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PROCEEDINGS

of the

Biological Society of Washington

VOLUME 95 1982

Vol. 95(1) published 13 April 1982 Vol. 95(2) published 11 August 1982 Vol. 95(3) published 5 October 1982 Vol. 95(4) published 20 December 1982

WASHINGTON PRINTED FOR THE SOCIETY

EDITOR

BRIAN KENSLEY

ASSOCIATE EDITORS Classical Languages Invertebrates GEORGE C. STEYSKAL THOMAS E. BOWMAN Plants Vertebrates DAVID B. LELLINGER RICHARD BANKS Insects

ROBERT D. GORDON

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 1981-1982

OFFICERS President RAYMOND B. MANNING

Vice President PAUL J. SPANGLER

Secretary MICHAEL A. BOGAN

Treasurer LESLIE W. KNAPP

COUNCIL Elected Members FREDERICK M. BAYER KRISTIAN FAUCHALD ISABEL C. CANET DAVID L. PAWSON

AUSTIN B. WILLIAMS

TABLE OF CONTENTS

Volume 95

Adis, Joachim, and Richard C. Froeschner. Notes on distribution of some Latin Amer- ican cotton-stainers (Dysdercus: Pyrrhocoridae: Hemiptera) and remarks on the biol- ogy of Dysdercustunbahnt- Schmidt. .2 ee ee ree ep ase pee e

Adkison, Daniel L. Description of Dactylokepon sulcipes n. sp. (Crustacea: Isopoda: Bopyridae)randsnotesiOmel) Cari Ge See cee era ee ee ae eee ee

Adkison, Daniel L., Richard W. Heard, and Guy T. Clark. Description of the male and notes on the female Argeiopsis inhacae (Crustacea: Isopoda: Bopyridae) ______________

Baker, H.R. A note on the genitalia of Potamothrix hammoniensis (Oligochaeta: Tubi- TGUAAE) i se a ee ec a

Barnard, J. Laurens, and Margaret M. Drummond. Redescription of Exoediceros fossor (Stimpson, 1856) an Australian marine fossorial amphipod, the type-genus of the new family EP xoedicerotidac \.. 2-5 2 Be ee

Barnard, J. Laurens, and Gordon S. Karaman. Classificatory revisions in gammaridean Amphipoda (Crustacea) 3 Pamt.2.¢ 202° 2k NS ae ie Mine ee eee dr se ee

Bayer, Frederick M. Some new and old species of the primnoid genus Callogorgia Gray, with a revalidation of the related genus Fanellia Gray (Coelenterata: An- tHOZOB) ee a wee eT Ne en Oe

Bowman, Thomas E., and Richard Franz. Anopsilana crenata, a new troglobitic ciro- lanid isopod from Grand Cayman Island, Caribbean Sea ______________________________-_-

Brooks, Daniel R., and Janine N. Caira. Atrophecaecum lobacetabulare, n. sp. (Di- genea: Cryptogonimidae: Acanthostominae) with discussion of the generic status of Paracanthostomum Fischthal and Kuntz, 1965, and Ateuchocephala Coil and Kuntz, | | en eee Ome eee Me Cen ee ee

Brown, Walter C., and Angel C. Alcala. A new cave Platymantis (Amphibia: Ranidae) from, the Philippine Islands)... 0 a eo

Bruce, Niel L., and Thomas E. Bowman. The status of Cirolana parva Hansen, 1890 (Crustacea: Isopoda: Cirolanidae) with notes on its distribution __________________________

Chernoff, Barry, and Robert Rush Miller. Mexican freshwater silversides (Pisces: Ath- erinidae) of the genus Archomenidia, with the description of a new species ____________

Child, C. Allan. Pycnogonida of the western Pacific islands I. The Marshall | E10 | ea ieee ee ee coe eR OAD Ay ge om cce te) Sat ee

Child, C. Allan, and Koichiro Nakamura. A gynandromorph of the Japanese pycnog- onid=Anoplodaciylus ge'stien's | (@rctriverramn)) seme eee eae ea ee een

Collette, Bruce B. South American freshwater needlefishes of the genus Potamorrha- phis (Beloniformes: Belomidac)y eae: eee ee ee ee See ee

Cressey, Roger F. A new genus of bomolochid copepods from Indo-West Pacific nem- UPterid fishes = Sebo os os EN ir ce an fk, Sole sks lee! lee

Crews, Celinda R., and Artie L. Metcalf. A new species of oreohelicid land snail from the: Sam “Agus Gime plains INS wi VIS xc sae a i ne

Cutler, Edward B., and Norma J. Cutler. A revision of the genus Siphonosoma (Stpuncula): 225 20.24 pack te ew SN pen ger angel een lr ie

Dawson, C. E. Review of the genus Micrognathus Duncker (Pisces: Syngnathidae), with descriptionrof iM. satanis me Spee aasake nee ae setae ee nse an eee te) eee

Dawson, C. E., and C. J. M. Glover. HAypselognathus horridus, a new species of pipefish (Syngnathidae) from South Australia _____.-________-_------ Sessa

Desbruyéres, Daniel, and Lucien Laubier. Paralvinella grasslei, new genus, new species of Alvinellinae (Polychaeta: Ampharetidae) from the Galapagos Rift geothermal VMS) ce ue Ee rai i a RT Soe ent ae oie gas ail

Downey, Maureen E. Evoplosoma virgo, a new goniasterid starfish (Echinodermata: Asteroidea) from the:Gulf of Mexico. 2 2s eee

Ewing, R. Michael. A partial revision of the genus Notomastus (Polychaeta: Capitel- lidae) with a description of a new species from the Gulf of Mexico ____-_________________

iv

371-376 702-708 334-337

563-566

610-620

167-187

116-160

522-529

223-231 386-391 325-333 428-439 270-28 1 292-296 714-747 495-504 256-264 748-762 657-687

403-407

484494 772-173

232-237

Fauchald, Kristian. Two new species of Onuphis (Onuphidae: Polychaeta) from Uru- ON hy NE NS Ned Nin Sp ates NY ed Bere A an Pk Ne as Na AG ooh ed | Se jt ard Fauchald, Kristian. Some species of Onuphis (Polychaeta: Onuphidae) from the Atlan- (WE QUESBIN » wae creitich ME tae 2 halen iia Beek SE Teal, ie epee N Nc, bs Bigs ABNER lige se Se Ree 1 ae WE Reea aN er a Fauchald, Kristian. A eunicid polychaete from a white smoker __________________-__________ Fauchald, Kristian. Description of Mooreonuphis jonesi, a new species of onuphid poly- chaete from shallow water in Bermuda, with comments on variability and population EC iy me ne EL eked ML EAN yl seweN Tee Ny I AEE 2 vera Enthe Usb net S Flint, Oliver S., Jr., and Joaquin Bueno-Soria. Studies on Neotropical caddisflies, XXXII: the immature stages of Macronema variipenne Flint & Bueno, with the division of Macronema by the resurrection of Macrostemum (Trichoptera: Hydropsychi- TEC) EL he apa) Su sl eve eae pe Nc babe Ll eee Boo dele Formas, J. R., and Alberto Veloso. Taxonomy of Bufo venustus Philippi, 1899 (Anura: Peomiodacmidae)mnom, central Chile... 2 22... 3.50 se Formas, J. R., and M. Ines Vera. The status of two Chilean frogs of the genus Eup- NepuemeAnunacmeeplodactylidac) .-.. 8.2. 22.2 ee ee a BL Fredette, Thomas J. Evidence of ontogenetic setal changes in Heteromastus filiformis duolehactan@apivellidae)irummr. Sa Soe meal aiel! yale 6 ok eb wl eat ol Garcia, Renato G., and Raymond B. Manning. Four new species of stomatopod crus- Tee SmnROMMecOe EOP pines 2.26 eee ee ee ee ees Garrido, Orlando H., and Albert Schwartz. A new species of Sphaerodactylus (Reptilia: SanasGekkonidae) from eastern Cuba 222) 2 George, Robert Y., and Noel A. Hinton. A new species of deep-sea isopod, Storthyn- gura myriamae, from the Walvis Ridge off South Africa __________-____-_--_--- Gleye, LindaG. Two new species of leptomysinid mysids (Crustacea: Mysidacea) from SOURS (CANINE: «SUIT RN sh oe eM eke tags BR et eemmeeane y cNeeS ORs bieaeeae aL seem ee Green, Karen D. Uncispionidae, a new polychaete family (Annelida) ____________________ Harasewych, M. G. Pterynotus xenos, a new species of muricid from off northern Teamraneancwionnscas Gastropoda), 22.2209 se2e8 8 ee ee Harding, Keith A. Courtship display in a Bornean frog _____________________-______-----_-_- Heaney, Lawrence R., and Gary S. Morgan. A new species of gymnure, Podogymnura (Mammalia: Erinaceidae) from Dinagat Island, Philippines ____________-____--__-__-- Hershkovitz, Philip. Subspecies and geographic distribution of Black-mantle Tamarins NOCnmUmsS Mericolis Spix (Primates: Callitrichidae) ___..._...-.-..2- 2-222. ee Heyer, W. Ronald. Two new species of the frog genus Hylodes from Caparao, Minas Gemiss braziy (Amphibia: Weptodactylidae) 2220..00 82.2). e ee ee Heyer, W. Ronald, Charles H. Daugherty, and Linda R. Maxson. Systematic resolution of the genera of the Crinia complex (Amphibia: Anura: Myobatrachidae) ______________ Hobbs, Horton H., Jr., and Daniel J. Peters. The entocytherid ostracod fauna of north- Saar 21 hl es SUES elie bin). 9a he be cy) pe do =O ht hae od DY ole | Hobbs, Horton H., Jr., and Henry W. Robison. A new crayfish of the genus Procam- Por mmOonmEsolihnwestern Arkansas =.f.2u28.5552).. eee tel gee ee Holt, Perry C. A new species of the genus Cambarincola (Clitellata: Branchiobdellidae) from Illinois with remarks on the bursa of Cambarincola vitreus Ellis, 1919, and the MRMISMONE NT OOKIMS AOI, L9G8. 0 ee Huddleston, Richard W. Comments on the nomenclatural status of the families Cau- cascliade and Favusellidae (Foraminiferida) .._...... = Huddleston, Richard W., and Drew Haman. Nomenclatural status of the foraminiferal MAMIE OOCITCIG SAIGON A 1980 Huddleston, Richard W., and Drew Haman. Jascottella, nom. nov. for Mamilla Scott, 1974 (Microproblematica) non Fabricius, 1823 (Mollusca) ____________________----- Hutchings, P. A., and C. J. Glasby. Two new species of Ceratonereis (Polychaeta: Nereididae) from estuarine areas of New South Wales, Australia ___-_ Kenk, Roman. Freshwater triclads (Turbellaria) of North America. XIII. Phagocata RUMDLONAG Me Was pe cles, from sNeVaday, werd sie eee A Kenk, Roman, and Anne M. Hampton. Freshwater triclads (Turbellaria) of North America. XIV. Polycelis monticola, new species, from the Sierra Nevada range in Pe EMOTE gee Selly ts ee eater yy ipl rl EEN RI coer) sarcu f wl ey inyrteesl bOI. ee oil

203-209 238-250 781-787

807-825

358-370

688-693

594-601

194-197

537-544

392-397

93-98

319-324 530-536

639-641 621-624

13-26

647-656

377-385

423-427

297-318

545-553

251-255

637-638

114-115

42]

515-521

161-166

567-570

Kensley, Brian, and Gary C. B. Poore. Anthurids from the Houtman Abrolhos Islands, Western Australia (Crustacea: Isopoda: Anthuridae)__.----- = Kornicker, Louis S. Alternochelata lizardensis, a new species of myodocopine ostra- code from the Great Barrier Reef of Australia (Rutidermatidae) __._- Kudenov, Jerry D. Redescription of the major spines of Polydora ligni Webster (Polychaeta::Spionidae) 2-2 8tosat Vines wi ea ee ee Rn geeeaee le Od 2 eee Kudenov, Jerry D., and John H. Dorsey. Astreptosyllis acrassiseta, a new genus and species of the subfamily Eusyllinae (Polychaeta: Syllidae) from Australia _____________ Kyte, Michael A. Ophiacantha abyssa, new species, and Ophiophthalmus displasia (Clark), a suggested new combination in the ophiuroid family Ophiacanthidae (Echi- nodermata: Ophiuroidea) fromoff Oregon, (USA. 2 ee eee eee Lewis, Julian J. A diagnosis of the Hobbsi group, with descriptions of Caecidotea teresae, n. sp., and C. macropropoda Chase and Blair (Crustacea: Isopoda: Aselli- aS) ona a a nN CI 2 A Sag SS Ao Louton, Jerry A. A new species of Ophiogomphus (Insecta: Odonata: Gomphidae) from thebwestern Highland Rit i Weminesse ec) See ae Lynch, John D., and Pedro M. Ruiz-Carranza. A new genus and species of poison-dart frog (Amphibia: Dendrobatidae) from the Andes of northern Colombia __________________ Manning, Raymond B., and Ch. Lewinsohn. Rissoides, a new genus of stomatopod crustacean from the east Atlantic and South Atnica |= 22 ee eee Manning, Raymond B., and Marjorie L. Reaka. Gonodactylus insularis, a new sto- matopod crustacean from Enewetak Atoll, Pacific Ocean ___________________________-___- Marshall, Harold G. Phytoplankton distribution along the eastern coast of the USA. IV. Shelf waters between Cape Lookout, North Carolina, and Cape Canaveral, Fikrig ee ee Rete eS aan er ae Ne ne Martin, Joel W., and Lawrence G. Abele. Naushonia panamensis, new species (De- capoda: Thalassinidea: Laomediidae) from the Pacific coast of Panama, with notes on the Semis wih.24 2 es Tee ee See i ESE SORA (Ta aae Un he ee McEachran, John D., and Janice D. Fechhelm. A new species of skate from Western Australia with comments on the status of Pavoraja Whitley, 1939 (Chondrichthyes: RaAjifOrmes 2 ttrts Sem Sn eee je 2 re et se RE es ae nD McEachran, John D., and Janice D. Fechhelm. A new species of skate from the western Indian Ocean, with comments on the status of Raja (Okamejei) (Elasmobranchii: Rajiformes)i 2.2 20. es a od A Mea Ane oe a ees McKaye, Kenneth R., and Catherine MacKenzie. Cyrtocara liemi, a previously un- described paedophagous cichlid fish (Teleostei: Cichlidae) from Lake Malawi, Atnica: 25 iieasee van! er he erie) ae iy Caer Be ee a

McKenzie, K.G. Description of a new cypridopsine genus (Crustacea: Ostracoda) from .

Campbell Island, with a key to the Cypridopsinae __________________________-__-__- Mendez, G. Matilde, and Mary K. Wicksten. Notalpheus imarpe: a new genus and species of snapping shrimp from western South America (Decapoda: Alpheidae) ______ Murdy, Edward O., and John D. McEachran. I[stigobius hoesi, a new gobiid fish from Australia (Perciformes !Gobudae)i aie. 1 nn ee ee ee Gas eee ee Nakamura, Izumi. Lateral line of Diplospinus multistriatus (Teleostei: Gempyli- GAS) ORL. oie PE oh OE oP Oe IM We oie cel Sais, RI a A I Nakamura, Koichiro, and C. Allan Child. Three new species of Pycnogonida from Sa- gami'Bay / Japa. oie bal a Opell, Brent D. A new Uloborus Latreille species from Argentina (Arachnida: Araneae: MWPOD ORAS Ry haa OU eo ee cp Parenti, Lynne R. Relationships of the African killifish genus Foerschichthys (Teleostei: €yprinodontitormes:/Aplocherliad ac), Mase a ae eee, Petuch, Edward J. Paraprovincialism: remnants of paleoprovincial boundaries in Recent marie molluscan' provinces! ._c.ONiiieh Me i ie eee ae Se eee eee Rausch, V. R., and R. L. Rausch. The karyotype of the Eurasian flying squirrel, Prero- mys volans (L.), with a consideration of karyotypic and other distinctions in Glaucomys spp? (Rodentia® SGnikidae) rl v* seer ee OL: AE A ee ee Rohr, David M., and Richard W. Huddleston. Yochelsoniella, nom. nov., a new name for Ellisella Rohr, 1980 (Gastropoda) non Gray, 1858 (Coelenterata) _____-_--___________

Vl

625-636

793-806

571-574

575-578

505-508

338-346

198-202

557-562

352-353

347-351

99-113

478-483

440-450

398-402

766-771

709-713

642-646

408-411

282-291

554-556

451-457

774-780

58-66

269

Rosewater, Joseph. A new species of the genus Echininus (Mollusca: Littorinidae: Pehinmumae) with a review of the subtamily ==> 2 i Schultz, George A. Amerigoniscus malheurensis, new species, from a cave in western @reconm(@nustacea: Isopoda: Trichomiscidae)).. Bs... eee Shelly, Roland M. A new xystodesmid milliped genus and three new species from the eastern Blue Ridge Mountains of North Carolina (Polydesmida) _________________________- Stauffer, Jay R., Jr., Brooks M. Burr, Charles H. Hocutt, and Robert E. Jen- kins. Checklist of the fishes of the central and northern Applachian Mountains ______ Sterrer, Wolfgang, and Thomas M. Iliffe. Mesonerilla prospera, a new archiannelid PROMI AnIMeNeAIVeStIN BEfMmmdar i096 ee Thomas, Richard. A new dwarf Sphaerodactylus from Haiti (Lacertilia: Gekkoni- Ty) eT NNR Ic ce 2 heh Td Oe eS ee ee es oe ied Thompson, Fred G. A new species of Euglandina from Peru (Gastropoda: Pulmonata: SHMUEL SCORE) a yl an al eR epee 0 ee Thompson, Fred G., and Jane E. Deisler. A new tree snail, genus Drymaeus (Buli- MUNA nOmMESOutMeastemm PTW 28 Uebelacker, Joan M. Review of some little known species of syllids (Annelida: Poly- chaeta) described from the Gulf of Mexico and Caribbean by Hermann Augener in I eran eaten erie a eter fect ee RT ed Po es te ee Vari, Richard P. Hemiodopsis ocellata, anew hemiodontid characoid fish (Pisces: Char- AcCHMeMetLoOmiwestern OUNIMAM oo se Vari, Richard P. Curimatopsis myersi, a new curimatid characiform fish (Pisces: Char- aciformes) from Paraguay scale), eM tars nee Gee Aemtee ah eee ROBO ON ee Vecchione, Michael. Morphology and development of planktonic Lolliguncula brevis eb a mClarm Vin OPSIGa)) a. 2 et ee ee ee -Wainright, Sam C., and Thomas H. Perkins. Gymnodorvillea floridana, a new genus and species of Dorvilleidae (Polychaeta) from southeastern Florida _____-___---__________ Wicksten, Mary K. New records of pinnotherid crabs from the Gulf of California REM OuicmeIIMOUNSHIGAG) <2 82. 6 St eS ee Wicksten, Mary K. Pinnixa costaricana, a new species of crab from Central America REM nay em EenTIMOUNCTIGAC) =. ot etl eae Ne on ye ee ee Zibrowius, Helmut, and Stephen D. Cairns. Remarks on the stylasterine fauna of the West Indies, with a description of Stylaster antillarum, a new species from the Lesser Paaiiilesnenmadania: Eydrozoa: Stylasterima) 20... 0 Zottoli, Robert. Two new genera of deep-sea polychaete worms of the family Amphar- etidae and the role of one species in deep-sea ecosystems ___________________----- Zusi, Richard L., and Gregory Dean Bentz. Variation in a muscle in hummingbirds and MiSEAMGniseSyStematic implications 2.2222... 21.2222 eee een es

vil

458-477

27-47

509-514

81-88

763-765

265-268

583-593

188-193

788-792

602-609

694-701

354-357

579-582

210-221

48-57

412-420

INDEX TO NEW TAXA

VOLUME 95

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

COELENTERATA Hydrozoa

Stvlaster antill arin sist eo aes Oe eR ee Mee

Callegorpia chariessa@..25 02's * i se ea ee Oe Se a tal Fanelliaxcoryimbosa@ 20022 es oe ee

PLATYHELMINTHES Trematoda

Atrophecaecum lobacetabulare wien 2 eat ee 2 eee

Phagocatashamiptonae 20235400) oe, ue a ies oe ee Polycelis, montiCola; 20x 20s. ey. sed ope Dee ga esa pi SS el eel ee ea

ANNELIDA Archiannelida

Mesonenilla prosper 25 aa SIE ae a Cambarincola illimoisensis 122502 es ee ee ie ne

AiSORE DOSS se SE a ae Tae teal CV ASSUSCLG [2.0 WNT. DRS DO Se I a Ss Ss ee Ore Ne

CELE VOWS ee ie hi este Dp Pl pe aE a ea Decemunciger’ ia. \ S002 aie be EO SS Se las Oe eee ne ne GPQNEGW Ec Fly WN RT Re CSR Re ahs a i cele A te ga real Brdecamera. 02:02. Rae. NU Ce oc ON eg) EAR Re 2 A eee Paleg 'x.weooy ee TY AYE SE: SAN ae Be 1 ad Eunice: pulvinopalpata. 2040) 22 PS 2 he RR on he ES Ee Se reece a GyMVWOU OPV CO a LEO ES AA ce ets) cea lied Jo eee oe len eg flonidang) Ga... LSA) EOE SUA of EE AR ea So a ee Haplosyilis floridamat.C.. fe eg ce Mooreonuphisyonesi 2000. 8 Re ee SIDE My SN Sept eee nd gd NG@tOmaStuls (Cai eHi oes os ise ae I a a a

Onuphiss(Nothnia)*australatlantica aa ee eee ee

heterodentata. 22. es a ae Ee

lithobiformis ¢ 2 to 2 0es) DS aie ie eM aaa Rae Ae ae

(Onuphis) declivorum 3204 ee ee ee ee eee

LiffiCilis sc eT) Ce a EINE ULES Tio ial te en aren LO ee ee

OV CWS QUIUZE 9 Sa he ee a we Pee | el i cis eh ee a

D011 ce Ee A TEA COW TM NL at MeN Me Meee NG aS Se A

Pavalvinehl@:, eo 0 fe ks i See le 5 TE ae (Te Pe en na ee GU GIST A. SRN ata Nera Iie A led Pc I sg aD UNCISPIONIDAE =) Mik ceck) Oe EO Ree ce Ies CR aa ee A A ee eS See

TWEE SIDUD ence telco eee al I IR SOP eA a) SE ae ce RRS DEL aes a SO 534

[AO TTGUUOID gee le TE lat a Tee TES a7 i ATI aR oO ot ne MY SRR SE ES 534 ARTHROPODA Crustacea

Meet Onen Se ARCMICCL CLETUS IS pee wate sche aos ver ne OW et eS ke a ee 793 PRIME TO OMS CUISMOULCHNCNSIS) waste ot Sie Bere Oy SN ee 89 ELSIE, CPA TACNT LeNS aaa ie ce t d SE SAAAn e niece Sy?) SCVMM LOCUPTO pe SO a PT me lia al Re SS et PSP en RO pe een eo eM 625 PAW ae) TCAEET CA) (0 ene TE Ge) on) Vu hee ee a Ne NU A oe iy ee Ne 171 ALAR D CDV TIOID SES sg ge a 8 SIE RI Re mR gi Se, a a a el 766

ROTA ce oe seeps Se pa OT cle ae A ek LT, Aveta oe ee OPE 768 BAA CLOTS IACI Camm mee a tic Re Ng ee I Ss as ES pe ge i Lo Oe RIL | Pv ge 339 WORE OL OLACMSUS 52 ee eee aig tne Renee nen re, Re ee PL se nee 538 Peer cnter MSIE SCI OSAMRI cae os Ne Ee A a a 319 OE GIEE DM CHCYB OS aks = Ae EEN TS bal SE i Ae lepetet Ear DNIeN ernee iee Eee 703 ESR HVE CS SMO CUACAD 1G Comer nme eee rnin 2 eet er Oy ie Ol a AAA OL ee pe nt Pode tsar 628 EO AID TION MCOMIID) AU ee ae ee I I Ee 610 LEENA TLIO! is ET are Pe 8 a le Se ne ee 182 SSAA CICS SLL SS A NS er ws Ne 08 Sb ea np ea Wee 347 PREM OSOMMAB IIL DUNEMSIS: Bi. 2 Vl a) 20 yet bie a 547 eer (ares) Fr eRe Eh ae) ee ahr ra) uh hele LO leet i i ea A ea as 495

NOUDISCUIS epee Sie 2 SRO NS ES Oe a oe eee eee ee ete Oe RE mS I nee 495

rea | | ee onl weit Ne pee SP MNES Ae BE ee a eS So Eas 502 EEE OU SADE METS, cena an A Bg AR SE PD 180 AWHONN ODE a eecsecien ae eS ep a YS PRT ctl le UDO ened Ad YS pk 182 erent RULER anh CLC EE een aS, ee PN NTE AD foes i age a ee ee 540 CLRPURGOVEOYET I Ba a ay Ne eer Deh BA Oe a REE Le ee pec ee aU 174

eh Coenen eR eS Ya eh Of Re uees Wl elie Wires tS eh eee ae Se 174 Re et) 7 nC Ree en mevmetn ns we ee My Sha iE re eS ete 176 MARLON OMIR ERI CHIOC) GC aemmne elmer ct Loe! eee ee 8g os se totes Sh NS ee 321 IE ERE VE ee tik Bs A sl a a Ee a ia en VP 170 fla MuSlivCUMD: (PUPUAILTAUCIASIIS cick Se NAA Sy BS Teg ar De pee NN te eal oe em ger ee ee ee 478 POSES CLERIG ASPROWIDE tec, sl tr ae Se ese ug esc ete ce ar aA OR cee IL a oe hE 709 ane easel eile Cah O)/) Spee ween me ee ye LE Me RE pels et ies eS ee 542 Sp SM SEGINTERE EPICUGW OND: 2 I TN I te ee eS 634 AUR COAL IESE MIT YAU S10 ne eT ln a ee Ee Sa 181 Me TINO CMC MSE MIM: Coree sep ew Ac See tt oe he ee ON ee Ne 170 MIRIAM TOR ST GAIT Lm ee ee el elles bam WE Nita be! OE ee le oe 579 EN eAMoanisn(Gimandiella) parasimulans 0 545 SABIE DERDCDEL AED, a dapat Sh Spee le eR D0 SS nce ae 168 ESN CHEGUEIS aan ancn cnet og ck lt A ce Pe ed, ERIN ete 352 ae ar Rte Tem Re A ei ee Vine on ie oa kN ee RT 169 MRC BGIER TCIM ANS LCLIIT LC eames Re Deon a eg ED oe fo a ei oe 93 IVI oo I ce a) a A ee pe I GE De es 176 fees BCU ELOISE ls ys tr cn ee eee eter one ERA 7) ee oc | ee eee Je RT Pgs ona bs ho es es le 184

Insecta ARTO MMOMUISH DOUGH CRG) (seeee ents! (2) ee ess ee ee ele oe a ah Pe 198 Myriapoda

EW OMGILI. pada SE aa ee a Rg SE AR a, 460

(AAA OTIS Seve Soe LA £9 ST NR SSE a we OIRO he SEN le BP 2) pe TN ONE AE Wes Cen ene hs 467

COMET OUUS! es a oS re 3 AOS IIT © PSE ep Neg ar Re Oe Os 463

fil RADIUS a 5 ps a NOT LUD LW NT NON nd Pc 0s yo ae eR ee ap See eee 470

Pycnogonida

Ammiothella:stawroniata 2 2.5 ce A Oe eae, 2 a ee eR ae eae kT RS, ee ae Dg Anoplodactylus wiarsiiall crisis’): 225. els sacs ale EG ues Usps etalon a er Oe etek 274 perforatus 200) so Bet aE Ei ane ne rs ee 289 SRUIODGEMSIS( co 5) AN a8 SO ASS TO SNe ep eg LA dR leg oe 285 ACS COVEY ICIS Lette OUIaEE $y 00 i ek WN AE ala a eG ig ol oh oe 283 Nymphon microne sicuni cee 6. 08 2a eer ets Tie ae ES URE ne ns ree ey 27 Arachnida Uloborus elongatus [Leite oes lye ti wena ea a I | a ete me ie 554 MOLLUSCA DrymMacus aurantiOstomus: 28 Bs ent Es PEN Ae ee eta Ces 265 Echinanus vivipariis 2.002) 20 ec) A ie SSN ee ee kee ee 69 Euplandinay easy | occ es ee SS a re 763 Oreohelix ‘litoralis 20.220 et ce a hE rte is Mee ars ye 256 Pterynotus (Pterynotus) xenwos” loc eat Le a 639 VOCHE] Sore dha 0500 ae ah oR eR IT Ha Lor Te Wate tt reper ea 269 ECHINODERMATA Evoplosomia: virgo ‘stot suet tec 6th SER is eee ea 0 Re Pole a en eo Te, Ophiacanthavaby sear 42 1k tk ee ee ee ee ee 505 Ophiophthalmus diplasia ac oe er ee ee ee 508 CHORDATA Pisces Archomentdiatwvarvel@e: 25! oe ag NT he Tre ea 430 GurimatopsisS miyerst Loo ee RE ee a 788 Gyrotocara dient < on. oe 398 Hemiodopsis ocellata (cs 2 tee a es 188 Hy pselognathus horridus | 2-22. jus 403 ISHPODIUS HOST 6 oe re es ce eae 643 Micro pnathuls: natans. 2068 el ae Mio ee ea te 682 Bay onayanallenigw qn. 2 2s 2 te ee NE So de eae 8 Rajan(Okamejel)eemstrat =... e 44] Amphibia AM OD OPTATVTUES aa il a NR IB RS > er 57) SVILOMMOPUS) chest a LN Ra ae A Ry le ec 0 ek Ai an a DSI) Piylodes baba@x tse hs ie 380 WOUTUZOLEIUTT Dees oo OA ue say 3 Tea 9 Onan aed Lt eg IT ee 382 Platyamantisispelaeuis: 26 o6 0.0 2 ts os, oi Ue A age I eS 386 Reptilia Sphaerodactylusseelicana.. oto 2 ss Se 392 VILO CV UO ep I alo Se we RN pS SIRI A SR 81 Mammalia Podogymmura: aureospinila@ a5 oo I Ta A a Sn ae ca oe 14 Sasuinus mignicollis*hernandezi. 25.2) a ne 649 MICROPROBLEMATICA TGS COPLSN 05 5h MI he Bah gO el A ie a Sc aR 421

S74, 0618

(ISSN 0006-324X)

Proceedings of the BIOLOGICAL SOCIETY of WASHINGTON

F QAO NIAA Libh AM Kt

Volume 95 13 April 1982 Number 1 ;

THE BIOLOGICAL SOCIETY OF WASHINGTON

1981-1982 Officers

President: Raymond B. Manning Secretary: Michael A. Bogan Vice President: Paul J. Spangler Treasurer: Leslie W. Knapp

Elected Council

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PROCEEDINGS Editor: Brian Kensley Co-editor: Stephen D. Cairns

Associate Editors

Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Richard Banks

Insects: Robert D. Gordon

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PROC. BIOL. SOC. WASH. 95(1), 1982, pp. 1-12

A NEW SPECIES OF SKATE FROM WESTERN AUSTRALIA WITH COMMENTS ON THE STATUS OF PAVORAJA WHITLEY, 1939

(CHONDRICHTHYES: RAJIFORMES)

John D. McEachran and Janice D. Fechhelm

Abstract.—The genus Pavoraja is resurrected for the Australian skates Raja nitida and Pavoraja alleni, n. sp., which differ from all recognized genera of skates in clasper and rostral structure, and by their combination of derived char- acter states. Pavoraja alleni is described and diagnostic characters are given for Pavoraja and P. nitida.

Raja nitida, which occurs off Tasmania, Victoria, and southern New South Wales (Whitley 1940), was placed in a new genus Pavoraja, along with another _ eastern Australian skate, Raja polyommata Ogilby, 1916 by Whitley (1939). Whit- ley (1939) only briefly described Pavoraja and gave no characteristics that dis- tinguish it from other skate genera. The following year Whitley (1940) offered little to augment his description, but stated that P. nitida resembles the South American skate, Malacorhina scobina (=Psammobatis scobina Philippi, 1857), that the type locality of P. nitida may be South America rather than Australia, and that the Australian specimen of P. nitida may represent a new species or subspecies. More recent authors (Fowler 1941, Bigelow and Schroeder 1953) considered Pavoraja synonymous with the genus Raja, because of Whitley’s lack of a generic diagnosis.

During investigations of the skate fauna of Western Australia a new species was discovered which closely resembles Raja nitida Gunther, 1880. Herein we describe the taxonomically important anatomical characters of R. nitida, com- ment on the taxonomic status of Pavoraja, describe the new species, and then comment on the relationship of R. polyommata to the other species.

Materials and Methods

Specimens of Raja nitida, R. polyommata, and the new species were obtained from the Australian Museum, Sydney (AMS); British Museum (Natural History) (BMNH); Tasmanian Museum, Hobart (TMH); and Western Australian Museum, Perth (WAM). Several individuals of R. nitida and one of the three specimens of the undescribed species were dissected to reveal the structure of the claspers, neurocrania, and scapulocoracoids. Several additional specimens of R. nitida and all three specimens of the new species were radiographed to verify the anatomical observations based on dissections and to count vertebrae, pectoral radials, and pelvic radials. Methods for making measurements and counts are described by McEachran and Compagno (1979, 1982).

Z PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1Icm

sl cf

SI.

a b

Fig. 1. Lateral view of right clasper, partially expanded to show components: a, R. nitida, AMS 1B.5275; b, Pavoraja alleni, WAM P19118 (Holotype). cf—cleft, hy—hypopyle, pr—pseudorhipidion, rh—rhipidion, sl—slit, sp—spike, sr—spur, st—sentinel.

Results

Claspers.—Raja nitida has very slender, short claspers which are constricted rather than expanded at origin of glans (Fig. la); without dermal denticles or pseudosiphon; inner dorsal lobe with pseudorhipidion extending from level of hypopyle to about distal one-third of glans, continuing distally as a fold of integ- ument; slit located lateral to pseudorhipidion; spur well developed; cleft medial to spur; rhipidion well developed, running from level of hypopyle to distal one- third of glans, distal section rotated laterally and running over base of sentinel; sentinel rod-shaped and covered with integument, extending from level of slit to near tip of glans; spike ventral to sentinel, located within sentina, disc-shaped with a sharp, naked lateral margin; axial cartilage forming a slender tip (Fig. 2a, b, c); dorsal marginal little expanded distally, distal margin truncate, with an inner distal extension entering glans and forming pseudorhidion; ventral marginal with a evenly convex distal margin; dorsal terminal 1 and ventral terminal mem- branous, broadly joined on ventral aspect of glans, forming a sheath-like covering of glans; dorsal terminal 2 broad, fused to distal and distolateral surface of dorsal marginal; dorsal terminal 3 fused with dorsal terminal 2, small with a distally pointed and laterally curved extension forming spur; ventral terminal V-shaped, lacking a sharp lateral margin, lying on dorsal surface of accessory terminal 1; accessory terminal 1 Y-shaped, fused with distal surface of ventral marginal, S-shaped distal extension forming sentinel; accessory terminal 2 short, attached to accessory terminal 1, with a dorsoventrally flattened, disc-shaped extension forming spike.

VOLUME 95, NUMBER 1 3

Fig. 2a, b, c. Right clasper cartilages of R. nitida, AMS 1B5275: a, Lateral view, partially ex- panded with dorsal terminal and ventral terminal cartilages shown separately; b, Dorsal view; c, Ventral view; d, e, f, right clasper cartilages of P. alleni, WAM P19118 (Holotype), d, Lateral view, partially expanded with dorsal terminal and ventral terminal cartilages shown separately; e, Dorsal view; f, Ventral view. Atr,—accessory terminal |, atr.—accessory terminal 2, ax—axial, dmg—dorsal marginal, dtr,—dorsal terminal 1, dtr,—dorsal terminal 2, dtr,—dorsal terminal 3, vmg—ventral mar- ginal, vtr—ventral terminal.

Neurocranium.—The neurocranium of R. nitida has a short, slender rostral Shaft, fused to flattened rostral node and appendices at tip of snout (Fig. 3a), widely separated from rostral base and neurocranium; rostral appendices free of rostral shaft posteriorly and possessing two foramina; propterygia of pectoral girdle reaching rostral appendices; nasal capsules of moderate size, set at about a 40° angle to transverse axis of neurocranium; foramen for profundus nerve on leading edge of nasal capsule; anterior foramen for preorbital canal on anterior margin of kidney-shaped basal fenestra of nasal capsule; precerebral space nar- row, inner walls of nasal capsules moderately bulging into precerebral space; interorbital region narrow (Table 1); preorbital processes poorly developed, sep- arated from supraorbital crests by shallow notch; postorbital processes well de- veloped; anterior fontanelle tear-shaped; posterior fontanelle trapezoid-shaped;

+ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Neurocranium of R. nitida, AMS 1B.5275: a, Dorsal view; b, Lateral view; c, Posterior view; d, Ventral view. ac—anterior cerebral vein foramen, af—anterior fontanelle, antc—antorbital condyle, bf—basal fenestra, end—endolymphatic foramen, es—efferent spiracular artery foramen, hf—hyomandibular facet, ic—internal carotid artery foramen, into—intercerebral vein foramen, ja— jugal arch, ibX—lateralis branch of vagus nerve foramen, nc—nasal capsule, obf—otic branch of facial nerve foramen, of—orbital fissure, onc—orbitonasal canal, os—optic stalk, peri—perilymphatic foramen, pf—posterior fontanelle, poc—preorbital canal foramen, postp—postorbital process, prep— preorbital process, prof—profundus nerve foramen, ra—rostral appendix, rb—rostral base, rn—ros- tral node, rs—rostral shaft, I—optic nerve foramen, I1J—oculomotor nerve foramen, 1!V—trochlear nerve foramen, VIJ—hyomandibular branch of facial nerve foramen, [X—glossopharyngeal nerve foramen, X—vagus nerve foramen.

foramen for anterior cerebral vein posterior to line connecting foramina for preor- bital and orbitonasal canals (Fig. 3b); trochlear nerve foramen posterior and dor- sal to optic nerve foramen; oculomotor nerve foramen situated above optic stock; orbital fissure located on posterior aspect of orbit, anterior to foramen for hyo- mandibular branch of facial nerve and posterior to foramen for interorbital vein; efferent spiracular artery foramen on ventral rim of orbit; jugal arches moderately slender; vagus nerve foramen immediately ventral to foramen for lateralis branch of vagus nerve and medial to foramen for glossopharyngeal nerve (Fig. 3c); basal plate moderately narrow (Fig. 3d).

Scapulocoracoid.—The scapulocoracoids of R. nitida are moderately short and anteroposteriorly elongated (Table 2), without an anterior bridge (Fig. 5a); an- terior fenestra little expanded; postdorsal fenestra moderately expanded; meso- condyle not expanded; three postventral foramina, first greatly enlarged; neo- pterygial ridge between mesocondyle and metacondyle incomplete.

Comments.—Raja nitida differs from all other species of Raja in clasper and rostral structure and differs from all other genera of skates in its combination of derived character states, suggesting that the genus Pavoraja should be resur- rected for this species. ‘‘Raja’’ nitida differs from Raja in possessing claspers with membranous dorsal terminal 1 and ventral terminal cartilages which are broadly joined along the ventral aspect of the glans and disc-like accessory ter- minal 2 cartilages; a reduced and incomplete rostral shaft which is widely sepa-

VOLUME 95, NUMBER 1

Fig. 4. Neurocranium of P. alleni, WAM P19117 (Paratype) Posterior view; d, Ventral view. Abbreviations as in Fig. 3.

“CTS a.

: a, Dorsal view; b, Lateral view; c,

rated from the rostral base, and propterygia of pectoral fins which reach rostral appendices. Reduction of the rostral base precludes this species from being clas- sified in Raja (Ishiyama 1958, Stehmann 1970, Hulley 1972, McEachran and Com-

pagno 1982).

Table 1.—Neurocranial measurements of P. alleni and P. nitida expressed as percentage of na-

sobasal length.

Nasobasal length (mm) Cranial length

Rostral cartilage length Prefontanelle length Cranial width

Interorbital width

Rostral base

Anterior fontanelle length Anterior fontanelle width Posterior fontanelle length Posterior fontanelle width Rostral appendix length Rostral appendix width Rostral cleft length Cranial height

Width across otic capsules Least width of basal plate Greatest width of nasal capsule Internasal width

P. alleni 278 mm TL @

26.7

185 23 84 32 26 13 33) 14 42 Il 2 16 15

6 24 58 24 37) 13

P. nitida 335 mm TL ¢

36.2

168 20 70 80 al 10 29

9 47 17 14 10

6 23 50 25 37 10

P. nitida 357 mm TL 3

40.1 Sil 24 70 80 ns) Ih 28 8 34 11 IU 1] >) 22 Sil 24 Sy 11

6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Scapulocoracoid measurements of P. alleni and P. nitida expressed as percentage of scapulocoracoid length.

P. alleni P. nitida P. nitida 278 mm TL ? 335 mm TL 3 357 mm TL 3 Scapulocoracoid length (mm) 16.3 Ds Dll Scapulocoracoid height 71 74 I) Premesocondyle 39 40 41 Postmesocondyle 61 60 58 Postdorsal fenestra length 32 3 33 Postdorsal fenestra height 24 WS) 19 Anterior fenestra length V7 15 1S) Anterior fenestra height DS 24 Dy Rear corner 53 59 58

Pavoraja Whitley, 1939

Type species.—Raja nitida Gunther, 1880.

Diagnosis.—Snout with small laterally compressed rostral process; caudal fin poorly developed, with an epichordal lobe and with or without a hypochordal lobe; claspers constricted at origin of glans; dorsal marginal with a distomedial extension forming pseudorhipidion; dorsal terminal 1 and ventral terminal mem- branous and broadly joined along ventral aspect of glans; dorsal terminal 3 with free distal tip forming spur; accessory terminal | with an S-shaped distal extension forming sentinel; accessory terminal 2 with a disc-shaped extension forming spike; rostral shaft slender and reduced, separated by a wide distance from nar- row rostral base; rostral appendices short, propterygia of pectoral girdle extending to rostral appendices; nasal capsules with basal fenestrae; anterior fontanelle and internarial plate moderately narrow; interorbital region narrow; preorbital pro- cesses poorly developed; foramen for anterior cerebral vein posterior to line

scp

pdfe

mtc

msc b pvf Fig. 5. Lateral view of scapulocoracoid: a, R. nitida, AMS 1B.8274; b, Pavoraja alleni, WAM

19117 (Paratype). af—anterior fontanelle, msc—mesocondyle, mtc—metacondyle, pdfe—postdorsal fenestra, prc—procondyle, pvf—postventral foramina, scp—scapular process.

VOLUME 95, NUMBER 1 7

connecting foramina for precerebral and orbitonasal canals; scapulocoracoid sub- rectangular to almost rectangular, moderately short and anteroposteriorly ex- panded; without anterior bridge; three postventral foramina; precaudal mono- spondylous vertebrae ranging from 26 to 29 and predorsal caudal diplospondylous vertebrae ranging from 66 to 79.

Remarks.—Arhynchobatis, Bathyraja, Breviraja, Gurgesiella, Psammobatis, Pseudoraja, “‘Raja’’ waitei, and Sympterygia also possess reduced rostra, a de- rived state (McEachran and Compagno 1979, 1982). Rostra of Arhynchobaitis, Bathyraja (in part), Psammobatis, ‘‘Raja’’ waitei, and Sympterygia are basally segmented and could presumably have evolved into the P. nitida state by distal retraction of the rostral shaft. The rostral shaft in Psammobatis 1s partially re- tracted while that of Pseudoraja is nearly absent (McEachran and Compagno 1979). However, all of these taxa differ from P. nitida in structure of the claspers, neurocrania and scapulocoracoids (Compagno and McEachran, in prep.) and thus it seems likely that rostral reduction has occurred separately several times within the skates. Pavoraja nitida shows a closer relationship to the genera with reduced but basally unsegmented rostra (Breviraja and Gurgesiella) in structure of the neurocranium and scapulocoracoid, and to a lesser degree, in clasper structure, and possibly could have been derived from a common ancestor of either of these taxa. Breviraja possesses a distally segmented rostral shaft which could have _ evolved into the P. nitida state by basal retraction of the proximal, unsegmented part of the shaft. However, uniqueness of the clasper and rostral structure of P. nitida precludes its classification with either Breviraja or Gurgesiella and sup- ports the resurrection of Pavoraja.

Based on the examination of several small immature specimens of R. polym- mata, this species is classified in Raja rather than Pavoraja. It possesses a stout rostral shaft and in all aspects agrees with the anatomical character states within Raja. Further comments on its relationships must await the procurement of large, mature specimens to make more extensive anatomical observations and to ex- amine the claspers.

Pavoraja nitida (Gunther, 1880) (Fig. 6, Table 3)

Diagnosis.—Disc width greater than 54% of total length; orbital length 1.7 to 2.1 times as long as spiracle; anterior margin of pelvic fins less than 75% of distance from origin of anterior lobe to extreme posterior margin of fin; second dorsal fin and epichordal lobe of caudal fin confluent; epichordal lobe shorter than base of second dorsal fin; hypochordal lobe of caudal fin absent; dorsal surface dark brown with small light spots, some of which are arranged into occelli; mature males range from 312 to 363 mm TL.

Comments.—It is very unlikely that the type locality of P. nitida is South America rather than Australia, as suggested by Whitley (1940). The holotype of P. nitida does not differ significantly from the specimens from Australia. No Specimens resembling P. nitida have been reported from South America, and Specimens resembling this species have not been discovered in the large samples of South American skates studied by McEachran.

Material examined.—BMNH 1879.5.14.417 (Holotype), Two Fold Bay, New

8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Pavoraja nitida, BMNH 1879.5.14.417 (Holotype).

South Wales, Australia, HMS Challenger, 6 April4 June 1874. AMS 1A.2493 (1), AMS 1A.3904.05 (1), AMS 1B.4324.25 (2), AMS 1B.5275 (2), AMS 1B.8274 (3), AMS E.2165 (1), AMS E.5453 (1), AMS I.10823.24 (2), AMS I.16564.001 (1), AMS 1A.3904.05 (1), TMH D816 (1).

Pavoraja alleni, new species

Holotype.—WAM P19118, 297 mm TL, mature male, collected off northwest- ern Australia, near Rowley Shoals, 17°17.0’S, 119°57.0’E, 350 m, 20 December 1969.

Paratypes.—WAM P19117, 278 mm TL, female, collected with holotype; WAM P8226, 159 mm TL, female, collected December 1963 or January 1964 in the eastern Indian Ocean, 17°05’S, 119°48’E, aboard the Umitaka Maru.

Diagnosis.—Disc width less than 52% of total length; orbital length 2.2 to 3.2 times as long as spiracle; anterior margin of pelvic fins 81 to 104% of distance from origin of anterior lobe of fin to extreme posterior margin of fin; second dorsal fin and epichordal lobe of caudal fin not confluent, distance from posterior margin of base of second dorsal fin to tip of tail considerably greater than length of base of second dorsal fin; hypochordal lobe of caudal fin small but present; dorsal surface light tan with minute dark spots loosely concentrated into ill de- fined blotches but not forming ocelli; males mature at 297 mm TL.

Description.—Disc 1.1 times as broad as long; maximum angle in front of spiracles 107° in holotype (110° to 112° in paratypes); margin of disc convex except concave opposite orbits and spiracles; outer corners of disc broadly round-

VOLUME 95, NUMBER 1 9

Table 3.—Proportional measurements and meristic values for P. alleni and P. nitida. Pro- portions are expressed as percentage of total length.

P.alleni P.alleni P.alleni P.alleni PP. nitida P. nitida P. nitida

(holotype) (paratype) (paratype) xy (holotype) n= 15 x Sex 3 © 2 3 Total length (mm) 297 278 159 209 242-358 Disc width 50.8 49.3 47.2. 49 59 54-62 59 Disc length 44.4 43.5 42.8 44 50 45-52 Si Snout length (preocular) 10.4 9.6 Ea eles 11.0 8.3-11.4 5) Snout length (preoral) 9.8 10.5 22s O57/ 11.8 9.3-12.6 10.6 Snout to maximum width 28.6 26.3 BY 2 29 25-31 28.7 Prenasal length 8.1 8.0 Oe 8.4 8.2 6.3-9.1 Teil Orbit diameter 4.7 3) 4.7 4.4 al 5.2-6.1 D5 Distance between orbits Doll Spl 3.3 3.0 31,2 2923.7, 353) Orbit and spiracle length 325 4.6 D2 5.0 6.3 6.2-8.9 6.6 Spiracle length LS) 1.8 1.4 1.6 B)522 2.7-6.3 Shy? Distance between spiracles 6.6 6.0 6.4 6.3 6.5 4.2-6.7 6.3 Mouth width 6.9 6.1 5.8 6.3 6.3 5.9-7.9 6.9 Nare to mouth SD) 3.5 4.5 oll gi) 3.24.7 4.2 Distance between nostrils 4.8 4.5 ayy 4.8 4.1 3.24.7 308) Width of first gill opening 0.9 1.0 0.8 0.9 7 1.6-2.1 ow Width of third gill opening 1.0 lest 0.8 1.0 159 1.5—2.0 lew Width of fifth gill opening 0.7 0.9 0.8 0.8 1.6 1.3-1.6 1.4 _ Distance between first gill openings 12 12 2 2 13 15-12 12 Distance between fifth gill openings 8 8 a 8 7 6-8 6 Length of anterior pelvic lobe 12 11 13 12 12 12-14 13 Length of posterior pelvic lobe 13 14 13 14 16 17-18 17 Distance—snout to cloaca 41 4] 40 41 44 41-45 43 Distance—cloaca to Ist dorsal fin 47 47 48 47 46 43-47 45 Distance—cloaca to caudal origin 55 57 5) 35) 54 51-56 53 Distance—cloaca to caudal tip 2) 59 61 60 a7 54-59 57 Number of tooth rows (upper jaw) 38 359) 45 39 34 31-36 3) Sample size of radiographs 3 4 Number of trunk vertebrae 26 28 26 26.7 ped 26-29 ies Number of predorsal caudal vertebrae 79 71 74 74.6 66 66-72 70.2 Number of pectoral radials 63 67 65.5 WZ 62-73 69.8 Number of pelvic radials 16 15 IS).5) 20 19-20 IF)

ed. Tip of snout with small, laterally flattened process. Axis of greatest width 76% (67 to 68%) of distance from tip of snout to axil to pectoral fins. Pelvic fins deeply incised, anterior lobe narrow and acutely pointed; anterior margin 85% (81 to 104%) as long as distance from origin of anterior lobe to posterior extreme of fin. Tail slender, little depressed, its width at midlength about two-thirds di- ameter of eye. Tail with narrow lateral fold along ventrolateral surface running from near tip of pelvic fins to origin of hypochordal lobe of caudal fin, widening near tip of tail fold to equal height of epichordal lobe of caudal fin. Length of tail from center of cloaca to distal tip 1.4 times (1.5 times) distance from tip of snout of center of cloaca.

Preocular length 2.5 times (2.2 to 2.5 times) as long as orbit; preoral length 2.0 times (2.4 to 2.5 times) internarial distance. Interorbital distance 0.6 (0.7 to 0.8) times length of orbit, orbit length 3.1 (2.2 to 3.3) times as long as spiracles.

10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Pavoraja alleni, WAM P19118 (Holotype): a, Dorsal view; b, Ventral view; c, Ventral view of head.

Fig. 8. Pavoraja alleni, WAM P19117 (Paratype).

VOLUME 95, NUMBER 1 11

Anterior and posterior nasal flaps without fringes; without nasal pits. Upper and lower jaws moderately arched (little arched). Teeth with pointed cusps near sym- physes of jaws, with rounded cusps near margin of jaws (with rounded cusps throughout jaws), teeth arranged in more or less transverse series (in quincunx).

Distance between first gill slits 2.5 (2.4 to 2.8) times as great as between nares; distance between fifth gill slits 1.6 (1.3 to 1.8) times as great as between nares; length of first gill slits 1.4 (1.1 to 1.2) times length of fifth gill slits and 0.1 (0.1 to 0.2) times mouth width. First dorsal fin slightly higher and longer than second; interspace between dorsal fins equal to or slightly shorter than base of first dorsal fin; second dorsal fin separated from epichordal caudal-fin lobe by distance equal to one-half base of second dorsal fin; epichordal lobe low, length of base about equal to that of second dorsal fin; distance from end of base of second dorsal to tip of tail considerably greater than length of base of second dorsal fin; hypo- chordal caudal lobe small.

Upper surface of disc, pelvics and tail densely covered with denticles. Ventral surface naked. Orbit with 3 thorns on anteromedial margin and 3 thorns on pos- teromedial margin; 3 prenuchal and 1 nuchal thorns; 3 irregular rows of thorns on dorsal surface of tail, no interdorsal thorns. Holotype with 5 alar spines on left and 4 on right lateral aspect of disc and a patch of malar thorns on antero- lateral aspect of disc.

_ The claspers of P. alleni are similar to those of P. nitida with the following

exceptions: claspers very slender; rhipidion poorly developed; sentinel curved laterally and extending to tip of glans (Fig. 1b); dorsal terminal | only loosely connected to ventral terminal along ventral aspect of glans; dorsal terminal 3 with a longer distal extension forming spur (Fig. 2d, e, f); ventral marginal with a truncated distal margin; accessory terminal | with a relatively longer and less curved distal extension forming sentinel; accessory terminal 2 with a more elon- gated shaft supporting disc-like extension forming spike.

The neurocranium of P. alleni is similar to that of P. nitida with the following exceptions: nasal capsules set at about a 30° angle to transverse axis of neuro- cranium (Fig. 4); rostral base better developed, extending nearly to leading edge of nasal capsules; precerebral space broader (Table 1), inner walls of nasal cap- sules not appreciably bulging into precerebral space; postorbital processes poorly developed; jugal arches relatively slender.

The scapulocoracoids of P. alleni are similar to those of P. nitida (Table 2) with the following exceptions: the anterior vertical margin is more perpendicular to horizontal axis (Fig. 5b); posterior corner more posteriorly located; postdorsal fenestra is oval-shaped rather than elliptical.

Color.—Dorsal surface uniformly light tan with minute dark spots loosely con- centrated into ill defined, symmetrically arranged blotches; tail darker with four obscure brown bands; dorsal fins tan with brown blotches. Ventral surface light tan.

Etymology.—Named after Gerald R. Allen (WAM) who furnished us with the specimens of the new species.

Acknowledgments

We wish to thank Alwyne Wheeler and Peter J. P. Whitehead for providing work space at the British Museum (Natural History); Gerald R. Allen (WAM),

12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

A. P. Andrews (TMH), and Doug Hoese (AMS) for providing specimens; and Matthias Stehmann for providing a radiograph and photograph of the holoype of Pavoraja nitida. Figures la, 2a, b, c and 3 were prepared by Debbie Allen, other figures by Janice D. Fechhelm. Helen Feney labelled and mounted the figures.

Literature Cited

Bigelow, H. B., and W. C. Schroeder. 1953. Fishes of the western North Atlantic, Part II. Sawfishes, guitarfishes, skates and rays, and chimaeroids.—Memoir Sears Foundation for Marine Re- search 1(2):588.

Fowler, H. W. 1941. The fishes of the groups Elasmobranchii, Holocephali, Isospondyli, and Ostar- ophysi obtained by the United States Bureau of Fisheries Steamer “‘Albatross’’ in 1907 to 1910, chiefly in the Philippine Islands and adjacent seas.—United States National Museum Bulletin 100(13): 1-879.

Hulley, P. A. 1972. The origin, interrelationships and distribution of southern African Rajidae (Chon- drichthyes, Batoidei).—Annals of the South African Museum 60: 1-103.

Ishiyama, R. 1958. Studies on the rajid fishes (Rajidae) found in the waters around Japan.—Journal Shimonoseki College of Fisheries 7: 1-394.

McEachran, J. D., and L. J. V. Compagno. 1979. A further description of Gurgesiella furvescens

with comments on the interrelationships of Gurgesiellidae and Pseudorajidae (Pisces, Rajoid-

e1).—Bulletin of Marine Science 29(4):530-553.

, and . 1982. Interrelationships of and within Breviraja based on anatomical structures

(Pisces: Rajoidei).—Bulletin of Marine Science, in press.

Stehmann, M. 1970. Vergleichend morphologische und anatomische Untersuchungen zur Neuord- nung der Systematik der nordostatlantischen Rajidae.—Archiv fur Fischeriewissenschaft 21:73-164.

Whitley, G. P. 1939. Taxonomic notes on sharks and rays.—Australian Zoologist 9:227—262, 2 plates.

. 1940. The fishes of Australia. Part I. The sharks, rays, devil fishes, and other primitive fishes

of Australia and New Zealand. Royal Zoological Society, New South Wales, Sydney. 279 pp.

Department of Wildlife and Fisheries Sciences, Texas A&M University, Col- lege Station, Texas 77843

PROC. BIOL. SOC. WASH. 95(1), 1982, pp. 13-26

A NEW SPECIES OF GYMNURE, PODOGYMNURA, (MAMMALIA: ERINACEIDAE) FROM DINAGAT ISLAND, PHILIPPINES

Lawrence R. Heaney and Gary S. Morgan

Abstract.—A new species, Podogymnura aureospinula, of Philippine gymnure is described from Dinagat, a small island off the northeast coast of Mindanao in the southern Philippines. This species is distinguished from other members of the subfamily by the possession of spiny pelage, inflation of the frontal region, and presence of a distinct cusp at the base of the talonid notch. This new species is second in size among the living Echinosoricinae only to Echinosorex gymnurus. The relationship of Podogymnura to the other extant genera of echinosoricines is discussed. Based on these comparisons, Podogymnura and Echinosorex are shown to share a number of cranial and dental characters and are considered to be more closely related to one another than either is to Hylomys or Neotetracus.

In 1972 and 1975, Dioscoro S. Rabor and a field party from Mindanao State _University collected mammals and birds on Dinagat and Siargao islands, which are located off the northeast coast of Mindanao in the southern Philippines. The itinerary, habitat descriptions, and a report on the birds from the 1972 trip may be found in duPont and Rabor (1973), and a report on the mammals in Heaney and Rabor (1982). Specimens from this collection are housed in the Delaware Museum of Natural History, University of the Philippines at Los Banos, and the U.S. National Museum of Natural History. Among the mammals collected on Dinagat are four specimens of a unique member of the family Erinaceidae. In this paper, we describe these specimens as a new species and discuss the relationships of the Philippine gymnures, Podogymnura, within the erinaceid subfamily Echi- nosoricinae.

Methods

External measurements were taken from collector’s labels. Cranial measure- ments (Table 1) were taken by Heaney with dial calipers graduated to 0.1 mm. Dental measurements (Table 2) were taken by Morgan using an Anderson cra- niometer attached to a Bausch and Lomb binocular microscope (Anderson 1968). All cranial measurements are as defined in DeBlase and Martin (1974) except the following: rostral length, from midline at anterior tip of nasals to orbital margin of infraorbital canal; rostral breadth, taken at labial edge of premaxillae just posterior to I?; post-palatal depth, depth of cranium measured at the point just posterior of palate to point above at 90° to occlusal plane of molars; I' to M°, maximum labial length from anterior edge of I' alveolus to posterior edge of M?® at alveolus; M? to M?, greatest width of palate taken at labial margin of alveoli; palatal width at M?, alveolar distance between lingual margins; height of coronoid, maximum vertical height from ventral edge of mandible to tip of coronoid process; depth of mandible, vertical height from ventral edge of mandible to alveolar

14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

surface between M, and M,; thickness of mandible, distance from lingual to labial edge of mandible taken between M, and M,. All dental measurements in Table 1 are dimensions of the teeth, not their alveoli, and represent the maximum length and/or width of a specified tooth or of a series of teeth. Dental nomenclature follows Szalay (1969:202). Specimens from the following museums were used in this study (standard acronym follows in parentheses): Delaware Museum of Nat- ural History (DMNH), Field Museum of Natural History (FMNH), University of the Philippines at Los Banos, Museum of Natural History (UPLB), and the U.S. National Museum of Natural History (USNM).

Podogymnura aureospinula, new species

Holotype.—DMNH 4386, adult female, skin and skull. Obtained 23 April 1972 by Dioscoro S. Rabor (original number 259) at Plaridel, Albor Municipality, Dinagat Island, Surigao del Norte Province, Republic of the Philippines. Skin well prepared and in good condition. Skull complete except for zygomatic arches, both of which are broken (Figs. 1, 2).

Referred specimens.—The holotype and one adult male from Balitbiton, Loreto Municipality, Dinagat (UPLB 3753) were examined and measured. Two additional specimens (one male, one female) from Kambinlio, Loreto Municipality, Dinagat, are in the collection at UPLB, but were not examined.

Diagnosis.—Size large; dorsal pelage short and spinous, golden brown color overall; temporal, sagittal, and nuchal crests prominent; frontal region conspic- uously inflated; interorbital region strongly constricted; external pterygoid pro- cesses large and separated at base from internal pterygoid processes by deep groove; mandibular rami robust; P? large; P* broad lingually; metaconule prom- inent on M! and M?; metacone present on M?®; P, with small, but distinct, meta- conid; distinct cusp at base of talonid notch between metaconid and entoconid on M,—M,.

Description.—Size (Tables 1 and 2) large for extant members of subfamily. Dorsal pelage composed of three types of hairs: slate-gray underfur ca. 5 mm in length; stiff, bristly, or spiny hairs, black at base and remainder golden yellow, many with black tips, ca. 15 mm in length; black, spiny hairs, ca. 12 mm in length. Golden spines twice as abundant as black hairs in middle of back. Black spines densest at mid-dorsum, decreasing in abundance laterally, disappearing on sides. Black-tipped golden spines especially common at mid-dorsum, also dis- appearing on sides. Only golden spines present on sides. Overall color of dorsum golden-brown, with black spines and black-tipped golden spines adding a black speckling. On holotype, golden color distinctly metallic when viewed at proper angle. Other specimens somewhat faded, not as metallic. Ventral pelage lacks spines, grades evenly from dorsal color to brownish-gray over most of venter; throat darker on some specimens. Ventral hairs of two types: soft, gray underfur ca. 5 mm in length, and slightly coarser guard hairs ca. 9 mm in length, gray at base and tipped with light brown. Pelage of rostrum and around eyes short, dense, and spiny. Upper and lower lips clothed in very short, moderately dense, white or light brown fur. Vibrissae dark at base, very light for most of length, up to 55 mm. Rhinarium long, naked, and distinctly bilobed, with nostrils opening later- ally. Ears relatively large, appearing naked, but with sparse covering of extremely

VOLUME 95, NUMBER 1 15

Fig. 1. From top to bottom—dorsal, ventral, and left lateral views of cranium and lateral view of mandible of Podogymnura truei truei (FMNH 61453) from Mt. Apo, Davao Prov., Mindanao (1-4) and Podogymnura aureospinula (DMNH 4386), holotype, from Plaridel, Dinagat, Surigao del Norte Prov. (5-8). Actual size.

Short, white hairs. Fore and hind feet with moderate covering of short, white or light brown hairs dorsally, nearly naked ventrally. Hind legs appear almost naked distal to knee joint. Dorsal base of tail with area sparsely furred, nearly naked, 15 mm in diameter. Tail with sparse covering of short hairs. Two pairs of mam- mae, one pair pectoral, one pair inguinal.

Skull large and robust. Temporal crests converge at or slightly anterior to

16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Stereophotographs of upper (1, 1’) and lower (2, 2') dentitions of Podogymnura aureo- spinula (DMNH 4836), holotype. x3.

interorbital constriction to form sagittal crest. Sagittal crest prominent, 0.5 to 1.0 mm in height anteriorly to 2.5 mm in height posteriorly where it meets nuchal crest. Nuchal crest well developed. Frontal region conspicuously inflated. Fron- tals expanded laterally into orbital fossa. Rostrum relatively broad posteriorly, resulting from inflation of posterodorsal portion of maxilla. Inflation causing con- vexity in dorsal profile of skull, beginning at anterior edge of frontals, approxi- mately dorsal to P?, and extending posteriorly to interorbital constriction. Highest point on skull dorsal to orbits. Interorbital region constricted. Braincase relatively long and not noticeably inflated. Paroccipital and mastoid processes prominent. Mastoid exposure on ventrolateral corner of skull gently concave and composed of thick bone. Post-tympanic process of squamosal large, broad posteriorly, and with well developed epitympanic sinus. Periotic not inflated. Periotic component of bulla reduced, having a distinct, rounded emargination in ventromedial edge. Basisphenoid component of bulla more vertically oriented, projecting ventral to

VOLUME 95, NUMBER 1 167

level of occipital condyles. Tympanic cavity broadly open ventrally. External pterygoid processes large, triangular-shaped, and separated at base from internal pterygoid processes by deep groove. Mandibular rami relatively thick.

All incisors single-rooted, I’ enlarged. I? and I? much smaller, nearly identical in size. Canine double-rooted, long, and flared laterally. P? single-rooted, small compared to other premolars. P? three-rooted, some specimens with slight lingual expansion and small, but distinct, hypocone. P* broad lingually, with hypocone and protocone parallel to palatal midline. Metaconule prominent on M? and M?, protoconule absent. M? relatively large, with small, but distinct metacone. I, and I, approximately same size, spatulate, and procumbent. I, considerably smaller. Canine large and vertical, highest tooth in lower tooth row. P, and P; compara- tively robust, P, one-half the size of P,. P, large, talonid basin moderately to well developed, and distinct metaconid present. Lower molars with small cusp at base of talonid notch between entoconid and metaconid, sometimes absent on Ms. Postcristid on M, and M, slopes lingually at hypoconulid to meet moderately to strongly developed postcingulid.

Etymology.—L. aureus, golden; L. spinula, diminutive of thorn. The specific name refers to the golden spines which characterize the dorsal pelage of this species. We suggest ‘“‘golden-spined gymnure’”’ as an English name.

Comparisons.—Podogymnura aureospinula is more closely related to Podo- gymnura truei than to any other species in the Echinosoricinae as judged by the combination of the following characters: long rostrum, absence of postorbital processes, constricted interorbital region, extreme anterior placement of upper molariform teeth relative to orbit and infraorbital foramen, relatively small I’, I? and I? equal in size, large laterally flaring upper canines, loss of P!, comparatively large P?, and M! and M? square in outline. On the other hand, P. aureospinula possesses at least three derived characters that are unique among the Echino- soricinae: spiny dorsal pelage, conspicuously inflated frontal region, and presence of a cusp at base of talonid notch on lower molars. These characters might justify generic distinction for P. aureospinula; however, rather than erect a monotypic genus, we choose to place this new species in the genus Podogymunura to indicate the presumed monophyletic nature of the two endemic Philippine erinaceids.

Podogymunura truei is the only echinosoricine which requires detailed compar- ison with P. aureospinula. The other extant echinosoricine genera are compared to Podogymnura in more general terms in the Discussion section. Podogymnura aureospinula and P. truei are quite different in external appearance. Besides its smaller size, P. truei has a longer, softer pelage, with no indication of spines, or even coarse hairs. Its underfur is particularly long, about twice as long as that of P. aureospinula. The dorsal surfaces of the fore and hind feet of P. truei have longer, darker hairs. The overall color of the pelage is darker in P. truei; dorsally it varies from reddish to chestnut brown compared to the golden brown color of the large species, and ventrally it is a medium brown, whereas the venter of P. aureospinula is gray, with a brownish tinge.

The most obvious difference between the skulls of Podogymnura aureospinula and P. truei is greater size of the former, its skull being 20% longer than the largest skull of P. truei measured (Table 1). Perhaps as a result of allometric changes correlated with increasing skull size, the temporal, sagittal, and nuchal crests and paroccipital and mastoid processes are more prominent in P. aureo-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

18

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VOLUME 95, NUMBER 1

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20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 2.—Selected dental and mandibular measurements (mm) of Podogymnura species.

Podogymnura Podogymnura Podogymnura Podogymnura

aureospinula truei truei truei truei truei minima Measurement holotype, UPLB 3753 _ holotype (N = 5) (N = 45)

Length of maxillary tooth row 20.6, 20.4 [52 15.5 + 0.44 15.0 + 0.65

(15.1—16.2) (14.4-15.9)

Length from M! to M? 9.8, 9.6 6.7 ies 0i20 7.0 + 0.33 (6.9-7.4) (6.6~7.5)

Length of P® Me PES) 1.8 19 1050 1.8 + 0.08 (1.8—2.1) (1.7—1.9)

Width of P® ORAS MTT, ell ese = 017. 1.4 + 0.07 (1.2-1.6) (1.3—1.5)

Length of P* 37, 310 D2 2.4 + 0.05 mse (Ibi (2.3-2.4) (2:3=2.6)

Width of P* 3.2, 3.0 DN 2.4 + 0.10 2.4 + 1.54 (2.3—2.5) (2.2—2.5)

Length of M! 3.8, 3.8 2.6 2.9 + 0.04 2.8 + 0.11 (2.82.9) (2.62.9)

Width of M! alls St! 2.3 Deh = On 2.6 + 0.08 (2.62.9) (2.5-2.7)

Length of M? BWo)5 Bias) 2.4 2°) 20S 2252-016 (2.42.7) (2.3—2.7)

Width of M? 3.7% 33 2.4 DEG, == 10205 Donne lel (2.6—2.7) (2.5—2.7)

Length of M? hs, 2D led 1.9) = O15 1.8 + 0.09 (1.7—2.1) (1.7—1.9)

Width of M?® DES OES 1.8 2.1 + 0.08 2.0r=,0 eli (2.0-2.2) (1.9-2.1)

Total length of mandible 41.2, 40.8 B30 e077. 28.8 + 0.58 (29.6—31.5) (28.2-29.6)

Height of coronoid 13)... 113).4! 9.2 9.7 + 0.50 9.3 + 0.59 (9.0-10.4) (8.6—10.0)

Depth of ramus between M, and M, 4.7, 4.8 Zall 3 op = OR2 Bop eas | )e75) (3.0-3.6) (2.8-3.5)

Breadth of ramus between M, and 2.6, 2.8 1.6 1.8 + 0.05 1.7 + 0.07 M, (1.7—1.8) (1.6+1.8)

Length of mandibular toothrow DNA 207. 15.8 1622) 310354 15.7 + 0.64 (15.7=17 1) (15.1—-16.6)

Length from M, to M, 11.2, 10.5 7.8 8.0 + 0.26 7.8) 10 (7.6-8.2) (7.5-8.3)

Length of P, 3.0, 2.6 1.9 2.0 + 0.10 2.0 + 0.19 (129-771) (1.8-2.3)

Width of P, IRS, les) ok 1.3 + 0.04 12008 (1.2=1.3) (1.1-1.3)

Length of M, 4.3, 4.1 3.0 Byaa | ree (0a 2.9 + 0.09 (39-3) (2.9-3.2)

Width of M, RSE on 5) loi 1.9+ 0.13 1.8 + 0.04 (1.7—2.0) (1.8-1.9)

VOLUME 95, NUMBER 1 21

Table 2.—Continued.

Podogymnura Podogymnura Podogymnura Podogymnura

aureospinula truei truei truei truei truei minima Measurement holotype, UPLB 3753 holotype (N = 5) (N = 4-5)

Length of M, 4105 Sal! Met 2.8 + 0.13 2.7 + 0.08 (2.62.9) (2.6-2.8)

Width of M, 2.4, 2.4 1.6 18+ 0.11 1.8 + 0.09 (1.7-1.9) (1.71.9)

Length of M, 33.54 De 2.3+0.11 2.3 + 0.15 (22214) (Qal=255)

Width of M, Delle, PAY 1.4 1255-70505) ls) ee LNG

(1.5-1.6) (1LZ=LD)

spinula, approaching the condition seen in the much larger Echinosorex. In most skulls of P. truei, the weak temporal crests meet near the anterior edge of the interparietal, forming a short, weak sagittal crest, while in larger skulls of the Same species, the temporal crests meet somewhat farther forward. The temporal crests of P. aureospinula converge at or slightly anterior to the interorbital con- striction to form the strong sagittal crest, which is particularly high posteriorly where it bisects the interparietal. The nuchal crest of P. aureospinula is more strongly developed than in P. truei.

Podogymuura truei shows no evidence of the frontal inflation characteristic of P. aureospinula. The dorsal margin of the skull of P. truei rises in a nearly straight line to reach a maximum height above the glenoid region, rather than dorsal to the orbits as in P. aureospinula. The inflation of the frontals and pos- terodorsal portion of the maxilla of P. aureospinula is not characteristic of any other modern echinosoricine, although some erinaceines (e.g. Paraechinus) have moderately inflated frontals. The interorbital region appears to be more constrict- ed in P. aureospinula, although this feature is certainly enhanced by the inflation anterior to the constriction. The more prominent development of the sagittal and nuchal crests gives the braincase of P. aureospinula the appearance of being longer, whereas the braincase of the smaller species appears shorter, broader, and more bulbous, particularly in smaller specimens.

The mastoid exposure on the posterolateral corner of the skull is inflated in Podogymuura truei and the bone is thin and nearly transparent. The mastoids of P. aureospinula are gently concave and composed of thicker bone. The periotic of P. truei is also inflated. The periotic is not inflated in P. aureospinula and the periotic component of the bulla has a small, rounded emargination in its ventro- medial edge which is lacking in the smaller species. The basisphenoid portion of the bulla of P. aureospinula is more vertically oriented, projecting ventral to the occipital condyles, whereas the basisphenoid bulla of P. truei is more horizontal, tending to enclose the tympanic cavity. Coupling the more vertical orientation of the basisphenoid bulla with the reduced periotic portion of the bulla, the tympanic cavity is more open ventrally in P. aureospinula. The post-tympanic process of the squamosal is relatively larger in P. aureospinula than in any other modern echinosoricine. Its external pterygoid processes are also relatively large, but these are smaller in P. truei. Podogymnura aureospinula has a deep groove at the base

22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of the external pterygoid processes which separates them from the internal pter- ygoid processes. There is no evidence of this groove in any specimen of P. truei examined. As with the skull, the mandible of P. aureospinula is larger and more robust than that of its smaller counterpart. The mandibular rami are particularly thickened in P. aureospinula.

The upper incisors and canines of Podogymnura aureospinula and P. truei are similar in morphology. The P? is somewhat longer and broader in P. aureospinula; it is more conical in P. truei. In most specimens of P. truei, the P® is two-rooted, thin, and blade-like, with no indication of a protocone or hypocone. Butler (1948) stated that the P? of Podogymnura had only two roots; however, the larger series of specimens now available shows this character to be variable. Most specimens of P. truei have a two-rooted P®, but the few that have a slight lingual expansion of P® have three roots. The P® of P. aureospinula is three-rooted, and bears a small hypocone. The P* of P. aureospinula is broader lingually, the hypocone is larger, and the protocone is higher and more lingually placed. In comparison, the P* in the smaller species is rather narrow lingually, the hypocone is small, and the protocone is located closer to the paracone. The M! and M? are similar in the two species, except for the more prominent metaconule and cingula in P. au- reospinula. The M? of P. aureospinula is larger and bears a small metacone, the latter being absent on the M? of P. truei.

As in the upper dentition, the P, and P, are relatively larger and more robust in P. aureospinula than in P. truei. The talonid basin is reduced on P, in P. truei and a metaconid is absent or tiny. The P, of P. aureospinula has a moderately to well developed talonid basin and the metaconid is distinct. The morphology of the lower molars is similar in the two species. P. aureospinula has a small, distinct cusp located at the base of the talonid notch between the entoconid and metaconid on M,, M,, and sometimes M,;. The entoconulid is normally located slightly anterior to, but close to the entoconid, and therefore this cusp does not appear to be homologous with an entoconulid. This cusp is absent in P. truei and seems to be unique among modern echinosoricines.

Ecology.—Virtually nothing is known of the natural history of Podogymnura aureospinula. The vicinity where the holotype of P. aureospinula was taken was described by duPont and Rabor (1973:4) as *‘. . . a logged area in rolling country and low hills where there were still many patches of remnant dipterocarp forests in the surrounding localities.’’ These forests are dominated by Dipterocarpus, Shorea, Hopea, Anisoptera, and Pentacme among the Dipterocarpaceae, and include members of at least eleven other plant families. Undergrowth consists mainly of rattan and ferns. Other terrestrial mammals taken at the type locality of P. aureospinula include Urogale everetti, Tarsius syrichta, Cynocephalus vo- lans, Sundasciurus mindanensis, Exilisciurus surrutilus, Batomys sp., Rattus ev- eretti, and Rattus rattus (Heaney and Rabor 1981).

Additional specimens examined.—Podogymnura truei truei (2 63, 4 2°). PHILIPPINES, Mindanao, Davao Province: E slope of Mt. McKinley, 5800 ft. elev., FMNH 56129, 56172, 56181; N slope of Mt. Apo, Lake Linau, 7800 ft. elev., FMNH 61435; E slope of Mt. Apo, Baclayan, 5400 ft. elev., FMNH 61453; Mt. Apo, 6000 ft. elev., USNM 125286 (holotype).

Podogymnura truei minima (2 66,3 2° ¢). PHILIPPINES, Mindanao, Bukid- non Province: Mt. Katanglad, near Malay Balay, DMNH 5949-5953.

VOLUME 95, NUMBER 1 23

Echinosorex gymnurus albus (1 6, 4 22). INDONESIA, Borneo, Sempang River: USNM 145581-582, 145584—586.

Hylomys suillus dorsalis (10 63, 1 2). MALAYSIA, Borneo, Sabah, Mt. Kinabalu, Bundu Tuhan: USNM 292337-339, 292341-342, 292350-354, 292356.

Neotetracus sinensis (1 6, 2 22, 42). CHINA: Yunnan, Ho mu shu Pass, USNM 241402; Szechuan, Kwan Shien, 3000 ft. elev., USNM 258124-129.

Discussion

The genus Podogymnura and its type species, P. truei, were described by Mearns (1905) from a single specimen collected on Mount Apo, south-central Mindanao, in the Philippines. Until the late 1940’s, this specimen, an adult female consisting of a complete body in alcohol and a skull lacking the zygomatic arches and braincase, was the only known specimen of the genus. Mearns provided a detailed description of the external characters of P. truei but he did not illustrate the cranium or mandible and his description of them was brief. He described the dentition of P. truei, comparing it with that of Hylomys, but his descriptions and comparisons are so general that they are of little use. Lyon (1909) noted that although Podogymnura and Hylomys appeared to be closely related with respect to size and external characters, they were distinct dentally. Cabrera (1925) pro- vided a key to the living echinosoricine genera in which he distinguished Podo- gymnura by the combination of loss of P}, P*® larger than P?, and larger upper canines. Butler (1948) regarded Podogymnura as intermediate between Echino- sorex and the smaller Hylomys and Neotetracus, although he pointed out that in the enlargement of the canines, relatively large P®, length of the rostrum, and position of P* and M' relative to the orbit and infraorbital foramen, Podogymnura is similar to Echinosorex.

Sanborn (1952) reported on 64 specimens of Podogymunura truei collected on Mount Apo and Mount McKinley on Mindanao. He compared Podogymuura to the other genera of modern echinosoricines, reaching the same general conclusion as did Butler. Sanborn (1953) described a new subspecies of Podogymnura truei, P. t. minima, from four specimens collected on Mount Katanglad, Bukidnon Province, north-central Mindanao. His diagnosis was based exclusively on the smaller size of P. t. minima. Detailed cranial and dental measurements of P. f. truei (including the type) and topotypic specimens of P. t. minima (Tables | and 2) indicate that, on the whole, the available specimens from Mount Katanglad are slightly smaller than specimens from Mount Apo and Mount McKinley, but there is broad overlap in size between them, especially in dental measurements. In fact, the teeth of the type specimen of P. t. truei from Mount Apo are actually smaller in many dental measurements than the teeth of the topotypic specimens of P. t. minima. Sanborn specifically noted that there was no difference in color between his series of P. t. minima and the nominate form. The DMNH specimens of P. t. minima, however, are lighter in color than topotypic specimens of P. f. truei, being slightly more reddish brown and having the tail uniformly lighter. Besides the references cited above, Hollister (1913), Taylor (1934), and Alcasid (1970) mentioned Podogymnura, but provided no new information.

Comparison of Podogymnura with other living echinosoricines.—Butler (1948) made exhaustive comparisons of three of the five extant genera of echinosori-

24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cines: Hylomys, Neotetracus, and Echinosorex. His comparisons of Podogym- nura were general, as they were based solely on published descriptions and fig- ures of the type of P. truei, the only specimen of the genus known at that time. The fifth living genus of echinosoricine, Neohylomys, was not described until 1959. Since 1948 many specimens of Podogymnura have been collected, but the genus has not been adequately compared with the other members of its subfamily. Our description of a new species of Podogymnura, which in some characters appears to bridge the morphological gap between Podogymnura and Echinosorex, calls for fuller comparisons and a reassessment of the phylogenetic relationships of the Philippine gymnures.

As Butler (1948) and others have noted, Hylomys and Neotetracus are closely related forms, although in our opinion they do represent distinct genera, contrary to Van Valen (1967). The status of Neohylomys is uncertain; the only specimens are in China, and unavailable for study. Many of the characters unique to Hy- lomys and Neotetracus, such as the reduced antemolar dentition, the palatal perforations, the presence of posterior processes of the maxillae and anterior processes of the parietals which extend across the frontals and meet or nearly meet dorsal to the orbits, the prominent flange on the anterior edge of the orbit, and the strongly concave anterior portion of the zygomatic arch for attachment of the lateral snout muscles, are derived characters which demonstrate a close phylogenetic relationship between the two genera. Conversely, many of the char- acters that are shared by Podogymnura and Echinosorex, but are not present in the other two genera, appear to be primitive for the Echinosoricinae. Of these characters, the longer rostrum, better developed maxillary dentition anterior to P*, widely separated maxillae and parietals dorsal to the orbits, and the nonper- forated palate are certainly primitive. In the remaining characters shared by Po- dogymnura and Echinosorex, it is difficult to determine the primitive character state for the Echinosoricinae because no appropriate outgroup exists. Butler (1948) felt that the two distinct groups of living echinosoricines represented two separate specialized offshoots from the primitive condition. Suffice it to say that Podogymnura and Echinosorex share so many cranial and dental features that they undoubtedly represent closely related genera.

Although they share many cranial characters, Podogymnura and Echinosorex are extremely different externally. Echinosorex is a larger animal with a tail equal to about 60% of the length of head and bedy. Podogymnura is medium-sized (P. aureospinula) or small (P. truei) and has a tail equal to about 35% of the length of head and body. Podogymuura is a chestnut brown (P. truei) to golden brown (P. aureospinula) color and has relatively short pelage. Echinosorex is either pure white (E. gymnurus albus) or predominantly black with white markings on the neck and face (all other forms of the species) and has long, coarse guard hairs on the dorsum. Based on its external appearance, especially its small size and short tail, Podogymnura truei more closely resembles Hylomys and Neotetracus than Echinosorex.

Additional cranial characters shared by Podogymnura and Echinosorex not listed above include: well developed canines and P®, small supraorbital crests, presence of two longitudinal grooves in maxillary component of hard palate which extend from incisive foramina to small foramina located medial to P? or P®, pres-

VOLUME 95, NUMBER 1 25

ence of a ventral process on maxillary portion of zygomatic arch, and more anterior placement of upper cheek teeth relative to orbit and infraorbital foramen. The tympanic bone is slender and not firmly attached to the bulla in Podogymnura and Echinosorex, whereas in Hylomys and Podogymnura the tympanic is broader and is firmly attached to the edge of the bulla, contributing to its formation and restricting the size of the opening of the tympanic cavity. Although Butler (1948) stated that Echinosorex lacked the anterior process of the tympanic, the large series of echinosoricines at our disposal reveals that both Echinosorex and Po- dogymnura definitely have an anterior process, but it is smaller than that of Hylomys and Neotetracus.

In addition to the differences in external morphology cited above, there are a number of cranial and dental characters that are diagnostic of Podogymnura and distinguish it readily from Echinosorex. These include: less prominent temporal, sagittal, and nuchal crests and mastoid and paroccipital processes; more concave maxillary portion of zygomatic arch; broader, more inflated braincase; more an- terior placement of upper molars relative to orbit and infraorbital foramen (M? is anterior to front edge of orbit and M! is below infraorbital foramen); smaller I; I? and I*® equal in size; canines relatively larger and flared laterally; loss of P}; M? square in outline; absence of protoconule on M!; smaller metaconule on M! and M?; reduced posterolingual apex of M?.

Podogymnura aureospinula is intermediate between Echinosorex and P. truei in many characters, although some of the characters on which this observation is based are probably related to its intermediate size. The more prominent tem- poral, sagittal, and nuchal crests and mastoid and paroccipital processes of P. aureospinula relative to P. truei are almost certainly correlated with its greater size. These features of the bony crests and processes are best developed in Echi- nosorex, the largest genus among living echinosoricines, and are likely to be allometric changes associated with increasing skull size. Other intermediate char- acters of P. aureospinula which are more difficult to ascribe to allometry are the larger P?, more prominent metaconule on M! and M?, the presence of a metacone on M?, and the deep groove separating the bases of the external and internal pterygoid processes. Whether these are derived characters indicating a closer relationship between Echinosorex and P.. aureospinula or whether these represent primitive characters shared by Echinosorex and P. aureospinula and lost by P. truei cannot be determined from available data. Regardless of the similarities between P. aureospinula and Echinosorex, the two species of Podogymnura are certainly more closely related to one another than either is to Echinosorex and have probably been isolated in the Philippines for a considerable period of time.

Acknowledgments

We wish to thank M. D. Carleton, P. Myers, and R. W. Thorington for valuable comments on the manuscript, and G. Lake and S. F. Campbell for assistance in preparation of the manuscript and tables. Photographic assistance was provided by D. Bay and V. Krantz. We thank P. Freeman, D. Niles, D. S. Rabor, R. W. Thorington, and R. M. Timm for permission to examine specimens under their care.

26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Alcasid, G. L. 1970. Checklist of Philippine mammals.—National Museum, Pee of Education (Manila). Museum Publication No. 5:1-51.

Anderson, S. 1968. A new craniometer and suggestions for craniometry.—Journal of Mammalogy 49:221-228.

Butler, P. M. 1948. On the evolution of the skull and teeth in the Erinaceidae, with special reference to fossil material in the British Museum.—Proceedings of the Zoological Society of London 118:446—S00.

Cabrera, A. 1925. Genera Mammalium—Insectivora, Galeopithecia——Museo Nacional Ciencia Natural, Madrid, 232 pp.

DeBlase, A. F., and R. E. Martin. 1974. A manual of mammalogy. Wm. C. Brown Co., Dubuque, Iowa, 329 pp.

duPont, J. E., and D. S. Rabor. 1973. Birds of Dinagat and Siargao, Philippines, an expedition report.—Nemouria (Occasional Papers of the Delaware Museum of Natural History) 10:1-111.

Heaney, L. R., and D. S. Rabor. 1982. An annotated checklist of the mammals of Dinagat and Siargao islands, Philippines.—(Occasional Papers of the Museum of Zoology, University of Michigan) (In press.)

Hollister, N. 1913. A review of the Philippine land mammals in the United States National Museum.— Proceedings of the United States National Museum 46:299-341.

Lyon, M. W., Jr. 1909. Remarks on the insectivores of the genus Gymnura.—Proceedings of the United States National Museum 36:449_456.

Mearns, E. A. 1905. Descriptions of new genera and species of mammals from the Philippine Is- lands.—Proceedings of the United States National Museum 28:425—460.

Sanborn, C. C. 1952. Philippine zoological expedition 1946-47: Mammalis.—Fieldiana: Zoology

33(2):89-158.

. 1953. Mammals from Mindanao, Philippine Islands collected by the Danish Philippine Ex-

pedition 1951—1952.—Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening

115:283-288.

Szalay, F. S. 1969. Mixodectidae, Microsyopidae, and the insectivore-primate transition.—Bulletin of the American Museum of Natural History 140:193-330.

Taylor, E. H. 1934. Philippine land mammals.—Monographs of the Bureau of Science, Manila 30: 1-548.

Van Valen, L. 1967. New Paleocene insectivores and insectivore classification.—Bulletin of the American Museum of Natural History 135:217-284.

(LRH) Museum of Zoology and Division of Biology, University of Michigan, Ann Arbor, Michigan 48109; (GSM) Division of Mammals, U.S. National Mu- seum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Present address of GSM: Florida State Museum, University of Florida, Gaines- ville, Florida 32611.

PROC. BIOL. SOC. WASH. 95(1), 1982, pp. 27-47

CHECKLIST OF THE FISHES OF THE CENTRAL AND NORTHERN APPALACHIAN MOUNTAINS

Jay R. Stauffer, Jr., Brooks M. Burr, Charles H. Hocutt, and Robert E. Jenkins

Abstract.—A table lists 398 forms and 5 intergrade populations in 28 families in an area on the Atlantic slope from the Susquehanna River south to the Peedee River, including Ohio River basin drainages from the Monongahela River in Penn- sylvania to the Tennessee River in Alabama and Tennessee.

The central Appalachians harbor a diverse fish fauna that includes numerous endemics. Jenkins, Lachner, and Schwartz (1972), as part of a zoogeographic analysis of this ichthyofauna, provided a table that lists the fishes of the central Appalachians by river drainage and general habitat. This table has been extremely valuable to ichthyologists, fisheries scientists, and environmental consultants throughout the past decade.

Numerous studies have substantially increased our knowledge of fish distri- bution throughout the central Appalachians (Hambrick et al. 1973, Hocutt and Hambrick 1973, Hocutt et al. 1973, Stauffer et al. 1975, Stauffer et al. 1976, Hocutt et al. 1978, Stauffer et al. 1978, Hendricks et al. 1979, Hocutt et al. 1979, Lee et al. 1980) and indicated that the original table should be updated. Moreover, it was thought that the addition of the Susquehanna, Licking, Green, and Ken- tucky rivers would enhance the usefulness of the faunal list.

The list (Table 1) includes 398 forms and 5 intergrade populations in 28 families. It covers an area on the Atlantic Slope from the Susquehanna River (New York and Pennsylvania) south to the Peedee River (North Carolina and South Caroli- na). Ohio River basin drainages that are included extend from the Monongahela River in Pennsylvania south to the Tennessee River in Alabama and Tennessee.

It should be noted that the list is conservative. If a question exists as to the current or historic presence of a species, it is not included. No attempt is made to distinguish species that were historically present in the drainage from those that currently occur. Trinomials are used only when the distribution of subspecies could be accurately determined.

The authors appreciate the encouragement of Dr. E. Lachner, who recognized the need for a revision of the original faunal list.

28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Fishes of the central and northern Appalachian drainages and their general habitat. KX = category. most frequently inhabited. Drainage occurrence: E = endemic, N = native, NP = probably present—native, NI = regarded as native but possibly introduced, I = introduced, IP = probably or possibly introduced, Ma = euryhaline or diadromous. Native extralimital distribution: So = south on Atlantic slope, No = north on Atlantic slope, O = predominantly Ohio basin form, M = lower and/or central Mississippi basin, G = Gulf of Mexico slope.

PETROMYZONTIDAE

Ichthyomyzon bdellium Ichthyomyzon castaneus Ichthyomyzon fossor Ichthyomyzon gagei Ichthyomyzon greeleyi Ichthyomyzon unicuspis Lampetra aepyptera Lampetra appendix Petromyzon marinus

ACIPENSERIDAE

Acipenser brevirostrum Acipenser fulvescens Acipenser oxyrhynchus Scaphirhynchus platorynchus

POLYODONTIDAE Polyodon spathula LEPISOSTEIDAE

Lepisosteus spatula Lepisosteus oculatus Lepisosteus osseus Lepisosteus platostomus

AMIIDAE

Amia calva

ANGUILLIDAE

Anguilla rostrata

CLUPEIDAE

Alosa aestivalis

Alosa alabamae

Alosa chrysochloris Alosa mediocris

Alosa pseudoharengus Alosa sapidissima Dorosoma cepedianum Dorosoma petenense

HIODONTIDAE

Hiodon alosoides Hiodon tergisus

SALMONIDAE

Coregonus artedii Coregonus clupeaformis

Lowland

x x KK

x mK x

x KK KKK OM

x xX

Upland

mK KK KK

x x

Habitat

Montane Big River

~*~ x

xxx KKK OK * x mK mK va x Km

x x

Stream

KK mK KK mK OK OM OM

Creek

x xX

Peedee

Cape Fear

Ma

Ma

Neuse

Drainage occurrence

Atlantic Slope

Tar

Roanoke

NN <Ne

Ma

Ma

Ma

Ma

Ma

Ma Ma Ma Ma

Ma

Ma

James

York

Rappahannock

Potomac

Susquehanna

Ne NENG ENGIN N N N N N Ma Ma Ma Ma Ma Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

NP

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

Ma

IP

VOLUME 95, NUMBER 1

Table 1.—Continued.

29

PETROMYZONTIDAE

Ichthyomyzon bdellium Ichthyomyzon castaneus Ichthyomyzon fossor Ichthyomyzon gagei Ichthyomyzon greeleyi Ichthyomyzon unicuspis Lampetra aepyptera Lampetra appendix Petromyzon marinus

ACIPENSERIDAE

Acipenser brevirostrum Acipenser fulvescens Acipenser oxyrhynchus Scaphirhynchus platorynchus

POLYODONTIDAE Polyodon spathula

LEPISOSTEIDAE

Lepisosteus spatula Lepisosteus oculatus Lepisosteus osseus Lepisosteus platostomus

AMIIDAE

Amia calva

ANGUILLIDAE

Anguilla rostrata

CLUPEIDAE

Alosa aestivalis

Alosa alabamae

Alosa chrysochloris Alosa mediocris

Alosa pseudoharengus Alosa sapidissima Dorosoma cepedianum Dorosoma petenense

HIODONTIDAE

Hiodon alosoides Hiodon tergisus

SALMONIDAE

Coregonus artedii Coregonus clupeaformis

(or) a OS Dayna ey o.6U6U = 5 NP N N N N N N WN Ma Ma Ma Ma Ma N N

Kanawha: below falls

Ma

Ma

Kanawha: above falls

Drainage occurrence

Ohio Basin & 6 6s > ep 3 Ory ye iS NP NP N N NP NP NP ING NN N NP N N N Ma Ma Ma N Ma Ma Ma Ma Ma Ma Ma N NP N

Kentucky

Lik LZ,

NP

Green

Z

Z ZZ Z

NP

Ma Ma

Ma

Ma

Cumberland: below falls

i 7

EG

Cumberland: above falls

Ze Tennessee

Ly Li Law Lie,

Native extralimital distribution

Atlantic Slope

NoSo

NoSo

NoSo

NoSo

Elsewhere

ie)

Hae 2

GM

GM

GM

GM

30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

x tab} ion =. p +

Lowland Upland Montane

Big River Stream Creek Peedee

Cape Fear Neuse Roanoke James

York Rappahannock Potomac Susquehanna

Tar

Oncorhynchus kisutch

Oncorhynchus nerka

Salmo gairdneri I Salmo trutta I I I I Salvelinus fontinalis XxX DS DP N N NI

Salvelinus namaycush OSMERIDAE

Osmerus mordax NI

UMBRIDAE

Umbra limi X Nea OX Umbra pygmaea 4 XX“ N-N N NUN ON SON NNN

ESOCIDAE Esox a. americanus xX xX xX N N N -“N N 2INSIN SN NSN Esox a. vermiculatus > XX Esox lucius

I

Esox masquinongy Xe XS WX’ I I I Esox niger xX XxX xX XN N N NeoiNeoN aN eNaaNeeN I

Esox reicherti

p= — — Za a” Ail en il onl oon lo

a

—

CY PRINIDAE

Campostoma a. anomalum x OX xX XX IP N N N N Campostoma oligolepis XE 20 OX Xx

Carassius auratus I I I I Clinostomus elongatus NI Clinostomus f. funduloides Clinostomus f. estor Clinostomus f. subsp. Xx Couesius plumbeus Ctenopharyngodon idella xX xX

Cyprinus carpio rT, 2 ~habe a oe eerie ieee Ericymba buccata Exoglossum laurae Exoglossum maxillingua Hemitremia flammea Hybognathus hayi Hybognathus n. nuchalis Hybognathus n. regius Hybopsis aestivalis hyostoma Hybopsis amblops

Hybopsis cahni

Hybopsis d. dissimilis Hybopsis hypsinotus Hybopsis i. insignis Hybopsis i. eristigma Hybopsis labrosa

Hybopsis monacha

Hybopsis storeriana XxX Hybopsis x-punctata

xx x x KKK

IP

* * *

x KK xX ~

x x x

~*~

Z,

Z,

iL,

xx x x KKK KKK KKK KKK KK

mK KKK KKK KKK

*

VOLUME 95, NUMBER 1 31

Table 1.—Continued.

Drainage occurrence

Ohio Basin a I 5 = Native a8 S$ Gistibution os 2 S 3) 5 8 Fs oO = . a EES at accrnrian aie se BS SRG 0 i omar 3 a x 3 5 5 > a 3 o n © he ae ag iN pe emg ae 2) a Seca Se Se Se en ci, Vee, Rok ree Sal ee a 5 So a a Be o o 2 =| =| 5 cs 2) Se pee Sa Oe eh A Omer Oe < cal Oncorhynchus kisutch Oncorhynchus nerka I I Salmo gairdneri I I I I I I i I I I I Salmo trutta I I I I I I I I Salvelinus fontinalis NG OEP NN IP I N NoSo Salvelinus namaycush No OSMERIDAE Osmerus mordax I UMBRIDAE Umbra limi N M Umbra pygmaea NoSo G ESOCIDAE _Esox a. americanus I NoSo G Esox a. vermiculatus New INE EN2 Ne ON N GM Esox lucius I I I I M Esox masquinongy No INN Y UIP ye LNG INE ON IN N Esox niger I I I N N NoSo GM Esox reicherti CY PRINIDAE Campostoma a. anomalum INS dN IND INL TUNG) CINE UING aN N NoSo O Campostoma oligolepis N, NE VNOOAIN GM Carassius auratus I I I I I I I I Clinostomus elongatus N N No OM Clinostomus f. funduloides N N N N NI So O Clinostomus f. estor N N Clinostomus f. subsp. N So Couesius plumbeus No Ctenopharyngodon idella Cyprinus carpio I I I I I I I I I I I I Ericymba buccata INI SENG Ne INE ONIN NO SING ING DON ONT IN efor GME Exoglossum laurae NI N O Exoglossum maxillingua NI No Hemitremia flammea N N G Hybognathus hayi N GM Hybognathus n. nuchalis N ING Ni N GM Hybognathus n. regius NoSo Hybopsis aestivalis hyostoma N Ne ONe IN! AN. EN IN N GM Hybopsis amblops IN NN NN Nie Ni SIN N M Hybopsis cahni E Hybopsis d. dissimilis No Ne) NON N N N N N O Hybopsis hypsinotus So Hybopsis i. insignis N N Hybopsis i. eristigma E Hybopsis labrosa So Hybopsis monacha E Hybopsis storeriana N N N NON) ANE Ne EN N GM

Hybopsis x-punctata N M

32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

Habitat

Lowland Upland Montane

Big River Stream Creek Peedee

Cape Fear Neuse

Tar

Roanoke James

York Rappahannock Potomac Susquehanna

x * Z, Z,

Hybopsis sp. cf. zanema Leuciscus idus I Nocomis biguttatus

Nocomis effusus

Nocomis |. leptocephalus

Nocomis 1. bellicus

Nocomis |. interocularis

Nocomis micropogon

Nocomis platyrhynchus

Nocomis raneyi

Notemigonus crysoleucas XxX Notropis albeolus

Notropis alborus

Notropis altipinnis 4

Notropis amnis xX XxX Notropis amoenus xX

Notropis analostanus

Notropis ardens

Notropis ariommus

Notropis atherinoides XxX Notropis baileyi

Notropis bellus

Notropis bifrenatus XxX Notropis blennius xX X Notropis boops

Notropis buchanani xX Notropis camurus x Notropis cerasinus

Notropis chalybaeus Xx Notropis chiliticus

Notropis chrysocephalus

Notropis coccogenis

Notropis cornutus

Notropis c. cummingsae xX Notropis d. dorsalis

Notropis emiliae XxX Notropis fumeus XxX Notropis galacturus

Notropis heterodon

Notropis heterolepis

Notropis hudsonius 4 Notropis leuciodus

Notropis lirus

Notropis lutipinnis

Notropis lutrensis XxX Notropis maculatus x Notropis mekistocholas

Notropis niveus

Notropis petersoni xX

IP. 7

x KK mK XK Z Z Z Z Z Z Z

x KX x x XK

NI

~ KKK KKK KK KK KKK KK mK KK Ze Z, Z Z Z,

xx KK ZZ,

YL, by Fy TA Zi ZL LL,

NI

<> > > > > mM td tO s >< >< Z Zz A WR We

x KK x x x KK XK

IP IP

xx Kx x Kx xx KK xx KKK ZA Zz LZ, eZ ZZZ ZZ

xe KKK KKK CK * xx KK OK FLT 5p TS

x x xX

x x * KKK mK KK Km KK KK ~

Pi, 4 74, ZA Nos) ZZ ZA Z

VOLUME 95, NUMBER 1 33

Table 1.—Continued.

Drainage occurrence

Ohio Basin = = : s Be tee SS oa a > distribution Es ee Bie = Ss SS 0) a) 53 a OReery ne e ee eu ey, a ic eae S a a a 5 5 > Ss SI 7) 2 ee i Wl ee ete FP SP Bee ine a x) s x 3 n as} £ © ro as) < S 3 SS se om a SRO, Gh BE ee Er eee. Nae, OMe i OMA a hy ms | ie < ea) Hybopsis sp. cf. zanema Leuciscus idus Nocomis biguttatus NI M Nocomis effusus N N N Nocomis |. leptocephalus N So Nocomis 1. bellicus N G Nocomis |. interocularis 2? SO Nocomis micropogon IN WEN? ON NP NEN UN Ni Ne SNiaeNo O Nocomis platyrhynchus E Nocomis raneyi Notemigonus crysoleucas N Nee NN > N) ¥elyeN) SNoSomGM Notropis albeolus N Notropis alborus So Notropis altipinnis So ‘Notropis amnis N N N GM Notropis amoenus No Notropis analostanus No Notropis ardens N NE INE ON) IN UN oNiN O Notropis ariommus N N N IND INP INT N O Notropis atherinoides N N NN NG Ny ON ON EN NE NogGivi Notropis baileyi N G Notropis bellus N G Notropis bifrenatus No Notropis blennius N N NN ENON UN @N N M Notropis boops NN NN N M Notropis buchanani N N NP NG SNew No IN N GM Notropis camurus N Notropis cerasinus N Notropis chalybaeus NoSo GM Notropis chiliticus IP Notropis chrysocephalus ING) NING LIBS ONO Ne ON UN ON Ae aNaeN GM Notropis coccogenis IP N So Notropis cornutus N N No M Notropis c. cummingsae So G Notropis d. dorsalis N Notropis emiliae N N N So GM Notropis fumeus N N N GM Notropis galacturus NI N N IP ON M Notropis heterodon No Notropis heterolepis N No M Notropis hudsonius N IP NoSo M Notropis leuciodus IP N N N So O Notropis lirus N G Notropis lutipinnis IP So Notropis lutrensis Notropis maculatus So GM Notropis mekistocholas Notropis niveus So Notropis petersoni So G

SSS

34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

a0) p g, ot feb} >

Lowland Upland Montane

Big River Stream Creek Peedee

Cape Fear Neuse

Tar

Roanoke James

York Rappahannock Potomac Susquehanna

x

Notropis photogenis Notropis procne procne Notropis p. longiceps Notropis pyrrhomelas Notropis rubellus Notropis rubricroceus XxX Notropis scabriceps Notropis scepticus Notropis semperasper Notropis shumardi X XxX Notropis spectrunculus

spectrunculus xX Notropis spectrunculus subsp. xX X= Notropis spilopterus

spilopterus Notropis stilbius

x x xx KK XK x x

No Nx N § NeeN

x «x x *

x KKK KKK KK ~

* *

IP N N

x x

Notropis stramineus stramineus Notropis telescopus Notropis umbratilus

cyanocephalus XxX Notropis volucellus Notropis whipplei Notropis sp. (paleband shiner) Notropis sp. (sawfin shiner) Notropis sp. (longirostris

group) xX Phenacobius crassilabrum XxX Phenacobius mirabilis Xx Phenacobius teretulus Phenacobius uranops Phoxinus cumberlandensis Phoxinus eos Phoxinus erythrogaster Phoxinus o. oreas Phoxinus o. subsp. Pimephales notatus Pimephales promelas x Pimephales vigilax XxX Rhinichthys atratulus Rhinichthys cataractae Semotilus atromaculatus Semotilus corporalis Semotilus lumbee N N Semotilus margarita Tinca tinca

xx x x * ~ ~*~

N N N NI IP

x KKK XK ~ x «KKK XK

x KK * x KKK XK

IP N N NN eo eNeeIe Sse

» OK XE Xi | J HZ HZ

x x > x rg

ING IN NG INE IN

> > > dO OOM Med i TAL VA PATA TAA ZZ PANERA ez ee

* * ~ = Z

CATOSTOMIDAE

Carpiodes carpio Xx XxX Carpiodes cyprinus XOX Xx N N N N N

Ss

VOLUME 95, NUMBER 1 35

Table 1.—Continued.

Drainage occurrence

Ohio Basin Z i = z Native 2 & ae 2 i: ee [) a pe) fas} o mo) ao) (oe) aoe; ae Sp. eS > ise dae Mee eeu ae see ee ee ee ae ee 2 gr Vere ee es es Se ier aes ee ee ee age eee OR eae eS ver vase ety SOT Bovey sa eh AG a 2 > ete ute RSs ICO EET EA RS) MOU Oy < za Notropis photogenis ING NC ONT) UNE ND ON’ ONL EN” ZN ON: N O Notropis procne procne No Notropis p. longiceps IP So Notropis pyrrhomelas So Notropis rubellus Ne NS Ne N N N N*N N Ne Ne eNe eNoseeM Notropis rubricroceus IP N So Notropis scabriceps E Notropis scepticus So Notropis semperasper Notropis shumardi N NP GM Notropis spectrunculus spectrunculus E Notropis spectrunculus subsp. E Notropis spilopterus spilopterus NEN ONE NON N: N ON GNU UN WN, 2Nieg, NOM Notropis stilbius N GM Notropis stramineus stramineus Ni UN. UNG ON: IN’ oN. UN: ON N N M Notropis telescopus Ieee N M Notropis umbratilus cyanocephalus N NN, ON oN ON- ON N M Notropis volucellus NOON =Ne UN UN ON ON: ON) FN EN. GN eN GM Notropis whipplei N N IN; ANG INP AN UN?) AN Nie N GM Notropis sp. (paleband shiner) N N Notropis sp. (sawfin shiner) N N Notropis sp. (longirostris group) N M Phenacobius crassilabrum E Phenacobius mirabilis N NN NTN 9 N] N N GM Phenacobius teretulus E Phenacobius uranops N N N Phoxinus cumberlandensis Ja. 7-18 Phoxinus eos No M Phoxinus erythrogaster N Ny aN UN NT UN] eNe ON UN M Phoxinus o. oreas N IP Phoxinus o. subsp. E Pimephales notatus Ne NS UNT EN ON ON? OND ON ENN UN NP Pe Nowwem™ Pimephales promelas I i TPs. ff NN NON ON | NING S| NortGM Pimephales vigilax N N NOON EN UN? UN NPN GM Rhinichthys atratulus N NS ON N NN N NN NO NN NoSoF mM Rhinichthys cataractae N N N N NoSo O Semotilus atromaculatus NEN Ne ON NOON ON NON N NN (NOENoSoUGM

Semotilus corporalis No Semotilus lumbee

Semotilus margarita N No M Tinca tinca

CATOSTOMIDAE Carpiodes carpio NS Ne Ne SIN] ON N GM Carpiodes cyprinus NNN NEN NG EN] NON N NoSo GM

36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Habitat Atlantic Slope

Lowland Upland Montane

Big River Stream Creek Peedee

Cape Fear Neuse

Tar

Roanoke James

York Rappahannock Potomac Susquehanna

x ~ Z

-Carpiodes velifer Catostomus catostomus Catostomus commersoni Cycleptus elongatus Erimyzon o. oblongus Erimyzon o. claviformis Erimyzon sucetta Hypentelium etowanum Hypentelium nigricans Hypentelium roanokense Ictiobus bubalus Xx Ictiobus cyprinellus x Ictiobus niger XxX Lagochila lacera Minytrema melanops xX Moxostoma anisurum XxX Moxostoma ariommum Moxostoma atripinne Moxostoma carinatum xX Moxostoma cervinum Moxostoma duquesnei Moxostoma erythrurum Moxostoma hamiltoni Moxostoma m. macrolepidotum Moxostoma m. breviceps Moxostoma pappillosum Moxostoma rhothoecum Moxostoma robustum Moxostoma rupiscartes

ICTALURIDAE

Ictalurus brunneus Ictalurus catus Ictalurus furcatus Ictalurus melas Ictalurus natalis Ictalurus nebulosus Ictalurus platycephalus Ictalurus punctatus Noturus baileyi Noturus elegans Noturus eleutherus Noturus exilis

Noturus flavipinnis Noturus flavus Noturus furiosus x Noturus gilberti Noturus gyrinus Noturus insignis

Noturus leptacanthus es

x xX x xX x x Zz, Z Z Z, Z,

Ne iN’ Ne aNeN

XxX «Ne N*IN: N- NPN ON @SNieNeN

x KK XK xxx KKK XK * xx KKK

x xX N N N N: N@eNeeNe ene aN Xx E

IP

x KX

ZZ Lae iz, ZA mM Z

xx XK

x KK MK KK KK KK KK OK OK ~ x x xX ~

* KKK mK KK KK KKK KK KK

~< Za

x «x «x «x X >< x «x x >< Spd vA Vd edz Z Z seed Ard Z Z Z Z HZ Z Zag Z

x x XK BP2Z2Z2Zz Z

IP JP IP IPP sneer

xm KKK KK KK KK mK OK *

x > x > Dede Dd DK DK DK Dd Dd Dd DK DK DK Dd De Dt Dx Zaz ZZ Zaz Zaz, Pa 74 Va ZzZZ ZZ,

VOLUME 95, NUMBER 1

Table 1.—Continued.

37

Drainage occurrence

Ohio Basin 3 ie 5 E Native a & 22 CGecutien awe Ee Bo : coe ee eo 5 ae Se CN sce Se eR > Soy 88e 08 A ee cee Se ee ee ae oe Ss 5 ¢ 2 & See ee Se Se Se Se ee aoe ee ER ops 0 Se ae ONS Suite aver ce oe OO TO oy Pe Se ai oS = & Zp ie ite Mie Oe ae Se IME Se (On Om ie < za Carpiodes velifer N N ING Ne OND OUNE. UNE N, N So GM Catostomus catostomus N No GM Catostomus commersoni N N N N N N N N N N N N NoSo M Cycleptus elongatus NP NP NP NP N NP N N N GM Erimyzon o. oblongus NoSo Erimyzon o. claviformis N N N GM Erimyzon sucetta N So GM Hypentelium etowanum NI G Hypentelium nigricans NN NN N NON N NO’ N NN ENoSoNGM Hypentelium roanokense Ictiobus bubalus N N NON NEN UN WN N GM Ictiobus cyprinellus N N N N N N GM Ictiobus niger N NESNES NS IN N GM Lagochila lacera N NP N N M . Minytrema melanops N N NC -N) SND UN UN, ON NON So GM Moxostoma anisurum N NP N N. No N UN - Ne ON NAN So M Moxostoma ariommum Moxostoma atripinne E Moxostoma carinatum N N N NN -N INO Ne N N GM Moxostoma cervinum Moxostoma duquesnei N N N NG UNE SN UN, EN aN ae Ni GM Moxostoma erythrurum IND eN SNe SSE TNT ENG ON Ne NING Ne eN GM Moxostoma hamiltoni Moxostoma m. macrolepidotum NoSo M Moxostoma m. breviceps NE NSN Ne NE ONS SN] UNG SN N O Moxostoma pappillosum So Moxostoma rhothoecum N Moxostoma robustum So Moxostoma rupiscartes So G ICTALURIDAE Ictalurus brunneus So G Ictalurus catus NoSo G Ictalurus furcatus N N N N N N N GM Ictalurus melas N N IP ND INS ENS ND END ONG EN GM Ictalurus natalis N N N NI N N N N N N N N NOoSo GM Ictalurus nebulosus N N NI N N N N IP N NoSo GM Ictalurus platycephalus So Ictalurus punctatus Ne NSN NIN Ne NSN] IN NS NEN So GM Noturus baileyi E Noturus elegans N N O Noturus eleutherus No ONe UN OND ON N M Noturus exilis N N N N M Noturus flavipinnis E Noturus flavus N N NEN NN NG Nn eN N M Noturus furiosus Noturus gilberti Noturus gyrinus NNN N NoSo GM Noturus insignis IP N IP NoSo Noturus leptacanthus NI So G

38

Table 1.—Continued.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Lowland

Habitat

Montane Big River

Stream

Creek

Drainage occurrence

Atlantic Slope

Peedee Cape Fear Neuse

Tar Roanoke

James

York Rappahannock Potomac Susquehanna

Noturus miurus Noturus nocturnus Noturus stigmosus Noturus stanauli

Noturus sp. cf. leptacanthus

Pylodictis olivaris

AMBLYOPSIDAE

Amblyopsis spelaea Chologaster agassizi Chologaster cornuta Typhlichthys subterraneus

APHREDODERIDAE

Aphredoderus sayanus

PERCOPSIDAE

Percopsis omiscomaycus

GADIDAE

Lota lota

CY PRINODONTIDAE

Fundulus albolineatus Fundulus catenatus Fundulus d. diaphanus Fundulus heteroclitus Fundulus lineolatus Fundulus notatus Fundulus olivaceus Fundulus rathbuni Fundulus stellifer Fundulus waccamensis Fundulus sp.

POECILIIDAE

Gambusia a. affinis Gambusia a. holbrooki Heterandria formosa

ATHERINIDAE

Labidesthes sicculus Menidia extensa

GASTEROSTEIDAE

Apeltes quadracus Culaea inconstans

COTTIDAE

Cottus baileyi Cottus b. bairdi

~ xX

x KK XM

x KK

x «x XK

x KK KK

x x

x KKK

xX

XxX XxX Xx

Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma N N N N N

Ma Ma Ma Ma Ma NP N

NI

Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma

Ma Ma Ma Ma

Ma Ma Ma IP IP

VOLUME 95, NUMBER 1 39

Table 1.—Continued.

Drainage occurrence

Ohio Basin | = Native Ss & & extralimital & & Ss distribution 3 2 Oo a) s s oS ae as a Oo S a) 3 oO no} "e aS) Be ee eS as > gs a 8 a ee Sa a cee ae ge aie le cae ees See Bae SS OS See ben Jer eg: SeecleS o 6k a 3 an ee) oO ¢ 5 5 5 = 2 ie Sy OMe EM tS) a ES aN SS Oy) a < a] Noturus miurus N N N N -Ne oN UN ON. ON? NTN GM Noturus nocturnus N N N N N GM Noturus stigmosus N Ni SNE NF =e N: M Noturus stanauli E Noturus sp. cf. leptacanthus Pylodictis olivaris Ne, ONY ON® ON: ONS ON: ON GN: UN] MINETIENE WAN GM AMBLYOPSIDAE Amblyopsis spelaea N O Chologaster agassizi N N N M Chologaster cornuta So Typhlichthys subterraneus N N N GM APHREDODERIDAE Aphredoderus sayanus N N N NoSo GM PERCOPSIDAE Percopsis omiscomaycus Nei N UN N N N N No M GADIDAE Lota lota NI NI No M CY PRINODONTIDAE Fundulus albolineatus E Fundulus catenatus N N N M Fundulus d. diaphanus I No Fundulus heteroclitus Fundulus lineolatus So G Fundulus notatus IN| IN| INT INT N GM Fundulus olivaceus N N GM Fundulus rathbuni So Fundulus stellifer IP G Fundulus waccamensis Fundulus sp. N N POECILIIDAE Gambusia a. affinis IP IP Ma Ma IP Ma GM Gambusia a. holbrooki NoSo G Heterandria formosa So G ATHERINIDAE Labidesthes sicculus N N N I N N N N N N N N So GM Menidia extensa GASTEROSTEIDAE Apeltes quadracus Culaea inconstans I COTTIDAE Cottus baileyi E

Cottus b. bairdi NN NaN ENE NS UNG EN N N No M

40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

en) ist) a = ped} oo

Lowland Upland Montane

Big River Stream Creek Peedee

Cape Fear Neuse Roanoke James

York Rappahannock Potomac Susquehanna

Tar

Cottus bairdi subsp. Cottus c. carolinae Cottus carolinae subsp. Cottus cognatus Cottus girardi

Cottus sp. (smoky sculpin) Xx Cottus sp. E

PERCICHTHYIDAE

Morone americana Morone chrysops Morone mississippiensis Morone saxatilis

CENTRARCHIDAE

Acantharcus pomotis Ambloplites cavifrons Xx Ambloplites r. rupestris en XN EX Centrarchus macropterus Elassoma evergladei Elassoma zonatum Elassoma sp.

Elassoma sp. Enneacanthus chaetodon Enneacanthus gloriosus Enneacanthus obesus Lepomis auritus

Lepomis cyanellus Lepomis gibbosus Lepomis gulosus

Lepomis humilis

Lepomis macrochirus Lepomis marginatus Lepomis megalotis Lepomis microlophus Lepomis punctatus Micropterus coosae Micropterus d. dolomieui Micropterus p. punctulatus Micropterus s. salmoides Pomoxis annularis Pomoxis nigromaculatus

PERCIDAE

Ammocrypta asprella

Ammocrypta clara

Ammocrypta pellucida

Ammocrypta vivax xX Etheostoma acuticeps

Etheostoma asprigene XxX

xx «KK x KK KK x KK KK

ZZ,

Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma I I I

xx x x x KK XK x KX

Ma Ma Ma Ma Ma Ma Ma Ma Ma Ma

* * Pd Z Za, Z

NN Ne UN

IP br area aT

x KK KK Z2ZZ5 ZZZ

Z Z Zi Z, Le Lie Za eZ,

x KKK KX KX > ZA ZZe AZ Vd WA Na VD Wa Vey 7 ZZ ZZ ZZ DID IZ 2 ZZ, Z Diva 2 Zane DD i ZZ ZL ad, ZZ4ZEZz BAe ZZZ Z,

x xX xX xx x KK ZZ ZZ Z5 ZH 5 5 cs cs 5 ty Z22ZHzZ

x x

NG NEI iP I ot

x xX x mK K mK KKK KKK KK

x xX xX xxx KX xx x x xxx KX

Ze

Zz,

Z,

Z

— Ll —_

Zaz —_ — —_ —

IP 1 ie eee

xx KK XK x KKK XK x KKK mK

VOLUME 95, NUMBER 1

Table 1.—Continued.

Drainage occurrence

4]

Ohio Basin ez) 2) = = Native ao as FS re extralimital > ‘is iS) is distribution ES ES ie 3a Te Sig, oo eat eae 2 ¢ é oes ee Wet Gee ee > SS elm Comey (2 Sy uN ME ee a ge Gen iS i es ae £2) on ee oe 80 a a a wa os = 5 2 24 2 e ES Se cae ee en Cae Came Ome oie cag caoey eee Pree Ge a OD ASS PE MRS BO TO) Ge < za Cottus bairdi subsp. N N Cottus c. carolinae NON SN N M Cottus carolinae subsp. N N Cottus cognatus No Cottus girardi Cottus sp. (smoky sculpin) N So Cottus sp. PERCICHTHYIDAE Morone americana NoSo Morone chrysops I NI NI IP INT N Pai Nee Nie) N GM Morone mississippiensis N N GM Morone saxatilis I I I I I I NoSo G CENTRARCHIDAE - Acantharcus pomotis NoSo G Ambloplites cavifrons Ambloplites r. rupestris IND IN SOND CERT IN IN} ING? ONG ON] SNR NS NT} BINom eM Centrarchus macropterus N N N So M Elassoma evergladei So Elassoma zonatum N N So GM Elassoma sp. E Enneacanthus chaetodon NoSo Enneacanthus gloriosus NoSo G Enneacanthus obesus NoSo Lepomis auritus IP I I I I NoSo G Lepomis cyanellus Ni OND SUNG) IPSN) ONE NE Ne) INT ING NON GM Lepomis gibbosus NI NIP IP IP IP NoSo M Lepomis gulosus NIP NN INS NG Se IN Som Givi Lepomis humilis te N N N N NON GM Lepomis macrochirus INS ING? NPS SINS ING IN) ON) © ONG NaN EN, So GM Lepomis marginatus N So GM Lepomis megalotis ING NING MT uN INS BING INI TINGS ING ee IN GM Lepomis microlophus IP [Pee NEN N So GM Lepomis punctatus N N So GM Micropterus coosae I G Micropterus d. dolomieui Ne ONG UNG TP? NY ON i DINO ONO ONG) FING goNfPr SiN! M Micropterus p. punctulatus INN, ENG INTIS IN| OND Nite Ny ONS Ne eNP IN M Micropterus s. salmoides Ne Ne ONY aR” UN: UN ON” UN) ON)’ UN) ee NIPSN So GM Pomoxis annularis NY INN IP NU SINY ING, UN IN: GN ANTS N GM Pomoxis nigromaculatus NON EN DP N N N N N N So GM PERCIDAE Ammocrypta asprella NN GM Ammocrypta clara ING N N GM Ammocrypta pellucida N’ ONY UN Ne Ne aNaeN) SN EN M Ammocrypta vivax N GM Etheostoma acuticeps E Etheostoma asprigene NigEN N GM

42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

Habitat

Lowland Upland Montane

Big River Stream Creek Peedee

Cape Fear Neuse Roanoke James

York Rappahannock Potomac Susquehanna

Tar

Etheostoma aquali Etheostoma atripinne Etheostoma barbouri Etheostoma bellum Etheostoma blennioides blennioides Etheostoma b. gutselli Etheostoma b. newmanii Etheostoma b.: newmanii X blennioides Etheostoma b.: n. X gutselli Etheostoma blennius Etheostoma boschungi Etheostoma caeruleum xX Etheostoma camurum xX Etheostoma chlorobranchium xX xX X Xx

x eM KKK XK xx x xx KK x KX

~ x *

NI

x KK XK

Etheostoma chlorosomum xX Etheostoma cinereum Etheostoma collis collis XxX Etheostoma c. lepidinion xX Etheostoma duryi Etheostoma etnieri Etheostoma flabellare Etheostoma fusiforme fusiforme Etheostoma f. barratti Etheostoma gracile Etheostoma histrio Etheostoma jessiae Etheostoma kanawhae Etheostoma kennicotti Etheostoma longimanum Etheostoma luteovinctum Etheostoma m. maculatum Etheostoma m. sanguifluum Etheostoma m. vulneratum Etheostoma mariae xX Etheostoma meadiae Etheostoma microlepidum Etheostoma neopterum x Etheostoma n. nigrum Etheostoma n. susanae Etheostoma n.: nigrum X

susanae Etheostoma obeyense Etheostoma olivaceum Etheostoma o. olmstedi xX Etheostoma o.: o. X

atromaculatum N Etheostoma o. atromaculatum xX X Xe 2X IN| INE INP INP INT IN

x KK xx KKK x x xX xm mK KK KKK XK Za Z, Z, Z,

x x

KKK KKK Km KK KK

N “Ne END aN

Km KKK KKK mK KK KK OK

x x x x

xx KX x x

N NN “N@N

VOLUME 95, NUMBER 1

Table 1.—Continued.

43

Drainage occurrence

Ohio Basin

Kanawha: below falls Kanawha: above falls

Monongahela Little Kanawha Guyandotte

Big Sandy Licking

Kentucky

Green

Cumberland: below falls

Cumberland: above falls

Tennessee

Native extralimital distribution

Atlantic Slope Elsewhere

Etheostoma aquali Etheostoma atripinne Etheostoma barbouri Etheostoma bellum Etheostoma blennioides blennioides Etheostoma b. gutselli Etheostoma b. newmanii Etheostoma b.: newmanii x blennioides Etheostoma b.: n. X gutselli Etheostoma blennius Etheostoma boschungi Etheostoma caeruleum ‘Etheostoma camurum Etheostoma chlorobranchium Etheostoma chlorosomum Etheostoma cinereum Etheostoma collis collis Etheostoma c. lepidinion Etheostoma duryi Etheostoma etnieri Etheostoma flabellare Etheostoma fusiforme fusiforme Etheostoma f. barratti Etheostoma gracile Etheostoma histrio Etheostoma jessiae Etheostoma kanawhae Etheostoma kennicotti Etheostoma longimanum Etheostoma luteovinctum Etheostoma m. maculatum Etheostoma m. sanguifluum Etheostoma m. vulneratum Etheostoma mariae Etheostoma meadiae Etheostoma microlepidum Etheostoma neopterum Etheostoma n. nigrum Etheostoma n. susanae Etheostoma n.: nigrum X susanae Etheostoma obeyense Etheostoma olivaceum Etheostoma o. olmstedi Etheostoma o.: 0. X atromaculatum Etheostoma o. atromaculatum

N°: AN GN ON, ON, No ON

ZZ, ZZ, ZZ,

NO UND ENG oN” oN, Ne ON

Ne NENG NG UN oN UN

esies

ZZ,

ZZ

Zm Zw

74, 74 \o| 74 74 Jeal teal leo

Os

GM

So

No GM

oT PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

Habitat

Lowland Upland Montane

Big River Stream Creek Peedee

Cape Fear Neuse

Tar

Roanoke James

York Rappahannock Potomac Susquehanna

Etheostoma o.: o. X vexillare Etheostoma o. vexillare Etheostoma o. maculaticeps Xx Etheostoma osburni Etheostoma parvipinne xX Etheostoma perlongum xX E Etheostoma podostemone Etheostoma proeliare Xx Etheostoma rufilineatum Etheostoma s. sagitta Etheostoma sagitta spilotum Etheostoma sellare Etheostoma serriferum x Etheostoma simoterum Etheostoma smithi Etheostoma s. spectabile Etheostoma squamiceps Etheostoma striatulum Etheostoma stigmaeum Etheostoma swaini XxX Etheostoma swannanoa xX Etheostoma tippecanoe Etheostoma tuscumbia Etheostoma variatum Etheostoma virgatum Etheostoma vitreum Etheostoma z. zonale Etheostoma sp. (duskytail darter) XxX X Etheostoma sp. (Elk darter) Etheostoma (Ulocentra) sp. A—emerald darter Etheostoma (Ulocentra) sp. B Etheostoma (Ulocentra) sp. C Etheostoma (Ulocentra) sp. D—golden snubnose darter X xX OK Etheostoma (Ulocentra) sp. E—(Green River) Etheostoma (Ulocentra) sp. F—(Barren River) splendid darter Perca flavescens XxX Percina aurantiaca Percina burtoni Percina c. caprodes Percina c. semifasciata Percina copelandi Percina crassa Percina e. evides Percina evides subsp. XxX

Za Zz

N- N

x KKK KK x KK x

x KK xx KX

N NN Nie N

xx KK KK x KKK KKK KX ~< KK KK

x mK mK

x Kx x

N -N N N GNNeN

x KK KK ~ x Km

x *

x * x x

x ~ *

NI NI NI NI NI NI NI NI NI NI

x KKK KK XX ~ x x xX

KKK KK KK XK Z Z

VOLUME 95, NUMBER 1 45

Table 1.—Continued.

Drainage occurrence Ohio Basin Native

extralimital distribution

Kanawha: below falls Kanawha: above falls Cumberland: below falls Cumberland: above falls

Monongahela Little Kanawha Guyandotte Big Sandy Licking Kentucky Tennessee Atlantic Slope Elsewhere

Green

Etheostoma o.: 0. X vexillare Etheostoma o. vexillare Etheostoma o. maculaticeps So Etheostoma osburni E Etheostoma parvipinne N Etheostoma perlongum Etheostoma podostemone Etheostoma proeliare N Etheostoma rufilineatum N N Etheostoma s. sagitta E Etheostoma sagitta spilotum E Etheostoma sellare Etheostoma serriferum So Etheostoma simoterum NI NI Etheostoma smithi Etheostoma s. spectabile NY OEN ON: Etheostoma squamiceps N Etheostoma striatulum Etheostoma stigmaeum N N Etheostoma swaini Etheostoma swannanoa Etheostoma tippecanoe N N NN N' 'N Etheostoma tuscumbia Etheostoma variatum Ne ON N N N N N O Etheostoma virgatum 18, Etheostoma vitreum No Etheostoma z. zonale NGS INGeN INGE INGOUN) NG ENE ON Etheostoma sp. (duskytail darter) N Etheostoma sp. (Elk darter) Etheostoma (Ulocentra) sp. N N N A—emerald darter Etheostoma (Ulocentra) sp. B N Etheostoma (Ulocentra) sp. C Etheostoma (Ulocentra) sp. D—golden snubnose darter N N Etheostoma (Ulocentra) sp. E—(Green River) E Etheostoma (Ulocentra) sp. F—(Barren River) splendid E darter Perca flavescens NI ree We Percina aurantiaca Percina burtoni N Percina c. caprodes Nae NMaINS ANE NG ON NEDIN SEN) <iN Percina c. semifasciata No Percina copelandi N NON IN IND INGEN N Percina crassa So Percina e. evides N NN ee NF) NL aN N Percina evides subsp. E I ne ee ee he a

ZZz ors

GM GM

Ze t ZeZ to ZeZ ZZ,

eal 74 4

Fz, = o

No

472 foal <

SB Ss

46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Drainage occurrence

Atlantic Slope

Habitat ~ =

E 2 5 . E a4 o 2 oe 5 ea = 2 3 Be a: BS es ith Bie Ta RB es oh a a pS) Rae Ot ab OZ ee 2S eee

Percina gymnocephala x. EX X

Percina macrocephala Xx xX xX

Percina maculata x > ia. <

Percina n. notogramma x Xx Ne Ne aN

Percina n. montuosa x x E

Percina ouachitae x Xx Xo xX

Percina oxyrhyncha xX UX PS

Percina p. peltata Xx 4 NN OSNY SIN SN

Percina p. nevisense Xx XxX Ni“ .N; =N

Percina peltata subsp. Xx x E

Percina phoxocephala Xx Xx

Percina rex Xx Xx E

Percina roanoka x x NG N= IN Ni

Percina s. sciera Xr eX MK IN

Percina shumardi DS DS XxX

Percina squamata 2h DK Xx

Percina tanasi x xX

Percina (Odontopholis) sp. XxX Xx

Stizostedion canadense Xr OX x

Stizostedion v. vitreum XxX Xee GX! IP NI NI NI Ih IN

SCIAENIDAE Aplodinotus grunniens Xe XxX XxX

Literature Cited

Hambrick, P. S., C. H. Hocutt, M. T. Masnik, and J. H. Wilson. 1973. Additions to the West Virginia ichthyofauna with comments on the distribution of other species.—Proceedings of the West Virginia Academy of Sciences 45:58—60.

Hendricks, M. L., J. R. Stauffer, Jr., C. H. Hocutt, and C. R. Gilbert. 1979. A preliminary checklist of the fishes of the Youghiogheny River.—Natural History Miscellanea 203:1-15.

Hocutt, C. H., R. F. Denoncourt, and J. R. Stauffer, Jr. 1978. Fishes of the Greenbrier River, West Virginia, with drainage history of the southern Appalachians.—Journal of Biogeography 5:59-80.

——_., , and . 1979. Fishes of the Gauley River, West Virginia.—Brimleyana 1:47-80.

, and P. S. Hambrick. 1973. Hybridization between the darters Percina crassa roanoka and

Percina oxyrhyncha (Percidae, Etheostomatini), with comments on the distribution of Percina

crassa roanoka in New River.—American Midland Naturalist 90:397—405.

; , and M. T. Masnik. 1973. Rotenone methods in a large river system.—Archiv fur

Hydrobiologia 72:245—252.

Jenkins, R. E., E. A. Lachner, and F. J. Schwartz. 1972. Fishes of the central Appalachian drainages: their distribution and dispersal, pp. 43-117. In: P. C. Holt (ed.). The distributional history of the biota of the southern appalachians. Part III: Vertebrates.—Research Division Monographs 4, Virginia Polytechnic Institute and State University. Blacksburg, Virginia.

VOLUME 95, NUMBER 1 47

Table 1.—Continued.

Drainage occurrence

Ohio Basin = = Native = S & = extralimital Se 2 > distribution Et Sela o «68 3 = 3 5 a < 2. oR a) es} o as) as) 2 = peat ake Mey > pa Agari oO a 2 oD vy =| a Eo] = an Ts 5 5 A — r 5 o 2 2 = w S 2 5 2 2 2 = es Be i= S > if ® = = I 5 es Oe ee 3 a | wo 88 ) 2 5 =| O s 2 Siete ee Os PE Se a OE eee. ie < ea] Percina gymnocephala E Percina macrocephala N N N NPN UN N N O Percina maculata N N N NPN NG NNN ON oN GM Percina n. notogramma No Percina n. montuosa Percina ouachitae N N GM Percina oxyrhyncha INN ONE SUN ENS NY ONS SIN? N Percina p. peltata No Percina p. nevisense Percina peltata subsp. Percina phoxocephala Nee Ne) N M Percina rex Percina roanoka IP Percina s. sciera N N Ne Ne NE Ne NN N GM Percina shumardi ING Ne NaeN N GM Percina squamata N N Percina tanasi E Percina (Odontopholis) sp. N N Stizostedion canadense N N N N NP N N UN N GM Stizostedion v. vitreum N INN NiN| Nie NIE NN N GM

SCIAENIDAE Aplodinotus grunniens N N N ING SuNi DAN INE SNL UN: N GM

Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr. 1980. Atlas of North American freshwater fishes.—North Carolina State Museum of Natural History, Raleigh, N. C. 854 pp.

Stauffer, J. R., Jr., K. L. Dickson, J. Cairns, Jr., and D. C. Cherry. 1976. The potential and realized influences of temperature on the distribution of fishes in the New River, Glen Lyn, Virginia.— Wildlife Monographs 50: 1-40.

—, C. H. Hocutt, and D. S. Lee. 1978. The zoogeography of the freshwater fishes of the Potomac River basin, pp. 44-54. In: K. C. Flynn and W. T. Mason (eds.). The freshwater Potomac: aquatic communities and environmental stresses.—Interstate Commission Potomac River Ba- sin. Rockville, Maryland.

——,, , M. T. Masnik, and J. E. Reed, Jr. 1975. The longitudinal distribution of the fishes of the East River, West Virginia-Virginia.—Virginia Journal of Science 26:121-125.

(JRS) Appalachian Environmental Laboratory, Center for Environmental and Estuarine Studies, University of Maryland, Frostburg State College Campus, Frostburg, Maryland 21532; (BMB) Department of Zoology, Southern Illinois University, Carbondale, Illinois 62901; (REJ) Department of Biology, Roanoke College, Salem, Virginia 24153; (CHH) Horn Point Environmental Laboratories, Center for Environmental and Estuarine Studies, University of Maryland, Cam- bridge, Maryland 21613.

PROC. BIOL. SOC. WASH. 95(1), 1982, pp. 48-57

TWO NEW GENERA OF DEEP-SEA POLYCHAETE WORMS OF THE FAMILY AMPHARETIDAE AND THE ROLE OF ONE SPECIES IN DEEP-SEA ECOSYSTEMS

Robert Zottoli

Abstract.—Two new ampharetid genera, Decemunciger and Endecamera, each with one new species, D. apalea and E. palea, are described from wood panels placed on the deep-sea floor by Turner (1973). The role of Decemunciger in deep- sea ecosystems is discussed.

Recently, I examined a collection of ampharetid polychaetes removed from pieces of wood collected by R. D. Turner, using the submersible DSRV Alvin, from four experimental bottom stations in the North Atlantic, at depths ranging from 1830 to 3995 meters. The wood had been placed at these sites by Turner to study molluscan wood borers, and to “‘test the hypothesis that wood is an im- portant source of nutrients and contributes to diversity in the deep sea’’ (Turner 1977:18).

Bivalve molluscs of the subfamily Xylophagainae (Family Pholadidae, Genera Xyloredo and Xylophaga) mechanically excavate burrows in wood (Turner 1973, 1977). The bivalves ingest wood particles, making wood by-products available within fecal pellets as food for detritus consumers. Adult and juvenile bivalves may be consumed by a variety of predators such as galatheid crabs. Turner (1977) suggests that young galatheid crabs may feed on recently settled Xylophaga lar- vae and, later, on larger invertebrates including adult borers. Stomach contents of older crabs contained sponge spicules, small pieces of wood, a nematode and polychaete setae (Turner 1977). Other groups in this deep-sea food chain include several families of polychaete worms, brittlestars, small sea urchins and predatory gastropods (Turner 1977).

Two new genera from the family Ampharetidae, each with one new species, have been found among the ampharetids associated with Turner’s wood panels. The external anatomy of these ampharetids, Decemunciger apalea n. sp. and Endecamera palea n. sp., is described, followed by a discussion of the role of the former species in deep-sea ecosystems.

Materials and Methods

Three experimental islands, each with 12 separate one foot cubes of spruce wood were placed by Turner (1977), for a period of five years, at the following locations:

1. Deep Ocean Station 1 (DOS-1), 39°46’N, 70°41’W, 110 miles south of Woods Hole, Mass., in 1830 m.

2. Deep Ocean Station 2 (DOS-2), 38°18.4’N, 69°35.6’W, 190 miles southeast of Woods Hole, Mass., in 3506 m.

VOLUME 95, NUMBER 1 49

3. Tongue of the Ocean, Bahama Islands (TOTO Tower 3), 24°53.2'N, 77°40.2'W, in 2066 m.

Each experimental island is encircled by wood panels, 24” x 5” x 1”, which are removed and replaced each time the islands are visited. At the time of re- trieval, panels are enclosed in mesh bags to prevent loss of crumbled wood and specimens. The mesh bags and their contents are then placed in retrieval boxes, carried on the DSRV Alvin basket. The contents of the bags may be preserved at the time the boxes are closed for return to the surface by puncturing plastic bags previously placed in the retrieval boxes, thus releasing gluteraldehyde. Al- ternatively, the contents in certain cases may be preserved immediately upon reaching the surface. Specimens were well preserved. The use of submersibles in studies of benthic communities is described by Grassle (1980a).

Systematics

Ampharetid polychaetes generally are wide anteriorly, tapering gradually to- wards the posterior end (Fig. 1A). The prostomium is generally trilobed. Seg- ments | and 2, which lie immediately behind the prostomium, are fused in most species, and ventrally form the lower lip. Segment 3, in some species, bears one lateral bundle of paleal setae on each side (Fig. 2A). The thorax begins at segment 4. The above segmental numbering system is that of Malmgren (1865-1866) and Fauvel (1927), who recognized two segments in front of the paleal.

Decemunciger, new genus

Type-species.—Decemunciger apalea n. sp. Gender, masculine.

Diagnosis.—Body short, of 13 thoracic setigerous segments, last 10 unciniger- ous, and of 14 abdominal uncinigerous segments. Segments | and 2 fused ventrally forming lower lip. Segment 3 lacking paleae. Prostomium lacking glandular ridges. Smooth, ventrally grooved, oral tentacles. Four pairs of smooth branchiae on dorsal surface of segments 3-5. Branchial groups separated mid-dorsally by nar- row space. Abdominal notopodia and notopodial and neuropodial cirri absent.

Remarks.—In comparison with other ampharetid genera with a similar distri- bution of uncinigerous thoracic segments, Decemunciger differs from Melinnata (Hartman, 1965), Melinnopsides (Day, 1964) and Muggoides (Hartman, 1965), among other characters, by having 4 rather than 3 pairs of branchiae; from Mel- innopsis (McIntosh, 1885) by lacking a fleshy ridge across dorsal surface of seg- ment 6; and from Mexamage (Fauchald, 1972) by having 4 pairs of branchiae inserted on 3 successive segments rather than 4.

Etymology.—Generic name derived from the Latin, refers to number of tho- racic uncinigerous segments.

Decemunciger apalea, new species Fig. 1A—C Material examined.—(asb = asbestos-backed panel; D = DSRV Alvin).

Description.—Maximum size 6.3 mm long, 0.9 mm wide. Sexually mature in- dividuals as small as 3.6 mm long, 0.54 mm wide. Holotype complete, 4.7 mm

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table of Material examined:

No. of specimens

Time Panel Submerged Removed submerged Adult Juveniles DOS-1 N28 8-30-75(D597) 9-28-77(D794) 2 yrs ) 1 N34 (asb) 8-30-75(D597) 7-29-77(D773) 2 yrs 2 32 N35 8-30-75(D597) 9-18-78(D834) 3 yrs 2 4 N47 6-15-76(D658) 7-30-77(D774) l yr 0 3 N65 8-17-76(D685) 7-29-77(D773) | yr 0 13 N67 8-17-76(D685) 8- 1-77(D776) 1 yr 0 7 N72 7-29-77(D773) 9-18-78(D834) 1 yr 0 3 N91 (asb) 9-28-77(D794) 9-18-78(D834) | yr ] 1 DOS-2 N31 9- 5-75(D601) 8- 3-77(D777) 2 yrs 16 10 Holotype (USNM #71545) and 4 paratypes (USNM #71546) TOTO TOWER 3 T56 (asb) 5-12-77(D755) 11-11-78(D851) 1 yr 0 1

long, 0.9 mm wide. Color in alcohol white to pale orange. Prostomium indistinctly trilobed, lacking glandular ridges. About 14 smooth oral tentacles, each with deep ventral groove. Segments | and 2 fused, ventral part forming lower lip. Segment 3 without paleae and not visually obvious. Four pairs of smooth branchiae, about ’% body length; 2 on segment 3, 1 on segment 4, and | on segment 5. Branchial groups separated mid-dorsally by narrow space. Notopodial lobes each bearing 7-11 winged capillary setae from segments 4—16. Each seta about 0.4 mm long, 7.5 wm wide basally, and 10 wm wide across the blade. Notopodia lobes (unci- nigerous pinnules) bearing toothed uncini from segment 7 to end of abdomen. Ten thoracic and 14 uncinigerous abdominal segments. Thoracic uncini in single transverse rows, 22-35 per row. Each with about 10 teeth, more or less in 3 transverse rows, above a rounded basal prow (Fig. 1B). Abdominal uncini in single tranverse rows, about 10-18 per row. Each with about 15 teeth in several transverse rows, above a rounded basal prow (Fig. 1C). Abdominal notopodia and notopodial, neuropodial, and anal cirri lacking. Pygidium rounded.

Remarks.—Mucus-lined tubes covered with particulate matter, about 3 times worm length. Female about 4 mm long and 0.5 mm wide with approximately 260 elliptical eggs in body cavity ranging from 25 to 150 wm across widest diameter. No gonoducts visible.

Etymology.—Specific name, derived from the Latin, refers to lack of paleae.

Endecamera, new genus

Type-species.—Endecamera palea n. sp. Gender, feminine.

Diagnosis.—Body short, of 14 thoracic setigerous segments, last 11 unciniger- ous and of 14 abdominal uncinigerous segments. Segments | and 2 fused, ventrally forming lower lip. Paleae present on segment 3. Prostomium lacking glandular ridges. Smooth, ventrally grooved, oral tentacles. Four pairs of smooth branchiae

VOLUME 95, NUMBER 1 51

A

Fig. 1. Decemunciger apalea: A, Lateral view of entire worm, 5.2 mm long; B, Mid-thoracic uncini, lateral and frontal views, length = 16 wm; C, Mid-abdominal uncini, lateral and frontal views, length = 14 um.

a2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

on dorsal surface of segments 3-4. No mid-dorsal space between branchial groups. Abdominal notopodia and notopodial and neuropodial cirri absent.

Remarks.—In comparison with other ampharetid genera with a similar distri- bution of thoracic uncinigerous segments, Endecamera differs from Ampharana Hartman, 1967; Amythasides Eliason, 1955, and Glyphanostomum Levinsen, 1884, by having 4 rather than 3 pairs of branchiae; from Amage Maimgren, 1865-1866, Grubianella McIntosh, 1885, and Phyllampharete Hartman and Fau- chald, 1971, by lack of abdominal notopodia; from Pterampharete Augener, 1918, and Sabellides Milne Edwards in Malmgren, 1865-1866, by having smooth rather than papillose oral tentacles, and from Paramage Caullery, 1944, by notosetae beginning on segment 4 rather than segment 6.

Etymology.—Generic name, derived from the Greek, and transcribed to the Latin with a feminine-singular ending, refers to number of thoracic uncinigerous segments.

Endecamera palea, new species Fig. 2A—C

Material examined.—St. Croix Station, 17°57.63'’N, 64°48.6’W, in 3995 m DSRV Alvin dive 876, 20 Dec 1978, ‘‘wild’’ wood about 6 feet long. Eighteen specimens. Holotype (USNM 71547); 3 paratypes (USNM 71548).

Description.—Maximum size 5 mm long, 0.75 mm wide. Holotype complete, 3.3 mm X 0.5 mm. Color in alcohol white to pale orange. Trilobed prostomium with middle lobe about same width as lateral lobes; lacking glandular ridges. Smooth, ventrally grooved, oral tentacles. Segments 1 and 2 fused ventrally form- ing lower lip. Two lateral groups of paleae, about 11 in each group on segment 3. Each palea approximately 0.38 mm long, 5 wm wide basally, tapering gradually to a fine point. Four pairs smooth branchiae, about %4 body length; 3 of each group in a straight line across dorsal surface of segments 3-4 with 4th inserted just anterior to most medial branchiae. No mid-dorsal gap between branchial groups. Notopodial lobes, each bearing 7-12 winged capillary setae from segment 4 to end of thorax. Setae about 0.43 mm long, 8 wm wide basally, and 10 um wide across the blade. Notopodia of segment 4 minute while those of segments 5 and 6 larger than those of segment 4, but smaller than those of subsequent segments. Fourteen thoracic setigerous segments. Neuropodial lobes bearing toothed uncini from segment 7 to end of abdomen; 11 thoracic and 14 uncinigerous abdominal segments. Thoracic uncini in single transverse rows, about 23-27 per row. Each with 10 teeth, more or less in 2 transverse rows, above a rounded basal prow (Fig. 2B). Abdominal uncini in single transverse rows, about 10-18 per row. Each with about 13 teeth in several rows above a rounded basal prow (Fig. 2C). Abdominal notopodia and notopodial, neuropodial and anal cirri lack- ing. Pygidium rounded.

Remarks.—Mucus-lined tubes covered with particulate matter, about 3 times body length. Sexually mature females were broken or twisted making it impos- sible to count the number of eggs in the body cavity. No gonoducts visible.

Etymology.—Specific name, derived from the Latin, refers to presence of pa- leae on segment 3.

VOLUME 95, NUMBER 1 53

Fig. 2. Endecamera palea: A, Lateral view of entire worm, 4 mm long; B, Mid-thoracic uncini, lateral and frontal views, length = 14 wm; C, Mid-abdominal uncini, lateral and frontal views, length = 12 um.

54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Feeding of the Two Species

Ampharetid polychaetes live in mucus-lined tubes covered by particulate mat- ter. The tube often extends some distance above the sediment surface. Jumars found that starved Hobsonia increased the length of their tubes more, over a period of several months, than those that were fed (Fauchald and Jumars 1979). They suggested that tube building may be a form of locomotion, allowing animals to enter new feeding areas. Fauchald and Jumars (1979) also suggested that hor- izontal or vertical tube orientation may depend on food availability; horizontal tube orientation would allow worms to cover a larger feeding area.

During feeding, worms emerge from the tube opening and evert their oral ten- tacles equal to about their body length over the sediment surface (Fauchauld and Jumars 1979; personal observations on Asabellides oculata [Webster], Hobsonia [Amphicteis| floridus [Hartmen] and Melinna cristata [Sars]). Food adheres to mucus produced on the ventral side of the tentacles and is carried by cilia to the mouth. The tentacular apparatus of Decemunciger apalea and Endecamera palea appears similar to that of shallow water ampharetids and it is assumed that the method of feeding is also similar. Pieces of wood, detritus and bivalve larvae (Xylophaga sp.) were found in the gut cavity of Decemunciger apalea, while only detritus was noted in the digestive tract of Endecamera palea.

The digestive tract of both species like most ampharetids consists of buccal cavity, oesophagus, stomach, intestine, and rectum. The buccal cavity houses about 14 ventrally grooved and ciliated, smooth oral tentacles.

Food available to deep-sea organisms includes plankton, remnants of marine macrophytes such as Sargassum, remnants of land plants, particulate residues of deteriorating nekton and chemo-autotrophic bacteria found in hydrothermal vent areas (George and Higgins 1979, Rowe and Staresinic 1979). Rowe and Staresinic (1979), using sediment traps, found that about 4 g C/m?/yr, mainly in the form of fecal pellets, reached the deep-sea bottom. Marine macrophyte re- mains were next in importance. Pieces of the brown alga Sargassum accounted for an average of 0.4 g C/m?/yr while the contribution from other sources was relatively small. |

Decemunciger apalea most likely feeds on fecal pellets produced by other animals on the “wood islands,”’ on bivalve larvae (Xylophaga sp.) that settle near them, and on any type of detrital particle that reaches them from the sur- rounding water column. Fungi and bacteria on small pieces of ingested wood may represent an added source of nutrients.

Life History/Recruitment/Growth Rate of Decemuniger

The rate of colonization in deep-sea sediments is about two orders of magnitude lower than in shallow water (Grassle 1977). In addition, larval recruitment and settlement, growth rates, and probably mortality rates, are generally lower in the deep sea (Grassle and Sanders 1973, Grassle 1977, Sanders 1979). Grassle (1980b) found fewer individuals and species in boxes of azoic sediment placed on the deep-sea floor, compared to samples taken from surrounding sediments. Because of these factors and others, deep-sea populations are commonly dominated by mature adults (Grassle 1977). Opportunistic wood boring, deep-sea bivalves (Subfamily Xylophagainae; Family Pholadidae) characterized by rapid growth,

VOLUME 95, NUMBER 1 55

Table 1.—Numbers of Decemunciger apalea (in parentheses) of various body lengths (in mm) for listed wood panels.

N31 DOS 2: (1) 2.0; (4) 2.7; (3) 3.0; (2) 3.2; (5) 3.6; (1) 4.0; (7) 4.5; (1) 5.4; C1) 6.0; (1) 6.3.

N32 DOS 1: (2) 0.4; (1) 0.6; (1) 0.65; (1) 0.85; (5) 1.0; (2) 1.2; (3) 1.4; (2) 1.75; C1) 1.8; C1) 1.9; (2) e038 (C)) 232 (8) e523 (1) 252 (Ob) S02 Gly S552 (10) S05

N34 and

NOS DOS me (0! 752 (1) 028; (2): 1:0; (2) 1.25; GB) 1.53 C1) 1:75; (1) 1.75: CG) 2.5; GC) 3:0.

ING DOSE) th2--() 153: (2) 1.5; CU) 1.75; (1) 2.5; C1) 2:65.

ING2Z DOS 1: (1) 1.5; (1) 3.0; (1) 4.0; (1) 4.5; C1) 35.25.

N91 DOS 1: (1) 1.0; (1) 4.5.

early sexual maturity, and relatively large numbers of eggs, appear to be an exception to the generality above (Turner 1973, 1977). The larger number of eggs and the resultant large number of motile larvae allow at least a few individuals to reach geographically scattered pieces of wood (Turner 1973, Sanders 1979).

Decemunciger appears to have a life cycle similar to these bivalves. The large number of small eggs in the body cavity implies a high reproductive rate. The apparent onset of sexual maturity within a year’s time, documented below, sug- gests early maturity and a rapid growth rate. Thus, also considering its ability to colonize a transient habitat, Decemunciger may be characterized as an oppor- tunistic species. Evidence for this is as follows, the only unknown aspect being the method of larval dispersal.

Decemunciger with well developed egg or sperm in the body cavity ranged from about 3.6 to 6.3 mm in total body length. Worms less than 3.6 mm long were therefore assumed to be juveniles. Eggs probably pass singly through certain nephridia and nephridiopores into the anterior part of the tube where they are fertilized by sperm, released by males in the same fashion. Hobsonia (Amphic- teis) floridus (Zottoli, 1974), Hypaniola kowalewski (Marinescu, 1964) and Mel- innexis artica (Annekova, 1931), which have eggs roughly the same diameter as Decemunciger, retain developing larvae in the maternal tube until they are able to crawl on the bottom. It is hypothesized that these brooding and colonization patterns are the same for Decemuniciger.

Sexually mature worms as well as juveniles were found on wood panels N72 and N91 which were submerged for one year. Thus, if one accepts the premise that wood panels are colonized solely by larvae, then sexual maturity is attained within a year (Table 1). The absence of sexually mature Decemunciger on wood panels N34, N65, and N67, which were also submerged for one year, suggests that larval worms settled on the wood late in the year and did not have sufficient time to reach sexual maturity. In addition, the presence of certain size classes of adults and juveniles on wood panels N31, N34, N72, and N91 suggests that this species reproduces only at certain times of the year. Rokop (1974) felt that deep- sea organisms generally reproduce throughout the year. Lightfoot, Tyler, and Gage (1979) suggest that cyclic seasonal breeding is more common in the deep- sea than previously supposed. Timing of reproduction most likely reflects sea- sonal abundance of food reaching the deep-sea floor (Lightfoot, Tyler, and Gage 1979).

56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Acknowledgments

Thanks are due to Dr. J. Fred Grassle, Charlene D. Long, Dr. Meredith L. Jones, and Dr. Ruth D. Turner for making specimens available and for reviewing the manuscript. The study, conducted by Turner (1977), is supported by the Office of Naval Research (ONR Contract No. 14-76-C-1281, NR 104-687 to Har- vard University). This paper is dedicated to my recently departed friend and colleague, Dr. Fred (Eric) Davis.

Literature Cited

Augener, H. 1918. Polychaeta. In: W. Michaelsen (ed.), Beitrage zur Kenntnis der Meersfauna Westafrikas 2(2):67—625.

Caullery, M. 1944. Polychetes sédentaire de L’Expedition du Siboga: Ariciidae, Spionidae, Chae- topteridae, Chloraemidae, Opheliidae, Owentidae, Sabellariidae, Sternaspidae, Amphictendiae, Ampharetidae, Terebellidae.—Siboga-Expeditie Monographie 24(2): 1-204.

Day, J. H. 1964. A review of the family Ampharetidae (Polychaeta).— Annals of the South African Museum 48:97-121.

Eliason, A. 1955. Neue oder wenig bekannte schwedische Ampharetiden (Polychaeta).—Goteborgs Kungelige Vetenskaps Handlingar 6B(17):1-17.

Fauchald, K. 1972. Benthic polychaetous annelids from deep water off western Mexico and adjacent

areas in the eastern Pacific Ocean.—Allan Hancock Monographs Marine Biology 7:1—575.

, and P. A. Jumars. 1979. The diet of worms: A study of polychaete feeding guilds —Ocean- ography Marine Biology Annual Review 17:193—284.

Fauvel, P. 1927. Polychetes sédentaires.—Faune de France 16:1—494.

George, R. Y., and R. P. Higgins. 1979. Eutrophic hadal benthic community in the Puerto Rico trench. In: The Deep-Sea Ecology and Exploitation —AMBIo Special Report, No. 6:51—58.

Grassle, J. F. 1977. Slow recolonization of deep-sea sediment.—Nature 265:618—619.

. 1980a. In Situ Studies of Deep-Sea Communities. In: Advanced Concepts in Ocean Mea-

surements For Marine Biology. F. P. Diemer, F. J. Vernberg, and D. Z. Mirkes (eds.).—The

Belle Baruch Library in Marine Science, No. 10:321-332.

. 1980b. Rates of colonization in deep-sea benthic communities [Abstract]—American Zool-

ogist 20(4):929.

, and H. L. Sanders. 1973. Life histories and the role of disturbance.—Deep-Sea Research

20:643-659.

Hartman, O. 1965. Deep-water benthic polychaetous annelids off New England to Bermuda and

other North Atlantic areas.—Allan Hancock Foundation Occasional Paper 28:1-378.

. 1967. Polychaetous annelids collected by the USNS ELTANIN and STATEN ISLAND

cruises, chiefly from Antarctic seas.—Allan Hancock Monographs in Marine Biology 2:1—387.

, and K. Fauchald. 1971. Deep-water benthic polychaetous annelids off New England to

Bermuda and other North Atlantic areas. Part 2.—Allan Hancock Monographs in Marine

Biology 6:1-327.

Levinsen, G. M. R. 1884. Systematik-geografisk. oversight over de nordiske Annulata, Gephyrea, Chaetognathi og Balanoglossi.—Videnskabelige Meddeleser Dansk Naturhistorisk Forening 1883:92-348.

Lightfoot, R. F., P. A. Tyler, and J. D. Gage. 1979. Seasonal reproduction in deep-sea bivalves and brittlestars.—Deep-Sea Research 26(8A): 767-773.

Malmgren, A. J. 1865-1866. Nordiska Hafs-Annulater.—Ofversigt Svenska Ventenskaps Akademiens Forhandlingar 22:181—192; 355-410.

McIntosh, W. C. 1885. Report on the Annelida Polychaeta collected by the HMS Challenger during the years 1873—76.—Challenger Reports 12:1—-554.

Nyholm, K. 1950. Contributions to the life-history of the ampharetid, Melinna cristata.—Zoologiska Bidrag fran Uppsala 29:79-91.

Okuda, S. 1947. On an ampharetid worm, Schistocomus sovjeticus Annekova, with some notes on its larval development.—Journal of the Faculty of Science Hokkaido Imperial University (6) 9:32 1—329)

VOLUME 95, NUMBER 1 57

Rokop, F. J. 1974. Reproductive patterns in the deep-sea benthos.—Science 186:743-745.

Rowe, G. T., and N. Staresinic. 1979. Sources of organic matter to deep-sea benthos. In: The Deep Sea Ecology and Exploitation.—AmBio Special Report No. 6:19-24.

Sanders, H. L. 1979. Evolutionary ecology and life-history patterns in the deep sea.—Sarsia 64: 1-7.

Turner, R. D. 1973. Wood-boring bivalves, opportunistic species in the deep sea.—Science

180: 1377-1379.

. 1977. Wood, molluscs, and deep-sea food chains.—Bulletin of the American Malacological

Union 1977:13-19.

Zottoli, R. A. 1974. Reproduction and larval development of the ampharetid polychaete Amphicteis floridus.—Transactions American Microscopical Society 93(1):78-89.

Fitchburg State College, Fitchburg, Massachusetts 01420.

PROC. BIOL. SOC. WASH. 95(1), 1982, pp. 58-66

THE KARYOTYPE OF THE EURASIAN FLYING SQUIRREL, PTEROMYS VOLANS (L.), WITH A CONSIDERATION OF KARYOTYPIC AND OTHER DISTINCTIONS IN GLAUCOMYS SPP. (RODENTIA: SCIURIDAE)

V.R. Rausch and R. L. Rausch

Abstract.—The karyotype of Pteromys volans (L.) Qn = 38; FN = 73) is de- scribed, based on material from Hokkaido and northeastern Siberia, and com- pared with those of Glaucomys volans (L.) (2n = 48; FN = 80) and Glaucomys sabrinus (Shaw) (2n = 48; FN = 78) from North America. Although the similar- ities of the fundamental numbers are suggestive of karyotypic evolution of the Robertsonian type, the fossil record of these flying squirrels is at present frag- mentary, and no conclusion is drawn as to their affinities. Definite differences were observed between the karyotypes of G. volans and G. sabrinus, which had been described as identical. In combination with karyotypic differences, other characteristics of the two North American species indicate both a divergence earlier and an age greater than have been considered on the basis of paleontologic evidence. Karyograms of the three species of flying squirrels are included.

In the course of investigations concerning northern mammals, we studied chro- mosomal preparations from two specimens of the Eurasian flying squirrel, Ptero- mys volans (L.), from Hokkaido and northeastern Siberia. Chromosomes of the two species of flying squirrels of the Nearctic genus Glaucomys were compared. The purpose of the present report is to define the karyotype of P. volans and to describe some previously unrecognized differences in the karyotypes of the two Nearctic species. Other distinguishing characteristics of Glaucomys spp. also are briefly discussed.

Materials and Methods

Two male specimens of P. volans were utilized. The first, P. volans orii Ku- roda, was provided in 1967 by Dr. H. Abe, who captured the animal at Koshimizu, Abashiri, Hokkaido Island (ca. 43°58’N, 144°30’E). Cells from bone marrow and from one testis were fixed and stained in acetic orcein, following standard meth- ods for mammalian karyology. The second, P. volans cf. incanus Miller, was trapped by us in August 1979 along the upper Kolyma River, Magadansk Oblast’, near the settlement of Sibik-Tiellakh (ca. 62°N, 149°30’E). Cells from marrow (femur) were prepared in the field and processed with Giemsa blood stain (Sea- bright 1972). In both cases, cells were treated with colchicine.

Preparations were made by the same methods for Nearctic flying squirrels, as follows: Glaucomys sabrinus yukonensis (Osgood), two males collected at An- chorage and near Fairbanks, Alaska, in October 1966 and April 1969, respective- ly; one male G. sabrinus bangsi (Rhoads), captured on Powwatka Ridge, Wallowa Mountains, Oregon (ca. 45°40’N, 117°27’W), in December 1980; and one male G. volans querceti (Bangs), collected in the vicinity of Tampa, Florida, in early 1980.

VOLUME 95, NUMBER 1 59

After chromosomes (in metaphase) were counted, representative cells were photographed on high-contrast film, and the component chromosomes in five cells from each animal were measured (excepting the squirrels from the upper Kolyma River and from Anchorage, in which only contracted chromosomes were ob- served in the preparations). Chromosomes exhibiting Giemsa-banding were seg- regated according to size and banding-pattern with use of photographs and light microscopy. The identification of the X-chromosome was based on measurements and pattern of banding. The Y-chromosome in Glaucomys spp. was selected subjectively, since the small submetacentric chromosomes in these animals, of which the Y is one, appeared to be similar in both size and banding. The termi- nology concerning the location of the centromere on individual chromosomes follows Levan et al. (1964). The number of major chromosomal arms (funda- mental number, or FN) was determined according to the procedure of Matthey (1945). The chromosomal components of 30 to 61 cells from each of the animals were counted, excepting those from the upper Kolyma River and Anchorage for which only 18 and 15 cells, respectively, were counted.

Results

I. Pteromys volans (2n = 38, n = 19).—The diploid complement was made up of 12 metacentric autosomes (arm-ratios ranged from 1.03 to 1.7), 22 submeta- centric-subtelocentric and 2 acrocentric autosomes (arm-ratios from 2.4 to 6.3), the submetacentric X-chromosome (arm-ratio from 1.6 to 1.8), and the acrocentric Y-chromosome (arm-ratio from 4.2 to 7.3). In 61 cells (53 mitotic and 8 meiotic) examined, 49 contained 38 chromosomes and 7 contained 19 bivalent elements. A karyogram from the male P. volans from Hokkaido is shown in Fig. 1.

The set of chromosomes (in all of the 18 cells examined) from the animal collected in northeastern Siberia was morphologically like that of P. volans from Hokkaido, consisting of 6 pairs of metacentric and 12 pairs of submetacentric- acrocentric autosomes, a submetacentric X, and an acrocentric Y-chromosome, as shown in Fig. 2.

The number of major autosomal arms was 70 in both animals. Two autosomes with thin and lightly stained short arms were considered to be acrocentric. The FN (autosomes + X and Y) was 73 for P. volans.

II. Glaucomys spp. (2n = 46).—Karyograms from G. volans (Florida) and from G. sabrinus (Oregon and Alaska) are shown in Figs. 3—5. Thirty bi-armed (meta- centric-subtelocentric) chromosomes (with arm-ratios ranging from 1.03 to 8.8) and 16 acrocentric chromosomes (with centromeres in the terminal region; arm- ratios ranged from 4.5 to 13.8) constituted the autosomal set in G. volans. In G. sabrinus, from both localities, the autosomes included 28 metacentric-subtelo- centric (arm-ratios ranging from 1.07 to 8.6) and 18 acrocentric (arm-ratios from 4.3 to 14.5) elements. The sex-chromosomes of the two species were similar morphologically and in pattern of banding: X submetacentric, of medium size (ranges of arm-ratios were 1.5 to 2.03 for G. volans, and 1.9 to 2.9 for G. sa- brinus), and Y submetacentric of small size (ranges of arm-ratios were 2.1 to 3.2 for G. volans, and 2.0 to 3.3 for G. sabrinus). The patterns of bands indicated that these two species have some equivalent, or homologous, chromosomes. However, significant morphologic differences also were evident.

60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Figs. 1-5. Karyograms of Pteromys volans (2n = 38) and Glaucomys spp. (2n = 48). 1, P. volans, male, from Hokkaido Island; Orcein stain. 2. P. volans, male, from the Magadansk Oblast; Giemsa blood stain. 3, G. volans with Giemsa-bands, male, from Florida. Arrow indicates metacentric ele- ments not present in G. sabrinus. 4, G. sabrinus with Giemsa-bands, male, from Oregon. Arrow indicates acrocentric elements not present in G. volans. 5, G. sabrinus, male, from Alaska; Orcein stain. Scale-lines have value of 5 wm.

VOLUME 95, NUMBER 1 61

Discussion

The gliding squirrels allocated to the genera Pteromys G. Cuvier, 1800, and Glaucomys Thomas, 1908, have had rather complex taxonomic histories and, as indicated by recent reviews (cf. Mein 1970, Black 1972), their origins and affinities are obscure. The grounds for acceptance of the genus Pteromys instead of Sci- uropterus F. Cuvier, 1825, for the flying squirrel in northern Eurasia have been discussed by Miller (1914) and by Ellerman and Morrison-Scott (1951), among others, but Simpson (1945) considered the latter to be probably the valid generic designation. In addition to the widely occurring P. volans, the genus includes a second species, P. momonga Temminck, 1846, which occurs on the Japanese Islands of Honshu, Kyushu, and Hondo. According to Ellerman and Morrison- Scott (1951), its cranial differences are so marked that it cannot be considered a race of P. volans.

In his revision of the genera and subgenera of the Sciuropterus-group of flying squirrels, Thomas (1908) recognized four subgenera in Sciuropterus, mainly on the basis of dental characters. These included Sciuropterus, with S. (Sciuropter- us) russicus Tiedemann, 1808 (=Sciurus volans Linnaeus, 1758) as type, and Glaucomys, which was erected for the Nearctic flying squirrels, with S. (Glau- comys) volans (=Mus volans Linnaeus, 1758) as type. Glaucomys was elevated to generic rank by Howell (1915), on grounds not stated, but he later (1918) defined cranial and dental characters that distinguish Glaucomys from Pteromys. These distinctions were confirmed by Ognev (1940), who also pointed out the marked differences in the structure of the os penis in P. volans and G. volans. Recently, from the study of both fossil and living flying squirrels, Mein (1970) placed Pteromys and Glaucomys in different generic groupings on the basis of dental characters.

The fossil record provides no clear indication of the origin of Pteromys volans. However, the comparison of serum proteins by means of double immunodiffusion by Zholnerovskaia et al. (1980) suggested that Pteromys has its closest affinities with Sciurus and Tamias, rather than with terrestrial sciurids. In North America, Glaucomys is known from the middle to late Irvingtonian. The stratigraphic range of Glaucomys volans extends from the late Irvingtonian, whereas remains of the northern G. sabrinus are known only from deposits of late Wurm age (Kurtén and Anderson 1980). Thenius (1972) considered that Glaucomys appears to have arisen from terrestrial sciurids, which had their center of radiation in North America.

The diploid number of 38 chromosomes, as reported by Tsuchiya (1979) and determined by us, alone distinguishes the karyotype of Pteromys volans from those of Glaucomys spp. (2n = 48) (Nadler and Sutton 1967, Schindler et al. 1973). In the former, 35 chromosomes were metacentric-subtelocentric, and only 3 (two autosomes and the Y-chromosome) were acrocentric. The relatively large, acrocentric Y of P. volans is markedly different from the male sex-chromosomes of Glaucomys spp. That the fundamental numbers in these squirrels are rather similar (73 in P. volans; 78 in G. sabrinus; and 80 in G. volans) suggests that karyotypic evolution involving chromosomal rearrangements of Robertsonian type might have occurred. However, no significance is attributed to the apparent similarities when nothing is known concerning the further relationships of the Eurasian and American species.

62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

The karyotype of a female specimen of P. momonga has been published by Tsuchiya (1979, fig. 7). It appears to be quite similar to that of P. volans, but the available details do not permit an extensive comparison. The presence of boreal forest far to the south in the Japanese Islands during the glacial maximum of Wurm time would seem to have favored the southward dispersal of Pteromys volans (cf. Kotani 1969).

With reference to the karyotypes of Glaucomys spp., the published information is discrepant. That G. volans and G. sabrinus are karyotypically identical has been generally accepted. However, for both, Schindler et al. (1973) considered the number of bi-armed (metacentric-subtelocentric) autosomes to be two more than did Nadler and Sutton (1967). Hsu and Benirschke (1973) concurred with Schindler et al. The karyotype defined by us for G. sabrinus yukonensis (Alaska) agreed more closely with that shown and discussed by Nadler and Sutton (1967), who studied preparations from G. sabrinus lascivus (Bangs) (California), than with the results of Schindler et al. (1973) for G. sabrinus from New Hampshire (an area where this species is marginally sympatric with G. volans). Accordingly, we examined new material from G. sabrinus as well as G. volans, from geographic regions far removed from areas of parapatry or sympatry (Oregon and Florida). The identities of the animals studied were confirmed from the distinctive char- acteristics of the os penis, in addition to other criteria.

Although the respective karyotypes of these flying squirrels appeared superfi- cially to be much alike, they exhibited distinct differences, principally in the presence in G. volans of two small metacentric autosomes that did not occur in G. sabrinus, and in G. sabrinus of two small acrocentric autosomes not observed in G. volans. As well, the total lengths of individual chromosomes differed be- tween the two species, including those in some presumed to be homologous; the relative length of the female haploid complement (autosomes + X) was recorded as 65 wm in G. volans and 86 um in G. sabrinus. Morphologically, the sex- chromosomes were similar.

Of the chromosomes of G. volans, the two small metacentric autosomes (see Fig. 3, arrow) were the most nearly metacentric, sensu stricto, with an arm-ratio ranging from 1.03 to 1.05. However, these and the two small acrocentric elements in G. sabrinus (see Fig. 4, arrow) were similar in total length; their value relative to the lengths of the respective haploid complements was the same (3.5%) in both species. The appearance of Giemsa-bands in these chromosomes suggests that they may be homologues, and that the small metacentric pair in G. volans arose as a result of a pericentric inversion involving the small acrocentric pair in G. sabrinus (or perhaps vice versa). The disparate total lengths of the respective complements further suggest quantitative differences in nuclear DNA, which were not defined by our methods.

Our findings indicate that the two species of Glaucomys are karyotypically distinct, and they are in contrast with the conclusions of others concerning these flying squirrels. The karyogram from G. volans from Florida (Fig. 3) appears to be identical with those shown by Schindler et al. (1973) as representative of both G. volans and G. sabrinus in New Hampshire, where the two species are sym- patric.

The two species of Glaucomys are distinguished additionally by major differ- ences in the form of the os penis, as is also Pteromys volans. Since detailed

VOLUME 95, NUMBER 1 63

Table 1.—Helminths of Glaucomys volans and G. sabrinus.

G. sabrinus G. volans

Midwest Midwest Oregon (n = 22) (n = 5) (n = 26) Species of helminth no. infected no. infected no. infected Nematoda: Capillaria americana Read, 1949 7 — — Syphabulea thompsoni (Price, 1928) 1 5 a= Syphabulea sp.* — — 17 Lemuricola sciuri (Cameron, 1932) 13 a a Citellinema bifurcatum Hall, 1916 2 D y Cestoda: Andrya sciuri Rausch, 1947 — 2 4 Monoecocestus thomasi Rausch and Maser, 1977 — — 18 Catenotaenia sp. — 3 — Hymenandrya sp. — — 3

* Description in preparation by J.-P. Hugot, Museum National d’Histoire Naturelle, Paris.

descriptions of these structures have been published, it suffices here to point out their diversity in the three species. The os penis of P. volans is small (ca. 4 mm long in our material) and consists of a slender, pointed bone with a well developed barb at the proximal end (cf. Ognev 1940, fig. 153). That of G. volans, first described by Pocock (1923), is long (ca. 14 mm in specimens from Florida) and very slender, whereas that of G. sabrinus is shorter (ca. 6.5 mm in our material), broader, and flattened (see Burt 1960, plate IV). Each differs markedly from the others, and we were unable to discern any fundamental similarities unless, per- haps, the barb near the distal end of the os penis of G. sabrinus is homologous with the basal barb in that of P. volans.

Certain differences in the helminth faunas of the two species of Glaucomys provide a further indication of dissimilarity. Of the nine species of helminths recorded by us from flying squirrels in North America (Table 1), six occur also in sciurids of other species (Rausch and Tiner 1948, Davidson 1976, McGee 1980). Host-specific cestodes of three species, Andrya sciuri, Monoecocestus thomasi, and Hymenandrya sp., are known only from Glaucomys sabrinus. The assem- blages involving these cestodes would seem to have arisen independently through coevolution of helminth and host (Rausch 1981), indicating comparatively early divergence of the two species of Glaucomys. These cestodes are Nearctic species, with no congeners known from Pteromys volans or other sciurids in Eurasia. The only helminth known from P. volans is Citellina petrovi Shul’ts, 1930, which is host-specific and has been recorded from European Russia, Japan, and Chukotka (Hugot 1980). Some of the characteristic ectoparasites of Pteromys volans and Glaucomys spp. are congeneric (Acarina and Mallophaga), but no species is shared. Consequently, these arthropods do not appear to provide any useful indications of the relationships of their hosts.

Various hypotheses have been proposed to account for the essential allopatry of the Nearctic flying squirrels, but the geographic ranges of these mammals seem simply to reflect differences in specific ecologic requirements. The two species occupy disjunct ranges over the greater part of North America where they occur,

64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

relative to the distribution of coniferous and deciduous forest. In the east, where their ranges are parapatric to sympatric, they appear as well to be segregated ecologically. As noted by Guilday (1962), G. sabrinus has a relict distribution in the Appalachian Mountains, where it occurs at elevations above ca. 1000 m as a consequence of climatic change (warming) during post-glacial time.

That the characteristic habitats of the two species of Glaucomys have a fun- damental relationship to their different dietary requirements is evident. Glauco- mys volans utilizes buds, fruits, and insects during spring and summer, but later feeds mainly on acorns and other nuts and seeds, some of which are stored in large quantities and consumed during winter (cf. Muul 1968). Glaucomys sabrinus depends mainly on hypogeous fungi during the warmer months, and utilizes epi- phytic lichens (Usnea and Alectoria) during winter (McKeever 1960, Maser ef al. 1978, Maser pers. comm.). The diet of Pteromys volans differs from these in that it feeds on sprouts or buds of Betula and Acer and on male flowers of conifers in summer, and on catkins or buds of Betula and Alnus in winter (Ognev 1940, Tavrovskii et al. 1971).

Suggestions have been made to account for the pattern of distribution of Glau- comys sabrinus on grounds other than ecologic requirements. Muul (1968) con- sidered that the southward spread of G. sabrinus is prevented by competitive interaction with G. volans, inasmuch as the latter breeds earlier seasonally and therefore would already occupy the limited number of tree-cavities available. However, as reported by Cowan (1936), and observed by us in northern Wiscon- sin, G. sabrinus commonly utilizes outside nests constructed mainly of twigs.

Weigl (1975) observed in captive animals that infection by a nematode of the genus Strongyloides was tolerated by G. volans, but was associated with high mortality in G. sabrinus. He considered that differential pathogenicity of this nematode might account in part for the segregation of the two species of Glau- comys where they are sympatric. Barbehenn (1978) suggested that this might constitute a mechanism by which the larger G. sabrinus would be excluded from habitat otherwise suitable for G. volans. However, Strongyloides robustus Chan- dler, 1942 exhibits little host-specificity, occurring in sciurids of various species (Rausch and Tiner 1948, McGee 1980). Davidson (1976) considered it to be the most pathogenic of the more common helminths in squirrels, and reported that infections involving more than 150 individuals often caused severe enteritis in gray squirrels, Sciurus carolinensis Gmelin. It might be expected that transmis- sion of this nematode would be enhanced by conditions of captivity, and that differences in feeding habits or other behavior could account for more massive infections in G. sabrinus. That Weig] did not find northern flying squirrels natu- rally infected by Strongyloides appears to be compatible with the results of other surveys of helminths in this sciurid.

Altogether, the differences between the two flying squirrels of the genus Glaucomys seem to indicate earlier divergence and greater age of these species than has been considered on the basis of paleontologic evidence. The origins and affinities of Glaucomys spp. and of Pteromys volans may be established if the earlier fossil record of these sciurids can be traced.

Acknowledgments

Specimens of flying squirrels were kindly provided by H. Abe, Faculty of Agriculture, Hokkaido University, Sapporo; Ms. E. L. Bull, Pacific Northwest

VOLUME 95, NUMBER 1 65

Range and Habitat Laboratory, U.S. Department of Agriculture, La Grande, Oregon; R. Anderson, Wallowa Valley Ranger District, U.S. Department of Ag- riculture, Joseph, Oregon; and I. L. Duncan, Center for Disease Control, Atlanta. Field work in the Soviet Union was carried out under the U.S.A./U.S.S.R. En- vironmental Protection Treaty, Area V, Subproject A-3. In the U.S.S.R., V. L. Kontrimavichus, D. I. Berman, and Ms. A. N. Leirikh, Institute of Biological Problems of the North, Academy of Sciences of the U.S.S.R., Magadan, kindly provided support and assistance. We express sincere thanks for these contribu- tions.

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Barbehenn, K. R. 1978. Concluding comments: From the worm’s view, *‘Eco-pharmodynamics”’ and 2000 A.D., pp. 231-236 in: D. P. Snyder, ed. Populations of small mammals under natural conditions. Volume 5, Special Publications Series.—Pymatuning Laboratory of Ecology, Uni- versity of Pittsburgh, Linesville, Pennsylvania. 237 pp. |

Black, C. C. 1972. Holarctic evolution and dispersal of squirrels (Rodentia: Sciuridae).—Evolutionary Biology 6:305—322.

Burt, W. H. 1960. Bacula of North American mammals. Museum of Zoology, University of Michigan, Miscellaneous Publications, No. 113. Ann Arbor. 75 pp. + 25 pl.

Cowan, I. McT. 1936. Nesting habits of the flying squirrel Glaucomys sabrinus.—Journal of Mam- malogy 17:58-60.

Davidson, W. R. 1976. Endoparasites of selected populations of gray squirrels (Sciurus carolinensis) in the southeastern United States.—Proceedings of the Helminthological Society of Washington 43:211-217.

Ellerman, J. R., and T. C. S. Morrison-Scott. 1951. Checklist of Palaearctic and Indian mammals. British Museum (Natural History), London. 810 pp.

Guilday, J. E. 1962. The Pleistocene local fauna of the natural chimneys, Augusta County, Virginia.— Annals of Carnegie Museum 36:87-122.

Howell, A. H. 1915. Descriptions of a new genus and seven new races of flying squirrels.—Pro-

ceedings of the Biological Society of Washington 28:109-114.

. 1918. Revision of the American flying squirrels. North American Fauna No. 44. Washington,

D.C. 64 pp.

Hsu, T. C., and K. Benirschke. 1973. Glaucomys volans (southern flying squirrel). Folio 312. Atlas of mammalian chromosomes, Volume 7. Springer-Verlag, New York.

Hugot, J.-P. 1980. Sur le genre Citellina Prendel, 1928 (Oxyuridae, Nematoda).—Annales de Par- asitologie (Paris) 55:97-109.

Kotani, Y. 1969. Upper Pleistocene and Holocene environmental conditions in Japan.—Arctic An- thropology 2:133-158.

Kurtén, B., and E. Anderson. 1980. Pleistocene mammals of North America. Columbia University Press, New York. 442 pp.

Levan, A., K. Fredga, and A. A. Sandberg. 1964. Nomenclature for centromeric position on chro- mosomes.—Hereditas 52:200-220.

Maser, C., J. M. Trappe, and R. A. Nussbaum. 1978. Fungal—small mammal interrelationships with emphasis on Oregon coniferous forests.—Ecology 59:799-809.

Matthey, R. 1945. L’évolution de la formule chromosomiale chez les vertébrés.—Experientia 1:50-60, 78-86.

McGee, S. G. 1980. Helminth parasites of squirrels (Sciuridae) in Saskatchewan.—Canadian Journal of Zoology 58:2040—2050.

McKeever, S. 1960. Food of the northern flying squirrel in northeastern California.—Journal of Mammalogy 41:270-271.

Mein, P. 1970. Les sciuropteres (Mammalia, Rodentia) néogenes d’ Europe occidentale.—Geobios 3:7-77.

Miller, G. S. 1914. The generic name of the common flying-squirrels.— Proceedings of the Biological Society of Washington 27:216.

Muul, I. 1968. Behavioral and physiological influences on the distribution of the flying squirrel, Glaucomys volans. Museum of Zoology, University of Michigan, Miscellaneous Publications, No. 134. Ann Arbor. 66 pp.

66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Nadler, C. F., and D. A. Sutton. 1967. Chromosomes of some squirrels (Mammalia: Sciuridae) from the genera Sciurus and Glaucomys.—Experientia 23:249-251.

Ognev,S.I. 1940. Zveri SSSR i prilezhashchikh stran. IV. Gryzuny. Akademiia Nauk SSSR. Moskva- Leningrad. 615 pp.

Pocock, R. I. 1923. The classification of the Sciuridae.—Proceedings of the Zoological Society of London (1923), pp. 209-246.

Rausch, R. L. 1981. Cestodes in mammals: The zoogeography of some parasite-host assemblages.

Symposium, La spécificité parasitaire des parasites de vertébrés. Muséum National d’ Histoire

Naturelle, Paris, 13-17 April 1981. (in press)

, and J. D. Tiner. 1948. Studies on the parasitic helminths of the north central states. I.

Helminths of Sciuridae.— American Midland Naturalist 39:728—747.

Schindler, A.-M., R. J. Low, and K. Benirschke. 1973. The chromosomes of the New World flying squirrels (Glaucomys volans and Glaucomys sabrinus) with special reference to autosomal heterochromatin.—Cytologia 38: 137-147.

Seabright, M. 1972. The use of proteolytic enzymes for the mapping of structural rearrangements in the chromosomes of man.—Chromosoma 36:204—210.

Simpson, G. G. 1945. The principles of classification and a classification of mammals.—Bulletin of the American Museum of Natural History 85:1-—350.

Tavrovskii, V. A., O. V. Egorov, V. G. Krivosheev, M. V. Popov, and Iu. V. Labutin. 1971. Mlekopitaiushchie Iakutii. Nauka, Moskva. 660 pp.

Thenius, E. 1972. Grundziige der Verbreitungsgeschichte der Saugetiere. Eine historische Tiergeo- graphie. Gustav Fischer Verlag, Jena. 345 pp.

Thomas, O. 1908. The genera and subgenera of the Sciuropterus group, with descriptions of three new species.—The Annals and Magazine of Natural History (8)1:1-9.

Tsuchiya, K. 1979. A contribution to the chromosome study in Japanese mammals.—Proceedings of the Japanese Academy (B)55:191-195.

Weigl, P. D. 1975. Parasitism as a possible biological weapon affecting the ranges and interactions of the flying squirrels, Glaucomys volans and G. sabrinus. American Society of Mammalogists, 55th Annual Meeting, 16-19 June 1975. No. 36, Abstracts of Technical Papers, p. 17.

Zholnerovskaia, E. I., N. N. Vorontsov, and O. K. Baranov. 1980. Immunologicheskii analiz sis- tematicheskikh vzaimootnoshenii letiag (Pteromys) c piat’iu rodami palearkticheskikh belich’ikh (Sciuridae, Rodentia).—Zoologicheskii Zhurnal 59:750—754.

Burke Memorial Washington State Museum, DB-10, and Division of Animal Medicine, SB-42, University of Washington, Seattle, Washington 98195.

PROC. BIOL. SOC. WASH. 95(1), 1982, pp. 67-80

A NEW SPECIES OF THE GENUS ECHININUS (MOLLUSCA: LITTORINIDAE: ECHINININAE) WITH A REVIEW OF THE SUBFAMILY

Joseph Rosewater

Abstract.—A new ovoviviparous species, Echininus viviparus, inhabiting high intertidal and supratidal areas in the Mariana Islands is described in the littorinid subfamily Echinininae which previously consisted of the polytypic Indo-Pacific species Echininus cumingi (Philippi, 1846) and the west Atlantic species Tectin- inus nodulosus (Pfeiffer, 1839). Characteristics of the subfamily are reviewed.

I received for identification from L. G. Eldredge of the University of Guam specimens of an Echininus which upon close examination proved to be not only a new species, but one which exhibits ovoviviparity. The species described here I originally thought to be Echininus cumingi spinulosus (Philippi, 1847), and the realization that it is not prompted me to take a fresh look at the subfamily Echin- ininae resulting in the following review.

Family Littorinidae Gray, 1840 Subfamily Echinininae Rosewater, 1972

Diagnosis.—The subfamily Echinininae is characterized by spinose to bluntly spinose shells; umbilicate or imperforate; opercula multispiral; radulae with mod- erately to greatly narrowed rachidians. Distribution is tropical west Pacific and west Atlantic.

The subfamily Echinininae was established by Rosewater (1972) to contain the genus Echininus which differs in matters of sculpture, radula, animal morphology, and ecology from either Littorininae or Tectariinae. This subfamily concept is strengthened by findings enumerated in the present paper. Members of the subfamily have a unique habitat among Littorinidae, for they inhabit highest shore levels, at times seeming almost to be terrestrial. The radula of Echininus, while said to be “‘not unusual’’ (Rosewater 1972:507), is distinctive when examined with the SEM (Fig. 1A—D). The radula of Tectininus is unique (Fig. 1E—F). The operculum is multispiral, a departure from all other known littorines. Character- istics of both radulae and opercula are here considered adaptations to a high shore habitat, as is the occurrence of ovoviviparity in the new species of Echin- inus described herein. All are indicative of a basic evolutionary trend in Echini- ninae toward a terrestrial habit. Shells of this group are also distinctive, especially that of E. cumingi cumingi, which is characterized by rather long, open spines and a deep umbilicus. These latter features, also, may be of significant adaptive value to an animal living on tropical shores enabling easier dissipation of heat.

Tectininus Clench and Abbott, 1942

Tectininus Clench and Abbott, 1942:4; type-species by original designation Echin- inus nodulosus (Pfeiffer, 1839).—Abbott, 1954:458-462.

68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Diagnosis.—Shells bluntly spinose, succeeding spiral rows of spines non-syn- chronous; imperforate; opercula multispiral; radula with very reduced rachidian, modified lateral, and marginals with few denticles. Distribution is subtropical to tropical west Atlantic.

As originally proposed by Clench and Abbott (1942) and elaborated by Abbott (1954) the monotypic subgenus, Tectininus, was distinguished from Echininus s.s. because its type, E. nodulosus, possesses a unique radula, although still exhibiting the formula 2-1-1-1-2 and a modified littorinid embayment (Fig. 1E—F). It is unlike any other radula in the family, having massive lateral teeth and a diminutive rachidian which appears almost vestigial. The operculum is multispir- al. It possesses a penis bearing a subterminal penial gland surrounded with fleshy papillae, which differs from the condition in the nominate subgenus and combines characteristics noted in the genera Littorina, Nodilittorina, and Tectarius. Aj- though Rosewater (1972) had not seen the paper and reported that nothing was known of reproduction in Echinininae, it had already been shown that 7. nodu- losus 1S Oviparous and produces a pelagic egg capsule (Borkowski 1971).

Until further evidence for its true relationships comes to light, I am satisfied to permit Tectininus to remain in its present position. As stated by Abbott, after careful study of the type-species of Tectininus, it appears to be **. . . a special- ization of the ancestral stock . . . inthe family Littorinidae’’ (Abbott 1954), whose apparent closest relations are in the Echinininae. Because of its unique radula Tectininus should be accorded full generic status.

Echininus Clench and Abbott, 1942

Echininus Clench and Abbott, 1942:3; type-species by original designation Tro- chus cumingii Philippi, 1846.—Rosewater, 1972:525.

Diagnosis.—Shells spinose to bluntly spinose; often with partly open spines; umbilicate to imperforate; opercula multispiral; radulae with moderately nar- rowed rachidians, laterals and marginals with moderately numerous denticles. Distribution is west Pacific from southern Japan through western Pacific arc to Cook Islands. |

The genus Echininus was delineated as consisting of 2 subgenera containing a total of 2 living species and | subspecies (Rosewater 1972). The nominate sub- genus, Echininus, inhabits the west Pacific and contains E. cumingi cumingi (Philippi, 1846) known from the southern Philippines southward through Mela- nesia (including New Guinea, Solomon Islands, New Hebrides, New Caledonia) eastward to the Cook Islands, and, E. cumingi spinulosus (Philippi, 1847) which occurs in southern Japan, the Ryukyu Islands, Taiwan and northern Philippines. Echininus adelaidensis (Cotton, 1947) is a Pliocene fossil from South Australia, which resembles certain trochaceans (Rosewater 1972). New evidence from pre- served specimens indicates that the Mariana Islands population, previously be- lieved to be part of E. cumingi spinulosus, is different from either of the previ- ously known Pacific subspecies and is a new species.

Echininus cumingi cumingi, type-species, is comparatively large, reaching 20.9 mm (Table 1), with 3 spiral rows of conspicuous projecting, open spines on the body whorl; it has a narrow, deep umbilicus; a multispiral operculum is present; the radula has a moderately narrow rachidian, lateral and inner marginal teeth

VOLUME 95, NUMBER 1 69

each having one large and several smaller cusps, and slender outer marginals having 4—5 cusps. Echininus cumingi spinulosus is very close in appearance to E. c. cumingi, although it never reaches as large a size (18.3 mm), is less spinose, and has a narrower umbilicus which occasionally is closed. The specimens from the Mariana Islands which were included with the former by Rosewater (1972) are consistently different from either E. c. cumingi or E. c. spinulosus. Available records indicate they are limited in distribution to the southern Mariana Islands, and, so far have been reported only on the islands of Saipan, Tinian, Rota and Guam. Specimens are smaller in size, the largest measuring 12.3 mm in length; have only 2 major rows of blunt spines on the body whorl. In addition, the shell shape is different; obesity of the Mariana specimens averages 0.76 compared with 0.87 in cumingi and 0.78 in spinulosus. There are a number of quite absolute anatomical and biological differences also. Females of the Mariana population reproduce ovoviviparously, and the expanded oviduct may contain large numbers of young. The embryos are visible through the transparent dorsal mantle tissue in various stages of development including young snails with shells of 1.25—1.5 whorls. This condition is very similar to that described in Littorina saxatilis by Thorson (1946), and Fretter and Graham (1962). The reproductive modes of E. c. cumingi and E. c. spinulosus are not known, but there is no evidence from examination of preserved specimens to indicate Ovoviviparity in them. There are also morphological differences between radulae and penes of the Mariana species and the more western and northern Echininus.

Echininus viviparus, new species Figs. 1-6, Table |

Description.—Shell: reaching 12.3 mm (about /% inch) in length, turbinate in shape, with blunt spines; average obesity (width/length) about 0.76 (76 specimens range from 0.65—0.84) relatively thick in structure, usually non-umbilicate or with only a thin slit between columella and parietal callus; suture usually evident although often partially obscured by spiral sculpture; whorls 3—6, moderately shouldered below suture, relatively flat-sided; usually with 2 major rows of blunt spines on body whorl and one on spire whorls; occasionally with one or more additional major rows of spines. External shell color varying from grayish to tannish orange, the darker color characteristic of shells retaining periostracum; periostracum frequently worn away on spines which then appear grayish white. Apertural coloration light to dark orange with occasional light or darker orange lines revolving within. Base moderately flattened, sculptured with weakly nod- ulose cords. Length of spire greater than half length of shell. Spire convex; spire angle quite variable: from 60—72°. Aperture subquadrate; outer lip thickened, smooth within, only slightly wavy at edge reflecting external sculpture; inner lip with small but distinct toothlike bulge. Suture evident when not partly obscured by sculpture. Primary sculptural feature: 2 spiral rows of irregular blunt spines on body whorl, and one row on spire whorls. Rows of spines not well aligned axially; body whorl spine count |1—16 in posterior row and 12—16 in anterior row; 13-14 on penultimate whorl of spire. Secondary spiral sculpture consists of 3-4 fat, non-spinose cords between each spinose row, which occasionally become nearly as spinose as spinose cords they separate. In addition, an overall micro-

70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

f ‘ 4 as : f ; ” i a ; te Z. | ‘ Ld i ath ‘ ie $ < 3

Fig. |. Radulae: A, B, Echininus viviparus, 2° from northeast coast of Tinian, Mariana Islands (USNM 796244): A, Showing 2 complete transverse rows, Bar = 10 wm, 875; B, Rachidian tooth, Bar = 5 um 1800. C, D, Echininus cumingi cumingi, 2 from Davao, Mindanao, Philippines (USNM 747765): C, Showing 2 complete transverse rows, Bar = 10 wm, 700; D, Rachidian tooth, Bar = 5 pm, 1400. E, F, Tectininus nodulosus, 2 from San Salvador, Bahamas (USNM 596683): E, Showing 1 complete transverse row; note small, narrow rachidian tooth and massive laterals, Bar = 10 wm, 510; F, Oblique view of rachidian tooth, Bar = 2 wm, 3500.

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VOLUME 95, NUMBER 1

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72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Shells of Echininus viviparus: A, B, Holotype (USNM 792356), 9.7 x 7.4 mm; C, D, Paratype (USNM 803296) 10.7 x 7.6 mm.

scopical sculpture of fine, closely-spaced threads. Axial sculpture consists of irregular lines of growth. Operculum small in size, multispiral (polygyrous spiral type of Fretter and Graham 1962) having S—6 volutions, chitinous, light brown in color. Nuclear whorls usually present, smooth, light brown to reddish brown; reaching about 1.25—1.50 volutions before developing spiral sculpture.

Animal: Radula littorinoid, 2-1-1-1-2; rachidian narrow, but with bulbous base having 3 basal denticles; lateral tooth with large central cusp and smaller lateral cusps and with distinct littorinoid embayment in which inner marginal tooth ar-

VOLUME 95, NUMBER | 73

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ww SZ

Fig. 3. Echininus viviparus: A, 2 with well developed young in expanded oviduct seen through dorsal mantle; B, Detail from A: shell of young snail of about 1.5 whorls; C, d showing arrangement of penial glands. All from northeast coast Tinian, Mariana Islands (USNM 796244).

ticulates; inner marginal with 4 cusps of which third from medial is largest; outer marginal with 4—5 cusps increasing in size laterally. Animal littorinoid; in the male penis is unbranched; wide and deep sperm groove runs along its posterior surface, the edges of which are brown pigmented; penis with 5-8 penial glands arranged in straight line along outer edge; well developed ctenidium present in

74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Fig. 4. Known distribution of Echininus species.

dorsal mantle cavity; leaflets well formed; efferent branchial vessel and osphra- dium present (similar to that shown in Fretter and Graham 1962, Fig. 7). Fecal pellets oval in shape, white with dark flecks. Female ovoviviparous with oviduct modified as brood pouch; containing as many as 50 or more embryos in various stages of development up to the crawling stage in which shell has 1.25-1.50 whorls.

Etymology.—Viviparus, an adjective, from the Latin, meaning ‘‘bearing active, living young.”’ }

Measurements.—See Table 1.

Types.—Holotype, USNM 792356; length 9.7 mm, width 7.4 mm; |5 paratypes, USNM 803296, ranging in length from 8.1—12.3 mm, and in width from 6.2—8.2 mm.

Type-locality.—Machong Point [=As Matmos or Dudu, personal communi- cation, L. G. Eldredge] (about 14°11’N, 145°18’E), Rota, Mariana Islands; in spray zone 30 feet landward from top of 50 foot cliff; 4 February 1979; L. G. Eldredge, collector.

Material examined.—Mariana Islands, West Pacific Ocean.

Guam: [1*] Agfayan Point (13°16'N, 144°44’E) high intertidal on limestone; July 1980, R. H. Randall, USNM 803297.

* Figure ‘‘1’’ refers to column | in Measurements Table 1; specimens from Guam are summarized together.

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Camel Rock (13°29’N, 144°42’E) at 20 foot elevation on limestone; 13 June 1980, L. G. Eldredge, USNM 803298.

Asan Point (13°29'N, 144°42’'E); 1951, D. B. Langford, USNM 613687.

Apra Bay (13°27'N, 144°38’E); November 1907, P. Bartsch, USNM PBB323.,

Guam, 29 August 1949, V. L. Haack, USNM 620383.

Guam, 1946, Capt. Draeger, USNM 707190.

Rota:

[2] Machong Point [=As Matmos or Dudu] (about 14°11'N, 145°18’E) in spray zone 30 feet landward from top of 50 foot cliff; 4 February 1979, L. G. Eldredge, type-lot, USNM 792356, 803296.

[3] West side Poniya Point (14°06'N, 145°10’E) above terraced bench in spray zone, 2 February 1979, R. K. Kropp, USNM 803299.

Tinian:

[4] Northeast coast (about 15°06’N, 145’39’E) in spray and splash zone of large blowhole, 100 feet landward from 20 foot cliff, 8 February 1979, L. G. Eldredge, USNM 796244.

Saipan:

[5] Puntan Agingan (15°07'N, 145°42’E) in depressions in limestone 6-10 feet landward from 25-30 foot cliff, 19 November 1980, L. G. Eldredge, USNM 803300.

[6] Puntan Magpi (15°16’N, 145°48’E) on limestone in spray of blowhole, 5-8 foot cliff, 21 November 1980, L. G. Eldredge, USNM 803301.

Ecology.—Lives in the high intertidal and supratidal areas on rough, pitted limestone apparently restricted in distribution by the extent of ocean spray; often in the vicinity of blowholes (Tinian) and sometimes found at considerable dis- tances landward from the tops of sea cliffs 20-50 feet high (Rota and Tinian) (L. G. Eldredge, personal communication) (Fig. 6).

Echininus viviparus lives highest above the sea of any littorinid that I have observed. It is described as living where it is only wet by spray from surf or the blowholes characteristic of coral shores, where it probably feeds on vegetation which is kept moist by the same spray. The ovoviviparous reproduction of this species appears to be in close accord with its high position on the shore and permits it to survive without a pelagic stage in its development. Careful studies need to be carried out before it can be stated unequivocally, however, that E. viviparus has no need to return to the sea for any part of its life history.

Geographical distribution.—Southern Mariana Islands: Guam, Rota, Tinian, and Saipan.

Comparative remarks.—Morphometrics of Echininus viviparus are compared with those of E. c. cumingi and E. c. spinulosus in Table 1. Note that E. viviparus is smaller than the other two species, reaching only 12.3 mm in maximum length versus 20.9 and 18.3 mm respectively. On the average it is less obese, 0.76 versus 0.87 and 0.78, although difficulties of measuring spinose shells may artifically enhance the obesity of E. c. cumingi. Numbers of postnuclear whorls and rows

76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig.5. A, Apex of Echininus viviparus, showing smooth protoconch of about 1.25 whorls (compare with Fig. 3B), specimen from northeast coast Tinian (USNM 796244), Bar = 100 um, 100x; B, Apex of Littorina saxatilis, another ovoviviparous littorine, showing smooth protoconch of about 1.5 whorls, specimen from Odiorne Point State Park, Rye, Rockingham County, New Hampshire (USNM 803282), Bar = 100 wm, 80x; C, Apertural view of young E. viviparus showing 2 major rows of blunt spines on body whorl separated by moderately strong intermediate row, same data as A, Bar = 250 pm, 40x; D, Apex of E. cumingi cumingi, showing decollate tip, specimen from Philippines (USNM 89449), Bar = 100 wm, 185x.

of tubercles are less in E. viviparus. The aperture length/shell length index is smaller in E. viviparus. These data show the new species is smaller in size and more slender than previously known species. It is also non-umbilicate where the other two usually have an umbilicus. There is a rather weak but persistent tooth- like bulge on the inner lip.

The animal of E. viviparus is typically littorinoid, having a blunt snout and tentacles with eyes at the outer bases. It is significant that females of FE. viviparus produce young ovoviviparously (Fig. 3A) because this has not been noted in other Echininus species. The penis of E. viviparus differs markedly from other species: in E. viviparus there are 5-8 penial glands in a straight line along the outer edge of the unbranched penis (Fig. 3C), while in E. c. cumingi, the glands occur only

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at the junction of the bulbous base and the penial extremity, laterally and pos- teriorly (Rosewater 1972: plate 406, fig. B). The arrangement in E. c. spinulosus is similar to the latter. Radulae of the three species of the genus Echininus are similar, with differences in some details. In all species the teeth are small (see magnifications, Fig. 1) and reminiscent of the ‘‘pick type’’ (Rosewater 1980). Rachidian teeth of all three species of Echininus are narrow distally, but in E. viviparus the base is bulbous with 3 denticles (Fig. 1B). The tip of the protoconch is frequently present in E. viviparus, but is smaller and often decollated in E. cumingi and E. spinulosus (Fig. 5).

Discussion.—When | initially mapped the distribution of Echininus (Rosewater 1972, plate 407), information from museum specimens indicated that the ranges of the two subspecies, FE. c. cumingi and E. c. spinulosus, were divided into northern and southern components by an east-west line drawn just south of Puerto Princessa, Palawan, Philippines (9°N). Some additional records have come to light for E. c. cumingi: Guimaras Island, Philippines [verifying the type-locality of E. cumingi] (10°35'N) (Kevin Marx, personal communication 1979); Lifou, Loyalty Islands (C. Lamb 1979, USNM) not very far from the Tana, New Heb- rides record reported by Rosewater (1972). The discovery of the new species, Echininus viviparus, changes the pattern of distribution of the genus Echininus in the Indo-Pacific (Fig. 4). Although obviously related generically, the Mariana species is quite distinct from the other Echininus taxa in its distribution, in certain aspects of its morphology and, so far as is known, in reproducing ovoviviparous- ly. In my previous evaluation of a few small, worn shells from Guam (Rosewater ibid.) I referred them to E. c. spinulosus because no anatomical evidence was available to the contrary. The additional freshly collected specimens provided by L. G. Eldredge have shown without doubt that E. viviparus is a separate species distinct from the subspecies group of E. cumingi. It is rather difficult to speculate upon the origin of FE. viviparus. Little fossil evidence exists to trace the evolu- tionary history of the genus Echininus or, in fact, of any members of the family Littorinidae. Some littorinid fossils may have been assigned to such families as Turbinidae and Trochidae which are almost indistinguishable as fossils. The most similar in appearance is the fossil, Tectarius rehderi Ladd, 1966, from Lower Miocene, Marshall Islands (Ladd 1966, Rosewater 1972, pl. 404, figs. 5-7). Ac- cording to Ladd (1960) much of the mollusk fauna now inhabiting west Pacific islands and the East Indies may have migrated in successive waves from east to west during Cretaceous and Tertiary times when an archipelago of islands existed through which species could move with relative ease. Such species as Tectarius rehderi may be the precursors of Recent Echininus and Tectarius species now inhabiting the western Pacific arc. It seems unlikely that the fossil species Echin- inus adelaidensis (Cotton, 1947) from the Pliocene of South Australia belongs in the Littorinidae and probably should be referred to the Trochacea. In the absence of data indicating a more widespread distribution, it appears that Echininus vi- viparus has evolved as an endemic species in the Mariana Islands where its ancestors became established during one of these earlier migrations. Other lit- torinid inhabitants of the Mariana Islands fail to show any indication of such endemicity (Roth 1976, Rosewater 1970, 1972).

Echininus viviparus joins the small but increasing group of mollusks whose life histories are known to include ovoviviparity, or brooding of young by the parent

78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ie oe Sa is

Fig. 6. A, Machong Point, Rota Island, Mariana Islands, habitat of Echininus viviparus 30 feet landward from top of 50 foot cliff; B, Same, showing spray reaching habitat; C, Same, showing pitted limestone; white dots are Echininus; Bar = 20 cm; D, Northeast coast Tinian, Mariana Islands, where Echininus lives 100 feet landward from top of 20 foot cliff. (Photos courtesy of L. G. Eldredge.)

through early stages of development. The occurrence of this form of reproduction in mollusks was reviewed by Van der Schalie (1936) who attributed its develop- ment to being of survival value for animals subjected to unfavorable environment. Ovoviviparity now is known to occur in several additional groups to those cited