Janeiro); S. Jewett, R. P. Vari, and S. Weitzman (USNM); and H.-J. Paepke (ZMB). Earlier versions of manu- script benefited from comments and sug- gestions by J. Albert, H. A. Britski, W. J. E. M. Costa, J. G. Lundberg, N. A. Menezes, T. A. Munroe, M. C. C. de Pinna, and R. P. Vari. A photograph of the lectotype of S. muelleri was taken by Mrs. A. Schumacher (NMW), and kindly provided by B. Herzig (NMW).
The author was financially supported by the Fundagao de Amparo a Pesquisa do Es- tado de Sao Paulo (FAPESP), Sao Paulo State Government, Brazil. The following in- stitutions also funded visits to study part of their gymnotiform collections: Academy of
Natural Sciences of Philadelphia (Philadel- phia), American Museum of Natural His- tory (New York), and National Museum of Natural History, Smithsonian Institution (Washington, D.C.).
Literature Cited
Begossi, A., & F. M.S. Braga. 1992. Food taboos and folk medicine among fishermen from the To- cantins River (Brazil).—Amazoniana 12:101- 118.
Boulenger, G. A. 1898. Onacollection of fishes from the Rio Jurua, Brazil.— Transactions of the Zoo- logical Society of London 14:421-428.
Britski, H. A. 1972. Peixes de agua doce do estado de Sado Paulo: Sistematica. Pp. 79-108 in Po- luigao e piscicultura.—Comissao Interestadual da bacia Parana-Paraguai, Sao Paulo.
Bullock, T. H., N. Fernandes-Souza, W. Graf, W. Hei- ligenberg, G. Langner, D. L. Meyer, F. Pimentel- Souza, H. Scheich, & A. Viancour. 1979. As- pectos do uso da descarga do orgao elétrico e eletrorrecepcao nos Gymnotoidei e outros peix- es amazonicos.— Acta Amazonica 9:549-572.
Campos-da-Paz, R. 1992. Revisaéo taxondmica do género Sternarchorhynchus Castelnau, 1855
(Ostariophysi Gymnotoidei, Apteronotidae).
Unpublished MLS. thesis, Universidade de Sao
Paulo, 159 pp.
, & H.-J. Paepke. 1994. On Sternarchorham- phus hahni, a member of the rhamphichthyid genus Rhamphichthys (Ostariophysi: Gymno- tiformes).—Ichthyological Exploration of Freshwaters 5:155-159.
Castelanu, F. 1855. Poissons nouveaux ou rares reé- cuillem pendant l’expedition dans les parties centrales de l’Amérique de Sud, de Rio de Ja- neiro a Lima, et de Lima au Para.—Paris, 1/2: 85-95, pls. 45-47.
Chappill, J. A. 1989. Quantitative characters in phy- logenetic analysis. — Cladistics 5:217-—234.
Costa, W. J.E.M. 1990. Andalise filogenética da fam- ilia Rivulidae (Cyprinodontiformes Aplochei- loidei)— Revista Brasileira de Biologia 50:65— 82.
Eigenmann, C. H. 1894. Notes on some South Amer-
ican fishes.— Annals of the New York Academy
of Sciences 7:625-637.
1910. Catalogue of the freshwater fishes of Tropical and South Temperate America. Pp. 375-511 in Report of the Princeton University Expedition to Patagonia, 1896-1899, Volume Three.
——, & W. R. Allen. 1942. Fishes of the Western South America, I: the Intercordilleran and Am-
VOLUME 108, NUMBER 1
azonian Lowlands of Peru; II: the High Pampas,
Bolivia, and Northern Chile; with a revision of
the Peruvian Gymnotidae, and of the genus Or-
estias. University of Kentucky, Lexington, 494
pp.
,& R.S. Eigenmann. 1891. A catalogue of the
fresh-water fishes of South America. — Proceed-
ings of the United States National Museum 14:
1-81.
—., & D. P. Ward. 1905. The Gymnotidae.— Proceedings of the Washington Academy of Sci- ences 7:157-186.
Ellis, M. M. 1912. Order Glanencheli. Family X.
Gymnotidae. Pp. 422-442 in C. H. Eigenmann,
The freshwater fishes of British Guiana, includ-
ing a study of the ecological groupings of species
and the relation of the fauna of the Plateau to that of the Lowlands. — Memoirs of the Carnegie
Museum 5:1-578.
1913. The gymnotid eels of Tropical Amer- ica.— Memoirs of the Carnegie Museum 6:109- 195.
Fink, S. V., & W. L. Fink. 1981. Interrelationships of the ostariophysan fishes (Teleostei).—Zoo- logical Journal of the Linnean Society 72:297- 353.
Fowler, H. W. 1939. A collection of fishes obtained
by Mr. William C. Morrow in the Ucayali river
basin, Peru.— Proceedings of the Academy Nat-
ural Sciences of Philadelphia 91:219-289.
1943. A collection of freshwater fishes from Colombia, obtained chiefly by Brother Nicéforo Maria. — Proceedings of the Academy of Natural Sciences of Philadelphia 95:260-264.
1945. Los peces del Peru: Catalogo siste- matico de los peces que habitan en aguas per- uanas.— Boletin del Museo de Historia Natural ‘Javier Prado,” Lima 7:176—185.
1951. Os peixes de agua doce do Brasil (3a Entrega).—Arquivos de Zoologia, Sao Paulo 6:405-628.
Géry, J., & Vu-Tan-Tué. 1964. Gymnorhamphi- chthys hypostomus petiti ssp. nov., un curieux poisson gymnotoide arénicole.— Vie et Milieu, Suppl. 17:485-498.
Gunther, A. 1870. Catalogue of the fishes in the col- lection of the British Museum. —British Muse- um Ed., 8:1-549, London.
Hoedeman, J. J. 1962. Notes on the Ichthyology of Surinam and other Guianas. 11. New Gymno- toid Fishes from Surinam and French Guiana, with additional records and a key to the groups and species from Guiana.— Bulletin of Aquatic Biology 3:97-107.
Ihering, R. von. 1907. Os peixes de agua doce do Brasil.— Revista do Museu Paulista 7:258—-336.
International Commission on Zoological Nomencla-
43
ture. 1985. International Code of Zoological Nomenclature. Third Edition. International Trust for Zoological Nomenclature & British Museum (Natural History), London. University of California Press, Berkeley, Los Angeles, 338 pp.
Jordan, D. S. 1920. The genera of fishes. Part IV. From 1881 to 1920, thirty-nine years, with the accepted type of each. A contribution to the stability of scientific nomenclature.— Leland Stanford Junior University Publications, Uni- versity Series, p. 512.
Kramer, B. 1990. Electro-communication in Teleost fishes: behavior and experiments. Springer- Ver- lag, Berlin, 240 pp.
Langner, G., & H. Scheich. 1978. Active phase cou- pling in electric fish: behavior control with mi- crosecond precision.—Journal of Comparative Physiology 128:235-—240.
Magalhaes, A. C. 1931. Monographia brazileira de peixes fluviaes. Graphicars Ed., Sao Paulo, 260 pp.
Mago-Leccia, F. 1976. Los peces Gymnotiformes de
Venezuela: un estudio preliminar para la revi-
sion del grupo en la America del Sur. Unpub-
lished Ph.D. thesis, Universidad Central de
Venezuela, Caracas, 376 pp.
1978. Los peces de la familia Sternopygidae de Venezuela.— Acta Cientifica Venezolana 29 (Supl. 1):1-89.
. 1994. Electric fishes of the continental waters
of America.—Biblioteca de la Academia de
Ciencias Fisicas Matematicas y Naturales 29:1-
ies
——., J. G. Lundberg, & J. N. Baskin. 1985. Sys- tematics of the South American freshwater ge- nus Adontosternarchus (Gymnotiformes, Apter- onotidae).— Contributions in Science, Natural History Museum, Los Angeles County, 358:1- 19.
Meinken, H. 1937. Beitrage zur Fischfauna des mit- tleren Parana.III].—Blatter fiir Aquarien und Terrarienkunde 48:73-80.
Miles, C. 1945. Some newly recorded fishes from the
Magdalena river system. Gymnotidae.—Cal-
dasia 3:461-464.
1947. Los peces del Rio Magdalena (‘‘A field book of Magdalena fishes’’). Ministerio de la Economia Nacional, Seccion de Piscicultura, Pesca y Caza, Bogota, 214 pp.
Nelson, G. 1973. Relationships of clupeomorphs, with remarks on the structure of the lower jaw in fishes. — Zoological Journal of the Linnean So- ciety 53 (Suppl. 1):333-349.
Ortega, H., & R. P. Vari. 1986. Annotated checklist of the freshwater fishes of Peru. —Smithsonian Contributions to Zoology 437:1—25.
44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Pimentel, R. A., & R. Riggins. 1987. The nature of cladistic data.— Cladistics 3:201-209.
Pinna, M. C. C. 1989. A new sarcoglanidine catfish,
phylogeny of its subfamily, and an appraisal of
the phylectic status of the Trichomycteridae
(Teleostei, Trichomycteridae).— American Mu-
seum Novitates 2950:1-39.
1991. Concepts and tests of homology in the cladistic paradigm. — Cladistics 7:367-394. Regan, C. T. 1911. The classification of the Teleos-
tean fishes of the order Ostariophysi. I. Cypri- noidea. Division 2. Gymnotiformes.— Annals and Magazine of Natural History, ser. 7, 11:23- 26.
Santos, G. M., M. Jégu, & B. Merona. 1984. Catalogo dos peixes comerciais do baixo rio Tocantins. — Projeto Tucurui. Eletronorte/CNPq/INPA. Ma- naus, 83 pp.
Schaefer, S. A. 1987. Osteology of Hypostomus ple-
costomus (Linnaeus), with a phylogenetic anal-
ysis of the loricariid subfamilies (Pisces: Silu- roidei). — Contributions in Science, Natural His-
tory Museum, Los Angeles County, 394:1-31.
1991. Phylogenetic analysis of the loricariid subfamily Hypoptopomatinae (Pisces: Siluro- idei: Loricariidae), with comments on generic diagnosis and geographic distribution. —Zoo- logical Journal of the Linnean Society 102:1- 41.
Starks, E. C. 1913. The fishes of the Stanford Expe- dition to Brazil.— Leland Stanford Junior Uni- versity Publications, University Series 12:1-77.
Steindachner, F. 1881. Beitrage zur Kenntniss der
Flussfische Sudamerika’s. IIJ.—Anzeiger der
Akademie der Wissenschaften, Wien 18:97—100.
1882. Beitrage zur Kenntniss der Flussfische
Sudamerika’s. III.—Denkschriften der Akade-
mie der Wissenschaften, Wien 44:1-18.
Taylor, R., & C. C. Van Dyke. 1985. Revised pro- cedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. — Cybium 9:107-119.
Travassos, H. 1960. Ictiofauna de Pirassununga. IV. Subordem Gymnotoidei Berg, 1940 (Actino- pterygii— Cypriniformes).— Boletim do Museu Nacional, Rio de Janeiro, nova série, Zoologia 217:1-34.
Triques, M. L. 1993. Filogenia dos géneros de Gym- notiformes (Actinopterygii, Ostariophysi), com base em caracteres esqueléticos.—Comunica- coes do Museu de Ciéncias da PUCRS, série zoologia, Porto Alegre, 6:85—130.
Vari, R. P. 1991. Systematics of the Neotropical char- aciform genus Steindachnerina Fowler (Pisces: Ostariophysi).—Smithsonian Contributions to Zoology 507:1-118.
Weitzman, S. H., & S. Fink. 1985. Xenurobryconin phylogeny and putative pheromone pumps in glandulocaudine fishes (Teleostei: Characi- dae).—Smithsonian Contributions to Zoology 421:1-121.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
108(1):45—-49. 1995.
A new species of Neoperla (Insecta: Pleocoptera: Perlidae) from Mississippi
Bill P. Stark
Department of Biology, Mississippi College, Clinton, Mississippi 39058, U.S.A.
Abstract. —Neoperla coxi, a new species, is described from male, female, egg and nymphal stages, and a holotype male is designated. Males differ from all known Nearctic Neoper/a in having most of the aedeagal tube spinulose. The new species is apparently endemic to southwest Mississippi.
During a recent survey to determine the status of Alloperla natchez Surdick & Stark, 1980, and Haploperla chukcho (Surdick & Stark 1980), two chloroperlid stoneflies en- demic to southwest Mississippi, specimens of an undescribed Neoperla were collected near the type localities of these species. A few additional specimens were found among unidentified material in my collection and from P. K. Lago of the University of Mis- sissippi, and a larger series was obtained from the Mississippi Entomological Muse- um (MEM), Mississippi State University. The material from the MEM was collected by malaise and blacklight traps at three sites in the Homochitto National Forest with the support of National Science Foundation grant DEB-9200856 awarded to Richard Brown.
Recent study of extensive Neoperla col- lections from throughout Mississippi and Alabama (Stark & Harris 1986, Stark & Lentz 1988.) suggests the new species is en- demic to the Homochitto National Forest and surrounding areas of southwest Missis- sippi and perhaps Louisiana.
The holotype is placed in the National Museum of Natural History (USNM) on in- definite loan from the Mississippi Ento- mological Museum. Paratypes are in the collections of the author (BPS), the Uni- versity of Mississippi (UM) and the Mis- Sissippi Entomological Museum. Termi- nology follows Stark & Lentz (1988).
Neoperla coxi, new species Figs. 1-8
Male. — Forewing length 8-10 mm. Gen- eral color yellow patterned with brown. Wings amber brown with darker veins; cos- ta and subcosta pale. Legs yellow except for diffuse dorsal brown spot near apex of fem- ora and a narrow longitudinal brown band extending from femur for *% of outer tibial margin. Cerci pale. Process of tergum 7 tri- angular, upturned in lateral aspect and armed ventrally with 4-5 prominent sen- silla basiconica. Mesal sclerite of tergum 8 with lateral margins parallel for most of length. Hemiterga slightly tapered apically, finger-like process of hemitergum slender and bent slightly ventrolaterad (Fig. 1). Tube of aedeagus slightly sinuate, 3.1 times as long as bulb; conspicuous spicules cover most of tube (Fig. 2). Sac sparsely armed in basal half by a pair of basolateral patches of slender spines (Figs. 2-3); apical arma- ture of scattered, thickened spines (Fig. 2), mid section unarmed (Figs. 2-3).
Female. —Forewing length 11-13 mm. Posterior margin of sternum 8 produced into a small rounded to slightly bilobed tab (Fig. 4). Spermathecal stalk membranous except for small apical section lined with fine brown setae (Fig. 5); stalk grotesquely inflated in some specimens.
Egg. —Length 0.31 mm; width 0.19 mm. Sessile collar surrounded by a single irreg-
46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-5. Neoperla coxi male and female genitalia. 1, Male terminalia; 2, Aedeagal tube, lateral; 3, Aedeagus, sac partially extruded, apical armature not shown; 4, Female sterna 8 and 9; 5, Vagina and spermathecal stalk, dorsal. T = tube; S = sac; B = bulb; H = hemitergum.
ular row of reticulation. Striae relatively straight, narrowed at each end and separat- ed by wide sulci. Sulci with 4—5 irregular aeropyle rows. Micropyles in sulci near equator (Figs. 7—8).
Mature nymph.—Body length 7—9 mm. General color yellow patterned with brown. Anterior of frons covered with a broad transverse brown pigment band; posterior margin of band sinuate, anterior margin
straight. Ocellar area covered with a small diffuse brown spot. Pronotal disk ringed by a sub-marginal pigment band. Each abdom- inal tergum with a narrow transverse pig- ment band (Fig. 6).
Etymology. —I am pleased to name this species for my friend and colleague, Prentiss G. Cox, in recognition of his interest and support of research at Mississippi College.
Types.— Holotype 6 and 104 6 and 48 2
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paratypes from Middleton Creek, Franklin County, Mississippi, TSN R4E Sec 31S, 1 Jun 1992, T. Schiefer, R. Fontenot (USNM, MEM). Additional paratypes, all from Mis- sissippi: Claiborne Co: Little Sand Creek, Rocky Springs, 14 May 1977, B. Stark, 1 2 reared (BPS). Sand Creek tributary, 0.5 mi W Hunt Road, 18 May 1978, B. Stark, 1 2 (BPS). Same location, 3 Jun 1986, B. Stark, J. Ballard, 1 2 (BPS). Copiah Co: Brushy Creek, Hwy 27, 0.5 mi E Hopewell, 20 May 1978, B. Stark, 1 6, 1 2 reared (BPS). Frank- lin Co: Clear Springs Lake, 4.5 mi SW Meadville, 30 May 1988, P. K. Lago, 1 6 (UM). Middleton Creek, TSN R4E Sec 31S, 9-15 Jun 1992, T. Schiefer, R. Fontenot, 6 6 (MEM). Same data except 29 Jun 1992, 4 6 (MEM). Same data except 30 Jun-—6 Jul 1992, 1 6 (MEM). McGehee Creek tribu- tary, TON R4E Sec 26SW, 1 Jun 1992, T. Schiefer, R. Fontenot, 88 6, 35 2 (MEM). Porter Creek, TSN R4E Sec 8NW, 1 Jun 1992, T. Schieffer, R. Fontenot, 6 6, 3 2 (MEM). Hinds Co: Sand Creek, Hunt Road, 24 Apr 1992, B. Stark, G. Gee, D. Kelly, 2 6 (BPS).
Discussion. —This description of N. coxi brings to 14 the number of Nearctic Neo- perla species. Using keys in Stark & Lentz (1988), males of NV. coxi are identified as N. stewarti Stark & Baumann, 1978, or N. os- age Stark & Lentz, 1988, and females are identified as N. stewarti. N. coxi males differ from all known Nearctic species in having most of the aedeagal tube spinulose (Fig. 2) and they also differ from both N. osage and N. stewarti in lacking prominent aedeagal armature in the basal half of the sac (Figs. 2-3). Separation of unassociated females of N. coxi from N. stewarti is more difficult, but the subgenital plate is slightly longer and the spermathecal stalk lining is less devel- oped in N. coxi; females of other Nearctic Neoperla differ from N. coxi in having most of the spermathecal stalk lined with brown setae. Nymphs are virtually identical in col- or pattern to N. robisoni Poulton & Stewart, 1986 (Ernst et al. 1986, Poulton & Stewart 1991), with perhaps a less extensive ocellar
47
Fig. 6. Neoperla coxi nymphal habitus.
pigment spot in N. coxi. The egg collar and posterior pole of N. coxi bear less prominent reticulations than do the eggs of N. stewarti.
Six Neoperla species are now known from Mississippi but only N. robisoni and N. carl- soni Stark & Baumann, 1978, have been
48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
a = Ait wie ee “Ai
a art m7 ie mere “asi” f 4 wth ertith rend te pansion uv
41,500 14rm WD?
Figs. 7-8. Neoperla coxi eggs. 7, Entire egg, lateral; 8, Detail of surface. St = striae, Su = sulci, M = micropyle.
collected from the same streams with N. August of 1992, and blacklight collections coxi. At the Middleton Creek site in Frank- were made 1 June, 29 June and 31 August. lin County, personnel of the Mississippi En- Early collections through 8 June contained tomological Museum operated a malaise exclusively N. coxi; N. robisoni males first trap continuously from 2 June through 31 appeared by 9 June and males of the 2 spe-
VOLUME 108, NUMBER 1
cies were collected together through 6 July. Thereafter, no males of N. coxi were taken but a single female was present in a mid- August malaise trap. N. robisoni males were collected through July and August and by late August, the first NV. carlsoni specimens were taken. Although elsewhere in Missis- sippi, NV. carlsoni has been collected as early as 22 June, at the Middleton Creek site a temporal emergence sequence which begins with N. coxi and ends with N. carlsoni may occur.
Acknowledgments
I thank R. L. Brown and T. Schiefer (Mis- sissippi State University) and P. K. Lago (University of Mississippi) for the loans of specimens. K. St. John (University of Mis- sissippi School of Dentistry) helped in the preparation of photomicrographs.
Literature Cited
Ernst, M. R., B. C. Poulton, & K. W. Stewart. 1986. Neoperla (Plecoptera: Perlidae) of the Ozark and
49
Ouachita Mountain region, and two new species of Neoperla.— Annals of the Entomological So- ciety of America 79:645-661.
Poulton, B. C., & K. W. Stewart. 1991. The stoneflies of the Ozark and Ouachita Mountains (Plecop- tera).— Memoirs of the American Entomologi- cal Society 38:1-116.
Stark, B. P., & R. W. Baumann. 1978. New species of nearctic Neoperla (Plecoptera: Perlidae), with notes on the genus.—Great Basin Naturalist 38: 97-114.
—., & S.C. Harris. 1986. Records of stoneflies (Plecoptera) in Alabama. — Entomological News 97:177-182.
—.,& D.L. Lentz. 1988. New species of Nearctic Neoperla (Plecoptera: Perlidae).— Annals of the Entomological Society of America 81:371-376.
Surdick, R. F., & B. P. Stark. 1980. Two new species of Chloroperlidae (Plecoptera) from Mississip- pi.— Proceedings of the Entomological Society of Washington 82:69-73.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 108(1):50—53. 1995.
Macrobrachium catonium, a new troglobitic shrimp from the Cayo District of Belize (Crustacea: Decapoda: Palaemonidae)
H. H. Hobbs III and Horton H. Hobbs, Jr.
(HHH, Jr) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. (deceased 22 March 1994); (HHH III) Department of Biology, P.O. Box 720, Wittenberg University, Springfield, Ohio 45501, U.S.A.
Abstract. —A new troglobitic shrimp, Macrobrachium catonium, is described from the Vaca Plateau, Cayo District of Belize. It is the third albinistic member of the genus known to occur along the Gulf of Mexico-Caribbean versant of Middle America. It may be distinguished from the Oxacan M. villalobosi Hobbs by the eye which is more pigmented and lacks an apical cleft, and by the shorter, often slightly upturned rostrum which does not reach the distal extremity of the antennal scale. It differs from the Tabascan M. acherontium in possessing a more attenuate (less vaulted) rostrum, and from both in that usually there are more than two ventral rostral teeth, and fewer subapical spines on the
appendix masculina.
The new shrimp described herein is the third albinistic member of the genus Mac- robrachium known to occur in caves on the Gulf of Mexico-Caribbean versant of Mid- dle America (Botosaneanu 1986). Macro- brachium villalobosi Hobbs (1973) has been reported from a single locality, Cueva del Nacimiento del Rio San Antonio, 10 km SSW of Acatan, Oaxaca, Mexico, and M. acherontium Holthuis (1977) from two caves in Tabasco, Mexico. Specimens of the new troglobite have been collected in two caves located approximately 46 km apart on the Vaca Plateau in the Cayo District of Belize close to Guatemala.
Macrobrachium catonium, new species
Description. —Rostrum (Fig. la, d) mod- erately high, weakly arched, and slightly de- flected or upturned anteriorly, tip not reach- ing distal extremity of antennal scale; dorsal margin with 6 to 9 teeth, as many as 3 bi- spinous (8 in holotype of which 3 bispi-
nous), and 1 or 2 epigastric; ventral margin with 2 to 5 teeth.
Carapace (Fig. la) with antennal spine arising slightly posterior to ventral part of orbital margin and hepatic spine almost di- rectly ventral to first epigastric tooth. Bran- chiocardiac groove prominent.
Abdomen (Fig. 1a) smooth, pleura of fifth abdominal somite with acute posteroven- tral angle, more anterior pleura rounded posteroventrally. Sixth somite 1.5 times as long as fifth, and telson 1.2 times longer than sixth; dorsal surface of telson (Fig. 1), 0) with anterior pair of spines situated at base of posterior third and posterior pair at about base of posterior sixth, slightly pos- terior to midway between anterior pair and median apex of telson; posterior margin of telson sharply contracted, forming broadly acute tip, and bearing 2 pairs of spines ven- tral to margin, more mesial pair decidedly overreaching apex of telson, lateral pair fall- ing short of apex; (holotype with row of 8 plumose setae between mesial pair of spines
VOLUME 108, NUMBER 1
and single simple submarginal seta dorsal- ly).
Eyes (Fig. la, d) moderately large, round- ed distally and with apical black to purplish pigment spot; cornea over pigmented area lacking facets.
Antennule (Fig. la, d) with proximal po- domere of peduncle longer than combined length of distal 2 podomeres, these subequal in length, and distal podomere falling short of base of lateral spine on antennal scale; distolateral spine on basal podomere reach- ing slightly beyond midlength of penulti- mate podomere; holotype with lateral long flagellum about 5 times length of postorbital length of carapace, mesial flagellum about 3 times as long. Antenna (Fig. la, d, 1) with peduncle as illustrated, basal segment with ventrolateral spine, flagellum about 7.3 times as long as postorbital carapace length. Antennal scale 2.5 times as long as broad, with lateral margin almost straight.
Gnathal appendages (Fig. 1b, e, h, 1, k, m) as figured. Third maxilliped reaching slightly beyond midlength of antennal scale.
First pereiopod (Fig. 1a) overreaching an- tennal scale by length of dactyl, latter sub- equal in length to mesial margin of palm of chela; carpus about twice length of chela and slightly shorter than merus. Second pereio- pod (Fig. la, n) overreaching antennal scale by only slightly more than length of chela; latter with fingers slightly longer than smooth palm; opposable margin of fixed finger with 1 very small corneous tubercle near base, otherwise fingers lacking tubercles, but both fingers with scattered fine setae and sub- apical clusters of curved stiff ones; carpus 1.2 times as long as either propodus or me- rus, and merus 1.4 times longer than ischi- um. Third pereiopod overreaching antennal scale by propodus and 3 length of carpus; latter almost half as long as merus, and me- rus 2.5 times as long as ischium. Fourth pereiopod overreaching antennal scale by dactyl and slightly less than half length of propodus; propodus approximately 2.7 times length of dactyl; carpus about half as
51
long as merus, and latter almost 2.5 times as long as ischium. Fifth pereiopod over- reaching antennal scale by dactyl and % length of propodus; propodus almost 5 times length of dactyl and 1.7 times length of car- pus, latter little less than 74 length of merus; merus 2.6 times length of ischium.
First pleopod (Fig. 1c) with exopodite 2.2 times as long as endopodite. Second pleo- pod (Fig. 1f) with exopodite 1.2 times length of endopodite and latter with appendix masculina (Fig. 1f, g) reaching distinctly be- yond its midlength. Lateral ramus of uro- pod (Fig. lo) with straight lateral margin bearing fixed spine and longer, slenderer, movable one at its mesial base.
Branchial formula typical of that of many, if not most, other members of Macrobra- chium: 5 pleurobranchs corresponding to pereiopods, 2 arthrobranchs at base of third maxilliped, podobranch on coxa of second maxilliped, and epipodites on second max- illa and first maxilliped.
Size. —Carapace length of male holotype 9.9 mm; that of 12 females, none of which Ovigerous, ranging from 8.8 to 14.4 mm.
Color. —Lacking pigment except for eye spot which purplish to black.
Type locality.—Lake in Actun Chapat (cave), Cayo District, Belize. This cave is located on the northern end of the Vaca Plateau and although it has not been fully explored it is estimated to be about 2 km in length. Shrimp were found in the lake passage, apparently a perched overflow route with a series of small lakes extending more than 150 m. The shrimp were numerous on the silt-covered substrate of the lakes which were as deep as 6 m. The troglobitic crab, Typhlopseudothelphusa acanthochela Hobbs, 1986, occurred in small numbers and the catfish with reduced eyes, Rhamdia laticauda typhla Greenfield, Greenfield, & Woods, were fairly abundant. Bats were roosting over the lakes and probably were insectivorous Natalus sp. and Mormoops sp. and frugivorous Corollia sp. and Glosso-
34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
if
g (i,ksm)
Fig. 1. Macrobrachium catonium, new species (all appendages from left side of holotype except c and h from female paratype from type locality): a, Lateral view; b, Third maxilliped; c, f, First pleopods; d, Dorsal view of anterior region of carapace, including cephalic appendages; e, Mandible; g, Appendices masculina and interna; h, First maxilla; i, Second maxilla; j, Dorsal view of caudal end of telson; k, First maxilliped; 1], Ventral view of basal part of antenna; m, Second maxilliped; n, Chela of second pereiopod; 0, Dorsal view of telson and right uropod. (Scales in mm.)
VOLUME 108, NUMBER 1
phaga sp. Temperature of the lake water in December 1991 was 25°C.
Specimens examined. —In addition to 1 6 and 9 2 from the type locality (Actun Cha- pat, 5 Dec 1992, R. Foster & W. R. Elliott, coll.), specimens were examined from Tunkul Cave, Chiquibal System, Cayo Dis- trict, Belize, 1 2, 24 Mar 1986, D. Coons, coll. and from ‘“‘Chiquibal System,’’ Cayo District, Belize, 2 9, 8 Mar 1986, D. Coons, coll. (This cave is located approximately 46 km south of Actun Chapat on the south end of the Vaca Plateau near Guatemala.)
Disposition of types. —The holotype (USNM 260328) and 10 of the paratypic females are deposited in the National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. A paratypic fe- male is in the collection of the Texas Me- morial Museum, Austin, and another in the Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands.
Relationships.—Macrobrachium caton- ium 1s Closely related to the two Mexican troglobitic shrimps assigned to the genus, M. villalobosiand M. acherontium. The three are markedly similar suggesting common ancestry. The apically notched eyes and long (reaching to or beyond the distal end of the antennal scale), slender, straight rostrum of M. villalobosi sets it apart from the other two, and the comparatively slender (nearly Straight rather than with a strongly convex ventral margin) rostrum with usually more than two ventral teeth in M. catonium dis- tinguishes it from M. acherontium. In ad- dition, there are fewer subapical setae on the appendix masculina of M. catonium than in the other two species. Holthuis (1977: 191) reported the presence of 5 pleuro-
53
branchs but no other gills, exopods, or epi- pods in M. acherontium whereas M. caton- lum possesses 2 arthrobranchs at the base of the third maxilliped, and epipodites on the second and first maxillipeds.
Etymology.—Catonium (L.), the lower world, noting the subterranean habitat of this shrimp.
Acknowledgments
Thanks are extended to T. Miller for lo- cating and sending us the misplaced shrimp collected from the Chiquibul Cave system in 1986, and to D. Coons who collected three of the specimens. We are also grateful to R. Foster and W. R. Elliott who collected the shrimp from the type locality, and to the latter who provided us with the infor- mation recorded in the description of the type locality. For their critical review of the manuscript appreciation is extended to C. W. Hart, Jr., B. F. Kensley, and A. B. Wil- liams.
Literature Cited
Botosaneanu, L. 1986. Stygofauna Mundi. A. Fau- nistic, distributional, and ecological synthesis of the world fauna inhabiting subterranean waters (including the marine interstitial). E. J. Brill, Leiden, Netherlands, 740 pp.
Hobbs, H. H., Jr. 1973. Two new troglobitic shrimps (Decapoda: Alpheidae and Palaemonidae) from Oaxaca, Mexico. — Association for Mexican Cave Studies Bulletin 5:73-80.
Holthuis, L. B. 1977. Cave shrimps (Crustacea De- capoda, Natantia) from Mexico. Part III. Fur- ther results of the Italian zoological missions to Mexico, sponsored by the National Academy of Lincei (1973 and 1975).—Problemi attuali di Scienza e di Cultura, Accademia Nazionale dei Lincei 171(3):173-195.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
108(1):54-60. 1995.
Procambarus (Ortmannicus) nueces (Decapoda: Cambaridae), a new crayfish from the Nueces River Basin, Texas
Horton H. Hobbs, Jr., and H. H. Hobbs III
(HHH, Jr.) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. (deceased 22 March 1994); (HHH III) Department of Biology, P.O. Box 720, Wittenberg University, Springfield, Ohio 45501, U.S.A.
Abstract. —Procambarus (Ortmannicus) nueces is described from a small, slug- gish tributary to the Nueces River in Atascoso County, Texas. It may be distinguished from its closest relative, P. (O.) zonangulus Hobbs & Hobbs, by the less attenuated distal part of the male first pleopod which lacks a cephal- omesial shoulder and bears a more robust mesial process.
Four specimens of this new crayfish, in- cluding a first form male and three juve- niles, were collected for the Smithsonian In- stitution by Bruce B. Collette and Oliver S. Flint from the type locality on 10 September 1960. While its distinct features were rec- ognized several years ago and futile searches made in several collections for additional specimens, we were reluctant to describe this new species until additional material be- came available. Not until July 1993, when the second author, assisted by Howard and Stephen Kronk, collected in the area where the original lot had been taken, were two additional specimens secured, both pro- cured with the aid of a seine in the same segment of stream in which Collette and Flint collected almost 39 years earlier. Al- though other aquatic habitats in the area were sampled in July and a search was made for burrows, no other crayfish were found. We offer the following description of this new species that is based on six specimens (3 adults and 3 juveniles) from the type lo- cality.
Procambarus (Ortmannicus) nueces, new species Fig. 1, Table 1
Diagnosis. —Body pigmented, eyes well developed. Rostrum of adults with well de-
fined marginal spines but lacking median carina. Carapace with single moderately strong cervical spines. Areola 14.7 to 17.3 times as long as wide and constituting 32.0 to 36.4% of total length of carapace and 44.0 to 47.9% of postorbital length. Suborbital angle very small to virtually obsolete; post- orbital ridge with small, corneous cephalic tubercle; hepatic area weakly tuberculate; branchiostegal spine well developed. An- tennal scale almost twice as long as broad, widest slightly proximal to midlength. Is- chia of third and fourth pereiopods with simple hooks overreaching basioischial ar- ticulation and lacking opposing tubercles on corresponding basis; coxa of fourth pereio- pod with strong bulbiform boss, that of fifth much smaller and flattened. First pleopods of first form male reaching coxa of third pereiopods, symmetrical but not conspic- uously tapering distally; cephalomesial margin lacking even rudiment of hump or shoulder. Terminal elements consisting of: (1) short, tapering mesial process directed caudodistally and inclined laterally, and (2) cephalic process obscuring much of central projection in cephalic aspect, corneous, sub- acute, tapering from broad base, inclined mesially and gently curved caudally, its apex lying slightly mesial to central projection; caudal element consisting of corneous, ta-
VOLUME 108, NUMBER 1
pering acute caudal process lying against caudal surface of central projection and in- conspicuous, non-corneous, setiferous cau- dal knob at lateral base of cephalic process; and corneous central projection, largest of terminal elements, inclined mesially and ta- pering to apex which directed caudodistally and slightly laterally. Annulus ventralis about 1.7 times as wide as long, dextral half elevated little more than sinistral; sinus originating on median line, disappearing under dextral wall and emerging on caudal flank of dextrally directed tongue, slightly posterior to midlength of annulus, where crossing median line and, in gentle arc, coursing caudally to margin of annulus. Sternum cephalic to annulus deeply cleft and bearing few prominent tubercles. Una- dorned postannulur sclerite more than half as broad and as long as annulus. First pleo- pod present in female.
Holotypic male, form I.—Cephalothorax (Fig. la, 1) subcylindrical in section, only slightly broader than high. Abdomen nar- rower than thorax (20.1 and 24.1 mm). Greatest width of carapace greater than height at caudodorsal margin of cervical groove. Cephalic section of carapace twice as long as areola, length of latter 33.1% of entire length of carapace (44.9% of postor- bital carapace length). Surface of carapace punctate dorsally, granulate to tuberculate laterally. Rostrum not noticeably deflected ventrally, with basally subparallel margins gently converging to level of distal margin of penultimate podomere of antennular pe- duncle where diverging and forming well developed marginal spines marking base of acumen, latter reaching distal extremity of ultimate podomere of antennular peduncle; dorsal surface concave with many small se- tiferous punctations particularly in basal half. Subrostral ridges evident in dorsal as- pect for very short distance at caudal margin of orbit. Postorbital ridges well developed, grooved dorsolaterally and bearing well de- veloped spiniform tubercles at cephalic ex- tremities. Suborbital angle very weak, vir- tually obsolete. Branchiostegal spines com-
55
paratively well developed. Single cervical spine on both sides of carapace rather small but well defined and acute.
Abdomen (Fig. 1j) slightly shorter than carapace. Pleura of third through fifth seg- ments subtruncate to rounded with caudov- entral extremities subangular. Cephalic sec- tion of telson (Fig. le) with 2 spines in each caudolateral corner, mesial one on each side movable; caudal margin of caudal section with shallow median excavation at end of dorsomedian groove extending length of section. Cephalic lobe of epistome (Fig. 10) subcylindrical with elevated (ventrally) ir- regular margins and bearing acute antero- median projection; central area subplane except for anteromedian groove; distinct anteromedian fovea and median longitu- dinal groove present on main body. Ventral surface of proximal podomere of antennular peduncle with strong spine slightly proxi- mal to midlength. Antennal peduncle with small spiniform tubercles on basis and is- chium; flagella reaching midlength of telson. Antennal scale (Fig. lh) 2.2 times as long as broad, widest slightly proximal to mid- length; greatest width of lamella more than twice width of thickened lateral part.
Third maxilliped extending cephalically to level of ultimate podomere of antennular peduncle; ischium not produced distolater- ally, its ventral surface densely studded with plumose setae.
Right chela (Fig. 1m) subovate in cross section, not strongly depressed. Mesial sur- face of palm with row of 6 tubercles sub- tended dorsolaterally by additional row of 6 and few scattered ones and ventrally by row of 4. Entire palm tuberculate but ven- trolateral surface more weakly so. Dorsal and ventral longitudinal ridges weak on both fingers. Opposable surface of fixed finger with dorsal row of 20 tubercles and lower one of 8; dorsal row, with fifth from base largest, extending from base of finger shght- ly beyond level of locking tubercle (more distal ones too small to be included in il- lustration); ventral row much shorter, ex- tending along middle third of finger and
56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
consisting of locking tubercle, which much larger than others and, not visible in dorsal aspect, 7 proximal and 3 distal to it; band of minute denticles present ventral to dorsal row of tubercles and suddenly broadening at level of locking tubercle, covering entire opposable surface distally to base of cor- neous tip; except for few small tubercles on proximodorsal and proximoventral sur- faces, finger otherwise with setiferous punc- tations. Opposable margin of dactyl with dorsal row of 24 small to minute, rounded tubercles along proximal % of finger and ventral row of 13 along second fourth of finger from base; band of minute denticles situated between tubercles of dorsal row proximally, broadening at level of proximal end of ventral row and covering opposable margin to base of corneous tip of finger; mesial surface with row of 6 tubercles along proximal third flanked by few others prox- imally but, except for few additional tiny tubercles proximally, finger with setiferous punctations. Both fingers with punctations forming rows flanking inconspicuous lon- gitudinal ridges and those with longer setae projecting toward opposable member. Car- pus of chela longer than broad, bearing prominent oblique furrow dorsally; tuber- culate mesially and on mesial half of dorsal surface; ventral surface with usual 2 distal tubercles, otherwise sparsely punctate. Me- rus tuberculate dorsally, distomesially, and ventrally; 2 premarginal tubercles larger than others on dorsodistal surface; ventral sur- face with mesial row of 16 tubercles and less clearly defined lateral one of 10. Ischium with ventromesial row of 5 tubercles.
Hooks on ischia of third and fourth per- eiopods (Fig. 1f) simple, both overreaching basioischial articulation and neither op- posed by tubercle on corresponding basis. Coxa of fourth pereiopod with prominent bulbiform boss; that of fifth with much smaller compressed one. Sternum between third, fourth, and fifth pere1opods compar- atively deep with mat of plumose setae ex- tending mesially from ventrolateral mar- gins.
First pleopods (Fig. 1b, c, d, g, n) as de- scribed in “Diagnosis.” Uropod (Fig. le) with both lobes of basal podomere bearing small acute spine, both rami with distola- teral spines, and distomedian spine on me- sial ramus situated distinctly proximal to boadly rounded distal margin.
Allotypic female. — Differing from holo- type, except in secondary sexual characters, as follows: 2 marginal spines on sinistral side of rostrum; acumen slightly overreach- ing antennular peduncle; cephalic section of telson with 3 spines in each caudolateral corner, more mesial pairs on each side mov- able, mesialmost spines minute; cephalic lobe of epistome more concave; antennal flagellum reaching tergum of sixth abdom- inal segment; 5 tubercles present in row ven- tral to mesial row on palm of chela (Fig. 1p); opposable margin of fixed finger with single dorsal row of 10 small tubercles along basal half, locking tubercle (only member of ventral row present) situated near mid- length of finger and minute denticles on both fingers confined to single row; opposable margin of dactyl with dorsal row of 12 small tubercles extending from base of finger to level of locking tubercle on fixed finger, ven- tral row represented by only 2 tubercles on proximal third of finger; merus with 14 tu- bercles in ventromesial row and 11 in ven- trolateral row (rows not distinct in part, so numbers approximate); ventromesial sur- face of ischium with row of 4 tubercles.
Annulus ventralis and adjacent sternal el- ements as described in “‘Diagnosis.”
Color notes. —Dorsum of carapace most- ly pale tan but rostrum darker and with red- dish suffusion; rostral and postorbital ridges bluish gray, latter flanked ventrolaterally by reddish splotch; mandibular adductor areas covered by reticulate dark brown pattern; dorsolateral part of branchiostegites with conspicuous black stripe extending from brownish-to-almost-black cervical groove to caudal margin of carapace; remainder of lateral surface of carapace pale gray with darker gray mottlings; cervical spine white and caudal flange of carapace edged in dark
VOLUME 108, NUMBER 1
Fig. 1.
S//
Procambarus (Ortmannicus) nueces, new species (all from holotype except i, p from allotype): a,
Lateral view of carapace; b, Mesial view of first pleopod; c, Caudal view of same; d, Cephalic view of same; e, Dorsal view of sixth abdominal segment, telson, and uropods; f, Basal podomeres of third, fourth, and fifth pereiopods; g, Lateral view of first pleopod; h, Antennal scale; i, Annulus ventralis and flanking sternal elements; j, Lateral view of abdomen; k, Caudal view of mandible; 1, Dorsal view of carapace; m, Dorsal view of distal podomeres of cheliped; n, Caudal view of first pleopods; 0, Epistome; p, Dorsal view of distal podomeres of
cheliped.
gray. Abdomen with broad dorsomedian black stripe extending from carapace through anterior third of sixth abdominal segment, most of which pinkish tan; stripe interrupted by narrow transverse bands of pinkish tan along caudal margins of terga; black stripe flanked laterally by narrower
pinkish tan stripe, and it, in turn, separated from mostly reddish pleura by concave splotches of gray-merging-to-black on bases of pleura, dark color continuing along cau- dodorsal margins of pleura; telson and uro- pods, particularly lateral ramus of latter, reddish brown. Antennae and antennules
58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.—Measurements (mm) of Procambarus (O.) neuces, new species.
Holotype Allotype Paratype
Carapace
Entire length 48.4 54.0 40.3
Postorbital length 35.6 39.3 40.3
Width 24.1 26.1 20.1
Height 22.0 24.0 19.0 Areola
Width 1.0 1.0 1.0
Length 16.0 Wes 14.7 Rostrum
Width 19 9.0 7.0
Length 14.5 17.7 11.0 Right chela
Length, palm
mesial margin 16.1 10.8 12.6 Palm width 13.6 2 10.4 Length, lateral margin 50.9 34.3 41.5*
Dactyl length 30.5 hell Dal Abdomen
Width 20.1 24.0 Wer
Length 47.0 53.0 41.0
* Approximate, tip broken.
greenish cream; antennal scales also pale laterally, lamella suffused with pink. Che- liped with distal part of merus and dorsal and distomesial parts of carpus olive tan, remainder of these and more proximal po- domeres pinkish cream; palm of chela pink- ish cream dorsally with olive to dark green- ish brown tubercles, deeper pink ventrally; fingers with greenish suffusion proximo- dorsally, and dactyl also proximomesially, fading quickly distally to very pale cream. Remaining pereiopods with merus and car- pus greenish dorsally, otherwise, cream to pinkish cream.
Size. —See Table 1 for measurements of the only three adult individuals available.
Type locality. —Sluggish stream tributary of the Nueces River 4.3 miles (6.9 km) south of Jourdanton at junction of State Routes 97 and 173, Atascosa County, Texas. About
5 m at its widest point, the stream was ex- tremely turbid when visited 24 July 1993 and very little flow was noted. The channel was slightly sinuous and tree limbs and oth- er vegetative debris impeded the flow in various sections. The substrate consisted of sand, gravel, and in the pooled areas that were as deep as 1.5 m, the bottom was cov- ered with silt. Sedges and grasses as well as composite plants dominated the riparian vegetation on the upstream (east) side of the bridge along St. Rte. 97 and the water re- ceived direct sunlight for much of the day- light hours.
Disposition of types.—The holotypic male, form I, and allotype (USNM 260326 and 260327) are deposited in the National Mu- seum of Natural History, Smithsonian In- stitution, as are the paratypes consisting of 1 6I, 1 juv 6, and 2 juv 2.
Relationships. —Procambarus (O.) nueces, new species, belonging to a closely allied group of species often treated as the blan- dingi “group” of the subgenus (see Ortmann 1905:100, Hobbs 1962:286), has its closest affinities with P. (O.) zonangulus Hobbs & Hobbs, 1990. Within this assemblage are the following currently recognized species: P. (O.) blandingii (Harlan 1830), P. (O.) acutus (Girard 1852), P. (O.) acutissimus (Girard 1852), P. (O.) lecontei (Hagen 1870), P. (O.) hayi (Faxon 1884), P. (O.) viaeviridis (Faxon 1914), P. (O.) cuevachicae Hobbs, 1941, P. (O.) caballeroi Villalobos, 1944, P. (O.) verrucosus Hobbs, 1952, P. (O.) lopho- tus Hobbs & Walton, 1960, P. (O.) texanus Hobbs, 1971, P. (O.) geminus Hobbs, 1975, P. (O.) zonangulus, and P. (O.) nechesae Hobbs, 1990. Four of these crayfishes typ- ically have symmetrical first pleopods: P. (O.) viaeviridis, P. (O.) texanus, P. (O.) zon- angulus, and P. (O.) nueces; at least one population of P. (O.) cuevachicae has nearly symmetrical ones. The new species differs from viaeviridis, which is the most marginal member of the group, in many conspicuous respects: among them, P. (O.) nueces has
VOLUME 108, NUMBER 1
well developed marginal spines on the ros- trum; the setiferous caudal knob of the first pleopod is laterally rather than cephalically situated, is not conspicuously produced, and the setae partly obscure other terminal el- ements; and the central area of the annulus ventralis is not deeply excavate. Distin- guishing it from P. (O.) cuevachicae are a more strongly tapering, subacute, less com- pressed cephalic process and a less twisted central projection of the first pleopod in first form males, and a straight, not reflexed, hook on the ischium of the fourth pereiopod that is not opposed by a tubercle on the corre- sponding basis. From fexanus it differs in that the cephalic process and central pro- jection are bent caudally at distinctly less than a right angle to the shaft of the ap- pendage, and there is not even a hint of a shoulder on the appendage proximal to the terminal elements; the cephalic lobe of the epistome is subcircular instead of subtrian- gular in outline; in the female there are fewer tubercles adorning the sternum anterior to the annulus, and the postannular sclerite is about *4 as long as broad and 1s as long as the annulus ventralis. The new crayfish dif- fers from P. (O.) zonangulus in that the first pleopod is not distinctly tapering distally and it lacks any trace of a shoulder on the cephalic or cephalomesial surface proximal to the terminal elements; the mesial process is less attenuate and comparatively stockier; in the female there are markedly fewer tu- bercles on the sternum anterior to the an- nulus, and the length of the postannular sclerite is as great as that of the annulus along its median longitudinal axis.
Acknowledgments
We are grateful to B. B. Collette and O. S. Flint for securing the first specimens of this new crayfish of which we are aware and to H. and S. Kronk for their assistance in obtaining additional specimens. For their critical reading of the manuscript we extend
59
thanks to J. F. Fitzpatrick, Jr., of the Uni- versity of South Alabama, and C. W. Hart, Jr., and R. B. Manning, of the National Mu- seum of Natural History.
Literature Cited
Faxon, W. 1884. Descriptions of new species of Cam-
barus; to which is added a synonymical list of
known species of Cambarus and Astacus. — Pro- ceedings of the American Academy of Arts and
Sciences 20:107-158.
1914. Notes on the crayfishes in the United States National Museum and the Museum of Comparative Zoology with descriptions of new species and subspecies to which is appended a catalogue of the known species and subspe- cies. — Memoirs of the Museum of Comparative Zoology at Harvard College 40:352—427. Girard, C. 1852. A revision of the North American
Astaci with observations on their habits and geographical distribution.— Proceedings of the Academy of Natural Sciences of Philadelphia 6:87-91.
Hagen, H. A. 1870. Monograph of the North Amer- ican Astacidae.—Illustrated Catalogue of the Museum of Comparative Zodlogy at Harvard College 3:viiit+ 109.
Harlan, R. 1830. Description of a new species of the genus Astacus.— Transactions of the American Philosophical Society 3(15):464—-465.
Hobbs, H. H., Jr. 1941. A newcrayfish from San Luis
Potosi, Mexico.— Zoologica 26(1):1-4.
1952. A new crayfish from Alabama with notes on Procambarus lecontei (Hagen).—Pro- ceedings of the United States National Museum 102(3297):209-219.
. 1962. Notes on the affinities of the members
of the Blandingii Section of the crayfish genus
Procambarus (Decapoda, Astacidae).— Tulane
Studies in Zoology 9(5):273-293.
1971. New crayfishes of the genus Procam- barus from Alabama and Texas (Decapoda: As- tacidae). — Proceedings of the Biological Society of Washington 84(11):8 1-94.
1975. New crayfishes (Decapoda: Cambari- dae) from the southern United States and Mex- ico.—Smithsonian Contributions to Zoology 201:1-34.
1990. On the crayfishes (Decapoda: Cam- baridae) of the Neches River Basin of Eastern Texas with the descriptions of three new spe- cies.— Proceedings of the Biological Society of Washington 103:573-597.
—., & H. H. Hobbs III. 1990. A new crayfish
60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
(Decapoda: Cambaridae) from southeastern persal over the United States.— Proceedings of Texas. — Proceedings of the Biological Society of the American Philosophical Society 44(180):91- Washington 103:608-613. 136.
—, & M. Walton. 1960. A new crayfish of the Villalobos, A. 1944. Estudios de los cambarinos mex- genus Procambarus from southern Alabama icanos, III: Una especie nueva de Procambarus, (Decapoda, Astacidae).— Proceedings of the Bi- Procambarus caballeroi n. sp.—Anales del In- ological Society of Washington 73(20):123-129. stituto de Biologia, Universidad Aut6noma de
Ortmann, A. E. 1905. The mutual affinities of the México 15(1):175-184.
species of the genus Cambarus, and their dis-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
108(1):61-67. 1995.
On a new Somanniathelphusa Bott, 1968, from Vietnam (Crustacea: Decapoda: Brachyura: Parathelphusidae)
Peter K. L. Ng and Takeharu Kosuge
(PKLN) Department of Zoology, National University of Singapore, Kent Ridge, Singapore 0511, Republic of Singapore;
(TK) % Department of Biology, Faculty of Science, University of the Ryukyus, Nishihara-cho, Okinawa 903-01, Japan
Abstract. —A new species of parathelphusid freshwater crab, Somanniathel- phusa pax, 1s described from Hanoi, Vietnam. The new species seems to be most closely allied to species from southern China, but differs in the form of the carapace, male abdomen and male gonopods.
Specimens of a freshwater crab recently obtained from a market in Hanoi, Vietnam, proved to belong to an undescribed species of Somanniathelphusa Bott, 1968 (family Parathelphusidae Alcock, 1910). The genus Somanniathelphusa s. str. aS now recog- nized (Bott 1970, Ng 1988, Ng & Naiyanetr 1993, Naiyanetr 1994), contains six de- scribed species: S. brevipodum Dai, Song, He, Cao, Xu & Zhong, 1975 [China], S. chongi (Wu, 1935) [China], S. falx Ng & Dudgeon, 1992 [China], S. sinensis (H. Milne Edwards, 1853) (type species) [Chi- na], S. taiwanensis Bott, 1968 [Taiwan], and S. zanklon Ng & Dudgeon, 1992 [Hong Kong]. Naiyanetr (1994) recently reviewed the genus Somanniathelphusa and showed that it should be split into four genera: So- manniathelphusa s. str., Sayamia Naiya- netr, 1994, Esanthelphusa Naiyanetr, 1994, and Chulathelphusa Naiyanetr, 1994.
The freshwater crab fauna of Vietnam is poorly known. Many of the Indo-Chinese species described by H. Milne Edwards (1853), A. Milne Edwards (1869), De Man (1904) and Rathbun (1904, 1905), did not have precise collection localities. With re- gards to the parathelphusids belonging or allied to Somanniathelphusa, Potamon (Parathelphusa) prolatus was described from “Mois Chero”? in northern Cochinchina
(Rathbun 1902:186), a location which is now supposed to be part of northern Vietnam (Turkay & Naiyanetr 1987:392). Potamon (Parathelphusa) prolatus, incorrectly syn- onymized with Sayamia dugasti (Rathbun 1902) by Bott (1970:112), is a valid species and should be transferred to Esanthelphusa (P. Naiyanetr, pers. comm.). Chulathelphu- sa neisi was described from somewhere in Cochinchina (Rathbun 1902:186), which may be somewhere in or near Vietnam. Balss (1914:408) reported Somanniathelphusa si- nensis from Tonkin in Vietnam, but Ng & Dudgeon (1992:757) noted that this record is doubtful and his specimens must be re- examined.
The present paper describes the new spe- cies, Somanniathelphusa pax. The abbre- viations G1 and G2 are for the male first and second pleopods respectively. Mea- surements are given in millimeters, in the sequence carapace width by carapace length. Specimens examined are deposited in the Zoological Reference Collection (ZRC), De- partment of Zoology, National University of Singapore.
Systematic Account
Family Parathelphusidae Alcock, 1910 Genus Somanniathelphusa Bott, 1968
62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
Somanniathelphusa pax, new species Figs. 1-3
Material examined. —Holotype male (carapace 41.9 by 32.8 mm) (ZRC), Hanoi, Vietnam, from Dong Xuan market, leg. T. Kosuge, 19 Oct 1993. Paratype female (car- apace 34.8 by 26.6 mm) (ZRC), same data as holotype. 2 males (larger, carapace 30.9 by 24.8 mm), 2 females (larger, carapace 31.2 by 25.0 mm) (ZRC), market in Hanoi, Vietnam, leg. A. U. Kara, September 1994.
Description of holotype male. —Carapace broader than long, surfaces smooth, trans- versely convex (Figs. 1, 2A). Epigastric cris- tae sharp, distinct, separated by deep lon- gitudinal groove; postorbital cristae sharp, distinct, reaching to beginning of shallow cervical grooves, inner edge of crista reach- ing to slightly below inner edge of epigastric crista; beyond cervical grooves, cristae are rounded, curving gently to meet bases of first epibranchial teeth; median H-shaped depression distinct, relatively deep (Figs. 1,
Somanniathelphusa pax, new species. Holotype male, carapace 41.9 by 32.8 mm (ZRC). Dorsal view.
2A, B). Frontal margin gently sinuous, con- fluent with sinuous, smooth supraorbital margin. External orbital angle triangular, inner margin distinctly shorter than outer, outer margin gently convex to straight. An- terolateral margin with 3 strongly devel- oped, sharp epibranchial teeth, first trian- gular, last spiniform, first and second teeth directed forwards, last tooth directed obliquely outwards (Fig. 2A, B).
Ischium of third maxilliped rectangular, much longer than wide, with deep subme- dian, longitudinal sulcus; basis separated from ischium by distinct suture; merus squarish, medially depressed; exopod long, reaching to middle of margin of merus, with pronounced subdistal tooth on inner margin and long flagellum (Fig. 2C).
Chelipeds strongly asymmetrical, left larger; outer surfaces of both chelae smooth (Figs. 1, 2F). Merus with small but distinct subterminal spine on dorsal margin. Carpus with strong, sharp spine on inner distal mar- gin; inner margin granulated (Fig. 1). Fin-
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Fig. 2. Somanniathelphusa pax, new species. Holotype male, carapace 41.9 by 32.8 mm (ZRC). A, carapace; B, frontal view; C, left third maxilliped (hairs omitted); D, anterior sternites; E, last left ambulatory leg; F, enlarged (left) chela; G, abdomen. Scales = 5.0 mm.
64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
gers of both chelae longer than palm, fingers of smaller chela not distinctly gaping when closed, with numerous denticles along cut- ting edges; fingers of larger chela strongly gaping when closed, cutting edges lined with several large and numerous smaller teeth (Riga):
Second pair of ambulatory legs longest. Merus with very strong, distinct dorsal sub- terminal spine (Fig. 2E).
Male abdominal cavity reaches an imag- inary line joining anterior edges of bases of chelipeds (Fig. 2D). Suture between ster- nites 2 and 3 only present medially, lateral parts obscured; suture between sternites 3 and 4 very shallow, indistinct (Fig. 2D). Male abdomen distinctly T-shaped, telson broad- ly triangular, lateral margins gently con- cave, gradually converging towards round- ed tip; segment 6 subequal in length to tel- son, strongly constricted at subproximal part, with low but distinct submedian tran- verse ridge, length 1.2 times greatest width, distal width 1.3 times proximal width; seg- ments 3-5 progressively more trapezoidal, segments 1 and 2 narrow (Fig. 2G).
G1 terminal and subterminal segments not demarcated (Fig. 3A—C); distal part slender, distal half gradually curving later- ally outwards, tip bent laterally; basal part very broad, outer margin distinctly convex when viewed ventrally (Fig. 3D-F). G2 with very short distal segment, 0.07 times length of elongated basal segment (Fig. 3G).
Paratype female.—The paratype female is fully mature. One ofits chelae is distinctly larger than the other, but the degree of en- largement of the major chela is not as pro- nounced as in the holotype male. The bran- chial regions of the female are distinctly in- flated and appear swollen, much more than in the holotype male. In some of the large species of Somanniathelphusa (e.g., S. bangkokensis and S. sexpunctata), larger specimens generally appear to be less in- flated than smaller ones. The holotype male of S. pax is distinctly larger than the female (carapace 41.9 by 32.8 mm against 34.8 by
26.6 mm). The paratype female agrees with the holotype male in all other non-sexual characters. The non-type specimens show some variation in the form of the frontal margin, from sinuous to almost straight.
Color.—Young males and females are beige to dull brown on all dorsal surfaces, the ventral surfaces being dirty white. Large males vary from brown to purple on the dorsal surfaces.
Discussion. —In characters such as cristae of the carapace, shape of the male abdomen and gonopods, S. pax clearly belongs to So- manniathelphusa (sensu Naiyanetr 1994) which occurs in China and Taiwan (Dai et al. 1975, Bott 1970, Ng & Dudgeon 1992). The carapace of S. pax is more rectangular compared to species like S. sinensis and S. zanklon which are more oval. The male ab- dominal segment 6 of S. pax is similar in shape to that of S. sinensis, S. zanklon and S. falx, but its telson is less triangular, with the distal part broader. Compared to S. chongi, the male abdominal segment 6 of S. pax is also proportionately less elongate (length to maximum width ratio 1.2 against 1.4). The deep and distinct longitudinal groove on the pollex of the enlarged male chela is diagnostic for S. pax. The Gls of S. pax however, differs markedly from all known congeners in having the tip bent lat- erally. In congeners, the tip is either straight, slightly folded laterally or hooked down- wards (fide Wu 1935, Dai et al. 1975, Ng & Dudgeon 1992).
As for allied genera and species known from Vietnam, S. pax differs from Esan- thelphusa prolatus in that the anterolateral margin of E. prolatus is more convex, with the smaller epibranchial teeth directed for- wards (margin less convex in S. pax, with the epibranchial teeth relatively larger and the last tooth directed obliquely outwards), the outer surface of the pollex of the en- larged male chela is smooth, without a lon- gitudinal groove (longitudinal groove dis- tinct in S. pax) and the male abdominal segment 6 is relatively shorter (length to
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65
D if F F:
Fig. 3. Somanniathelphusa pax, new species. Holotype male, carapace 41.9 by 32.8 mm (ZRC). A-C, left G1; D-F, distal part of left G1; G, left G2. A, D, ventral views; C, F, dorsal views; B, E, ventro-marginal views. Scales = 1.0 mm.
maximum width ratio 0.9 against 1.2) with the distal width much wider than the prox- imal width (distal to proximal width ratio 1.6 in E. prolatus, 1.3 in S. pax) (fide Rath- bun 1905:245, fig. 59, pl. XI fig. 4). The G1 of E. prolatus s. str. is not known.
The dorsal surface of the carapace of Chu- lathelphusa neisi is not only distinctly less convex than all other Somanniathelphusa pax, the distal male abdominal segments are not slender and there is no distinct constric- tion on segment 6 (length to maximum width
66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ratio 1.0; distal to proximal width ratio 1.1) (cf. Rathbun 1905:249, fig. 61, pl. XI fig. 5). The first author has examined the G1 of C. neisi, and it is straighter, with the distal parts not curved or hooked, and the terminal seg- ment distinctly demarcated compared to S. Dax
The type specimens of S. pax were ob- tained by the second author from a market in Hanoi which had almost certainly come from rice-fields near the city. The first au- thor subsequently received additional spec- imens (non-types) from Hanoi and the spe- cies 1s apparently frequently sold in mar- kets. The very low prices and abundance of the crabs strongly suggest that they came from the neighborhood of Hanoi. The col- lection of rice-field crabs for food is a com- mon practice in Indo-China. The first au- thor has observed this often in Thailand, and in every instance, the crabs had been collected from slow-flowing streams or rice- fields on the outskirts of the town and sold as a cheap source of protein. In Thailand, crabs are kept dried (the individuals often caked in mud) and can be kept alive for long periods provided they are kept cool (see Ng & Naiyanetr 1993:43). They are eaten in a variety of ways, but are usually fried with spices until they are crispy.
The study of the freshwater crab fauna of Vietnam is still very much in the explora- tion and survey stage. The number of taxa known is likely to increase substantially in the coming years as the country becomes more open to the scientific community.
Etymology.—The species name is de- rived from the Latin for peace. Used as a noun in apposition.
Acknowledgments
The first author is grateful to Anna Ursula Kara for passing the Vietnamese crabs to him for study and Phaibul Naiyanetr for information on FE. prolatus. The study has been partially supported by a research grant (RP 900360) to the first author from the National University of Singapore.
Literature Cited
Alcock, A. 1910. Brachyura I. Fasc. II. The Indian Freshwater Crabs— Potamonidae. Catalogue of the Indian Decapod Crustacea in the collection of the Indian Museum, Calcutta, pp. 1-135, Pls. 1-14.
Balss, H. 1914. Potamonidenstudien.—Zoologisch Jahrbucher (Systematics) 37:401-—10, pl. 15.
Bott, R. 1968. Parathelphusiden aus Hinterindien
(Crustacea, Decapoda, Parathelphusidae). —
Senckenbergiana Biologica, Frankfurt 49(5):403—
422.
1970. Die Siisswasserkrabben von Europa,
Asien, Australien und ihre Stammesgeschichte.
Eine Revision der Potamoidea und Parathel-
phusoidea (Crustacea, Decapoda).—Abhand-
lungen der Senckenbergischen Naturforschen-
den Gesellschaft, Frankfurt 526:1-338, pls. 1-
58.
Dai, A.-Y., Y.-Z. Song, L.-Y. He, W.-J. Cao, Z.-B. Xu, & W.-L. Zhong. 1975. Description of several new species of freshwater crabs belonging to the intermediate hosts of lung flukes.—Acta Zoo- logica Sinica, Beijing 21(3):257—264, pls. 1-3.
Man, J. G., De. 1904. Décapodes d’eau douce rap- portes par M. A. Pavie, avec 2 pls. Mission Pa- vie, Indo-Chine 1879-1895. Etudes diverses. III.— Recherches sur |’Histoire Naturelle, Paris 1904:316-331, 2 pls.
Milne-Edwards, A. 1869. Revision du genre Thelphuse et description de quelques especés nouvelles faisant partie de la collection du Muséum.—Nouvelles Archives du Muséum d’Histoire Naturelle Paris 5:161-191, pls. 8-11.
Milne-Edwards, H. 1853. Mémoire sur la famille des Ocypodiens.—Annales du Sciences naturelle, Zoologie (3)20:163-228, pls. 6-11.
Naiyanetr, P. 1994. On three new genera of Thai ricefield crabs allied to Somanniathelphusa Bott, 1968 (Crustacea: Decapoda: Brachyura: Para- thelphusidae).— Raffles Bulletin of Zoology, Singapore 42(3):695-700.
Ng, P. K. L. 1988. The Freshwater Crabs of Penin-
sular Malaysia and Singapore. Department of
Zoology, National University of Singapore,
Shinglee Press, Singapore, pp. i—viii, 1-156, 4
color pls.
, & D. Dudgeon. 1992. The Potamidae and
Parathelphusidae (Crustacea: Decapoda:
Brachyura) of Hong Kong.—Invertebrate Tax-
onomy, Melbourne 6:741-768. .
Ng, P. K. L., & P. Naiyanetr. 1993. New and recently described freshwater crabs (Crustacea: Decap- oda: Brachyura: Potamidae, Gecarcinucidae and Parathelphusidae) from Thailand.—Zoolo- gische Verhandelingen, Leiden 284:1-117, figs. 1-68.
VOLUME 108, NUMBER 1
Rathbun, M. J. 1902. Description des nouvelles es- péces de Parathelphusa appartenant au Muséum de Paris. — Bulletin du Muséum national d’His- toire naturelle, Paris 1902(3):184—187.
1904. Les crabes d’eau douce.— Nouvelles Archives du Muséum d’Histoire naturelle, Paris (4)6:225-312, pls. 9-18.
1905. Les crabes d’eau douce.— Nouvelles Archives du Muséum d’Histoire naturelle, Paris (4)7:159-323, pls. 13-22.
67
Turkay, M., & P. Naiyanetr. 1987. The identity of Potamon rangoonense Rathbun 1904 and Thel- phusa larnaudii A. Milne-Edwards 1869, with introduction of Neolarnaudia botti n. g. n. sp. (Crustacea: Decapoda: Potamidae).—Sencken- bergiana biologica, Frankfurt 67(4/6):389-396.
Wu, H. W. 1935. Enumeration of the river-crabs (Potamonidae) of China with descriptions of three new species. —Sinensia 4(11):338-352.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
108(1):68-75. 1995.
A new species of Goreopagurus McLaughlin (Decapoda: Anomura: Paguridae) from the Pacific, and a comparison with its Atlantic counterpart
Patsy A. McLaughlin and Janet Haig
(PMcL) Shannon Point