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NVINOSHLINS SSIYVYEEIT_ LIBRARIES SMITHSONIAN _ NO (Je) = w = were w” = w = = = y a WN = < = = ra —_| ey J, = SS \ \, =| <= = 9 3 Of 5 QS z o -— 9 z g Ugg = oN 3 = oa > = > = ios Ss > = (2p) es 7p) i z W) ae Jia (SMITHSONIAN INSTITUTION NOILNLILSNI_ _ tt z ul ie uw a ul & a ow (sa w a o ce 3 oe a a — = al < a < ra < c = oe s cc S oc = fee o 2 rs e ra a fe) ae ina Ven zm a Zz See & SM ON Licrel : catpemt UPL AC GN A 6 oes A ee OAR UARI TT INCRE ON, NO ; ria } Wisk ee PROCEEDINGS of the Biological Society of Washington VOLUME 103 1990 Vol. 103(1) published 21 March 1990 Vol. 103(3) published 25 September 1990 Vol. 103(2) published 28 June 1990 Vol. 103(4) published 19 December 1990 WASHINGTON PRINTED FOR THE SOCIETY EDITOR C. BRIAN ROBBINS ASSOCIATE EDITORS Classical Languages Invertebrates GEORGE C. STEYSKAL STEPHEN D. CAIRNS FRANK D. FERRARI RAFAEL LEMAITRE Plants Vertebrates DAvip B. LELLINGER G. DAvip JOHNSON Insects WAYNE N. MATHIS All correspondence should be addressed to the Biological Society of Washington, Smithsonian Institution Washington, D.C. 20560 ALLEN PREss INC. LAWRENCE, KANSAS 66044 OFFICERS AND COUNCIL of the BIOLOGICAL SOCIETY OF WASHINGTON FOR 1990-1991 OFFICERS President LESLIE W. KNAPP President-Elect STORRS L. OLSON Secretary G. DAVID JOHNSON Treasurer MICHAEL VECCHIONE COUNCIL Elected Members ROGER F. CRESSEY, JR. JANET W. REID JANET R. GOMON WAYNE C. STARNES ROBERT HERSHLER JEFFERY T. WILLIAMS TABLE OF CONTENTS Volume 103 Adkison, Daniel L. and Sneed B. Collard. Description of the cryptoniscium larva of Entophilus omnitectus Richardson, 1903 (Crustacea: Isopoda: Epicaridea) and records fromthe: Grullf Of MIGX1CO css 0 a ee er ta Alvarez, Fernando and Jose Luis Villalobos. Pseudothelphusa galloi, a new species of freshwater crab (Crustacea: Brachyura: Pseudothelphusidae) from southwestern Anderson, William D., Jr., N. V. Parin, and John E. Randall. A new genus and species of anthiine fish (Pisces: Serranidae) from the eastern South Pacific with comments on anthime ‘relationships2..c304 0 a2 eee ee ee ee ee ee Baba, Keiji and Janet Haig. A new species of chirostylid crustacean (Decapoda: Ano- mura) from off the west coast of North America ..........c.ccccccccsccccccsecssccccsecseeseeseeseeeseseesesseeseeseeeeeeseseee Baba, Keiji and Sang-Chul Oh. Galathea coralliophilus, a new decapod crustacean (An- omura: Galatheidae) from Singapore, Gulf of Thailand, and West Irian... Banta, William C. and John C. Redden. A checklist of the Bryozoa of the Galapagos... Barnard, J. L. and James Darwin Thomas. Ensayara jumane, a new species from Belize, Caribbean Sea (Amphipoda, Lysiamassic ae) 2... ccceeeeeceeeeeeeeeeeeeeeeeeeeeeseeeeveeeteeseeeteeeeteteteteenceeneene Bayer, Frederick M. A new isidid octocoral (Anthozoa: Gorgonacea) from New Cale- donia, with descriptions of other new species from elsewhere in the Pacific Ocean..... Bayer, Frederick M. The identity of Fannyella rossii J. E. Gray (Coelenterata: Octoco- MANTA atten ee 2a MF IN ae ec OP RRM SRE Ect ae oe eee ee Bernard, Ernest C. New species, clarifications, and changes in status within Eostentomon Berlese (Hexapoda: Protura: Eosentomidae) from the United States... eeececccccccnee Blake, James A. A new genus and species of Polychaeta commensal with a deep-sea Gy aASar il Cl earn he Oe ei Se Da aia eee aoa Us aR Rr ep Bogle, Mary Ann. A new species of Cladocarpus (Cnidaria: Hydroida: Plumulariidae) fron the: Strarts rots Elo ric ae ese ae ee Pein eee oa ea eon oak ce Browning, M. Ralph. Taxa of North American birds described from 1957 to 1987 ...... Browning, M. Ralph. Erroneous emendations to names proposed by Hekstra (Strigidae: Cadle, John E. and Roy W. McDiarmid. Two new species of Centrolenella (Anura: Centrolenidae) from northwestern Peru e........-cccccccccccccccccccecceccesceeeeeseeeeeeeeeeseeeseeeeeesesseesseesesteteessestesstttevestesteee Camp, David K. and Hans G. Kuck. Additional records of stomatopod crustaceans from Isla del Coco and Golfo de Papagayo, east Pacific OC an i.....ccccccescccccecseeeesecceeeeeeeeees Campos, Ernesto. Calyptraeotheres, a new genus of Pinnotheridae for the limpet crab Eabiajeranti Glassell) 1933) (Erustacea:) Brachyilra) see Campos, Ernesto and Alma Rosa de Campos. Taxonomic remarks on Schizobopyrina Markham, 1985, with the description of S. bruscai (Crustacea: Isopoda: Bopyridae) Casanova, Jean-Paul. A new species of Paraspadella (Chaetognatha) from the coastal waters Of Japan 02a sues on aye Pi Rt eee Oe Re DERE nna nes et Pere OR Castro, Ricardo M. C. and Richard P. Vari. Moojenichthys Miranda-Ribeiro (Pisces: Ostariophysi: Characidae), a phylogenetic reappraisal and redescription .....----s------- Child, C. Allan. Prototrygaeus jordanae, a new species of pycnogonid from Monterey Bay 5 Catach rer eh es 0 eA ela RS oe a Child, C. Allan. Pycnogonida of the Western Pacific Islands, VIII. Recent collections from islands of the Great Barrier Reef, Australia cc ccccccccccccccccccccecceceeeeceeeceneceeeteeseeeeee Cutler, Norma J. and Edward B. Cutler. A revision of the subgenus Phascolosoma (Sipunculas Phascolosoma)e. en ee oe Nae eee ee Delaney, Paul M. Tridentella williamsi, a new species of isopod crustacean from the British Virgin Islands, western Atlantic (Flabellifera: Tridentellidae)... ccc Donnelly, Maureen A., Craig Guyer, and Rafael O. de Sa. The tadpole of a dart-poison frog Phyllobates lugubris (Anura: Dendrobatidae) 649-654 103-105 922-930 854-860 358-363 789-802 120-126 205-228 773-783 861-890 681-686 229-234 432-451 452 746-768 847-853 364-371 633-642 907-912 525-542 157-160 311-335 691-730 643-648 427-431 Ernst, Carl H. and Jeffrey E. Lovich. A new species of Cuora (Reptilia: Testudines: Emydidae)iiromithesRyukyullsland see Erséus, Christer, Jarl K. Hiltunen, Ralph O. Brinkhurst, and Don W. Schloesser. Re- definition of Teneridrilus Holmquist (Oligochaeta: Tubificidae), with description of two new species fromm North America i cececceceeeseeeeeeeeeeeeeeeteeenennnnnnnnnnnnnncnnnnnneneeceseneceeneeeeeeeeeeeseseee Escobar-Briones, Elva and Luis A. Soto. Heteromysis mexicana, a new species from Campeche Bank, Gulf of Mexico (Crustacea: Mysidacea) 0. ceeeeceecescseeeeececeeceneeeeeeeeeeennnees Fitzhugh, Kirk. Fabricinuda, a new genus of Fabriciinae (Polychaeta: Sabellidae).............. Gardner, Alfred L. and Carolyn S. Ferrell. Comments on the nomenclature of some neotropical bats (Mammalia: Chiroptera) 22... ccc ceecccecenneeeeeeeeeeeeeeeeesnnennnnteteeeceneeeneeeeeneee Gonzalez, Sergio A. and Mario E. Edding. Achelia assimilis (Haswell, 1884) in the Heterozostera bed of Puerto Aldea, Coquimbo: first record from the northern Chilean coast (Pycnogomida: Armmotheicae) cece cceeceeeeeeeeeeeeeeeeeneeeenttententnnttnttttecectntnnte Graves, Gary R. Systematics of the ““Green-Throated Sunangels” (Aves: Trochilidae): ave ta xen Relay ED LS). sash SI a ld oe i el wa Es ae el Graves, Gary R. A new subspecies of Diglossa gloriosissima (Aves: Thraupinae) from theswesternvAndes;ofiColombiai kei se ee Handley, Charles O., Jr., and Alfred L. Gardner. The holotype of Natalus stramineus Gray (Mammalia: Chiroptera: Natalicace) 2200000000 eect eeeeeeeeeeeeenennnneeeeeeeneneeettnnnnnnneeeeeee Harold, Antony S. Redescription of Polyipnus fraseri Fowler, 1934 (Teleostei: Stomi- iformes: Sternoptychidae), with remarks on paedOmMoOrphosis eee eee ee eeeeeeeeee Harrison-Nelson, Elizabeth and Thomas E. Bowman. A range extension to the north for Macrochiridotea giambiagiae Torti and Bastida (Crustacea: Isopoda: Valvifera).... Hendrickx, Michel E. Range extension and host record for Dissodactylus ususfructus Griffith, 1987 (Crustacea: Brachyura: Pinnotheridae) cece eeceeeeeeceeecceeeeeeeeeeeeeeteee Hershler, Robert. Pyrgulopsis bruneauensis, a new springsnail (Gastropoda: Hydrobi- idae) from the Snake River Plain, southern [abo occ ececeecccneeeececeeedeneeeeseseeseneeseeeeeeeeeeeeees Hershler, Robert and William L. Pratt. A new Pyrgulopsis (Gastropoda: Hydrobiidae) from southeastern California, with a model for historical development of the Death Walleyiry drographic System ses a eee Hershler, Robert and Fred G. Thompson. Antrorbis breweri, a new genus of hydrobiid cavesnail (Gastropoda) from Coosa River Basin, northeastern Alabama... Hershler, Robert, J. Malcom Pierson, and R. Stephen Krotzer. Rediscovery of Tulotoma magnifica (Conrad) (Gastropoda: Viviparida) occ cceeecece cece ceeeeeeeeeeeeeeeeeeeeceeeeeeeeeeeetetcnnnnnee Hobbs, Horton H., Jr. On the crayfishes (Decapoda: Cambaridae) of the Neches River basin of eastern Texas with the descriptions of three new SPeCieS eee eee Hobbs, Horton H., Jr. and H. H. Hobbs III. A new crayfish (Decapoda: Cambaridae) frommsoutheastermmMlexas ees esl ec UE NE ee oe James, Samuel W. Diplocardia kansensis, a new earthworm from Kansas, with redes- criptions of D. riparia Smith and D. fuscula Gates (Annelida: Oligochaeta: Megascole- Jazdzewski, Krzysztof. A redescription of Tiron antarcticus K. H. Barnard, 1932 (Crus- tacea: Amphipoda: Synopiidae) with an updated key to the species of Tiron Liljeborg, ARS (Gs pe eae rt eae soe i ae a als ROR ee ee ACS IR Kathman, R. Deedee. Some tardigrades from Colorado, with a description of a new species of Macrobiotus (Macrobiotidae: Eutardigrada) 2.0... Kensley, Brian and Richard Heard. The genus Axianassa (Crustacea: Decapoda: Thalas- Sinidea) ringthep ATM Eni ca Sp seas a UI hee al EY SE ae Kropp, Roy K. and Jane H. Dominguez. Mesacturus dicrurus, new species, an unusual stomatopod from Micronesia (Stomatopoda: Gonodactylidae) eee eeeeeeceeeeeeee Lambert, Philip. A new combination and synonymy for two subspecies of Cucumaria fisheri Wells (Echinodermata: Holothuroidea) .......::-::::ccssss-ssssssssecsssseeeeeeeesseeveeeceeseeeeeeeeeeesutneneceeeeeeee Lynch, John D. A new ocellated frog (Centrolenidae) from western Colombia..................... Macpherson, E. The status of the caridean shrimp Pandalina modesta (Bate, 1888) (Crustacea: Decapoda: Pandalidae) with redescription of the SpeCl@S 222.2222. Mahadeva, Madhu N. and Thomas A. Munroe. Three new species of symphurine tongue- fishes from tropical and warm temperate waters of the eastern Pacific (Symphurus: Gynoglossidac- Pleuroneciformes) = eee 26-34 839-846 131-139 161-178 501-508 151-156 6-25 962-965 966-972 509-515 127-130 106-107 803-8 14 279-299 197-204 815-824 573-597 608-613 179-186 110-119 300-303 558-572 372-375 913-921 35-38 353-357 Malinky, John M. Solenotheca, new hyolitha (Mollusca) from the Ordovician of North PAGTV ET CAs! Ne aN a IE eet ei fe Mangum, Charlotte P. The Fourth Annual Riser Lecture: The role of physiology and biochemistry in understanding amimal phylogeny ..........eeccceecesssssesssseeeeseeeeseeeteeceeccecceceeeeeecececceceeeeeeeeee Manning, Raymond B., Elliot W. Dawson, and W. Richard Webber. A new species of Chaceon from New Zealand (Crustacea: Decapoda: Geryomnidae) ........ccccceeccceeeeeeeeeceeeeeeeeee Mathis, Wayne N. and Tadeusz Zatwarnicki. Taxonomic notes on Ephydridae (Dip- Menezes, Naércio A. and Stanley H. Weitzman. Two new species of Mimagoniates (Teleostei: Characidae: Glandulocaudinae), their phylogeny and biogeography and a key to the glandulocaudin fishes of Brazil amd Paraguay. ccesscssccsccccccccccccccceeccseeeeeeeeeeeeee Mooi, Rich. Living cassiduloids (Echinodermata: Echinoidea): a key and annotated Nakamura, K. and C. Allan Child. Pycnogonida of the Western Pacific Islands, VII. On some rare species from the Flores Sea, [mGomesia a... ccc eeeeeeeeeeeeeeeeeneeceeeececnnnnennenenteeeee Ng, Peter K. L. Luteocarcinus sordidus, new genus and species, from mangrove swamps in peninsular Malaysia (Crustacea: Decapoda: Brachyura: Pilumnidae: Rhizopinae).. Ohwada, Takashi. Redescriptions and synonymy of Nephtys imbricata Grube, 1857 (RolvchaetaniNe ph tye 24a ee UN ca ale a oi OA Cao Perrin, William F. Subspecies of Stenella longirostris (Mammalia: Cetacea: Delphin- Pettibone, Marian H. New species and new records of scaled polychaetes (Polychaeta: Polynoidae) from the Axial Seamount Caldera of the Juan de Fuca Ridge in the northeast Pacific and the east Pacific Ocean off northern Califormia.. Poss, Stuart G. and Bruce B. Collette. Scorpaenodes immaculatus, a new species of scorpionfish (Osteichthyes: Scorpaenidae) from Walters Shoals, Madagascar Ridge... Randall, John E., Thomas H. Fraser, and Ernest A. Lachner. On the validity of the Indo-Pacific cardinalfishes A4pogon aureus (Lacepéde) and A. fleurieu (Lacepéde), with description of a related new species from the Red Sea o.............:-::cscseseeeseeseeeeeeeeeeeeeeeeeeeeeeseceseettee Reed, Edward B. Tachidius incisipes Klie and other harpacticoids from northwestern Canada '\(Crustacea:;Copepodal) ii c2) ee ee ee ee Reid, Janet W. Redescription and new records of Trichodiaptomus coronatus (G. O. Sars) (Copepoda: Calanoida: Diaptomidae) from Brazil ccccccccccesccecececeseceeeeeeeeeeeeeeeeeeeeeeeee Reid, Janet W. Canthocamptus (Elaphoidella) striblingi, new species (Copepoda: Har- pacticoida))fromeC@ostavRica esd is Sich aN le os linea la Reid, Janet W., Javier A. Molina Arevalo, and Manuel M. Fukushima. Metacyclops leptopus totaensis, new subspecies (Crustacea: Copepoda) from Lago de Tota, Colom- NA Ee a ay UG a a A yl ae ER Cc wera a Reiswig, Henry W. Correction of Ijima’s (1927) list of Recent hexactinellid sponges (Reo reise ean) ea Se I Le a ae EE a cL i es Robinson, Harold. Six new combinations in Baccharoides Moench and Cyanthillium Blumer (WVernonieaeVASte race ae) ie ie ei Nile a sen On elton posse eI Robinson, Harold. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). VII. Wbheseenussivepidap oa se AC Gs ole ee Ross, Charles A. Crocodylus raninus S, Miller and Schlegel, a valid species of crocodile (Reptilia: Crocodylidae) from Bormeo 2... eeeteeneeseesesnessene Roux, Ana Marta and Ricardo Bastida. The occurrence of Sphaeroma serratum (Fa- bricius, 1787) in the western South Atlantic (Crustacea: [sopoda) .....::::cscccssssecessseeseeeeeeee San Martin, Guillermo and Julio Parapar. Exogone acerata (Exogoninae: Syllidae: Poly- chaeta), a new species without antennae from the Mediterranean Sea... Schuster, R. O., T. L. Tyler, J. A. Skinner, and E. A. Sugden. The primary types of the Richard M. Bohart Museum of Entomology, I. Tardigrad a... cccc-cccccccsseessnsssseeeeeseeeeeeeees Spangler, Paul J. A new species of halophilous water-strider, Mesovelia polhemusi, from Belize and a key and checklist of New World species of the genus (Heteroptera: Mesove- ING Ge) RA AMR reaper f pue 1 oes LE NR AIT ase ONE Ree mee LE Ea ee 265-278 235-247 602-607 891-906 1-5 380-426 63-85 304-310 95-99 187-196 453-463 825-838 543-549 39-62 341-349 140-150 336-340 674-680 731-745 248-253 464-498 955-961 350-352 687-690 376-379 86-94 Stock, Jan H. and Young Won Jo. The Japanese amphipod genus Eoniphargus, redis- Covercdiinva SOUTHNOKeAaN CAV Ce ye ee ee ee Thomas, James Darwin and J. L. Barnard. Gitana dominica, a new species from the Caribbean Sea (Amphipoda: Amphilochidae) cc eeeeeeeeeceeeececceeeeeeeeennnnnnnnnneceseeseeeseeeeee Vari, Richard P. and Ramiro Barriga S. Cyphocharax pantostictos, a new species (Pisces: Ostariophysi: Characiformes: Curimatidae) from the western portions of the Amazon Vari, Richard P. and Darrell J. Siebert. A new, unusually sexually dimorphic species of Bryconamericus (Pisces: Ostariophysi: Characidae) from the Peruvian Amazon............ Westheide, Wilfried. Meiopriapulus fiyiensis Morse (Priapulida) from South Andaman, another example of large-scale geographic distribution of interstitial marine meiofauna Wetzer, Regina. A new species of isopod, Aega (Rhamphion) francoisae (Flabellifera: Aegidae), from the cloaca of an ascidian from the Galapagos Islands... Wicksten, Mary K. On the status of Alpheus barbara Lockington (Crustacea: Caridea: JANID 0) GY EEG FAYE) a ie a a IL EC ee DL Pee SNC Williams, Austin B. and E. E. Boschi. Panopus margentus, a new crab from the Argentine warm temperate subregion (Decapoda: Kamthidae) eee cecececeeeeeecnnntttetee Williams, Austin B. and John J. McDermott. An eastern United States record for the western Indo-Pacific crab, Hemigrapsus sanguineus (Crustacea: Decapoda: Grapsi- Williams, Austin B. and Nguyen Ngoc-Ho. Pomatogebia, a new genus of thalassinidean shrimps from western hemisphere tropics (Crustacea: Upogebiidea)... Ziatarski, Vassil N. Porites colonensis, new species of stony coral (Anthozoa: Scleractinia) olfsthetCaribbeantcoastiofeRanania se eee Zullo, Victor A. Supplementary description and phylogenetic significance of Arcoscal- pellum conradi (Gabb) (Cirripedia: Scalpellidae) from the Paleocene Vincentown for- FOREN AKON AY: COPE INCE TALS STEN I eS ee aod cet tae 624-632 617-623 550-557 516-524 784-788 655-662 100-102 598-601 108-109 614-616 257-264 663-673 INDEX TO NEW TAXA Volume 103 (New taxa are indicated in italics; new combinations designated n.c.) COELENTERATA Hydrozoa (CLAM OCA US HLS TCE tis a NI el OO Poe 229 Anthozoa Acame ll ar@ispar fees ek 220 Ne IMT J au oe Ne ed Ri ea AU Mh ee rae ven ade ed UE 215 Tsidelllaesnt Ch OL rye ee en Sra ee ec 208 OOVSEOTTIIS TSP a Ea 8 ah CSA kB Nr a 219 COST ULC vcs tee Ns Bag Bk le JON A Sa sa RR eee ee RR 219 J XO) wt 66) (0) 71-1 KY kee AN Tbe EL A at Po sal 1 cel RNA re en ne De\T MOLLUSCA Hyolitha SS OLCTIOL MCC ee oe NN I I TO BE UM eh, ep 0 a lA ee A sR Se AEE ee 269 DOK C VC ae NIA BU ASE Oa eo IE ae fgets CLEARER EOE NP 269 Gastropoda PAT AEIO TID US essere ae TORT IP GUE SB ee BEATIN Weal ets 192) ep 197 DY OWC If] Pe a etd LD LEE et ee ee ee a SSS ETD 198 Py reul OpSiSvOrUuneaue nists 0s 2 es ese ee ei a eae a tee Stee 803 SESUVNEVO WH ites tee NIA lel Re et tro «RUE Ione erase EOE AY NO Dan eno 8 IU co 2 IN A oa 279 ANNELIDA Polychaeta EX OP OMEN CCH ee Ne TN MES Be fs eS Bg NDR I el en 687 Fabricinuda 162 pseudocollaris 172 IDSCULOD GID BaP Ne SFO ETN AN Tk 200 87 MRO CR te a UR eRe 171 [ODUCT OUI oY eM eaeeyeren oot encore Wesediittin Uses ERM nec lee ne Connie nN enter es eS pt ete lee ee 165 Inminicolasnte: syrah 0a ge te TAS ere aes se 163 trilobata n.c. ....... 170 825 828 834 IP AT AD GUY NO CMR oe Tee Sta EN let es pe ca ce 830 brisinga 831 (PCV CCCQ ae : 681 thyasira 684 IDiplocardiatkanseris isin seco. wees sty Pa eh EMT acted se ele eee IA deca ld 179 ETRY Tad GL TeV TUS CCL V 10S se et aa ae URN SUE ee bee ev a eet Oe 841 841 842 TARDIGRADA Eutardigrada ARTHROPODA Pycnogonida PNrrTnn yo lve | heap 79 Ode Ce Mea ee ere UN A Ee eit re aN ee eal 313 PATO DIO GAC ATV IUS HDC Ce Meee tei eee eA Set ec cae Ree wae era ce ee IE 327 iN TymenpeL ONNS CACY I pean Rae ee 324 Prototrygaeus jordanae........... 157 Rhynchothorax vallatus 333 Seguapallene crassa.................... 321 BIRATAY SV UUITIRE 2 CLS WVCLL Lees a na a en ree RE eee occ en oA OA See Ste rae clin eee 317 Crustacea Negan(hamphiom) if iQNCOUS Ce xe tee ee eer laa 655 Axianassa arenaria 563 OT IYOH ES 5B 3S IS Ae eh at ko re ee 567 Wj LEY UCLUGOTAS US foe sene ale rs ante ee cee enc A I TOU PD a0 a oa acre acta ee eats 566 CIV DEF ACOLE TCS meantime ee eaten ILO EL aE MLE MRE Sg IR See Ie ee ae oO eee at ee ON DD 365 FEENEY SOUS a OSI ad ek Ro A een ee 365 Canthocamptus (Elaphoidella) striblingi.. 336 Chraceon ald wy 1 cece ceeeeeeeeveeeveveeseeverererrnenn ’ : 602 EOniphangusyelandulatus ee ee 624 | EW ANSEE SVEN EY jf UA (LOM AUS x eee LR ie ee 121 Galatheatcoral top iil tis Beets aU Ha IEE ane Sok oe ee he ete ee Sd ee 358 (TUTE OY YIN KEL DNS GZ AYN lA a 854 Gi CATAL OF TT TLTC Cl eet tesco nc ese) eRe sed, SOUR Root rhe UO Pas fee ere ahd ep Hie 618 Heteromysis mexicana 131 PETE CO COU CUILUUS fe tere er tle TIES Sat Scr RD Sad ae NaS taser tae en 95 IS CO CLL UES Mame cee PN aE PER ROE Nise ek EO ne en ns ea eae 98 INAS SA CUUTUIS CLE CITE TEU Syme estaate aes RSAC re ON cee rec etn Nee eda ste Sn ase aor 372 IMetacy clopsileptopuSitOLae rts ts esc e esse ee ee ete rece 674 Panopeus margentus 598 IROMMALO LCDI Gee ene ae 615 Procambarus (Girardiella) kensleyi 583 TUL SHOCLTACLUS es ae ea cre AN el eae el ee AU gO i pe ei eee Ie eee Rete eT See ia 577 Procambarus (Ortmannicus)) 7echesae ee 590 ZOTIA TLS ULES tess eae EARL ek, et hon ung aie Neen et ne Na wie! ew ES Aen ah se VA a Ree me a are, 608 Pseudothelphusa galloi 103 Schozobopyrina platylobata n.c..... 637 DYGUES C Claman ie are Uhre MRD Rie ode AIEEE Se RN Dec LR a ee Bae, Er been a 638 Mera erated ay y71 0 [car ras ieee se ete ances eet aoe ee ned tea le Ie a ee 643 Insecta Athyroglossa (Parathyroglossa) QitOr ate .cc..........-vevevvvvvvvnnnvnvnnvvvnsovnvvonssosesssvsssseseeseseseseeseeeseseeseseseesesssessssessisstssssnsoee 893 Atissa subnuda n.c............... 896 Eosentomon crypticum ..... ie 863 CLV; OLY CL rae ate ces aa SLE er re a cine bd ee 868 TULL LDV GLLLT Yate eset caatoest ere eater etre aa oe) Ri ee Ne ee hs Ce 870 OSL COT UT rare ties a ane ta ed We sehd gs oR a i ce Bh 872 CLIC C Cp een ae ee eI onion Ma ol ERNE Eee AAR TERRE es enh oe ge Ute 877 richardi 879 IS CLV CLTLTACLITC TAS Cem ie ee ora a ae et eons Oe ae ie Pe 881 IS TLLCLC], [amore met POM R Ase De MeO RS i Bree at Aoi, Se ie fia ah 884 ECHL TTY S | CX em ae as Loca OO ot SMA hi «Let Leas ena he a pee ta 885 INTESOVELIARD OLE TYTTLS meee amnesic rah ets LAL Naa ace cee es ea MC SOL NC MEARE. cree ea 87 Es Eant hy Serpe Wy at Trl oe eee eae eee eres eRe ese oe ae te ND i a ae 901 S.C hem asain ur ria pr ce eeeee aera eee ete ete eens ee Ce 896 CHAETOGNATHA PaaS (A LL AB 2 C77 a as ee Mek PANU se NS blotches 907 ECHINODERMATA IPSeud OCMUSsaSi gy eat US Me Coes ae ee 915 CHORDATA Pisces ANGLOLANETATAS: Bosca oP coo OS eR ce Be sat SR Ta Re en ee 923 CUPUO TOY CLO Ire ih ka cp Sa ae iE ee 924 ND OS OMG SCLLO TA sik ea oT ea ool 57 BEY COMAIMEDICUS: PCCLUT GLUES ec. deacc rte IER cs Pe UI alse or le ee 517 Cyphocharax pantostictos 551 Mimagoniates rheocharis 399 SY VEC OU Mee ea ei ai 8 yee Nak EO cs RMN Eee ear 387 SCORPACTIOG ESTE T11LCUE AETES ae zr Ne 2 oS ek ecto ac 543 Syrup S1C/2 GDA 1a ia eat a cna Ia ace 939 (1101 0X0) | 161) Ke .. 949 oligomerus 932 Centrolenella euhystrix ................... Fe eM oat ce 1 AOL ee re oA a 748 hesperia 753 DS NOHO ce Wace eta ELA AE Wh eed ER eco Med, ACER co Pa BE SN 35 WO nate Veliy 11 eye oe has ied oe are ie Oe ee aT A= EE etc eg a et 31 Aves Di glossaxelOrlOSiSSiTMa-DOy ete eet ee 962 Mammalia StenellaWOnginostrsKCes17OCA rier CCT Clee eee eee a rele evn ee 460 LONPITOStEIS KORTE TALL iS is Ae Pe i Naa eR ee a SUF GUL ee a On ee 457 PLANTAE ACC AL OLA CS sears Nee eI NS aN SA ee) a 248 (for three new combinations see p. 250) @y ext ten NT Se steer NS Ue AeA, Sy 6 at OO ee 248 (for three new combinations see p. 252) Wepidaploarbec kit eee se Sn oe oe : 482 SOL ONION A erect nner so M RINGS STS SILI BS 5 a i a Ne ee 493 (for 115 new combinations see pp. 480-496) _ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ” THE BIOLOGICAL SOCIETY OF WASHINGTON 1988-1989 Officers President: Kristian Fauchald Secretary: G. David Johnson President-elect: Leslie W. Knapp Treasurer: Michael Vecchione Elected Council Gary R. Graves Meredith L. 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Correspondence dealing with membership and subscriptions should be sent to The Treasurer, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, National Museum of Natural History, Washington, D.C. 20560. |___THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 1-5 THE EXTINCT SLOTH, MEGALONYX (MAMMALIA: XENARTHRA), FROM THE UNITED STATES MID-ATLANTIC CONTINENTAL SHEELF,) THS H. Gregory McDonald and Clayton E. Raj Abstract. —A radius of the ground sloth, Megalonyx, fro 3 New Jersey, is the first evidence that ground sloths, along with ma‘ fy & f 1AM 2 pe O( 5 MAR 5 0 (990 y, aff thenepanteOl istodorns anid mammoths, inhabited this region during glacial lowering of sea level in the Pleistocene. During the Pleistocene large areas of the continental shelf were exposed by the low- ering of sea level and were inhabited by the terrestrial Pleistocene fauna. Subsequent sea level rise has made recovery and documen- tation of the species that inhabited this re- gion difficult. Most discoveries of vertebrate remains from the continental shelf are ac- cidental; hence, the currently known diver- sity of terrestrial vertebrates from this area is low. Evidence of a ground sloth from the continental shelf contributes to our knowl- edge of the faunal diversity of this area dur- ing the Pleistocene. In July 1966, Ronald Stires of the trawler Kingfisher recovered a partial left radius of the ground sloth Megalonyx while dredging on the continental shelf off Sandy Hook, New Jersey. Location of the specimen was 40°16.5'N latitude and 73°54.5’W longi- tude. The specimen was recovered from a bottom depth of 19 meters. Description of specimen. —The radius (USNM 25175) (Fig. 1) lacks the distal end; the proximal end is eroded. The specimen is dark in color and well permineralized. Breakage of the distal end is angular, with some rounding of the edges of the breaks. Part of the articular surface of the proximal end is preserved but the perimeter of this area has weathered, exposing trabecular bone. There is a slight amount of weathering of the overall surface of the bone. These modifications preclude satisfactory mea- surements of original dimensions, but the following permit an approximation of the size of the bone: anteroposterior diameter of the proximal articular surface—53 mm, mediolateral diameter of proximal end—51 mm, anteroposterior width of shaft, distal- ly—88 mm. Despite the fragmentary nature of the specimen, enough is preserved to show the distinctive sigmoid curvature of the shaft, characteristic of the radii of megalon- ychid sloths, in which the distal end is offset anteriorly relative to the proximal end. The distal end of the shaft is compressed me- diolaterally relative to the proximal end, which is roughly equidimensional. The me- dial surface is slightly concave with a com- plementary convexity on the lateral surface. Preserved on the medial surface are the muscle scars of the m. pronator quadratus. Proximal to the offset of the shaft the bone is circular in cross section. The radial tuber- osity is large and well developed and offset medially on the posterior edge. There is no indication of an epiphyseal line on the prox- imal end, which, along with the general massiveness of the bone and rugosity of the muscle scars, suggest that it is probably from an adult. Biogeography. —Whitmore et al. (1967), Oldale et al. (1987) and Gallagher et al. (1989) have documented the distribution of mammoths and mastodons on the Atlantic 2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON S Cm Fig. 1. Megalonyx cf. jeffersonii, partial left radius, USNM 25175. A. Lateral view. B. Medial view. continental shelf of North America. Al- definitely can be referred to the genus Meg- though ground sloths have been mentioned alonyx, and is probably M. jeffersonii al- as part of the fauna recovered from the con- _ though, given the fragmentary nature of the tinental shelf, no genus or species has ever specimen, identification to species is ten- been given. The specimen described here uous. Megalonyx jeffersonii is the only late => Fig. 2. A. Approximate locality at which Megalonyx radius was recovered. Depth in meters. B. Cross section along 40°6.5’N latitude showing relationship of location of Megalonyx radius to shore and the submarine Hudson Canyon. Map modified from Veatch and Smith (1939). (Original depths for map and cross section from Veatch and Smith [1939] were in feet and fathoms.) VOLUME 103, NUMBER 1 Sloth Radius Sea Level Sloth Radius Vv Hudson Canyon = PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pleistocene species of the genus (McDonald 1977), and the size of the specimen is within the range of that species. Megalonyx is the most widespread of the North American ground sloth genera and was present in the northeastern United States at least as early as the Irvingtonian (Middle Pleistocene). Late Pleistocene records for the genus in this region are not uncommon (Parris 1983). so it is not unexpected that it would be among the megafauna migrating onto the continental shelf exposed by the lowering of sea level. The ecology of Mega/onyx commonly has been interpreted as that of a forest edge browser (Stock 1925). Utilization of gallery forests along rivers also has been postulated (McDonald & Anderson 1983). although the pattern could be biased by the fact that ver- tebrate remains are more readily buried and recovered along rivers than in upland hab- itats. As sea level lowered and rivers ex- tended onto the continental shelf Wega/on- yx may have utilized the expanding gallery forests if this was indeed its preferred hab- itat. The proximity of the location of the specimen described here to the Hudson River and its canyon on the continental shelf (Fig. 2) fits this pattern. As noted by Dillon & Oldale (1978). outflow of glacial meltwa- ter in the Hudson River Channel eroded a broad valley across the continental shelf un- til about 12.000 years B.P. The fragmentary and abraded nature of the specimen does suggest the alternative interpretation that the specimen does not represent primary burial of an individual inhabiting and dying on the exposed con- tinental shelf. The proximity of the speci- men to the mouth of the Hudson River and nearness to shore make secondary transport of the specimen a factor to consider. Because of the permineralization of the specimen no absolute date is possible. How- ever if the specimen represents primary de- position on the continental shelf and not secondary transport. then some inferences regarding the time of its presence on the continental shelf can be made. Although Megalonyx was the only genus of ground sloth to range as far north as Alaska (Stock 1942). this was probably during an inter- glacial phase and does not necessarily in- dicate that the genus was capable of better thermoregulation than other ground sloths (McNab 1985). Bloom (1983) states that the Atlantic Coastal Plain from lower New York Harbor eastward and northward was gla- ciated north of latitude 40°30'N. The prox- imity of the location of this specimen (40°16.5'N longitude) to Bloom’s boundary would place it rather close to the edge of the continental glacier during glacial makxi- mum. The location of the specimen is prob- ably too close to the ice front at this time for the animal’s thermoregulatory capabil- ities, based on our current understanding of ground sloth physiology. It is more likely that the presence of .\Vegalonyx on the con- tinental shelf post-dates the retreat of the continental glacier from the region. Whit- more et al. (1967) provide dates of around 11,000 years for intertidal salt marsh peat deposits on the Atlantic continental shelf. indicating subaerial exposure until this time. so the sloth remains may be as young as 11,000 years. Further confirmation of the presence and timing of ground sloths on the Mid Atlantic Continental Shelf will require the recovery of additional specimens. Acknowledgments This discovery would have remained un- known if Ronald Stires had not presented the specimen to the National Museum of Natural History. His generosity is greatly appreciated. We wish to thank William B. Gallagher et al. (1989) for allowing us access to their manuscript when “in press’ and for permission to cite it. The photos for Figure 1 were made by Victor Krantz and the figure prepared by Mary Parrish. Anita Buck kind- ly read early drafts of the manuscript and made many helpful editorial suggestions. Robert J. Emry and David C. Parris read and improved the penultimate draft. VOLUME 103, NUMBER 1 Literature Cited Bloom, A. L. 1983. Sea level and coastal morphology of the United States through the Late Wisconsin glacial maximum. Pp. 215-229 in H. E. Wright, Jr., ed., Late-Quaternary environments of the United States, Vol. 1. The Late Pleistocene (S. C. Porter, ed.), University of Minnesota Press, Minneapolis, 407 pp. Dillon, W. P., & R. N. Oldale. 1978. Late Quaternary sea-level curve: Reinterpretation based on gla- ciotectonic influence.— Geology 6:56-60. Gallagher, W. B., D. C. Parris, B. S. Grandstaff & C. DeTample. 1989. Quaternary mammals from the continental shelf off New Jersey.—The Mo- sasaur 4:101-110. McDonald, H.G. 1977. Description of the osteology of the extinct gravigrade edentate Megalonyx with observations on its ontogeny, phylogeny and functional anatomy. Unpublished M.S. Thesis, University of Florida, 328 pp. —., & D.C. Anderson. 1983. A well-preserved ground sloth (Vegalonyx) cranium from Turin, Monona County, Iowa.— Proceedings Iowa Academy Science 90(4):134—140. McNab, B. K. 1985. Energetics, population biology, and distribution of xenarthrans, living and ex- tinct. Pp. 219-236 in G. G. Montgomery, ed., The evolution and ecology of armadillos, sloths and vermilinguas, Smithsonian Institution Press, Washington, D.C., 451 pp. Oldale, R. N., F. C. Whitmore, Jr., & J. R. Grimes. 1987. Elephant teeth from the western Gulf of Maine, and their implications. — National Geo- graphic Research 3(4):439-446. Parris, D.C. 1983. New and revised records of Pleis- tocene mammals of New Jersey.—The Mosa- saur 1:1-21. Stock, C. 1925. Cenozoic gravigrade edentates of western North America.— Carnegie Institution of Washington Publication 331: 206 pp. . 1942. A ground sloth in Alaska.—Science 95: $52-553. Veatch, A. C., & P. A. Smith. 1939. Atlantic sub- marine valleys of the United States and the Con- go submarine valley.—Geological Society America Special Paper No. 7: 101 pp. Whitmore, F. C., Jr., K. O. Emery, H. B. S. Cooke, & D. J. P. Swift. 1967. Elephant teeth from the Atlantic continental shelf.—Science 156(3781): 1477-1481. (HGM) Cincinnati Museum of Natural His- tory, Cincinnati, Ohio 45202; (CER) De- partment of Paleobiology, National Mu- seum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 6-25 SYSTEMATICS OF THE “GREEN-THROATED SUNANGELS” (AVES: TROCHILIDAE): VALID TAXA OR HYBRIDS? Gary R. Graves Abstract. —Four species of hummingbirds, Heliangelus squamigularis Gould, 1871, Heliangelus barrali Mulsant & Verreaux, 1872, Heliotrypha speciosa Salvin, 1891, and Heliotrypha simoni Boucard, 1892, were described from 19th century commercial “‘Bogota’’ collections. The systematic status of these taxa, which I collectively refer to as ““green-throated sunangels”’ (GTS), is unresolved, but they have been variously treated as one or more valid species or as hybrids. I examined three systematic hypotheses—that GTS specimens represent (1) one or more valid species; (2) genetic variants of other species; or (3) hybrids. Plumage and mensural characters of GTS (n = 14) suggest they represent hybrids of Heliangelus amethysticollis x Eriocnemis cupreoventris from the Eastern Cordillera of the Colombian Andes. Alternate hypotheses of hybridity are discussed. Discrimination of hybrids and identifying their parental species depends upon an efficient “hybrid diagnosis.”’ The current method of diagnosing hybridity is often insufficient in that the materials, methods, and results are not properly documented. I address these problems and suggest guidelines for hybrid diagnoses. Untold thousands of hummingbird skins were exported from northwestern South America in the 19th century for the milli- nery trade and collectors of natural history specimens. Systematists sorted through some of the massive shipments of ““Bogota”’ trade skins and described dozens of new species, a few from unique specimens (e.g., Boucard 1892). Most were subsequently verified by the discovery of populations; others were determined to be of hybrid or- igin (Berlioz & Jouanin 1944). However, the validity of more than a dozen taxa remains indeterminate (Morony et al. 1975). These represent some of the most challenging problems in avian taxonomy. Resolving their systematic status depends on the me- chanics of discriminating avian hybrids from valid biological species. This paper has two aims that are ad- dressed concurrently. I evaluate the system- atic status of an enigmatic group of hum- mingbird taxa known only from a handful of 19th century specimens. Of more general interest, I examine the assumptions, ma- terials, and methods of the hybrid diagnosis in avian taxonomy. Taxonomy of the ““Green-throated Sunangels”’ Four species of hummingbirds, that I col- lectively refer to as “‘green-throated sun- angels” (hereafter abbreviated as GTS), were described from 19th century ““Bogota”’ col- lections: Heliangelus squamigularis Gould, 1871; Heliotrypha barrali Mulsant & Ver- reaux, 1872; Heliotrypha speciosa Salvin, 1891; and Heliangelus simoni Boucard, 1892. Heliotrypha Gould, 1853 is now con- sidered a junior synonym of Heliangelus Gould, 1848. Taxonomic uncertainty within the group began with Gould (1871), who was initially inclined to consider the type specimen of H. squamigularis a sport or variant of some VOLUME 103, NUMBER 1 other Heliangelus species, but who after fur- ther investigation characterized it as a new species related to Heliangelus exortis and H. amethysticollis. H. barrali and H. squa- migularis were not compared with one another before being described. Salvin (1892) considered these taxa as identical, but distinct from his newly described H. speciosa. In the first review of all four taxa, Boucard (1895) followed Salvin’s synony- my of H. barrali and H. squamigularis, but treated H. simoni (Boucard 1892) and H. speciosa as valid species, while noting the possibility that both were varieties of H. squamigularis. Cory (1918) lumped H. spe- ciosa and H. simoni and initiated the two- species taxonomy for the group adopted by Simon (1921), Peters (1945), and provi- sionally by Morony et al. (1975). Hartert (1922), perhaps cued by Boucard (1895), proposed that H. simoni and H. speciosa were aberrations of a single valid GTS species (=H. squamigularis). The possible hybrid origin of GTS was first raised by Berlioz (1936), who suggested that a spec- imen in Paris, which had previously been identified as H. simoni (discussed later), represented a hybrid of Heliangelus exortis x Haplophaedia aureliae. This opinion was endorsed by Jouanin (1950) and Greenway (1978), but Berlioz & Jouanin (1944) were less conclusive, stating simply that GTS were hybrids between either H. exortis or H. amethysticollis and some species of Erioc- nemis (including the closely related genus Haplophaedia). Meyer de Schauensee (1949) at first doubted the notion of hybridity but later agreed with Berlioz & Jouanin (1944) and supposed that GTS were hybrids of Er- iocnemis sp. X Heliangelus sp. (Meyer de Schauensee 1966). Hilty & Brown (1986) listed H. speciosa and H. squamigularis as presumed hybrids without mentioning pa- rental species. Syntypes of Heliotrypha simoni Boucard’s (1892) use of the plural “spec- imens”’ in his description of H. simoni im- 1 plied that the description was based on two Or more syntypes. Cory (1918) and Lord Rothschild (Hartert 1922) each obtained a syntype of H. simoni, presumably from Boucard before his death in 1905. The la- bels of both specimens (FMNH 46294, AMNH 483683) are marked, “Heliotrypha simoni, 6, Typical specimen, Colombia,” in what appears to be Boucard’s handwriting (fide Greenway 1978). Simon (1921) stated in a footnote that he could not find the type of H. simoni in Boucard’s collection, which suggests that only two syntypes existed, both of which were sold or exchanged by Boucard to other museums. Berlioz (1936), however, argued that a specimen in the Boucard Col- lection labeled ‘‘Heliotrypha speciosa, 6, Co- lombia” (now deposited in MNHN, Paris), was in fact the type and only existing spec- imen of H. simoni (see Berlioz & Jouanin 1944, Jouanin 1950). He apparently based his conclusion on the close resemblance of the specimen to Boucard’s description of H. simoni, and on the fact that Boucard (1895) did not mention possessing a specimen of H. speciosa. Several explanations are possible for this discrepancy. Assuming that there were orig- inally three examples of H. simoni, Boucard may have attached a new label to the re- maining syntype after disposing of the other two. Other possibilities are that Boucard (1895) obtained the specimen of H. speciosa after the publication of his monograph or that the specimen was labeled by Boucard as H. speciosa before he described H. simoni and never relabeled afterwards. In any event, only specimens designated by Boucard Mu- seum labels as ““H. simoni” should be re- garded as syntypes. Materials and Methods The type specimens of GTS are deposited in three different museums and are not available for loan. This prevented me from comparing all type specimens simulta- neously. I examined three type specimens: (1) the type of Heliotrypha barrali (AMNH 8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Ventral view of “green-throated sunangels” deposited in the American Museum of Natural History (from left to right: 483680, 483681, 483678, 483684, 483682, syntype of ““Heliotrypha simoni”’ [483683], type of ‘‘Heliotrypha barrali”’ [37655)). 37655) and (2) a syntype of Heliotrypha si- mont (AMNH 483683) in the American Museum of Natural History (see Hartert 1922, Greenway 1978); and (3) a syntype of Heliotrypha simoni (FMNH 46294) in the Field Museum of Natural History. The AMNH types were compared directly with five additional specimens (AMNH 483678, 483680, 483681, 483682, 483684) that have been variously identified as one or more of the GTS taxa (Fig. 1). The FMNH type was compared directly with FMNH 46286 (identified as H. barrali on its Boucard Mu- seum label). I examined one additional GTS specimen (ANSP 160344 [formerly AMNH 483679]). These specimens were compared with mensural data and color transparencies of the following specimens: one labeled H. speciosa (considered by Berlioz [1936] as a syntype of H. simoni) in the Museum Na- tional D’ Histoire Naturelle (MNHN), Paris; the types of H. squamigularis (BM 88.7.25.178) and H. speciosa (BM 87.3.22.889), and an unnumbered speci- men of H. barrali (photographs only) in the British Museum of Natural History (see Ap- pendix). GTS specimens and color transpar- encies were compared with series of all hummingbird species in the National Mu- seum of Natural History (USNM) and the American Museum of Natural History. In addition to the GTS specimens examined in this study, at least one other specimen exists (Berlioz 1964). Color comparisons of specimens were made under Examolites® (Macbeth Corp.). Measurements (wing chord, tail from in- sertion of central rectrices to tip of the outer and innermost rectrices, and culmen from anterior extension of feathers) were taken VOLUME 103, NUMBER 1 Table 1.—Measurements (mm) of “‘green-throated sunangels.”’ Age* Type specimens (1) squamigularis (BM) adult (2) barrali (AMNH) immature (3) speciosa (BM) adult (4) simoni (AMNH) immature (5) simoni (FMNH) adult Other specimens (6) “speciosa” (MNHN) adult (7) FMNH 46286 immature (8) AMNH 483678 adult (9) AMNH 483680 immature (10) AMNH 483681 adult (11) AMNH 483682 adult (12) AMNH 483684 adult (13) ANSP 160344 immature Outermost Wing chord rectrix Central rectrix Culmen 63.5° 43.2° 33.0° 16.6° 63.1 42.8 34.5 17.3 63.5° 40.6° 36.8" 19.4¢ 60.4 43.4 33.6 18.5 64.5 43.3 34.7 16.9 64.04 42.54 32.04 19.04 64.5 43.9 35.7 16.8 64.6 43.9 35.9 17.1 64.5 43.8 35.6 15.7 63.7 41.4 33.6 16.3 56.5 — — — 62.3 40.8 — 16.8 60.5 39.8 33.6 16.8 a Tmmatures have corrugations on the ramphothecum of the upper jaw. > Measurement from Salvin 1892. ¢ Measurement courtesy of J. Becker. 4 Measurement courtesy of C. Jouanin. with digital calipers and rounded to the nearest 0.1 mm (Table 1). I used principal components analysis (PCA) on untransformed variables to re- duce the dimensionality of data and to fa- cilitate the analysis of morphology in two dimensions. Unrotated principal compo- nents were extracted from correlation ma- trices (SYSTAT). Systematic Status of Green-throated Sunangels Investigations of GTS have engendered a remarkable variety of systematic opinions. If nothing else, this strongly suggests that multiple hypotheses of origin must be ad- dressed. Accordingly, I considered three possibilities. GTS may represent one or more of the following entities: (1) rare ge- netic variants of other Heliangelus species (Gould 1871); (2) hybrids (Berlioz 1936; Berlioz & Jouanin 1944; Meyer de Schauen- see 1949, 1966; Jouanin 1950; Greenway 1978; Hilty & Brown 1986), or (3) popu- lation samples of one or more valid biolog- ical species (Gould 1871, Salvin 1892, Bou- card 1895, Cory 1918, Peters 1945, Morony et al. 1975). Do green-throated sunangels represent rare genetic variants of other species?—Several examples of intra-population variation in plumage are known in Heliangelus species. Polymorphism in the number of iridescent gorget feathers in females has been well doc- umented, particularly in Heliangelus exortis (Chapman 1917; Zimmer 1951; Bleiweiss 1985a, b), and melanism involving part or the entire plumage is known in a number of Andean genera (Hartert 1922, Greenway 1978, Graves, pers. obs.). Intrasexual color polymorphism, however, does not appear to be significantly correlated with size. GTS closely resemble some species of Helian- gelus (e.g., H. exortis), but differ in body proportions from all species and by having lengthened tibial plumes and a green or sil- very-green gorget in combination with bril- liantly reflective plumage on the posterior part of the body. These qualitative char- acters indicate that GTS are not plumage variants of any other species of humming- bird. 10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hybrids or species? — As demonstrated by a century of equivocal taxonomy, it is dif- ficult to determine whether GTS are hybrids or valid species. This is due primarily to two factors. GTS specimens were collected in the 19th century and are unaccompanied by ecological, sexual, or locality data. They are believed to have originated from the Andean region of northwestern South America, an area of high species diversity where new species of hummingbirds are still being discovered (e.g., Eriocnemis mirabi- lis). The large number of GTS specimens (n = 15), presumably collected in a biotically diverse but poorly-known region, favors the valid species hypothesis. On the other hand, the plumage color and morphology of GTS are variable and intermediate between sun- angels (Heliangelus) and pufflegs (Erioc- nemis and Haplophaedia). This suggests that hybridization is involved. As hybrids have no standing in zoological nomenclature, the burden of proof is on taxonomists to reject the hybrid origin of GTS conclusively be- fore conferring species status on them. The process of discriminating avian hy- brids and their parental species can be termed the “‘hybrid diagnosis.’ Most tax- onomists consider the pathways of hybrid diagnosis to be self-evident and the docu- mentation of methods and diagnostic as- sumptions to be unnecessary. However, omissions of these crucial data obscure the diagnoses of all but the most obvious cases of hybridity. Beyond calling attention to an “unusual” specimen, a hybrid report based on an incomplete diagnosis is of little value to taxonomists and evolutionary biologists. As a minimum, the following points (not mutually exclusive), should be explicitly ad- dressed in hybrid diagnoses. 1. Potential parental species: What species were considered as possible parental species and why? 2. Diagnostic assumptions of character analysis: What operational assumptions were made concerning the inheritance of plumage and morphological characters of hybrids? How were characters defined and apomorphies identified? 3. Documentation of results: Can the hy- pothesis of hybridity be rejected? Ifnot, how were the parental species identified to the exclusion of all others? How were alternate hypotheses (e.g., valid taxon; genetic or de- velopmental variant) rejected? Hybrid Diagnosis Potential parental species. —For any hy- brid of unknown parentage, the pool of po- tential parental species (species hypotheti- cally or actually available for hybridization) can be defined taxonomically and geograph- ically. Interordinal hybridization is un- known in birds (Gray 1958); interfamilial hybridization has been reported in captivity (e.g., turkey < guinea fowl) but is unknown in nature. Thus, the taxonomic pool can be narrowed considerably if the hybrid can be identified to a particular family-level group (e.g., hummingbird or duck), which is al- ways the case. The taxonomic pool may be further restricted to a subfamily, genus, or a single pair of species when the rationale for doing so can be vigorously supported. For example, Parkes (1984) properly re- stricted the pool of potential species of a hybrid cuckoo collected in Pennsylvania to the only pair of Coccyzus species that occur sympatrically in North America north of the Gulf coast. In the interest of compre- hensiveness, however, he could have also addressed the six other species of Coccyzus in a few sentences in much the same way a taxonomist would mention other species in the differential diagnosis of a new species. As suggested by the cuckoo example, the pool of potential parental species can be limited geographically. The degree of limi- tation depends on knowledge of the migra- tory habits of the potential parental species and the geographic origin of the hybrid. For instance, the taxonomic pool of potential parental species of a hybrid hummingbird is defined by the family Trochilidae (345+ VOLUME 103, NUMBER 1 species). A hybrid hummingbird originating from Arizona could have no more than 20 potential parental species (190 species com- binations). On the other hand, a hybrid from an unspecified area of northwestern South America could have 150+ potential paren- tal species (10,440 species combinations). Clearly, the difficulty of hybrid diagnosis is directly proportional to taxonomic species diversity and geographic scope. Diagnostic assumptions of character anal- ysis. —In diagnosing putative hybrids, I as- sumed that mensural characters, such as wing and bill length, were polygenic and additive and that the morphology of hybrids does not exceed that of the parental species (Falconer 1981). Plumage characters in hy- brids may resemble a mosaic of the parental species or be inherited intact from one par- ent, depending on the number of encoding genes and their interaction (Hutt 1949, Buckley 1982). Hypothetically, hybrids may exhibit a wide range of plumage phenotypes. The major pigments in bird plumage, mel- anins, carotenoids, and porphyrins, appear to be under separate genetic control and mu- tually independent between feather tracts. The inheritance of structural colors, which dominate the plumage of hummingbirds, is poorly understood (Fox & Vevers 1960, Lu- cas & Stettenheim 1972), but the complex- ity of color-producing structures suggests a polygenic mode of inheritance. What little is known about inheritance in hybrid hummingbirds is summarized by Banks & Johnson (1961) and Short & Phil- lips (1966). Strongly contrasting patterns of non-structural color (e.g., rufous and black rectrices of Selasphorus sp.) are expressed in some fashion in all crosses. There is rea- son to doubt that the same is always true, however, for plumage characters exhibiting brilliant structural color. For example, the coronal iridescence found in species of Ca- lypte is evident in five examples of C. anna x Selasphorus sasin and one specimen of C. anna X Stellula calliope (although it is never as extensive on the hybrids as it is on 11 C. anna), but is lacking in the single known C. anna X Archilochus alexandri hybrid (Banks & Johnson 1961). This suggests either that few genes control the color of coronal plumage or that phenotypic expression is controlled by a modifer in these species. The shape of gorget feathers, rectrices, and re- miges of hybrids is generally intermediate between those of the parental species, re- flecting a polygenic mode of inheritance. Banks & Johnson (1961) assumed that hybridization in hummingbirds does not produce traits of species or genera other than those involved in the particular cross. This assumption rules out the possibility of ata- vistic characteristics—those not found in either parental species but which reflect a pattern postulated to be ancestral or the re- sult of mixed alleles encoding polygenic traits. Although atavism is well known in certain anseriform hybrids (e.g., Harrison & Harrison 1963), it has not been documented in hybrid hummingbirds. This study was geared toward the iden- tification of apomorphic character states in putative hybrids. The mosaic expression of parental autapomorphies in a number of characters is the best indicator of hybridity of a unique specimen and provides the only direct evidence of parentage. However, be- cause many plumage characters are poly- genic, the expression of parental apomor- phies may be obscured in hybrids. When parental apomorphies are not identifiable, the parentage of a hybrid may be indicated, although less conclusively, by the expres- sion of a combination of plesiomorphic characters unique to a single pair of parental species. Results The original labels of GTS specimens are marked ‘“‘Columbia”’ (sic), ““Colombia,”’ ‘“‘New Grenada,” or “Bogota Collection.” Thus, the geographic pool can only be de- fined in general terms. Berlioz & Jouanin (1944) showed that the vast majority of skins 12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON b Fig. 2. Feet and tarsi of “green-throated sunangels” (a = FMNH 46286; b = FMNH 46294), Heliangelus amethysticollis (c, male), and Eriocnemis cupreoventris (d, male). Note downy leg puffs on tibias of a, b, and d. => Fig. 3. Ventral view of “green-throated sunangels’’: (a) type of ““Heliangelus squamigularis”; (b) FMNH 46286; (c) syntype of ““Heliotrypha simoni” (FMNH 46294); (d) ‘““Heliotrypha speciosa’ (MNHN). 13 VOLUME 103, NUMBER 1 eT VOLUME 103, NUMBER 1 prepared in the ““Bogota”’ method were col- lected in the northern Andes and adjacent lowlands, a region roughly encompassed by the present boundaries of Colombia. In the absence of unequivocal locality data for any of the ““green-throated sunangels,” the geo- graphic pool of potential parental species must initially include all hummingbirds re- corded from Colombia, a total of 143 species in 61 genera (Hilty & Brown 1986). There are two major lineages of hum- mingbirds, the Phaethornithinae and Tro- chilinae. As Heliangelus and related genera belong to the Trochilinae, I treat the subfamily Phaethornithinae as an outgroup. I identified all species of hummingbirds that occur in Colombia that shared with GTS one or both of the following characters that are apomorphic with respect to species in the outgroup: (1) lengthened downy tibial piumes (leg puffs) (Fig. 2); and (2) a brilliant gorget that contrasts with adjacent plumage and extends from the chin posteriorly to the upper breast (Figs. 3, 4). With the exception of GTS, no taxon exhibits both of these characters. It follows that if GTS are hy- brids, then one parental species contributed leg puffs and the other the brilliant gorget. Species representing four genera of hum- mingbirds, Boissonneaua, Eriocnemis, Haplophaedia, and Ocreatus, have downy tibial plumes that exceed those of GTS in length. Narrowing the pool of potential pa- rental contributors of the brilliant gorget is more difficult. Including taxa with brilliant- ly reflective throats that do not contrast with adjacent plumage, species representing most of genera of the subfamily Trochilinae could be the gorgeted parent. However, only species of Heliangelus have gorgets that are similar in structure, shape, and size to those of GTS. Thus, a first review based on two apormophic characters limits the potential _— 15 parental species to five of the 61 genera of Colombian hummingbirds. Reduction of the species pool is support- ed by comparison of general morphology. The remiges, rectrices, body plumage, and bill of GTS are unspecialized and lack many of the elaborations that are common within trochiline hummingbirds. Assuming poly- genic inheritance of these structures, if GTS are hybrids, then their parental species must be morphologically unspecialized. Sexual dichromatism within the series of GTS specimens, if any, is minor (Fig. 1). Collec- tions of sexually dichromatic species of hummingbirds from Bogota collections are often sexually skewed in favor of brightly colored adult males (Graves, in prep.). Plumage of immature males of these species may resemble that of females. Because im- mature GTS specimens do not differ sig- nificantly in appearance from adults (which suggests that the sexes are similar), sexual bias in collecting due to appearance of GTS specimens is unlikely. Assuming a 1:1 sex ratio the probability of finding only males or females in a random sample of 13 (num- ber examined) individuals is P < 0.0002 (Binomial test). However, as pointed out by Haldane (1922), the heterogametic sex (2 in birds) may be rare or absent in F, hybrids. Therefore, for diagnostic purposes I enter- tained the possibility that the sample of GTS specimens was exclusively male. By structural criteria alone, 106 species (47 genera including Ocreatus and Boisson- neaua) may be eliminated from the pool of potential parental species. These include species with specialized bills (e.g., Ensifera ensifera, Schistes geoffroyi), remiges (e.g., Campylopterus falcatus, Aglaeactis cupri- pennis), rectrices (e.g., Ocreatus underwoo- dii, Acestrura mulsant), and body plumage (e.g., Colibri coruscans, Lophornis stictolo- Fig. 4. Type of “Heliotrypha speciosa’: (a) ventral view; (b) enlargement of upper breast showing white pectoral band; (c) side view of head showing sloping profile forehead and thick bill. 16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pha). External morphology of the remaining 37 species representing 14 genera (K/ais, Chlorestes, Lepidopyga, Chrysuronia, Gold- mania, Goethalsia, Amazilia, Adelomyia, Anthocephala, Urosticte, Phlogophilus, He- liangelus, Eriocnemis, Haplophaedia) is rel- atively unspecialized. These bear further scrutiny as potential parental species of GTS. GTS have unpatterned rectrices and uni- formly dark bills. This suggests that species with spotted or patterned rectrices (e.g., Amazilia sp., Anthocephala floriceps, Ade- lomyia melanogenys, Phlogophilus hemi- leucurus) or markedly bicolor lower ram- photheca (e.g., Amazilia spp.) can be eliminated, leaving species from three gen- era (Heliangelus, Eriocnemis, Haplophae- dia) as potential parental species of GTS. Excepting these, all other species may be rejected from the species pool by two or more criteria (available from the author). In sum, rejection of species whose distinctive characters (some of which are apomorphic) are not found in GTS, reduces the species pool to nearly the same subset of species that share apomorphic characters with GTS. Berlioz (1936) noted the downy tibial plumes of the MNHN specimen, Heliotry- pha speciosa, and suggested that some species of Colombian puffleg (Eriocnemis spp., Haplophaedia spp.) was one of its par- ents. He concluded that the entirely green body plumage and non-brilliant undertail coverts of this specimen seemed to preclude the possibility of a cross with a species of Eriocnemis (with brilliant violet or blue un- dertail coverts) and that only one hybrid combination was possible, Haplophaedia aureliae xX Heliangelus exortis. Berlioz’s statement on undertail coverts may be ex- tended to other GTS specimens—none of the specimens I examined exhibit the struc- Fig. 5. tural brilliance found in many species of puffleg (e.g., Eriocnemis vestitus, E. luciani, E. cupreoventris, E. mirabilis, E. alinae, E. derbyi). Undertail coverts of the potential Heliangelus parental species vary from pure white to gray with white margins. Contrary to Berlioz’s reasoning, the hybrid progeny of Eriocnemis sp. X Heliangelus sp. might be expected to have green undertail coverts with white or grayish-white margins, simi- lar to those of GTS. One character of GTS that has not re- ceived mention by previous investigators is the extensive distribution of brilliant green reflections from the body plumage. These reflections extend posteriorly to the vent and upper tail coverts and are particularly ap- parent when specimens are viewed head-on in direct light. Barbule modifications of this type are well-developed in several species of Eriocnemis (e.g., E. vestitus, E. cupreo- ventris) but are weakly developed or lacking in both species of Haplophaedia. Several species of Heliangelus (e.g., H. exortis, H. amethysticollis, H. viola) exhibit brilliant reflections on the upper breast but lack them posteriorly when viewed head-on in direct light. The presence of brilliantly reflective plumage (similar to that of GTS specimens) on the flanks and abdomen of hypothetical Haplophaedia spp. x Heliangelus spp. hy- brids would represent a clear case of “‘ata- vism,” a phenonemon that has not been demonstrated in trochiline hybrids. Several other characters contradict Ber- lioz’s hypothesis that Haplophaedia aure- liae is involved in the parentage of GTS. Both sexes of H. aureliae have bronze crowns and uppertail coverts that contrast with the green back and rump. The dorsum of GTS specimens lacks such contrast, and, in fact, some specimens are brightest on the — Head profiles of ““green-throated sunangels”’ (a = FMNH 46286, b = FMNH 46294) and Haplophaedia aureliae (c). Nasal operculum of H. aureliae is more inflated and exposed than in “green-throated sunangels”’ and Heliangelus spp. VOLUME 103, NUMBER 1 17 18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON -2 al 0 PE Al Fig. 6 Bivariate plot of Principal Component factor scores of ““green-throated sunangels” and some of their potential parental species. Diamonds = Heliangelus exortis. Circles = Eriocnemis cupreoventris. Triangles = Haplophaedia aureliae. Squares = Heliangelus amethysticollis. Hollow and solid symbols represent males and females, respectively. Numbers represent “green-throated sunangel’”’ specimens from Table 1. Lines envelop groups of males and females of Heliangelus amethysticollis x Eriocnemis cupreoventris. uppertail coverts. Northern races of Hap- lophaedia aureliae (e.g., H. a. aureliae, H. a. caucensis), especially females, have white or grayish white abdomens and lower breasts speckled peripherally with green. Feathers of the ventral midlines of females and im- mature males of Colombian species of He- liangelus have wide buffy margins. Conse- quently, the venters of hypothetical female hybrids of H. aureliae x Heliangelus spp. would be extensively buffy, not green as in GTS. Another important character is the rela- tionship between the nasal operculum and the anterior extension of feathering on the bill of GTS specimens (Fitzpatrick et al. 1979) (Fig. 5). In Heliangelus exortis and H. amethysticollis, feathering extends an- teriorly to the distal edge of the nasal oper- culum but does not cover it. Feathering in Haplophaedia spp. does not reach the distal edge of the operculum, which is inflated and exposed. In Eriocnemis, feathers extend to the distal edge of the nasal operculum, or slightly beyond, imparting a sloped appear- ance to the forehead in profile. Feathering and forehead profile of GTS is somewhat intermediate between that found in Helian- gelus exortis or H. amethysticollis and sev- eral species of Eriocnemis, but differs from VOLUME 103, NUMBER 1 a Fig. 7. 19 Li | Li A i Males of the two most probable parental species of ‘“‘green-throated sunangels’’ examined in this study: (a) Heliangelus amethysticollis; (b) Eriocnemis cupreoventris. that of H. aureliae or H. lugens. This and the other characters mentioned above in- dicate that both species of Haplophaedia may be rejected as potential parental species of GTS and support the hypothesis that the puffleg parent is some species of Eriocne- MIs. Of the seven species of Eriocnemis (ex- cluding E.. godini, which is of uncertain sta- tus) that occur north of Ecuador, all but EZ. cupreoventris can be rejected as a parental species of GTS. Both sexes of E. mosquera (see Bleiweiss [1988] for measurements) and E. luciani have deeply forked tails and are significantly larger than GTS specimens or any of its possible Heliangelus parents. Nei- ther of these puffleg species occur in the Eastern Cordillera in sympatry with He- liangelus amethysticollis and hypothetical hybrids between these species and 7. exortis would have tails more deeply forked than in any GTS specimen. In addition, GTS lack white bases to throat feathers found in those species and the greenish outer rectrices of 20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Ranges of measurements (mm) of potential parental species of ““green-throated sunangels.”’ Spec- imens from a number of localities were chosen in order to incorporate the range of intraspecific variation found in Colombian populations. Species Sex n Wing chord Outermost rectrix Innermost rectrix Culmen Heliangelus exortis ‘) 10 64.0-67.6 44.8-50.0 32.1-34.5 13.3-15.0 Q 10 56.9-60.9 38.9-41.3 29.7-33.9 14.2-16.8 Heliangelus amethysticollis 3 10 65.1-71.8 43.6-48.8 41.4-46.0 14.8-16.0 g 10 60.5-65.6 35.9-42.4 36.9-39.6 15.5-17.1 Eriocnemis cupreoventris 3 15 58.8-65.2 40.6-45.3 24.3-28.4 17.8-19.6 Q 7 58.2-60.6 38.7-42.6 23.2-26.3 16.7-17.9 Haplophaedia aureliae r) 10 59.1-62.8 35.4-41.3 32.6-37.6 17.3-19.6 Q 10 55.3-58.9 32.8-37.8 32.3-36.1 16.2-20.1 E. mosquera. E. mirabilis and E. alinae are small, have restricted ranges in the northern Andes, and possess several apomorphic characters that are not expressed in GTS. E. derbyi is sexually dichromatic and di- vergent in tail structure and plumage (e.g., black leg puffs) and can be conclusively re- jected as a possible parent of GTS. E. ves- titus is moderately sexually dichromatic but both sexes possess a small violet (male) or blue (female) gorget on the upper throat. Hypothetical Heliangelus spp. x E. vestitus hybrids of both sexes would probably have a small gorget of some shade of purple or pink. Female hybrids would be bufher and less green on the breast and lower belly than are GTS. E. cupreoventris exhibits weak sexual di- chromatism, lacks a contrasting gorget, and is similar in size to both GTS and possible Heliangelus parents. The nasal operculum is partially covered with feathers in both sexes. When viewed head-on in direct light, the body plumage of adult males reflect a brilliant golden-green anteriorly changing to bluish-green on the upper tail coverts and to coppery-gold on the breast and belly. Fe- males are slightly duller below. These char- acters make E. cupreoventris the most prob- able parental puffleg species of GTS. The probable sunangel parent can be lim- ited to the only two species with extensive distributions in the Colombian Andes, H. exortis and H. amethysticollis. (H. mavors, H. strophianus, and H. spencei possess apo- morphies not found in GTS.) Both species occur sympatrically with E. cupreoventris in the Eastern Cordillera (see Hilty & Brown 1986). H. amethysticollis differs from H. ex- ortis primarily in having a well-defined white pectoral band below the throat of both sexes, a larger, more extensive gorget in males, and a less deeply forked tail. Intergeneric hy- brids involving these species may best be distinguished by mensural characters of the parental species (Table 2). Additional de- scriptions of potential parental species can be found in Salvin (1892), Zimmer (1951), Bleiweiss (1985a, b), and Hilty & Brown (1986). Measurements of the most probable pa- rental species of GTS (H. exortis, H. ame- thysticollis, E. cupreoventris; H. aureliae in- cluded for comparison) overlap extensively (Table 2). I compared the measurements of these species and GTS specimens. Under the diagnostic assumptions used in this study, a GTS specimen could not be the hybrid progeny of a pair of species if the measurements of the specimen occurred outside the cumulative range (+0.5 mm for wing and tail; +0.2 mm for culmen) of their measurements. Statistics were not per- formed because the reference samples were chosen to maximize ranges of measure- ments. This procedure is conservative be- cause it assumes that the inheritance of quantitative characters is mutually inde- VOLUME 103, NUMBER 1 21 Table 3.—Comparison of “green-throated sunangel” (GTS) measurements with the cumulative range of measurements for combinations of potential parental species (Heliangelus amethysticollis, H. exortis, Eriocnemis cupreoventris, Haplophaedia aureliae). A male or female symbol indicates that all measurements of a particular “‘sreen-throated sunangel” specimen fall within the range of measurements (by sex) for that combination of species. Numbers in parentheses refer to specimens listed in Table 1 for which all measurements were available. Ratios at the bottom of each column denote the minimum number of males and maximum number of females possible assuming that all “green-throated sunangels” represented hybrids of those species. Binomial P-values are given, assuming a 1:1 sex ratio. GTS H. amethy. x E. cupreo. H. amethy. x H. aureliae H. exortis x E. cupreo. H. exortis x H. aureliae (1) 3 ) ) ) (2) de fo fs) 6 (3) ) $2 — ) (4) é fe) fe) é (5) 3 3 3 3 (6) 3 9 3 3 (7) 3 3 = 8 (8) é $ = 3} (9) ) $ _ $ (10) 3e 39 3 3 (13) 9 89 89 89 3:9 8:3 6:5 6:1 10:1 P < 0.08 P= 0.20 P < 0.05 P = 0.005 pendent. Unfortunately, this procedure rules out few hybrid possibilities (Table 3). None of the possible pairs of parental species can be rejected for 7 of 11 of the GTS speci- mens. Multivariate morphological relationships (Table 4) of potential parental species and GTS specimens are illustrated by the first two axes of a Principal Components Anal- ysis in Fig. 6. Inspection of factor scores revealed that most GTS specimens are clus- tered near the center of the bivariate plot. Only one specimen (type of H. speciosa) falls within the envelope outlining the factor scores for H. aureliae and H. amethysticollis and none occurs in the H. exortis x E. cu- preoventris envelope. Assuming that the in- heritance of polygenic size and shape char- acters is reflected in the spread of factor scores, these pairs of species are not in- volved in the parentage of GTS, with the possible exception of the type of H. speciosa. All GTS specimens fall within the factor score envelope of H. amethysticollis x E. cupreoventris, and ten of eleven specimens fall within the H. exortis x H. aureliae en- velope. However, as noted previously with univariate comparisons, if GTS are the progeny of H. exortis x H. aureliae, they would be predominately male. This fact, and a variety of plumage characters previously discussed suggest that GTS are not hybrids of the latter two species. The scatter of GTS factor scores within the H. amethysticollis x E. cupreoventris envelope shows that an even sex ratio is possible. With the excep- tion of H. speciosa, plumage characters and measurements of GTS specimens are con- Table 4.— Factor loadings for the first two principal components from analysis of “green-throated sunan- gels” and potential parental species (see Fig. 6). I II Variable Wing chord 0.91 0.10 Outermost rectrix 0.85 —0.39 Central rectrix 0.67 0.72 Culmen =), 7/7 0.32 Variance explained Percent 62.5 19.6 Cumulative 62.5 82.1 22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sistent with the hypothesis that they rep- resent hybrids of H. amethysticollis and E. cupreoventris (Fig. 7). Note, however, that the wing chord of AMNH 483682 (Table 1), which is probably a female, is signifi- cantly shorter than any specimen in the sample of these two species. The position of H. speciosa on the bivar- iate plot is well removed from the other GTS specimens. H. speciosa differs from other GTS specimens in having a broad white pectoral band (Fig. 4) instead of a few semi-concealed spots, a shallowly-forked tail, and a slightly longer bill. The presence of a white pectoral band and the confor- mation of the gorget of H. speciosa indicates that a white-banded species of Heliangelus (e.g., H. amethysticollis) is one of the pa- rental species. The well-developed leg puffs of H. speciosa (fide M. P. Walters) indicate that the other parental species is a puffleg. The anterior extension of feathering over the nasal operculum of H. speciosa, how- ever, indicates that the other parent could not be H. aureliae which is similar to it in size and shape. H. speciosa lies within the PCA envelope for H. cupreoventris x H. amethysticollis. Despite the difference in ap- pearance of H. speciosa from other GTS specimens, it seems probable that they rep- resent the same hybrid cross. Additional study of the specimen may be required to verify this fact. If true, then H. speciosa rep- resents an extreme hybrid phenotype that resembles its sunangel parent much more than its puffleg parent. Conclusions With the possible exception of H. spe- ciosa, GTS specimens examined in this study, for which measurements were avail- able, are probably hybrids of Heliangelus amethysticollis x E. cupreoventris. Berlioz’s (1936) hypothesis that the MNHWN speci- men and perhaps others were hybrids of Haplophaedia aureliae and Heliangelus ex- ortis 1s not supported by the data. Because the hypothesis of hybridity cannot be re- jected, GTS cannot be considered as valid taxa. Additionally, the data (especially the variability and inconsistency of plumage characters) do not support the hypothesis that GTS, taken as a whole, represent pop- ulation samples of one or more valid species. Geographic origin. —H. amethysticollis and E. cupreoventris are sympatric in An- dean forests and shrublands (2000-3000 m elevation) in the northern half of the Eastern Cordillera. Thus, GTS specimens could ac- tually have been collected in the environs of ‘Bogota.’ Mulsant & Verreaux (1872) reported that the type of Heliotrypha barrali was collected on the Rio Saldana, Depart- ment of Tolima, in the Central Cordillera of the Colombian Andes. However, as pre- viously mentioned, the original labels of H. barrali and other specimens lack specific lo- cality data and in the absence of corrobo- rating evidence, the possible Central Cor- dilleran origin of the type of H. barrali can be dismissed. Nomenclature. — Hybrids are individuals and not taxa. Thus, the names Heliangelus squamigularis Gould, 1871, Heliotrypha barrali Mulsant & Verreaux, 1872, Heli- otrypha speciosa Salvin, 1891, and Heli- otrypha simoni Boucard, 1892, are available only for the purposes of homonymy in tax- onomy and should not be used in the pop- ular literature. For the purposes of field guides, these hybrids may be referred to col- lectively as ““green-throated sunangels.”’ Discussion Hybrid diagnoses can be simple or ex- tremely complex depending on circum- stances. Factors that affect the success of hybrid diagnoses include: (1) the number of hybrid individuals and their age, sex, and hybrid composition (e.g., F,, backcross); (2) the number of distinctive plumage and mor- phological characters on the hybrid; (3) the number of species in the putative hybrid’s taxonomic group; (4) and knowledge of the hybrid’s taxonomic group and of the re- gional avifauna where the hybrid originat- VOLUME 103, NUMBER 1 ed. (The most challenging diagnoses are of hybrids represented by a unique, unsexed, possibly immature specimen without spe- cific locality data, which belongs to a spe- ciose, dull-plumaged, and poorly known taxonomic group from a poorly collected region of high species diversity!) Ideally, parental species are identified with certainty, but failing this, what result jus- tifies the considerable effort expended in the average hybrid diagnosis? In terms of value to future researchers, it is far better to have a short list of species that includes the cor- rect pair of parents, than an exact deter- mination of parental species that may be wrong (errors of this sort are frequently per- petuated in the literature; see Graves 1988). Rejection of any species from the pool of potential parental species must be based on the unequivocal violation of diagnostic as- sumptions, and there is no logical reason for reducing the species pool beyond the limits suggested by the data. Conclusive knowledge of what species or species com- binations are not parents, as well as those that might be parents of a hybrid is far more valuable than perhaps recognized by most taxonomists, especially when the majority of species fall into the former category. Species in the latter category constitute the nucleus for future analyses. Acknowledgments R. Banks, M. R. Browning, P. Cannell, F. Gill, S. Olson, and R. Zusi commented on an earlier version of the manuscript. Banks, R. Bleiweiss, T. Schulenberg, and Zusi re- viewed a later incarnation. I thank them for their suggestions and criticism. I thank J. Becker, H. James, C. Jouanin (MNHN), M. Robbins (ANSP), and M. Walters (BM) for providing measurements or answering de- tailed questions about specimens, Becker and Jouanin for providing excellent color transparencies of specimens for my use, and V. Krantz for preparing illustration prints. M. Farmer deserves special thanks for translating French. I am also grateful to the 23 curators and staff of the American Museum of Natural History and the Academy of Nat- ural Sciences of Philadelphia (ANSP) for providing access to collections and the Field Museum of Natural History for loaning GTS specimens. Lastly, I would like to thank R. L. Zusi for patiently answering untold thou- sands of my questions about hummingbird systematics over the past few years. Museum research was supported by grants from the Smithsonian Research Opportu- nities Fund and the Frank M. Chapman Memorial Fund of the American Museum. I dedicate this paper to Jacques Berlioz and Christian Jouanin in recognition of their pioneering work on hummingbird hybrids. Literature Cited Banks, R. C., & N. K. Johnson. 1961. A review of North American hybrid hummingbirds. —Con- dor 63:3-28. Berlioz, J. 1936. Note sur l’identite probable du type d’Heliotrypha simoni Boucard (Trochilides). — Bulletin du Museum National D’Histoire Na- turelle, Paris 14:233-236. . 1964. Lacollection de Trochilides A. L. But- ler.—Oiseau 34:91-105. , and C. Jouanin. 1944. Liste de Trochilides trouves dans les collections commerciales de Bogota. — Oiseau 14:126-155. Bleiweiss, R. 1985a. Variation and population struc- ture of the Tourmaline Sunangel, Heliangelus exortis exortis (Aves, Trochilidae).— American Museum Novitates 2811:1-14. 1985b. Iridescent polychromatism in a fe- male hummingbird: is it related to feeding strat- egies?— Auk 102:701-713. 1988. Systematics and geographic variation in the Golden-breasted Puffleg Eriocnemis mos- quera (Aves, Trochilidae).— American Museum Novitates 2913:1-8. Boucard, A. 1892. A complete list up to date of the hummingbirds found in Columbia, with de- scriptions of several supposed new species. — The Humming Bird 2:73-87. . 1895. Genera of humming birds. Vol. 5, Part 1. Bournemouth, London. Buckley, P. A. 1982. Avian genetics. Pp. 21-110 in M. Petrak, ed., Diseases of cage and aviary birds. 2nd ed. Lea and Febiger, Philadelphia, 680 pp. Chapman, F. M. 1917. The distribution of bird-life in Colombia.—Bulletin of the American Mu- seum of Natural History 36:1-729. 24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cory, C. B. 1918. Catalogue of birds of the Americas. Part 2, No. 1.—Field Museum of Natural His- tory Zoological Series 13:1-313. Falconer, D. S. 1981. Introduction to quantitative genetics. 2nd ed. Longman, New York, 340 pp. Fitzpatrick, J. W., D. E. Willard, and J. W. Terborgh. 1979. A new species of hummingbird from Peru.— Wilson Bulletin 91:177-186. Fox, H. M. and G. Vevers. 1960. The nature of an- imal colours. Macmillan, New York, 246 pp. Gould, J. 1871. Descriptions of six new humming- birds. — Proceedings of the Zoological Society of London 1871:503—505. Graves, G.R. 1988. Evaluation of Vermivora x Opo- ronis hybrid wood-warblers.— Wilson Bulletin 100:285-289. Gray, A. P. 1958. Bird hybrids. Commonwealth Ag- ricultural Bureaux, Bucks, England, 390 pp. Greenway, J. C., Jr. 1978. Type specimens of birds in the American Museum of Natural History. Part 2.—Bulletin of the American Museum of Natural History 161:1-305. Haldane, J. B.S. 1922. Sex-ratio and unisexual ste- rility in hybrid animals.—Journal of Genetics 12:101-109. Harrison, J. M., & J. G. Harrison. 1963. Comments on a hybrid Red Shoveler x Northern Shovel- er.— Bulletin of the British Ornithological Club 83:21-25. Hartert, E. 1922. Types of birds in the Tring Mu- seum. B. Types in the general collection. Trochi- li.—Novitates Zoologicae 29:403-412. Hilty, S. L., & W. L. Brown. 1986. A guide to the birds of Colombia. Princeton University Press, Princeton, New Jersey, 836 pp. Hutt, F.B. 1949. Genetics of the fowl. McGraw-Hill, New York, 590 pp. Jouanin, C. 1950. Catalogue systematique des types de Trochilides du Museum National d’Histoire Naturelle de Paris.—Bulletin du Museum Na- tional d’Histoire Naturelle de Paris 22:1—27. Lucas, A. M., & P.R. Stettenheim. 1972. Avian anat- omy—integument. Agricultural handbook 362. United States Department of Agriculture, Washington, D.C. Meyer de Schauensee, R. M. 1949. Birds of the Re- public of Colombia. Part 2.—Caldasia 23:38 1- 644. . 1966. The species of birds of South America. Livingstone Press, Narberth, Pennsylvania, 577 pp. Morony, J. J., Jr., W. J. Bock, & J. Farrand, Jr. 1975. Reference list of the birds of the world. Amer- ican Museum of Natural History, New York. Mulsant, E., & J. Verreaux. 1872. Description d’une espece nouvelle d’oiseau moche.— Annales So- ciete Linneenne de Lyon 18:106—107. Parkes, K.C. 1984. An apparent hybrid Black-billed x Yellow-billed Cuckoo.— Wilson Bulletin 96: 294-296. Peters, J. 1945. Check-list of birds of the world. Vol. 5. Museum of Comparative Zoology, Cam- bridge, Massachusetts, 306 pp. Salvin, O. 1891. Descriptions of new species of Upu- pae and Trochili in the collection of the British Museum.—The Annals and Magazine of Nat- ural History (6th series) 7:374-378. 1892. Catalogue of birds in the British Mu- seum, Vol. 16. Short, L. L., and A. R. Phillips. 1966. More hybrid hummingbirds from the United States.—Auk 83:253-265. Simon, E. 1921. Histoire naturelle des Trochilidae (synopsis et catalogue). Paris, 416 pp. Zimmer, J. T. 1951. Studies of Peruvian Birds. No. 61. The genera Aglaeactis, Lafresnaya, Pteroph- anes, Boissonneaua, Heliangelus, Eriocnemis, Haplophaedia, Ocreatus, and Lesbia. —Ameri- can Museum Novitates 1540:1-55. Department of Vertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. Appendix Comparative Description of ““Green-throated Sunangels”’ Descriptions of structural colors are unusually sub- jective and actual color varies with the angle of in- spection and direction of light. For this reason I use general color descriptions. Numbers in parentheses re- fer to specimens in Table 1. The crown, nape, back, and rump are medium green. Uppertail coverts are medium green to bluish-green. There is no contrast between crown and back. When viewed head-on in direct light, scattered feathers on crown (5), back, wings, and upper tail coverts show brilliant golden-green to green reflections. A brilliant green frontlet, variable in intensity and definition, is found in most adults (absent in 10), but is faint or lacking in immatures. When present, the frontlet is small (1, 2, 4, 6, 8, 11, 12), similar in size to that found in H. exortis or H. amethysticollis. Prominence of the frontlet is affected by variations in skin preparation. Lores, auriculars, and neck at the sides of the throat are medium green, but appear much darker when viewed head-on. A small white postocular spot is present. A brilliant gorget is found in all specimens but is variable in size, color, and degree of contrast with adjacent plumage. Gorget margins are somewhat irregular and indistinct in all adult specimens. Gorget color is vari- VOLUME 103, NUMBER 1 able and can be characterized (viewed head-on) as bluish-green (4) and silvery-green (2, 9, 13) to silvery bluish-green (7) in immatures, and from bluish-green (5) and pale green (12) to silvery-green (1, 3, 6, 8, 10, 11) in adults. In indirect light, gorgets of some speci- mens emit faint coppery or pinkish reflections. Color variation appears to be fairly continuous and color characterizations are arbitrary. For example, “silvery- green” includes various shades of pale metallic green (= “leaden” of Salvin 1892). Gorgets of immatures are oval in shape, wider posteriorly and may be surround- ed by dull, lax plumage (7). Gorgets of adults are larger, contrast less with adjacent plumage, and may have irregular margins (5). Brilliant gorget feathers are rounded and about the size of those in male Helian- gelus sp. (e.g., H. exortis), but become progressively smaller toward the chin and malar regions in adults. The upper and lower breast, abdomen, and flanks (ex- cept 2) are medium green. Feathers along the midline of immatures may have narrow buffy margins. Brilliant golden-green or green reflections (or faint coppery in 5) are scattered over the breast, abdomen, and flanks when viewed head-on. Plumage of immatures is duller than in adults. The type of H. speciosa (3) differs from other GTS specimens in having a large white pectoral patch spotted with green discs and darker underparts with few brilliant reflections. White or buffy spots, 25 mostly subterminal, are present on two to ten feathers of the upper pectoral area on most specimens (1, 2, 7, 8, 9, 10, 11, 13). Undertail coverts are variable in length (not exceeding half the length of the tail) and are medium green or bluish-green with narrow to broad white or grayish white margins; basal barbs in some specimens are long and downy. Central rectrices are dark green to bronzy green; outer retrices are bluish- black. Depth of tail fork varies from 3.8 to 10.5 mm. Outer web of outermost rectrix is well-developed (>12 width of inner web). The outermost rectrix ranges from 8.0 to 9.5 mm in width and is more acuminate in shape than in H. exortis or H. amethysticollis. Tibial feathers (leg puffs) are white or buffy (2), variable in length (2.5 to 8 mm), and more or less downy in texture. [The presence of leg puffs on (1) cannot be determined from photographs of specimens or published descriptions]. The remiges are unmodified (e.g., not emarginate) and dull dark brown in color. Outer webs of primaries of some specimens have a faint bronzy-green sheen. Bills are unmodified and straight and dark blackish-brown. The upper mandible of (3) is broken (Fig. 4). Feathering extends anteriorly on the bill to the distal edge of the nasal flange (not inflated), somewhat obscuring it. Feet are dark brown or dark blackish brown. Additional notes on plumage can be found in Berlioz (1936). PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 26-34 A NEW SPECIES OF CUORA (REPTILIA: TESTUDINES: EMYDIDAE) FROM THE RYUKYU ISLANDS Carl H. Ernst and Jeffrey E. Lovich Abstract.—A new species of Asian box turtle, Cuora evelynae, from the Ryukyu Islands is described and compared with C. flavomarginata from Taiwan and southern China. Recognition is based on its pattern of very large light colored pleural blotches (length of blotch at its medial height 49-72% of medial length of pleural) that at their dorsal border coalesce to form lateral stripes in 52% of adults examined, its large light brown plastral blotch which is indented at the bridge, and the usual presence of less than 10 rows of large scales on the anterior surface of the forelimbs. A discriminant function is given that separates C. evelynae from C. flavomarginata. Geological history of the Ryukyu Islands is discussed, in view of speciation of C. evelynae. The yellow-margined box turtle, Cuora flavomarginata (Gray, 1863) occurs in southern China, Taiwan, and the Ryukyu Islands (Iverson 1986). Even though these localities have been long isolated (Inger 1947), no consistent differentiation of these turtle populations has been reported pre- viously. Hsu (1930) described a subspecies, Cuora f. sinensis, from Tungting Lake, Hunan, China, as differing from the nominate Tai- wanese population on the basis of having the anterior border of the plastron obtusely emarginate, each plastral scute with deeply cut parallel lines, each marginal with its pos- terior angle slightly overlapping the next with the degree of overlap most pronounced in the posterior third of the marginals so that the posterior carapacial rim is somewhat serrated, a small notch between the anal scutes, and a much shorter tail. Pope (1935) commented that with the exception of the longer tail (which may be sexual dimor- phic), all the other characters used by Hsu are either common variants expected in most emydid turtles or differences generally cor- related with age, but Pope thought that until direct comparison between Chinese and Taiwanese C. flavomarginata could be made, it was best to consider the two pop- ulations as distinct subspecies. Tanaka & Sato (1983) have referred to the turtles on the Ryukyu Island of Iriomote as Cuora f. flavomarginata indicating an affinity with the population on Taiwan. However, Fang (1934) had critically compared specimens from Taiwan and the Ryukyu Islands with Hsu’s diagnostic characters and found that the mainland Chinese turtles could not be differentiated from the insular populations. Recent examination and comparison of C. flavomarginata from these three popu- lations has shown that turtles from the Ryu- kyu Islands can be distinguished from those of the other two populations and represent an undescribed species. Methods and materials. —Sixty-six turtles were examined (Ryukyu Islands, 38; Tai- wan, 14; China, 14). Sexes were determined by the characters given by Ernst & Barbour (1989). Straight-line measurements of each specimen were taken with dial calipers ac- curate to 0.1 mm. Variables included: the greatest carapace length, carapace width and depth at the level of the seam separating vertebrals 2 and 3, marginal width (the dif- VOLUME 103, NUMBER 1 ference between the carapacial width and the width across the pleurals taken between the points of juncture of the marginals and pleurals at the level of the seam between vertebrals 2 and 3), greatest plastron length, greatest width and length of both plastral lobes, least bridge length, greatest width and length of the cervical scute and all vertebrals and medial seam lengths of all plastral scutes (Gul., Hum., Pect., Abd., Fem., An.). Care- ful notes and drawings were made of head, neck, limb, carapacial and plastral patterns. Colors were recorded from living turtles and color transparencies. Shell proportions were expressed as ratios of one measurement to another. Several ratios proved useful in our description (abbreviations used in text are given in parentheses): width/length of cer- vical scute (W/L CS), width/length of des- ignated vertebrals (W/L Ist, V W/L 2nd V, etc.), marginal width/carapacial width (MW/ CW), marginal width/carapacial length (MW/CL), carapacial width/carapacial length (CW/CL), carapacial depth/carapa- cial length (D/CL), carapacial depth/cara- pacial width (D/CW), length of light-col- ored blotch at its medial heights on pleural scute 2/total length of pleural scute 2 at the same point (PBL/PSL), plastral length/car- apacial length (PL/CL), bridge length/plas- tral length (B/PL), bridge length/carapacial length (B/CL), length of the anterior plastral lobe/plastral length (APL/PL), width of an- terior plastral lobe/plastral length (APW/ PL), width of anterior plastral lobe/length of anterior plastral lobe (APW/APL), length of posterior plastral lobe/plastral length (PPL/PL), width of posterior plastral lobe/ plastral length (PPW/PL), and width of pos- terior plastral lobe/length of posterior plas- tral lobe (PPW/PPL). The number of rows of large scales at the lateral edge of the an- tebrachium between the claw of digit 5 and the first horizontal skin fold proximal to the elbow (presented in text as FLSR) was re- corded. Statistical techniques were executed using SYSTAT (Wilkinson 1986) and STAT- 27 GRAPHICS (STSC 1986). Males and fe- males were combined for analysis since sta- tistically significant size differences were not detected (x male CL, 137.6 mm; x female CL, 143.3 mm: ANOVA, F = 0.98; df = 1,39; P = 0.33). Juveniles (23) were includ- ed in all analyses due to small total sample size (66). Males, females and juveniles were evenly distributed between population sam- ples. Variables were transformed for para- metric analysis as follows: those based on proportions, such as the interanal seam length (An) and PBL/PSL, were arcsine square root transformed, FLSR was square root transformed, and natural logarithms were taken of all others to reduce variance (Lewontin 1966, Moriarty 1977). Principal components analysis (PCA) was used as a data reduction technique to identify or- thogonal factors and their important vari- ables. Interanal seam length (An) was ex- cluded from this step because it had previously been identified to vary indepen- dently of population. Variables with high factor loadings were used in subsequent analyses as were factor scores (Kachigan 1986). Following PCA, differences among populations were tested using multivariate analysis of variance (MANOVA) as sug- gested by Willig et al. (1986). Following identification of differences among these putative populations, a three-group dis- criminant function analysis was conducted using the raw variables selected with PCA. Discriminant scores were calculated by multiplying these variables by their asso- ciated unstandardized canonical coefh- cients and summing the products. Each specimen was then plotted along the axes providing maximal separation of the a priori groups. Levels of significance were set a priori at alpha = 0.05. Specimens from the following collections were examined: American Museum of Nat- ural History (AMNH), California Academy of Sciences (CAS), Field Museum of Natural History (FMNH), George Mason Univer- sity (GMU), Museum of Comparative Zo- 28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Carapace and plastron of Cuora evelynae, new species (AMNH 50804). VOLUME 103, NUMBER 1 29 Table 1.—Summary statistics for significant variables by populations. Means (mm) are followed by one standard error in parentheses. Probabilities are given for univariate F-tests between localities. Refer to text for abbreviations. Ryukyu Islands Variable (35) CW 90.6 (3.2) APW 63.7 (2.5) PPW 71.4 (3.0) Gul 19.7 (0.7) Hum 6.0 (0.4) Abd 32.3 (1.5) FLSR 8.8 (0.2)* PBL/PSL (%) 64.6 (0.5) Population (n) Taiwan China (13) (14) Probability 74.9 (6.7) 92.0 (5.7) 0.03 50.8 (5.3) 67.2 (4.5) 0.01 57.0 (6.3) 76.3 (5.3) 0.02 16.4 (1.7) 19.6 (1.2) 0.05 6.6 (0.6) 8.8 (0.8) 0.01 24.4 (2.8) 28.9 (2.4) 0.02 10.8 (0.6) 8.9 (0.2) <0.001 34.2 (1.0) 35.3 (0.4) <0.001 * Includes four additional specimens not included in other statistical analyses. ology, Harvard University (MCZ), Natur- historisches Museum Wien, Vienna (NHMW), United States National Museum of Natural History (USNM), William P. McCord, Hopewell Junction, New York, personal collection (WPM), and William H. Randel, Hatboro, Pennsylvania, personal collection (WHR). Results and discussion. — Turtles from the Ryukyu Islands showed two pattern differ- ences from the other two populations. The pleural blotch in these specimens was larger (PBL/PSL; x = 65%) than those of turtles from Taiwan (34%) and mainland China (35%). The pleural blotch accounted for sig- nificantly different proportions of the pleu- ral scute length (PBL/PSL) between popu- lations (ANOVA; F = 703.08; df = 2,59; P < 0.0001). In 14 of 27 adults (52%) from the Ryukyu’s the blotches were so large as to coalesce dorsally, forming two lateral light longitudinal stripes in addition to the nor- mal medial stripe (Fig. 1). No specimens from either Taiwan or China had lateral stripes. The shape and coloration of the dark medial blotch on the plastron also varied (Fig. 1). Turtles from Ryukyu usually had a large, hourglass-shaped blotch, indented at the bridge, which was light brown in col- or; those from the other two populations had large dark brown to black blotches with straight sides or only a very shallow inden- tation at the bridge. Summary statistics and univariate test probabilities between pop- ulations are given for all significant vari- ables in Table 1. Eight variables, about six more than expected at alpha = 0.05, show significant differences among populations. The first three factors generated by PCA for 28 variables accounted for 93 percent of the total variance (Factor I—84%, eigen- value = 23.49, Factor II—6%, eigenvalue = 1.65, Factor III—3%, eigenvalue = 0.96). The highest loadings in Factor I are all size related mensural variables. PPL had the highest loading in Factor I (0.997). The pro- portion of pleural blotch coloration (PBL) had the highest loading in Factor II (0.790). FLSR had a very high negative loading (—0.729) in Factor II. MW had the highest loading in Factor III (0.920). All other vari- ables were highly correlated with these four and add little to the analysis, so they were not analyzed further. Factor scores for Fac- tor I did not differ significantly between populations (ANOVA; F = 2.45; df = 2,50; P=0.10), nor did scores for Factor III (AN- OVA; F = 0.46; df = 2,50; P = 0.64). How- ever, Factor II scores did (ANOVA; F = 55.85; df = 2,50; P < 0.001). Differences among populations were suggested by MANOVA when all three sets of factor 30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON DISCRIMINANT FUNCTION 2 —9 Gal 3 7 DISCRIMINANT FUNCTION 1 Fig. 2. Discriminant function analysis of turtle populations from the Ryukyu Islands(1), Taiwan(2), and southern China(3) for the characters PPL, PBL/PSL, MW, and FLSR. Polygons define extreme values of each cluster. Numbers represent individual turtles. scores were included (Wilks’ Lambda = 0.23; F = 17.76; df= 6,96; P < 0.001). Univariate F-tests for the highest loading variable in each factor were as follows: Factor I—PPL (ANOVA; F = 2.83; df = 2,59; P = 0.07), Factor II—PBL (ANOVA; F = 703.08; df = 2,59; P < 0.001), Factor WI—MW (AN- OVA; F = 0.92; df = 2,59; P = 0.40). Dif- ferences were again demonstrated using MANOVA for these three variables (Wilks’ Lambda = 0.04; F = 82.64; df = 6,114; P < 0.001). FLSR differed significantly between pop- ulations (ANOVA; F = 9.07; df = 2,56; P < 0.001). Specimens from Taiwan had a mean of 10.8 while those from the Ryukyu Islands and China had means of 8.8 and 8.9 respectively. Variables with high loadings (PPL, PBL, FLSR, MW) were entered into a discrimi- VOLUME 103, NUMBER 1 nant function analysis and each specimen was classified into a predicted population. The first and second discriminant functions provide significant (P < 0.001) discrimi- nation between populations (Fig. 2). The function correctly classified 100% of the Ryukyu specimens, 92% of those from mainland China, and 77% of the specimens from Taiwan. Discriminant scores (DS) are >—1 for specimens from the Ryukyus and <—1 for specimens of C. flavomarginata from China and Taiwan based on the fol- lowing discriminant function: DS = 3,603 ar) 33,737 SCRVLJPSIY) ae 0.02580(FLSR) + 0.01970(MW) + 0.00960(PPL). This function correctly clas- sified all specimens of both species. Although closely related to C. flavomar- ginata, the turtle population in the Ryu- kyu’s can be distinguished by the combi- nation of characters listed above. Since it is allopatric, and has apparently been isolated for more than a million years (see discussion below), we believe it to be at least an incip- ient species and have treated it as such, rath- er than asa subspecies of C. flavomarginata. Cuora evelynae, new species Fig. | Holotype. —CAS 26113, adult male; Ishi- gaki Shima, Ryukyu Islands, Japan; Victor Kuhne, 5-11 May 1910. Paratypes.—CAS 21026-21029, 26102- 26112, 26801 (five adult males, eleven adult females); AMNH 50804 (adult female); MCZ 56064 (adult male); USNM 34076- 34079 (adult male, adult female, two ju- veniles) from the type-locality. Diagnosis.—A domed species of Cuora with a yellow head; a dark brown to black carapace with a large yellow to light brown blotch which at its medial height extends 49-72% (X = 64.6) across the middle of each pleural, and at its dorsal border coalesces to form lateral stripes in 52% of adults; a large pale brown plastral blotch, which is in- dented at the bridge; the plastron lacking a 31 medial posterior notch; most adults with at least a partially obliterated interanal seam, and usually less than 10 (¥ = 8.8) rows of large scales on the anterior surface of the foreleg (a character shared with the main- land Chinese population). Description (from all specimens exam- ined).—Carapace length to 164 mm (males 159, females 164), elliptical, domed (D/CL 0.41-0.52, x = 0.468; D/CW 0.57-0.71, x = 0.648; CW/CL 0.67-0.80, x = 0.725); wid- est at marginals 8, highest at posterior of vertebral 2 or vertebral 3. Carapace sides straight, anterior marginals flared, posterior marginals at best only slightly serrated with no medial notch. Marginals over bridge downturned (MW 6.4—18.7 mm, x = 10.86; MW/CW .056-.16, x = 0.108; MW/CL 0.04— 0.12, x = 0.078). Marginals large, and ap- proximately the same width throughout. Carapacial scutes in young individuals are rugose because of growth annuli; those of older turtles are worn smooth. Cervical scute rectangular to triangular, longer than wide (W/LCS 0.46-0.95, x = 0.782). Vertebrals 3-5 wider than long, vertebral 2 usually longer than wide (79% of adults) and ver- tebral 1 may be either wider than long (53% ofadults) or longer than wide (47% of adults). Vertebrals 4 and 5 widest, vertebral 5 pos- teriorly flared, vertebral | slightly flared an- teriorly and only contacting marginal 1. Three low longitudinal keels present; me- dial keel most pronounced and extending along all 5 vertebrals, lateral keels extend from posterior of pleural 1 through pleural 4 and may disappear with age. Color black to dark brown; rim of marginals yellow. Me- dial yellow stripe always present. Areolae of pleurals and vertebrals yellow or light brown. Light areolae of pleurals large (PBL/PSL 0.49-0.72, x = 0.646), and tend to coalesce at their dorsal borders to form lateral light stripes in 52% of adults (Fig. 1). Undersides of marginals yellow. Plastron length to 160 mm (males 154, females 160), shorter than carapace in 23(85%) specimens (PL/CL 0.93-1.01, ¥ = 32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.973), movable hinge between the pectoral and abdominal scutes. Posterior lobe longer and wider than anterior lobe (APL/PL 0.40- 0.44, x = 0.420; PPL/PL 0.57-0.61, x = 0.586; APW/PL 0.49-0.57, x = 0.524; PPW/ PL 0.55-0.62, X = 0.592; APW/CW 0.67- 0.75, X = 0.712; PPW/CW 0.75-0.87, x = 0.800). Anterior lobe rounded in front; pos- terior lobe rounded in rear, usually without a medial notch. Bridge moderate (B/CL 0.27-0.37, xX = 0.311; B/PL 0.28-0.39, x = 0.318); axillary and inguinal scutes gener- ally lacking, or small. Average plastral for- mula Abd. > An. > Pect. > Gul. > Fem. > Hum.; 14 (40%) had this formula, but 4 other formulae occurred; 23 (66%) had Abd. > An., 11 (31%) had An. > Abd., 21 (60%) had Fem. > Hum., 14 (40%) had Hum. > Fem. Intergular and interanal seams often obliterated in large individuals. Plastron and bridge yellow, with a large pale brown me- dial blotch occurring on all scutes and in- dented at the bridge (Fig. 1). Head narrow, snout slightly projecting; upper jaw with slight medial hook. Dorsal- ly, the head is lemon-yellow to olive. Lat- erally behind the orbit and jaws, is a large yellow blotch that encloses the tympanum. This blotch is dark bordered dorsally. A sec- ond smaller yellow blotch lies dorsally and behind the first, and extends across the top of the head to touch a similar blotch from the other side just in front of the cervical skin fold. Jaws and chin immaculate yellow. Neck yellow to olive with no pattern. Digits partially webbed. Forelimbs with large scales (6-11 rows, X = 8.8), 29 of 34 individuals (85.3%) had less than 10 rows; outer surface olive brown or yellow to red- dish-brown, inner surface and sockets yel- low. Hindlimbs with smaller scales, colored similar to forelimbs; hindfoot olive to brown with large scales at heel and ankle. Tail yel- low dorsally and bordered by two olive or brown stripes, tip yellow. Males with moderately concave plastra, and longer, thicker tails with the vent be- yond the carapacial rim. Females with flat plastra and smaller tails with the vent be- neath the posterior marginals. Cuora evelynae is known only from the islands of Iriomote, Ishigaki, and Okinawa (Iverson 1986); the Okinawa specimen (MCZ 55838) may represent an escaped captive. These turtles probably originated from mainland China or Taiwan at a time when the Ryukyus were connected by a land bridge, or only separated by shallow water. Subsequent submergence of the land bridge isolated the Ryukyu population thus allow- ing for speciation to occur. Based on the reports of Yabe & Aoki (1923), Yabe (1929a, b), and Hanzawa (1935), Inger (1947) has presented a sum- mary of the geology of these islands. The Ryukyu cordillera arose in the late Permian or early Mesozoic Era. It seems to have bro- ken into several mountain masses between the Permian and upper Eocene. After the Eocene there have been many fluctuations in sea level great enough to have altered radically the area available for use by ter- restrial or semi-terrestrial animals such as C. evelynae. From the upper Eocene through the lower Oligocene, Iriomote and Ishigaki were probably submerged (Yabe & Aoki 1923, Hanzawa 1935), but the islands emerged again during the middle and upper Oligocene. Three additional submergences took place, finally ending in the middle Pleistocene, during which at least the high- est points of Iriomote and Ishigaki were above sea level. Between these latter sub- mergences, Iriomote and Ishigaki probably had direct land connections with China through Taiwan, and it was then that the parent stock of C. evelynae may have reached the islands. However, the possibility of tur- tles rafting from Taiwan on the Kuroshio Current can not be excluded (Lovich et al. 1985). Final inundation of this land bridge during the early to middle Pleistocene, about 1.5 million years ago, presumably isolated the turtles on the islands and subsequently VOLUME 103, NUMBER 1 separated Iriomote and Ishigaki. Inger (1947) proposed a similar scenario to ex- plain present variation in the amphibian fauna of the Ryukyus, which also shows strong Oriental affinities. Ye (1985) summarized the fossil record of Cuora in China, Taiwan and Japan, em- phasizing specimens of C. flavomarginata. The earliest records for the genus are from the late Miocene (approximately 8 million years ago) of Yunnan, and Pleistocene re- mains are known from Kyushu and Hon- shu, Japan. This raises the possibility that the Ryukyu Islands population of C. eve- lynae may have reached there by island hopping along the Japanese archipelago. However, since no fossils of the genus are known from Korea and no living species of Cuora reaches there, it is more likely that movement was in the opposite direction and that the Kyushu and Honshu turtles came from mainland China by way of Taiwan and the Ryukyus. Other material.—Cuora evelynae: CAS 21015-24, 26114-15; GMU 730-31; MCZ 7997, 55838. Cuora flavomarginata: AMNH 110181; CAS 10834-40; FMNH 121225-26, 127324, 216515; GMU 975; NHMW 29515, 29518; USNM 140825; WPM 1-S(live); WHR 1-7(live). Etymology.—The genitive noun used as the specific epithet honors Dr. Ernst’s wife Evelyn for her years of encouragement and help, and for her contributions to the study of turtle helminths. Acknowledgments Thanks are given to those individuals who allowed examination of turtles in their col- lections. Dr. George R. Zug and two anon- ymous reviewers offered valuable sugges- tions for improvement of the manuscript. Manuscript preparation was supported by DOE grant DE-AC09-76SR00819 to the University of Georgia, Savannah River Ecology Laboratory. 33 Literature Cited Ernst, C. H., & R. W. Barbour. 1989. Turtles of the world. Smithsonian Institution Press, Washing- ton, D.C., 313 pp. Fang, P. W. 1934. Notes on some chelonians of China.—Siensia 4:145-199. Gray, J. E. 1863. Observations on the box tortoises, with the descriptions of three new Asiatic species. — Proceedings of the Zoological Society of London 1863:173-179. Hanzawa, S. 1935. Topography and geology of the Riukiu Islands.—Tohoku Imperial University Science Report, (2) Geology 18:1-61. Hsu, H.-F. 1930. Preliminary note on a new variety of Cyclemys flavomarginata from China. —Con- tributions from the Biological Laboratory of the Science Society of China, Zoological Series 6:1-7. Inger, R. F. 1947. Preliminary survey of the am- phibians of the Ryukyu Islands. — Fieldiana:Zo- ology 32:296-350. Iverson, J.B. 1986. A checklist with distribution maps of the turtles of the world. Privately printed, Paust Printing, Richmond, Indiana, 283 pp. Kachigan, S. K. 1986. Statistical analysis. Radius Press, New York, 589 pp. Lewontin, R. C. 1966. On the measurement of rel- ative variability.—Systematic Zoology 15:141- 142. Lovich, J. E., C. H. Ernst, & S. W. Gotte. 1985. Geo- graphic variation in the Asiatic turtle Chinemys reevesii (Gray) and the status of Geoclemys gran- geri Schmidt.— Journal of Herpetology 19:238- 245. Moriarty, D. J. 1977. On the use of the variance of logarithms.—Systematic Zoology 26:92-93. Pope, C. J. 1935. Natural history of Central Asia, Volume X. The reptiles of China. American Museum of Natural History, New York, 604 pp. 1986. Statgraphics users guide. STSC Inc., Rockville, Maryland. Tanaka, S., & F. Sato. 1983. Brief observation of the mating behavior of the box turtle Cuora flavo- marginata in nature.— Biological Magazine, Okinawa 21:75-76. Wilkinson, L. 1986. SYSTAT: the system for statis- tics. SYSTAT Inc., Evanston, Illinois. Willig, M. R., R. D. Owen, & R. L. Colbert. 1986. Assessment of morphometric variation in nat- ural populations: the inadequacy of the univar- iate approach. —Systematic Zoology 35:195-203. Yabe, H. 1929a. The latest land connection of the Japanese Islands to the Asiatic continent. — Pro- ceedings of the Imperial Academy of Tokyo 5: 167-169. . 1929b. Geologic age of the latest continental STSC. 34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON stage of the Japanese Islands. — Proceedings of the Imperial Academy of Tokyo 5:430-433. ——. & R. Aoki. 1923. A summary of the strati- graphical and paleontological studies of the Cai- nozoic of Japan.—Proceedings of the Pan-Pa- cific Science Congress (Australia) 1:954-969. 1985. Studies on fossils of Cuora of China and Japan.—Acta Herpetologica Sinica 4:81— 87. Ye, X. (CHE) Department of Biology, George Mason University, Fairfax, Virginia 22030; (JEL) Savannah River Ecology Laboratory, P.O. Drawer E, Aiken, South Carolina 29801. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 35-38 A NEW OCELLATED FROG (CENTROLENIDAE) FROM WESTERN COLOMBIA John D. Lynch Abstract. — A new species of Centrolenella (C. ignota) is described from cloud forests of the Cordillera Occidental in Colombia. The new species is most closely allied to the Ecuadorian C. anomala, the only other brown centrolenid known. Knowledge about the Neotropical frog family Centrolenidae has increased from a view of a small family of one or two dozen species seemingly centered in Central America in 1950 to a modest-sized family of at least 65 species centered in Colombia. Our current ideas of relationships within the family are much in need of study and largely reflect the perspectives of the Costa Rican fauna (Savage 1967, Savage & Starrett 1967, Starrett & Savage 1973). Lynch & Duell- man’s (1973) arrangement closely paral- leled that from Costa Rica but emphasized the fact that many species from northwest- ern South America seem to strain the Costa Rican arrangement. Lynch & Duellman (1973) named one pe- culiar species from the Amazonian slopes of the Andes in Ecuador as Centrolenella anomala because it was tan, not green, in life. They treated the species as the sole member of a species group (op. cit.:58). While collecting on the western flank of the Farallones de Cali in western Colombia in July 1979, I obtained a series of a small centrolenid that initially I thought to be C. anomala because it was tan at night. During the day the animals changed to olive-brown. The Colombian frogs are intermediate, at least in part, between C. anomala and the other ocellated Centrolenella. Centrolenella ignota, new species Fig. 1 Holotype.—ICNMNH 14748, an adult male, 24.4 mm SVL, from a series collected at Penas Blancas, Farallones de Cali, ca 6 km by road SW Pichinde, Depto. Valle de Cauca, Colombia, 1900 m, 4 Jul 1979 by Humberto Carvajal and John D. Lynch. Paratypes. -ICNMNH 14749-77, KU 209763-65, taken with the holotype. Diagnosis. — 1) vomerine teeth and odon- tophores absent; 2) bones very pale green in life; 3) parietal peritoneum white, visceral peritoneum clear; 4) color in life tan-brown to very pale olive with black ocelli contain- ing orange or yellow centers; in preserva- tive, very pale lavender with black ocelli; 5) outer fingers with basal webbing III 3*-37 IV; 6) webbing on foot I 2--2+ IT 1'%2-2% III 142-275 IV 3 -1% V; 7) snout truncate in dorsal and lateral profiles; 8) dorsal skin shagreened with elevated warts correspond- ing to ocelli; 9) arms and legs lacking dermal fringes; 10) humeral spine absent; 11) lower three-fourths of tympanum visible, directed anterolaterally with slight posterior incli- nation. Centrolenella ignota is most similar to C. anomala but differs in lacking the dark brown flecks that are interspersed among the ocelli and in lacking the spicules on the skin of the dorsum; furthermore, C. ignota has subanal tubercles. The two species may differ as well in that C. 7gnota has pale green bones (white in the only specimen of C. anomala ever captured) and has traces of green pigmentation (C. anomala never ex- hibited any green cast in its dorsal pattern). Description. — Adults small, snout-vent length in males 22.3-25.4 mm (x = 23.9 [S.E. = 0.12], n = 31), in two females 24.2 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ee ieee Fig. 1. Centrolenella ignota, topotype (ICNMNH 18007, male, 23.8 mm SVL). and 24.4 mm; head wider than body; width of head 31.5-34.8% (x = 33.4 + 0.1l,n = 35) of snout—vent length; snout short, high, truncate in dorsal and lateral profiles; can- thus rostralis round; loreal region flat; lips not flared; nostrils at tip of snout; internarial region concave, nostrils proturberant, di- rected anterolaterally; eye to nostril distance 53.1-75.0% (X = 62.5 + 1.0, n = 35) eye length; eyes large, directed anterolaterally; width of upper eyelid 76.0—1 14.3% (X = 96.7 + 1.4, n = 35) interorbital distance; supra- tympanic fold obsolete; tympanic annulus distinct, tympanum round to slightly higher than long, directed dorsolaterally with slight posterior inclination; length of tympanum 18.8-27.6% (xX = 22.7 + 0.4, n = 35) eye length; choanae large, round, not concealed by palatal shelf of maxillary arch; no vo- merine odontophores or teeth; tongue round to ovoid, bearing a shallow notch poste- riorly, posterior edge not adherent to floor of mouth; males with vocal slits postero- lateral to tongue; males with median subgu- lar vocal sac. Forelimb moderately slender; no humer- al spine or hook; no ulnar tubercles or folds; palmar tubercle round to ovoid, larger than oval thenar tubercle; no supernumerary tu- bercles on palm; subarticular tubercles low, basal tubercles broader than long, more dis- tal ones round; fingers with lateral keels; median edge of fourth and lateral edge of third fingers bearing fleshy ridge (confluent with basal webbing); webbing formula III 3+-(2%4-3-) IV; first finger longer than sec- ond; all fingers bearing discs, discs rounded apically but broader than long (subtrun- cate); discs of fingers II-IV largest, but all discs larger than tympanum; males with swollen base of thumb, non-spinous nuptial pad on dorsal surface of thumb (metacarpal section only); hind limbs slender; length of shank 52.1-58.7% (X = 55.0 + 0.3, n = 35) snout—vent length; tarsal tubercles and folds absent; inner metatarsal tubercle oval, flat; outer metatarsal tubercle apparently absent; no supernumerary plantar tubercles; subar- ticular tubercles small, round; toes about one-half webbed; webbing formula I (2—2>)— (2*-2%) IT (1 '4-1%4)-(2'2-2%) TM (14-12) (2%-3°-) IV (2%-3*)41%2-1%) V; discs of toes smaller than those of fingers, round to subtruncate. Skin of dorsal surfaces smooth to very finely shagreened, lacking spicules; white spots are elevated flat warts; venter and pos- terior surfaces of thighs bearing flat areo- lations; anal opening under a short trans- verse flap at upper level of thighs; pair of enlarged flat warts on posteroventral surface of thighs (subanal warts). Color in preservative: Cream above with pale violet stippling over head, dorsum, and upper surfaces of limbs (this stippling is very fine and provides a pale lavender wash to the dorsal surfaces); dorsum bearing dense violet stippling around bases of white warts (forming ocelli); ocelli generally on top of head, back, and on shank but, in some in- dividuals, also on side of head, top of thigh, and tarsus near heel; ventral surfaces cream. Color in life: Pale tan to olive-brown above with black ocelli having orange (or VOLUME 103, NUMBER 1 yellow) centers; tips of digits yellow; parietal peritoneum white; bones very pale green; iris whitish gray with a gold cast and black reticulation. Remarks. — When the majority of speci- mens were collected (4 Jul 1979), the area was receiving a light rain. During the rain the frogs were calling actively on vegetation 0.5 to 2 m above a sluggish stream (average width 0.5 m). Females were found sitting on vegetation. Some individuals were found as much as two meters away from the stream in dense vegetation. The call of C. ignota is a series of chirps. The discovery of C. ignota, with its non- green coloration, provides evidence that the coloration of C. anomala is probably nat- ural (some colleagues have doubted that brown centrolenids exist). The nearest rel- atives of C. ignota appear to be C. anomala and C. cochranae. These three species have small ocelli on the dorsum bearing reddish centers (orange-tan in C. anomala, red in C. cochranae, orange or yellow in C. ignota) and exhibit virtually the same head shape and degree of webbing; differences among the three species are slight. In addition to coloration differences, they differ in the presence of subanal warts (absent in C. anomala), vomerine dentition (usually present in C. cochranae), skin texture (spic- ules in C. anomala and C. cochranae), and adult size (C. cochranae is larger than the other two, cochranae males are 23.8—26.7 mm snout—vent length and females are 27.2— 30.0 mm snout-—vent length). At present, I consider C. ignota and C. anomala to be sister species and consider C. cochranae the sister species of the pair of tan/brown species. The conjectured syn- apomorphies are (1) brown pigmentation (shared by anomala and ignota) and (2) small ocelli on elevated warts. The plesiomorphic conditions are (1) green pigmentation (true for all other centrolenid frogs) and (2) no ocelli (true for nearly all other centrolenids, see below). There are only two other species of cen- 37 trolenids having ocelli (C. ocellata and C. ocellifera). Each has more webbing on the hand than the three previously cited and each has larger ocelli that are not on ele- vated warts (also cream or pale yellow in life). It is tempting to assert that the ocelli constitute a synapomorphy for these five species but a case could be made for arguing that the ocelli of C. ocellata and C. ocellifera are not homologous to the ocelli of C. anomala, C. cochranae, and C. ignota but rather are homologous to the open reticu- lation found in the dorsum of centrolenids of the fleischmanni group. The published illustration of C. grandisonae in Cochran and Goin (1970) shows ocelli, although in their description it is apparent that they did not find ocelli in that species. Centrolenella grandisonae has red (flat and elevated) warts on the dorsum. Upon preservation the red disappears leaving cream spots. Etymology.—The trivial name is Latin (ignotus), for strange, and is used to reflect the annectent coloration of C. ignota be- tween that of C. anomala and the green col- oration seen in most species of the genus. Acknowledgments I thank Humberto Carvajal for assistance in the field and Stephen C. Ayala, William E. Duellman, and Pedro M. Ruiz for pro- vision of working space. Museum abbre- viations used in the text refer to collections at the Instituto de Ciencias Naturales in Bo- gota (ICNMNH) and the Museum of Nat- ural History at the University of Kansas (KU). Literature Cited Cochran, D. M., & C. J. Goin. 1970. Frogs of Co- lombia.— Bulletin of the United States National Museum 288:1-655. Lynch, J. D., & W. E. Duellman. 1973. A review of the centrolenid frogs of Ecuador, with descrip- tions of new species.—-University of Kansas Mu- seum of Natural History, Occasional Papers 16: 1-66. Savage, J. M. 1967. A new tree-frog (Centrolenidae) from Costa Rica.—Copeia 1967(2):325-331. 38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON —, & P. H. Starrett. 1967. A new fringe-limbed Southern California Academy of Science 72:57— tree-frog (family Centrolenidae) from lower 78. Central America.—Copeia 1967(3):604-609. Starrett, P. H., & J. M. Savage. 1973. The systematic School of Biological Sciences, The Uni- status and distribution of Costa Rican glass-frogs, : : genus Centrolenella (family Centrolenidae), with eee of Nebraska, Lincoln, Nebraska description of a new species.— Bulletin of the PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 39-62 ON THE VALIDITY OF THE INDO-PACIFIC CARDINALFISHES APOGON AUREUS (LACEPEDE) AND A. FLEURIEU (LACEPEDE), WITH DESCRIPTION OF A RELATED NEW SPECIES FROM THE RED SEA John E. Randall, Thomas H. Fraser, and Ernest A. Lachner Abstract. —Two large species of Indo-Pacific cardinalfishes of the genus Apo- gon, subgenus Ostorhinchus, previously identified either as A. aureus (Lacepéde) or A. fleurieu (Lacepéde), are shown to be distinct. Their most characteristic color marking is a black bar encircling the caudal peduncle. They are separated by gillraker counts (22-27 for aureus, 19-23 for fleurieu) and the form of the black peduncular marking (a spot on juvenile fleurieu, expanding to a bar in adults; always a bar in aureus, typically broader dorsally and ventrally to a slight hourglass shape. Both species occur from East Africa (only fleurieu in the Red Sea) to the western Pacific; fleurieu is known only from continental shelf localities except for the Seychelles. A third species with a black peduncular bar, A. pselion, is described as new from specimens from the northern Red Sea. It is more slender (depth 2.8-3.4 in SL), small (largest 41.3 mm SL), and distinctive in life color (four dusky yellow stripes on head separated by blue lines, one stripe continuing as a yellow band midlaterally on body). The Apogonidae (popularly known as cardinalfishes) is one of the largest families of tropical fishes; the great majority of these fishes are found in the marine environment. Apogonids are small (only a few species ex- ceed 20 cm total length), with two separate dorsal fins (the first of VI to VIII spines), II anal spines, a double-edged preopercle, large eyes, and a large oblique mouth. Most are nocturnal, and those for which the repro- ductive strategy is known are mouth brood- ers. Fraser (1972) recognized three subfami- lies and 20 genera in the family. Nelson (1984) wrote that there are about 192 species, but it is clear from the number of unde- scribed species on museum shelves and probably more that remain to be discovered in the sea that well over 200 species will eventually be recorded. There has long been confusion over the correct specific name for a large species of Apogon with a broad black bar posteriorly on the body which occurs in the Indian Ocean and western Pacific. Most authors have used the name Apogon aureus (La- cepéde) for this fish, but some (Gon 1987, gave 12 references') have called it A. fleurieu (Lacepéde). Lacepéde (1802:23) described the latter as Ostorhinchus fleurieu; his illus- tration from a drawing by Commerson was published as fig. 2 of pl. 32 in volume 3 of Histoire Naturelle des Poissons (1801) (re- produced by Gon 1987: fig. 2). Lacepéde’s description, obviously based on Commer- son’s drawing and not a specimen, appeared in volume 4 (1802:23). Because no teeth are apparent on the drawing, Lacepéde as- sumed that they were fused to form dental plates like those of scarids, diodontids, and tetraodontids. This led Whitley (1959) to ' Weber & de Beaufort (1929:319) used Apogon au- reus, not A. fleurieu. 40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON regard fleurieu as an oplegnathid. Smith (1961) realized that Lacepéde’s figure is an apogonid. He wrote, ““Most workers have refused to accept Lacepede’s fleurieu on the grounds that it is not an Apogonid fish. To me his 1802, Pl 32, fig. 2 of fleurieu rep- resents nothing else, ...” Fraser (1972) discussed the controversy in detail. He opted for the use of Apogon aureus (which was described by Lacepéde, 1802:253, 273, 275, as Centropomus aureus) because the description of Ostorhinchus fleurieu “does not agree with any apogon- id’’. He was influenced in his decision by Lacepéde’s description of the dentition, the apparent lack of a lateral line, and the high number of rays of the second dorsal fin (14). He concluded, “‘Perhaps the best course of action would be the rejection of the name on the basis that it is an unidentifiable tax- on.” The second author suspected as early as 1974 that two species had been confused under the presumed synonymous names Apogon aureus and A. fleurieu. About five years ago both junior authors had deter- mined that there were indeed two species and differentiated them by gillraker count and the form of the dark peduncular bar. In one species both juveniles and adults have a well-defined dark peduncular bar that is usually expanded dorsally and ventrally to form a slight hourglass shape. Juveniles of the other species have a dark spot centro- posteriorly on the peduncle which expands with growth to form a solid bar, but without dorsal and ventral expansions. At that time the junior authors were inclined to preserve the name Apogon fleurieu for one species and describe the other as new. They were subsequently joined by the senior author, and with the analysis of more specimens and the original Lacepéde accounts of the two species, it was concluded that both the names fleurieu and aureus should be rec- ognized. In the meantime Gon (1987) re- viewed the problem of Lacepéde’s apogonid names fleurieu and aureus. Like Smith (1961), he regarded Lacepéde’s illustration of fleurieu as identifiable and placed aureus in its synonymy. He designated a neotype for fleurieu (BPBM 15821, 94 mm SL, from Papua New Guinea) and illustrated it. Lacepéde’s figure of fleurieu exhibits a pe- duncular bar that is narrower dorsally and ventrally, thus typical of a specimen that had a peduncular spot as a juvenile but had not yet developed a complete bar. It is clear from Gon’s (1987) description of fleurieu and his material that he had both species. Unfortunately he chose a specimen with the hourglass peduncular bar as the neotype of fleurieu. If we were to follow Gon in his neotype designation, we would need to de- scribe the other species as new. We prefer to link fleurieu to the species with the pe- duncular bar like Lacepéde’s figure bearing this name, thus leaving the species with the hourglass bar as aureus. It may be argued that Lacepéde’s description of Centropomus aureus 1s not diagnostic for either of these species. However, the type localities of Mauritius and Réunion strongly suggest that the one with the hourglass bar was the species described by Lacepéde. We have examined specimens of this species from six collec- tions from Mauritius. Unfortunately, all are in poor condition and none merits neotype designation. The other species, A. fleurieu, is known thus far only from continental shelf localities except for three lots from the Sey- chelles which are stranded continental frag- ments (for discussion see Springer 1988:128-— 129). In retaining the name Apogon aureus for the species which is the best represented of the two in museum collections and most often reported in the literature by this name, we are being less disruptive to nomencla- tural stability than if we were to adopt the name fleurieu for this fish. Because of the great similarity of Apogon aureus and A. fleurieu we have included lit- erature records of these two species and plotted the distributions of Fig. 1 only from specimens reported in sufficient detail to VOLUME 103, NUMBER 1 permit identification (generally this meant a good illustration) or from the examination of specimens. As pointed out by both Fraser (1972) and Gon (1987), the recognition of Apogon fleu- rieu as a valid taxon will result in Ostorhin- chus Lacepéde (type series, O. fleurieu La- cepéde) replacing Nectamia Jordan as a subgenus of the genus Apogon. See Fraser (1972) for a diagnosis of Ostorhinchus (as Nectamia). Bleeker (1874) recorded Dipterodon hex- acanthus Lacepéde (1801:pl. 30, fig. 2; 1802: 166, 168) as a junior synonym of Amia (=Apogon) aurea Lacepéde. He was fol- lowed by Day (1875) and Weber & de Beau- fort (1929) (though questioned). Barnard (1927) stated that this nominal species can- not be included in the synonymy of aureus because the description of the dentition is not that ofan Apogon. Fowler & Bean (1930), however, did not agree with Barnard, point- ing out that both Ostorhinchus fleurieu and Dipterodon hexacanthus have a dark trans- verse band across the caudal peduncle. They added, “The large teeth shown in the figure of the latter we think an error in engraving.” Fraser (1972) admitted that D. hexacanthus could be an apogonid, but at best a Chei- lodipterus, not an Apogon. He wrote, “I treat this name as a nomen dubium, perhaps in- volving a member of the Apogonidae.”’ Gon (1987) accepted Fraser’s opinion, and we also concur. Specimens of a third species of Apogon with a dark peduncular bar that were col- lected by the senior author and colleagues in the northern Red Sea represent an un- described species. Though notably smaller than either aureus or fleurieu, this species could easily be misidentified as the young of the former. The purpose of the present paper is to provide descriptions of all three species and to differentiate them. Apogonid material for the present study has been examined at or obtained from the following institutions: Australian Museum, Sydney (AMS); Academy of Natural Sci- 41 ences of Philadelphia (ANSP); British Mu- seum (Natural History), London (BMNH); Bernice P. Bishop Museum, Honolulu (BPBM); California Academy of Sciences, San Francisco (CAS, SU); Hebrew Univer- sity, Jerusalem (HUJ); Museum National d’Histoire Naturelle, Paris (MNHN); Na- tional Science Museum, Tokyo (NSMT); Queensland Museum, Brisbane (QM); Rijksmuseum van Natuurlijke Historie, Leiden (RMNH); J. L. B. Smith Institute of Ichthyology, Grahamstown (RUSI); Tel Aviv University, Ramat Aviv (TAU); U.S. National Museum of Natural History, Washington, D.C. (USNM); and Western Australian Museum, Perth (WAM). Lengths given for specimens are standard length (SL), measured from the front of the upper lip to the base of the caudal fin (pos- terior end of hypural plate); body depth is the depth from the base of the anterior dor- sal spines; body width is measured just pos- terior to the gill opening; head length is mea- sured from the front of the upper lip to the end of the opercular membrane, and snout length from the same anterior point to the fleshy edge of the orbit; orbit diameter is the greatest fleshy diameter and interorbital width the least bony width; caudal peduncle depth is the least depth, and caudal pedun- cle length the horizontal distance between verticals at the rear base of the anal fin and the caudal-fin base; lengths of fin spines and soft rays are measured to their extreme base; caudal concavity is the horizontal distance between verticals at the tips of the shortest and longest caudal rays. Pectoral-ray counts include the rudimen- tary upper ray; lateral-line scale counts are made to the base of the caudal fin (hence do not include the pored scales posterior to the hypural plate); gillraker counts are made on the first gill arch and include all rudi- ments; the count of the upper-limb rakers is given first; the raker at the angle is con- tained in the lower-limb count. Counts of the rays of the median fins are the same for all three species. Pectoral-ray 42 Table 1.—Gillraker counts of species of Apogon. Total gillrakers Lower limb Upper limb 2) 26 25 24 23 22 21 20 18 16 14 40 59 84 10 93 73 aureus (total) 31 35 Comoros 40 59 84 10 58 13, Other localities 74 16 23 49 129 30 fleurieu (total) 38 24 28 Red Sea PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Seychells 28 13 36 15 52 25 19 15 Other localities 20 pselion counts are nearly always 14. Gillraker counts, however, are useful in distinguish- ing the new species and Apogon fleurieu from A. aureus (Table 1). Tables 2-4 present proportional mea- surements of specimens of the three species as percentages of the standard length. Body and fin proportions are given in the text to the nearest 0.05. Paratypes of the new species are listed chronologically by date of collection. Data in parentheses in the description of the new species refer to paratypes. Apogon aureus (Lacepéde) Figs. 1-4; Tables 1, 2 Centropomus aureus Lacepéde, 1802:253, 273, 275 (type localities, Mauritius and Réunion). Apogon roseipinnis Cuvier in Cuvier & Va- lenciennes 1829:490 (type locality, Sr Lanka).— Valenciennes in Cuvier & Va- lenciennes 1830:553 (Ambon). Apogon annularis var. roseipinnis (non Cu- vier) Gunther, 1859:239 (in part) (Hong Kong). Apogon annularis (non Ruppell) Playfair & Gunther, 1867:20 (Zanzibar).— Boulen- ger, 1887:655 (Muscat, Oman). Amia aurea Bleeker, 1874:48 (Indonesian localities). — Bleeker, 1873-1876:92, pl. 3375 10S, II. Apogon aureus Macleay, 1883:236 (Port Moresby, New Guinea).— Sauvage, 1891: 142 (Sulawesi, Sri Lanka, Mauritius, and Madagascar).— Weber & de Beaufort, 1929:319 (Indonesian localities, northern New Guinea). — Munro, 1955:120, pl. 21, fig. 329 (Sri Lanka). — Woodland & Slack- Smith, 1963:31 (Heron Island, Great Bar- rier Reef).—Ida & Moyer, 1974:114, fig. 5B (Miyake-jima, Japan). — Masuda et al., 1975:204, pl. 37G (southern Japan).— Burgess & Axelrod, 1975:1447, fig. 10 (Vanuatu). —Fourmanoir & Laboute, 1976: 288 (figure with erroneous color) (New Caledonia).— Burgess & Axelrod, 1976: 43 VOLUME 103, NUMBER 1 GiGe 6°€7 €°97 8ST 6 9b p97 (5 74 O'€7 ANG yisug] Uy TA[ag 9°91 Si TLI 9°91 CSI 6S L9l €91 091 6r1 yisug] outds otAjadg GEG b'97 6°87 L'87 187 GEC WG GSC 6°97 6°97 yisus] uy [e10190g mo) Z Ol = 6 TI lel O7I 6 TI = = a AjIABoUOD [epne) vTE CEE uayo1q LSE CSE 7 9E TSE udyo1q uayoiq OSE yisug] uy [epne) uayx01q O'€T (On 16 9°€7 lv 0°97 6 V7 6 €°S7 CET Avi [eue jsasuo] Jo ysuaT] CSI Lvl GEI 79 091 6S Cr LSI 091 Lvl auids [vue puodas jo YyIsUaT LY 9°€ eS v'Y 9'€ 9b tL gE (52 lv oulds [eue jsiy jo ysUaT L'97 ASG 917 8°LTZ SG 67 187 0°97 udyo1q v'ST7 Avi [esiop jsaduo] Jo yIsuaT SI CSI 6 LI OLI 191 € 91 691 091 9°91 all uy [esIOp puodsas jo sulds jo ysuaT GGG uadyo1q L‘07 PCC IE 9°07 p17 uayo1q 707 Sal auids [esIop YLINoj Jo yIsuaT CET lialic 917 1€7 EG 8°07 VIZ uayoiq 9°07 LI auids [esiop pily) Jo ysuIT 9°01 0°6 Gl ZO vit 8°01 101 9°01 8°6 9°6 aulds [esiIop puodsas jo yIsUIT 6'€ TE lv Or ev 6'€ rs €v TE ce aulds [esiop is1y JO ysUIT L6€ 0'0r Lv CIP b' Or 7 Or 9°0r O'8E 7 Or €°6€ yysug] otAjadaig 19 1$9 6'S9 L°S9 ps9 C79 L'€9 0'€9 8°79 €°€9 yisug] [eurolg SIP Lv cep Liv TEP GGh, €€V OIr L‘Ov € Cr yisug] [esiopoid 617 v€7 O'€Z OCC CET i 4 PCC LET 9°€7 I'v yisua] sjounpad jepned 9LI LI OLI pL 9°91 GE ei CLI 8°91 THI yidap sjounpad [epne) €°07 8°61 LIZ O17 v'07 v0 1°07 9°61 707 6°07 yysua] mef ioddy) €6 16 8°6 66 L6 66 9°6 68 L6 9°6 YIPIM [11Q1O191U] Cel 97TI Cri trl pr er SEI Or Trl 8°SI J9]9WeIp 11q10 8°6 L8 16 66 68 €6 €6 68 68 88 yi3ug] Nous €6€ LSE € Or v'0r Liv 6'6£ Ir 9°8E 9°6€ € Or yisusg] peoH Sr TLI 6'SI €SI OSI € 9] 6S v'9l 0°91 LSI yipIm Apog VIP Orr 47 7 Or vir € Or b'6€ 0'0r 88E TSE yidap Apog ¢80I 9°66 0'r6 0'16 718 8°9OL SOL 79S 6'6P Tee (uu) YdUI] PrepuLIg TS107Z LOELT 76S 1 8r90E 8P90E LOST OIr6l ESELT €SELT 88ECT Wddd ns Wadd Wddd Wddd SVO Wadd Ns ns Wadd ‘yZus] Prepueys oy} Jo odejUs0I0d e se possoIdxo snainv UuOsoOdP JO sudUIIOSdS JO SJUSWIoINSsvoW [eUuOTIIOdOIg—"Z IQuL PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 44 9°€7 LEZ €S7 vv 8'r7 8°SZ 797 9°€7 9°€Z (4 yisug] Uy OTATag Sr CSI 9S 0°91 LSt E91 L9l CSI vy Ol Ga yysug] outds otajod 9°SZ €97 BLT LLZ 797 €°97 PLE 1°97 1°97 9°97 yisug] uy [e10}00g = € 01 001 OTT 601 L’6 611 8°6 = vl AjIABOUOO [epNneD usyxO1q TCE 9 EE TCE TCE €€€ CEE SiSG uayoiq pee yisuz] uy [epned usxOI1qG 8°77 L'€T 9°b7 v€T €°€7 8°77 LET uayoiq (a X6 Avi [Bue jsosuo] Jo yIsUuIT Lvl 79 Ot 9°SI Srl 8cI LSM or 791 GI suids [eue puodsas jo ysusT Or LY Sr 61 ev LY SL uayxo1q 8b I'v oulds yeue js1y Jo yIsUIT 697 VST 187 €L7Z 8°57 V'L7 GEG 6°97 v'LT ELT ABI [BSIOP }sosUO] JO YISUIT BSI 191 p91 L9l 6ST Lest €91 791 Gl LOI uy [esIop puooas jo oulds jo y3UueT 607 LAG SIZ O17 uayo1q 661 917 C61 9°81 761 auids [es1op YyVINOJ Jo ysUIT 761 L:07 9°07 O17 0°02 707 617 £61 0°61 €°07 ouids [esiop pil} Jo yIsUTT 06 v6 00! 601 v'Ol €01 ane uayo1q OTT 8°01 aulds [esIop puodas fo yIsUIT €€ Ie 1€ Iv Ip gE €€ GE 6'€ 9°€ aulds [esiop jsiy JO yBUIT 8°6€ 8°6E vir 0'0r b Or 6°6€ 7 6E CBE OLE 8°6£ yysug] otAjadaig 669 v9 199 g°S9 b'€9 169 peo 119 0°79 ps9 yisug] [eurolg OTP TEV TEP elv Clr (4 7 8°r OEP TIP Lty yisug] [esiopaig O17 0°77 8°07 617 O'€7 17 917 COG €€7 LET yidug] ajounped jepneD L91 8°91 191 LOI TOI 8°91 Ieee 191 9°91 8ST yidap ajounpad jepneD 661 L'07@ E17 €'07 707 LOG 661 8°61 0°07 917 yisuz] mel add 68 8°8 16 €6 68 9°6 66 8°6 9°6 66 YIPIM [B1IGIOIO} UT 671 Vel 9 €I TEI CEI 9EI EE 9°€1 orl LOI J2]9WIPIP 11G1O 66 v6 L'6 €6 ¢:6 9°6 €6 66 b°6 16 yisua] nous 9°6€ 9°6€ vir 8°6E 6 0b ¢ 0b vir L6€ 7 Or Ctr yisus] peo L91 9°91 PLT LI ELI pL 691 bOI 7 OI p91 yipim Apog Llp Clr 60b 9°0r OSE 6 °6€ bp 8€ L9€ 7 9E TSE yidop Apog 1ZOl 796 GG Lape 9°OL 7S9 €°09 67S OTP 6°CE (Wu) YSU] PrepuLIS PL8Ie T1861 OI PTIT 9IPTITZ OIPTITZ 8E9LT 8E9LT CCO6II pOLI6! 8S78I Wadd Wddd WNSn WNSn WNSn. 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(western Sulu Sea, Philippines). — Russell, 1983:47 (southern Great Barrier Reef). — Hayashi & Kishimoto, 1983:27, pl. 5, fig. 21 (iriomote Island, Ryukyus and Ose- zaki, Shizuoka Prefecture).—Shen, 1984: 48, figs. 304-6a, 6c (Taiwan).— Masuda et al., 1984:147, pl. 131 I (southern Ja- pan).—Gloerfelt-Tarp & Kailola, 1984: 145, 327, fig. on 144 (off Bali).—Gon in Smith & Heemstra, 1986:549, pl. 48, fig. 175.3 (East Africa south to Durban).— Shao & Chen, 1986:93, fig. 38 (Tai- wan).—Allen & Steene, 1987, pl. 35, fig. 2 (Christmas Island, Indian Ocean).— Al- Baharna, 1986:71, fig. (Bahrain). Amia fleurieu (non Lacepéde) Fowler, 1918: 65 (Philippines).—Fowler, 1927:274 (Philippines).— Fowler & Bean, 1930 (Philippine and Indonesian localities and Borneo). Apogon fleurieu (non Lacepéde) Smith, 1949:207, pl. 22, fig. 481 (Natal and Mo- zambique).—Fourmanoir, 1957:83, fig. 59 (Comoro Islands).—Shen & Lam, 1977:177, fig. 21 (Taiwan) (misspelled fleurien).— Gon, 1987:140, fig. 1 Gn part). Ostorhynchus fleurieu (non Lacepéde) Smith, 1961:399, pl. 46D (in part).—Smith & Smith, 1963:20, pl. 60D (Seychelles). Gronovichthys aureus Munro, 1967:244, 251, pl. 29, fig. 432 (New Guinea). Diagnosis.—A species of the genus Apo- gon, subgenus Ostorhinchus, with dorsal rays VII-1,9; pectoral rays 14 (rarely 13 or 15); lateral-line scales 24; median predorsal scales 5; total gillrakers 22—27; preopercular ridge smooth, the posterior margin and most of ventral margin serrate; body depth 2.25- 2.85 in SL; interorbital width 4.05—4.35 in head; posterior nostril usually only slightly larger than anterior; pale (coppery with ir- idescence in life) with a black bar encircling posterior caudal peduncle in both young and adult, its midlateral width one-half to two- thirds orbital diameter, its upper and lower edges expanded to form a slight hourglass shape, a broad blackish stripe from front of snout to orbit, continuing behind eye, this stripe bordered by a blue line in life; a nar- row blackish streak (blue in life) on maxilla and continuing a short distance posterior; a small blackish spot on each lateral-line scale; a row of dark dashes basally in anal fin; maximum standard length about 125 mm. Description. — Dorsal rays VII-I,9, all rays branched, the last to base; anal rays II,8, all rays branched, the last to base; pectoral rays 13(2), 14(64), 15(4), the upper two and low- er one or two unbranched; pelvic rays I,5; principal caudal rays 17, the upper and low- er unbranched; upper and lower procurrent caudal rays 8 or 9, the posterior two seg- mented; lateral-line scales 24 (plus 4 pored scales posterior to caudal-fin base, the last long and pointed); scales above lateral line to origin of dorsal fin 2; scales below lateral line to origin of anal fin 6; median predorsal scales 5, the fourth and fifth deeply indented posteriorly in the midline; circumpedun- cular scales 12 (minimum zigzag count); gillrakers 6-8+ 16-20, 1-2 upper and 0-1 lower as rudiments, the total count 22-25: pseudobranchial filaments 16 in a 33 mm specimen, 27 in a 92 mm one, and 32 ina 108.5 mm one; branchiostegal rays 7; pre- dorsal bones 3; vertebrae 10+ 14. Body moderately deep, the depth 2.25— 2.85 in SL, and compressed, the width 2.1— 2.8 in depth; head length 2.4—2.6 in SL; dor- sal profile of head slightly convex to straight except for rounded front of snout; snout length 4.0-4.65 in head; eye large; the orbit diameter 2.55-3.05 in head; interorbital space flat to slightly convex with median and lateral longitudinal ridges, the least width 4.05-4.35 in head; caudal peduncle depth 2.2—2.85 in head; caudal peduncle length 1.65-1.85 in head. Mouth large, the maxilla reaching to or slightly posterior to a vertical at rear edge of pupil, the upper jaw length 1.9-2.1 in head; mouth oblique, forming an angle of VOLUME 103, NUMBER 1 about 35° to horizontal axis of head and body; posterior margin of maxilla slightly rounded to slightly concave, the corners rounded; supramaxilla not present; lower jaw projecting, knob-like; upper jaw with a broad band of villiform teeth except for gap at symphysis, the teeth of the outer rows slightly incurved and inwardly depressible, the teeth of the inner rows at side of jaw very small; a narrow band of villiform teeth in lower Jaw, with three or four rows at front of jaw and a single row posteriorly, the inner teeth longer than outer; vomer with one to two rows of small teeth forming a V-shape with rounded apex; palatines with a single irregular row of small teeth. Tongue broadly triangular, the tip moderately pointed. Anterior nostril a short membranous tube located in front of center of orbit nearly half distance to anterior end of snout; posterior nostril ovate to slit-like, without a rim, lo- cated obliquely dorsoposterior to anterior nostril, the internarial distance about 1.5 times greater than distance from posterior nostril to orbit; posterior nostril usually only slightly larger than anterior; large openings, one in front of anterior nostril, one below internarial space and one at edge of inter- orbital space, leading to broad subsurface channels; a pair of prominent pores on tip of lower jaw; numerous small sensory pores over dorsal surface of head, side of snout, suborbital region, naked part of preopercle, and mandible. A single, poorly developed, flat, obtuse spine on opercle at level of upper edge of pupil; preopercular ridge smooth; posterior margin, rounded corner, and most of ven- tral margin of preopercle serrate (35 serrae on 33 mm specimen, 73 on 92 mm fish). Lateral line conspicuous, very slightly arched anteriorly, then paralleling dorsal contour of body and ending midlaterally a short distance onto base of caudal fin; scales ctenoid; head naked except for cheek, oper- cle, subopercle, and nape; no scales on dor- sal and anal fins except for a very low sheath of small scales on extreme base of second 47 dorsal and anal fins; basal part of caudal fin with progressively smaller scales that ex- tend about half distance to posterior mar- gin; paired fins naked except for triangular scaly process basally on lower surface of pel- vic fins. Origin of first dorsal fin above base of third lateral-line scale; first dorsal spine slender and short, about one-third length of second spine; second dorsal spine 3.6—4.3 in head; third dorsal spine longest (though only slightly longer than fourth), 1.65-—2.3 in head; first dorsal soft ray longest, 1.45— 1.6 in head; origin of anal fin below base of third dorsal soft ray; anal spine small, 3.4— 4.45 in length of second spine; second anal spine 2.35-2.85 in head; first anal soft ray longest, 1.55-1.75 in head; caudal fin forked, the caudal concavity 3.15-3.9 in head, the fin length 2.75-3.1 in SL; third or fourth pectoral rays longest, 1.4—1.5 in head; origin of pelvic fins slightly anterior to upper base of pectorals; pelvic fin tips extending well beyond anus, often to origin of anal fin, the first soft ray longest, 1.5-1.85 in head. Color in alcohol pale to light brown with a dark brown bar encircling posterior caudal peduncle, its midlateral width about one- half to two-thirds orbit diameter, its upper and lower edges usually expanded to pro- duce a slight hourglass shape; a dark brown stripe from front of snout, broadening as it passes to orbit, and continuing diffusely a short distance posterior to eye, its edges often darker brown; tip of snout dark brown; a narrow brown streak on maxilla and con- tinuing a short distance beyond it; lower opercle, subopercle, thorax, and lower ab- domen of some specimens partly silvery; tubular anterior nostril pale; a small dark brown spot on underside of each lateral-line scale, usually visible externally (more evi- dent on anterior than posterior scales); fins pale except anterior part of first dorsal fin, proximal upper and lower edges of caudal fin and lateral edge of pelvic fins which are dusky, and a dark brown line basally in anal fin, interrupted by each ray, which diverges 48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON NEW. ZEALAND a 160° ; 160° Fig. 1. Distributions of Apogon aureus (@) and A. fleurieu (®). outward from midbase of fin to end in a streak on last ray; peritoneum pale; diges- tive tract black; gillrakers dusky. Color in life coppery with iridescence, paler posteriorly, becoming golden on post- orbital head and side of body; a black bar across posterior caudal peduncle, broader dorsally and ventrally; a blackish stripe, edged in bright blue, from front of snout through eye, continuing diffusely a short distance behind eye where the blue margins tend to break up into dashes and spots; some individuals with a few indistinct bluish spots anteriorly on side of body; tip of lower jaw blackish; a bright blue line on upper lip, extending across maxilla, and continuing a short distance posterior to maxilla; lateral line pale with a series of small blackish spots, one per scale (spots progressively fainter, and often absent, posteriorly); median fins with pale orange-yellow membranes and salmon pink rays, the anterior part of the first dorsal fin dusky orange; distal part of lobes of second dorsal, anal, and caudal fins sometimes red; base of anal fin with a nar- row Orange band separated from an adjacent outer narrow blue band by a black line on membranes of fin; pectoral fins pale yellow- ish with light orange rays; pelvic fins with yellow membranes and orange rays, the lat- eral edge dusky bluish. Remarks. — As mentioned above, Apogon aureus 1s very similar to A. fleurieu; previ- ously these two taxa were considered syn- onymous. See the Remarks of the following account (of fleurieu) for discussion of their similarities and differences. Apogon aureus occurs from the coast of East Africa to the western Pacific where it ranges from southern Japan to Sydney, New South Wales; it is known from the following islands: Madagascar, Réunion, Mauritius, Seychelles, Sri Lanka, Christmas, Indone- sia, Philippines, Taiwan, New Guinea, Va- nuatu, and New Caledonia. With the excep- tion of the Tonga Islands where the senior author has observed and photographed the species underwater, it is not known from the islands of Micronesia and Polynesia. In spite of much collecting, it has not been taken at the atolls of the Maldives or Chagos Archipelago. It has not been observed or collected in the Red Sea or Persian Gulf and is as yet unknown from the coasts of India, Burma, and Thailand. It has, however, been recorded from the Gulf of Oman (as Apogon annularis) by Boulenger (1887); Boulenger’s specimen from Muscat was examined at the British Museum (Natural History). Collections of this species have been made in the depth range of 1-30 m from coral reefs or rocky substrata where there are caves VOLUME 103, NUMBER 1 49 Fig. 2. Adult of Apogon aureus, BPBM 30648, 92 mm SL, Port Moresby, Papua New Guinea. or crevices to provide shelter by day. The senior author photographed the species off northeastern Bali in 40 m. It does not nor- mally occur in areas subject to much wave action. The localities where collections have been made in only a few meters of water are all in well-protected bays or lagoons. Apogon aureus is generally encountered in small aggregations, sometimes mixed with A. apogonides (Bleeker). As noted by Ida & Moyer (1974), it is unusually bold for an apogonid and will venture a short distance from shelter to approach a diver if he re- mains still. It leaves the shelter of the reef shortly before sunset for nocturnal feeding, mainly on the larger zooplankton. In 1975 John E. McCosker and associates collected 39 specimens (CAS 35490) of a cardinalfish identified as Apogon aureus at Grande Comore, Comoro Islands, in caves in 20-30 m. These fish are immature and range in SL from 37-55 mm. We took rou- tine meristic data on all 39 specimens. The lower-limb gillraker count is high, 18-20 (see Table 1, under aureus). Since no other differences could be found between the Comoro fish and typical aureus, we provi- sionally identify them as A. aureus. It is hoped that future collecting at these islands will result in specimens of the adult of this form and a record of its life color. Material examined. —East Africa: RUSI 3205, 54 mm. Kenya: Mombasa, BMNH 1913.4.7.51, 92 mm. Zanzibar: BMNH 1867.3.7.555, 84 mm; RUSI 3167, 86 mm. Mozambique: Bazaruto Island, RUSI 3168, 80 mm. Pinda, RUSI 3164, 86.5 mm. Mo- zambique Island, RUSI 3165, 3: 64.5-87 mm. Natal: Durban, BMNH 1916.9.23.14, 81 mm; RUSI 11655, 8: 86-92 mm; SU 31278, 68 mm. Madagascar: MNHN 8762, 73 mm; USNM 17104, 3: 78-83 mm. Nossi Be, USNM 212408, 2: 62-79 mm. Mauri- 50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Juvenile of Apogon aureus, BPBM 23388, 32 mm SL, Taiwan. 1937.5.26.9, 98 mm; BMNH 1941.4.18.17, 97 mm; BPBM 20151, 5: 105-118 mm; MNHN 8759, 3: 66-81 mm; RUSI 1158, 3: 99-108 mm. Seychelles: Mahé, RUSI 3166, 71 mm. Gulf of Oman: Muscat, BMNH 1887.11.11.65, 99 mm; BMNH 1901.1.30.48, 64 mm. Sri Lanka: Trinco- malee, USNM 212422-24, 10: 61-88 mm; USNM 212587, 86.5 mm; MNHN 8700, 74.5 mm, holotype of Apogon roseipinnis. Sumatera (Sumatra): Weh, RMNH 12730, 4: 43-84 mm. Viet Nam: Nha Trang Bay, CAS 62567, 52: 74-95 mm; CAS 62568, 7: 90.5—98 mm. Jawa (Java): Jakarta, RMNH 12731, 86.5 mm. Java Sea, RMNH 12733, 77 mm. Sulawesi (Celebes): MNHN 60, 56.5 mm. Gulf of Tomini, Dodepo and Pasejogo Islands, USNM 171278, 2: 64-71 mm. Simba Strait, USNM 171290, 48.5 mm. Makassar Island, USNM 171294, 8: 24-51 mm. Buton Strait, Labuan Blanda Island, USNM 171292, 3: 35-46 mm. Selayar, RMNH 10020, 84 mm. Bali, BPBM 32247, 3: 45-48 mm. Lombok, BPBM 30047, 87 mm. Molucca Islands: Ambon, BPBM 19410, 5: 70.5-86 mm; MNHN 8699, 2: 79-80 mm; MNHN 8706, 73 mm; RMNH 8158, 5: 70-89 mm; USNM 210441, 26 mm. Buru (Bouro), USNM 171281, 4: 55-78 mm; Tomahu Island (near Buru), USNM_ 171283, 62-77 mm; USNM 212425, 72 mm. Seram (Ceram), BMNH 1858.4.21.175,180, 2: 70-81 mm. Bacan (Batjan), RMNH 13056, 76 mm. Gillolo Island, USNM 171280, 78 mm. Makyan Island, USNM 171291, 73 mm; USNM 171276, 7: 75-87 mm. Saparua Island, USNM 210076, 4: 75-81 mm; USNM 210357, 44: 22-81 mm. Tidore Island (south of Ternate), USNM 171282, 3: 80-83 mm; Dowarra Island, USNM 171290, 47 mm. Banda Islands: Banda Neira, USNM VOLUME 103, NUMBER 1 51 Fig. 4. Underwater photograph of Apogon aureus, about 95 mm TL, Bali. 213077, 49 mm. Western Australia: Ash- more Reef, AMS I.26742-024, 2: 32-33 mm. Northwest Cape, AMS I.19641-004, 2: 97- 97.5 mm. Dampier Archipelago, Rosemary Island, AMS I.19688-006, 2: 41-44.5 mm. Warroora, QM I.10263, 103 mm. Borneo: Sabah, Darvel Bay, Danawan and St. Amil Islands, USNM 171268, 2: 72.5-73 mm. Philippines: Sulu Archipelago, Singaan Is- land, USNM 171279, 3: 57-73 mm. Sima- luc Island and Simaluc Sibi Sibi Island, USNM 171275, 7: 41-80 mm. Jolo, SU 27367, 6: 92.5-107 mm. Tumindao Island, USNM 171277, 85 mm. Mindanao: Basilan Island, USNM 171288, 86 mm. Tonquil Island, USNM 171269, 2: 77-80 mm. Ma- lanipa Island, USNM 171273, 2: 64-81 mm. Tulnalutan Island, USNM 171270, 64 mm. Inamucan Bay, USNM 171274, 90 mm. Murcielagos Bay, USNM 171284, 3: 71-75 mm. Negros: SU 27353, 7: 50.5-98 mm. Dumaguete, BPBM 26513, 82 mm; BPBM 28579, 81 mm; SU 53318, 3: 53.5-93 mm. Mindoro: Galera Bay, USNM 171267, 2: 43-73 mm; USNM 171293, 3: 42-80 mm. Marinduque: Santa Cruz Island, USNM 171285, 2: 45-51 mm. Luzon: Batangas, USNM 171272, 6: 42-93 mm. Hong Kong: BMNH 1856.11.17.73, 67 mm. Taiwan: BPBM 23388, 32 mm. Papua New Guinea: southeast New Guinea, AMS 1.266, 95.5 mm. Port Moresby, AMS I.17537-003, 3: 88.5-92 mm; BPBM 15921, 94 mm (in- valid neotype of Apogon fleurieu, see Re- marks, under 4. fleurieu); BPBM 30648, 7: 85-92 mm. Madang, AMS I.17086-008, 2: 55-65 mm; BMNH 1974.5.25.1608-1609, 2: 52-63 mm; USNM 212426-27, 8: 30.5- 62 mm. Queensland: Great Barrier Reef, Palm Island, ANSP 122321, 80 mm. One Tree Island, AMS I.20206-006, 3: 53-60.5 mm. New South Wales: Sydney Harbor, 52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON AMS 1I.16777.001, 57 mm; AMS 1.17734- 001, AMS I.18850-004, 47.5 mm; ANSP 135507, 5: 52-61 mm. Apogon fleurieu (Lacepéde) Figs. 1, 5-7; Tables 1, 3 Ostorhinchus fleurieu Lacepéde, 1801:pl. 32, fig. 2; 1802:23 (type locality, Pacific Ocean). Apogon annularis var. roseipinnis (non Cu- vier) Gunther, 1859:239 (in part) (Am- bon). Apogon annularis (non Ruppell) Klunzin- ger, 1870:713 (Red Sea). Apogon aureus (non Lacepéde) Day, 1875: 61, pl. 16, fig. 8 (Madras, India).— Ran- dall, 1983:63, fig. 82 (Red Sea).— Allen & Steene, 1987:pl. 55, figs. 1, 2 (Similan Is- lands, Andaman Sea). Apogon (Amia) aureus (non Lacepéde) Klunzinger, 1884:22 (Quseir, Egypt).— Botros, 1971:296 (Red Sea). Ostorhynchus fleurieu Smith, 1961:399 (in part). —Kotthaus, 1970:62, figs. 245, 246, 250 (southern Red Sea). Apogon (Nectamia) fleurieu Dor, 1984:112 (Red Sea).—Kuronuma & Abe, 1986:99, pl. 10 (Persian Gulf). Apogon (Ostorhinchus) fleurieu Gon, 1987: 140 (an part). Apogon sp. Allen & Steene, 1987:pl. 35, fig. 5 (Phuket, Thailand). Diagnosis.—A species of the genus Apo- gon, subgenus Ostorhinchus, with dorsal rays VII-I,9, pectoral rays 14 (rarely 13 or 15); lateral-line scales 24; median predorsal scales 5; gillrakers 19-23; preopercular ridge smooth, the posterior margin and most of ventral margin serrate; body depth 2.4—2.85 in SL; interorbital space 4.05—4.6 in head; posterior nostril usually only slightly larger than anterior; pale (coppery with irides- cence in life), the young with a blackish spot midlaterally on posterior caudal peduncle which expands to a broad blackish bar in adults (dark bar not distinctly broader dor- sally and ventrally); a broad blackish stripe from front of snout to orbit and continuing diffusely behind eye (stripe bordered above and below by a blue line in life); a narrow brown streak (blue in life) on maxilla and a short distance posterior to it; a small black- ish spot on each lateral-line scale (may be faint or absent posteriorly); a line of dark brown dashes basally in anal fin. Maximum size about 105 mm SL. Description. — Dorsal rays VII-I,9, all rays branched, the last to base; anal rays IJ,8, all rays branched, the last to base; pectoral rays 14 (rarely 13 or 15); the upper two and lower two or three unbranched; pelvic rays I,5; principal caudal rays 17, the upper and low- er unbranched; upper and lower procurrent caudal rays 7-8, the posterior two seg- mented; lateral-line scales 24; scales above lateral line to origin of dorsal fin 2; scales below lateral line to origin of anal fin 6; median predorsal scales 5, the fourth and fifth deeply indented posteriorly in median line; circumpeduncular scales 12; gillrakers 5-6 (rarely 7)+15-16 (rarely 14 or 17), 1- 2 upper and O-1 lower as rudiments, the total count 19-23; pseudobranchial fila- ments increasing in number with size from 17 in 29 mm specimen to 29 in 102 mm specimen; branchiostegal rays 7; predorsal bones 3; vertebrae 10+ 14. Body moderately deep, the depth 2.4—2.85 in SL, and compressed, the width 2.15—2.5 in depth; head length 2.35—2.5 in SL; dorsal profile of head straight except for rounded front of snout; snout length 4.15-4.65 in head; eye large, the orbit diameter 2.55-3.1 in head; interorbital space flat to slightly convex with median and lateral longitudi- nal ridges, the width 4.05—4.6 in head; cau- dal peduncle depth 2.35-—2.7 in head; caudal peduncle length 1.7—2.0 in head. Mouth large, the maxilla reaching to or posterior to a vertical at rear edge of pupil, the upper jaw length 1.9—2.1 in head; mouth oblique, forming an angle of about 35° to horizontal axis of head and body; posterior edge of maxilla slightly rounded to slightly VOLUME 103, NUMBER 1 concave, the corners rounded; supramaxilla not present. Dentition, nostrils, pores, and scales es- sentially as described for Apogon aureus. A single, poorly developed, flat, opercular spine at level of upper edge of pupil, the dorsal and ventral margins of the spine forming an angle of 90° or more at the tip; preopercular ridge smooth; posterior mar- gin and most of ventral margin of preopercle serrate (28 serrae on 29 mm specimen and 94 on a 96 mm fish). Origin of first dorsal fin above base of third lateral-line scale; first dorsal spine slender and short, about one-third length of second spine; second dorsal spine 3.45—4.4 in head; third or fourth dorsal spine longest, 1.8—2.1 in head; first dorsal soft ray longest, 1.45-1.6 in head; origin of anal fin below base of third dorsal soft ray; first anal spine very small, 3.25-3.85 in length of second anal spine; second anal spine 2.45-2.75 in head; first anal soft ray longest, 1.6—1.75 in head; caudal fin forked, the caudal concav- ity 3.5—4.15 in head, the fin length 2.95-3.1 in SL; third or fourth pectoral rays longest, 1.45-1.6 in head; pelvic fin tips reaching posterior to anus but not beyond origin of anal fin, the first soft ray longest, 1.55-1.75 in head. Color of adults in alcohol light brown with a dark brown bar about three-fourths orbit diameter in width posteriorly on caudal pe- duncle, the upper and lower edges of bar not broader than central part; a dark brown stripe from front of snout, broadening as it passes to orbit, and continuing diffusely a short distance behind eye; a dark brown streak usually present on side of maxilla of adults which extends diagonally downward behind end of maxilla; tubular anterior nos- tril pale; a small dark brown spot on un- derside of each lateral-line scale, the spots progressively smaller and less pigmented posteriorly; edge of eye dark brown except ventrally; dorsal fins pale or slightly dusky anteriorly, particularly the first dorsal fin; anal fin pale, the leading edge sometimes 53 dusky, with a narrow dark brown line of dashes at base which diverges outward from midbase to end in a streak on last ray; caudal fin usually with some dusky pigment proxi- mally on upper and lower edges; paired fins pale except for a dusky lateral edge on pel- vics; peritoneum pale; digestive tract black; gillrakers dusky. Some specimens largely silvery over lower opercle, subopercle, tho- rax and ventral part of abdomen. Juveniles of about 30 mm SL are pale with a round diffuse dark brown spot mid- laterally on posterior part of caudal pedun- cle about the size of pupil or slightly larger; the dark stripe on the side of the snout is only faintly developed. With growth the pe- duncle spot gradually expands dorsally and ventrally to form the bar typical of adults (this attained at a SL of about 75 to 80 mm); the stripe on the head becomes more darkly pigmented, and the dark spots along the lat- eral line develop and become progressively darker. Color in life coppery with iridescence, be- coming golden on side of body and post- orbital part of head; peduncular spot of ju- veniles and bar of adults black; edges of blackish stripe on head bright blue, these margins often breaking into dashes or spots posterior to eye; occasional individuals with a few blue spots anteriorly on side of body; a bright blue line on side of maxilla and a short distance beyond; lower edge of eye orange; median fins pale salmon, more of this color on rays than membranes, the lead- ing edges of dorsal and anal fins and prox- imal upper and lower edges of caudal fin slightly dusky; an orange line at base of anal fin (broader posteriorly), separated from an outer adjacent blue line by a row of blackish dashes, one per membrane; pectoral fins pale salmon; pelvic fins with pale yellowish membranes and light orange rays, a narrow dusky leading edge and orange submarginal band. Remarks. —Apogon fleurieu shares many characters with A. aureus: essentially the same body and fin proportions (compare PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Adult of Apogon fleurieu, BPBM 19811, 96 mm SL, Gulf of Aqaba, Red Sea. Juvenile of Apogon fleurieu, BPBM 18258, 33 mm SL, Gulf of Aqaba, Red Sea. VOLUME 103, NUMBER 1 55 Fig. 7. Tables 2 and 3), meristic data except gill- raker counts, relatively large size, and such color features as the coppery-golden ground color, black peduncular bar in adults, blue- edged blackish stripe on head; blue line on maxilla, row of blackish spots along lateral line, row of dark dashes basally in the anal fin, dusky leading edges of dorsal, anal, and pelvic fins, and dusky proximal upper and lower edges of the caudal fin. The two species differ in gillraker counts (19-23 for fleurieu, compared to 22-27 for aureus) and the nature of the dark pedun- cular marking. Juveniles of aureus have a solid bar across the posterior part of the caudal peduncle (Fig. 3) in contrast to a dif- fuse dark spot for fleurieu (Fig. 6). The dark peduncular bar of aureus is sharply defined and usually expanded dorsally and ventrally to form a slight hourglass shape. The bar of adult fleurieu is not as well defined, is slight- ly broader laterally than that of aureus, and Underwater photograph of Apogon fleurieu, about 100 mm TL, Gulf of Aqaba, Red Sea. lacks the expanded upper and lower parts usually seen on aureus. There appears to be a difference in the maximum size attained by these two car- dinalfishes. A. aureus often exceeds 90 mm SL, the largest reported, 121 mm SL (RUSI 12345, from off Durban). Only 28 speci- mens of fleurieu of 423 examined exceed 90 mm SL, the largest, BPBM 31874, 102 mm SL, from the northern end of the Gulf of Aqaba. A specimen of 98 mm SL taken in the Persian Gulf was lost in shipment to the Bishop Museum, but a color photograph of it is on file at the Museum. Two specimens of 4. fleurieu from Hong Kong (SU 60820, 61-62 mm SL) and two from off Somalia (USNM 212415, both 63 mm SL) are fully mature females. Apogon fleurieu is known from the Red Sea, coast of East Africa south to Durban, Seychelles, Persian Gulf, India, Sri Lanka, Andaman Sea (Similan Islands and off Bur- 56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ma), Ambon, southern Malaysia, Hong Kong, and Philippines. Gerald R. Allen sent us an underwater photograph of this species from Flores, Indonesia. As mentioned, all of the collection sites except the Seychelles are on continental shelves (if the islands of Indonesia and the Philippines may be re- garded as on the Asian continental shelf), and the Seychelles are continental in origin. Six lots of A. fleurieu were collected by trawling from the R/V Anton Bruun, three off western India (17°41’N-20°N, 70° 71°33'E) in 71-97 m, two off Somalia in 25-— 31 m, and one in the Andaman Sea at 9°54'N, 97°42'E in 73 m. The shallowest collection of the species was made in 0-7 m off Kovalam, Kerala, southwestern India by the senior author and William F. Smith- Vaniz. The specimens of A. fleurieu from the Seychelles have a higher average number of gillrakers than other localities, and those from the Red Sea have slightly higher counts than other localities (Table 1). Seychelles specimens also attain larger size, in general, than those from other localities. All but five of the twenty-eight specimens which are longer than 90 mm SL were collected from these islands. Although there are many localities where Apogon fleurieu and A. aureus seem not to coexist, they do overlap in such localities as East Africa, Seychelles, Sri Lanka, Indo- nesia, Philippines, and Hong Kong. Both species were collected together at one sta- tion in Sri Lanka (USNM 212411 and 212424). The International Code of Zoological No- menclature, 3rd Edition, 1985, Article 75, Neotypes, p. 157, (b), Circumstances ad- mitted, states, ““A neotype is to be desig- nated only in connection with revisory work, but only in exceptional circumstances. .. .” Article 75, (b)(i), p. 157 states, ““The expres- sion ‘revisory work’ refers to a critical study of the nominal species-group taxon in ques- tion, regardless of the scope of the work in which it is published.’ Other statements of the ““Code”’ not satisfied by the account of Gon (1987) are listed on p. 159, (d), Qual- ifying conditions, ““A neotype is validly des- ignated only when it is published with the following particulars:”’ (2) “data and de- scription sufficient to ensure recognition of the specimen designated.” Material examined.—Red Sea: Gulf of Aqaba, BPBM 19811, 96 mm; BPBM 18258, 2: 28.5-33 mm; BPBM 31874, 102 mm; HUJ 11321, 62 mm; HUJ 11909, 2: 32.5-55.5 mm; HUJ 11912, 2: 32.5-34.5 mm; HUJ 11920, 2: 33-34 mm; HUJ 11922, 41:29.5-69 mm; MNHN 1988-689, 2: 38.5— 44 mm; NSMT-P.29514, 2: 38.5—43.5 mm; RUSI 3170, 9: 34.5—64.5 mm; RUSI 27672, 2: 37.5-44 mm; USNM 191657, 22: 49-87 mm; USNM 191704, 51: 30-50 mm; WAM P.29707-001, 2: 37-41 mm. Somalia: 11°14— 18’N, 51°8’E, USNM 212415, 28: 61.5-82 mm; USNM 212416, 13: 59-82 mm. Ke- nya: Mombasa, BPBM 27315, 63 mm. Zan- zibar: USNM 212405, 86 mm; USNM 212406, 81 mm; USNM 212407, 34: 23- 29 mm. Mozambique: Inhaca, RUSI 1733, 9: 29-77 mm; RUSI 1835, 2: 79-83 mm; RUSI 3169, 2: 67-95.5 mm. Delagoa Bay, ANSP 97470, 2: 37-43 mm. Natal: Sodwa- na Bay, RUSI 9207, 15: 29-46 mm. Mad- agascar: Nossi Be, USNM 212408, 2: 59- 76 mm. Seychelles: Mahé, ANSP 153774, 91.5 mm; ANSP 153775, 23: 41-101 mm; RUSI 3175, 51 mm; USNM 212409, 75: 34-100 mm. Cosmoledo Group, As- sumption Island, RUSI 3176, 44.5 mm. In- dia: off Bombay, USNM 212418, 74 mm; USNM 212419, 3: 77-80.5 mm; USNM 212420, 9: 65.5—75 mm. Kerala, Kovalam, BPBM 27638, 5: 60.5-65.5 mm. Wadge Banks, USNM 212410, 79 mm. Sri Lanka: Trincomalee, USNM 212411-14, 15: 21- 77 mm. Andaman Sea: off southern end of Burma, USNM 212421, 10: 52.5-67 mm. Indonesia: Ambon, BMNH 1855.3.24.41, 92 mm. Malaysia: Johore, SU 30409, 90.5 mm. Hong Kong: 22°18’N, 114°23’E, SU 60820, 3: 61-63.5 mm. Philippines: Jolo Island, USNM 171287, 10: 76-89 mm; VOLUME 103, NUMBER 1 USNM 171289, 73 mm. Masbate, Catain- gan Bay, USNM 212417, 9: 19-26 mm. Apogon pselion, new species Figs. 8, 9; Tables 1, 4 Holotype. —BPBM 21515, male, 36.2 mm, Red Sea, Gulf of Aqaba, Sinai Pen- insula, El Himeira, coral knoll, 12 m, ro- tenone, J. E. Randall and O. Gon, 25 Apr (OWT. Paratypes.—TAU 9670, 26.5 mm, Red Sea, Gulf of Suez, Abu Zneiman, rotenone, L. Fishelson, 22 Sep 1967; TAU 9669, 2: 29.0-—30.5 mm, Gulf of Aqaba, Eilat, rote- none, D. Popper, 22 Jan 1969; USNM 213381, 36.0 mm, Gulf of Aqaba, Sinai Peninsula, El Himeira, 0-18 m, rotenone, V. G. Springer et al., 16 Jul 1969; USNM 213382, 2: 33.5-35.5 mm, El Himeira, 8—- 16 m, V. G. Springer et al., 19 Jul 1969; USNM 213383, 7: 39.3-41.3 mm, Sinai Peninsula, east coast, Ras Burqa, 9-15.5 m, rotenone, V. G. Springer et al., 21 Jul 1969; USNM 213385, 38.0 mm, just north of Ras Burqa, to 11 m, rotenone, V. G. Springer et al., 23 Jul 1969; USNM 213387, 11: 20.6- 36.8 mm, El Himeira, 21.5—27.5 m, V. G. Springer et al., 9 Sep 1969; CAS 60679, 2: 24.6—35.7 mm, NSMT-P..44623-24, 2: 24.5- 35.1 mm, RUSI 27056, 2: 22.0-34.4 mm, WAM P.29386-001, 2: 25.2-35.0 mm—all with same data as USNM 213387; BPBM 13381, 32.7 mm, Gulf of Aqaba, Eilat, off marine biological laboratory, reef in 43 m, rotenone, J. E. Randall and D. Popper, 6 Jun 1972; BPBM 18261, 6: 23.2-36.9 mm, Sinai Peninsula, east coast, Coral Island, east side, 30 m, quinaldine, J. E. Randall and O. Gon, 24 Sep 1974; BPBM 31988, 6: 33.8- 39.6 mm, MNHN 1977-827, 5: 32.5-37.5 mm-—both lots with same data as holotype. Diagnosis.—A species of the genus Apo- gon, subgenus Ostorhinchus, with dorsal rays VII-I,9; pectoral rays 13-14 (usually 14); lateral-line scales 24; median predorsal scales 4; gillrakers 19-22; preopercular ridge smooth, the posterior margin and rear half 57 of ventral margin of preopercle serrate; body depth 2.8-3.4 in SL; interorbital space 4.55- 5.4 in head; posterior nostril more than twice as large as anterior; four dusky golden yel- low stripes on head separated by blue lines, the midlateral stripe passing from snout, through eye, and continuing along side of body as a brassy yellow stripe; a black bar encircling posterior caudal peduncle. Description. — Dorsal rays VII-I,9, all rays branched, the last to base; anal rays II,8, all rays branched, the last to base; pectoral rays 14(13-14), the upper two and lower three unbranched; pelvic rays I,5; principal cau- dal rays 17, the upper and lower unbranched; upper and lower procurrent caudal rays 7, the posterior two segmented; lateral-line scales 24 (plus 3 scales posterior to end of hypural plate); scales above lateral line to origin of first dorsal fin 2; scales below lat- eral line to origin of anal fin 5'2; median predorsal scales 4, the fourth scale deeply indented medially on the posterior edge and the third scale slightly indented posteriorly; circumpeduncular scales 12; gillrakers 6+ 15 (5-6+ 14-16), 2-3 upper and O-1 lower as rudiments; pseudobranchial filaments 15 (10-16); branchiostegal rays 7; predorsal bones 3; vertebrae 10+ 14. Body moderately elongate, the depth 2.95 (2.8-3.4) in SL, and compressed, the width 2.35 (1.9-2.5) in depth; head length 2.55 (2.45—2.6) in SL; dorsal profile of head straight except for rounded front of snout; snout length 3.85 (3.95—4.3) in head; eye large, the orbit diameter 2.8 (2.5-3.0) in head; interorbital space flat to slightly con- vex, the width 4.9 (4.55—5.4) in head; caudal peduncle about twice as long as deep, the least depth 2.6 (2.5-3.3) in head. Mouth large, the maxilla reaching to or slightly posterior to a vertical through rear edge of pupil, the upper jaw length 1.95 (1.9- 2.05) in head; mouth oblique, forming an angle of about 35° to horizontal axis of head and body; posterior edge of maxilla slightly concave; no supramaxilla present; lower jaw projecting, the tip dorsoventrally thickened; 58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. Holotype of Apogon pselion, BPBM 21515, 36.2 mm SL, Gulf of Aqaba, Red Sea. villiform teeth in a narrow band in jaws, with two rows at front of upper jaw (except for a gap at symphysis) and about five rows at side of jaw (teeth of inner rows on side of jaw very small); front of lower jaw with about three rows of teeth narrowing to one at side of jaw; a single irregular row of very small teeth forming a V on vomer; a single row of very small teeth on palatines; tongue broad at base tapering to lanceolate form with expanded rounded tip. Anterior nostril a short membranous tube directly anterior to center of eye half the distance to front of snout; posterior nostril ovate without a raised rim, its largest di- ameter two or more times greater than di- ameter of anterior nostril, located dorso- posterior to anterior nostril closer to edge of orbit than anterior nostril; numerous small pores readily visible dorsally on head, scattered over surface of preopercle and along its anterior margin, on suborbital rim, and mandible. A single, poorly developed, flat opercular spine, its tip forming an angle slightly great- er than 90°; posterior margin of preopercle serrate (33 serrae on holotype), the serrae smaller on upper margin than the broadly rounded corner; lower margin of preopercle serrate about half distance from middle of rounded corner to end of free edge; pre- opercular ridge not serrate. Lateral line conspicuous, very slightly arched anteriorly, then paralleling dorsal contour of body and ending midlaterally a short distance posterior to caudal-fin base (three pored scales posterior to end of hy- pural); scales weakly ctenoid; head naked except operculum and nape; no scales on fins except for a low sheath at extreme base of dorsal and anal fins and small scales on base of caudal fin which extend at most half distance to posterior margin. Origin of first dorsal fin above base of third lateral-line scale; first dorsal spine slender and short, about one-fourth length of second spine, 11.9 (9.8-14.5) in head; third dorsal spine longest, 2.1 (1.9—2.15) in VOLUME 103, NUMBER 1 59 Fig. 9. Underwater photograph of Apogon pselion, about 40 mm TL, Eilat, Gulf of Aqaba, Red Sea. head; first dorsal soft ray longest, 1.6 (1.5— 1.65) in head; origin of anal fin below base of second dorsal soft ray; first anal spine small, about one-third length of second anal spine, 8.35 (7.3-10.7) in head; second anal spine 3.0 (2.45—2.95) in head; first anal soft ray longest, 1.9 (1.7—1.9) in head; caudal fin forked, its length 3.2 (3.1-3.6) in SL, the caudal concavity 4.7 (4.1—4.9) in head; third and fourth pectoral rays longest, 1.55 (1.5— 1.75) in head; first and second pelvic rays longest, 1.75 (1.65—1.85) in head. Color of holotype in alcohol: pale with a black bar encircling posterior caudal pe- duncle and extending slightly onto caudal- fin base, this bar widest on side of peduncle (bar width about half orbit diameter); a very diffuse and faint dusky midlateral stripe on body; a narrow dusky stripe from front of upper lip to middle of front edge of orbit; a narrow dusky stripe from front of lower jaw to lower edge of eye and merging with broad dusky area on side of postorbital head; fins pale except for a dusky stripe near base of second dorsal and anal fins; peritoneum pale to slightly dusky; digestive tract black. Color of holotype when fresh: body pale pinkish gray, overlaid with iridescent bluish silver ventrally, with a midlateral brassy yellow stripe faintly edged in pale iridescent blue, and a black bar encircling posterior caudal peduncle and adjacent caudal-fin base; head with four dusky golden yellow stripes separated by pale blue lines; median fins transparent bluish, the rays edged in pale salmon, the second dorsal and anal fins with a pale blue-edged dusky light orange band near base; paired fins transparent with pale salmon rays. Remarks. — This species is given the spe- cific name pselion from the Greek noun for bracelet or anklet, in reference to its most distinctive color marking, the ring of black around the posterior caudal peduncle. 60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Apogon pselion coexists in the northern Red Sea with the related A. fleurieu. The former is a much smaller species (the largest specimen 41.3 mm SL). At this size A. fleu- rieu has a diffuse dark spot posteriorly on the side of the peduncle whereas 4. pselion at all sizes represented by our material has a solid black bar. A. pselion could be con- fused by its color pattern in preservative with the young of A. aureus (though the lat- ter is not known from the Red Sea), which already has a completely formed black pe- duncular bar at sizes as small as 33 mm SL. Most specimens of these two species can be distinguished by gillraker counts (see Table 1). Apogon pselion may be differentiated from both aureus and fleurieu by its more elon- gate body (the depth 2.8-3.4 in SL, com- pared to 2.25—2.85 for the other two species), narrower interorbital space (4.55-5.4 in head, compared to 4.05-4.6 for the other two species), shorter fins, in general (com- pare Tables 2-4), three instead of four lat- eral-line scales posterior to end of hypural plate, and life color. A. aureus and A. fleu- rieu lack the four yellow stripes on the head and lateral yellow stripe on the body, and A. pselion lacks the series of small dark spots along the lateral line. Apogon pselion is at present known only from the Gulfs of Aqaba and Suez; it may be confined to the northern Red Sea where the water temperature is distinctly cooler than the central and southern part of the Sea. No specimens have been collected in rotenone stations from the Red Sea coasts of Sudan, Ethiopia, or Saudi Arabia. The type material has been obtained from reefs in the depth range of about 10 to 43 m. Acknowledgments We thank the following persons for the loan of specimens and/or pertinent infor- mation for the present study: Gerald R. Al- len, M. Eric Anderson, Marie Louise Bau- chot, David Catania, Martine Desoutter, Lev Fishelson, Anthony C. Gill, Daniel Go- lani, Ofer Gon, Menachem Goren, Doug- lass F. Hoese, Susan L. Jewett, Alwyne C. Wheeler, and Jeffrey T. Williams. Richard L. 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Ten new records of 62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cardinalfishes from Taiwan, with a synopsis of the Family Apogonidae.—Journal of the Tai- wan Museum 39(2):61-104. Shen, S. C. 1984. Coastal fishes of Taiwan. Shih- chieh Shen, private publisher, Taipei, 190 pp. —., & C. Lam. 1977. A review of the cardinal- fishes (Family Apogonidae) from Taiwan.— Acta Oceanographica Taiwanica Science Reports of the National Taiwan University 7:154-192. Smith, J. L. B. 1949. The sea fishes of Southern Af- rica. Central News Agency Ltd., Cape Town, xvi + 550 pp. . 1961. Fishes of the family Apogonidae of the Western Indian Ocean and the Red Sea.—Ich- thyological Bulletin, Department of Ichthyolo- gy, Rhodes University 22:374-418. —, & M. M. Smith. 1963. The fishes of Sey- chelles. The Department of Ichthyology, Rhodes University, Grahamstown, 215 pp. Smith, M. M., & P. C. Heemstra, eds. 1986. Smiths’ sea fishes. 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WASH. 103(1), 1990, pp. 63-85 LIVING CASSIDULOIDS (ECHINODERMATA: ECHINOIDEA): A KEY AND ANNOTATED LIST Rich Mooi Abstract.—A key to the 30 known living species of cassiduloids is provided which includes abbreviated geographic and bathymetric data. All species are illustrated for the first time in a single work. An annotated list of genera and species includes basic taxonomic information, remarks upon systematic affin- ities, more detailed discussions of geographic and bathymetric ranges, and summaries of general biology and habitat preferences. The order Cassiduloida (sensu Kier 1962) consists of some 800 species of irregular echinoids, most of which are known only as fossils. Kier (1962:1) characterized this order as comprising “‘all those ‘irregular’ echinoids having petals, phyllodes, and bourrelets (the floscelle).”” However, this general description also applies to such forms as clypeasteroids, which have not only well- developed petaloids, but also what can be interpreted as floscelles consisting of inter- ambulacral bourrelets and reduced phyl- lodes with buccal and large food groove po- dia (sensu Mooi 1986). This, along with great variation in the major features used to char- acterize the families within the order Cas- siduloida (for example, the periproct varies in position from aboral and in a deep groove to submarginal), renders it difficult to un- ambiguously define the order Cassiduloida without using features common to other major irregular echinoid groups. Lack of unique unifying characters leads to suspi- cions that the group is not monophyletic. I am at present preparing a phylogenetic re- vision of the living cassiduloid species that should shed light on this problem. In spite of the possibility that the Cassiduloida as defined by Kier (1962) may not be a natural group, it remains convenient to refer to this assemblage as the “‘cassiduloids.’’ For the purposes of this paper, this group includes all those shallow burrowing irregular echi- noids with a relatively high test, short spines, a posteriorly placed periproct, and well-de- veloped floscelle. Although abundant as Mesozoic and Ce- nozoic fossils, the cassiduloids have dra- matically decreased in number since the Eocene (Kier 1974, Suter 1988). Studies of the pattern of decline of the cassiduloids (Suter 1988) and the evolution of other ma- jor irregular echinoid groups (Mooi 1987) have made desirable a review of the distin- guishing features of, and basic biological in- formation on, the Recent taxa of the Cas- siduloida. In his monograph of the echinoids, Mortensen (1948) listed 27 Re- cent species of cassiduloids, including the 5 species in the Neolampadidae which he thought constituted a separate family within the order. Since Mortensen’s work, three new species of cassiduloids have been de- scribed (Krau 1954, Baker 1983, Mooi 1990), and additional morphological and biogeographic information has become available, particularly for the poorly known neolampadids (McKnight 1968). Kier (1962) never mentioned the Neolampadidae in his revision of the Cassiduloida and did not offer a reason for their exclusion. Although Philip (1963) raised the family Neolampa- didae to subordinal rank (Neolampadina) within the Cassiduloida, only its familial 64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON standing is recognized here, pending phy- logenetic revision of the living taxa in the order. The most recent key to the cassiduloids is that of Mortensen (1948). In light of in- formation accumulated over the past 40 years, a new, illustrated key to the living species of the group is needed. The key and annotated list include the neolampadids and the cassiduloid genus Oligopodia, which Kier (1962) also omitted from his revision. For the first time in a single work, all the known Recent species are figured. I have included brief comments on taxonomy, affinities, and geographic and bathymetric ranges as well as a summary of what has become known of the biology of some of the species since Mortensen (1948). List of Recognized Taxa Phylum Echinodermata Bruguiére, 1789 Subphylum Echinozoa Haeckel in Zittel, 1895 Class Echinoidea Leske, 1778 Subclass Euechinoidea Bronn, 1860 Infraclass Acroechinoidea A. B. Smith, 1981 Cohort Irregularia Latreille, 1825 Superorder Microstomata A. B. Smith, 1984 Series Neognathostomata A. B. Smith, 1981 Order Cassiduloida Claus, 1880 Family Echinolampadidae Gray, 1851 Genus Echinolampas Gray, 1825 Echinolampas ovata (Leske, 1778) Echinolampas alexandri de Loriol, 1876 Echinolampas chuni (Déderlein, 1905) Echinolampas crassa (Bell, 1880) Echinolampas depressa Gray, 1851 Echinolampas keiensis (Mortensen, 1948) Echinolampas koreana H. L. Clark, 1925 Echinolampas rangii Desmoulins, 1837 Echinolampas sternopetala A. Agassiz & H. L. Clark, 1907 Echinolampas sumatrana (Doderlein, 1905) Genus Conolampas (A. Agassiz, 1883) Conolampas sigsbei (Agassiz, 1878) Conolampas diomedea Mortensen, 1948 Conolampas malayana Mortensen, 1948 Conolampas murrayana Mortensen, 1948 Family Cassidulidae L. Agassiz & Desor, 1847 Genus Cassidulus Lamarck, 1801 Cassidulus caribaearum Lamarck, 1801 Cassidulus infidus Mortensen, 1948 Cassidulus mitis Krau, 1954 Cassidulus malayanus (Mortensen, 1948) Genus Eurhodia Haime in d’Archiac & Haime, 1853 Eurhodia relicta Mooi, 1990 Genus Oligopodia Duncan, 1889 Oligopodia epigonus (van Martens, 1865) VOLUME 103, NUMBER 1 Genus Rhyncholampas A. Agassiz, 1869 Rhyncholampas pacificus (A. Agassiz, 1863) Genus Studeria Duncan, 1891 Studeria recens (A. Agassiz, 1879) Family Apatopygidae Kier, 1962 Genus Apatopygus Hawkins, 1920 Apatopygus recens (Milne Edwards, 1863) Apatopygus occidentalis H. L. Clark, 1938 Genus Porterpygus Baker, 1983 Porterpygus kieri Baker, 1983 Family Neolampadidae Lambert, 1918 Genus Neolampas A. Agassiz, 1869 Neolampas rostellata A. Agassiz, 1869 Genus Anochanus Grube, 1868 Anochanus sinensis Grube, 1868 Genus Aphanopora de Meijere, 1902 Aphanopora echinobrissoides de Meijere, 1902 Genus Nannolampas Mortensen, 1948 Nannolampas tenera (de Meijere, 1902) Genus Tropholampas H. L. Clark, 1923 Tropholampas loveni (Studer, 1880) Key to the Living Cassiduloids The following key relies on morphology of the test and external appendages, partic- ular aspects of which are illustrated in Figs. 1-5. The species themselves are illustrated at the end of the key, in Figs. 6-12 (see Annotated List). When a species is encoun- tered within the key, an abbreviated de- scription of its range is given in square brackets, followed by a statement of its known bathymetric range. Nomenclatural authorities and other taxonomic informa- tion can be found in the annotated list of species that follows the key. Anatomical ter- minology is that of Mooi (1989). 1. Petaloids strongly developed (Figs. 6-11); respiratory podia with well-developed inner and outer pores; phyllodes conspicuous; bourrelets moderately to strongly developed ae epee ae y, — Petaloids completely absent (Fig. 12); when present, aboral podia are single-pored, and not developed into respiratory podia; phyllodes inconspicuous, usually lacking in- ner series of phyllopores; bourre- lets poorly developed or lacking . Apical system monobasal, without separate genital plates (Fig. 1a); na- ked zone present medially on oral surface (except in Studeria recens [Fig. 10d]), although not always strongly developed (Figs. 6-10); anal sulcus absent, or typically (ex- cept in Cassidulus malayanus [Fig. 9c]) short and shallow (Figs. 6-10); buccal podia present; globiferous pedicellariae absent ............ Apical system tetrabasal, with sep- arate genital plates (Fig. 1b); naked zone absent (Fig. 11); anal sulcus long and very deep (Fig. 11); buc- cal podia absent; globiferous ped- icellamnaciprescnite aan ee . Periproct just submarginal (Figs. 6—8); anal sulcus absent; three large, triangular plates along adoral edge of periproctal membrane (Fig. 2a); 65 26 24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ophicephalous pedicellariae with distal, closed oval ring of gripping teeth (Fig. 3a); many calcite spic- ules in stem of non-respiratory po- Giad kee Seek tee i ee Periproct aboral or marginal (Figs. 9, 10); anal sulcus present, often weakly developed; more than four or five large plates in adoral part of periproctal membrane (Fig. 2b); ophicephalous pedicellariae with distal, U-shaped row of distal teeth (Fig. 3b); no spicules in stem of non-respiratory podia, but some spicules may occur in sucker tip of podium : . Test margin oval in outline with periproct on slight posterior pro- jection, or rostrum (Figs. 6, 7); oral surface concave (Figs. 6, 7); peri- stome slightly displaced anterior- ly; primary aboral spination rela- tively dense, average distance between spine tubercles less than 1.5 times primary spine tubercle diameter (Echinolampas) ....... Test margin almost circular in out- line, rostrum very short, or absent (Fig. 8); oral surface flat (Fig. 8); peristome central, or slightly dis- placed posteriorly; primary aboral spination relatively sparse, aver- age distance between spine tuber- cles more than twice primary spine tubercle diameter (Conolampas) . . Columns of respiratory podia in each petaloid very unequal in length, shorter column less than 60% length of longer in petaloid of ambulacra I and V (Figs. 6e, 7a, b, (GD tented eee awe ee Rahs aitan SA Columns of respiratory podia in each petaloid not very unequal in length, shorter column more than 65% length of longer in petaloid of ambulacra I and V (Figs. 6a—d, 7c, es ED ae ae eee 6. Peristome oval, bourrelets not 1Y/ 14 projecting into peristome; peri- stome width more than 17% test width Peristome pentagonal, bourrelets slightly projecting into peristome; peristome width less than 15% test width . Test width greater than 85% test length; ambitus broadly rounded in cross section; oral surface tumid [northeastern Central America, Yucatan, southeastern U.S., Greater and Lesser Antilles to northeastern South America; 30- 33 Oma ise. . 2 5 2 eee hteont Echinolampas depressa (Fig. Test width less than 85% test length; ambitus relatively sharp in cross section; oral surface some- what flattened [Japan; 150-500 m] . Echinolampas sternopetala (Fig. . Apical system more than 40% test length away from anterior edge of test; test almost conical; ambitus relatively sharp in cross section; oral surface flattened [Kepulauan Kai, southern Philippines; 245- AQO MM) ocdceess oko 3 ee ent Ic Echinolampas keiensis (Fig. Apical system less than 35% test length away from anterior edge of test; test smoothly arched in side view; ambitus broadly rounded in cross section; oral surface tumid [Korean Strait; 73 m] ce Echinolampas koreana (Fig. . Petaloids wide, eight or more pri- mary spine tubercles in row across interporiferous zone half-way down petaloid of ambulacrum V; peristome pentagonal, bourrelets slightly projecting into peristome Petaloids narrow, fewer than eight primary spine tubercles in row across interporiferous zone half- way down petaloid of ambulacrum V: peristome oval, bourrelets not projecting into peristome ....... 6e) 7d) 7a) 7b) VOLUME 103, NUMBER 1 10. Test smoothly arched antero-pos- 11. 1? teriorly; ambitus broadly rounded in cross section; oral surface tu- mid; tuberculation of aboral sur- face fine, more than 100 primary spine tubercles in 25 mm7?; outer pore of each respiratory podial pore pair circular or subcircular [Red Sea, northern Indian Ocean to western Australia; 9-75 m] Test with relatively sharp ambitus; oral surface somewhat flattened; tuberculation of aboral surface coarse, fewer than 100 primary spine tubercules in 25 mm7?; outer pore of each respiratory podial pore pair strongly elongate Test high, height greater than 50% test length; tuberculation of aboral surface very coarse, fewer than 50 primary spine tubercules in 25 mm? [South Africa; 25-500 m] Test low, height less than 50% test length; tuberculation of aboral sur- face not very coarse, more than 70 primary spine tubercles in 25 mm? [west coast of Africa, Cape Verde Islands; shallow to 1670 m] ed Echinolampas rangii (Fig. Apical system almost central, more than 45% test length from anterior edge of test; width of periproct less than 15% test width; more than five primary spine tubercles in row across interporiferous zone half- way down petaloid ofambulacrum V [Sumatra; 371 m] .. Echinolampas sumatrana (Fig. Apical system slightly anterior, less than 45% test length from anterior edge of test; width of periproct more than 16% test width; five or fewer primary spine tubercles in row across interporiferous zone half-way down petaloid of ambu- lacrum V Echinolampas ovata (Fig. Echinolampas crassa (Fig. 6a) 6d) 7c) Te) 3s 14. 1S): 16. Test relatively wide, width greater than 85% test length; phyllodes widened, interrupting taper of am- bulacra as they approach peri- stome [Red Sea, Indian Ocean, Malaysia; 8-365 m] ... Echinolampas alexandri (Fig. Test relatively narrow, width less than 85% test length; phyllodes narrow, ambulacra _ tapering smoothly as they approach peri- stome [Sumatra; 371 m] Spine tubercles on oral surface sparsely distributed, fewer than seven in 25 mm? in interambula- crum 5 next to peristome; outer pore of each respiratory pore pair round [West Indies; 120-800 m] PR atom Conolampas sigsbei (Fig. Spine tubercles on oral surface rel- atively densely distributed, 10 or more in 25 mm? in interambula- crum 5 next to peristome; outer pore of each respiratory pore pair elongate Peristome approximately central; periproct approximately half its own length away from ambitus; in- terporiferous zone of petaloids rel- atively wide, more than twice as wide as single column of respira- tory podia as measured at a point half-way down anterior petaloid [Maldives; 230 m] ... Conolampas murrayana (Fig. Peristome slightly posterior; peri- proct almost as far from ambitus as it is long; interporiferous zone of petaloids relatively narrow, less than twice as wide as a single col- umn of respiratory podia as mea- sured at a point half-way down an- terior petaloid Longest column of respiratory po- dia in each petaloid reaches am- bitus when viewed from above Echinolampas chuni (Fig. 67 6b) 6c) 8a) 15 8d) 16 68 19% 20) De PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON [Philippines; 265 m] .... Conolampas diomedeae (Fig. 8b) Longest column of respiratory po- dia in each petaloid ends well short of ambitus when viewed from above [eastern Indonesia, south- ern Philippines; 245-400 m] .... Scpsirdy 2 98 Conolampas malayana (Fig. 8c) Peristome longer than wide ..... 18 Peristome circular, or wider than LOM Pe wae errecu cates Aenea a hat, 19 . Periproct slightly longer than wide [Indian Ocean, Malaysia; 35-140 m] ... Oligopodia epigonus (Fig. 10b) Periproct wider than long [north- ern South America; 57-112 m] We SR alae Eurhodia relicta (Fig. 10a) Naked zone absent from medial area of oral surface; three gono- pores; columns of respiratory po- dia in each petaloid equal in length [Arafura Sea; 236 m] A See Studeria recens (Fig. 10d) Large naked zone in medial area of oral surface; four gonopores; columns of respiratory podia in each petaloid unequal in length, with columns Ia, IIb, IIIb, [Va, and Vb the shortest of each pair Large forms, test of adults over 40 mm in length; more than 70 res- piratory podia in anterior petaloid of adults; more than 20 podial pores in anterior phyllode [Gulf of California to west coast of Pana- ma, Galapagos; 2-130 m] .. Rhyncholampas pacificus (Fig. 10c) Relatively small forms, test of adults less than 30 mm in length; fewer than 60 pore pairs in anterior petaloid of adults; fewer than 15 podial pores in anterior phyllode (CaSSIdULUS ess) ae eer Periproct longer than wide, located aborally, about half the distance from the apical system to the pos- terior edge; anal sulcus long, nar- row, without conspicuous hood 20 74 MD De 24. De over periproct [Indonesia; 250-290 m] ...Cassidulus malayanus (Fig. Periproct round, or wider than long, located aborally, consider- ably more than half the distance from the apical system to the pos- terior edge; anal sulcus short, wide, with small but distinct hood over periproct Test height greater than 50% test length; periproct subcircular; few- er than 45 respiratory podia in an- terior petaloid of adults [East coast of South America?; shallow water, precise locality and depth data un- available] Cassidulus infidus (Fig. Test height less than 50% test length; periproct distinctly wider than long; more than 45 respira- tory podia in anterior petaloid of adults Apical system less than 40% test length from anterior edge; test rel- atively wide, more than 80% test length [Belize, Bahamas to Bar- bados; very shallow, 2-10 m] ... .... Cassidulus caribaearum (Fig. Apical system approximately 40% test length from anterior edge; test relatively narrow, less than 80% test length [Sepetiba Bay, Brazil; shallow water, precise depth data unavailable] vaste hee atanae Cassidulus mitis (Fig. Three gonopores; petaloids slight- ly reduced, fewer than 16 pore pairs in ambulacrum III of large adults; peristome roughly pentagonal [Three Kings Islands, New Zea- land; 90-300 m] Four gonopores; at least 17 pore pairs in ambulacrum III of large adults; peristome transversely oval, or roughly triangular (Apatopygus) Test large, adults commonly reaching 40 mm in length; more 9c) 2D) 9b) 28) 9a) 9d) Porterpygus kieri (Fig. 11c) De) VOLUME 103, NUMBER 1 26. Quik 2S 29. than 20 hydropores in madreporic plate; valves of globiferous pedi- cellariae with short blade and a single pair of long, distal, fang-like teeth (Fig. 4a) [New Zealand; 5- 145 m] . Apatopygus recens (Fig. 11a) Test small, adults usually less than 20 mm in length; fewer than 15 hydropores in madreporic plate; valves of globiferous pedicellariae with long, narrow blade and two pairs of long, distal, fang-like teeth (Fig. 4b) [southern and southwest- ern Australia; 17—40 m] ... Apatopygus occidentalis (Fig. 11b) Periproct on ambitus, or slightly submarginal, not in deep anal sul- cus ai} cus 29 Apical system sunken to form ab- oral ““marsupium” in females; po- dial pores entirely absent from ab- oral ambulacra; four gonopores [South Africa; 135-350 m] Reh: Tropholampas loveni (Fig. 12e) Apical system not sunken in fe- males; podial pores present in ab- oral ambulacra; two or three gono- pores (four in rare individuals) .. 28 Apical system monobasal, without separate genital plates; typically three gonopores (rarely four, never two); valves of ophicephalous ped- icellariae lacking thorns on proxi- mal “‘handle” (Fig. 5a) [West In- dies, Mediterranean; 145-1260 m] ks Neolampas rostellata (Fig. 12a) Apical system tetrabasal, with sep- arate genital plates; two gono- pores; valves of ophicephalous pedicellariae with prominent thorns on proximal part of “han- dle”’ on largest valve (Fig. 5b) [East Indies, Timor Sea; 390 m] Nannolampas tenera (Fig. 12d) Apical system sunken to form ab- oral ““marsupium”’ [China Sea; 69 depthiunknowmnl|es ae sae se. ote Anochanus sinensis (Fig. 12b) — Apical system not sunken [East In- dies, northern and northeastern Australia; 350-710 m] Bi ie hs Aphanopora echinobrissoides (Fig. 12c) Annotated List of Cassiduloid Genera and Species This list of genera is arranged by family according to Kier & Lawson (1978), which is in turn largely based on Kier (1962). I depart from their arrangement only in not recognizing the family Pliolampadidae Kier, 1962, pending phylogenetic revision of that group to resolve systematic problems that Kier (1962) himself acknowledged. As a re- sult, Eurhodia (which Kier placed in the Pliolampadidae) is here placed in the Cas- sidulidae, following Mooi (1990). Studeria is also provisionally recognized as a cassi- dulid. Oligopodia, which Kier & Lawson (1978:125) listed among the “‘doubtful nominal genera,”’ is here considered to be a cassidulid as well, as its similarity to some members of the genus Eurhodia suggests. Species described after 1948 are placed in the genera and families to which they were assigned by the original authors. Generic synonymies can be found in Kier (1962), and complete species descriptions and syn- onymies in Mortensen (1948), Krau (1954), Baker (1983), and Mooi (1990). The type species for each genus is indicated by an asterisk (*). Within families, genera are list- ed alphabetically after the type genus, and species are listed alphabetically within gen- era. Illustrations of these taxa (Figs. 6-12) follow this same arrangement. Order Cassiduloida Claus, 1880 Family Echinolampadidae Gray, 1851 Echinolampas Gray, 1825 Echinolampas ovata (Leske, 1778)* Fig. 6a Geographic range.— A widespread species occurring in the Red Sea and the tropical 70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Apical systems of cassiduloids: a, Mono- basal system of Cassidulus caribaearum Lamarck, 1801 [Anegada, British Virgin Islands]; b, Tetrabasal system of Apatopygus recens (Milne Edwards, 1863) [USNM E16325]. Hydropores represented by open circles, gonopores in solid black, ocular plates shaded, genital plates unshaded, anterior towards top of page. Scale bar is 1 mm long. Indian Ocean eastward to the northwestern coast of Australia. Bathymetric range. —9 to 75 m (Morten- sen 1948). Remarks. —Several attempts have been made to subdivide the genus Echinolampas into separate genera and subgenera (Mor- tensen 1948). None of these schemes have become universally accepted, and Kier (1962:107) finally decided “‘that all these sections and subgenera are based on char- acters too variable to be of generic distinc- Fig. 2. Periproctal plate patterns of echinolampa- dids and cassidulids: a, Echinolampas depressa Gray, 1851 [USNM E12929]; b, Cassidulus caribaearum La- marck, 1801 [Anegada, British Virgin Islands]. Peri- proct in solid black, primary spine tubercles repre- sented by two concentric circles, miliary spine tubercles by single open circle, aboral is towards top of page. Scale bars are 1 mm long. Fig. 3. echinolampadids and cassidulids: a, Conolampas sigs- bei (A. Agassiz, 1878) [USNM E12941] showing closed oval of distal teeth; b, Cassidulus caribaearum La- marck, 1801 [Anegada, British Virgin Islands] showing U-shaped, open row of distal teeth. Scale bars are 50 um long. Valves from ophicephalous pedicellariae of tion.’ In spite of Kier’s (1962) attempt to revive the name Echinolampas oviformis (Gmelin, 1789) for Echinolampas ovata, re- cent authors (Clark & Rowe 1971, Dollfus & Roman 1981) have argued convincingly for the retention of Leske’s (1778) old name. This species apparently can be found in the littoral zone of the Red Sea (Dollfus & Ro- man 1981). This is supported by observa- tions of this species in the intertidal, on fine carbonate sand without mud at Ashmore Reef, N. W. Australia (Lyle Vail, pers. comm.). At this locality, E. ovata also ap- pears to have a diurnal activity cycle, “bur- rowing through the sand with about half of their test exposed” at night, but remaining more deeply burrowed during the day (Lyle Vail, pers. comm.). McNamara & Philip (1980) speculated that this species lives slightly inclined, with the posterior deeper in the substrate, buried in the sand up to the level of the petaloids. Echinolampas alexandri de Loriol, 1876 Fig. 6b Geographic range.—The Red Sea and throughout the Indian Ocean and Malaysia. VOLUME 103, NUMBER 1 Fig. 4. Valves from globiferous pedicellariae of apatopygids: a, Apatopygus recens (Milne Edwards, 1863) [USNM E11089] showing single pair of fang- like teeth; b, Apatopygus occidentalis H. L. Clark, 1938 [after Baker 1983] showing two pairs of fang-like teeth. Scale bars are 100 um long. Bathymetric range.—8 to 365 m. Remarks.—Mortensen (1948) reported two subspecies (E. alexandri sibogae and E. alexandri forcipulata) to which Dollfus & Roman (1981) added a third (E. alexandri arctambulacrum). Mortensen (1948) felt that E. ovataand E. alexandri were very distinct, but because of variation in both species, Dollfus & Roman (1981) suggested that there is considerable overlap in morphology. Al- though more material from all parts of their ranges will be necessary before a complete comparison can be made, based on differ- ences in petaloid width and peristome shape (see key, above), it would appear that E. ovata and E. alexandri are good species. Echinolampas chuni (Déderlein, 1905) Fig. 6c Geographic range.—Known only from two denuded specimens from Sumatra (Doderlein 1906). Bathymetric range. —From a single “‘Val- divia”’ station at 371 m. Remarks. —No new information has come to light since Déderlein’s (1906) and Mor- tensen’s (1948) descriptions. Since the only known specimens were dead when collect- 71 Valves from ophicephalous pedicellariae of Fig. 5. neolampadids: a, Neolampas rostellata A. Agassiz, 1869 [USNM E20529], illustrating absence of thorns on ‘handle’; b, Nannolampas tenera (de Meijere, 1902) [after Mortensen 1948] showing presence of thorns on “handle.” Scale bars are 50 um long. ed, nothing is known of the biology of this species. Echinolampas crassa (Bell, 1880) Fig. 6d Geographic range. —Known only from the South African coast. Bathymetric range.—Mortensen (1948) says 25 to 500 m, but Thum & Allen (1975) record it from as shallow as 12 m. Remarks. —The ecology of this species is amongst the best known of all cassiduloids. Thum and Allen (1975:362) reported that this echinoid prefers the ripple slopes of rip- ple beds composed of biogenic substrates (75-95% calcium carbonate), and that it is ‘“‘an indirect deposit feeder and habitually burrows during feeding,” being frequently overlain by 30 to 50 mm of substrate. They also estimated a growth rate of approxi- mately 5 mm of test length per year. Their largest specimen was about 125 mm in length, suggesting (p. 373) “that this lamp urchin must be rather long-lived.”” Thum & QL &- ovata | b E. alexandri == Fig. 6. Family Echinolampadidae, genus Echinolampas: a, E. ovata (Leske, 1778) [after Mortensen 1948]; b, E. alexandri de Loriol, 1876 [after Mortensen 1948]; c, E. chuni (Déderlein, 1905) [after Déderlein 1906]; d, E. crassa (Bell, 1880) [after Mortensen 1948]; e, E. depressa Gray, 1851 [Florida Department of Natural Resources 13562]. From left to right for each species: lateral view, aboral view, oral view. Anterior end is to the left. Petaloids, peristomes, and periprocts in solid black, naked zone stippled. All scale bars are 10 mm long. 72 VOLUME 103, NUMBER 1 73 OL E. keiensis = b E. koreana == d E. sternopetala @ E. sumatrana | Fig. 7. Family Echinolampadidae, genus Echinolampas (cont'd): a, E. keiensis (Mortensen, 1948) [after Mortensen 1948]; b, E. koreana H. L. Clark, 1925 [after Clark 1925 and Mortensen 1948]; c, E. rangii Des- moulins, 1837 [after Agassiz 1872]; d, E. sternopetala A. Agassiz & H. L. Clark, 1907 [after Mortensen 1948]; e, E. sumatrana (Déderlein, 1905) [after Déderlein 1906]. Conventions as in Fig. 6. All scale bars are 10 mm long. 74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON b C. diomedeae = Fig. 8. Family Echinolampadidae (cont’d), genus Conolampas: a, C. sigsbei (A. Agassiz, 1878) [after Kier 1962]; b, C. diomedeae Mortensen, 1948 [after Mortensen 1948]; c, C. malayana Mortensen, 1948 [after Mortensen 1948]; d, C. murrayana Mortensen, 1948 [after Mortensen 1948]. Conventions as in Fig. 6. All scale bars are 10 mm long. Allen (1976:27) later analyzed the breeding behavior of E. crassa, discovering a “‘re- markable synchrony of gonad tissue mass both within and between sexes.”’ They also compared ambient substrate and gut con- tent particle dimensions, and suggested that larger particles are excluded from the diet because of the fixed mouth size, and smaller ones because the animal has difficulty ma- nipulating them (Thum & Allen 1976). These echinoids feed using the oral podia to lift particles to the peristome and can also VOLUME 103, NUMBER 1 75 Q C. caribaearum —>>E ee b C. infidus == C C. malayanus es Fig. 9. Cassidulidae, genus Cassidulus: a, C. caribaearum Lamarck, 1801 [Anegada, British Virgin Islands]; b, C. infidus Mortensen, 1948 [after Mortensen 1948]; c, C. mitis Krau, 1954 [after Krau 1954]; d, C. malayanus (Mortensen, 1948) [after Mortensen 1948]. Conventions as in Fig. 6. All scale bars are 10 mm long. use the circum-oral bourrelet spines to el- Echinolampas depressa Gray, 1851 evate particles into the mouth (Thum & AlI- Figs. 2a, 6e len 1976). Cram (1971) described the early Geographic range.—From eastern Cen- life history and larval morphology of E. tral America as far north as the north coast crassa, reporting that metamorphosis oc- of the Yucatan, the southeastern U.S., both curred at about 40 days after fertilization. coasts of Florida, the Greater and Lesser 76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0 Eurhodia relicta ———— C Rhyncholampas pacificus —— d Studeria recens aa | Fig. 10. Cassidulidae (cont'd): a, Eurhodia relicta Mooi, 1990 [Holotype, USNM E20480]; b, male Oligopodia epigonus (van Martens, 1865) [USNM E35684]; c, Rhyncholampas pacificus (A. Agassiz, 1863) [after Kier 1962]; d, Studeria recens (A. Agassiz, 1879) [after Agassiz 1881]. Conventions as in Fig. 6. All scale bars are 10 mm long. Antilles south to northeastern South Amer- Remarks. —Markel (1978) described the ica as far east as French Guyana. Aristotle’s lantern from juveniles of this Bathymetric range. —30 to 310 m (Serafy species. The lantern disappears as the ani- 1979). mal approaches 5 mm in length and does VOLUME 103, NUMBER 1 6) Apatopygus recens sl 4 G Porterpygus kieri Fig. 11. 77 Apatopygidae: a, Apatopygus recens (Milne Edwards, 1863) [USNM E16325]; b, Apatopygus occi- dentalis H. L. Clark, 1938 [after Baker 1983]; c, Porterpygus kieri Baker, 1983 [after Baker 1983]. Conventions as in Fig. 6. All scale bars are 10 mm long. not appear to function at all in the feeding of the echinoid. Serafy (1979) documented the occurrence of this species in the Gulf of Mexico on carbonate sands consisting large- ly of calcareous algal fragments. In spite of the apparent frequency with which this species is collected, little else is known of its ecology. Echinolampas keiensis (Mortensen, 1948) Fig. 7a Geographic range.—Known only from Kepulauan Kai (Kei Islands) and from off Zamboanga in the southern Philippines (Mortensen 1948). Bathymetric range. —245 to 400 m (Mor- tensen 1948). Remarks.—This species has apparently not been encountered since Mortensen’s (1948) original description. E. keiensis was originally placed in a separate genus, Plani- lampas Mortensen, 1948 along with Echi- nolampas sternopetala (see below). Echinolampas koreana H. L. Clark, 1925 Fig. 7b Geographic range.—From the Korean Strait (Clark 1925) and western Japan (N1- siyama 1968). Bathymetric range.—73 to 100 m. Remarks. —Clark (1925) compared his new species with EF. sternopetala, and Ni- siyama (1968) compared FE. koreana with E. alexandri. E. koreana differs from both 78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ‘6h Neolampas rostellata b Anochanus sinensis | BeOS VOLUME 103, NUMBER 1 of the other species in having a much higher test and a pentagonal peristome. It also lacks the distinctive inequality in the length of the respiratory podial columns so strongly developed in the petaloids of E. sternope- tala. For these reasons, I would agree with Mortensen (1948) in suggesting that these species are not particularly closely related. Nisiyama (1968) appears to be the only au- thor to have published on E. koreana since Clark (1925) described it from a single spec- imen. Nothing has been published on the biology of this species. Echinolampas rangii Desmoulins, 1837 Fig. 7c Geographic range. — Apparently from the west coast of Africa (Senegal) and the Cape Verde Islands (Mortensen 1948). Bathymetric range.—Mortensen (1948) recorded it from 1570 to 1670 m, but stated that it is undoubtedly also known from “shallow” water without suggesting an up- per limit. Remarks. —No work has been published on this species since Mortensen (1948). Echinolampas sternopetala A. Agassiz & H. L. Clark, 1907 Fig. 7d Geographic range.—Known only from Japanese waters from Sagami Bay to Ka- goshima Bay (Shigei 1986). Bathymetric range. —100 to 500 m (Shi- gei 1986). Remarks. — Mortensen (1948) made this the type species of a new genus, Planilam- pas, which he erected largely on the basis of the flatness of the oral surface. Kier (1962) —_— 79 synonymized this genus with Echinolam- pas. Shigei (1986) described Echinolampas sternopetala from Sagami Bay, Japan, but provided no comments on substrate type or ecology. Echinolampas sumatrana (Déoderlein, 1905) Fig. 7e Geographic range. —Known from a single denuded test collected off Sumatra. Bathymetric range. —From a single “‘Val- divia”’ station at 371 m. Remarks. — Mortensen (1948) agreed with Doderlein (1906) in suggesting that more specimens of E. sumatrana were necessary before it could be firmly established that it was distinct from E. crassa. However, dif- ferences in peristome shape and petaloid width (see key, above) seem to indicate that E. sumatrana is distinct from E. crassa. Conolampas (A. Agassiz, 1883) Conolampas sigsbei (A. Agassiz, 1878)* Figs. 3a, 8a Geographic range.—Throughout the Greater and Lesser Antilles, west coast of Florida, and the north and northeastern coasts of the Yucatan. Bathymetric range. —120 to 800 m. Remarks.—Mooi (1990) recently sum- marized what little is known of the biology of this species, suggesting that it lives on the surface of the sediment, not burrowed, and feeds on relatively fine carbonate substrates. Judging from the flatness of the oral surface and overall similarity in spination and test shape, other species of Conolampas prob- ably have similar habits. Fig. 12. Neolampadidae: a, Neolampas rostellata A. Agassiz, 1869 [MCZ 2739]; b, presumed female Ano- chanus sinensis Grube, 1868 [reconstructed from Mortensen 1948 and McKnight 1968]; c, presumed male Aphanopora echinobrissoides de Meijere, 1902 [after McKnight 1968]; d, Nannolampas tenera (de Meijere, 1902) [after Mortensen 1948]; e, male Tropholampas loveni (Studer, 1880) [after Mortensen 1948 and MCZ 4507]. Conventions as in Fig. 6. All scale bars in mm. 80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Conolampas diomedeae Mortensen, 1948 Fig. 8b Geographic range. —Mortensen (1948) recorded a single specimen (the holotype) from off Mindoro Island in the Philippines. David & de Ridder (1989) describe an ad- ditional seven specimens from the Philip- pines. Bathymetric range. —Mortensen’s (1948) specimen was dredged from 265 m, David and de Ridder’s (1989) from stations be- tween 181 and 195 m. Remarks. —David & de Ridder’s (1989) specimens are the only ones collected since Mortensen’s (1948) description. They pro- vide some biometrics and figures, but no information on the biology of this species. Conolampas malayana Mortensen, 1948 Fig. 8c Geographic range.—Known only from Kepulauan Kai (Kei Islands) and off Zam- boanga in the southern Philippines (Mor- tensen 1948). Bathymetric range. —245 to 400 m. Remarks.—No new information since Mortensen (1948) discovered the species. Conolampas murrayana Mortensen, 1948 Fig. 8d Geographic range. —Only two specimens known from off the Maldive Islands in the Indian Ocean. Bathymetric range.— Dredged from 229 m. Remarks.—No new information since Mortensen (1948). Family Cassidulidae L. Agassiz & Desor, 1847 Cassidulus Lamarck, 1801 Cassidulus caribaearum Lamarck, 1801* Figs. la, 2b, 3b, 9a Geographic range. —Belize, Central America, and from the Bahama Islands south through the Virgin, Leeward, and Windward Islands to Barbados. Bathymetric range. — Very shallow water, from less than 1 m to probably no deeper than 10 m. Agassiz (1872) reported frag- ments from 197 m, but it is not at all certain whether these actually represent C. cari- baearum. Remarks. —Some confusion has arisen on the spelling of the species name, recent pa- pers reporting it as C. cariboearum (Kier 1975) and C. caribbearum (Gladfelter 1978). As Mortensen (1948) recorded, Lamarck’s original and correct spelling is C. caribaea- rum. Kier (1975) described this species as living buried in coarse sand at Carrie Bow Cay, Belize. Gladfelter (1978) reported that C. caribaearum broods its young among the aboral spines and was locally abundant in coarse, oolitic carbonate sand in very shal- low water. He also described locomotion in this species, which uses “‘ditaxic waves, passing from the front to the rear of large movable spines on the lateral portions of the ventral surface; this mechanism is unique among echinoids’” (p. 149). C. caribaearum feeds in much the same way as described for Echinolampas crassa (see above), but apparently does not rely as much on activ- ities of the circum-oral spination (Gladfelter 1978). Gladfelter (1978) also studied repro- ductive biology and seasonal variation in population density in this widely distrib- uted, but surprisingly seldom encountered, shallow water cassiduloid. Cassidulus infidus Mortensen, 1948 Fig. 9b Geographic range.—Known only from a single specimen labeled “Bahia” (Morten- sen 1948). Probably from the east coast of South America. Bathymetric range. —No depth record ex- ists for the holotype, but Mortensen (1948) felt that it probably occurs in shallow water. Remarks. — Although Krau (1954) com- pared her new species, C. mitis, with C. in- fidus, she relied on Mortensen’s (1948) de- scription, as no new information has been published on the latter, poorly known species. VOLUME 103, NUMBER 1 Cassidulus mitis Krau, 1954 Fig. 9c Geographic range.—Known only from Sepetiba Bay, near Rio de Janeiro, Brazil. Bathymetric range. —Krau (1954) did not provide precise depth data, but said that the new species was collected from “shallow waters.” Remarks. —Tommasi & Lima-Verde (1970) synonymized Cassidulus delectus Krau, 1960 with C. mitis and noted that C. mitis broods its young in a manner similar to that of C. caribaearum (Gladfelter 1978). Cassidulus malayanus (Mortensen, 1948) Fig. 9d Geographic range. —Only two known specimens, both from Kepulauan Kai (Kei Islands), according to Mortensen (1948). Bathymetric range.— Approximately 250- 290 m (Mortensen 1948). Remarks. —Mortensen (1948) described this species as a member of the genus Pro- cassidulus Lambert, 1918. The latter genus was erected by Lambert (1918) to replace Cassidulus, which he thought was preoc- cupied (Mortensen 1948). Kier (1962) syn- onymized Procassidulus with Rhynchopy- gus d’Orbigny, 1856. Rhynchopygus is stated by Kier (1962) to have a tetrabasal apical system. According to Mortensen (1948), Procassidulus malayanus has a ““compact”’ (=monobasal) apical system and so cannot be a Rhynchopygus. Since this species is so much like other species in Cassidulus, it ap- pears best to place C. malayanus in this genus, at least until more material becomes available. Mortensen (1948:225) reported that the two specimens he studied were tak- en from a “sandy bottom.” Eurhodia Haime in d’Archiac & Haime, 1853 Eurhodia relicta Mooi, 1990 Fig. 10a Geographic range. —Only two specimens known, the holotype from off western Suri- 81 nam, paratype from off Venezuela (Mooi 1990). Bathymetric range.—Dredged from 57 and 112 m, respectively. Remarks. —The type species of the genus is Eurhodia morrisi Haime in d’Archiac & Haime, 1853. E. relicta apparently occurs in terrigenous, siliceous substrates (Mooi 1990), but little else is known of its biology. Oligopodia Duncan, 1889 Oligopodia epigonus (van Martens, 1865)* Fig. 10b Geographic range. —Mortensen (1948) recorded this species from off the east coast of Africa, the Malaysian region, off Jolo in the Philippines, and the Bonin Islands. Be- cause specimens from off the northeast coast of Somalia, South Africa, Tonga, and New Zealand can be found in the U.S. National Museum (USNM), it would appear that this species ranges from the western Indian Ocean eastward through Indonesia and the Philippines all the way to the South Pacific Ocean. This means that O. epigonus has the largest range of any known living species of cassiduloid. Bathymetric range. —Live specimens are recorded from 35 to 141 m. Mortensen re- ported dead specimens from 5 to 390 m, suggesting that the bathymetric range could be much greater than the live specimens indicate. Remarks. —Females of this species have gonopores many times larger than those of the male (Mortensen 1948). Although once thought to be closely related to Apatopygus, Oligopodia differs in having a monobasal apical system and in ambulacral plate pat- tern and overall test shape. This genus ap- pears to be much more closely related to Eurhodia, as noted by Mooi (1990). Mor- tensen (1948:232) said that “‘it is not rare in places with a suitable bottom, viz. a coarse sandy or even gravelly bottom” and re- ported that it occasionally is found in sub- strates with a high percentage of calcareous algal fragments. 82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rhyncholampas A. Agassiz, 1869 Rhyncholampas pacificus (A. Agassiz, 1863)* Fig. 10c Geographic range. —The tropical eastern Pacific Ocean, from the Gulf of California to the Gulf of Panama. Also known from the Galapagos Islands (Mortensen 1948). Bathymetric range.—Clark (1925) said that it occurs from 7 to 130 m, but in his description of Rhyncholampas, Agassiz (1872) recorded it from water less than 2 m deep. Remarks.—The biology of this large species of cassidulid is surprisingly poorly known. Agassiz (1872) mentioned that the living echinoid lives partially buried in the substrate up to the level of the petaloids, but no other direct observations of living specimens have since been reported. Studeria Duncan, 1891 Studeria recens (A. Agassiz, 1879) Fig. 10d Geographic range.— Agassiz (1881) re- corded the species from the Arafura Sea, south of Papua-New Guinea. Bathymetric range. — Dredged from 236 m. Remarks. — According to Kier (1962), the type species of the genus is Studeria elegans (Laube, 1869) because this was the only species included in the original description of the genus. Many previous authors had considered S. recens to be the type of Stu- deria. There is some uncertainty that S. re- cens is a Studeria, but there are strong sim- ilarities (except in overall test shape) between this species and Kier’s (1962) figure of S. subcarinatus (Goldfuss, 1928). S. recens also fits the description of the genus (Kier 1962: 216), particularly with respect to the 3 gono- pores, strongly developed bourrelets, and absence of an oral, medial naked zone. Kier (1962) synonymized Hypselolampas H. L. Clark, 1917 (which included H. recens) with Studeria because of a lack of significant characters separating S. recens from other members of Studeria Duncan, 1891. No other occurrences of this poorly known species have been published since Agassiz’s (1881) description and nothing is known of its biology apart from the fact that it was collected from mud. Family Apatopygidae Kier, 1962 Apatopygus Hawkins, 1920 Apatopygus recens (Milne Edwards, 1863)* Figs. 4a, lla Geographic range.—Known only from New Zealand. Bathymetric range. —6 to 146 m. Remarks. —The behavior and general bi- ology of this well-known species have been described by Higgins (1974). The echinoid burrows to a depth of approximately 30 mm into coarse sands or fine gravels composed of terrigenic material and high proportions of broken shell. Higgins (1974:513) sug- gested that A. recens is “‘an almost contin- uous deposit feeder,’ but did not describe the manner by which food is collected. Apatopygus occidentalis H. L. Clark, 1938 Figs. 4b, 11b Geographic range.—Five specimens are known from along the western and southern coasts of Australia (Baker 1983). Bathymetric range.—17 to 40 m (Baker 1983). Remarks.— Although Baker (1983) pro- vided a detailed redescription of this very rare species, almost nothing is known of its general biology. Porterpygus Baker, 1983 Porterpygus kieri Baker, 1983* pam lalc Geographic range.—Known only from five specimens collected near Three Kings Islands, New Zealand (Baker 1983). VOLUME 103, NUMBER 1 Bathymetric range.— Dredged from ap- proximately 90 to 300 m. Remarks. —Baker (1983:172) suggested that the discovery of this second genus at- tributable to the family Apatopygidae “‘sup- ports the isolation of Kier’s Apatopygidae as a separate family.”’ Nothing is known of the biology of this species. Family Neolampadidae Lambert, 1918 Neolampas A. Agassiz, 1869 Neolampas rostellata A. Agassiz, 1869* Figs. 5a, 12a Geographic range.— West Indian waters and the Eastern Atlantic, particularly Flor- ida, across the Atlantic to the Moroccan coast, and the Mediterranean. Bathymetric range.—Mortensen (1948) reported the species from 145 to 1260 m. Remarks. —Examination of specimens in the National Museum of Natural History (USNM) and the Museum of Comparative Zoology, Harvard (MCZ) reveals that, as in Oligopodia, females of this species have much larger gonopores than the males. Al- though this species is fairly well known from North American waters, its preference for deep waters has prevented direct observa- tion of habits. Anochanus Grube, 1868 Anochanus sinensis Grube, 1868* Fig. 12b Geographic range.— Reported only from the China Sea. Bathymetric range.—No depth record available for the only known specimen. Remarks. —Both Mortensen (1948) and McKnight (1968) have suggested that 4An- ochanus sinensis may be the female of Aphanopora echinobrissoides (see below). Because the only known specimen has never been completely figured, the drawings pre- sented here have been based on the similar Aphanopora and the description given in Mortensen (1948). 83 Aphanopora de Meijere, 1902 Aphanopora echinobrissoides de Meijere, 1902* Fig. 12c Geographic range. —Mortensen (1948) recorded the species from the Timor and Sulu Seas, and McKnight (1968) described two specimens from an area approximately 150 miles north of Norfolk Island. Bathymetric range.—The Timor and Sulu specimens were found at 350 to 390 m, and McKnight’s (1968) specimens were dredged from 710 m. Remarks. —This species might actually be sexually dimorphic, with the females (pos- sibly represented by Anochanus sinensis, see above) possessing an aboral brood cavity at the apical system. Therefore, the males would probably not have a sunken apical system, as figured by McKnight (1968). McKnight (1968) redescribed the species from material representing a range exten- sion. However, his specimens were denuded tests, which prevented any speculation on the biology of the species. Nannolampas Mortensen, 1948 Nannolampas tenera (de Meijere, 1902)* Figs. 5b, 12d Geographic range. — Known only from the Timor Sea. Bathymetric range.—From a single “*Si- boga’”’ station, at 390 m. Remarks.—Mortensen (1948) assigned Neolampas tenera de Meijere, 1902 toa new genus, Nannolampas, because the latter species has a tetrabasal apical system. Noth- ing is known of the biology of N. tenera. Tropholampas H. L. Clark, 1923 Tropholampas loveni (Studer, 1880)* Fig. 12e Geographic range. —South African coast, Cape Peninsula to Cape St. Francis (Mor- tensen 1948). 84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bathymetric range. — Dredged from 135- 350 m (Mortensen 1948). Remarks.—The females of this species have a strongly developed aboral “‘marsu- plum” at the apical system, and the males have only a very slightly sunken apical sys- tem. In spite of the markedly different po- sitions of the periproct, Mortensen (1948) thought that 7ropholampas was closely re- lated to Anochanus on the basis of common possession of an adapical brood cavity. Acknowledgments I would like to thank David Pawson, Mal- colm Telford, and an anonymous reviewer for reading early versions of the paper, and Cindy Ahearn for her tireless efforts in lo- cating collection data. Personal support was provided by a Natural Sciences and Engi- neering Research Council of Canada Post- doctoral Fellowship and additional funding by the Smithsonian Institution. Literature Cited Agassiz, A. 1872-1874. Revision of the Echini.— Memoirs of the Museum of Comparative Zo- ology 3:383-762. 1881. Report on the scientific results of the voyage of the H.M.S. “Challenger” during the years 1873-1876, Zoology, Vol. III, Part IX, Report on the Echinoidea. London, England, Longmans & Company, 321 pp. Baker, A.N. 1983. A new apatopygid echinoid genus from New Zealand (Echinodermata: Cassidu- loida).—Records of the National Museum of New Zealand 2(15):163-173. Clark, A. M., & F. W. E. Rowe. 1971. Monograph of the shallow-water Indo-west Pacific echino- derms. London, British Museum (Natural His- tory), 238 pp. Clark, H. L. 1925. A catalogue of the Recent sea- urchins (Echinoidea) in the collection of the British Museum (Natural History). London, En- gland, Oxford University Press, 250 pp. Cram, D. L. 1971. Life history studies on South Af- rican echinoids (Echinodermata) 2. Echinolam- pas (Palaeolampas) crassa (Bell) (Echinolam- padidae).— Transactions of the Royal Society of South Africa 39(3):339-352. David, B., & C. de Ridder. 1989. Echinodermes: Echinides irréguliers. Résultats des campagnes MUSORSTOM, volume 4.— Mémoires du Mu- séum Nationale d’Histoire Naturelle Paris (Serie A) 143:203-227. Doéderlein, L. 1906. Die Echinoiden der deutschen Tiefsee-Expedition. — Wissenschaftliche Ergeb- nisse der deutschen Tiefsee-Expedition auf dem Dampfer “Valdivia” 1898-1899 5(2):61-290. Dollfus, R. P., & J. Roman. 1981. Les échinides de la Mer Rouge. Paris, Bibliothéque Nationale, 145 pp. Gladfelter, W. B. 1978. General ecology of the cas- siduloid urchin Cassidulus caribbearum [sic].— Marine Biology 47(2):149-160. Higgins, R. C. 1974. Observations on the biology of Apatopygus recens (Echinoidea: Cassiduloida) around New Zealand.— Journal of Zoology Lon- don 173:505-516. Kier, P. M. 1962. Revision of the cassiduloid echi- noids.— Smithsonian Miscellaneous Collections 114(3):1-262. 1974. Evolutionary trends and their func- tional significance in the post-Paleozoic echi- noids.—Journal of Paleontology 48 (supple- ment) Paleontological Society Memoirs 5:1-95. 1975. The echinoids of Carrie Bow Cay, Be- lize.—Smithsonian Contributions to Zoology 206:1-45. —., & M. H. Lawson. 1978. Index of living and fossil echinoids 1924-1970.—Smithsonian Contributions to Paleobiology 34:1—182. Krau, L. 1954. Ona new species of Recent sea urchin, Cassidulus mitis, order Cassiduloida, Echinoi- dea, found on the coast of Sepetiba Bay.— Me- morias do Instituto Oswaldo Cruz 52(2):455- 475. Lambert, J. 1918. Considerations sur la classification des échinides atelostomes. — Mémoires de la So- ciete Academique de |’Aube ser., 3 55:9-54. Leske, N.G. 1778. Additamenta ad Jacobi Theodori Klein naturalem dispositionem Echinoderma- tum et lucubratiunculam de aculeis echinorum marinorum. Lipsiae, 214 pp. Markel, K. 1978. On the teeth of the Recent cassi- duloid Echinolampas depressa Gray, and on some Liassic fossil teeth nearly identical in structure (Echinodermata, Echinoidea).— Zoomorphologie 89:125-144. McKnight, D. G. 1968. Some echinoids and ophi- uroids from off Norfolk Island and from Wan- ganella Bank.— New Zealand Journal of Marine and Freshwater Research 2:204—213. McNamara, K. J., & G. M. Philip. 1980. Tertiary species of Echinolampas (Echinoidea) from southern Australia.—Memoirs of the National Museum, Victoria 41:1-14. Mooi, R. 1986. Non-respiratory podia of clypeaster- oids (Echinodermata, Echinoides): II. Diversi- ty.—Zoomorphology 106:75-90. VOLUME 103, NUMBER 1 . 1987. Acladistic analysis of the sand dollars (Clypeasteroida: Scutellina) and the interpreta- tion of heterochronic phenomena. Unpublished Ph.D. dissertation, University of Toronto, Can- ada, 208 pp. . 1989. Living and fossil genera of the Clypeas- teroida (Echinoidea: Echinodermata): An illus- trated key and annotated checklist.—Smithson- ian Contributions to Zoology 488:1-51. 1990. A new “living fossil’? echinoid (Echi- nodermata) and the ecology and paleobiology of Caribbean cassiduloids.— Bulletin of Marine Science 46:(in press). Mortensen, T. 1948. A monograph of the Echinoidea, 4(1). Holectypoida, Cassiduloida. Copenhagen, Denmark, C. A. Reitzel, 471 pp. Nisiyama, S. 1968. The echinoid fauna from Japan and adjacent regions, Part 2.— Palaeontological Society of Japan Special Papers 13:1-491. Philip,G. M. 1963. Two Australian Tertiary neolam- padids and the classification of cassiduloid echi- noids.— Palaeontology 6(4):718-726. Serafy,D.K. 1979. Memoirs of the Hourglass cruises. St. Petersburg, Florida, Florida Department of Natural Resources, 120 pp. Shigei, M. 1986. The sea urchins of Sagami Bay col- 85 lected by His Majesty the Emperor of Japan. Tokyo, Japan, Maruzen, 204 pp. Suter, S. J. 1988. The decline of the cassiduloids: Merely bad luck?— Proceedings of the Interna- tional Echinoderms Conference, Victoria, 1987, pp. 91-95. Thum, A. B., & J. C. Allen. 1975. Distribution and abundance of the lamp urchin Echinolampas crassa (Bell) 1880 [sic] in False Bay, Cape.— Transactions of the Royal Society of South Af- rica 41(4):359-373. ——,,& . 1976. Reproductive ecology of the lamp urchin Echinolampas crassa (Bell), 1880 [sic] from a subtidal biogenous ripple train. — Transactions of the Royal Society of South Af- rica 42(1):23-33. Tommasi, L. R., & J. S. Lima-Verde. 1970. Obser- vacoes sObre Cassidulus mitis Krau, 1960 [sic, should be 1954] (Cassiduloida, Echinoidea).— Boletim Instituto Oceanografico Sao Paulo 18(1): 1-9. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 86-94 A NEW SPECIES OF HALOPHILOUS WATER-STRIDER, MESOVELIA POLHEMUST, FROM BELIZE AND A KEY AND CHECKLIST OF NEW WORLD SPECIES OF THE GENUS (HETEROPTERA: MESOVELIIDAE) Paul J. Spangler Abstract. —A new species of water strider, Mesovelia polhemusi, from man- grove cays in the Stann Creek District of Belize is described, illustrated by line drawings and scanning electron micrographs, and compared with M. hamble- toni Drake & Harris, a similar species, and M. halirrhyta Polhemus from Costa Rica, the only other described halophilous species included in Mesovelia. A key to males and a checklist of the 10 species presently described from the Western Hemipshere are included. According to the classification of the Mesoveliidae by Andersen & Polhemus (1980), there are 23 species and subspecies of Mesovelia described for the world fauna; 14 from the Old World and 9 from the New World. Because it is necessary to examine many specimens of the widely distributed M. mulsanti to confirm or refute the infra- specific categories cited by Jaczewski (1930), only the nominate subspecies (with bisig- nata as a synonym) is included in the check- list below. The tenth New World species, Mesovelia polhemusi, new species, is de- scribed below. The new species of Mesovelia was col- lected during a survey of aquatic and semi- aquatic Hemiptera and Coleoptera of man- grove, coral reef, and estuarine habitats in the Stann Creek District of Belize. Collec- tions of a variety of water bugs were made in the Sittee River estuary at Sittee Point, in Placencia Lagoon, and on numerous cays—on Bread and Butter Cay, the reef crest at Carrie Bow Cay, Man of War Cay, Round Cay, Twin Cays, Wee Wee Cay, and cays on the Blue Ground Range. Specimens of Mesovelia polhemusi, new species, were found only on the mangrove cays where they occurred on mud flats among the prop-roots of red mangrove and the pneumatophores of black mangrove during ebb tide and on the surface of the salt water or resting on emergent vegetation when the tide was incoming or full. The first specimen of this undescribed species was found in April 1986 while I was examining mud flats between the dense tan- gle of roots of the red mangrove Rhizophora mangle L. That find led to a concerted search for more specimens and some were ulti- mately found. They did not seem to be widely dispersed but some were found among pneumatophores of black mangrove, Avicennia nitida Jacquin. In order to obtain some information on the abundance of this species, counts of specimens were made for 8 plots of 1 m among the pneumatophores. The number of specimens from the plots varied from 0 to 18 with 6 being the mean ofthe sample. During the fieldwork in 1986, the bugs were difficult to find but abundant specimens were found on numerous cays during subsequent trips as listed under the type data below. During limited collecting on the mainland in freshwater estuarine habitats Mesovelia mulsanti White was found to be abundant but no specimens of M. polhemusi were obtained. This new species probably occurs widely on the many cays along the coast of Belize and perhaps VOLUME 103, NUMBER 1 it will be found on the mainland if more intensive collecting efforts are made there. The specimens of the type series were found on mud flats and the water surface among the prop roots of red mangrove and pneumatophores of black mangrove. Some were hiding in small cracks, fiddler crab bur- rows, and similar niches until flushed out of hiding. During subsequent field trips oth- er specimens were found running about openly, usually on shaded, damp to muddy soil. Mesovelia polhemusi, new species Figs. 1—20 Holotype 6 (Figs. 1, 2).—Body form and size: Elongate; slightly widening to mid- length of abdomen then converging to end of genital capsule. Length, 2.21 mm; width (across abdomen), 0.64 mm. Color: Ground color yellowish brown; with indication of a yellowish longitudinal vitta on midline of dorsum. Connexivia light yellowish brown. Legs light yellowish brown except each femur with darker brown sub- apical band. Antennal segments dark red- dish brown except basal segment lighter yel- lowish brown. Venter yellowish except brown laterally. Body, legs, and antennae with very short yellowish brown, decum- bent pubescence. Head (Figs. 3-5): Length, 0.46 mm; broadened anteriorly to tubercles; width be- tween tubercles, 0.35 mm. Median sulcus posteriorly reaching caudal margin, becom- ing evanescent anteriorly between eyes. Eyes large (width of eye/interocular space: 0.15/ 0.21 mm); length, 0.21 mm; with many om- matidia, converging anteriorly. Rostrum extending almost to anterior margin of metacoxae. Antenna (Fig. 19) long, slender; segment | stoutest; lengths of segments 1- 4: 0.50 mm, 0.34 mm, 0.62 mm, 0.62 mm. Antennal segment | with numerous short setae and one long, anteriorly directed, thin seta at distal third. Antennal segments 2, 3, and 4 set with many long, thin setae. Cuticle 87 with dense plastron setae and peg plates (Figs. 3-6). Thorax (Fig. 16): Pronotum and meso- notum granulate along hind margins; meta- notum granulate on each side of midline. Pronotum short; posterior margin straight. Midline lengths of pronotum, mesonotum, metanotum = 0.21, 0.20, 0.12 mm. Pro- sternum with dense plastron setae (Fig. 6). Legs with numerous longer, thin, hairlike setae; metatibia with scattered stiff brown setae. Protibia with grooming comb on apex (Figs. 7, 8). Protarsus (Fig. 9) with long, slender claws (Fig. 10). Right profemur armed beneath with 11 black spines; left profemur with 9 black spines; length of spines equal 4 to 2 the width of the femur where they arise. Right mesotibia with grooming comb apically (Fig. 11). Right mesofemur armed beneath with 20 black spines (Fig. 12); left mesofemur with 15 black spines of similar length as spines of profemora. All femora moderately stout; tibia and tarsi slender. Abdomen (Fig. 13): Long, slender. Seg- ment 3 widest. Lengths of abdominal terga 1-7 = 0.12 mm, 0.15 mm, 0.12 mm, 0.15 mm, 0.14 mm, 0.17 mm, 0.23 mm. Median scent pore at anterior third of tergum 4 (Figs. 14, 15). First genital segment (8th abdom- inal) without spine or cluster of black spi- nules. Cuticle with peg plates (Fig. 17) and spiracles (Fig. 18). Genitalia: As illustrated (Fig. 20). Female. —Length, 2.75 mm; width (across abdomen), | mm. Fore and middle femora armed beneath with 9 and 16 spines, re- spectively. Variations. —The number of spines on the femora was found to be variable. The fem- oral spines varied as follows (n = 20 for each sex). Profemoral: males, 5-12 (x = 10); fe- males, 1-12 (< = 7). Mesofemoral: males, 14-21 (x = 17); females, 6-17 (* = 11). Males (n = 20) ranged in length from 2.1 1- 2.40 mm (xX = 2.14) and in greatest width from 0.64—0.78 mm (X = 0.71). Females (n = 20) ranged in length from 2.33-2.75 mm PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON er | Figs. 1-6. Mesovelia polhemusi, n. sp. 1, habitus, dorsal, x23; 2, habitus, ventral, x22; 3, head, dorsal, 150; 4, head, ventral, x170; 5, plastron setae, ventral surface of head, x 1100; 6, plastron on prosternum, x 2800. VOLUME 103, NUMBER 1 Figs. 7-12. Mesovelia polhemusi, n. sp. 7, protibial grooming comb, x 700; 8, protibial grooming comb, x 800; 9, protarsus, x 500; 10, protarsal claws, x 1300; 11, mesotibial grooming comb, x 1000; 12, mesofemur, x 350. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 13-18. Mesovelia polhemusi, n. sp. 13, abdominal terga, x 60; 14, abdominal tergum 4 and scent gland orifice, x 250; 15, abdominal scent gland orifice, x 600; 16, head and thorax, x60; 17, peg plates and spiracle, side of abdomen, =x 300; 18, spiracle, x 5000. VOLUME 103, NUMBER 1 Figs. 19-23. Mesovelia polhemusi, n. sp. 19, antenna; 20, genital capsule with clasper. 21, Mesovelia amoena Uhler, male clasper. 22, Mesovelia hambletoni Drake & Harris, male clasper. 23, Mesovelia cryptophila Hun- gerford, male clasper. (X = 2.32) and in greatest width from 0.80- 1.01 mm (X = 0.93). Type data.—Holotype, apterous male: Belize: Stann Creek District: Twin Cays, 18 May 1986, P. J. Spangler and R. A. Fai- toute; deposited in the U.S. National Mu- seum of Natural History, Smithsonian In- stitution. Allotype: Same data as holotype. Paratypes: Belize: Stann Creek District: Same data as holotype, 37 specimens. Blue Ground Range, 9 Nov 1987, P. J. Spangler and R. A. Faitoute, colln 20, 35 specimens. Bread and Butter Cay, 25 Mar 1988, R. A. Faitoute, colln 25, 15 specimens. Man of War Cay, 16 May 1986, P. J. Spangler and R. A. Faitoute, colln 6, 1 specimen; 8 Nov 1987, P. J. Spangler and R. A. Faitoute, colln 16, 31 specimens. Round Cay (S of Coco Plum), 23 Mar 1988, R. A. Faitoute, colln 19, 7 specimens. Twin Cays: Aanderaa Flats, 19 May 1986, P. J. Spangler and R. A. Faitoute, colln 11, 38 specimens; Aan- deraa Flats, 7 Nov 1987, P. J. Spangler and R. A. Faitoute, colln 12, 12 specimens; Aan- deraa Flats, 8 Nov 1987, 20 specimens; Aanderaa Flats, 17 Mar 1988, R. A. Fai- 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Mesovelia polhemusi, lengths of segments of legs (in millimeters). Tarsal Tarsal Tarsal Femur Tibia 1 2 3 Anterior 0.73 O.58 0.04 0.04 0.06 Middle 0.85 0.85 0.08 0.14 0.21 Posterior 1.16 1.35 0.10 0.19 0.15 toute, colln 2, 25 specimens; Aanderaa Flats, 21 Mar 1988, R. A. Faitoute, colln 11, 20 specimens; Hidden Lake, 17 Mar 1988, R. A. Faitoute, colln 4, 36 specimens; north- east side of island, 21 Mar 1988, R. A. Fai- toute, colln 12, 1 specimen; West Pond, 18 Mar 1988, R. A. Faitoute; colln 5, 26 spec- imens. Wee Wee Cay, 24 Mar 1988, R. A. Faitoute, colln 20, 19 specimens. Paratypes will be deposited in the Museo de Zoologia, Sao Paulo; the American Mu- seum of Natural History, New York, New York; the British Museum (Natural Histo- ry), London; the California Academy of Sci- ences, San Francisco, California; the Ca- nadian National Collection, Ottawa; Institute Royal de Histoire naturelle de Bel- gique, Bruxelles; Instituto de Zoologia Agri- cola, Facultad de Agronomia, Maracay, Venezuela; Laboratorium voor Zoolo- gische, Oecologie en Taxonomie, Utrecht; the Museum National de Histoire Natural, Paris; Museo Argentina de Ciencias natu- rales, Buenos Aires; Universidad Nacional de La Plata, La Plata; the National Museum of Natural History, Smithsonian Institu- tion, Washington, D.C.; the Snow Ento- mological Museum, University of Kansas, Lawrence, Kansas; Zoologische Sammlung Bayerischen Staates, Munchen; and the col- lection of John T. Polhemus, Englewood, Colorado. Etymology. —This new species, Mesove- lia polhemusi, is dedicated to my colleague and indefatigable aquatic heteropterist, John T. Polhemus, who has greatly increased our knowledge of these fascinating insects. Comparative notes. —The male of Meso- velia polhemusi, new species, may be sep- arated easily from its congeners by the fol- lowing combination of characters. Middle femur with a row of black spines; first gen- ital segment (8th abdominal) without spine or cluster of black spinules; length of anten- nal segment 2 distinctly less than the width of the vertex and one eye. Mesovelia pol- hemusi is most similar to the somewhat variable Ecuadorian species Mesovelia hambletoni Drake & Harris (1946) but dif- fers from it by its smaller size (length less than 3 mm vs. more than 3.5 mm), and the male clasper (Fig. 20) not hooklike as on M. hambletoni (Fig. 22). Males of Mesovelia polhemusi may be distinguished from the only other described strictly halophilous species of the genus, Mesovelia halirrhyta Polhemus (1975), by the absence of the two clusters of spinules on the first genital segment (8th abdominal) and the presence of only 15—20 black spines on the mesofemur vs. 25 spines reported for the 9 specimens in the type series of M. halirrhyta. The only key to the species of Mesovelia of the Western Hemisphere is that by Jac- zewski (1930) for four of the five species known at that time. In the intervening 59 years, the number of species of Mesovelia from the New World has doubled and the following key is provided to the males of the 10 species presently recognized from this hemisphere. Because of similarities between M. polhemusi and M. hambletoni as well as between M. cryptophila and the wide-rang- ing M. amoena, the diagnostic male claspers of these taxa are illustrated and referred to in the key. A complete clasper (Fig. 23) of M. cryptophila is also illustrated because Hungerford’s (1924b) “‘freehand sketches of the left genital claspers of male Mesovelias”’ did not include the base and is misleading. Key to Males of Mesovelia of the Western Hemisphere 1. Mesofemur with a row of prom- inent black spines ventrally ..... D, Mesofemur without row of black SPINES’ 2. aceed oh eee d/ VOLUME 103, NUMBER 1 Table 2.—Checklist of New World Mesovelia Mulsant & Rey. Species Locality amoena Uhler, 1894:218 (=douglasensis Hungerford, 1924a:142) bila Jaczewski, 1928:77 blissi Drake, 1949:146 cryptophila Hungerford, 1924b:454 halirrhyta Polhemus, 1975:245 hambletoni Drake & Harris, 1946:8 mulsanti White, 1879:268 (=bisignata Uhler, 1884:274) polhemusi Spangler, 1990:87 thomasi Hungerford, 1951:33 zeteki Harris & Drake, 1941:276 2(1). 3(2). 4(3). 5(4). Ecuador Belize First male genital segment (eighth abdominal) with subbasal black- tipped median spine ventrally or two clusters of black spinules ... 3 First male genital segment (eighth abdominal) without spine or cluster of black spinules First male genital segment (eighth abdominal) with a subbasal black- tipped median spine ventrally blissi Drake, 1949 First male genital segment (eighth abdominal) with two clusters of black spinules First male genital segment (eighth abdominal) with two patches of widely separated black spinules ventrally; each patch with about ZOO ONSpinules ye) json) eee Re dere Rie re thomasi Hungerford, 1951 First male genital segment (eighth abdominal) with two tightly packed clusters of coarse or thin spinules Mesofemur with 25 black spines ventrally. Males with two widely separated rows of thin spinules on first genital sternum; each row with three to five spinules. Wenethy2=s—2-7is) TM) ae halirrhyta Polhemus, 1975 Mesofemur with 8 to 15 black spines. Males with two tightly Guatemala; Mexico Brazil; Panama 7(1). 8(7). 9(7). USA: Michigan to Florida, Nevada, California, Texas, Hawaiian Is- lands; Belize; Bonaire; Brazil; Costa Rica; Curacao; Dominican Republic; Mexico; Panama Argentina; Brazil Mexico; Panama USA: Florida, Iowa, Kansas, Michigan, Mississippi, New Jersey Colombia; Costa Rica Southern Canada; eastern half of USA, Hawaiian Islands; neotropi- cal region to Argentina packed clusters of stout black spi- nules on first genital sternum. Kenethas—4amnmiee nee cee mulsanti White, 1879 Small species; length less than 3.0 mm. Male clasper, in lateral view, not hooklike; apex blunt ante- riorly (Fig. 20) Silas ar WENT Be polhemusi, new species Large species; length more than 3.5 mm. Male clasper, in lateral view, strongly hooklike; apex acute and directed anteroven- trally (Fig. 22) .. hambletoni Drake & Harris, 1946 Sixth male abdominal sternum with posteromedial fringe of black SpInUles) Lessee ses Va er 8 Sixth male abdominal sternum without posteromedial fringe of blackispinulesienee ei hoe a. 9 Seventh male abdominal ster- num with two clusters of black spinules bila Jaczewski, 1928 Seventh male abdominal ster- num without black spinules Oe AAP zeteki Harris & Drake, 1941 Prothoracic and mesothoracic pleura and nota (at least laterally) broadly pruinose, with numerous minute shiny black spots. Ante- rior lobe of pronotum without distinct depressions on each side 94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of midline. Propleuron without longitudinal light streak behind eye. Male clasper, in lateral view, short and robust (Fig. 23) pad eethn cryptophila Hungerford, 1924b Prothoracic and mesothoracic pleura and nota not broadly prui- nose, without numerous, minute, shiny, black spots. Anterior lobe of pronotum with a distinct depression (apodeme) on each side of midline. Propleuron usu- ally with longitudinal light streak behind eye. Male clasper, in lat- eral view, long and slender (Fig. DAY(S) pen eae ete ts BER amoena Uhler, 1894 Acknowledgments I thank Klaus Ruetzler and the other ad- ministrators of the Smithsonian Institu- tion’s Caribbean Coral Reef Ecosystem Project for financing the survey of the halo- philic Hemiptera and Coleoptera in Belize during which time this undescribed species was discovered. I also thank Young T. Sohn for the line drawings; Robin A. Faitoute for SEM photomicrographs and assistance with the fieldwork in 1986 and 1987 and all of the specimens collected in March 1988; Su- sann Braden for SEM photomicrographs; John T. Polhemus for constructively re- viewing the manuscript; and Phyllis M. Spangler for typing the manuscript into a word processor. This is contribution 284, Caribbean Coral Reef Ecosystems (CCRE) Program, Smithsonian Institution. References Cited Andersen, N. M., & J. T. Polhemus. 1980. Four new genera of Mesoveliidae (Hemiptera; Gerromor- pha) and the phylogeny and classification of the family.—Entomologica Scandinavica 11:369- 392. Drake, C.J. 1949. Two new Mesoveliidae with check list of American species (Hemiptera). —Boletin Entomologia Venezolana 7(3—4):145-147. —, & H. M. Harris. 1946. A new mesoveliid from Ecuador (Hemiptera; Mesoveliidae).— Bulletin of the Brooklyn Entomological Society 41(1):8-9. Harris, H. M., & C. J. Drake. 1941. Notes on the family Mesoveliidae (Hemiptera) with descrip- tions of two new species.—Iowa State College Journal of Science 15(2):275-277. Hungerford, H. B. 1924a. A new Mesovelia with some biological notes regarding it (Hemiptera-Meso- veliidae).—The Canadian Entomologist 56:142— 144. 1924b. A second new species of Mesovelia from the Douglas Lake, Michigan region (He- miptera-Mesoveliidae).—Annals of the Ento- mological Society of America 17(4):453-456. 1951. A new Mesovelia from Mexico and Guatemala.—Journal of the Kansas Entomo- logical Society 24(1):32-34. Jaczewski, T. 1928. Mesoveliidae from the state of Parana.— Annales of the Musei Zoologici Po- lonici 7(2—3):75-8 1. 1930. Notes on the American species of the genus Mesovelia Muls. (Heteroptera, Mesove- liidae).— Annales Musei Zoologici Polonici 9(1): 3-15. Polhemus, J. T. 1975. New estuarine and intertidal water-striders from Mexico and Costa Rica (He- miptera: Gerridae: Mesoveliidae). — Pan-Pacific Entomologist 51:243-247. Uhler, P. R. 1884. Order IV. Hemiptera. Pp. 204— 206 in J. S. Kingsley, ed., Standard natural his- tory. S. E. Cassin & Co., Boston, vii + 555 pp. 1894. On the Hemiptera-Heteroptera of the island of Grenada, West Indies.— Proceedings of the Zoological Society of London 1894:167- 224. White, F. B. 1879. List of the Hemiptera collected in the Amazons by Prof. J. W. H. Trail, MA., M.D.., in the years 1873-1875, with descriptions of the new species. — Transactions of the Ento- mological Society of London 267-276. Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 95-99 LUTEOCARCINUS SORDIDUS, NEW GENUS AND SPECIES, FROM MANGROVE SWAMPS IN PENINSULAR MALAYSIA (CRUSTACEA: DECAPODA: BRACHYURA: PILUMNIDAE: RHIZOPINAE) Peter K. L. Ng Abstract.— A new genus and species of mangrove-dwelling pilumnid crab of the subfamily Rhizopinae, Luteocarcinus sordidus, is described from mangrove swamps in western peninsular Malaysia. It can be separated from other littoral mud-dwelling genera in having a distinctively structured third maxilliped. It is the first rhizopine reported from littoral mangrove substrates. Crabs of the Indo-West Pacific pilumnid subfamily Rhizopinae Stimpson, 1858 sen- su Ng (1987) are small, usually mud-dwell- ing crabs that are complex taxonomically. Ng (1987) recognized 20 genera with 74 species in the subfamily. However, several of the species, and some of the genera, were only provisionally classified in the Rhizo- pinae. Almost all occur in sublittoral waters. In the present paper, a new genus and species, Luteocarcinus sordidus, is reported from littoral mangrove areas in western peninsular Malaysia. A diagnosis is provid- ed for the genus and species, and affinities with other taxa discussed. The abbrevia- tions G1 and G2 are used for the male first and second pleopods respectively. Mea- surements are included for the carapace width and length. Type specimens, consist- ing of two males and two females, are de- posited in the Zoological Reference Collec- tion of the Department of Zoology, National University of Singapore. Family Pilumnidae Samouelle, 1819 Subfamily Rhizopinae Stimpson, 1858 Luteocarcinus, new genus Diagnosis. — Anterolateral margin ar- cuate, without distinct teeth, lobes or in- dentations, slightly crested, confluent with slightly divergent posterolateral margins. Posterior margin of epistome distinctly sep- arated into four rectangular lobes by three deep clefts; median lobes small, squarish, outer lobes very broadly rectangular. Third maxilliped broad, almost completely cov- ering buccal cavity when closed, not form- ing any rhomboidal gap; merus large, dis- tinctly wider than ischium, outer anterolateral angle lobiform, rounded, not strongly produced outwards, inner angle produced, tip rounded; ischium quadrate, inner margin gently and evenly rounded; exopod stout, inner margin smooth, with- out sharp subterminal tooth, distal area ex- panded to form broad cristiform lobe, distal flagellum well developed, extending beyond width of merus. Male gonopore sternal, opening from coxa of last ambulatory leg via very narrow channel formed by fused adjacent sternites. Male abdomen with sev- en freely articulating segments; first very narrow, not reaching base of last ambula- tory coxa; second narrow; third trapezoidal, lower margin sinuous, upper margin slightly concave, lateral margins slightly concave; fourth to sixth progressively less trapezoi- dal, more squarish, lateral margins slightly concave to almost straight; seventh distinct- ly triangular, tip rounded, lateral margins slightly convex. Type species. —Luteocarcinus sordidus, new species, designated herein. 96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON G H Fig. 1. Luteocarcinus sordidus holotype male: A, Carapace; B, Face and dorsal border of epistome; C, Sternum; D, Left third maxilliped; E, Right chela; F, Left chela; G, Left cheliped carpus; H, Right cheliped carpus; I, Dactylus, propodus and carpus of left last ambulatory leg. VOLUME 103, NUMBER 1 97 Fig. 2. Luteocarcinus sordidus A-F, holotype male; G, H, paratype male. A, Abdomen; B, Left gonopore (arrow); C, E, G, Left G1 (dorsal view); D, F, H, Left G1 (ventral view). 98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Etymology.—The generic name is de- rived from the Latin “‘luteum”’ for mud; al- luding to the muddy mangrove habitat of the type species. The gender is masculine. Luteocarcinus sordidus, new species Figs. 1, 2 Typhlocarcinus sp. Sasekumar, 1974:63, 65. Thyphlocarcinus sp. Sasekumar 1980:22 (spelling erroneous). Diagnosis. —Carapace transverse, dis- tinctly broader than long; appears slightly trapezoidal, posterolateral margins slightly diverging; surfaces smooth, without gran- ules, almost glabrous except on anterolat- eral margins, which have numerous short, simple hairs; regions poorly defined, central H-shaped depression distinct but neither deep nor prominent. Eye fitting tightly into orbit, cornea distinct, pigmentation restrict- ed to median part. Frontal margin entire, slightly sinuous, deflexed. Chelipeds asym- metrical in larger males, outer surfaces smooth, almost glabrous; inner angle of car- pus with distinct blunt tooth; propodus without distinct projection partly overlap- ping base of dactylus; fingers shorter than palm. Ambulatory legs not distinctly elon- gate, segments smooth, unarmed, covered with scattered hairs, second pair longest, upper margin of median part of dactylus of last pair concave, distal part curves gradu- ally downwards, dactylus appearing sin- uous. G1 very sinuous, slender, distal part strongly tapered, tip slightly curved up- wards; G2 very short, sinuous, without trace of flagellum, distal part strongly dilated forming cup-like structure. Littoral man- grove-dwelling species. Material.—Holotype male (6.7 by 5.0 mm), Sementa mangroves, Selangor, pen- insular Malaysia, leg. K. Sagathevan, 1988; paratype male (4.7 by 3.4 mm), 2 paratype females (5.9 by 4.2 mm, 5.7 by 4.1 mm), mangrove mud, near Batu Pahat, Johore, peninsular Malaysia, leg. P. K. L. Ng, May 1983. Etymology.—The species name is de- rived from the Latin “‘sordida’’ for dirty, alluding to the appearance of the uncleaned animals. Remarks.—The characters of Luteocar- cinus are unusual in their combination, al- though no one character is unique to it. The most apparent difference is the form of the third maxilliped. The outer anterolateral angle of the merus is not distinctly expanded and appears rounded. This character affili- ates it with Typhlocarcinus Stimpson, 1858, and Typhlocarcinops Rathbun, 1909, gen- era that are separated from the closely allied Rhizopa Stimpson, 1858, Ceratoplax Stimpson, 1858, and Rhizopoides Ng, 1987, by having the anterolateral angle of their third maxilliped merus rounded and not produced. In the structure of the exopod of the third maxilliped, however, Luteocarci- nus 1s closer to the condition of Rhizopa, Ceratoplax and Rhizopoides, being stout, with the inner margin smooth. In Typhlo- carcinus and Typhlocarcinops, the exopod is more slender and the inner margin has a distinct subterminal tooth. In 7yphlocar- cinus and Typhlocarcinops, the ischium is distinctly larger and wider than the merus, the proportions of which are reversed for Luteocarcinus, Rhizopa, Ceratoplax and Rhizopoides. A new genus has thus been established to accommodate the unusual suite of characters possessed by Luteocar- cInUS. Sasekumar (1974, 1980) reported an un- identified “‘ 7 yphlocarcinus” species from the mangrove swamps of the Kapar Forest Re- serve and the Port Klang river bed, west of Port Swettenham and east of the Strait of Klang in Selangor, peninsular Malaysia. These specimens had been identified and examined by the late Raoul Seréne (Saseku- mar 1974), but are no longer extant, having been used for experiments (Sasekumar, pers. comm.). K. Sagathevan (pers. comm.), who is studying the ecosystem of the Selangor mangrove swamps, has informed the author that the *“‘7yphlocarcinus”’ fide Seréne is not common and 1s only occasionally seen. Two VOLUME 103, NUMBER 1 of the sites where “7 yphlocarcinus sp.” were collected (Sasekumar 1974) are stream beds in the Kapar Forest Reserve. In one (Station 4), a density of 4.3 individuals per square meter was recorded. In another station on the river bed of Port Klang, a density of five individuals per square meter was reported. Both sites in the Kapar Forest Reserve were located in the higher parts of the mangrove. Sagathevan recently sent the author a col- lection of small mangrove crabs he had ob- tained at a Selangor mangrove swamp. Most were Xenophthalmus pinnotheroides White, 1846. One male, however, proved to be the unknown Selangor “‘7yphlocarcinus.” The Selangor specimen proved to be al- most identical with three unidentified rhi- zopine specimens collected by the present author from a mangrove swamp adjacent to Batu Pahat in Johore, southwestern pen- insular Malaysia, in 1983. The three Johore specimens are smaller than the Selangor male. Their external carapace and append- age features, however, agree very well. The Selangor male is here made the holotype of Luteocarcinus sordidus, the three Johore specimens, paratypes. The G1 of the Johore male, however, does not have the tip dis- tinctly upcurved. This difference is not re- garded as significant since the Johore male is smaller than the holotype. The female specimens appear mature, the pleopods being fully setose, but the abdomen only partially covers the sternum. Uncleaned crabs are covered with a thin layer of “‘pile,”’ resembling pubescence which is easily scraped off. The rounded outer anterolateral angle of the third maxilliped merus and structure of the first male abdominal segment (not reaching the base of the last pair of am- bulatory legs) of Luteocarcinus sordidus almost certainly led Seréne to identify the earlier Selangor specimens as “7 yphlocar- cinus.”’ Although L. sordidus is close to Ty- Dhlocarcinus, the form of its third maxil- liped argues against its inclusion in that genus. 99 The genus 7Jyphlocarcinus was estab- lished for 7. villosus Stimpson, 1858, from Hong Kong. Since then, additions and transfers have resulted in six species being currently recognized in the genus (Ng 1987). All the known Typhlocarcinus species have been described from deeper sublittoral waters, usually in muddy or silty substrates. Luteocarcinus sordidus was obtained during low tide on littoral mangrove mud. The concave dorsal margin of the me- dian part of the last ambulatory dactylus in L. sordidus, which makes it appear sinuous, is useful as a species character if used with other external characters. Ng (1987) has commented that since this character (an up- curved or sinuous last dactylus) is also pres- ent in other genera (and non-rhizopine taxa as well), and is probably associated with burrowing, it is not useful generically. Acknowledgments Thanks are due to Mr. Tan Hong Kim for leading the author through the mangrove swamps of Johore, and for his hospitality and kindness while he was in Batu Pahat. The author is also grateful to Dr. A. Sase- kumar and Mr. K. Sagathevan (both Zool- ogy Department, University of Malaya) for their help. Literature Cited Ng, P. K. L. 1987. The Indo-Pacific Pilumnidae II. A revision of the genus Rhizopa Stimpson, 1858 and the status of the Rhizopinae Stimpson, 1858 (Crustacea: Decapoda: Brachyura).—Indo-Ma- layan Zoology 4(1):69-111, pl. 1. Sasekumar, A. 1974. Distribution of macrofauna on a Malayan mangrove shore. — Journal of Animal Ecology 43:51-69. . 1980. The present state of mangrove ecosys- tems in Southeast Asia and the impact of pol- lution. Malaysia.—South China Sea Fisheries Development and Coordinating Programme, Manila, Philippines, FAO/UNEP 15, SCS/80/ WP/94b:i-vi, 1-80. Department of Zoology, National Uni- versity of Singapore, Kent Ridge Campus, Singapore 0511, Republic of Singapore. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 100-102 ON THE STATUS OF ALPHEUS BARBARA LOCKINGTON (CRUSTACEA: CARIDEA: ALPHEIDAE) Mary K. Wicksten Abstract. — Alpheus barbara, described by Lockington (1878) based ona single specimen from California, has not been reported since its description. The type material has been lost. One of the major distinguishing features between A. barbara and A. clamator, the presence of a spine on the basicerite of the second antenna in the latter species, is inconsistent. Spines on alpheids may be lost, blunt or reduced in old or large individuals. Based on the evidence, A. barbara Lockington, 1878 is considered to be a junior subjective synonym of A. clamator Lockington, 1877. In 1878, W. Lockington described a new species of snapping shrimp, Alpheus bar- bara, from Santa Barbara, California. The description was based on the unique type material, a specimen originally reported by J. Kingsley in 1878. Kingsley, however, re- ported the specimen as Alpheus clamator Lockington, 1876. According to Lockington (1878), Kingsley’s specimen differed from A. clamator by the dissimilar proportions of the articles of the carpus of the second pereopods and the absence of spines on the basicerites of the second antennae and meri of the posterior three pereopods. Neither writer figured the specimen. Both men- tioned that it was damaged, missing the ros- trum and frontal region of the carapace. The type specimen, “in the collections of the Peabody Academy of Science at Salem, Massachusetts” or “the Peabody Museum of Yale College” (Kingsley 1878:189), ap- parently was lost. Neither Holmes (1900) nor Schmitt (1921) reported seeing speci- mens of the species. No additional material of the supposed species has been reported since the species was described. In an attempt to compile an accurate list of decapod crustaceans of southern Califor- nia, I checked records of all species reported in the area. Extensive collections have been made in southern California from the shore- line to the lower continental slopes—the Al- lan Hancock Foundation, University of Southern California, has over 19,000 spec- imens of decapod crustaceans, and large col- lections also exist at Scripps Institution of Oceanography, the California Academy of Sciences and the National Museum of Nat- ural History, Smithsonian Institution. Decapod crustaceans have been collected and studied by environmental agencies, the California Department of Fish and Game, students and researchers from many insti- tutions, park personnel and divers. Despite all of this activity, no further material of A. barbara has been reported in the literature or deposited in collections, nor has it been found off western Mexico. It seems odd that a species in such a well- studied area as southern California would not be found unless it lived in a very in- accessible habitat or occurred in a fresh- water site that had been destroyed by hu- man activity. Although exact collecting data was not provided, Alpheus barbara proba- bly came from an intertidal or shallow sub- tidal rocky area. The collector, W. G. W. Harford, collected other alpheids known to live in such habitats (Kingsley 1878). Al- though there is the remote possibility that the species became extinct, it seems unlike- ly. Most coastal environmental degradation VOLUME 103, NUMBER 1 in southern California has occurred in bays, estuaries and marshlands (Wicksten 1984a). Kingsley originally referred his specimen to A. clamator. This common species is well represented in collections. To determine if A. barbara might be misidentified among such material, I examined all of the speci- mens of A. clamator in the collections of the Allan Hancock Foundation. These 1030 specimens have been taken throughout the entire geographic and bathymetric range of the species, and include individuals of both sexes and a wide range of sizes. When examining the specimens of A. cla- mator, 1 looked for the differences which Lockington (1878) indicated as different be- tween A. barbara and A. clamator: the pro- portions of the articles of the carpus of the second pereopod and absence of spines from the basicerite and meri of the third—fifth pe- reopods. The proportions of the carpal ar- ticles vary slightly and can be difficult to measure exactly without a micrometer. Supposedly, in A. barbara, the first two seg- ments of the carpus of the second pereopod are equal in size, while in A. clamator, the first segment is 1.3 x as long as the second (Kim & Abele 1988). Such a slight difference probably could be easily confused, and is not a useful character for distinguishing be- tween two species. Ofthe 1030 specimens of A. clamator that I examined, a spine was absent from at least one basicerite in 14 specimens. In 10, the spine was absent on the right but present on the left, and in four, it was absent on the left but present on the right. In addition, in three specimens the spine was blunt or re- duced on the right, and in another two it was blunt or reduced on the left. In all, about 2% of individuals of A. clamator lack a spine on the basicerite on at least one side. All individuals had spines on the meri of the posterior three pereopods, but these spines could be blunt in large individuals. Animals that were regenerating appendages often had abnormally slender, spineless legs. At present, three valid species of Alpheus 101 are known from southern California: A. cla- mator, A. bellimanus Lockington and A. californiensis Holmes (Wicksten 1984b). The major chelae of the latter two species differ greatly in the pattern of spines and sulci from those described by Kingsley for his specimen, so it is unlikely that A. bar- bara could have been confused with either species. The distinction between A. barbara and A. clamator was based on three differences. The proportions of the segments of the car- pus of the second pereopod could have been misinterpreted without accurate measuring devices: there is little difference between segments “‘equal’’ and one being 1.3 as long as the other. The spine on the basicerite is missing in a low proportion of the pop- ulation of A. clamator. In a “damaged” specimen, meral spines might be missing from the posterior pereopods. In the account by Lockington (1878) de- scribing A. barbara, there is no evidence that Lockington ever actually examined Kingsley’s specimen. The short description differentiates between A. clamator, A. bel- limanus and A. barbara, but provides no description of the specimen except what was reported by Kingsley. Lockington (p. 472) ended the description by stating, ““As Kings- ley had only an imperfect specimen, and does not describe the rostrum and front, I cannot be sure that this species belongs to this section”’ (of alpheids). Based on the de- scription and the evidence of variability within A. clamator, I suggest that Alpheus barbara Lockington is a junior subjective synonym of 4. clamator Lockington. Snapping shrimp of the genus A/pheus are known to show considerable within-species variation. The major chelae of males and females often differ in size and shape, as do those of juveniles from adults. Widespread species also can have regional variants. (See, for example, the discussion of A. floridanus by Chace 1972:66; and the remarks on vari- ation in the genus A/pheus by Banner & Ban- ner 1982:21.) Such within-species variabil- 102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ity has confused systematists working on the family and has resulted in many species having extensive lists of synonyms. Kim & Abele (1988:21) treated A. bar- bara as a valid species ““because the absence of a lateral spine on the basicerite is a quite distinct and important character.” From my study, it appears that the presence of the spine of the basicerite usually, but not al- ways, is consistent within a single species. I have observed similar variation in spines on the carpal segments of the major chelae of A. armillatus Milne Edwards and A. ca- nalis Kim & Abele— most individuals of the population have distinct spines, but a few very large (and probably old) individuals have small knobs or no spines at all. Anyone attempting to identify or describe alpheids should be cautious about using presence or absence of spines alone as a distinguishing feature between species, especially when ex- amining a large or old individual. Literature Cited Banner, D. M., & A. H. Banner. 1982. The alpheid shrimp of Australia. Part III: The remaining al- pheids, principally the genus A/pheus, and the family Ogyrididae.— Records of the Australian Museum 34:1-357. Chace, F. A., Jr. 1972. The shrimps of the Smith- sonian-Bredin Caribbean expeditions with a summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia).— Smithsonian Contributions to Zoology 98:1-179. Holmes, S. J. 1900. Synopsis of California stalked- eyed Crustacea. — Occasional Papers of the Cal- ifornia Academy of Sciences 7:1-262. Kim, W., & L.G. Abele. 1988. The snapping shrimp genus Alpheus from the eastern Pacific (Decapo- da: Caridea: Alpheidae).— Smithsonian Contri- butions to Zoology 454:1-119. Kingsley, J. S. 1878. A synopsis of the North Amer- ican species of the genus A/pheus.—Bulletin of the U.S. Geological and Geographical Survey of the Territories, U.S. Department of the Interior 4:189-199. Lockington, W. N. 1877. Description of seventeen new species of Crustacea.— Proceedings of the California Academy of Sciences 7:41-48. 1878. Remarks on some new Alphei, with a synopsis of the North American species.— An- nals and Magazine of Natural History ser. 5, 1: 465-480. Schmitt, W. L. 1921. The marine decapod Crustacea of California.— University of California Publi- cations in Zoology 23:1-—470. Wicksten, M. K. 1984a. Early twentieth century rec- ords of marine decapod crustaceans from Los Angeles and Orange counties, California. —Bul- letin of the Southern California Academy of Sci- ences 83(1):12-42. 1984b. New records of snapping shrimps (family Alpheidae) from California. — Proceed- ings of the Biological Society of Washington 97: 186-190. Department of Biology, Texas A&M Uni- versity, College Station, Texas 77843. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 103-105 PSEUDOTHELPHUSA GALLOI, A NEW SPECIES OF FRESHWATER CRAB (CRUSTACEA: BRACHYURA: PSEUDOTHELPHUSIDAE) FROM SOUTHWESTERN MEXICO Fernando Alvarez and Jose Luis Villalobos Abstract. —A new pseudothelphusid crab from the State of Guerrero, Mexico, is described. Its affinities with four other species present in the Mexican Pacific slope are discussed. Since the publication of Rodriguez’ (1982) monograph on the Pseudothelphusidae, two new species belonging to the genus Pseu- dothelphusa have been described (Alvarez 1987, 1989). The third species, described in this paper, comes from the State of Guer- rero in southern Mexico. Pseudothelphusa galloi, new species, lacks a defined marginal process on the gonopod like some of its con- geners from the Pacific slope of Mexico. This new species was collected by J. P. Gallo while studying the feeding habits of the river otter Lutra longicaudis annedectens, which feeds on this and other freshwater crusta- ceans. Types are deposited in the Carcino- logical Collection, Instituto de Biologia, Universidad Nacional Autonoma de Méxi- co (IBUNAM). Pseudothelphusa galloi, new species Fig. 1 Description. —Superior frontal border of carapace formed by small tubercles, divided medially by deep notch posteriorly forming the median groove. In frontal view, front inclined towards central portion. Inferior frontal border well defined, formed by small blunt tubercles, sinuous in frontal view. Dorsal surface of carapace with deep me- dian groove, narrow anteriorly, broader posteriorly. Cervical grooves slightly arched, not reaching anterolateral margin. Carapace regions ill defined, gastric and branchial re- gions elevated; epigastric lobes well marked. Area between epigastric lobes and superior frontal border inclined anteriorly. Antero- lateral margin with 9 denticles between or- bit and cervical groove, with 22 to 27 be- tween cervical groove and epibranchial region. Pterygostomian region bearing setae around third maxillipeds. Ratio ischium/ exopod of third maxilliped 0.4 to 0.57. Che- lipeds unequal, fingers of major chelae gap- ing, granulated and curved. Palm of che- liped with fine granulation, becoming more dense dorsally and ventrally. Gonopod with well developed lateral lobe, covering frontal portion of apical cavity; in cephalic view is roughly triangular, ending in two tips. Superior margin of mesial pro- cess (mesial crest) curving proximally, forming a rounded, axe-shaped lobe. Mar- ginal process reduced to two small bumps, located anteriorly on mesial crest. Apex cavity elongated along a caudo-cephalic axis, bearing 27 setae with terminal pore restrict- ed to lateral part of cavity. Caudal border thick, decreasing in thickness laterally. Material examined. —Mexico. Rio La Parota, small tributary of Pinela River (16°46’N, 99°13’W) Municipio Ayutla de los Libres, Estado de Guerrero; 15 Apr 1985; J. P. Gallo; 1 male holotype, c.b. 65.9 mm, c.l. 39.8 mm; 1 male paratype, c.b. 67.8 mm, c.l. 40.4 mm (IBUNAM-EM-3479). Rio Pinela, Municipio Ayutla de los Libres, Estado de Guerrero, 28 Dec 1983; J. P. Gal- 104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1 Fig. 1. Pseudothelphusa galloi, holotypic male, A—G, left gonopod: A, cephalic view; B, lateral view; C, caudal view; D, mesial view; E, apical view; F, detail of cephalic view; G, detail of mesial view; H, left third maxilliped. VOLUME 103, NUMBER 1 lo; 1 male, c.b. 57.0 mm, c.l. 35.8 mm, 2 females, c.b. 28.0 and 37.3 mm, c.l. 18.2 and 23.6 mm (IBUNAM-EM-3478). Etymology.—This species is named in honor of Juan Pablo Gallo. Remarks. —The species shares with Pseu- dothelphusa jouyi, P. lophophallus and P. sonorae the vestigial character of the mar- ginal process of the gonopod. However, in P. jouyi and P. lophophallus these vestiges appear on the mesial crest as a series of acute denticles (Rodriguez 1982, figs. 94a and 96a) and in P. sonorae there is only a small pro- tuberance with no denticles (Rodriguez 1982, fig. 97c). In general, the gonopod mor- phology of P. galloi is closer to that of P. jouyi. Acknowledgments We thank Juan Pablo Gallo for collecting and donating the type specimens and An- tonio Cantu for the illustrations. The second 105 author was supported by CONACyT Grant PCCNCNA-03 1542. Literature Cited Alvarez, F. 1987. Pseudothelphusa mexicana, a new freshwater crab from the State of Guerrero, Mexico (Brachyura: Pseudothelphusidae).— Proceedings of the Biological Society of Wash- ington 100:1-3. . 1989. Smalleyus tricristatum, new genus, new species, and Pseudothelphusa parabelliana, new species (Brachyura: Pseudothelphusidae) from Los Tuxtlas, Veracruz, Mexico.— Proceedings of the Biological Society of Washington 102:45— 49. Rodriguez, G. 1982. Les Crabes d’eau douce d’Amé- rique. Famille des Pseudothelphusidae. — Faune Tropicale 22:1-223. (FA) Department of Zoology, University of Maryland, College Park, Maryland 20742; (JLV) Coleccion de Carcinologia, Instituto de Biologia, Universidad Nacional Auto- noma de Mexico, Apdo. Postal 70-153, Mexico, D. F. 04510, México. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 106-107 RANGE EXTENSION AND HOST RECORD FOR DISSODACTYLUS USUSFRUCTUS GRIFFITH, 1987 (CRUSTACEA: BRACHYURA: PINNOTHERIDAE) Michel E. Hendrickx Abstract. — Dissodactylus ususfructus Griffith (Pinnotheridae) was found for the first time in the Gulf of California, Mexico, about 19° of latitude north of its previous northernmost record. Among the five specimens collected, two males and one female were found associated with Clypeaster speciosus Verrill. During sampling activities in 1982 and 1985 in the Gulf of California, Mexico, aboard the R/V E/ Puma of the Universidad Nacional Autonoma de Mexico (UNAM; CORTES Cruises), a small series of speci- mens of Dissodactylus Smith (Crustacea: Pinnotheridae) was found among inverte- brates and on freshly captured specimens of Clypeaster (Echinoidea: Clypeasteridae). Later examination of these specimens dem- onstrated that they belonged to an unde- scribed species of Dissodactylus and repre- sented the first positive record of the genus on Clypeaster for the Pacific. A preliminary description of this species of pinnotherid was prepared at that time (Hendrickx 1987), but was never published. The genus Dissodactylus was recently re- viewed by Griffith (1987a) who described two new species for the Pacific coast of America (D. schmitti and D. ususfructus). On reading Griffith’s paper, it became ev- ident that the material collected in the Gulf of California belonged to D. ususfructus, a species already recognized as new by the late S. A. Glassell in the 1930s in a manuscript that he never published (see Griffith 1987a: 402). This species is known only from three localities between Costa Rica and Ecuador, and has not yet had a host species positively identified. The discovery of D. ususfructus in the Gulf of California provides new information on its distribution and ecology. All the speci- mens reported herein are held in the refer- ence collection of the Estacion Mazatlan, UNAM (EMU). Dissodactylus ususfructus Griffith, 1987 Dissodactylus ususfructus Griffith, 1987a: 401, figs. 3, 8K, 10B, 12D, 141; 1987b: figs. 7C, 9C, 13B, 17D. Material examined. —CORTES 1 Cruise, station 19, 6-V-1982, 28°09'30’N, 112°46'30”W, off Cabo San Miguel, Baja California, Mexico, trawling at 30-35 m, sand, 2 6¢c.w. 6.7 and 7.6 mm, | 2 c.w. 6.2 mm (EMU-2635).—CORTES 2 Cruise, station 49B, 19-III-1985, 26°59’N, 111°53'30"W, off Bahia Santa Inés, Baja California, Mexico, trawling at 68 m, 1 ¢ c.w. 5.5 mm (EMU-2636A).—CORTES 2 Cruise, station 50, 20-III-1985, 25°46'N, 109°35'W, off Rio Fuerte, Sinaloa, Mexico, trawling at 96-98 m, muddy sand, | 2 c.w. 7.6 mm (EMU-2636B). Previous records. —SSE of Judas Point, Costa Rica (Zaca station 214; type locality), off Santa Elena Bay, Ecuador, and SW of Secas Islands, Panama (Griffith 1987a). Remarks.—The present records extend the known distribution of D. ususfructus northward about 19° of latitude, to Cabo San Miguel and to off Rio Fuerte, respec- tively, on the west and on the east coasts of the Gulf of California. The positive identification of Clypeaster VOLUME 103, NUMBER 1 speciosus Verrill, 1870 as a host of D. usus- fructus (CORTES 1 Cruise, station 19) part- ly confirms the hypothesis of Griffith (198 7a: 403), in that this pinnotherid is associated with Clypeaster. The possible association of D. ususfructus with another species of Clyp- easter, C. europacificus H. L. Clark, 1944, was suggested by Griffith (1987a) because of the presence of this species of irregular echinoid in successive samples taken by Zaca at station 214, the type locality of D. ususfructus, but this had not been con- firmed. According to Caso (1980, 1986), both Clypeaster speciosus and C. europaci- ficus are commonly found throughout the Gulf of California in similar habitats (shal- low water to 90 m for C. speciosus, 18 to 165 m for C. europacificus; mostly on sand). They have also been collected at least once in the same trawl (Velero III, station 699- 37, Canal Angeles, Gulf of California) (Caso 1980:9, 24), which suggests that they are, at least occasionally, sympatric. The hypo- thetical association of D. ususfructus with two species of Clypeaster does not seem un- likely. Indeed, many species of Dissodac- tylus are known to occur on several species or even genera of irregular echinoids (Grif- fith 1987a, Jangoux 1987). The bathymetry provided by Griffith (1987a) for D. ususfructus is rather impre- cise (80-120 m). The present material was found between 30-35 m, at 68 m, and be- tween 96-98 m, on sandy bottom (62 to 100% sand). Other environmental condi- tions at bottom level were as follows: water temperature, 13.2 to 14.2°C; dissolved oxy- gen, 1.33 to 3.5 ml O,/1. Acknowledgments I am grateful to Dr. Maria Elena Caso, dean of the Instituto de Ciencias del Mar y Limnologia, UNAM, who called my atten- 107 tion to the capture of specimens of Disso- dactylus ususfructus during the CORTES Cruises. I also thank Drs. J. Bouillon and M. Jangoux for the facilities provided dur- ing my stay at the Laboratoire de Biologie Marine, Université Libre de Bruxelles, and Mercedes Cordero H. for aid in preparing and typing the manuscript. This study was partly supported by CONACyT (ICECXNA-021926) and the DGPA, UNAM. This is Contribution 602 of the Instituto de Ciencias del Mar y Limnologia. Literature Cited Caso, M. E. 1980. Los equinoideos del Pacifico de México. Parte Tercera. Orden Clypeasteroi- da.— Publicacion Especial, Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autonoma de México 4:1-252. 1986. Los equinodermos del Golfo de Cali- fornia colectados en las Campanas SIPCO I-II- III a bordo del B/O “El Puma.”—Anales del Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autonoma de México 13 (1):91-184. Griffith, H. 1987a. Taxonomy of the genus Disso- dactylus (Crustacea: Brachyura: Pinnotheridae) with descriptions of three new species. — Bulle- tin of Marine Science 40(3):397-422. 1987b. Phylogenetic relationships and evo- lution in the genus Dissodactylus Smith, 1870 (Crustacea: Brachyura: Pinnotheridae).—Ca- nadian Journal of Zoology 65:2292-2310. Hendrickx, M. E. 1987. La faune de crustacés (Sto- matopoda et Decapoda) du Golfe de Californie: Taxonomie, aspect écologique et distribution. Thése de Doctorat, Université Libre de Bru- xelles. Vol. I, i-xvii, 1-741; Vol. II, tables 1- 210, figs. 1-428. Jangoux, M. 1987. Diseases of Echinodermata. III. Agents metazoans (Annelida to Pisces).—Dis- eases of Aquatic Organisms 3(1):59-83. Instituto de Ciencias del Mar y Limnolo- gia, Estacion Mazatlan, UNAM, Apdo. Postal 811, Mazatlan, Sinaloa 82000, Mex- ico. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 108-109 AN EASTERN UNITED STATES RECORD FOR THE WESTERN INDO-PACIFIC CRAB, HEMIGRAPSUS SANGUINEUS (CRUSTACEA: DECAPODA: GRAPSIDAE) Austin B. Williams and John J. McDermott Abstract. — An ovigerous female of a western Indo-Pacific grapsid crab, Hemi- grapsus sanguineus, is reported from New Jersey, U.S.A. During a field trip on 24 Sep 1988, a stu- dent member (R. A. Nusbickel) of an in- vertebrate biology course at Franklin and Marshall College, Lancaster, Pennsylvania, discovered a live ovigerous grapsid crab at Townsends Inlet, Cape May County, New Jersey (39°07'06’N, 74°43’00”W). The crab was taken during an ebbing tide from a Fu- cus covered boulder in the mid-intertidal zone, under the south end of the highway bridge over the inlet. There was slight, but not recent, damage to the crab’s left frontal margin. The crab was transported to Lan- caster on the same day, where it was main- tained at room temperature (ca. 23°) in a one gallon aquarium provided with rocks, Fucus, and shallow (ca. 2.5 cm) sea water (ca. 30 ppt). Water was changed daily, but the crab was not fed (however, it may have eaten some of the algae). On 27 Sep some of the embryos were re- moved for microscopical examination. They were in the eyed stage (eye pigment 0.057 mm long X 0.029 mm wide), had consid- erable body pigment, a beating heart, and the yolk was ca. '4—'4 of the embryo volume. The eggs were nearly round—mean diam- eter 0.363 mm, SD 0.015, n = 20. On 3 Oct the crab was discovered out of the water and oscillating its abdomen. It subsequently entered the water (as it usually did when JJMcD was about to change the water), where it released some embryos, aborted some “‘prezoeae”’ (no dorsal spines), and released one zoea having a dorsal spine. At this time the eyes were larger than on 27 Sep (0.114 mm long x 0.057 mm wide), and the yolk was reduced to ca. 4 of the embryo volume. Released embryos and zoeae were preserved in glycerine alcohol as were some from 4 Oct. On the morning of 6 Oct the crab was found dead and was preserved. The crab and larvae are now deposited in the crus- tacean collection of the National Museum of Natural History, Smithsonian Institution (USNM 239154). The crab was identified as Hemigrapsus sanguineus (de Haan, 1853) by reference to Sakai (1976) and by comparison with spec- imens in the USNM. The first zoea fits the description of the first zoea of that species (Kurata 1968) and resembles figures of that developmental stage redrafted by Rice (1980). The carapace of the adult was distinctly patterned or mottled, there were small red- dish rounded spots on the upper parts of the chelipeds, and the walking legs were band- ed. The following morphometric data were obtained with a dial caliper to the nearest 0.05 mm: carapace width 35.8, length 30.7; width between outer orbital teeth 24.95; ab- domen width 4th segment 26.2, 5th 25.85, 6th 23.6, telson 14.1; right chela length 17.6, height 9.45; left chela length 17.65, height 9.35. Hemigrapsus sanguineus is listed by Sa- kai (1976) as one of the commonest crabs of Japan, having a western Pacific distri- VOLUME 103, NUMBER 1 109 Fig. 1. bution extending from Sakhalin, Korea, and north China to Hong Kong, and on all the coasts of Japan from Hokkaido to Okinawa. Occurrence of an ovigerous female in New Jersey may be one of the interesting but potentially distressing introductions that re- sult from world shipping (Carlton 1979). If insemination had taken place before the crab was transported, it may have been a lone immigrant, but if not, at least one male of the species would have been introduced into New Jersey waters with it. The Cape May area should be monitored to see if a pop- ulation has become established. No other records of this species in the United States are known to us. Acknowledgment We thank J. L. Richardson for reviewing the manuscript. Literature Cited Carlton, J.T. 1979. Introduced invertebrates of San Francisco Bay. Pp. 427-444 in T. J. Conomos, Hemigrapsus sanguineus: mature female collected in New Jersey, U.S.A., dorsal view. ed., San Francisco Bay the urbanized estuary. Pacific Division, American Association for the Advancement of Science, San Francisco, Cali- fornia, 493 pp. Kurata, H. 1968. Larvae of decapod Brachyura of Arasaki, Sagami Bay. Il. Hemigrapsus sangui- neus (de Haan) (Grapsidae).—Bulletin of the Tokai Regional Fisheries Research Laboratory, No. 56:161-165. Rice, A. L. 1980. Crab zoeal morphology and its bearing on the classification of the Brachyura. — Transactions of the Zoological Society of Lon- don 35:271-424. Sakai, T. 1976. Crabs of Japan and the adjacent seas. — Kodansha Ltd., Tokyo, 773 pp. (English text), 251 plates (many colored), 461 pp. (Japanese text), as 3 separate volumes. (ABW) National Marine Fisheries Ser- vice-NOAA Systematics Laboratory, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560; (JJMcD) Department of Biology, Franklin and Marshall College, Lancaster, Pennsy]l- vania 17604-3003. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 110-119 A REDESCRIPTION OF TIRON ANTARCTICUS K. H. BARNARD, 1932 (CRUSTACEA: AMPHIPODA: SYNOPIIDAE) WITH AN UPDATED KEY TO THE SPECIES OF TIRON LILJEBORG, 1865 Krzysztof Jazdzewski Abstract. — Tiron antarcticus K. H. Barnard (Amphipoda, Synopiidae) is re- described; drawings of the appendages of the type specimen are presented. A new key for 7iron Liljeborg is proposed. The description of 7iron antarcticus by K. H. Barnard (1932) is rather superficial and, with its single figure presenting pereo- pod 7, cannot satisfy the needs of amphipod taxonomists. This description was sufficient to regard the species as an undoubted mem- ber of the genus 7iron Liljeborg, but J. L. Barnard (1972) in his survey of the family was unable to put 7. antarcticus in the key to the species of Tiron owing to this inad- equate description. The present redescrip- tion based on the type material aims at fill- ing this gap. Material and Methods The type material, the only available for the species, consists of two female speci- mens in the collection of the British Mu- seum. They were kindly loaned to the au- thor by this institution. Appendages of the right side of the holotype have been dis- sected and mounted on slides in polyvinyl lactophenol stained with lignin pink. Spec- imens and slides are deposited in the British Museum (Natural History) of London. Drawings of the appendages were done from temporary glycerol mounts prior to mount- ing in permanent slides that were used in final study of some details. Holotype.—? 7.5 mm with bristled oos- tegites, coll. R.R.S. William Scoresby, St. WS 33, South Georgia, 21 Dec 1926, 54°59'S, 35°24'W; tow-net (horizontal haul 130 m, bottom 135 m). Fourteen amphipod species in the sample. Paratype. —?? immature 6.5 mm (neither oostegites nor penes traceable), coll. R.R.S. Discovery, Sta. 175, Bransfield Strait (South Shetlands), 2 Mar 1927, 63°17'S, 59°48'W; large dredge, 200 m, night. Thirty-three am- phipod species in the sample. The only oth- er species in both these two samples was Epimeria excisipes K. H. Barnard. Tiron antarcticus K. H. Barnard Figs. 1-5 Tiron antarcticus K. H. Barnard, 1932:148- 149, fig. 86. Body slender; head galeate and keeled, rostrum and lateral cephalic lobes sharp. Eyes present, rounded, dorsally appressed. Accessory eye not visible, possibly due to the long preservation of the material but note that accessory eyes were not mentioned by K. H. Barnard (1932: “‘lower eyes not traceable’’). Last four (type) or three (paratype) tho- racic segments as well as pleon segments excepting the last one, keeled; last thoracic segment and the said five pleon segments produced dorsally as incised teeth, the last two being the largest. Antennae rather slender, comparatively VOLUME 103, NUMBER 1 111 E Fig. 1. Tiron antarcticus, female, holotype: A, Outline of body, thoracic and abdominal appendages omitted; a, Tooth of urosomite 2, dorsal view; B, Antenna 1; C, Distal part of antenna | primary flagellum; D, Antenna 2; E, Distal part of antenna 2 flagellum. 112 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Tiron antarcticus, female, holotype: A, Mandible (place of insertion of mandibular palp indicated by dotted circle); a, Lacinia mobilis; a’, Tip of spine of mandibular spine row; B, Mandibular palp; b, Seta of mandibular palp; C, Pars molaris of mandible; c, Edge of pars molaris; D, Outline of maxilla 1; E, Tip of maxillary palp; F, Tip of outer lobe of maxilla 1; G, Tip of inner lobe on maxilla 1. VOLUME 103, NUMBER 1 short. Antenna | as long as about one-fourth of body length, antenna 2 as long as about one-third of body. Antenna 1 very weakly setose. Lengths of peduncular articles of an- tenna | in proportions of 14:11:9. Accessory flagellum long, 4-articulate; primary flagel- lum with 10 articles, several bearing long, slender aesthetascs (6 in type specimen). Antenna 2 with elongate articles 4 and 5, their proportion as 3:2. Flagellum with 12 articles (in K. H. Barnard’s description 10- 11 articles) some of which distally armed with long, slender spines. Mandible with columnar, triturative mo- lar. Edge of molar provided with three rows of variously shaped teeth. Incisor with four teeth, lacinia mobilis with three teeth fol- lowed by row of about eight distally dentate parallel sided spines. Mandibular palp in- serted at level of molar, 3-articulated; article 2 three times as long as article 1 and nearly four times as long as article 3; articles 2 and 3 armed with five and three long, delicately feathered setae, respectively. Lower lip (not dissected) with well de- veloped fleshy inner lobes, outer lobes widely separated. Maxilla 1 with 2-articulate palp ending with marginal row of seven heavy, dentate spines and submarginal row of six curved setae. Inner plate with marginal row of 11 feathered setae, outer plate with row of nine apically dentate spines. Maxilla 2: inner plate with medial, sub- marginal row of feathered setae and one more or less parallel marginal row of such setae; apically these rows ending with some curved, smooth setae and other apically forked and delicately serrated on one side. Outer plate apically with double row of mostly smooth and occasionally curved se- tae; some feathered setae bordering these rows from inner and outer sides. Maxilliped with 5-articulate palp. Inner and outer plates apically with several char- acteristic robust, falcate spines. Coxae 1-4 suboval to trapezoidal; coxa 4 dominated by coxa 3; coxae 5 and 6 divided by excavation into anterior and posterior 113 lobes, coxa 7 small, pear-shaped. Coxae 1-— 3 with ventral margins rather richly setose, remaining coxae only with several short set- ules. Pereopods | and 2 (gnathopods) simple. Article 6 (propus) narrow and elongate with posterior margin armed by several stout, feathered setae. Article 5 (carpus) especially long, nearly twice as long as article 6, dense- ly setose on posterior margin but nearly na- ked anteriorly. Posterior margin of propus with row of spiny spines. Article 2 (basis) long and narrow, rather densely setose on both margins. Pereopods 3 and 4 slender, weakly setose, with articles 4, 5 and 6 (merus, carpus and propus) armed posteriorly with some spines. Pereopods 5—7 subequal in length, but pereopod 6 is longest, these appendages shiny, especially their articles 4, 5 and 6. Articles 2 and 3 of pereopod 6 rather dense- ly setose anteriorly. A distinct increase in the size (width) of articles 2 (basis) and 4 (merus) observed in the sequence from pe- reopod 5 to pereopod 7; these articles in pereopod 7 with large, overhanging postero- distal lobes. Article 2 (basis) in pereopod 7 posteriorly crenulate with short setules; ar- ticle 4 (merus) richly armed posteriorly with several groups of spines. Dactyls of all pereopods ordinary, elon- gate, with subapical spine and short wire seta. Six pairs of subovate coxal gills (II-VII pereon segments) and four pairs of ooste- gites (II-V). Epimera 1-3 on their posterior margin with increasing number of notches with short setules. Surface of epimera near lower and posterior margins with delicate pilosity. Such pilosity also observed in some other places like outer margin of telson, distal parts of pereopods, mouthparts, etc. Postero-infe- rior angles of epimera distinctly pointed, this being not exactly consistent with orig- inal description of K. H. Barnard (1932: ‘“‘with a very slightly produced point’). Pleopods normally developed with mul- tiarticulated rami fringed with long feath- 114 ered setae. Retinaculum composed of two hooked spines and robust feathered seta. Uropod | distinctly longer than uropod 2; in both uropods peduncle and rami armed with spines; exopodites shorter than endo- podites by 4 to '. Uropod 3 with subequal lanceolate rami twice as long as peduncle. Exopodite 1-ar- ticulate, armed mainly with spines; inner margin with some short, feathered setae. Outer margin of endopodite naked, inner margin proximally with long feathered setae (majority of these setae broken in the type specimen but regular row of long feathered setae present in paratype), distally with spines. : Telson fully cleft, long, as long as two last pleon segments and nearly reaching apex of third uropods. Each lobe of telson with row of eight small spines, last one inserted api- cally in notch. Some few small setules pres- ent on upper surface. Discussion The nomenclature of Tiron species is dis- ordered. The gender of this genus is mas- culine, the generic name coming from Greek “reapwv (Liljeborg 1865). Therefore, the proper endings of adjectives are ““-us” and not “-um” or “-a.” According to the Inter- national Code of Zoological Nomenclature (1961: art. 30 on the agreement in gender) one should amend the following names of species of Ziron: biocellata to biocellatus intermedia to intermedius spiniferum to spiniferus. Fig. 3. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON One feature that is considered to be of taxonomic importance was wrongly de- scribed for 7. antarcticus by K. H. Barnard (1932) and then repeated in the diagnosis of this species by J. L. Barnard (1972), namely, the dorsal telson armament. It was said to consist of 4-5 “‘setules” whereas in fact on the upper surface of telson in T. antarcticus holotype there are seven small spines (excepting the apical eighth one) ar- ranged in a somewhat irregular row. According to J. L. Barnard (1972) T. ant- arcticus resembles T. biocellatus, which, on the other hand, has mouthparts very similar to T. tropakis. The features of T. antarcticus distinguishing it most clearly from T. bio- cellatus are the setose second article of the mandibular palp, the lack of dorsal crenu- lation of pleonites, and the armament of telson (only weak dorsal setules in T. bio- cellatus). In my opinion, the species most similar to 7. antarcticus is T. spiniferus; it follows both from the description and from the figures by G. O. Sars (1895). Good dis- criminating characters are here the pleonite serration in 7. spiniferus lacking in T. ant- arcticus, and smooth hind margin of pereo- pod 7 basis in 7. spiniferus versus crenu- lated and setulose hind margin of this article in 7. antarcticus. I follow the opinion of Just (1981) that the lack of mandibular palp is not a suff- cient reason to create a new genus Metatiron Rabindranath (Rabindranath 1972, Ledoy- er 1979). Just’s survey table of features of eight stubby-legged Tiron species clearly shows that one cannot find any other im- portant character to be shared by Tiron species lacking mandibular palp. ~ Tiron antarcticus, female, holotype: A, Maxilla 2; B, Maxilliped; C, Pereopod 1 (=gnathopod 1); D, Pereopod 2 (=gnathopod 2); E, Propus and dactylus of pereopod 2; e, Spine on propus of pereopod 2. Fig. 4. Tiron antarcticus, female, holotype: A, pereopod 3 with oostegite; B, Pereopod 4 with coxal gill and oostegite; b, Dactylus of pereopod 4; C, Pereopod 5; D, Pereopod 6; E, Pereopod 7; e, Dactylus of pereopod 7. P. 116. Fig. 5. Tiron antarcticus, female, holotype: A, Retinaculum of pleopod 2; B, Uropod 1; C, Uropod 2; D, Uropod 3; E-G, Epimera 1-3; F, Posterodistal part of epimeron 2; H, Telson (half). P. 117. 115 VOLUME 103, NUMBER 1 ae ee) NER ee SRE PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 116 117 VOLUME 103, NUMBER 1 118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON On the other hand the diagnosis of Tiron by J. L. Barnard (1972, p. 83) should be amended by changing the information: ““mandible with palp” to ““mandible with or without palp,” since 5 of 12 known species (T. bellairsi, T. brevidactylus, T. caecus, T. triocellatus and T. tropakis) lack mandib- ular palp. I propose also two other small amenda- tions to J. L. Barnard’s diagnosis of Tiron (p. 83): 1) to add “usually” in the note that “hands (propus—K. J.) elongate, linear, lacking dis- tinct spines” — since distinct spines are pres- ent in T. brevidactylus (see Rabindranath 1972); 2) to add “‘or overreaching” in the note: “uropod | reaching apex of uropod 2”— since this is the case at least in 7. antarc- ticus, T. bellairsi, T. brevidactylus and T. spiniferus. Key to the Genus Tiron The key presented below is based on the literature data and mainly on the key by J. L. Barnard (1972) and the above mentioned table of Just (1981). The key is updated in the sense that four species are added— namely T. caecus Ledoyer, 1979, T. bellairsi Just, 1981, T. triocellatus Goeke, 1982 and the herein redescribed T. antarcticus K. H. Barnard. I have avoided using the feature of accessory eye since I believe that it can be overlooked in long preserved material due to fading. The key presented here will surely need further improvements since for many species we lack data on their mor- phological variability and sexual dimor- phism. One should mention, for instance, that even such a seemingly conservative fea- ture as the condition of the inner lobe of maxilla 1 can be unexpectedly sexually di- morphic as in 7. brevidactylus where fe- males have four setae and males none (Ra- bindranath 1972). 1. Pereopods 3-7 with stubby dactyls 2 Pereopods 3—7 with ordinary, claw- shapedudactyls™. 5 545 ao eee 10 . Mandibular palp present ....... 3 Mandibular palp absent ........ 4 . Rami of uropod 3 pointed; outer plate of maxilliped normal, ovate 8 Rami of uropod 3 truncate; outer plate of maxilliped with apical ex- cavation guarded by falcate wings TRUER be Se ee T. thompsoni . Telson with at least several sub- apicalispines. -. 42.1 hae eee 5 Telson lacking subapical spines 6 . Posterior margin of basis of pe- reopod 7 with setules; accessory flagellum with five articles ...... Uh PO Roles ad Medel T. tropakis Posterior margin of basis of pe- reopod 7 smooth, without setules; accessory flagellum with two arti- cles Lei. jet gt tees eee T. caecus . Palp of maxilla 1 slender, much shorter than outer plate; apical spines of uropod 3 exopodite long, nearly one-half of this article length ey Err er enc 5 a6 T. bellairsi Palp of maxilla 1 stout, longer than outer plate; apical spines of uropod 3 exopodite short, many times shorter than this article length .. 7 . Maxillary palp armed with spines and setae at tip only; basis of pe- reopod 7 lacking special double row of facial setules ..... T. brevidactylus Maxillary palp armed with setules along major part of its outer mar- gin; basis of pereopod 7 with spe- cial double row of facial setules . ED tN xen N,N T. triocellatus . Telsonic medial spine rows absent re Tn ee eo be T. altifrons Telsonic medial spine rows pres- eNite. ha ck aks See Oe 9 . Basis of pereopods 6 and 7 with posterior submarginal row of long, OUTOROSS SHES so0c00- T. intermedius Basis of pereopods 6 and 7 lacking special row of setae ..... T. australis VOLUME 103, NUMBER 1 10. Posterior margin of basis of pe- reopod 7 crenulate and setulose 11 — Posterior margin of basis of pe- reopod 7 smooth T. spiniferus 11. Telsonic lobes with row of small spines; article 2 of mandibular palp with several long setae .. 7. antarcticus — Telsonic lobes with at most one subapical spine; article 2 of man- dibular palp lacking setae T. biocellatus Acknowledgments I am greatly indebted to the staff of the Department of Zoology, British Museum (Natural History), London, and namely to Dr. R. Lincoln, Mr. S. H. Halsey, Mrs. J. Ellis and Mrs. A. Gurney, first for entrusting me with the study of the type specimens of Tiron antarcticus and second for their pa- tience in vast but courteous correspondence caused by my too long keeping of the ma- terial. Thanks are due to Dr. J. L. Barnard, Smithsonian Institution, Washington, D.C., for his precious critical comments to this text and for his friendly help. Literature Cited Barnard, J. L. 1972. A review of the family Syno- piidae (= Tironidae), mainly distributed in the 119 deep sea (Crustacea: Amphipoda).—Smithson- ian Contributions to Zoology 124:1-94. Barnard, K. H. 1932. Amphipoda.—Discovery Re- ports 5:1-326. Goeke, G. D. 1982. Tiron triocellatus, a new species of amphipod (Gammaridea: Synopiidae) from the western Atlantic and Gulf of Mexico. —Jour- nal of Crustacean Biology 2:148-153. International Code of Zoological Nomenclature. 1961. International Trust for Zoological Nomencla- ture, London, 176 pp. Just, J. 1981. Tiron bellairsi sp. n. (Amphipoda, Syn- opiidae) from coral sand in Barbados with notes on behaviour.— Zoologica Scripta 10:259-263. Ledoyer, M. 1979. Les gammariens de la pente ex- terne du Grand Récif de Tuléar (Madagascar) (Crustacea Amphipoda).— Memorie del Museo Civico di Storia Naturale di Verona (2)2:1—150. Liljeborg, W. 1865. Bidrag till kinnedomen om un- derfamiljen Dysianassina inom underordningen Amphipoda bland kraftdjuren.— Upsala Univ- ersitets Arsskrift 1865, Upsala, Edquist and Ber- glund, 25 pp. Rabindranath, P. 1972. Three species of Gammari- dean Amphipoda (Crustacea) from the Trivan- drum Coast, India.— Zoologischer Anzeiger 188: 84-97. Sars, G. O. 1895. An account of the Crustacea of Norway. Volume 1. Amphipoda. Cammermey- ers, Christiania, 711 pp. Department of Invertebrate Zoology and Hydrobiology, University of Lodz, 12/16 Banacha s-t, 90-237 Lodz, Poland. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 120-126 ENSAYARA JUMANE, A NEW SPECIES FROM BELIZE, CARIBBEAN SEA (AMPHIPODA, LYSIANASSIDAE) J. L. Barnard and James Darwin Thomas Abstract. — Ensayara jumane is described from Belize. It differs from the west Mexican Ensayara ramonella J. L. Barnard (1964) in the excavate postero- ventral margins of article 2 on pereopods 5-7, and the sparse (9 versus 25) setae on articles 5-6 of gnathopod 1. A new diagnosis for Ensayara, a key to species, list of species, references to original descriptions and geographic codes for each species are provided. This is the first record of the genus from the Caribbean Sea. We present the genus Ensayara in the form to appear in Barnard & Karaman (1990), with a key to species, list of species, references to original descriptions and geo- graphic distribution codes from Barnard & Barnard (1983) for each species. Ensayara J. L. Barnard Ensayara J. L. Barnard, 1964:79 (Ensayara ramonella J. L. Barnard, 1964, original designation). Diagnosis. —Flagella of antennae short. Mouthparts forming quadrate bundle. La- brum and epistome each produced sepa- rately from the prebuccal complex, epi- stome blunt and weakly dominant in projection. Incisor weakly toothed at op- posite corners; molar simple, large, palp at- tached opposite molar. Inner plate of max- illa 1 poorly setose (0-1); palp biarticulate, large. Inner and outer plates of maxilliped well-developed, palp strongly exceeding outer plate, dactyl well-developed. Coxa | slightly shortened and partly cov- ered by coxa 2, scarcely tapering. Gnathopod | short, simple, article 6 long- er than 5, dactyl small, gnathopod 2 mi- nutely chelate, article 6 slightly shorter than article 5. Pereopod 3 strongly prehensile, parach- elate, hand very broad, palm crenulate or spinose. Inner ramus of uropod 2 without notch. Uropod 3 short, peduncle elongate, inner ramus slightly shortened, outer ramus 2-ar- ticulate. Telson short, entire. Variables. —Peduncle of antenna 1 with three distinct articles (Jara); coxa 1 large and ordinary (dentaria; dactyl of pereopod 7 slender and even (ramonella); stunted (car- pinei). Remarks. —Better microscopy has re- solved article 3 of antenna | which is small and mostly obsolescent medially, but has 1.5 rows of facial aesthetascs present. Relationship. —Like Endevoura in the en- larged prehensile pereopod 3 but dactyl of maxilliped unguiform, not bulbous. Species. —angustipes Ledoyer, 1978, 1986 [693]; carpinei Bellan-Santini, 1974 [340B]; dentaria Hirayama, 1985 [391]; iara Lowry & Stoddart, 1983 [776s]; jumane Barnard & Thomas, new species, herein [471]; mi- crophthalma Ledoyer, 1986 [698]; ramo- nella J. L. Barnard, 1964 [376]. Marine, cosmopolitan in low latitudes, 1— 1900 m, 7 species. Key to the Species of Ensayara 1. Article 2 of gnathopod 1 toothed, mandibular palp article 3 with 7 Desetae? ss -.uionitie te bal ieee dentara — Article 2 of gnathopod 1 smooth, VOLUME 103, NUMBER 1 mandibular palp article 3 with 1-2 IDESELACIE A. ces ee Am Pe AM 2 2. Inner rami of uropods 1-2 with 1 Spimexeachy Sy ea Roel TMT) oe 3 — Inner rami of uropods 1-2 without SPITSSWES sya ees ee ene UA LES, 4 3. Mandibular palp article 3 with 2 D-setae, inner ramus of uropod 3 reaching apex of article 1 on outer ramus, dactyl of pereopod 7 thick REE SPR Eat mente Airs Ae Sat 9 BE carpinel — Mandibular palp article 3 with 1 D-seta, inner ramus of uropod 3 not reaching apex of article 1 on outer ramus, dactyl of pereopod 7 ordi- TV AIayApeMe ee KU RCRI EEE LENAG S682, Aa) 4. Lateral cephalic lobe very sharp, coxae 2—3 with dense, short setae atl, ised I) SRE De ea Soe microphthalma — Lateral cephalic lobe blunt, coxae 2-3 with sparse, tiny setae ....... 5 5. Article 2 of pereopod 7 excavate . 2 a ALPERT EER SNe jJumane n. sp. — Article 2 of pereopod 7 not excavate 6. Carpus of pereopod 3 slender (L x W = 18:6), article 3 of antenna | free angustipes — Carpus of pereopod 3 stout (L x W = 18:11), article 3 of antenna 1| tel- escoped into article 2 ramonella Ensayara jumane, new species Figs. 1-3 Etymology.—Named for a tribe of Uto- Aztecan Indians from Central America, name a noun in apposition. Diagnosis. — Lateral cephalic lobe stubby, not subacute; eyes large, deeply pigmented; flagellum of antenna 2 not stubby; article 3 of mandibular palp with only 2 inner (C) setae; palp of maxilla | uniarticulate; dactyl of maxilliped with strong nail; coxae 1-4 with 0-1 ventral seta or sparse tiny setules, no dense setal clusters; articles 4—5 of pereo- pod 3 elongate, article 4 expanded (not lin- 121 ear); article 2 of pereopods 5-7 with pos- terior setule notches, of pereopods 6-7 posteroventrally excavate; inner ramus of uropods 1-2 naked. Description. — Lateral cephalic lobes lack- ing cavity below for insertion of antenna 2; eyes black, one row of clear ommatidia ex- posed peripherally. Antennae 1-2 very short, reaching equally, article 3 of antenna | ob- solescent medially, armed with 1.5 rows of aesthetascs in callynophore, primary flagel- lum with 5 articles, accessory flagellum with 3 articles. Gland cone weak, flagellum of antenna 2 with 5 articles. Prebuccal mass weakly humped anterior- ly. Incisors smooth in middle, convex, la- ciniae mobiles absent, 2 rakers present, mo- lars massive, subconical with weak spinoserrate distomedial margin, otherwise non-triturative; palp article 1 scarcely elon- gate, article 2 with 2 apicolateral setae, article 3 with 1-2 inner setae (probably C-type), apex obliquely truncate, with 4 E-setae. Lower lip with fused inner lobes forming broad truncate line, outer plates widely spread, lacking cones, mandibular lobes large. Inner plate of maxilla 1 large, sub- conical, naked; outer plate with 7 weakly serrate spines, medial margin with thick se- tules, palp l-articulate, apex with 3-4 thick and one thin spines. Inner plate of maxilla 2 very short, with 2 apical setae, outer plate more extended but small, with 5 apical setae and 2 basomedial setae on dorsal face. Inner plates of maxillipeds long, slender, with one apicofacial setule and weak, almost fully fused tooth spines on apices, outer plates subfalcate, with several nearly fused tooth- spines on medial margins; palp slender, poorly armed, article 3 with apical comb, dactyl unguiform, with thick apical nail and longitudinal comb. Coxae 1-4 increasingly elongate, anterior margin of coxa | strongly convex, with dis- tal notch and seta, one short midventral but submarginal setule; coxa 2 with 7 tiny and 1 long posteroventral setules; coxa 3 with same; coxa 4 with 15 tiny setules, no corner 122 seta. Article 3 of gnathopod 1 swollen, ar- ticle 4 tiny, article 5 scarcely lobate, article 6 strongly tapering, simple, dactyl stubby and setulate. Article 3 of gnathopod 2 elon- gate, articles 5-6 covered with straw-se- tules, palm weakly produced. Pereopod 3 grossly subchelate, articles 5—6 attached in eusirid fashion (tenuously), humped base of article 6 flexing into hollow of article 5, palm lined with partially chisel-shaped tooth spines, dactyl fitting palm. Pereopod 4 of normal gammaridean structure, with one unlocking setule. Article 2 of pereopod 5 broadly pyriform, posteriorly lobate, mar- gin weakly crenulosetulate; pereopods 6—7 slightly longer than 5, article 2 more nar- rowly pyriform, posteroventral margin weakly concave, weakly lobate. Posteroventral corner of epimera 2—3 mi- nutely extended as tooth. Uropods 1-2 poorly spinose, one spine each on dorsolat- eral and dorsomedial apex of peduncles, uropod | with basodorsal spine laterally, each outer ramus with one dorsal spine at midlength, apical nails on rami almost fully immersed. Uropod 3 with one apical spine on peduncle, inner ramus not reaching apex of article 1 on outer ramus, latter with ap- icomedial tooth, rami serrate apicomedi- ally, article 2 of outer ramus prominent. Telson ovate, with 2 pairs of dorsodistal penicillate setules. Female “‘w.’’—One large egg (room for 2 more). Oostegites vestigial or absent, coxa 5 with tiny broad flap-lobe tightly appressed to proximal base of coxa, lobe bearing 4 Fig. 1. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON vestigial setules; coxa 4 with similar lobe lacking setules, no oostegites found on cox- ae 2-3. Coxae 1-4 with ventral submarginal setules much longer than in male, whip-like, formula for coxae 1—4 = 1-6-6-14. Gills sac- like, sharply tapering apically, not plaited, lacking basal lobes. Color.—Ten minutes after preservation, body white, each pereonite with concen- trated orange blotch laterally. Holotype. —USNM 242012, male “t’”’ 2.38 mm. Type-locality. —Carrie Bow Cay, Belize, Central America, 18 Jun 1982; south side of channel between Carrie Bow Cay and South Water Cay, formalin wash of coral rubble from overhangs 6 m, J. D. Thomas, collector, station JDT-Bel 75C. Paratypes.— Type locality, male “‘v”’ 2.20 mm, female “w” 2.21 mm and 15 other specimens. Relationship. — Differing from the west Mexican Ensayara ramonella J. L. Barnard (1964) in the excavate posteroventral mar- gins of article 2 on pereopods 5-7, and the sparse (9 versus 25) setae on articles 5-6 of gnathopod 1. Closely similar to E. angustipes Ledoyer, 1978, from Mauritius, but differing in the excavate article 2 of pereopod 7, the lack of spines on the inner rami of uropods 1-2, the uniarticulate palp of maxilla 1, and the presence of a strong nail on the dacty! of the maxilliped. We are not certain about the exactitude of observations in the literature on the con- ~— Ensayara jumane, unattributed figures = holotype, male “t” 2.38 mm; u = male “v” 2.20 mm. Legend: Capital letters in figures refer to parts; lower case letters to left of capital letters refer to specimens and to the right refer to adjectives as described below: A, antenna; B, body; C, coxa; D, dactyl; G, gnathopod; H, head; I, inner plate or ramus; L, labium; M, mandible; O, outer plate or ramus; P, pereopod; S, maxilliped; T, telson; U, upper lip; Y, oostegite; r, right; s, setae removed; t, left. Fig. 2. Ensayara jumane, holotype, male “‘t” 2.38 mm, p. 124. Fig. 3. Ensayara jumane, unattributed figures = holotype, male “t’? 2.38 mm; v = male “v” 2.20 mm; w = female “w” 2.21 mm, p. 125. 123 VOLUME 103, NUMBER 1 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 124 _-|---, LLLE iz Le LZA, LZ 125 VOLUME 103, NUMBER 1 126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dition of the nail on the maxillipedal dactyl and the presence or absence of a second (basal) article on the palp of maxilla 1 and these must be confirmed in several of the species by better flattening and higher power microscopy. Distribution. — Belize, 6 m. Acknowledgments Fieldwork for this report was supported by Smithsonian’s CCRE committee; this is CCRE Contribution No. 261; we thank Dr. Klaus Ruetzler, head of this program, and Mike Carpenter for their assistance. The second author was supported by NSF Grant 8515186. Linda Lutz of Vicksburg, Missis- sippi, inked our drawings. Literature Cited Barnard, J. L. 1964. Marine Amphipoda of Bahia San Quintin, Baja California. — Pacific Natural- ist 4:55-139. —., &C. M. Barnard. 1983. The freshwater Am- phipoda of the world. Mt. Vernon, Virginia: Hayfield Associates, 2 volumes, 849 pp. , & G. S. Karaman. 1990. The families and genera of marine gammaridean Amphipoda, ex- cept Gammaroidea.— Records of the Australian Museum, Supplement (in press). Bellan-Santini, D. 1974. Amphipodes bathyaux de Méditerranée.— Bulletin de l'Institut Océano- graphique, Monaco 721(1427):20 pp., 8 figs. Hirayama, A. 1985. Taxonomic studies on the shal- low water gammaridean Amphipoda of west Kyushu, Japan V. Leucothoidae, Lysianassidae (Prachynella, Aristias, Waldeckia, Ensayara, Lepidepecreum, Hippomedon and Anonyx.— Publications of the Seto Marine Biological Lab- oratory 30:167-212. Ledoyer, M. 1978. Amphipodes gammariens (Crus- tacea) des biotopes cavitaires organogenes re- cifaux de I’Ile Maurice (Océan Indien).—The Mauritius Institute Bulletin 8:197—332. 1986. Crustacés amphipodes gammariens familles des Haustoriidae a Vitjazianidae.— Faune de Madagascar 59:599-1112. Lowry, J. K., & H. E. Stoddart. 1983. The shallow- water gammaridean Amphipoda of the subant- arctic islands of New Zealand and Australia: Lysianassoidea.— Journal of the Royal Society of New Zealand 13:279-294. (JLB) NHB-163, Department of Inver- tebrate Zoology, Smithsonian Institution, Washington, D.C. 20560; (JDT) Reef Foun- dation, P.O. Box 569, Big Pine Key, Florida 33043. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 127-130 A RANGE EXTENSION TO THE NORTH FOR MACROCHIRIDOTEA GIAMBIAGIAE TORTI AND BASTIDA (CRUSTACEA: ISOPODA: VALVIFERA) Elizabeth Harrison-Nelson and Thomas E. Bowman Abstract.—Macrochiridotea giambiagiae is reported from Rio de Janeiro, Brazil, and slight differences from the original description by Torti and Bastida (1972) are noted. This new record extends the known range of the species more than 1000 km to the north. The genus Macrochiridotea Ohlin, 1901 comprises | 1 species confined to the South- ern Hemisphere (New Zealand, Falkland Is- lands, east and west coasts of South Amer- ica from 23°12'S to 49°29’S): M. michaelseni Ohlin, 1901, M. stebbingi Ohlin, 1901, M. kruimeli Nierstrasz, 1918, M. setifer Men- zies, 1962, M. uncinata Hurley & Murray, 1968, M. robusta Bastida & Torti, 1969, M. giambiagiae Torti & Bastida, 1972, M. marcusi Moreira, 1973, M. lilianae Mor- eira, 1973, M. mehuinensis Jaramillo, 1977, M. australis (Richardson, 1911) (trans- ferred from Chiriscus by Poore 1984). It has been been found at depths ranging from the intertidal to 309 meters, and on substrates varying from mud with fine sand to coarse quartz and calcareous sands. Specimens of this genus may reach 15 mm in length. Mor- eira (1973) reviewed the genus and provid- ed a revised diagnosis which includes the statement that the inner lobe of maxilla 1 bears 2 apical setae. In fact this lobe bears a single seta in M. giambiagiae and the number of setae is unknown in M. kruimeli, M. michaelseni, M. setifer, and M. stebbingi. The remaining 6 species all have 2 setae. We report herein a new record for M. giambiagiae, which extends this species’ known range north by about 5° (1000 km), and slightly increases the known depth range, from 2 to 7 m. Its distribution thus extends from the northern to the southern limits of the Eastern South American Warm Tem- perate Region as defined by Briggs (1974). It is the most northerly occurring member of the genus. It is somewhat surprising that M. giam- biagae, published in 1972 and recorded in the Zoological Record for that year (pub- lished in 1978), is not listed in subsequent papers on Macrochiridotea by other authors (Moreira 1973, Epelde-Aguirre & Lopez 1975, Jaramillo 1977, Poore 1984), and is not included in Moreira’s 1972 review of the genus. Macrochiridotea giambiagiae Torti & Bastida Fig. 1 Macrochiridotea giambiagiae Torti & Bas- tida, 1972: passim. (Mar del Plata, Ar- gentina; Chuy, Uruguay; Cassino, Brazil). Material.—Copacabana Beach, Rio de Janeiro, Brazil, 150-300 m from shore in coarse quartz sand, 5-7 m, leg. James D. Thomas, 3—4 May 1985, | 6 (3.5 mm), 2 °° (1 ovigerous—4.5 mm, | non-ovigerous— 5.5 mm), 4 juveniles, USNM 241981.—Vil- la Gesell, Buenos Aires, Argentina, 2 para- type of M. giambiagiae (5.5 mm), USNM 139314. Description. —Length 1.75 to 5.5 mm. Our specimens have the features diagnostic of M. giambiagiae, i.e., dorsal surface of body smooth; head without tubercles, setose on lateral margins, slightly convex dorsally, 128 lateral incisions shallow, eyes small, faintly pigmented (Fig. 1a); 1 seta on endopod of maxilla | (Fig. 1c); pereopods 2 and 3 with- out dactyl (Fig. 1g); male pleopod 2 like that shown by Torti & Bastida (Fig. 11). Our material differs from Torti & Basti- da’s description as follows: article 4 of an- tenna | about *%4 rather than '2 as long as article 2 and with 4 rather than 2 setae on distal margin (Fig. 1b); terminal article of flagellum with 3 rather than 2 apical setae. Exopod of maxilla 1 with 12 spines (Fig. lc), type material with 9. Palp of maxilla 2 with 5 setae (Fig. 1d), type material with 4; endopod with 8 setae, type material with 6. Tip of pleotelson with 2 small setae medial to posterior pair of long setae (Fig. 1h). The small setae are not shown by Torti & Bas- tida and could not be seen in our paratype. Torti & Bastida (1972:21-—22) mention that pleonite 1 of M. giambiagiae tends to telescope into pereonite 7 and may not be visible in dorsal view. This was true for the 2 largest (4.5 and 5.5 mm) specimens from Rio de Janeiro, but the first segment was visible on the smaller specimens (1.75—3.5 mm). Torti & Bastida did not note any sexual dimorphism in their 19 females and 3 males, except the appendix masculina on the male pleopod 2, nor did they note any differences between the adult male holotype and another adult male and juvenile male included with their material. The female paratype of M. giambiagiae (USNM 139317, 5.5 mm) from Villa Ge- sell, Province of Buenos Aires, Argentina (37°28'S, 57°07'W) appears to conform to the description and figures of Torti & Bas- tida except that the distal process of the carpus of pereopod 1 bears several articu- lated setae as do our Rio de Janeiro speci- Fig. 1. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mens (Fig. If). Torti & Bastida’s illustra- tions, which depict the holotype male (4.3 mm) collected at Mar del Plata, Argentina (38°03'18"S, 57°32'30”W), show nonarticu- lated setae. Our male (3.5 mm) bears artic- ulated setae, thus this characteristic does not appear to be size/age/sex related. The differences between our specimens and those of Torti & Bastida are slight com- pared to those separating species of Macro- chiridotea. If, however, additional collec- tions should show them to be constant, the possibility of assigning distinct specific or subspecific status to the Rio de Janeiro form would have to be considered. Acknowledgments We thank James D. Thomas of the Reef Foundation, Big Pine Key, Florida, for pro- viding us with the specimens. Literature Cited Bastida, R., & Torti, M.R. 1969. Un nuevo isopodo del genero Macrochiridothea de las costas Ar- gentinas (Valvifera, Idoteidae).— Neotropica 15(47):65-72. [In Spanish.] Briggs, J.C. 1974. Marine zoogeography. New York, McGraw Hill, 475 pp. Epelde-Aguirre, A., & Lopez, M. T. 1975. Zonacion en el sustrato arenoso de playa Blanca, Bahia de Coronel y observaciones sobre crustaceos poco frecuentes.—Boletin de la Sociedad Biol- ogica de Concepcion 49:161-170. [English sum- mary.] Hurley, D. E., & Murray, R.H. 1968. A new species of Macrochiridothea from New Zealand, with notes on the idotheid subfamily Chaetilinae (Crustacea Isopoda: Valvifera.— Transactions of the Royal Society of New Zealand (Zoology) 10(26):241-249. Jaramillo, E. 1977. Macrochiridotea mehuinensis (Valvifera, Idoteidae), nuevo isopodo marino de las costas del sur de Chile.—Studies on the Neotropical Fauna and Environment 12(1):71- 80. [English summary.] =) Macrochiridotea giambiagiae, female, USNM 241981: a, Head, dorsal view; b, Antenna 1; c, Maxilla 1; d, Maxilla 2; e, Maxilliped; f, Pereopod 1, tip of carpus enlarged (medial view); g, Pereopod 2; female, USNM 241982: h, Tip of pleotelson; male, USNM 241983: i, Pleopod 2, endopod with appendix masculina. 130 Menzies, R. J. 1962. The Zoogeography, ecology, and systematics of the Chilean marine isopods. Re- ports of the Lund University Chile Expedition 1948-49:42.—Lunds Universitets Arsskrift, N.F. Avd. 2, 57(11):1-62. Moreira, P. S. 1972. On the distribution of species of Macrochiridothea Ohlin, 1901 in southern Brazil (Crustacea: Isopoda: Valvifera). —Revis- ta Brasileira de Biologia (Rio de Janeiro) 32(3): 395-399. 1973. Species of Macrochiridothea Ohlin, 1901 (Isopoda, Valvifera) from southern Brazil, with notes on remaining species of the genus. — Boletim Instituto Oceanografico, Sao Paulo 22: 11-47. Nierstrasz, H. F. 1918. Alte und neue Isopoden.— Zoologische Mededeelingen 4(2):103-142, pls. 9, 10. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ohlin, Axel. 1901. Isopoda from Tierra del Fuego and Patagonia. 1. Valvifera.—Svenska Expedi- tionen till Magellanslanderna 2(11):26 1-306. Poore, G. C. B. 1984. Clarification of the monotypic genera Chiriscus and Symmius (Crustacea: Isop- oda: Idoteidae).— Proceedings of the Biological Society of Washington 97:71-77. Torti, M. R., & Bastida, R. 1972. Presencia del genero Macrochiridothea Ohlin, 1901 en Uruguay y Brasil: M. giambiagiae sp. nov. (Crustacea, Is- opoda).—Neotropica 18(55):16-22. [In Spanish with English summary.] Department of Invertebrate Zoology (Crustacea), NHB Stop 163, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 131-139 HETEROM YSIS MEXICANA, A NEW SPECIES FROM CAMPECHE BANK, GULF OF MEXICO (CRUSTACEA: MYSIDACEA) Elva Escobar-Briones and Luis A. Soto Abstract. —Heteromysis mexicana is described from waters of Campeche Bank, southern Gulf of Mexico. Its general morphology closely resembles the eurytopic species H. formosa Smith, 1873, which is distributed along the west- ern northern Atlantic in subtropical and temperate coastal waters. Available morphologic and ecologic data for 24 species of Heteromysis occurring in the Tropical Western Atlantic show no specific latitudinal distribution or host selectivity. The Heteromysis occurring in the Trop- ical Western Atlantic comprise 26 species, most of which have cryptic habits (Modlin 1987c) and are associated with coralheads, anemones and sponges. The group exhibits considerable adaptive radiation. In the Gulf of Mexico 11 species have been described and 13 more occur in the Caribbean-Antil- lean region and the Tropical Western At- lantic (Tables 1, 2). Most records in the Gulf are limited to the eastern and northern parts. This paper presents the description of a new species of Heteromysis, the first re- corded for the southwestern Gulf of Mexico and summarizes the zoogeography of the Heteromysis species recorded in the Trop- ical Western Atlantic. Materials and Methods The specimens reported herein were col- lected during the research program “‘Ocean- ography of the Gulf of Mexico” (OGMEX) conducted aboard the R/V Justo Sierra in March 1987. Material collected has been coded as follows: OGMEX.I is the project name and cruise number, in order the fol- lowing digits represent the year of collection and the station number, and in parentheses is the depth in meters. The number and sex of the individuals captured is designated as lf (= female), 1m (= male), and 1j (= ju- venile). Hydrographic parameters and sediment texture were determined at the station. Bot- tom salinity and temperature were recorded with a Niels-Brown CTD sensor. The sed- iments obtained with a Smith-MclIntyre grab were analyzed according to Folk (1969) for the sand and mud size fractions using sed- iment sieves and the pipette method re- spectively. Organic content of sediment was determined by the loss on ignition tech- nique, and burnt to 550°C. Samples were fixed and stored in 70% ethanol. Total length (T.L.), measured with a calibrated ocular micrometer in a dis- secting microscope, is the distance along the dorsal midline from the tip of the rostrum to the posterior margin of the telson, ex- cluding apical spines. Illustrations were made with the aid of a camera lucida. Results Heteromysis mexicana, new species Figs. lA-E, 2A-E, 3A—H, 4A, B Type material.—Holotype: Male (T.L. 10.2 mm) collected at Campeche Bank, southern Gulf of Mexico, March 1987, ob- tained from a night trawl sample [OG- 132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Heteromysis species occurring in the Gulf of Mexico. (Additional distributional information in brackets.) $$$ Species i H. beetoni Modlin, 1984 H. bermudensis cesari Bacescu, 1968 H. dispar Brattegard, 1970 H. filitelsona Modlin, 1984 H. formosa, Smith, 1873 gomezi Bacescu, 1970 . quitarti Bacescu, 1968 . hopkinsi Modlin, 1984 . mariani Bacescu, 1970 . nouveli Brattegard, 1969 _ rubrocineta Bacescu, 1968 aTertrTs MEX.I-87-52 (168) Im]. Dissected and mounted on two slides and deposited in USNM, USNM 241592. Allotype: Female (T.L. 10.3 mm) from same sample. Deposited in USNM, USNM 241593. Paratypes: Juvenile (T.L. 5.4 mm) from same sample. Deposited in USNM, USNM 241594. Type locality.—Campeche Bank, south- ern Gulf of Mexico, Mexico, between 19°31'N, 92°37'W and 19°33'N, 92°37'W. Description. —Body robust. Carapace (Fig. Source Quadrant Modlin (1984) NE Bacescu (1968) SE (Carib.) Brattegard (1970) SE Modlin (1984) NE Stuck et al. (1979b) NE (eurytop.) Tattersall (1951) NE Bacescu (1970) SE Bacescu (1968) E (Keys) Modlin (1984) NE Bacescu (1970) SE Brattegard (1969) SE (Keys) Bacescu (1968) SE 1A) with anterior margin produced into small triangular rostrum, posterior margin deeply emarginate, exposing part of thoracic segment 7 and all of 8, anterolateral lobes rounded. Eyes large, oval; cornea large, brown. Antennular peduncle (Fig. 1B) segment | as long as segment 3, with four long setae on conical distolateral process and two setae near middle of distal margin. Segment 2 short with two plumose setae distodorsally. Segment 3 with male lobe located midven- trally near distal margin and crowned by a Table 2.—Heteromysis species occurring in the Caribbean Sea and the Western Tropical Atlantic. Species ——— . actinae Clarke, 1955 agelas Modlin, 1987 bermudensis Sars, 1885 bredini Brattegard, 1970 . coralina Modlin, 1987 . disrupta Brattegard, 1970 elegans Brattegard, 1974 floridensis Brattegard, 1969 kensleyi Modlin, 1987 mayana Brattegard, 1970 mureseanui Bacescu, 1986 . siciliseta Brattegard, 1970 . tuberculospina Modlin, 1987 . venezuelensis Bacescu, in press Heteromysis sp. A Brattegard, 1974 TEES SEPT TTTATS Source Distribution Clarke (1955) Carib.-Antil. Modlin (1987c) Bahamas Tattersall (1951) Bermuda-Carib. Brattegard (1970) Antilles Modlin (1987b) Fla. Keys Brattegard (1970) Caribbean Brattegard (1974a) Caribbean Brattegard (1969) Baham.-Fla. Keys Modlin (1987b) Caribbean Brattegard (1970) Caribbean Bacescu (1986) Brazilian Brattegard (1970) Caribbean Modlin (1987a) Caribbean Bacescu, in press Caribbean Brattegard (1974a) Caribbean VOLUME 103, NUMBER 1 133 Fig. 1. Heteromysis mexicana (male 10.2 mm). A. Dorsal view; B. antennular peduncle; C. antennal scale; D. left mandlible and mandibular palp; E. detail of left mandible lacinia mobilis and accessory blades. 134 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dense tuft of hairs, two simple setae ven- trally at midlength, one plumose seta me- dially at midlength and seven setae disto- medially. Antennular flagellum with stout spinules along proximal half. Antennae (Fig. 1C) peduncle segment 1 short, compressed; segment 2 twice length of segment 1 and 1.6 times length of seg- ment 3, with three plumose setae at middle of distal margin; segment 3 with four setae near middle of distal margin; flagellum with two rows of setules along entire length. An- tennal scale as long as or slightly longer than peduncle, 2.3 times as long as maximum width with distal tip articulated; lateral margin nearly straight, medial margin con- vex, entire margin setose, dorsal surface with nine proximal setae. Mouthparts (Figs. 1D, E, 2A—D), man- dibular palp 3-segmented, segment 1 nar- row, segment 2 broad, lateral margin with 19 simple setae, one setae in proximal dor- sal position, medial margin with curved row of 10 simple setae and three simple setae distally, segment 3 lateral margin sinuous, with four long submarginal plumose setae proximally, nine surface pectinate spinules, 23-26 pectinate setae diminishing in size proximally, distally one longer robust seta. Left mandible with short bladelike inci- sor, lacinia mobilis crownlike with 17 cusps longer on both ends, accessory blades in spine row with five blades, all slightly ser- rate on inner margin, molar process well developed with 14 rows of triturative teeth. Right mandible with three bladelike inci- sors, two setose accessory blades, molar process well developed with 12 rows of tri- turative teeth and lateral plumose setae. Maxillule well developed, outer plate with 14 apical spines and five subapical setae, middle inner margin setose, inner plate glo- bose with four pectinate spinules, four sim- ple setae and one hooked serrated spine on distal margin, seven plumose setae subdis- tally and two simple setae on lateral process. Maxillae (not figured) typical of the genus, exopod elongate with 22 plumose setae on distolateral margin. Paragnath typical of the genus with inner and outer lobe shoulders setose, mandibular process short, setose. Labrum tyical of the genus with apical mar- gin setose. Thoracic endopod | (Fig. 2E), short, ro- bust, carpopropodus with six marginal setae and eight dorsodistally; dactylus rounded with nine pectinate setae. Thoracic endopod 2 (Fig. 3A), ischium with 10-11 simple setae on anterior margin, merus anterior margin with seven setae; car- popropodus distally expanded with 10 spines on posterior margin, three distal se- tae, and four robust setae distally on ante- rior margin; dactylus small, rounded mit- ten-shaped, setose all around. Thoracic endopod 3 (Fig. 3B), strong, short, merus and carpopropodus inflated, lacking spines characteristic of genus; dac- tylus ending in terminal claw. Thoracic endopod 4 (Fig. 3C), carpopro- podus with six segments, distal one with two serrated flexible spines; dactylus small with long terminal setae and ending in curved clawlike spine. Thoracic endopod 5 (Fig. 3D) to 8, car- popropodus with five segments, increasing slightly in size distally. Female with oostegites in thoracic seg- ments 6, 7 and 8 well developed, increasing in size distally. Pleopods 1 to 5 (Fig. 3E—-H) not sexually dimorphic. Well developed, similar in form, unsegmented, increasing in size distally. Pleopod 1, five setae and one terminal spine on anterior surface, five plumose setae on lateral margin, five setae on pseudobran- chial lobe. Pleopod 2, six setae on anterior surface, one setae on distal margin, three setae on pseudobranchial lobe, four setae on lateral margin. Pleopod 3 with 11 setae on anterior surface, two setae on distal mar- gin, four setae on pseudobranchial lobe, four on lateral margin. Pleopod 4, 13 setae on anterior surface, three setae on distal mar- gin, three setae on pseudobranchial lobe, four setae on lateral margin, one distally. VOLUME 103, NUMBER 1 135 Fig. 2. Heteromysis mexicana (male 10.2 mm). A. Mandibular palp third segment; B. right mandible; C. maxillule; D. paragnath; E. thoracic endopod 1. Fig. 3. Heteromysis mexicana (male 10.2 mm). A. Thoracic endopod 2; B. thoracic endopod 3; C. thoracic endopod 4; D. thoracic endopod 5; E. pleopod 1; F. pleopod 2; G. pleopod 3; H. pleopod 4. VOLUME 103, NUMBER 1 Pleopod 5, 16 setae on anterior surface, four setae on distal margin, three setae on pseu- dobranchial lobe, four setae on lateral mar- gin. Uropods (Fig. 4A), exopod subequal in length to endopod, lateral margin straight, apex rounded, medial margin slightly con- vex, setose all around. Endopod lateral mar- gin straight, medial margin concave with 23 stout spines, equal in size along entire mar- gin, apex rounded, both margins with plu- mose setae. Telson (Fig. 4B) 1.7 times longer than wide at base, lateral margins straight, distal half of lateral margins with 14-15 spines increasing slightly in length posteriorly; one longer apical spine on each side. Telsonal cleft completely lined with 22-23 small spines, increasing slightly in size distally; depth 0.17 times length of telson. Remarks.—The new species closely re- sembles the eurytopic species Heteromysis formosa Smith (1873) and can be distin- guished from it by morphological differ- ences listed in Table 3 from which the most important features that separate them are the well developed male lobe on antennular peduncle segment 3; the absence of three pairs of spines on the carpopropodus of the thoracic limb 3; medial margin of uropodal endopod with 20-23 spines (14-19 in H. formosa) and 22-23 spines in the cleft of the telson (8—20 in H. formosa). Both species have in common well developed but not sexually dimorphic pleopods (Modlin, pers. comm.). Ecological notes.—This species was an occasional component of the macrocrusta- cean epibenthic community from the inner continental shelf. Specimens were obtained associated with empty Strombus gigas shells between 19°31’N, 92°37’'W and 19°33’N, 92°37'W at 168 m depth. Bottom salinity was 36.28%o and temperature was 23.08°C. The sediment at the site was fine sand mud with an organic matter content of 23% and carbonates 11%. The behavior of hiding in 137 Sym 3 pm Fig. 4. Heteromysis mexicana (male 10.2 mm). A. uropodal endopod and exopod; B. telson. gastropod shells closely resembles that re- ported for Heteromysis formosa (Clarke 1955, Brattegard 1969) and other species morphologically dissimilar, e.g., H. ber- mudensis, H. mayana, H. tuberculospina (Modlin, 1987a). Etymology.—Named for the country in which it was found, Mexico. 138 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.—Comparison of Heteromysis formosa Smith (1873) and H. mexicana n. sp. Character H. formosa H. mexicana Antennular peduncle, male lobe Antennular peduncle segment 3 Antennal scale length/breadth ratio 34 Third thoracic limb carpopropodus Uropodal endopod rudimentary! with 2 plumose setae* with 3 pairs of spines*> with 14-16 spines? well developed with 1 plumose seta 2.3 without spines with 20-23 spines with 17-19 spines* with 16-19 spines> Telson lateral margin with 12-14 spines! with 14-17 spines with 14-16 spines?? with 11 spines* with 10-18 spines> Telson cleft with 20 spines? with 22—23 spines with 8-10 spines* with 16-13 spines? ' Smith, 1873. 2 Sars, 1882. 3 Tattersall, 1951. 4 Brattegard, 1969. 5 Stuck et al., 1979a. Discussion This new species is the first report of Het- eromysini for the southwestern Gulf of Mexico. The peracaridean shelf fauna in this region is largely unknown. Knowledge of it is limited to the investigations of epibenthic macroinvertebrates in Laguna de Términos (Escobar & Soto 1988) and Alvarado (Soto et al. 1986). Twenty-six species of Heteromysids have been described in the West Tropical Atlan- tic (Bacescu 1968, 1970, 1986; Brattegard 1969, 1970, 1973, 1974a, b; Clarke 1955; Modlin 1982, 1984, 1987a, b, c; Sars 1885; Smith 1873; Stuck et al. 1979a, b), of which 11 have been reported from the northern and southeastern parts of the Gulf of Mex- ico (Table 1), and 13 have been reported from the Caribbean and the Western Trop- ical Atlantic (Table 2). At least 12 species are cryptic and are associated with sponges and corals. Heteromysids mostly inhabit shallow waters, but the present report of Heteromysis mexicana from the Campeche Bank at 168 m is among the deepest. Re- ports for the closely related species, H. for- mosa, reach up to 203 m in the Gulf of Mexico and 227 min the southeastern coast of the United States (Tattersall 1951). Acknowledgments The authors wish to express their appre- ciation to Dr. Thomas Bowman of the Smithsonian Institution, Dr. Mihail Baces- cu from the Musee D’ Histoire Naturelle ‘““Grigore Antipa,’’ Rumania and Dr. Rich- ard F. Modlin of the University of Alabama in Huntsville for their critical reviews on an earlier version of this manuscript. For their support in the field gratitude is expressed to the team of the Benthic Ecology Lab. This is Contribution #587 from the Instituto de Ciencias del Mar y Limnologia, UNAM. Literature Cited Bacescu, M. 1968. Heteromysini nouveaux des eaux cubaines: trois especes nouvelles de Hetero- mysis et Heteromysoides spongicola n.g. n. sp.— Revue Roumaine de Biologie—Zoologie 13:221- 237. . 1970. New spongicolous Heteromysis of the Caribbean sea (H. gomezi n. sp. and H. mariani VOLUME 103, NUMBER 1 n. sp.).—Revue Roumaine de Biologie—Zoolo- gie 15:11-16. 1986. Heteromysis mureseanui n. sp. and Kalliapseudes viridis, ssp. brasiliensis n. ssp., from the Brazilian littoral waters. — Revue Roumaine de Biologie—Biologie Animale 31:93-97. Brattegard, T. 1969. Marine biological investigations in the Bahamas. 10. Mysidacea from shallow water in the Bahamas and southern Florida. Part I.—Sarsia 39:17-106. 1970. Mysidacea from shallow water in the Caribbean Sea.—Sarsia 43:111-154. 1973. Mysidacea from shallow water on the Caribbean coast of Colombia.—Sarsia 54: 1-66. 1974a. Additional Mysidacea from shallow water on the Caribbean coast of Colombia.— Sarsia 57:47-86. . 1974b. Mysidacea from shallow water on the Caribbean Coast of Panama.—Sarsia 57:87-108. Clarke, W. D. 1955. A new species of the genus Het- eromysis (Crustacea, Mysidacea) from the Ba- hama Islands, commensal with a sea-anemo- ne.—American Museum Novitates 1716:1-13. Escobar, E., & L. A. Soto. 1988. Mysidacea from Términos Lagoon, Southern Gulf of Mexico, and description of a new species of Taphromysis. — Journal of Crustacean Biology 8:335-367. Folk, R. L. 1969. Petrologia de las rocas sedimen- tarias. Universidad Nacional Autonoma de Mexico, Mexico, 405 pp. Modlin, R. F. 1982. Contributions to the ecology of the mysid crustaceans in the shallow waters of Dauphin Island, Alabama.—Northeast Gulf Science 5:45—-49. 1984. Mysidacea from the Florida Middle Ground, northeast Gulf of Mexico, with de- scriptions of three new species of Heteromysis and a key to the Heteromysini of the western Atlantic. — Journal of Crustacean Biology 4:287- 297. . 1987a. Mysidacea from shallow waters in the vicinity of Carrie Bow Cay, Belize, Central America, with descriptions of two new species. — Journal of Crustacean Biology 7:106-121. . 1987b. Heteromysis kensleyi and H. coralina, 139 new species from the shallow waters off Looe Key, Florida (Mysidacea: Heteromysini).—Pro- ceedings of the Biological Society of Washington 100:653-658. 1987c. Heteromysini from Grand Bahama Island: description of Heteromysis agelas, new species, first description of male H. floridensis, and notes on H. guitarti (Crustacea:Mysida- cea).— Proceedings of the Biological Society of Washington 100:296-301. Sars,G.O. 1882. Oversight af Nerges Crustaceer med forelobige Bemaerkninger over de nye eller mindre bekjendte Arter. I Podophthalmata-Cu- macea-Isopoda-Amphipoda.—Christiania Vi- denskabsselskabs Forhandlinger 18:54—55. 1885. Reports on the Shizopodda collected by H.M.S. Challenger during the years 1873- 1876.—Challenger Reports—Zoology 13:1-225. Smith, S. I. 1873. Report upon the invertebrate an- imals of Vineyard Sound and the adjacent waters. — Report of the Commission of Fisheries and Fish 1871-1872:295-747. Soto, L. A., et al. 1986. Ecologia poblacional de los camarones peneidos de los principales sistemas lagunares del Golfo de México. Segundo In- forme Técnico. CONACyT-PCEBNA 021436, 65 pp. Stuck, K. C., H. M. Perry, & R. W. Heard. 1979a. An annotated key to the Mysidacea of the north central Gulf of Mexico.—Gulf Research Re- ports 6:225-238. ' , & . 1979b. Records and range extensions of Mysidacea from coastal and shelf waters on the eastern Gulf of Mexico.—Gulf Research Reports 6:239-248. Tattersall, W. M. 1951. A review of the Mysidacea of the United States National Museum.—Bul- letin of the United States National Museum 201: 1-292. Instituto de Ciencias del Mar y Limnol- ogia, Universidad Nacional Autonoma de Mexico, A.P. 70-305, 04510 Mexico City, Mexico. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 140-150 REDESCRIPTION AND NEW RECORDS OF TRICHODIAPTOMUS CORONATUS (G. O. SARS) (COPEPODA; CALANOIDA; DIAPTOMIDAE) FROM BRAZIL Janet W. Reid Abstract. — Trichodiaptomus coronatus (G. O. Sars), originally described from Sao Paulo, Brazil, has been recorded recently only from Brazilian Amazonia and the Orinoco Delta. This article reports new records from four localities in the Brazilian highlands, including the first record from the Rio Sao Francisco Basin. A redescription of representatives from these populations is furnished. Morphological variations reported for this species in Amazonia were not ob- served in the southern populations examined. T. coronatus inhabits clear or blackwater, extremely oligotrophic rivers, ponds and reservoirs. G. O. Sars (1901) described a distinctive freshwater calanoid copepod, Diaptomus coronatus, from cultures of dried mud from the State of Sao Paulo, Brazil. Wright fur- nished additional morphological details of populations of this species from Santarém, State of Para (1927), and the State of Sao Paulo (1937). Thomasson (1954, 1955) re- ported Diaptomus melini as a new, similar species from Manaus, State of Amazonas. Noting morphological variability in Ama- zonian populations of D. coronatus, Bran- dorff (in Brandorffet al. 1982) synonymized D. melini with Sars’ species and proposed the new genus Trichodiaptomus for it. Al- though 7. coronatus has been recorded sev- eral other times from Brazilian Amazonia (Andrade & Brandorff 1975, Brandorff 1972, Cipolli & Carvalho 1973, Matsumura-Tun- disi 1986) and once from the Orinoco Delta, Venezuela (Dussart 1984), it has not been encountered again in southeastern Brazil. This article reports new records from the Distrito Federal, Goias, and Minas Gerais in the central Brazilian highlands, including the first record from the Rio Sao Francisco Basin; redescribes the species; and sum- marizes knowledge of its ecological require- ments. Genus Trichodiaptomus Brandorff (in Brandorff et al., 1982) Trichodiaptomus coronatus (G. O. Sars, 1901) Figs. 1-21 Diaptomus coronatus G. O. Sars, 1901:14— 16, pl. II, figs. 9-17.—Daday, 1905: 151.—Tollinger, 1911:66, fig. A2.—Pes- ta, 1927:80.—Wright, 1927:74, 75, 90- 91, 100, pl. VI, figs. 7-9; 1937:66, 77-79, pl. Il, figs. 5-8; 1938:562.—Brandorff, 1972:8, 9, 20-23, 50, figs. 19-22.—Ci- polli & Carvalho, 1973:95, 97, 98, 100, 101, 108. ‘**‘Diaptomus” coronatus G. O. Sars.—An- drade & Brandorff, 1975:97, 103.—Bran- dorff, 1976:618, 619, 622, fig. 3; 1978: IO. Diaptomus melini Thomasson, 1954:193- 194, fig. III 1 a-c; 1955:214.— Brandorff, 1972:20-21, 48. ‘‘Diaptomus’’ melini Thomasson.—An- drade & Brandorff, 1975:102. Trichodiaptomus coronatus (G. O. Sars).— Brandorff et al., 1982:76, 104-106, figs. 104-110.—Dussart & Defaye, 1983: 134.—Arcifa, 1984:143.—Dussart, 1985: 201.—Reid, 1985:78-79, fig. 1.—Mat- VOLUME 103, NUMBER 1 sumura-Tundisi, 1986:547, 551, figs. 89- 94, 100. Notodiaptomus coronatus (G. O. Sars).— Dussart & Defaye, 1983:134.—Dussart, 1984:34, 39. Rhacodiaptomus Melini (Thomasson). — Brehm, 1965:15. Rhacodiaptomus Mileni (Thomasson). — Brehm, 1965:15. Material. — Brazil: 1 6 and 1 °, Santarém, Para (undated), USNM 58920, 58921, col. S. Wright. 100+ adults, Lagoa Bonita (La- goa Mestre d’Armas), Distrito Federal, 15°34'S, 47°10'W, 19 Jan 1979, 24 Jan 1979, 9 Sep 1980, 23 Sep 1980, 21 Oct 1980, 4 Nov 1980; 15 adults, Lagoa Formosa, Goias, 15°30’S, 47°36’W, 26 Oct 1980, 4 Jul 1982; 100+ adults, Represa (Rep.) Santo Anténio do Descoberto, Distrito Federal/Goias, 15°44’S, 48°10’W, 25 Oct 1980, 8 May 1982, all USNM 241951; 200+ adults, Lagoa Bonita, 23 Sep 1980, Museu de Zoologia da Universidade de Sao Paulo (MZUSP) 9651; 200+ adults, Lagoa Bonita, 9 Sep 1980, MZUSP 9652; 200+ adults, Rep. Santo Antonio do Descoberto, 8 May 1982, MZUSP 9653; 16 adults, Lagoa Formosa, 4 Jul 1982, MZUSP 9654; col. J. W. Reid and/or L. El-Moor Loureiro. 12 2, Lagoas Tacho, Paiano, and Cipo (combined sam- ples), Pirapora, Minas Gerais, 17°20'55’S, 44°57'00’W, 1988; M. B. G. e S. Dabés col- lection. Female.—Median lengths (and ranges), excluding caudal setae, of specimens from Lagoa Bonita 1.13 mm (1.05-1.28 mm; n = 40); from Rep. Descoberto 1.20 mm (1.12-1.28 mm; n = 14); from Lagoa For- mosa 1.20 mm (1.08-1.30 mm; n = 18); from Pirapora ponds 1.28 mm (1.22-1.35 mm; n = 12). Body widest at pedigers 1-2 in dorsal view (Fig. 1). Pedigers 4 and 5 (Figs. 1-3) separated; pediger 5 produced on each side into posteriorly directed, point- ed wing reaching past midlength of genital segment. Few females with one wing bear- ing large dorsally directed spine, wing some- 141 times also not fully pointed (Figs. 4, 5); op- posite wing always normally pointed. Prosomites with 1-5 pairs of papillae each tipped with fine hair, and rows of hairs near posterior margins, hairs on pedigers 2—4 coarser. Pediger 4 (Figs. 1-5) with lateral row of 5-13 large spines, usually a similar number on each side. Urosome (Figs. 1, 2, 6) of 2 segments; anterior portion of genital segment slightly expanded laterally; lateral sensillae on genital segment similar to pro- somal ones. Genital field as in Fig. 6. Inner margins of caudal rami hairy. Rostral points acute. Antennule (Figs. 2, 7) reaching past end of caudal rami; ar- mament similar to that of Scolodiaptomus corderoi (Wright) as redescribed by Reid (1987), particularly in having only | seta on article 11. Antenna (Fig. 8) with 2-3 setae on anterior margin of endopod 1, posterior margin of this article lacking ornament; an- tenna otherwise normal for family. Gnathal lobe of mandible (Fig. 9) with acute apical and subapical teeth, dentition otherwise similar to that of S. corderoi. Remaining mouthparts and legs 14 also similar to those of S. corderoi and typically diaptomid; Schmeil’s organ present on posterior surface of leg 2 endopod article 2. Leg 5 (Figs. 10-12) slender; exopod ar- ticle 3 continuous with article 2. Endopod of 2 distinct articles, reaching slightly be- yond midlength of exopod article 1; endo- pod 2 with subterminal oblique row of 5-8 spinules. Male. — Median lengths of specimens from Lagoa Bonita 0.85 mm (range 0.82—1.00 mm; n = 24); Rep. Descoberto 0.90 mm (0.72-0.95 mm; n = 20); Lagoa Formosa 0.90 mm (0.88—-1.06 mm; n = 3). Habitus (Fig. 13) as in female, except urosome 5-segmented, pediger 4 with 4-8 spines on each side, wings of pediger 5 short. Left an- tennule, mouthparts, and legs 1—4 as in fe- male. Right antennule (Figs. 14-16) with articulated spines on articles 8 and 12; spine on article 11 longer than spine on article 10; spine on article 13 equal to or longer than 142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Trichodiaptomus coronatus, females (1 and 2 from Pirapora ponds; 3-6 from Lagoa Bonita): 1, Habitus, dorsal; 2, Habitus, right lateral; 3, Pedigers 4-5 of normal specimen; 4, Pedigers 4—5 and genital segment of specimen with abnormal spine on left wing; 5, Pedigers 4—5 of specimen with abnormal spine on rounded left wing; 6, Urosome and caudal rami, ventral. VOLUME 103, NUMBER 1 ai \ Figs. 7-12. 143 Trichodiaptomus coronatus, females from Lagoa Bonita: 7, Antennule; 8, Antenna; 9, Gnathal lobe of mandible; 10, Leg 5; 11 and 12, Detail of end of leg 5 endopod 2 from different specimens. spine on article 11. Spines on articles 10, 11, and 13 more developed in specimen from Amazonia (Fig. 16). Articles 14-16 without spines. Antepenultimate article without distal process. Left leg 5 (Figs. 17-21) slender; posterior mammiform process on basipod | small, ending in short spine. Margins of basipod 2 nearly straight. Endopod of 2 indistinctly separated articles, distal article with few hairs near conical tip. Exopod of 2 articles, each with hairy pad on anteromedial sur- face, proximal pad slightly developed. Ar- ticle 2 rotated inward, bearing acute, finely pectinate digital process, and slender spine with fine hairs. Right leg 5 (Figs. 17, 20), with basipods 1 and 2 similar to basipod of left leg; en- dopod of 2 indistinctly separated articles, oblique terminal surface hairy. Exopod 1 with inner and outer distal corners slightly extended. Exopod 2, lateral spine shorter 144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 13-16. Trichodiaptomus coronatus, males (13-15 from Lagoa Bonita; 16 from Santarém): 13, Habitus, dorsal; 14, Right antennule; 15 and 16, Articles 8-14 of right antennule (most setae and esthetascs not indicated). than breadth of exopod; terminal claw equal in length to exopod, tip of claw slightly re- curved in some specimens. Variation. —Lengths of females from southern Brazil are less than lengths (1.19- 1.46 mm) reported by Brandorff et al. (1982) for Amazonian populations. Wright (1937) gave the length of the female as 1.28 mm and of the male as 0.97 mm; Sars (1901) as 1.30 mm for females. The southern popu- VOLUME 103, NUMBER 1 v Figs. 17-21. 145 ag Trichodiaptomus coronatus, males (17, 18, 20, 21 from Lagoa Bonita; 19 from Represa Des- coberto): 17, Leg 5, posterior; 18, Detail of left endopod, posterior; 19, Detail of left exopod, posterior (stippling indicates area of proximal hairy pad); 20, Leg 5, right-lateral; 21, Left exopod 2, medial. lations did not include individuals with short thoracic wings and/or prominences on the left side of the genital segment, originally named D. melini by Thomasson (1954) and termed “‘Females II-IV’’ by Brandorff (Brandorff et al. 1982). All southern females observed, both by Wright (1937) and in the present study, correspond to Brandorffs “Female I,” with symmetrical genital seg- ment and long, posteriorly directed, pointed wings; except for the few anomalous indi- viduals noted here. Brandorff in Brandorff et al. (1982) noted that Type I females tended to have more lateral spines on pediger 4 than did other morphs. Among the southern populations, females from Lagoa Formosa had signifi- cantly (P < 0.01; t = 5.5, df= 114) fewer spines on each side (4-10, mean 7.1) than females from Lagoa Bonita (6-13, mean 8.8). Lagoa Formosa females also had fewer, al- though not significantly fewer spines than females from Rep. Descoberto (7-11, mean 8.6) and Pirapora (9-11, mean 9.3). These latter three populations did not differ sig- nificantly among themselves in spine num- ber. Males from the three populations ex- amined (Bonita, Descoberto, Formosa) did not differ significantly in spine number, having 4-8 spines on each side. In both sexes, 146 spine number tended to be about equal on each side, differing by no more than 4. On females from S40 Paulo, Sars (1901) ob- served “‘about 12 denticles,” while Wright (1937) noted 8-12 spines. Matsumura-Tundisi (1986, Amazon) fig- ured the antennule of the female as reaching only to midlength of the genital segment; Wright (1927, Amazon) and Sars (1901, Sao Paulo) to midlength of the anal somite. Brandorff (1972), Matsumura-Tundisi (1986), and Wright (1927) figured the exo- pod 3 of leg 5 of the female as distinct from exopod 2. Matsumura-Tundisi (1986) and Thomasson (1954) figured the endopod of leg 5 of the female as a single article. These are probably lapses of observation. It appears that the unarticulated spines on articles 10, 11, and 13 of the right an- tennule of males are more prominently de- veloped in Amazonian populations. Wright (1937:78) mentioned differences “particu- larly in the spination of the right antenna of the male” between Sao Paulo and Am- azonian specimens. His figure ofa male from Sao Paulo (op cit.: pl. II, fig. 6) shows rel- atively short spines on articles 10, 11 and 13, similar to those from the southern high- lands (Fig. 15). Sars (1901: pl. III, fig. 15) also figured a short spine on article 13. Al- though Wright did not figure these spines from Amazonian males, he mentioned (1927:90) that the process of “‘the fifteenth segment” (probably article 13, since more distal articles have no processes) extends well past the middle of the succeeding ar- ticle, similar to the structure of the male from Santarém deposited at NMNH (Fig. 16). Figures of Amazonian specimens by Brandorff (1972) and Matsumura-Tundisi (1986) also show long unarticulated spines similar to Fig. 16. Distribution and ecology. —Brandorff (1976) noted the broad distribution of T. coronatus from the Brazilian Amazon to Sao Paulo State, a range further enlarged north- wards by Dussart’s (1984) record from the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Orinoco Delta (Fig. 22). In spite of this un- usual latitudinal distribution, 7. coronatus seems to be a species of narrow habitat re- quirements (Table 1). Although it has been found in both lentic and lotic waters, avail- able habitat data clearly indicate that it is restricted to warm, acid waters of low con- ductivity and extreme oligotrophy. Thus it has been most frequently collected from, but is not limited to Amazonian “black”’ waters (high in humic acids). Most of the localities, particularly in Amazonia, have been shallow streams and floodplain (var- zea) lakes; similarly, Lagoas Bonita and Formosa are small, shallow ponds less than 10 ha in area, with extensive macrophyte beds. The ponds at Pirapora are also shal- low, with much macrophyte coverage (M. B. G. e S. Dabés, pers. comm.). Santo An- tonio do Descoberto and Guarapiranga seem to be unusual habitats for this species, being large reservoirs; at least the former, in which T. coronatus was the only diaptomid species at the time of collection, has no significant macrophytes. Possibly 7. coronatus is a more littoral than pelagic species, particu- larly when in competition with other di- aptomids. Occurring only in Guarapiranga Reser- voir out of 41 water bodies examined in eastern Sao Paulo in 1935 (Wright 1937), the species appears to have disappeared from Guarapiranga and has been recorded re- cently from none of the now mainly eutro- phic reservoirs in that state (Sendacz et al. 1985). Sao Paulo reservoirs are now char- acterized by, among other species, Scolo- diaptomus corderoi and Thermocyclops de- cipiens (Kiefer), which are typical inhabitants of highly productive systems (Reid 1987, 1988, 1989; Sendacz et al. 1985). Of numerous water bodies examined in the central Brazilian highlands, those in which T. coronatus is present include sev- eral of the most oligotrophic but none of the more productive systems. Therefore, it seems to be a useful indicator species, and VOLUME 103, NUMBER 1 147 Fig. 22. Records of Trichodiaptomus coronatus (G. O. Sars) in eastern South America. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 148 ‘(p861) Youn, Wor sus » -oned ORS ‘dS ‘eed ‘Vd ‘sIBIOD SPU “OW ‘setop ‘OD ‘eiope, oMsiq “Ad ‘SeUOZEUY ‘JV ‘SUONLIAIIQGE 9381S ULITIZEIE q ‘(soqed ‘S$ 9°D ‘a ‘W Aq porddns spuod viodeig 10j eyep jeyuswuostAUs) 10daI Juasoid *] | ‘(9961) IsIpunL-einuins}eW “OT “(pR6l) Wessnd ‘6 (7861) Te 19 Hropuesg 8 (8161) HropuLI ‘LZ (EL6I) OUTPAIED F MIODID “9 “(ZL61) H1OPURIA “6 (C61 ‘PS61) UOSsEMIOYL “p (LET) THBEIAA “€ “(LZ61) TSM “C (1061) SIPS ‘T= o1ues10 — — 09 87 MoOTTeYS spuod ule[dpooy (11) OW ‘erodestg ‘odiD ‘ourred ‘oyoe, Svose Ty Aejo/pues Ieg]o Ol — C7 Z puod (11) OD ‘esouri0,j voseT o1ues10 regjo Ol 69 _ iG puod (11) Ad ‘e1uo0g voseT Aeyo prqiny Ol = Gil Ol HOAIOSOI (11) AG/OD ‘O1seqoosaq Op orugiuy o1URS “doy = yoriq 8-€ «MOL,, = = oP] (01) NV ‘OureIsSLID 037 = - = _ _ = WIevdI1S (6) BjanzousA “e1[9qd ODOULIO ‘elent) OUR) pues yortq 6 ORS = Fig. 2. Fabricinuda bikinii from the Bikini Atoll (AHF 207): A-C, Dorsal, lateral (right side) and ventral views, respectively, of the anterior end; D, Posterior end, dorsal view; E, Left half of branchial crown, inner margin; F, Frontal view of anterior peristomial margin, dorsal margin at bottom; G, Pseudospatulate setae from setiger 7; H, Thoracic uncinus from setiger 3; I, Abdominal uncinus from setiger 9. VOLUME 103, NUMBER 1 167 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 168 VOLUME 103, NUMBER 1 9 ee &\ Ay) @, A 169 0.1 mm A Bees) 0.02 mm B-D Fig. 4. Fabricinuda bikinii from the Aldabra Atoll (USNM 121995): A, Posterior end, dorsal view; B, Pseudospatulate seta from setiger 6; C, Thoracic uncinus from setiger 5; D, Abdominal uncinus from setiger 10. elongate, narrowly hooded; 4—5 per fascicle. Inferior thoracic notosetae of setiger 2 short elongate, narrowly hooded; 2 per fascicle. Setigers 3-8 with pseudospatulate setae; 2 per fascicle (Fig. 2G). Abdominal neuro- setae modified, elongate, narrowly hooded; 3-4 per fascicle. Thoracic acicular uncini in straight or irregular rows; 5—11 per fascicle (Fig. 2H). Abdominal uncini with 8-9 teeth in profile, 3-4 teeth per row; manubrium slightly expanded proximally, base qua- drangular, about same length as dentate re- gion (Fig. 21). All specimens examined un- —_— pigmented; body opaque. Tubes loosely constructed of calcareous sand grains; about same length as animals. Methyl green stain- ing darkest on ventrum of posterior peri- stomial ring, ventral anterior margin of se- tiger 1, ventrally and laterally on setigers 2— 5; staining distinctly increasing on setigers 6-8; abdominal setigers and pygidium staining darkest; all other areas staining very lightly. Remarks. —The material of Fabricinuda bikinii examined here extends the species distribution from the Bikini Atoll in the Pa- Fig. 3. Fabricinuda bikinii from the Aldabra Atoll (USNM 121995): A-C, Dorsal, lateral (left side) and ventral views, respectively, of the anterior end; D, Lateral view (left side) of slightly contracted specimen; E, Proximal region of dorsalmost radiole, inner margin; F, Frontal view of anterior peristomial ring margin, dorsal margin at bottom. 170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cific Ocean to the Aldabra Atoll in the In- dian Ocean. Because there are minimal dif- ferences between specimens from both areas (cf., Fig. 2, based on type material, and Figs. 3, 4, based on Aldabra material), I am re- luctant to view them as separate species. In general, specimens from the Aldabra Atoll tend to be more slender and elongate than what is seen in the type material. Sim- ilarly, the branchial crown of specimens from Aldabra may be relatively shorter, comprising only one-fifth the total body length. The number of individuals in the type series is, however, considerably fewer. The only marked difference between specimens from both areas is seen with methyl green staining: there are no distinct staining patterns on the anterior peristomial ring in the type material, whereas in the Aldabra material there are a pair of dark rectangular patches laterally and a dark patch ventrally on the anterior peristomial ring. This condition in the Aldabra material is similar to what has been seen in some spec- imens of Fabricinuda trilobata (Fitzhugh 1983; see below). While differences in stain- ing patterns may be indicative of regional, populational, or species-specific variation, the differences seen in the present material may also be a function of age and preser- vation. Facricinuda bikinii is unique in that the mouth has been shifted dorsally and in front of the attachment point of the branchial lobes to the peristomium. In addition, the point of attachment for the branchial lobes is reduced to a narrow, transverse ridge just behind the mouth. As a result of this latter condition, the branchial lobe bases sit large- ly unattached over the anterior end; only the dorsal margins of the lobes are con- nected to the peristomium. An intermediate condition is seen in F. trilobata (Fitzhugh 1983; see below) in that the crown attach- ment has been shifted dorsally to a lesser degree but with lobes completely attached to the peristomium and the mouth in the typical sabellid position. As a result of this shift in crown attachment in these two species, the anterior margin of the anterior peristomial ring is exposed ventrally to some extent as a horizontal or oblique shelf. This shelf-like condition was described in part by Hartman (1954) for F. bikinii. The abdominal uncinus figured by Hart- man (1954: fig. 178B; compare with Figs. 21 and 4D herein) is unusual in that the manubrium is slender and gently curved back, similar to the handle of thoracic un- cini. This misinterpretation of the manu- brium was probably due to examining only uncini which were still lying side by side in the fascicle and not in true profile. Fabricinuda trilobata (Fitzhugh, 1983), new combination Fig. 5 Fabriciola trilobata Fitzhugh, 1983:276- 284, figs. 1, 2, 3a—c.— Uebelacker, 1984a: 54-20, 54-22, figs. 54-15, 54-16. Additional description. —Dorsal and ven- tral lips absent. Ventral filamentous ap- pendages vascularized, unbranched (Fig. 5A), slightly wider than pinnules; width uni- form except for basal swelling of margin adjacent to dorsal radiole; distal end blunt; surface with minute transverse wrinkles. Anterior peristomial ring margin available for branchial lobe attachment reduced in size, attachment shifted dorsally (Fig. SB). Branchial lobes completely attached to peri- stomium. Mouth in usual sabellid position relative to branchial lobes. Narrow, exposed shelf-like region present below crown at- tachment area. Inferior thoracic notosetae of setiger 2 short, elongate, narrowly hood- ed; 1 per fascicle. Inferior thoracic notosetae of setigers 3-8 pseudospatulate setae (Fig. 5C); 1-2 per fascicle. Remarks. —Fabricinuda trilobata is dis- tinctive in the partial shifting dorsally of the branchial crown and mouth, resulting in an exposed, narrow shelf. As noted in Remarks under F. bikinii, this condition in F. trilo- bata is intermediate to what is seen in F. bikinii on the one hand and remaining species on the other. This suggests a possible VOLUME 103, NUMBER 1 171 Fig. 5. Fabricinuda trilobata (paratypes, A, C from USNM 74689, B from USNM 74691): A, Left half of branchial crown, inner margin; B, Frontal view of anterior peristomial ring margin, dorsal margin at bottom; C, Pseudospatulate seta from setiger 7. transformation series in which dorsal shift- ing of the branchial crown occurred prior to the mouth being displaced above the branchial lobes. Together with various workers in the past, Fitzhugh (1983) erroneously referred to the ventral filamentous appendages as “‘palps.”’ Fabricinuda pseudopalpa, new species Figs. 6, 7 Material examined.—California: Holo- type (USNM 122021) and 24 paratypes (USNM 122022), Dutch Harbor, San Nic- olas Island, Channel Islands, low intertidal, among roots of Phyllospadix and rocks, coll., R. Seapy, 5 May 1977. Description. —Holotype complete with 8 thoracic and 3 abdominal setigers; length 3.10 mm (0.70 mm comprising branchial crown), maximal width 0.23 mm. Branchial crown one-fourth to one-sixth total body length. Three pairs of radioles; distal ends filamentous, same width as pinnules. Ra- dioles each with 8-10 pairs of pinnules, all terminating at same height as radioles. Dor- sal lips erect (Fig. 6E), longer than wide, slightly tapered, distally blunt. Ventral fil- 172 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON amentous appendages absent. Branchial crown broadly attached to peristomium. Body cylindrical, elongate, tapering slightly in abdomen. Peristomial eyes black, round- ed to crescentic; pygidial eyes black, round- ed. Anterior rim of anterior peristomial ring with lobe on either side of dorsal midline; length and thickness of lobes variable; lobes on most specimens small (Fig. 6A), about same size as conical lobe-like structure above mouth (located between lobes); with vas- cular loops visible in peristomium below paired lobes; specimens with slightly longer lobes (Fig. 7A) with vascular loops extend- ing through most of length, lobes slightly thicker, surface smooth; less commonly, lobes very long, digitiform (Fig. 7B—D), with vascular loops extending most of length, surfaces relatively smooth, width uniform, distal ends rounded. Remainder of anterior peristomial ring margin a low ridge, slightly higher ventrally, smooth. Anterior and pos- terior peristomial rings of equal length or anterior ring slightly longer; each ring about as long as wide. Annulation between rings distinct ventrally and laterally. Setiger 1 slightly shorter than posterior peristomial ring, less than one-half length of setiger 2; setigers 3-7 successively longer, setigers 4— 7 each about 2—2.5 times longer than setiger 2; setiger 8 shorter, same length as setiger 3. Setiger 9 about one-halflength of 8, about as long as wide; setigers 10-11 successively shorter, narrower. Pygidium about same length as setiger 11; conical, bluntly round- ed (Fig. 6D). Superior thoracic notosetae elongate, narrowly hooded; 4—5 per fascicle. Inferior thoracic notosetae of setiger 2 short, elongate, narrowly hooded; 2 per fascicle. Setigers 3-8 with pseudospatulate setae; 1- 3 per fascicle (Fig. 6F). Abdominal neuro- setae modified, elongate, narrowly hooded; 4-6 per fascicle. Thoracic acicular uncini in straight single or irregular rows; 6-13 per fascicle (Fig. 6G). Abdominal uncini with 11-12 teeth in profile, proximal tooth slightly larger (Fig. 6H); 4-5 teeth per row; 17-22 uncini per fascicle; manubrium slightly expanded proximally. Branchial crown unpigmented; anterior peristomial ring light to dark brown, anterior margin of posterior peristomial ring similar; pigment diminishing posteriorly; posterior thoracic setigers and abdomen unpigmented, cream colored. Tubes loosely constructed with quartz sand grains, about same length as animals. No distinct methyl green staining patterns. Etymology. —The specific name refers to the extensible, palp-like quality of the dor- sal, anterior peristomial ring lobes. Remarks. —Fabricinuda pseudopalpa is very distinctive in that the vascular loops can extend into the dorsal lobes of the an- terior peristomial ring, with the lobes lengthening to varying degrees, resembling ventral filamentous appendages or palps. It is not known to what extent this condition might be a preservation artifact. Specimens with vascular loops not extending into the lobes resemble F. /imnicola (Hartman, 1951) and F. pseudocollaris, new species, with re- gard to the branchial crown not being shift- ed dorsally. Fabricinuda pseudopalpa dit- fers from these and other species of the genus in lacking ventral filamentous appendages but in having dorsal lips. Fabricinuda pseudocollaris, new species Figs. 8, 9 Material examined. —Florida: Holotype (USNM 122023) and five paratypes (USNM 65903), entrance to East Lagoon, Seahorse Key, silty sand, coll., J. H. Taylor, 24 Apr 1960. Description. —Holotype complete with 8 thoracic and 3 abdominal setigers; length 10.00 mm (branchial crown comprising 1.00 mm), maximal width 0.24 mm. Branchial crown one-ninth to one-tenth total body length. Crown broadly attached to peris- tomium, not displaced dorsally (Fig. 8B); with mouth between branchial lobes. Three pairs of radioles; distal ends filamentous, same width as pinnules. Radioles each with VOLUME 103, NUMBER 1 173 E G Fig. 6. Fabricinuda pseudopalpa (paratypes, USNM 122022): A-C, Dorsal, lateral (right side) and ventral views, respectively, of anterior end; D, Posterior end, dorsal view; E, Left half of branchial crown, inner margin; F, Pseudospatulate seta from setiger 4; G, Thoracic uncinus from setiger 4; H, Abdominal uncinus from setiger 9. 174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON B C D \ Fig. 7. Fabricinuda pseudopalpa (paratypes, USNM 122022): A, Dorsal view of anterior end of specimen with dorso-lateral lobes partially expanded due to extension of vascular loops; B-D, Dorsal, lateral (right side) and ventral views, respectively, of anterior end of specimen with extreme extension of vascular loops into dorso- lateral lobes. 6-7 pairs of pinnules, all terminating at same height; longest about two-thirds total ra- diole length. Dorsal and ventral lips absent. Ventral filamentous appendages vascular- ized, unbranched (Fig. 9A); distinctly wider than pinnules, width uniform throughout except for slight basal swelling along margin adjacent to dorsal radiole; distally blunt; same length as radioles or slightly shorter. Ventral filament surface relatively smooth or with minute transverse wrinkles; blood vessel very large. Body cylindrical, uniform- ly slender, thread-like, tapering slightly pos- teriorly. Peristomial eyes black, rounded to crescentic; pygidial eyes black, rounded. Anterior margin of anterior peristomial ring a low ridge all around except for low round- ed lobe to either side of dorsal midline (Fig. 8A-C); dorsal lobes smaller than median conical lobe above mouth; ridge of same height ventrally and dorsally, concave lat- erally (Fig. 8B). Anterior peristomial ring VOLUME 103, NUMBER 1 0.2 mm Fig. 8. views, respectively, of the anterior end; D, Posterior end, dorsal view. about 4 times longer than posterior ring; annulation between rings visible ventrally and laterally. Large pair of membranous lappets on ventrum of anterior peristomial ring (Fig. 8C), originating ventro-laterally on anterior half of ring; separated by wide gap, with posterior margin of gap developed as narrow Shelf. Lappets directed anteriorly with distal margins broadly rounded; ter- minating below anterior rim of anterior 175 B 0.1 mm A-C C SSeS) Fabricinuda pseudocollaris (holotype, USNM 122023): A-C, Dorsal, lateral (right side) and ventral peristomial ring; area of ring anterior to lap- pets slightly narrower in lateral view (Fig. 8B). Setiger | as wide as long, slightly longer than posterior ring; setigers 2-3 each about twice as long as setiger 1; setigers 4-5 each twice as long as 3; setigers 6—7 each twice as long as 4 or 5: setiger 8 slightly longer than 6 or 7. Setiger 9 about one-half length of setiger 8; setigers 10-11 successively shorter. Pygidium same length as setiger 11, 176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON B-D 0.02 mm Fig. 9. Fabricinuda pseudocollaris (paratypes, USNM 65903): A, Proximal region of right half of branchial crown, inner margin; B, Pseudospatulate seta from setiger 5; C, Thoracic uncinus from setiger 5; D, Abdominal uncinus from setiger 9. tapering to rounded end (Fig. 8D). Superior thoracic notosetae elongate, narrowly hood- ed; 4-6 per fascicle. Setiger 2 inferior tho- racic notosetae short, elongate, narrowly hooded; 2 per fascicle; setigers 3-8 with pseudospatulate setae; 2 per fascicle (Fig. 9B). Abdominal neurosetae modified, elon- gate, narrowly hooded; 3-4 per fascicle. Thoracic acicular uncini in irregular double rows; 8-12 per fascicle (Fig. 9C). Abdomi- nal uncini with 9-10 teeth in profile, 3-4 teeth per row; manubrium constricted be- VOLUME 103, NUMBER 1 low dentate region, expanded proximally with quadrangular base (Fig. 9D); 20-25 un- cini per fascicle in setigers 9-10; 18 per fas- cicle in setiger 11. Ventral filamentous ap- pendages yellowish to light brown; remainder of branchial crown unpigmented or dark brown on inner lobe margins. Con- ical lobe above mouth dark brown, pigment extending posteriorly along dorsal midline of anterior peristomial ring; region between ventral lappets of anterior ring dark brown; remainder of body cream colored. Tubes loosely constructed with quartz sand grains; same length as animals. Methyl green stain- ing greatest in ventral rectangular area of anterior peristomial ring below lappets, through posterior peristomial ring to ante- rior margin of setiger 1; ventrum of setigers 2—4 or 5 uniformly dark, remaining thoracic setigers lightly stained; abdominal setigers and pygidium dark. Etymology. —The specific name refers to the ventro-lateral anterior peristomial ring lappets, which give the impression of a col- lar. Remarks. — Fabricinuda pseudocollaris is very distinctive with its paired, ventral, membranous lappets on the anterior peri- stomial ring. The concomitant narrowing of this ring anterior to these structures appears somewhat similar to the more pronounced narrowing of the far anterior region seen in F. trilobata and F. bikinii. At this time I do not consider the two conditions to be ho- mologous. Relative to other species in the genus, F. pseudocollaris also has the longest anterior peristomial ring. Species Possibly Referable to Fabricinuda Day (1957) described the species, Fabri- cia mossambica, later referred to as Fabri- ciola mossambica by Day (1963, 1967), from South Africa. In the original description, Day (1957: fig. 8k) noted that contracted speci- mens have a truncated anterior end. The figure shows the anterior end of a specimen lacking the branchial crown and with the 177 anterior peristomial ring margin smooth. The attachment area for the crown is dis- placed dorsally somewhat and the anterior peristomial ring is indicated as longer than the posterior ring. Day’s figures of relaxed specimens (Day 1957: fig. 8g-j), however, show a rounded ventral lobe and no dorsal lobes on the anterior peristomial ring mar- gin; ventral filamentous appendages are present, but degree of vascularization is not mentioned. Unlike typical Fabriciola, this species lacks the membranous anterior peri- stomial ring collar and the manubrium of abdominal uncini is the same length as the dentate region. The general features de- scribed by Day (1957) for F. mossambica suggest that it is a species of Fabricinuda. Acknowledgments I gratefully acknowledge the following in- dividuals for comments on earlier drafts of this paper: Kristian Fauchald, Robert E. Knowlton, Diana L. Lipscomb, Meredith L. Jones, and Vicki A. Funk. I thank Brian Kensley and Kristian Fau- chald for supplying material collected from the Aldabra Atoll, and for allowing me to participate in the 1985 expedition. Leslie H. Harris, Collections Manager, Allan Han- cock Foundation, kindly provided loans of specimens. I thank Thomas H. Perkins for bringing the specimens of Fabricinuda pseu- docollaris to my attention and for his very thorough review of the manuscript. Literature Cited Banse, K. 1956. Beitrage zur Kenntnis der Gattungen Fabricia, Manayunkia und Fabriciola (Sabel- lidae, Polychaeta).—Zoologische Jahrbucher, Abteilung ftir Systematik, Okologie und Geo- graphie der Tiere 84:415-438. 1957. Die Gattungen Oriopsis, Desdemona und Augeneriella (Sabellidae, Polychaeta). — Vi- denskabelige Meddelelser fra Dansk Naturhis- torisk Forening, Kobenhavn 119:67—105. 1970. The small species of Euchone Malm- gren (Sabellidae, Polychaeta).— Proceedings of the Biological Society of Washington 83:387- 408. 178 Day.Jobe 19572 ihe polychaete fauna of South Af- rica. Part 4: New species from Natal and Mo- cambique.— Annals of the Natal Museum 14: 59-129. 1963. The polychaete fauna of South Africa. Part 8: New species and records from grab sam- ples and dredgings. —Bulletin of the British Mu- seum (Natural History) Zoology 10:383-445. 1967. A monograph on the Polychaeta of southern Africa. Part 2: Sedentaria.— British Museum of Natural History Publications 656: 459-878. Fitzhugh, K. 1983. New species of Fabriciola and Fabricia (Polychaeta: Sabellidae) from Belize.— Proceedings of the Biological Society of Wash- ington 96:276-290. —_._ 1988. A phylogenetic systematic analysis of several hierarchical levels within the Order Sa- bellida (Polychaeta: Annelida). Unpublished Ph.D. Dissertation, George Washington Uni- versity, Washington, D.C., xxvi + 604 pp. _ 1989. Asystematic revision of the Sabellidae- Caobangiidae-Sabellongidae complex (Anneli- da: Polychaeta). — Bulletin of the American Mu- seum of Natural History 192:1-104. Gitay, A. 1970. Areview of Augeneriella (Polychaeta: Sabellidae) and a new species from Northern Sinai.—Israel Journal of Zoology 19:105-109. Hartman, O. 1951. Fabricinae (feather-duster poly- chaetous annelids) in the Pacific. — Pacific Sci- ence 5:379-391. 1954. Marine annelids from the northern PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Marshall Islands.—Professional Papers of the U.S. Geological Survey 260-Q:619-644. __ 1959. Catalogue of the polychaetous annelids of the world. Part 2.—Allan Hancock Founda- tion Occasional Paper 23:355-628. _ 1969. Atlas of sedentariate polychaetous an- nelids from California.— Allan Hancock Foun- dation, University of Southern California, Los Angeles, 812 pp. Perkins, T. H. 1984. Revision of Demonax Kinberg, Hypsicomus Grube, and Notaulax Tauber, with a review of Megalomma Johansson from Flor- ida (Polychaeta: Sabellidae). — Proceedings of the Biological Society of Washington 97:285-368. Rullier, F. 1954. Essai de révision du genre Fabricia (Annélides Polychétes).— Bulletin Société Zool- ogique de France 79:14—-29. Uebelacker, J. M. 1984. Chapter 54. Family Sabel- lidae Malmgren, 1867, pp. 54-1, 54-43 in J. M. Uebelacker and P. G. Johnson, eds., Taxonomic guide to the Polychaetes of the Northern Gulf of Mexico. Vol. VII. Final Report to the Mineral Management Service, contract 14-12-001- 29091. Barry A. Vittor & Associates, Inc., Mo- bile, Alabama. Department of Invertebrates, American Museum of Natural History, Central Park West at 79th St., New York, New York 10024. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 179-186 DIPLOCARDIA KANSENSIS, A NEW EARTHWORM FROM KANSAS, WITH REDESCRIPTIONS OF D. RIPARIA SMITH AND D. FUSCULA GATES (ANNELIDA: OLIGOCHAETA: MEGASCOLECIDAE) Samuel W. James Abstract.—The definition of Diplocardia riparia is changed to include an intestinal origin in the seventeenth segment. Consequently D. prosenteris is synonymized with D. riparia. Diplocardia fuscula is now understood to have genital markings in the male field and a variable circulatory system. Diplocardia kansensis, closely related to D. riparia and D. fuscula, is described from Kansas and Arkansas material. It is proposed to refer to the posterior extension of the extra-esophageal vessels of Megascolecidae as clitellar vessels, rather than as posterior latero-parietals. Diplocardia riparia F. Smith was de- scribed from material collected in the flood- plain of the Illinois River near Havana, II- linois. It has since been recorded from several locations around the American Midwest, from Ohio to Oklahoma and Ne- braska (Olson 1928; Gates 1955, 1977; McNab & McKey-Fender 1955; Teotia et al. 1950). It is of minor economic impor- tance in southern Kansas and western Mis- souri, where it is collected and sold for bait. There have been questions about the status of certain specimens (see Gates 1955). In the course of examining material identified by the late W. R. Murchie and deposited in the National Museum of Natural History (USNM), it became clear that the original description (F. Smith 1895a) and Eisen’s (1899) additions contained information that was incorrect or misleading. Further, there were three distinct taxa represented in the USNM lots labelled Diplocardia riparia, one of which was clearly that species. One lot labelled D. riparia actually be- longs to D. fuscula Gates. In the course of examining these specimens and paratypes of D. fuscula, it was determined that the definition of D. fuscula needed to be altered. Another group is similar to worms pre- viously identified as Diplocardia prosenteris McNab & McKey-Fender in James (1984, 1988) and James & Cunningham (1989). These worms are described here as D. kan- sensis. Diplocardia prosenteris was de- scribed as a subspecies of D. riparia (McNab & McKey-Fender 1955) and was elevated to specific rank in Gates (1977). The species described below was deter- mined to be new based on Gates (1977) and examination of D. riparia collected by F. Smith. A complete description of D. riparia is included to provide a single source for morphological characteristics of this species. No other museum collections were searched for representatives of this or any related species. In placing these earthworms in subfamily Acanthodrilinae of the Megascolecidae, I follow the classification of Jamieson (1971) and Brinkhurst & Jamieson (1971). Diplocardia kansensis, new species Fig. 1A—C Diplocardia prosenteris. —James, 1982:38.— James, 1984:91.—James, 1988:479.— James & Cunningham, 1989:79. Material. —USNM 42137, limestone quarry east of Myron, Izard Co., Arkansas, 16 Apr 1962, W. R. Murchie; Konza Prairie 180 Research Natural Area, 1 km north of In- terstate 70 in Geary Co., Kansas, 25 May 1981, S. W. James; West Kimball Road, Manhattan Riley Co., Kansas, 12 May 1986, S. W. James; Schoolyard at Browning and Dickens Sts., Manhattan, Riley Co., Kan- sas; 27 May 1987; upland tallgrass prairie in Fort Riley, Geary Co., Kansas, 16 Sep 1986, S. W. James; Big Basin, Clark Co., Kansas, 7 Oct 1986, S. W. James. Description. — External characteristics: 55 to 100 by 2 to 3.5 mm, width at segment xxx, body cylindrical in cross section throughout, widest at segments vi to 1x, seg- ments 120 to 145. Setae closely paired throughout; setal formula AA:AB:BC:CD = 3:1:3.5:1.5. Prostomium tanylobous, peri- stomium smooth, biannulate segments iv plus one of iii or v, triannulate posterior to iv or v. Medium to dark brown anterior pigmentation, sometimes present along mid- dorsal line and on hindmost dorsal seg- ments, nephridiopores near D. First dorsal pore 9/10 or 10/11, spermathecal pores on small bumps on leading edges of viii and ix, just lateral to setae a. Ovipores in xiv, presetal; male pores in xix in male grooves which are in AB in xviii to xx; prostatic pores and penial setae at ends of grooves. Clitellum 1/2 xii, xiii to xviii or xix, saddle- shaped to annular. Unpaired mid-ventral genital markings viii, ix or 1x, x; rarely x, x1 or all of viii to xi; unpaired mid-ventral genital markings xvii, xx; paired genital markings postsetal in xvii, xx (all), paired, postsetal in one or more of xx to xxi, pre- setal in some Of xvili, xxi, XxX11; some paired 20/21. Penial setae bowed, 750 to 850 um by 12 to 17 um, tips tapering to a point slightly flattened around the edges (Fig. 1C). Internal characteristics: Septum 5/6 faintly muscled, 6/7 to 12/13 muscular, thickest in 7/8 to 9/10; last set of trans- septal muscles originating in xii. Alimen- tary canal with gizzards in v, vi; esophagous smooth externally with some chloragogen, vascularized in xii to xiii, internal texture PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pebbly changing to low longitudinal folds in Xv, xvi; no calciferous lamellae. Small glands on dorsal and lateral surfaces of esophagus in vii-ix. Intestinal origin at 16/17 or within XVii, intestinal constriction usually present just anterior to origin of simple typhlosole in xxi or xxii; typhlosole half to one-third of lumen diameter. Nephridia 2 per segment, stomate, with- out bladders, duct meeting body wall near D. Ascending tubule from nephrostome with blind “caecum.” Vascular system with subintestinal trunk, single dorsal trunk, these connected by lat- eral trunks in v to ix, lateral-esophageal hearts in x to xiii. Lateral trunks have lateral and septal branches from a point above at- tachment to subintestinal trunk, in 1x a large vessel from lateral trunk to body wall of segments ix to xii. Supra-esophageal vessel x to xiii, extra-esophageal v to xili, in xili branching out as clitellar vessel over x1i to XVii. Small vessels from extra-esophageal to septa and body wall in v to xil. Male sexual system holandric, testes and funnels free, coagulum usually filling x and xi; vasa deferentia from funnels under peri- toneum, entering body wall in xvii; seminal vesicles of ix divided into 2 to 4 lobes, of xii divided into numerous small ovoid lobes, vesicles of xii larger. Paired prostates in xviii and xx, slender, 2 to 3 times longer than the delicate kinked ducts, extending through several segments. Ovaries and funnels in xiii, free, ovaries fan-shaped or globular cluster of numerous strings. Spermathecae (Fig. 1B) in viii, 1x, variable ampulla and diverticular shape. Diverticulum 4 to 5 lobed internally, or lobes slightly incised, sessile or ectal end free, di- verticulum attached to duct below base of ovoid ampulla. Holotype. —USNM 122285. Konza Prai- rie Research Natural Area, 1 km north of Interstate 70 in Geary Co., Kansas, 25 May 1981, S. W. James. Paratypes (3).—USNM 122286. Konza VOLUME 103, NUMBER 1 Fig. 1. 181 1 OTR TAA huni tne X eee nnn Wiens XV D Diplocardia kansensis new species: A, Ventral view of segments vii—xxiii; B, Spermatheca; C, Tip of penial seta, scale line equals 10 wm. D. riparia: D, Tip of penial seta, same scale as C. D. fuscula: E. Spermatheca; F, Ventral view of segments vii to xxxi. Prairie Research Natural Area watershed 1D, eroded waterway, 8 Oct 1988, S. W. James. Remarks.—Diplocardia kansensis differs from D. riparia in the slightly spatulate tip of the penial setae, smaller body size, pres- ence of pre-clitellar genital markings, the large number of genital markings, the pres- ence of unpaired mid-ventral genital mark- ings, greater typhlosolar development and a more anterior typhlosolar origin. This species can be distinguished from other members of the genus by the following set of characters: spermathecae in vil, ix; last hearts in xiii, no calciferous lamellae, gen- ital markings pre- and post-clitellar, a typh- 182 losolar origin in or before xxii, and a tanylo- bous prostomium. In Gates (1977) or James (1990) it will key to D. fuscula Gates, from which it differs in size, disposition of genital markings, typhlosole, presence of clitellar vessels, and prostomium characteristics. Diplocardia kansensis inhabits upland areas rather than river banks, though on Konza Prairie it is only abundant in eroded areas at the upper ends of water courses. However, none of the drainages along which it has been collected bear water except after heavy rains. The Big Basin material was ob- tained on prairie ridgetops under Bison bi- son dung pats. The nature of the habitat of Murchie’s Arkansas material is unknown. James & Cunningham (1989) give the di- etary habits of this species, under the name D. prosenteris. Diplocardia riparia F. Smith, 1895 Fig. 1D Diplocardia riparia. —Smith, 1895a:138.— Smith, 1895b:286.— Eisen, 1899:166.— Eisen, 1900:175, figs. 143, 144.—Mi- chaelsen, 1900:325.—Smith, 1900:442.— Gates, 1955:242.—-McNab & McKey- Fender, 1955:128.— Murchie, 1967:534. Diplocardia riparia prosenteris. —McNab & McKey-Fender, 1955:128. Material.—Section 23, Range 18&19, Ohio Twp., Franklin Co., Kansas, in cattle feedlot near spring, 17 Apr 1987, S. W. James; bank of the Le Moine River near Ripley, Brown Co., Illinois, 6 Sep 1987, S. W. James; bank of the Chariton River near U.S. Hwy. 136, Schuyler Co., Missouri, 9 Apr 1988, S. W. James; bank of the Walnut River at fairgrounds, Winfield, Cowley Co., Kansas, 19 Sep 1987, S. W. James; bank of Cedar Creek, SW of Fairfield, Jefferson Co., Iowa, 11 Oct 1987, S. W. James; River Rat bait shop, Manhattan, Kansas (allegedly from SE Kansas), 16 May 1987; bank of Grand River, near Freeman, Cass Co., Mis- souri, 12 Jun 1987, S. W. James; Minnesota Bait and Fly shop, Kansas City, Kansas, 12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Jun 1987; Angler’s Market, Harrisonville, Missouri, 12 Jun 1987; bank of Stranger Creek at Kansas Hwy 92, Leavenworth Co., Kansas, 22 Jun 1987, S. W. James; USNM 42142, Newlight, Tensas Parish, Louisiana, 16 Apr 1966, R. E. Tandy; USNM 42139, Illinois Hwy 149 W of Murphysboro, Jack- son Co., Illinois, 19 Apr 1963, W. R. Mur- chie; USNM 42143, Beaucoup Creek, N of Murphysboro, Jackson Co., Illinois, 11 Apr 1967, W. R. Murchie; USNM 42144, Prai- rie Creek, Benton Co., Iowa, 15 Apr 1963, W.R. Murchie; USNM 42145, Osage Fork at Rt. 5, Laclede Co., Missouri, 18 Apr 1963, W.R. Murchie; USNM 26409, Dirty Creek Bottoms, near Warner, Oklahoma, Apr 1954, V. L. Rounds; USNM 47995, Kansas, ‘‘various localities,» Nov 1917, Mr. Harrah (F. Smith collection); USNM 48035, Ha- vana, Mason Co., Illinois, 24 May 1894, F. Smith; USNM 47993, Havana, Mason Co., Illinois, Apr 1895, F. Smith; USNM 47994, bottomlands near Rankin Crossing Bridge, White Heath, Piatt Co., Illinois, 10 Apr 1915, F. Smith. Types lost. Description. — External characteristics: 85 to 170 by 4 to 6 mm, width at segment xxx, body cylindrical in cross section through- out, widest at segments vi to 1x, segments 126 to 206. Setae closely paired throughout; setal formula AA:AB:BC:CD = 3:1:4:1.4. Prostomium tanylobous, numerous longi- tudinal furrows in peristomium, biannulate ili or v, triannulate posterior to iv or v. Medium to dark brown segments iv plus one of anterior dorsal pigmentation, some- times present along mid-dorsal line and on hindmost segments, nephridiopores near D. First dorsal pore 9/10 to 12/13, spermathe- cal pores on small bumps on leading edges of viii and ix, just lateral to setae A. Ovi- pores in xiv, presetal; male pores in xix in male grooves which are in AB in xviii to XX; prostatic pores and penial setae at ends of grooves. Clitellum xii to xviil or xix, sad- dle-shaped or interrupted at mid-ventral line, paired genital markings postsetal in xvii (all), paired, postsetal in one or both of xx— VOLUME 103, NUMBER 1 xxl. Penial setae bowed, 700 to 900 um by 22 to 28 um, tips tapering to a conical, un- sculptured point (Fig. 1). Internal characteristics: Septa 5/6 to 12/ 13 muscular, thickest in 7/8 to 9/10; last set of trans-septal muscles originating in x1iil. Alimentary canal with gizzards in v and vi, esophagous smooth externally with some chloragogen, vascularized in xi to xiii, in- ternal texture pebbly changing to low lon- gitudinal folds by xv, xvi; no calciferous la- mellae. Intestinal origin at 16/17 or within XVli, intestinal constriction usually present just anterior to origin of simple low to ru- dimentary typhlosole in xxii to xxv. Nephridia 2 per segment, stomate, with- out bladders, duct meeting body wall near D. Vascular system with subintestinal trunk, single dorsal trunk, these connected by lat- eral trunks in v to ix, lateral-esophageal hearts in x to xii. Lateral trunks have lateral and septal branches from a point above at- tachment to subintestinal trunk, in ix a large vessel from lateral trunk to body wall of segments 1x to x11. Supra-esophageal vessel ix or 9/10 to xiii, extra-esophageal iii to x1i1, in xil1 branching out to body wall over xii to xvi. Small vessels from extra-esophageal to septa and body wall in v to ix. Male sexual system holandric, testes and funnels free, coagulum usually filling x and x1; vasa deferentia from funnels, under peri- toneum, entering body wall in xvii. Seminal vesicles of 1x divided into two or three lobes, of xii divided into numerous small ovoid lobes; vesicles of xii larger. Paired prostates in xviii and xx, long and slender, several times longer than the kinked ducts, extend- ing through several segments. Stout ducts lacking muscular sheen, meeting body wall just posterior to penial setal follicles. Ovaries and funnels in xii, free, ovaries fan-shaped or globular cluster of numerous strings. Spermathecae in viii, 1x, variable ampulla and diverticular shape. Divertic- ulum 3 to 6 lobed internally, slightly in- cised, ental end sessile, ectal end free, di- 183 verticulum attached to thickened portion of duct below base of ampulla. Remarks.—Types of D. riparia are not mentioned in Reynolds & Cook (1976:163). The most important difference between the material examined here and earlier de- scriptions is the intestinal origin in xvii, rather than xviii. It was repeatedly observed that relaxed specimens and those that had been starved prior to preservation appeared to have an intestinal origin posteriorly in XVll Or apparently in xvili. Specimens fixed with full guts, as were all of Murchie’s and mine, clearly showed an intestinal origin within xvii or at 16/17, which is commonly interpreted as in xvii. Sections made by F. Smith and now at USNM have collapsed intestines, the first segments of which have longitudinal furrows resembling the val- vular condition of the last portions of the esophagus. McNab & McKey-Fender (1955) used the segment of intestinal origin and spermathe- cal characters to define Diplocardia riparia prosenteris, but spermathecal variation within individuals and within populations is equal to that deemed of subspecific rank. This is not to say that consistent geographic variations in spermathecal details do not exist. However, the consistency of somatic morphology within the material examined argues against naming subspecific variants. Since D. riparia prosenteris McNab & McKey-Fender was distinguished from D. riparia primarily on the basis of an intes- tinal origin in xvii, I suggest that this sub- species, elevated to specific rank by Gates (1977), be synonymized with D. riparia. Diplocardia riparia prosenteris is also un- represented by types. Diplocardia fuscula Gates 1968 Fig. 1E, F Diplocardia fuscula. —Gates, 1968:22.— Gateswo7 7-216 Material.—USNM 42134, Greenwood, Caddo Parish, Louisiana, 16 Nov 1954, D. 184 Brumfield coll. USNM 38789, Negreet, Sa- bine Parish, Louisiana, 28 Mar 1965, R. E. Tandy, G. E. Gates, and J. M. Byrd, coll. (paratypes). Description (Greenwood material). —Ex- ternal characteristics: 154 to 193 by 5 to 7 mm, width at segment xxx, body cylindrical in cross section throughout, widest at seg- ments vi to ix, segments 120 to 145. Setae closely paired throughout; setal formula AA: AB:BC:CD = 4:1:5:2. Prostomium proepi- lobous, peristomium furrowed, segments biannulate ili, iv or iv only, triannulate posterior to iv. Traces of medium brown anterior dorsal pigmentation (bleached by preservative?), sometimes present along mid-dorsal line and on posterior segments, nephridiopores near D. First dorsal pore 10/ 11, spermathecal pores on small bumps on leading edges of viii and ix, just in the line of or median to setae a. Ovipores in xiv, presetal; male pores in xix in male grooves which are in AB in xviii to xx; prostatic pores at ends of grooves. Clitellum 1/2 xii, Xlii to xix, interrupted at mid-ventral line. Unpaired mid-ventral sucker-like genital markings 10/11, 11/12, 12/13; paired gen- ital markings postsetal in xvii (all), some- times in xx (Fig. 1). Penial setae vestigial, sigmoid, not markedly different from am- bulatory setae. Internal characteristics: Septum 5/6 faintly muscled, 6/7 to 12/13 muscular, thickest in 7/8 to 9/10; last set of trans- septal muscles originating in xiii or xiv. Al- imentary canal with gizzards in v, vi; esoph- agous smooth externally, covered with brown chloragogen, vascularized in xii to Xv, internal texture pebbly changing to low longitudinal folds by xv, xvi; ventral esoph- ageal typhlosole viii to xiv; no calciferous lamellae. Intestinal origin in xviii, intestinal constriction anterior to origin of simple typhlosole in xxiv or xxv; typhlosole one quarter of lumen diameter or less. Nephridia 2 per segment, stomate, with- out bladders, duct meeting body wall near PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON D. Ascending tubule from nephrostome with blind ‘‘caecum”’ at point of joining with main nephridial tubule. Vascular system with subintestinal trunk, single dorsal trunk, these connected by lat- eral trunks in v to ix, lateral-esophageal hearts in x to xiii. Segmental vessels of xiv to xvi not connected to ventral trunk; pos- terior latero-parietal vessels lacking. Lateral trunks of vi to viii have body wall and septal branches from a point above attachment to subintestinal trunk, of v, vi with body wall branches and vessels to gizzard; in ix a large vessel from lateral trunk to body wall of segments 1x to x11. Supra-esophageal vessel x tO Xili, extra-esophageal v to xii, origi- nating as small sub-pharyngeal vessels in i or i. Small vessels from extra-esophageal to septa and body wall in vi to xu, to giz- zards in Vv, V1. Male sexual system holandric, testes and funnels free, coagulum usually filling x and xi; vasa deferentia from funnels superficial, passing lateral to prostatic ducts, entering body wall at 18/19; seminal vesicles of 1x divided into 2 to 4 lobes, of xii divided into numerous small ovoid lobes, vesicles of xii larger. Paired prostates in xviii and xx, slen- der, 2 to 5X longer than stout, muscular ducts, glands in segments of origin or also extending into one or two adjacent seg- ments; duct diameter increasing ectally. Ovaries and funnels in xii, free, ovaries fan-shaped or globular cluster of numerous strings. Spermathecae (Fig. 1E) in viii, 1x, ovoid ampulla, variable diverticular shape. Diverticula | to 5 lobed internally, attached at middle by a short, broad stalk to lateral or anterior face of spermathecal duct, a muscle attaching ectal diverticular end to base of the preceding septum. Remarks. —These worms were identified as Diplocardia riparia by W. R. Murchie, probably on the basis of pigmentation and intestinal origin. Since D. riparia is now understood to have an intestinal origin in XVli, placing these specimens in D. riparia VOLUME 103, NUMBER 1 is inappropriate. There are other differ- ences, such as the locations of genital mark- ings and the size of penial setae. One specimen of D. fuscula reported by Gates (1968) has the same location and date as the material described herein, but the collector is (erroneously?) listed as “‘D. Brumble.”’ No mention was made of vari- ation that may be related to location of col- lection, and the holotype was chosen from the Negreet specimens. Differences between the Greenwood specimens and the para- types are (characteristics of the former are given): an intestinal origin in xvili, extra- esophageal trunks not extending to xiv and, segmental vessels of xiv to xvi not meeting the ventral trunk. However, the three para- types were variable in several characteris- tics, and two disagreed with the description given by Gates (1968): These two have pos- terior latero-parietal trunks in xiil and su- pra-esophageal vessels terminating in xiii. The third lacks posterior latero-parietals and its supra-esophageal vessel ended in xiv, and agrees with the species description in these and most other particulars. However it has post-clitellar genital markings, a character- istic shared with the other paratypes, but not with Gates’ description. The great majority of characteristics usu- ally considered of systematic importance in this genus are similar in the Greenwood and Negreet lots. The major difference—and in view of the vascular system variability in the D. fuscula paratypes—the only consis- tent difference, is the segment of intestinal origin. The collection locations are approx- imately 100 km apart and in separate river drainages. The two lots may be considered subspecifically distinct, but until more data can be collected on geographic variation in D. fuscula, they should be considered the same variable species. In view of the change in the definition of D. riparia proposed above, and the new data on D. fuscula, D. fuscula is now to be dis- tinguished from D. riparia by reduction or 185 loss of penial setae, frequent replacement of posterior latero-parietal vessels with large segmental connections from the dorsal ves- sel to the body wall and the ventral vessel, presence of unpaired pre-clitellar genital markings and a greater number of segments. Contrary to the original description, D. fus- cula are not characterized by lack of genital markings in segments xvii and xx, and vari- ability in intestinal origin and the blood cir- culation in clitellar segments are to be ex- pected. For the sake of clarity in future discus- sions of the vascular system in megascolec- ids, I propose to replace the cumbersome term “‘posterior latero-parietal vessel” with “‘clitellar vessel.’ This pair of vessels is the posterior extension of the extra-esophageal vessels. The posterior extent of branches of the clitellar vessel coincides closely with the termination of the clitellum. In the Puerto Rican Trigaster longissimus (Borges & Mo- reno 1990), the clitellum begins in xx and the clitellar vessel in xviii. This clearly in- dicates a functional connection between ex- ternal and internal characteristics, and no- menclature should reflect this. Acknowledgments This research was supported by a short- term visiting fellowship at the Division of Worms, National Museum of Natural His- tory, Smithsonian Institution. The assis- tance of Meredith Jones and Chery! Bright was essential to the project. The Maharishi International University Department of En- gineering generously made its scanning elec- tron microscope available for my use. Literature Cited Borges, S., & A. G. Moreno. 1990. Nuevas especies del genero Trigaster Benham 1886 (Annelida: Oligochaeta) para Puerto Rico.—Bolletino del Museo Regionale de Ciencia Naturale de Torino (in press). Brinkhurst, R. O.,& B.G. M. Jamieson. 1971. Aquatic 186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Oligochaeta of the world. University of Toronto Press, Toronto, xi + 860 pp. Eisen, G. 1899. Notes on North American earth- worms of the genus Diplocardia. — Zoological Bulletin 2:161-172. . 1900. Research in American Oligochaeta, with especial reference to those of the Pacific coast and adjacent islands.— Proceedings of the Cal- ifornia Academy of Sciences, Third Series 2:85- 276. Gates, G. E. 1955. Notes on several species of the earthworm genus Diplocardia Garman 1888.— Bulletin of the Museum of Comparative Zool- ogy at Harvard 113:229-259. . 1968. Louisiana earthworms. V. Diplocardia fuscula, n. sp.,an addition to an American genus (Acanthodrilidae, Oligochaeta, Annelida). — Proceedings of the Louisiana Academy of Sci- ences 31:22-26. 1977. More on the earthworm genus Diplo- cardia. —Megadrilogica 3:1—47. James, S. W. 1982. Effect of fire and soil type on earthworm populations in a tallgrass prairie. — Pedobiologia 24:37—40. . 1984. New records of earthworms from Kan- sas (Oligochaeta: Acanthodrilidae, Lumbrici- dae, Megascolecidae). — Prairie Naturalist 16:9 1- 95. 1988. The postfire environment and earth- worm populations in tallgrass prairie. — Ecology 69:476-483. . 1990. Oligochaeta: Megascolecidae and other earthworms from southern and midwestern North America. in D. L. Dindal, biology guide. Wiley, New York (in press). —., &M.R.Cunningham. 1989. Feeding ecology of some earthworms in Kansas tallgrass prai- rie.—American Midland Naturalist 121:78-83. Jamieson, B.G.M. 1971. A review of the megascole- coid earthworm genera (Oligochaeta) of Austra- lia. Part I— Reclassification and checklist of the megascolecoid genera of the world.—Proceed- ings of the Royal Society of Queensland 82:75- 86. McNab, J. A., & D. McKey-Fender. 1955. Studies in the genus Diplocardia (Oligochaeta: Megasco- lecidae). —The Wasmann Journal of Biology 13: 113-143. Michaelsen, W. 1900. Oligochaeta. Das Tierreich 10. R. Friedlander and Son, Berlin. xxix + 575 pp. Murchie, W. R. 1967. Chromosome numbers of some Diplocardian earthworms (Megascolecidae— Oligochaeta). — American Midland Naturalist 78: 534-537. Olson, H. W. 1928. The earthworms of Ohio.— Ohio Biological Survey 4:47-90. Reynolds, J. W., & D. G. Cook. 1976. Nomenclatura Oligochaetologica. University of New Bruns- wick Press, Fredericton, New Brunswick, x + 217 pp. Smith, F. 1895a. A preliminary account of two new Oligochaeta from IIlinois.— Bulletin of the IIli- nois State Laboratory of Natural History 4:138— 148. 1895b. Notes on species of North American Oligochaeta.— Bulletin of the Illinois State Lab- oratory of Natural History 4:285-297. 1900. Notes on species of North American Oligochaeta. III. List of species found in Illinois, and descriptions of Illinois Tubificidae. — Bul- letin of the Illinois State Laboratory of Natural History 5:441-459. Teotia, S. P., F. L. Duley, & T. M. McCalla. 1950. Effect of stubble mulching on number and ac- tivity of earthworms.—Nebraska Agricultural Experimental Station Research Bulletin 165, 20 pp. Department of Biology, Maharishi Inter- national University, Fairfield, Iowa 52556. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 187-196 REDESCRIPTION AND SYNONYMY OF NEPHTYS IMBRICATA GRUBE, 1857 (POLYCHAETA: NEPHTYIDAE) Takashi Ohwada Abstract.—The polychaets Nephtys imbricata Grube, 1857, Nephtys serra- tifolia Ehlers, 1897 and Nephtys serrata Hartman, 1953 are synonymized. The holotype of N. imbricata is redescribed. Nephtys imbricata was originally de- scribed by Grube (1857) as Nephtys, ap- pearing like polynoids due to the scale-like structures on the dorsum. These structures are exceptional in the family Nephtyidae, and N. imbricata has been known only by its short original Latin description which lacked a figure. Although its possible syn- onymy with Nephtys squamosa Ehlers, 1887 was suggested by Hartman (1950, 1959), its identity remained obscure until now. The holotype of N. imbricata, however, was found in the Zoological Museum, Univer- sity of Copenhagen by Dr. Mary E. Petersen, and she kindly provided me an opportunity to examine that specimen. Nephtys imbricata Grube, 1857 Figs. 1-4 Nephthys imbricata Grube, 1857:11. Nephthys serratifolia Ehlers, 1897:24—25, tab 1, fig. 13.— Ehlers, 1901:68.— Mon- ro, 1930:114-115, fig. 41a, b.—Monro, 1936:139-140.— Wesenberg-Lund, 1962: 89-94, figs. 35-39.— Hartman, 1964:106, pl. 32, fig. 7.—Hartmann-Schroder, 1965: 141-145, figs. 110-113.—Rozbaczylo & Castilla, 1974:202-203, fig. 7. Nephtys serratus Hartman, 1953:33-34, fig. 10a—g.— Hartman, 1964:106, pl. 32, figs. 55 ©5 Nephtys imbricata Wesenberg-Lund, 1962: 95.—Rozbaczylo & Castilla, 1974:195. Material examined. — Holotype of Neph- tys imbricata: Valparaiso, Chile, 18.1x.1841, H. Kroyer, coll. (Zoological Museum, Uni- versity of Copenhagen = ZMC unnum- bered; for measurement see the description below). Syntypes of Nephtys serratifolia: Be- tween Falkland Islands and Argentina (49°35’S, 64°43’W), 113 m, Rophaniel coll., 2 specimens (Zoological Museum, Univer- sity of Hamburg = ZMH V-1198; both in- complete posteriorly, 15.1 mm long for 44 setigers, 2.9 mm wide with parapodia, and 16.2 mm long for 51 setigers, 2.5 mm wide with parapodia). Holotype of Nephtys ser- rata: Port William, Falkland Islands (51°40’S, 57°41'W), 40 m, sand, small stones and algae, 4.vii.1902, Swedish Antarctic Expedition 1901-1903, (Swedish Museum of Natural History = SMNH 620; 53-setiger anterior fragment, 47.3 mm long, 5.8 mm wide with parapodia, and | 1-setiger middle fragment, 11.7 mm long). Description. —Holotype of Nephtys im- bricata incomplete posteriorly, 48.1 mm long for 95 setigers; 3.2 mm wide with para- podia, 1.5 mm wide without parapodia at widest part of the body, about setiger 17, narrowing gradually to setiger 26. Body brown, lacking pigmentation in alcohol. Proboscis partly everted. Eyes absent (Fig. la). Anterior margin of prostomium nearly straight, thin, spatulate (Fig. 2a); lateral margins rounded, broadest between second antennae, extending into setiger | to nuchal organs; posterior margin with V-shaped middorsal projection extending to near pos- terior border of setiger 1; posterolateral 188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. only outlined): a, Anterior part, dorsal view (anterior end stretched laterally due to partially everted proboscis); b, 85th to 89th setiger, dorsal view (neuropodia omitted); c, Same, ventral view (notopodia omitted). margins joining lateral margins behind nu- chal organs. Anterior part between first an- tennae somewhat translucent, marked by intricate pattern (Fig. 2a) of opaque tissue. First antennae tapered, pointed, continuous with frontal margin of prostomium, direct- ed anterolaterally. Second antennae, stout, Nephtys imbricata, holotype (ZMC unnumbered; setae omitted; b, c, translucent scale-like lamellae conical, pointed, much stouter than first ones, nearly twice as long, well behind Ist antennae and halfway on ventral surface of prostomium near lateral margins, directed anterolaterally. Nuchal organs everted at posterolateral corners of prostomium. Proboscis (on the basis of the holotype of VOLUME 103, NUMBER 1 189 Fig. 2. Nephtys imbricata, holotype (ZMC unnumbered; a, setae omitted; b—d, tips of setae omitted): a, Anterior end, dorsal view (stretched laterally due to partially everted proboscis; anterior part of prostomium bending downward. dc—dorsal cirrus); b, Left neuropodium, setiger 7, dorsolateral view; c, Same, setiger 15, ventral view; d, Same, setiger 90, dorsolateral view. Nephtys serrata) terminally with 20 bifid papillae and single small triangular mid- dorsal and midventral papillae. Subtermi- nal papillae rapidly decrease in size towards base of proboscis, distalmost 2 or 3 arranged in 22 longitudinal rows, longer than or sim- ilarly long as bifid papillae, thereafter re- duced to 14 rows proximally, 3—5 in each row. No middorsal or midventral papilla. Proximal surface smooth. Setiger 1 modified (Fig. 2a); notopodia reduced, with short rounded postacicular lobe in front of, and rounded preacicular lobe behind rather rounded acicular lobe; acicular lobe supported by acicula whose tip curves outward. Dorsal cirri tiny, slender, pointed, arising anterolaterally from outer side of acicular lobes near base between preacicular and postacicular setal fascicles (Fig. 2a—dc), hardly recognizable. Neuro- podia of setiger | on anterior margin, lateral to prostomium, with thin rudimentary acic- ular lobes projecting anteriorly; left acicular lobe supported by thin acicula whose tip curves inward. Ventral cirri large, long, ta- pered, pointed, lateral to neuropodia, con- tinuous with anterior setigerous margin, di- rected posterolaterally. Notosetae of setiger 1 include preacicular barred (laddered) cap- illaries and postacicular very finely serrated slender capillaries; neurosetae long, thin, smooth capillaries, surrounding acicular lobes. Parapodia generally similar throughout from setiger 2 (Figs. 1, 3) except that bran- chiae (interramal cirri) lacking on setiger 2. Both rami with preacicular and postacicular setal fascicles (Fig. 3). Both notopodial and neuropodial acicular lobes conical and pointed (Figs. 1, 2b—-d, 3). Acicula termi- 190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Nephtys imbricata, holotype (ZMC unnumbered; tips of setae omitted): a, Left parapodium, setiger 7, anterior view (tip of notopodial postsetal lamella bending posteriorly); b, Right parapodium, setiger 28, anterior view (neuroacicula broken near its tip); c, Same, posterior view (preacicular setae omitted); d, Right parapodium, setiger 78, anterior view. VOLUME 103, NUMBER I nates dorsal to the tip of acicular lobe in notopodia, and in the tip of acicular lobe in neuropodia. Notopodial presetal lamellae thin and broad, extending from lower mar- gins of notopodia, shorter than acicular lobes, projecting almost to the level of the tip of notoacicula, distally rounded on an- terior parapodia (Fig. 3a, b), triangularly pointed on posterior parapodia (Fig. 3d). Small lamellae dorsal to notopodial presetal lamellae (Figs. la, 3a, b), thin and rounded, posteriorly reduced and becoming much smaller and hardly recognizable (Figs. 1b, 3d). Preacicular setae appear mostly beyond dorsal margins of presetal lamellae, usually slightly cover small suprapreacicular la- mellae (Fig. 3a, b, d). Neuropodial presetal lamellae thin, extending from upper mar- gins of neuropodia, rather triangularly on anterior parapodia (Figs. 2b, 3a) and rather truncately on middle and posterior para- podia (Fig. 3b, d), similarly long as acicular lobes on anterior neuropodia (Figs. 2b, 3a) and shorter than acicular lobes posteriorly (Figs. 2d, 3b, d). Inconspicuous thin broad lamellae ventral to neuropodial presetal la- mellae (Figs. lc, 2c, 3a, b, d), much shorter than presetal lamellae. Preacicular setae ap- pear mainly beyond ventral margins of pre- setal lamellae on anterior and middle neu- ropodia (Figs. 2c, 3a, b), partly cover infrapreacicular lamellae on anterior para- podia (Fig. 3a). Notopodial postsetal lamellae thin, ex- tending further than acicular lobes from seg- mental wall above and behind acicular lobes; small on setigers 2—5 or 6 following the small size of parapodia, increasing in size on an- terior setigers (Fig. 3a—c), thereafter becom- ing smaller and narrower posteriorly (Fig. 3d). Notopodial postsetal lamellae ligulate with rather rounded tip on setigers 2-10 on right side and setigers 2—13 on left side (Figs. la, 3a), especially rounded on setigers 2—4 (Fig. la); ligulate with triangular tip from setiger 11 on right side and setiger 14 on left side (Fig. 3b, c); becoming tapered pos- 191 teriorly as lamellae becoming narrower (Fig. 3d). On ligulate notopodial postsetal la- mellae with triangular tip, often a few (mostly one, sometimes two, rarely three) incisions on dorsal margin and/or one (rare- ly two) incision on ventral margin (Fig. 3d); postsetal lamellae with incision(s) occurring more often posteriorly among entire ones (Fig. 1b); no clear regularity in the change of the number of incisions along body. Neu- ropodial postsetal lamellae thin, larger than notopodial ones, extending from behind acicular lobes. Neuropodial postsetal la- mellae ligulate, entire with rather triangular tip on setigers 2—1 1 on right side and setigers 2-14 on left side (Fig. 3a); one incision de- velops from setiger 12 on right side and setiger 15 on left side (Figs. 2c, 3b, c); second incision occurs from setiger 36 on right side and setiger 43 on left side (Fig. 3d), incisions more conspicuous posteriorly; third inci- sion appears from setiger 87 on right side and setiger 83 on left side (Fig. Ic); inci- sions, however, inconspicuous thereafter. Although the number of incisions tends to increase along the body, postsetal lamellae without incision and with two incisions among those with one incision, and postse- tal lamellae with one incision and with three incisions among those with two incisions. Small lobe on superior edge of neuropodia (Figs. la, 2b, d, 3a—d). Dorsal cirri thin, long and narrow, triangularly elongate, extend- ing laterally from basal part of branchiae behind acicular lobes, no longer than neu- ropodial postsetal lamellae on anterior se- tigers (Figs. la, 3a), longer than any lamella on middle and posterior setigers (Figs. 1b, 3b-d). Branchiae first present on setiger 3, fully developed by setiger 8 (Fig. 3a), thereafter not reduced (Fig. 3b, c) even on last segment of the incomplete holotype (Fig. 3d). On right side, branchia appears as short process on setiger 3, elongates on setiger 4, begins to recurve on setiger 5, and completely re- curves on setiger 6, thereafter gradually in- 192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.1mm a b C Fig. 4. Nephtys imbricata, holotype (ZMC unnum- bered): a, Tip of rather short finely serrated neuropodial preacicular capillary seta, 78th right parapodium, an- terior view; b, Subdistal part of long finely serrated notopodial postacicular capillary seta without spur, same, anterior view; c, Basal part of long finely serrated notopodial postacicular capillary seta with spur, same, anterior view. creases in size to setiger 8. On left side, bran- chia emerges as very small, conical process on setiger 3, elongates but still very short on setiger 4, abruptly develops into re- curved form on setiger 5, thereafter gradu- ally increases in size to setiger 8. Scale-like thin translucent lamellae (Figs. la, b, 3) on dorsolateral surface of body from setiger 6, not fully developed until middle setigers where lamellae extend mainly posterolaterally over notopodia and partly overlap successive segments in im- bricate arrangement (Fig. 1b). Similar but smaller lamellae (Figs. 1c, 3b, c) on ventro- lateral surface of body from setiger 4, not fully developed until middle setigers where lamellae extend somewhat posterolaterally and slightly cover neuropodia and succes- sive segments (Fig. Ic). Both preacicular and postacicular setae occur in spreading fascicles, with postacicu- lar fascicles wider than preacicular ones (Fig. 3a, b, d); preacicular setae much fewer than postacicular setae; postacicular setae mostly thicker than preacicular ones. Preacicular setae mostly consist of barred capillaries; a few finely serrated capillaries (Fig. 4a) also occur at upper and lower end of preacicular fascicles (Fig. 3a, b, d), usually as thick as barred setae and similarly long or longer than barred setae. Postacicular setae consist of very long finely serrated capillaries (Fig. 4b), and in addition, on middle and pos- terior setigers, are several equally long finely serrated capillaries with spur on basal one- third to one-fourth (Fig. 4c) in middle of setal fascicles, and some short finely serrat- ed capillaries on anterior side of ventral half of neuropodial postacicular setal fascicles, thick as barred setae and similarly long or slightly longer than barred setae. No lyrate setae found. Remarks. —Nephtys imbricata is closely related to N. acrochaeta Hartman, 1950 and N. squamosa Ehlers, 1887. Nephtys imbri- cata, however, differs from N. acrochaeta in that branchiae are present from setiger 3 in the former and setigers 9-10 in the latter. Neuropodial postsetal lamellae are laciniate in N. acrochaeta and N. imbricata whereas they are entire in N. squamosa. The capil- lary setae with basal spur are present in the postacicular fascicles both in N. acrochaeta and N. imbricata, but absent in N. squa- mosa. Nephtys glossophylla Schmarda, 1861 from Chile also resembles N. imbricata in the possession of scale-like dorsal lamellae, laciniate neuropodial postsetal lamellae and finely serrated capillary setae with basal spur. Although it is probable that N. glossophylla is synonymous with N. imbricata, the for- mer is known only by its original descrip- tion (Schmarda 1861:90) and that is insuf- ficient to distinguish it from other species. VOLUME 103, NUMBER 1 193 Fig. 5. Anterior end and neuropodia of Nephtys serratifolia and N. serrata (a, b, N. serratifolia, syntypes (ZMH V-1198), c-e, NV. serrata, holotype (SMNH 620); tips of setae omitted): a, Anterior end, dorsal view; b, Left neuropodium, setiger 26, ventral view; c, Anterior end, dorsal view; d, Right neuropodium, setiger 46, ventral view; e, Left neuropodium, middle setiger, dorsal view. The type of N. glossophylla is deposited in the British Museum (Natural History), but the specimen has collapsed and its identity could not be clarified. Distribution.—Southern part of South America (both Atlantic and Pacific Coasts), 20-309 m, fine sand (Fig. 6). Synonymy of Nephtys imbricata Examination of the type specimens of Nephtys imbricata, N. serratifolia, and N. serrata revealed that all of them have scale- like lamellae, laciniate neuropodial postse- tal lamellae, and postacicular capillary setae with basal spur. Branchiae occur from se- tiger 3 in all type specimens, although JN. serrata was originally described to have branchiae from setiger 4 (Hartman 1953: 33). Geographically, N. serratifolia has been reported from both Pacific and Atlantic Coasts of southern South America, and the only known occurrences of N. imbricata and N. serrata (their type localities) are within 194 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 80°W __70°W 60° W 50° W 20°S SOUTH AMERICA 30°S 40°S 50°S Fig. 6. Geographical occurrence of Nephtys imbri- cata (asterisk—type locality of Nephtys imbricata, open circle—type locality of N. serratifolia, solid circle— occurrence reported as N. serratifolia, triangle—type locality of N. serrata). the range of distribution of N. serratifolia (Fig. 6). There are few differences between the type specimens of the three species (Table 1). While the presence of the tiny dorsal cirri on the 1st notopodia has been found in the holotype of N. imbricata (Fig. 2a), they have not been recognized in the syntypes of N. serratifolia or the holotype of N. serrata. This type of tiny dorsal cirri is barely rec- ognizable and is supposed to be easily bro- ken off (Ohwada 1989). Their apparent ab- sence in the syntypes of N. serratifolia and the holotype of N. serrata does not neces- sarily imply their absence in live animals. The most distinct difference between the type specimens examined lies in the mor- phology of the erect lobes on the superior edge of neuropodia. While these small lobes extend further than presetal lamellae on the anterior neuropodia of the holotype of N. imbricata (Fig. 2b), they are short through- out the length of the holotype of N. serrata (Fig. Se), and they are apparently absent in the syntypes of N. serratifolia. The previous descriptions of N. serratifolia show that the erect lobes develop to various extent in this species, and the degrees of their develop- ments in the type specimens of N. imbricata and N. serrata are within the range of their variations reported for N. serratifolia. This indicates that the difference in the mor- phology of the erect lobes between N. im- bricata and N. serrata has no specific im- portance. Their apparent absence in the syntypes of N. serratifolia is also explained Table 1.—Differences among the type specimens of Nephtys imbricata, N. serratifolia, and N. serrata. N. imbricata N. serratifolia N. serrata Ist dorsal cirri _ tiny, slender, pointed (Fig. 2a) small, ligulate, extend- ing beyond presetal lamellae in the ante- rior setigers (Fig. 2b, d) cirriform on anterior parapodia, thereafter foliaceous with ta- pered tip (Figs. lc, Erect lobes on neuropodia Ventral cirri not recognized not recognized cirriform on anterior parapodia, thereafter rather flattened in basal half, decreasing in width to vary- ing extent, distally becoming con- 2c) ical, tips pointed (Fig. 5b) not recognized small, short throughout the body (Fig. Se) rather flattened in basal half, suddenly decreas- ing in width to about %, distally becoming conical, terminating in fine columnar tip (Fig. 5d) SSS SSS SSS SSS SSS VOLUME 103, NUMBER 1 by the small size of the specimens. There appear to be some differences in the mor- phology of the ventral cirri among the type specimens examined (Table 1), but they are thought to be nothing more than intraspe- cific variations. Based on the close agreement among the type specimens of N. imbricata, N. serrati- folia, and N. serrata in the other characters, and the geographically overlapping ranges of these three species (Fig. 6), it is appro- priate to synonymize N. serratifolia and N. serrata with N. imbricata, the latter the se- nior synonym. The similarity in the prosto- mium morphology has been suggested to reflect phylogenetic closeness (Ohwada 1985), and the possession of the similar pat- terns of opaque tissue in the anterior trans- lucent part of prostomium in these three species (Figs. 2a, 5a, c) denies the possible morphological convergence between the species of different phylogenetic origins. Acknowledgments I am grateful to Dr. J. B. Kirkegaard and Dr. M. E. Petersen, Zoological Museum, University of Copenhagen, Dr. G. Hart- mann-Schroder, Zoological Institute and Zoological Museum, University of Ham- burg, and Dr. R. Olerod, Swedish Museum of Natural History, for the loan of the type specimens. Mr. A. I. Muir, British Museum (Natural History) kindly permitted me to examine the specimens in that museum. The manuscript benefited from the helpful sug- gestions by Dr. M. E. Petersen, and Dr. T. H. Perkins, Florida Department of Natural Resources, U.S.A. This work was supported by a Grant-in-Aid for Scientific Research from the Ministry of Education, Science and Culture, Japan. Literature Cited Ehlers, E. 1887. Report on the annelids of the dredg- ing expedition of the U.S. coast survey steamer Blake. Memoirs of the Museum of Comparative Zoology of Harvard College 15:vi + 335 pp., 60 pls. 195 1897. Polychaeten. Hamburger Magelhae- nische Sammelreise. Hamburg, Friedrichsen & Co., 148 pp., 9 pls. 1901. Die Polychaeten des magellanischen und chilenischen Strandes. Ein faunistischer Versuch. Festschrift zur Feier des Hundert- funfzigjahrigen Bestehens des K6niglichen Ge- sellshaft der Wissenschaften zu G6ttingen (Abh. Math.-Phys.). Berlin, Wiedmannsche Buch- handlung, 232 pp., 25 pls. Grube, E. 1857. Annulata Orstediana. Enumeratio Annulatorum, quae in itinere per Indiam occi- dentalem et Americam centralem annis 1845-— 1848 suscepto legit cl. A. S. Orsted, adjectis spe- ciebus nonnullis a cl. H. Kréyero in itinere ad Americam meridionalem collectis.— Videnska- belige Meddelelser fra Dansk naturhistorisk Forening 1 Kjobenhavn 1857:1-29. Hartman, O. 1950. Polychaetous annelids. Goniad- idae, Glyceridae, Nephtyidae.— Allan Hancock Pacific Expeditions 15:1-181. . 1953. Non-pelagic Polychaeta of the Swedish Antarctic Expedition 1901-03.—Further Zoo- logical Results of the Swedish Antarctic Expe- dition, 1901-03 4(2):1—83, 21 figs, 1 chart. . 1959. Catalogue of the polychaetous annelids of the world. Part 1.—Allan Hancock Founda- tion Publications, Occasional Paper 23:1-353. 1964. Polychaeta Errantia of Antarctica. — Antarctic Research Series 3:1-131. Hartmann-Schroéder, G. 1965. Die Polychaeten des Sublitorals. Zur Kenntnis des Sublitorals der chilenischen Kiiste unter besonderer Beriick- sichtung der Polychaeten und Ostracoden.— Mitteilungen aus dem Hamburgischen zoolo- gischen Museum und Institut 62 (Suppl.):59- 305. Monro, C. C. A. 1930. Polychaete worms.—Discov- ery Report 2:1—222. 91 figs. . 1936. Polychaete worms. IJ.— Discovery Re- port 12:59-198, 34 figs. Ohwada, T. 1985. Prostomium morphology as a cri- terion for the identification of nephtyid poly- chaetes (Annelida: Phyllodocida), with refer- ence to the taxonomic status of Aglaophamus neotenus. — Publications of the Seto Marine Bi- ological Laboratory 30(1/3):55-60. 1989. Redescription of Nephtys squamosa Ehlers (Polychaeta: Nephtyidae).— Proceedings of the Biological Society of Washington 102(1): 124-130. Rozbaczylo, N., & J. C. Castilla. 1974. La familia Nephtyidae en Chile (Annelida, Polychaeta). — Studies on the Neotropical Fauna 9:179-206. Schmarda, L. K. 1861. Untersicht der Ordnung Abranchia. Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise um die Erde 1853 196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON bis 1857. 1. Turbellarien, Rotatorien und An- Ocean Research Institute, University of neliden, Part 2, 164 pp., 22 pls., Leipzig. Tokyo 15-1. 1-Chome Minamidan Nakes b) b) b) bp} Wesenberg-Lund, E. 1962. Polychaeta Errantia. Re- ports of the Lund University Chile Expedition no-ku, Tokyo, 164 Japan. 1948-1949.—Lunds Universitets Arsskrift, N.F. Afdelning [2] 57(12):1-137. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 197-204 ANTRORBIS BREWERI, A NEW GENUS AND SPECIES OF HYDROBIID CAVESNAIL (GASTROPODA) FROM COOSA RIVER BASIN, NORTHEASTERN ALABAMA Robert Hershler and Fred G. Thompson Abstract. — An aquatic cavesnail from Coosa River Basin, Alabama, repre- senting a monotypic genus, is described (Antrorbis breweri, new genus, new species). Diagnostic features of the genus include a minute, planispiral to low- trochoid shell with apical microsculpture of spirally arranged low tubercles; blind, unpigmented animal; few ctenidial filaments; intestine with coils on lateral surface of style sac and in pallial roof; simple penis; capsule gland with ventral channel; and two sperm pouches. Antrorbis and three other North American cavesnail genera that also have a simple penis and capsule gland with ventral channel herein are placed in the Lithoglyphinae. Among the numerous discoveries of North American nonmarine mollusks made by Leslie Hubricht is a minute planispiral snail from subterranean stream in Manitou Cave, northeastern Alabama, which he as- signed (Hubricht 1940:35) to Horatia Bour- guignat, 1887. Hubricht did not describe the snail, and its affinities were uncertain in light of the determination by Hershler & Longley (1986) that other putative North American Horatia are not congeneric with this Euro- pean group. As part of ongoing review of North American cavesnails, the senior au- thor recently collected live examples of the Manitou cavesnail which we describe below as a new genus and species of Hydrobiidae. Antrorbis, new genus Diagnosis. — A minute-sized (2.7—3.0 mm) North American group characterized by a planispiral—low-trochoid shell with apical microsculpture of spirally arranged low tu- bercles. Operculum paucispiral, without ventral peg. Animal blind, unpigmented. Ctenidial filaments few (7-10). Central rad- ular teeth with single pair of basal cusps. Intestine coiling on right-lateral style sac and in pallial roof. Male with a simple penis. Females oviparous. Oviduct entering cap- sule gland, which has a ventral channel. Two sperm pouches present. Type species. —Antrorbis breweri, new species (by monotypy). Etymology. —Masculine, from the Clas- sical Greek, antrum, a cave, and orbis, a circle, and referring to subterranean habitat and discoidal shell of the snail. Discussion. —Among North American freshwater hydrobiids, the Lithoglyphinae group of epigean genera (Thompson 1984) and three subterranean forms (Pterides Pils- bry, 1909; Phreatodrobia Hershler & Long- ley, 1986; Holsingeria Hershler, 1989) share with Antrorbis the simple penis and capsule gland with ventral channel. Antrorbis re- sembles some Phreatodrobia which have planispiral to low-trochoid shells, but a number of other highly unusual features (i.e., wrinkled protoconch sculpture, absence of basal cusps on the central radular teeth, presence of an anterior coil of the capsule gland) occur among this group of Texas en- demics, mitigating against a close relation- ship with Antrorbis or any other known ge- 198 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nus. Holsingeria and Pterides, local endemics from the Powell River Basin in southwestern Virginia and Panuco River Basin in northeastern Mexico, respectively, share with Antrorbis the spirally arranged, tubercular protoconch sculpture; and al- though these three genera are well differ- entiated by various combinations of fea- tures from shell, operculum, intestinal coiling, and pallial oviduct complex, they appear to represent a natural group. The heterogeneous assortment of unusual character-states exhibited by these cave- snails makes it difficult to assess affinities among them, or with other hydrobiid groups, but nevertheless we are placing them in the Lithoglyphinae, the only subfamily that agrees with them in general soft anatomical features.! The lithoglyphines, as previously conceived (Davis & Pons da Silva 1984, Thompson 1984), were a morphologically compact group of genera characterized by a squat shell with large aperture, which ac- comodates a broad foot required for holding onto hard substrate in swift current. In light of recent studies on the anatomy of hydro- biid snails, and the discovery of additional new genera, shell shape (reflecting special- ization for a particular habitat) no longer can be considered a defining feature of the Lithoglyphinae. In the wake of the taxo- nomic changes proposed herein, the Lith- oglyphinae now parallels the hydrobiid subfamilies Nymphophilinae and Littori- dininae in that included genera vary from nearly planispiral to broadly trochoidal to elongate-slender in shell shape. The Litho- glyphinae are represented in South America and Europe, and are widely deployed on the North American continent where they in- habit lentic and lotic epigean habitats as well as subterranean streams. No litho- glyphines are known from brackish-water habitats. We suspect that the Lithoglyphi- ' The Hydrobuinae differ anatomically from this group in having a small pallial tentacle; a penial lobe; and a complex, pigmented renal oviduct. nae are an ancient freshwater group, but the fossil record offers no useful information in this regard because the subfamily is not identifiable on the basis of shell features. Antrorbis breweri, new species Manitou cavesnail Figs. 1-5 Horatia sp.—Hubricht 1940:35. Horatia micra. —Stein 1976:21. “Horatia.”’ —Burch 1982:270.—Hershler & Longley 1986:153, figs. 23k, 1, 28a. Material examined. -USNM 860429, holotype; University of Florida (UF) 135984 (4 specimens), USNM 860430 (9), para- types, RH and party coll., 6 Jun 1988.— USNM 860431 (16), RH coll., 16 Sep 1988.—USNM 860432 (10), L. Hubricht coll., 19 Jun 1957. All material from Man- itou Cave (Fig. 1), Little Wills Valley, Coosa River Basin, Fort Payne, AL (7.5 minute series), T. 7S, R. 9E, NE % section 18. Description. —Shell (Fig. 2; Table 1) dis- coidal, clear, transparent, 1.5—1.7 mm wide; height about half of width. Surface usually covered by moderately thick, yellow-orange periostracum. Whorls, 2.5-3.0, well round- ed, sutures deeply indented. Whorl expan- sion rate moderate. Translation of proto- conch and first teleoconch whorl highly variable, yielding impressive diversity of spire development. Aperture near-circular in outline, often slightly longer than wide, with adapical edge advanced. Inner lip thin, less curved than outer, adnate to small por- tion of body whorl, very slightly flared. Um- bilicus broadly open. Protoconch (Fig. 3a— d), 1.25 whorls, sculptured with raised tu- bercles arranged in numerous spiral rows. Tubercular sculpture weak or absent on tel- eoconch. Teleoconch having strong collab- ral growth lines. Operculum (Fig. 3e) thin, paucispiral, with 4.5 whorls. Ventral surface of operculum slightly convex, lacking peg development. Animal with 2.5 whorls. Pigment absent, except for some small black granules scat- VOLUME 103, NUMBER 1 199 Alabama 4000 _| Kilometers Fig. 1. Map (from Fort Payne Quadrangle, USGS 7.5 minute series, 1946 [revised 1983]) showing location of Manitou Cave. Radular (Fig. 4) formula: 5-1-4(5)/1-1, 4-1-4(5), 20, 13-15 (from SEM micrographs of paratypes). Central teeth (Fig. 4a) broadly trapezoidal with deeply excavated basal tered on stomach and digestive gland pos- terior to gonad. Ctenidium with 7—10 small, triangular filaments. Osphradium filling about a third of ctenidium length. 200 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. SEM micrographs of shells of Antrorbis breweri, new species: shell at upper left, USNM 860429, holotype (shell width, 1.53 mm); all others, USNM 860430, paratypes (printed to same scale). processes. Basal cusps of central teeth elon- gate, arising from edges of lateral angles. Style sac and stomach about equal in length (Fig. 5c). Stomach chambers poorly distin- guishable externally; posterior caecal cham- ber absent. Digestive gland of 0.5-0.75 whorls, consisting of two elongate masses covered with small, lobate swellings. Pos- teriormost lobe terminates slightly proxi- mal to tip of animal. Intestinal (In) coil on right lateral style sac surface a simple, “U- shape.” Coil in anterior pallial roof a more complex, “reversed-S shape” (when viewed dorsally). Testis a single, unlobed mass filling 0.5 whorl, slightly overlapping posterior stom- VOLUME 103, NUMBER 1 201 v Fig. 3. Shells, operculum, penis of Antrorbis breweri, USNM 860430: a—-d, SEM micrographs showing shell apex and microsculpture (scale bars = 200 um, 86 um, 176 um, 27 um); e, operculum (bar = 0.5 mm); f, penis (bar = 0.5 mm). 202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Radula of Antrorbis breweri, USNM 860431: a, Centrals (scale bar = 6.0 wm); b, Laterals (bar = 3.8 um); c, Inner marginals (bar = 4.3 um); d, Outer marginals (bar = 4.3 wm). ach. Seminal vesicle short, largely anterior to testis; connected just proximal to anterior tip of testis. Prostate gland elongate, largely pallial (ca. 80% of length); anterior vas def- erens exits from prostate tip. Penis (Fig. 3f) simple, blade-like, with terminal papilla; specialized penial glands absent. Ovary a single, unlobed mass (usually wholly posterior to stomach) filling <0.25 whorl, orange in color. Pallial oviduct (Fig. 5a) bipartite, overlapping style sac. Capsule gland (white) slightly smaller than albumen gland (clear). Oviduct (Ovi) with single, small coil on posterior left-lateral surface of albumen gland. Bursa copulatrix (Bu, Fig. 5b) pear-shaped, large, pressed against and partly posterior to albumen gland. Seminal receptacle (Sr) smaller, narrower, posi- VOLUME 103, NUMBER 1 Fig. 5. b, Left lateral aspect of bursa copulatrix and seminal receptable (rotated and slightly enlarged relative to “‘a’’); c, Right lateral aspect of stomach and associated structures. Ag = albumen gland; Bu = bursa copulatrix; Cg = capsule gland; In = intestine; Oes = oesophagus; Ov = ovary; Ovi = oviduct; Pst = posterior stomach; Sr = seminal receptacle; Sts = style sac; Vc = ventral channel of pallial oviduct. tioned along posterior edge of bursa copu- latrix. Ducts of sperm pouches short, en- tering oviduct just distal to coil. Ventral channel narrow relative to capsule gland width; opening of capsule gland broad, sub- terminal. Etymology.—Named after Dr. Stephen Brewer, the current owner of Manitou Cave, in recognition of his cooperation with and interest in this study. Habitat. —Manitou Cave, formerly a commercial cave, has been closed to the public since 1980. A large opening fitted 203 Morphology of Antrorbis breweri, USNM 860431: a, Left lateral aspect of female reproductive system; with a metal door serves as the entrance to the cave on the side ofa small hill. On ground level below the door a small (<1 m across), cool stream emerges amongst limestone rubble. Antrorbis breweri was absent from the epigean spring, but was collected from the uppermost portion of cave stream. Snails were found at a point where water cascaded from several narrow openings into a shallow (1-2 cm) pool in a small (ca. 4 m*), cement- lined, rectangular structure (which formerly served as the municipal water source for Fort Payne). Snails were collected (rarely) 204 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Shell measurements (mm) of adults of Antrorbis breweri. WH = number of whorls; SH = shell height; SW = shell width; LBW = length of body whorl; WBW = width of body whorl; AL = aperture length; AW = aperture width. USNM 860429 Das 0.98 1.53 0.83 1.07 0.68 0.53 (holotype) USNM 860430 3.0 0.85 1.69 0.72 1.19 0.62 0.63 (paratypes) DoD 0.88 1.67 0.77 1.15 0.61 0.61 2.75 0.80 1.66 0.68 1.17 0.59 0.57 Ded 0.80 1.79 0.63 1.22 0.62 0.62 from bottoms of loose bricks and natural breakdown littering the pool bottom. Other accessible reaches of cave stream were not searched. Scarcity of the snail also was men- tioned by Stein (1976). Acknowledgments We thank Dr. Brewer of Fort Payne for permission to collect in Manitou Cave. The assistance of staff of the Scanning Electron Microscopy Laboratory at National Mu- seum of Natural History (NMNH) also is appreciated. Ms. M. Ryan, Department of Invertebrate Zoology, NMNH, drafted the map. Literature Cited Burch, J. B. 1982. North American freshwater snails: identification keys, generic synonomy, supple- mental notes, glossary, references, index.— Walkerana No. 4:217-365. Davis, G. M., & M. C. Pons da Silva. 1984. Pota- molithus: morphology, convergence, and rela- tionships among hydrobioid snails. — Malacolo- gia 25:73-108. Hershler, R. 1989. Holsingeria unthanksensis, a new genus and species of aquatic cavesnail from east- ern North America. — Malacological Review 22: 93-100. —, & G. Longley. 1986. Phreatic hydrobiids (Gastropoda: Prosobranchia) from the Edwards (Balcones Fault Zone) Aquifer Region, south- central Texas. — Malacologia 27:127-172. Hubricht, L. 1940. A subterranean snail from an arte- sian well.—Nautilis 54:34-35. Stein, C. B. 1976. Gastropods. Jn H. Boschung, ed., Endangered and threatened plants and animals of Alabama.—Alabama Museum of Natural History, Bulletin Number 2:21-41. Thompson, F. G. 1984. North American freshwater snail genera of the hydrobiid subfamily Litho- glyphinae. — Malacologia 25:109-141. (RH) Department of Invertebrate Zool- ogy, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (FGT) Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 205-228 A NEW ISIDID OCTOCORAL (ANTHOZOA: GORGONACEA) FROM NEW CALEDONIA, WITH DESCRIPTIONS OF OTHER NEW SPECIES FROM ELSEWHERE IN THE PACIFIC OCEAN Frederick M. Bayer Abstract.—The status of the genera Jsidella, Acanella, and Lepidisis in the subfamily Keratoisidinae is discussed and the new species [sidella trichotoma and Acanella dispar are described and illustrated. New records of Acanella sibogae Nutting are presented and description of the species amplified and supported by new illustrations of colony, polyps and sclerites. Orstomisis cros- nieri, a new genus and species of Keratoisidinae, is described and illustrated. A new key to genera of Isidinae and Keratoisidinae is presented. Among the gorgonians from New Cale- donia obtained by M. Georges Bargibant of ORSTOM, Nouméa, is a large isidid colony superficially resembling a robust Keratoisis but branched dichotomously from the nodes rather than laterally from the internodes. The characters usually considered “‘diag- nostic’’ agree with those attributed to Jsi- della, but its growth form and other features are so divergent that assignment of the spec- imen to that genus must be considered ques- tionable. The first specimen of this species was re- ceived after a manuscript describing New Caledonian Isididae (Bayer & Stefani 1987a) had been submitted for publication. A de- scription of it was prepared for inclusion in another paper on Isididae (Bayer & Stefani 1987b) but was withdrawn when six more specimens were discovered among numer- ous isidids in a rich collection of deep-water gorgonians from New Caledonian waters re- ceived from the Muséum National d’His- toire Naturelle, Paris. As it is now clear that lack of research funding precludes further study of that collection, the unpublished original description has been revised for publication with supplemental data from the additional specimens and comparative in- vestigation of species of Isididae from else- where in the Pacific. In order to ascertain a taxonomically jus- tiflable position in the family Isididae for this unusual species, it was necessary to sur- vey all the genera of the subfamily Kera- toisidinae. These are: Keratoisis Wright, 1869; Isidella Gray, 1858; Acanella Gray in Wright, 1869; Lepidisis Verrill, 1883; and, possibly, Tenuisis Bayer & Stefani, 1987; the subfamilial affinities of Australisis Bayer & Stefani, 1987, remain to be determined and may not lie with the Keratoisidinae. The genera Keratoisis and Tenuisis, all species of which branch from the inter- nodes, need no further consideration in the present context. The genus I[sidella Gray, represented by two nominal species in the North Atlantic and the Mediterranean Sea, shares many morphological features with other genera of the subfamily Keratoisidinae, including Ac- anella Gray, Lepidisis Verrill and Keratoisis Wright. Keratoisis differs by branching from the calcareous internodes, and Lepidisis by its unbranched, whiplike, often spiral growth form. Acanella differs from Jsidella chiefly by its bushy colonial form that results from branching in whorls of 3—6 from the horny 206 nodes. Both species of Isidella are charac- terized by flattened, openly flabellate growth form produced by sparse, distant, dichot- omous branching from the horny nodes mostly in one plane. Muzik (1978:737) refers all species of Keratoisidinae branched from the organic nodes to the genus Jsidella Gray, 1858, thus synonymizing Acanella Gray, 1869, and in- corporating the branched Lepidisis longi- flora Verrill, 1883. This treatment is con- sistent with the comment made by Verrill (1883:18) that Lepidisis differs from Acanel- la “only in having the external layer of small scale-like spicula, both in the coenenchyma and on the calicles,”’ as the development of the superficial layer of small scales is in- consistent. Deichmann (1936:242), in com- menting about Verrill’s type specimen of L. longiflora, does not even mention scales, reporting only “‘a large number of flat, short, blunt rods, especially in the tentacles,” but does describe (p. 241) “‘small, flattened, nar- row rods or scales with rounded ends” in both coenenchyme and polyps of Lepidisis caryophyllia Verrill, the unbranched type species. Therefore, it seems quite certain that Verrill’s “scales” are the small, flat rods ob- served in the present material. The polyps of all species of Keratoisidi- nae have large spindles and/or rods in the body wall, often conspicuously projecting between the bases of the tentacles, and small, more or less flattened and sometimes thorny rods and/or double stars in the pharyngeal walls. In some species of Keratoisis there is a superficial layer of small, flattened, scale- like rods in the polyps and coenenchyme, so this character does not reliably distin- guish Lepidisis from Keratoisis. The quality of retractability of the polyps has long been used as a primary character distinguishing the subfamily Isidinae from the Keratoisidinae and Mopseinae (Studer [& Wright] 1887; Kiikenthal 1915, 1919, 1924; Bayer 1956). The polyps of Keratoisis, Acanella, Isidella, and Lepidisis (Kerato- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON isidinae) form permanently protruding, often prominent, columnar or conical ver- rucae incapable of retraction into the thin intervening coenenchyme. Although the polyps of Isis hippuris can be completely retracted within the thick coenenchyme, leaving only minute pores in the coenen- chymal surface, this is accomplished by folding the virtually sclerite-free tentacles inward over the mouth and closing the coe- nenchymal aperture over them. The polyps of Chelidonisis, the other genus comprising the Isidinae (sensu Kukenthal), cannot re- tract because the coenenchyme is very thin; the body wall, stiffened by numerous scler- ites, forms a permanently projecting ver- ruca. In contraction, the tentacles fold in- ward over the mouth, closing the verrucal aperture with the sclerite-filled bases of the tentacles (see Bayer & Stefani 1987b: fig. 30). In Kukenthal’s system, both are treated as retractile, comprising the subfamily Isi- dinae. The polyps of Muricellisis Kukenthal also are retractile, but are armed with sclerites arranged as a transverse crown (=collaret) and points (=operculum), hence “‘retrac- tile’ in the more restricted sense of with- drawal of anthocodia within anthostele (Bayer et al. 1983:11; Verseveldt & Bayer 1988:8). Because of this difference, Ktiken- thal (1915) proposed a separate subfamily Muricellisidinae to accommodate this one genus. When the character of “retractility” is considered in light of the morphology of Isis, Chelidonisis, Muricellisis, and Kera- toisis (+ Acanella, Isidella, Lepidisis), it is clear that the subfamily Isidinae as so de- fined is artificial. However, the genera Jsis and Chelidonisis share a different character that sets them apart from all other isidids and may, indeed, justify the subfamily Isidi- nae. Those two genera alone in the family have sclerites of basically radiate capstan form, with tubercular sculpture. In /sis they are 8-radiates, some larger at one end, form- VOLUME 103, NUMBER 1 ing clubs, some elongated to form spindles that may be coarse and pebble-like; in Chel- idonisis they are typical 6-radiates. At the risk of redundance in dealing with isidid classification, a revised subdivision of the family is proposed as follows: Key to the Genera of Isidinae and Keratoisidinae Sclerites in the form of 6- and 8-radiates, clubs, and tuber- culate spindles ....... ISIDINAE Coenenchyme thick, polyps not projecting ..... Isis Linnaeus Coenenchyme thin, polyps projecting as conical verrucae AN Mpa Chelidonisis Studer Sclerites in the form of more or less prickly rods or spindles Re eccsage awe Uf KERATOISIDINAE Colonies unbranched ight aed Beacons eae Lepidisis Verrill Colonies branched Branches originate on inter- NOGES Ea en) Keratoisis Wright Branches originate on nodes Polyps forming short, cylin- drical verrucae into which ten- tacles and oral part can be fully retracted; branching dichoto- mous, internodes short (up to 2 cm but mostly | cm or less); colonies multiplanar, flabel- late, compressed, trunk form- ing a massive calcified holdfast Pe peat ak cid Orstomisis, new genus Polyps forming cylindrical, conical or trumpet-shape ver- rucae, tentacles folding over the mouth but not retractile Branching dichotomous, tri- chotomous, or lateral, pre- dominantly planar, internodes hollow, long or very long (3.5- SHCT) een tan. nee Isidella Gray Branching verticillate, at least 1(4). 2(3). 3): 4(13). 5(6). 6(5). 7(8). 8(7). 9(10). 10(9). 11(12). 12(11). 207 in upper parts, colonies dense- ly or openly bushy, internodes solid, shorter, up to 2cm ... Verge ce Me ee eee Acanella Gray Sclerites in the form of flat plates, sometimes elongate and spindle-like but never with complex tubercular sculpture .... MOPSEINAE. (See Bayer & Stefani, 1987a:51; 1987b: 941.) 13(4). Isidella Gray, 1858 Isidella Gray, 1858:283; 1870:14.—Studer [& Wright], 1887:44.—Kukenthal, 1915: 118; 1919:564; 1924:414.—Deichmann, 1936:239.—Bayer, 1956:F222; 1981:941 (in key).—Carpine & Grasshoff, 1975: 107.— Bayer & Stefani, 1987a:51 (in key); 1987b:941 (in key). Isis. —G. von Koch, 1887:90. Diagnosis. —Isididae sparsely branched dichotomously or trichotomously from the nodes, usually in one plane, forming spindly colonies often of candelabrum form; inter- nodes long (up to 8 cm), hollow, longitu- dinally grooved, straight or nearly so; base of main stem forming a lobed, rootlike cal- careous holdfast anchored in soft substrate. Polyps non-retractile, cylindrical, armed with stout, faintly prickly needles and/or rods longitudinally placed in body wall; wall of pharynx with abundant small prickly rods. Type species.—Isis elongata Esper (by monotypy). Remarks.—Gray’s original (1858) and subsequent (1870) descriptions of /sidella do not provide sufficient detail for ade- quately defining the genus. Verrill (1883:13) considered the genus doubtful, but Studer [& Wright] (1887:44) accepted it on the ba- sis of Koch’s (1887) account of Jsis nea- politana Koch (=Isidella elongata [Esper]) and defined it essentially as diagnosed above. Koch (1887), Verrill (1883), Studer [& Wright] (1887), Nutting (1910), and Bayer 208 (1956) maintained that the large rods or spindles of the polyps do not project be- tween the bases of the tentacles as they do in Acanella, but Kiikenthal (1924) stated that the large septally placed spindles can “ein klein wenig zwischen den Tentakeln vorragen,” and Carpine & Grasshoff (1975: 108, 109, fig. 59) showed that the spindles may project conspicuously (even though this fact contradicted the generic characteristics stated on page 107). Three lots of J. elongata, among them one from the Zoological Station at Naples, the authenticity of which can hardly be doubt- ed, clearly show the distinctly projecting spindles described and illustrated by Car- pine & Grasshoff. A fourth, received from Prof. C. C. Nutting without locality (but possibly from the Zoological Station at Na- ples, as the polyps were skilfully relaxed be- fore fixation), shows strong, septally placed spindles that do not project between the ten- tacles—because the tentacles are fixed in more or less extended attitudes. It is difficult to reconcile these observations with Koch’s careful illustrations (1887:text-fig. 51; pl. 5, fig. 8), which show polyps of J. elongata without such strong needles, so the identity of “‘Isis elongata’’ seems open to question. Nevertheless, the salient generic features of the genus, i.e., dichotomous branching from the horny nodes, more or less in one plane, with prominent, non-retractile polyps armed with rods and needles, are sufficiently dis- tinctive that validity of the genus Jsidella is not in question. Distribution. — Heretofore, the genus Jsi- della has been reported only from the Med- iterranean and eastern Atlantic. Isidella trichotoma, new species Figs. la, 2-4 Material.—Southeast of Hawaii: 18°33.1’N, 155°26.1’W, 6300 feet (=1920 m), coll. J. G. Moore, serial 1723b, 17 Oct 1962. One colony much broken and lacking holdfast, alcohol, USNM 56715 (holotype). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Diagnosis.—Isidella with dichotomous and trichotomous branching, internodes long (to 8 cm), hollow, nearly straight. Pol- yps uniserially placed at wide intervals, cy- lindrical, short, armed with longitudinally placed needles the largest of which may project slightly between bases of tentacles; pharyngeal walls with small, flat rodlets. Description. —The colony (Fig. la) is sparingly branched from the nodes, trichot- omous and dichotomous, the internodes snow white, smooth, not longitudinally ribbed, hollow, nearly straight, long, the longest intact internode 8 cm long and 1.15 mm in diameter about mid-length; an in- ternode broken at one end is 8.5 cm long and only 0.8 mm in diameter. The stoutest internode is 2 mm in diameter. The nodes of the largest branches are about 2 mm long, brown, those of the thinner branches | mm or less, yellowish brown. The polyps (Fig. 2) are uniserially placed, mostly about 8 mm apart, occasionally as close as 5 mm and as far as 10 mm. They stand vertically or obliquely slanted toward the branch tips, cylindrical, mostly about 4-5 mm tall with tentacles folded over the mouth, but a few smaller individuals occur in the spaces between larger ones, indicating that new polyps do not arise exclusively at the branch tips; they are about 3 mm in diameter at mid-height, widening toward the base and sloping into the coenenchyme. Slender, nearly smooth pointed needles (Fig. 3a) about 1.5 mm long are longitudinally arranged in groups of 2 or 3 along the mes- enterial insertions, projecting little if at all between the tentacle bases; smaller needles are scattered among and between the mes- enterial groups, mostly lying deep in the thick, translucent mesogloea of the polyp body. Converging groups of still smaller needles lie in the bases of the tentacles, be- coming more or less longitudinal along the rachis; clusters of small, flat, blunt rods (Fig. 3d) lie longitudinally in the pinnules. The pharyngeal wall contains scattered small, flattened rodlets, with a median waist and <7 Lotae_.s 4 8 = S 3) > i=) oe) fo) os) oN \Oo nN on + Z 72) D> S 8o ‘S) a a 3 — ss) re S S NE He} 7.) o a=) oO =| ZS u fe) wo — ™~ \oO Vay Z N =) 8 £ is) ~ is) <= WS x ~ is} 3 aS) Ne os a * Ss es co \O a) Z 22) =) S 2 aS} x se} “a o =| Oo i=) is] I fe) oS Y a No) Ne} co \o a) Z N =) 3 a as) VOLUME 103, NUMBER 1 Fig. 1. 210 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Isidella trichotoma USNM 56715: polyps. tapered to blunt ends, mostly 0.11 mm in length, with a few straight-sided flat rods up to 0.15 mm in length (Fig. 3d). The extremely thin coenenchyme con- tains slender, delicate needles (Fig. 3b, c) up to 4 mm in length and 0.05 mm in diameter lying longitudinally along the axis. They are straight or slightly curved, not entirely uni- form in diameter, and ornamented with scattered, low granules. There is clear evi- dence that their length is achieved by fusion of adjacent sclerites (Figs. 3c, 4). The nature of the holdfast is unknown but, considering the shape of the colony and the depth of the habitat, it most likely was a lobate, rootlike structure similar to that of other species of [sidella. Etymology.—From Greek tricha = in three parts + tomos = a cutting, in allusion to the manner of branching. Remarks. —Even though the branching of this species is trichotomous in some cases and thus not uniplanar, it is not verticillate in the manner of Acanella. The very long, slender, nearly straight, hollow internodes and widely separated ramifications result in a spindly, openly branched colony of gross aspect closer to Isidella than to Acanella with its short, solid, more or less conspic- uously curved internodes. Acanella Gray, 1870 Acanella Gray, 1870:16.—Studer [& Wright], 1887:44.—Nutting, 1910:14.—Kuken- thal, 1919:573; 1924:414.—Deichmann, 1936:243.— Bayer, 1956:F222; 1981:941 (in key).—Bayer & Stefani, 1987a:51 Gn key); 1987b:941 (in key). Tsidella. —Muzik, 1978:737 (part). Diagnosis.—Keratoisidinae branched in whorls from nodes, at least in upper parts of colonies, forming bush-like colonies of moderate size (rarely more than 20 cm) when anchored in soft substrate by lobate hold- fast, larger and compressed (possibly to 1 m in height) when attached to hard bottom; internodes solid, short (to 2 cm). Polyps non- retractile, cylindrical, armed with sparsely prickly needles and/or rods longitudinally or obliquely placed in body wall; pharyngeal wall with small thorny stars or short rods. Remarks. —The consolidation of Isidella and Acanella as a single genus of Kerato- isidinae with branching originating at the nodes, as proposed by Muzik (1978) has practical advantages. However, the verti- cillate branching of Acanella is so distinc- tive that it seems preferable to retain it pending a thorough review of all species in- volved. e. d, Tentacles, pinnules, Coenenchym 3) Ma) oS 3 cal Mo) e} jaa) ow “a o ~~ q x oO n Tal — ~ \o va) Z 22) =) iS} = i) ~ is} <= aS nN n = S x 2 ix 1S) ”n = = & i=) a) 5 : fo] (3) n 8 2 \o ce oO oS os : Oo] a a, ue) q (] Fig. 3. 212 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Isidella trichotoma USNM 56715: Large coenenchymal needle produced by fusion of smaller sclerites. Acanella sibogae Nutting, 1910 Figs. 1b, 5 ?Acanella rigida Wright & Studer, 1889:31, pl. 9, fig. 4.—Thomson & Henderson, 1906:33. ?Acanella robusta Thomson & Henderson, 1906:33. Acanella sibogae Nutting, 1910:14, pl. 3, fig. 2, 2a, pl. 5, fig. 4.—Ktikenthal, 1919:575; 1924:419. ?Acanella japonica Kikenthal, 1915:120; 1919:582, pl. 44, fig. 76. Material.—Japan, off Mizimoko-shima Light: 32°36'N, 132°23’E, 437 fathoms (=799 m), bottom greenish brown mud, fine grey sand and foraminifers, USFC steamer Albatross sta. D-4957, 23 Aug 1906. One large colony with holdfast, alcohol, USNM 49792. Japan: off Shio Misaki Light: 33°25'20’N, 135°36'20’E, 244-290 fathoms (=446-530 m), bottom brown mud, sand and foramin- ifers, USFC steamer A/batross sta. D-4966, 29 Aug 1906. Two damaged colonies with holdfasts, alcohol, USNM 49619. Japan: off Shio Misaki Light: 33°23'40’N, 135°33’E, 587 fathoms (=1074 m), bottom brown mud, sand and stones, USFC steam- er Albatross sta. D-4969, 27 Aug 1906. Damaged branch, in alcohol, USNM 49468. Japan: off Shio Misaki Light: 33°23’30’N, 135°34’E, 649 fathoms (=1187 m), bottom brownish green mud and foraminifers, 214 USFC steamer Albatross sta. D-4971, 30 Aug 1906. Two colonies, one with holdfast, in alcohol, USNM 49466. Philippines: east coast of Luzon, off Batag I.: 12°44’42’N, 124°59'50’E, 383 fathoms (=700 m), green mud and sand, USFC steamer Albatross sta. D-5445, 3 Jun 1909. One colony with holdfast, in alcohol, USNM 50156. Philippines: off Balicasag I. between Siquijor and Bohol Is.: 9°22'30’N, 123°42'40’E, 392 fathoms (=717 m), bot- tom globigerina ooze, USFC steamer A/- batross sta. D-5527, 11 Aug 1909. Seven specimens more or less complete with hold- fast, 3 lacking holdfast, and several frag- mentary colonies, in alcohol, USNM 49917. Molucca Passage, off Mareh I.: 0°37’'00’N, 127°15'00’E, 417 fathoms (=763 m), bot- tom grey mud, USFC steamer A/batross sta. D-5618, 27 Nov 1909. Two colonies with holdfast, one nearly complete, alcohol, USNM 49896; 2 colonies, one with hold- fast, alcohol, USNM 50147. Buton Strait, off North I.: 5°35'00’S, 122°20'00’E, 559 fathoms (=1022 m), bot- tom green mud, USFC steamer Albatross sta. D-5648, 16 Dec 1909. Two colonies, alcohol, USNM 49969. Indonesia: Flores Sea: 5°36'30’S, 120°49'00"E, 692 fathoms (=1266 m), grey mud and sand, bottom temperature 39.2°F, USFC steamer Albatross sta. D-5660, 20 Dec 1909. One verticillate branch, in al- cohol, USNM 49959. Diagnosis. — Acanella with tall, vertically or obliquely placed polyps armed with sev- eral very large spindles longitudinally or di- agonally placed in the body wall, and pro- jecting conspicuously as 8 strong points around the folded tentacles. Holdfast a lobed, rootlike structure; secondary branch- ing commonly in whorls of 2. Remarks. —Several colonies taken by the USFC steamer A/batross agree in all essen- tial features with the original description of an incomplete colony obtained by the Si- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON boga Expedition north of Ceram (2°40’S, 128°37.5’E, 835 m) (Nutting 1910:14) and provide data to supplement the description of the species. One fully developed colony from Buton Strait southeast of Celebes (USNM 49969), lacking its basal holdfast, is about 15 cm tall (Fig. 1b); an immature specimen from the same station is 6 cm tall including the rootlike holdfast. Other specimens from Ja- pan (USNM 49466, 48468, 49619, 49792), the Philippines (USNM 49917, 50156), Flores Sea (USNM 49959), and Molucca Strait (USNM 50147) are in close agree- ment in colonial form, and size and arma- ture of polyps. The type specimen (Nutting 1910:pl. 3, fig. 2, 2a), evidently a branch from a larger colony, branched in pairs from the nodes but two pairs showed evidence of a third branch. Whereas the present specimens show a tendency to branch in pairs (..e., “whorls” of 2), whorls of 3 and 4 are com- mon and some nodes produce as many as 6 branches in a whorl. The internodes are solid, white, smooth but weakly ribbed lon- gitudinally, those of the principal branches 13-17 mm long, of the small side branches about 10 mm. The nodes are reddish brown, short, 1.5 mm to less than 1 mm in length. A few anastomoses between branches are present, as well as branchlets originating from internodes. Although Nutting (1910:15) described the polyps as mostly columnar and vertically placed on the branches, his photographs of the type branch and a twig with 3 polyps clearly show that they tend to slant distad toward the twig tips. The same condition prevails in the present specimens, on which vertically placed polyps certainly occur, but oblique ones predominate. The polyps (Fig. 5) generally conform in size with the dimensions given by Nutting, and show the “‘crown of eight well-marked points around the margin, each point con- sisting of the distal end of a single spicule”’ VOLUME 103, NUMBER 1 (Nutting 1910:15). Several large, more or less obliquely placed, curved spindles about 3 mm long surround the body of the polyps, projecting between the bases of the tentacles to produce the crown. Comparisons.—The polyps of Acanella verticillata Kiikenthal, 1915, from off Sib- erut Island have strongly projecting mar- ginal spines but are smaller (2 mm tall) than in the Siboga specimen and those reported here (4—5 mm). Moreover, according to Ku- kenthal’s description and illustration (1919: 584, fig. 258), a layer of obliquely placed, curved spindles covers the proximal part of the vertically placed projecting spindles. The large number of branches in the whorls de- scribed for verticillata is present, if not com- mon, also in some of the specimens of si- bogae now reported. Without a reexamination of type mate- rial, it is impossible to determine with cer- tainty whether or not Acanella rigida Wright & Studer from off Banda, A. robusta Thom- son & Henderson, 1906, from the Andaman Islands, and A. verticillata Kukenthal, 1915, from off Siberut Island west of Sumatra are identical with sibogae, but all characters mentioned in the original descriptions are consistent with that interpretation. Distribution.—Japan south and west to the Philippines and Indonesia, 446-1266 m. Acanella dispar, new species Figs. lc, d, 6-8 ?Lepidisis longiflora. —Nutting, 1908:572. Not Lepidisis longiflora Verrill, 1883:19. Material. —Hawatian Islands, off Maka- puu: 21°18’N, 157°32'W, 1200 feet (=366 m), Star IJ submersible, dive 2, 1 Feb 1978, K. Muzik, coll. One large colony now much broken, of which 3 branches are preserved in alcohol, the main trunk and large branch- es as well as the greater part of the secondary branches dry, USNM 56816, holotype. Hawaiian Islands, off Kaena Point: 21°35.85'N, 158°24.55'W, 275-445 m, 215 “Sango XII’ Haul 1, 27 Jul 1971, R. Grigg, coll. Damaged branches, identification un- certain, USNM 56719. Hawaiian Islands, off French Frigate Shoal: 23°53.1'N, 165°31.9'W, 326-363 m, “Sango XIV” Haul 2, 27 Aug 1971, R. Grigg, coll. Damaged branches, identification un- certain, USNM 56726. Hawaiian Islands, off NW coast of Oahu: Kahuku Point N. 79°, E. 10.1’, 216-251 fathoms (=395-—459 m), USFC steamer 4/- batross sta. D-4121, 25 Jul 1902. Frag- ments, identification uncertain; reported as Lepidisis longiflora by Nutting (1908), USNM 25358. Diagnosis. —Acanella with principal branching commonly in whorls of 2, rough- ly planar, secondary branching in whorls of 3 or more, bushy. Polyps with longitudi- nally placed needles of body wall developed asymmetrically, those of abaxial side stronger and more or less projecting be- tween bases of tentacles; pharyngeal walls with thorny stars. Description. — The intact colony was large, probably | m or more in height, in the shape of a profusely branched, strongly com- pressed bush. Branching arises from the nodes predominantly in one plane, singly and in opposite pairs, 1.e., whorls of 2; branching becomes verticillate at several places in the colony resulting in branches of typical Acanella aspect; here the nodes may give rise to 3 or more branchlets as well as 1 or 2, and in some places are abnormally overgrown by internodal material, possibly induced by the presence of a small actinian. The internodes of the principal branches are solid, cylindrical, slightly expanded at each end, weakly ribbed longitudinally, 15-— 20 mm long and up to 15 mm in diameter; all the subordinate branches bend upward so their proximal internodes are slightly curved. The distalmost internodes, straight or nearly so, are up to 20 mm in length but only 0.5—0.7 mm in diameter. The holdfast proper is not preserved, but the proximal PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON fe 6. Acanella dispar USNM 56816: Distal branches with twigs in whorls. Stereoscopic pairs. Scale bars = 1 cm. VOLUME 103, NUMBER 1 ANY Fig. 7. Acanella dispar, polyps: a, b, USNM 56816. c, 56719. d, 56726. 10 cm of the main trunk is thickened by irregular massive deposits of internodal ma- terial that obscure many of the nodes and form a strong basal support about 2 cm in diameter. The polyps are closely crowded on the smaller branches and twigs, concentrated on the two sides in the plane of branching but not in a Strict biserial arrangement; on the major branches they are more distantly placed, commonly 4 mm apart or even more. They are cylindrical, weakly curved and di- rected more or less distally, somewhat wider apically across the bases of the infolded ten- tacles, about 2.25 mm tall and 1 mm in diameter, 1.5 mm across the bases of the tentacles. Spiculation of the polyps varies with re- spect to size and number but the arrange- ment is generally consistent. Clusters of small needles lie along the mesenterial in- sertions, converging toward the margin of the tentacular crown (Fig. 8a); a few of the needles in one or more rows, sometimes only on the abaxial side, sometimes all around, are much larger than the rest (Fig. 8b). These project little, if at all, in some polyps, but in others one or more needles Eat ane TRS oe js ia Ale. an > eal od Z O i= c g 2a Zp) < 3 fy S > an = 1S) fe N Se) o } | ° ~Q ea bs cal fy 0 N 0 Z, A jaa ea OQ © ee AY VOLUME 103, NUMBER 1 may project conspicuously. Interseptal groups of small needles converge toward the base of each tentacle, extending upward along the rachis more or less longitudinally or obliquely; groups of much smaller rodlets extend into the pinnules (Fig. 8c, d). The walls of the pharynx contain numerous thorny [6-radiate?] stars (Fig. 8e, f), mostly about 0.06 mm long, but a few may reach 0.12 mm. The large needles of the polyps are about | mm long, the smaller ones about 0.5 mm. The internodes are snow white, the distal nodes brown, the proximal ones nearly black; in the alcoholic material the polyps are light brown to dark brown, depending upon conditions of preservation; upon drying the soft tissue becomes dark brown, almost black. Etymology. —Latin dispar = different. Comparisons.—This is the only species of Acanella known so far that inhabits hard bottom and develops a stout trunk. It is also unique in its planar primary branching that in the smaller branches becomes bushy, forming a flattened colony with side branch- es of typical Acanella aspect. Its polyps are smaller and less strongly armed than in any species heretofore recorded. This combi- nation of characters may eventually require generic reallocation of A. dispar. Remarks. —It is probable, but not certain, that the fragments from Hawaii reported by Nutting (1908) as the western Atlantic Lep- idisis longiflora Verrill are this species. Orstomisis, new genus Diagnosis. —Isididae dichotomously branched from the nodes, predominantly in one plane, forming compressed, multipla- nar flabellate colonies; internodes solid, short (up to 10 mm), often strongly curved, not longitudinally grooved; base of main —_— 219 stem developed as a massive calcareous trunk attached to solid substrate. Polyps forming prominent, cylindrical verrucae in- vested by thick, skinlike epithelium into which tentacles can be retracted completely; calycular rods not projecting beyond the bases of the infolded tentacles; walls of pharynx with small, sparsely tuberculate rodlets. Type species. —Orstomisis crosnieri, new species, here designated. Etymology.—From the acronym ‘“‘OR- STOM” for Office de la Recherche Scien- tifique et Technique Outre-Mer, now known as the Institut Francais de Recherche pour le Developpement en Cooperation, the agency responsible for collection of the ma- terial here described. Orstomisis crosnieri, new species Figs. 9-14 Material examined. —South of New Cal- edonia: 22°59.5’S, 167°22'E, 542 m; N/O Vauban, ORSTOM, Georges Bargibant coll., 7 Feb 1986. HGP-49. One damaged, in- complete colony, USNM 78372 (syntype). South of New Caledonia: 22°59'5S, 167°22'0E, 490-515 m, N/O Vauban sta. CP-216, 29 Sep 1985. Two colonies, MNHN Paris (syntypes). Southeast of New Caledonia: 23°05'792S, 167°46'544E, 600 m, N/O Jean Charcot, ORSTOM, sta. CP-52, 31 Aug 1985. Two colonies MNHN Paris (syntypes), one USNM 84774 (syntype). Loyalty Islands: 20°35'078S, 166°53' 990E, 460 m, N/O Jean Charcot sta. DW- 83, 6 Sep 1985. One colony MNHN Paris (syntype). Diagnosis. — As for the genus. Description. —The colonies (Figs. 9, 10a) are 20-45 cm in height, mostly not as wide as high but in two cases approximately so. Fig. 8. Acanella dispar USNM 56816, sclerites: a, Small needles of body wall. b, Large needles of body wall. c, d, Flat rodlets from pinnules. e, f, Thorny stars from pharynx. a-c, e, 0.3 mm scale; d, f, 60 um scale. 220 The main stem of all specimens forms a massive calcareous trunk (Fig. 10a), broken from the substrate so the holdfast proper is unknown. In the lower parts of the trunk, the thickening internodes grow over the nodes, ultimately obliterating all external evidence of their presence. Several primary branches arise from the main stem, subse- quently branching from the nodes mostly in one plane from which the terminal twigs irregularly diverge; the resulting colonies consist of several roughly parallel planes forming a compressed, complex, multipla- nar flabellum. Branching typically is di- chotomous, two branches arising from a node at 90°-180°; the widely diverging branches usually curve upward producing conspicuously lyre-shaped bifucations (Fig. 10b). The distal and intermediate internodes (Fig. 10c) are round in cross-section, not longitudinally grooved, the surface weakly granular and crowded with desmocyte pits clearly visible by SEM at low magnification; at higher power, vestiges of pits in various stages of infilling are abundant along with sharply defined pits where the desmocytes presumably were still functional at the time of collection (Fig. 10d). At the apex of the distalmost internodes the tubercles are more prominent and closely interspersed with desmocyte pits (Fig. 14). The thinnest in- ternodes are only 0.5 mm in diameter. The lowermost internodes before the onset of secondary thickening are nearly straight, about | cm in diameter and 13-18 mm long, gradually decreasing in thickness distad where the narrower internodes are about 1 mm in diameter and 7-10 mm long. On the larger branches, the nodes are very short, appearing as a dark brown line about 1 mm wide; those of the smaller branches may be up to 2.5 mm long. Although suc- cessive nodes may bifurcate, more com- monly branches are separated by 2—4 un- branched internodes, i.e., they arise from every 3rd to 5th node. In some cases, three PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON branches arise from a single node, one con- tinuing the direction of the preceding inter- node, the others diverging to each side at about 90°; where three successive nodes produce two lateral and one axial branch a ““pinnate” arrangement results. Rarely one node produces more than three branches, but this does not occur regularly to produce a bushy colony as in Acanella. The polyps are cylindrical, about 0.9-1.3 mm in diameter and 1.5—2.0 mm tall, de- pending upon the degree of contraction (Fig. 1 1b). In contraction the tentacles are folded over the mouth and are withdrawn into the verrucal aperture. The verrucal margin may be closed more or less completely over the retracted tentacles. When undamaged, the verrucae are covered by a smooth, thick ep- ithelium through which rodlike sclerites can be seen faintly or not at all. They have eight more or less distinct longitudinal furrows that distally divide the verrucal margin into low, rounded lobes. The smooth epithelium also covers the coenenchyme, in some places with faint longitudinal grooves following the course of the stem canals. Terminal branch- lets may have two polyps opposed at the apex and the older polyps may be arranged approximately in pairs, but they become more or less widely separated by unequal growth of intervening coenenchyme on the more proximal internodes, where they are scattered on all sides. Cylindrical rods (Figs. 12a, 13a) reaching about 1 mm in length and 0.1 mm in diameter are placed longi- tudinally or diagonally in the verrucal wall, with smaller rods interspersed among them. The rachis of the tentacles contains smaller rods, decreasing in size in the pinnules (Figs. 12b, 13b). Twinned rods in the shape of crosses are not uncommon (Figs. 12a, 13a). The pharyngeal wall contains small, sparse- ly knobby or thorny rodlets of basically 8-radiate form (Fig. 12c). The coenenchyme between the polyps is practically devoid of sclerites, although a few somewhat flattened rods occur widely VOLUME 103, NUMBER 1 Fig. 9. Orstomisis crosnieri USNM 84774: syntype colony. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 10. Orstomisis crosnieri USNM 78372: a, Colony severely decorticated. b, Detail of branches. c, Part of axial internode. d, Surface of axial internode. Scale at a = 5 cm; b = | cm. 223 VOLUME 103, NUMBER 1 Orstomisis crosnieri, terminal branchlets. a, USNM 78372, with verrucae flayed. b-e, USNM 84774: Fig. 11. with verrucae intact or partly flayed. Pharyngeal Twinned rodlet. c, b, a, Rods of verrucae. n oO = = oO = i>) a an ~ faa) (o2) ~ a =) = aS) S = a NZ ) = 7 O n Et © Q al 7S or Mie Oe g ~ Oe (Bs Se 38 Ge °o o oO se - =) 72) 3) o uU E oO ~~ + ir - ae | ue) io ° Ee xs Ay ra) “a o ~ q x oO n im Oo = = o > os Se ~~ ~~ + (oe) Z N =) & WS iS io) 9 RS S <2 a) S S a 2 e) el ~~ OS se s yee VOLUME 103, NUMBER 1 Fig. 13. distal internode. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON BA Fig. 14. Orstomisis crosnieri USNM 84774: Surface of distal internode immediately below tip. Stereoscopic pairs. VOLUME 103, NUMBER 1 scattered, especially in the vicinity of pol- yps. Brownish pigmented streaks, usually discontinuous, mark the course of the stem canals. Remarks. — Although thick, the smooth, skinlike epithelium seems to be very deli- cate, as it had been completely stripped off the first specimen received. This exposed the rod-shaped sclerites of the verrucae (Fig. 11a) and obscured the fact that the tentacles can be folded within a fleshy verrucal mar- gin (Fig. 11b), leading to provisional clas- sification of the specimen as a species of Isidella on the basis of its branching from the horny nodes, in spite of some morpho- logical discrepancies. Specimens in better condition received later immediately re- vealed the unique nature of the verrucae (Fig. 1 1b—e), necessitating the establishment of a new genus. This new genus cannot be accommodated in either of the subfamilies Isidinae and Muricellisidinae established by Ktikenthal on the basis of retractability of the polyps. It provides evidence that retractability alone is an insufficient basis for subfamilial dis- tinction. Among the Isididae, /sis, Cheli- donisis, and Muricellisis are the only genera having non-scalelike sclerites with tuber- cular ornamentation; that they are “‘retrac- tile’? can only be regarded as coincidental, as the polyps of each retract in different ways. The polyps of Jsis are virtually devoid of sclerites and retract into a thick coenen- chyme by folding the tentacles over the mouth and closing the rim of the coenen- chymal aperture. The polyps of Chelidonisis have blunt, conical or hemispherical ver- rucae filled with sclerites that extend onto the tentacles, which merely fold inward dur- ing contraction, and therefore are not “‘re- tractile’ in the same sense. The polyps of Muricellisis are divided into a proximal, projecting anthostelar part into which the distal, anthocodial part armed with sclerites arranged as a crown and points can retract more or less completely—at least theoreti- 227 cally. The only character shared by these three genera is the tubercular ornamenta- tion of the sclerites: capstans, clubs and spindles in /sis, sexradiates in Chelidonisis, and sharp spindles in Muricellisis. Orstomisis, undeniably “‘retractile,”’ does not share this spicular character. Its sclerites are definitely rods of the Keratoisis type. It does not clearly fit into either Isidinae or Muricellisidinae, but it does not justify still another subfamily. A solution to the subfamily problem is to define the Isidinae on the basis of tubercular ornamentation of sclerites, not retractability, and merge Mur- icellisidinae with it. Then Keratoisidinae can be defined on the basis of sparsely prickly or thorny rod-shaped sclerites, with Or- stomisis the only genus with retractile pol- yps. Etymology. —Dedicated to Dr. Alain Crosnier, in recognition of his tireless efforts to obtain financial support for a compre- hensive study of the rich collection of Oc- tocorallia obtained by expeditions of OR- STOM. Comparisons. — This species is unlike any isidid heretofore described. The retracta- bility of the tentacular part of the polyps into a firm, proximal calyx is unique in the Keratoisidinae and is paralleled only in Muricellisis in the Isidinae (Muricellisidi- nae of Kukenthal). The truly dichotomous manner of brancing predominant in this species occurs elsewhere only in Chelidoni- sis (Isidinae). Acknowledgments I here express thanks to Dr. B. Richer de Forges and Mr. Georges Bargibant of the Noumeéa office of ORSTOM for making available the gorgonians from New Cale- donia described in this paper. Dr. Manfred Grasshoff of the Natur-Museum Sencken- berg, Frankfurt, West Germany, and Mr. Phil Alderslade of the Northern Territory Museum of Arts and Sciences, Darwin, 228 Australia, kindly reviewed the manuscript and offered helpful comments. The scan- ning electron micrographs were made by Mr. W. R. Brown, chief of the SEM Laboratory, National Museum of Natural History, Smithsonian Institution. Ms. Molly Kelly Ryan provided lettering of the illustrations. Literature Cited Bayer, F. M. 1956. Octocorallia. Jn Treatise on In- vertebrate Paleontology F:166-231. Lawrence, University of Kansas Press. 1981. Key to the genera of Octocorallia ex- clusive of Pennatulacea (Coelenterata: Antho- Zoa), with diagnoses of new taxa. — Proceedings of the Biological Society of Washington 94(3): 902-947, figs. 1-80. —., M. Grasshoff, & J. Verseveldt, eds. 1983. II- lustrated trilingual glossary of morphological and anatomical terms applied to Octocorallia. E. J. Brill/Dr. W. Backhuys, Leiden, 75 pp., 20 pls. ——., & J. Stefani. 1987a. Isididae (Gorgonacea) de Nouvelle-Calédonie. Nouvelle clé des genres de la famille.—Bulletin du Muséum National d’ Histoire Naturelle Paris (4)9(A no. 1):47-106, figs. 1-4, pls. 1-30. ——, & . 1987b. Newand previously known taxa of isidid octocorals (Coelenterata: Gorgo- nacea), partly from Antarctic waters. — Proceed- ings of the Biological Society of Washington 100(4):937-991, figs. 1-31. Carpine, C., & M. Grasshoff. 1975. Les gorgonaires de la Méditerranée occidentale.—Bulletin de l'Institut Océanographique Monaco 71(1430): 1-140, figs. 1-62. Deichmann, E. 1936. The Alcyonaria of the western part of the Atlantic Ocean.—Memoirs of the Museum of Comparative Zodlogy at Harvard College 53:1-317, pls. 1-37. Gray, J.E. [1858]. Synopsis of the families and genera of axiferous zoophytes or barked corals. —Pro- ceedings of the Zoological Society of London 1857:278-294. 1870. Catalogue of the lithophytes or stony corals in the collection of the British Museum. London, British Museum, [iv] + 40 pp. Koch, G. von. 1887. Die Gorgoniden des Golfes von Neapel und der angrenzenden Meeresabschnit- te.— Fauna und Flora des Golfes von Neapel 15: i-x + 1-99, pls. 1-10. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Kikenthal, W. 1915. System und Stammesgeschichte der Isididae.—Zoologische Anzeiger 46:116- 126. 1919. Gorgonaria.—Wissenschaftliche Er- gebnisse der deutschen Tiefee-Expedition auf dem Dampfer “Valdivia” 1898-1899 13:1-946, pls. 30-89. . 1924. Gorgonaria.— Das Tierreich 47:i—xxvili + 1-478, figs. 1-209. Muzik, K. 1978. A bioluminescent gorgonian, Lep- idisis olapa, new species (Coelenterata: Octo- corallia), from Hawaii.— Bulletin of Marine Sci- ence 28(4):735-741, figs. 1-4. Nutting. C. C. 1908. Descriptions of the nice collected by the U.S. Bureau of Fisheries steam- er Albatross in the vicinity of the Hawaiian Is- lands in 1902.— Proceedings of the United States National Museum 34:543-601, pls. 41-51. 1910. The Gorgonacea of the Siboga Expe- dition V. The Isidae.—Siboga-Expeditie Mon- ographie 13b2:1-24, pls. 1-6. Studer, T. [& E. P. Wright.]. 1887. Versuch eines Systemes der Alcyonaria.— Archiv fur Naturge- schichte 53(1):1—74, pl. 1. Thomson, J. A., & W. D. Henderson. 1906. An ac- count of the alcyonarians collected by the Royal Indian Marine Survey Ship Investigator in the Indian Ocean. Part 1. The alcyonarians of the deep sea. i-xvi + 1-132 pp., pls. 1-10. Calcutta: Indian Museum. Verrill, A. E. 1883. Report on the Anthozoa, and on some additional species dredged by the ““Blake”’ in 1877-1879, and by the U.S. Fish Commis- sion steamer “‘Fish Hawk” in 1880-82.—Bul- letin of the Museum of Comparative Zoology at Harvard College 11:1-72, pls. 1-8. Verseveldt, J., & F. M. Bayer. 1988. Revision of the genera Bellonella, Eleutherobia, Nidalia and Ni- daliopsis (Octocorallia: Alcyoniidae and Nida- liidae), with descriptions of two new genera.— Zoologische Verhandelingen 245:i-vi + 1-131 pp., figs. 1-64. Wright, E. P., & Th. Studer. 1889. Report on the Alcyonaria collected by H.M.S. Challenger dur- ing the years 1873—1876.—Report on the Sci- entific Results of the Voyage of H.M.S. Chal- lenger during the Years 1973-76, Zoology 31 (part 64):i-Ixxvi + 1-314, pls. 1-43. Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 229-234 A NEW SPECIES OF CLADOCARPUS (CNIDARIA: HYDROIDA: PLUMULARIIDAE) FROM THE STRAITS OF FLORIDA Mary Anna Bogle Abstract. —Cladocarpus delicatus, a new species of plumulariid hydroid, is described, bringing to nine the number of species of Cladocarpus known from the Straits of Florida. The delicate appearance of the colony and spike-like projection from the medial abcauline wall of the hydrotheca distinguishes C. delicatus from other species of this genus. A key to the species of Cladocarpus known from the tropical western Atlantic is provided. In May 1962, the staff of the Institute of Marine Sciences [now the Rosenstiel School of Marine and Atmospheric Sciences (RSMAS)] of the University of Miami be- gan a faunal survey of the Straits of Florida and its approaches. During the course of the project, conducted intermittently over a 10- year period, a large number of thecate hy- droids was collected, including many mem- bers of the family Plumulariidae. In this collection a new plumulariid species, Clado- carpus delicatus, belonging to the subfamily Aglaopheniinae was discovered. The holo- type is described and illustrated in this pa- per. An unpublished account of the species appears elsewhere (Bogle 1975). The holotype of Cladocarpus delicatus is deposited in the collections of the National Museum of Natural History, Smithsonian Institution (USNM). The paratype is locat- ed in the invertebrate museum of the Ro- senstiel School of Marine and Atmospheric Sciences. Subfamily Aglaopheniinae Stechow, 1911 Genus Cladocarpus Allman, 1874 Cladocarpus delicatus, new species (Figs. 1-3) Material examined. — Material in the USNM: Holotype, USNM 60212, 1 colony, R/V Gerda Sta. G-237, Straits of Florida (25°15'’N, 79°14’W-25°17'N, 79°14'W), 393-397 m, 30 Jan 1964. Colony with phy- lactogonia and gonothecae present (Table 1); 18 mm high. Material at RSMAS: Paratype, UMML 5: 151, one colony, R/V Gerda Sta. G-239, Straits of Florida (25°20'N, 79°15'’W- 25°22'N, 79°16'W), 256-348 m, 30 Jan 1964. Fragments of colony with phylacto- gonia and gonothecae present. Description. —Trophosome: Holotype colony with unbranched hydrocaulus, at- taining a height of 18 mm; anchored in the substrate by a tangle of rhizoidal filaments. Hydrocaulus slender and nonfascicled, car- rying hydrocladia along the distal half. Hy- drocauline nodes indistinct. Three very strong, oblique, hydrocauline constrictions just proximal to the first hydrocladium with one nematotheca present between succes- sive constrictions. Proximal half of hydro- caulus with a single longitudinal row of nematothecae along the anterior face. An- terior face of distal half of hydrocaulus with two tubular nematothecae between succes- sive hydrocladia, one in the axil of the apophysis and the other proximal to it. Unmodified hydrocladia arranged alter- nately, few in number and distant (~0.75 mm apart); divided into long (~ 1 mm), sin- uous, often weakly delineated internodes. Distal fourth of each internode arched 230 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Measurements of nematothecae and hydrothecae of the holotype of Cladocarpus delicatus. nn eee UTES EEE ESSE San an Measurements (in micrometers) Range (mean) Number* i Distance from apex of supracalycine nematotheca to apex of preceding supracalycine nematotheca Hydrotheca, total depth Maximum diameter Diameter at margin Mesial nematotheca, total length Supracalycine nematotheca, total length Length above hydrothecal margin Gonotheca, length Width 966-1092 (1016) 10 522-605 (574) 11 231-314 (298) 11 138-176 (162) 11 132-165 (151) 11 143-176 (164) 11 33-60 (44) 11 396-418 (411) 2) 170-226 (209) 2 * Number of measurements made. slightly over the hydrotheca of that inter- node. Hydrothecae distant, located approxi- mately midway along each hydrocladial in- ternode. Each hydrotheca narrow at the proximal end, greatly inflated toward the middle and narrowing again slightly near the hydrothecal margin. Medial abcauline wall noticeably thickened and often pro- truding out and upward so as to form a spike-like projection originating approxi- mately two-thirds up the hydrothecal face. Length of spike as much as 0.1 mm. Pro- jection usually with a small cavity formed by the continuation of the hydrothecal cav- ity into it but without a terminal aperture. Hydrothecal margin with a single large me- dial tooth and 4 to 5 smaller lateral teeth on each side. Strong, posterior intrathecal ridge extending obliquely upward across one-third to one-half of both lateral faces of the hydrotheca. Number of internodal septa opposite hydrotheca usually 2, occa- sionally 3. Septa very short and weak; lo- cated proximal to the posterior intrathecal ridge. No other septa regularly present per internode except one at the base of the me- sial nematotheca. A single long, tubular mesial nematotheca springing from the hydrocladium as much as 100 um below the base of the hydrotheca. Mesial nematotheca with an aperture run- ning from the top to midway along the free upper surface. A pair of long, tubular supracalycine nematothecae projecting obliquely outward and upward from their hydrocladial origin; slightly geniculate near their distal end. Gonosome: Gonothecae narrow, length- ened, obovate sacs arising in small clusters near the base of the apophyses of unmodi- fied and modified hydrocladia that bear phylactogonia. Gonothecae with oval ter- minal aperture. Phylactogonia arising from the most dis- tal hydrocladia, each of which is usually modified into a short, nematophorous spike bearing no hydrotheca. A single phylac- togonium arises from an area lateral to the proximal nematothecae of each of these hy- drocladia and arches medially over the front of the hydrocaulus, thus protecting the go- nangia borne there. Phylactogonium usually composed of 3 to 4 very slender nematoph- orous branchlets formed by the bifurcation of alternate branchlets, with the entire struc- ture resembling stag antlers. Phylactogonial nematothecae long and tubular, directed outward and upward away from the gonan- gia. Etymology. —Delicatus (Latin) = deli- cate, alluding to the delicate appearance of the colony. Geographic range. —Cladocarpus delica- VOLUME 103, NUMBER 1 231 Fig. 1. tus is known only from the northern Straits of Florida. Discussion. —This small delicate hydroid closely resembles in size and in the general form of the trophosome and gonosome two other species of Cladocarpus reported from the Straits of Florida. They are C. doli- chotheca Allman, 1977, and C. tenuis Clarke, Cladocarpus delicatus, new species: Holotype colony, USNM 60212, 18 mm high. 1879. The distinctive hydrotheca of C. deli- catus with its inflated appearance and its unusual spine-like projection from the ab- cauline wall, however, readily distinguishes it from either C. dolichotheca or C. tenuis as well as from other members of the genus Cladocarpus. This new hydroid brings to nine the number of species of Cladocarpus 232 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Cladocarpus delicatus, new species: A, Hy- drotheca of holotype, lateral view, USNM 60212 (northern Straits of Florida), scale 0.5 mm; B, Hydro- theca of holotype, lateral view showing spike-like pro- jection from medial abcauline wall, USNM 60212 (northern Straits of Florida), scale 0.5 mm. reported from the Straits of Florida and its approaches and to thirteen the number known from the tropical western Atlantic. Currently, approximately 60 species are as- signed to the genus Cladocarpus (see Mil- lard 1975, Rees & Vervoort 1987, Vervoort 1966) though several are questionably in- cluded in the genus since no gonosome has been described. The following key is to as- sist in the identification of species of Clado- carpus known from the tropical western At- lantic. Those species of Cladocarpus reported from the Straits of Florida and its approaches and treated in detail by the au- thor (Bogle 1975) are indicated in the key with an ‘*’, A Key to the Tropical Western Atlantic Species of Cladocarpus la. Hydrothecal margin entire C. carinatus Nutting, 1900* 1b. 2a. 2b. 3a. 3b. 4a. Ab. Sa. Sb. 6a. 6b. Ta. Toy 8a. 8b. 9a. One or more teeth on hydrothe- CalVinian pin Ret ee) eens cia 2 Hydrothecal margin with | or 2 prominent median teeth; others, hE OREN SONG, OGY soos cuvaemades 3 Hydrothecal margin with several subequal teeth; none prominent Gah rs ee ediper er u eae 2s eae 11 Hydrothecal margin with 1 prominent median tooth ...... 4 Hydrothecal margin with 2 prominent median teeth ...... 12 Posterior (adcauline) intrathecal Tadgel absentee eee ee 5 Posterior (adcauline) intrathecal Pde presente one eee 7 Hydrotheca obconic (like an in- verted cone); face of hydrotheca straight .. C. flexuosus Nutting, 1900 Hydrotheca long and slender, not cone-like; anterior profile of hy- drothecaiconcavier. ee eee 6 Anterior profile of hydrotheca weakly concave; hydrotheca not appreciably narrowed (ratio of lateral width of hydrotheca at its midpoint to the length approxi- mately 1:4) seen C. dolichotheca Allman, 1877* Anterior profile of hydrotheca strongly concave; lateral and frontal aspect of hydrotheca ap- preciably narrowed (ratio of lat- eral width of hydrotheca at its midpoint to the length approxi- mately 1:6) C. tenuis Clarke, 1879* Hydrocaulus fascicled ......... Balser C. longipinna Fraser, 1945 Hydrocaulus nonfascicled Intrathecal ridge present but short and weak; anterior (abcauline) profile of hydrotheca slightly COMVOXE C. flexilis Verrill, 1885 Intrathecal ridge short but strong; anterior (abcauline) profile of hy- drotheca convex or sinuous ... 9 Hydrocladial internode with no VOLUME 103, NUMBER 1 Fig. 3. Florida), height 1.3 mm. 9b. 10a. 10b. more than 3 septa; mesial ne- matotheca not adnate to the hy- drotheca but arising as much as 100 um below the base of the hy- drotheca C. delicatus, new species* Hydrocladial internode with 5 or more septa; mesial nematotheca adnate to or arising just below the base of the hydrotheca Anterior (abcauline) profile of hydrotheca sinuous RA Ae C. ventricosus Allman, 1877* Anterior (abcauline) profile of lla. iL 115), Was 233 Cladocarpus delicatus, new species: Phylactogonium of paratype, UMML 5:151 (northern Straits of hydrotheca convex See Pas C. obliquus Nutting, 1900* Hydrotheca with a conspicuous, sigmoid, posterior intrathecal ridge extending to anterior (ab- cauline) face hee tai C. sigma (Allman, 1877)* Hydrotheca with a very short and straight posterior intrathecal ridge a Oe C. compressus Fewkes, 1881 Hydrotheca deep with 3 or more internodal septa opposite it; su- pracalycine nematothecae with 234 strong posterior constriction; ne- matothecal margin not noticea- bly Serrate = Aa Lee ee AE cakes C. paradiseus Allman, 1877* 12b. Hydrotheca relatively short and squat with usually only | or 2 internodal septa opposite it; su- pracalycine nematothecae lack- ing a strong posterior constric- tion and with a finely serrated margin .. C. grandis Nutting, 1900* Acknowledgments This paper is a result of research under- taken while at the Rosenstiel School of Ma- rine and Atmospheric Sciences, University of Miami, and is part of a series resulting from the National Geographic Society—Uni- versity of Miami Deep-Sea Biology Pro- gram. I wish to thank Dr. Dennis M. Opresko and Dr. Richard B. McLean, both of Oak Ridge National Laboratory, for their sug- gestions and comments on the manuscript. I would also like to express my gratitude to Dr. Stephen D. Cairns of the National Mu- seum of Natural History for his assistance in depositing the holotype in the collections of the USNM. Oak Ridge National Laboratory is oper- ated by Martin Marietta Energy Systems, Inc., under contract DE-AC05-840R21400 with the U.S. Department of Energy. Pub- lication No. 3238, Environmental Sciences Division, Oak Ridge National Laboratory. Literature Cited Allman, G.J. 1874. Report on the Hydroida collected during the expedition of H.M.S. ‘Porcupine.’— Transactions of the Zoological Society of Lon- don 8:469-48 1. . 1877. Report on the Hydroida collected dur- ing the exploration of the Gulf Stream by L. F. de Pourtalés.—Memoirs of the Museum of Comparative Zoology at Harvard College 5(2): 1-66. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bogle, M.A. 1975. Areview and preliminary revision of the Aglaopheniinae (Hydroida: Plumulari- idae) of the tropical western Atlantic.— Unpub- lished M.S. Thesis, University of Miami, 307 pp. Clarke, S. F. 1879. Report on the Hydroida collected during the exploration of the Gulf Stream and the Gulf of Mexico by Alexander Agassiz, 1877— 1878.—Bulletin of the Museum of Comparative Zoology at Harvard College 5:239-252. Fewkes, J. W. 1881. Report on the Acalephae, in Reports on the results of dredging, under su- pervision of Alexander Agassiz, in the Carib- bean Sea, in 1878, 1879, and along the Atlantic coast of the United States during the summer of 1880, by the U. S. Coast Survey Steamer Blake.— Bulletin of the Museum of Compara- tive Zoology at Harvard College 8:127-140. Fraser, C. M. 1945. Notes on some recently collected hyroids in the United States National Museum, with descriptions of three new species. —Journal of the Washington Academy of Sciences 35:21- 23% Millard, N.A.H. 1975. Monograph on the Hydroida of southern Africa.— Annals of the South Afri- can Museum 68:1-513. Nutting, C. 1900. American hydroids. Part I. The Plumularidae.—Special Bulletin United States National Museum 4(1):1-285. Rees, W. J., & W. Vervoort. 1987. Hydroids from the John Murray Expedition to the Indian Ocean, with revisory notes on Hydrodendron, Abieti- nella, Cryptolaria and Zygophylax (Cnidaria: Hydrozoa).—Zoologische Verhandelingen 237: 1-209. Stechow, E. 1911. Uber Hydroiden der Deutschen Tiefsee-Expedition. Ein neues Genus thecater Hydroiden.— Zoologischer Anzeiger 37:193- 197. Verrill, A. E. 1885. Results of the explorations made by the Steamer Albatross off the northern coast of the United States in 1883.—Report of the United States Fish Commission 1 1:503-699. Vervoort, W. 1966. Bathyal and abyssal hydroids.— Galathea Report 8:97-174. Environmental Sciences Division, Oak Ridge National Laboratory, Building 1505 MS 6036, P.O. Box 2008, Oak Ridge, Ten- nessee 37831-6036. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 235-247 THE FOURTH ANNUAL RISER LECTURE: THE ROLE OF PHYSIOLOGY AND BIOCHEMISTRY IN UNDERSTANDING ANIMAL PHYLOGENY Charlotte P. Mangum Abstract. —On several occasions in the history of physiology and biochem- istry, claims have been made of uniquely superior approaches to understanding animal phylogeny. Perhaps the most infamous episode was the putative di- chotomy between the muscle phosphagens found in protostomes and deutero- stomes, which later proved to be factually incorrect. I argue that physiology and biochemistry can be powerful tools in understanding animal evolution, and I give an example in which the common quaternary structure of the oxygen carrier hemocyanin in the three groups of living arthropods is strong evidence for acommon origin and the integrity of the Phylum Arthropoda. The evidence is compelling, not because the character is molecular but because molecular structure can be interpreted in the context of its physiological setting in living, breathing animals. Thus the argument can be constructed in Darwinian terms and the evidence would be extremely difficult to interpret alternatively. G. Evelyn Hutchinson has defined the phylum as a group of organisms which is not very clearly related to any other group. This definition may seem to be only hu- morous but in fact it is fairly accurate, both historically and conceptually. Throughout the modern history of zoology taxonomists have devised subsuming sets until they can no longer do so, and have then stopped and called the most general set a phylum. Thus it is the height of absurdity when we turn around and ask ourselves why we do not understand the relationships of the animal phyla. Ignorance has been cleverly built into the process (to keep ourselves in business?). Nevertheless I, like many others, find the In 1985 the annual Riser Lecture was initiated by members, alumni and friends of the Marine Science Center, Northeastern University of Nahant, Massa- chusetts. The occasion was the official retirement of Professor Nathan W. Riser. As teacher, biologist and founder of the facility, ‘Pete’ Riser endowed the lab- oratory with a legacy—the importance of considering the whole organism regardless of one’s special focus. We dedicate these annual lectures to that principle. temptation irresistable. I believe that the quest makes us think about how animals work in ways that we would not if we fail to undertake it. And understanding how an- imals work is what physiological and bio- chemical zoology is all about. Do physiology and biochemistry have anything to offer to understanding animal phylogeny that other subdisciplines do not? In one sense the answer must be affirmative because the logic is essentially tautologous. If physiology and biochemistry had nothing unique about them they would not be rec- ognized as coherent disciplines. The more meaningful question is do physiology and biochemistry have anything especially co- gent to offer because of their unique subject matter? My answer is yes, but my reasons may surprise the reader. Historical Episodes of Physiological and Biochemical Inquiry into Animal Phylogeny Muscle phosphagens. — First I must illus- trate the reasons why physiology and bio- 236 chemistry have not lived up to their poten- tial in contributing to phylogenetic knowledge. The most celebrated example of an attempt to use biochemical data in the study of animal evolution involves the dis- tribution of the phosphagens, the guanidine derivatives found in muscle and a variety of other tissues that store high energy PO,s. In the 1920s and 30s there was something of a revolution in thinking about animal phylogeny. The early metabolic biochem- ists were rapidly unravelling pathways of intermediary metabolism, concentrating on somatic muscle. A “dichotomy” was per- ceived to distinguish “vertebrates”! and ‘“‘invertebrates’’!: while the vertebrates con- tain creatine PO,, it was said, the inverte- brates contain the structurally distant com- pound arginine PO, (Needham et al. 1932, Kutscher & Ackermann 1933, Baldwin & Needham 1937). This theory also supposed that phosphoarginine was the original, now relict form and that phosphocreatine is a relatively new molecule that arose late in animal evolution. Even more excitement was generated by the first exceptions be- cause they were found in the deuterostomes, some of which contain both phosphagens (e.g., Baldwin & Yudkin 1950). The co- existence was regarded as an example of transition from the relict to the modern form. A number of reviews and monographs were written using these relationships to in- troduce the idea that biochemistry under- taken in a comparative framework can make important contributions to phylogenetic knowledge. When Baldwin published the 1949 edition of his essay entitled Compar- ative Biochemistry, the evidence cited in support of a phylogenetic distribution of the muscle phosphagens represented six of the ' I remain amazed by our persistence in employing, not only in our biological curricula but also in our original researches, this particular pseudo-dichotomy, the fallacy of which has been recognized for a century and a half! PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON several dozen animal phyla. Ten years later in the final edition of his textbook, the da- tabase included about 50 species represent- ing 9 phyla, which was considered a large sample (Baldwin 1959). By the middle 1950s van Thoai & Robin (1969) had begun to report a number of disturbing exceptions of phosphocreatine in the wrong (viz. protostomate) animals. Nonetheless the notion that these com- pounds are distributed along phylogenetic lines persisted for another decade. Of some 118 species of annelids and related groups examined by van Thoai & Robin (1969), for example, more than twice as many con- tain the wrong phosphagen as the right one (albeit an equally large number contains compounds closely related to phosphoar- ginine, which were unknown to the early workers). Phosphocreatine has been found in the Porifera as well. Thomas Kuhn (1970) wrote a book on the heuristic role of what he called para- digms in advancing scientific knowledge. Paradigms are “‘universally recognized sci- entific achievements that for a time provide model problems and solutions to a com- munity of practitioners”’ (p. viii). And if the communication facilitated by such wide- spread concurrence is fruitful, the truth of the paradigm matters very little. In support of his thesis Kuhn (1970) noted that he would use examples from physics and chemistry because he knew them best. But he also maintained that he could have used examples from biology had he so chosen. I have always thought that Kuhn was wrong on that point, primarily because in biology we almost never concur so widely that we communicate without a lot of ar- gument. But the phosphagen theory was about as close to a paradigm as biology ever gets.” ? The only other examples that come to my mind of paradigms that proved to be incorrect are the bioge- netic law and perhaps, insofar as the principles are regarded as generally true, Drosophila genetics. VOLUME 103, NUMBER 1 As early as 1946 Greenwald had found the wrong phosphagen in annelids and echiuroids. Instead of concluding that something might be wrong with the theory, however, he concluded that the annelids and echiuroids are more closely related to the chordates than previously supposed! When later investigators failed to confirm his orig- inal report of both phosphagens in the hemi- chordate Balanoglossus, Baldwin (1953) ac- knowledged that the original finding might be wrong (the field had always been plagued by inadequate analytical methods). But, he also suggested, in the intervening 20 years perhaps a mutation has occurred and the relict compound has now been replaced by the modern form. Finally, in their 1958 re- view Ennor & Morrison wrote: “This suggestion (of a phylogenetic dis- tribution) . . . had an unfortunate and in- hibitory effect on biochemical thought in this field for many years, and was made in spite of the fact that there was evidence to show that some invertebrates possess creatine” (p. 664, parenthesis mine). They then hedged by concluding that the theory might be valid within the deutero- stomes! It is difficult for those of us who were not around at the time to appreciate the impact made by the phosphagen theory. Consider, for example, the status of inquiry into an- imal phylogeny in the 1920s. The deuter- ostome grouping was by no means univer- sally accepted, and the notion that acorn worms might be more closely related to ver- tebrates than to other Vermes was also dis- tasteful to some. The phosphagen theory was welcomed by proponents of modern phy- logeny as especially cogent evidence be- cause it was biochemical. Evelyn Hutchin- son has reminded me that the introduction of biochemical data irreversibly expanded zoological thinking about phylogeny and so, in that sense, it is a good example of Kuhn’s (1970) heuristic though erroneous para- digm. But, I suggest, physiologists and biochemists would be remiss if we failed to 237 PENGUIN KANGAROO CHICKEN RATTLESNAKE DUCK | TURTLE FLY Rag OT PIG HORSE DONKEY DOG BIRD MAN MONKEY MAMMA BULLFROG RERTILES at AMPHIBIANS INSECTS VERTEBRATE Fig. 1. Phylogenetic tree of the animal kingdom based on the primary structural similarities of cyto- chrome c. Redrawn from Dickerson & Geis (1969). conclude that there is a better way of think- ing about phylogeny. The theory has become a whipping boy for classical zoologists who do not want to be told that morphological data may not always take precedence. And biochemists have written a number of fairly messianic papers on how to go about the task in the right way. In my opinion, few have. Alternative biochemical approaches. —In the 1960s the comparison of primary struc- ture of proteins was advocated as a bio- chemical approach to phylogeny (e.g., Zuck- erkandl & Pauling 1965). Indeed there have now been a number of such comparisons, perhaps the best known of which culmi- nated in the phylogenetic tree based on the amino acid sequences of cytochrome c (Fig. 1). This approach, though certainly less sim- plistic than the phosphagen theory still has two major difficulties. One is technical and hopefully transient and the other episte- mological and therefore more persistent. Until recently, sequencing macromole- cules has remained a formidable task. Cy- tochrome c was a good choice because it is 238 B Mexico @ Jamaica Venezuela Tanzania PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON co a fn L g Australia Fig. 2. A map of the world based on the frequency with which a nation’s scientific journals are cited. Redrawn from Kidron & Segal (1984). ubiquitous and it is at least not the biggest or most complex of proteins. But note the scope of the phylogenetic tree (Fig. 1). In no way does it truly depict the relationships of the animal phyla. If one were to map the nations of the world based not on their land masses but on the frequency with which their scientific journals are cited (Fig. 2), one would distort the geographic truth by a fac- tor averaging about eight. In contrast, bas- ing animal phylogeny on nine species of mammals, three birds, two reptiles, one am- phibian, one teleost, and two insects distorts zoological truth by a factor of eighteen! Scarcity of information is no reason to reject an entire approach, but it does underscore the need to explore alternatives, unless one can suppress curiosity more than I can. The second problem is that the nature of a putatively causal relationship between protein structure and phylogenetic related- ness is by no means clear. I will argue that the comparison of primary structures, made outside of a physiological context, tells us more about the evolution of the molecule than the evolution of the animal phyla. Once again the annelids, that perhaps most bio- chemically diverse of multicellular animals, can be used as an example. The primary structure of the monomeric hemoglobin in the red cells of the bloodworm Glycera di- branchiata has more in common with that of the hemoglobins of the distantly related vertebrates than it does with the monomeric hemoglobin in the red cells of the terebellid Enoplobranchus sanguineus, a member of the same class (Imamura et al. 1972, Weber et al. 1977)! Still different molecular approaches have also been introduced. The technique of DNA pairing remains in limbo and at this writing the prospects do not appear to be good. Nei- ther it nor immunological techniques prom- ise to be very useful in understanding the relationships of higher taxa, for the simple reason that they require a minimum amount of relatedness that is essentially precluded by our definition of the phylum. The very recent introduction of compar- isons of sequences of nucleic acids rather than amino acids should prove to be useful. Only time and extensive validation of methodology will tell. I cannot resist the temptation to report a sense of the déja vu, however, when I read sentences such as “‘the use of methods discussed here make it pos- VOLUME 103, NUMBER 1 239 Table 1.—Phylogenetic relationships based on fractional structural similarity of 5S rRNA. Data from Lane et al. (1985). Mean + SD. Sabellastarte japonica Sabellastarte japonica Perinereis brevicirrus 0.90 + 0.03 Solemya velum 0.90 + 0.03 Lingula anatina 0.86 + 0.03 sible to define the phylogenetic afhliations of any organism” (Pace et al. 1986). In fact the use of those very methods led to patently absurd conclusions, apparently unnoticed by Lane et al. (1985). The mean values for fractional ““homology”’ (read: structural similarity) of 5S RNA indicate that the mol- lusc Solemya velum is more closely related to the annelids Perinereis brevicirrus and Sabellastarte japonica than the two annelids are to one another (Table 1)! To be more accurate by taking into account the error around the mean values, Table | suggests that S. velum, P. brevicirrus, S. japonica and Lingula anatina are all more or less equally related to one another. (At this point I will confess parentheti- cally that I eagerly await further explication and application of the methods used in re- lating sequences of the 16-18S ribosomal RNA fractions, a better choice, by Field et al. (1988). The exact way in which the ma- trix analysis of the data was performed was unclear in the original, presumably due to space constraints specified by the journal, and consultation of the literature cited clar- ified only the general approach, not details that would help me understand discrepan- cies between the relationships depicted in the various figures. I hope that my enthu- siasm for this work does not arise entirely from the agreement of the conclusions reached by Field et al. (1988) with my own phylogenetic predilections, but I am not sure.) Why have we gone wrong?—I describe these historical incidents to illustrate what I believe are at best limited and at worst Perinereis brevicirrus Solemya velum Lingula anatina 0.87 + 0.03 0.92 + 0.03 0.88 + 0.04 0.93 + 0.03 0192730203 0.92 + 0.02 0.96 + 0.02 0.91 + 0.03 0.96 + 0.02 poor approaches to animal phylogeny. The databases remain small. But the compari- son of muscle phosphagens and protein structures were also unsound because they lacked a Darwinian component. The prac- titioners never asked the question why a particular phosphagen or heme protein was selected in a particular group. The early metabolic biochemists noted that they real- ly did not know why phosphocreatine ap- peared to have been selected over phos- phoarginine in the deuterostomes, but there was so little discussion that they must not have regarded the point as serious. To this day the relative advantages of the various phosphagens remain a subject of specula- tion (Hird 1986, W. R. Ellington pers. comm.). Centipedal Hemocyanin and Its Phylogenetic Implications Finally, I arrive at my own subject, an example of biochemical data that are suf- ficiently large in number and clear in rela- tion to physiological function that they can make a contribution in deciding a contro- versial question about animal phyla. The biochemical and physiological data pertain to the molecular structure and respiratory function of the hemocyanins (Hes), the Cu- containing O, carriers; the phylogenetic question is the status of the Phylum Ar- thropoda. Distribution and quaternary structures of the hemocyanins. —In contrast to the hemo- globins the Hcs have always been regarded as coherent in taxonomic distribution, being 240 found only in arthropods and molluscs. Moreover, their higher order structure dif- fers so fundamentally that there is little doubt that arthropod and molluscan Hcs had a separate origin. Arthropod hemocyanins: Arthropod Hcs are built as multiples of anywhere from 6 to 48 monomeric subunits of about 67-80 x 103 d, each with a single active site (Van Holde & Miller 1982). The monomers are put together loosely. The first aggregate in assembly of the native polymer is a hex- amer, which exists alone in the blood of some species. In the laboratory a hexamer can be made of only one kind of polypeptide chain, although in nature there are always two or more. If the 450 x 10? d hexamers contain an immunologically distinctive kind of chain that tends to dimerize, two hex- amers pair to form 900 x 10* d dodecamers, the most common aggregate. The further formation of still larger aggregates remains somewhat unclear but some hypotheses suppose that a still different kind of chain is required to make the 1.5 xX 10° d 24-mers (icosatetramers, found in a number of che- licerates and a single group of crustaceans) and another still must be present to make 3 x 10° d 48-mers (tessaracontaoctamers, found in Limulus). In the arthropods especially, the progres- sive formation of higher order polymers is a physiologically important process because it lowers the number of osmotically active macromolecules without diminishing the O, carrying capacity of the blood, which allows the animal to maintain an excess of blood hydrostatic over colloid osmotic pressure and thus to form its primary urine (Snyder & Mangum 1982). Polymerization also makes more compact molecules, which minimizes viscosity and makes it easier for the animal to push its blood around. Arthropod Hes have a distinctive elec- tron dense image. In high resolution micro- graphs hexamers look like 12.5 um wide hexagons in top view and squares in side view. In lower resolution micrographs they appear more or less spherical. The hexagons PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON are actually trigons of kidney-shaped mono- mers and there are two stacks of them in a hexamer. With only one exception the two hexamers in a dodecamer are rotated 45° with respect to one another, so they look like one hexagon and one square. 24-mers look like two pairs of spherical hexamers divided by a 2 um cleft. The orientation of the 24-mers in a 48-mer is still somewhat uncertain. But a native 48-mer can disso- ciate to an aggregate that looks like a native 24-mer, which can dissociate to a native- looking dodecamer, etc. The dissociation products never include intermediate mul- tiples, 18- or 36-mers. One would not ex- pect to find 18-mers if the assembly process entails pairing first hexamers and then do- decamers. But there is no known structural constraint on putting a third dodecamer onto a 24-mer to make a triantahexamer. Molluscan hemocyanins: Molluscan Hcs are built of very large (450-500 x 103 d) monomers that have been likened to a string of beads because each contains not one but 7-8 active sites (Van Holde & Miller 1982). The 7-8 O, binding domains are covalently linked to one another, although the giant monomers are loosely linked together in higher order aggregates. The native polymer exists in the blood as 10 (4.5 x 10° d), 20 (no less than 9 x 10° d) or sometimes even more of these chains. The blood of many gastropods is opaque, not because the Hc is so highly concentrated but because it scat- ters so much light. We routinely prepare these molecules, incidentally, by spinning them down in a not especially powerful ul- tracentrifuge. In electron micrographs they look like cylinders of various heights de- pending on degree of polymerization, and the ends of the cylinders have ‘collars’ and ‘caps.’ Distribution of the hemocyanins: In the arthropods Hcs have long been known to occur in the crustaceans and chelicerates and several years ago I suggested that they might also have occurred in the trilobites, because trilobites inhabited the O, poor aquatic hab- itat (Mangum 1980). In the molluscs they VOLUME 103, NUMBER 1 have been found in four of the five classes examined (Polyplacophora, Bivalvia, Gas- tropoda and Cephalopoda), and my recent preliminary examination of the scaphopods was inconclusive (Mangum et al. 1987). So a few years ago I was surprised to run across a report of Hc ina centipede, a mem- ber of the largely terrestrial uniramian ar- thropods. Sundara Rajulu (1969) had the interesting idea that an O, carrier might be needed in the relatively primitive scutiger- omorph centipedes because it is the only group of Uniramia in which the tracheal system ends blindly in the blood rather than proceeding all the way to the tissues. His evidence was very suggestive but it did not include reversible O, binding, the critical property of an O, carrier. Perhaps for this reason the importance of the possibility had not been recognized by proponents of var- 10us points of view in the ongoing debate about arthropod phylogeny. At the time I was visiting Drs. Robert and Nora Terwil- liger at the University of Oregon Institute of Marine Biology. We went out into the woods and picked up the first centipede we saw and performed an electrophoretic anal- ysis of its blood. Sure enough the blood con- tained a Cu-binding polypeptide. But when we examined its absorption spectrum, it was totally different from that of a Hc. We had chosen a group of centipedes in which the tracheal system does terminate at the tis- sues, making a blood O, carrier redundant. When I returned to Virginia I asked the resident expert where one must go to collect scutigeromorph centipedes and he pointed to the ceiling and I grabbed an individual of Scutigera coleoptrata and started working on it.? Centipedal Hemocyanin Structural and functional properties. —The blood of the common house centipede has the typical absorption spectrum of Hc, with 3 later learned that my bathtub is a far better col- lecting site. 241 a protein band at 280 and a Cu-O, maxi- mum at 339 wm (Mangum et al. 1985). Fol- lowing deoxygenation the absorption peak at the active site disappears and following re-oxygenation the peak reappears. Already this means that a Hc is present. Detailed measurements of O, binding revealed an exceptionally low oxygen affinity, which may prove to be related to high blood PO, in this terrestrial animal, and exceptionally great cooperativity that increases markedly with oxygenation state (Fig. 3). The Bohr shift is typically arthropod in magnitude and also normal, unlike that of some chelicerates. However, the direction of the Bohr shift varies within fairly closely related species so it cannot be regarded as taxonomically useful. Centipedal Hc is fairly concentrated in the blood, which is typical of terrestrial ar- thropods, and the concentration almost cer- tainly varies with hydration state in these desiccation-prone animals. In our original report I said that the blood, mercifully, fails to clot but this statement may slightly mis- represent the case. At least in small bore capillary tubes and at the termini of severed antennae, the blood becomes sticky and ceases to flow. In electrophoretic procedures intended to estimate molecular size the centipede monomers co-migrate almost exactly with those of the blue crab Callinectes sapidus and slightly behind the smaller ones found in Limulus. Centipedal Hc has monomers of two different sizes, 72 and 80 x 10° d, figures very typical of the arthropod Hcs. In our early electron micrographs of the native polymer we saw only four hexamers and we thought that the native polymer might be a 24-mer, which would also be very conventional. But in the meantime our collaborators Drs. K. E. Van Holde and K. I. Miller of Oregon State University per- formed sedimentation equilibrium mea- surements, and they arrived at a molecular weight estimate of 2.8 x 10° d, in between what we would expect for a 24- and a 48- mer. We had noticed on our gels that, unlike 242 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 100 HcOo, 50 /o 0 20 40 Po,,mm Hg Fig. 3. Data from Mangum et al. (1985). C. sapidus and Limulus, centipede blood contains a lot of material that differs in size from typical arthropod monomers. The amount this material, 30-35% of the total, would be more than enough to bias the mo- lecular weight estimates. We thought that centipedal Hc might consist of garden-va- riety arthropod monomers somehow com- plexed with a uniquely uniramian matrix. But when Dr. Miller purified the Hc (which was no mean feat; I feel fortunate when I collect as much as 10 wl blood from an in- dividual, and I rarely find more than one or two animals per week and only from about May into October) and re-electrophoresed it, all of the extra material disappeared, leaving behind only the garden-variety ar- thropod monomers. By this time our mi- crographs fairly clearly showed more than four hexamers. The centipede polymer is a triantahex- amer, a multiple of 36 monomers. Initially O, binding of centipedal Hc at pH 7.5 and 25°C. 0.05 M Tris maleate buffered physiological saline. we were quite excited about this result be- cause, aS indicated above, Hc triantahex- amers had never before been found. But after contemplating it for just a little while we recognized that the aggregation of three dodecamers violates no known structural constraint. The real question is why there are not more of them. There is one structural feature of persis- tent interest. The electrophoretic procedure to which I alluded above is useful in char- acterizing molecular weight but it is not the most sensitive. For example, C. sapidus Hc contains monomers of only two different sizes but it can have as many as six chains separable in the native conformation by charge (Mason et al. 1983, Mangum & Rai- ner 1988). But centipedal Hc still contains only two monomers when we separate the native chains by charge. So, according to some of the current models of assembly, it should not have enough different chains to VOLUME 103, NUMBER I 243 Fig. 4. Electron dense images of centipedal Hc. See text for explanation. make the native polymer! This is of no phy- logenetic interest but it does mean that the process of assembly must be rethought. In electron micrographs we see nothing about centipedal Hc that fundamentally dis- tinguishes it from other arthropod Hcs (Fig. 4). One image shows two pairs of spheres, each sphere measuring about 12 um in di- ameter and each separated by a 2 um cleft. Garden variety hexamers. One pair is in sharper focus than the other and one pair is slightly skewed with respect to the other. A second view shows two pairs of hexamers with a fifth in the middle. A third view shows a triangle of substructures surrounding a fourth. Two of the three peripheral units are in sharper focus than the third and, on closer inspection, additional material even more out of focus can be seen on the flat sides. More complex images, suggesting larger numbers of hexamers are also common. All of these images can be produced by the model shown in Fig. 5. The six hexamers Fig. 5. are arranged in an octahedral array. Two of the three pairs are lined up side by side and the third is located in an axis perpendicular to the other two. When viewed from the top or from one side only four hexamers can be seen. In this view the two pairs are not in exactly the same focus because one pair is on top and the other is underneath. When viewed from the other side five of the six can be seen. Slight tilt results in the virtual disappearance of the fifth and further tilting results in the virtual disappearance of a fourth. This is the most closed possible structure, permitting maximal interactions between the various subunits, which is con- sistent with the very great cooperativity of O, binding. In an alternative, less closed arrangement all three pairs might be lined up side by side. But this arrangement cannot yield the image of five symmetrically placed hexamers and the image of four would show them un- skewed and in the same focus. Finally, the Postulated arrangement of hexamers in centipedal triantahexamer. See text for explanation. 244 triangular image would not have additional material on the flat sides. Arthropod phylogeny. — What is the phy- logenetic significance of uniramian Hc? Again I must digress, to summarize briefly the various views of arthropod phylogeny. For more than two decades Manton (e.g., 1973, 1979) argued forcefully that the so- called Phylum Arthropoda is in fact a poly- phyletic assemblage of at least three and possibly four unrelated taxa. She considered the three extant groups to be of clearly dif- ferent origins and noted that the extinct Tri- lobita may be so. Her conclusion was based on the functional morphology of the limbs, both locomotor and masticator. The sheer volume of morphological evidence she amassed in support of this inference was so formidable that few non-experts such as myself dared attempt to assess it. Her view of the living groups was supported by An- derson (1973), who compared their devel- opmental morphology and concluded that features such as the fate map of the blastula are quite different. Because the develop- mental biologists among my colleagues usu- ally decline comment on the fundamental- ity of blastular fate maps, I cannot assess this evidence either. The arguments were articulated so persuasively that they have even begun to appear in recent textbooks (e.g., Barnes 1987). Recently, however, more conservative views have been revived, also persuasively. Hessler & Newman (1975) considered all of the living species of primitive crustaceans and mentally dissected away their special- ized features, leaving behind only the prim- itive ones. From them they reconstructed a hypothetical primitive crustacean and con- cluded that it could have descended from a trilobite, which they also regarded as a vi- able candidate for the ancestral chelicerate. They argued that the Phylum Arthropoda is at most diphyletic and at least one reason for not favoring monophyly is that they did not even address the question of the origin of the Uniramia. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Most of the contributors to a volume on the subject (Gupta 1979) favored mono- phyly. While many of the arguments were based on a single character, taken together they are fairly cogent. Still more so are chap- ters by Clarke (1979), who broadly surveyed internal organ systems and came down in favor of monophyly, and Boudreaux (1979), who tried to show that 17 both derived and shared characters are not convergent. Boud- reaux (1979), however, went to some lengths to emphasize that many of the 17 characters are the same ones used by Manton (1973, 1979) to support her case, which seems to me to indicate only the ease of alternative interpretation and thus to weaken his case. The importance of Uniramian Hc in ar- thropod phylogeny. — By far the simplest in- terpretation of our findings is that the native arthropod Hc polymer is an 18th derived and shared character with a common, single origin. In each of the three living groups the native polymer is built of polypeptide chains of about the same size and linked to one another in the same way. In the uniramians, the crustaceans and all chelicerates except Limulus, the size of the chains is essentially identical; thus the monomers may be prim- itive features. In each group the hexamer, the basic functional unit that exhibits many of the critical respiratory features has the same size and the same appearance. The hexamer may also be a primitive character. But, the arrangement of hexamers in larger aggregates is distinctive and characteristic of the particular group. Recently, the ho- mology (in the strict evolutionary sense) of the crustacean and chelicerate Hcs has been further documented by elegant and quite compelling structural evidence (Linzen et al. 1985, Markl 1986). My inference does not clearly predict the condition of the blood in the onycophorans, which are variously regarded as more or less peripheral to the mainstream of uniramian evolution. H. D. Ellerton (Ellerton et al. 1983 and pers. comm.) could find none but he does not believe that his evidence is com- VOLUME 103, NUMBER 1 245 Table 2.—The relationship of the respiratory properties of selected arthropod and molluscan Hcs. Data from Mangum & Lykkeboe (1979), Mangum (1983), Mason et al. (1983), Diefenbach & Mangum (1983), Mangum & Burnett (1986) and Mangum (unpubl.). Busycon canaliculatum Bohr shift reversed O, affinity at physiological pH moderate Cooperativity moderate Inorganic ion sensitivity broad L-lactate sensitivity none Urate sensitivity none CO, sensitivity indirect pletely decisive. Sundara Rajulu (pers. comm.) suspects that Hc may be there. If so, it will be most exciting to examine its quaternary structure. My hypothesis does require the existence of Hc in the Trilobita. The elemental com- position of trilobite fossils would be most interesting to learn, if it is technically pos- sible. The role of biochemical evidence in a physiological context. —Is this character more persuasive than others because it en- tails molecular structure and is therefore “closer to the gene’? Of course not—cer- tainly not quaternary structure. But I sug- gest that it is more persuasive than Man- ton’s and Boudreaux’s morphological characters for two reasons. First, as I will show below, on good Darwinian principles this evidence cannot be interpreted alter- natively. Second, we know a great deal about these molecules and how they work in liv- ing, breathing animals and so we know which features are and are not critical to respira- tory function and hence subject to natural selection. The database on which my sum- mary of structure is based consists of almost 200 Hes. We have O, binding data obtained under fairly physiological conditions for about 150 arthropod and 80 molluscan Hes. We have information on in vivo blood PO,, pH and oxygenation state for probably sev- eral dozens of arthropods though, regrett- ably, only perhaps one dozen molluscs. We know how much oxygenation occurs at the Limulus polyphemus Callinectes sapidus reversed normal moderate moderately low moderate very great broad narrow none great none great indirect direct gill and why, how much deoxygenation oc- curs at the tissues and why, and what en- vironmental and physiolgical challenges perturb the system and how much and why. We know how big a role the He plays in total aerobic metabolism, that most fun- damental process in life. We can make quantitative predictions of the hypothetical consequences of altering particular respi- ratory properties of the molecule. Let us suppose for the moment that my conclusion is wrong, that the arthropod Hcs arose independently on three separate oc- casions and that their common quaternary structures are the products of convergence. If that were true, there must have been an overwhelming selection pressure for that particular structure, almost certainly to dic- tate particular respiratory properties. And yet no such selection pressure exists. It 1s very clear that the quaternary structure of a He does not constrain respiratory prop- erties. I remind the reader once more that we have an actual example of a convergent Hc, the molluscan variety. And the O, bind- ing properties of molluscan and arthropod Hcs, convergent molecules, can be more alike than those of two arthropod Hcs, ho- mologous molecules (Table 2). So very sim- ilar respiratory properties can be associated with fundamentally dissimilar quaternary structures and one cannot argue that the ar- thropod structure was evolved repeatedly to preserve any particular respiratory proper- ties because it simply does not do so. By far 246 the more probable interpretation is that the arthropod structure arose only once and that it is an inherited feature that (along with others) unites at least the three living groups in a natural, monophyletic taxon. There must be other examples of mole- cules understood so well at both structural and physiological levels. I shall welcome the time when physiologists and biochemists once again regard animal phylogeny as a proper subject of inquiry. Acknowledgments This essay was written while the author was supported by the National Science Foundation (DCB 88-16172, Physiological Processes). Literature Cited Anderson, D. F. 1973. Embryology and phylogeny of annelids and arthropods. Oxford, Pergamon Press, 495 pp. Baldwin, E. 1949. An introduction to comparative biochemistry. Cambridge University Press, Cambridge, 164 pp. 1953. Biochemistry and evolution.—Society for Experimental Biology, Symposium 7:22-30. 1959. Dynamic aspects of biochemistry. Cambridge University Press, Cambridge, 526 pp. ——, & D. M. Needham. 1937. A contribution to the comparative biochemistry of muscular and electric tissues.— Proceedings of the Royal So- ciety, London B 122:197-205. ——.,, & W.N. Yudkin. 1950. The annelid phos- phagen, with a note on phosphagen in Echino- dermata and Protochordata. — Proceedings of the Royal Society of London B136:614-631. Barnes, R. D. 1987. Invertebrate zoology. W. D. Saunders & Co., Philadelphia, 893 pp. Boudreaux, H. B. 1979. Significance of intersegmen- tal tendon system in arthropod phylogeny and a monophyletic classification of Arthropoda. Pp. 551-556 in A. P. Gupta, ed., Arthropod phy- logeny. Van Nostrand Reinhold Co., New York. Clarke, K. U. 1979. Visceral anatomy and arthropod phylogeny. Pp. 467-549 in A. P. Gupta, ed., Arthropod phylogeny. Van Nostrand Reinhold Co., New York. Dickerson, R. E., & I. Geis. 1969. Structure and ac- tion of proteins. Harper & Row, New York, 120 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Diefenbach, C.O. daC., & C. P. Mangum. 1983. The effects of inorganic ions and acclimation salinity on hemocyanin-oxygen binding in the horse- shoe crab Limulus polyphemus. —Molecular Physiology 4:197-206. Ellerton, H. D., N. F. Ellerton, & H. A. Robinson. 1983. Hemocyanin—a current perspective. — Proceedings of Biophysics and Molecular Bi- ology 41:413-248. Ennor, A. H., & J. F. Morrison. 1958. Biochemistry of the phosphagens and related guanidines. — Physiological Reviews 38:631-674. Field, K. G. et al. 1988. Molecular phylogeny of the animal kingdom.—Science 239:748-753. Greenwald, I. 1946. The presence of creatine in the testes of various invertebrates. — Journal of Bi- ological Chemistry 162:239-250. Gupta, A. G. 1979. Arthropod phylogeny. Van Nos- trand Reinhold Co., New York, 762 pp. Hessler, R. R., & W. A. Newman. 1975. A trilobito- morph origin for the Crustacea.— Fossils and Strata 4:437-459. Hird, F. J. R. 1986. The importance of arginine in evolution.—Comparative Biochemistry and Physiology 85B:285-288. Imamura, T., T. O. Baldwin, & A. Riggs. 1972. The amino acid sequence of the monomeric hemo- globin component from the bloodworm Glycera dibranchiata. —Journal of Biological Chemistry 247:2785—2797. Kidron, M., & R. Segal. 1984. The new state of the world atlas. Simon & Schuster, New York, no pagination. Kuhn, T. S. 1970. The structure of scientific revo- lutions. University of Chicago Press, Chicago, 210 pp. Kutscher, F., & D. Ackermann. 1933. The compar- ative biochemistry of vertebrates and inverte- brates. — Annual Review of Biochemistry 2:355— 390. Lane, D. J., D. A. Stahl, G. J. Olsen, & N. R. Pace. 1985. Analysis of hydrothermal vent-associ- ated symbionts by ribosomal RNA se- quences. — Biological Society of Washington Bulletin 6:389—400. Linzen, B. et al. 1985. The structure of arthropod hemocyanins.—Science 229:519-524. Mangum, C. P. 1980. Respiratory function of the hemocyanins.— American Zoologist 20:19-38. . 1983. The effect of hypoxia on hemocyanin- oxygen binding in the horseshoe crab Limulus polyphemus.— Molecular Physiology 3:217—224. , & L. E. Burnett. 1986. The CO, sensitivity of the hemocyanins and its relationship to Cl- sensitivity. — Biological Bulletin 171:248-263. —.,, & G. Lykkeboe. 1979. The influence of in- organic ions and pH on the oxygenation prop- erties of the blood in the gastropod mollusc Bu- VOLUME 103, NUMBER 1 sycon canaliculatum. — Journal of Experimental Zoology 207:417-430. ——., KI. Miller, J. L. Scott, K. E. Van Holde, & M. P. Morse. 1987. Bivalve hemocyanin: Structural, functional and evolutionary rela- tionships. — Biological Bulletin 173:205-221. , & J. S. Rainer. 1988. The relationship be- tween subunit composition and oxygen binding of blue crab hemocyanin.—Biological Bulletin 174:77-82. ——., J. L. Scott, R. E. L. Black, K. I. Miller & K. E. Van Holde. 1985. Centipedal hemocyanin: Its structure and its implications for arthropod phylogeny. — Proceedings of the National Acad- emy of Science USA 82:3721-3725. Manton, S. M. 1973. Arthropod phylogeny—a mod- ern synthesis. — Journal of Zoology 171:111-130. 1979. Uniramian evolution with particular reference to the pleuron. Pp. 317-343 in M. Ca- matini, ed., Myriapod biology. Academic Press., London. Markl, J. 1986. Evolution and function of structurally diverse subunits in the respiratory protein he- mocyanin from arthropods.— Biological Bulle- tin 171:90-115. Mason, R. P., C. P. Mangum, & G. Godette. 1983. The influence of inorganic ions and acclimation salinity on hemocyanin-oxygen binding in the blue crab Callinectes sapidus. —Biological Bul- letin 164:104—123. Needham, D. M., J. Needham, E. Baldwin, & J. Yud- kin. 1932. A comparative study of the phos- phagens with some remarks on the origins of vertebrates. — Proceedings of the Royal Society of London B110:260-272. Pace, N. R., D. A. Stahl, D. J. Lane, & G. J. Olsen. 247 1986. The analysis of natural microbial pop- ulations by rRNA sequences. — Advances in Mi- crobial Ecology 9:1-56. Snyder, G. K., & C. P. Mangum. 1982. The rela- tionship between the size and shape of an ex- tracellular oxygen carrier and the capacity for oxygen transport. Pp. 173-188 in J. Bonaven- tura, C. Bonaventura, & S. Tesh, eds., Physi- ology and biology of horseshoe crabs. Allan Liss Co., New York. Sundara Rajulu, G. 1969. Presence of haemocyanin in the blood of a centipeded Scutigera longi- cornis (Chilopoda: Myriapoda).—Current Sci- ence 38:168-169. Van Holde, K. E., & K. I. Miller. 1982. Haemocy- anins.— Quarterly Review of Biophysics 15:1- 129. van Thoai, N., & Y. Robin. 1969. Guanidine com- pounds and phosphagens. Pp. 163-204 in M. Florkin & B. T. Scheer, eds., Chemical zoology Vol. IV, Academic Press., New York. Weber, R. E., C. Mangum, H. Steinman, C. Bonaven- tura, B. Sullivan, & J. Bonaventura. 1977. Hemoglobins of two terebellid polychaetes: En- oplobranchus sanguineus and Amphitrite orna- ta. Comparative Biochemistry and Physiology 56A:179-187. Zuckerkandl, E., & L. Pauling. 1965. Molecules as documents of evolutionary history.— Journal of Theoretical Biology 8:357-366. Department of Biology, College of Wil- lam and Mary, Williamsburg, Virginia 23185. PROC. BIOL. SOC. WASH. 103(1), 1990, pp. 248-253 SIX NEW COMBINATIONS IN BACCHAROIDES MOENCH AND CYANTHILLIUM BLUME (VERNONIEAE: ASTERACEAE) Harold Robinson Abstract.—Three species, Vernonia adoensis Schultz-Bip. ex Walp., V. gui- neensis Benth., and V. lasiopus O. Hoffm. in Engl., are transferred to the genus Baccharoides Moench, and three species, Conyza cinerea L., C. patula Ait., and Herderia stellulifera Benth. are transferred to the genus Cyanthillium Blume. The present paper provides six new com- binations of Old World Vernonieae that are known to belong to the genera Baccharoides Moench and Cyanthillium Blume. The ap- plicability of these generic names to these species groups was first noted by the author almost ten years ago (Robinson et al. 1980), and it was anticipated that other workers more familiar with the paleotropical mem- bers of the Vernonieae would provide the necessary combinations. A recent study of eastern African members of the tribe by Jef- frey (1988) also cites these generic names as synonyms under his Vernonia Group 2 subgroup C and Vernonia Group 4, al- though he retains the broad concept of Ver- nonia. At this point it is not certain when anyone will undertake more detailed studies of these groups. However, the author does need to refer to a few members of these genera and does not intend to use older, incorrect names. Instead of making isolated combinations as needed, as was done with Gymnanthemum coloratum (Robinson & Kahn 1986), an effort is made here to place needed combinations together in one paper. Detailed monographic studies and the ma- jority of the combinations are left to future workers. In spite of the limited scope of the present effort, some explanation is offered. The summary of the Vernonieae by S. B. Jones (1977) showed that the Eastern Hemi- sphere species placed in the genus Vernonia were all chemically and cytologically dis- tinct from the Western Hemisphere mem- bers of that genus. Although generic limits were not discussed by Jones, his study placed the Old World Vernonia in a group on the opposite side the basic division in the genus from typical Vernonia in the eastern United States. Subsequent studies by Jones (1979b, 1981) showed that certain pollen types also were restricted to Old World members of Vernonia s.1., types that are shared by some Old World members of the tribe tradition- ally placed in other genera. The characters noted by Jones have been treated by the present author as evidence of a basic divi- sion in the Vernonieae between groups that have included many genera in each hemi- sphere, and traditional Vernonia s.l. can be seen as an excessively paraphyletic core ge- nus that is defined only by what “‘it is not” rather than by what “‘it is.”’ In progressing toward definitive generic concepts in the Vernonieae, the pattern seen by Jones jus- tifies removing at least all of the Eastern Hemisphere members of the tribe from Ver- nonia. This segregation of the Old World genera was partially put into practice by the use of the names Baccharoides Moench, Gymnanthemum Cass., and Cyanthillium Blume, by Robinson et al. (1980) and in the transfers made in the study of Distephanus Cass. (Robinson & Kahn 1986). At present there is good reason not to transfer all the Old World species of Ver- nonia out of the genus. Morphological and VOLUME 103, NUMBER 1 anatomical studies show the problem is more complex than one Old World genus versus one New World genus. Several gen- era should be recognized in both the neo- tropical and paleotropical elements of Ver- nonia s.l1. More detailed studies of the type under way in neotropical species (Robinson 1987a, b, c, 1988a, b, c, 1989; Robinson & Funk 1987) are needed before all transfers can be made properly. Nevertheless, the po- sition of some of the species is fully re- solved, especially of species belonging to older segregate genera such as Baccharoides and Cyanthillium. The lack of total resolution of the tribe is not a reason to retain under Vernonia s.1. names and concepts known to be obsolete. The author believes that Vernonia will ul- timately be restricted to Western Hemi- sphere species with non-lophate type A pol- len and a chromosome number of N = 17 and will perhaps be restricted to only the eastern North American element of that group. The detailed variations of pollen in the tribe are to be more fully compared and illustrated elsewhere. The generic synony- mies follow mostly those of Jeffrey (1988), but some additional synonymy is provided under Cyanthillium. Baccharoides Moench Baccharoides Moench, Methodus. 528. 1794. Type: Conyza anthelmintica L. (=B. anthelmintica (L.) Moench). Ascaricida Cass., Dict. Sci. Nat. 3, suppl. 38. 1817, nom. superfl. Type: Conyza an- thelmintica L. Candidea Tenore, Atti Reale Accad. Sci. Sez. Soc. Reale Borbon 4 (Cl. Botan.): 104, t. 1, 2. 1839. Type: Candidea senegalensis Tenore. Vernonia subsect. Stengelia Schultz-Bip. ex Walp., Repert. Bot. Syst. 2: 946. 1843. Vernonia adoensis Schultz-Bip. ex Walp. Stengelia Steetz in Peters, Naturw. Reise Mossambique, Bot. 360. 1864. Type: Vernonia schimperi DC. 249 Vernonia sect. Stengelia (Schultz-Bip. ex Walp.) Benth. in Benth. & Hook. f., Gen. AL, Be WD Wea Si, This group of species has been imper- fectly recognized in the past, primarily on the basis of expanded foliose appendages on the involucral bracts. The character is well developed in the genus but is not a reliable basis for distinction. Smith (1971) studied a group titled the “‘stengelioid species” of Vernonia, but many species included in the study were not Baccharoides. Smith recog- nized the heterogeneity in the group he treated, especially in the form of the pollen and in the shape of the corolla, but these were not detailed for all of the species. More recently, Kingham (1976), Jones (1981), Jeffrey (1988), and the present au- thor have studied Baccharoides pollen in more detail. It is lophate and distinctive among Old World tricolporate Vernonieae by the presence of polar areoles. The grains fall into Jones’ (1981) Type C, although pol- lens of all species studied in detail are dif- ferent from most Type C pollen of the Neo- tropics by the 3 equatorial areoles of the intercolpi and the strong basal columellae under the crests of the exine. Jeffrey (1988) emphasized the pollen, the flattened inner pappus bristles, and the co- rollas with long, slender, basal tubes and abruptly expanded cylindrical limbs and comparatively short erect lobes as definitive characters of Baccharoides within Vernonia s.1. Ignoring some older counts, the genus seems to have a consistent chromosome number of N = 10 (Smith 1971, Jones 1979a). The group is also notable for the presence of epoxy resins and has been stud- ied for possible commerical exploitation. Neither the chromosome number nor the chemistry is unique to the genus. Many of the species are understood well enough to justify new combinations, but most of these, and others not known to the present author, are left for final resolution by other workers. The species listed by Smith 250 (1971) are an artificial group and the Smith paper should not be used without reference to the discussion by Jeffrey (1988). Except for its discussion of the Smith paper, the Jeffrey study itself is limited to East African species. It seems that a monograph of the genus would be comparatively simple for someone with better resources to study pa- leotropical plants, for most of the species occur in Africa. One species was described from Arabia, and the widely distributed type species was described from Sri Lanka. The three species combinations needed at present are as follows: BACCHAROIDES ADOENSIS (Schultz-Bip. ex Walp.) H. Robinson, comb. nov. Vernonia adoensis Schultz-Bip. ex Walp., Repert. Bot. Syst. 2:946. 1843. For ex- tensive synonymy see Smith (1971) and Jeffrey (1988). Distribution is subsaharan Africa with concentration in the east (Smith 1971, map p. 19). BACCHAROIDES GUINEENSIS (Benth. in Hook. f. & Benth.) H. Robinson, comb. nov. Vernonia guineensis Benth. in Hook. f. & Benth., Niger Fl. 427. 1849. For extensive synonymy see Smith (1971). Distribution is subsaharan Africa from Sierra Leone in the west eastward to western Tanzania and south to southwestern Angola and northern Zimbabwe (Smith 1971, map p. 48). BACCHAROIDES LASIOPUS (O. Hoffm. in Engl.) H. Robinson, comb. nov. Vernonia lasiopus O. Hoffm. in Engl., Pflan- zew. Ost. Afrika 403. 1895. For extensive synonymy see Smith (1971). The species ranges from western Ethiopia to north- eastern Angola and northeastern Zim- babwe (Smith, 1971, map p. 38). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cyanthillium Blume Cyanthillium Blume, Bijdr. Fl. Ned. Ind. 889. 1826. Lectotype (designated here): Cyanthillium villosum Blume. Isonema Cass., Bull. Soc. Philom. Paris 1817:152. 1817, nom. illeg., non R. Br. (1810). Type: Isonema ovata Cass. Cyanopis Blume ex DC., Prodr. 5:69. 1836, nom. illeg. et superfl., non Cass. (1817). Lectotype (Jones 1980): Cyanthillium vil- losum Blume. Vernonia sect. Tephrodes DC., Prodr. 5:24. 1836. Lectotype (Jones 1981): Conyza ci- nerea L. Claotrachelus Zoll. & Mortiz ex Zoll., Na- tuur-Geneesk. Arch. Ned Indié 2:263, 565. 1845. Type: Claotrachelus rupestris Zoll. & Moritz ex Zoll. Seneciodes L. ex Post & O. Kuntze, Lex. Gen. Phan. 2:515. 1903. Type: Conyza cinerea L. Triplotaxis WHutch., Bull. Misc. Inform. 1914:355. 1914. Lectotype: Herderia stel- lulifera Benth. in Hook. Vernonia subsect. Orbivestus S. B. Jones, Rhodora 83:61. 1981. Lectotype (Jeffrey 1988): Vernonia karaguensis Oliv. & Hiern. Vernonia subsect. Hilliardianae S. B. Jones, Rhodora 83:66. 1981. Lectotype (Jeffrey 1988): Webbia oligocephala DC. Vernonia subsect. Tephrodes (DC.) S. B. Jones, Rhodora 83:70. 1981. The designation of a new lectotype for Cyanthillium rejects the earlier choice of C. moluccense that seems to date from Jones (1980). Jones’ selection was totally arbi- trary, being the first species listed by Blume without any evident study of the group in- volved, and thus can be rejected under the Code. The lectotype cited by Jeffrey (1988) evidently follows Jones. Jones’ lectotypifi- cation is unsuitable because the species is virtually unknown, being available to most botanists only as a microfiche of the frag- ment in the DeCandolle herbarium at Ge- neva. The fragment C. moluccense as seen VOLUME 103, NUMBER 1 in the microfiche may be a Cyanthillium. The choice in this paper brings the typifi- cation into conformity with those of the synonyms, the superfluous generic name Cyanopis Blume and the homonym I[sone- ma Cass., which both equal Cyanthillium patula. It is unfortunate that many Flora formats now require types to be selected for all ge- neric names, even those treated as syn- onyms. Although the intention is noble, the result is too often poorly chosen lectotypes, made by unqualified investigators. Any study of Cyanthillium should in- clude a study of members of Erlangea Schultz-Bip. (1853). The type, E. plumosa Schultz-Bip., of Gabon, is known to the au- thor only from its description. Erlangea is technically distinguished from Vernonia s.1. and that part recognized here as Cyanthil- lium by the five, deciduous pappus seg- ments that have been referred to as plu- mose. The type of Erlangea is described with sessile amplexicaule leaves, a feature that would be unusual in the usually nar- rowly petiolate Cyanthillium. As described, the type of Erlangea has a habit similar to Vicoa (Inula) auriculata Cass. and it may be close to Erlangea schinzii O. Hoffm. of northernmost Namibia, which also has ses- sile leaves. Examination of material match- ing the description of the type, and further evaluation of the pappus distinction, may show that the genus falls fully into the syn- onymy of Cyanthillium. The genus Cyanthillium owes its early recognition to the presence of a deciduous pappus in the species group that includes the lectotype species. The broad achene and short deciduous pappus of C. patula places this species outside of typical Vernonia. Other pappus variations occur in Cyanthil- lium and have been the basis of further ge- neric distinctions. The type species of Trip- lotaxis has a very distinctive, collar-like outer pappus that persists after the inner pap- pus has completely fallen. Other species from Africa that are described as having a pappal 251 ring don’t have the highly distinctive struc- ture seen in 7riplotaxis but have only a moderately expanded upper callus on the achene of a form seen in many Asteraceae. The latter species are not closely related to Cyanthillium. Other members of Cyanthil- lium, such as C. cinereum, have achene and pappus structures like those of typical Ver- nonia, without unusual rings or fragility. Be- cause of the differences in pappus structure listed above, the species now placed in Cyanthillium were not suspected of forming arelated group separate from Vernonia until the pollen study by Kingham (1976) and the recent study by Jeffrey (1988). Nevertheless, the species seem to share a generally her- baceous habit, herbaceous and usually nar- rowly petiolate leaves, and broadly to nar- rowly ovate, slightly acuminate involucral bracts, often with greenish outer surfaces and purplish margins. Jeffrey (1988) men- tions various forms of T-shaped hairs in members of the group. The pollen type of Cyanthillium is one of the many paleotropical types in the Ver- nonieae in which the colpus is not evident. Studies by the author show that the polar areas of the grains have a polar areole sur- rounded by a tier of five to seven areoles that variously match or straddle or fall be- tween the pores below them. The polar or- ganization is basically different from forms in the tribe that have well developed colpi. In the present group there is not even a consistent single areole above or below the pores. The major organizational feature of the grain is centered on the poles rather than the pores. SEM studies show that members of Cyanthillium are distinctive within the non-colporate species of the Old World by the modification of the basal columellae of the exine to form bridges under the crests. Points of attachment of the exine to the footlayer seem to be restricted to positions under the intersections of the crests. The precise pollen form is not presently known outside the genus, although similar forms without modified basal columellae are 252 known in species belonging to the group cit- ed by Jeffrey (1988), for which the name Crystallopollen Steetz in Peters seems to have priority. The few available chromosome numbers in Cyanthillium fall within the range noted for paleotropical members of the tribe. Jones (1979a) reported N = 20 for C. stelluliferum (as Triplotaxis). This seems to be based on X = 10. Jones reported N = 9, 18, and ca. 18 for the widespread adventive C. cine- reum. Jones also reported N = 10 for two species of the closely related genus Erlan- gea, including E. remifolia Wild & Pope, which I suspect is a member of Cyanthil- lium. Cyanthillium has problems not seen in Baccharoides, and a monograph will be more difficult. The distribution is throughout the paleotropical region, and there are many variations within the group in obvious char- acters such as the achene and pappus. The three combinations needed at present are as follows: CYANTHILLIUM CINEREUM (L.) H. Robinson, comb. nov. Conyza cinerea L., Sp. Pl. 862. 1753. Vernonia cinerea (L.) Less., Linnaea 4:291. 1829. Senecioniodes cinerea (L.) Post & O. Kuntze, KexsiGens Phan 225155 1903) The species is the most widely distributed member of the genus. It occurs throughout the paleotropical region and is widely ad- ventive in the Neotropics. CYANTHILLIUM PATULUM (Ait.) H. Robinson, comb. nov. Conyza patula Ait., Hortus Kew. 3:184. 1789. Isonema ovata Cass., Bull. Soc. Philom. eA WSIS Ws TGs Conyza chinensis Lam., Encycl. 2:83. 1786, hom. illeg., non L. Cyanthillium villosum Blume, Bijdr. FI. Ned. Ind. 889. 1826. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cyanthillium pubescens Blume, Bidr. FI. Ned. Ind. 890. 1826. Centratherum chinense [Lam.] Less., Lin- naea 4:320. 1829. Vernonia chinensis (Less.) Less., Linnaea 6: 105, 674. 1831. Cyanopis madagascariensis DC., Prodr. 5: 69. 1836. Vernonia pratensis Klatt, Ann. K. K. Na- turhist. Hofmus. 7:99. 1892. Cacalia patula (Less.) O. Kuntze, Rev. Gen. Pl. 324. 1894. Cyanthillium chinense (Lam.) Gleason, Bull. Torrey Bot. Club 40:306. 1913. The species is cited by Gleason (1922) as occurring in tropical Asia, the East Indies, and being introduced into Guadeloupe. The species is now also credited to Madagascar on the basis of the newly synonymized Cy- anopsis madagascariensis and Vernonia pratensis. CYANTHILLIUM STELLULIFERUM (Benth.) H. Robinson, comb. nov. Herderia stellulifera Benth. in Hook. f. & Benth., Niger Fl. 425. 1849. Triplotaxis stellulifera (Benth.) Hutch., Bull. Misc. Inform. 1914:356. 1914. The distribution given by Jeffrey (1988) is Uganda, west and central tropical Africa and Angola. Literature Cited Gleason, H. A. 1922. Tribe 1. Vernonieae.— North American Flora 33:47-110. Jeffrey, C. 1988. The Vernonieae in East Tropical Africa.— Kew Bulletin 43:195-277. Jones, S. B. 1977. Vernonieae—Systematic review. Pp. 503-521 in V. H. Heywood et al., eds., The biology and chemistry of the Compositae. Ac- ademic Press, London, New York, 1189 pp. . 1979a. Chromosome numbers of Vernonieae (Compositae).— Bulletin of the Torrey Botanical Club 106:79-84. . 1979b. Synopsis and pollen morphology of Vernonia (Compositae: Vernonieae) in the New World.— Rhodora 81:425-447. VOLUME 103, NUMBER 1 1980. Tribe Vernonieae. Jn J. F. Macbride, ed., Flora of Peru.—Fieldiana, Botany n. s. 5: 22-69. . 1981. Synoptic classification and pollen mor- phology of Vernonia (Compositae: Vernonieae) in the Old World.—Rhodora 83:59-75. Kingham, D. L. 1976. A study of the pollen mor- phology of tropical African and certain other Vernonieae (Compositae). —Kew Bulletin 31:9— 26. Robinson, H. 1987a. Studies of the Lepidaploa com- plex (Vernonieae: Asteraceae). I. The genus Stenocephalum Sch.Bip.—Proceedings of the Biological Society of Washington 100:578-583. 1987b. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). II. A new genus, Echinocoryne.—Proceedings of the Biological Society of Washington 100:563-589. 1987c. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). III. Two new genera, Cyrtocymura and Eirmocephala. — Proceedings of the Biological Society of Washington 100: 844-855. 1988a. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). IV. A new genus, Les- singianthus. — Proceedings of the Biological So- ciety of Washington 101:929-951. 1988b. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). V. A new genus, Chrysolaena.—Proceedings of the Biological Society of Washington 101:952-958. 253 1988c. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). VI. A new genus, Ay- nia. — Proceedings of the Biological Society of Washington 101:959-965. —., & V.A. Funk. 1987. A phylogenetic analysis of Leiboldia, Lepidonia, and a new genus Stra- mentopappus (Vernonieae: Asteraceae).—Bo- tanische Jahrbiicher fiir Systematik, Pflanzen- geschichte und Pflanzengeographie 108:213-228. ——., & B. Kahn. 1986. Trinervate leaves, yellow flowers, tailed anthers, and pollen variation in Distephanus Cassini (Vernonieae: Astera- ceae).— Proceedings of the Biological Society of Washington 99:493-501. , F. Bohlmann, & R. M. King. 1980. Che- mosystematic notes on the Asteraceae. III. Nat- ural subdivisions of the Vernonieae.—Phyto- logia 46:421-436. Schultz-Bip., C. H. 1853. Triga novorum Cassiniace- arum generum.—Flora 36:33-38. Smith, C. E. 1971. Observations on stengelioid species of Vernonia. Agriculture Handbook no. 396. U.S.D.A., Agricultural Research Service. Department of Botany, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560. 254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE % BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON, SW7 5BD Applications Published in the Bulletin of Zoological Nomenclature The following applications were published on 29 September 1989 in Vol. 46, Part 1 of the Bulletin of Zoological Nomenclature. Comment or advice on these appli- cations is invited for publication in the Bulletin and should be sent to the Executive Secretary, ICZN, British Museum (Natural History), Cromwell Road, London SW7 5BD, U.K. Case No. 2652 CHORISTIDAE Verrill, 1882 (Mollusca, Gastropoda) and CHORISTIDAE Esben-Petersen, 1915 (Insecta, Mecoptera): a proposal to remove the homonymy. 2682 Fryeria Gray, 1853 and F. rueppelii Bergh, 1869 (Mollusca, Gastropoda): proposed conservation. 2662 Aphonopelma Pocock, 1902 (Arachnida, Araneae) proposed precedence over Rhechostica Simon, 1892. 2696 Ixodes angustus Neumann, 1899 and I. woodi Bishopp, 1911 (Arachnida, Acari): proposed conservation by the replacement of the holotype of J. angustus by a neotype. Castiarina Gory & Laporte, 1837 (Insecta, Coleoptera): proposed conser- vation. Helophorus brevipalpis Bedel, 1881 (Insecta, Coleoptera): proposed prece- dence over Helophorus creticus Kiesenwetter, 1858. Helophorus obscurellus Poppius, 1907 (Insecta, Coleoptera): proposed pre- cedence over Helophorus fausti Kuwert, 1887. Ceratopogon puncticoollis Becker, 1903 (currently Culicoides puncticollis; Insecta, Diptera): proposed precedence over Ceratopogon algecirensis Strobl, 1900. Micropterus patachonicus King, 1831 and Anas pteneres Forster, 1844 (both currently in Tachyeres Owen, 1875; Aves, Anseriformes): proposed conservation of the specific names. VOLUME 103, NUMBER 1 INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE Opinions Published in the Bulletin of Zoological Nomenclature The following Opinions were published on 29 September 1989 in Vol. 46, Part 1 of the Bulletin of Zoological Nomenclature. Opinion 1549 1550 1551 1552 1553 1554 ISDS 1556 LOST) 1558 1559 1560 1561 1562 1563 1564 1565 1566 EUGLENIDAE Stein, 1878 (Protista, Flagellata) and EUGLENIDAE Seid- litz, 1875 (Insecta, Coleoptera): homonymy removed, and ADERIDAE Winkler, 1927 (Insecta, Coleoptera): given precedence over EUGLE- NESIDAE Seidlitz, 1875. Dysidea Johnston, 1842 (Porifera, Keratosa): conserved. Hypsibius Ehrenberg, 1848 (Tardigrada): Macrobiotus dujardini Doyére, 1840 designated as the type species. Dioctophyme Collet-Meygret, 1802 (Nematoda): spelling confirmed. ATYIDAE De Haan, [1849] (Crustacea, Decapoda) and ATYIDAE Thiele, 1925 (Mollusca, Gastropoda): homonymy removed. Cryptocoeloma Miers, 1884 (Crustacea, Decapoda): Cryptocoeloma haswelli Rathbun, 1923 designated as the type species. Parasigara Poisson, 1957 (Insecta, Heteroptera): Corisa transversa Fieber, 1848 confirmed as the type species. Dytiscus ater De Geer, 1774 (currently I/ybius ater) and Dytiscus planus Fabricius, 1781 (currently Hydroporus planus; Insecta, Coleoptera): specific names conserved. Elachista Treitschke, 1833 (Insecta, Lepidoptera): conserved, and E. bifas- ciella Treitschke, 1833 confirmed as the type species. Dacus parallelus Wiedemann, 1830 (currently Anastrepha parallela; Insecta, Diptera): lectotype replaced. Ludita Nagy, 1967 (Insecta, Hymenoptera): Tiphia villosa Fabricius, 1793 designated as the type species. Asterias squamata Delle Chiaje, 1828 (currently Amphipholis squamata; Echinodermata, Ophiuroidea): specific name conserved. Climacograptus manitoulinensis Caley, 1936 (currently Paraclimacograptus manitoulinensis, Graptolithina): specific name conserved. Anabas oxyrhynchus Boulenger, 1902 (currently Ctenopoma oxyrhynchum; Osteichthyes, Perciformes): specific name not conserved. Heliases ternatensis Bleeker, 1856 (currently Chromis ternatensis; Osteich- thyes, Perciformes): specific name conserved, and Chromis viridis (Cu- vier, 1830): name adopted for the fish formerly known as C. caerulea (Cuvier, 1830). Neamia octospina Smith & Radcliffe in Radcliffe, 1912 (Osteichthyes, Per- ciformes): specific name conserved. Platanista Wagler, 1830 (Mammalia, Cetacea): conserved. Megaloceros Brookes, 1828 (Mammalia, Artiodactyla): original spelling emended. 255 256 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON REVIEWERS The following people reviewed manuscripts for the Proceedings in 1989. A. Alvarinio; W. D. Anderson, Jr.; C. Baldwin; R. C. Banks; K. Banse; J. Barlow; A. Bauer; A. R. Beaumont; R. L. Bezy; R. Bleiweiss; W. Blow; J. Bouillon; T. E. Bowman; G. Boxshall; G.-O. Brandorff, R. O. Brinkhurst; B. Brown; W. C. Brown; N. L. Bruce; R. C. Brusca; J. S. Buckeridge; J. E. Cadle; S. D. Cairns; D. R. Calder; D. K. Camp; A. F. Campaner; E. Campos; D. Canatella; M. D. Carleton; J. Carlton; R. S. Carney; J. L. Carr; F. A. Chace, Jr.; B. Chernoff; E. Coan; S. L. Coats; J. S. H. Collins; B. C. Coull; J. Cracraft; R. F. Cressey; R. I. Crombie; A. Crosnier; G. M. Davis; H. J. Dumont; R. J. Emry; S. Farkus, K. Fauchald; D. L. Felder; R. M. Feldman; S. V. Fink; W. L. Fink; J. F. Fitzpatrick, Jr.; B. A. Foster; L. R. Franz; R. C. Froeschner; V. A. Funk; A. L. Gardner; J. Garth; R. D. Gordon; G. R. Graves; K. Green; H. Griffith; C. Hand; A. S. Harold; G. Hartmann; L. R. Heaney; R. Heard; J. W. Hedgpeth; G. F. Hevel; R. W. Heyer; R. P. Higgins; A. Hirayama; J. Ho; J. T. Hoeg; R. S. Hoffmann; J. R. Holsinger; W. D. Hope; P. Hutchins; T. It6; K. Izawa; N. K. Johnson; J. Just; Z. Kabata; S. J. Keeley; B. Kensley; P. Knight-Jones; K. F. Koopman; R. K. Kropp; J. D. Kudenov; D. B. Lellinger; R. B. Manning; J. C. Markham; J. W. Martin; W. N. Mathis; R. W. McDiarmid; P. A. McLaughlin; J. G. Mead; M. Milligan; C. Mitchell; T. Miura; R. F. Modlin; B. L. Monroe; H. Morino; K. M. Muzik; C. W. Myers; P. Myers; N. A. Neff, D. Nelson; W. A. Newman; S. L. Olson; J. M. Orensanz; K. C. Parks; D. C. Parris; E. C. Peters; R. H. Pine; D. Polhemus; C. W. Potter; H. D. Pratt; R. J. Raikow; S. Rainer; J. V. Remsen; K. Reutzler; J. W. Reynolds; K. Riemann-Zueneck; C. B. Robbins; C. E. da Rocha; G. Rodriguez; R. H. Rosenblatt; D. Russell; J. M. Savage; S. A. Schaefer; H.-K. Schminke; R. T. Schuh; M. Schwartz; N. J. Scott; M. E. Seidel; W. F. Smith-Vaniz; V. G. Springer; D. H. Staples; W. C. Starnes; G. C. Steyskal; J. H. Stock; R. E. Strauss; J.-O. Stromberg; M. Telford; A. Tsukagoshi; M. Tiirkay; J.-W. Wagele; T. R. Waller; L. Watling; M. J. Wetzel; A. B. Williams; G. Wilson; W. W. Wirth; T. Yamaguchi; H. C. Yeatman; G. R. Zug; V. A. Zullo; R. L. Zusi. INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (botany, zoology, and paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with a summary in an alternate language when appropriate. Submission of manuscripts. —Submit manuscripts to the Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History NHB Stop 108, Smithsonian Institution, Washington, D.C. 20560. Review.—One of the Society’s aims is to give its members an opportunity for prompt pub- lication of their shorter contributions. Manuscripts are reviewed in order of receipt by a board of Associate Editors and appropriate referees. 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Literature citations in the text should be in abbreviated style (author, date, page), except in botanical synonymies, with unabbreviated citations of journals and books in the Literature Cited sections. Direct quotations in the text must be accompanied by author, date, and pagination. The establishment of new taxa must conform with the requirements of the appropriate international codes of nomenclature. When appropriate, accounts of new taxa must cite a type specimen deposited in an institutional collection. Examples of journal and book citations: Eigenmann, C. H. 1915. The Cheirodontidae, a subfamily of minute characid fishes of South America.— Memoirs of the Carnegie Museum 7(1):1-99. Ridgely, R. S. 1976. A guide to the birds of Panama. Princeton, New Jersey, Princeton University Press, 354 pp. Olson, S. L. 1973. The fossil record of birds. Pp. 79-238 in D. Farner, J. King, and K. Parkes, eds., Avian biology, volume 8. Academic Press, New York. Figures and tables with their legends and headings should be self-explanatory, not requiring reference to the text. Indicate figure and table placement in pencil in the margin of the manu- script. Plan illustrations in proportions that will efficiently use space on the type bed of the Proceedings. Original illustrations should not exceed 15 x 24 inches. Figures requiring solid black backgrounds should be indicated as such when the manuscript is submitted, but should not be masked. CONTENTS The extinct sloth, Megalonyx, (Mammalia: Xenarthra), from the United States Mid-Atlantic continental shelf H. Gregory McDonald and Clayton E. Ray Systematics of the “Green-Throated Sunangels” (Aves: Trochilidae): valid taxa or hybrids? Gary R. Graves A new species of Cuora (Reptilia: Testudines: Emydidae) from the Ryukyu Islands Carl H. Ernst and Jeffrey E. Lovich A new ocellated frog (Centrolenidae) from western Colombia John D. Lynch On the validity of the Indo-Pacific cardinalfishes Apogon aureus (Lacepéde) and A. fleurieu (Lacepéde), with description of a related new species from the Red Sea John E. Randall, Thomas H. Fraser, and Ernest A. Lachner Living cassiduloids (Echinodermata: Echinoidea): a key and annotated list Rich Mooi A new species of halophilous water-strider, Mesovelia polhemusi, from Belize and a key and checklist of New World species of the genus (Heteroptera: Mesoveliidae) Paul J. Spangler Luteocarcinus sordidus, new genus and species, from mangrove swamps in peninsular Malaysia (Crustacea: Decapoda: Brachyura: Pilumnidae: Rhizopinae) Peter K. L. Ng On the status of Alpheus barbara Lockington (Crustacea: Caridea: Alpheidae) Mary K. Wicksten Pseudothelphusa galloi, a new species of freshwater crab (Crustacea: Brachyura: Pseudothel- phusidae) from southwestern Mexico - Fernando Alvarez and Jose Luis: Villalobos Range extension and host record for Dissodactylus ususfructus Griffith, 1987 (Crustacea: Brachyura: Pinnotheridae) Michel E. Hendrickx An eastern United States record for the western Indo-Pacific crab, Hemigrapsus sanguineus (Crustacea: Decapoda: Grapsidae) Austin B. Williams and John J. McDermott A redescription of Tiron antarcticus K. H. Barnard, 1932 (Crustacea: Amphipoda: Synopiidae) with an updated key to the species of Tiron Liljeborg, 1865 Krzysztof Jazdzewski Ensayara jumane, a new species from Belize, Caribbean Sea (Amphipoda, Lysianassidae) J. L. Barnard and James Darwin Thomas A range extension to the north for Macrochiridotea giambiagiae Torti and Bastida (Crustacea: Isopoda: Valvifera) Elizabeth Harrison-Nelson and Thomas E. Bowman Heteromysis mexicana, a new species from Campeche Bank, Gulf of Mexico (Crustacea: Mys- idacea) Elva Escobar-Briones and Luis A. Soto Redescription and new records of Trichodiaptomus coronatus (G. O. Sars) (Copepoda: Cal- anoida: Diaptomidae) from Brazil Janet W. Reid Achelia assimilis (Haswell, 1884) in the Heterozostera bed of Puerto Aldea, Coquimbo: first record from the northern Chilean coast (Pycnogonida: Ammotheidae) Sergio A. Gonzalez and Mario E. Edding Prototrygaeus jordanae, a new species of pycnogonid from Monterey Bay, California C. Allan Child Fabricinuda, a new genus of Fabriciinae (Polychaeta: Sabellidae) Kirk Fitzhugh Diplocardia kansensis, a new earthworm from Kansas, with redescriptions of D. riparia Smith and D. fuscula Gates (Annelida: Oligochaeta: Megascolecidae) Samuel W. James Redescriptions and synonymy of Nephtys imbricata Grube, 1857 (Polychaeta: Nephtyidae) Takashi Ohwada Antrorbis breweri, a new genus and species of hydrobiid cavesnail (Gastropoda) from Coosa River Basin, northeastern Alabama Robert Hershler and Fred G. Thompson A new isidid octocoral (Anthozoa: Gorgonacea) from New Caledonia, with descriptions of other new species from elsewhere in the Pacific Ocean Frederick M. Bayer A new species of Cladocarpus (Cnidaria: Hydroida: Plumulariidae) from the Straits of Florida Mary Ann Bogle The Fourth Annual Riser Lecture: The role of physiology and biochemistry in understanding animal phylogeny Charlotte P. Mangum Six new combinations in Baccharoides Moench and Cyanthillium Blume (Vernonieae: Aster- aceae) Harold Robinson international Commission on Zoological Nomenclature: Applications and Opinions Reviewers— 1989 100 103 106 108 110 7 120 ¥ 27 131 1404 151 Sy 161 179 187 197 205 229 235 248 254 256 - PROCEEDINGS e OF THE BIOLOGICAL SOCIETY OF ~ WASHINGTON VOLUME 103 NUMBER 2 28 JUNE 1990 ISSN 0006-324X THE BIOLOGICAL SOCIETY OF WASHINGTON 1990-1991 Officers President: Leslie W. Knapp Secretary. G. David Johnson President-elect: Storrs L. Olson Treasurer: Michael Vecchione Elected Council _Roger F. Cressey, Jr. Janet W. Reid Janet R. Gomon Wayne C. Starnes Robert Hershler Jeffery T. Williams Custodian of Publications: Austin B. Williams PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: George C. Steyskal Invertebrates: Stephen D. Cairns Frank D. Ferrari Plants: David B. Lellinger Rafael Lemaitre Insects: Wayne N. Mathis Vertebrates: G. David Johnson Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $15.00 ($20.00 to non-USA addresses) include subscription to the Proceedings of the Biological Society of Washington. Library subscriptions to the Proceedings are: $25.00 within the U.S.A., $30.00 elsewhere. The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued sporadically) are available. Correspondence dealing with membership and subscriptions should be sent to The Treasurer, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, National Museum of Natural History, Washington, D.C. 20560. THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 257-264 PORITES COLONENSIS, NEW SPECIES OF STONY CORAL (ANTHOZOA: SCLERACTINIA) OFF THE CARIBBEAN COAST OF PANAMA Vassil N. Zlatarski Abstract.—A new species of Porites (P. colonensis) from off the Caribbean coast of Panama is described. It is common and has very thin, undulated, foliaceous colonies; pali; and has dark brown or red tissues with white or green polyp centers. New species of shallow water corals are unusual, particularly in an area frequently visited by reef investigators. The new scler- actinian from Panama, described below, is common, and very easily distinguished un- derwater and in the laboratory by color, col- ony shape, and skeletal elements. Order Scleractinia Bourne, 1900 Suborder Fungiida Duncan, 1884 Superfamily Poritioidea Gray, 1842 Family Poritidae Gray, 1842 Genus Porites Link, 1807 not Porites Cuvier, 1798:678-679 (=Gal- axea Oken, 1815; Dendrophyllia Blain- ville, 1830; Mussa Oken, 1815 sensu Ve- ron and Pichon, 1982:141). (A proposal to suppress Porites Cuvier, 1798 is nec- essary in order to validate the priority of Link’s authorship.) Porites Link, 1807:162. Neoporites Duchassaing & Michelotti, 1866: 191. Cosmoporites Duchassaing & Michelotti, 1866:193. Type species. — Porites polymorphus Link, 1807 (by tautonomy) = Madrepora porites Pallas, 1766 (in part). This genus is cosmopolitan. Eocene-Re- cent. Abundant reef coral since Miocene. Porites colonensis, new species Figs. 1-20 Etymology. — From the name of the town of Colon, Panama. Diagnosis. — Porites with very thin, folia- ceous, undulated colonies. Lower surface with concentric holotheca. Corallites dis- tinct to poorly distinguishable. Axial struc- ture absent (empty fossa) or present (from weak columella to massive columella with a central tubercle). Septal plan bisymmet- rical, more or less clear; a dorsal directive septum, a ventral directive septum with two other septa fuse in triplet and four lateral pairs of septa arranged symmetrically in re- lation to the dorso-ventral axis. Five pali (one on the triplet, and one on each lateral pair), seldom six pali. Tissue color: ordi- narily polyps are very dark brown with small bright white centers; polyps may be dark red with green centers. Distinguishing char- acters: foliaceous colony shape, presence of pali, and the dark (brown or red) color of the polyp with contrasting white or green centers. Holotype. —USNM 82020 (Figs. 1, 2 and 7): Length 112 mm, width 93 mm, height 74 mm, thickness of the folia at peripheral edge 1-4 mm; collected 20 Aug 1987 by V. Zlatarski and H. Guzman. Paratypes. -USNM 82021-82054 (Figs. 258 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-2. Porites colonensis, holotype, USNM 82020: 1, Side view, colony growing on Mussa angulossa (Pallas, 1766) Oken, 1815; 2, Top view, both x0.77. VOLUME 103, NUMBER 2 NO NA \O Figs. 3-6. Porites colonensis, paratypes, colony morphology: 3, USNM 82022; 4, Corallite size variation, USNM 82024; 5, USNM 82023; 6, USNM 82021, all figures <0.77. 260 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 7-8. Porites colonensis, pali, variation of skeletal structure and corallite diameter: 7, Holotype, USNM 82020, x 2.24; 8, Paratype, USNM 82024, x3.2. VOLUME 103, NUMBER 2 ‘ ' “4 ‘es 4 F 4 — % ms . es ; j Sen oe - F “ <2 a ‘ ns ; ws ’ i ae : $e S 7 * j (14) aoe ‘A: % : Figs. 9-14. Porites colonensis, paratype, USNM 82028, variability within one colony: 9, USNM 82028D, in the upper part of the photo—small corallites with thin septa, with axial fossa or with weak columella; in the lower part of the photo—much larger corallites with thicker septa and with weak or strong columella, x11; 10, USNM 82028A, thin skeletal elements, not well distinguished corallites, presence of axial fossa or weak columella, x19; 11, USNM 82028C, moderate thick skeletal elements, well distinguished corallites, presence of axial fossa or weak columella, x19; 12, USNM 82028C, thick skeletal elements, well distinguished corallites, presence of axial fossa, weak or strong columella, x 19; 13-14, Vertical and oblique views of one columella, USNM 82028B, x 64. 262 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ga é , 7 ath Figs. 15-20. Porites colonensis, paratype, USNM 82027, variability within one colony—axial structure and ornamentation of septa and pali: 15-16, Vertical and oblique views of one axial fossa, USNM 82027B, x38; 17-18, Vertical and oblique views of a weak columella, USNM 82027B, x 38; 19, Oblique view of weak columella, columella, USNM 82027A, x45; 20, Massive columella with central tubercule, clear septal plan (down left—a dorsal directive septum, up right—a ventral directive septum) fused with two septa in a triplet, and four lateral pairs of septa arranged symmetrically in relation to the dorso-ventral axis, USNM 82027A, x45. VOLUME 103, NUMBER 2 3-6, 8-20), collected from 26 Jun 1987 to 23 Sep 1987, by V. Zlatarski and H. Guz- man. Type locality. — Off the southeast coast of the island of Largo Remo, east of Colon, Bahia Las Minas; depth 3 m. Description. —Colony with irregular fo- liaceous form (Figs. 1-6), generally attached by a small central area to coralla of other Scleractinia (Fig. 1) or growing on sponges. Maximal colony diameter ranges from 4 to 13 cm. Peripheral edge of colony rounded, generally undulated. Folias of colony thin (generally 1-4 mm, seldom 6 mm). Colonies sometimes form foliaceous layers (Figs. 1, 3, 6) of which only the uppermost is alive. Lower surface of corallum covered by dis- tinct rings of holotheca, concentric to the attachment of the colony. Calicular surface cerioid. Corallites vary from indistinguish- able to quite discrete, their diameter mea- sured from the middle of the wall varies from 0.6 to 1.3 mm. Skeletal elements not compact but porous, thus they do not have constant width and thickness, and therefore all measurements correspond only to the plane of the measurements. Corallites on concave areas of colony surface generally smaller than corallites on convex parts of the colony. In the first case calices not well distinguished, small (diameter 0.6—-0.7 mm), and skeletal elements thin, but in the convex parts calices are well distinguished, well sep- arated, larger (diameter 1.0—1.3) and wall and septa are thicker (Figs. 4, 8-12). Septal plan bisymmetrical, varying from unclear (Figs. 9, 10) to clear (Fig. 20). In clearly bisymmetrical plans dorsal directive sep- tum is independent. Opposite to it is the ventral directive septum, which is fused with 2 other septa in triplet. Four lateral pairs of septa arranged symmetrically in relation to dorso-ventral axis. Septa ornamented by more or less inclined to vertical angular granules, varying in shape, size, number, distribution, and orientation (Figs. 9—20). Generally 5 pali surround corallite center (Figs. 7, 8), located on the top of triplet and 263 on axial edge of each lateral pair. Sometimes there 1s a smaller 6th palus on axial edge of dorsal directive septum. The pali are ver- tical (Figs. 15-19) or inclined (Figs. 13, 14, 20); their granulation is variable in shape, number and distribution (Figs. 9-12). The axial structure (columella) is the most vari- able skeletal element, being present or ab- sent in neighboring corallites (Figs. 10-12). When it is absent, the 5 (Sometimes 6) pali are connected in a ring circumscribing the deep axial fossa (Figs. 8—center, 15, 16). When present, the columella may be pari- etal, formed by 4 or more continuations of the septa, not on the same level, toward the center of the calice (Figs. 17-19). Elsewhere, the columella may be considerable and even become massive, occupying all the space in- side the pali, and furnished with a central tubercule (Fig. 20). The wall is a synapticu- lotheca. The color of the living tissues is very dark brown with very bright white polyp centers or dark red with green centers. Ecology. — Found on tilted reef slopes, at depths of 3 to 27 m; usually in small cavities (with approximately diameter 20 cm), fixed on dead parts of other Scleractinia or on Porifera. Distribution. —Caribbean coast of Pana- ma, from Colon to Isla Grande and in San Blas area. Localities of collected specimens: Largo Remo Island, depth 3 m, USNM 82020-82029, 82032, 82033, 82039-82054; Payardi (in front of the Refinery, depth 4.5 m, USNM 82030, 82031, 82034; Naranjo Abajo Island, depth 7 m, USNM 82038; Palina (El Mamey), USNM 82035, depth 8 m, USNM 82036, depth 2.8 m, USNM 82037, depth 3 m. Also observed at the last locality at 13 m and 17 m depths. Observed also near Portobelo, and in Holandes, San Blas area (depth 3—27 m). Comparison and discussion. —P. colonen- sis differs from P. astreoides Lamarck, 1816 by a combination of three characters: the presence of pali, the foliaceous colony form, and the dark polyp color with white or green 264 centers. It is distinguished from all branch- ing Porites by its colony form and by the two colors of its living tissues. The poor knowledge of P. branneri Rathbun, 1888 in Caribbean waters makes it difficult to com- pare that species to other Porites. Nonethe- less, the material described here is well dis- tinguished by its colony shape, corallite and colony size, skeletal characters and tissue colors from the data existing on Brazilian P. branneri and the characters of its type specimens (USNM 10961, 10962). In the area where P. colonensis was found, P. astreoides and the branching Porites were also observed, but colonies having inter- mediate characters between P. colonensis and the other Porites were not found. This observation and the presence of the char- acter combination described previously are the reasons for describing it as a new species. Future studies will bring additional infor- mation about the polyps and the skeletal characteristics on different levels of biolog- ical organization, about the life history and the distribution of this new taxon, and will undoubtedly improve the taxonomy of the genus. It is clear that until now, no Porites has been described with these characters and that its representatives do not show inter- mediates with the representatives of the other living Porites species. The only similar fossil species to P. co- lonensis is P. trinitatis Vaughan, 1926 (in Vaughan & Hoffmeister, 1926), from the Miocene of Trinidad. The types of P. trin- itatis (holotype USNM M053674, and five paratypes USNM 68302) do not show suf- ficient details of the skeletal structures for a good comparison. Nevertheless, P. colo- nensis has thinner, undulated colony plates, and smaller corallites. (In P. trinitatis the diameter of the corallites is 1.25 mm-—1.8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mm, in average 1.6 mm). Future studies should be done on more and better pre- served fossil material. Acknowledgments This work was made possible by the sup- port of the Smithsonian Institution. I would like to thank especially Chip Clark for the photography, and Hector Guzman and Er- nesto Weil for the field assistance. The scan- ning electron photomicrographs were taken in S. E. M. laboratory of the National Mu- seum of Natural History, Smithsonian In- stitution by Brian E. Kahn. I thank George C. Steyskal for advice on zoological no- menclature and K. R. Moore for technical assistance. I am grateful to Stephen D. Cairns, Stephen C. Jameson, Thomas E. Bowman and William J. Sando for review- ing the manuscript. Literature Cited Rathbun, R. 1888. Annotated catalogue of the species of Porites and Synaraea in the United States National Museum, with a description of a new species of Porites.— Proceedings of United States National Museum 10:354—366, pls. 15-19. Vaughan, T. W., & J. E. Hoffmeister. 1926. Miocene corals from Trinidad.— Carnegie Institution of Washington, Publication No. 344, Papers of the Department of Marine Biology 23:107—134, pls. 1-7. Veron, J. E. N., & M. Pichon. 1982. Scleractinia of Eastern Australia. Part IV. Family Poritidae.— AIMS Monograph Series 5:1-159. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 265-278 SOLENOTHECA, NEW HYOLITHA (MOLLUSCA) FROM THE ORDOVICIAN OF NORTH AMERICA John M. Malinky Abstract.—The first well-preserved hyoliths from the Ordovician of North America are here described as Solenotheca bakerae n. gen., n. sp., family Hy- olithidae, order Hyolithida. This species seems to have been a rare component of soft-bottom, normal marine assemblages in Ordovician carbonate rocks, and thus far it appears to be endemic to North America. Poor preservation of the types of the North American Ordovician species Hyolithes baconi Whitfield, H. multicinctus Bradley, H. pinniformis Ruedemann, H. pumilus Ruedemann, H. rhine Ruedemann, and H. versaillensis Miller & Faber renders the generic identification of these species uncertain. H. baconi and H. multicinctus are included under Solenotheca with question; the other species cited above are retained tentatively in Hyolithes. Their names should only be used for the type specimens. The holotype of H.? dubia Miller & Faber is a triobite spine, and this species is here removed from the Hyolitha. Hyolitha are locally abundant in Cam- brian strata of marine origin, but are rela- tively rare in Ordovician and younger Pa- leozoic rocks. Cambrian hyoliths in the Soviet Union and China are currently the focus of considerable interest because of their utility in biostratigraphic subdivision of the Lower Cambrian (see Missarzhevsky 1969 and Qian et al. 1985 for detailed list of ref- erences). In contrast, except for several re- cent works (Marek 1967, 1983a, b; Malinky et al. 1987; Houbrick et al. 1988), post- Cambrian Paleozoic hyoliths have been lit- tle studied. This report constitutes the first study of North American Ordovician Hy- olitha in which modern taxonomic methods are used. The first hyolith described, Hyolithes acutus Eichwald (1840), was from the Or- dovician of the eastern European Baltic re- gion. Eichwald (1860) later described sev- eral other Ordovician species from that area. The earliest study of the Hyolitha to include detailed consideration of stratigraphic dis- tribution in addition to taxonomy was that of Barrande (1867). Barrande’s work was the first monograph ever devoted to hy- oliths and other conical problematica, and he included Ordovician as well as younger hyolith species from central Europe. Other Ordovician species from central Europe and Scandinavia were described in later works by Novak (1891), Holm (1893), Reed (1909), and Zazvorka (1928). Thoral (1935) de- scribed a fauna of Tremadocian and Ar- enigian hyolith species from France (Marek 1983b), which included the first known or- thothecid hyolith with preserved casts of the intestine and other internal structures (Houbrick et al. 1988). Thoral’s specimens served in part as the basis for removing the Hyolitha from the Mollusca and assigning them to the extinct phylum Hyolitha by Runnegar et al. (1975). Marek & Yochelson (1976) continued to regard them as mol- luscs. Until recently, species named by those early workers represented nearly one-third of all known Ordovician species. The other Ordovician species were discovered at widely scattered localities throughout the world, and were usually named incidentally 266 in studies devoted to other organisms or to entire faunas. By the mid-20th century, a total of one hundred eleven Ordovician hy- olith species had been named (Sinclair 1946). Compared to Europe, hyoliths in the Or- dovician of North America seem to be quite rare. In nearly a century of study, only nine North American species had been named (Sinclair 1946), and only one occurrence of a previously known European species had been reported (Matthew 1895). None of these species have ever been restudied, and with rare exceptions their names have not been used in the literature for any material except the types. Little is known of their stratigraphic or biogeographic distribution, and nothing of their phylogenetic relation- ships. Marek (1963, 1966, 1967, 1983a, b) brought a new perspective to the study of the Hyolitha in general and to Ordovician hyoliths in particular. He reevaluated many of the European species named by Barrande and other early workers, and supplemented the original descriptions with new material from Europe and North Africa. The local abundance of hyoliths in Ordovician car- bonate rocks of Baltoscandia was noted by Jaanusson (1984) and Jaanusson & Mutvei (1982) but they did not describe any taxa. With one recent exception (Malinky 1987), no attempt has been made to reevaluate any hyoliths from the Ordovician of North America until now. Restudy of Ordovician hyoliths from North America indicates that specimens from the Middle Ordovician of the Upper Mississippi Valley region represent Solen- otheca bakerae n. gen. n. sp., in the family Hyolithidae, order Hyolithida. Morphology of the types of the Ordovician species Hy- olithes baconi Whitfield (1878) and H. mul- ticinctus Bradley (1930) suggests affinity to Solenotheca, to which they are here reas- signed with question. In contrast, poor pres- ervation of the types of H. pinniformis Rue- demann (1912), H. pumilus Ruedemann (1926), H. rhine Ruedemann (1901), and H. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON versaillensis Miller & Faber (1894) renders the original descriptions of these species in- adequate and their generic identifications uncertain; they are retained under Hyolithes with question. The Ordovician hyolith species Hyolithes vanuexmi Walcott (1884) from the Eureka district in Nevada was reassigned to Chel- sonella Malinky (1987), order Orthotheci- da, class Hyolitha. The type or types of the Ordovician species H. crowelli Roy (1941) and H. parviusculus (Hall 1862) cannot be located at present. These names should not be used for any other material until the types are located or until well preserved topotypes become available for study. Type specimens of H.? dubius Miller & Faber (1894) consist of trilobite spines, and that species is here reassigned to the Arthropoda. The holotype of H. miseneri (Foerste, 1917) is presently under study. Distribution and Paleoecology Marek’s (1976) analyses of the geographic and stratigraphic distribution of fifteen Or- dovician hyolith genera was the first and only study of its kind. He recognized two distinct hyolith assemblages in the Ordo- vician; one occurs in Baltoscandia whereas the other is common in central Europe and North Africa with rare representatives in Australia and South America. None of the genera in these assemblages have been iden- tified unequivocally in North America. However, poor preservation complicates the recognition of any European hyolith taxa in North America. A specimen questionably assigned to the Ordovician genus Elegan- tilites Marek (1967) from Czechoslovakia was reported by Harrison & Harrison (1975) from the Silurian Brassfield Formation in Ohio. A species of the Ordovician genus Joachimilites Marek (1967), previously known only in Czechoslovakia has been identified tentatively from the Middle Cam- brian Maryville Limestone in Alabama (Malinky 1988). Nonetheless, preliminary VOLUME 103, NUMBER 2 observations support the notion that North American hyoliths comprise a third distinct Ordovician assemblage characterized by Solenotheca. Published reports and illustra- tions of hyoliths from outside North Amer- ica seem to indicate that Solenotheca is endemic to North America. This genus has not been identified yet in any of the well-preserved and relatively abundant Or- dovician hyolith assemblages of central Eu- rope or North Africa, or from older hyolith- bearing strata in the Soviet Union and China. The stratigraphic distribution of Soleno- theca is poorly known. The types of this genus were collected more than 75 years ago from the Upper Mississippi Valley region by an unknown worker, and geographic and stratigraphic details about those occur- rences are not available. The age of Soleno- theca cannot be determined more precisely than Middle Ordovician (Webers & Austin 1972) because specific localities and asso- ciated faunas are not known with certainty. Assignment of Hyolithes baconi Whitfield and H. multicinctus Bradley to Solenotheca does not extend the stratigraphic range of the genus because both of those species were also discovered in the Middle Ordovician. In addition, their inclusion under Soleno- theca does not extend the geographic range significantly for they also were discovered in the Mississippi Valley region. Solenotheca has only been reported thus far from carbonate rocks. Middle Ordovi- clan carbonate strata of the Upper Missis- sippi Valley represent shallow, normal ma- rine environments inhabited by soft-bottom benthic assemblages (Webers 1972). These assemblages are dominated by brachiopods, bryozoans and molluscs; hyoliths seem to have a sporadic distribution in these rocks and are rare at any given locality. Direct examination of type specimens and pub- lished accounts in the literature show that most other Ordovician hyoliths in North America occur in carbonates, although a few specimens are known from shale. This may 267 be largely a bias of collection and preser- vation. Hyoliths in shale and other clastic rocks tend to preserve less well than lime- stone specimens and may have been over- looked or simply not collected because of preservation. Fisher (1962) noted that hyoliths occur in a wide range of facies in the marine en- vironment except for hypersaline and reef facies. His observations seem to be based primarily upon Cambrian occurrences. Ma- terial described herein and undescribed specimens in collections of the Smithsonian Institution suggest that Ordovician hyoliths were widely distributed geographically and in a variety of marine facies, but were rare overall within the marine environment. To date, assemblages of hundreds of individ- uals on bedding surfaces in sandstone or more rarely in limestone have not been ob- served in the Ordovician. Such occurrences are locally known in Cambrian sandstones of the Upper Mississippi Valley (Marek & Yochelson 1976) and in Cambrian lime- stones of west-central Montana. The rela- tive scarcity of hyoliths in Ordovician and younger Paleozoic rocks may be related to their apparent mechanical inefficiency in comparison to gastropods or other more mobile benthos (Yochelson 1984). Mode of life of the Hyolitha has been a matter of controversy until recently. Fisher (1962) suggested several possibilities, in- cluding benthic with venter down, benthic with the apex embedded in sediment, and nektic and planktic. Marek & Galle (1976) used the presence of an encrusting tabulate coral on the dorsum only of several Devo- nian hyoliths from central Europe to suggest that a benthic mode of life with venter rest- ing upon the seafloor seemed most likely. Several Late Ordovician specimens from southwestern Ohio (USNM 50107A, USNM 50102A) lend support to their in- terpretation. These specimens are also en- crusted by an epibiont which, in this in- stance is a bryozoan. Like the Devonian tabulate coral, the bryozoan also is confined 268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. A, E, Hyolithes? multicinctus Bradley: A, Paratype FMNH 5916B showing dorsum, x3. E, Holotype FMNH 5916A showing dorsum, <1.8, Missouri. B, G, Solenotheca? baconi (Whitfield): B, Paralectotype UCMP 34360A showing dorsum. G, Lectotype UCMP 35219 showing venter, Wisconsin. Both x1.1. C, D, External VOLUME 103, NUMBER 2 to the dorsum (Fig. 1C, D, F, H). Had the hyoliths lived with the dorsum down or with the apex imbedded in the sediment, both dorsum and venter would be covered with epibionts. Repository designations used in this manuscript are: National Museum of Nat- ural History, USNM; Field Museum of Nat- ural History, FMNH; New York State Mu- seum, NYSM; and Museum of Paleontology, University of California, Berkeley, UCMP. Systematic Paleontology Phylum Mollusca Class Hyolitha Marek Order Hyolithida Matthew Family Hyolithidae Nicholson Solenotheca, new genus Type species. — Solenotheca bakerae, new species. Included species.—Solenotheca bakerae. Hyolithes baconi Whitfield (1878), and H. multicinctus Bradley (1930) are possible representatives of this genus. Diagnosis.—Hyolithid which has _ nar- rowly rounded dorsum with concavo-con- vex slopes; ligula short and aperture or- thogonal (see Marek 1983a). Remarks. —Knowledge of Solenotheca is derived from six well preserved specimens from the Upper Mississippi Valley region which formerly were listed as hypotypes of Hyolithes baconi Whitfield (1878) by an un- known worker. Thirty other specimens from widely scattered localities in the Mississippi Valley region are possible representatives of this genus, but incomplete preservation pre- cludes confident identification of those specimens. Solenotheca differs from all other genera currently included under the Hyolithidae — 269 (see Malinky 1988 for listing of genera in that family) by the presence of concavo- convex slopes on the dorsum. The only oth- er hyoliths to possess a similar feature are members of the Family Similothecidae Malinky (1988) from the Lower Cambrian of Newfoundland. However, the dorsum on similothecid hyoliths terminates in a pro- truding keel on the longitudinal ridge. So- lenotheca \acks a keel. Furthermore, So- lenotheca has a cancellate pattern of ornament created by closely spaced trans- verse lirae and widely spaced longitudinal lirae; on the similothecid hyoliths the trans- verse Ornament as well as longitudinal or- nament is widely spaced. The features which characterize Solenotheca have not been re- ported on any Soviet or Chinese hyolith (Syssoiev 1962, 1968, 1974; Val’kov 1975; Qian 1977, 1978). Stratigraphic range. —Middle Ordovi- cian. Etymology.—The name of the genus is derived from the Greek solen, meaning fur- row, in reference to the concave slopes. Solenotheca bakerae, new species Fig. 2E-G Diagnosis. —Solenotheca with widely spaced longitudinal lirae superimposed upon closely spaced transverse lirae and trans- verse threads which create a cancellate pat- tern on shell and internal mold. Description. — Venter slightly inflated with narrowly rounded lateral margins. Dorsum low and narrowly rounded in middle. Ad- jacent slopes convex next to lateral margins, but are markedly concave elsewhere. Trans- verse outline of shell subtriangular; sides of triangle concave. Ventral ligula short and straight, broadly rounded anterior edge. mold and dorsum of shell, respectively, of unidentifiable hyolith USNM 50107A showing bryozoan encrustation, x2, Ohio. F, H, Latex cast and external mold, respectively, of unidentifiable hyolith USNM 50102A showing bryozoan encrustation; F, x4, H, x 4.5, Ohio. I, External mold of unidentifiable hyolith USNM 50107B showing dorsum, x2, Ohio. J, External mold of unidentifiable hyolith USNM 6833 showing venter(?), x2, Ohio. 270 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. A-C, Hyolithes? rhine Ruedemann: venter, dorsum and left lateral view, respectively, of holotype NYSM 3514, original shell, x2, New York. D, H, Hyolithes? pumilus Ruedemann: D, paralectotype NYSM 9425B, dorsal(?) external mold, x 5.5, New York. H, lectotype NYSM 9425A, dorsal(?) external mold, x 1.6, New York. E-G, Solenotheca bakerae n. gen., n. sp: holotype USNM 50086A, right lateral, dorsal and ventral VOLUME 103, NUMBER 2 Sides of ligula dip steeply away from ante- rior edge. Apertural rim orthogonal with shallow sinus along aperture on each lateral margin but not on dorsal margin. Apertural rim not flared. Apical end appears straight. Shell covered with alternation of trans- verse threads and transverse lirae. Lirae parallel outline of aperture; on dorsum lirae transverse, and curve on lateral margins to follow apertural sinuses. On venter lirae fol- low edge of ligula. Fine, longitudinal lirae superimposed upon transverse ornament create cancellate pattern on shell. Internal mold also with cancellate pattern of orna- ment defined by transverse threads and lirae with longitudinal lirae superimposed upon them. Operculum unknown. Remarks. —The holotype (Fig. 2E-G) is a steinkern covered with scattered frag- ments of shell; it measures 25.5 mm in length and has an apertural width and height of 9.5 mm and 5.0 mm respectively. Matrix adheres to the apertural end but enough de- tail is exposed so that the apertural rim can be seen. A small portion of the apical end is broken. Several other specimens are suf- ficiently well preserved to serve as para- types. The types of this species were origi- nally designated as hypotypes of Hyolithes baconi Whitfield (1878) by an unknown worker, probably because of a general re- semblance in conch form between them. However, the types of H. baconi lack all details of the aperture and ligula, as well as ornament on the shell. No further compar- ison between the two species is possible. Material.—All specimens reposited at USNM. Holotype 50086A and paratypes under 50086B, 50086C, 25269 and 2 para- types under 25270. Specimens questionably referred to Solenotheca species indetermi- nate, are catalogued as follows: 11 under 15764A and 2 under 15764B; 11 under — 271 50088; | under 25272, 4 under 50087A and 1 under 50087B. Occurrence. —Geographic and strati- graphic information for all specimens is scant. Knowledge of occurrences is derived entirely from labels associated with the specimens. The holotype and paratypes 50086B and 50086C are from ‘“‘Minne- apolis, Minnesota.” Paratype 25269 is from “5 m. n. Monroe, Green Co., Wis.” This presumably means 5 miles north of Mon- roe. Paratypes 25270 are from “Black River (Platteville), Belleville, Wisconsin.”’ The 11 specimens catalogued as 15764A, the 2 un- der 15764B and those under 50088 are all from “Beloit, Wis.’’; specimen 25272 is from “Platteville, Wisconsin”; specimens 50087A and 50087B are in a container labelled “Cannon Falls, Minnesota’ but labels en- closed with the specimens say “chiefly from beneath sponge bed Platteville-Upper Dix- on, Illinois.”” Labels with those specimens indicate that they were collected by E. O. Ulrich and R. S. Bassler in 1906, although there is no indication as to which locality is correct (Fig. 3). Webers & Austin (1972) recognized five Middle Ordovician formations which en- compass several hundred meters of carbon- ate rock in the Upper Mississippi valley re- gion. The Platteville, Decorah and Galena formations contain layers of limestone or dolostone as well as other lithologies. In the absence of any detailed data on localities or associated faunas, confident assignment of any of the specimens cited above to for- mation is impossible at this time. The age of strata in these areas is Middle Ordovician (Webers & Austin 1972). Stratigraphic range.—Middle Ordovi- cian. Etymology.—The species is named for Cathy Baker, Simpson College. views, respectively, original shell, x 2.8, Minnesota. I, Slab under USNM 78431A showing venters of uniden- tifiable Ordovician hyoliths, x2, Missouri. J, Unidentifiable Ordovician hyolith, USNM 50087B, venter, x 2, Ohio. K, Hyolithes? pinniformis Ruedemann. Holotype(?) NYSM 9423, venter(?), x 5.3, New York. Die MINNESOTA MINNESOTA =| Ordovician Outcrop Belt Minneapolis WISCONSIN e , = Minneapolis Cailnon Falls? WISCONSIN Ls) CO) @ Cannon Falls Fig. 3. Generalized locality map for holotype and paratypes of Solenotheca bakerae n. gen., n. sp. Class Hyolitha incertae sedis Solenotheca? baconi (Whitfield, 1878) Fig. 1B, G Hyolithes baconi Whitfield, 1878:77; 1882: 225, pl. 6, figs. 9-11.—Sinclair, 1946:74. Description. — Venter flat toward middle but becomes slightly inflated toward nar- rowly rounded lateral margins. Dorsum low and narrowly rounded in center; adjacent slopes appear nearly flat to slightly concave. Ligula appears long and straight. Apical sin- gle small and apical end straight. Complete dorsum, aperture, shell and operculum un- known. Remarks. —Type material of this species consists of dolomitized steinkerns from which all details except the proportions of the conch have been obscured by diagenesis. All specimens are embedded in matrix, from which extraction intact would probably be impossible. Specimen UCMP 32519 (Fig. 1G) is here designated the lectotype; it mea- sures 34.0 mm in length and is 11.0 mm wide near the apertural end. The paralec- totypes (Fig. 1B) are comparable to the lec- totype in size. This species was named and described but not illustrated by Whitfield (1878). His initial description was repeated nearly ver- batim slightly later (Whitfield 1882) and was accompanied by drawings (pl. 6, figs. 9-11). The drawings show this species to have two longitudinal sulci on the dorsum and trans- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON verse lirae on both dorsum and venter. The drawings also depict a short ligula with a broad anterior edge, and a low dorsum. No specimens in the type lot match these illus- trations; there is no evidence for the sulci or lirae shown. The drawings are apparently reconstructions of how Whitfield thought complete, well preserved specimens of this species should appear. The gently concave dorsal slopes suggest affinity to Solenotheca, to which this species is referred with ques- tion. Material. —Lectotype UCMP 32519 with paralectotypes on specimens under UCMP 34360-A and UCMP 34360-B. Occurrence. — According to Whitfield (1878:77), these specimens were discovered ‘in the hard bluish layers of the Trenton group, below Carpenter’s quarry, near Be- loit, Wisconsin.’ Beloit is located in south- ern Rock County near the border of Illinois and Wisconsin. The age of the species is Middle Ordovician. Solenotheca? multicincta (Bradley, 1930) Fig. 1A, E Hyolithes multicinctus Bradley, 1930:240, pl. 25, fig. 13.—Sinclair, 1946:78. Description.—Conch with small apical angle and apical end appears straight. Dor- sum low and narrowly rounded; slopes on dorsum appear slightly concave. Lateral margins narrowly rounded and dorsal aper- tural rim appears orthogonal. Shell covered with closely spaced transverse lirae; lirae curve slightly in middle of dorsum to create shallow sinus but are transverse elsewhere. Dorsum covered with fine, closely spaced longitudinal lirae; on lateral margins lirae also curved to create shallow sinus. Venter, complete apertural end and operculum un- known. Remarks. —Knowledge of this species is derived from the holotype (Fig. 1E) and a specimen included in this species as a para- type (Fig. 1A). The holotype measures 26.0 mm in length, and has an apertural width VOLUME 103, NUMBER 2 of 9.6 mm; the paratype is comparable in size. Both specimens retain the original shell, but each is embedded partly in matrix so that only the dorsum is exposed. Any at- tempt to extract these specimens from the matrix would probably destroy them. Bradley (1930) noted comparison be- tween this species and H. baconi Whitfield (1878), but he seems to have compared his specimens to Whitfield’s illustrations rather than to Whitfield’s specimens. No compar- ison between the types of these species is possible because of preservation. Bradley (1930) also reported minor differences of ornament between H. multicinctus and H. muiseneri (Foerste, 1917) as a basis for sep- arating these species. Foerste (1917) stated that H. miseneri has a cancellate pattern on the dorsum formed by the intersection of transverse and longitudinal lirae, whereas Bradley (1930) stated that all lirae on H. multicinctus are transverse. Bradley (1930) was mistaken on that point, for a reexam- ination of the types of H. multicinctus dem- onstrates that fine longitudinal lirae occur on the dorsum of those specimens, suggest- ing that both forms may belong to the same species. Concave slopes on the dorsum of H. multicinctus suggest placement under Solenotheca, to which this species is now assigned with question. Assignment to ge- nus for H. miseneri (Foerste, 1917) awaits further study of that specimen. Material. —Holotype FMNH 5916A and paratype FMNH 5916B. Occurrence.—Bradley (1930:241) cited the occurrence as ““Kimmswick limestone near Glen Park, Missouri.” The age of the species is Middle Ordovician. Hyolithes? pinniformis Ruedemann, 1912 Fig. 2K Hyolithes pinniformis Ruedemann, 1912: 111, pl. 7, figs. 12, 13.—Sinclair, 1946: 79. Description.—Conch with subtriangular outline; apical angle appears small and api- 273 cal end appears straight. The shell, and all other details of conch, and operculum, un- known. Remarks. —The holotype (Fig. 2K) is the only known specimen of this species. It con- sists of a subtriangular impression on a bed- ding surface in black shale. That specimen measures 11.3 mm in length, and has a max- imum width near the apertural end of 3.2 mm. Preservation of this specimen is ex- tremely poor; even the most elementary morphologic features, such as the ligula or any distinguishing features of dorsum and venter are lacking. Ruedemann’s (1912) drawings of this species (pl. 7, figs. 11, 12) depict a shell with a complete aperture and lirae on the exterior. They bear no resem- blance to the specimen described and illus- trated herein. Perhaps the drawings are re- constructions of how Ruedemann thought complete specimens of this species should appear, or perhaps the illustrated specimen is lost. This species cannot be properly di- agnosed using only the specimen described herein. Material. —Holotype(?) NYSM 9423. Occurrence. —Ruedemann (1912:111) cited the occurrence as: “‘Canajaharie shale at Canajaharie, N. Y.’’ Canajaharie is lo- cated in Montgomery County in central New York. The age of the species is Middle Or- dovician. Hyolithes? pumilus Ruedemann, 1926 Fig. 2D, H Hyolithes pumilus Ruedemann, 1926:77, pl. 10, figs. 5, 6.—Sinclair, 1946:79. Description. —Dorsum inflated and nar- rowly rounded in center; lateral margins narrowly rounded. Apical angle small and apical end appears straight. Complete aper- tural end, shell, venter and operculum un- known. Remarks. — This species is represented by two poorly preserved external molds on a slab of dark gray shale that also contains ostracods and a “‘syntype”’ of Rafinesquina 274 alternata centristriata Ruedemann. Speci- men NYSM 9425A (Fig. 2H) is here des- ignated the lectotype; it measures 9.3 mm in length and has an apertural width of 4.0 mm. The paralectotype (NYSM 9425B) (Fig. 2D) is similar in size and preservation to the lectotype. Both specimens are smooth and featureless, and neither retains a shell or an operculum. Ruedemann (1926, pl. 77, figs. 5, 6) il- lustrated specimens of this species that he refers to as the holotype (fig. 5) and paratype (fig. 6). In one drawing, the holotype pos- sesses a Short ligula which is broadly round- ed at the anterior edge, and it appears to have a longitudinal sulcus near the middle of the dorsum. Another drawing depicts a paratype that is less well preserved and lacks the ligula and sulcus. None of the existing specimens match Ruedemann’s illustra- tions. Either his drawings are reconstruc- tions of how he thought well preserved spec- imens of this species should appear, or the illustrated specimens are lost. Material. —Lectotype NYSM 9425A and paralectotype NYSM 9425B. Occurrence. —Ruedemann (1926:78) re- ported that the slab of shale containing both specimens was discovered in “‘zone I of the lower Lorraine (Whetstone Gulf) shale, in the Whetstone and Lorraine gulfs.”” The age of the species is Late Ordovician. Hyolithes rhine Ruedemann, 1901 Fig. 2A—C Hyolithes rhine Ruedemann, 1901:36, pl. 2, figs. 12-15.—Sinclair, 1946:79. Description. — Venter slightly inflated with narrowly rounded lateral margins. Dorsum inflated and narrowly rounded in center; slopes on dorsum slightly inflated. Apical end appears straight, and apical angle small. Faint longitudinal sulcus near each edge of the dorsum; sulcus extends along entire length of dorsum but becomes shallow near apical end. Shell covered with faint trans- verse lirae on dorsum. Lirae curve slightly in middle of dorsum to form shallow sinus; PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON apertural rim may be orthogonal. Internal mold of venter smooth. Complete apertural end and operculum unknown. Remarks.—The holotype (Fig. 2A—C) is the only known specimen of this species. It measures 21.0 mm in length, and has an apertural width and height of 10.2 mm and 6.2 mm respectively. The shell is preserved locally on the dorsum, but is absent on the venter. The holotype was broken out of limestone, and an impression of the outer surface of the shell remains in the matrix as a counterpart of the holotype. The counter- part is an impression of the dorsum rather than the venter as Ruedemann (1901) be- lieved. The dorsal lirae of H. rhine resemble those of H. multicinctus Bradley (1930), but the lirae on H. rhine are less conspicuous. Rue- demann (1901) compared this species to drawings of H. baconi Whitfield and H. multicinctus Bradley, but detailed compar- ison between these species is precluded by the absence of important taxonomic fea- tures on each. Hyolithes rhine possesses longitudinal sulci on the dorsum, a feature shared with only three other hyolith species: Hyolithes acutus Eichwald (1840), the type species of that genus, from the Ordovician of Estonia, H. crebescens Resser & Endo (1937) from the Middle Cambrian of China, and Hal- lotheca aclis (Hall, 1876) from the Devo- nian of New York. Nonetheless, various other features of the conch of each form suggest that each form probably represents a different species. Material.— Holotype NYSM 3514. Occurrence. —Ruedemann (1901) did not cite a specific occurrence for this species, but he named it in a study of the fauna of the Trenton conglomerate at Rysedorph Hill, Rensselaer County, New York. The introduction of the work includes general- ized information about a number of local- ities on Rysedorph Hill, and several out- crops are marked with arrows in a photograph of the hill. The specific outcrop which yielded the type of this species is not VOLUME 103, NUMBER 2 275 Fig. 4. A, B, D, Hyolithes? versaillensis Miller & Faber: dorsal, ventral and right lateral views, respectively, of lectotype FMNH 8876A, x17, Indiana. C, specimen FMNH 8881 formerly assigned to Hyolithes? dubia Miller & Faber, lateral view, x 12, Indiana. indicated. The age of the species is Middle Ordovician. Hyolithes? versaillensis Miller & Faber, 1894 Fig. 4A, B, D Hyolithes versaillensis Miller & Faber, 1894: 15; pl. 8, figs. 20-22.— Cummings, 1908: 965, pl. 42, figs. 4, 4a.—Sinclair, 1946: 81. Description.— Venter flat to slightly in- flated, with narrowly rounded lateral mar- gins. Dorsum inflated and broadly rounded, and transverse shape subtriangular. Apical end appears straight, and apical angle small. Several widely spaced ribs on internal mold of dorsum; internal mold of venter smooth. Complete aperture, shell, and operculum unknown. Remarks. —Type material of this species consists of eight specimens; specimen FMNH 8876-A (Fig. 4A, B, D) is here des- ignated the lectotype. That specimen mea- sures 2.9 mm in length, and has an apertural width and height of 2.2 mm and 1.3 mm respectively. All paralectotypes are com- parable to the lectotype in size. Every spec- imen of this species is a partially weathered limonitized steinkern. No specimen is com- plete and none is operculate. Miller & Faber (1894) thought that these specimens were casts rather than internal molds, and based their species diagnosis on the supposed “smoothness” of the shell. They also noted that “there are not many characters to ascribe to this species” (p. 155) and as a result, their description is gener- alized and lacks detail. The diagnosis of this species and the illustrations (pl. 8, figs. 20- 22) are inadequate to characterize this species because few features are preserved. Material. —Lectotype FMNH 8876A and seven paralectotypes under 8876B. Occurrence. — Miller & Faber (1894:155) reported that this species “‘is quite common in the upper part of the Hudson River Group, at Versailles, Indiana, associated with Palaeoconcha faberi, Cyclora pulcella and other small fossils.’’ Versailles is locat- ed in Ripley County in southeastern In- diana. The age of the Hudson River group is Late Ordovician. Class Trilobita incerate sedis Genus and species indeterminate Fig. 4C Hyolithes? dubius Miller & Faber, 1894:155, pl. 8, fig. 23.—Cummings, 1908:964, pl. 42, fig. 3.—Sinclair, 1946:75. Description. —“‘Conch”’ tubular and api- cal angle small. Taper of conch so slight that both ends appear of equal diameter. Shell 276 and internal mold smooth; both ends un- known. Remarks.—The holotype (Fig. 4C) and only specimen of this species measures 4.2 mm in length with a diameter of 1.0 mm. The original skeletal material is largely in- tact, although at the narrower end a small section of the internal mold is exposed. The specimen is broken at both ends, and no other features than those cited above are preserved. Miller & Faber (1894) did not recognize that the original skeletal material is pre- served on this specimen, for they cited the specimen as being a “‘cast.”” They were ap- parently uncertain about the affinity of this species, and assigned it to Hyolithes with question. They suspected affinity to calyme- nid trilobites which occur in the same strata as this species. However, because all the trilobite fossils of which they were aware were intact, they ruled out the possibility that the type of H.? dubius might be a frag- ment of a trilobite spine. They noted that this species is “like Hyolithes versaillensis, with which it is associated, and it would seem, therefore, impossible that it (H.? du- bius) should represent the broken spines of trilobites” (p. 156). The two species are quite dissimilar in terms of both morphology and preservation. H.? dubius lacks any evidence for hyolith affinity such as a shell with trans- verse growth lirae or an operculum. It is without question the spine of a trilobite. Material. —Holotype FMNH 8881. Occurrence. — Miller & Faber (1894:156) cited the occurrence as “‘the upper part of the Hudson River Group, at Versailles, In- diana, associated with Palaeoconcha faberi, Cyclora pulcella, Hyolithes versaillensis, and other small fossils.”” The age is Late Ordo- vician. Acknowledgments I wish to thank the National Museum of Natural History for support of this project in the form of a Postdoctoral Fellowship. I PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON wish to thank R. E. Grant, J. Pojeta, E. L. Yochelson, R. H. Mapes and an anonymous reviewer for their helpful comments on this manuscript at various stages of completion. I am grateful to J. Thompson for her assis- tance in arranging loans of specimens for this study. I would also like to thank the curatorial staffs at the Field Museum of Nat- ural History, Museum of Paleontology, University of California, Berkeley, and the New York State Museum for their generous loan of specimens. Literature Cited Barrande, J. 1867. System silurien du centre de la Boheme. Ordres des Pteropodes, 3. Published by the author, Prague and Paris, 179 pp. Bradley, J.H. 1930. Faunas of the Kimmswick Lime- stone of Illinois and Missouri.— Walker Mu- seum Contributions 2:465-479. Cummings, E. R. 1908. The stratigraphy and pa- leontology of the Cincinnati series of Indiana. — Indiana Geological Survey Annual Report 32: 605-1189. Eichwald, E. 1840. Ueber das silurische Schichten- system in Estland. Zeitschrift fiir Natir- und Heilkunde, 1 and 2, St. Petersburg, 210 pp. 1860. Lethaea rossica ou paleontologie de la Russie. E. Schweizerbart, Stuttgart, 1657 pp. Fisher, D. W. 1962. Small conoidal shells of uncer- tain affinities. Pp. W98—W143 in R. C. Moore, ed., Treatise on invertebrate paleontology. Part W. Miscellanea. Geological Society of America and University of Kansas Press, New York and Lawrence, Kansas. Foerste, A. G. 1917. Notes on the Richmond and related fossils.—Journal of the Cincinnati So- ciety of Natural History 22:42—55. Hall, J. 1862. Contributions to paleontology; com- prising descriptions of new species of fossils, from the Upper Helderburg, Hamilton and Che- mung Groups.—New York State Cabinet of Natural History 15:29-198. . 1876. Illustrations of Devonian fossils: Gas- teropoda, Pteropoda and Cephalopoda (plates only).—New York State Geological Survey, 74 pl. Harrison, W. B., & L. K. Harrison. 1975. A Maquo- keta-like molluscan community in the Brassfield Formation (Early Silurian) of Adams County, Ohio.— Bulletins of American Paleontology 67: 193-234. Holm, G. 1893. Sveriges kambrisk-siluriska Hyolith- VOLUME 103, NUMBER 2 idae och Conulariidae. Sveriges Geologiska Un- dersorning, C, 112, Stockholm, 172 pp. Houbrick, R.S., W. Sturmer, & E. L. Yochelson. 1988. Rare Mollusca from the lower Devonian Huns- ruck Slate of southern Germany.—Lethaia 21: 395-402. Jaanusson, V. 1984. Ordovician benthic macrofaunal associations. Pp. 127-139 in D. L. Bruton, ed., Aspects of the Ordovician System. —., & H. Mutvei. 1982. Ordovician of Oland. IV International Symposium, Ordovician System. Guide to Excursions, 3. Section of Palaeozool- ogy. Swedish Museum of Natural History, 23 pp. Malinky, J. 1987. Taxonomic revision of lower and middle Paleozoic Orthothecida (Hyolitha) from North America and China.—Journal of Paleon- tology 61:942-959. 1988. Early Paleozoic Hyolitha from North America: Reexamination of Walcott’s and Res- ser’s type specimens. — Journal of Paleontology 62:218—233. , R. M. Linsley, & E. L. Yochelson. 1987. Taxonomic revision of Hyolitha from the Mid- dle Paleozoic of North America.—Journal of Paleontology 61:1173-1186. Marek, L. 1963. New knowledge on the morphology of Hyolithes.—Sbornik Geologickych Ved Pa- leontology 1:53-72. 1966. New hyolithid genera from the Or- dovician of Bohemia.—Casopis Narodniho Musea 135:89-92. 1967. The class Hyolitha in the Caradoc of Bohemia.— Sbornik Geologickych Ved Pa- laeontology 9:51-113. . 1976. The distribution of the Mediterranean Ordovician Hyolitha. Pp. 491-499 in M. G. Bassett, ed., The Ordovician system: Proceed- ings of a Palaeontological Society symposium. University of Wales Press and National Mu- seum of Wales, Cardiff. 1983a. The Ordovician hyoliths from Anti- Atlas (Morocco).— Sbornik Naradniho Muzea v Praze, Acta Musei Nationalis 39, 35 pp. . 1983b. The hyoliths in the Arenigian of Mon- tagne Noire. Pp. 57-62 in R. Courtessole, L. Marek, J. Pillet, G. Ubaghs, & D. Vizcaino, eds., Calymenina, Echinodermata et Hyolitha de Y’Ordovicien inferieur de la Montagne Noire (France Meridionale).—Memoire de la Societe Scientifiques de l’Aude. —.,&A.Galle. 1976. The tabulate coral Hyostra- gulum, an epizoan with bearing on hyolithid ecology and systematics.— Lethaia 9:51-64. —., & E.L. Yochelson. 1976. Aspects of the bi- ology of Hyolitha.— Lethaia 9:65-82. Matthew, G. F. 1895. Traces of the Ordovician sys- Dai tem on the Atlantic Coast.— Transactions of the Royal Society of Canada | (2nd series):253-271. Miller, S. A., & C. L. Faber. 1894. Description of some Cincinnati fossils. — Journal of the Cincin- nati Society of Natural History 17:137-158. Missarzhevsky, V. V. 1969. Paleontologic part. Pp. 112-320 in M. E. Raaben, ed., The Tommotian Stage and the Lower Cambrian boundary prob- lem. Nauka Publishers, Moscow. Novak, O. 1891. Zur Kenntniss der Fauna der Etage F-fl, in der palaeozischen Schichtengruppen Bohmens. Abhandlungen der k6niglische Ge- sellschaft der Wissenschaften 7, 48 pp. Qian, Y. 1977. Hyolitha and some problematica from the Lower Cambrian Meishucun Stage in central and S.W. China.—Acta Palaeontologica Sinica 16:255-275. . 1978. The Early Cambrian hyolithids in cen- tral and southwest China and their stratigraph- ical significance.— Memoirs of the Nanjing In- stitute of Geology and Palaeontology, Academia Sinica 11:1-38. —., Wen Yu, Diyong Liu, & Zongzhe Wang. 1985. Restudy of the Precambrian—Cambrian bound- ary section at Meishucun of Jinning, Yunnan.— Kexue Tongbao 30:1086-1090. Reed, F. R. C. 1909. Lower Palaeozoic Hyolithidae from Girvan.— Transactions of the Royal So- ciety of Edinburgh 47:203-222. Resser, C. E., & R. Endo. 1937. Descriptions of the fossils. Pp. 103-301 in R. Endo & C. E. Resser, The Sinian and Cambrian formations and fos- sils from southern Manchukuo.— Manchurian Science Museum Bulletin 1. Roy, S. K. 1941. Contributions to paleontology. — Field Museum of Natural History, Geology se- ries 4:203-220. Ruedemann, R. 1901. Trenton conglomerate of Ry- sedorph Hill, Rensselaer County, New York, and its fauna.—New York State Museum Bulletin 49:3-114. 1912. The Lower Siluric shales of the Mo- hawk Valley.—New York State Museum Bul- letin 162, 151 pp. 1926. The Utica and Lorraine formations of New York.—New York State Museum Bulletin 272, 227 pp. Runnegar, B., J. Pojeta, J. Neal, J. D. Taylor, M. E. Taylor, & G. McClung. 1975. Biology of the Hyolitha.—Lethaia 13:21-25. Sinclair, G. W. 1946. Notes on the nomenclature of Hyolithes.—Journal of Paleontology 20:72-85. Syssoiev, V. A. 1962. Cambrian hyolithids from the lower slope of the Aldan Shield. Akademiya Nauk S.S.S.R., 63 pp. . 1968. Stratigraphy and hyolithids of the old- 278 est Cambrian beds of the Siberian Platform. Nauka Publishers, Yakutsk, 67 pp. 1974. Biostratigraphy and paleontology of the lower Cambrian of Europe and western Asia. Akademiya Nauk S.S.S.R., 248 pp. Thoral, M. 1935. Contribution a l’etude paleonto- logique de l’Ordovicien inferieur de la Mon- tagne et revision sommaire de la faune cam- brienne de la Montagne Noire. Montpellier, 362 pp. Val’kov, A. K. 1975. Biostratigraphy and hyolithids of the lower Cambrian of the northeast of the Siberian Platform. Akademiya Nauk S.S.S.R.., 188 pp. Walcott, C. D. 1884. Paleontology of the Eureka Dis- trict, Nevada.— U.S. Geological Survey Mono- graph 8, 298 pp. Webers, G. F. 1972. Paleoecology of the Cambrian and Ordovician strata of Minnesota. Pp. 474— 484 in P. K. Sims & G. B. Morey, eds., Geology of Minnesota: A centennial volume. Minnesota Geological Survey, Minneapolis. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON , & G. S. Austin. 1972. Field trip guidebook for Paleozoic and Mesozoic rocks of southem Minnesota. Minnesota Geological Survey Guidebook Series No. 4. Whitfield, R. P. 1878. Preliminary descriptions of new species of fossils from the lower geological formations of Wisconsin.— Wisconsin Geolog- ical Survey Annual Report for 1877, 50-89. . 1882. Paleontology. Geology of Wisconsin. — Wisconsin Geological Survey, 163-363. Yochelson, E. L. 1984. Speculative functional mor- phology and morphology that could not func- tion: The example of Hyolithes and Biconu- lites. —Malacology 25:255-264. Zazvorka, V. 1928. Revision of the Hyolithi from dy.—Palaeontographica Bohemiae 13:1—22. University of Maryland, European Di- vision, APO New York, 09102. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 279-299 A NEW PYRGULOPSIS (GASTROPODA: HYDROBIIDAE) FROM SOUTHEASTERN CALIFORNIA, WITH A MODEL FOR HISTORICAL DEVELOPMENT OF THE DEATH VALLEY HYDROGRAPHIC SYSTEM Robert Hershler and William L. Pratt Abstract. —Pyrgulopsis giulianii, new species, from a few, mid-elevation streams in southern Sierra Nevada (Indian Wells and Kern River Valleys), is described and new records for three other local congeners also are given. Details of shell form and penial morphology distinguish the new snail from related forms found in Owens Valley. Zoogeographic patterns of Death Valley System Pyrgulopsis support the hypothesis that a highly integrated pluvial drainage was present in the region, and also suggest that the system had various historic connections (possibly non-contemporaneous with the above) to adjacent areas. A model of drainage evolution of the Death Valley System is presented, based on distributional data of hydrobiids and other organisms of perennial waters. This is the third and final paper in a series on systematics of springsnails (Gastropoda: Hydrobiidae) from Death Valley System, southeastern California and southwestern Nevada. Earlier contributions dealt with fauna from the Ash Meadows spring oasis in Amargosa Desert (Hershler & Sada 1987), and Owens and Amargosa Basins (exclusive of above, Hershler 1989). Results of survey of remaining portions of the system and ad- jacent areas are summarized herein, includ- ing description of a new Pyrgulopsis and new records for three other congeners. A model for historical development of drain- age in the Death Valley System is presented, largely based on zoogeography of resident Pyrgulopsis. Materials and Methods Localities visited during this portion of the survey are listed in Appendix 1, and consisted of low- to mid-elevation springs and perennial streams. Fuller treatments of taxa (other than the new species) are in Hershler (1989) and Hershler & Sada (1987). Shell morphometric methodology is that of Hershler (1989). Dots on distribution maps represent one or several closely spaced lo- calities. Repositories of material examined are indicated in the text as follows: LACM — Los Angeles County Museum of Natural History; SBMNH-— Santa Barbara Museum of Natural History; UNLVM-— University of Nevada at Las Vegas Museum of Natural History; USNM—National Museum of Natural History; WBM— Walter Miller per- sonal collection. Systematics Family Hydrobiidae Troschel, 1857 Genus Pyrgulopsis Call & Pilsbry, 1886 Pyrgulopsis giulianii, new species Southern Sierra Nevada springsnail Figs. 1-4 Pyrgulopsis cf. stearnsiana. — Hershler, 1989:194 (Sage, Sand Canyons; figs. 37— 40). Material examined. —California. Kern County: Stream in Sage Canyon, USNM 853520, 857975; Stream in Sand Canyon, USNM 860444 (holotype), 853519 (para- 280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Shell parameters for Pyrgulopsis giulianii. UNSM catalog number and number of specimens (in parentheses) are given beneath locality name. WH = number of whorls, SH = shell height, SW = shell width, LBW = length of body whorl, WBW = width of body whorl, AL = aperture length, AW = aperture width, W = whorlexpansion rate, D = distance of generating curve from coiling axis, T = translation rate, SA = aperture shape. Locality Sand Canyon Sage Canyon Ninemile Canyon Cow Canyon Parameter 853519 (10) 853520 (15) 860446 (6) 860448 (8) WH mean 4.13 4.15 4.25 4.44 SD 0.13 0.16 0.27 0.18 range 4.00-4.25 4.00-4.50 4.00-4.50 4.00-4.50 SH 2.43 (mm) 2.58 3.33 3.16 0.19 0.14 0.41 0.15 2.06-2.74 2.39-2.87 2.82-3.99 3.01-3.48 SW 1.68 1.67 2.26 2.17 0.11 0.13 0.25 0.12 1.45-1.81 1.52-2.04 2.26—-2.60 1.98-2.37 LBW 1.90 1.99 2.55 2.38 0.16 0.11 0.23 0.10 1.61-2.13 1.83-2.22 2.27-2.78 2.24—2.49 WBW 1.43 1.48 1.82 1.77 0.08 0.08 0.18 0.08 1.27-1.53 1.37-1.61 1.56—2.08 1.65-1.89 AL 1.21 1.18 1.59 1.55 0.11 0.08 0.13 0.07 1.03-1.36 1.06-1.39 1.41-1.76 1.35-1.55 AW 0.97 0.96 1.31 1.21 0.07 0.05 0.13 0.06 0.88—1.06 0.88-1.13 1.16-1.49 1.08—1.26 W 1.95 1.99 1.88 1.65 0.15 0.24 0.18 0.19 1.69-2.23 1.59-1.89 1.61-2.14 1.45-2.03 D 0.60 0.62 0.58 0.56 0.04 0.06 0.04 0.03 0.54-0.68 0.51-0.74 0.50-0.62 0.51-0.61 T 4.90 6.29 4.84 4.65 0.47 0.47 0.91 0.51 4.27-5.65 4.32-8.61 3.84-5.96 4.04—-5.31 SA 1.25 1.23 1.22 1.21 0.06 0.06 0.04 0.06 1.14—-1.36 1.13-1.34 1.19-1.29 1.15-1.30 types), 857974, SBMNH 35140 (paratypes); Stream in Sand Canyon, 3.7 km up canyon from US 6, WBM 4230, 4387; Small streamlet, S Fork Short Canyon, USNM 860445; Stream in Ninemile Canyon, USNM 860446; Stream in Grapevine Can- yon, USNM 860447; Stream in Grapevine Canyon, 7.2 km up canyon from US 6, WBM 4229, 4360; Stream in Cow Canyon, USNM 860448. Diagnosis. —A small- to moderate-sized species with ovate conic shell. Penis small relative to head/foot; penial lobe reduced, filament elongate relative to remaining pe- nis. Penial glandular ridges 1-4; ventral ridge sometimes borne on low swelling. VOLUME 103, NUMBER 2 Fig. 1. holotype (standard and side views); c, USNM 860447; d, USNM 860446; e, USNM 860448; f, USNM 860445. The holotype is 2.75 mm tall (other micrographs are printed at the same scale). Description. —Shell morphometric data are in Table 1. Shell (Fig. 1) 2.0-4.0 mm high, height/width, 130-170%. Apex pro- truding (Fig. 2e). Whorls, 4.0-4.5, slightly to moderately convex, with indented su- tures and slight sub-sutural shoulders. Ap- erture ovate, angled above, about half as tall as body whorl. Inner lip straighter than out- er, slightly to moderately thickened and re- Scanning electron micrographs of shells of Pyrgulopsis giulianii, new species: a, b, USNM 860444, flected, adnate to small portion of or slightly separated from body whorl. Outer lip thin; apertural plane slightly tilted relative to coiling axis. Umbilicus slit-like to moder- ately open. Shell surface usually encrusted with brown-black deposits. Operculum (Fig. 2f) paucispiral, with ec- centric nucleus; whorls, 3—4. Opercular sur- face attached to foot bearing elongate (ca. 282 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Pyrgulopsis giulianii: a-d, Radula, USNM 860448 (a, Centrals,bar = 12.0 um; b, Laterals and inner marginals, bar = 8.6 wm; c, Inner marginal, bar = 7.5 um; d, Outer marginals, bar = 8.6 um); e, Shell apex (bar = 136 um), USNM 860447; f, Dorsal view of operculum showing thickened callus (bar = 1.0 mm), USNM 853519. VOLUME 103, NUMBER 2 a 283 Fig. 3. Penes (whole mounts) of P. giulianii: a—c, Dorsal aspects; d, e, Ventral (a, e, USNM 857974, bar = 0.25 mm [others to scale]; b-d, WBM 4229). Note stained glandular ridges (both dorsal and ventral examples visible) and black subepithelial pigment (mostly in penial filaments). 50% of operculum length), gently curved, thickened, non-calcareous, amber callus (similar to those described for congeners by Taylor 1987). Dark, grey-black epithelial pigment on most of snout (except distal tip), proximal portion of cephalic tentacles, along anterior and posterior edges of sides of head/ foot, and part or all of operculigerous lobe. Pigment on central portions of sides of neck absent to dark (dense subepithelial pigment cluster present in area). Ctenidial filaments, ca. 20. Osphradium ca. 25% of ctenidium length. Style sac and remaining stomach ca. equal in length. Small caecal chamber present. Radular (Fig. 2a— d) formula: 5(6)-1-5(6), 2(3,4)-1-3(4,5), 23- 29, 23-35 (from two populations). Central tooth broadly trapezoidal; basal cusps short, triangular; basal process moderately exca- vated. Penis (Fig. 3) rarely protruding be- yond edge of mantle collar, usually of stunt- ed appearance, relatively flat (apart from ventral swelling), longer than wide. Fila- ment slender, sub-equal to remaining penis length. Reduced lobe short relative to fila- ment length. Tip of lobe usually ornament- ed with small glandular ridge; somewhat larger, single ridges on dorsal penial surface 284 Kilometers Fig. 4. Distribution of P. giulianii. at central position ca. halfway between base of penis and base of filament, on ventral surface near (sometimes on) inner edge. A single specimen (from Cow Canyon) had a small fourth ridge located on dorsal surface between ridge on lobe and central dorsal ridge. Filament with dark sub-epithelial pig- ment streak; pigment granules also scattered throughout small area surrounding base of filament. Albumen gland sub-equal to cap- sule gland. Seminal receptacle small, posi- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tioned anterior to bursa copulatrix. Bursa copulatrix small relative to capsule gland, positioned partly posterior to gland. Type locality. — A moderate-sized stream in floor of Sand Canyon, Kern County (cen- ter of section 7, T 25S, R 38E; ca. 1068 m elevation) (Hershler 1989:fig. 8d). Snails common in shallow water (<0.5 m), on wa- tercress. Distribution and habitat.—Known from six small- to moderate-sized streams in VOLUME 103, NUMBER 2 southern Sierra Nevada, of which five are in western Indian Wells Valley and one is in adjacent Kern River Valley (Fig. 4). Etymology. —Named after Derham Giu- liani, an avid student and indefatiguable collector of Great Basin biota, who discov- ered a number of new localities of Pyrgu- lopsis in Owens Valley area. Remarks.—Hershler (1989) tentatively assigned this snail to P. stearnsiana (Pilsbry) based on similarity of shell, but later com- parison with near-topotypical, preserved material of the latter revealed striking dif- ferences in their penes and indicated sepa- rate species status for the populations in southern Sierra Nevada. Pyrgulopsis giuli- anii 1s distinguished from close allies found in Owens Valley (to the north) by its com- bination of large size, moderately elongate shell, and small penis with small lobe and very reduced ventral swelling. Pyrgulopsis micrococcus (Pilsbry, 1893) Fig. 5 Amnicola micrococcus Pilsbry in Stearns, 1893:277 (Small spring in Oasis Valley, Nevada; fig. 1). Pyrgulopsis micrococcus. — Hershler & Sada, 1987:788 (numerous localities, mostly in Ash Meadows; figs. 8a, 9-16). Pyrgulopsis micrococcus. —Hershler, 1989: 182 (numerous localities; figs. 17c, d, 20— Drs)’ Paludestrina stearnsiana. —Berry, 1909:78 (Rill near mouth of Mill Creek Canyon). Amnicola stearnsiana. —Berry, 1948:59 (Mill Creek Canyon). Paludestrina longinqua. —Hannibal, 1912: 34 (Spring branch, mouth of Mill Canon, fide Berry 1909). Hydrobia sp.— Taylor, 1954:69 (Old Wom- an Springs, Inyo Co.). Material examined. —California. San Bernardino County: Mohave River at Bar- stow, USNM 713484; Old Woman Springs, 20.8 km SE of Lucerne Valley, LACM 106639, 107741; USNM 526395; Cushen- 285 bury Springs, 16 km SE of Lucerne Valley, LACM 106640, 106660, 106661, WBM 4270, USNM 860449; Box S Spring, 2.6 km SE of Lucerne Valley, LACM 106641; Box Spring (probably same as above), WBM 4266; Broadbent Spring, 10.2 km ESE of Lucerne Valley, LACM 106639, WBM 4264; Bear Lake, USNM 175096; Roadside spring between N shore highway and Big Bear Lake at point 1.2 km E of road which crosses lake, WBM 4274; Spring-fed pond between N shore road and Big Bear Lake, at point 1.6 km W of road which crosses lake, WBM 4276; Spring zone SW of Big Bear Ranger Station, USNM 860450; Spring, S side of CA Highway 18, N side Big Bear Lake, LACM 106644; Rill near mouth of Mill Creek Canyon, LACM 106646, 106647; Small stream, Mill Creek Canyon, SE of CA Highway 38, 0.34 km NE of Power House Canyon Bridge, LACM 106645; Mill Creek at Thurman Flats Picnic Area, USNM 860451; Spring 4.5 km up Mill Creek Rd. from junction with Yucaipa Rd., WBM 4263. Nevada. Clark County: Cold Creek, UNLVM 3212; Willow Creek, UNLVM 3451. Diagnosis.—A small-sized species, with globose to ovate-conic shell. Penis with moderate-sized lobe; distal edge of lobe usu- ally ornamented with small glandular ridge. Remarks. — This snail is very similar, even in general form of penis and disposition of glandular ridge, to P. stearnsiana (Pilsbry), which occurs in Central California, ““from Sonoma County to Monterey County along the coast and inland in the foothills of the Sierra Nevada” (Taylor 1981:152). The sole reliable distinguishing feature appears to be the larger and longer penial lobe of P. mi- crococcus, but study of additional material of P. stearnsiana will be necessary to deter- mine generality of this difference. Populations in San Bernardino Moun- tains are separated from remaining range of P. micrococcus to the north by poorly watered (and apparently snailless) Mohave Desert (but note that an old collection was 286 taken from an intermediate locale at Bar- stow). Populations in the headwaters of the Willow Creek drainage (which enters Indian Springs Valley) on the northeast slope of Spring Range are apparently relicts from pre- Wisconsin time, since the snail is absent from well-studied (26 samples) late Wis- consin sediments from the valley floor at the mouth of Willow Creek (Quade & Pratt 1989). Pyrgulopsis owensensis Hershler, 1989 Fig. 6 “Undescribed form of Fontelicella’’(?).— Taylor 1985:318 (Owens Valley, E Fork Walker River; unfigd.). Pyrgulopsis owensensis Hershler, 1989:187 (numerous localities in eastern Owens Valley; figs. 26a—d, 27-32). Material examined. —California. Mono County: Spring, W side East Fork Walker River, USNM 860452.—Nevada. Lyon County: Spring at Wiley Ranch, USNM 860453. Diagnosis.—A small- to moderate-sized species with globose to ovate-conic shell. Penis large relative to head/foot; lobe en- larged, filament short. Penial glandular ridges, 2-6; ventral ridge borne on pro- nounced swelling. Remarks. —Populations in Walker Basin represent significant range extension of species (previously known only from Owens Valley) into pluvial Lahontan System. Pyrgulopsis wongi Hershler, 1989 Fig. 7 Pyrgulopsis wongi Hershler, 1989:196 (nu- merous localities in Owens Valley area; figs. 41-47). Material examined.—California. Mono County: Springs SW of Conway Summit, USNM 860454; Spring, Pizona, USNM 869034; Upper Pizona Spring, USNM 869035; Spring in West Queen Canyon, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON USNM 860511; Truman Spring, USNM 860512. Inyo County: Spring NW corner of Round Valley, USNM 860455; Spring, SW corner of Round Valley, USNM 860456; ‘“‘Smoke Spring,” USNM 853926; Springs, Marble Canyon, USNM 860457 (upper spring), 860458 (lower); Spring, 1.2 km NW of Big Pine Spring, USNM 860459; Spring at McMurry Meadows, USNM 860513; Spring on N side of Red Mountain, USNM 860514; Spring in canyon N of McGann Springs, USNM 860515; McGann Springs, USNM 869036; Tub Springs, USNM 869037; Spring, 0.1 km N of Independence Creek, USNM 860460; Springs SW of Lone Pine, USNM 869038 (North), USNM 869039 (South); Spring at Lower Diaz Creek, USNM 869040; Springs at Upper Diaz Creek, USNM 869041 (North), USNM 869042 (South); Spring S of Carrol Creek, USNM 869043; Stream, Talus Canyon, USNM 860461; Canyon N of Johnson Can- yon, USNM 853925; Johnson Canyon, USNM 853928; Stream, Tunawee Canyon, USNM 860462; Canyon S of Tunawee Can- yon, USNM 853927; Stream, Sacatar Can- yon, USNM 860463 (N_ Fork), USNM 860464 (S Fork). Nevada. Esmeralda Coun- ty: Springs at Dyer Ranch, USNM 860465. Mineral County: Huntoon Spring, USNM 869033. Diagnosis. —A small- to moderate-sized species with globose to low conical shell. Penis elongate and broad relative to head/ foot; filament large and lobe moderate-sized. Glandular ridges, 7-12, of which two are borne on prominent ventral swellings lo- cated near distal edge of lobe. Remarks. —The Inyo County popula- tions fall into previously known range of species. Localities in Fish Lake Valley and Mono Valley represent significant range ex- tensions. The latter area, with a recent his- tory of significant volcanic activity (Gilbert et al. 1968, Kilbourne et al. 1980), previ- ously had been considered devoid of peren- nial water mollusks (Taylor 1985:318). VOLUME 103, NUMBER 2 Kilometers Fig. 5. Distribution of P. micrococcus (Pilsbry). Discussion In an earlier paper, Hershler (1989) con- cluded that springsnail zoogeography (in part) supports the frequently advocated hy- pothesis that a highly integrated pluvial drainage was present in Death Valley Sys- tem (Fig. 8) during late Quaternary, pluvial period (10,000—100,000 B.P.). These data also provide evidence of historic connec- 287 tions between the system and adjacent re- gions, some of which were probably pre- Quaternary. Results of this study support this conclusion insofar as additional ex- amples of both of the above features have been discovered, which are synthesized with the earlier data below. Pyrgulopsis is the most diverse (13 spp.) and among the most widespread genera of 288 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Kilometers Fig. 6. Distribution of P. owensensis Hershler. freshwater mollusks in the Death Valley System, and therefore is a highly suitable subject for zoogeographic inquiry. Other prosobranchs associated with perennial waters, 7ryonia Stimpson, 1865 (Hydro- biidae), and Assiminea Fleming, 1828 (As- simineidae), are less diverse and consider- ably more localized in the system (Taylor 1985, Hershler 1987, Hershler & Sada 1987, Hershler 1989), and largely are excluded from further discussion. Distributions of Death Valley System Pyrgulopsis are summarized in Table 2. For purposes of discussion, it is assumed that non-aquatic dispersal of these snails is in- significant, on a large scale, and that distri- bution of a species reflects past or present continuity or near-continuity of aquatic habitat. Evidence for pluvial integration of Death VOLUME 103, NUMBER 2 Kilometers Fig. 7. Distribution of P. wongi Hershler. Valley System is provided by distributions of two of the more widespread forms. The range of Pyrgulopsis wongi (Fig. 7) closely approximates pluvial Owens River drain- age, as this snail extends from Mono Valley through both Adobe and Long Valleys into Owens Valley. This supports inclusion of the former valley (currently without surfi- cial water connection to Owens Basin and considered of uncertain historic relation- 289 ships by Hubbs & Miller 1948:79) in pluvial Owens River drainage. Distribution of P. muicrococcus (Fig. 5) similarly supports in- tegration of now-isolated sub-units of plu- vial Amargosa River drainage (i.e., sites along modern course of river, Ash Mead- ows, northern Death Valley). This species also has been found (Quade & Pratt, pers. comm.) in Wisconsin pluvial sediments of Chicago Valley, a now dry tributary of the 290 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (@) 25 50 75 [Ee a PiKilometens L. Manly YL. Panamint L. Searles L. Mojave to Colorado R. Base map from Snyder et al.1964 118° Fig. 8. Pleistocene Death Valley System showing drainage relations (not necessarily contemporaneous). Stippled areas encircled by dashed lines may not have contained lakes for significant portions of the pluvial period. VOLUME 103, NUMBER 2 Amargosa Basin, together with two other snails from Amargosa drainage, 7ryonia variegata Hershler and Sada and an unde- scribed Assiminea species now living at Ash Meadows and elsewhere in the drainage. In addition, presence of P. micrococcus in Pan- amint Valley and San Bernardino Moun- tains more generally supports pluvial inte- gration of the three rivers comprising Death Valley System (Owens, Amargosa, Mo- have). Snail distributions also imply the follow- ing historic drainage connections: a) be- tween Owens Basin and both Lahontan Sys- tem, and Deep Springs and Fish Lake Valleys; b) between lower Owens Basin and adjacent Kern River Valley; c) between Mo- have and Los Angeles Basins; d) between Amargosa Basin and both Frenchman Flat and Indian Springs Valley; and e) between the system and Saline Valley. Implied connections between Amargosa Basin and both Frenchman Fiat and Indian Springs Valley (which do not currently drain into the system) are intriguing because these areas are currently separated by low, pre- sumably labile divides of alluvial material and could have drained to the Amargosa possibly during the Quaternary (“alluvial cone [drainage] connection” of Hubbs & Miller 1948:147). However, other examples are clearly untenable within the framework of modern regional topography and prob- ably reflect Neogene conditions. For in- stance, distribution of P. wongi implies for- mer aquatic connections between Owens Basin and Deep Springs and Fish Lake Val- leys to the east. These areas are profoundly separated by White Mountains, and surfi- cial water connections between them during pluvial period is extremely unlikely. (The latter two valleys drained [if at all] into La- hontan System [Miller 1928, Hubbs & Mil- ler 1948].) Note, however, that inception of Basin and Range was a relatively recent event (ca. 2-6 million years B.P.) in Owens Valley area (Bachman 1979, Giovannetti 1979, St. Amand-Roquemore 1979) and 291 Table 2. Distributions of Pyrgulopsis spp. Basins or valleys considered part of the Death Valley System by Hubbs & Miller (1948:table 1) are indicated by asterisks. Species Distribution P. aardahli Northern Owens Valley* P. amargosae Lower Amargosa Basin* P. crystalis Ash Meadows (Amargosa Ba- sin)* P. erythropoma Ash Meadows (Amargosa Ba- sin)* P. fairbanksensis Ash Meadows (Amargosa Ba- sin)* P. giulianii Indian Wells Valley,* Kern River Valley P. isolatus Ash Meadows (Amargosa Ba- sin)* P. micrococcus Frenchman Flat, Indian Springs Valley, Amargosa Basin,* Northern Death Val- ley,* Panamint Valley,* Sa- line Valley, San Bernardino Mtns. (Mohave,* Los Ange- les Basins) P. nana Ash Meadows (Amargosa Ba- sin)* P. owensensis Owens Valley,* E Fork Walker Basin P. perturbata Northern Owens Valley* P. pisteri Ash Meadows (Amargosa Ba- sin)* P. wongi Mono Valley,* Long Valley,* Adobe Valley,* Owens Val- ley,* Huntoon Valley, Deep Springs Valley, Fish Lake Valley distribution of P. wongi could reflect prior drainage relationships. A second example involves P. micrococcus, whose distribution suggests a former aquatic connection be- tween Saline Valley and pluvial Owens drainage. Saline Valley is a small, but deeply downthrown basin that probably was closed to external drainage during pluvial times (Hubbs & Miller 1948). The valley is sep- arated from Panamint Valley (which was part of the pluvial Owens System) by the Nelson Range, which connects the Inyo and Cottonwood Ranges. However, Grapevine Canyon, at the eastern end of the Nelson 292 Range, is the trace of a predominantly slip- strike fault (Cemen et al. 1985:fig. 1), and restoring 10 km of movement along this fault would open Saline Valley to Panamint Valley drainage. Taxonomic affinities of Death Valley Sys- tem Pyrgulopsis can only be addressed in a general sense owing to relative paucity of data on adjacent extra-limital faunas, but are consistent with the above in that they point toward a diverse origin of resident forms. Pyrgulopsis wongi from western portion of system (and Lahontan System) is not closely related to other local forms, but is very sim- ilar to P. californiensis (Gregg & Taylor), from west and south of the system (Taylor 1981:152). Also present in Owens Basin is an apparent species flock, comprising P. ow- ensensis and three local endemics, without obvious affinities to other snails found in the system. The presence of P. owensensis in Lahontan System suggests possible der- ivation of the Owens Basin group from the north via Neogene Owens River. Note that P. nevadensis (Stearns), from Lahontan Sys- tem, has a pattern of penial glandular ridges (discerned in rehydrated material; Hershler & Thompson 1987) suggesting a close re- lationship with this group. Pyrgulopsis mi- crococcus has affinities with P. amargosae Hershler from lower Amargosa Basin, but is much more similar to P. stearnsiana from west of the system. Ash Meadows endemics include (at least) two species flocks com- posed of pairs of very similar forms: a) P. erythropoma (Pilsbry) and P. crystalis Hershler & Sada, having trochoid shells and unlobed penes bearing single, central ven- tral glandular ridge; and b) P. nanus Hersh- ler & Sada and P. isolatus Hershler & Sada, having globose-conic shells and large-lobed penes bearing single glandular ridge at ter- minus of lobe. Inclusion of P. pisteri in the former group and P. fairbanksensis Hershler & Sada in the latter is conjectured on basis of general penial similarity, but these dis- play other significant morphological differ- ences from respective group members and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON may represent additional lineages. Ash Meadows snails are dissimilar to an unde- scribed form of the nearby Las Vegas and Pahrump Valleys, which is closely related to P. deserta (Pilsbry) from Virgin Basin; and preliminary observations suggest that the fauna, except P. micrococcus, instead was derived from that of White River drain- age. For example, P. fairbanksensis is sim- ilar to undescribed taxa from Pahranagat, White River, and Railroad Valleys in south- eastern Nevada, whereas both P. nana and P. isolatus closely resemble P. avernalis (Pilsbry) of Moapa warm springs complex (also in southeast Nevada). A hypothetical model of the history of Death Valley System and adjacent drain- ages is described below, based on both geo- logical evidence and distributions of springsnails and other fauna of perennial waters. The model, which partly conforms to interpretations advanced by Hubbs & Miller (1948), Taylor (1985), and Minckley et al. (1986), is heuristic in intention, and presented as a framework for testing hy- potheses generated from study of zoogeog- raphy of regional aquatic organisms. In early Miocene time (Fig. 9), the Mo- have River region probably drained to the Pacific, in the Los Angeles Basin area, as did most of the region immediately east of the present Sierra Nevada.' Fauna in these drainages included progenitors of the pres- ent Pacific coastal Pyrgulopsis and snails in Mohave System which gave rise to P. mi- crococcus. The White—Virgin River region, ‘ Occurrence of P. micrococcus in San Bernardino Mountains, Los Angeles Basin, suggests former drain- age of the Mohave through this area. Three other fresh- water mollusks (Anodonta californiensis Lea, Pisidium compressum Prime, Valvata humeralis Say) also occur (or occurred until recently) in both Los Angeles Basin and Death Valley System and not in coastal California immediately to the north (Taylor 1981, 1985). Addi- tionally, one of the three primary freshwater fishes of Los Angeles Basin, Pantosteus santaanae (Snyder), may have originated within the ancestral Colorado River drainage (Smith 1966, Minckley et al. 1986). VOLUME 103, NUMBER 2 (0) 100 l ! J] Kilometers Fig.9. Death Valley System and adjacent drainages during early Miocene, >16 Ma, before the develop- ment of basin and range structure. Screened areas are Pacific Ocean. Drainage lines are diagrammatic. It is dcubtful whether any drainage, possibly excepting the San Joaquin on west slope of Sierra Nevada (Huber 1981) even approximately followed its modern course at this time. 1 = Future area of Death Valley; 2 = Future area of Owens drainage; 3 = Discharge of Ow- ens—Kern drainage to Pacific; 4 = Ancestral Mohave River; 5 = Los Angeles Basin area; 6 = Amargosa River (included for convenience, existence in early Miocene uncertain); 7 = Ash Meadows; 8 = White-Virgin River system; 9 = Pahrump Valley (present location, which dates from late Miocene); 10 = Ancestral San Joaquin River. including the ancestral Amargosa and Col- orado Rivers, drained eastward across northern Arizona and southeastern Utah to the Gulf of Mexico (Nilsen & McKee 1979), and was then inhabited by progenitors of the present Moapa River and Ash Meadows species complexes. Shift to an extensional tectonic regime and uplift of the basin ranges in the late Miocene (Dickinson 1979) produced major alter- ations in drainage relations (Fig. 10). Uplift of the Spring, Sheep, Las Vegas, Pahranagat, and associated minor ranges, severed the Amargosa Basin; transferring its drainage to the Death Valley System and introducing progenitors of the Ash Meadows Pyrgulop- 293 {0} 100 l ! | Kilometers Fig. 10. Death Valley System and adjacent drain- ages during later Miocene, <15 Ma, after development of basin and range structure in southern Nevada. | = Death Valley region; 2 = Owens drainage region; 3 = Discharge of Owens—Kern drainage to Pacific; 4 = An- cestral Mohave River; 5 = Los Angeles Basin area; 6 = Amargosa River (see note in Fig. 9); 7 = Ash Mead- ows; 8 = White-Virgin River system; 9 = Pahrump Valley (see note in Fig. 9); 10 = Ancestral San Joaquin River; 11 = Bouse Embayment; 12 = “Lake Huala- pai.” sis to the system.” Pyrgulopsis micrococcus penetrated upstream into the Amargosa drainage and presumably gave rise to the local P. amargosae. Direction of flow of Colorado River was reversed, with the river coursing westward in approximately its modern path. Ponding in the present Lake Mead—Lake Mohave area produced the Hu- alapai limestones. These have been inter- preted as marine (Blair & Armstrong 1979), but Taylor (1983) showed that the late Mio- cene molluscan fauna, as far downstream as Parker Dam, consisted of typically fresh- water species (and not the marine taxa char- acteristic of the Bouse Embayment). Ac- cordingly, we map the area of the Hualapai 2 Progenitors of the endemic goodeid fish Empetrich- thys (sister group to Crenichthys of White River drain- age; Parenti 1981) also may have been introduced by this event. 294 (s) 100 L_1| J Kilometers Fig. 11. Death Valley System and adjacent drain- ages during middle Pliocene, ca. 5 Ma, after initiation of major tilting of Sierra Nevada block fault. 1 = Death Valley region; 2 = Owens River; 3 = South Fork Kern River; 4 = Ancestral Mohave River; 5 = Los Angeles Basin area; 6 = Amargosa River; 7 = Ash Meadows; 8 = White-Virgin River system; 9 = Pahrump Valley; 10 = Ancestral San Joaquin River; 11 = Bouse Em- bayment. member as a lake, informally termed “‘Lake Hualapai.”’ Note that Taylor (1983:294) re- corded P. avernalis, presently surviving in the Moapa River drainage, in the Hualapai sediments. During Pliocene time, tilting of the Sierra Nevada fault block beheaded trans-mon- tane drainage, which was diverted south- wards and formed the Owens River, which drained to the Pacific via Mohave River. Uplift began at the south, moving north- wards (Huber 1981). Various evidence in- dicates that Owens Valley was forming by 6 Ma (see above), and the present Owens drainage probably was formed by not long after that time (Fig. 11). Coastal Pyrgulopsis of the P. californiensis group were trans- ferred with the beheaded source areas, and gave rise to P. wong. Severence of the trans- montane headwaters of the San Joaquin River is dated at 3.2 Ma; thereafter drainage was to the Mono Basin, and over the south- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (+) 100 l i ] Kilometers Fig. 12. Death Valley System and adjacent drain- ages during late Pliocene, ca. 4 Ma, after uplift of Trans- verse Ranges and before formation of Death Valley graben. 1 = Death Valley region; 2 = Owens River; 3 = South Fork Kern River; 4 = Mohave River; 5 = Los Angeles Basin; 6 = Amargosa River; 7 = Ash Mead- ows; 8 = White-Virgin River system; 9 = Pahrump Valley; 10 = Ancestral San Joaquin River; 11 = Bouse Embayment. ern sill to the Owens River (Huber 1981). This drainage change may have allowed progenitors of the Pyrgulopsis owensensis group to invade the system from the north. The South Fork Kern River was tributary to the Owens Basin.* Walker Pass, inter- 3 In addition to P. owensensis and P. wongi, two other freshwater mollusks (Helisoma newberryi [Lea], Vor- ticifex effusus [Lea]) occurred in Lahontan and Death Valley Systems during historic times (Taylor 1981). These and another three Lahontan species (Pisidium ultramontanum Prime, Valvata utahensis Call, Stag- nicola kingi [Meek]) were widespread within the latter during the Quaternary (Taylor 1985). The Tui Chub, Gila bicolor (Girard), also occurs in both areas (al- though differentiated in Owens and Mohave drainages; Miller 1973), and several other fishes of Death Valley System are closely related to Lahontan fauna (Minckley et al. 1986). 4 As evidenced by distribution of P. giulianii. Schreck and Behnke (1971) suggested a similar drainage con- nection based on relationships of endemic Upper Kern River Valley golden trout, Salmo aguabonita Jordan. VOLUME 103, NUMBER 2 {e) 100 L__1 _J Kilometers Fig. 13. Death Valley System and adjacent drain- ages during late Pliocene, <3 Ma, after formation of Death Valley graben and beheadal of ancestral San Joaquin drainage. 1 = Death Valley region; 2 = Owens River; 3 = South Fork Kern River; 4 = Mohave River; 5 = Los Angeles Basin; 6 = Amargosa River; 7 = Ash Meadows; 8 = White-Virgin River system; 9 = Pah- rump Valley; 10 = San Joaquin River; 11 = Bouse Embayment. preted as a wind gap in this model, is ap- proximately 700 m above the margin of In- dian Wells Valley, suggesting that drainage severence occurred no earlier than 2.5 Ma (based on uplift rates in the Deadman’s Pass area; Huber 1981), and possibly more re- cently as uplift rates increased southwards. During the late Pliocene (Fig. 12), uplift of the Transverse Ranges, which enclose the present internal drainage of the Mohave Desert, resulted from compression accom- panying the northward movement of the crustal block west of the San Andreas Rift zone (Woodburne 1975). Drainage direc- tion of the Mohave River was reversed, probably with considerable dislocation, re- routing, and ponding. Drainage of the Death Valley System was shifted from the Los An- geles Basin to the Bouse Embayment. De- position in basins east of the San Bernar- dino Mountains apparently shifted from eastern-derived to western-derived sedi- 295 ments around the Hemphillian-Blancan transition (Woodburne 1975). Late Plio- cene tilting of the Sierra Nevada block (Cole & Armentrout 1979) transferred South Fork Kern River from the Owens to the Kern system. Development of internal drainage in the Death Valley rift occurred in early Pleistocene, establishing a new base level and transferring the drainage sump of the Death Valley System from the Bouse Em- bayment to Death Valley (Fig. 13), thereby completing development of the present sys- tem.° Acknowledgments Fieldwork was supported (in part) by Cal- ifornia Fish and Game (Contract FG7342). Bureau of Land Management (Ridgecrest Resource Area) and California Fish and Game (Bishop Office) loaned vehicles and provided other logistic support. Collecting permits were provided by State of Califor- nia and Joshua Tree National Monument. Assistance in the field was provided by J. Aardakl, M. Blymyer, J. Farrell (BLM); D. Wong (California Fish and Game); W. Cas- sidy (Fort Irwin); R. Moon and associates (Joshua Tree National Monument); D. Giu- liani, S. Denton-Pratt, and D. Sada. Mate- rial was loaned by Dr. J. McLean and C. Coney (LACM), and Drs. W. B. Miller and E. Hochberg (personal collection of former, loaned through SBMNH). The assistance of the Scanning Electron Microscopy Labo- ratory (NMNH) is greatly appreciated. M. 5 Although extensional tectonics of the Death Valley region have been extensively studied (e.g., Wernicke et al. 1988 and references cited therein), little has been published regarding drainage. Cemen et al. (1985) re- corded a late Pliocene (to 4 Ma) basin oriented roughly from north of the Black Mountains to Salt Creek, at approximately a 45° angle to the present valley. Hunt and Mabey (1966) made a convincing case for depo- sition of the “‘no. 2 gravel”’ alluvial fans, estimated at 0.5 Ma, under an external drainage regime. They con- sidered major movement of the normal fault along which the valley has been downthrown to be late Qua- ternary in age. 296 Ryan and P. Greenhall (both NMNH, In- vertebrate Zoology) drafted maps, and pre- pared and digitized camera lucida drawings of shells, respectively. The section on his- tory of the Mohave—Death Valley region benefited from comments by D. L. Weide. Literature Cited Bachman, S. B. 1979. Pliocene—Pleistocene break-up of the Sierra Nevada—White-Inyo Mountains block and formation of Owens Valley.—Geol- ogy 6:461-463. Berry, E. G. 1948. Snails collected for the schisto- somiasis investigations.— United States Na- tional Institute of Health Bulletin 189:55-69. Berry, S. S. 1909. The known Mollusca from San Bernardino County, California.— Nautilus 23: 73-79. Blair, W. N., & A. K. Armstrong. 1979. Hualapai Limestone member of the Muddy Creek For- mation: The youngest deposit predating the Grand Canyon, southeastern Nevada and northwestern Arizona.— United States Geolog- ical Survey Professional Paper 1111:1-14. Cemen, I., L. A. Wright, & K. L. Verosub. 1985. Ce- nozoic sedimentation and sequence of defor- mational events at the southeastern end of Fur- nace Creek strike-slip fault zone, Death Valley region, California. Pp. 127-141 in K. T. Biddle & N. Christie-Beck, eds., Strike-slip deforma- tion, basin formation and sedimentation. —So- ciety of Economic Palaeontologists and Min- eralogists Special Publication 37. Cole, M. R., & J. M. Armentrout. 1979. Neogene paleogeography of the western United States. Pp. 297-323 in J. M. Armentrout, M. R. Cole, & H. TerBest, eds., Cenozoic paleogeography of the western United States. — Society of Econom- ic Paleontologists and Mineralogists, Pacific Section, Pacific Coast Paleogeography Sympo- sium 3. Dickinson, W.R. 1979. Cenozoic plate tectonics set- ting of the Cordilleran region in the United States. Pp. 1-13 in J. M. Armentrout, M. R. Cole, & H. TerBest, eds., Cenozoic paleogeography of the western United States. — Society of Econom- ic Paleontologists and Mineralogists, Pacific Section, Pacific Coast Paleogeography Sympo- sium 3. Gilbert, C. M., M. N. Christensen, Y. Al-Rawi, & K. R. Lajoie. 1968. Structural and volcanic his- tory of Mono Basin, California—Nevada. — Geo- logical Society of America Memoir 116:275- 329. Giovannetti, D. 1979. Volcanism and sedimentation PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON associated with the formation of southern Ow- ens Valley.— Geological Society of America An- nual Meeting Abstracts 11:79. Hannibal, H. 1912. The aquatic molluscs of southern California and adjacent regions, a transition fau- na.— Bulletin of the Southern California Acad- emy of Sciences 1 1:18—46. Hershler, R. 1987. Redescription of Assiminea infi- ma Berry 1947 from Death Valley, California. — Veliger 29:274-288. 1989. Springsnails (Gastropoda: Hydrobi- idae) of Owens and Amargosa (exclusive of Ash Meadows) River drainages, Death Valley Sys- tem, California-Nevada.—Proceedings of the Biological Society of Washington 102:176-248. ——, & D. W. Sada. 1987. Springsnails (Gastro- poda: Hydrobiidae) of Ash Meadows, Amar- gosa Basin, California-Nevada.— Proceedings of the Biological Society of Washington 100: 776-843. —., & F.G. Thompson. 1987. North American Hydrobiidae (Gastropoda: Rissoacea): Rede- scription and systematic relationships of Tryo- nia Stimpson, 1865 and Pyrgulopsis Call and Pilsbry, 1886.— Nautilus 101:25—32. Hubbs, C. L., & R. R. Miller. 1948. Correlation be- tween fish distribution and hydrographic history in the desert basins of western United States. Pp. 17-166 in The Great Basin, with emphasis on Glacial and Postglacial times.—Bulletin of the University of Utah 38, Biological Series 10. Huber, N. K. 1981. Amount and timing of uplift and tilt of the central Sierra Nevada— Evidence from the upper San Joaquin Basin.— United States Geological Survey Professional Paper 1197:1— 28. Hunt, C. B., & D. R. Mabey. 1966. Stratigraphy and structure, Death Valley, California.— United States Geological Survey Professional Paper 494A:1-162. Kilbourne, R. T., C. W. Chesterman, & S. H. Hood. 1980. Recent volcanism in the Mono Basin— Long Valley region of Mono County, Califor- nia.—California Division of Mines and Geol- ogy, Special Report 150:7—22. Miller, R. R. 1973. Two new fishes, Gila bicolor sny- deri and Catastomus fumeiventris, from the Ow- ens River basin, California.— Occasional Papers of the Museum of Zoology, University of Mich- igan 667:1-19. Miller, W. J. 1928. Geology of Deep Spring Valley, California.— Journal of Geology 36:510-525. Minckley, W. L., D. A. Hendrickson, & C. E. Bond. 1986. Geography of western North American freshwater fishes: Description and relationships to intracontinental tectonism. Pp. 519-613 (+ bibliography) in C. H. Hocutt & E. O. Wiley, VOLUME 103, NUMBER 2 eds., The zoogeography of North American freshwater fishes. John Wiley and Sons, New York. Nilsen, T. H., & E. H. McKee. 1979. Paleogene pa- leogeography of the western United States. Pp. 257-276 in J. M. Armentrout, M. R. Cole, & H. TerBest, eds., Cenozoic paleogeography of the western United States. —Society of Econom- ic Paleontologists and Mineralogists, Pacific Section, Pacific Coast Paleogeography Sympo- sium 3. Parenti, L.R. 1981. A phylogenetic and biogeograph- ic analysis of cyprinodontiform fishes (Teleos- tei, Atherinomorpha).— Bulletin of the Ameri- can Museum of Natural History 168:335—557. Quade, J.. & W. L. Pratt. 1989. Late Wisconsin groundwater discharge environments of the southwestern Indian Springs Valley, southern Nevada.— Quaternary Research 31:35 1-370. St. Amand, P., & G. R. Roquemore. 1979. Tertiary and Holocene development of the southern Sier- ra Nevada and Coso Range, California. —Tec- tonophysics 52:409—410 (abstract). Schreck, C. B., & R. J. Behnke. 1971. Trouts of the upper Kern River Basin, California, with ref- erence to systematics and evolution of western North American Sa/mo.—Journal of the Fish- eries Research Board of Canada 28:987-998. Smith, G. R. 1966. Distribution and evolution of the North American catostomid fishes of the genus Pantosteus subgenus Catostomus. —Miscella- neous Publications of the Museum of Zoology, University of Michigan 129:1-132. Stearns, R.E.C. 1893. Report on land and freshwater shells collected in California and Nevada by the Death Valley Expedition, including a few ad- ditional species obtained by Dr. C. Hart Mer- riam and assistants in parts of the southwestern United States.— North American Fauna 7:269- 283. Taylor, D. W. 1954. Nonmarine mollusks from the upper Miocene Barstow Formation, Califor- nia.— United States Geological Survey Profes- sional Paper 254C:67-80. 1981. Freshwater mollusks of California: A distributional checklist.—California Fish and Game 67:140-163. . 1983. Late Tertiary mollusks from the lower Colorado River Valley.— University of Michi- gan, Museum of Paleontology Contribution 26: 289-298. . 1985. Evolution of freshwater drainages and molluscs in western North America. Pp. 265-— 321 in C. J. Hocutt & A. B. Leviton, eds., Late Cenozoic history of the Pacific Northwest.— American Association for the Advancement of Science, San Francisco, California. 297 . 1987. Fresh-water molluscs from New Mex- ico and vicinity.— New Mexico Bureau of Mines & Mineral Resources Bulletin 116:1—50. Wernicke, B., G. J. Axen, & J. K. Snow. 1988. Basin and range extensional tectonics at the latitude of Las Vegas, Nevada.—Geological Society of America Bulletin 100:1738-1757. Woodburne, M. O. 1975. Cenozoic stratigraphy of the Transverse Ranges and adjacent areas, southern California.—Geological Society of America Special Paper 162:1-91. (RH) NHB STOP 118, Department of In- vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (WLP) Museum of Natural History, University of Nevada, Las Vegas, 4505 Maryland Parkway S, Las Vegas, Nevada 89154. Appendix Localities visited. Data are name of site, county, state, township and range coordinates, elevation of site, and date of visitation. I. Walker Basin and environs. Cedar Creek (un- named on topo sheet), Slinkard Valley, Mono Co., CA; T 8N, R 22E, NE % sec. 9, 2025 m, 7-12-88.—‘“‘Tin Cup” Spring, Mono Co., CA; T 9N, R 22E, NW sec. 20, 1928 m, 7-12-88.—Spring in unnamed canyon, Slinkard Valley, Mono Co., CA; T 9N, R 22E, SE % sec. 19, 2050 m, 7-12-88.—Springs by cabin, Slinkard Valley, Mono Co., CA; T 8N, R 22E, NE '% sec. 9, 1983 m, 7-12-88.—Springs in SE corner of Slinkard Valley, Mono Co., CA; T 8N, R 22E, SW 4 sec. 15, 2135 m, 7-12-88.—Springs in SE corner of Slinkard Valley (ca. 75 m SE of above), Mono Co., CA; T 8N, R 22E, SW %4 sec. 15, 2135 m, 7-12-88.—Springs en- tering small creek in northern Little Antelope Valley, Mono Co., CA; T 8N, R 22E, SE 4 sec. 13, 1684 m, 7-12-88.—Springs in meadow in northern Little An- telope Valley, Mono Co., CA; T 8N, R 23E, NW 4 sec. 19, 1684 m, 7-12-88.—Spring N of Camp Ante- lope, Mono Co., CA; T 8N, R 23E, NE % sec. 16, 1708 m, 7-14-88.—Leakage from holding tank (of spring), Spring Creek Canyon, Mono Co., CA; T 8N, R 23E, SW 4 sec. 23, 1952 m, 7-13-88.—Spring zone along E side of West Walker River in canyon, Mono Co., CA; T 6N, R 23E, NE 4 sec. 9, 2013 m, 7-13-88.— Spring zone along W side of Little Walker River up- stream from above, Mono Co., CA; T 6N, R 23E, SW Ys sec. 15, 2050 m, 7-12-88.—Brownie Creek, Mono Co., CA; T 6N, R 22E, SW ‘4 sec. 27, 2184 m, 7-13- 88.—Spring by Cloudburst Creek, Mono Co., CA; T 6N, R 22E, NE % sec. 27, 2159 m, 7-13-88.—Spring 298 at Marine Corps Center, Mono Co., CA; T 6N, R 22E, NW 14 sec. 24, 2080 m, 7-13-88.—Seeps along Junc- tion Creek, Mono Co., CA; T 6N, R 23E, NE '% sec. 20, 2098 m, 7-12-88.—Spring along E side of Little Walker River, Mono Co., CA; T 5N, R 23E, NE % sec. 4, 2263 m, 7-14-88.—Burcham Creek, Mono Co., CA; T 6N, R 23E, NW '% sec. 11, 2257 m, 7-12-88.— Stream along Burcham Flat Road, Mono Co., CA; T 6N, R 23E, SW % sec. 14, 2211 m, 7-12-88.—Fales Hot Springs, Mono Co., CA; T 6N, R 23E, SE % sec. 24, 2227 m, 7-12-88.—Stream, Yaney Canyon, Mono Co., CA; T 5N, R 24E, SE % sec. 15, 2098 m, 7-12- 88.—Spring W of Bridgeport Ranger Station, Mono Co., CA; T 5N, R 24E, NW '% sec. 23, 2098 m, 7-12- 88.—Spring just S of above, Mono Co., CA; T 5N, R 24E, NW 4 sec. 23, 2074 m, 7-13-88.— Buckeye Hot Spring (seepage), Mono Co., CA; T 4N, R 24E, NE % sec. 4, 2166 m, 7-13-88.—Seepage along Buckeye Creek, Mono Co., CA; T 4N, R 24E, NW 4 sec. 4, 2166 m, 7-13-88.—Spring N of Twin Lakes, Mono Co., CA; Matterhorn Peak, CA (15), T 4N, R 24E, NW % sec. 28, 2318 m, 7-13-88.—Summers Creek and springs in Summers Meadows, Mono Co., CA; T 3N, R 25E, NW % sec. 6, 2172 m, 7-13-88.—Stream from Cameron Canyon, Mono Co., CA; T 3N, R 25E, NE % sec. 6, 2135 m, 7-13-88.—Spring SW of Little Bodie Mine, Mono Co., CA; T 3N, R 25E, SW % sec. 1, 2306 m, 7-11-88.—Spring ca. 0.4 km NE of above, Mono Co., CA; T 3N, R 25E, SW % sec. 1, 2306 m, 7-11-88.— Warm Springs, Mono Co., CA; T 4N, R 26E, SW 4 sec. 16, 2342 m, 7-11-88.—The Hot Springs, Mono Co., CA; T 4N, R 25E, NW 4 sec. 9, 2013 m, 7-11- 88.— Travertine Springs, Mono Co., CA; T 5N, R 25E, SW '4 sec. 34, 2050 m, 7-11-88.—Small springs along East Walker River NW of Bridgeport, Mono Co., CA, between 1952-2013 m: seep with pool, E side of river, T 6N, R 25E, NW 4 sec. 26; piped spring on W side of river, T 6N, R 25E, NW '% sec. 23, 7-11-88; seep on E side of river, T 6N, R 25E, SE % sec. 14, 7-13- 88; spring W of river and just N of Murphy Creek, T 6N, R 25E, NE % sec. 14, 7-14-88; seeps (2) on steep hill along E side of river, T 6N, R 25E, SW % sec. 12, 7-13 and 14-88; small seeps on W side of river, T 6N, R 25E, SW % sec. 12.—Spring at Wiley Ranch, Lyon Co., NV; T 8N, R 25E, NW % sec. 5, 1922 m, 7-11- 88. II. Mono Basin. Springs SW of Conway Summit, Mono Co., CA; T 2N, R 25E, NE % sec. 2, 2318 m, 11-8-88. III. Adobe Valley and environs. Spring, Pizona, Adobe Valley, Mono Co., CA; T 1N, R 31E, SE % sec. 4, 2135 m, 5-27-89.—Upper Pizona Spring, Adobe Valley, CA; T 1N, R 31E, SW % sec. 11, 2227 m, 5-27-89.—Huntoon Spring, Huntoon Valley, NV; Huntoon Valley, NV—CA (15) (sections unplated), 11.4 km NW from SE corner of quadrangle, 1891 m, 5-28- 89. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON IV. Fish Lake Valley. Springs ca. 2.4 km SW of The Crossing, Esmeralda Co., NV; T 1S, R 36E, SW 4 sec. 16, 1449 m, 7-16-88.—Spring just NE of The Crossing, Esmeralda Co., NV; T 1S, R 36E, NE % sec. 10, 1449 m, 7-16-88.—Springs at Dyer Ranch, Esmeralda Co., NV; T 2S, R 35E, SE % sec. 13, 1440 m, 7-16-88.— Spring W of cemetery, Esmeralda Co., NV; T 2S, R 35E, SW 4 sec. 24, 1458 m, 7-16-88.—Seep in canyon WSW of Dyer School, Esmeralda Co., NV; Mt. Bar- croft, CA—NV (15), 12.5 km SE of NW corner of quad- rangle, 1684 m, 7-16-88.—Cottonwood Creek, Inyo Co., CA; T 6S, R 37E, NW % sec. 5, 1708 m, 7-16- 88. V. Owens Basin. Spring in West Queen Canyon, Mineral Co., NV; T 1N, R 32E, SW 4 sec. 16, 2013 m.— Truman Spring, Mineral Co., NV; T 1N, R 32E, SW 4 sec. 7, 2166 m.—Spring NW corner of Round Valley, Inyo Co., CA; T 5S, R 31E, NW % sec. 25, 1800 m.—Spring, SW corner of Round Valley, Inyo Co., CA; T 6S, R 31E, SE % sec. 31, 1586 m, 8-12- 88.—Springs, Marble Canyon, Inyo Co., CA; T 7S, R 35E, SE % sec. 35, SW 4 sec. 36, 1891-2074 m.— Spring, 1.2 km NW of Big Pine Spring, Inyo Co., CA; T 9S, R 33E, NW % sec. 16, 2074 m, 5-14-88.—Spring at McMurry Meadows, Inyo Co., CA; T 10S, R 33E, NW ‘% sec. 22, 2044 m.—Spring on N side of Red Mountain, T 11S, R 34E, SE % sec. 31, 1403 m.— Spring in canyon north of McGann Springs, Inyo Co., CA; Mt. Pinchot, CA (15), 13.6 km NW of SE corner of quadrangle.— McGann Springs, Inyo Co., CA; T 13S, R 34E, NW 4 sec. 4, 1708 m, 5-21-89.—Tub Springs, Inyo Co., CA; T 13S, R 34E, SE % sec. 17, 1952 m, 4-30-89.—Spring, 0.1 km N of Independence Creek, Inyo Co., CA; T 13S, R 34E, SW % sec. 21, 1830 m, 6-21-88.—Springs SW of Lone Pine, Inyo Co., CA; T 16S, R 36E, SE % sec. 2, 1952 m, 4-29-89.—Spring at Lower Diaz Creek, Inyo Co., CA; T 16S, R 36E, SE 4 sec. 5, 1312 m, 4-29-89.—Springs at Upper Diaz Creek, Inyo Co., CA; Lone Pine, CA (15), 10.3 km NE from SW corner of quadrangle, 1830 m, 5-27-89.—Spring in canyon 1.6 km S of Carrol Creek, Inyo Co., CA; Lone Pine, CA (15), 7.5 km SW from NE corner of quadrangle, 1617 m, 5-9-89.—Stream, Talus Canyon, Inyo Co., CA; Monache Mtn., CA (15), 10 km NNW from SE corner of quadrangle, 1830 m, 10-2-88.— Stream, Tunawee Canyon, Inyo Co., CA; Monache Mtn., CA (15), 7.5 km NNW from SE corner of quad- rangle, 1769 m, 3-28-88.—Stream, Sacatar Canyon, Inyo Co., CA; T 23S, R 37E, secs. 3-4, 1769-1830 m, 5-18-88. VI. Indian Wells Valley. Stream, Ninemile Canyon, Inyo Co., CA; Lamont Peak, CA (7.5), 2.1 km SSW from NE corner of quadrangle, 1769 m, 7-18-88.— Stream, Grapevine Canyon, Inyo Co., CA; T 25S, R 37E, NE % sec. 23, 1281 m, 4-14-88.—Stream, Short Canyon, Inyo Co., CA; T 25S, R 38E, SW % sec. 31. VII. Mohave Basin and envions. Spring, Cow Can- VOLUME 103, NUMBER 2 yon, Kern River Valley, Kern Co., CA; Walker Pass, CA (7.5), 5.0 km WSW from NE corner of quadrangle, 1330 m, 11-4-88.—Canebrake Creek, Kern River Val- ley, Kern Co., CA; T 25S, R 36E, SW 4 sec. 14, 1086 m, 11-4-88.—Spring, Spring Canyon, Kern River Val- ley, Kern Co., CA; T 25S, R 36E, SE % sec. 34, 1171 m, 11-4-88.—Brown Spring, Kern Co., CA; T 27S, R 35E, NW 4 sec. 5, 946 m, 11-4-88.— Kelso Creek, SW of Rocky Point, Kern Co., CA; T 27S, R 35E, NW 4 sec. 20, 988 m, 11-4-88.—Horse Canyon Spring (and spring just to E), Kern Co., CA; Horse Canyon, CA (7.5), 7.6 km SSW of NW corner of quadrangle, 1464 m, 7-18-88.—Bird Spring (dry), Kern Co., CA; Horse Canyon, CA (7.5), 4.1 km ENE of SW corner of quad- rangle, 1208 m, 7-18-88.— Frog Spring, Kern Co., CA; T 28S, R 35E, NE % sec. 10, 1196 m, 11-4-88.—Shoe- maker Spring, Kern Co., CA; T 28S, R 35E, center of section 12, 1452 m, 11-4-88.— Butterbredt Spring, Kern Co., CA; T 29S, R 36E, SE % sec. 28, 1281 m, 11-3- 88.—Spring, Poleline Canyon, Kern Co., CA; T 30S, R 37E, NE % sec. 8, 854 m, 11-3-88.—Alphie Spring, Kern Co., CA; T 29S, R 36E, SE 4 sec. 35, 964 m, 11-3-88.—Spring, Hoffman Canyon, Kern Co., CA; T 30S, R 36E, SW '% sec. 3, 1098 m, 11-3-88.—Sweet- water Spring, Kern Co., CA; T 32S, R 34E, SE % sec. 14, 1388 m, 7-20-88.—Springs S of Proctor Lake, Kern Co., CA; T 32S, R 34E, S sec. 32, 1257 m, 7-20-88.— Spring, Bean Canyon, Kern Co., CA; T 10N, R 14W, SW '%4 sec. 4, 1238 m, 11-3-88.—Indian Spring, Los Angeles Co., CA; T 7N, R 14W, SE % sec. 18, 891 m, 7-21-88.—Little Rock Creek, ca. 8 km upstream from dam, Los Angeles Co., CA; Juniper Hills, CA (7.5), 5.7 km NNE from SW corner of quadrangle, 1159 m, 11- 5-88.— Pallet Creek, Big Rock Creek, Los Angeles Co., CA; T 4N, R 9W, SW 4 sec. 6, 1086 m, 7-21-88.— Icy Springs, Los Angeles Co., CA; T 3N, R 9W, NE %4 sec. 1, 1757 m, 11-5-88.—Spring, Sawmill Canyon, Los Angeles Co., CA; T 3N, R 8W, NE % sec. 11, 2074 m, 11-12-88.—Spring, Grapevine Canyon, San Ber- nardino Co., CA; T 4N, R 2W, NW 1% sec. 26, 1147 m, | 1-6-88.— Rabbit Springs, San Bernardino Co., CA; T 4N, R 1W, NW % sec. 11, 885 m, 7-21-88.—Broad- bent Spring (unnamed on topo sheet), San Bernardino Co., CA; T 3N, R 1E, NW % sec. 3, 1144 m, 7-21- 88.—Cushenbury Springs, San Bernardino Co., CA; T 3N, R 1E, NE % sec. 10, 1251 m, 7-21-88.—Cotton- wood Spring (dry), San Bernardino Co., CA; T 4N, R 2E, SE % sec. 25, 970 m, 7-21-88.—Spring NW of Old Woman Springs (dry), San Bernardino Co., CA; T 4N, R 3E, NE 4 sec. 31, 976 m, 7-22-88.—Old Woman Springs, San Bernardino Co., CA; T 4N, R 3E, NE % sec. 31, 985 m, 7-22-88.—Two Hole Spring, San Ber- nardino Co., CA; Old Woman Springs, CA (15), 11.1 km NE of SW corner of quadrangle, 1165 m, 7-22- 88.—Spring zone SW of Big Bear Ranger Station, San Bernardino Co., CA; T 2N, R 1E, NW 4 sec. 17, 2074 m, 7-21-88.—Springs SW of Fawnskin, San Bernar- 299 dino Co., CA; T 2N, R 1W, SW % sec. 14, 2089 m, 7-21-88.— Mill Creek at Thurman Flats Picnic Area, San Bernardino Co., CA; T 1S, R 1W, NE % sec. 8, 1074 m, 7-27-88.—Little Morongo Creek, below Mo- rongo Lakes, CA; T 1S, R 4E, SE % sec. 24, 756 m, 11-11-88.—Spring, Big Morongo Canyon (upper por- tion), San Bernardino Co., CA; T 1S, R 4E, SE % sec. 18, 1110 m, 11-6-88.—Springs, Big Morongo Canyon (lower portion), San Bernardino Co., CA; T 1S, R 4E, SE %4 sec. 28, 744 m, 11-10-88.—Spring, Smith Water Canyon, Riverside Co., CA; T 2S, R 7E, NW % sec. 5, 1293 m, 11-10-88.—Fortynine Palms Oasis, San Bernardino Co., CA; Queen Mtn., CA (7.5), 3.1 km SE of NW corner of quadrangle, 854 m, 11-10-88.— Garlic Spring, San Bernardino Co., CA; T 13N, R 3E, SW '4 sec. 11, 708 m, 4-9-86.— Mohave River, Afton Canyon, San Bernardino Co., CA; T 11N, R 5E, SW Y% sec. 13, 428 m, 4-10-86.—Seepage, W side of Soda Lake, San Bernardino Co., CA; T 12N, R 8E, NW 4 sec. 2, 290 m, 7-22-88.—Old Mormon Spring (dry), San Bernardino Co., CA; Avawatz Pass, CA (15), 1.8 km NNW of SE corner of quadrangle, 634 m, 11-9- 88.—Snake Spring, Granite Mtns., San Bernardino Co., CA; T 8N, R 13E, SE % sec. 5, 1220 m, 11-7-88.— Cottonwood Spring, Granite Mtns., San Bernardino Co., CA; T 8N, R 13E, NW % sec. 7, 1366 m, 11-7- 88.—Arrowhead Spring, Providence Mtns., San Ber- nardino Co., CA; T 9N, R 13E, SE % sec. 22, 1208 m, 11-7-88.—Van Winkle Spring, San Bernardino Co., CA; T 8N, R 13E, NW % sec. 23, 1098 m, 11-8-88.— Goldstone Spring, Mid Hills, San Bernardino Co., CA; T 10N, R 14E, SW % sec. 31, 1415 m, 11-8-88.—Live Oak Spring, Mid Hills, San Bernardino Co., CA; T 13N, R 13E, NW % sec. 19, 1568 m, 11-7-88.—Key- stone Spring, Mid Hills, San Bernardino Co., CA; T 14N, R 16E, NW 4 sec. 29, 1781 m, 7-23-88.—Dove Spring (dry), New York Mtns., San Bernardino Co., CA; T 15N, R 17E, SW 4% sec. 19, 1452 m, 7-23-88.— Pachalka Spring, Clark Mtn., San Bernardino Co., CA; T 17N, R 12.5E, NE % sec. 36, 1488 m, 7-23-88.— Ivanpah Springs, Clark Mtn., San Bernardino Co., CA; T 17N, R 13E,SE% sec. 24, 1269 m, 7-23-88.—Bonan- za Spring, Clipper Mtns., San Bernardino Co., CA; T 7N, R 15E, NW % sec. 22, 634 m, 11-14-88.—Piute Spring, San Bernardino Co., CA; T 12N, R 18E, NW Y% sec. 24, 915 m, 7-25-88. VIII. Amargosa Basin and environs. Choppo Spring, Inyo Co., CA; T 21N, R 7E, SE % sec. 2, 610 m, 7-25-88.—Tule Spring (dry), California Valley, San Bernardino Co., CA; Tecopa, CA (15), 8.8 km NNW of SE corner of quadrangle, 720 m, 7-25-88.— Beck Spring, Kingston Range, San Bernardino Co., CA; T 20N, R 10E, NE % sec. 31, 1354 m, 7-25-88.—Cold Creek, Clark Co., NV; T 18S, R 55E, NE '% sec. 1, 1867 m, 9-5-77.— Willow Creek, Clark Co., NV; T 18S, R 55E, NE % sec. 2, 1830 m, 7-31-77. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 300-303 SOME TARDIGRADES FROM COLORADO, WITH A DESCRIPTION OF A NEW SPECIES OF MACROBIOTUS (MACROBIOTIDAE: EUTARDIGRADA) R. Deedee Kathman Abstract. — Fourteen species of tardigrades were found above 3200 m on Mt. Evans and an additional species was found at 3963 m at Guenella Pass, Col- orado. One of the species from Mt. Evans, Macrobiotus caelicola, is new to science. It differs from other Macrobiotus species by having a combination of the following characters: the presence of ventral and dorsal pores covering its body, the large size and shape of the claws and dentate lunules, two macro- placoids and no microplacoid, and the shape and size of the eggs. Thirty-one species of tardigrades have been reported from Colorado (Higgins 1959, Baumann 1960, Landreth & Thomas 1970, Anderson et al. 1984, Beasley 1989). The most recent of these publications (Beasley 1989) increased the number from 17 species to the present 31. During the present study, 15 species were collected: five are new to Colorado and one is new to science. Materials and Methods Five samples of moss were collected on Mt. Evans between 3262 m and the summit at 4348 m, and one sample each was col- lected at 3659 m and 3963 m at Guenella Pass, both in Clear Creek County, Colo- rado, U.S.A. All seven samples were col- lected on 11-12 Aug 1986. Each sample of moss was placed in a pa- per bag and air-dried for several months, then removed from the bag, placed in a stoppered funnel and allowed to soak in water for eight hours, after which the moss was removed and shaken in a separate con- tainer of water several times. The water and its contents were poured through a 45 wm sieve to retain the tardigrades, which were placed in a gridded petri dish and extracted under a stereomicroscope. Each tardigrade was placed directly into Hoyer’s mounting medium on a microscope slide and overlain with a cover slip. After complete drying of the mountant, the cover slip was ringed with nail polish to prevent further air penetra- tion. Identifications were made using a phase- contrast compound microscope with oil im- mersion. All measurements were made us- ing a calibrated eyepiece micrometer. All drawings were done with a drawing tube attached to the compound microscope. Results All seven samples contained tardigrades, with a total of 263 individuals belonging to 8 genera and 15 species, distributed among the sites as shown in Table 1. One of these species, Isohypsibius pappi, is new to North America, and three of the species, Hebe- suncus conjungens, Diphascon nodulosum and D. pingue, are reported from Colorado for the first time. The new species, Macro- biotus caelicola, is described below. Taxonomic Account Eutardigrada Marcus, 1927 Macrobiotidae Thulin, 1928 Macrobiotus Schultze, 1834 Macrobiotus caelicola, new species Fig. | Description. — Holotype. Total length 620 um; colorless; eyes present (Fig. 1A). Entire ventral and dorsal surfaces covered with ir- VOLUME 103, NUMBER 2 Table 1. 301 Numbers of individuals in each species found at two locations in Clear Creek County, Colorado. General distributions and descriptions for each species are given in Ramazzotti and Maucci (1983). Guenella Pass 3659 m Echiniscus blumi Richters, 1903 Echiniscus wendti Richters, 1903 Milnesium tardigradum Doyére, 1840 1 Hypsibius convergens (Urbanowicz, 1925) 1 Ramazzottius oberhaeuseri (Doyére, 1840) Isohypsibius pappi (tharos, 1966) TIsohypsibius landalti (Iharos, 1966) Hebesuncus conjungens (Thulin, 1911) Diphascon nodulosum (Ramazzotti, 1957) Diphascon pingue (Marcus, 1936) Diphascon recamieri Richters, 1911 Macrobiotus caelicola new species Macrobiotus harmsworthi Murray, 1907 Macrobiotus hufelandi Schultze, 1834 Minibiotus intermedius (Plate, 1888) regularly-shaped, equal-sized pores (exam- ple of pores in posterior end shown in Fig. 1A). Buccal lamellae present. Buccal ring with distinct dentation. Buccal tube with ventral tube support; buccopharyngeal tube 66 wm long, 5 wm wide. Pharyngeal bulb large, wider than long; apophyses large; 2 macroplacoids, the first 8 wm long, the sec- ond 6 um long; no microplacoid (Fig. 1B). Furcae as shown in Fig. 1C. Doubleclaws large, Y-shaped; 2 large accessory points on each primary branch; primary branch long and thin, 17.5 um in leg II, 32.5 um in leg IV; secondary branch short and close to base, 12.5 um in leg II, 20 um in leg IV; lunules large and dentate (Fig. 1D, E). Sclerotized bar below the claws on the first 3 pairs of legs (Fig. 2D). USNM 235439. Paratypes.— Total length up to 668 um. Buccopharyngeal tube length 66-70 um, width 5-6 um, distance between stylet sup- port insertion and end of tube (= pharyngeal tube length) 13-15 wm. First macroplacoid length 8-10 um, second macroplacoid length 6-8 um, with the first always longer than the second. Sclerotized bar below the claws in all specimens. 45 specimens: USNM 235440-235442; 2 specimens in Dastych Mt. Evans 3963 m 3262 m 3811 m 4299 m 4348 m 19 1 1 91 15 ee NS) = 46 collection (Hamburg, West Germany); 2 specimens in Kristensen collection (Uni- versity of Copenhagen, Denmark); remain- ing specimens in Kathman collection (Sid- ney, British Columbia). Eggs. —Eggs round, up to 124 um in di- ameter; covered with projections up to 34 um long (Fig. 1 F); projections thin, with dark patches on exterior, apices divided into multi-tipped points, some with tiny setae projecting from them (Fig. 1G); egg surface smooth between projections. USNM 235443 1-235444. Type locality.— All specimens were col- lected on 11 Aug 1986 at 4299 m, Mt. Ev- ans, Clear Creek County, Colorado, U.S.A. Etymology.—Caelicola is a masculine Latin word meaning dweller in heaven; this species is thus named because it was found at such a high altitude as well as pertaining to the surname of the person in charge of the Mt. Evans Research Station, Dr. Robert Angell. Discussion This is the first report of a Macrobiotus species with a sclerotized bar below the claws. It is generally thicker and larger than PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. those bars found in species of Diphascon or Isohypsibius, at least superficially more closely resembling the leg plate of some echiniscids. The lower portion of the bar appears to have two distinct elliptical areas with fine spine-like granulation, but these could not be clearly discerned using light microscopy, even at 2000 magnification and using phase contrast lenses. This species is most similar to Macrobi- otus islandicus Richters, 1904 but differs in the following characters. Macrobiotus islan- dicus has small pores arranged in transverse rows on the dorsal and lateral surfaces, while M. caelicola has irregularly-spaced pores not in transverse bands covering both the dorsal and ventral surfaces. The width of the buc- cal tube for M. islandicus is much larger than for M. caelicola, with the width to length ratio being approximately 14% for M. islandicus and 7.5-8.5% for M. caelicola. There is a curve in the rostral part of the Macrobiotus caelicola. A, Entire animal, ventral view, pores cover entire ventral and dorsal surfaces; B, Buccopharyngeal apparatus; C, Furca; D, Claws of 2nd leg; E, Claws of 4th leg; F, Egg (split); G, Projections on egg. Scale bars in ym as follows: A, 80; B, 24; C, D, 10; E, 12; F, 40; G, 12. buccal tube for M. islandicus, while the buc- cal tube for M. caelicola is straight. Both macroplacoids are longer in M. islandicus than in M. caelicola, and their profiles are smooth in M. islandicus but rough-edged in M. caelicola. In M. islandicus the secondary branch of each doubleclaw is inserted half- way or slightly more than halfway up the primary branch, while in M. caelicola the secondary branch is inserted approximately one-third of the distance from the base of the primary branch; both branches are long- er and thinner in M. caelicola. The lunules are always obvious and dentate in M. cae- licola. The eggs of M. caelicola are larger (mean diameter of 120 wm) than for M. is- landicus (90-100 wm diameter). The pro- jections on the eggs of M. islandicus are 11— 12 um maximum length, while those of /. caelicola are much longer (mean length = 20 um for 9 eggs), reaching 34 wm in some eggs. No other Macrobiotus species have eggs VOLUME 103, NUMBER 2 with the characters of M. caelicola. If eggs are not used for differentiation from other species, there are only two other moss- dwelling Macrobiotus species with the com- bined characters of cuticular pores, two ma- croplacoids, and no microplacoid. They are M. islandicus and Macrobiotus annae Richters, 1908. M. caelicola differs from M. islandicus as discussed above, and from /. annae in that M. annae is a small tardigrade (length to 370 um) and has a narrow buccal tube (3 wm), a small oval pharynx, small claws, and small smooth lunules. The 15 species of tardigrades collected during this study increase the number known from Colorado from 31 to 37. Although all 15 species were collected at high altitudes, none of them appear to be restricted to these altitudes, since all of them (except the newly reported species) have been reported else- where at much lower altitudes. Acknowledgments I wish to thank R. M. Kristensen and H. Dastych for examining several of the species and identifying /. caelicola as a new species. Thanks are also due R. O. Brinkhurst for collecting the moss while teaching at the Mt. Evans Research Station of the University of Denver. 303 Literature Cited Anderson, R. V., R. E. Ingham, J. A. Trofymow, & D. C. Coleman. 1984. Soil mesofaunal distribu- tion in relation to habitat types in a shortgrass prairie.— Pedobiologia 26:257-261. Baumann, H. 1960. Beitrag zur Kenntnis der Tar- digraden in Nord-Amerika.— Zoologischer An- zeiger 165:123-128. Beasley, C. W. 1989. Taradigrada from Gunnison County, Colorado, with the description of a new species, Diphascon craigi.—The Southwestern Naturalist. (In Press) Higgins, R. P. 1959. Life history of Macrobiotus is- landicus Richters with notes on other tardi- grades from Colorado.—Transactions of the American Microscopical Society 78:137-154. Landreth, K., Jr., & B.O. Thomas. 1970. Studies of the egg laying process in Tardigrada [Hypsibius (Isohypsibius) augusti] in Weld County.—Jour- nal of the Colorado—Wyoming Academy of Sci- ence 7:9. Ramazzotti, G., & W. Maucci. 1983. Il phylum Tar- digrada.—Memorie dell’Istituto Italiano di Id- robiologia 41:1-1012. Department of Biology, P.O. Box 1700, University of Victoria, Victoria, British Co- lumbia, Canada V8W 2Y2; (Present ad- dress) 10651 Blue Heron Road, Sidney, British Columbia, Canada V8L 3X9. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 304-310 PYCNOGONIDA OF THE WESTERN PACIFIC ISLANDS, VII. ON SOME RARE SPECIES FROM THE FLORES SEA, INDONESIA K. Nakamura and C. Allan Child Abstract. —During a fisheries research cruise by the Japanese fisheries re- search vessel Hakuho Maru, in the Flores Sea, Indonesia, a joint cruise with LON-LIPI, Indonesia, WESTPAC, IOC-UNESCO, and the Ocean Research Institute of the University of Tokyo, two trawling stations resulted in the capture of several rare species of pycnogonids. Two of these species were taken only for the second time since their original descriptions in 1908; Bathyzetes setiger (Loman), and Ascorhynchus levissimus Loman. They are refigured and discussed in light of current morphological knowledge. Three other species rare in lit- erature were taken and are discussed; Hemichela micrasterias Stock, Pallenopsis tydemani tydemani Loman, and Colossendeis leptorhynchus Hoek, while the cosmopolitan species C. macerrima Wilson was also taken. Pigrogromitus tim- sanus Calman, collected at a shore station near the Celebes, has also been included. During the course of joint research cruise KH-85-1 in the Flores Sea, Indonesia, in- volving LON-LIPI, Indonesia, and WEST- PAC, IOC-UNESCO with the vessel Ha- kuho Maru, Ocean Research Institute of the University of Tokyo, two trawl stations were made in about 600 meters which resulted in the capture of five rare or little known pycnogonids. Two of these species, Asco- rhynchus levissimus Loman and Bathyzetes setiger (Loman), were taken only for the sec- ond time since their description in 1908. Three other species, Hemichela micrasteri- as Stock, Pallenopsis tydemani tydemani Loman, and Colossendeis leptorhynchus Hoek, have few specimens recorded in lit- erature and are thus relatively rare. The very common Colossendeis macerrima Wilson was also captured at one of the stations. This frequency of capture suggests a rich faunal assemblage, particularly from so few sta- tions. This richness is in keeping with the literature of Indonesian pycnogonids which has, since the first large monograph by Lo- man in 1908, consistently proved that there are great numbers of pycnogonid species and specimens to be found in this huge archi- pelago. Stock (1953) recorded additional pycno- gonid fauna in his Indonesia—Philippine re- port and there have been other Indonesian faunal listings in subsequent reports on that region (Stock 1954, 1968, 1983, 1985). The rarity of the many species known to inhabit Indonesia may be in part due to the past minimal collecting frequency for the whole archipelago, particularly in the deeper slope and basin areas of its various seas. The distribution pattern of the two rarest species, known from their two captures only, is, of course, central Indonesia. That of Hemichela micrasterias is along a corridor from Indonesia to central Japan, including the Philippines. The distribution of Palle- nopsis t. tydemani is only known from In- donesia and Japan, from 100 to 800 meters. Members of the genus Colossendeis have usually been found to be cosmopolitan in VOLUME 103, NUMBER 2 deep waters. This may hold true for C. /ep- torhynchus which is known from the Atlan- tic and Pacific Oceans and is certainly true for C. macerrima. Another cosmopolitan or at least pantropical species, Pigrogromitus timsanus Calman, was taken incidentally at a shore station just prior to the Flores Sea stations. Systematics Family Ammotheidae Genus Ascorhynchus Sars Ascorhynchus levissimus Loman Fig. 1A-D Ascorhynchus levissimus Loman, 1908:33- 34, pl. IV, figs. 46-5 1.—Stock, 1953:304 [key]; 1975:130 [text]. Material examined. — Flores Sea: 05°56.0'S, 119°29.0'E, 630-657 m, sta. KH85-1, B-1, 12 Feb 1985 (1 9). Distribution. —This is the second speci- men of this species known since Loman de- scribed the first, also from the Flores Sea (07°24’S, 118°15.2'E). His specimen was from slightly deeper water, 794 m. It is thus confined, to our knowledge, to the Flores Sea, but since the entire archipelago is so rarely collected, particularly its deeper slopes and basins, the species may have a wider distribution when other collections from adjacent slopes and basins become known. Remarks. —From examination of the above specimen, Loman’s type specimen also appears to be a female, judging from the oviger sixth segment which lacks the usual tuft of setae common to males of the larger species of Ascorhynchus. The ocular tubercle is less a broadly rounded hump than in Loman’s type figures. In the Hakuho Maru specimen, it is more abbreviated with its anterior surface joined to the neck at a sharper angle and it has unpigmented eyes which are readily discernible, at least in their upper halves. The only other major difference between the two specimens is that the anterior pair 305 of propodal claws in this specimen are tiny triangular points not as long as the propodal diameter. In Loman’s figure of the entire specimen (pl. IV, fig. 48), the propodal claws are indistinctly illustrated as approximately equal in size. The remaining six claws of the specimen in hand are about 0.3 the pro- podal length and the tarsus and propodus of each are almost equal in length. A set of figures are provided herein as the type fig- ures are somewhat diagrammatic and do not provide details of currently recognized critical characters. This giant species was inadvertently omitted from a recent key (Child 1987:906- 907) to the very large species of this genus. It can be followed to couplet three, second part, where it would be listed as having a tarsus equal to or slightly longer than the propodus, a very shortened propodal claw on the first pair of legs, and a second scape segment longer than the first, both of which form the chelifore, along with a tiny chela of very reduced size. Genus Bathyzetes Stock, 1955 Bathyzetes setiger (Loman, 1908) Fig. 1E-G Eurycyde setigera Loman, 1908:29-30, pl. V, figs. 52-58. Bathyzetes setiger.—Stock, 1955:261, 262 [key], fig. 24. Material examined. — Flores Sea: 05°54.9’S, 119°29.5’E, 558-593 m, sta. KH85-1, B-2, 12 Feb 1985 (1 9). Distribution. —This record, like the last one, marks only the second time this species has been collected, but like other species in this report, it is possibly not as rare as the frequency of collections from deeper In- donesian seas. Loman’s four type specimens came from further north, in the Molucca Sea, off northeastern Sulawesi (Celebes) Is- land, in 1165-1264 meters. This second capture extends its distribution south into the Flores Sea, and into shallower waters at a minimum depth of 558 meters. 306 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | by ual y x1 {i l ‘ ae nee Og bow Na SS SEE \ ‘ G ao oa Fig. 1 Ascorhynchus levissimum Loman, female: A, Trunk, dorsal view; B, Trunk, lateral view; C, Terminal segments of first leg, with terminal claw enlarged; D, Terminal segments of fourth leg. Bathyzetes setiger (Loman), female: E, Trunk, dorsal view; F, Trunk, lateral view; G, Terminal segments of third leg. VOLUME 103, NUMBER 2 Remarks. — Thanks to J. H. Stock, Insti- tute for Taxonomic Zoology, University of Amsterdam, we were permitted to examine Loman’s syntypes of this species and of B. virago (Loman). The two species are very closely related and could be synonymized except for the differences in size, setae size and arrangement, palp segment length ra- tios, and differences in the strigilis segments of the oviger. Loman’s B. setiger is twice the size of B. virago, a difference which is not a valid species diagnostic character in itself (e.g., the great size differences in specimens of Anoplodactylus lentus Wilson). Bathy- zetes setiger has a seventh palp segment longer than the eighth while this segment is shorter than the eighth in B. virago. As Stock (1955:262, fig. 24) illustrated the terminal oviger segments differ in the two species. B. setiger has a longer terminal claw extending well beyond the enlarged distal denticulate spine, while the claw of B. virago is hardly longer than the denticulate spine with which it is paired. In the two specimens we examined from the type series, B. virago appears notably more setose with many more long slender setae on the chelifores and legs than in B. setiger. The above female agrees in most all characters with B. setiger except that it appears to have even fewer setae than the syntype. The very long dor- sodistal setae of the trunk and lateral pro- cesses are even more prominent with fewer short setae around each. It would be difficult to confuse either of these setose species with a species of another genus, particularly with the absence of any form of ocular tubercle on either of the Bathyzetes species. Genus Hemichela Stock, 1954 Hemichela micrasterias Stock Hemichela micrasterias Stock, 1954:90-94, figs. 42-43; 1985:153-155, figs. 1-11.— Staples, 1982:464—-465 [text].—Child, 1988a:10.—Nakamura & Child (1990). Material examined. — Flores Sea: 05°56.0'S, 119°29.0’E, 630-657 m, sta. KH85-1, B-1, 12 307 Feb 1985 (1. subadult); 05°54.9’S, 119°29.5’E, 558-593 m, sta. KH85-1, B-2, 12 Feb 1985 (1 ovig 2 1, juv). Distribution. —This species was originally described from one Indonesian specimen and has recently been recorded from the Philippines (Child 1988a:10) and Japan (Nakamura & Child 1990). This distribu- tion more or less follows a corridor theory (Child 1983:699, 713, 1988b:55) proposed as relating western Pacific species distribu- tions to a pathway extending from at least as far south as New Zealand to as far north as the Japanese Islands. The known distri- bution of several genera (Austrodecus, Hemichela, Cheilopallene, and others) and a number of species (this species being one example) appear to follow the western Pa- cific island archipelagoes north and south, presumably in accordance with north-south current trends among these islands. The previously known depths at which this species was taken were 20-84 meters, but the above specimens are the first from much greater depths; 558-657 meters. Remarks.—This genus is closely related to Paranymphon in most characters except for the loss in adults of the immovable che- lae fingers and a few other differences in less conspicuous characters. The lateral process tubercles, for instance, are very similar in the two genera but the tubercles, for Hemi- chela, are much larger in these specimens from the Flores Sea than those illustrated by Stock (1954:91, fig. 42a) for the type. They are at least equal in length to or even larger than the tubercles of P. spinosum Caullery, and much taller than those of P. magnidigitatum Hong & Kim (1987). Both genera have lateral dendritic spines or tu- bercles on the anterior and posterior of their lateral processes, at least in the male, and have very tall ocular tubercles and abdom- ina. The above specimens have no suggestion of eyes, in keeping with their deeper water habitats, and their ocular tubercles are much more slender distally, with prominent lat- 308 eral sensory papillae. The ocular tubercle of Stock’s type specimen is distally broader and has darker areas suggesting eyes. The chelae develop their single finger as- pect very late in the subadult stage. The juvenile has two well-developed chela fin- gers on each chelifore with teeth equal to those of the type specimen. The subadult from station B-1 has fully formed ovigers but an immovable finger is still present on both chelae, but it is reduced in size relative to the movable finger. Only in adults with fully developed sexual characters is the im- movable finger missing or fully atrophied. Family Callipallenidae Hilton Genus Pigrogromitus Calman, 1927 Pigrogromitus timsanus Calman Pigrogromitus timsanus Calman, 1927:408— 410, fig. 104a-f.—Child, 1988a:21 [lit- erature]. Material examined. —Sulawesi (Celebes) Island: Samalona Island, just off Ujung Pan- dang (Makassar), coll. Dr. T. Miura, 2 m, 8 Feb 1985 (1 2). Distribution. —This species is pantropical and in the Indo-Pacific, follows a north— south distribution from Australia through the island archipelagoes to at least as far north as the Ryukyu Islands. It has only been taken in shallow localities. Genus Pallenopsis Wilson, 1881 Subgenus (Bathypallenopsis) Stock, 1975 Pallenopsis (Bathypallenopsis) tydemani tydemani Loman Pallenopsis tydemani Loman, 1908:65-66, pl. 10, figs. 139-145.—Hedgpeth, 1949: 277, fig. 361, j.—Utinomi, 1951:160; 1971:323. Pallenopsis (Bathypallenopsis) tydemani ty- demani. —Stock, 1975:1032 [text], fig. 31c, 1035-1036 [text]. Material examined. — Flores Sea: 05°54.9'S, 119°29.5’E, 558-593 m, sta. KH85-1, B-2, 12 Feb 1985 (1 9). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Distribution. —This is another species represented in Loman’s pioneering mono- graph on the Indonesian fauna. He origi- nally described it from specimens taken in the Flores Sea in 694 and 794 m, and it has since been collected off Kyushu and Honshu Islands in Japan in about 800 and 100—200 m respectively. This female was collected just northeast of the syntypes. Remarks.—The specimen has only one leg which survived the rigors of the trawl, but the ovigers are much like those figured by Stock (1975:1033, fig. 31c) of the syn- type. There are apparently only six specimens of this species recorded in the literature, making it another rarity, but it is adequately figured by Hedgpeth, Loman, and Stock. Family Colossendeidae Hoek Genus Colossendeis Jarzynsky Colossendeis leptorhynchus Hoek, 1881 Colossendeis leptorhynchus Hoek, 1881:64— 65, pl. VIII, figs. 3-7.—Stock, 1978:402, 406-408, fig. 21; 1981:454-455; 1983: 299-300; 1986:417. Colossendeis pennata Pushkin, 1970:1490- 1492, fig. 2. Material examined. —Flores Sea: 05°56.0’S, 119°29.0’E, 630-657 m, sta. KH85-1, B-1, 12 Feb 1985 (1 9); 05°54.9’S, 119°29.5’E, 558— 593 m, sta. KH85-1, B-2, 12 Feb 1985 (1 2). Distribution.—This species is found in predominantly Southern Hemisphere deep- water localities and could be cosmopolitan in distribution if enough specimens were known from a few other collecting sites. It has not been taken yet in the Indian Ocean, for instance. Hoek’s types were taken from off Valpariso, Chile, and the species has been taken in the Walvis Basin off southern Af- rica, in Indonesia, the Philippines, and the Caribbean Basin in depths of 531-3675 m. Remarks. —The tarsus of the species and these specimens is almost twice the pro- podal length and the sixth palp segment is VOLUME 103, NUMBER 2 very slightly shorter than the slender sev- enth. The seventh segment is slightly longer than the combined length of the terminal three segments. The tarsus is notably long in this species, longer than that of its nearest relation, C. macerrima Wilson, but the ter- minal claw is longer than described for the syntype by Stock (1978:402). The claws of the above specimens are less than four times shorter than the propodus while those de- scribed by Stock are slightly more than six times shorter. All of these characters, except for claw length, conform to Stock’s descrip- tion of Hoek’s syntypes and tend to rein- force the separation of this species from oth- ers with which it has been synonymized in the past. Colossendeis macerrima Wilson, 1881 Ccolossendeis macerrima Wilson, 1881:246— 247, pl. I, figs. 9-12, pl. V, fig. 32.—Fry & Hedgpeth, 1969:53, figs. 7, 8 [litera- ture].—Stock, 1978:400—401, fig. 2M. Colossendeis villegentei A. Milne-Edwards, 1881:933. Colossendeis leptorhynchus var. septen- trionalis Caullery, 1896:362-363. Material examined. — Flores Sea: 05°54.9’S, 119°29.5'E, 558-593 m, sta. KH85-1, B-2, 12 Feb 1985 (1 9). Distribution. —This is a cosmopolitan deep-water species found from about 400 to almost 4000 m. Remarks.—This is another species with a very long slender proboscis which is usu- ally slightly upturned distally. The third seg- ment of the palp, as emphasized by Stock (1978:400—401), is notably shorter than the fifth segment, shorter in this ratio than with any other related species. Palp segments 6 and 7 are not as long as in C. /eptorhynchus, and are approximately equal in length while lacking the slender proportions of Hoek’s species. The two species are otherwise dif- ficult to separate as they tend to be of similar sizes along with having slender proboscides, the most immediately obvious character of 309 the species. The comparative figures given by Stock (1978:fig. 2) are excellent for sep- arating these two species along with C. cu- curbita Cole. Acknowledgments We wish to thank Dr. Masumi Horikoshi, chief scientist of this cruise and Dr. Suguru Ohta of the Ocean Research Institute, the University of Tokyo, for their generosity and kindnesses to one of us (KN). We also thank Dr. Shigeo Gamo, Yokohama Na- tional University, for the loan of a net and Dr. Tomoyuki Miura, Kagoshima Univer- sity, for his collaboration in shore collect- ing. We acknowledge with thanks the help of Dr. J. H. Stock, Institute of Taxonomic Zoology, University of Amsterdam, for loaning syntype specimens of Loman’s Ba- thyzetes. Literature Cited Calman, W.T. 1927. 28. Report on the Pycnogonida. Zoological Results of the Cambridge Expedition to the Suez Canal, 1924.—Transactions of the Zoological Society of London 22(3):403-410, 3 figs. Caullery, M. 1896. Pycnogonides. Resultats scienti- fiques de la campagne du Caudan dans le golfe de Gascogne, aout-sept. 1895.—Annales de Universite de Lyon 26:361-364, pl. XII. Child, C. A. 1983. Pycnogonida of the Western Pa- cific Islands, II: Guam and the Palau Islands. — Proceedings of the Biological Society of Wash- ington 96(4):698-714, 5 figs. 1987. New and little known Pycnogonida from Antarctic and Subantarctic waters. — Pro- ceedings of the Biological Society of Washington 100(4):902-9 16, 6 figs. 1988a. Pycnogonida of the Western Pacific Islands, III: Recent Smithsonian—Philippine ex- peditions.—Smithsonian Contributions to Zo- ology 468:i-iv, 1-32, 12 figs. 1988b. Pycnogonida from Aldabra Atoll.— Bulletin of the Biological Society of Washington 8:45-78, 9 figs. Fry, W. G., & J. W. Hedgpeth. 1969. Pycnogonida. 1. Colossendeidae, Pycnogonidae, Endeidae, Ammotheidae.—Fauna of the Ross Sea, 7. Memoirs of the New Zealand Oceanographic Institute 49:1-139, 1 pl., 209 figs., 16 tabs. Hedgpeth, J. W. 1949. Report on the Pycnogonida 310 collected by the Albatross in Japanese waters in 1900 and 1906.—Proceedings of the United States National Museum 98(3231):233-321, figs. 18-51. Hoek, P. P. C. 1881. Report on the Pycnogonida dredged by HMS Challenger 1873-76.—Re- ports of the Scientific Results of the Exploring Voyage of HMS Challenger 3(10):1-167, 21 pls., 2 figs. Hong, J.-S., & I. H. Kim. 1987. Korean pycnogonids chiefly based on the collections of the Korea Ocean Research and Development Institute. — Korean Journal of Systematic Zoology 3(2);137- 164, 17 figs., 1 tab. Loman, J. C. C. 1908. Die Pantopoden der Siboga- Expedition.—Siboga Expeditie Monographie 40: 1-88, I-XV, 4 figs. Milne-Edwards, A. 1881. Compte rendu sommaire d’une exploration zoologique faite dans |’Atlan- tique, a bord du navire le Travailleur.— Comptes Rendus Hebdomadaires des Seances de |’Aca- demie des Sciences, Paris 93:93 1-936. Nakamura, K., & C. A. Child. 1990. Pycnogonida from waters adjacent to Japan.—Smithsonian Contributions to Zoology (in press). Pushkin, A. F. 1970. [New species of the genus Co- lossendeis (Pantopoda).]— Zoologicheskii Zhur- nal 49(10):1488-1496, 4 figs. [In Russian with English summary.] Staples, D. A. 1982. Pycnogonida of the Calliope River and Auckland Creek, Queensland. — Memoirs of the Queensland Museum 20(3):455— 471, 5 figs., 1 pl. Stock, J. H. 1953. Contributions to the knowledge of the pycnogonid fauna of the East Indian Archi- pelago. Biological Results of the Snellius Ex- pedition, XVII.—Temminckia 9:276-313, 18 figs. 1954. Pycnogonida from Indo-West Pacific, Australian and New Zealand waters. Papers from Dr Th. Mortensen’s Pacific Expedition 1914— 1916.— Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn 116:1- 168, 81 figs. 1955. Pycnogonida from the West Indies, Central America and the Pacific Coast of North America. Papers from Dr Th. Mortensen’s Pa- cific Expedition 1914-1916.—Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn, 117:209-266, 26 figs. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1975. Pycnogonida from the continental shelf, slope, and deep sea of the tropical Atlantic and East Pacific. Biological results of the University of Miami deep-sea expeditions, 108.— Bulletin of Marine Science 24(4):957-1092, 59 figs. . 1978. Abyssal Pycnogonida from the North- eastern Atlantic Basin, Part IJ.—Cahiers de Biologie marine 19:397-413, 3 figs. . 1981. Abyssal Pycnogonida from the Walvis Basin, Southern Atlantic.— Cahiers de Biologie marine 22:453-471, 8 figs. 1983. Pycnogonides des campagnes COR- INDON et MUSORSTOM II (detroit de Mak- assar et Philippines), avec description de Pal- lenopsis dentifera sp. nov.— Bulletin du Museum National d’Histoire Naturelle, Paris (4)5, sec. A(1):299-305, 10 figs. . 1985. Deux Pycnogonides rares, recoltes pen- dant la campagne CORINDON II dans le de- troit de makassar, Indonesie.— Bulletin Zoolo- gisch Museum, Universiteit van Amsterdam 10(18):153-158, 22 figs. 1986. Pycnogonida from the Caribbean and the Straits of Florida. Biological results of the University of Miami deep-sea expeditions. — Bulletin of Marine Science 38(3):399-441, 15 figs. Utinomi, H. 1951. On some pycnogonids from the sea around Kii Peninsula.— Publications of the Seto Marine Biological Laboratory 1(4):159-168, 2 figs. . 1971. Records of Pycnogonida from shallow waters of Japan.— Publications of the Seto Ma- rine Biological Laboratory 18(5):317-347. Wilson, E. B. 1881. Report on the Pycnogonida. Re- ports on the results of dredging ... Blake.— Bulletin of the Museum of Comparative Zool- ogy, Harvard 8(12):239-256, pls. I-V. (KN) Faculty of Education, Yokohama National University, 156 Tokiwadai, Ho- dogaya-ku, Yokohama 240, Kanagawa Pre- fecture, Japan; (CAC) Department of In- vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 311-335 PYCNOGONIDA OF THE WESTERN PACIFIC ISLANDS, VIII. RECENT COLLECTIONS FROM ISLANDS OF THE GREAT BARRIER REEF, AUSTRALIA C. Allan Child Abstract.—Eleven pycnogonid species were known from the Great Barrier Reef of Australia, from Torres Strait to the vicinity of Gladstone. Fifteen species are added to the known number with nine species previously known from other localities and six species new to science. These were taken from islands in the northern section (Lizard Island) and the southern section (Heron Island) of the Barrier Reef. The six new species are Ammothella prolixa, Tanystylum haswelli, Seguapallene crassa, Nymphon draconis, Anoplodactylus brucei, and Rhyn- chothorax vallatus. The new species are described and figured, the distribution of all species is included without conclusions concerning distribution patterns due to the paucity of this material and other records of Barrier Reef species, and remarks on species affinities are included. This report treats two small collections of about 100 specimens comprising fifteen named species plus two unnamed for lack of suitable material in eleven genera rep- resenting six families of the Pycnogonida. The collections were taken from two small island groups; Lizard Island (14°38’'S) in the northern section of the Barrier Reef, and Heron Island (23°27'S) at the southern end of the Reef, plus one species taken at Or- pheus Island approximately in the middle of the Reef archipelago. Collecting on Liz- ard and Heron Islands was greatly facilitat- ed by the presence of marine laboratories on both islands, while a resort on Orpheus Island made collecting there much easier. Among the fifteen named species report- ed herein from these islands are six new species; Ammothella prolixa, Tanystylum haswelli, Seguapallene crassa, Anoplodac- tylus brucei, Nymphon draconis, and Rhyn- chothorax vallatus. This high incidence of new species (40%) reflects the sparse amount of collecting done in these islands for such microorganisms. This is not the first report on pycnogonids from the Great Barrier Reef, although it is the first to treat Reef species exclusively. Haswell (1885) was the first to report on a species (Nymphopsis armatus) from the Queensland coast, although it was not strict- ly from a Barrier Reef locality. Carpenter (1892, 1893) was the next specialist to re- port on Reef species with four taken off Murray Island in Torres Strait. His species were; Parapallene australiensis (Hoek), P. haddoni Carpenter, Rhopalorhynchus cla- vipes Carpenter, and Ascorhynchus tenui- rostre Carpenter. His work was followed by that of Flynn (1918, 1919a, b, 1929) who redescribed the types of Haswell and described five other species from Australia, only four of which were taken in the Barrier Reef area. The new species described by Flynn (1929) from near the Barrier Reef are: Ascorhynchus melwar- di Flynn (near Cape York, northernmost Queensland), Nymphopsis armatus Haswell (Lindeman Island, Whitsunday Passage), Pallenopsis hoeki (Miers) (near Cape Y ork), and Parapallene famelica Flynn (Lindeman Island). 312 Other Australian species have been de- scribed and the distribution of known species extended, but since Flynn’s 1929 paper, no other Barrier Reef species were described until Stock (1954) listed Anoplodactylus longiceps Stock from Lindeman Island. Clark (1963) described Pycnogonum torresi Clark from Torres Strait on the northern- most Barrier Reef and listed Ascorhynchus minutum Hoek from a coral reef off Port Curtis at Gladstone. Staples (1982) described several species from the Calliope River at Gladstone, but there have been no reports of pycnogonids from the Barrier Reef itself since Clark’s paper reporting on the total Australian pyc- nogonid fauna known at that time. Staples also noted that there were at the time twen- ty-eight species known from Queensland, but only eleven of these were taken on or in close proximity to the Great Barrier Reef. This report adds fifteen species to the eleven species known for a total of twenty- six now known from the Reef. Only one of the previously known species from the Reef was taken in the collections represented in this report while nine species are known from the Australian mainland or from more distant localities and six are reported as new species. The most probable reason for this lack of similarity in collections is undoubt- edly due to differences in collecting meth- ods. While many specimens were previ- ously taken over the years fortuitously by trawl or other macrocollecting devices, the Lizard and Heron Island collections were made for their microorganism contents, thereby greatly enriching our knowledge of these tiny organisms of the Great Barrier Reef. The previously known species compris- ing most of this report have a tendency to be distributed in a northern array above the equator, suggesting only that there has been more collecting to the north than in most Australian and New Zealand waters. One species is known to be pantropical—temper- ate while two have a broad Indo-west Pa- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cific distribution. Another has only been taken before in New Zealand, while two are known from either one or both sides of Aus- tralia to Indonesia or the Philippines, and three are known only from the Philippines or from there plus other western Pacific lo- calities. One species has been taken previ- ously only in Japan and Korea, providing a very disjunct picture of its distribution. All are shallow water inhabitants. Very little can be inferred from these dis- tributional bits and pieces and no clear pic- tures of distribution can be taken from such small groups of species. The only clear pic- ture we get from the data in this report is that the Great Barrier Reef is apparently rich in species judging by this small sam- pling from two of the hundreds of islands. The presence of six previously undescribed species out of the fifteen in these collections suggests that very little collecting of minute forms has been done in these localities, and that intensive collecting over most of the length of the Barrier Reef might increase the number of new and known species by ten- fold. Systematics Family Ammotheidae Dohrn Genus Achelia Hodge Achelia assimilis (Haswell) Ammothea assimilis Haswell, 1884:1026- 1027, pl. LIV, figs. 5—9. Achelia (Ignavogriphus) assimilis.—Fry & Hedgpeth, 1969:106, figs. 152, 153, 156, tab. 13, 14 [literature]. Achelia assimilis. —Child, 1988b:2 Material examined. —Heron Island: southern reef flat, on alcyonarian, coll. A. J. Bruce, 16 Jan 1977 (1 juv); rubble on SW reef crest, coll. N. Bruce, 2 Jun 1978 (1 juv); rubble on reef flat, coll. N. Bruce, 6 Jun 1978 (2, sam), Lizard Island: off Casuarina Beach, coral with Halimeda and red algae clumps, 2 m, coll. B. Kensley, sta. K-L1, 27 May 1980 (2 VOLUME 103, NUMBER 2 juv); southern tip, coral rubble on patch reef, 1 m, sta. JDT/LIZ-3, 23 Jan 1989 (1 3, 1 juv); S of Lizard Head Peninsula, rubble in 2 m, sta. JDT/LIZ-14, 29 Jan 189 (6 4, 2); Lizard Head, rubble bank in 2 m, sta. JDT/ LIZ-15, 31 Jan 1989 (10 4, 2, juv); Lizard Head, small rubble in sand, 2 m, sta. JDT/ LIZ-19, 2 Feb 1989 (4 4, Q). Distribution. —Haswell’s long-known species has been taken in many southern hemisphere localities along with records from Indonesia and the Philippines. It has a mainly littoral to shallow depth range. Remarks. — There may possibly be more than one species hiding under the umbrella of this name. It is an extremely variable species with figures in the literature having almost no tubercles to specimens figured with extreme tuberculation. Genus Ammothella Verrill Ammothella prolixa, new species Fig. 1 Material examined. —Lizard Island: Watson’s Bay, SW border, reef and sand flat rubble in 1-7 m, sta. JDT/LIZ-7, 25 Jan 1989 (1 3, holotype, 1 2, 2 juv, paratypes). Other material. —Orpheus Island, just off Townsville: cove S of resort, rubble in 1-2 m, sta. JDT/OPH-1, 12 Feb 1989 (2 9). Lizard Island: North Point, rubble at bot- tom of cliff, 12 m, sta. JDOT/LIZ-13, 28 Jan 1989 (1 2); S of Lizard Head Peninsula, rub- ble in 2 m, sta. JDT/LIZ-14, 29 Jan 1989 (1 2, 1 juv); off Casuarina Beach, coral with Halimeda clumps and red algae, 2 m, sta. K-L1, 27 May 1980 (1 juv). Description. —Size very small, leg span 8 mm. Trunk and lateral processes slender, elongate, fully segmented, glabrous except for tiny paired or single dorsodistal tuber- cles on lateral processes. Lateral processes 2 to 2.5 times longer than their maximum diameters, constricted. proximally, swollen distally, separated by slightly more than twice their diameters distally. Neck mod- erately narrow, expanded distally, without 313 setae or tubercles. Ocular tubercle very slen- der proximally, expanded to club-shape dis- tally, slightly more than 4 times longer than maximum diameter, eyes fully distal, large, well pigmented. Abdomen very long, slen- der, longer than proboscis, erect, curving toward posterior, armed with median and distal fields of tubular and pointed spines. Median field with 6 tubular spines, 2 dorsal are longer than those lateral and ventral, and 2 shorter lateral pointed spines. Distal field with 2 long tubular spines and 2 shorter pointed spines, with 2 very short setae lat- eral to anus. Proboscis with extreme basal and distal constrictions and very swollen median sec- tion, originating from extended proximo- ventral pedestal on trunk anterior. Chelifores slender, slightly longer than proboscis, lightly spinose. First scape seg- ment only half length of second segment, first armed with single lateral seta distally, second with short proximal tubular spine, 2 median tubular spines and | long pointed spine, one distal tubular spine with 4 long pointed spines on inflated terminus of scape. Chela with atrophied movable finger on bulbous palm lacking immovable finger but with single long lateral seta. Palp very long, slender, segments only moderately armed with ventral setae. First segment not longer than wide, armed with single lateral seta longer than segment di- ameter. Second segment almost as long as fourth, the longest segment, second armed with 2 moderately long lateral setae. Fourth with 3 long lateral and distal setae, several shorter endal setae, and a lateral elevated pore at median length. Fifth and sixth seg- ments subequal, half length of fourth, armed moderately with short setae, some slightly longer than segment diameter. Seventh and eighth increasingly shorter, with few ventral setae, ninth longer, almost as long as fifth and sixth segments, armed with moderate number of short ventral and distal setae. Entire palp about 1.5 times longer than pro- boscis. 314 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Ammothella prolixa, new speices, holotype male: A, Trunk, dorsal view; B, Trunk, lateral view; C, Third leg, with enlargement of terminal segments; D, Oviger; E, Strigilis segments, enlarged. VOLUME 103, NUMBER 2 Oviger typical of genus, longest segments with | to several short lateral setae. Second and fourth segments longest, second almost as long as fourth. Strigilis weak, seventh seg- ment armed with 3 long ectal setae and 1 endal denticulate spine smaller than those of more distal segments. Eighth with 2 ectal setae and 1 endal denticulate spine, ninth without ectal setae but 1 endal denticulate spine. Tenth segment a tiny knob with 2 very broad denticulate spines having at least 8 serrations per side. Legs slender, well armed with tubular and pointed spines. First and second coxae with paired dorsolateral tubular spines, paired lateral long pointed spines, and fringe of short setae laterally and ventrally on second coxae. Third coxae without long spines but with several short ventral setae. First tibiae the longest segment, longer than second tib- iae which are longer than femorae. All ma- jor segments with 3-9 tubular spines ran- domly placed and 4—8 long pointed spines, those on dorsal tibiae up to 6 times longer than diameter of segment. Femoral cement gland tube placed dorsodistally, a robust cylinder almost twice femoral diameter. Tarsus very short, subtriangular, with 2 se- tae and | spine ventrally. Propodus mod- erately short, slightly curved, with 3 well separated heel spines, 6 small sole spines, and few dorsal and distal setae longer than segment diameter. Claw relatively short, only little over half propodal length, aux- iliary claws almost as long as main claw. Female: very similar to male but of slight- ly larger size. Oviger second and fourth seg- ments notably shorter than those of male. Sexual pores on all second coxae ventrodis- tally. Measurements (holotype, in mm).— Trunk length (chelifore insertion to tip 4th lateral processes), 0.85; trunk width (across 2nd lateral processes), 0.59; proboscis length, 0.53; abdomen length, 0.62; third leg, coxa 1, 0.12; coxa 2, 0.35; coxa 3, 0.31; femur, 0.74; tibia 1, 0.85; tibia 2, 0.8; tarsus, 0.09; propodus, 0.31; claw, 0.14. 315 Distribution. —The species is known from the type locality, Lizard Island, in the north- ern sector of the Barrier Reef, and from Or- pheus Island, in the middle section of the Reef, all in shallow depths to 12 m. Etymology. — This species is named (Lat- in: stretched out, long) for its elongate slen- der trunk and appendages. Remarks. — This species was at first glance reminiscent of Ammothella elegantula Stock, another slender attenuated species. Further examination brought many small differences to light, such as the very slender proboscis of A. elegantula, its abdomen without large spines, the lack of any tubular spines on Stock’s species, the long lateral spines of the lateral processes and the sim- ilar spines of the first two coxae, the broad neck, and the very short terminal palp seg- ments, all characters quite different from this new species. This species is another addition to the appendiculata—rugulosa group of species in this genus, species having tubular and pointed spines, long ocular tubercles, long curved abdomina with spine fields, and legs with similar segment length ratios and spi- nation. This species group is larger, with 18 species, than the group of 15 species without these spines. Of the 18 species having tu- bular spines, this species is closest to the known Pacific species; A. indica Stock, A. alcalai Child, and possibly A. pacifica Hil- ton. It can be compared least with A. pacifica of the three because Hilton’s species is much more compact with lateral process setae, shorter ocular tubercle and abdomen, and many more tubular spines on each major leg segment. The new species is very similar to the more slender A. indica, particularly in a comparison of the legs, ovigers, and palps of both species. The differences are found in the more widely separated lateral pro- cesses of the new species along with its shorter oviger segments, fewer tubular leg spines, lack of a tubular spine on the first scape segments, rounded ocular tubercle 316 apex, a shorter propodus, and fewer dentic- ulate spines on the female oviger, while hav- ing a denticulate spine on the eighth male oviger segment which is lacking on that of A. indica. This new species appears to be closest to A. alcalai, recently described from the Phil- ippines (Child 1988a:2—4, fig. 1). The lateral processes of the new species are placed far- ther apart than in A. alcalai (and are farther apart than those of all other species dis- cussed herein), but both species have small lateral process dorsodistal tubercles. The tu- bular and pointed spines of A. alcalai are predominantly much shorter than those placed in corresponding positions on A. pro- lixa, and the distal palp segments are no- tably shorter than those of the new species. The one character which separates this new species from almost all others of the appendiculata—rugulosa group is that of the anteroventral extension of the trunk which acts as a pedestal for the proboscis base. In most species, the proboscis originates di- rectly from the flat trunk anterior between and ventral to the palp and chelifore inser- tions, and there is no anterior trunk exten- sion or pedestal present. Ammothella stauromata Child Ammothella stauromata Child, 1982:271- 273, fig. 1; 1988a;5, 7; 1988c:809-810. Material examined. —Lizard Island: S of Lizard Head Peninsula, rubble in 2 m, sta. JDT/LIZ-14, 29 Jan 1989 (1 6, 2 9). Distribution. —This species is known from Enewetak Atoll, Marshall Islands, the northern and southern Philippines, and American Samoa, in depths of 0-3 m. This additional record further extends the broadening distribution known for this species to a more southern locality. Few specimens have been taken in any known locality, but A. stauromata appears to have a particularly wide western Pacific islands distribution. It has not yet been taken in the Indian Ocean. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Remarks. — The only other species of Am- mothella with large mediandorsal trunk tu- bercles found in Australia is A. thetidis Clark, a species with many striking differences from A. stauromata. Clark’s species lacks the me- diandorsal tubercle on the posterior rim of the first trunk segment, a tubercle which is present on this species. Clark’s species is apparently blind while this species has prominent eyes, the proboscis of Clark’s species is bulbous and without constrictions while that of this species has a modified tripartite shape similar to proboscides of many species of Ascorhynchus. This species has elongate palp segments in comparison with those of Clark’s, and there are large paired first coxae tubercles on this species while Clark’s is without these tubercles. There are a number of other small differ- ences between the two species. There are few other Ammothella species known with large dorsal tubercles, and this species shares even fewer characters with these other species than with A. thetidis. Genus Eurycyde Schioedte Eurycyde setosa Child Eurycyde setosa Child, 1988a:8—10, fig. 3. Material examined. —Lizard Island: S of Lizard Head Peninsula, rubble in 2 m, sta. JDT/LIZ-14, 29 Jan 1989 (1 4, 1 9). Distribution. —This species was recently described from a male collected in the northern Philippines at Batan Island, in O-— 4 m. This second and third specimen greatly extend its known distribution southerly to the northern Great Barrier Reef, but within the known depth range of the type. This is perhaps another of the growing list of species with known distribution along a corridor extending from Australia and New Zealand north to at least the Philippines and sometimes to Japan. There are undoubtedly many more species sharing this extremely rich corridor and their capture must await extended and close collecting on the Barrier Reef, New Guinea, and particularly in In- VOLUME 103, NUMBER 2 donesia and the Philippines where the ben- thic microfauna remains little known. Remarks.—These two specimens have only slight differences from the type speci- men, a male. The major difference is in the first coxae tubercles which in this male are slightly shorter and bear only very few tiny setae instead of the longer and heavily se- tose tubercles of the type. It should be noted that the coxae tubercles of the female are only slightly smaller than those of the male, although the female is a good bit larger in most measurements than the male. The abdomen of the type is missing, but can be described from the two specimens in hand. It is only slightly longer than the ocu- lar tubercle, is swollen distally, and armed with six long distal feathered spines, each longer than the abdomen itself, and placed in a dorsolateral fan arrangement with one slightly shorter feathered spine placed ven- trally and more proximal than the dorso- lateral fan. There are also two short lateral setae adjacent to the anus. The remaining characters agree very well with the type male. Genus Tanystylum Miers Tanystylum haswelli, new species Fig. 2 Material examined. — Lizard Island: Liz- ard Head, rubble bank in 2 m, sta. JDT/ LIZ-15, 31 Jan 1989 (1 4, holotype). Description. —Size very tiny, leg span 6 mm. Trunk subcircular in dorsal outline, slightly wider than long, unsegmented. Lat- eral processes contiguous, even squeezed to- gether. Ocular segment anterior rim ex- tending only slightly beyond lateral process rims, armed with single lateral setae, ocular tubercle slightly posterior to anterior rim of segment, little wider than tall, with small slender apical tubercle anteriorly and two low bumps posterolaterally. Eyes large, darkly pigmented. Proboscis widest at base, moderately in- flated, tapering gradually to cylindrical tip circling flat lips. Abdomen short, carried al- 317 most horizontally, extending only slightly beyond tips of fourth lateral processes, orig- inating from a low bulge between bases of third lateral processes, armed with distal fringe of six short setae. Chelifores single-segmented, tubercle-like, blunt, curved inward, carried at very ele- vated angle, each armed with 4—5 short dis- tal setae. Palp four-segmented, little longer than proboscis, originating with broad basal seg- ment only as long as its proximal diameter. Second segment consisting of coalesced sec- ond and third segments, the juncture evi- dent beneath integument, armed with 4—5 ectal setae and 2-3 ventrodistal setae, none as long as segment diameter. Third segment only little longer than its diameter, armed with 2-3 short ventrodistal setae. Fourth segment a curved cylinder about 3.5 times longer than its diameter, armed with many ventral and distal setae mostly longer than its diameter. Oviger fourth and fifth segments subequal in length, armed with few short recurved spines laterally. Second segment very slight- ly shorter, armed with 1-2 short lateral spines. Sixth and seventh segments short, their combined length not as long as fifth segment, armed with several stout recurved spines, in greater numbers on sixth segment, and 1-2 short distal setae on seventh seg- ment. Seventh lacks spinose distal apoph- ysis. Terminal three segments each shorter than last, eighth rounded, armed with 3 dis- tal setae longer than segment diameter, without spines. Ninth a short curved cyl- inder without spines, with single distal seta longer than segment diameter. Tenth seg- ment a tiny bud, broader than long, armed with 2 stout straight spines lacking any form of denticulation. Legs very stout, the major segments armed with dorsal bulges bearing several setae each. First coxae not as long as their diameters, armed with low laterodistal tubercle on an- terior surface bearing 2 short setae. Poste- rior surface armed with 1-4 short latero- 318 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Third leg, with enlargement of cement gland tube; D, Oviger; E, Strigilis segments, enlarged. distal setae. Second coxae with 2 laterodistal setae, third with few ventrodistal setae. Fe- mur with large ventromedian bulge bearing 1-2 short setae, a large dorsomedian bulge with 4—5 longer setae, and a fringe of 7-8 distal setae. Cement gland orifice a short tiny tube tapering to constricted tip. Femur slightly longer than tibiae which are equal in length. Tibiae armed with many short setae mostly on dorsal bulges. Tarsus very short, heavily setose ventrally. Propodus stout, only slightly curved, armed with 3 Tanystylum haswelli, new species, holotype male: A, Trunk, dorsal view; B, Trunk, lateral view; C, stout heel spines, about 5 shorter sole spines, many short flanking setae, and few short dorsal and distal setae. Claw broad, short, less than half propodal length, auxiliary claws more slender, almost as long as main claw. Female of the species unknown. Measurements (in mm).—Trunk length (chelifore insertion to tip 4th lateral pro- cesses), 0.6; trunk width (across 2nd lateral processes), 0.66; proboscis length, 0.35; ab- domen length (from bulge anterior), 0.28; VOLUME 103, NUMBER 2 third leg, coxa 1, 0.19; coxa 2, 0.26; coxa 3, 0.22; femur, 0.5; tibia 1, 0.44; tibia 2, 0.44; tarsus, 0.06; propodus, 0.39; claw, 0.16. Distribution. —The species is known only from its type locality, Lizard Island on the northern Great Barrier Reef, in 2 m. Etymology.—The new species is named for Dr. William A. Haswell who, following Hoek’s offshore records of Australian species taken by the Challenger, provided the first substantial knowledge of Australian pyc- nogonids. Remarks. — This specimen appeared to be T. orbiculare Wilson at first examination. Specimens identified as Wilson’s species have been found along the littoral of eastern Australia, although there is some question as to whether or not they are indeed this far flung species (Clark 1977:332). The unique type of Wilson’s 7. orbiculare (unfortu- naiely a female without ovigers—the male ovigers bear the diagnostic characters) was examined for purposes of comparison. A set of figures of the type is presented herein (Fig. 3), along with a male oviger from a speci- men taken near the type locality by Wilson himself. The type specimen of this species has not been figured adequately for many years. The new species differs from T. orbiculare mainly in the palps and abdomen. The ab- domen of Wilson’s species is notably longer and extends almost the length of the first coxae of the fourth leg pair, it has no basal bulge and is swollen in dorsal view rather than having a median constriction as in the new species. The lateral processes are larger in girth in relation to the trunk size in the new species, and the terminal palp segment is notably longer than that of Wilson’s species. The palp of 7. orbiculare has a vari- able number of segments, ranging some- times from four to as many as seven in some specimens. The large ventral bulge on each femur of the new species is not present on the femorae of 7. orbiculare, but most of the remaining characters of these two species are very similar. Both species have male 319 Ovigers without the characteristic seventh segment apophysis. This new species has some similarities to T. hooperi Clark (1977:325—327, figs. 20— 30), another species found on the New South Wales coast of Australia. The major differ- ence between these two species is that 7. hooperi has the male oviger bearing a sev- enth segment apophysis. The two species have very similar abdomina, proboscides, and legs, but the palps are quite different. Perhaps a species with greater similarities to the new species than 7. hooperi is T. bre- dini Child (1970:296—299, fig. 3) in which males of both species have ovigers without a seventh segment apophysis. There are a number of other similarities between these species including the proboscis, four-seg- mented palps, ovigers, legs, and ocular tu- bercle and abdomen. The differences lie in T. bredini having anterior and posterior lat- eral process tubercles, larger first coxae tu- bercles, a slightly more tapering proboscis, shorter auxiliary claws, and a much shorter terminal palp segment. It is possible that these differences might be reduced or elim- inated by examination of a large suite of specimens, but with only one male available of the new species, I will keep the Australian specimen separate from the Society Islands species until more Lizard Island specimens can be collected for examination. Family Callipallenidae Hilton Genus Callipallene Flynn Callipallene species indeterminate ?Callipallene emaciata subsp. Stock, 1954: 46-48, figs. 19h, 1, 20c-e. Material examined. —Heron Island: SW reef crest, rubble in 0-1 m, coll. N. Bruce, 2 Jun 1978 (1 8). Lizard Island: S of Lizard Head Penin- sula, rubble in 2 m, sta. JDT/LIZ-14, 29 Jan 1989 (1 9); Lizard Head, rubble bank in 2 m, sta. JDT/LIZ-15, 31 Jan 1989 (1 2); SW of Lizard Head, rubble zone in 1.5 m, sta. JDT/LIZ-17, 1 Feb 1989 (1 2); Lizard 320 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON N A, ) Gl eos, Fig. 3. Tanystylum orbiculare Wilson, holotype female: A, Trunk, dorsal view; B, Trunk, lateral view; C, Third leg, with ova indicated within. Nontype male: D, Oviger. Head, small rubble in sand, 2 m, sta. JDT/ LIZ-19, 2 Feb 1989 (2 9). Distribution. —In spite of the very shallow water in which these specimens were taken, they agree in almost all respects with Stock’s figures for his unnamed subspecies. His ma- terial came from 65 fathoms (119 m) off Three Kings Island, New Zealand. Remarks.—The figures given by Stock emphasize the statement he made (1954:46) VOLUME 103, NUMBER 2 that “the specimens have almost no striking features.” The above specimens also agree with this comment and I must therefore re- frain from naming them. Having only one male among five females does not constitute an adequate number of specimens to justify adding another new species in this highly variable genus. These specimens agree in the neck and trunk habitus figured by Stock (1954:47, fig. 20d), but have a few more long setae on the chelae and laterally on the legs. The propo- dus is almost exactly as that figured (fig. 191), and the ocular tubercle has the same small conical apex. The legs of the above specimens bear the low swellings of the fem- orae and first tibiae as in Stock’s fig. 20e. There is nothing unique about any of these characters among the many species in this genus to warrant erecting another species from this scant material. Genus Cheilopallene Stock Cheilopallene nodulosa Hong & Kim Cheilopallene nodulosa Hong & Kim, 1987: 153-155, fig. 12.—Nakamura & Child (1990). Material examined. —Lizard Island: North Point, rubble at bottom of cliff, 12 m, sta. JDT/LIZ-13, 28 Jan 1989 (2 9). Distribution. —Hong & Kim recently de- scribed this species from a female and a juvenile found on the west coast of Korea in floating Sargassum weed. Nakamura & Child (1990) list it among the fauna of the Japanese Islands, and it appears herein on the Great Barrier Reef as a third recorded locality. The above record at 12 m is the deepest recorded capture. This new record extends the distribution for this species greatly to the south and into the southern hemisphere, suggesting that it is another of the growing list of species inhabiting a north— south corridor of related localities from Australia through the western Pacific Is- lands and to Japan and Korea. Remarks.—The female figured by Hong 321 & Kim (1987:fig. 12) is apparently a sub- adult with incompletely developed ovigers. The above females appear to be fully adult and agree in almost all respects with the characters of the type except for the ovigers and the placement of several small tuber- cles. The ovigers of the females above have segment lengths in agreement with those of the type, but each segment from the fourth onward has a stout lateral spine on the dor- sodistal surface. The strigilis has fully den- ticulate spines in the formula 3:3:3:4, with several tiny teeth on both the ectal and endal surfaces of the terminal claw imparting a frayed or worn appearance. The denticulate spines each bear 3-4 lateral serrations per side and are broad and short. The peculiar lateral bulges or ‘“‘baloons”’ at the proximal margins of each scape are present on both these specimens, suggesting that they are a normal character of the anat- omy rather than an artifact. Hong & Kim did not figure the tiny setose tubercles over the insertion of each scape nor the similar tubercles of the first lateral processes (single) and the first coxae (double). These tubercles are relatively inconspicuous and could eas- ily be overlooked. Genus Seguapallene Pushkin Seguapallene crassa, new species Fig. 4 Material examined. — Lizard Island: Liz- ard Head, rubble bank in 2 m, sta. JDT/ LIZ-15, 31 Jan 1989 (1 6, holotype, 1 3, 2 9, paratypes). Other material. — Lizard Island: S of Liz- ard Head Peninsula, rubble in 2 m, sta. JDT/ LIZ-14, 29 Jan 1989 (1 6 with eggs, 1 6, 2 2); Lizard Head, small rubble in sand, 2 m, sta. JDT/LIZ-19, 2 Feb 1989 (1 6 with eggs, 1 °); Palfrey Island, near SW point, rubble in 1 m, sta. JDT/LIZ-21, 6 Feb 1989 (1 juv); Lizard Island, off Casuarina Beach, coral with Halimeda clumps and red algae in 2 m, sta. K-L1, 27 May 1980 (1 4 with eggs, 1 9). 322 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Seguapallene crassa, new species, holotype male: A, Trunk, dorsal view; B, Trunk, lateral view; C, Third leg; D, Chela; E, Oviger; F, Strigilis segments, enlarged. Paratype female: G, Oviger. VOLUME 103, NUMBER 2 Description. —Size very small, leg span about 6 mm. Trunk fully segmented, stout, lateral processes crowded, touching to slightly separated, each only as long as its diameter or slightly shorter, glabrous, su- ture line at base of each lateral process and neck prominent. Neck moderately short, flanked by extremely large oviger bases and extending to broad expansion at chelfore insertions. Ocular tubercle at neck midpoint over Oviger bases, about as tall as wide, eyes prominent, well pigmented, with large sen- sory papillae on horn-like tubercles at apex. Abdomen moderately short, carried at low oblique angle, inflated in dorsal view, armed with 6 short distal setae. Proboscis short, rotund, a broad cylinder proximally, tapering abruptly distally to very small oral surface. Lips flat, without setae or other adornment. Chelifores 2-segmented, inserting into raised crests or cowls on neck anterior. Scape robust, moderately setose with several endal and dorsodistal setae shorter than segment diameter. Chela palm triangular, armed with distal fringe of setae with greater numbers at junction of palm and immovable finger. Fingers moderately slender, curved only at tips, glabrous, with 12 block-shaped teeth on each finger. Oviger fourth segment slightly curved, armed with 2-3 short endal setae, segment about 0.6 as long as fifth. Fifth segment with 6-7 ectal setae and small distal apophysis with apical seta. Sixth segment very short, shorter than any strigilis segment, armed with 2 lateral setae. Strigilis segments each shorter than last, armed with dorsodistal seta each and endal denticulate spines in the formula 6:5:5:5, with a terminal claw hav- ing 7-8 endal serrations. Denticulate spines with 2—5 serrations per side. Legs stout, moderately setose, second tib- iae the longest segment, femorae longer than first tibiae, terminal leg segments small in relation to proximal leg segments. First cox- ae with 2 short laterodistal setae, second 323 coxae with 2 long lateral setae per side and fringe of shorter ventrodistal setae, third coxae with few ventral and ventrodistal se- tae. Femorae with fringe of short setae around the median length, several distal se- tae with single longer dorsodistal seta, ce- ment gland pores not evident. First tibiae with 2 long proximolateral setae, a fringe of short median and distal setae, with long proximal, median, and distal setae on dorsal surface. Second tibiae with several dorsal and dorsolateral setae, those along median line longest, several short lateral setae, and few ventrodistal setae. Tarsus very short, semitriangular, with 3—4 ventral setae and 1 spine. Propodus very small, slightly curved, with 2 heel spines, 4—5 slender sole spines, and many distal and dorsodistal se- tae longer than segment diameter. Claw ro- bust, short, less than half propodal length, slightly curved, auxiliaries more slender, al- most as long as main claw. Female slightly larger in most measure- ments, oviger segments four and five much shorter than those of male, fifth shorter than fourth and without distal apophysis. Oviger strigilis spine formula: 6:5:4:5, with 9 ter- minal claw serrations. Major oviger seg- ments without setae or recurved spines. Neck equal in length to that of male. Measurements of holotype (in mm).— Trunk length (chelifore insertion to tip 4th lateral processes), 0.7; trunk width (across lst lateral processes), 0.46; proboscis length, 0.26; abdomen length, 0.16; third leg, coxa 1, 0.18; coxa 2, 0.32; coxa 3, 0.15; femur, 0.54; tibia 1, 0.5; tibia 2, 0.64; tarsus, 0.07; propodus, 0.27; claw, 0.12. Distribution. —This species is known from its type locality, Lizard Island, from a rub- ble bank at Lizard Head, in 2 m, and from other Lizard Island localities. Etymology.—The specific name (Latin: thick, stout) refers to the stout trunk with its closely crowded lateral processes. Remarks. —There were only two species known in this genus, one of the many genera 324 in this cumbersome family. These are S. insignatus Pushkin, the type of the genus, and S. micronesica Child. The distribution of these species is extremely disjunct with Pushkin’s species found in the subantarctic Crozet Islands, and S. micronesica found in the tropical Palau Islands. This third species is another from the tropics and is closest to S. micronesica. Pushkin’s species has lateral process tu- bercles and an oviger with shorter fourth and fifth segments, the latter lacking a distal apophysis, both characters which are dif- ferent in the two tropical species. The new species differs from S. micro- nesica in having much more compacted lat- eral processes which are more robust, short- er leg segments which are also rather robust, a smaller propodus with a more typical claw rather than the short one of S. micronesica, and a chela with block-like closely set teeth rather than well separated triangular teeth. The new species is about 0.3 smaller than the Palauan species, but both have very sim- ilar ocular tubercles, abdomina, probos- cides, chelifores, and sometimes prominent suture lines on the trunk. Family Nymphonidae Wilson Genus Nymphon Fabricius Nymphon draconis, new species Fig. 5 Material examined. —Lizard Island: S of Lizard Head Peninsula, rubble in 2 m, sta. JDT/LIZ-14, 29 Jan 1989 (1 6, holotype, 3 6 with eggs, 1 juv, | larva, paratypes). Other material.—Lizard Island: Lizard Head, rubble bank in 2 m, sta. JDT/LIZ- 15, 31 Jan 1989 (1 2); SW of Lizard Head, rubble zone in 1.5 m, sta. JDT/LIZ-17, 1 Feb 1989 (1 2); Lizard Head, small rubble PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON in sand, 2 m, sta. JDT/LIZ-19, 2 Feb 1989 (1 6, 2 2, 3 juv); off Casuarina Beach, rubble with encrusting and red algal turf in 2 m, sta. K-L2, 27 May 1980 (1 juv). Description. —Size moderately small, leg span 11.4 mm. Trunk fully segmented, seg- ments fairly short, bulbous, not longer than wide, lateral processes short, slightly longer than wide, separated by more than their di- ameters in anterior pairs, less than their di- ameters in posterior pairs, glabrous. Neck short, without parallel sides, oviger implan- tation bulges just anterior to first lateral pro- cesses, occupying half neck length, neck armed with short seta over insertion of each chelifore. Ocular tubercle low, only slightly taller than basal width, eyes large, anterior pair larger than posterior pair, ocular tu- bercle capped with pair of short lateral sen- sory tubercles. Abdomen very short, semi- erect, a tapering truncate cone armed with pair of short lateral setae. Proboscis barrel-shaped, widest swelling at midpoint, tapering distally to cylinder, lips almost flat. Chelifores very large, chelae massive, globular, moderately setose. Scape as long as proboscis, slightly swollen distally, armed with few short dorsal and lateral setae. Che- la very swollen, palm ovoid, much larger than short fingers. Fingers armed with closely crowded unevenly bifurcate teeth, 11 on each finger, fingers carried at sharp lateral angle to palm. Palp segments relatively short, second segment longest, third and fifth subequal in length, first and fourth little longer than their diameters. Second segment with few short lateral and distal setae, third with many ventral setae longer than segment diameter, fourth and fifth with many long ventral and lateral setae, some about 3 segment diam- = Fig. 5. Nymphon draconis, new species, holotype male: A, Trunk, dorsal view; B, Trunk, lateral view; C, Chela, with enlargement of four teeth; D, Terminal leg segments, enlarged; E, Femur, showing cement gland and tubes; F, Oviger with egg attached; G, Strigilis segments, enlarged, with proximal and distal denticulate spines further enlarged. Paratype female: H, Trunk anterior, dorsal view. VOLUME 103, NUMBER 2 325 326 eters in length. Terminal segment slightly curved. Oviger fifth segment longest, swollen dis- tally, without apophysis, fourth segment about 0.6 length of fifth. Fourth armed with few distal setae, 2 longer than segment di- ameter. Fifth armed with many setae in- creasing in numbers and length distally. Sixth segment with many long setae mostly endal, with greater numbers distally. Stri- gilis segments increasingly shorter with ter- minal segment shortest, armed with 2 to many long ectal setae, some twice as long as segment diameter, and endal denticulate spines in the formula 10:10:10:7, with ter- minal claw only slightly shorter than tenth segment. Claw with few tiny distal setules, without teeth. Denticulate spines with pair of wider lateral serrations proximally and many smaller distal serrations, distal spines of each segment longer than proximal spines. Legs moderately slender, with few dorsal setae, more ventral setae, second tibiae 0.3 longer than first tibiae, femorae slightly shorter than first tibiae. Femorae with 10- 11 conspicuous ventral cement gland tubes, each slender, less than half femoral diam- eter in length, placed in row along 0.7 of femoral length. Each major segment with 3 long distal setae. Tarsus cylindrical, distally inflated, about half propodal length, armed with 4—5 long ectal setae and many endal setae along with 2 slender spines. Propodus slender, very slightly curved, armed with 7— 8 slender sole spines alternating with 2-3 sole setae between spines, without heel or heel spines, but with row of short lateral setae and several long dorsal and distal se- tae, the longest about 3 times segment di- ameter. Claw very reduced, broad, very curved, without endal setules. Auxiliaries about 0.3 longer than main claw, well curved distally, armed with few tiny endal setules proximally. Female slightly larger in most measure- ments than male, neck equal in length to male’s. Chelae not inflated, palm a curved PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cylinder, fingers slightly longer than those of male and armed with 12-13 unevenly bifurcate teeth. Oviger fourth and fifth seg- ments shorter, fifth not clubbed distally, none of the segments with the long setae of the male oviger, but rather few short setae. Measurements of holotype (in mm).— Trunk length (chelifore insertion to tip 4th lateral processes), 1.23; trunk width (across 2nd lateral processes), 0.55; proboscis length, 0.56; abdomen length, 0.15; third leg, coxa 1, 0.17; coxa 2, 0.49; coxa 3, 0.22; femur, 1.04; tibia 1, 1.4; tibia 2, 1.66; tarsus, 0.21; propodus, 0.4; claw, 0.07; auxiliaries, 0.11. Distribution. — This species is known from its type locality, south of Lizard Head Pen- insula on Lizard Island in 2 m, and from other Lizard Island localities in 1.5—2 m. Etymology.—The species name (Latin: draconis, a dragon-like lizard) is a play on the name Lizard Island, its type locality. Remarks. —This new species is the elev- enth reported of the NV. aequidigitatus group discussed recently by Child (1988b:67—68). It fits well in the key of the ten known species at couplet 8 with N. megacheles Child, and couplet 9 with N. biformidens Stock and N. aequidigitatus Haswell, the three species with which it is most closely allied. Mem- bers of the aequidigitatus group have, be- sides the diagnostic characters of the genus, another set of characters more or less in common which involve chelae teeth usually being bifurcate, a longer than usual terminal palp segment, often very inflated chelae in the males, an oviger claw lacking the usual endal teeth, and auxiliary claws often longer than the short main claw with any or all of these claws bearing endal setules or rugos- ities. Some members of the group may be without one or more of these characters. This new species differs from N. mega- cheles in having a slightly longer trunk with the lateral processes wider apart as a con- sequence, somewhat shorter legs with more setae, and a much longer tarsus in relation to propodal length and a much shorter main VOLUME 103, NUMBER 2 claw. The tarsus of N. megacheles is ex- tremely short and both it and the propodus have robust heel spines. The type of N. me- gacheles is listed as a female, but the large inflated chelae and long setose fifth oviger segment suggest that it is actually a male. As far as is known, only males have the very inflated chelae in this group while those of the female are the more typical semicylin- drical shape. The strigilis terminal claw dif- fers between these two species in that the new species claw has few endal setules while that of N. megacheles has an endal lamina. None of the species of this group have the conspicuous cement gland tubes of the new species. There are many similarities between this new species and N. biformidens, including those of neck and trunk habitus, leg segment length ratios, tarsus and propodus relative lengths, main and auxiliary claw lengths, and palp and chelifore similarities. The dif- ferences are in a shorter ocular tubercle lack- ing the lateral “horns” of N. draconis, the slender strigilis claw and shorter strigilis segments (those of a female specimen were compared with Stock’s species figures which are also of a female), the very different che- lae teeth which are all bifurcate in the new species while not all are bifurcate in N. bi- formidens, and the very different oviger denticulate spines which, in the new species, are more like those of the genus Callipallene in having many tiny distal serrations. The denticulate spines of Stock’s species have 7-8 larger lateral serrations per side. The new species has fewer characters in common with Haswell’s N. aequidigitatus. It has longer chelae fingers in relation to palm length and the fingers bear many more teeth. It also has slightly longer leg segments with far fewer setae, particularly the longer setae, a third palp segment longer than the terminal segment, again also lacking the long setae of the new species, and a very different oviger which lacks most of the long setae present on the oviger of the new species. 327 Males of several of the aequidigitatus group are unknown, but where they are known, none display the conspicuous fem- oral cement gland tubes of this new species. Family Phoxichilidiidae Sars Genus Anoplodactylus Wilson Anoplodatylus batangensis (Helfer) Pycnosoma batangense Helfer, 1938:174— 176, fig. 6a—c. Anoplodactylus batangensis. —Stock, 1968: 54 [early literature].—Child, 1988a:14 [later literature]. Material examined. —Lizard Island: be- tween Palfrey and South Islands, dead coral rubble with algal turf in 3 m, sta. K-L11, 30 May 1980 (1 6 with eggs, 1 2); S of Lizard Head Peninsula, rubble in 2 m, sta. JDT/ LIZ-14, 29 Jan 1989 (3 4, 1 2); SW of Lizard Head, rubble zone in 1.5 m, sta. JDT/LIZ- 17, 1 Feb 1989 (1 9). Distribution.—This species has a pan- tropical distribution in littoral and shallow depths. The above records mark the first time it has been taken in Australian waters. Anoplodacatylus brucei, new species Fig. 6 Material examined. — Heron Island: rub- ble from 6.2 m, sta. HI/26/5 coll. A. J. Bruce, 28 Apr 1978 (1 4, holotype). Description. —Size moderately small, leg span 14.6 mm. Trunk lightly segmented, first segmentation line fully encircling trunk, second lighter, difficult to discern laterally, third only a hint, not encircling trunk. An- terior pair of lateral processes longest, each succeeding posterior pair shorter to last pair which are wider than long. Lateral processes separated by their maximum diameters or slightly less, armed with short dorso- and laterodistal setae numbering 4 on anterior pair, 3 on middle pairs, and | on posterior pair. Neck fairly short, flanked by lateral expansions or “wings” bearing rudimentary 328 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Anoplodactylus brucei, new species, holotype male: A, Trunk, dorsal view; B, Trunk, lateral view; C, Third leg, with cement gland and tube, enlarged; D, Chela; E, Oviger; F, Terminal two strigilis segments, enlarged. palp buds which point laterally rather than little taller than wide, with broad rounded anteriorly. Neck armed with single seta per apical cone. Eyes large, anterior pair slightly side. Ocular tubercle moderately large with larger than posterior pair, darkly pigment- broad base occupying most of neck dorsally, ed, occupying most of ocular tubercle at its VOLUME 103, NUMBER 2 midpoint. Abdomen rather short, erect, swollen from constricted base, tip tapering, armed with 6-7 short lateral and dorsal se- tae. Proboscis a cylinder curving anteriorly, slightly swollen distally, lips rounded. Chelifores moderately short, scape not as long as proboscis, with slight distal infla- tion, armed with few dorsodistal and lateral setae. Chelae long, slender, palm a cylinder twice as long as its diameter, armed with few setae encircling finger bases. Movable finger longer than immovable finger, well curved, overlapping immovable finger at tip, armed with 4 ectal setae longer than finger diameter. Immovable finger not as curved, about equal in length to palm. Fingers with- out teeth. Ovigers moderately slender, rather se- tose. Third segment longest, almost twice length of second segment, with proximal constriction, both armed with few short lat- eral setae. Fourth segment with several short ectal setae, fifth and sixth equal in combined length to fourth, fifth with many recurved setae, sixth with pad of 8—9 similar setae on one side only, segment about 0.3 as long as fifth segment. Legs lightly setose, moderately long. Sec- ond coxae of third and fourth pair with long ventrodistal sexual tubercle equal in length to segment diameter, armed with many short setae. Femur the longest segment, with short dorsodistal tubercle half as long as segment diameter, armed with seta as long as seg- ment diameter. Cement gland a long tube slightly over half femoral diameter, arising from low bulge at midpoint of segment. Tib- iae subequal in length, with short dorsal, lateral, and ventral setae and single longer dorsodistal seta. Tarsus very short, sub- triangular, with several short ventral setae. Propodus robust, moderately long, with well marked heel bearing 2 broad spines, the dis- tal one larger than proximal spine, 3 distal stout setae, and 9-10 short sole spines with very short lamina at claw base. Claw stout, 329 curved at tip, 0.6 as long as propodus, with tiny curved auxiliaries at base. Female of the species unknown. Measurements (in mm).—Trunk length (chelifore insertion to tip 4th lateral pro- cesses), 1.87; trunk width (across Ist lateral processes), 1.06; proboscis length, 0.89; ab- domen length, 0.35; third leg, coxa 1, 0.29; coxa 2, 0.83; coxa 3, 0.4; femur, 1.39; tibia 1, 1.3; tibia 2, 1.31; tarsus, 0.16; propodus, 0.66; claw, 0.46. Distribution. —The new species is known only from Heron Island, its type locality, in 6.2 m. Etymology.—I take pleasure in naming this species for Alexander J. (Sandy) Bruce, sometime collecting partner, expedition cabin mate, collector of this specimen, pro- lific author and systematist of pontoniid shrimp of the world, and Director of the Natural Sciences section of the Northern Territories Museum of Arts and Sciences, Darwin, Australia. Remarks. —This new species has little to differentiate it from many others in this ge- nus and there are several species quite sim- ilar to this one among the Anoplodactylus legions. Similar species are A. digitatus (Bohm) and its cousin A. paradigitatus Child, A. erectus Cole, A. californicus Hall, and A. allotrius Child. All have some characters agreeing with those of the new species. Most of the similarities are found in the legs as all share a long sexual tubercle on the second coxae of the posterior legs, a single cement gland tube, sometimes a dorsodistal tuber- cle on the femorae, and other similarities of the propodus. The legs of A. erectus are almost exactly like those of the new species except that Cole’s species does not have the small dor- sodistal tubercle on the femur. One of Cole’s paratype specimens was examined for com- parison and this specimen has small slender dorsodistal lateral process tubercles which are lacking in the new species. Many spec- imens of A. erectus from Panama do not 330 have these tubercles or else have low bumps in their place, so this character is inconsis- tent and not a good diagnostic feature. The lateral processes of A. erectus are notably longer than those of the new species, as is the third oviger segment and the ocular tu- bercle. The laterally projecting palp buds of the new species are apparently unique as other species with this character have the buds pointing toward the anterior of the specimen. Both A. digitatus and A. paradigitatus have legs with very similar characters to those of A. brucei, except that the second coxae sex- ual tubercles are not as long as those of the new species, the lateral processes are longer in the two species, and the proboscis is of a very different shape with females of the two species. Females of these species have small alar processes on the proximoventral surfaces of their proboscides. Unfortu- nately, a female of the new species is not available for comparison. Neither of the other species have palp buds suggesting any similarity to those of A. brucei. Hall’s species (with the recently synony- mized A. portus Calman) also has very sim- ilar legs, trunk habitus, chelae, and ocular tubercle, but the chelifore scapes are much shorter, the lateral processes are much clos- er together, the oviger terminal segment is much shorter and has quite different seta- tion, and the femoral cement gland tube is a truncate cone rather than a slender tube as in A. brucei. The new species has even less similarity to A. allotrius, but has legs which agree in most characters. The sexual tubercle is much shorter than in the new species and the lat- eral processes are closer together, the chelae have longer fingers with several teeth on the movable finger, the scapes are shorter, there are low lateral process tubercles, and the Oviger terminal segments (fifth and sixth) are shorter and with fewer setae than the same segments of the new species. Although A. brucei has no outstanding character to set it off from all others, I be- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lieve that the shades of difference in the several characters discussed are sufficient to separate this species from all others known in the genus. Anoplodactylus chamorrus Child Anoplodacatylus chamorrus Child, 1983: 705-707, fig. 3; 1988a:16. Material examined. —Lizard Island: be- tween Palfrey and South Islands, dead coral rubble with algal turf in 3 m, sta. K-L11, 30 May 1980 (1 6 with eggs, | 4, 2 2); same locality, coral rubble with algal turf on reef crest in 0.3 m, sta. K-L12, 31 May 1980 (1 6 with eggs, 3 2); S of Lizard Head Peninsula, rubble in 2 m, sta. JDT/LIZ-14, 29 Jan 1989 (1 8). Distribution. —This species was recently described from specimens taken in Guam, and it has more recently been taken in sev- eral localities in the southern Philippines. These records greatly extend its range to the south but add nothing to the known depth range. Remarks. —The lateral processes are closely crowded in this small compact species and there are few other such com- pact species known in the Pacific. Such com- pact species of this genus were previously grouped under the genus Halosoma, now synonymized under Anoplodactylus. The only Australian species known with this compact character is A. haswelli (Flynn). The Guamian species differs from Flynn’s by having broad lateral ““wings” between the neck and first lateral processes, small lateral process dorsodistal tubercles, a conspicuous cement gland tube on the femorae, and sev- eral other smaller differences. The species differs from other more northerly distributed compact species such as A. crassus Nakamura & Child, A. viri- dintestinalis (Cole), and perhaps A. mono- trema Stock, in having the lateral neck wings mentioned above. Some compact species have conspicuous oviger bulges of the male in this location, but few have these broad VOLUME 103, NUMBER 2 anterior wings with tiny palp vestiges along their first lateral process margins. Anoplodactylus glandulifer Stock Anoplodactylus glandulifer Stock, 1954:80— 84, fig. 36.—Child, 1982:273-274 [liter- ature]; 1988b:58-59; 1988c:813. Material examined. —Lizard Island: North Point, rubble at bottom of cliff, 12 m, sta. JDT/LIZ-13, 28 Jan 1989 (1 2); Liz- ard Head, small rubble in sand, 2 m, sta. JDT/LIZ-19, 2 Feb 1989 (1 4, 1 question- able larva). Distribution. —This species has been tak- en in many localities from the Red Sea and Indian Ocean to as far east as the Samoa Islands. These records mark its first capture on the Great Barrier Reef. It is known from 0-12 m. Remarks. —The conspicuous cement gland cups number from 2 to 4 per femur, and serve as a good recognition character in males in a genus where the cement gland is much more often served by a single orifice per femur. Females, of course, are difficult to distinguish as a particular species from other similar species without accompanying males, but both sexes in this species have very long propodal lamina which sets them off from those many species with shorter lamina. The female from station 13 is only provisionally placed in this species due to lack of an accompanying male. This female propodus agrees well with 4. glandulifer as does the ocular tubercle and lateral process placement, and is little different from other females examined of this species. Anoplodactylus longiceps Stock Anoplodactylus longicollis Williams, 1941: 36-38, figs. 2-5 (preoccupied). Anoplodactylus longiceps Stock, 1951:16 [footnote]; 1954:83 [text]; 1956:97-98, fig. 14c, d.—Clark, 1963:56 [key].—Child, 1975:20, fig. 9f. Material examined. —Lizard Island: North Point, rubble at bottom of cliffin 12 331 m, sta. JDT/LIZ-13, 28 Jan 1989 (1 dam- aged 2); S of Lizard Head Peninsula, rubble in 2 m, sta. JDT/LIZ-14, 29 Jan 1989 (1 2). Distribution. — This species is known from the east and west coasts of Australia in trop- ical to temperate waters, and from the Kei Islands in eastern Indonesia in depths of 28- 134 m. These records establish the species at Lizard Island on the Barrier Reef and in much shallower water at 2 and 12 m. Remarks. —This very long slender species is easily recognized with its very widely sep- arated lateral processes, tall ocular tubercle with a long slender apical cone, slim legs with a long femoral dorsodistal tubercle and two low cement gland cups, and its mod- erately short proboscis. This species is reminiscent of another very elongate species, 4. attenuatus Child, from the Philippines. The Philippine species has lateral processes spaced even wider than in this species and also lacks the femoral dor- sodistal tubercle, has a shorter oviger, has a crenulate major heel spine on the propo- dus, lacks the usual tiny auxiliary claws, and has a very different ocular tubercle which is shorter than its width. Both species, though, have a widely spaced pair of cement gland cups on each femur. They are probably quite closely related, having possibly originated from a single parent stock. Anoplodactylus species indeterminate Material examined.—Lizard Island: off Casuarina Beach, coral with Halimeda clumps and red algae in 2 m, sta. K-L1, 27 May 1980 (1 larva); between Palfrey and South Islands, dead coral rubble with algal turf in 3 m, sta. K-L11, 30 May 1980 (1 larva); point between Anchor Bay and Wat- son’s Bay, coral rubble in 2 m, sta. JDT/ LIZ-5, 24 Jan 1989 (1 9); Lizard Head, small rubble in sand, 2 m, sta. JDT/LIZ-19, 2 Feb 1989 (1 juv). Remarks. —The juvenile and larvae are too immature for determination and the fe- male is unlike other species in these collec- 332 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Rhynchothorax vallatus, new species, holotype female: A, Trunk, dorsal view; B, Trunk, lateral view; C, First leg; D, First leg terminal segments, enlarged; E, Oviger, greatly enlarged. tions and shows little similarity to published figures of other species from Australia. Genus Endeis Philippi, 1843 Endeis biserata Stock Endeis biserata Stock, 1968:57-60, fig. 21; 1970:1; 1974:17; 1979:28-30, fig. 9.— Child, 1988a:20. Phoxichilus meridionalis Loman (non Bohm, 1879), 1908:78-79. Material examined. —Lizard Island: Watson’s Bay, SW border, reef and sand flat rubble in 1-7 m, sta. JDT/LIZ-7, 25 Jan 1989 (1 2, 1 juv); S of Lizard Head Penin- sula, rubble in 2 m, sta. JDT/LIZ-14, 29 Jan 1989 (1 8). Distribution. —This is a wide-ranging species having been taken from Brazil, the Red Sea, Indian Ocean, Indonesia, Hawaii, and the Philippines in 0-37 m. These rec- VOLUME 103, NUMBER 2 ords place it to the south of its nearest known capture in Indonesia, on the northern Great Barrier Reef and well within its known depth range. Remarks.—The male from station 14 lacks the long dorsodistal tubercles of the femorae, but has a moderately long and very robust spine in its place. This femoral spine is flanked by two shorter broad spines. Most of the small spines of the trunk and legs are short but very broad, a character not found or at least not remarked on by Stock in his description. The male is otherwise very like the type and other specimens figured in Stock’s fig. 21 (1968:57). The leg length ra- tios, the propodal configuration, trunk habi- tus, oviger, and the leg lateral spines are all very like those shown in the figures. The female has the dorsodistal femoral tuber- cles, but those of the above specimen are slightly longer than that shown in Stock’s fig. 21e. Family Rhynchothoracidae Thompson Genus Rhynchothorax Costa Rhynchothorax vallatus, new species Fig. 7 Material examined. — Lizard Island: S of Lizard Head Peninsula, rubble in 2 m, sta. JDT/LIZ-14, 29 Jan 1989 (1 2, holotype). Description. —Size very tiny, leg span only 3.2 mm across second lateral processes. Trunk fully segmented, covered with tiny granular papillae as are appendages, with greater numbers of papillae on extremities. Trunk moderately slender, without other adornment. Lateral processes very short, not as long as their diameters, widely separated, tubercular, armed with short dorsomedian seta each. First lateral processes with pair of posterior tubercles as broad as their length, second processes with similar pair of pos- terior tubercles and smaller anterior tuber- cle at lateral process bases. Third lateral processes with small anterior tubercle and smaller posterior tubercle, fourth with only small anterior tubercle. Ocular tubercle and 33313. eyes lacking, with pair of broad, low, an- terior-pointing tubercles in their place. Neck moderately narrow, flanked by short cylin- drical tubercles as bases for palps, each bearing low ectal tubercle. Abdomen short, with broad base, tapering to small rounded anus, extending to less than half second cox- al lengths on fourth legs. Proboscis typical for genus, with 2 paired dorsolateral swellings, constricted at base and distally at lips, carried horizontally, without labial fringe or lamina on anti- meres. Palps 4-segmented, with prominent pa- pillae. First segment cylindrical, as long as combined length of terminal 3 segments, armed with few distal setae. Second segment short, cylindrical, not as long as twice its diameter. Third segment slightly more than half length of first, armed with few short setae and dorsal tubercle slightly longer than segment diameter, placed slightly distal to dorsal midpoint of segment. Fourth seg- ment short, little longer than second, re- curved dorsally to height of third segment tubercle, armed with few short ectal setae. Oviger also typical, tiny, fourth and sixth segments subequal. Seventh segment swol- len distally, armed with 2 short broad endal spines, eighth and ninth segments shorter, subequal in length, armed with single small endal spine each. Tenth segment broader than long, armed with ectal seta, tiny endal basal spine, curved endal lamina, and broad curved claw which closes onto lamina. Legs of approximately equal size except for second coxae on fourth leg pair which are 0.3 shorter than coxae 2 of other legs. Second coxae of anterior 3 pairs twice length of first coxae, femorae the longest segments with first tibiae longer than second. Three major segments armed with dorsodistal se- tae longer than segment diameters and sev- eral short setae increasing in numbers on distal leg segments. Tarsus very short, with single sole seta. Propodus short, inflated, slightly curved, armed with 4—5 sole setae, several short lateral setae, and several dorsal 334 setae almost as long as propodal diameter. Claw short, broad, well curved, auxiliaries present, short, slender, less than half main claw length. Male of species unknown. Measurements (in mm).—Trunk length (proboscis insertion to tip 4th lateral pro- cesses), 0.91; trunk width (across Ist lateral processes), 0.44; proboscis length, 0.57; ab- domen length, 0.19; first leg, coxa 1, 0.1; coxa 2, 0.16; coxa 3, 0.09; femur, 0.26; tibia 1, 0.25; tibia 2, 0.23; tarsus, 0.04; propodus, 0.18; claw, 0.05. Distribution. — This species is known from its type locality only, south of Lizard Head Peninsula, Lizard Island, in 2 m. Etymology. —The name proposed for this species is a play on the Great Barrier Reef (Latin: surrounded by a wall or rampart) which surrounds Lizard Island with a ram- part of coral. Remarks.—This new species has a few characters close to the juvenile type of R. malaccensis Stock (1968:20-22, fig. 6), a species known from the Straits of Malacca, Indonesia. The similarities are mostly found in the proboscis and trunk. The lateral pro- cesses are similar, but in Stock’s species they are placed much closer together. The pro- boscides have the dorsolateral bulges in both species and are of similar lengths. The new species is closest to R. philop- sammum Hedgpeth (1951:111-115, fig. 3) from the west coast of North America. Here again, the lateral processes of Hedgpeth’s species are similar to the new species, in- cluding the tubercles size and placement, but the lateral processes themselves are placed very close together rather than well separated as in the new species. There are similarities in the appendages of both species, but the third palp segment is shorter in the new species than in Hedgpeth’s species. Also in the palp, the first and third segments (second and fourth segments ac- cording to his text, mistaking the palp bases as segments instead of lateral extensions of the trunk anterior) are subequal in Hedg- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON peth’s species while this does not hold true for R. vallatus. The tubercle of the third palp segment is much more distally placed in R. philopsammum than in the new species. The main and auxiliary claws differ in lengths between the two species, being much shorter in R. vallatus, and the femur is long- er than the tibiae in the new species, but femur and first tibia are equal in length in Hedgpeth’s species. Finally, the first coxae of R. philopsammum have several small dorsodistal tubercles, particularly on the second and third pairs, while the first coxae of this new species have a single low tubercle only on the anterior surface of the first three leg pairs. The fact that these two species are sepa- rated in their distribution by the breadth of the Pacific Ocean is not in itself a valid rea- son to separate these species, but this fact combined with the various subtle differ- ences discussed above should serve to maintain the two as separate species. Acknowledgments I wish to thank the collectors; J. D. Thom- as, B. F. Kensley, and J. Clark for their spec- imens from Lizard and Orpheus Islands, and A. J. (Sandy) Bruce and Niel Bruce, his son, for the Heron Island collections. Holotype specimens are deposited in the Australian Museum, Sydney, while other specimens are deposited in the National Museum of Nat- ural History, Smithsonian Institution, un- der catalog numbers of the old U.S. Na- tional Museum. Literature Cited For the literature published prior to 1978, see the complete bibliography published by Fry & Stock 1978. A Pycnogonid Bibliography, in Sea Spiders (Pycno- gonida).— Zoological Journal of the Linnean Society of London, 63(1 +2):197—238. Child, C. A. 1982. Pycnogonida of the Western Pa- cific Islands, I. The Marshall Islands. —Pro- ceedings of the Biological Society of Washington 95:270-281, figs. 1-3. 1983. Pycnogonida of the Western Pacific VOLUME 103, NUMBER 2 Islands, II. Guam and the Palau Islands. — Pro- ceedings of the Biological Society of Washington 96:698-714, figs. 1-5. 1988a. Pycnogonida of the Western Pacific Islands, III. Recent Smithsonian—Philippine ex- peditions.—Smithsonian Contributions to Zo- ology 468:1—32, figs. 1-12. —. 1988b. Pycnogonida from Aldabra Atoll.— Bulletin of the Biological Society of Washington 8:45-78, figs. 1-9. 1988c. Pycnogonida of the Western Pacific Islands, V. A Collection by the Kakuyo Maru from Samoa. — Proceedings of the Biological So- ciety of Washington 101:809-816, figs. 1-2. Clark, W. C. 1977. The genus Tanystylum Miers, 1879 (Pycnogonida) in the southern oceans. — Journal of the Royal Society of New Zealand 7(3):313-338, figs. 1-67. 335 Hong, J.-S., I.-H. Kim. 1987. Korean pycnogonids chiefly based on the collections of the Korea Ocean Research and Development Institute. — The Korean Journal of Systematic Zoology 3(2): 137-164, figs. 1-17. Nakamura, K., & C. A. Child. 1990. Pycnogonida of waters adjacent to Japan.—Smithsonian Con- tributions to Zoology (in press). Stock, J. H. 1979. Pycnogonida from the mediolit- toral and infralittoral zones in the tropical west- ern Atlantic.— Studies on the Fauna of Curacao and other Caribbean Islands 59(184): 1-32, figs. 1-9. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 336-340 CANTHOCAMPTUS (ELAPHOIDELLA) STRIBLINGI, NEW SPECIES (COPEPODA: HARPACTICOIDA) FROM COSTA RICA Janet W. Reid Abstract. —The male of Canthocamptus s. |. (Elaphoidella) striblingi, a new species of harpacticoid copepod from the Monteverde Cloud Forest Preserve, Costa Rica, is distinguished within its subgenus by the biarticulate endopods of legs 1 and 4; the exopod of leg 5 and the endopod of leg 2 each having 4 setae; and the unmodified spines of the exopod of leg 4. The female is unknown. In a collection of aquatic invertebrates from bromeliads in Costa Rica made by Dr. James B. Stribling, there occurred a single male specimen of a previously undescribed species of harpacticoid copepod belonging to the large cosmopolitan genus Cantho- camptus Westwood, 1836, subgenus Ela- phoidella Chappuis, 1928. Order Harpacticoida Sars, 1903 Family Canthocamptidae Sars, 1906 Genus Canthocamptus Westwood, 1836 Subgenus Elaphoidella Chappuis, 1928 Canthocamptus (Elaphoidella) striblingi, new species Figs. 1-15 Type material.—1 6, from water con- tained in narrow-leaved epiphytic brome- liad in Monteverde Cloud Forest Preserve, Costa Rica, altitude about 750 m, 27 Aug 1988, dissected on one slide (USNM 243304), deposited in the National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. Male. —Body (Figs. 1—4) cylindrical; length, excluding caudal setae, in lactic acid 0.64 mm. Cephalosome with oblong nuchal organ. Posterior hyaline frills of all somites smooth; urosomites 3-5 each with single row of uniform, slender spinules on lateral and ventral posterior margins, each spinule row continuous ventrally. Anal somite with row of 4 spinules on each side and ventral row of 10 spinules above each caudal ramus; anal operculum with 4 large, somewhat ir- regular teeth. Caudal ramus 2 times longer than broad, with little-developed dorsal ridge ending in dorsal seta inserted just an- terior to insertion of posterior lateral seta; anterior lateral seta slightly longer than pos- terior lateral seta. Small innermost terminal seta of ramus about half length of ramus. Inner margin of ramus without ornament. Rostrum (Fig. 2) projecting in lateral view, with rounded apex bearing 2 subapical ros- tral filaments. Antennule (Fig. 5) about half length of cephalosome, geniculate, with es- thetascs on articles 4 and 8. Antenna (bro- ken in dissection) with allobasis; uniarti- culate exopod (Fig. 6) bearing 4 setae. Mandible (Fig. 7) with uniarticulate palp bearing 3 terminal setae. Maxillule (Fig. 8) with precoxal arthrite having 4 teeth and 3 setae; coxa with | stout seta bent at mid- length and | slender seta; basis with | stout seta bent at midlength and 4 slender setae. Maxilla (Fig. 9) with 2 endites each with 3 terminal setae; basis with claw and 2 setae. Maxilliped (Fig. 10) prehensile; basis with- out seta; endopod with comb of 4 or 5 spi- nules on anterior and posterior surface, plus few spinules on dorsal surface. Swimming legs 1-4 (Figs. 11-14) each with exopod of 3 articles and endopod of 2 articles, except endopod of leg 3 consisting VOLUME 103, NUMBER 2 337 > =a =~ ( [ A IS Gala Figs. 1-10. Canthocamptus (Elaphoidella) striblingi, new species, male: 1, Habitus, dorsal; 2, Habitus, lateral; 3, Posterior urosomites and caudal rami, ventral; 4, Anal somite and caudal rami, dorsal; 5, Antennule, some setae not drawn; 6, Exopod of antenna; 7, Mandible; 8, Maxillule; 9, Maxilla; 10, Maxilliped. Scale for Figs. 1 and 2, 200 um; remaining figures not to same scale. of 3 articles of which article 2 bears apoph- Leg 2 _ basis 0-1 exopod 0-1; 1-1; 1,2,2 ysis. Setation formula for major armament endopod 0-0; 2,2,0 as follows: Leg 3 basis O-1 exopod 0-1; 1-1; 2,2,2 endopod 0-0; 1-0; 0,2,0 Leg 1 basis 1-1 exopod 0-1; 0-1; 0,2,2 Leg 4 basis 0-1 exopod O-1; 1-1; 2,2,2 endopod 1-0; 1,2,0 endopod 0-0; 0,2,0 338 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 11-15. Leg 4; 15, Leg 5. (Not to same scale.) Leg 5 (Figs. 2, 15) with exopod bearing 4 setae; 2 innermost setae coarsely plumed, 2 outermost setae naked. Basipods of both fifth legs fused, lacking armament. Leg 6 (Fig. 2) represented only by simple, slightly elevat- ed plate. Female. —Unknown. Etymology. —The new species is named in honor of its collector, Dr. James B. Strib- ling. Remarks.—Hamond (1987 [1988]) re- turned 18 superspecific taxa within the fam- ily Canthocamptidae, including Elaphoi- della Chappuis, to the status of subgenera of the genus Canthocamptus Westwood, pending eventual revision of the family. Ap- proximately 140 species are presently as- Canthocamptus (Elaphoidella) striblingi, new species, male: 11, Leg 1; 12, Leg 2; 13, Leg 3; 14, signed to the genus (now subgenus) E/a- phoidella Chappuis, which was recently revised and split into four genera by Apos- tolov (1985). Two of Apostolov’s proposed new genera correspond to Lang’s (1948) Elaphoidella Group X, which included species having biarticulate endopods of leg 1; Apostolov’s proposed Elaphoidellopsis is further distinguished primarily by having biarticulate endopods of leg 4. There are 18 previously known species and subspecies in this group; Apostolov’s inclusion of Ela- pDhoidella sewelli Chappuis, 1928 and its subspecies in Elaphoidellopsis is inexplica- ble, since these have leg 1 endopods of 3 articles and therefore fall in Apostolov’s Elaphoidella. VOLUME 103, NUMBER 2 The new Costa Rican species falls within Elaphoidellopsis Apostolov. However, Ha- mond (1987 [1988]) argued convincingly for retention of most canthocamptids within the broad category Canthocamptus until the morphology of most species, particularly type-species of the various superspecific taxa, can be described by contemporary standards. Apostolov (1985) partly rede- scribed Canthocamptus elaphoides Chap- puis, 1923, type species of Elaphoidella Chappuis; unfortunately, C. elaphoides, ac- cording to Apostolov, is extremely poly- morphic and the taxon may even represent a collective species! Apostolov resolved the problem of great variability in setation of swimming legs, and in other characters, of many species of Elaphoidella by proposing a simple scheme relying primarily on the number of articles of the swimming legs. However, he failed to present any argument as to why this arrangement might represent a more natural grouping than, for example, Lang’s (1948) ten-group division. Aposto- lov’s brief diagnoses of his proposed new genera contain mutual inconsistencies, omissions, and errors, and are seriously in- complete; for instance, his entire diagnosis of the male of Elaphoidellopsis consists of pointing out the ““marked sexual dimor- phism” in legs 3-5. It remains to be seen whether these proposed subgroups will be sustained after examination of the subgenus Elaphoidella by modern systematic meth- ods. On the other hand, Apostolov has fur- nished a valuable service in providing a list of species of this very large group. His keys contain a number of errors, such as the placement of E. sewelli, mentioned above; and some apparent typographical mistakes. However, the keys are more or less me- chanically useful in species discrimination, as I have employed one in the following discussion, without wishing to imply accep- tance of these proposed groupings in a sys- tematic sense. In Apostolov’s key to Elaphoidellopsis, Canthocamptus striblingi keys to E. (now 339 Canthocamptus) siolii (Kiefer, 1967); but C. striblingi is easily distinguished by having a longer caudal ramus without spinules on the inner margin; the ramus of C. siolii is only slightly longer than broad and has a group of spinules on the distal part of the inner margin. Kiefer also reported only 3 setae on leg 2 endopod 2, but his figure (Kiefer 1967: fig. 25) seems to show a socket on the outer distal corner of that article. The 3 large oper- cular teeth of C. siolii also may not be a reliable distinguishing character, since the number of opercular teeth may vary within a species, and Kiefer described C. siolii from only two male specimens. Males of the three other neotropical species in this group all have 2 modified spines on leg 4 exopod 3, only 3 setae on leg 5 exopod, and differ oth- erwise in setation of swimming legs. These are C. crenobia (Petkovski, 1973), C. einslei (Petkovski, 1973), and C. subcrenobia (Pet- kovski, 1980), all from Cuba. C. siolii is known only from the upper Rio Negro, an affluent of the Amazon River, Brazil. Acknowledgments The University of Wisconsin (Madison) is thanked for funding Dr. J. B. Stribling’s field work in Costa Rica. A Smithsonian Senior Postdoctoral Fellowship supported the preparation of this article. Several anon- ymous reviewers made valuable suggestions for improvement of the manuscript. Literature Cited Apostolov, A. 1985. Etude sur quelques copépodes harpacticoides du genre Elaphoidella Chappuis, 1929 de Bulgarie avec une révision du genre. — Acta Musei Macedonici Scientiarum Natural- ium 17(7/145):133-167. Chappuis, P. A. 1928. Zur Kenntnis der Mikrofauna von Britisch Indien. III. Copepoda Harpacti- coida.—Records of the Indian Museum 30(4): 375-385. Hamond, R. 1987(1988). Non-marine harpacticoid copepods of Australia. I. Canthocamptidae of the genus Canthocamptus Westwood s. lat. and Fibulacamptus, gen. nov., and including the de- scription of a related new species of Cantho- 340 camptus from New Caledonia.—Invertebrate Taxonomy 1(8):1023-1247. — Kiefer, F. 1967. Neue Copepoda Harpacticoida aus dem Amazonasgebiet.—Crustaceana 13(1):114— 122. Lang, K. 1948. Monographie der Harpacticiden. Nordiska Bokhandeln, Stockholm, 1682 pp. Petkovski, T. K. 1973. Subterrane Siisswasser-Har- pacticoida von Kuba (Vorlaufige Mitteilung). — Résultats des Expéditions Biospéologiques Cu- bano-Roumaines a Cuba 1:125-141. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1980. Finf neue Elaphoidella-Arten (Cope- poda, Harpacticoida) aus den subterranen Ge- wassern von Kuba.—Acta Musei Macedonici Scientiarum Naturalium 16(2/135):33-70. Department of Invertebrate Zoology, NHB-163, National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C. 20560. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 341-349 TACHIDIUS INCISTIPES KLIE AND OTHER HARPACTICOIDS FROM NORTHWESTERN CANADA (CRUSTACEA: COPEPODA) Edward B. Reed Abstract.—Twelve species of harpacticoid copepods are reported from the shores of fresh and brackish lakes and ponds in northern Northwest Territories and Yukon Territory. Attheyella ussuriensis Rylov is a new record for North America and Tachidius incisipes Klie a new record for Canada. The Southern Party of the Canadian Arc- tic Expedition 1913-1918 collected marine and freshwater Copepoda from localities between Vancouver Island, B.C. and Coro- nation Gulf, N.W.T. (Johansen 1922). Marsh (1920) and Johansen (1922) com- mented on the importance of Canadian Arc- tic Expedition collections as the first from a region that was little known in regard to micro-crustacean fauna. In their reports neither Johansen nor Marsh mentioned harpacticoid copepods, although both dis- cussed species of calanoid and cyclopoid co- pepods. Willey (1920) recorded eight known species of harpacticoids and described an additional three as new to science; most were marine species but at least one was found in fresh-to-brackish water. Records for har- pacticoids have lagged behind those for calanoids and cyclopoids although several investigations have added to the knowledge of freshwater copepods in the northern reaches of Yukon and Northwest Territories since the Canadian Arctic Expedition. The purpose of this paper is to furnish specific site records for several species of harpacti- coids; two of which may be the first for Ca- nadian waters. Participation in an excursion to Yellow- knife and Inuvik, N.W.T. following the XIXth Congress of the International As- sociation of Limnology (SIL) in 1974, af- forded me the opportunity to collect along margins of lakes and ponds. Specimens were examined and dissected in glycerin or lactic acid (Humes & Gooding 1964). Insofar as possible, examination and drawings were made without cover slips. Species were determined primarily by the key of Wilson & Yeatman (1959). Prelim- inary identifications were confirmed by comparing specimens with the descriptions found in Gurney (1932), Lang (1948), Bo- rutzky (1952), and Dussart (1967). Waters Sampled Because the harpacticoid collections were opportunistic grab samples, there are few data on the waters sampled. Frame and Grace Lakes (Table 1) occupy rock basins with emergent vegetation in protected areas. Grace Lake is about 63 ha in surface and 9 m in mean depth (Brunskill, pers. comm. 1974). The Fort Franklin sample came from a pond about 0.1 ha in area and 20 cm deep in a muskeg at the edge of the village. This pond and the roadside ditch on the Hay River road at Yellowknife contained brown water and much vegetation. The Tuktoyaktuk ponds are beach ponds in the sense of Johansen (1922), that is, they are only a little above sea level and situated quite close to the seashore but are not di- rectly connected to the sea. These ponds undoubtedly receive Arctic Ocean water during onshore storm surge, as indicated by the presence of stranded driftwood. Meijer- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 342 x x puod M.SToLET “N,9S089 “LA ‘JWiog 9surys x xX puod yorog ‘9 xX x puod yorog “q x x x x x puod yovog ‘8 MZ ET “Ni L769 ynqyesovyn AeMYSIP{ 19}SdUloq UO YI0ID x yTeqdwieg Jesu puod [jews MPHoEET “Ni 1789 “LAN Anu xX Xx OBeT[IA IeoU puUOd [[euIS M.S7oETT “NZ 10S9 “UMN ‘ulpjuely 1104 ayTUyMOT[E A JO X JSOMYLIOU WY L[ “BO PROT IOATY AeyY UO YOUP spIspeoy x dYVT IPIL) xX xX dYeT suiely MALTS TE “N,0€079 “LN ‘9sTMOTS A SS SS ods (1681 (€681 6761 7061 9681 7E6I (ZO61 (F681 r981 7881 €16l $9}BUIPI0OD AqjeooT pApoiq § ‘preyory “PIZeI) “JJarY “NOL ‘1095 “AOTAY ‘Z10q = ‘JIawYydS) ‘yxos0g “qYyo01q IID -opnasd 7 pie dys 1uos DQZDIM “WY WL, Sisuatans —-afvT) suaidpa — Sadiuids “SII sad1siou1 “ds -yourlg) -aoplaa ‘g -u1yoiny W 1a1yinp -n “Vy 11pjo1ys DAyo DAIONUN Sadiasip snipiyov I, Duosou pau snjduvo DIUDIOW -uapiou —- -oSay OF -12aJDFT = - uD youl -0dg pjjad snjduivo -OYyuy -0yrtuE ee EE ‘pL6| ‘SOWOMLLIT UOYNA pue IsoMYWON Woy pojda][09 spodsadoo proonoediey Jo satsedg “] 31QeL VOLUME 103, NUMBER 2 Table 2. Comparison of three species of Tachidius. T. discipes 22 antennule 7 articles antenna exp 2 setae seg. 3, P1-4 5,6,6,5 spines & setae seg. 3 enp P-4 5,5,5,5 P5 9 setae and spines anal operculum 12 or more spinules 36 spinules on chitinous ridge on antennule article | antenna exp 2 setae P2 enp article 2 modified; article 3 with apical setae P5 5 to 8 usually 7 setae and spines anal operculum with spinules ing (1975) found electrical conductivity of 4776 and 2370 uwSiemens in water from two Tuktoyaktuk ponds. Meijering and I likely sampled different ponds, although both sampled the same day. Shingle Point is also on the coast of the Arctic Ocean; however, the pond sampled here lies on top of a low bluff above the beach and probably does not receive storm surge seawater. Species Identified Nine of 16 samples contained 12 species of harpacticoid copepods (Table 1). In the following accounts, N.W.T. and Y.T. forms that fell within descriptions provided by the above manuals are listed without comment. Tachidiidae Several males and females plainly refer- able to the genus JZachidius were collected from Tuktoyaktuk beach ponds (Table 1). The females were clearly 7. incisipes on the basis of antennule of 9 articles, setae and spine formula of P1—P4 and U-shaped canal in the genital segment (Table 2). The presence of a modified P2 indicated that the males were not incisipes. The close 343 T. incisipes T. spitzbergensis 9 articles 7 articles at least 5 setae 3 setae 5,6,5,5 5,6,5,5 6,6,6,6 5,5,5,5 9 8 several small spinules smooth genital field with prominent U-shaped canal spinules lacking spinules present at least 5 setae 3? P2 not modified P2 modified less than 9, usually 7 6 spinules spinules similarities of males of discipes and spitz- bergensis are evident from Table 2. Olofs- son (1917) noted the morphological close- ness of male discipes and spitzbergensis and distinguished between them on the number of setae and spines on PS. Given the vari- ability in the armament of P5 in discipes, (Lang 1948, Dussart 1967) this does not seem a reliable characteristic. Lang (1948) synonomized spitzbergensis with discipes; however Wilson and Yeatman (1959) re- tained spitzbergensis in their key, using se- tation of P5 to separate the males and ab- sence of spinules on the anal operculum of spitzbergensis to separate females. On the basis of setation of P5, the Tuktoyaktuk males were assigned to 7. discipes. Tachidius incisipes Klie, 1913 Seven females ranged in length from 0.75 to 0.82 mm. Metasome slightly flattened, widest at second thoracic segment. Rostrum small, pointed. Nuchal organ and lateral discs oval and discs present on all thoracic segments (Fig. 1). Genital field with U-shaped canal leading from genital pore to seminal receptacle (Fig. 2). Anal oper- culum with many spinules on free margin, row of fine spinules extending from oper- 344 culum base down each side. Caudal rami longer than broad (Fig. 3). Oblique row of spinules on dorsal and mesial surfaces, two dorsal setae set close together, outer caudal seta spine-like and about twice length of inner seta. Both medial setae jointed and having straight-sided basal sections which constrict abruptly, longer about 2.5 times the length of the shorter. Antennule of 9 segments (Fig. 4). Antenna of 4 segments, exopodite 2-segmented with a total of 5 se- tae (Fig. 5). Mandible with biramus palp, blade with one large blunt tooth and smaller teeth (Fig. 6). Outer surface with a process which Gurney (1932, fig. 403) depicted as a bump is clearly hook-like when seen in side view. Maxillule of two segments (Fig. 7) and maxilla of 3 endites, coxa-basis and an exopod, although segmentation not al- ways distinct (Fig. 8). Maxilliped 3-seg- mented (Fig. 9), with a terminal claw. Both rami of legs 1-4 of three articles, basopodites with prominent mounds dense- ly fringed with long hairs, outer mound ap- pearing to project between rami when legs are viewed anteriorly. Pl basipod spines large, article 3 of exopod with three spines and three setae, spines of exopod smooth (Fig. 10). P2 lateral spines of exopod articles 1 and 2, smooth; article 3, one marginal spine smooth and one toothed (Fig. 11), an- terior face of endopod article 3 with two rows of spinules. P3 with prominent blade- like spinules on outer margin of exopodite segments and and article 2 of endopod, en- dopod article 3 with a total of six spines and setae (Fig. 12). P4 with rows of spinules on anterior face of all exopodite articles, en- dopod article 3 with a total of five spines and setae (Fig. 13). The numbers and ar- rangement of spines and setae agree with those given by Lang (1948:282). PS a single Figs. 1-15. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON plate with a total of eight spines and setae (Fig. 2). Tachidius discipes Giesbrecht, 1882 The Tuktoyaktuk males were compatible with the specific characters as given by Oloffson (1917), Gurney (1932), Lang (1948), and Wilson & Yeatman (1959) in three regards: modified endopodite P2, an- tennule, and P5. The second segment of P2 endopodite of Tuktoyaktuk animals has a stout projection which overlaps a notch on segment three and bears two setae that are set close together at its inner distal angle (Fig. 14). The first article of the antennule enlarged and bearing a rounded prominence, armed with small marginal spinules (Fig. 15). The inner lobe of P5 with 3 spines and the outer, 2 spines and 2 setae (Fig. 16). The occurrence of a sample with two species of Tachidius each represented by one sex only may not be as strange as it appears at first view. Olofsson (1918) reported 7. longicornis (= T. incisipes) and T. spitzber- gensis occurring together in several samples from four lakes on Spitsbergen. Moreover, the numbers of each sex were often quite unequal. Seventeen individuals, all female, of 7. brevicornis (= T. discipes) were found in one sample from a creek mouth at Ber- nard Harbour, N.W.T. (Willey 1920). Atteyella (Mrazekiella) ussuriensis Rylov, 1932 Rylov (1932) described Attheyella ussuri- ensis from females collected in small pud- dle-like ponds on the shore of the Ussuri River near Khabarovsk (135°5’E, 48°30'N). Rylov did not find males and the species seems not to have been reported since the => Female Tachidius incisipes: 1, Habitus. 2, Genital field and PS. 3, Caudal ramus and anal operculum. 4, Antennule, setation omitted. 5, Antenna. 6, Mandible blade. 7, Maxillule. 8, Maxilla. 9, Maxilliped. 10, Pl. Feathering on setae omitted. 11, P2. 12, P3. 13, P4. Scale D: Fig. 1. Scale A: Figs. 2-13. 346 original description, since both Lang (1948) and Borutzky (1952) used Rylov’s figures. One female was found in a collection from a pond at Campbell Creek near Inuvik. This female was readily identified in the keys of Lang and Borutzky and agreed with Rylov’s description in regard to rami, enlarged base of caudal seta 2 (Fig. 17), form, size and number of teeth on the anal operculum (Fig. 17) and spines on P5 (Fig. 18). Rylov figured the spines of the basal article of PS without sockets. This was also the case with the Campbell Creek female. Borutzky noted that the characteristic structure of PS differen- tiates ussuriensis from all other species of Attheyella. Moraria duthiei T. & A. Scott, 1896 One female was taken from a pond at Shingle Point (Table 1). Ramus with a dor- sal chitinous ridge which terminated pos- teriorly in a sharp point (Fig. 19); anal oper- culum roughly triangular with a blunt point and eroded margin (Fig. 19); PS exopodite jointed, feathering on setae sparse, some smooth (Fig. 20). This individual had a pe- culiar knob on the outer of the middle cau- dal setae (Fig. 19). Bryocamptus (Bryocamptus) hutchinsoni Kiefer, 1929 A few specimens were collected from a roadside ditch near Yellowknife. These an- imals were like the typical form in possess- ing non-bifid spinules on the anal opercu- lum and in characteristics of the ramus as given by Kiefer (1929) and Wilson & Yeat- man (1959). A form in which the spinules are bifid occurs widely in Alaska, western Figs. 16-27. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Canada and western United States (Wilson & Yeatman 1959). Bryocamputus (Bryocamptus) vejdovskyi (Mrazek, 1893) Five males were collected at Fort Frank- lin and one male at Shingle Point (Table 1). B. vejdovskyi is a member of the mor- phologically variable minutus group (Wil- son 1956). In addition to sexual dimor- phism there is geographical variation. Typical females possess only one well de- veloped caudal seta, non-bifid anal oper- culum spines, the ramus terminates in a spi- nous process on its outer distal corner; males possess 3 well developed caudal seta, lack spinous process, and endopod P4 article 2 with 4 setae (Wilson & Yeatman 1959). Wil- ley (1925) described as Canthocamptus minusculus a form in which the female lacked the spinous process on the outer, dis- tal margin of the ramus. Willey had only four females, some of which had bifid oper- culum spines and some, simple spines. Wil- ley believed minusculus to be the North American form of Canthocamptus vejdoy- skyi Mrazek (Willey 1925:157). Kiefer (1934) gave the name Bryocamptus vejdov- skyi forma minutiformis to specimens with bifid anal operculum spinules, but which were otherwise typical B. vejdovskyi (Wilson & Yeatman 1959). The Shingle Point male appeared to be typical B. vejdovskyi and had simple oper- culum spines (Fig. 21). This male possessed large teeth-like spines on the ventral surface of the last abdominal segment at the base of the caudal ramus; these spines were not mentioned by Lang (1948) or Wilson & — Male Tachidius discipes: 14, Endopod P2. 15, Antennule. 16, P5 and P6. Female Attheyella ussuriensis: 17, Caudal ramus and anal operculum. 18, P5. Female Moraria duthiei: 19, Caudal ramus and operculum. 20, P5. Male Bryocamptus vejdovskyi: 21, Caudal ramus and anal operculum. 22, P2. 23, P3. 24, P4. 25, Female Halectinosoma, Pseudobradya sp.? 26, Mandible blade. 27, P5. Scale A: Figs. 14-18. Scale B: Figs 19 and 20. Scale C: Figs. 26 and 17. 347 VOLUME 103, NUMBER 2 348 Yeatman (1959). The Fort Franklin males differed from the Shingle Point male only in having bifid operculum spinules, resem- bling B. v. forma minutiformis in this char- acter. The large spines on the outer margins of exopodites of P2 (Fig. 22), P3 (Fig. 23) and the terminal article of P4 (Fig. 24) were completely smooth in the Shingle Point male. The terminal article of P2 endopodite (Fig. 22) and P5 of the Shingle Point male were similar to figures of these appendages in Lang (1948) except that the two spines on the inner lobe of P5 appeared relatively shorter and stouter (Fig. 25). Laophontidae Onchyocamptus mohammed (Blanchard & Richard, 1891) Several males and females were collected from pondsa, b, and c, Tuktoyaktuk. Willey (1923) described as a new species, Lao- phonte calamorum from Lake St. John, Quebec. Lang (1948) regarded L. calamo- rum as a synonym of O. mohammed. Ectinosomatidae Halectinosoma? The Tuktoyaktuk pond sample contained two female harpacticoids, 0.59 and 0.66 mm in length, that were clearly referable to Ec- tinosomatidae. In addition to familial and generic fea- tures, the Shingle Point animals possessed these features: markedly conical caudal rami, length 1.5 and 1.8 times the width at base; abdominal segment of larger female densely covered both dorsally and ventrally with very tiny spinules; abdominal segment of smaller appeared covered with tiny pits. Hyaline lappets which project from the rear margin of the last abdominal segment over the bases of the caudal setae in many ecti- nosomatids could not be seen on either fe- male. Both possessed a hyaline pseudo- operculum with smooth margin. The larger PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON female had an egg sac containing 6-8 fairly large ova. Lang (1965) stated that the mandibles re- veal useful specific characters in the genera Ectinosoma and Halectinosoma. The cut- ting edge of the mandibles of the Tuktoyak- tuk females had a heavily sclerotized bi- dentate pars incisiva and six-dentate lacina (Fig. 26). All marginal setae of PS stout and spine-like, the longest reaching to posterior margin of genital segment; surface setae in- conspicuous; two rows of small spinules on anterior surface of basopodite (Fig. 27). The maxillae were destroyed during dis- section; the structure of this appendage would have permitted distinctions between Halectinosoma and Pseudobradya. At least four species of Ectinosomatidae have been reported from coastal waters of N.W.T. and northern Alaska. The Canadian Arctic Expedition collected Pseudobradya minor (T. & A. Scott, 1896) and Ectinosoma neglectum Sars, 1904 at Bernard Harbour on Dolphin and Union Straits, N.W.T., and E. finmarchicum from Camden Bay, Col- linson Point, Alaska (Willey 1920). Wilson (1973) identified Pseudobradya major (Olofsson, 1917) from Nuwuk Lake, a land- locked brackish water pond, Point Barrow, Alaska. On the basis of characters of P5, caudal rami, antenna and other features, the Tuk- toyaktuk females did not seem to conform to descriptions of ectinosomatids reported from northwestern coastal waters. Too little is known about these ectinosomatids to de- cide if they represent undescribed species; this must await more specimens. Discussion These records of Bryocamptus vejdovskyi extend the ranges of forms with bifid and simple opercular spines but do not clarify their relationship. The harpacticoids taken at Tuktoyaktuk are members of genera with well-recognized euryhaline distributions (Lang 1948, Wil- VOLUME 103, NUMBER 2 son & Yeatman 1959). Bryocamptus hutch- insoni appears to be known from North America only; otherwise, with the possible exception of Attheyella ussuriensis, the har- pacticoids found in N.W.T. and Y.T. sam- ples are species of wide geographical distri- bution particularly in the Palearctic and Nearctic Regions. Some species such as Tachidius incisipes and Moraria duthiei seem mostly northern in occurrence; others such as Onychocamptus mohammed and Nitocra spinipes extend into Africa and Asia Minor. Ackowledgments I extend special thanks to Dr. Janet W. Reid, National Museum of Natural History, Smithsonian Institution, for advice, useful comments on a draft manuscript and for providing pieces of literature from the C. B. Wilson copepod library. Drs. C. J. Brunskill and M. C. Healey organized and ably lead the post-congress excursion. Two anony- mous reviewers provided helpful com- ments. Literature Cited Borutzky, E. V. 1952. Fauna of U.S.S.R. Crustacea. Vol. 3. no. 4. Freshwater Harpacticoida. Zoo- logical Institute, Academy of Sciences, Moscow, Israel Program of Scientific Translation, 1964. Dussart, Bernard. 1967. Les Copepodes Des Eaux Continentales D’Europe Occidentale. Tome I: Calanoides et Harpacticoides. N. Boubee et Cie, Paris. Gurney, Robert. 1932. British fresh-water Copepoda. Vol. 2. Ray Society, London, 336 pp. Humes, A. G., & R. U. Gooding. 1964. A method for studying the external anatomy of cope- pods. —Crustaceana 6:238-240. Johansen, Frits. 1922. The crustacean life of some arctic lagoons, lakes and ponds. Report of the Canadian Arctic Expedition 1913-18. Vol. VII. Part N. F. A. Acland, Ottawa, 25 pp. Kiefer, Friedrich. 1929. Zur Kenntnis der freileben- den Copepoden Nordamerikas. — Zoologischer Anzeiger 86:97-100. . 1934. Neue Ruderfusskrebse aus Nordamer- ika.— Zoologischer Anzeiger 108:206-297. Lang, Karl. 1948. Monographie der Harpacticiden. Hakan Ohlsson, Lund, 1682 pp. . 1965. Copepoda Harpacticoida from the Cal- 349 ifornia Pacific coast.—Kungliga Svenaka, Ve- tenskoppsahademiens Handlingar Series 4, 10(2): 1-560 + 6 pls. Marsh, C. D. 1920. Freshwater Copepoda. Report of the Canadian Arctic Expedition 1913-18. Vol. VIII. Part J. J de Labroquerie Tache, Ottawa, 25 pp. Meijering, M. P. D. 1975. Notes on the systematics and ecology of Daphnia pulex Leydig in north- ern Canada.—Internationale Revue der Ge- samten Hydrobiologie und Hydrographie 60: 691-703. Olofsson, Ossian. 1917. Beitrag zur Kenntnis der Harpacticiden—Familien Ectinosomidae, Can- thocamptidae (Gen. Maraenobiotus) und Tachi- diidae nebst Beschreibungen einiger neuen und wenig bekannten, arktischen Brackwasser-und Susswasser-Arten.— Zoologiska Bidrag fran Uppsala 6:1-39 + 7 pls. 1918. Studien tiber die Susswasserfauna Spitzbergens. — Zoologiska Bidrag fran Uppsala 6:183-648. Rylov, V. M. 1932. Resultats scientifigues de ’Ex- pedition hydrofaunistique du Musee zoologique dans Siberia orientale en 1927. IV. Les Euco- pepodes d’eau douce de la region de |’Oussauri (Crustacea). Travaux de L’Institute Zoologique, Leningrad, 1:248-280 + 6 pls. Willey, A. A. 1920. Marine Copepoda. Report of the Canadian Arctic Expedition 1913-18. Vol. VIII. Part K. Thomas Mulvey, Ottawa, 47 pp. . 1923. Notes on the distribution of free-living Copepoda in Canadian waters.— Contributions to Canadian Biology and Fisheries. New Series 1:303-334. 1925. Northern Cyclopidae and Cantho- camptidae.— Transactions of the Royal Society of Canada, 3rd Series, 19:137-158 + 3 pls. Wilson, M. S. 1956. North American harpacticoid copepods. 1. Comments of the known fresh- water species of the Canthocamptidae. 2. Can- thocamptus oregonensis n. sp. from Oregon and California.— Transactions of the American Mi- croscopical Society 75:290-307. 1973. North American harpacticoid cope- pods 10. Pseudobradya major (Olofsson, 1917) n. comb. from Nuwuk Lake, Alaska, with a checklist of copepod associates. — Transactions American Microscopical Society 92:657-662. —, & H.C. Yeatman. 1959. Harpacticoida. Pp. 815-861 in W. T. Edmondson, ed., Fresh-water biology. John Wiley & Sons, Inc., New York. Department of Biology, Colorado State University, Fort Collins, Colorado 80521; (Postal address) 1901 Stover Street, Fort Collins, Colorado 80525. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 350-352 THE OCCURRENCE OF SPHAEROMA SERRATUM (FABRICIUS, 1787) IN THE WESTERN SOUTH ATLANTIC (CRUSTACEA: ISOPODA) Ana Marta Roux and Ricardo Bastida Abstract. —The European species Sphaeroma serratum is extending its range on the Argentine coast. It was probably introduced by ships in the harbor of Mar del Plata (Argentina) and subsequently spread to adjacent rocky intertidal communities and other harbors of the Buenos Aires Province. A key is given for identification of known southwestern Atlantic species of Sphaeroma. Sphaeroma serratum was first recorded from the Atlantic coast of Europe, later from the Mediterranean and Atlantic coasts of northern Africa and more recently from the Black Sea, southern Africa and western Australia (Kensley 1978, Jacobs 1987). Sphaeroma serratum has been known from Buenos Aires Province since 1964, where it was known only from Mar del Plata harbor (38°08’S, 57°31’W) having been found in samples taken during fouling studies (Bas- tida 1968, 1971; Bastida et al. 1980). It has since been found as part of fouling com- munities in other harbors in Argentina, e.g., Puerto Quequén (38°36’S, 58°40'W), Inge- niero White (38°47’S, 62°14'W) and Puerto Belgrano (38°54’S, 62°06’W) (Bastida 1972, Bastida & Torti 1973, Bastida & Branke- vich 1982, Martinez et al. 1984). Having colonized these harbor areas, in recent years Sphaeroma serratum has spread to natural rocky areas around Mar del Plata and has become a member of the local intertidal community (Bastida & L’Hoste 1976). Sphaeroma serratum has not yet been found on the Patagonian coasts, but its chances of colonization are thought to be high. The wide sandy beaches of Buenos Aires probably acted as a natural barrier to the spread of the species southward (Escofet et al. 1979). Sphaeroma serratum has not been recorded from the coast of Uruguay and Brazil, althought the genus is repre- sented in Brazil by three other species, Sphaeroma terebrans Bate, 1866, S. anan- dalei Stebbing, 1911, and S. walkeri Steb- bing, 1905 (Loyola e Silva 1960, Pires 1982). The distribution of Sphaeroma serratum north of Argentina may be limited by the estuary of the Rio de la Plata due to its great extension and the influence of its waters on Argentine and Uruguayan coasts. The method of introduction of Sphaeroma ser- ratum was probably as part of the fouling community on the hull of ships arriving in Argentina. This transportation was ob- served for Sphaeroma walkeri in Victoria Harbor (Hong Kong) and its dispersal to areas around Hong Kong (Mak et al. 1985). Other methods of transportation could have been by ballast water (Carlton & Iverson 1981) and less probably by pieces of wood floating adrift. Burrowed pieces of wood found in coastal areas are frequently inhab- ited by specimens of Sphaeroma serratum, which obtain protection in the teredine gal- leries and holes (Bastida & Torti 1972). Key to the Southwestern Atlantic Species of Sphaeroma 1. Pleotelson triangular, apex pointed. Pereon with tubercles. Mandible without lacinia mobilis ...S. terebrans VOLUME 103, NUMBER 2 — Pleotelson spoon-shaped, apex markedly rounded. Body with or without tubercles. Mandible with or without lacinia mobilis 2. Pereon and pleon without tubercles. Left mandible with lacinia mobilis Si HL ets ea Cale IM ea a S. serratum — Body with tubercles. Left mandible with or without lacinia mobilis... 3 3. Pereon with numerous round tu- bercles. Pleotelson with 2 subme- dian rows of tubercles, diverging from the base and almost reaching the apex. Left mandible with true lacinia mobilis S. walkeri — Pereon with transversally elongate tubercles. Pleotelson with 2 pair of tubercles situated on either side of the midline, followed by a single median tubercle. Mandible without true lacinia mobilis ..... S. anandalei Material One hundred fifty specimens of Sphae- roma serratum from different localities of the Buenos Aires Province were analyzed. The studied material is deposited in the col- lection of the Laboratory of Benthic and Fouling communities of the INIDEP (Mar del Plata, Argentina) and 12 specimens from the port of Mar del Plata are deposited in the National Museum of Natural History, Washington, D.C., USNM 139316. Acknowledgments The authors wish to express their grati- tude to Dr. Brian Kensley and Dr. Thomas Bowman for their kind suggestions. This is contribution No. 611 of INIDEP. Literature Cited Bastida, R. 1968. Preliminary notes of the marine fouling at the Port of Mar del Plata, Argentina. — Proceedings of the 2nd International Congress on Marine Corrosion and Fouling. Compte- Rendu, Athens, Greece, pp. 557-562. 1971. Las incrustaciones biologicas en el 351 Puerto de Mar del Plata. Periodo 1966/67.— Revista del Museo Argentino de Ciencias Natu- rales “Bernardino Rivadavia,” Hidrobiologia 3(2):203-285. 1972. Studies of fouling communities along Argentine Coasts.— Proceedings of the 3rd In- ternational Congress on Marine Corrosion and Fouling, Gaithersburg, Maryland, U.S.A., pp. 847-864. 1973. Estudio preliminar sobre las incrus- taciones biologicas de Puerto Belgrano (Argen- tina).—Corosion y Proteccion 5:3-11. —., & G. Brankevich. 1982. Estudios ecologicos sobre las comunidades incrustantes de Puerto Quequén (Argentina). II. Caracteristicas del Macrofouling.—Cidepint Anales 1982:156-193. —, & S. L’Hoste. 1976. Relaciones troficas de las comunidades incrustantes (‘‘Fouling”’) del Puerto de Mar del Plata. —LEMIT Anales 3-1976, Serie II 329:159-203. ——,&M.R. Torti. 1972. Organismos perforantes de las costas argentinas. I. La presencia de Ly- rodus pedicellatus (Quatrefages, 1849) (Mollus- ca, Pelecypoda) en el Puerto de Mar del Plata. Clave para el reconocimiento de los Teredinidae Sudamericanos.—Physis 31(82):39-SO0. ———,, M. Trivide Mand)n1, V. Lichtschein de Bastida, & M. Stupak. 1980. Ecological aspects of ma- rine fouling at the Port of Mar del Plata (Ar- gentina).—V Congreso Internacional de Cor- rosion Marina e Incrustaciones. Seccion Biologia Marina, Barcelona, Espana, pp. 229-320. Carlton, J. T., & E. W. Iverson. 1981. Biogeography and natural history of Sphaeroma walkeri Steb- bing (Crustacea: Isopoda) and its introduction to San Diego Bay, California.— Journal of Nat- ural History, London 15:31-48. Escofet, A., N. Gianuca, S. Maytia, & V. Scarabino. 1979. Playas arenosas del Atlantico Sudocci- dental entre los 29° y 43°LS. Consideraciones generales y esquema biocenologico.— Memorias del Seminario sobre Ecologia Bentonica y Se- dimentacion de la Plataforma Continental del Atlantico Sur, Montevideo, Uruguay, UNES- CO:245-258. Jacobs, B. J. M. 1987. A taxonomic revision of the European, Mediterranean and NW African species generally placed in Sphaeroma Bosc, 1802 (Isopoda: Flabellifera: Sphaeromati- dae).— Zoologische Verhandelingen 238:1-71. Kensley, B. 1978. Guide to the marine isopods of southern Africa.—South African Museum, Cape Town, 173 pp. Loyola e Silva, J. 1960. Sphaeromatidae do litoral Brasileiro (Isopoda—Crustacea).—Boletim da Universidade do Parana Zoologia 4:1-182. Mak, P. M. S., Z. G. Huang, & B. S. Morton. 1985. Sphaeroma walkeri Stebbing (Isopoda, Sphae- 352 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON romatidae) introduced into and established in dos Estados de Sao Paulo e Rio de Janeiro.— Hong Kong.—Crustaceana 49(1):75-82. Boletim do Instituto Oceanografico, Sao Paulo Martinez, D., R. Bastida, & G. Brankevich. 1984. 31(2):43-55. Ecological aspects of marine fouling at the Port of Ingeniero White (Argentina). — Proceedings Instituto Nacional de Investigacion y of the VI International Congress on Marine Cor- Desarrollo Pesquero (INIDEP), C.C. 175 rosion and Fouling, Marine Biology, Athens, Greece, pp. 521-537. Playa Grande, 7600 Mar del Plata, Argen- Pires, A. M.S. 1982. Sphaeromatidae (Isopoda Fla-__ tina. bellifera) da zona entre marés e fundos rasos PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 353-357 THE STATUS OF THE CARIDEAN SHRIMP PANDALINA MODESTA (BATE, 1888) (CRUSTACEA: DECAPODA: PANDALIDAE) WITH REDESCRIPTION OF THE SPECIES E. Macpherson Abstract. — The identity of Pandalus modestus Bate, 1888 from South Africa is studied. The species belongs to the genus Pandalina Calman, 1899. Com- parison with the other species of the genus revealed that Bate’s species is closely related to Pandalina profunda Holthuis, 1946 from European and west African waters. However, both species are easily distinguished by the length of the pereopods, clearly longer in P. modesta. Bate (1888) described a new species of pandalid shrimps under the name Pandalus modestus from South Africa (Agulhas Bank, 274 m [= 150 fms] depth). The species was later cited in the same area, under the same name, by Stebbing (1914) and by Barnard (1950), who questionably synonymized the species with Pandalina brevirostris (Rathke, 1843), a common species in the European coasts. The same author suggested that the species should be synonymous with Pan- dalina profunda Holthuis, 1946, from Eu- ropean and west African waters. The differences between Pandalina bre- virostris and Pandalina profunda were def- initely established by Greve (1967). This author indicated that there is really only one reliable difference separating the two species: the spinose extent on the ambulatory dac- tyls. However, the identity of the specimens from southern Africa remained unresolved (Crosnier & Forest 1973). Unfortunately, Bate’s type is in a poor condition; the rostrum is broken and the pereopods are lost (Crosnier & Forest 1973). Therefore, in this study specimens collected around the type locality (Cape area) have been used for a comparison with the two species of Pandalina. This comparison revealed that the spec- imens from southern Africa actually rep- resent a different species of the genus Pan- dalina Calman, 1899; therefore, Bate’s species should be validated. Because of the problems of identity and distribution that have existed with Pandalus modestus, the species is here redescribed. Specimens used for this study came from the collections of the South African Mu- seum, Cape Town (SAM), Muséum natio- nal d’Histoire naturelle, Paris (MNHN) and Instituto de Ciencias del Mar, Barcelona (ICM). The abbreviation CL indicates car- apace length excluding rostrum. Pandalina modesta (Bate, 1888) Figs. 1, 2 Pandalus modestus Bate, 1888:670, pl. 114, fig. 4.—Stebbing, 1910:392.—Stebbing, 1914:36. Pandalina brevirostris. —Barnard, 1947: 384.—Barnard, 1950:676, fig. 126a—e.— Kensley, 1981:28 (not Pandalina brevi- rostris Rathke, 1843). Pandalina profunda.—Macpherson, 1983: 64 (not Pandalina profunda Holthuis, 1946). Material examined. —Off Cape area and East London: 12 2 (CL = 5.0-6.8 mm), 2 ovig. 2 (CL = 5.4—5.5 mm), 4 6 (CL = 5.7- 6.5 mm), R/V Pieter Faure, 268-360 m 354 (several samples together), SAM A8396- 8398.—1?2(CL=4.5 mm), SAM A13280.— 12(CL=6.1 mm), SAM A13642.—1 2? (CL = 5.7 mm), SAM A1280.—South of Na- mibia: 1 @ (CL = 7.1 mm), 26°42’S, 14°06’E, 395 m, ICM D1047. Description. —Rostrum horizontal, not nearly reaching level of end of penultimate antennular segment, around 0.5 times length of carapace; dorsal margin armed with 8 to 10 teeth, 5 proximals being movable; lower margin bearing 2 to 4 teeth. Antennal spine stronger than pterygostomian spine. Abdomen with 3rd somite rounded pos- teriorly, unarmed, slightly overhanging 4th segment. Pleura of 3 anterior somites broadly rounded, of 4th and 5th bearing sharp tooth. Sixth somite slightly less than 2 times as long as 5th somite (length mea- sured on dorsal margin) and less than 2.5 times maximum height. Telson as long as 6th somite, with 7 or 8 pairs of dorso-lateral spines and 2 longer pairs on apex. Eyes broadly subpyriform, maximum di- ameter about 0.2 times carapace length; ocellus present, slightly constricted at junc- ture with cornea. Antennular peduncle with small tooth on inner margin of basal segment. Ultimate and penultimate segment of similar length. Antennal scale with lateral margin nearly straight, about 0.7 to 0.8 times as long as carapace, 5.0 times as long as wide, disto- lateral tooth not overreaching blade. Mouth parts as illustrated. Third maxil- liped with epipod; endopod slightly over- reaching antennal scale, armed terminally with long, slender apical and few subapical spines; penultimate segment about 0.5 times as long as terminal segment. Exopod absent. Pereopods with well-developed epipods on 4 anterior pairs. First pereopod slender, overreaching antennular peduncle by length of propodus. Second left pereopod longer than right. Right overreaching antennular peduncle by somewhat less than length of chela; merus occurring at level of anterior border of basicerite; carpus with 4 to 6 seg- ments and about 1.8 times long as chela. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Left overreaching antennular peduncle by somewhat less than half carpus length; car- pus composed of 16 to 21 articles and some- what less than 5 times chela length. Third pereopod with carpus clearly overreaching antennular peduncle; ischium with 1 or 2 posterior spines; merus occurring pterygo- stomian angle by 0.4 of merus; merus about 2 times as long as carpus, armed with 11 to 14 posterior and outer spines; carpus bear- ing 6 to 9 posterior and outer spines; propo- dus almost two times length of carpus and 3.0 to 3.8 times length of dactylus, having 10 to 18 posterior spines; dactylus armed with 3 or 4 spines on proximal half of pos- terior margin. Fourth pereopod with carpus clearly occurring penultimate segment of antennular peduncle. Ischium with 2 or 3 posterior spines; merus about 2 times length of carpus, armed with 14 to 18 posterior and outer spines; merus occurring ptery- gostomian angle by 0.2 of merus length; car- pus having 5 to 7 posterior and outer spines; propodus 1.7 to 2.0 times length of carpus and about 4.5 times length of dactylus, bear- ing 12 to 14 posterior spines; dactylus armed with 3 or 4 spines on the proximal half of posterior margin. Fifth pereopod with me- rus clearly not reaching pterygostomian an- gle; ischium with 1 or 2 small posterior spines; merus about 1.7 times length of car- pus, having 10 to 13 posterior and outer spines; carpus armed with 4 outer spines; propodus almost two times carpus length and 4.3 to 5.0 times as long as dactylus and bearing 9 or 10 posterior spines; dactylus with 3 spines on proximal 2/3 of posterior margin. Uropods overreaching end of telson; ex- opodite longer than endopodite and bearing movable spine. Remarks. —Pandalina brevirostris is closely related to P. profunda but it is readily differentiated by the length of the dactylus of 3rd—Sth pereopod and the part of the dactylus bearing posterior spines. In P. pro- funda the dactylus is long and slender with the posterior margin carrying spines only in the proximal half. In P. brevirostris the dac- VOLUME 103, NUMBER 2 355 Fig. 1. Pandalina modesta (Bate, 1888), 2, CL = 4.5 mm, SAM A13280. A, Anterior carapace and anterior appendages; B, Right third maxilliped, distal end; C, Second right pereopod; D, Second left pereopod; E, Third right pereopod; F, Same, dactylus. 356 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Imm —_—I ABC 2mm —— D 05mm EFGHI Fig. 2. Pandalina modesta (Bate, 1888). A—D, 2, CL = 4.5 mm, SAM A13280. A, Right antennula; B, Right antenna, ventral view; C, Telson; D, Abdomen. E-I, 2, 5.7 mm (SAM A8396-8398). E, Right mandible; F, Right first maxilla; G, Right second maxilla; H, Right first maxilliped; I, Right second maxilliped. VOLUME 103. NUMBER 2 tylus is much shorter and the spines of the posterior margin always are regularly dis- persed over the entire length (Holthuis 1946, Greve 1967, Zariquiey-Alvarez 1968). Pandalina modesta has a long and slender dactylus, with spines only in the proximal half; thereby it resembles P. profunda (ma- terial examined of P. profunda: 1 2 [CL = 5.7mm], Travailleur, st. 58, 440 m [MNHN Nal213].—2 2 [CL = 3.1-4.3 mm], 47°34.8'N, 07°18.1'W, 825 m [MNHN Nal0319].—1 2 [CL = 3.2 mm], 48°40.8’S, 09°47.7'W, 373 m [MNHN Nal0321].—4 2 [CL = 2.8-3.8 mm], 48°41.6’S, 09°52.9'W, 350 m [MNHN Nal10322].—1 6 [CL = 3.8 mm], 7 2 [CL = 3.6-4.4 mm], 47°36’S, 07°16.8'W, 330 m [MNHN Na10320)]). The comparative analysis of these specimens shows that they are easily distinguished by the length of the pereopods, clearly longer in P. modesta than in P. profunda. The merus of the 2nd left pereopod of P. modesta almost reaches the end of the basal segment of the antennular peduncle, while in P. profunda it slightly overreaches the basicerite. In P. modesta the 3rd pereopod has the merus reaching or overreaching the termi- nal border of the cornea. In P. profunda it reaches only to the level of the proximal border of the cornea. The merus of the 4th pereopod in P. mo- desta reaches or overreaches the distal mar- gin of the basicerite, while in P. profunda it only slightly overreaches the pterygosto- mian border. Also, the carpus of the left second per- eopods has generally more articles in P. mo- desta (16 to 21) than in P. profunda (14 to 16). Distribution. —Southern Africa, from East London to Saldanha Bay and south of Na- mibia. Depth range: 265 to 360 m. Acknowledgments Iam grateful to Dr. L. B. Holthuis (Riyks- museum van Natuurlijke Historie, Leiden) for reading a draft of the manuscript. [ thank Dr. F. A. Chace, Jr. (National Museum of 357 Natural History, Washington) and Dr. A. Crosnier (Museum national d’Histoire na- turelle, Paris) for the revision of the manu- script. Thanks are also extended to Dr. For- est (Museum national d’Histoire naturelle, Paris) and Mrs. M. van der Merwe (South African Museum, Cape Town) for the loan of the material of Pandalina. Literature Cited Barnard, K. H. 1947. Descriptions of new species of South African decapod Crustacea, with notes on synonymy and new records.— Annals and Mag- azine of Natural History, ser. 11, 13:361-392. . 1950. Descriptive catalogue of South African decapod Crustacea (crabs and shrimps).—An- nals of the South African Museum 38:1-837. Bate, C. S. 1888. Report on the Crustacea Macrura collected by H.M.S. Challenger during the years 1873-76.—Report on the Scientific Results of the Voyage of H.M.S. “Challenger” during the Years 1873-76, 24:1-942. Crosnier, A., & J. Forest. 1973. Les crevettes pro- fondes de |’Atlantique oriental tropical. — Faune Tropicale 19:1-409. Greve, L. 1967. Notes on the genus Pandalina in Norwegian waters (Crustacea Decapoda).—Sar- sia 26:1-5. Holthuis, L. B. 1946. Note on the genus Pandalina (Crustacea Decapoda), with the description of a new species from European waters.—Zoolo- gische Mededeelingen 26:28 1-286. Kensley, B. 1981. On the zoogeography of southern African decapod Crustacea, with a distribution- al checklist of the species. Smithsonian Con- tributions to Zoology 338:1-64. Macpherson, E. 1983. Crustaceos Decapodos cap- turados en las costas de Namibia. — Resultados Expediciones Cientificas (supl. Investigacion Pesquera) 11:3-79. Stebbing, T. R. R. 1910. General catalogue of South African Crustacea (Part V of S.A. Crustacea, for the marine investigations in South Africa).— Annals of the South African Museum 6:281- 593. 1914. South African Crustacea (Part VII of S.A. Crustacea, for the marine investigations in South Africa).—Annals of the South African Museum 15:1-55. Zariquiey-Alvarez, R. 1968. Crustaceos Decapodos Ibéricos.— Investigacion Pesquera 32:i-xv + 1- 510. Instituto de Ciencias del Mar, P° Nacional s/n. 08003 Barcelona, Spain. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 358-363 GALATHEA CORALLIOPHILUS, A NEW DECAPOD CRUSTACEAN (ANOMURA: GALATHEIDAE) FROM SINGAPORE, GULF OF THAILAND, AND WEST IRIAN Keiji Baba and Sang-Chul Oh Abstract. —Galathea coralliophilus, a new galatheid crustacean, is described and illustrated. The new species is common in association with reef corals in Singapore and the Gulf of Thailand. Galathea spinosorostris reported by John- son (1970:6) from Singapore and Galathea aff. consobrina described by Gordon (1935:5) from West Irian are synonymized with the present species. During ecological studies on the coral communities of the Gulf of Thailand in 1984, Dr. Y. Nakasone collected galatheid crustaceans commensal with several species of reef corals and made them available to one of us (KB) for identification; they were immediately identified as an undescribed species of Galathea. Recently, Singaporean galatheids in the collection of the National University of Singapore (Zoological Refer- ence Collection [ZRC]), taken mostly on coral reefs, rarely from crinoids or fouling rubbles on a ship or among fouling com- munity, were made available for study. They comprise two species: one of them is A/lo- galathea elegans (Adams & White), a com- mon crinoid associate in the Indo-West Pa- cific, and the other is identical with the Thailand species, as well as with the species reported by Johnson (1970:6) as Galathea spinosorostris, which, according to him, is the only shore galatheid in Singapore. How- ever, Johnson’s identification is incorrect, as the identity of Galathea spinosorostris Dana, 1852 has recently been established (Baba 1988:78). In addition, examination of the literature discloses that Galathea aff. consobrina from West Irian (Gordon 1935: 5) should also be merged with the new species. Most of the material from Thailand is retained in the collection of Kumamoto University, and part will be sent to the Smithsonian Institution. Locations of sta- tions where the Thailand survey was carried out can be found in Nakasone et al. (1986: fig. 1). The measurements of the specimens examined are given in parentheses under ‘“‘Material,” showing minimum and maxi- mum carapace lengths in millimeters taken between the rostral tip and midpoint of the posterior margin of the carapace. The host corals from Thailand have been identified by Kazuhiko Sakai (see Nakasone et al. 1986:155), and those, as well as a host cri- noid, from Singapore have been determined by Beverly Goh (Peter Ng, pers. comm.). Galathea coralliophilus, new species Fig. 1 Galathea aff. consobrina.—Gordon, 1935: 5, figs. 2, 3a, b. Galathea spinosorostris. —Johnson, 1970:6, fig. 1b. [Not G. spinosorostris Dana, 1852.] Material.—Thailand: Khang Khao Is- land, sta. B, 26 Oct 1984, on Pavona fron- difera Lamarck: 1 ovig. 2 (3.5); on Pocillo- pora damicornis Linnaeus: 4 6 (3.1—5.5), 2 ovig. 2 (3.7, 5.1); 15 Nov 1984, on P. dami- cornis: 1 6 (4.5), 1 ovig. 2 (3.8), 1 2 (3.2); sta. C, 25 Oct 1984, on P. damicornis: | 6 (3.0). Samui Island, sta. 1, 10 Nov 1984, on Acropora sp.: 1 29(3.5); on Psammocora con- VOLUME 103, NUMBER 2 tigua (Esper): 4 6 (2.8-4.8). Singapore: Pulau Hantu, North, 12 Aug 1986, coll. B. Goh, on Acropora (Acropora) tenuis (Dana): 1 3 (4.7) (ZRC 1987.2555); Pulau Hantu, South, 19 Aug 1986, on Pavona frondifera: 2 2 (4.1, 4.3) (ZRC 1987.2558-2559); Pulau Hantu, West, 19 Sep 1986, on Pocillopora dami- cornis: 3 6 (3.7-5.3), 1 2 (4.9) (ZRC 1987.2582-2585). Cyrene Reef, 15 Aug 1986, coll. B. Goh, on Pavona decussata (Dana): | ovig. 2 (4.3+), 1 2 (3.7+) (ZRC 1987.2556-—2557). Pulau Semakau, 22 Aug 1986, coll. B. Goh, on Pavona frondifera: 1 3 (4.5), 2 ovig. 2 (4.4, 4.1+), 1 2 (cephalo- thorax missing) (ZRC 1987.2560—2563); on Cyphastrea serailia (Forskal): 1 6 (3.9), 1 2 (carapace missing) (ZRC 1987.2564—2565); on Pavona cactus (Forskal): 1 6 (4.8) (ZRC 1987.2566); coll. G. Lim, on crinoid Co- master gracilis (Hartlaub): 1 6 (5.3) (ZRC 1987.2596). Pulau Jong, 26 Aug 1986, coll. B. Goh, on Pocillopora damicornis: | ovig. 2 (4.4+) (ZRC 1987.2567). Pulau Jong, South, 24 Oct 1986, coll. B. Goh, on Pavona frondifera: | ovig. 2 (5.5) (ZRC 1987.2595). Raffles Lighthouse, 2 Sep 1986, coll. B. Goh, on Acropora (Acropora) willisae Veron & Wallace: 1 6 (5.1) (ZRC 1987.2568); on Po- cillopora damicornis: 3 6 (3.7—4.8), 1 ovig. 2 (4.2), 1 2 (4.7) (ZRC 1987.2569-2573); 16 Oct 1986, on Pocillopora damicornis: 1 6 (4.9), 1 ovig. 2 (5.9) (ZRC 1987.2586-2587); on Pavona frondifera: 3 6 (4.5—6.3), 1 2 (6.0) (ZRC 1987.2588-2591) [largest male is ho- lotype]; on Pavona decussata: 2 6 (4.5, 5.6), 1 ovig. 2 (4.5) (ZRC 1987.2592-2594). Ter- embu Pempang Laut, South, 9 Sep 1986, coll. B. Goh, on Pocillopora damicornis: 5 6 (4.0-6.4), 3 ovig. 2 (3.7-5.2) (ZRC 1987.2574—2581). Southern Islands, Dec 1986, coll. J. B. Sigurdson, from fouling rubble on ship: 1 6 (6.4), 1 ovig. 2 (4.8) (ZRC 1987.2597—2599). Sentosa Reef, Dec 1985, coll. P. Ng: 2 6 (3.7, 4.1), 2 ovig. 2 (3.9, 4.1) (ZRC 1987.2600-2603). Pulau Busing, 1 May 1987, coll. J. B. Sigurdson, from foul- ing community: 2 ovig. 2 (4.5, 5.2) (ZRC 1987.2604—2605). 359 Description of holotype.—Carapace ex- cluding rostrum and lateral spines nearly as long as wide, dorsal surface with distinct transverse ridges as figured, cervical groove indistinct; first transverse ridge with pair of submedian spines, each spine accompanied by feathered seta mesial to it, also lateral to it on left side only; scale-like ridge present directly behind second transverse ridge, bearing 4 pits for roots of stiff feathered se- tae. Lateral margins convex, bearing 7 spines, first (anterolateral) pronounced, sec- ond small and somewhat dorsal in position, third to seventh located behind end of cer- vical groove, fifth largest. Another spine ventral to level of second lateral marginal spine and somewhat anterior to anterior end of linea anomurica. Outer angle of orbit strongly produced, ending in sharp point. Rostrum broadly triangular, 1.4 times as long as wide when measured between in- cisions formed by 2 proximal teeth, dorsal surface with several pronounced feathered setae in addition to fine simple ones, lateral margin with 4 sharply incised teeth, basal lateral tooth smaller, ultimate lateral tooth terminating opposite midlength of rostral median spine. Eyestalks with few feathered setae near mesial limit of terminal fringe of very fine setae proximal to cornea. Abdominal segments sparsely provided with feathered setae as illustrated; second and third segments each with 4 transverse ridges, first (anterior) ridge elevated, second ridge medially interrupted by considerable space, third ridge uninterrupted, accompa- nying groove anterior to it, fourth ridge faint and medially interrupted. Pterygostomian flap anteriorly ending in sharp spine, bearing small spine on second ridge. Basal segment of antennule with 3 well developed terminal spines, distal segment with tuft of pronounced terminal setae. First (proximal) segment of antennal peduncle with rather long distomesial spine nearly reaching end of third segment; second seg- 360 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ment with distomesial and distolateral spines of subequal size; third segment with small distomesial spine. Third maxilliped having ischium with small distoventral and short but stout dis- todorsal spines, mesial ridge with 23 den- ticles; merus with 2 well developed ventral spines on distal half of length, distal one present at midlength of space between prox- imal spine and distal end of ventral margin; dorsal margin with distodorsal spine of small size and eminence proximal to it; carpus unarmed but with 3 eminences on dorsal margin; dactylus elongate. Third thoracic sternum depressed below level of following sternum, anteriorly nar- rowed, anterior margin with distinct me- dian notch. Chelipeds 2.7 times as long as carapace, provided with coarse setae in moderate den- sity and very spinose as illustrated; merus with 5 rows of spines, distal 2 of mesial marginal spines prominent; carpus with 5 rows of spines, penultimate of mesial mar- ginal spines very strong; palm about 3 times as long as wide, with 4 rows of spines on dorsal surface and mesial and lateral mar- gins; fingers gaping, touching each other with several intermeshing teeth at tip when closed; movable finger 0.57 as long as palm, bearing few spinules slightly dorsal to prox- imal mesial margin, opposable margin with prominent but short basal process proximal to Opposing process at midlength of fixed finger. Walking legs sparsely provided with pro- nounced feathered setae. First and second walking legs subequal in size, third one shorter; meri dorsally provided with 10 spines on first leg, 9 or 10 on second leg, 4 on third leg proximal to distodorsal spine, distolaterally with pronounced spine on all these legs, accompanying tiny one proximal and ventral to it on first and second legs; carpi with few tiny spines on dorsolateral surface parallel to 4 dorsal marginal spines. Dorsal margins of propodi with 2 proximal spines, ventral margins with 4 spinelets on first leg, 3 on second leg and 3 or 4 on third leg; dactyli ending in strongly curved sharp claw, ventrally bearing 5 teeth each with stiff seta at base, all these teeth and setae de- creasing in size toward base of segment. Epipods present on chelipeds, absent from walking legs. Variation.—The scale-like ridge imme- diately behind the second transverse ridge is absent in only two of the 66 specimens examined. In these two, the second stria is interrupted at midpoint and both of their mesial limits are directed posteriad. The feathered seta mesial to each epigastric spine is consistently present, and an additional one lateral to it is present in two specimens including the holotype. The spine on the pterygostomian flap is usually very tiny, oc- casionally discernible only under high mag- nification, and very rarely obsolete (in two specimens). The propodal ventral spinelets vary from two to five (mostly four) on the first walking leg, two to four (mostly three) on the second walking leg, one to four (mostly three, occasionally two) on the third walking leg. The merus of the third walking leg bears three to five (mostly four) spines on the dorsolateral surface near dorsal mar- gin, in addition to a distodorsal one. The dorsal margin of the merus of the third max- iliiped bears at most three spines; the me- dian one is mostly tiny and rarely missing; the proximal one tends to be obsolete; and the distal one is constantly present. Habitat.—The specimens examined are mostly reef associates, very frequently taken from Pocillopora damicornis Linnaeus, oc- casionally from Pavona frondifera Lamarck, rarely from Pavona decussata (Dana), P. cactus (Forskal), Acropora tenuis (Dana), A. willisae Veron & Wallace, Cyphastrea serai- lia (Forskal) and Psammocora contigua (Es- per); a few lots are taken from a crinoid Comaster gracilis (Hartlaub), fouling rubble on a ship, and among fouling community. Parasites. —Six of the 50 specimens from VOLUME 103, NUMBER 2 361 Fig. 1. Galathea coralliophilus new species, male holotype, ZRC1987.2588: a, Carapace and abdomen. dorsal view, fine setae on transverse ridges omitted; b, Left pterygostomian flap; c, Basal segment of right antennule, ventral view; d, Right antennal peduncle, ventral view; e, Endopod of right third maxilliped, lateral view; f, Anterior part of sternal segments; g, Left cheliped, dorsal view; h, Left first walking leg, lateral view; i, Left third walking leg, lateral view. Scales = 1 mm; scale 1 for g; scale 2 for a; scale 3 for b, h, i; scale 4 for f; scale 5 for c, d, e. 362 Singapore have externas of rhizocephalan parasites; no externas on the Thailand spec- imens. Remarks. —Galathea aff. consobrina, re- ported by Gordon (1935:5) from West Irian, the identity of which was questioned be- cause ofa lack of two spinules on the hepatic region (Baba 1988:73), is in all probability identical with the present species. The rounded outer orbital angle in G. aff. con- sobrina, which Gordon stressed as one of the characteristics that differentiates her unique specimen from G. consobrina, is the only one that does not agree with the present specimens. Very possibly she must have overlooked a sharp tip, because, in my ex- perience, such a sharply produced angle as illustrated by Gordon (1930:fig. 2) has not been observed in any of the known species that have a rounded outer orbital angle. The differences between G. coralliophilus and G. consobrina De Man, 1902 are ap- parent in the following particulars that were verified by examination of both the male holotype in the collection of the Sencken- berg Museum at Frankfurt am Main (SMF 4556) and additional specimens from the Philippines (Baba 1988:73) of G. consobri- na: 1) the epipods on the chelipeds are pres- ent in G. coralliophilus, absent in G. conso- brina,; 2) the hepatic region bears two spinules other than the second of the seven lateral marginal spines of the carapace in G. consobrina, none in G. coralliophilus; 3) the third thoracic sternum is much wider in G. consobrina than in G. coralliophilus (see Baba 1988:fig. 30e); 4) a spine on the anterior surface of the pterygostomian flap is usually present though mostly tiny in G. corallio- Dhilus, absent in G. consobrina. Johnson (1970:6) mistakenly reported G. spinosorostris from Singapore. The true G. spinosorostris originally known from the Hawauian Islands (Dana 1852:480) is char- acterized by scale-like ridges on the bran- chial region, the outer orbital angle un- armed, three hepatic spinules including the second lateral marginal spine of the cara- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pace, and lack of a spine on the anterior portion of the pterygostomian flap (Baba 1988:78). The brief account of morphology with an illustration, as well as a note on the habitat, of G. spinosorostris provided by Johnson, suggests that his specimens should be referable to the present new species. He stated that the merus of the third walking leg bears seven or eight dorsal spines. This is the only one to be at variance with the present species. However, it is most likely that his note is based on the first or second walking leg, because most of the known species including the present new species has the third walking leg with at most four or five dorsal and one distolateral spines. The presence of a spine on the anterior stria of the pterygostomian flap and two epi- gastric and seven lateral marginal spines on the carapace link the species to Galathea orientalis Stimpson, 1858 (see Miyake & Baba 1967:233). However, they differ in that: the carpus of the third maxilliped bears a few but usually distinct dorsal spines in G. orientalis but lacks them in G. corallio- philus; the scale-like ridge immediately be- hind the second transverse ridge is distinct in G. coralliophilus, absent in G. orientalis; and the rostrum has a few to several feath- ered setae on the dorsal surface in G. cor- alliophilus but lacks them in G. orientalis. Etymology. —The specific name is a noun in apposition from the Greek, “‘corallion,”’ coral, and “‘philos,” loving or fond of, for the association of the species mostly with corals. Acknowledgments We thank Yukio Nakasone of the Uni- versity of the Ryukyus and Peter Ng of the National University of Singapore, for mak- ing the interesting specimens available for study. We owe a Special debt of gratitude to Raymond B. Manning and Fenner A. Chace, Jr. of the Smithsonian Institution for re- viewing the manuscript. The holotype of Galathea consobrina De Man in the collec- tion of the Senckenberg Museum was ex- VOLUME 103, NUMBER 2 amined on loan by the courtesy of Michael Turkay. Literature Cited Baba, K. 1988. Chirostylid and galatheid crustaceans (Decapoda: Anomura) of the “‘Albatross” Phil- ippine Expedition, 1907—1910.— Researches on Crustacea, Special Number 2:v + 203 pp. Dana, J.D. 1852. Crustacea, Part 1. Jn United States exploring expedition, during the years 1838, 1839, 1840, 1841, 1842, under the command of Charles Wilkes, U.S.N.. 13:vili + 685 pp., C. Sherman, Philadelphia. De Man, J. G. 1902. Die von Herrn Professor Ku- kenthal im Indischen Archipel gesammelten Dekapoden und Stomatopoden.—Abhandlun- gen herausgegeben von der Senckenbergischen Naturforschenden Gesellschaft 25:467-929, pls. 19-27. Gordon, I. 1935. Anomura (excluding Paguridea), in Résultats scientifiques du Voyage aux Indes ori- entales Néerlandaises de LL.AA.RR. le Prince et la Princesse Leopold de Bélgique. Volume III, 363 Fascicule 15.—Mémoires du Musée Royal d’Histoire naturelle de Belgique, hors série, 12 pp. Johnson, D. S. 1970. The Galatheidea (Crustacea, Decapoda) of Singapore and adjacent waters. — Bulletin of the National Museum Singapore 35(1): 1-44. Miyake, S., & K. Baba. 1967. Galatheids of the East China Sea (Chirostylidae and Galatheidae, De- capoda, Crustacea).—Journal of the Faculty of Agriculture, Kyushu University 14(2):225-246. Nakasone, Y., M. Tsuchiya, V. Manthachitra, & M. Nishihira. 1986. Species composition of deca- pod crustaceans associated with living corals in the Gulf of Thailand. —Galaxea, Publications of Sesoko Marine Science Center, University of the Ryukyus 5:141-156. (KB) Faculty of Education, Kumamoto University, Kumamoto, 860 Japan; (SO) Department of Natural Sciences, Cheju Col- lege of Education, Cheju, 590 Korea. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 364-371 CALYPTRAEOTHERES, A NEW GENUS OF PINNOTHERIDAE FOR THE LIMPET CRAB FABIA GRANTI GLASSELL, 1933 (CRUSTACEA, BRACHYURA) Ernesto Campos Abstract.— Fabia granti Glassell, 1933 is incorrectly assigned to the genus Fabia Dana, 1851. Its unique morphological features include: (1) lack of two longitudinal sulci on the carapace; (2) the anterolateral margins of the carapace are sharp-edged; (3) the third maxilliped has a palp of two articles, carpus longer and wider than propodus, and (4) abdomen, in both sexes, of seven free somites, and these support the placement of F. granti as type species of a new pinnotherid crab genus, Calyptraeotheres. In addition to the morphological differences, this genus is ecologically different from others in the Pinnotheridae, since it is a symbiont of gastropods, family Calyptraeidae. Restimen. — Fabia granti Glassell, 1933 ha estado incorrectamente asignada dentro del género Fabia Dana, 1851. Sus caracteristicas exclusivas incluyen: (1) ausencia de dos surcos longitudinales sobre el caparazon; (2) margenes del caparazon agudos; (3) tercer maxilipedio con su palpo formado de dos artejos, el carpus mas ancho y largo que el propodus y (4) el abdomen, en el macho y la hembra, formado de siete somitos libremente articulados. Lo anterior apoya la asignacion de F. granti como especie tipo de un nuevo género de cangrejo en la familia Pinnotheridae, Calyptraeotheres. Adicional a las caracteristicas morfologicas, Calyptraeotheres es ecologicamente diferente de otros géneros en los Pinnotheridae dado que es simbionte de Mollusca— Gastropoda, familia Calyptraeidae. The pinnotherid crab genus Fabia was es- tablished by Dana, 1851 with F. subquada- rata Dana, 1851, from the northeastern Pa- cific, as its type species. Subsequently, eleven species were assigned within this genus (Schmitt et al. 1973, Rodrigues da Costa 1970, De Melo 1971, Cobb 1973, Dai 1980, Gore 1986). Of these, two species have been transferred to other genera. Pregenzer (1979) assigned F. hickmani to Pinnotheres Bosc, 1802, and Campos (1989a) included F. un- quifalcula in Orthotheres Sakai, 1965. Re- cent examination of males and females of F. granti Glassell, 1933, and their compar- ison with specimens and descriptions of F. subquadrata (Rathbun 1918, Davidson 1968) revealed several morphological dif- ferences of taxonomic importance. In F. subquadrata two longitudinal sulci, which arise from the upper margin of the orbit, are present on the carapace of the female, the palp of the third maxilliped is composed of three articles in both sexes and the ab- domen of the male has somites 3—5 fused (Rathbun 1918, pers. obs.). In contrast, F. granti does not show the sulci, the palp is composed of two articles in both sexes, and the abdomen of the male is formed by seven free somites. These morphological differ- ences suggest that (1) F. granti should not be considered as congeneric with F. subquadrata; (2) its inclusion within genus VOLUME 103, NUMBER 2 Fabia was incorrect; and (3) as F. granti is morphologically unique in the Pinnotheri- dae, a new genus should be proposed to re- ceive this singular species. Systematic Results Family Pinnotheridae Calyptraeotheres, new genus Diagnosis. — Female: Carapace with an- terolateral margin arcuate and sharp-edged, dorsal regions ill-defined. Third maxilliped obliquely placed in buccal cavity; ischium and merus indistinguishably fused, widen- ing distally, inner margin concave, distal one almost straight; palp with two articles, car- pus wider and longer than propodus, latter digitiform and obliquely truncated. Abdo- men with seven free somites, covering ster- num but not reaching to buccal area. Male: Carapace subpentagonal, regions ill-defined, dorsal surface even and with short and separate setae; anterolateral mar- gins with fringe of hair-like setae. Third maxilliped similar to that of female. Ab- domen with seven free somites, widest at third somite and narrowing toward seventh. Type species.—By present designation, Fabia granti Glassell 1933. The genus is monotypic. Etymology. —Calyptraeotheres has been selected to emphasize the symbiotic rela- tionship between the limpet-crabs and gas- tropods, family Calyptraeidae. Gender masculine. Distribution. —East Pacific: Golfo de Cal- ifornia and West coast of Baja California Sur at Bahia Magdalena (Glassell 1933, 1935). Hosts. —Mollusca: Gastropoda: Calyp- traeidae: Crepidula and Crucibulum. Ac- maeidae: Acmaea. Remarks. —At present five genera in the Pinnotheridae contain one or more species with the palp of the third maxilliped com- posed of two articles. These are, Dissodac- tylus (sensu Griffith 1987), Durckheimia (sensu Seréne 1967), Ostracotheres (sensu 365 Tesch 1918, Pregenzer 1988), Pinnotheres (sensu Roberts 1975), and Xanthasia (sensu Burger 1895). The genus Calyptraeotheres can be distinguished from all of these genera because its carapace is smooth, with the re- gions ill-defined and the anterolateral mar- gins sharp-edged, and the carpus of the third maxilliped being larger than the digitiform and obliquely truncated propodus. Further- more, Calyptraeotheres is also different eco- logically. This genus is an ecosymbiont of limpets, Gastropoda—Calyptraeidae, while the genera listed above are symbionts of echinoids, Bivalvia, and/or Tunicata, which suggest divergent evolutionary trends. Only one species is now included within Calyp- traeotheres; however, two additional ones, Pinnotheres politus (Smith, 1870) and P. garthi Fennuci, 1975, possibly should be included in this new genus. The comparison between C. granti and descriptions and fig- ures of P. politus and P. garthi recorded by Rathbun (1918), Garth (1957) and Fennuci (1975) suggest that both males and females of C. granti share a morphological similarity to the South American species which are symbionts of limpets, family Calyptraeidae, too. The major difference observed in P. politus and P. garthi is the presence of a minute and rounded dactylus on the palp of the third maxilliped (Fig. 2B, C). A de- cision on the status of these Austral species and whether they should be included in Cal- yptraeotheres awaits a revision of the type series or of voucher specimens. Calyptraeotheres granti (Glassell, 1933), new combination Figs. 1, 2A, D-F Fabia granti Glassell, 1933:342, pl. 26, fig. 1-3; 1934:301; 1935:105; 1938:452, pl. 33, fig. 12.—Balss, 1957:1421.—De Melo, 1971:202.—Schmitt, McCain & David- son, 1973:24.—Campos-Gonzalez & Ma- cias-Chavez, 1988:241.—Campos, 1989a: De Material examined. —Eighty-four males, 366 carapace width from 1.7 to 3.1 mm and 296 females (196 ovigerous), carapace width from 2.2 to 8.8 mm, collected between Au- gust 1983 to July 1984, Laguna Percebu, about 23 km south of San Felipe, Baja Cal- ifornia, Mexico, (lat 30°49'00’N, long 114°41'40”W), E. Campos coll. Distribution. —Golfo de California, and West coast of Baja California Sur at Bahia Magdalena (type locality), (Glassell 1933, 119335): Female. —Carapace (Fig. 1A) with front little projected, anterior third arcuate, lat- eral margins subparallel and posteriorly convergent; posterior margin straight or lit- tle concave in middle. Carapace margins, specially of frontal and anterolateral re- gions, sharp-edged. Gastric, cardiac and in- testinal regions ill-defined, limited by straight cervical depressions arising from orbital region. Antennular grooves large. Orbits small, completely filling sockets, cor- nea not visible in dorsal view. Buccal cavity crescentic. Third maxilliped placed obliquely, ischium and merus indistin- guishably fused, widening distally, inner margin concave, distal margin straight; palp placed in outer angle of merus, with two articles, carpus longer and wider than prop- odus, latter obliquely truncated (Fig. 2A). Chelipeds stout, palm widening distally, outer and inner surfaces even; dactylus with proximal, small, blunt tooth, occasionally other smaller tooth present; propodus with two proximal blunt teeth separated by notch where tooth of dactylus fits. Fingers little deflexed, without gap when closed; ventral margin of palm almost straight, with fringe of hair arising from distal third reaching to end of pollex; both fingers curved inward to tip. In decreasing order, relative length of walking legs 3 > 2 > 4 > 1. Third slightly longer then second, latter a little longer than fourth. Legs 1 to 3 of similar shape, their height increasing towards third leg, fourth being the most slender. First three pairs of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON walking legs with margins of propodi con- verging distally, those of fourth pair sub- parallel; relative length of above-mentioned propodi similar to those of walking legs. Dactyli acute and curved at tip, in decreas- ing order the relative length is 4 > 3 > 2 le Abdomen with seven free somites, cov- ering the sternum and not reaching to buccal cavity, fourth somite widest, fifth longest. Male. —Carapace subpentagonal (Fig. 1B), as long as wide, occasionally a little wider than long, regions ill-defined, dorsal surface even, with short, separate setae. With ex- ception of posterior margin in small speci- mens, carapace margins with closely-set short hair-like setae. Frontal area from widely arcuate to subrectangular, with me- dial sulcus covered with setae. Cervical depressions similar to the female, but shal- lower. Eyes dorsally visible. Antennular grooves filling ventral surface of frontal re- gion. Antenna small but visible. Third maxilliped covered with feathery setae; similar to that of female except for relatively wider merus (Fig. 2D). Chelipeds stout, longer than first leg, me- rus and carpus evenly, with hair; chela as long as, or longer than, the preceding articles together; palm widening suddenly in prox- imal third, dorsal margin almost convex, ventral one rather straight, both with fringe of hair-like setae; palm surfaces covered with scattered hair, inner surface inflated; dac- tylus curved, with dorsal fringe of hair and proximal, acute and triangular tooth on cut- ting edge; pollex without hair but with small blunt teeth along cutting surface. Fingers curved inwardly to tip. Walking legs similar in shape, surface with small and scattered hair-like setae; dorsal and ventral margins of meri, propodi and dactyli with fringe of short and very close setae; legs 2 and 3 with an additional fringe of natatory setae. Propodus spatulate, dac- tylus curved, more slender than preceding articles. In decreasing order relative length VOLUME 103, NUMBER 2 367 Fig. 1. Calyptraeotheres granti (Glassell, 1933): A, female; B, male, right side without hair-like setae. of walking legs 2 > 3 > 4 > 1; fourth leg reaching to proximal third of propodus of lee Be Abdomen (Fig. 2E) with tomentum on margin, widest at third somite, narrowing towards triangular seventh somite; fourth somite longest. Gonopods (Fig. 2F) disto- laterally acute, curved the tip, forming right angle. Taxonomic remarks. —Calyptraeotheres 368 Z | PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 103, NUMBER 2 granti was incorrectly assigned to the genus Fabia. This species has two longitudinal depressions on the carapace arising behind the orbital margin. These differ from the longitudinal sulci of Fabia, which invari- ably arise from the upper margin of the orbit (see fig. 54a in Rathbun 1918). In C. granti the palp of the outer maxilliped is composed of two articles, the carpus being longer and wider than the digitiform propodus. In all species of Fabia the palp of the outer max- illiped has three articles, the carpus being shorter or subequal than the trapezoidal propodus and the dactylus inserted invari- ably in the middle of the ventral margin of the propodus. In addition, the abdomen of the male in C. granti has seven free somites. In contrast males of Fabia sensu stricto have two or more abdominal somites fused. As regards to the male’s abdominal morphol- ogy in the genus Fabia, Gore (1986) pointed out that F. subquadrata and F. concharum have seven free abdominal somites. This supposition herein is considered as a mis- interpertation of Davidson’s (1968) figures. Rathbun (1918:86) describing the male and the hard stage female of F. subquadrata [as Pinnotheres choncharum (Rathbun), see Wells (1928) and Davidson (1968) for cor- rections] noted that the abdomen of both adult males and inmature females possesses the fourth and fifth somites partially fused, contradicting Gore’s (1986) interpretation. With respect to F. concharum, examination of two hard stage females from the Allan Hancock Foundation (males were not found in the United States National Museum or elsewhere) revealed that abdominal somites 2—4 are fused. Previous accounts about the morphology of the stages of development in Pinnotheridae (Christensen & Mc- —_— 369 Dermott 1958, Pearce 1966, Jones 1977, Campos 1989b) have pointed out that an adult or hard stage male and an immature or hard stage female are almost identical morphologically; this applies to the male and inmature female of Fabia subquadrata (Rathbun, 1918:86). These observations permit me to infer that males of F. con- charum have, like the hard stage female, the abdominal somites 2—4 fused. This infer- ence, which must be confirmed, in my opin- ion is more believable than the interpreta- tion made by Gore (1986). His are based neither on specimens nor in descriptions of the above-mentioned species of Fabia but on the Davidson’s (1968) figures, which do not reflect adequately the morphology of the male abdomen in both F. subquadrata and F. concharum. Additional data that support my above-mentioned inference about the adbominal fusion in F. concharum is that males of F. emiliai (De Melo, 1971) (per- haps a junior synonym of F. byssomiae Say, 1818), F. tellinae Cobb, 1973 and a new species of Fabia from the Golfo de Califor- nia have two or more abdominal somites fused. This is similar to the condition ob- served in males and females in the hard stage of F. subquadrata and females in the hard stage of F. concharum. The remaining four species of the genus Fabia, F. canfieldi Rathbun, 1918, F. sebastianensis Rodrigues da Costa, 1970, F. obtusidentata Dai, 1980, and F. delderi Gore, 1986 are incorrectly or questionably assigned to Fabia (Campos, in prep). Ecological remarks.—C. granti is com- mon on the intertidal area of the peninsular coast of the upper Golfo de California. However, the greatest number was found in Laguna Percebu area. There I have recorded Fig. 2. A, D-F: Calyptraeotheres granti (Glassell, 1933). B: Pinnotheres garthi Fennuci, 1975, third maxilliped of male. C: P. politus (Smith, 1870), third maxilliped of male. A, third maxilliped of female; D, third maxilliped of male; E, Abdomen of male; F, first pleopod of male. A, slightly modified from Glassell (1933); B, redrawing from Fennuci (1975); C, redrawing from Garth (1957). 370 infestation up to 35%. According to Glassell (1935), C. granti was found as symbiont of Crucibulum spinosum (Sowerby, 1824), Crepidula nieva C. B. Adams, 1852 (= C. striolata Menke, 1851), and Acmaea me- soleuca Menke, 1851 [= Nomaeopelta me- soleuca (Menke, 1851)]. Although I have collected the above three species of limpets, I found C. granti living in C. spinosum only, a species that I consider as the preferred host. My conclusion is based on the fact that C. spinosum 1s the only of the above species of limpets that possesses a suitable space between the cephalic area and the shell for the growth and development of C. granti. The other two species, which have the ce- phalic-shell area very reduced, perhaps may harbor occasionally and temporarily very young stages of the crabs. This agrees with the observations recorded by Glassell (1935). Acknowledgments I am indebted to Raymond B. Manning and Alma Rosa de Campos for the support given to my work on systematics of Pin- notheridae. My great appreciation is due to Lon McClanahan, Roger Seapy and Lady Janie Yeo Kirk who helped me out during my stay at California State University, Ful- lerton. My gratitude is for George Steyskal who assisted me in the derivation of the generic name Calyptraeotheres. This inves- tigation was sponsored by the project “‘Sis- tematica y Bioecologia de los Arthropoda dominantes en areas selectas del municipio de Ensenada” of the Escuela Superior de Ciencias, Universidad Autonoma de Baja California and by agreement SEP-UABC 089-01-0352. Voucher material has been deposited in the following institutions: Es- cuela Superior de Ciencias, Universidad Autonoma de Baja California (uncata- loged); National Museum of Natural His- tory (Smithsonian Institution); School of Zoology, University of New South Wales; and San Diego Museum of Natural History. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Balss, H. 1957. Decapoda, VIII: Systematik, in H. G. Bronn, Klassen und Ordungen des Tier- reichs, Band 5, Abteilung 1, 7(12):1505-1672. Birger, O. 1895. Ein Beitrag zur Kenntniss der Pin- notherinen.— Zoologische Jahrbiicher, Abthei- lung fiir Systematik, Geographie und Biologie der Thiere 8:361-390. Campos, E. 1989a. Comments on taxonomy of the genus Orthotheres Sakai, 1969 (Crustacea, Brachyura, Pinnotheridae).— Bulletin of Marine Science 44(3):1123-1128. 1989b. Tumidotheres, a new genus for Pin- notheres margarita Smith, 1869 and Pinnothe- res maculatus Say, 1818 (Brachyura, Pinnothe- ridae). — Journal of Crustacean Biology 9(4):672— 679. Campos-Gonzalez, E., & L. J. Macias-Chavez. 1988. Fases posplanctonicas de Petrolisthes armatus (Gibbes) (Decapoda, Porcellanidae) comensales con la lapa Crucibulum (Crucibulum) spinosum (Sowerby) (Gastropoda, Caliptraeidae) en el alto Golfo de California, Mexico.— Revista de Bio- logia Tropical 35(2):241-244. Christensen, A. M., & J. J. McDermott. 1958. Life history and biology of the oyster crab, Pinnothe- res ostreum Say.—Biological Bulletin 114(2): 146-179. Cobb, S. P. 1973. Fabia tellinae, a new species of commensal crab (Decapoda, Pinnotheridae) from the northeastern Gulf of Mexico. —Crus- taceana 25(1):70-74. Dai, Ai-yun. 1980. Fabia obtusidentata Dai, sp. nov. in Ai-yun Dai, Zhong-qi Feng, Yu-zhi Sung, & Guo-xiao Chen, New species and new records of family Pinnotheridae from Hainan Island. — Acta Zootaxonomica Sinica 5(2):129-142 (In Chinese with English summary). Davidson, E. S. 1968. The Pinnotheres concharum complex (Crustacea, Decapoda, family Pin- notheridae).— Bulletin Southern California Academy of Sciences 67(2):85-88. De Melo, G. A. S. 1971. Duas novas espécies de Pinnotheridae (Crustacea, Brachyura) do litoral brasileiro.— Papeis Avulsos de Zoologia 23(22): 197-203. Fennuci, J. L. 1975. Los cangrejos de la familia Pin- notheridae del litoral argentino (Crustacea, De- capoda, Brachyura).—Physis, seccion A 34(88): 165-184. Garth, J. S. 1957. The Crustacea Decapoda Brachy- ura of Chile.—Reports of the Lund University Chile Expedition 1948-49 29:1-127. Glassell, S. A. 1933. Description of five new species of Brachyura collected on the west coast of Mex- VOLUME 103, NUMBER 2 ico.— Transactions of the San Diego Society of Natural History 7(28):331-334. 1934. Affinities of the brachyuran fauna of the Gulf of California.—Journal of the Wash- ington Academy of Sciences 24(7):296-302. 1935. New or little known crabs from the Pacific coast of northern Mexico.—Transac- tions of the San Diego Society of Natural His- tory 8(14):91-106. 1938. New and obscure decapod Crustacea from the west American coast.— Transactions of the San Diego Society of Natural History 8(33): 411-454. Gore, R.H. 1986. Fabia felderi species novum, a new pinnotherid crab from the central eastern coast of Florida (Crustacea: Decapoda: Brachyura).— Northeast Gulf Science 8(2):143-148. Griffith, H. 1987. Taxonomy of the genus Dissodac- tylus (Crustacea: Brachyura: Pinnotheridae) with description of three new species.— Bulletin of Marine Science 40(3):397—422. Jones, J.B. 1977. Post-planktonic stages of Pinnothe- res novaezelandie Filhol, 1886 (Brachyura: Pin- notheridae).—New Zealand Journal of Marine and Freshwater Research 11(1):145-158. Pearce, J. B. 1966. The biology of the mussel crab, Fabia subquadrata from the waters of the San Juan Archipelago, Washington.—Pacific Sci- ence 20(1):3-35. Pregenzer, C., Jr. 1979. A redescription of Pinnothe- res hickmani (Guiler) and comparison with Pin- notheres novaezelandiae Filhol and Pinnotheres pisum (L.) (Decapoda Brachyura).—Crusta- ceana Supplement 5:22-30. 1988. A redescription of Pinnotheres holo- thuriensis Baker, 1907 and Pinnotheres subglo- bosus Baker, 1907 with a reassignment to the genus Ostracotheres (Decapoda, Brachyura).— Crustaceana 55(1):17—28. 371 Rathbun, M. J. 1918. The grapsoid crabs of Amer- ica.— United States National Museum Bulletin 97:1-461. Roberts, M. H., Jr. 1975. Description of a pea crab, Pinnotheres chamae, sp. nov. from the jewel box, Chama congregata. —Chesapeake Science 16(4):238-241. Rodrigues da Costa, H. 1970. As species brasileiras da familia Pinnotheridae (Crustacea, Reptantia) com descricao de uma nova espécie (Fabia se- bastianensis).—Trabalhos Oceanograficos, Uni- versidade Federal da Pernambuco (Recife), 9/11 [1967/1969]:255-264. Schmitt, W. L., J. C. McCain, & E.S. Davidson. 1973. Decapoda I, Brachyura I. Family Pinnotheridae, in H.-E. Gruner & L. B. Holthuis, eds., Crus- taceorum Catalogus, 3:1-160, W. Junk, Den Haag. Seréne, R. 1967. Sur deux espéces nouvelles de brachyoures (Crustacés Décapodes) et sur une troisiéme peu conne, récoltées dans la région Malaise. — Bulletin du Muséum National d’His- toire Naturelle, Paris 38(6):8 17-827. Tesch, J. J. 1918. Goneplacidae and Pinnotheridae. The Decapoda of the Siboga Expedition II.— Siboga-Expeditie 39 c!:1-295. Wells, W. W. 1928. Pinnotheridae of Puget Sound— Publications Puget Sound Biological Station, University of Washington 6:283-314. Department of Biological Sciences, Cal- ifornia State University Fullerton, Fuller- ton, California 92634; (Postal address) Es- cuela Superior de Ciencias, Universidad Autonoma de Baja California, Apartado Postal 2300, Ensenada, Baja California, México. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 372-375 MESACTURUS DICRURUS, NEW SPECIES, AN UNUSUAL STOMATOPOD FROM MICRONESIA (STOMATOPODA: GONODACTYLIDAE) Roy K. Kropp and Jane H. Dominguez Abstract. —Mesacturus dicrurus, the third species in the genus, is described. The new species is distinguished from other species in the genus by having an apically bifurcate median projection of the telson that is about as long as the base of the telson and that is covered with relatively long setae. The apical bifurcation is V-shaped. Manning (1969), in a discussion of the Gonodactylus-like stomatopods in the Gon- odactylidae, presented a diagnosis and dis- cussion of the genus Mesacturus and a key to species then included in the genus. He noted that there were two general types of telson morphology in the genus, in one the telson is similar to that of Gonodactylus, whereas in the other the telson is charac- terized by a median bifurcate process. Later, Manning (1978) erected a new genus, Mes- acturoides, to include the species of Mes- acturus having Gonodactylus-like telsons. Two species, Mesacturus furcicaudatus (Miers, 1880) and M. kempi (Odhner, 1923), remained in Mesacturus. Among crustaceans collected from con- solidated coralline algae at the seaward mar- gin of an erosion bench at Guam, Mariana Islands, we found specimens of Mesacturus not referable to either known species. Fur- ther examination of stomatopods collected at Saipan, Mariana Islands, by the senior author revealed specimens of the new species. We examined stomatopods in the collection of the National Museum of Nat- ural History, Smithsonian Institution (USNM) that were collected from Yap, Car- oline Islands, and had been identified as M. kempi and found them to be referable to the new species. Primary types have been deposited in the USNM, Bernice P. Bishop Museum, Ho- nolulu (BPBM), and the Natural History Museum of Los Angeles County, Los An- geles (LACM), as indicated in the material examined. We thank the University of Guam Ma- rine Laboratory and the Division of Crus- tacea, USNM, for support and R. B. Man- ning for reviewing the manuscript. This is Contribution No. 278 from the University of Guam Marine Laboratory. Family Gonodactylidae Giesbrecht, 1910 Mesacturus dicrurus, new species Figs. 1, 2 Material.—Holotype: Mariana Islands: Guam; Pago Bay, Taogam Point [13°25’N, 144°48’E]; from consolidated coralline al- gae at seaward edge of erosion bench; 7 June 1986; 1 male (USNM). Paratypes: Mariana Islands: Guam; same locality and habitat as holotype; 3 Sep 1984; 3 females (USNM, LACM). Saipan; sand + coral; 25 Dec 1945; Coll. A. H. Banner; 1 male, 1 female (USNM). Saipan; SSW side of island; in coral heads; 1945; Coll. A. H. Banner; 2 males (USNM). Saipan; Wing Beach, near Matuis [15°16’N, 145°48’E]; from reef rock bearing coralline algae col- lected just shoreward of reef margin; 0.5 m; 21 Nov 1980; 3 females (BPBM). Caroline Islands: Yap; Ifalik (formerly Ifaluk) Atoll; south end of Falarik Islet VOLUME 103, NUMBER 2 Fig. 1. 373 Dyce) F TN j IN Mesacturus dicrurus, male holotype: a, Anterior part of body; b, Sixth abdominal somite, telson (dorsal view); c, Same (lateral view); d, Left claw; e, Left uropod (ventral view); f, Median projection of telson, showing setae. Scale: 1 mm. [O7°15'N, 144°27’E]; Sta. 13-C-3, 14-B-1, 19-D, 20-C, 706; 25 Aug 1953; Coll. D. P. Abbott and Bates, 4th Pacific Atoll Survey Team, Pacific Science Board; 5 males, 2 fe- males, 1 juvenile (USNM 104692, 104694— 7). Ifalik Atoll; north end of Falarik; Sta. 359), SIZ, OBS Ail, ZA Seo, 7 Ore WOS3e Coll. F. M. Bayer; 3 males, 2 females (USNM 104689-91). Description of holotype. —Eyestalks cylin- drical, slightly inflated proximally, length about twice width. Ocular scale small, blunt, erect. Anterior margin of rostral plate concave, anterolateral angles bluntly rounded or acute, but not spiniform, width of plate greater than median length, anterior spine twice as long as basal part of plate. Dactylus of raptorial claw serrated on dis- tal half of inner margin. Opposable margin 374 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Mesacturus dicrurus, juvenile paratype, total length 12.9 mm: a, Eyes, rostral plate; b, Sixth abdominal somite, telson (dorsal view). Female paratype, total length 33.3 mm (USNM 104692): c, Eyes, rostral plate. Scale: a = 1 mm, b = 0.5 mm, c = 2 mm. of propodus with 10 articulated teeth along midlength, with 4 rounded bosses proxi- mally fitting into corresponding sockets on dactylus. Anterior 5 abdominal somites smooth. Dorsal surface of sixth abdominal somite with 4 swollen, unarmed carinae, subme- dian carinae confluent medially. Telson slightly wider than long, posterior medial projection half total length of telson, narrowing slightly distally, with V-shaped fork at % projection length, surface covered with very stout simple setae. Dorsal surface of telson with 3 broadly inflated carinae, median largest. Posterior margin with 4 clusters of 4-5 dorsally curved teeth, base of median projection with 5 irregularly placed teeth. Base of uropod with single lobe or spine distally near inner margin, outer margin of proximal segment of exopod with 7 spines, distal 3 recurved. Variations. —Some of the paratypes dif- fered slightly from the holotype. Variation was noticed in the shape of the rostral plate. In some specimens from Falarik the ante- VOLUME 103, NUMBER 2 rior margin of the plate was very concave and the anterolateral angles of the plate more sharply pointed than in specimens from the Marianas (compare Fig. 2c to Fig. la). The number of teeth on the margin of the propo- dus of the raptorial claw ranged from 6 to 13. Size-related variation in the develop- ment of carinae on the sixth abdominal seg- ment and the telson, spination of the pos- terior margin of the telson, and the degree of setal development on the median pro- jection of the telson was noticeable. In larger individuals the carinae were much more swollen than in small individuals (compare Fig. 1b to Fig. 2b). In large individuals the posterior margin of the telson had 4 clusters of 4—5 teeth, but in small individuals the posterior margin had 6 individual teeth. Small individuals had fewer setae on the median projection of the telson than large specimens. The number of teeth on the out- er margin of the proximal segment of the uropod varied from 7 to 8. The larger num- ber resulted from the presence of an addi- tional very small tooth proximally. Measurements.— Total length of males 11.4 to 33.2 mm, of females 10.7 to 33.3 mm. Measurements of holotype: total length 22.4 mm; carapace length 4.6 mm; eyestalk length 1.9 mm; cornea length 0.9 mm; ros- tral plate length 1.2 mm, width 1.6 mm; sixth abdominal somite width 3.6 mm; tel- son length (excluding median projection) 1.6 mm, width 3.6 mm; median projection of telson length 1.4 mm. Color. —Body brown fading to tan at base of telson, median process of telson olive- green. Antennae, walking leg, maxillipeds, pleopods, and uropods pale olive-green. Meral spot clear with black spot anteriorly. Habitat. —Guam specimens were collect- ed from consolidated coralline algae at sea- ward edge of erosion bench. Saipan speci- mens were collected from coralline- encrusted reef rock just shoreward of fring- 375 ing reef margin or from coral heads. Yap specimens from Station 706 (USNM 104692) were collected from inside coral heads on the outer slope of the reef plat- form. The bathymetric range of the new species is from the intertidal zone to 0.5 m. Remarks. — The telson of Mesacturus fur- cicaudatus (Miers, 1880) differs from that of the new species and M. kempi (Odhner, 1923) by having a deeply bifurcate median projection; the median projection is shal- lowly bifurcate in the latter two species. The new species differs from M. kempi in the relative lengths of the base of the telson and the median projection. In M. dicrurus the median projection is about as long as the base of the telson, whereas in VM. kempi the median projection is about twice the length of the base. Also, the median projection is covered by relatively long setae in the new species, but in M. kempi the projection is covered by relatively short setae. The bi- furcation of the median projection of the telson is broadly concave in M. kempi, but V-shaped in the new species. Etymology. —From the Greek dikros, meaning forked, in combination with oura, meaning tail, in reference to the forked me- dian projection of the telson. Distribution. —Mesacturus dicrurus is known only from Yap, Guam, and Saipan. Literature Cited Manning, R. B. 1969. Notes on the Gonodactylus section of the family Gonodactylidae (Crusta- cea, Stomatopoda), with descriptions of four new genera and a new species.— Proceedings of the Biological Society of Washington 82:143-166. . 1978. Anew genus of stomatopod crustacean from the Indo-West Pacific region.— Proceed- ings of the Biological Society of Washington 91: 1-4. Battelle Ocean Sciences, 1431 Spinnaker Drive, Ventura, California 93001. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 376-379 THE PRIMARY TYPES OF THE RICHARD M. BOHART MUSEUM OF ENTOMOLOGY, I. TARDIGRADA R. O. Schuster, T. L. Tyler, J. A. Skinner, and E. A. Sugden Abstract. —This paper lists the primary types of Tardigrada deposited in the Richard M. Bohart Museum of Entomology, University of California, Davis, California (UCD). A brief history of the museum is also given. This is the first in a series of articles listing the primary type specimens in the collection of the Richard M. Bohart Museum, De- partment of Entomology, University of Cal- ifornia, Davis, California (UCD). All of the primary types listed here are holotypes. The Bohart Museum is presently responsible for 1438 primary types. Infor- mation was compiled from type labels, orig- inal descriptions, field notes and reexami- nation of type specimens. Names listed for types and host species are as in the original description of the type. Changes in the name ofa host species before original publication of the type are noted. Zoological or botanical synonomies are not established in this paper. History of the Bohart Museum Collections The research collections of the Depart- ment of Entomology, University of Cali- fornia, Davis were established in 1946. Mr. A. T. McClay, the first curator, enlarged the holdings from a Schmidt box each of Cal- liphoridae and honey bees to about 600,000 specimens, housed in California Academy style cabinets. The collection was particu- larly rich in Coleoptera donated from his personal collection. Mr. McClay retired in 1966 and mana- gerial responsibility was assumed by Mr. Robert O. Schuster. At that time the deci- sion was made to maintain a type collection, and to broaden holdings of non-insectan ar- thropods because an invertebrate collection did not exist at the University of California, Davis. Prior to 1966 primary types were deposited with the California Academy of Sciences. The collections have grown partly through active California-based field research pro- grams, and the result is a rich selection of material from the Western United States and Mexico. In addition, the research in- terests of faculty and graduate students have led to significant cosmopolitan holdings in many taxonomic groups. In recognition of the rapid destruction of neotropical habi- tats, the collection and preparation of spec- imens from northern South America, Cen- tral America, and the West Indies is a high priority of the museum. The various collections of insects and re- lated arthropods were officially dedicated on 11 October 1983 as the Richard M. Bo- hart Museum of Entomology. The museum holds in excess of four million prepared specimens, includes from 50,000 to 100,000 uncataloged allotypes and paratypes, and is supported by the Department of Entomol- ogy and the Bohart Museum Society, found- ed in 1986. The acronym used for the Rich- ard M. Bohart Museum of Entomology is UCD. This list, the Tardigrada, contains 21 types resulting from the studies of Albert A. Gri- garick, Robert O. Schuster, and Elizabeth C. Toftner at UC Davis, Donald S. (Woody) Horning, Jr., presently at Mcleay Museum, VOLUME 103, NUMBER 2 University of Sydney, Australia, the late Franz Mihelcic, and Diane R. Nelson at East Tennessee State University. The tardigrade collection comprises 60,000 slides and is cosmopolitan with specimens from all con- tinents excluding Asia. We felt it appropri- ate to list the Tardigrada separately from the rest of the primary types, as these in- vertebrates are not currently thought to be arthropods. The Bohart Museum Tardigrada Type Collection Class Heterotardigrada Family Oreellidae Oreella breviclava Grigarick, Schuster, & Nelson, 1983:66. Holotype is a female, UCD 1269; type locality La Carbonera, Campo Elias, Merida, Venezuela, 2000 ft, collected by R. W. Brooks, A. A. Grigar- ick, J. McLaughlin, & R. O. Schuster on 30 Jun 1979. Family Echiniscidae Bryodelphax crossotus Grigarick, Schuster, & Nelson, 1983:69. Holotype is a female, UCD 1270, from Rancho Grande, Ara- gua, Venezuela, 1130 m, collected by R. W. Brooks, A. A. Grigarick, J. Mc- Laughlin, & R. O. Schuster on 14 Jul 1979. Echiniscus (Bryodelphax) dominicanus Schuster & Toftner, 1982:225. Holotype is a female, UCD 1215, from Constanza, La Vega Province, Dominican Republic, collected by R. O. Schuster on Groutiella mucronifolia (Hooker & Greville, 1824: 116) Crum & Steere 1950:146. Crum & Anderson 1981:738 state that ““The type of Orthotrichum mucronifolium Hook. & Grev. 1824, collected by Guilding in the West Indies, is clearly the same as the Mexican type of O. apiculatum Hook. 1818, collected by Humboldt and Bon- pland. It is obvious, therefore, that Grou- tiella mucronifolia belongs in the synon- ymy of G. apiculata.” The label data for 377 the host plant for E. (Bryodelphax) do- minicanus 1s probably incorrect, and Groutiella apiculata (Hooker, 1818:plate 45) Crum & Steere 1950:146 is the ap- propriate name for this moss. Echiniscus aliguantilus Grigarick, Schuster, & Nelson, 1983:73. Holotype is a female, UCD 1266, from La Carbonera, Campo Elias, Merida, Venezuela, collected by R. W. Brooks, A. A. Grigarick, J. Mc- Laughlin & R. O. Schuster on 30 Jun 1979. Echiniscus becki Schuster & Grigarick, 1966: 127. Holotype sex is uncertain, UCD 209, from Mountaineer Mine, south of Vidal Junction, Riverside Mountains, River- side County, California, collected by A. J. Beck & R. O. Schuster. Echiniscus cavagnaroi Schuster & Grigar- ick, 1966:321. Holotype sex is uncertain, UCD 94, from Old Bella Vista trail, Santa Cruz Island, Galapagos Islands, Ecuador, 90 m, collected by R. O. Schuster, on 4 Feb 1964, in mixture of Frullania sp., Parmelia sp., and Ramalina sp. Echiniscus horningi Schuster & Grigarick, 1971:105. Holotype sex is uncertain, UCD 322, from Silver Falls State Park, Marion County, Oregon, collected by D. S. (Woody) Horning, Jr., on 10 Jan 1970. Echiniscus knowltoni Schuster & Grigarick, 1971:109. Holotype sex is uncertain, UCD 401, from Black Pine, Oneida County, Idaho, collected by G. F. Knowlton on 29 Sep 1969. Echiniscus kofordi Schuster & Grigarick, 1966:321. Holotype sex is uncertain, UCD 95, from Darwin Research Station, Santa Cruz Island, Galapagos Islands, Ecuador, less than 10 m, collected by R. O. Schus- ter on 21 Jan 1964, on Ramalina sp. Echiniscus (Echiniscus) laterculus Schus- ter, Grigarick, & Toftner, 1980:265. Ho- lotype is a female, UCD 1264, from Riv- erton, El Dorado County, California, collected by E. C. Toftner & R. O. Schus- ter, on 1 Feb 1974. Echiniscus marginoporus Grigarick, Schus- ter, & Nelson, 1983:76. Holotype is a fe- 378 male, UCD 1268, from La Mucuy, Cam- po Elias, Merida, Venezuela, 2000 m, collected by R. W. Brooks, A. A. Grigar- ick, J. McLaughlin, & R. O. Schuster, on 30 Jun 1979. Echiniscus mosaicus Grigarick, Schuster, & Nelson, 1983:76. Holotype is a female, UCD 1267, from La Carbonera, Campo Elias, Merida, Venezuela, collected by R. W. Brooks, A. A. Grigarick, J. Mc- Laughlin, & R. O. Schuster, on 30 Jun 1979. Echiniscus robertsi Schuster & Grigarick, 1965:56. Holotype sex is uncertain, UCD 93, from bog at Sand Point, Popof Island, Alaska, collected by Warren G. Roberts, in mid June 1963. Parechiniscus armadilloides Schuster, 1975: 335. Holotype is a female, UCD 904, from Logan Canyon near Logan, Cache Coun- ty, Utah, collected by D. R. Miller & A. S. Menke, on 5 Apr 1973. Pseudechiniscus goedeni Grigarick, Mihel- cic, & Schuster, 1964:5. Holotype sex is uncertain, UCD 90, from 6 mi SSW of Breitenbush Hot Springs, Marion Coun- ty, Oregon, collected by K. Goeden, on 18 Oct 1962, in lichens on Douglas fir and cedar. Pseudechiniscus raneyi Grigarick, Mihelci¢, & Schuster, 1964:6. Holotype sex is un- certain, UCD 91, from 46 mi E of Fresno on Hwy 189, Tulare County, California, 4300 ft, collected by A. A. Grigarick, on 31 Oct 1962, in mixed lichen and moss on pine. Class Eutardigrada Family Hypsibiidae Hypsibius (Diphascon) iltisi Schuster & Gri- garick, 1965:32. Holotype sex is uncer- tain, UCD 92, from Mt. St. Helena, Napa County, California, collected by J. S. Buckett & M. E. Irwin, on 13 Mar 1962, on a rotting cone of Pseudotsuga menzie- sil (Mirbel, 1825:63,70) Franco 1950:74. Tsohypsibius saltursus Schuster, Toftner, & PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Grigarick, 1977:126. Holotype is a fe- male, UCD 960, from Pope Beach, Lake Tahoe, El Dorado County, California, collected by R. O. Schuster & E. C. Toft- ner, on 10 Oct 1975. Family Macrobiotidae Macrobiotus grandipes Schuster, Toftner, & Grigarick, 1977:118. Holotype is a fe- male, UCD 959, from Pope Beach, Lake Tahoe, El Dorado County, California, collected by R. O. Schuster & E. C. Toft- ner, on 12 Sep 1975. Macrobiotus tridigitus Schuster, 1983:254. Holotype sex is uncertain, UCD 1217, from Sierra Martial, Tierra del Fuego, Ar- gentina, 2000-2500 ft, collected by A. M. Shapiro, on 19 Jan 1979, at tree line, in cryptograms. Pseudodiphascon dubium Schuster & Toft- ner, 1982:228. Holotype sex is uncertain, UCD 1216, from Cabrera, Sanchez Prov- ince, Maria Trinidad, Dominican Repub- lic, collected by R. O. Schuster, on 1 Aug 1978, on Lejeunea sp. Acknowledgments The authors thank R. M. Bohart for com- ments on the manuscript, G. L. Webster and K. A. Chambers for assistance with botan- ical nomenclature. Literature Cited Crum, H. A., & L. E. Anderson. 1981. Mosses of eastern North America, volume 2. Columbia University Press, New York, New York, 663 pp. ——, & W.C. Steere. 1950. Additions to the moss flora of Panama.—The Bryologist 53(2):139- 152. Franco, J.D. A. 1950. Cedrus libanensis et Pseudo- tsuga menziesii. —Boletim da Sociedad Broteri- ana, 2 serie, 24:73-77. Grigarick, A. A., F. Mihel¢ié, & R. O. Schuster. 1964. New Tardigrada from western North America: I. Pseudechiniscus. — Proceedings of the Biolog- ical Society of Washington 77:5-8. , R. O. Schuster, & D. R. Nelson. 1983. Het- VOLUME 103, NUMBER 2 erotardigrada of Venezuela. — Pan-Pacific Ento- mologist 59(1—4):64—77. Hooker, W. J. 1818. Musci exotici. Containing fig- ures and descriptions of new or little known foreign mosses and other cryptogamic subjects, volume |. Longman, Hurst, Rees, Orme and Brown, London, viii + pl. 1-96. —., & R. K. Greville. 1824. Sketch of the char- acters of the species of mosses belonging to the genera Orthotrichum (including Schlotheimia, Micromitrion and Ulota), Glyphomitrion and Zygodon.— Edinburgh Journal of Science 1:110— 133. Mirbel, M. 1825. Essai sur la distribution géogra- phique des coniféres.— Mémoires du Muséum National d’Historie Naturelle, Paris 13:28-76. Schuster, R.O. 1975. A new species of Parechiniscus from Utah (Tardigrada: Echiniscidae).— Mem- orie dell’Istituto Italiano di Idrobiologia—Ver- bania Pallanza 32(Supplemento): 105-110. 1983. A new species of Macrobiotus from Tierra del Fuego (Tardigrada: Macrobioti- dae).— Pan-Pacific Entomologist 59(1—4):254— 255. , & A. A. Grigarick. 1965. Tardigrada from western North America with emphasis on the fauna of California.—University of California Publications in Zoology 76:1-67. ——, & 1966a. Tardigrada from the Ga- lapagos and Cocos Islands. — Proceedings of the California Academy of Sciences 34:315-328. ——_., & . 1966b. New Tardigrada from west- erm North America: II. Echiniscus.—Proceed- 379 ings of the Biological Society of Washington 79: 127-130. ——., & . 1971. Two new species of Echinis- cus from the Pacific Northwest (Tardigrada: Echiniscidae).— Proceedings of the Entomolog- ical Society of Washington 73(1):105-110. , A. A. Grigarick, & E. C. Toftner. 1980. A new species of Echiniscus from California (Tar- digrada: Echiniscidae).— Pan-Pacific Entomol- ogist 56(4):265-267. —., & E.C. Toftner. 1982. Dominican Republic Tardigrada. Pp. 221-236 in D. R. Nelson, ed., Proceedings of the Third International Sym- posium on Tardigrada, East Tennessee State University Press, Johnson City. , E. C. Toftner, & A. A. Grigarick. 1977. Tar- digrada of Pope Beach, Lake Tahoe, Califor- nia.— Wasmann Journal of Biology 35(1):333- 336. (ROS! & TLT) Department of Entomol- ogy, University of California, Davis, Cali- fornia 95616; (JAS) Department of Ento- mology and Plant Pathology, University of Tennessee, Knoxville, Tennessee 37901; (EAS) USDA-ARS Honeybee Lab, 507 W. 4th St., Weslaco, Texas 78596. 1 Mr. Robert O. Schuster unexpectedly died on 15 August 1989, after this paper was submitted. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 380-426 TWO NEW SPECIES OF MIMAGONIATES (TELEOSTEI: CHARACIDAE: GLANDULOCAUDINAE), THEIR PHYLOGENY AND BIOGEOGRAPHY AND A KEY TO THE GLANDULOCAUDIN FISHES OF BRAZIL AND PARAGUAY Naércio A. Menezes and Stanley H. Weitzman Abstract.—Two new species of freshwater glandulocaudine characid fishes are described from coastal streams of eastern Brazil. The first, Mimagoniates sylvicola, is from southeastern Bahia, the second, Mimagoniates rheocharis, from southeastern Santa Catarina and northeastern Rio Grande do Sul. Mima- goniates sylvicola is hypothesized to form an unresolved trichotomy with Mi- magoniates lateralis and a monophyletic line leading to the more derived Mimagoniates microlepis and Mimagoniates rheocharis. Mimagoniates rheo- charis most parsimoniously appears to be a sister species of M/. microlepis but its phylogenetic relationships may be quite complex and its possible introgres- sion with M. inequalis and/or its origin by introgression between M. inequalis and M. microlepis is discussed. Solution to this problem awaits additional! data based on more appropriate population samples than available to us and on genetic information. The biogeography of all these species is briefly discussed on the basis of the limited available phylogenetic information. The descriptions of the new fish species herein continue a series of contributions to the knowledge of the fauna of the streams of the east coast of Brazil. As discussed by Weitzman et al. (1986:344—345), the fresh- water fish fauna of the coastal or littoral streams of eastern Brazil appears to contain many new species, a fact unrecognized until the last 15 or 20 years. The descriptions provided here are a product of a continuing study by us of the Glandulocaudinae, a characid subfamily distributed from Costa Rica south to northern Argentina in Atlan- tic drainages and to the Guyas basin in Pa- cific drainages. We describe these new fishes here because we wish to have them available for a phylogenetic analysis of the Glandu- locaudinae and especially the Glandulocau- dini in progress. The Glandulocaudini, one of apparently four tribes of the Glandulocaudinae (Weitz- man et al. 1988:383, Weitzman et al. in Weitzman & Fink 1985:112—117), currently consists of two genera, Glandulocauda Ei- genmann (1911) and Mimagoniates Regan (1907). Use of the generic names Mima- goniates, Coelurichthys Miranda-Ribeiro (1908) and Glandulocauda in the tribe has changed frequently, with little agreement among various authors. Weitzman & Fink (1985:1, 2, 109) reviewed some of the no- menclatural and phylogenetic problems re- garding the generic names of the tribe. We follow their usage until ongoing studies al- low us to evaluate the present preliminary phylogeny of the tribe based on more abun- dant data than shown in Fig. 1. The Glandulocaudini contains eight known species, including the two described here. Schultz (1959), the last author to re- view the species of the Glandulocaudini in any detail, accepted five species. He consid- ered M. lateralis (Nichols) a synonym of M. inequalis (Eigenmann). These two species VOLUME 103, NUMBER 2 eC & x 3 5 «‘ LORS i) Oy ey y i Ne) OS S & » < S O° ~ & IS eg © wd s ° es oS Sede Sache vil S & « ES e 6 * 8 ¥ 8 ‘ N wi Hooks strongly developed on caudal-fin rays 8-12. Hooks weakly developed on rays IO-II. 7 Hooks at least on caudal-fin rays |O-II. Hooks absent on caudal-fin rays. a Caudal-fin-ray pump chamber well developed. Caudal-fin-ray pump chamber poorly developed. IN Caudal-fin-ray pump present. Gland cells concentrated around pump opening. Caudal-fin-ray pump absent. Gland cells spread in rows along fin rays. NN Modified caudal-fin squamation from dorsal caudal-fin lobe. Caudal pump formed from terminal lateral-line scale or from squamation on ventral lobe. Modified caudal fin with hypertrophied glandular tissue present. Females internally fertilized in at least three tribes. Fig. 1. Diagram of tentative phylogenetic relationships of the species of the Glandulocaudini based on an analysis of synapomorphies found in the caudal skeleton of males. Filled in squares = apomorphies and empty squares = plesiomorphies at the levels indicated. are recognized as distinct and valid here. Géry (1966:228—230) constructed a key to the tribe and recognized seven species. He appeared doubtful that the group as pre- sented by Schultz or himself was mono- phyletic. Six of the species tentatively in- cluded as valid by Géry are among those accepted here. Inclusion of Géry’s seventh species, Glandulocauda terofali Géry (1964), would make polyphyletic the monophyletic Glandulocaudini as defined below. This species lacks the modified dorsal lobe cau- dal-fin squamation that is synapomorphic for the adults of both sexes of all species included by us in the Glandulocaudini. It also lacks the ventrally bowed form of cau- dal-fin rays 11 and 12 or the particular kind of modification into caudal fin-ray pump structures which form a synapomorphic morphological transformation series con- fined to members of this tribe. See the key below for a summary exposition of these glandulocaudin characters. We follow Weitzman & Fink (1985:103, 109) in as- signing Géry’s species to Diapoma Cope and accept Diapoma terofali (Géry) as a member 382 of the glandulocaudine tribe Diapomini be- cause it has the derived ventral lobe caudal- fin squamation in both sexes closely match- ing that of Diapoma speculiferum Cope. This squamation pattern (see Weitzman & Fink, 1985:fig. 15) is synapomorphic for the Dia- pomini, which also includes the species of Acrobrycon Eigenmann & Pearson and Planaltina Bohlke. The keys to the species of the Glandu- locaudini presented by Schultz (1959) and Géry (1966) were based on information from previous literature, some of the types and a few additional specimens. Because we are describing two new species and have ex- amined all of the types and have new in- formation from over 200 recently collected population samples, we provide an almost entirely new key to the species of the Glan- dulocaudini. See Weitzman et al. (1988:384— 390, figs. 5, 6) for distribution maps and a summarized comparison of the collections available to Schultz and to us. A prelimi- nary discussion of the Glandulocaudini, its phylogeny and biogeography were pre- sented by Weitzman et al. (1988:384—419). See also Weitzman & Fink (1985:98—99) for a discussion of the morphology and possible function of the fin-ray pheromone pump mechanism of the species of Mimagoniates. The two new species described below were designated as “new species A”’ (here = M. sylvicola) and “‘new species B” (here = M. rheocharis) in Weitzman et al. (1988). A third putative species, “‘new species C,”’ rec- ognized as possibly new by them, appears closely related to M. microlepis (Steindach- ner) and may be one of several relatively distinct populations of this species. The re- lationships among and distinctness of these populations is a complex problem and will require extensive analysis, research not un- dertaken here. The importance of well-documented phy- logenies to biogeographic hypotheses is em- phasized by the changes noted below in the section on biogeography. A change in our tentative understanding of the phylogenetic PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON relationships of M. sylvicola based on a pop- ulation sample unavailable to Weitzman et al. (1988) resulted in a consequent change in our hypotheses regarding the biogeo- graphic history of the species of Mimagoni- ates. Compare the tentative phylogenetic diagram presented here, Fig. 1, with that of Weitzman et al. (1988:fig. 10). The phylo- genetic and biogeographic accounts by Weitzman et al. (1988:414-419) of M. rheo- charis remain essentially unchanged, but questions are raised about of phylogenetic relationships of this species that could have extensive biogeographic consequences, de- pending on the nature of their answers. The phylogenetic relationships of all species of the Glandulocaudini need further investi- gation. Methods and Materials Counts and measurements are those de- scribed by Fink & Weitzman (1974:1-2). In the counts given in the text of the descrip- tions, the holotype is given first followed by the mean, range and number of paratypes counted given in parentheses. The posterior terminal ray of the dorsal fin is not divided to its base and is counted as one. However, the posterior terminal ray of the anal fin is so divided and thus the apparent two pos- teriormost rays of the anal fin are counted as one. Note that “branched rays” refers to all rays posterior to the anterior unbranched rays of a given fin even if, as with the pos- terior terminal ray of the dorsal and pectoral fins, the posterior rays are unbranched at their bases or their distal segments. Body depth was measured vertically (orthogonal to the longitudinal body axis) from the dor- sal-fin origin. All measurements other than standard length (SL) are expressed as a per- centage of SL except subunits of the head which are recorded as a percentage of head length. Total vertebral counts (including the Weberian apparatus) were taken from ra- diographs and from cleared Alizarin red and Alcian blue stained preparations. These VOLUME 103, NUMBER 2 preparations are called “‘cleared and stained” in the text. The terminal “‘half centrum,” hypural bones and associated vertebral ele- ments, usually designated as PU,+U,, but not necessarily consisting only of those ele- ments (Schultze & Arratia 1989:203) was counted as one vertebral element. Statistical comparisons and basic statis- tics of characters taken from population samples were computed using SYSTAT (version 4.0, Systat, Inc., 1989) and BIOM- PC (version 2, Exeter Publishing Ltd, 1988). All hypotheses of statistical significance are two-tailed. Common logarithms were used for transformations of original data used in covariance analyses. In the morphometric scatter plots, both axes are logarithmic and the regression lines are based on log trans- formed data plotted in the figures. Regres- sion equations given in the figure captions are based on logarithmic transformations of original data and correspond to their re- spective logarithmic transformed covari- ance analyses in the text. The regression lines and equations are given only to present a concept of the comparative approximations of the growth lines of sexually maturing and mature population samples within the lim- its of the data. In no instances are regression lines, regression equations or covariance analyses to be construed as predictors of body shape beyond the data shown in the appropriate plots. Logarithmic regression lines and semilogarithmic scatter plots were prepared using Sigma-Plot (version 3.10, Jandel Scientific, 1988). In cases where a value of a t-statistic was computed for com- parisons of population means of counts, square-root transformations were utilized. In the key, and abbreviated parts of the di- agnoses, the ranges, means and other sta- tistical parameters that may be given are for all of the paratypes and the holotype of each respective new species. In the statistical comparisons (statistical hypotheses of dif- ferences), ranges, means and other statisti- cal parameters are naturally limited to the population samples being compared. 383 In our discussions, comments on phylog- eny are based on the concepts of phyloge- netic systematics of Hennig (1966) as re- viewed and discussed by Wiley (1981). Maddison et al. (1984) are followed for out- group considerations. Biogeographic prin- cipals follow Humphries & Parenti (1986). Specimens examined for this study are deposited in the Museu Anchieta, Porto Alegre (MAPA), Museu de Zoologia da Universidade de Sao Paulo (MZUSP); the Smithsonian Institution, National Museum of Natural History (USNM); Museu de Ciéncias, Pontificia Universidade Catolica do Rio Grande do Sul (MCP); Museum of Zoology, University of Michigan (UMMZ) and the California Academy of Sciences (CAS). Specimens examined in detail, other than those listed or cited here, were listed in Weitzman & Fink (1985:102-108. Geographic entities. (such as rivers) and place names (except those of countries) are in the language of the country of origin. Countries are in English because there is an English name for the various countries. Therefore rio (rio in Portuguese, rio in Spanish) is not capitalized when referring to a river (thus rio Jordao and rio Grande), but it is capitalized when referring to a place name (thus Rio de Janeiro and Rio Grande do Sul). We do this because we are attempt- ing to avoid the transferal of English style and grammar onto foreign localities and geographical units. Key to the Species of the Glandulocaudini Although this key employs many char- acters that are apparently synapomorphic at their level of use, the key is not a hy- pothesis of phylogeny for any of the includ- ed taxa except for the Glandulocaudini di- agnosed in the first half (la) of the first couplet. Because the primary purpose of this key is to identify species, we use any ap- propriate character that either clusters and/ or distinguishes species. Thus, except for couplet la where the characters are synapo- 384 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON morphies, a given character may be either a plesiomorphy, a synapomorphy or an au- tapomorphy at its point of use. At this time we lack corroborated polarity hypotheses for many of the characters that are not associ- ated with the secondary sexual features of adult males and our hypotheses for the sec- ondary sexual characters cannot be consid- ered well-supported. Homoplastic charac- ters are used where they serve the purpose of the key. The reader should note that when using couplet 4, many maturing males of species of Mimagoniates that have a well- developed caudal pump chamber when ful- ly mature, will key to 4a rather than 4b (where they belong) because of the incom- plete development of the specimen’s caudal fin-ray pump. This will often be the case in relatively small and some moderate sized specimens even in the presence of appar- ently active testes. la. Adult males and females with modified caudal-fin squamation extending posteriorly onto caudal fin from base of ventral portion of dorsal caudal-fin lobe; principal caudal-fin rays 11 and 12 of adult males somewhat bowed or curved ventrally, these and adjacent rays modified into a chamber or sup- port for a chamber of a fin-ray pump organ in some species (com- pare Figs. 2-6)... .Glandulocaudini (2) 1b. Adult males and females without modified caudal-fin squamation or, if such scales present, then these derived from both caudal-fin lobes, only from ventral caudal-fin lobe or primarily from terminal lateral- line scale; principal caudal-fin rays 11 and 12 of adult males straight or if modified, not bowed ventrally (as in Fig. 6) or modified into a fin- ray pump organ (as in Figs. 2—5) although they may provide origins for muscular and ligamentous at- tachments to a scale-diaphragm 2a. 2b. 3a. 3b. PUMP HaaNe ChamiS mies ee Siphes Mat ee other tetragonopterine or glandulocaudine characid groups Caudal fin-ray pump absent in all sexes at all ages (see Fig. 6); hy- pertrophied glandular tissue wide- spread along caudal-fin rays, es- pecially those of ventral caudal-fin lobe, but most dense along proxi- mal portions of rays 11 and 12; more than | hook on each anterior anal-fin ray that bears hooks .... POR S MERE | NORTE Glandulocauda (3) Caudal fin-ray pump present in mature males, modified primarily from caudal-fin rays 10-12, some- times relatively simple and repre- sented by expanded, flattened an- terior portions of ray halves, sometimes by an anterior fin-ray pump chamber; hypertrophied glandular tissue confined to area immediately around and on caudal pump region of gland (see Figs. 2, 5); no more than | hook on any anal-fin ray that bears hooks .... peg Oa TAS Pen Yee. Mimagoniates (4) Branched anal-fin rays 20—23; per- forated lateral line scales 11-21; scale rows between dorsal-fin ori- gin and anal-fin origin 14—17; scale rows around caudal peduncle 17— 18, usually 18; dorsal-fin origin somewhat anterior to vertical line drawn from anal-fin origin ..... ..Glandulocauda melanogenys Eigen- mann (upper rio Tieté, in Sao Paulo, Brazil) Branched anal-fin rays 15-18; per- forated lateral-line scales 4—6; scale rows between dorsal-fin origin and anal-fin origin 11-13; scale rows around caudal peduncle 16; dor- sal-fin origin somewhat posterior to vertical line drawn from anal- finyOpisini | Wie aN eae ary iene operas Glandulocauda melanopleura Eigenmann VOLUME 103, NUMBER 2 385 (upper rio Iguacu, in Parana and Santa Catarina, Brazil) anal-fin origin 13-15; dorsal-fin origin at line drawn vertically from 4a. Caudal fin-ray pump little devel- base of seventh to ninth branched oped in males of completed sexual anal-fin rays; lateral dark body maturity and without obvious stripe of adult males nearly black, pump chamber enclosed by mod- clearly distinct; body depth at dor- ified proximal portions of caudal- sal-fin origin about 4.0 to 4.5 times fin ray halves 11 and 12; ray halves in SL ...Mimagoniates barberi Regan of this region of these rays modi- (tributaries of rio Paraguay in fied into expanded, flattened struc- parts of Brazil and Paraguay) tures parallel to one another; these 6a. Hooks absent on all principal cau- modified ray halves of each side of dal-fin rays of adult males (see Figs. caudal fin developed so that a ava [Ud WA) A EE Nay ea eee EC) groove exists between them; groove 6b. Hooks present on at least principal and accompanying hypertrophied caudal-fin rays 10 and 11 of adult glandular tissue constitute a prim- males, hooks also frequently pres- itive pump organ (see Figs. 2,3) (5) ent on ray 12 (see Figs. 5, 21)... (8) 4b. Caudal fin-ray pump well-devel- 7a. Lateral series scales 49-56; scale oped in males of completed sexual rows between dorsal-fin and anal- maturity, consisting of bilateral fin origins 16-18; body depth of chambers, one on each side of fin adult males 3.4—3.8 in SL; in wild and each chamber enclosed by flat- caught specimens body silvery tened expanded portions of ray blue, when black horizontal stripe halves of rays 11 and 12; pump present, located at and partly dor- chamber with an obvious posterior sal to mid-lateral region of body opening, lateral slit and anterior from tip of snout to central caudal- opening (see Figs. 4,5) ........ (6) fin rays, stripe diffuse and broad 5. Branched anal-fin rays 23-29, usu- and often obscured by silvery blue ally 25-27, rarely 28 or 29; scales coloration posteriorly and by sil- in lateral series (including lateral very pigment anteriorly; stripe line scales) 36-41, usually 37-40; ranelvaclcanlyadenned snr scale rows between dorsal-fin ori- = 35 ............ M. sylvicola, new species gin and anal-fin origin 16-18; dor- (tributaries of Atlantic Ocean in sal-fin origin at vertical line drawn southern Bahia, Brazil) from base of second or third 7b. Lateral series scales 37-44; scale branched anal-fin ray; mid-lateral rows between dorsal-fin and anal- dark body stripe of adult males dif- fin origins 12-15; body depth of fuse, poorly developed, often not adult males 3.8—5.1 in SL; in wild apparent; body depth approxi- caught specimens body mahogany MAAK ZITO S31 MG eesscose brown, especially dorsally, with See Mimagoniates inequalis dark brown, nearly black, horizon- (Eigenmann) tal lateral body stripe below mid- dos Patos and lagoa Mirim, in Rio region of body prominent and ex- Grande do Sul, Brazil and tending from tip of lower jaw, northeastern Uruguay) posteriorly ventral to eye, across 5b. Branched anal-fin rays 30-36; opercle and body sides just ventral to mid body region onto caudal pe- duncle and across central portion scales in lateral series 41—48; scale rows between dorsal-fin origin and 386 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Mimagoniates barberi, unstained central basal region of caudal fin adjacent to caudal peduncle of a preserved adult male, lateral view, left side, anterior at left. SL 32.7 mm, UMMZ 205420, Paraguay, rio Aguary- mi, tributary to rio Paraguay, San Pedro. Illustrates external features of a caudal gland without complex pump chamber. Darkly pigmented glandular groove between principal rays 11 and 12 indicated by arrow. Modified dorsal caudal-fin lobe scales, although present, are transparent and not visible. of, caudal Mine akn cera iecre lca he c= 2s | INURE area Mimagoniates rheocharis, new . Mimagoniates lateralis (Nichols) species (tributaries of Atlantic Ocean from (tributaries to Atlantic Ocean of Santos, Sao Paulo south to Joinville, southern Santa Catarina and northern Santa Catarina, Brazil) Rio Grande do Sul, Brazil) 8a. Branched anal-fin rays 23-29, usu- 8b. Branched anal-fin rays 26-33, usu- ally 24-26; branched dorsal-fin ally 28-31; branched dorsal-fin rays 8-12, rarely 8; scale rows be- rays 7-9, rarely 9; scale rows be- tween dorsal-fin origin and anal- tween dorsal-fin origin and anal- fin origin 17-22, usually 19-20; fin origin 14-17, usually 15-16; scale rows around caudal peduncle scale rows around caudal peduncle 19-23, usually 20-22; branched 15-18; numerous strong hooks on dorsal-fin rays 8-12, rarely 8; very at least caudal-fin rays 11-12 small hooks on caudal-fin rays 1 1- (sometimes 7-12) of fully adult 12 of fully adult males (Fig. 21); males (see Fig. 5); dorsal-fin origin dorsal-fin origin at vertical line at a vertical line drawn from base drawn from anal-fin origin or from of branched anal-fin rays 5 or 6 . base of2tto 4iraysiposteriomtoanal=)1)) 09) ess ne ae eee Mimagoniates microlepis fiNtOn Sin ea he eal hone (Steindachner) (tributaries to VOLUME 103, NUMBER 2 387 Fig. 3. Mimagoniates barberi, osteology of central basal region of caudal-fin skeleton, principal fin-rays 6— 15, of adult male, lateral views, left side, anterior at left. SL 35.6 mm, UMMZ 205420, same locality as Fig. 2. Principal ray 12 indicated by arrow. (A) Illustrates glandular groove between rays 11 and 12. (B) Illustrates relationship of modified dorsal caudal-fin lobe squamation to glandular groove. Note that modified scales together with their epidermis form a movable flap just lateral to surface of rays and glandular groove. Free border of this flap occurs along ventral margin of ventral row of scales and at posterior edge of posterior scale of ventral scale row. Atlantic Ocean from southern Bahia south to northern Rio Grande do Sul, also in upper rio Iguacu, Parana) Mimagoniates sylvicola, new species Figs. 7-16, Table 1 Species A.— Weitzman et al., 1988:figs. 6, 10 [phylogeny and biogeography]. Holotype. —MZUSP 36612, male, SL 30.2 mm, Brazil, Bahia, Municipio de Prado, forest stream tributary to Atlantic Ocean, near Fazenda Embacuaba, approximately 8-9 km NW of Cumuruxatiba, 17°05’S, 39°13’W, 20 Mar 1985; N. Menezes, R. M. C. Castro, M. Weitzman, and S. Weitzman. Paratypes. —Following 2 lots of imma- tures to adults collected with holotype: MZUSP 28817, spms. 42, SL 15.1-30.2 mm; USNM 276557, spms. 42, SL 14.7- 33.5 mm, | male SL 29.3 mm and | female SL 26.6 mm [both cleared and stained]. Fol- lowing lots of immature to adult paratypes all collected 20 Mar 1985 by N. Menezes and party unless otherwise noted: MZUSP 28815, spms. 77, SL 11.0-—27.4 mm; USNM 276547, spms. 77, SL 14.4—27.4 mm, Bra- zil, Bahia, Municipio de Prado, first stream (locally called rio do Sul) south of rio Cai, on road between Cumuruxatiba and Ita- maraju, 17°00’S, 39°12’W. MZUSP 28816, spms. 28, SL 12.7-25.1 mm; 6, spms. 25, SL 13.2-24.0 mm, Brazil, Bahia, Municipio de Prado, small stream NW of Cumurux- atiba, about 17°01’S, 39°12'’W. USNM 300633, spms. 5, SL 22.3-31.3 mm and USNM 300634, spm. 1, cleared and stained, SL 31.8 mm, Brazil, Bahia, Municipio de Porto Seguro, riacho Ronca Agua, tributary to right margin of rio Camurugi, tributary to rio Joao de Tiba drainage, 15 km NW of Porto Seguro, approximately 16°20’S, 30°07'W, 19 Feb 1986, I. Rosa and party. Diagnosis. —Mimagoniates sylvicola may be separated from all other species of Mi- magoniates by use of the key to the species provided above. It is distinguished from its morphologically most similar relative, !. lateralis, by the following characters: lateral series scales 49-56 (37-44 for M. lateralis), scale rows between dorsal-fin and anal-fin origins 16—18 (12-15 for M. /ateralis). Cer- tain body measurement ratios differ signif- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON o Wiha Fig. 4. Mimagoniates microlepis, unstained central basal region of caudal fin adjacent to caudal peduncle of a preserved adult male, lateral view, left side, anterior at left. SL 51.7 mm, USNM 279876, Brazil, Santa Catarina, rio Itapocu. Illustrates external features of a caudal gland with a complex pump chamber. Arrow at left indicates anterior intake opening, middle arrow indicates lateral slit of chamber and arrow at right designates posterior exit opening of chamber. These openings surrounded by glandular tissue, especially exit opening. Modified dorsal caudal-fin scales appear as rather poorly focused “ghost” images in upper half of picture, above main body of pump chamber. icantly between adults of these species but overlap broadly in young and juveniles. For example, body depths in adult males and females of M. sylvicola diverge considerably from those in adult males and females of M. lateralis (body more elongate and slen- der). See “Discussion” below and Figs. 15, 16. Preserved and live colors differ between the species. Preserved males of M. sylvicola with dark lateral body stripe relatively pale and diffuse, occurring mostly at and partly dorsal to mid-lateral body region. Approx- imately dorsal half of opercle dark, nearly black (relatively pale in M. lateralis). Mima- goniates lateralis with a dark, relatively nar- row, Clearly defined lateral body stripe that lies mostly ventral to mid-lateral body re- gion. Dark stripe continues onto ventral one- third of opercle. Males of M. sylvicola with distal one-fourth to one-fifth (less poste- riorly) of anal-fin rays black (distal two- thirds to one-half black in M. /ateralis). Males of M. /ateralis with distal one-fourth of most elongate anterior unbranched ray and branched portions of anterior five to six branched rays hyaline or with a thin scat- tering of dark chromatophores, never black as in M. sylvicola. Numerous other, but less obvious, color differences occur in pre- served males of both species. These best discerned by comparing respective color de- scriptions given below. Life color of these VOLUME 103, NUMBER 2 —— Ze Toe = 389 ELLE are enema BAIA RD Ay ON Re WU EG REA CH EE SEP BP IS SoM HE a RP BE OT EE CII SS — Se ae A. oe vam nate ae ee ae, Ono a ear a a ee as sos ars cos aon oe a= — at ——— Sa — —_—— a =——S—., ——=—— Fig. 5. Mimagoniates microlepis, osteology of central and basal region of caudal skeleton of adult male, lateral view, left side, anterior at left. SL 45.3 mm, USNM 236089, Brazil, Parana, rio Nhundiaquara at Morretes. Relationship of modified dorsal lobe caudal-fin squamation to caudal gland shown in inset at left. Distal regions of ventral and posterior scales and their epidermis form a flap partly covering intake openings of pump chamber. species quite different. Male M. sylvicola with dorsally located black lateral “‘stripe”’ obscured by silvery blue reflective color, es- pecially anteriorly. Ventrally located black stripe of male M. lateralis deep black except at its mid-length ventral to dorsal-fin origin where partly obscured by blue to silvery pig- ment in some population samples. Wild caught males of M. /ateralis with a yellow- orange stripe just ventral to black lateral stripe, absent in M. sy/vicola, although both species often with anal-fin base yellow to orange. Note, in aquaria at least, MW. /ateralis loses yellow or orange coloration but black stripe always present. See live color descrip- tions below for a more complete account of M. sylvicola. Description.—Table 1 presents morpho- metrics of holotype and paratypes. Except where noted, entire description refers to lots from near Cumuruxatiba. These collections are treated statistically as one population sample since no statistical differences were found among them and all lots were col- lected from only a few kilometers apart. Counts for specimens from rio Camurugi are given only when they differ from those from near Cumuruxatiba. Body compressed, moderately elongate; body deepest about midway between snout tip and dorsal-fin origin, near anal-fin ori- gin. Predorsal body profile gently convex to snout tip. Body profile slightly elevated at dorsal-fin origin, straight along dorsal-fin base and nearly straight to origin of dorsal procurrent caudal-fin rays. Dorsal-fin origin 390 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON = SS ee SS = se Ses aaa ap eee ace oa) Sp BaSS Ear <=>) HD ES AN SG ee ee <> lay || mea (GY) IZ << SS SOS cs Seer SS= LOS ta 8 S WOES Fig. 6. Glandulocauda melanogenys, osteology of central and basal region of caudal skeleton of adult male, lateral view, left side, anterior at left. SL 40.2 mm, USNM 236093, Brazil, Sao Paulo, headwater stream of rio Tieté near Campo Grande (=Alto da Serra, type locality), about 3.5 km west of Paranapiacaba. Decurved principal caudal-fin rays 11 and 12 are labeled and inset at left illustrates relationship of modified dorsal caudal- fin squamation to rays 11 and 12. Note that modified caudal-fin squamation of females of in all species of Glandulocauda and Mimagoniates is very similar to that illustrated here but may reach only about half the relative dimensions depicted here. nearer to caudal-fin base than to snout tip. Ventral body profile convex from anterior tip of lower jaw to point on abdomen about midway between pectoral- and pelvic-fin bases. Belly profile abruptly becomes con- cave and then straight to anal-fin origin. Body profile slightly convex along anal-fin base to anal-fin insertion. Ventral profile of caudal peduncle slightly convex, especially in adult males where profile formed by ven- tral procurrent caudal fin rays. In females and juveniles this profile nearly straight. Head and snout of moderate size in pro- portion to body length. Lower jaw protrud- ing, anterior to upper jaw. Lower jaw of males thick and heavy compared to that of females. Mouth angled posteroventrally from anterior tip of snout to posterior part of mandibular joint. Maxilla extending pos- teriorly to a point anterior of a vertical line drawn through anterior border of pupil of eye. Dorsal-fin rays 11,8 (unbranched rays 11 in all specimens, branched rays x = 8.0 [3 spms. with 9], range = 8-9, n = 90); of 6 specimens from rio Camurugi (not included in n = 90) VOLUME 103, NUMBER 2 391 Table 1. Morphometrics of Mimagoniates sylvicola, new species. Standard length is expressed in mm; mea- surements through head length are percentages of standard length; the last four entries are percentages of head length. Specimens are from the area near Cumuruxatiba, Bahia, MZUSP 28815, 28816, 28817, 36612 and USNM 276547, 276556, 276557. Holotype n Range x Standard length 30.2 88 14.5-30.2 20.9 Depth at dorsal-fin origin = 88 20.7-29.3 24.9 Males 29.1 44 21.5-29.3 25.7 Females — 44 20.7-27.2 24.0 Snout to dorsal-fin origin 61.6 88 57.1-63.8 60.1 Snout to pectoral-fin origin 28.1 88 24.3-29.0 26.3 Snout to pelvic-fin origin 43.4 88 39.8-46.9 43.3 Snout to anal-fin origin 55.6 88 54.4-61.0 57.1 Caudal peduncle depth — 88 8.5-14.3 10.8 Males 13.2 44 8.7-14.3 11.6 Females — 44 8.5-11.7 10.1 Caudal peduncle length 12.3 88 10.2-13.8 12.1 Pectoral-fin length PIB \7) 88 20.2-24.8 22.6 Pelvic-fin length 14.9 88 11.1-16.1 13.3 Dorsal-fin base length 14.2 88 11.8-16.6 13.8 Dorsal-fin height 19.2 88 14.4-21.0 18.1 Anal-fin base length 33.4 84 29.7-35.2 32.6 Anal-fin lobe length 19.2 87 17.2-22.6 20.1 Eye to dorsal-fin origin 46.4 88 42.9-51.0 46.3 Dorsal-fin origin to caudal-fin base 43.7 88 39.4-46.9 42.4 Bony head length 27.8 88 24.8-28.6 26.3 Horizontal eye diameter 35.7 90 33.7-47.4 40.6 Snout length 22.6 90 20.8-26.5 23.8 Least interorbital width 33.3 90 31.6-40.5 35.3 Upper jaw length 45.2 90 34.4-49.3 45.7 2 with 8 and 4 with 7 branched rays; pos- terior ray not split to its base and counted as 1. Adipose fin present, slender. Anal-fin rays iv,25 (unbranched rays iv in all spec- imens, branched rays X = 24.8, range = 23- 26, n = 90); posterior ray split to its base and counted as 1. Anal fin with moderately developed lobed anterior portion including fourth unbranched ray and first 5-6 branched rays. Anal fin of sexually mature males with bilateral blunt hooks on anterior 6 branched fin rays, | set of hooks for each ray (see Fig. 9). Pectoral-fin rays 1,10 (un- branched rays i in all specimens, branched rays X = 9.7, range = 9-11, n = 90); all 6 specimens from rio Camurugi with 10 branched rays. Posterior tips of longest pec- toral-fin ray extend posteriorly beyond or- igin of pelvic fin; of about equal length in both sexes. Pectoral-fin rays without hooks. Pelvic-fin rays 8 (8 in all specimens except 1 with 9, n = 90). Pelvic fin with anterior most ray branched in all specimens (see Fig. 10). Adult males with total of over 100 small to tiny hooks present on rays of pelvic fin, distributed as in Fig. 10. Each ray bears 9 to over 40 hooks, depending on the matu- rity of the specimen and/or the fin ray ex- amined. Principal caudal-fin ray count 10/9 in all specimens (n = 90). Fin rays modified in association with caudal pheromone pump as in Figs. 11 and 12. Fig. 11 illustrates a relatively immature pump, while Fig. 12 shows a presumably mature pump in which pump chamber has well-developed water 392 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Mimagoniates sylvicola, new species, holotype, MZUSP 36612, male, SL 30.2 mm; Brazil, Bahia, Municipio de Prado, unnamed forest rivulet near Fazenda Embacuaba, 8-9 km northwest of Cumuruxatiba. entrances and exit. Fin rays modified very much like those of M. /ateralis. Caudal-fin rays without bony hooks. See also Weitz- man & Fink (1985:98—99), Weitzman et al. (1988:384—-413) and ‘Discussion’? below, regarding phylogeny of M. sylvicola for hy- pothetical function of secondary sexual characters and phylogeny for glandulocau- dins as indicated by their caudal pump mor- phology. Scales cycloid, almost deciduous, with few radii along posterior border; smallest scales often nearly without or without radii. Ter- minal scale of modified caudal series with exaggerated radii appearing as incisions of posterior scale borders (see Figs. 11b, 12b). Lateral line incomplete, perforated scales 8 (x = 7, range = 6-8, n = 34); 2 specimens from rio Camurugi with 9 perforated scales. Lateral series scales 53 (x = 52.7, range = 49-56, n = 34). Predorsal scales 25 (x = 25.9, range = 24-28, n = 39). Scale rows between dorsal-fin origin and anal-fin origin 17 (x = 16.7, range = 16-18, n = 64). Scale rows around caudal peduncle 20 (x = 19.7, range = 19-20, n = 21); 1 specimen from rio Camurugi with 22 scale rows around caudal peduncle. Premaxillary teeth in 2 distinct rows al- though this is not clear in Fig. 13. Larger teeth tricuspid, smaller teeth tricuspid or bicuspid, smallest ones unicuspid. Outer row teeth 6 (Xx = 5.4, range = 3-7, n = 90). Inner row teeth 3 (x = 3.0, range 2-5, n = 90). Outer and inner row premaxillary teeth somewhat compressed compared to most “‘tetragonopterine” characid teeth which often appear almost circular in cross sec- tion. Maxillary teeth 6 (* = 6.8, range = 5— 10, larger specimens usually with highest counts, n = 90); two specimens from rio Camurugi with 11 maxillary teeth. Maxil- lary teeth show an increase in number with increasing SL from a mean of 5.9 in 9 spec- imens between 15.5 and 16.5 mm SL toa mean of 7.6 in 14 specimens between 25.0 and 30.5 mm SL. Anterior 4—5 maxillary teeth tricuspid and larger than remaining teeth with 2 or 1 cusps. Dentary with 4 large tricuspid teeth in all specimens, n = 90; smaller posterior dentary series unicuspid except anterior tooth which is tricuspid, 10 (X = 8.9, range = 6-12, n = 90); 1 specimen from rio Camurugi had 13 dentary teeth. Maxillary and dentary teeth shaped much like premaxillary teeth as described above. At any given SL considerable variation in tooth count occurs and nearly any tooth count within ranges given may be expected. No significant differences in tooth number found between males and females. Vertebrae 40 (x = 39.9, range = 39-41, VOLUME 103, NUMBER 2 393 Fig. 8. Mimagoniates sylvicola, new species, paratype, USNM 276547, female, SL 25.7 mm, Brazil, Bahia, Municipio de Prado, rio do Sul. n = 88). Dorsal limb gill-rakers 6 (X = 6.0, range 6-7, n = 90, two specimens from rio Camurugi with 5 dorsal limb gill-rakers); ventral limb gill-rakers 12 (¥ = 11.7, range = 11-13, n = 90). Branchiostegal rays 4 in 3 cleared and stained specimens, 3 rays orig- inating from anterior ceratohyal and 1 ray from posterior ceratohyal. Color in alcohol. —See Figs. 7 and 8 for preserved color pattern in males and fe- males. Body pale to medium brown, almost white ventrally, darkest dorsally. Lateral body stripe pale, diffuse, best developed in males. Stripe extending from darker oper- cular spot on dorsal half of opercle poste- riorly to a dark, spot-like region on caudal peduncle. Immediately posterior to this spot, caudal gland region enveloped in black pig- ment forming triangular-shaped area with its posterior apex continuous onto ray 11 and to a certain extent ray 12. Remainder of caudal fin dusky due to scattering of dark chromatophores, especially along ventral border of 19th principal caudal ray. Dorsal body surface dark dusky, especially in area of predorsal scales. Pectoral, pelvic, dorsal and anal fins dusky from scattered dark chromatophores along fin rays. Pelvic fins considerably darker than pectoral fins. Anal fin with a dark, elongate stripe running length of fin. Width of stripe about '4—% height of fin. Stripe borders dis- tal ends of fin rays posteriorly; anterior por- tion of dark stripe separated from distal ends of first five or six fin rays by relatively hya- line area on anterior lobe of fin. Dorsal fin with horizontal dark stripe extending pos- teriorly from about mid-length of anterior elongate undivided ray to posterior tips of two terminal dorsal-fin rays. Adipose fin dusky with scattered dark chromatophores. Head dark brown around mouth and on dorsal surface of snout, between eyes, dor- sum of cranium and nape. Iris dorsal to pu- pil dark brown to black, most of remainder ofiris silvery with some dark brown or black areas ventrally. Circumorbitals pale brown or silvery with evenly scattered dark chro- matophores. Ventral area of opercle, pre- opercle and posterior region of branchio- stegal rays silvery, without much dark brown pigment. Color in life. —Life color patterns taken from color slides and color notes made while collecting specimens listed above from clear and black waters near Cumuruxatiba. Sides of body silvery deep blue with back dark brown and abdominal area silvery white. All fins translucent, lemon yellow with dark brown pigment described above under pre- served color description appearing brown to black. Females with similar color pattern 394 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 9. Mimagoniates sylvicola, new species, paratype, anterior 11 anal-fin rays of an adult male, lateral view, left side, anterior at left, S1 32.5 mm, USNM 300634, Brazil, Bahia, riacho Ronca Aqua. For explanation see Description of M. sylvicola. but blue, yellow, and dark pigment patterns much paler. In life caudal-fin rays 13 and 14 considerably darkened with black pig- ment. Some male specimens display a con- siderable but rather diffuse lateral dark brown stripe below lateral mid-region of body, suggesting elongate lateral stripe of M. lateralis. Specimens from rio do Sul, Cu- muruxatiba area (USNM 276547), gold sil- very in color and without blue coloration. Some of these specimens with black pig- ment considerably reduced, absent or cov- ered in patches by guanine, especially on body sides. These specimens may have been infested by metacercaria of a trematode as noted for other similar appearing characids by Geéry & Delage (1963). Sexual dimorphism. —Females lack the caudal pheromone pump organ, anal-fin and pelvic-fin hooks of males (Figs. 9, 10) and display a more subdued live body color- ation as noted above. Figure 14 presents graphic evidence that the caudal peduncle depth is usually deeper in adult males than in adult females and that males reach a greater adult length than females, so far as known. Below in covariance analyses we compare males and females of unequal length ranges, although the range of the fe- males is included within that of the males. We do this because we believe these ranges are expressions of their natural differences. In our population samples of various species of Mimagoniates the length of the largest males always exceeds that of the largest fe- males. If in nature the females reached the same lengths as the males our results could be biased, but longer females were not rep- resented in our samples. We hypothesize that most of our samples fairly represent the adult lengths of both sexes. Figure 14 in- dicates that caudal peduncle depth diver- gence between males and females begins around 18 mm in SL. Even though we have few female specimens in the size range be- tween 23.0 to nearly 28.0 or longer, we are inclined to accept that 28.0 mm SL may be the approximate adult size limit for females. Also, at least some glandulocaudine species undergo delayed sexual maturation in males, VOLUME 103, NUMBER 2 395 Fig. 10. Mimagoniates sylvicola, new species, paratype, pelvic-fin osteology of an adult male, ventral view, left side, anterior at left. SL 29.3 mm, USNM 276557, same locality data as holotype, Fig. 7. Medial fin-ray at bottom of picture. Weitzman & Fink (1985:38, 42). If present in species of Mimagoniates, this kind of growth pattern might affect the regression slope shown for sexually maturing and sex- ually mature males towards that of the fe- males in Fig. 14, if the large males of latent sexual maturity were included in the male population sample graphed and analyzed. Fig.11. Mimagoniates sylvicola, new species, paratype, osteology of central basal region of caudal-fin skeleton, principal rays 6-15, of an adult but still developing male, lateral views, left side, anterior at left. SL 29.3 mm, USNM 276557, same locality data as holotype, Fig. 7. Principal ray 12 indicated by arrow. (A) Illustrates area of developing rays associated with caudal pump. Note that pump chamber not yet fully developed. (B) Illustrates modified dorsal caudal-fin lobe squamation in natural position. 396 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 12. Mimagoniates sylvicola, new species, paratype, osteology of central basal region of caudal-fin skeleton, principal rays 6-15, of an adult male, lateral views, left side, anterior at left. SL 32.5 mm, USNM 300634, Brazil, Bahia, riacho Ronca Aqua. Principal ray 12 indicated by arrow. (A) Illustrates area of fully developed caudal pump. (B) Illustrates relationship of modified dorsal caudal-fin lobe squamation to pump skeleton. In the two slopes plotted in Fig. 14, latent males, if present, are included in that of the females. Thus Fig. 14 appears to demon- strate that as males mature sexually their caudal peduncles increase in depth at a fast- er rate relative to body length than it does in juveniles or females. With the above res- ervations and explanations in mind, we find that divergence 1n caudal peduncle depth on SL by apparent sex in an F-ratio test for homogeneity of slopes in an analysis of covariance were significantly different (Fo.05,a, 36) = 9-42, P < 0.002) between 45 males and 45 females from the area near Cumuruxatiba (MZUSP 28815, 28816, 28817, 36612 and USNM 276547, 276556, 276557). Etymology. —The name sylvicola is from the Latin silva (forest) and colo (dwell or inhabit) and is in reference to the forested nature of the streams in which this fish is found. Discussion. — Weitzman et al. (1988:404, 414-419, fig. 10), as new species A, tenta- tively hypothesized that M. sylvicola was a relatively primitive species of Mimagonia- tes with respect to caudal-fin pump evolu- tion. They placed M. sy/vicola in an unre- solved polytomy in their phylogenetic diagram along with M. barberi and M. in- equalis and a monophyletic line leading to M. lateralis, M. microlepis and their new species B and C. This hypothesis was based on the information that apparently fully adult males of all three species, VW. barberi, M. inequalis and their new species A (here = M. sylvicola), lack the more derived chambered, caudal-fin ray pump of the oth- er four species. Compare caudal pump structures in Figs. 2 and 3 with those in Figs. 4 and 5. Evidence taken from a specimen of M. sylvicola, USNM 300633, SL 32.5 mm, sub- sequently available to us from near Porto Seguro, Bahia, showed that the caudal gland matures with a well-developed pump cham- ber, (compare Figs. 11 and 12). However, this species lacks the caudal-fin hooks found in the more derived species, M. rheocharis, described below, and in M. microlepis (com- pare Figs. 5, 11, 12, and 24). This infor- mation would place M. sylvicola in a tri- chotomy, Fig. 1, with M. Jateralis and a monophyletic line leading to the two species with caudal-fin hooks. This hypothesis would leave M. inequalis, M. barberi and a monophyletic line leading to the species of Mimagoniates with more derived caudal pumps in a trichotomy at a lower level in the phylogenetic diagram (Fig. 1). If this relationship can be supported by further phylogenetic evidence, then the biogeo- VOLUME 103, NUMBER 2 Fig. 13. Mimagoniates sylvicola, new species, paratype, jaws and dentition of an adult male, lateral view, right side, anterior at right. SL 29.3 mm, USNM 276557, same locality data as holotype, Fig. 7. For explanation see text under Description of M. sylvicola. graphic comments of Weitzman etal. (1988: 412, 413) need some alteration (see section on biogeography below). It was noted in the diagnosis above that adult males and females of M. sy/vicola from Cumuruxatiba (MZUSP 28815, 28816, 28817, 36612 and USNM 276547, 276556, 276557), differ in body depth from adult males and females of M. /ateralis from the Santos region (CAS 36634, MZUSP 40276, 40277, USNM 226468, 254268, 257200, and 257202). Forty five males of M. sylvi- cola and 22 males of M. Jateralis were sig- nificantly different (Fo.05, 4,63) = 49.75, P < 0.000) in an analysis of covariance for body depth on SL in an F-ratio test for homo- geneity of slopes, Fig. 15. Body depth ratios of SL useful for identification of fully or near fully mature males (at or above 23.0 mm SL) are as follows: m = 12 specimens of sylvicola from Cumuruxatiba and near Por- to Seguro, X = 3.6, range = 3.4-3.8 and n = 23 specimens of M. J/ateralis from near Santos and Municipio de Cananéia, «= 4.3, 398 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON - o) al fo) Ww fo) es 30.0 Tod F CAUDAL PEDUNCLE DEPTH (mm) (log) 20.0 STANDARD LENGTH (mm) (log) CUMURUXATIBA, BAHIA, BRAZIL Mimagoniates sylvicola, Fig. 14. Mimagoniates sylvicola, new species, caudal peduncle depth as a function of SL by sex. Plot axes are logarithmic. Logarithmic regression equation for 45 males: (Y = —4.089 + 1.625X), for 45 females: (Y = —3.377 + 1.368X). Immature males and juveniles may be included in the category designated female. For explanation see under Sexual dimorphism in text of M. sylvicola. range = 3.8—5.1. The shorter males in these size ranges accounted for most of the over- lap. In a similar covariance analysis for slopes, 45 adult females of M. sylvicola and 26 fe- males of M. lateralis from the same popu- lation samples as the males discussed above, also displayed significant difference in body depth (Fo.0s, a, 67) = 16.3, P < 0.001) (Fig. 16). The number of branched anal-fin rays in the two species was significantly different (¢ = 22.012, P < 0.00 in a two-sample, two- tailed t-test), although there was some over- lap in counts: n = 90 specimens of M. syl- vicola from near Cumuruxatiba (same lots as listed above), range = 23-26, x = 24.8, SD = 0.7728 and n = 91 specimens of M. lateralis from near Santos and Municipio Cananéia (same lots as listed above), x = 27.9, range = 25-31, SD = 1.0899. Ecology. — Little is known about the ecol- ogy of this species. Most of the streams in which it was captured were relatively slow moving, with little gradient. They were ap- proximately 4 to 6 meters wide, to 1.5 me- ters deep and surrounded by vegetation, usually trees of a few to many meters high. The water varied from clear to black (tea color). The fish occurred in depths of 0.1 to about 0.5 meters usually in areas of little current over white sandy, rocky or dark mud bottoms. They occurred in both sunlight or shaded areas, most often near shore, espe- cially near emergent or submerged vegeta- tion where almost immediate cover could VOLUME 103, NUMBER 2 Mimagoniates sylvicola, a=cr BODY DEPTH (mm) (log) _| E Nee 399 CUMURUXATIBA, BAHIA, BRAZIL 20.0 —— 30.0 STANDARD LENGTH (mm) (log) Mimagoniates lateralis, SANTOS, SAO PAULO, BIREVAIL A=C Fig. 15. Mimagoniates sylvicola and Mimagoniates lateralis, body depth as a function of standard length for young to adult males. Plot axes are logarithmic. Logarithmic regression equation for 45 males of M. sylvicola: (Y = —2.650 + 1.420X) and 22 males of M. lateralis: (Y = 0.473 + 0.695X). For explanation see Discussion under M. sylvicola. be taken from predators such as large cich- lids or the characid Oligosarcus. Specimens of M. sylvicola, now MZUSP 28817 and USNM 276557, were collected 22 March 1985 ina well-shaded forest rivu- let less than 1 meter wide and about 20-30 cm deep in most places. This creek was in a ravine of a mostly uncut, undisturbed tall forest, 1 or 2 km from the Atlantic coast, about 17 km from Cumuruxatiba, Bahia. The stream bottom consisted of forest litter, rocks, soil and sand with a mild nearly 0° to 30° gradient, well covered by riparian vegetation in many places. The water was tea colored. Other fish species taken at this site were Rachoviscus graciliceps, species of Astyanax, Characidium, Aspidoras, Hep- tapterus, a gobiid and a hypoptopomine lor- icariid catfish. Mimagoniates rheocharis, new species Figs. 17-28, Table 2 Species B.— Weitzman et al., 1988:figs. 6, 10, 23 [phylogeny and biogeography]. Holotype.—MZUSP 40278, male, SL 47.3 mm, Brazil, Santa Catarina, Municipio de Praia Grande, rio Faxinalzinho at Mae dos Homens, near Praia Grande, approximately 400 Mimagoniates sylvicola, Oo =So° 8.0 7.0 6.0 5.0 4.0 BODY DEPTH (mm) (log) 20.0 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON CUMURUXATIBA, BAHIA, BRAZIL ———=== 1 30.0 STANDARD LENGTH (mm) (log) Mimagoniates lateralis, SANTOS, SAO PAULO, BRAZIL | A= Fig. 16. Mimagoniates sylvicola and Mimagoniates lateralis, body depth as a function of standard length for young to adult females. Plot axes are logarithmic. Logarithmic regression equation for 45 females of M. sylvicola: (Y = —2.567 + 1.385X) and 26 females of M. Jateralis: (Y = —1.807 + 1.118X). For explanation see Discussion under M. sylvicola. 29°20’S, 14°40’W, 9 Jun 1985; C. A. S. de Lucena, R. E. Reis, and L. R. Malabarba. Paratypes. — Following 2 lots of imma- tures to adults collected with holotype: MCP 13616, spms. 3, SL 24.3-40.8 mm; USNM 279878, spms. 3, SL 33.3-—47.8 mm. Fol- lowing 3 lots of immature to adult paratypes all collected 7 Jun 1985 by C. A. S. de Lu- cena and party: Brazil, Santa Catarina, Municipio de Nova Veneza, rio Jordao at Jordao Alto, approximately 28°36’S, 49°28'W, MCP 13617, spms. 83, SL 21.9- 39.0 mm; MZUSP 40279, spms. 81, SL 22.4— 42.8 mm; USNM 270879, spms. 89, SL 22.8-49.0 mm. Following 3 lots of imma- ture to adults all collected by R. M. C. Cas- tro 13 Feb 1988: Brazil, Santa Catarina, Municipio de Praia Grande, tributary of rio Grande approximately 29°20’S, 49°40’W, MZUSP 40280, spms. 29, SL 13.5-38.4 mm; USNM 306339, spms. 18, SL 17.0- 40.0; spms. 2 to R. M. C. Castro, SL 28.0- 31.30 mm. MZUSP 40281, SL 41.8 mm (mature male), Brazil, Rio Grande do Sul, Municipio de Osorio, arroio das Pedras, ap- proximately 29°52’S, 50°19’W, 11 Feb 1988, R. M. C. Castro. MCP 13613 spms. 7, SL 27.5-31.3 mm, Brazil, Municipio de Oso- rio, arroio Agua Parada, tributary to rio Ma- quiné, in Maquiné, approximately 29°40’S, 50°11’W, 10 Oct 1989, R. E. Reis, S. O. Kullander, L. R. Malabarba and J. Pozzi. MCP 10806, SL 39.5 mm (mature male), Brazil, Rio Grande do Sul, Municipio de VOLUME 103, NUMBER 2 Torres, tributary of rio Trés Furquilhas, Chapéu, approximately 29°19’S, 49°44’W, 25 May 1986, C. A. S. Lucena, R. E. Reis, and L. R. Malabarba. Diagnosis. —Mimagoniates rheocharis may be separated from all other species of Mimagoniates by use of the key provided above and the characters described and dis- cussed below. Mimagoniates rheocharis is sympatric with and apparently at least part- ly syntopic with M. microlepis in south coastal Santa Catarina. It is also sympatric and syntopic with M. microlepis in the rio Maquiné drainage. The southern distribu- tion of M. rheocharis (rio Maquiné drainage and streams immediately surrounding Oso- rio in Rio Grande do Sul) is adjacent to the northeastern limit (rio Gravatai drainage) of M. inequalis where these two species abut and appear to be allopatric (see the Bio- geography section and phylogenetic discus- sion in the Discussion section below). The possible relationships of M. rheocharis to M. inequalis or M. microlepis is uncertain and perhaps complex but the following se- ries of characters distinguish M. rheocharis from one or the other of these species or, in some cases, both of them. Because the com- parisons are somewhat complex, for clarity the two compared species are treated sep- arately. Mimagoniates rheocharis may be distin- guished from M. microlepis, by the follow- ing characters (see also the statistical anal- yses of overlapping characters under Statistical Comparisons below). Scale rows around caudal peduncle 19-23 (15-18 for M. microlepis); scale rows between dorsal- fin and anal-fin origins 1 7—22, rarely 17 (14— 17, rarely 17 for M. microlepis); total lateral series scale count shows broad overlap be- tween both species, 41-48 (42-49 for M. microlepis) but in at least one population comparison of M. rheocharis and M. mi- crolepis showed a statistically significant dif- ference (see the statistical comparisons be- low); branched dorsal-fin ray count 8-12, usually 9-10, rarely 8 (7-9, rarely 9 for M. 401 microlepis); branched anal-fin ray count 23- 29, usually 24—26 and rarely 28 or 29 (26- 33, usually 28-31 and rarely 26 or 27 for M. microlepis); total vertebral count 35—40, usually 36-38, rarely 39 or 40 (38-41, usu- ally 39-40, rarely 38 in M. microlepis); tenth and eleventh principal caudal-fin rays of adult males with small, little developed hooks posterior to caudal organ but with sturdy hooks on ray 11 along ventral border of expanded ray segments of caudal organ, Fig. 24 (adult male M. microlepis with well- developed hooks on caudal-fin rays 7-12 and no sturdy hooks on principal ray 11 along ventral border of expanded ray seg- ments of caudal organ, note, some popu- lations with few or no hooks on rays 7 and/ or 12); preserved and live colors in part dif- ferent between both species, but variation in live colors among various population samples of M. microlepis make compari- sons difficult. A few life color characters appear consis- tently different in fully mature males: pel- vic-fin rays and membranes of adult males distally white (adult male M. microlepis with yellow and/or black pigment of pelvic fins continuous to edge of fin where fin bordered by narrow band of white); anal fin posterior to anterior lobe bordered by broad band of deep yellow pigment, very little to no black pigment on fin (adult male /. microlepis with posterior portion of anal fin ventrally bordered by narrow band of black pigment, none or very little yellow pigment). As treated below in the Discussion, there are some, perhaps plesiomorphic, body shape similarities shared between M. rheo- charis and M. inequalis. Furthermore, /. rheocharis appears to have the caudal pe- duncle and body depths for both sexes in- termediate between those of M. microlepis and M. inequalis; see Figs. 17 and 27. Cer- tain aspects of the caudal pump morphology of M. rheocharis could also be interpreted as intermediate. This suggests the possibil- ity that M. rheocharis might be closely re- lated to M. inequalis rather than to M. mi- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Mimagoniates inequalis, MORUNGAVA, RIO GRANDE DO SUL, BRAZIL Mimagoniates rheocharis, RIO GRANDE, SANTA CATARINA, BRAZIL 402 =o 12) o=co a =Q —N D (e) <=, 100 9.0 =~ & 8.0 £ 70 aE KF 6.0 ae LJ OQ 5.0 > OQ O Mm 4.0 3.0 + @ 2.8 4 ———. 13.0 20.0 30.0 STANDARD LENGTH (mm) (log) Mimagoniates microlepis, PARANAGUA, PARANA, BRAZIL Oo Sc e=9 Fig. 17. Mimagoniates rheocharis, M. microlepis and M. inequalis, body depth as a function of SL by species and sex. Plot axes are logarthmic. Logarithmic regression equation for 31 males of M. rheocharis: (Y = —1.846 + 1.199X), 25 females or this species: (Y = —2.115 + 1.271X); for 39 males of M. microlepis: (Y = —2.130 + 1.235X) and for 31 females of the later species: (Y = —2.705 + 1.399X); and for 26 males of M. inequalis: (Y = —2.230 + 1.3499X), 14 females of this species: (Y = —1.812 + 1.2187X). For explanation see Diagnosis, Sexual dimorphism, Discussion and Description under M. rheocharis. crolepis or even derived by introgression from ancestral sympatric populations of M. microlepis and M. inequalis in a manner suggested by Hubbs (1955:19) for fresh- water fishes in North America. However, not all counts or body proportions of M. rheocharis are intermediate between the two species and we question the intermediacy of M. rheocharis based on tentative polarity and parsimony analyses of the available data (see Discussion below). In any case, the sim- ilarities of many features, especially of im- mature or just maturing specimens, shared between M. rheocharis and M. inequalis makes it a practical necessity to describe the differences in some detail between the pop- ulation samples at hand of these two species. The phylogenetic polarities of most of these characters are relatively uncertain for those species possessing them. Therefore their use as synapomorphies suggesting relationships to and/or among species of Mimagoniates that may share them remains to be more fully investigated. Tentatively, many of these VOLUME 103, NUMBER 2 403 Fig. 18. Mimagoniates rheocharis, new species, holotype, MZUSP 40278, male, SL 47.3 mm; Brazil, Santa Catarina, rio Faxinalzinho at Mae dos Homens, near Praia Grande. characters appear plesiomorphic for M. in- equalis when the latter is compared to M. rheocharis or M. microlepis. For convenience the following same char- acters as those treated above to separate M. rheocharis and M. microlepis are considered for comparison between M. rheocharis and M. inequalis even though not all these char- acters are diagnostic for separation of the latter two species (see also the section below on Statistical Comparisons). Scale rows around caudal peduncle = 19-23 for M. rheocharis (16-19 for M. inequalis) (both species commonly with a count of 19); scale Fig. 19. Mimagoniates rheocharis, new species, paratype, USNM 306339, female, SL 34.3 mm; Brazil, tributary to rio Grande, Santa Catarina. 404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 20. Mimagoniates rheocharis, new species, paratype, MZUSP 40281, male, SL 41.8 mm; Brazil, Rio Grande do Sul, arroio das Pedras, near Osorio. rows between dorsal-fin and anal-fin origins = 17-22 (16-18 for M. inequalis); lateral series scale count = 41-48, rarely 41 (36- 41 for M. inequalis, 41 being uncommon); branched dorsal-fin ray count 8-12, usually 9 or 10, only occasionally 8 (8-9, rarely 9 for M. inequalis); branched anal-fin ray count 23-29, usually 24-26, rarely 28 or 29 (24-30, rarely 24 for M. inequalis), these counts significantly different statistically be- tween some population samples but not in others (see statistical analyses below); total vertebral count 35-40, rarely 39-40 (36-39 for M. inequalis); principal caudal-fin rays 10 and 11 of adult males with small, little developed hooks posterior to caudal-fin or- gan, see Fig. 24 (adult male of M. inequalis without hooks on caudal-fin rays 10 and 11); preserved color patterns essentially in- distinguishable between the two species and we have not been able to study the live color patterns of adult males where differences between the species would most likely be present. Description.—Table 2 presents morpho- metrics of the holotype and paratypes. Ex- cept where noted, the entire description re- fers to the population sample from near Praia Grande, southern Santa Catarina. These collections were treated statistically as one population sample in an attempt to represent the species as a whole from this area. Counts and ratios of measurements for other population samples taken from a trib- utary of the rio Grande are given only when they differ from those from near Praia Grande. Body compressed, relatively deep, espe- cially near dorsal-fin origin; body deepest at vertical line through anal-fin origin. Pre- dorsal body profile relatively arched in adult males, less so in adult females and imma- tures which have predorsal profile gently convex to tip of snout. Body profile elevated at dorsal-fin origin, still strongly arched in males, less so in females and juveniles. Dor- sal body profile nearly straight along dorsal- fin base to adipose fin. Body profile poste- rior to adipose fin somewhat concave dorsal to caudal peduncle, ending at origin of pro- current caudal-fin rays. Dorsal-fin origin nearer to caudal-fin base than to snout tip. VOLUME 103, NUMBER 2 Fig. 21. 405 Mimagoniates rheocharis, new species, paratype, anterior 13 anal-fin rays of an adult male, lateral view, anterior is at left. USNM 279879, SL 45.4 mm; Brazil, Santa Catarina, rio Jordao at Jordao Alto. For explanation see Description under M. rheocharis. Ventral body profile strongly convex in adult males from anterior tip of lower jaw to or- igin of pelvic fins, less strongly convex in females and juveniles. Belly profile in adult males slightly concave to anal-fin origin, straight or nearly so in females and juve- niles. Body profile along anal-fin base in males slightly concave at anterior base in region of anterior lobe of anal fin; straight along base of remainder of fin in males and along entire anal-fin base in females and juveniles. Ventral profile of caudal peduncle convex in adult males when anterior 5 or 6 strongly developed procurrent ventral cau- dal-fin rays are included in that profile; slightly concave or nearly straight in females and juveniles. Head and snout of moderate size in pro- portion to body length. Lower jaw protrud- ing, slightly anterior to upper jaw. Lower jaw of adult males thick and heavy com- pared to that of females and juveniles. Mouth angled posteroventrally from ante- rior tip of snout to posterior part of man- dibular joint. Maxilla long, extending to a point ventral to a horizontal line drawn from ventral border of eye in juveniles and adults of both sexes. Maxilla extends posteriorly to a point anterior to vertical line drawn through anterior border of pupil of eye. Dorsal-fin rays 11,9 (unbranched rays 11 in all specimens, branched rays x = 8.9, range = 8-10, n = 56); posterior ray not split to its base and counted as 1. Adipose fin pres- ent, slender. Anal-fin rays iv,26 (iv or v, usually iv, X = 26.8 for branched rays, range = 25-29, n = 56); posterior ray split to its base and counted as 1. Anal fin with mod- erately developed lobe anteriorly (Figs. 18, 19). Lobe includes fourth or fifth undivided ray and first 3 divided rays. Anal fin of sex- ually mature males with bilateral hooks, 1 on each side, on anterior unbranched ray iv or v, whichever occurs just before branched rays (Fig. 21). Usually anterior 7 branched fin rays with bilateral hooks, 1 set for each ray. Pectoral-fin rays i,10 (unbranched ray iin all specimens, branched rays x = 10.1, range 10-11, n = 56). Posterior tips of lon- gest pectoral-fin rays extend posteriorly be- yond origin of pelvic fins; fins of about equal extent in both sexes. Pectoral-fin rays with- 406 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 22. Mimagoniates rheocharis, new species, paratype, pelvic fin of adult male, ventral view, left side, anterior at left, medial fin ray at bottom. USNM 279879, SL 45.4 mm; same locality data as specimen in Fig. 21. out hooks. Pelvic fin rays 8 in all specimens. Pelvic fin with anterior (first) ray branched once, branches remaining close together and entire ray tapering as in most characiforms that have anterior ray unbranched (Fig. 22). Sexually mature, large adult males with over 400 hooks on each pelvic fin. Principal caudal-fin ray count 10/9 in all specimens, (” = 56). Fin rays modified in association with caudal pheromone pump as in Figs. 23 and 24. Fin rays modified more like those in M. microlepis than any other species of Mimagoniates (Fig. 5). Cau- dal-fin rays 10 and 11 with small bony hooks. Ventral borders of 4 anterior ex- panded ray segments of ray 11, which form anterior external wall of dorsal portion of pump chamber, with about 4 short but large hooks, 2 middle hooks often bicornate, oth- ers | hooked. Scales cycloid, with few radii along pos- terior border. Terminal scale of modified caudal-fin series without exaggerated radii (Fig. 24b). Lateral line incomplete, perforated scales 7 (X = 6.7, range 5—9, n = 48). Lateral series scales 45 (x = 44.3, range = 41-48, n = 48). Predorsal scales = 22 (xX = 22.3, range = 21-24, n = 46). Scale rows between dorsal- fin and anal-fin origins 19 (x = 18.8, range = 17-21, n= 55). Scale rows around caudal peduncle 20 (X = 19.7, range = 19-22, n = 46). Premaxillary teeth in 2 distinct rows, Fig. 25. Larger and smaller teeth tricuspid in all large specimens, sometimes in smaller spec- imens small teeth bicuspid or conical. Outer row teeth 5 (x = 6.2, range 5-8, n = 55). Inner row teeth few, 3 (X = 2.1, range = 1- 3, nm = 55). Outer and inner row premaxil- VOLUME 103, NUMBER 2 407 Fig. 23. Mimagoniates rheocharis, new species, paratype, unstained central basal region of caudal fin and _ adjacent caudal peduncle of adult male, lateral view, left side, anterior at left. USNM 279879, SL 49.0 mm; same locality data as specimen in Fig. 21. Arrow at left indicates anterior intake region of pump chamber. Middle arrow indicates region of lateral slit with ray half of principal ray 11 just dorsal to arrow tip. Arrow at right points to opening of posterior exhaust vent of pump chamber. Note that in this species pump chamber not formed into an obvious cylindrical tube as that of M. microlepis, Fig. 3. Glandular tissue surrounding chamber openings in this photograph obscure details of pump. lary teeth somewhat compressed compared to most “‘tetragonopterine” characid teeth which are often circular in cross section. Maxillary teeth 8 (X = 5.4, range = 3-8, larger specimens usually with higher counts, n= 55). All maxillary teeth usually tricuspid in large sepcimens; small specimens with posterior maxillary teeth often conical. Dentary with 4 large tricuspid teeth in all specimens, smaller posterior teeth 12 (X = 8.4, range = 5-12, almost always greater number of teeth in largest specimens, an- terior small maxillary teeth tricuspid, pos- terior ones conic, n = 54). Maxillary and dentary teeth shaped much like premaxil- lary teeth described above. No significant difference in tooth number found between males and females. Vertebrae 38 (X = 38, range = 37-40, n = 107). Dorsal limb gill rakers 7 (X = 6.6, range = 6-8, n = 54); ventral limb gill rakers 12 (x = 11.7, range = 11-13, n = 54). Bran- chiostegal rays 4, in 3 cleared and stained specimens, 3 rays originating on anterior ceratohyal and | ray from posterior cera- tohyal. Color in alcohol.—See Figs. 18-20 for preserved color pattern of males and fe- males. Body pale to medium brown, pale yellowish brown ventrally, much darker dorsally. Lateral body stripe diffuse in both sexes, especially anteriorly. Stripe extending from vertical humeral spot posteriorly to caudal fin and onto dorsal region of ventral caudal-fin lobe and small part of dorsal cau- dal-fin lobe. Lateral stripe extends over all 408 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bee de = eee o . pene 28 6 cay oa ees Pe ee Se EE Tes Coca ee) EAS = === 9 OE eet ND ees P [ee = Saueeagnacces = Le etic “lp ee LLB ae renee ae LD Tee = ae Sy ss ap BSB eo GE 85 eye 2S FE Coes er SS oS TS WG rT ——— SSS oss —=—— [So eee ae cs SAD ee a 8 PE es a ayy pee ee i ee ——— PEED Gea > Fig. 24. Mimagoniates rheocharis, new species, paratype, osteology of central basal region of caudal skeleton, principal fin rays 6-15, of an adult male, lateral views, left side, anterior at left. USNM 279879, SL 45.4 mm, same locality data as specimen in Fig. 21. Principal ray 12 indicated by arrow. (A) Illustrates caudal pump region. (B) illustrates relationship of modified dorsal caudal-fin lobe squamation to caudal pump. VOLUME 103, NUMBER 2 409 Fig. 25. Mimagoniates rheocharis, new species, paratype, jaws and dentition of adult male, lateral view, right side, anterior at right. USNM 279879, SL 45.4 mm, same locality data as in Fig. 21. For explanation see text Description under M. rheocharis. caudal gland structures, including those de- rived from dorsal caudal-fin lobe such as modified caudal squamation. Stripe espe- cially dark on principal rays 10, 11 and 12, less so on ray 13. Humeral spot vertically elongate, especially in sexually mature males. Remainder of caudal fin dusky be- cause of scattered dark chromatophores. Dorsal border of first principal caudal-fin ray and ventral border of nineteenth prin- cipal caudal-fin ray black. Utmost dorsal body surface black, forming a narrow stripe extending from supraoccipital region to base of dorsal procurrent rays of caudal fin. Re- mainder of dorsal body surface ventral to lateral body stripe pale brown. Pectoral, pelvic, dorsal and anal fins dusky with scattered dark chromatophores along fin rays and membranes. Anal fin with a dark elongate stripe running length of fin. 410 20.0 7 ™N D ° WS os S ie =~ ae te an LJ Soro > fa) © 0 co N © un oO 30.0 STANDARD LENGTH (mm) Mimagoniates rheocharis, RIO JORDAO, SANTA CATARINA, BRAZIL 26.0 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON O —+—— 1 40.0 50.0 (log) Fig. 26. Mimagoniates rheocharis, new species, paratypes, body depth as a function of SL by sex. Plot axes are logarithmic. Logarithmic regression equation for 27 males of M. rheocharis: (Y = —2.190 + 1.300X) and for 23 females of this species: (Y = —1.831 + 1.183X). For explanation see Sexual dimorphism under ™. rheocharis. Stripe width about one-fourth height of anal fin anteriorly and about one-half fin’s height posteriorly. Stripe of nearly uniform width throughout its length, but fin height changes, being considerably shorter posteriorly. In sexually mature males stripe considerably darker anteriorly, especially dorsal to an- terior anal-fin lobe which appears relatively hyaline. Dorsal-fin with a horizontal dark stripe in adult males and females extending posteriorly from about mid-length of an- terior elongate undivided ray to posterior tips of terminal two dorsal-fin rays. Stripe usually narrow, less than one-eighth height of dorsal fin at latter’s longest measurement. Width and density of stripe variable de- pending on sex and sexual maturity. Pre- served males sometimes with posterior por- tion of stripe diffuse. Adipose fin dusky with scattered dark chromatophores. Head dark brown around mouth and on dorsal surface of snout, between eyes, dor- sum of cranium and nape. Mental area of lower jaw dark brown. Head area posterior to circumorbitals and extending ventrally from parietal region, across dorsal opercular region dark brown. Dark area continues ventrally across posterior region of oper- cular bone to just reach interopercular bone; looks like an anteriorly misplaced humeral spot. Iris dorsal to pupil dark brown, most of remainder of iris silvery. Circumorbitals silvery if guanine preserved, pale yellowish brown if guanine destroyed by formalin. Dark brown chromatophores scattered evenly through circumorbital area. Anterior area of opercle, all of preopercle, and bran- chiostegal rays silvery or pale brown, with- out much dark brown pigment. Color in life. —Life colors described here taken from a 35 mm color slide made by Ricardo M. C. Castro of an adult male 41.8 mm SL (MZUSP 40281). See also black and white photograph of this specimen in pre- VOLUME 103, NUMBER 2 411 Mimagoniates inequalis, MORUNGAVA, RIO GRANDE DO SUL, BRAZIL Mimagoniates rheocharis, RIO GRANDE, SANTA CATARINA, BRAZIL iy =6f =o = BO on) (of D i SO 7/0) IS oa CAUDAL PEDUNCLE DEPTH (mm) 1.0 + 13.0 20.0 STANDARD LENGTH (mm) 30.0 (log) Mimagoniates microlepis, PARANAGUA, PARANA, BRAZIL © =of e=9 Fig. 27. Mimagoniates rheocharis, M. microlepis and M. inequalis caudal peduncle depth as a function of SL by species and sex. Plot axes are logarithmic. Logarithmic regression equation for 31 males of M. rheocharis: (Y = —4.222 + 1.588X), for 26 females of the same species: (Y = —3.311 + 1.374X); for 40 males of M. microlepis: (Y = —4.151 + 1.592X) and for 30 females of the later species: (Y = —3.239 + 1.374X); and for 26 males of M. inequalis: (Y = —3.021 + 1.3375X), 14 females of this species; (Y = —2.625 + 1.2116X). For explanation see Diagnosis, Sexual dimorphism, Discussion and Description under M. rheocharis. servative, Fig. 20. Specimen photographed just after capture from a clear water stream surrounded by vegetation. Site located im- mediately north of city of Osorio, Rio Grande do Sul. Sides of body pale silvery blue. Broad lateral body stripe somewhat deeper silvery blue from humeral spot to caudal peduncle termination. Just dorsal to silvery blue color of body sides, back with a narrow dark brown line extending from parietal region of head to just ventral to adipose fin. Lateral portion of back between narrow brown line and dorsomedian nar- row dark brown line extending across dor- salmost portion of back, a brownish yellow green color. Dorsal region of caudal pedun- cle nearly yellow. Ventral abdominal area, most of lower jaw, ventral opercular area, branchiostegal rays and their membranes silvery white. Dark pigment of head similar to that described for preserved specimens except that dorsal region of opercle appears silvery blue. Dorsal caudal-fin lobe and principal caudal-fin rays 14-16 on ventral caudal-fin lobe bright yellow, except for black proximal half of ray 14. This black pigment continuous with black pigment surrounding structures of caudal phero- 412 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON D Oo 8.0 o a“™N iS 7.0 & NY Sle = 6.0 oO LJ Q Ld rr 5.0 S) FZ =) SQ) Lo 4.0 au =| < Q =) Al P2-> A2~— P3. By stage 28, all tooth rows are keratinized. In our single stage 41 specimen, the A2 and P3 rows are reduced in length. The tail comprises 56% of the total length in stage 24 larvae and 60% of the total length in stage 41. The ratio of tail height to total length changes from 17% in stage 24 to 13% in stage 41. In the early stages (24 and 25), the vent tube is medial and the spiracle is located at mid-body; in later stages, the vent tube is dextral and the spiracle is low on the left side of the body. Changes in body length, body depth, body width, tail length, and total length are summarized in Table 1. Comparisons Among Species We have not examined the tadpole of Phyllobates bicolor but we can compare the tadpole of P. /ugubris to tadpoles of the re- maining species in the genus. All Phyllo- bates tadpoles examined have: depressed bodies (width > depth), sinistral spiracles, dorsal eyes and nostrils, low-finned tails with 430 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Measurements (mm) of various stages of tadpoles of Phyllobates lugubris collected in Costa Rica and Panama. Values are means + one standard deviation. Stage n Body length Body depth Body width Tail length Total length 24 3 4.30 + 0.361 2.00 + 0.300 Desa OMS 5.47 + 0.611 9.77 + 0.379 25 15 4.16 + 0.519 1.80 + 0.288 2.82 + 0.459 6.23 + 0.466 10.39 + 0.844 28 2 7.10 + 0.566 2.95 + 0.071 4.60 + 0.141 — — 37 2 10.30 + 0.000 4.35 + 0.212 6.90 + 0.141 14.05 + 0.636 24.35 + 0.636 41 1 9.0 4.0 6.5 13.7 Doll rounded tips, emarginate oral discs, two an- terior and three posterior labial tooth rows, with the second anterior row interrupted medially, serrated jaw sheaths, and rounded papillae on the posterior and lateral margins of the oral disc. The mouthparts of Phyllobates tadpoles we examined undergo ontogenetic changes similar to those observed in P. /ugubris and P. terribilis. In some stage 25 larvae of P. vittatus, the third posterior row of labial teeth is not keratinized and the jaw sheaths lack serrations. Although Silverstone (1976) stated that Phyllobates vittatus larvae have two rows of labial papillae, all stage 25 tad- poles we examined have a single row of mar- ginal papillae. By stage 28, Phyllobates vit- tatus larvae have two rows of papillae and all labial tooth rows are keratinized. In stage 25 Phyllobates aurotaenia tadpoles, the la- bial tooth rows are not keratinized, the jaw sheath edges are smooth, and there is one row of marginal papillae on the posterior labium and one row of submarginal pa- pillae. By stage 37, Phyllobates aurotaenia tadpoles have keratinized labial teeth, ser- rated jaw sheaths, and two rows of papillae. Ontogenetic changes in oral papillae have been reported for some hylids (Donnelly et al. 1987, Zweifel 1961). The vent tube location changes ontoge- netically in the four species. In early stages (24 and 25) the vent tube is medial; it be- comes dextral in Phyllobates lugubris and P. vittatus by stage 28. The developmental series of P. aurotaenia and P. terribilis lack specimens in stage 28; both species have dextral vent tubes by stage 37. Myers et al. (1978) described changes in the relationship between tail height and to- tal length for Phyllobates terribilis; tail height equals 12-15% of total length in stages 24— 27 and increases to a maximum of 20% of total length in later stages. Similar changes in tail height relative to total length occur in Phyllobates aurotaenia and P. vittatus. In Phyllobates lugubris, tail height does not in- crease relative to increasing total length. Phyllobates tadpoles collected from male nurse frogs have been in stage 24 or 25; at this stage of development, the mouthparts of the species we examined were not com- pletely developed. Measurements of stage 25 larvae for P. aurotaenia, P. terribilis, and P. vittatus are summarized in Table 2. The tadpole of P. vittatus is larger than tadpoles Table 2. Measurements (mm) of stage 25 Phyllobates tadpoles. The following abbreviations identify the species: Pa = P. aurotaenia, Pt = P. terribilis, Pv = P. vittatus. Values are means + one standard deviation. Species n Body length Body depth Body width Tail length Total length Pa 6 3.75 + 0.138 1.57 + 0.103 2.37 + 0.121 6.73 = 0.175 10.48 + 0.306 Pt 12 4.38 + 0.175 1.89 + 0.131 2.98 + 0.171 7.302 + 0.258 11.652 + 0.334 Pv 7 4.40 + 0.883 2.04 + 0.395 3.07 + 0.170 8.08° + 0.898 12.70" + 1.138 n= 10. bn=5 VOLUME 103, NUMBER 2 Table 3. Tadpole size (mean total length) at stage 25 and adult mean snout—vent length (SVL) in Phyl- lobates. Body size data for P. terribilis from Myers et al. (1978), for all other species from Silverstone (1976). Tadpole total Male SVL Female SVL Species length (mm) (mm) (mm) P. aurotaenia 10.48 26.9 30.4 P. lugubris 10.39 19.2 DD) P. terribilis 11.65 41.1 43.2 P. vittatus 12.70 24.4 DSP of the other species (Tables 1, 2); and its oral disc is wider (P. vittatus, 1.22 mm, P. aurotaenia, 0.80 mm; P. /ugubris, 1.17 mm; P. terribilis, 1.09 mm). Although absolute size varies among the stage 25 tadpoles, the relationship between body width and body depth is similar (body width/body depth = 1.51 in Phyllobates aurotaenia, 1.57 in P. lugubris, and 1.58 in P. terribilis and P. vit- tatus). The relationship between tail length and total length in stage 25 tadpoles is also similar among species; the tail accounts for 64% of total length in Phyllobates auro- taenia and P. vittatus, 60% of total length in P. lugubris, and 63% in P. terribilis. No clear relationship exists between tadpole size and adult body size in Phyllobates (Table 3). Our observations agree with those of Myers et al. (1978); accurate identification of Phyllobates tadpoles in early (stage 24 or 25) developmental stages requires nurse frog identification. Phyllobates tadpoles in these stages exhibit little variation in larval char- acteristics other than absolute size. We ex- pect that the tadpole of Phyllobates bicolor will resemble those of the other species. Acknowledgments We thank the Organization for Tropical Studies for supporting our research through Jessie Smith Noyes pre-doctoral fellowships granted to MAD and CG. We thank C. W. Myers for permission to report on Phyllo- bates tadpoles he raised in the laboratory and those he collected in Panama and Co- lombia. We thank J. M. Savage for allowing us to examine tadpoles of Phyllobates vit- 431 tatus. R. Altig, B. I. Crother, D. M. Hillis, C. W. Myers, J. M. Savage, and two anon- ymous reviewers read and commented on the manuscript. Literature Cited Altig, R., & G. F. Johnston. 1986. Major character- istics of free-living anuran tadpoles.—Smith- sonian Herpetological Information Service 67: 1-75. Donnelly, M. A., C. Guyer, D. M. Krempels, & H. E. Braker. 1987. The tadpole of Agalychnis cal- carifer (Anura: Hylidae).—Copeia 1987:247- 250. Gosner, K. 1960. A simplified table for staging an- uran embryos and larvae with notes on identi- fication. — Herpetologica 16:183-190. Myers, C. W. 1987. New generic names for some Neotropical poison frogs (Dendrobatidae).— Papeis Avulsos de Zoologica 36:301-306. —., J. W. Daly, & B. Malkin. 1978. A dangerously toxic new frog (Phyllobates) used by Embera indians of western Colombia, with discussion of blowgun fabrication and dart poisoning.— Bulletin American Museum Natural History 161: 307-366. Savage, J. M. 1968. The dendrobatid frogs of Central America.—Copeia 1968:745-776. —, & J. Villa. 1986. Introduction to the herpe- tofauna of Costa Rica/Introduccion a la Her- petofauna de Costa Rica.—Contributions to Herpetology. Number 3. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 220 pp. Silverstone, P. A. 1976. A revision of the poison- arrow frogs of the genus Phyllobates Bibron in Sagra (family Dendrobatidae).—Los Angeles County Museum of Natural History Science Bulletin 27:1-53. Starrett, P. 1960. Descriptions of tadpoles of Middle American frogs.—Miscellaneous Publications Museum Zoology University Michigan 110:1- 37 + 1 plate. Zweifel, R.G. 1961. Larval development of the tree frogs Hyla arenicolor and Hyla wrightorum. — American Museum Novitates 2056:1-19. (MAD) Department of Herpetology and Ichthyology, American Museum of Natural History, New York, New York 10024-5192; (CG) Department of Zoology and Wildlife Science, Auburn University, Auburn, Ala- bama 36849; (ROS) Department of Zool- ogy, University of Texas, Austin, Texas 78712-1064. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 432-451 TAXA OF NORTH AMERICAN BIRDS DESCRIBED FROM 1957 TO 1987 M. Ralph Browning Abstract. — Ninety-nine names proposed from 1957 to 1987 for North Amer- ican birds are evaluated. Of these, 35 are judged taxonomically distinct; seven are recognized provisionally; 54 are considered synonyms of established names; and three forms cannot be identified positively. Five names are taxonomic changes, providing new names for forms previously described. Since the publication of the fifth edition of the A.O.U. Check-list (American Orni- thologists’ Union 1957), more than 800 forms of birds, world-wide, have been new- ly described; 99 of these are from North America (sensu A.O.U. 1957). About half the names of these North American forms already have been considered, by various authors, to be synonyms of previously ex- isting names, but others have received little or no taxonomic attention. In the following accounts I summarize the taxonomic status of the taxa of North American birds for which names have been proposed since 1957. With minor excep- tions I evaluated each form using the com- parative material available to the original author, including, where possible, the ho- lotypes and paratypes. In evaluating the forms I emphasized the range of variation more than the average difference in any giv- en character. My standard for recognizing subspecies primarily on color has been more stringent than the so-called “75 percent rule’; statements in the accounts that a pop- ulation differs from another means that at least 95 percent of the specimens I com- pared could be identified. Each account includes the original name with the authority and year; complete ci- tations are given in the Literature Cited. The type locality, modified when required, follows. The depository of holotypes or syn- types, when known, is abbreviated and ap- pears in parentheses if I examined the spec- imens or in brackets if not. Abbreviations are: California Academy of Sciences (CAS); Cleveland Museum of Natural History (CMNH); Delaware Museum of Natural History (DMNH); Field Museum of Natu- ral History (FM); Louisiana State Univer- sity Museum of Natural Science (LSUMNS); National Museum of Canada (NMC); pri- vate collection of Amadeo Rea (AMR); Mu- seum of Vertebrate Zoology, University of California at Berkeley (MVZ); Royal On- tario Museum (ROM); San Diego Museum of Natural History (SDMNH); Texas Co- operative Wildlife Collection, Texas A&M University (TCWC); James Ford Bell Mu- seum of Natural History, University of Minnesota (UMM); University of Michigan Museum of Zoology (UMMZ); Utah Mu- seum of Natural History (UMNH); and U.S. National Museum of Natural History (USNM). The present taxonomic status of each new name is next. References with the present name include sources that have provided data or an opinion on the taxonomic status of that name. I summarize characters and ranges, including a more detailed charac- terization as warranted by my study. Com- ments are omitted for forms discussed in my earlier papers (Browning 1974, 1977, 1978, 1979a). Names of genera (except PI- VOLUME 103, NUMBER 2 coides) and species, and sequence of the families, follow the sixth edition A.O.U. check-list (1983) and A.O.U. (1989). Podicipedidae Aechmophorus clarkii transitionalis Dick- erman, PBSW 99(3), p. 436, 17 Oct 1986.—Silver Lake, Lake County, Ore- gon (USNM). =Aechmophorus clarkii transitionalis. Birds from northern North America are larger (Dickerman 1986a) than nominate clarkii of Mexico (Dickerman 1963, 1973). Hydrobatidae Oceanodroma leucorhoa cheimomnestes Ainley, 1980.—Guadalupe Island, Mex- ico (USNM). =Oceanodroma (?leucorhoa) cheimom- nestes. The birds breeding on Guadalupe Island during the summer were assigned to socor- roensis by Ainley (1980), who differentiated cheimomnestes, the subspecies breeding there during winter, by physiological, mor- phological, and vocal characteristics. Bourne & Jehl (1982) synonymized cheimomnestes with O. /. beali. Power & Ainley (1986) fur- ther demonstrated temporal and morpho- logical differences between the summer (So- corroensis) and winter (cheimomnestes) birds breeding on Guadalupe Island. I agree with Jehl & Everett (1985) that further study is required to determine the status of the two Guadalupe Island populations. Ardeidae Leucophoyx thula arileuca Oberholser, 1974.—mouth of Bear River, North Bay (=Bear River Bay), Great Salt Lake, Box Elder Co., Utah (USNM). =Egretta thula brewsteri Thayer & Banks, 1909 (see Browning 1974, Behle 1985). 433 Anatidae Anser albifrons elgasi Delacour & Ripley, 1975.—“‘Sacramento, California” = Sac- ramento National Wildlife Refuge, Cali- fornia (USNM). =Anser albifrons elgasi (see Cramp 1977, Krogman 1979). This subspecies differs from A. a. gam- belli (sensu Delacour and Ripley 1975) and frontalis by its larger size and darker col- oration. Anser a. elgasi winters in the Sac- ramento Valley, California. The breeding range was not known at the time of the de- scription, but birds conforming to the de- scription of e/gasi breed in Alaska’s Cook Inlet (Timm et al. 1982). Palmer (1976) and Bellrose (1976) synonymized e/gasi with gambelli but Johnsgard (1979) and Godfrey (1986) recognized the subspecies. Anas platyrhyncha neoboria Oberholser, 1974.—Athabaska River, La Lasine, Al- berta (USNM). =Anas platyrhynchos platyrhynchos Lin- naeus, 1758 (see Browning 1974, 1978). Falconidae Falco peregrinus tundrius White, 1968.— near NW Sherman Basin, Adelaide Pen- insula, Northwest Territories, Canada [NMC]. =Falco peregrinus tundrius (see Palmer 1988, White & Boyce 1988). This widely accepted, northern subspe- cies differs most from F. p. anatum by its smaller size, paler color, narrower malar stripe and in immatures by the narrower ventral linear stripes (White 1968). It breeds on the tundra from Alaska to Labrador and western Greenland and migrates as far south as Argentina. Phasianidae Callipepla squamata hargravei Rea, 1973.— Pepper Ranch, 7 miles N, 32 miles E of 434 Folsom, Union Co., New Mexico [LSUMNS, ex G. M. Sutton]. =Callipepla squamata hargravei. The population from the northeastern portion of the range of pallida was described as paler than other examples of the species. I examined over 100 specimens from the range of pallida (sensu A.O.U. 1957) and 9 specimens from Rea’s (1973) comparative series. I found that the throat and sides of the head of eastern specimens are paler gray and less brownish and the belly is paler and usually less buffy than specimens from Ar- izona and western New Mexico. In fresh fall plumage specimens of hargravei also differ ventrally from pallida by having the ter- minal bars of the “‘scaled’’ areas more brownish and less black. Post-mortem color changes and birds possibly introduced from the western pop- ulation (Rea 1973) complicated compari- sons of specimens from the range of har- gravel. Callipepla s. hargravei is resident from southeastern Colorado to western Oklahoma, southwestern Kansas, northern New Mexico and northwestern Texas. Gruidae Grus canadensis rowani Walkinshaw, Jul— Aug 1965.—10 miles west of Fawcett, Al- berta [FM]. =Grus canadensis rowani (see Johnson & Stewart 1973, Aldrich 1979). This northern interior subspecies is in- termediate in size to nominate canadensis and the more southern tabida (see Johnson & Stewart 1973). Color of the primary shafts, described by Walkinshaw (1965) as paler than in nominate canadensis is subject to individual variation (Aldrich 1979). Grus c. rowani breeds from the Mackenzie District to central Alberta, Saskatchewan, northern Ontario, and possibly central British Co- lumbia; it winters in New Mexico, Okla- homa, and Texas. Grus canadensis pulla Aldrich, 1972.—Cap- tive bird hatched from egg taken 7 miles PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON northwest of Fontainebleau, Jackson Co., Mississippi (USNM). =Grus canadensis pulla (see Walkinshaw 1973). This taxon was described as the darkest subspecies of Grus canadensis. It is similar in overall size to pratensis, smaller (except tarsus) than tabida, larger than nominate canadensis, and has longer tarsi than ro- wani. Grus c. pulla formerly occurred on the Gulf Coastal Plain of Louisiana, Mississip- pi, and Alabama (see Aldrich 1972), but it is now resident only in Jackson County, Mississippi. Scolopacidae Actitis macularia rava Burleigh, 1960.— Lewiston, Nez Perce Co., Idaho (USNM). =Actitis macularia (Linnaeus, 1766). Burleigh (1960a) described rava, from the western range of A. macularia, as darker gray (less brown) above, with the ventral spots more intensely black and less densely distributed than in eastern specimens of macularia. Although rava was recognized by Wetmore (1965a, b) and Sutton (1967), both authors characterized the form as paler above (contra Burleigh 1960a). My exami- nation of 52 specimens from the breeding grounds of A. macularia revealed no geo- graphic variation. Laridae Larus californicus albertaensis Jehl, 1987.— Frog Lake (53°55'N, 110°15’W), Alberta [UMMZ]. =Larus californicus albertaensis. Although only four adults from the north- ern population of this species were exam- ined, I found them easily distinguishable from 24 adults of nominate californicus to the south by its greater overall size, espe- cially the bill, and paler gray mantle. It nests from the Northwest Territories to Alberta, Saskatchewan, and North Dakota. The win- ter distribution is presumably throughout VOLUME 103, NUMBER 2 the species’ range, mainly along the Pacific coast from British Columbia to Baja Cali- fornia (Jehl 1987). Columbidae Zenaida asiatica grandis Saunders, 1968.— Near Ruidosa, Presidio Co., Texas, alti- tude about 3000 ft (USNM). = Zenaida asiatica mearnsi (Ridgway, 1915). Saunders (1968) described this popula- tion from western Texas as larger than mearnsi of the northwestern range of the species, asiatica of southern Texas and northern Mexico, and monticola Saunders, 1968 (type locality 11 miles S Acatan, Pueb- lo, Mexico). He also described grandis as paler than nominate asiatica, and slightly browner on the back and head than mon- ticola. The subspecies monticola, from the interior plateau of Mexico, was described by Saunders (1968) as having a longer wing chord and tail than asiatica and mearnsi, and to be grayer with a shorter bill than the latter. This species is subject to considerable in- dividual variation in color and size (Aldrich 1981). Some populations measured by Saunders are statistically different in some characters, but overlap is nearly complete in the mensural ranges. Most of the speci- mens I compared were collected by Saun- ders including about 60 specimens of mearnsi, 22 monticola, 15 grandis, and 50 nominate asiatica. I found that both dorsal and ventral coloration varies individually. I place grandis and monticola in the syn- onymy of mearnsi. Saunders (1968) also named other subspecies of Z. asiatica from regions geographically beyond the scope of this paper. Cuculidae Geococcyx californianus dromicus Ober- holser, 1974.—Brownsville, Cameron Co., Texas (USNM). =Geococcyx californianus (Lesson), 1829 (see Browning 1978, but see Rea 1983). 435 Rea (1983) suggested that the overlap in measurements of dromicus and birds from California and Arizona given by Oberholser (1974) might be the result of the latter’s use of missexed specimens and that dromicus might prove to be a “useful” subspecies. I find no reason to believe that Oberholser’s series were missexed. Strigidae Otus flammeolus borealis Hekstra, 1982.— Penticton, Okanagan Valley, shore of Okanagan Lake, British Columbia [MVZ]. =Otus flammeolus idahoensis Merriam, 1892. Hekstra (1982) described borealis as slightly larger, duller gray, and less white than idahoensis (type locality Ketchum, Blaine Co., Idaho). Specimens from the stated range of borealis, from interior Brit- ish Columbia (east of the Cascade Moun- tains) to northeastern California, are within the range of individual variation of ida- hoensis 1n size and plumage characters (J. T. Marshall, pers. comm.). Otus flammeolus frontalis Hekstra, 1982.— Estes Park, Larimer Co., Colorado (USNM). =Otus flammeolus frontalis. This subspecies was described as much darker with more extensive chestnut brown facial discs than O. f flammeolus, the sub- species found to the south. Differences be- tween frontalis and other populations of the species were not given. My series consisted of 11 specimens of frontalis and 10 speci- mens of idahoensis Merriam. The new sub- species differs from flammeolus and ida- hoensis by its wider ventral streaks and darker facial discs (J. T. Marshall, pers. comm.). The range of frontalis extends from the Rocky Mountains and, probably, most of the Great Basin. Bubo virginianus scalariventris Snyder, 1961.—Elsas, on the upper Kapsuskasing 436 River, in Algoma District, Ontario [ROM]. =Bubo virginianus subsp.? Snyder (1961) proposed scalariventris for the population from part of the eastern range of subarcticus Hoy, 1852 (=wapacuthu Gmelin of A.O.U. [1957] but see Manning [1952]). He distinguished scalariventris as ‘“‘more coldly grey with bolder bars below” than subarcticus and that it differs from nominate virginianus by the lack of rufous and “‘virtual lack of large, vague, finely ver- miculated spots superimposed over the barred ventral pattern... .’’ Johnsgard (1988) recognized scalariventris, but he stat- ed that it “possibly should be included in subarcticus.” 1 join Godfrey (1986), who stated that he was unable to express an opin- ion on the validity of scalariventris. Caprimulgidae Chordeiles minor divisus Oberholser, 1974.— Loveland, Larimer Co., Colorado (USNM). =Chordeiles minor hesperis Grinnell, 1905 (see Browning 1978). Trochilidae Phasmornis mystica Oberholser, Texas birds, 1974:485.—Boot Springs, Chisos Mountains, Texas (holotype unknown). =hybrid or abberant individual (see Brown- ing 1978, Mayr & Vuilleumier 1983). Lampornis clemenciae phasmorus Ober- holser, 1974.—northeastern side of Chi- sos Mountains, Pine Canyon, 6000 ft, Brewster Co., Texas (USNM). =Lampornis clemenciae phasmorus (see Browning 1978). Picidae Dendrocopos scalaris soulei Banks, 1963.— Ruffo’s Ranch, Cerralvo Island, Baja Cal- ifornia, Mexico [CAS]. =Dendrocopos scalaris soulei. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The form was described as similar in size to /ucascanus of southern Baja California, and intermediate to eremicus of northern Baja California and sinaloensis of mainland Mexico. Dendrocopos s. soulei was de- scribed as more grayish ventrally and more extensively black on the three outer rectrices and with larger and more abundant ventral spots than /ucasanus. From cactophilus of the southwestern United States, the sub- species was diagnosed as darker above with spotted rather than streaked flanks. Eight specimens of the endemic popula- tion from Cerralvo Island are distinctly pal- er and less buffy below than larger series of other subspecies of this species. Also, the ventral spots are more rounded than those of sinaloensis. I did not detect the described differences in the amount of black on the three outer rectrices between /ucasanus and soulei. The feathers immediately above the bill are blackish in sou/ei and brownish in lucasanus. Short (1968) reviewed the species’ southwestern forms but neglected to men- tion the birds from Cerralvo Island, and later he (Short 1982) synonymized soulei with /ucasanus. Dendrocopos s. soulei is en- demic to Cerralvo Island, Baja California. Dendrocopos pubescens parvirostris Bur- leigh (1960) = 1961.—Moscow. Latah Co., Idaho (USNM). =Dendrocopos pubescens leucurus (Hart- laub. 1852) (see Short 1982). Burleigh (1961) separated birds from southern British Columbia to eastern Ore- gon and most of Idaho from the wide rang- ing /eucurus. He characterized parvirostris as smaller, or similar in size to the darker western furati. The characters given for par- virostris are within the range of individual variation of /eucurus (sensu Short 1982). Tyrannidae Contopus sordidulus amplus Burleigh, 1960[b].—Havre, Hill Co., Montana (USNM). VOLUME 103, NUMBER 2 =Contopus sordidulus vieilei Coues, 1866 (see Browning 1974). Contopus sordidulus siccicola Burleigh, 1960[b].— Potlatch, Latah Co., Idaho (USNM). =Contopus sordidulus vieilei Baird, 1866 (see Browning 1974). Empidonax oberholseri spodius Oberholser, 1974.—Gray, Bonneville Co., Idaho (USNM). =Empidonax oberholseri Phillips, 1939 (see Browning 1974, 1978). Hirundinidae Progne subis arboricola Behle, 1968.—Pay- son Lakes, 8300 ft elevation, 12 miles southeast Payson, Utah Co., Utah [UMNH]. =Progne subis arboricola (see Phillips 1986). Behle (1968) gave the range of arboricola as Utah, northern Arizona, and northern Mexico. According to Phillips (1986), ar- boricola breeds from southwestern British Columbia to central Baja California, locally in northwestern Mexico, and to the Trans- Pecos region of Texas, and winters in South America. I compared 43 specimens from the east- ern United States, 15 from Arizona, and 9 from Utah and conclude that specimens of arboricola are larger than those of hesperia and nominate subis, and, in females, usually whiter than nominate subis. Hubbard (1972) considered birds from southeastern Arizona to be intermediate between arboricola and nominate subis. | agree with Hubbard (1972) that a broader review is necessary to estab- lish the validity of this subspecies, but I recognize arboricola provisionally. Hirundo albifrons ganieri Phillips, 1986.— Swallow Bluff, Decatur Co., Tennessee [LSU]. =Hirundo pyrrhonota ganieri. Phillips (1986) briefly described ganieri as shorter in wing chord than nominate pyr- 437 rhonota (=albifrons of Phillips), but he did not otherwise differentiate the two taxa. My evaluation of ganieri was based on 20 spec- imens collected before the reported (Phillips 1986:33; see also A.O.U. 1931, 1957) spread of this species to the southeastern United States. There is overlap in wing chord (see Phillips 1986) between pyrrhonota and ga- nieri. 1 found that the extent of black on the throat and the pale chestnut wash of the upper breast are noticeably reduced in ga- nieri. The chestnut of the undertail coverts is also darker than in pyrrhonota. In females the rump of ganieri averages paler than in pyrrhonota. Hirundo p. ganieri breeds west of the Appalachians from Tennessee to south-central Texas and migrates to Mexico and Middle and South America. Corvidae Cyanocitta cristata burleighi Bond, 1962.— South Brook, Newfoundland (USNM). =Cyanocitta cristata cristata (Linnaeus, 1758) (see Phillips 1986). The darker northern birds described by Bond (1962) represent the end of a cline, and many northern individuals are similar in coloration to populations to the south. Aphelocoma coerulescens suttoni Phillips, 1965.—Scroogs’ Arroyo, 35 km S Pueblo, Colorado (DMNH). =Aphelocoma coerulescens suttoni (Phil- lips) (see Browning 1974, 1978). Aphelocoma coerulescens mesolega Ober- holser, 1974.— Fort Davis, Jeff Davis Co., Texas (USNM). =Aphelocoma coerulescens suttoni Phillips, 1964 (see Browning 1978, Phillips 1986). Corvus cryptoleucus reai Phillips, 1986.—3 km NE Nogales, southern-most Arizona. =Corvus cryptoleucus Couch, 1854. This subspecies was described as differing from nominate cryptoleucus of the Laredo region of Texas and adjacent northeastern Mexico by having a greater wing chord and 438 longer tail. Most of the range of the species (see A.O.U. 1983) was included in reai. Phillips’ (1986) brief description shows overlap in wing chord and tail for the two forms and included specimens that he stated might not be correctly sexed and fully adult. A study of the variation among definitely sexed and aged birds is needed before sub- species can be recognized. Paridae Parus rufescens caliginosus Burleigh, 1959.—Twenty miles northeast of Mos- cow, Latah Co., Idaho (USNM). Preoc- cupied by Parus wollweberi caliginosus van Rossem, 1947. Parus rufescens levyi Burleigh, 1968. New name for P. r. caliginosus Burleigh, 1959. =Parus rufescens rufescens Townsend, 1837 (see Phillips 1986). Burleigh (1959) characterized the pop- ulations from the southeastern part of the range of nominate rufescens, which he later renamed /evyi, as darker and less reddish than the northwestern populations. I agree with Phillips (1986) that the considerable individual variation of rufescens (sensu A.O.U. 1957) precludes recognition of /evyi. Parus wollweberi vandevenderi Rea, in Phil- lips 1986.—Sycamore Creek, 1.2-—3.3 miles (1.9—5.3 km) N Sunflower, NE Mar- icopa Co., Arizona [SD]. =Parus wollweberi vandevenderi. Based on examination of over 150 spec- imens of the species, I conclude that spec- imens of vandevenderi are darker and duller than phillipsi of southeastern Arizona and New Mexico. The subspecies is resident from central (Yavapai Co.) and eastern counties in Arizona to southwestern (Ca- tron Co.) and western (Grant Co.) New Mexico but occasionally occurs in winter in the Lower Sonoran Zone of its range and to southern Arizona. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sittidae Sitta canadensis clariterga Burleigh, 1960{a]. — Headquarters, Clearwater Co., Idaho (USNM). =Sitta canadensis Linnaeus, 1766 (see Banks 1970, Phillips 1986). Banks (1970) found that the supposed variation between eastern and western spec- imens compared by Burleigh (1960a) was the result of seasonal differences between the series, sooting of birds in eastern in- dustralized areas, and variation in the qual- ity of the prepared specimens. Certhiidae Certhia americana alascensis Webster, in Phillips 1986.—Ft. Wainwright, Fair- banks, Alaska (USNM, ex Univ. Alaska). =Certhia americana alascensis. Eight available specimens from south- central Alaska are paler dorsally with larger grayish white streaks on the back than ex- amples of montana from the northwestern interior and than occidentalis, the western coastal subspecies. Webster (in Phillips 1986) believed that specimens from the southern coast of Alaska are intermediate to alascensis and occidentalis, but I regard this as an area of intergradation between occidentalis and montana. Certhia americana stewarti Webster, in Phillips 1986.— Masset, Queen Charlotte Is., British Columbia [FM]. =Certhia americana stewart. The population from the Queen Charlotte Islands, separated from occidentalis, was described as “more orange or brighter or- ange dorsally than any other race.” I ex- amined 29 specimens of occidentalis and found that dorsal.coloration appears to be clinal. Seven specimens from coastal British Columbia and a specimen from Juneau are dorsally browner than the three available specimens of stewarti but those from west- VOLUME 103, NUMBER 2 ern Washington are slightly more orange. Specimens from western Oregon are less or- ange dorsally than those from Washington and also differ from those to the north by their darker heads. Specimens from the cen- tral coast of California are similar to those from western Oregon, but are buffy below (Webster, in Phillips 1986:200). Certhia americana idahoensis Webster, in Phillips 1986.—Coolin, N Bonner Co., Idaho [FM]. =Certhia americana montana Ridgway, 1882. Webster’s description of idahoensis (in Phillips 1986) did not provide a clear com- parison with the adjoining subspecies. The breeding range of idahoensis was given as central Alberta, extreme northern Idaho, and northwestern Montana. I found that the bill length and color of 50 specimens of ida- hoensis from Idaho and northwestern Mon- tana are similar to other specimens of mon- tana. I agree with Webster that the northern limits of ze/otes include the eastern slope of the Cascade Mountains in Oregon. How- ever, the name caurina Aldrich, 1946 (type locality Mt. Adams, Yakima County, Washington), listed as a synonym of mon- tana by Webster, may refer to pale individ- uals of zelotes of California. The name /eu- costicta, synonymized with montana by Webster, is a recognizable subspecies (con- tra Johnson 1965, see Austin & Rea 1976) and, according to Behle (1985), ranges as far north as west-central Utah. Certhia familiaris iletica Oberholser, 1974.—The Bowl, Guadalupe Moun- tains, Culberson Co., Texas (USNM). =Certhia americana montana Ridgway, 1882 (see Browning 1978, Webster, in Phillips 1986). Troglodytidae Campylorhynchus brunneicapillum sandi- egense Rea, in Phillips 1986.—3.7 km W 439 San Pasqual, west-central San Diego Co., California [SD]. =Campylorhynchus brunneicapillus sandi- egensis. This geographically isolated subspecies has overall coloration somewhat interme- diate to anthonyi of east-central California and the subspecies of Baja California, and differs from other subspecies as character- ized by Rea (in Phillips 1986). Campylo- rhynchus b. sandiegensis is a resident sub- species found from coastal Orange, Los Angeles and Ventura counties of California south to northwestern Baja California (Car- rizo Val., Val. Palmas). Catherpes mexicanus pallidior Phillips, 1986.—Green River, Wyoming [CM]. =Catherpes mexicanus pallidior. Comparisons of ten available specimens from the range of pallidior and a larger series of other C. mexicanus revealed that the sub- species is the palest taxa of the species. Catherpes m. pallidior breeds from eastern Montana to northwestern Colorado and northeastern Utah; the winter range is not known. Catherpes mexicanus croizati Phillips, 1986.—La Laguna, Sierra Laguna, Baja California. =Catherpes mexicanus croizati. This resident subspecies of southern Baja California was described by Phillips (1986) as “‘like the preceding races” but the “red- dest” brown dorsally, and the “‘warmest,” especially ventrally, form of the species. I compared 60 specimens from the south- western range of the species and found that croizati is darker dorsally than conspersus from southern California and central Baja California and pallidior, and is similar in this character to the northwestern subspe- cies, griseus, and punctulatus Ridgway, 1882 (type locality Forest Hill, Placer Co., Cali- fornia). Below, croizati approaches pallidior 440 in paleness, and is paler ventrally than the other northern subspecies. Thryothorus ludovicianus nesophilus Ste- venson, 1973.— Dog Island, Franklin Co., Florida (USNM). =Thryothorus ludovicianus nesophilus (see Phillips 1986). Known only from limited topotypical material, nesophilus was described by Ste- venson (1973) as intermediate to burleighi and nominate /udovicianus in color. I rec- ognize it provisionally. Troglodytes bewickii pulichi Phillips, 1986.—Dallas, Texas. =Thryomanes bewickii pulichi. As characterized, this south-central form is paler than calophonus, less reddish brown than nominate bewickii and more reddish below than the other southern and western forms of the species. Thryomanes b. pulichi is resident in Oklahoma and most of Kan- sas, and also winters south to south-central Texas. Troglodytes bewickii anthonyi Rea, in Phil- lips 1986. New name for Thryothorus leu- cophrys Anthony, 1895, preoccupied by Troglodytes leucophrys Tschudi, 1844 (=Henicorhina leucophrys). =Thryomanes bewickii leucophrys (Antho- ny, 1895). This new name is necessary only if the genus Thryomanes is merged with Troglo- dytes as by Phillips (1986). Troglodytes troglodytes ochroleucus Rea, in Phillips 1986.—near Killisnoo, Admiral- ty Island, SE Alaska (USNM). Troglodytes troglodytes muiri Rea, in Phil- lips 1986.— Navarro River, 7 km inland, Mendocino Co., California [AMR]. Troglodytes troglodytes obscurior Rea, in Phillips 1986.—North Fork Cosumnes River, 977 m, Grizzly Flats area, El Do- rado Co., California [SD]. =Troglodytes troglodytes pacificus Baird, 1864. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON These three forms were named from the range of pacificus (sensu A.O.U. 1957). Rea (in Phillips 1986) confined the range of pa- cificus to Prince of Wales Island, south- eastern Alaska, and probably the Queen Charlotte Islands, British Columbia. My re- view of the variation in some of Rea’s ma- terial suggests that these taxa are valid. However, recognition of these putative sub- species cannot be confirmed until a thor- ough analysis of the geographic variation of the western populations is presented. Troglodytes troglodytes salebrosus Burleigh, 1959.— Dismal Lake, Shoshone Co., Ida- ho (USNM). =Troglodytes troglodytes salebrosus (see Rea, in Phillips 1986). This northern interior subspecies is less rufescent (Burleigh 1959) than pacificus (sensu A.O.U. 1957; Rea, in Phillips 1986). Troglodytes t. salebrosus breeds from south- ern British Columbia, east of the coast range, to southwestern Alberta, northeastern Washington, eastern Oregon, northern Ida- ho, and western Montana. It winters in its breeding range and casually southward. Telmatodytes palustris canniphonus Ober- holser, 1974.—4 miles N Sandusky, Bay Point, Ottawa Co., Ohio (CMNH). =Cistothorus palustris dissaeptus (Bangs) 1902 (see Browning 1978, Phillips 1986). Telmatodytes palustris cryphius Oberholser, 1974.—Blackmer, Richland Co., North Dakota (USNM). =Cistothorus palustris iliacus Ridgway, 1903 (see Browning 1978, Phillips 1986). Cistothorus palustris browningi Rea, in Phil- lips, Known birds, 1986:114.—Pitt Meadows, SW _ British Columbia [DMNH]. =Cistothorus palustris browning. This is the grayest and least rufescent of the Pacific Northwest coastal subspecies. Thirty-three specimens of browningi are darker throughout than an equal number of paludicola of the southern coast of Wash- VOLUME 103, NUMBER 2 ington and central coast of Oregon. The black of the crown of browningi is greatly reduced to nearly obsolete. It breeds from southwestern British Columbia, including Vancouver Island (formerly ?), to the Puget Sound region and to Thurston Co. in central western Washington, and winters south- ward to northwestern Pacific County in southwestern Washington. Cistothorus palustris deserticola Rea, in Phillips 1986.— New River, 3.2 km NNW Seeley, Imperial Co., central-southern California [SD]. =Cistothorus palustris deserticola. This subspecies of the southwestern U.S. is distinguished from the coastal subspecies by its wider dorsal streaks. Based on 7 spec- imens, I conclude that deserticola is more rufescent than aestuarinus of central Cali- fornia and is less intensely brown than pal- udicola of coastal Washington and Oregon. Cistothorus p. deserticola is resident in the Salton Sea area of southeastern California and south-central Arizona (extirpated from the Gila River drainage, Rea in litt.), and north along the lower Colorado River to southern Nevada. I provisionally recognize deserticola. Muscicapidae Regulinae Regulus calendula arizonensis Phillips, 1965.—vicinity of Phelps Ranger Station, White Mountains, Arizona (DMNH). =Regulus calendula calendula Linnaeus, 1766 (see Hubbard & Crossin 1974, Browning 1979a). Turdinae Hylocichla fuscescens subpallida Burleigh and Duvall, 1959.—Moscow, Latah Co.., Idaho (USNM). =Catharus fuscescens subpallidus (see Wet- more et al. 1984). Based on comparisons of over 500 spec- imens of the species, I conclude that sub- pallidus is somewhat duller and more gray 441 above than other forms of the species, and further differs from salicicola by its slightly darker crown. The buff of the throat is slightly paler than in other subspecies. Ca- tharus f. subpallidus breeds from northern Washington east of the Cascades to north- eastern Oregon, northern and central Idaho, and western Montana. The winter range is not known. When naming subpallidus, Bur- leigh and Duvall (1959) restricted the type locality of salicicola to Fort Garland, Col- orado. Catharus guttatus munroi Phillips, 1962.— Nulki Lake, British Columbia (DMNH). =Catharus guttatus munroi. Phillips (1962) described munroi from northwestern Canada as differing from the ‘“‘small’’ western subspecies by the distinct hue of brown on the sides and flanks, and the more reddish and paler dorsal color. The range of munroi was said to include most of that of euborius Oberholser, 1956, from Yukon Territory. Aldrich (1968) synony- mized munroi with euborius and the A.O.U. (1957) synonymized both names with nom- inate guttatus. Ripley (1964) synonymized munroi with nanus of coastal Alaska and western British Columbia. I compared 800 specimens of C. guttatus in fresh fall plumage and agree with Phillips (1962) that munroi is distinguishable. Dor- sally, munroi is darker than nanus, paler and grayer than faxoni, the northeastern subspecies. The sides and flanks are paler than in nominate guttatus. I also compared the type of euborius to specimens of munroi. Although the type is a worn individual collected in July, I dis- agree with Phillips (1962) that it is useless for subspecific identification. The holotype of euborius differs from eastern specimens by lacking the buffy coloration typical of faxoni, and it is darker above than either munrol, faxoni, or nominate guttatus. The sides of the holotype of ewborius, although very worn, are also grayer than those of munroi. Five USNM migrants identified by Phillips in 1987 as “‘euborius’”’ are similar 442 to the holotype of euborius. The range of euborius appears to be restricted to central southern Yukon. The range of C. g. munroi is central and northern British Columbia, and, according to Phillips (1962), possibly the extreme northeastern part of British Co- lumbia and probably extreme southwestern Yukon. Catharus guttatus jewetti Phillips, 1962.— Hurricane Ridge and Elwha River, Olym- pic Mountains, Clallam Co., Washington (USNM). =Catharus guttatus jewetti (see Phillips et al. 1964 and below). The population of the Olympic Penin- sula, Washington, was described as more reddish brown and less gray than oromelus Oberholser, 1932 (type locality 15 miles NE Lakeview, N base of Crook Peak, Warner Mountains, Lake Co., Oregon) of the moun- tains from southern British Columbia through the Cascades to northern Califor- nia, and as darker than s/eveni Grinnell, 1901 (type locality Point Sur, Monterey Co., Cal- ifornia) of coastal Washington, Oregon, and California (Phillips 1962). Specimens of jJewetti were characterized as paler than nominate guttatus and verecundus (type lo- cality Queen Charlotte Islands) of south- eastern Alaska and coastal British Colum- bia. Aldrich (1968) considered the size and coloration of the Olympic Peninsula birds intermediate between s/eveni and oromelus. I found that the immature syntype of jewetti is indistinguishable from specimens from southeastern Alaska. However, on the basis of the adult syntype and other specimens, I conclude that jewetti is a recognizable sub- species as described by Phillips (1962). Judged from less than a dozen specimens, Jewetti migrates or winters to southern Cal- ifornia and Nevada, northern Arizona (Monson & Phillips 1981), and New Mex- ico. Turdus migratorius aleucus Oberholser, 1974.—South Yollo Bolly Mountain, Trinity Co., California (USNM). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON =Turdus migratorius propinquus Ridgway, 1877 (see Browning 1974, 1978). Mimidae Toxostoma lecontei macmillanorum Phil- lips, 1965.—13 km E Buttonwillow, Kern Co., California (DMNH). =Toxostoma lecontei macmillanorum. Specimens in fresh fall plumage from the isolated population in the San Joaquin Val- ley, California, were described as darker than other populations of the species. Although my series of macmillanorum was small, I found that it differs from specimens of ar- enicola as described. Dumetella carolinensis meridianus Burleigh (1959) = [1960].—Athens, Clarke Co., Georgia (USNM). =Dumetella carolinensis carolinensis (Lin- naeus, 1766) (see Monroe 1968, Phillips 1986). Birds from the southern range of the species named meridianus by Burleigh (1960c) are within the range of variation of nominate carolinensis. Bombycillidae Bombycilla cendrorum larifuga Burleigh, 1963.— Headquarters, Clearwater Co., Idaho (USNM). =Bombycilla cedrorum larifuga (see Behle 1985, and below). Burleigh (1963) divided B. cedrorum and described the population from southwest- ern and central Canada and the western United States as paler than the other pop- ulations of the species. I examined 45 specimens from the west- ern range of B. cedrorum and found that the palest specimens from the type series of /ar- ifuga are those Burleigh collected and pre- pared. In the description of /arifuga, Bur- leigh (1963) remarked that specimens from Oregon and Washington are darker brown than birds from Idaho. All but one of the specimens from Oregon and Washington VOLUME 103, NUMBER 2 were collected earlier than Burleigh’s series from Idaho. The exceptional bird, from Washington, was collected by Burleigh and resembles his Idaho series in pallor. Spec- imens collected by persons other than Bur- leigh from Idaho, Washington, Oregon, and Montana, although darker than Burleigh’s series of /arifuga, are noticeably grayer and less reddish brown above and on the upper breast, and have paler crowns than speci- mens from the eastern range of the species. These western specimens are more similar to the Idaho series prepared by Burleigh than to specimens collected elsewhere. I provi- sionally agree with Oberholser (1974) and Behle (1985) that the population named /ar- ifuga should be recognized. Bombycilla cedrorum aquilonia Burleigh, 1963.—Searston, Newfoundland (USNM). =Bombycilla cedrorum cedrorum Vieillot, 1808. Burleigh (1963) named the population from Newfoundland to northern Canada and Alaska aquilonia and described it as the grayest subspecies. I examined 23 of the specimens studied by Burleigh; except for the holotype of aquilonia, I lacked males from Canada. I conclude that there is too much individual variation to recognize an additional eastern subspecies (contra Behle 1985). Vireonidae Vireo solitarius jacksoni Oberholser, 1974.— 16 miles south of Roundup, Musselshell Co., Montana (USNM). =Vireo solitarius plumbeus Coues, 1866 (see Browning 1978). Vireo gilvus petrorus Oberholser, 1974.— Fort Steele, Carbon Co., Wyoming (USNM). =Vireo gilvus brewsteri (Ridgway, 1903). In an earlier paper (Browning 1974) I con- cluded that the population from southern British Columbia to Sonora and the Trans- Pecos region of Texas, named petrorus, be- longed to the western subspecies /eucopo- 443 lius. Allan R. Phillips (pers. comm.) sug- gested that petrorus is a northward extension of brewsteri from Mexico. Upon reexami- nation, I find that specimens of petrorus are slightly darker crowned than swainsoni, leu- copolius, and, especially, nominate gilvus. A series of 21 specimens from the Rocky Mountains are darker and duller on their upper parts than a larger series of the other northern forms, are larger than /eucopolius (see Browning 1974), and are most similar to brewsteri in size and coloration. I now consider petrorus to be a synonym of brew- steri, the subspecies breeding from Nayarit north to the Sierra Madre Occidental, southern Arizona (Phillips et al. 1964), and the Rocky Mountains to at least Montana. Vireo olivaceus caniviridis Burleigh, 1960[a]. — Moscow, Latah Co., Idaho (USNM). =Vireo olivaceus caniviridis (see Wetmore et al. 1984, Behle 1985). This pale subspecies breeds in Washing- ton, Idaho, and northern Oregon. The win- ter range is unknown. Emberizidae Parulinae Dendroica petechia hypochlora Oberholser, 1974.—3 miles N Fort Whipple (near Prescott), 3000 ft, Yavapai Co., Arizona (USNM). =Dendroica petechia sonorana Brewster, 1888 (see Browning 1974). Dendroica dominica axantha Oberholser, 1974.—Lucasville, Scioto Co., Ohio (CMNBH). =Dendroica dominica albilora Ridgway, 1873 (see Browning 1978). Oporornis formosus umbraticus Oberholser, 1974.— 1.25 miles down Ohio River from Vanport, near mouth of Four-mile Run, Beaver Co., Pennsylvania (USNM). =QOporornis formosus (Wilson, 1811) (see Browning 1978). Icteria virens danotia Oberholser, 1974.— 20 miles W Mountain Home, Kerr Co., Texas (USNM). 444 =Icteria virens virens (Linnaeus, 1758) (see Browning 1978). Thraupinae Piranga ludoviciana zephyrica Oberholser, 1974.—Santa Rita Mountains, Madera Canyon, Santa Cruz Co., Arizona (USNM). =Piranga ludoviciana (Wilson, 1811) (see Browning 1978). Piranga rubra ochracea Phillips, 1966.— Trout Creek just above its mouth, near Cane Springs, lat. 34°57’'N, long. 113°37'W, western Arizona (DMNH). =Piranga rubra ochracea. Females in first basic plumage from northwestern Arizona were described by Phillips (1966) as similar in size but darker and duller (less yellow) than the large-billed western subspecies cooperi and as paler and less green than nominate rubra. Adult males of ochracea were described as more purple and less orange than cooperi. Nine specimens, including adult females and immatures from the type series of ochracea, are brighter green above than the dull grayish-backed cooperi, and the dark olive backed nominate subspecies. Adult fe- males also differ from the other subspecies by their ochraceous rumps and heads and grayish ochraceous sides and flanks. Im- mature males are intermediate to nominate rubra and cooperi in back color. Piranga r. ochracea is known to breed in the central part of Big Sand Valley in Mojave County, Arizona, and to winter in Sinaloa and Coli- ma, Mexico. Cardinalinae Richmondena cardinalis clintoni Banks, 1963.—Ruffo’s Ranch, Cerralvo Island, Baja California, Mexico [CAS]. =Cardinalis cardinalis clintoni. Described as similar in size to ignea of southern Baja California but smaller than PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON superba of the southwestern U.S. and north- western Mexico, the males of clintoni were characterized as less intensely red with paler gray edges to the dorsal feathers than ignea. Compared to superba, the red areas of clin- toni were described as more pink and not orangish. Females of clintoni were charac- terized as grayer dorsally and less brown than ignea. I agree with Banks (1963) and also found the crown coloration of four males of clin- toni 1s more orange (less red) than the afore- mentioned forms. The population of central Baja California, seftoni, was not mentioned by Banks (1963). I compared two males of seftoni and found they are ventrally similar to 26 specimens of ignea and 27 of clintoni, and dorsally similar to clintoni in the gray edges of the back feathers, but the red feath- ers of the back and crown are more similar to superba. Paynter (1970b) recognized clin- toni only provisionally. Guiraca caerulea mesophila Oberholser, 1974.—Lipscomb, Lipscomb Co., Texas (USNM). =Guiraca caerulea caerulea (Linnaeus, 1758) (see Browning 1978). Emberizinae Hortulanus fuscus aimophilus Oberholser, 1974.—Fort Davis, Jeff Davis Co., Texas (USNM). =Pipilo fuscus texanus van Rossem, 1934 (see Browning 1978). Use of the specific name follows Zink (1988) and A.O.U. (1989). Pipilo aberti voorhiesi Phillips, 1962.—ca. 15 km S Tucson, Pima Co., Arizona [DMNH]. =Pipilo aberti dumeticola van Rossem, 1946 (see Hubbard 1972). Baird’s (1852) description of P. aberti could apply to either the western (dumetico- la) or the eastern (nominate aberti) popu- VOLUME 103, NUMBER 2 lation. Phillips (1962), believing that the ho- lotype of aberti was collected from the western population, renamed the eastern population. Evidence presented by Hub- bard (1972) and McKlevey (1955) concern- ing the itineraries of the possible collectors of the type of P. aberti does not support Phillips’ (1962) speculation (contra Paynter 1970a). Spizella pusilla perissura Oberholser, Texas birds, 1974:941.— Valentine, Cherry Co., Nebraska (USNM). Spizella pusilla vernonia Oberholser, 1974.—Japonica, Kerr Co., Texas (USNM). =Spizella pusilla arenacea Chadbourne, 1886 (see Browning 1978). Pooecetes gramineus altus Phillips, in Phil- lips, Marshall, & Monson 1964.—no type designated. Pooecetes gramineus altus Phillips, 1965.— extreme southern part of Kenrick Park, San Francisco Mountains, Arizona (DMNB). =Pooecetes gramineus altus Phillips, 1964 (see Rea 1983). This dorsally dark brown subspecies was described as less buffy than affinis and sim- ilar in size to confinis. Paynter (1970a) syn- onymized altus with confinis. It occurs from northern Arizona to southern Utah, western Colorado, and New Mexico. The authorship of a/tus has been ques- tioned by Parkes (1966) because Marshall (who was not specifically given as the author of the name) may have written the section on P. gramineus in Phillips et al. (1964); he suggested that the author of this subspecies be given as “Phillips”? =Marshall, in Phil- lips, Marshall, & Monson. Monson & Phil- lips (1981) cited Marshall as the author for the taxon, but Rea (1983) believed that Phillips is the true author of a/tus. It is spe- cifically noted in Phillips, Marshall, & Mon- son (1964:1x) that Phillips “is responsible for the scientific names and classification 445 used.” According to Phillips (pers. comm.), he submitted the manuscript describing a/- tus before 1964, but its publication (Phillips 1965) was delayed. Chondestes grammacus quillini Oberholser, 1974.—Cotulla, La Salle Co., Texas (USNM). =Chondestes grammacus strigatus Swain- son, 1827 (see Browning 1978). Amphispiza bilineata dapolia Oberholser, 1974.—Chisos Mountains, Pine Canyon, 6000 ft, Brewster Co., Texas (USNM). =Amphispiza bilineata opuntia Burleigh & Lowery, 1939 (see Browning 1978). Amphispiza bilineata belvederei Banks, 1963.—east side Cerralvo Island, Baja California, Mexico [CAS]. =Amphispiza bilineata belvederei. This form was characterized as darker dorsally than bangsi of southern Baja Cal- ifornia, and more gray (less brown) than deserticola of central Baja California and mainland Mexico. The maxilla of belvederei was described as slightly curved compared to the straighter maxilla of bangsi. Paratypes of be/vederei are noticeably dark vinaceous above, a color lacking in 20 spec- imens of bangsi and a larger series of deser- ticola. The crown of be/vederei is also grayer than in bangsi. I did not compare the shape of the maxilla. Paynter (1970a), who did not examine specimens from Cerralvo Island, provisionally recognized belvederei. Banks (1963) suggested that carmenae van Ros- sem, 1945, from Carmen Island, Baja Cal- ifornia, may also warrant recognition. A single specimen from Carmen Island ex- amined by me is paler dorsally than bangsi and belvederei. Ammodramus henslowi houstonensis Ar- nold, 1983.—south-central Houston, Harris County, Texas [TCWC]. =Ammodramus henslowi henslowi (Audu- bon, 1829). This local population was described as 446 darker than the eastern subspecies, susur- rans, and the western nominate hens/owi. All but the holotype of houstonensis were available to me for comparison. The male paratype collected in June is actually browner (not more black) dorsally than most specimens of the nominate subspecies, and two worn July males from the type series are indistinguishable in this characteristic. Nine specimens of nominate henslowi from Wisconsin, Ohio, and Missouri are blacker, and therefore, agree with the description of houstonensis in dorsal coloration. The nape of the new form was described as duller than nominate hens/owi, but this character is not useful in separating the two populations. The yellow lore was said to be absent in hous- tonensis. This character is subject to indi- vidual variation in the nominate subspe- cies, and, in fact, the lores of the June male from Houston are yellow. Rump coloration, not mentioned by Arnold (1983), is darker and more richly chestnut in susurrans than in nominate hens/owi and houstonensis. The amount of individual variation in color- ation within the western form precludes rec- ognition of additional subspecies. Passerella iliaca chilcatensis Webster, 1981.—near Tsirku River, about 250 m elevation, 7 miles (airline) SSW Klukwan, Alaska [CAS]. =Passerella iliaca chilcatensis. Swarth (1922) was the first to notice that birds from the southern part of the range of fuliginosa, as then recognized, differ from those to the north, but he declined to name the population for lack of sufficient speci- mens. Webster (1981) characterized chil- catensis as similar below but more reddish and duller above than /fuliginosa. I found that chilcatensis differs from the holotype and 30 other specimens of fuligi- nosa, including one bird from Vancouver Island, as described and that the subspecies is blacker and less reddish both dorsally and on the ventral spots than 35 specimens of townsendi, the form breeding farther to the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON north. The new subspecies breeds from the Chilkat River area of southeastern Alaska to the Stewart area of British Columbia and winters on the coasts of Oregon and Cali- fornia south to San Francisco. Melospiza melodia callima Oberholser, 1974.—West Point, Orange Co., New York (USNM). =Melospiza melodia melodia (Wilson, 1810) (see Browning 1978). Melospiza melodia melanchra Oberholser, 1974.—Bay Point, 3 miles N Sandusky, Ohio (CMNB). =Melospiza melodia euphonia Wetmore, 1936 (see Browning, 1978). Zonotrichia leucophrys aphaea Oberholser, 1974.—Caribou Mountain, Bonneville Co., Idaho (CMNH). =Zonotrichia leucophrys leucophrys (For- ster) 1772 (see Browning 1974, 1978). Junco hyemalis henshawi Phillips, 1962.— Bennett, British Columbia (USNM). =Junco hyemalis cismontanus Dwight, 1918. The name henshawi is a new name for the population that the A.O.U. (1957) called cismontanus Dwight, 1918. The name cis- montanus was originally proposed by Dwight (1918) for convenience to discuss hybrids between the oreganus and hyemalis groups. It was later applied by Miller (1941: 343, 402) as the name of the stabilized hy- brid population that breeds from south-cen- tral Yukon to central interior British Co- lumbia and west-central Alberta, but Miller proposed that it should be applied to similar appearing hybrids as well as for the true subspecies. The name cismontanus 1s valid as applied to that breeding population (Mil- ler 1941; contra Phillips 1962, Browning 1974, Rea 1983). Junco hyemalis simillima Phillips, 1962.— Pringle Falls, 4245 ft, Experiment Station (Deschutes National Forest), Deschutes Co., Oregon (DMNH). VOLUME 103, NUMBER 2 =Junco hyemalis simillimus (see Browning 1974, 1978, 1979b). Junco oreganus eumesus Oberholser, 1974.— Blue Mountains, 3500 ft, ridge on east fork of Touchet River, 21 miles SE Dayton, Columbia Co., Washington (USNM). =Junco hyemalis shufeldti Coale, 1887 (sen- su Phillips 1962, see Browning 1974, 1978). Calcarius pictus roweorum Kemsies, 1961.— Anaktuvik, Alaska (USNM). =Calcarius pictus (Swainson), 1832 (see Jehl 1968). Calcarius pictus mersi Kemsies, 1961.— Little Cape, Ontario [NMC]. =Calcarius pictus (Swainson), 1832 (see Jehl 1968). Kemsies’ (1961) descriptions of the two forms of C. pictus were based on compari- sons of inadequate samples and character- izations of geographic variation using sea- sonally incomparable specimens (Jeh! 1968). Icterinae Agelaius phoeniceus stereus Oberholser, 1974.—Barr, Adams Co., Colorado (USNM). =A gelaius phoeniceus stereus (see Browning 1978). I recognize stereus provisionally. Agelaius phoeniceus zastereus Oberholser, 1974.—Boise, 2700 ft, Ada Co., Idaho (CMNH). =A gelaius phoeniceus zastereus (see Brown- ing 1974, 1978). I recognize zastereus provisionally. Agelaius phoeniceus heterus Oberholser, 1974.— Fort Wingate, McKinley Co., New Mexico (USNM). =Agelaius phoeniceus fortis Ridgway, 1901 (see Browning 1978). Rea (1983) stated that heterus may prove to be recognizable. 447 Quiscalus major alabamensis Stevenson, 1978.—4.5 km E Mobile, Baldwin Co., Alabama (USNM). =Quiscalus major alabamensis. Stevenson (1978) recognized four sub- species of Quiscalus major: torreyi from New Jersey to northeastern Florida, westoni on the Florida Peninsula, alabamensis along the coast of Alabama and southeastern Missis- sippi, and nominate major from south- western Mississippi to Texas. The subspe- cies westoni Sprunt, 1934 (type locality St. Johns River marshes, Indian River Co., Florida), was included in nominate major by the A.O.U. (1957) as was the range at- tributed to alabamensis. The eye is dark in westoni and nominate major, but it is pale in torreyi and alaba- mensis (Stevenson 1978). I compared three males and four females of alabamensis and about 15 specimens each of the other sub- species. I found that males of westoni are slightly larger than torreyi and alabamensis, with measurements of the wing chord av- eraging less than in nominate major. The subspecies alabamensis differs in size from the other taxa as implied. Females of a/a- bamensis are similar to torreyi in color, and paler than westoni and nominate major. Icterus bullockii eleutherus Oberholser, 1974.—Del Rio, Val Verde Co., Texas (USNM). =Icterus galbula bullockii (Swainson, 1827) (see Browning 1978). Carduelinae Leucosticte tephrocotis irvingi Feinstein, 1958.—Anaktuvuk Pass, Brooks Range, Alaska (USNM). =lTeucosticte arctoa tephrocotis (Swainson, 1832). Feinstein (1958) divided the subspecies tephrocotis and named the population of Alaska’s Brooks Range, confining fephro- cotis to the northern Rocky Mountains. He 448 described irvingi as similar in size, but more brightly colored, above and below. The red- dish regions of irvingi were described as av- eraging deeper red and more purplish than in tephrocotis. Specimens of irvingi were de- scribed as showing “a slight inclination to- wards Jittoralis in coloration” (Feinstein 1958:12), but with less gray in the malar and auricular region than in typical exam- ples of that subspecies. The type series of irvingi was reported by Feinstein (1958) to consist of 7 males and 5 females. These, and five other specimens from the Brooks Range are well within the range of individual variation in size and col- or found in 40 specimens of tephrocotis from the Rocky Mountains and bear no resem- blance to /ittoralis. The characters of irvingi are attributable to individual variation, and I fully agree with R. E. Johnson (in litt.) that irvingi does not warrant recognition. Erythrina mexicana anconophila Oberhol- ser, 1974.--Chinati Mountains, Presidio Co., Texas (USNM). =Carpodacus mexicanus frontalis (Say, 1823) (see Browning 1978). Loxia curvirostra vividior Phillips, in Mon- son and Phillips 1981.—in “El Paso Co.,”’ Colorado [=about 8-15 km from Mon- ument] (sic) (USNM). =Loxia curvirostra vividior. This subspecies was described as overall similar in size to neogaea Griscom, 1937, but often having a thinner bill. Phillips (in Monson & Phillips 1986) also described vi- vidior as more ochraceous than neogaea and gave its range as “usually the high moun- tains of the western United States from Montana to Colorado and probably west to Deschutes Co., Oregon. .. .”’ For the use of the name neogaea see Dickerman (1986b, 1987). I provisionally recognize vividior. Loxia curvirostra reai Phillips, in Monson & Phillips 1981.—Dismal Lake, SE Shoshone Co., Idaho [AMR]. =Loxia curvirostra reai (but see Payne 1987). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON This subspecies was described as having a bill often heavier than the otherwise sim- ilarly sized minor. Males were described as less reddish, and females as rich yellow on the rump and more deeply ochraceous be- low than minor. The range of reai was given as the mountains of northern Idaho (typical series) to southern British Columbia and Sheridan, Wyoming, and casually in west- ern Oregon and Arizona, Minnesota, Mich- igan, and Kansas. Payne (1987) considered reai indistin- guishable from minor, but he (1987:28) based his conclusion mainly on a compar- ison of the holotypes of pusilla and minor. Payne designated AMR 3627 as the lecto- type of reai but this specimen is not from the original series (Rea, pers. comm.). For the correct use of minor see Dickerman (1986b, 1987). Although Dickerman listed both vividior and reai in his summary, he did not comment on the validity of either name. My acceptance of reai and vividior is provisional, recognizing that the taxonomy of L. curvirostra still requires a comprehen- sive review. Passeridae Passer domesticus plecticus Oberholser, 1974.—Gray, Bonneville Co., Idaho (USNM). =Passer domesticus domesticus Linnaeus, 1758 (see Browning 1978). Acknowledgments I thank Allan R. Phillips for the many useful discussions about the taxonomy of North American birds and for the oppor- tunity to examine the long series of speci- mens available to him during his visits to the U.S. National Museum. I thank the Cal- ifornia Academy of Sciences, Delaware Mu- seum of Natural History, Field Museum of Natural History, Texas A&M University, and San Diego Museum of Natural History for loaning specimens, and extend appre- VOLUME 103, NUMBER 2 ciation to the many museums that loaned specimens to Dr. Phillips. Appreciation is extended to David Niles for his encourage- ment and assistance in locating some of the types described by Phillips. I am grateful to Robert W. Dickerman, Allan R. Phillips and Amadeo M. Rea for their critical reading of an earlier draft of the manuscript. 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Variation of Ladder-backed Woodpeckers in southwestern North Ameri- ca.—Proceedings of the Biological Society of Washington 81:1-10. 1982. Woodpeckers of the world. Delaware Museum of Natural History Monograph Series No. 4, 676 pp. Snyder, L. L. 1961. On an unnamed population of the Great Horned Owl.—Royal Ontario Mu- seum Contribution No. 54, 7 pp. Stevenson, H. M. 1973. An undescribed insular race of the Carolina Wren.— Auk 90(1):35-38. 451 . 1978. The populations of Boat-tailed Grack- les in the southeastern United States.—Pro- ceedings of the Biological Society of Washington 91(1):27-51. Sutton, G. M. 1967. Oklahoma birds: Their ecology and distribution, with comments on the avifau- na of the southern Great Plains. University of Oklahoma Press, Norman, 674 pp. Swarth, H. S. 1922. Birds and mammals of the Sit- kine River region of northern British Columbia and southeastern Alaska.— University of Cali- fornia Publications in Zoology 24(2):125-314. Timm, D. E., M. L. Wege, & D. S. Gilmer. 1982. Current status and management challenges for Tule White-fronted Geese.— Transactions of the North American Wildlife and Natural Resource Conference 47:453-463. Walkinshaw, L.H. 1965. A new Sandhill Crane from central Canada.—Canadian Field-Naturalist 79(3):18 1-184. . 1973. Cranes of the world. Winchester Press, New York, 370 pp. Webster, J.D. 1981. A new subspecies of Fox Spar- row from Alaska.— Proceedings of the Biologi- cal Society of Washington 96(4):664-668. Wetmore, A. 1965a. Additions to the list of birds of the Republic of Colombia.—L’Oiseau 35(special): 156-162. 1965b. The birds of the Republic of Pana- ma.—Smithsonian Miscellaneous Collections 150, part 1, 483 pp. , R. Pasquier, & S. L. Olson. 1984. The birds of the Republic of Panama.— Smithsonian Mis- cellaneous Collections 150, part 4, 670 pp. White, C. M. 1968. Diagnosis and relationships of the North American tundra-inhabiting Pere- grine Falcons.— Auk 85(2):179-191. , & D. A. Boyce, Jr. 1988. An overview of Peregrine Falcon subspecies. Pp. 789-810 in T. J. Cade, J. H. Enderson, C. G. Thelander, & C. M. White, eds., Peregrine Falcon populations, their management and recovery. The Peregrine Fund, Inc., Boise, Idaho. Zink, R. M. 1988. Evolution of Brown Towhees: Al- lozymes, morphometrics and species limits. — Condor 90(1):72-82. U.S. Fish and Wildlife Service, National Ecology Research Center, National Mu- seum of Natural History, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(2), 1990, p. 452 ERRONEOUS EMENDATIONS TO NAMES PROPOSED BY HEKSTRA (STRIGIDAE: OTUS) M. Ralph Browning Abstract. —Four emendations (Browning 1989) of names of New World owls proposed by Hekstra (1982) are incorrect subsequent spellings. The spelling “yruguaii’ of the proposed subspecific name of Otus choliba (sensu Hekstra) is chosen as correct. I earlier (Browning 1989) misinterpreted Articles 31c and 32 d (11) and applied Ap- pendix D, IV, of the International Code of Zoological Nomenclature (ICZN 1985) to four subspecific names of New World owls proposed by Hekstra (1982). According to Article 32, the four emendations (Browning 1989) are incorrect subsequent spellings and have no nomenclatural standing. The correct names as proposed by Hek- stra (1982) and the incorrect emendations (in parentheses) are: Otus atricapillus morelius (not morelien- SIS) Otus atricapillus inambarii (not inam- bariensis) Otus guatemalae peteni (not petenensis) Otus choliba urugaii [sic] (not uruguaien- SIS) Hekstra (1982) first used Otus choliba uruguail [sic] as a nomen nudum in a list of species (1982:54). He spelled the sub- specific name “uwrugaii’’ in the formal proposal, but again as uruguaii in a dis- cussion of the etymology, from the Ar- royo Urugua-i. The repeated use of uru- guail indicates his intended spelling, and as first revisor (ICZN 1985, Art. 24[b] and [c]) I select uruguaii as the correct spell- ing. Konig & Straneck (1989:3) listed this taxon as uruguaii and did not refer to other spellings. Acknowledgments I thank S. L. Olson, J. M. Savage, and G. C. Steyskal for pointing out the erro- neous emendations and for reading the manuscript. I also thank R. C. Banks, who read a later version and offered helpful suggestions. Literature Cited Browning, M. R. 1989. The type specimens of Hekstra’s owls.— Proceedings of the Biolog- ical Society of Washington 102:515-519. Hekstra, G. P. 1982. Descriptions of twenty-four new subspecies of American owls (Aves: Strigidae).— Bulletin Zoologisch Museum Universiteit Amsterdam 9(7):49-63. ICZN. 1985. International code of zoological no- menclature. 3rd ed. International Trust for Zoological Nomenclature, British Museum (Natural History), London, 338 pp. Konig, C., & R. Straneck. 1989. Eine neue Eule (Aves: Strigidae) aus Nordargentinien. — Stuttgarter Beitrage zur Naturkunde, Series A (Biologie) Nr. 428, 205:1-20. Biological Survey, National Ecology Research Center, U.S. Fish and Wildlife Service, National Museum of Natural History, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 453-463 SUBSPECIES OF STENELLA LONGIROSTRIS (MAMMALIA: CETACEA: DELPHINIDAE) William F. Perrin Abstract. —Three subspecies of Stenella longirostris are named, described and compared: S. /. /ongirostris from the world’s tropical oceans, S. /. orientalis from tropical oceanic and coastal waters of the eastern Pacific, and S. /. cen- troamericana from Pacific coastal waters of Central America. In a review of variation in Stenella spp. in eastern Pacific and Hawaiian waters (Perrin 1975a) I provisionally delineated four forms of the spinner dolphin: 1. “Costa Rican Spinner Porpoise, Ste- nella longirostris subsp. A (unnamed?),”’ 2. “‘Eastern Pacific Spinner Porpoise, Stenella longirostris subsp. B (unnamed?),”’ 3. “Whitebelly Spinner Porpoise, Ste- nella longirostris subsp. C (unnamed?),” and 4. *‘Hawaiian Spinner Porpoise, Stenella longirostris subsp. D (unnamed?).” I did not assign trinomial names to the forms because of a lack of data for spinner dolphins in other parts of the world and uncertainty about the affinities of the ho- lotype specimen of the species. (The holo- type is a skull from an unknown locality.) Sufficient data have now accumulated to al- low formal description and naming of three subspecies. Two of the subspecies corre- spond to Nos. | and 2 above, and the third includes No. 4. No. 3, the “whitebelly spin- ner,” has proved to be a highly variable hybrid/intergrade between Nos. 2 and 4. Skull measurements summarized below were taken as described in Perrin (1975a). Museum acronyms. —ANSP, Philadel- phia Academy of Natural Sciences, Pitts- burgh, Pennsylvania; BMNH, British Mu- seum (Natural History), London; CAS, California Academy of Sciences, San Fran- cisco; DKC, personal collection of D. K. Caldwell, Marineland of Florida, St. Au- gustine; EDM, collection of Edward D. Mitchell; LACM, Los Angeles County Mu- seum of Natural History; LSUMZ, Louisi- ana State University Museum of Zoology, Baton Rouge; MCZ, Museum of Compar- ative Zoology, Harvard University, Cam- bridge, Massachusetts; MMBL, National Marine Mammal Laboratory, Seattle, Washington; MNHN, Museum National d’Histoire Naturelle, Paris; MVZ, Museum of Vertebrate Zoology, University of Cali- fornia, Berkeley; NSM, National Science Museum, Tokyo; RMNH, Riksmuseum van Natuurlijke Historie, Leiden; SDMNH, San Diego Museum of Natural History, San Diego, California; SWFC, Marine Mammal Synoptic Collection, Southwest Fisheries Center, La Jolla, California; TCWC, Texas Cooperative Wildlife Collection, Texas A&M University, College Station; UCMP, Museum of Paleontology, University of California, Berkeley; UF, Florida State Mu- seum, University of Florida, Gainesville; USNM, National Museum of Natural His- tory, Washington, D.C.; ZMA, Zoologisch Museum, University of Amsterdam. Stenella longirostris longirostris (Gray, 1828) Holotype. —RMNH 8676, skull only, un- known locality. Referred specimens. —Hawaiian Islands: CAS 10529, 16455, 16456, 16457, 16458; LACM 27093, 27095, 54049, 54050, 54056, 454 54057, 54060, 72296; SWFC WFP605, WFP606, WFP623, WFP669, WFP670, WEFP671; MCZ 51700; MMBL 1194; NSM 24615, 24815; USNM 339649, 504140, 504470. Christmas Island (Central Pacific): ANSP 19194, 19195. Washington Island: USNM 504251. Rangiroa (Tuamoto Is- lands): USNM 504252. Hiva Oa (Marquesa Islands): USNM 504253. Enewetok (Mar- shall Islands): USNM 395404. Western Pa- cific between | and 9°S, 147 and 160°E: NSM M24928, M24929, M24930, M24931, M24933, M24934, M25373. Western Pa- cific between 3 and 5°N, 142 and 150°E: NSM M25376, M24932. Japan: NSM MO05100, M24800. Sri Lanka: BMNH 1891.10.13.2, 1948.4.20.1, 1949.10.27.1 and 2. Maldive Islands: BMNH 1959.7.9.1, 1959.7.9.3, 1959.7.9.4, 1959.7.9.5 and 1959.7.9.6. Ivory Coast: USNM 470557. Liberia: RMNH 21.720. Senegal: ZMA 12.263, 13.146, 13.147, IFAN2; EDM 784. North Carolina: USNM 291352, 504525. South Carolina: USNM 500859. Florida: DKC R-4-SLS; UF 7861, 18719; LSUMZ 17017; Texas: TCWC 28286, 29035; USNM 504224, 504233, 504433-504456. St. Vin- cent (Lesser Antilles): UF 18720-18724. Venezuela: ZMA 15.138. Diagnosis. —A subspecies of Stenella lon- girostris characterized by distinct tripartite color pattern (Fig. 1), erect (in adult male) to falcate dorsal fin, relatively small post- anal hump, relatively large adult size (to about 215 cm; average about 180-190 cm) and relatively long (394-464 mm) and broad (zygomatic width 150-170 mm) adult skull. Distribution.—Tropical and subtropical waters of the world. Replaced in the eastern Pacific by S. /ongirostris orientalis and S. longirostris centroamericana (described be- low) and possibly in Southeast Asia/north- ern Australasia by an undescribed dwarf subspecies (Perrin et al. i989) with lower average tooth and vertebral counts and in the northwestern Indian Ocean/Red Sea by another locally adapted form (Robineau & PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rose 1983). Most common near coasts and islands (Gilpatrick et al. 1987). Description of holotype. —Measurements (in mm) of the holotype skull: condylobasal length 426, length of rostrum 277, width of rostrum at base 76, width of rostrum at '2 length 48, width of rostrum at *%4 length 36, greatest preorbital width 142, greatest post- orbital width 159, zygomatic width 156, pa- rietal width 125, length of left temporal fos- sa 50, height of left temporal fossa 43, length of left upper toothrow 241, length of left ramus 368, height of left ramus 57. Teeth: upper left 54, upper right 53, lower left 52, lower right 54. External size, shape and col- oration unknown. Variation.—The color pattern is stable except in a broad zone of hybridization/ intergradation with S. /ongirostris orientalis in the offshore eastern tropical Pacific (range of the “whitebelly spinner’’—Perrin et al. 1985), where coloration and shape may be intermediate between those of the two forms (Fig. 1; Perrin 1972, Perrin, Sloan & Hen- derson 1979). In these specimens, the cape may be less distinct and the ventral margin of the dorsal overlay (terminology of Perrin 1972) stepped, ragged or grading into ven- tral spots. Adult size may vary geographically (Per- rin etal. 1989). The largest individual known was from Hawaii (CAS 16456, 213 cm). An adult male hybrid/intergrade from the east- ern Pacific (SWFC BGBO08) was 235 cm long, but this may have been a case of en- docrinally induced gigantism; the next larg- est in the sample of 262 specimens from the population (including several from the same school) was over 30 cm shorter (Perrin & Gilpatrick 1990). The largest body of ex- ternal measurements from a single region is from the tropical Atlantic (Perrin et al. 1981): 17 adult females ranged from 177 to 204 cm (average 188.9 cm) and 17 males from 173 to 208 cm (average 192.5 cm); four physically mature males were 192-208 VOLUME 103, NUMBER 2 455 Fig. 1. Typical appearance of adult males of Stenella longirostris longirostris (top), S. 1. orientalis (bottom) and a hybrid/intergrade (center). S. /. centroamericana is not illustrated, because the color pattern is not well known. cm long and two females 196 and 201 cm _calities will be necessary to demonstrate this long. adequately. The same is true of tooth counts Skull size may also vary geographically (Table 1). Skulls from the hybrid/intergra- (Table 1), but larger series from more lo- dation zone discussed above may be inter- 456 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Means, standard deviations (for sample sizes =30), sample sizes (in parentheses), and maximum and minimum values for selected skull measurements and tooth counts for geographical series of cranially adult specimens of Stenella longirostris longirostris (from Perrin et al. 1989). Specimens of dwarf form from Southeast Asia excluded. Indian Ocean Western Pacific Central Pacific Atlantic Condylobasal length 409.2 420.1 436.9 427.0 + 13.10 (7) 394-430 (7) 411-431 (24) 417-464 (41) 395-458 Length of rostrum 264.9 272.1 282.6 276.8 + 11.00 (7) 250-281 (7) 262-281 (24) 263-304 (41) 251-304 Width of rostrum: At base 74.0 78.3 79.3 76.6 + 3.41 (7) 71-76 (7) 73-84 (24) 74-86 (42) 68-83 At 4 length 43.3 47.9 47.0 44.4 + 2.30 (7) 42-45 (7) 44-54 (24) 42-56 (41) 41-50 At % length — SID 31.4 32.3 31.9 + 2.63 (7) 28-34 (5) 26-36 (24) 27-37 (39) 25-39 Width of PMXs at '2 length 19.4 21.0 21.1 21.0 + 2.32 (7) 18-20 (6) 19-23 (24) 17-23 (37) 17-32 Preorbital width 141.6 144.6 150.8 145.6 + 3.86 (7) 135-146 (5) 140-150 (24) 140-158 (42) 137-153 Postorbital width 155.4 160.8 165.2 161.1 + 4.03 (7) 153-160 (6) 155-169 (25) 158-172 (41) 152-169 Zygomatic width 154.0 156.2 163.5 159.2 + 3.92 (7) 151-160 (5) 152-161 (25) 154-171 (40) 150-167 Parietal width 128.7 127.6 131.4 130.5 + 4.25 (7) 122-133 (5) 125-131 (25) 122-140 (41) 121-140 Length of temporal fossa 50.0 47.4 53.3 48.1 + 3.30 (7) 46-58 (5) 44-51 (25) 45-61 (42) 42-56 Height of temporal fossa 41.4 41.0 42.9 38.3 + 3.52 (7) 39-45 (5) 36-44 (25) 37-50 (42) 30-49 Length of upper toothrow 232.3 237.2 245.6 243.5 + 10.30 (7) 224-242 (6) 219-246 (24) 224-263 (41) 221-265 Length of ramus 352.0 366.3 372.4 368.4 + 11.68 (7) 366-370 (7) 360-371 (23) 352-399 (40) 343-399 Height of ramus 55.9 55.8 58.4 56.4 + 2.28 (7) 55-57 (6) 55-57 (23) 53-64 (41) 51-61 Upper teeth (per row) 59) 53.3 55.0 55.4 + 3.01 (15) 49-59 (12) 48-61 (29) 50-62 (41) 48-64 Lower teeth (per row) 50.6 51.4 52.5 53.9 + 3.32 (8) 42-49 (15) 45-58 (29) 48-57 (43) 47-62 mediate in size and form between those in this subspecies and in S. /. orientalis (Perrin 1975a, Perrin, Sloan & Henderson 1979). Remarks. —This subspecies includes the ‘““Hawaliian Spinner Porpoise, Stenella lon- girostris subsp. D (unnamed?)”’ (Perrin 1975a). Hybrids/intergrades with S. /. ori- entalis (described below) correspond to the ““Whitebelly Spinner Porpoise, Stenella lon- girostris subsp. C (unnamed)”’ (Perrin 1975a). Genetic as well as morphological evi- VOLUME 103, NUMBER 2 dence support the hypothesis that the highly variable “‘northern whitebelly spinner” and “southern whitebelly spinner” (Perrin, Sloan & Henderson 1979, Perrin et al. 1985) rep- resent a zone of hybridization or intergra- dation between the two distinct forms to the west and the east; “‘whitebelly spinners” possessed no unique mitochondrial DNA haplotypes when compared with animals to the east (Dizon & Perrin 1987). Hewitt (1988) noted that the width of many hybrid zones is about one hundred times the estimate of dispersal for the in- dividual organisms. The zone of hybridiza- tion/intergradation for S. /. Jongirostris and S. I. orientalis (described below) is about 2000 km wide (Perrin et al. 1985). The for- mer tends to inhabit coastal waters and waters around islands (Gilpatrick et al. 1987, Perrin et al. 1983), but tagged oceanic spin- ner dolphins in the eastern Pacific have moved at least 400 km (Perrin, Evans & Holts 1979). However, as pointed out by Hewitt, gene dispersal rate is likely to vary greatly with population density, structure and patchiness, and more information is needed on these before the significance of the relatively great dispersal distance of in- dividuals relative to the width of the hy- bridization/intergradation zone can be weighed. The holotype specimens of two junior synonyms of S. /ongirostris are probably of this form; both are relatively long and broad skulls from unknown localities. These are the holotypes of Delphinus alope Gray, 1846 (BMNH 847a; condylobasal length 412 mm, zygomatic width 166 mm) and D. steno- rhynchus Gray, 1866 (BMNH 1471a; con- dylobasal length ca. 447 mm, zygomatic width ca. 167 mm). A third holotype spec- imen, that of D. microps, also from an un- known locality (BMNH 349a), is at the low- er end of the known range of adult skull length (394 mm) and below (at 147 mm) the range of skull width (150 mm) for this 457 subspecies, suggesting that it belongs to a different form. It seems likely that the very broadly dis- tributed populations of this form will even- tually prove to include several additional locally adapted races sufficiently distinct to merit recognition as subspecies. This seems to almost certainly be the case in Southeast Asia (Perrin et al. 1989) and is possibly the case in the northwestern Indian Ocean (Ro- bineau & Rose 1983). The problem is sim- ply one of material; cetacean specimens are very costly and difficult to collect, prepare and store. In the case of S. /ongirostris the species 1s known from several very large regions from only a single or a very few specimens; these areas include the Atlantic coast of South America, both coasts of Af- rica and the South Pacific. If, as seems very likely, there proves to be a distinct (dwarf) form of the species in Southeast Asia, the name [Delphinus] ro- seiventris Wagner, 1846 may apply; the probable holotype specimen! MNHN 1882- 104 is a small skull (condylobasal length 384 mm, zygomatic width 153 mm) from the Moluccas. Use of the common name “Gray’s spin- ner dolphin” is proposed for this, the nom- inate subspecies of S. /ongirostris. Specimens examined. —In addition to the specimens designated as referred specimens above, the specimens listed in Perrin (1972, 1975a), Perrin, Sloan & Henderson (1979), and Perrin et al. (1981, 1989). Stenella longirostris orientalis, new subspecies Holotype. —SDNHM 21427, physically mature male, total length 186 cm, from 12°20'N, 92°53'W (ca. 120 km off the coast ‘The holotype specimens of Delphinidae at the MNNH are currently under review by D. Robineau of the Centre National d’Etude des Mammiféres Marins. 458 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON San Diego | 30° ee Hawaiian |e. ° 20° ns Acapulco : ee ocoesers a Ea Pe op ee er aa ee a3 Te Se 33¢7+-a74- 199 303 , 559 | 1064 ia an 10°| ; : ees ee es BERS 8 ; ———— ae SERGE T ga {sos 4 389 | 362 | 355 0° 10° 20° 30° | : it 160° 150° 140° 130° 120° 110° 100° go° go° 70° 20° 15° HONDURAS EL : i SALVADOR i NICARAGUA § centroamericana io: Be l. orientalis 5° . COLOMBIA Y ECUADOR 105° 100° 95° 90° 95° g0° 75° Fig. 2. Distribution of subspecies of Stenella longirostris in the eastern Pacific. Modified from Perrin et al. (1985). A. Schools sighted from tuna seiners and research vessels, identified to subspecies on the basis of predominant appearance of dolphins in school. Numbers indicate total number of schools sighted in square. Open portion of square = proportion of total sightings in the square that were of schools of the eastern spinner, S. 1. orientalis or (on Central American coast, tisted in parentheses) S. /. centroamericana. Shaded portion = S. 1. longirostris (in west, southwest, and far south) and intergrades/hybrids between S. /. /ongirostris and S. 1. orientalis. B. Approximate distributions of S. /. orientalis and S. 1. centroamericana in Central American waters, based on records with voucher specimens. VOLUME 103, NUMBER 2 of Guatemala in the eastern tropical Paci- fic), 10 Mar 1968, killed incidentally in tuna purse-seine fishery, complete skeleton, cast and external photographs. External photo- graph in Perrin (1972:989, fig. 9) and Leath- erwood et al. (1988:151, fig. 191). Referred specimens. -USNM 395396, 395531, 39533-34 from 12°51'N, 93°18'W; USNM 395260, 395270-73, 395526, 395593, 7°11'N, 90°32'W; USNM 395274- 75, SWFC 0025 (CV245), SWFC 0027 (CV240), UF18726, 7°20'N, 92°30'W; LACM 54052 (WAWSO), ca. 21°16'N, 106°16’'W; LACM 54036 (WAWS54), ca. 18°N, 102°12’W; LACM 54039 (WAW60), 14°30'N, 99°35'W; LACM 64062 (RLB40S5), ca. 14°20'N, 107°20’W; USNM 324974-75, 12°32'N, 91°4'W; USNM 395930, 16°15'N, 98°55'W; CAS 15665, 13°30’ to 14°6'N, 98° to 190°56'W; CAS 15666, 13°30’ to 14°2’W, 99°47' to 102°2’W; MVZ 140641, 13°33’ to 14°2’N, 99° to 108°W; CAS 15667-69, MVZ 140642 and 140645, UCMP 86287, 9°50’ to 10°36’N, 99°47’ to 102°2’'W; USNM 395026, ca. 3°N, 87°W; USNM 396169, T°21'N, 87°14’W; USNM 23302, Pacific Ocean between Panama and Galapagos Is- lands; USNM 88976, ca. 21°35'N, 106°40’'W; LACM 54038 (WAWS8), ca. 17°N, 100°45'W; SDMNH 21200, ca. 18°20'N, 106°40'W; CAS 13821, 13828, ca. 12°22’N, 80°50’W; MMBL 1967-102, ca. 21°43'N, 10°47'W. Diagnosis. —A subspecies of Stenella lon- girostris characterized by uniform gray col- oration dorsally and laterally (Fig. 1), with patches of white in axillary and genital areas, erect to forward-canted dorsal fin in adult male, relatively large post-anal hump, rela- tively small adult size (to 199 cm in adult males and 193 cm in adult females), and relatively short (351-407 mm) and narrow (zygomatic width 139-153 mm) skull. Distribution. —Tropical coastal and oceanic waters of the eastern Pacific (Fig. 2) in a large triangular region, with the base of the triangle extending from about the tip of Baja California (ca. 24°N) in the north to ca. 10°S off the coast of Peru and the tip of 459 the triangle at about 145°W, 10°N. Most common off Mexico and Central America, in a region extending about 1000 km off- shore (Perrin et al. 1985). Replaced south of the Gulf of Tehuantepec and north of the Bay of Panama in inshore coastal waters of extreme southern Mexico, Guatemala, El Salvador, Honduras, Nicaragua and Costa Rica by Stenella longirostris centroameri- cana (described below). Description of holotype.—The holotype specimen, a physically mature 186-cm male (all vertebral epiphyses fused to the centra) was dark gray dorsally and laterally, with the gular region slightly lighter gray and with patches of white in the axillary and genital regions (photograph in Perrin 1972:989, fig. 9). The dorsal fin was large and canted for- ward, and the post-anal hump was promi- nent; these features are preserved in the cast of the specimen in the SDMNH collection. Measurements (in mm) of the holotype skull: condylobasal length 401, length of rostrum 259, width of rostrum at base 74, width of rostrum at '2 length 46, width of rostrum at ¥4 length 33, greatest preorbital width 133, greatest postorbital width 152, zygomatic width 150, parietal width 129, length of left temporal fossa 54, height of left temporal fossa 39, length of left upper toothrow 226, length of left ramus 346, height of left ramus 54. Teeth: upper left 49, upper right 48, lower left 52, lower right 52. Variation. —The white axillary and gen- ital patches may be confluent, with dark spots or patches between them. In a broad zone of hybridization/intergradation with S. 1. longirostris in the offshore eastern trop- ical Pacific (the range of the “‘whitebelly spinner’ —Perrin et al. 1985), coloration may be intermediate between those of the two forms (Fig. 1; Perrin 1972, 1975a; Per- rin, Sloan & Henderson 1979). Range in total length in 431 adult males (right testis =100 g) was 160-199 cm (ayv- erage 176.1 cm, standard deviation 6.12 cm); 697 adult females (at least one corpus in the Ovaries) were 152-193 cm long (average 171.3 cm, standard deviation 5.99 cm) (Per- 460 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Means, standard deviations (for sample sizes =30), sample sizes (in parentheses), and maxi- mum and minimum values for selected skull mea- surements and tooth counts for Stenella longirostris orientalis and S. |. centroamericana. From Perrin et al. (1989). S. 1 centro- S. 1. orientalis americana Condylobasal length 386.5 428.6 (26) 351-407 (5) 416-439 Length of rostrum 245.5 279.0 (26) 218-262 (5) 268-288 Width of rostrum At base V2 72.4 (29) 66-77 (5) 70-77 At 4 length 41.6 42.0 (26) 37-47 (5) 39-43 At ¥% length 30.5 29.8 (26) 26-36 (5) 28-32 Width of PMXs at 19.6 21.4 % length (26) 16-22 (5) 21-22 Preorbital width 132.3 134.2 (29) 127-138 (5) 131-138 Postorbital width 148.1 151.2 (29) 140-154 (5) 149-152 Zygomatic width 146.4 150.4 (29) 139-153 (5) 149-152 Parietal width 125.9 128.8 (29) 119-132 (5) 127-130 Length of temporal 48.5 49.6 fossa (29) 41-57 (5) 46-57 Height of temporal 36.6 36.4 fossa (29) 29-47 (5) 34-42 Length of upper 2A2 2 245.0 toothrow (25) 192-299 (5) 238-255 Length of ramus 329.1 369.6 (27) 301-348 (5) 358-379 Height of ramus 52.3 55.2 (29) 47-57 (5) 53-57 Upper teeth 52.6 + 3.18 55.3 (per row) (32) 46-61 (5) 50-59 Lower teeth 50.5 + 2.39 56.3 (per row) (37) 45-56 (5) 51-59 rin et al. 1985). Ranges and averages for skull measurements and tooth counts are given in Table 2. Additional data on ontogenetic, sexual, individual and geographic variation in size, shape, coloration and skeleton are given in Perrin (1972, 1975a, b), Perrin, Sloan & Henderson (1979), Perrin et al. (1985), Schnell et al. (1982, 1985), Douglas et al. (1986), and Leatherwood et al. (1988). Remarks.—This subspecies corresponds to the “Eastern Spinner Porpoise, Stenella longirostris subsp. B (unnamed?)”’ (Perrin 1975a). Hybrids/intergrades with S. /. lon- girostris correspond to the “Whitebelly Spinner Porpoise, Stenella longirostris subsp. C (unnamed?)”’ (Perrin 1975a). Two alternative hypotheses have been proposed for the origin of this distinctive dolphin in the eastern Pacific: differentia- tion during isolation ofa portion of the trop- ical eastern Pacific by a temperate-water barrier extending across the equator during a glacial period, or parapatric differentiation due to the peculiar oceanographic structure of the eastern Pacific (very shallow mixed layer and thick oxygen minimum layer that create an unique oceanic dolphin habitat— Au & Perryman 1985) and facilitated by complex social structure (Perrin et al. 1985). The holotype specimen of the junior syn- onym Delphinus microps Gray, 1846 (BMNH 349a) is a small adult skull from an unknown locality; its measurements are within the range for S. /. orientalis. It may have come from the eastern Pacific, or it may have come from a different region where spinner dolphins are relatively small, e.g., Southeast Asia (Perrin et al. 1989). If future analyses were to determine that it came from the eastern Pacific, then the subspecies would bear the name microps rather than orien- talis. Continued use of the common name ‘eastern spinner dolphin” is proposed for this subspecies. Specimens examined. —In addition to the specimens designated as referred specimens above, the specimens listed in Perrin (1972, 1975a), Perrin, Sloan & Henderson (1979), and Perrin et al. (1981, 1989). VOLUME 103, NUMBER 2 Stenella longirostris centroamericana, new subspecies Holotype. —USNM 395933, skull and ex- ternal measurements, adult male, one of three specimens collected at 9°47'N, 85°42'W (Gulf of Nicoya, Pacific coast of Costa Rica) on 26 Mar 1963 by D. W. Wal- ler; original field number BCFBL,SD XII-9. Paratypes. —-USNM 395931 and 395932 (same collection data as holotype). Referred specimens. -USNM 395934 from 9°58'N, 86°3’W (outer side of Penin- sula de Nicoya, Pacific coast of Costa Rica); SWFC 0025 (DAB100), 14°19’N, 93°10’W (ca. 80 km off coast of southern Mexico near border with Guatemala, at southern end of Gulf of Tehuantepec over continental shelf). Diagnosis. —A subspecies of Stenella lon- girostris characterized by uniform gray col- oration dorsally and laterally (ventral col- oration poorly known, but photographs of animals in the wild and sketches by field observers do not show the well-defined ax- illary and genital white patches typical of S. l. orientalis), erect to forward-canted dorsal fin in adult male, relatively large post-anal hump, relatively large adult size (to 216 cm), and relatively long (416-439 mm) and nar- row (zygomatic width 149-152 mm) skull. Distribution. —Coastal waters to ca. 80 km offshore over the continental shelf south of the Gulf of Tehuantepec and north of the Bay of Panama off extreme southern Mex- ico, Guatemala, El Salvador, Honduras, Nicaragua and Costa Rica (Fig. 2). Replaced to the north, south and west by S. /. ori- entalis. Description of holotype. —Measurements (in mm) of the holotype skull: condylobasal length 416, length of rostrum 269, width of rostrum at base 71, width of rostrum at 2 length 43, width of rostrum at % length 30, greatest preorbital width 135, greatest post- orbital width 152, zygomatic width 152, pa- rietal width 129, length of left temporal fos- 461 sa 48, height of left temporal fossa 36, length of upper left toothrow 239, length of left ramus 358, height of left ramus 56. Teeth: upper left 58, upper right 57 (estimated), lower left 57, lower right 57. External mea- surements (taken in field by D. W. Waller, following Norris 1961, in cm) of the carcass: total length 216.4, beak tip to melon 14.3, to angle of gape 28.3, to center of eye 32.0, to blowhole 32.1, to origin of flipper 46.3, to tip of dorsal fin 100.0, to umbilicus 99.7, axillary girth 69.2, posterior length of flip- per 16.8, span of flukes 39.9. Variation. —Three adult males (testis weight = 100 g) were 212-216 cm long and 12 adult females (at least one corpus in ova- ries) 193-211 cm (average 198.8 cm) (Perrin et al. 1985). Ranges and averages for skull measurements and tooth counts are given in Table 2. Additional data on osteological variation are given in Douglas et al. (1986), Perrin (1975a, b), Perrin et al. (1985), and Schnell et al. (1982, 1985). Remarks. —This subspecies corresponds to the “Costa Rican spinner porpoise, Ste- nella longirostris subsp. A (unnamed?)” (Perrin 1975a). It differs from S. /. orientalis primarily in its relatively greater body size and its relatively long and narrow skull. The few skulls available for this form can be easily discriminated from skulls of the other forms, based on simple scatterplots of length and width measurements or on scat- terplots of scores for discriminant functions (Perrin 1975a), but it can be expected that overlap will be encountered when larger se- ries of specimens become available. The paucity of specimens is due to an apparent ecological difference between this form and S. /. orientalis. Large numbers of the latter are killed by tuna purse seiners because they are captured to catch yellowfin tuna (Thunnus albacares) that accompany them (Perrin 1975b); the former do not often “carry fish’? and thus are only very rarely captured and killed. The common name “Costa Rican spinner 462 dolphin” has been used for this form since 1975, but its range is now known to include the coastal waters of several nations. There- fore it is proposed that this subspecies be called the ‘““Central American spinner dol- phin.”’ Specimens examined. —In addition to the specimens designated above as paratypes, the specimens listed in Perrin (1972, 1975a), Perrin, Sloan & Henderson (1979), and Per- rin et al. (1981, 1989). Acknowledgments I wish to thank R. L. Brownell, Jr., J. G. Mead, C. W. Potter, D. Robineau, G. D. Schnell and I. Barrett for reading the manu- script and suggesting useful changes. Cu- rators who provided access to specimens in their charge are listed in Perrin (1975a), Per- rin, Sloan & Henderson (1979), and Perrin etal. (1981, 1989); I also thank C. W. Potter for assistance with specimens at the USNM. Literature Cited Au, D. W. K., & W. L. Perryman. 1985. Dolphin habitats in the eastern tropical Pacific. — Fishery Bulletin 83:623-643. Dizon, A. E., & W. F. Perrin. 1987. Genetic distances among and within species, populations, and schools of spinner and spotted dolphins. — Ab- stracts of the Seventh Biennial Conference on the Biology of Marine Mammals, December 5- 9, 1987, Miami, Florida, p. 17. Douglas, M. E., G. D. Schnell, & D. J. Hough. 1986. Variation in spinner dolphins (Stenella longi- rostris) from the eastern tropical Pacific Ocean: Sexual dimorphism in cranial morphology.— Journal of Mammalogy 67:537-544. Gilpatrick, J. W., Jr., W. F. Perrin, S. Leatherwood, & L. Shiroma. 1987. Summary of distribution records of the spinner dolphin, Stenella longi- rostris, and the pantropical spotted dolphin, S. attenuata, from the western Pacific Ocean, In- dian Ocean and Red Sea.— U.S. Department of Commerce, NOAA Technical Memorandum, NOAA-TM-NMES-SWEFC-89, 42 pp. Hewitt, G. M. 1988. Hybrid zones—natural labora- tories for evolutionary studies. — Trends in Ecol- ogy and Evolution 3:158-167. Leatherwood, S., R. R. Reeves, W. F. Perrin, & W. E. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Evans. 1988. Whales, dolphins, and porpoises of the eastern North Pacific and adjacent Arctic waters. A guide to their identification. Dover Press, New York, 245 pp. Norris, K. S. (ed.). 1961. Standardized methods for measuring and recording data on the smaller cetaceans. — Journal of Mammalogy 42:47 1-476. Perrin, W. F. 1972. Color patterns of spinner por- poises (Stenella cf. S. longirostris) of the eastern Pacific and Hawaii, with comments on delphi- nid pigmentation.—Fishery Bulletin 70:983- 1003. . 1975a. Variation of spotted and spinner por- poise (genus Stenel/a) in the eastern Pacific and Hawaii.— Bulletin of the Scripps Institution of Oceanography 21:206 pp. 1975b. Distribution and differentiation of populations of dolphins of the genus Stenella in the eastern tropical Pacific.—Journal of the Fisheries Research Board of Canada 32:1059- 1067. —., & J. W. Gilpatrick, Jr. 1990. Spinner dolphin Stenella longirostris (Gray, 1828). In S. H. Ridgway & R. J. Harrison, eds., Handbook of marine mammals, volume 5. Academic Press, New York (in press). —, W. E. Evans, & D. B. Holts. 1979. Move- ments of pelagic dolphins (Stenella spp.) in the eastern tropical Pacific as indicated by results of tagging, with summary of tagging operations, 1969-76.—U.S. Department of Commerce, NOAA Technical Report, NMFS SSRF-737, 14 pp. ——., N. Miyazaki, & T. Kasuya. 1989. A dwarf form of the spinner dolphin (Stenella longiros- tris) from Thailand.--Marine Mammal Science 5:213-227. ,P.A. Sloan, & J.R. Henderson. 1979b. Taxo- nomic status of the ‘southwestern stocks’ of spinner dolphin Stenella longirostris and spot- ted dolphin S. attenuata. —Report of the Inter- national Whaling Commission 29:175-184. ——., M.D. Scott, G. J. Walker, & V. L. Cass. 1985. Review of geographical stocks of tropical dol- phins (Stenella spp. and Delphinus delphis) in the eastern Pacific.—U.S. Department of Com- merce, NOA Technical Report, NMFS SSRF- 28, 28 pp. , E. D. Mitchell, J. G. Mead, D. K. Caldwell, & P. J. H. van Bree. 1981. Stenella clymene, a rediscovered tropical dolphin of the Atlantic. — Journal of Mammalogy 62:583-598. —., M. D. Scott, G. J. Walker, F. M. Ralston, & D. W. K. Au. 1983. Distribution of four dol- phins (Stenella spp. and Delphinus delphis) in the eastern tropical Pacific, with an annotated catalog of data sources.—U.S. Department of VOLUME 103, NUMBER 2 Commerce, NOAA Technical Memorandum, NOAA-TM-NMEFS-SWFC-38, 65 pp. Robineau, D., & J.-M. Rose. 1983. Note sur le Ste- nella longirostris (Cetacea, Delphinidae) du golfe d’Aden.— Mammalia 47:237-245. Schnell, G. D., M. E. Douglas, & D. J. Hough. 1982. Geographic variation in morphology of spotted 463 , & 1985. Further studies of morphological differentiation in spotted and spinner dolphins (Stenella attentuata and S. lon- girostris) from the eastern tropical Pacific. Southwest Fisheries Center Administrative Re- port LJ-85-04C, 30 pp., Southwest Fisheries Center, P.O. Box 271, La Jolla, California 92038. and spinner dolphins (Stenella attenuata and S. longirostris) from the eastern tropical Pacific. : : : ; : Southwest Fisheries Center Administrative Re- National Marine Fisheries Service, port LJ-82-15C, 213 pp., Southwest Fisheries Southwest Fisheries Center, 8604 La Jolla Center, P.O. Box 271, La Jolla, California 92038. Shores Drive, La Jolla, California 92038. ~ PROC. BIOL. SOC. WASH. 103(2), 1990, pp. 464-498 STUDIES IN THE LEPIDAPLOA COMPLEX (VERNONIEAE: ASTERACEAE) VII. THE GENUS LEPIDAPLOA Harold Robinson Abstract.—The Neotropical Vernonian genus Lepidaploa is recharacterized, nomenclaturally clarified, and its 116 species are listed. Lepidaploa has seriate- cymose inflorescences and lophate pollen surfaces of a type found in the related genera of the Lepidaploa Complex. Lepidaploa is distinct in the Complex by the sessile heads, the heads with 1.3-3.0 times as many involucral bracts as flowers, nodular style bases, sometimes glanduliferous achenes, and lack of glands on the anther appendages. Variations in exine patterns in Lepidaploa are compared with the three other genera of the Complex with rhizomatous crests, Echinocoryne, Stenocephalum and Chrysolaena, and the two genera with non-rhizomatous crests, Lessingianthus and Mattfeldanthus. New combina- tions are provided for 114 species, and two species, L. beckii and L. solomonii are described as new. The present paper concludes a series of papers aimed at the resolution of the Neo- tropical Lepidaploa group in the tribe Ver- nonieae. Previous papers in the series have dealt with the five closely related genera of the informally defined Lepidaploa Com- plex, Stenocephalum, Echinocoryne, Les- singianthus, Chrysolaena and Aynia (Rob- inson 1987a, b, 1988a, b, c) that differ from Vernonia s. str. by having consistently lo- phate pollen, and with two less closely re- lated genera, Cyrtocymura and Eirmoceph- ala (Robinson 1987c), that usually have true scorpioid cymes. The remaining genus Lep- idaploa of the Lepidaploa Complex having lophate pollen is treated in the present pa- per. Lepidaploa was named originally by Cas- sini (1817b) as a subgenus of Vernonia with brief comments on distinctions from the other two subgenera accepted at that time. Six species were mentioned in association with the subgenus, Vernonia glauca, V. fas- ciculata, V. arborescens, V. divaricata, V. scorpioides and V. albicaulis, in that order. The genus was treated more fully five years after the original description (Cassini 1823), but was still referred to as a subgenus. Ge- neric status of Lepidaploa is sometimes er- roneously dated from the 1823 treatment (Gleason 1906:174) because of the combi- nations under Lepidaploa included in that treatment. Lepidaploa was not given generic status until it was listed in a series of genera by Cassini (1825) in his treatment of the genus Oliganthes. The latter listing men- tioned no included species. The proper characterization of Lepida- ploa dates from the treatment of the sub- genus by Cassini in 1823. The subgenus was given the alternative French name Lépi- daple, and was described as having invo- lucral bracts “‘régulierement imbriquees, appliquées, subcoriaces, lancéolées, acu- minées et presque spinescentes au sommet; les intérieures étrécies de bas en haut, ter- minées en pointe, nullement élargies, ar- rondies, ni colorées au sommet.” Cassini (1823) clearly distinguished his subgenus from typical Vernonia by the lack of a sub- ulate appendage and lack of coloration on the bracts of the involucre. The lack of bract VOLUME 103, NUMBER 2 appendages also distinguished Lepidaploa from the paleotropical subgenus Ascaricida (=Baccharoides Moench) based on Vernon- ia anthelmintica L. (Cassini 1817a, b). In 1823 Cassini included seven species in the subgenus Lepidaploa, L. scorpioides (from Vernonia scorpioides), L. phyllostachya (same as the earlier V. arborescens), L. aris- tata, L. albicaulis (from V. albicaulis), L. lanceolata (=?V. longifolia), L. canescens (from V. canescens) and L. buxifolia in that order. The most notable difference between the 1823 and 1817 listings was the elimi- nation of the two species of Vernonia from eastern North America, V. glauca and V. fasciculata. The latter belong to the Ver- nonia s. str. close to the type, V. novebo- racensis. Cassini further cited additional species of Vernonia mentioned by Kunth (Humboldt et al. 1818) that were “‘presque indubitables” members of the subgenus Lepidaploa, V. gracilis, V. tournefortioides (=V. scorpioides), V. geminata, V. mollis, V. pellita, V. micrantha and V. frangulae- folia. None of the Cassini names or com- binations in Lepidaploa were valid as of the 1823 publication since Lepidaploa was not at that time recognized at the generic level. A precise typification of the subgenus Lepidaploa was not attempted until almost 100 years after the original description. Gleason (1906:165) rather incidentally stat- ed that Lepidaploa was based on V. scor- pioides. To the extent that this can be re- garded as a serious lectotypification, this appears to have been an arbitrary selection of the first species listed by Cassini in 1823. The incidental and arbitrary nature of the selection allows the rejection of the unrepre- sentative species as lectotype. Robinson et al. (1980:428), in a review of subdivisions of the Vernonieae, listed Lepidaploa with the lectotype Vernonia albicaulis, which is far more representative of the species given by Cassini in 1823. The value of the latter lectotypification has been confirmed in the present series of studies, since Lepidaploa can now represent a broader group follow- 465 ing its traditionally broader interpretation by such authors as Baker (1873). Vernonia scorpioides, which has traditionally repre- sented a narrower concept as a section or series Scorpioideae, has now been treated as the type species of a smaller well-defined segregate genus Cytocymura (Robinson 1987c). Shortly after the lectotypification by Rob- inson et al. (1980), Jones (1980) listed Lep- idaploa in the synonymy of Vernonia with Vernonia glauca (L.) Willd. cited as the type. At present, no earlier history for this typi- fication is known. The choice was evidently again an arbitrary selection of the first species listed by Cassini in 1817(b). Such an arbi- trary lectotypification would not have to be rejected, but V. glauca is an undesirable se- lection for Lepidaploa in view of Cassini’s subsequent failure to include the species in the subgenus (1823). The choice is further undesirable because V. glauca is an im- mediate relative of the type species of Ver- nonia, V. noveboracensis, and it usually has shortly appendaged involucral bracts of the type that Cassini (1823) pointedly indicated were lacking in his subgenus Lepidaploa. One must conclude that Cassini’s (1817b) original inclusion of V. glauca in Lepidaploa was based on an inadequate understanding of that species. The specimen no. 8428 on the Jussieu herbarium (sheet no. 8428, IDC microedition 6206, fiche 620) that was seen by Cassini seems to have essentially no ap- pendages on the involucral bracts. The lectotypifications by Gleason (1906) and Jones (1980) share one particularly un- wholesome aspect associated with their ar- bitrary nature. Both attempts were treating Lepidaploa only as a synonym of Vernonia with no concern for its possible importance as a taxon in its own right. The arbitrary nature of the selections allows for rejection, and the poor choices involved make rejec- tion desirable, even if both had priority. A major part of the confusion regarding the concept of Lepidaploa arises from Cas- sini’s reliance upon the involucral bracts as 466 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a character. Some typical Vernoniae such as V. fasciculata lack appendages on the bracts, while many species now recognized as Lep- idaploa possess narrowly pointed involucral bracts. Appropriately, Cassini seemed to place little confidence in his 1817 subgenus. The changed listing of species by Cassini in 1823 created a more closely related group with only one species, of those identifiable with modern concepts, lacking seriate-cymes and only two of those species proving, on later examination, to have non-lophate pol- len. Unfortunately, Cassini did not accom- pany his improved grouping with an im- proved description. Nevertheless, Cassin1’s confidence in the group had risen sufficient- ly by 1825 to recognize Lepidaploa at the generic level. During the period from 1825 to 1841 a number of authors recognized Lepidaploa at the generic level: Cassini (1825), Reichenbach (1828), Bartling (1830), Cassini (1830), and Spach (1841). DeCandolle (1836) and later major au- thors did not treat Lepidaploa at the generic level. DeCandolle (1836) retained Lepida- ploa as his section VII of Vernonia con- taining 195 species including some from Asia and Africa. The type species of Ver- nonia was placed in the section IX. Ascar- icida (including Baccharioides). Bentham (Bentham & Hooker 1873) broadened the concept of Lepidap/loa within Vernonia to include a series Paniculatae which included the type species of Verno- nia, but he restricted the subgenus Lepi- daploa geographically to, “Species Ameri- canae ultra 200.’ The Lepidaploa concept of Bentham and his followers such as Baker (1873), Gleason (1922), and Jones (1979b) came to include all American Vernonia with unappendaged involucral bracts and a dou- ble pappus. As such, the concept included the type of Vernonia and fell into the syn- onymy of the subgenus and section Ver- nonia. The resurrection of Lepidaploa in the present paper is nearly in the sense of Cas- sini in 1823, though it traces nomenclatur- ally to the confused concept of 1817. As defined herein, the genus Lepidaploa in- cludes 116 species distributed throughout the Neotropical Region, many species oc- curring in the West Indies, the Andes, and Brazil, and a smaller number occurring in Central America. The genus includes most of the Neotropical species formerly placed in Vernonia that have heads in well-devel- oped seriate-cymes as described below. The genus does not include some species with markedly scorpioid seriate-cymes such as Cyrtocymura scorpioides and Eirmocephala brachiata (Robinson 1987c) or many seri- ate-cymose members of Lessingianthus (Robinson 1988a) and Mattfeldanthus (Robinson & King 1979). The distinguish- ing features such as the sessile heads, re- duction in size of inflorescence bracts, styles with well-developed basal nodes, fusion of veins in the corolla lobes, glands on the achenes, ‘tack of glands on the anther ap- pendages, and details of the rhizomatous crests on the pollen are discussed more fully under separate headings. Inflorescence Form The inflorescences of the Asteraceae are generally cymose; the great majority of the species, and most Vernonieae, including those in typical Vernonia, have a distinctly cymose structure. The inflorescence forms in the tribe have been discussed by many authors (Gleason 1923, Cabrera 1944, Jones 1979b). Some of the inflorescence branches become aligned in a series and may achieve a scorpioid form. The branches are usually marked by heads borne singly or in groups at a series of nodes. In the extreme, each branch appears superficially spicate or ra- cemose in a form that has been referred to by most as a scorpioid cyme. In this series of studies the use of the latter term is re- stricted to inflorescences such as those of Cyrtocymura and Eirmocephala which are actually scorpioid, and the more common non-scorpioid forms in the Vernonieae are VOLUME 103, NUMBER 2 referred to with the more broadly applicable term seriate-cyme (Figs. 61, 62; Robinson 1987c, 1988a). The extreme forms of seri- ate-cymose inflorescences in the Vernonie- ae with heads appearing lateral and sessile are nearly restricted to the few related groups of almost exclusively Neotropical distri- bution that are treated in the present series of papers on the Lepidaploa Complex. In these inflorescences each branch ap- pears spicate but is actually a series of lateral proliferations. As noted by Gleason (1923) and Cabrera (1944), each head is develop- mentally terminal and only appears lateral as a result of its displacement by a lateral branch produced at its base. The resulting structure in the Neotropical members of the tribe varies in appearance from a spike to an obvious scorpioid cyme. The most ob- viously scorpioid forms occur in the more remotely related Neotropical genera with type A pollen, Cyrtocymura, Eirmocephala (Robinson 1987c), and Dipterocypsela, and in one African species Vernonia peculiaris Verdcourt, but the seriate cymes with heads appearing lateral and sessile are otherwise almost completely restricted to the imme- diate Lepidaploa Complex of genera having lophate pollen. Within the Complex, the seriate-cymes with apparently sessile heads occur in Lep- idaploa itself, as well as in Chrysolaena Robinson (1988b), some of Lessingianthus Robinson (1988a), Mattfeldanthus Robin- son & King (1979), and Stenocephalum Sch.Bip. (Robinson 1988a). Variations of the inflorescence with the heads raised on short or long peduncles occur in Aynia Rob- inson (1988c), Echinocoryne Robinson (1987b), and many other species of Lessin- gianthus. One of the most distinctive fea- tures of Lepidaploa appears to be the con- sistency of the sessile heads. The only distinctly pedunculate heads in the genus are those that are terminal on the branches and a few rare non-terminal pedunculate heads in such species as V. cleocalderonae and V. macahensis, and specimens such as 467 Wilbur 13260 from Panama illustrated by Keeley (1982). The terminal peduncles real- ly differ from the other heads only in the lack of a subsequent lateral branch, but the presence of a peduncle on a lower head, which is exceedingly rare in Lepidaploa, can be interpreted developmentally as a down- ward displacement of the succeeding lateral branch. The presence of apparent peduncles is comparatively common throughout most of the diversity of the related genus Lessin- gianthus. Thus, while seriate-cymes occur in both Lepidaploa and Lessingianthus, the ““*pedunculate’”” forms are essentially re- stricted to the latter genus in the pair. The ““non-pedunculate’’ seriate-cyme that is characteristic of Lepidaploa and Mattfel- danthus is closer to the condition in the ses- sile-headed but less seriate inflorescences in many other Vernonieae, and it occurs to some extent in all but one genus of the Lep- idaploa Complex where it appears to be primitive. The “peduncles” in Lessingian- thus and Echinocoryne are rather individ- ually distinctive in the tribe and are con- sidered as derived separately in each of those genera. As noted by various authors (Baker 1873, Gleason 1923), there is variation in the de- gree of development of the bracts that are located at the series of nodes in the seriate- cyme. The genera Cyrtocymura, Diptero- cypsela, and Eirmocephala have minute bracts that are scarcely visible under the heads. Members of the more restricted Lep- idaploa Complex usually have more ob- vious foliose bracts subtending the heads at each node of the series, but the bracts may vary in size from slightly larger than minute to essentially the size of the vegetative leaves. The bracts of Mattfeldanthus and Steno- cephalum are essentially like vegetative leaves. The bracts of Chrysolaena are very small. The bracts in Lepidaploa and Les- singianthus show contrasting trends in their relative size. Species of Lessingianthus with sessile heads subtended by foliose bracts in a seriate-cyme have the bracts similar in 468 size to the vegetative leaves that are below the inflorescence. There is little or no abrupt change in size of foliose elements at the base of the inflorescence. In Lepidaploa, the bracts at the base of the inflorescence are almost always at least slightly discontinuous in size from the vegetative leaves immedi- ately below them (Figs. 61, 62), and the in- florescence often has very reduced bracts. In only a few species of Lepidaploa, such as the Brazilian V. obtusifolia, is there no ev- ident reduction in the size of the foliar struc- tures starting at the base of the inflores- cence. Involucral Bracts The involucre of Lepidaploa is significant for both the ratio of bracts to flowers in the heads and for the differentiation of the bracts. There has been no systematic study of the ratio of involucral bracts to flowers in the heads of the Vernonieae. Nevertheless, sig- nificant differences in the ratios have be- come evident from counts made during the description of various species and genera. Differences in the ratio are sometimes sim- ply a reflection of the overall number of flowers. In Stenocephalum (Robinson 1987a), the higher percentage of bracts is primarily related to the reduced number of flowers, whereas in Lepidonia (Robinson & Funk 1987), the nearly equal number of bracts and flowers is related to the compar- atively large number of flowers in the heads. In the case of Echinocoryne and the type species of Mattfeldanthus, the ratio is al- tered by the extremely high number of in- volucral bracts in the heads. In Chrysolaena the imbalance of the ratio derives from the unusually small number of bracts in the head. Nevertheless, there are also differ- ences in the ratio in some groups that lack extremes in numbers of bracts or flowers. In the type species of Vernonia and its im- mediate relatives, the number of bracts and flowers seems nearly equal, but in Lepida- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ploa and Lessingianthus the bracts are usu- ally one and a half to three times as nu- merous as the flowers. Many species of Lepidaploa have strong differentiation between the outer and inner involucral bracts. The outer bracts are more pointed and are often aristate. The inner bracts are usually blunt with at most a short mucro. A common pattern in the genus is a lower cluster of spreading, slender bracts contrasted with the more erect and ap- pressed, less pointed, inner bracts. Such a pattern occurs in a few species of Lessin- gianthus such as L. niederleinii, but the larg- er heads usually distinguish the members of the latter genus with such involucres from Lepidaploa. Veins of Corolla Lobes During various studies of the family I have noted that the veins in the corolla lobes of the Vernonieae are more completely fused at the tip than those of many other tribes of the Asteraceae, and in the present study most members of the Lepidaploa Complex have been seen to have veins that fuse at the apex more broadly than those in most other species of the Vernonieae. In Aynia and Mattfeldanthus the fusion is extreme and the lobe tip is filled with a large scleri- fied shield (Robinson 1988c). Species of Chrysolaena have the narrowest veins dis- tally and the weakest fusion of any members of the Complex. Lepidaploa and Lessin- gianthus both show a moderate thickening of lobe veins and a further moderate en- largement in the apical fusion that falls far short of the extreme form. Still, the latter is greater than that in most genera of the Ver- nonieae outside of the Complex such as Ver- nonia s str. Basal Stylar Node In Lepidaploa the style base characteris- tically has a large zone of spreading, thick- walled cells that result in a very large disc in dried plants. It is this structure that is VOLUME 103, NUMBER 2 here referred to as a basal stylar node. The sclerified disc is sharply angled inward under the base of the style and it leaves a comparatively narrow central area of at- tachment. Styles of Lessinginathus, Steno- cephalum, Echinocoryne, and Chrysolaena show little or no zone of thick-walled cells at the base and they have no large basal disc. The disc is not sharply angled under the base of the style and the area of attachment is not or scarcely narrower than the style shaft. There is no functional basis known for the differences in style bases of the members ofthe Lepidaploa Complex, and the changes in style base form must be considered rather rare and essentially random. Nevertheless, style base differences correlate closely with the generic limits in the Lepidaploa Com- plex. This pattern shows the comparative continuity within each genus compared to the broad phyletic gaps between the genera. Glands on Achenes and Anther Appendages Short-stalked, capitate glands that pro- duce sesquiterpene lactones are common in the Vernonieae as glandular punctations on the surface of various structures in different genera. Lepidaploa often has glands on the achene surface, while Lessingianthus never has such glands. In most Lepidaploa, the glands occur among the long setulae that cover the achenes, but a few species such as V. macahensis from the Rio de Janeiro re- gion, have glanduliferous achenes without setulae. Glands occur on the anther appendages of various Vernonieae, including three ele- ments among the genera treated in the pres- ent series of studies: one of the three species of Eirmocephala, E. cainarachiensis (Hi- eron.) H. Robinson (Robinson 1987c), one specimen of Lepidaploa that was identified as possibly V. helophila, and six of the seven species of the genus Chrysolaena. The re- maining related genera, Aynia, Echinoco- ryne, Lepidaploa, Lessingianthus, Mattfel- 469 danthus, and Stenocephalum, seem to lack glands except in the one case mentioned above. The various species mentioned in which the glands occur on the appendages, including the species of Ezrmocephala, are all in groups that also have glands on the achenes. Pollen The Lepidaploa Complex is characterized by pollen with crests in a lophate pattern, and it includes most of the Neotropical species with lophate pollen that are tradi- tionally placed in the genus Vernonia (Figs. 1-54). A lophate pattern occurs also in the phyletically distinct Old World element of the Vernonieae (Figs. 55-58) and ultimately in the related tribe Lactuceae. It has been concluded in this series of studies that the complex and widely occurring lophate form is more primitive than the non-lophate, spi- nose grains called the Lychnophora type by Stix (1960) and Type A by Jones (1979b). Cases of reversion to Type A pollen from lophate forms have been cited in the Old World Distephanus Cassini (Robinson & Kahn 1986) and in the Neotropical Lessin- gianthus (Robinson 1988a); other such re- versions I presume have occurred many times. Accordingly, the pollen of the Lepi- daploa Complex is considered to have re- tained the more primitive pattern in the tribe. Within the Lepidaploa Complex there are further distinctive details of the pollen that can be used in the classification. The vari- ation in structure of the basal columellae under the crests partially correlates with ge- neric concepts, but a SEM is necessary to observe the character. In Lepidaploa the columellae show a type that is referred to here as rhizomate or rhizomiform (Figs. 1- 51), where the columellar material is weakly attached to the foot layer and is often nearly completely diverted at the base into a hor- izontal structure (Figs. 6, 15, 16, 25, 30, 33). The rhizomatous crests of this Lepidaploa 470 type are known only from the Neotropical Lepidaploa Complex in the genera Lepi- daploa, Chrysolaena, Echinocoryne, and Stenocephalum (Robinson 1987a, b, 1988b) and from one species of Eirmocephala (Robinson 1987c). The contrasting form, with separate columellae strongly attached to the foot-layer (Figs. 52-56), occurs in di- verse elements of the tribe including the Neotropical Lessingianthus (Figs. 52, 53), Aynia, and Mattfeldanthus (Fig. 54) of the Lepidaploa Complex, and the African gen- era such as Baccharoides Moench. (Figs. 55, 56). Horizontal structures under the crests in paleotropical Vernonieae such as Cyan- thillium Blume (Figs. 57, 58) are totally dif- ferent, being bridges between strongly at- tached points at the intersections of the crests. Because of the comparatively re- stricted nature of the rhizomate form, it is regarded as a derived condition in this study. Within the restricted Lepidaploa Com- plex, a few complications in the pattern of the basal columellar structure are seen, but these are more apparent than real. Firstly, one species of Mattfeldanthus has distinct horizontal structures under its pollen crests, but these are intermittent bridges like those of Cyanthillium. Secondly, some species of Lepidaploa have the rhizomes absent or poorly developed, but these differ from the typical Lessingianthus type by the tendency for the columellae under the crests to be narrow and less strongly attached to the foot- layer (Figs. 3, 8, 12, 18, 31, 37-39, 42, 51). The less rhizomate forms in Lepidaploa are commonest within the genus in species with Type D pollen. A taxonomically useful characteristic of the pollen that can be seen with a light mi- croscope is the pattern of the areoles on the surface of the grain. The pollen types that are of significance in the Lepidaploa Com- plex are considered in the following order: Type B, Type C, Aynia Type, Type G, and Type D. Of the pollen Types, only B and D seem to characterize related groups, and none characterize a whole genus. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Type B pollen. —The pollen areolation that has been called the Vernonia argyrophylla- type by Stix (1960) and Type B by Jones (1979b) is distinguished by colpi that extend from pole to pole with no separate polar areoles and by three equatorial areoles across the intercolpar area in a 1:2:1:2:1 or rarely 1:2:3:2:1 pattern. Type B pollen occurs pri- marily in Lessingianthus subg. Lessingian- thus (Fig. 52), but grains in some species of Lepidaploa also have a third equatorial role in some of their intercolpar regions (Figs. 23, 26, 45, 50). None of the latter has the third areole characteristically in all the in- tercolpi, and none has strong basal colu- mellae like those of Lessingianthus. The in- tercolpar region in Lepidaploa usually differs by having a 1:2:2:1 pattern (Figs. 5, 11, 14, 29, 36, 46) in the C, D, G, and Aynia Types. Type C pollen.—Many species of Lepi- daploa, including the type species, show pollen grains that have a polar areole. The areole is positioned at the apices of the three colpi. Such grains were called the Vernonia cognata-type by Stix (1960) and Type C by Jones (1979b). Type C pollen with rhizo- matous crests occurs in Chrysolaena (Rob- inson 1988b) and Stenocephalum (Robin- son 1987a) of the Lepidaploa Complex, and a Type C pollen with non-rhizomatous crests is found in the small subgenus Oligocepha- lus of Lessingianthus (Robinson 1988a). There is a significant variation within the Type C pollens that occur in Lepidaploa. The Andean and West Indian elements of the genus are sometimes erratic in the expression of the polar areole. One pole may have the areole while the other end does not. This is often a consistent feature of grains in a species, and the difference prob- ably reflects the relative outward or inward facing positions of the poles in the original pollen tetrads. It would seem a slight loss for neither pole to. have an areole, and. the Central American V. tortuosa, with tech- nically Type G pollen, might have Type C ancestry. Some of the species showing vari- able poles also have the crests particularly VOLUME 103, NUMBER 2 thick on each side of the pores (Figs. 2, 4, 10), a condition reminiscent of Type A grains. The species of Lepidaploa with Type C grains that are found in Brazil have char- acteristically narrower crests (Fig. 17), and they consistently have areoles at both poles. A number of species with Type C pollen, Vernonia cleocalderonae (Figs. 17, 18) from central Brazil, and V. cotoneaster (Figs. 12, 18) of a group found in eastern Brazil and northern South America, have basal colu- mellae that are partially non-rhizomatous. Such columellae have been noted elsewhere in Lepidaploa only in the members with Type D pollen areolation that is discussed below. In contrast, the West Indian V. seri- cea (Fig. 6), the widely distributed V. salz- mannii (Figs. 15, 16), and to a lesser extent, V. canescens (Fig. 8) of the northern Andes have crests with rhizomiform baculae that peel away from the footlayer comparatively easily. Aynia Type pollen. — Vernonia tovarensis (Figs. 19-24) seems to have a pollen form that is unique in Lepidaploa in having the intercolpar areoles nearest the poles actually reaching the poles (Fig. 19). The colpi ter- minate about half way between the pores and the poles. The pattern of areoles is known from only the one species in Lepi- daploa, but is the same areolation found in Aynia and the comparatively unrelated Central American genus Harleya Blake. As in Aynia, the crosswalls above and below the pores show a sutural line in the middle, but none of the grains of either Aynia or V. tovarensis show separation at the median suture of the crosswalls, and it is not pos- sible to say at this time whether they only meet or actually fuse. In spite of the similar areolation, the exine differs from that of Ay- nia by the modification of the basal colu- mellae into a rhizomiform structure of the type found in most of Lepidaploa. Type G pollen.—The majority of species of Lepidaploa, especially in Brazil, have Type G pollen that does not have areoles centered at the poles of the grains and does 471 not have crosswalls in the colpi (Figs. 25-— 33). In these characters the pollen is like Type B of Jones (1979b) or Vernonia ar- gyrophylla-type of Stix (1960), and no dis- tinction was made in those earlier studies. More recently the Jones and Stix terms have been applied more narrowly to pollen of the type found in Lessingianthus (Robinson 1988a), with three equatorial areoles across the intercolpi and nonrhizomatous colu- mellae. The Type G pollen in Lepidaploa usually has intercolpar areoles in a 1:2:2:1 pattern correlated with obviously rhizomi- form baculae as seen under the SEM (Figs. 27, 30, 31, 33). The seemingly slight differ- ence has proven one of the more phyleti- cally significant ones in the Lepidaploa Complex. The closest approach to the Type B pollen areolation in a species of Lepida- plea is in V. psilostachya DC. where the colpi reach the poles and there is a variable number of 2 or 3 areoles across the inter- colpus in | or 2 tiers (Figs. 25-27). Exam- ination shows that even in this species very few of the grains have 3 areoles across all the intercolpi simultaneously. A third in- tercolpar areole is even less common in oth- er species with Type G pollen. Type G pollen was earlier distinguished by the author as the Vernonia geminata- type (Robinson 1980). Unfortunately, the use of the name geminata-type now appears inappropriate as a result of the present study. The Colombian name Vernonia geminata H.B.K. was misapplied to a Brazilian species with Type G pollen. The Colombian species is a synonym of V. canescens having Type C pollen. The Brazilian species with Type G pollen should have been identified as V. subsquarrosa. The present paper adopts the term Type G for pollen that has been pre- viously called the geminata-type. The letter is the next that would follow in the series used by Jones for his two studies (1979b, 1981). Type G pollen is closest to Type C pollen in its structure, and the species involved seem to be interrelated. Type G pollen seems 472 to occur in Brazilian species that are rea- sonably close in relationship to some others of that area with Type C pollen. Some species of the Andes of Peru and Bolivia such as V. retrosetosa and V. tristis seem to be possible relatives of Brazilian species. However, the Type G pollen in the central American V. tortuosa and the Colombian V. sclareaefolia and V. trilectorum seems separately derived from more closely related Andean members of Lepidaploa having Type C pollen. A brief survey of some andean species shows those with Type G pollen lack resiniferous idio- blasts on the achene surface, whereas those with Type C pollen often have such idio- blasts. Type G grains are found outside of Lep- idaploa in the related genus Echinocoryne, and non-rhizomatous grains with similar areolation are seen in the less closely related Mattfeldanthus. Type D pollen. —A final pollen type with- in Lepidaploa is that referred to by Stix (1960) as the Vernonia arenaria-type and by Jones (1979b) as Type D (Figs. 34-51). The intercolpus has a 1:2:2:1 or sometimes a 1:2:1:2:1 areolation pattern and there are no areoles centered on the poles, but the colpus is not continuous. The grains are dis- tinctive by the presence of crosswalls above and below the pore which divide each col- pus into three sections (Figs. 35, 40, 41, 44, 49). These are not the partial intrusions into the colpus that are seen in some grains of other pollen types; rather, these crosswalls are complete, unbroken crests. Type D pol- len is almost restricted to Lepidaploa, but grains with similar crosswalls occur in two species of Lessingianthus. Pollen of the two Lessingianthus species (Robinson 1988a) differs from that of Lepidaploa by the con- sistency of the 1:2:1:2:1 intercolpar pattern and the strong unconstricted basal colu- mellae. The weak columellae of the Type D pollen in Lepidaploa are stronger than those of most members of their genus, but they are not mistakeable for the type found in Lessingianthus. The two species of Lessin- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gianthus have long-pedunculate heads and they are not particularly like Lepidaploa in any feature except the pollen. The group of species in Lepidaploa with Type D pollen is one of the most natural subdivisions in the genus. The distinctive areolation and the trend toward stronger basal columellae in the pollen is correlated in many of the species with a distinctive pubescence of T-shaped hairs on their stems and leaves. The appearance of the pubes- cence varies from densely sericeous to to- tally appressed, and it is not matched in other members of Lepidaploa. The essentially non-rhizomatous colu- mellae of the Type D pollen in Lepidaploa is interpreted here as a reversion from the rhizomatous condition. There is an irregu- larity and partial fusion in those columellae and constriction of the bases (Figs. 37-39, 42, 47, 51) not seen in Lessingianthus. Only a few other species, such as V. cotoneaster and V. cleocalderonae, with Type C pollen, have been noted as having similar basal col- umellae (Figs. 12, 18). In all of these, the columellae show at least some tendency to- ward the rhizomatous condition, and in a few parts of some grains rhizomiform col- umellae are present (Figs. 37-39, 42). Limitations and exclusions. — Although some pollen types evidently reflect related groups within the genus Lepidaploa, a sub- division of the genus by pollen alone would be artificial. Only the species with Type D pollen are considered to constitute a phy- letic group among themselves. Species with the Type G pollen appear to relate to at least two different elements of the species group having pollen Type C, but these pollen types reflect large groups of species, and they pre- dominate in different geographical areas. Taxonomic subdivisions in the latter group are not expected to coincide completely with the limits of the pollen types. No special status is foreseen for the one Venezuelan species with Aynia Type pollen. In the following undivided alphabetical list of the species of Lepidaploa, the pollen VOLUME 103, NUMBER 2 473 Figs. 1-9. Lepidaploa, Type C pollen. 1-3. L. arborescens from St. Vincent, Morton 5721. 1. Polar view. 2. Colpar view. 3. Broken grain showing baculae. 4-6. L. sericea from Haiti, Leonard 8990. 4. Colpar view. 5. View showing intercolpus. 6. Crest of grain showing rhizomiform baculae broken away from footlayer. 7, 8. L. canescens from Ecuador, Hitchcock 20342. 7. Polar view. 8. Broken grain showing baculae. 9. L. cotoneaster from Brazil, King & Bishop 8620, polar view. types are indicated under each of the species as C, D, G, Aynia Type, B/G for G Types with a tendency for Type B areolation, and C/G for pollen with polar areoles usually at only one pole. One Neotropical species examined in this study, Acilepidopsis echitifolia (Mart. ex DC.) H. Robinson (1989), synonym Vernonis ar- arana Gardn., of Brazil and Paraguay, has lophate pollen and shows some features that suggest relationship to Lepidaploa. How- ever, examination of the pollen (Figs. 59, 474 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 12.0um 4.3um 1.76um Figs. 10-18. Lepidaploa, Type C pollen. 10-12. L. cotoneaster from Brazil, King & Bishop 8620. 10. Colpar view. 11. Intercolpar view. 12. Detail of crest showing baculae. 13-16. L. salzmannii from Peru, Ferreyra 9327 (13, 14); from Brazil, Hatschbach 43889 (15, 16). 13. Polar view. 14. Intercolpar view. 15. Broken grain showing rhizomiform baculae weakly attached to footlayer. 16. Detached exine showing rhizomiform baculae from below. 17, 18. L. cleocalderonae from Brazil, Calderon et al. 2689, holotype. 17. Polar view. 18. Detail of crest showing baculae. VOLUME 103, NUMBER 2 475 Figs. 19-24. Lepidaploa tovarensis pollen, from Venezuela, Pittier 12794, isotype. 19. Polar view showing three intercolpar areoles meeting at pole. 20-22. Colpar views showing crosswalls with distinct median sutures. 23. Intercolpar view. 24. Detail of crests showing modified baculae. 60) shows a type that is called Type E by Jones (1981) in his survey of the O!d World members of the tribe. Such pollen is not otherwise known from any Vernonia sensu lato native to the Neotropical region. The form of the pollen strongly suggests that the species is actually most closely related to Old World members of the tribe such as the African Vernonia polysphaera Baker and the Asian Acilepis squarrosa D. Don that have the same type of pollen (Kingham 1976; Jeffrey 1988). Large reddish glands, branched inflorescences, and procumbent bases of the stems help to distinguish the South American Acilepidopsis from possible relatives in both hemispheres (Robinson 1989). Geography Lepidaploa has the most extensive geo- graphical range of any genus in the related Complex, occurring from Mexico and the West Indies southward through most of montane or savanna areas of South Amer- ica. In contrast, the other genera are all pre- dominantly or exclusively Brazilian, except the monotypic Aynia of Peru. Chromosome Number All available counts (Jones 1979a, Keeley 1978) indicate that the Lepidaploa Complex has the New World pattern of n = 17 chro- mosomes (Jones 1977). Eleven species of Lepidaploa have been counted, including 476 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2. Jum 12.5um Figs. 25-33. Lepidaploa, Type G pollen. 25-27. L. psilostachya from Brazil, Riedel & Lund 1745/590. 25. Polar view. 26. Intercolpar view. 27. Detail of crests showing rhizomiform baculae. 28-31. L. subsquarrosa from Brazil, Rose 20194 (28-30) and Mimura 491 (31). 28. Polar view. 29. Intercolpar view. 30, 31. Broken grains showing rhizomiform baculae. 32, 33. L. persericea from Brazil, Mori et al. 13700. 32. Polar view. 33. Broken grain showing rhizomiform baculae weakly attached to footlayer. VOLUME 103, NUMBER 2 477 15. 8um ca? Leaf 16.7um Figs. 34-42. Lepidaploa, Type D pollen. 34-39. L. aurea from Brazil, Irwin et al. 14595. 34. Polar view. 35. Colpar view showing crosswalls. 36. Intercolpar view. 37-39. Details of crests showing modified baculae. 40-42. L. tombadorensis from Brazil, Harley et al. 22906. 40, 41. Colpar views showing crosswalls. 42. Broken grain showing crest with modified baculae. 478 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON »oum Figs. 43-51. Lepidaploa, Type D pollen. 43-47. L. chalybaea from Brazil, Mori & Benton 13094. 43. Polar view. 44. Colpar view showing crosswalls. 45, 46. Intercolpar views showing variation in areole pattern. 47. Detail of crest showing modified baculae and weak attachment to footlayer. 48-51. L. rufogrisea from Brazil, Cuatrecasas & Duarte 26641. 48. Polar view. 49. Colpar view showing crosswalls. 50. Intercolpar view. 51. Detail of crest showing modified baculae. VOLUME 103, NUMBER 2 479 a.4um 17.6um Figs. 52-60. Various lophate pollen types in the Vernonieae. 52, 53. Lessingianthus rubricaulis from Co- lombia, Pennell 10657. 52. Polar view. 53. Detail of crests showing baculae. 54. Mattfeldanthus nobilis from Brazil, Hatschbach 50443, broken grain showing baculae. 55, 56. Baccharoides adoensis from Rhodesia, Best 336. 55. Colpar view. 56. Detail of crest showing baculae. 57, 58. Cyanthillium patula from the Philippines, Steiner 468. 57. View showing pore. 58. Broken grain showing partial rhizomiform bridges under crests. 59, 60. Acilepidopsis echitifolia from Paraguay, Pedersen 3269. 59. View showing pore. 60. Detail of crests showing baculae. 480 one hexaploid, V. trinitatis. The base num- ber has also been reported for other genera of the Complex, Lessingianthus (Robinson 1988a) and Chrysolaena (Robinson 1988b), but the latter genera both show an incidence of tetraploidy that has not yet been reported from Lepidaploa. The known chromosome numbers are given with the pollen type under the species in the taxonomic summary. Further Distinctions A number of Brazilian species such as Vernonia subsquarrosa have insect galls of a type mentioned in the original description of V. alvimii (Robinson 1980). The galls are actually achenes infested with Dipteran lar- vae so that they grow larger than normal. They are fat and rather fusiform with a much reduced pappus. Such achenes have not been seen in the related genera such as Lessin- gianthus. None of the species of Lepidaploa have the generally larger size of all floral parts that is common in Lessingianthus. Only one species belonging to Lepidaploa, Vernonia eriolepis, was placed in the Vernonia section Lepidaploa Macrocephalae of Baker (1873) that included many species now placed in Lessingianthus. Lepidaploa eriolepis has some of the smallest floral parts of any species that was included in that section. The difference in size of floral parts in the genera seems to extend to the width of the style shaft and to some extent to size of the pollen. The somewhat larger pollen of Les- singianthus may be a factor in the common occurrence of extra intercolpar areoles in that genus. The difference in average size of floral parts in Lepidaploa and Lessingianthus is like that which sometimes results from higher chromosome numbers. However, the basic numbers of the two genera both seem to be n = 17. Some of the specimens of Lepidaploa with larger pollen, approaching that of Lessingianthus in size, may have PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON higher ploidy levels as in V. trinitensis, but the larger size in Lessingianthus seems to be achieved without increase in chromo- some number. Future studies should at- tempt to measure actual DNA content. Taxonomic Summary The genus Lepidaploa is delimited in the present series of studies in the following manner. Lepidaploa (Cassini) Cassini Lepidaploa (Cassini) Cassini in G. Cuvier, Dict. Sci. Nat. 36:20. 1825; 60:586. 1830. Rchb., H. G. L., Consp. Regn. Veg. 1:99. 1829 [1828]; Bartl., F. G., Ord. Nat. PI. 136. 1830; Spach, E., Hist. Nat. Veg. 10: 39. 1841. Vernonia subg. Lepidaploa Cassini, Bull. Sci. Soc. Philom. 1817:66. 1817. Vernonia sect. Lepidaploa (Cassini) DC., Prodr. 5:26. 1836. Annual or perennial herbs or shrubs, erect to spreading, to 1.5(—3.0) m tall; stems, leaves, and involucral bracts sparsely scab- rid to sericeous or weakly tomentose, often glandular-punctate. Leaves alternate (op- posite in L. canescens var. opposita), sessile to shortly petiolate. Inflorescence cymose with short or long seriately cymose branch- es, subcapitular bracts slightly to markedly smaller than nearest vegetative leaves. Heads sessile, broadly campanulate; bracts of involucre 1.3—3.0 times as numerous as florets in the head, unappendaged, in 3-6 series, outer bracts often slender and more spreading with aristate tips, inner bracts persistent, usually more erect and acute. Flowers (8—)10—-35 in a head; corollas usu- ally lavender, narrowly funnel-form, throats short but distinct, lobes bearing long hairs, spicules, glands, or with combinations, veins broadly fused at tips of lobes; anther ap- pendages non-glanduliferous (except pos- sibly in L. helophila); style base with an expanded node and with sclerified cells be- VOLUME 103, NUMBER 2 low. Achenes prismatic, mostly 8—10-ribbed, usually densely long-setuliferous, often with glands on the surface, with or without resin- iferous cells; carpopodia stopper-shaped or turbinate, extending upward among the bas- es of the lowest setulae, the cells oblong with porose lateral walls; inner pappus of nu- merous long capillary oristles, about as long as the corolla, outer pappus of distinct shorter scales. Pollen grains 45—50(—60) um in diameter, lophate with crests rhizoma- tous or incompletely columellate, columel- lae narrower at base when present, crest pat- tern usually with two equatorial intercolpar areoles, sometimes with polar areoles (type C), with crosswalls above and below pores (type D), or with colpi continuous to the poles (type G). Lectotype: Vernonia albicaulis Vahl ex Pers. [=Lepidaploa glabra (Willd.) H. Rob- inson]. The genus Lepidaploa can be credited with the following 116 species. The synonymy given below includes basionyms of the ac- cepted species, basonyms of species placed in synonymy, and any subsequent combi- nations of these names at the species level in Vernonia. Synonymies generally follow Keeley (1978) for the West Indies, Ariste- guieta (1964) for Venezuela, Jones (1980) for Peru, Cabrera (1944) for Argentina, and have been influenced by Baker (1873) in some of the older names from Brazil. Lepidaploa acutiangula (Gardn.) H. Robinson, comb. nov. Vernonia acutiangula Gardner, London J. Bot. 5:225. 1846. Brazil (Maranhao). Pol- len C. Lepidaploa acuminata (Less.) H. Robinson comb. nov. Vernonia acuminata Less., Linnaea 6:663. 1831. V. expansa Gleason, Bull. New York Bot. Gard. 4:186. 1906. Jamaica. Pollen C; N = 17 (Keeley 1978). 481 Lepidaploa alvimii (H. Robinson) H. Robinson, comb. nov. Vernonia alvimii H. Robinson, Phytologia 45:168. 1980. Brazil (Bahia). Pollen G. Lepidaploa araguensis (Badillo) H. Robinson, comb. nov. Vernonia araguensis Badillo, Bol. Soc. Ve- nez. Ci. Nat. 10:283. 1946. Venezuela. Pollen C. Lepidaploa araripensis (Gardn.) H. Robinson, comb. nov. Vernonia araripensis Gardner, London J. Bot. 5:222. 1846. Brazil (Para). Pollen C. Lepidaploa arborescens (L.) H. Robinson, comb. nov. Conyza arborescens L., Syst. Nat. ed. 10, 2: 1213. 1759. Vernonia arborescens (L.) Swartz, Fl. Ind. Occid. 3:1320. 1806. V. divaricata Swartz, Fl. Ind. Occid. 3:1319. 1806. V. divaricata Less., Linnaea 4:306. 1829. V. icosantha DC., Prodr. 5:49. 1836. V. albicoma Gleason, Bull. New York Bot. Gard. 4:185. 1906. V. intonsa Gleason, Bull. New York Bot. Gard. 4:182. 1906. V. permollis Gleason, Bull. New York Bot. Gard. 4:179. 1906. V. ventosa Gleason, Bull. New York Bot. Gard. 4:179. 1906. V. amarantina Gleason, Bull. Torrey Bot. Club 40:307. 1913. V. parvuliceps Ek- man, Ark. Bot. 13(15):71. 1914. Antilles, Pollen C, Figs. 1-3; N = 17 (Keeley 1978). The present concept differs from that of Keeley (1982) by its restriction to Antillean material. Separate specific status is restored for the continental material that was placed in synonymy by Keeley. The continental species is discussed under the name Lepi- daploa canescens. Lepidaploa arbuscula (Less.) H. Robinson, comb. nov. Vernonia arbuscula Less., Linnaea 6:664. 1831. V. arcuata Gleason, Bull. Torrey 482 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bot. Club 33:185. Gleason, Bull. Torrey Bot. Club 33:187. 1906. V. bahamensis Griseb., Fl. Brit. W. Ind. 352. 1861. Bahamas. Pollen C; N = 17 (Keeley 1978). Lepidaploa arenaria (Mart. ex DC.) H. Robinson, comb. nov. Vernonia arenaria Mart. ex DC., Prodr. 5: 54. 1836. V. sarmentiana Gardner, Lon- don J. Bot. 5:221. 1846. Brazil (Bahia, Piauhy). Pollen D. Lepidaploa argyrotricha (Sch.Bip. ex Baker) H. Robinson, comb. nov. Vernonia argyrotricha Sch.Bip. ex Baker in Mart., Fl Bras. 6(2):96. 1873. Brazil (Goias, Espirito Santo, Minas Gerais, Rio de Janeiro). Pollen C; N = 17 (Keeley 1978). Lepidaploa aristosquamosa (Britton) H. Robinson, comb. nov. Vernonia aristosquamosa Britton, Bull. Torrey Bot. Club 18:332. 1891. Bolivia. Pollen G. Lepidaploa aronifolia (Gleason) H. Robinson, comb. nov. Vernonia aronifolia Gleason, Bull. Torrey Bot. Club 40:323. 1919. Cuba. Pollen C/G. Lepidaploa aurea (Mart. ex DC.) H. Robinson, comb. nov. Vernonia aurea Mart. ex DC., Prodr. 5:58. 1836. Brazil (Bahia, D.F., Goias, Minas Gerais). Pollen D, Figs. 34-39. Lepidaploa auyantepuiensis (Aristeg.) H. Robinson, comb. nov. Vernonia auyantepuiensis Aristeg., Acta Bot. Venez. 2(5—8):362. 1967. Venezuela. Pol- leng@: 1906. V. obcordata The species seems close to Lepidaploa eh- retiifolia Benth. but the pubescence of the stems and leaf undersurfaces is coarser and more erect. Also, the pollen is larger (nearly 60 wm in diam.), and there are usually three intercolpar areoles as in Type B pollen. The size of the grains and the coarseness of the hairs may indicate the species is a polyploid. Lepidaploa barbata (Less.) H. Robinson, comb. nov. Vernonia barbata Less., Linnaea 4:287. 1829. Brazil (Bahia, Mato Grosso, Minas Gerais). Pollen D. Lepidaploa bakerana (Britton) H. Robinson, comb. nov. Vernonia bakerana Britton, Bull. Torrey Bot. Club 18:331. 1891. Bolivia. Pollen C. Lepidaploa beckii H. Robinson, sp. nov. Plantae suffrutescentes et interdum sub- volubiles 1.4—3.0 m longae laxe ramosae. Caulis brunnei et atrescentes dense sordide velutini. Folia alterna, petiolis 5-10 mm longis dense velutinis; laminae ovato-lan- ceolatae 15—27 cm longae et 6-11 cm latae base rotundatae ad quintum basilares latis- simae margine integrae vel minime remote crenulatae apice breviter anguste acumi- natae supra virides in nervis insculptae ten- uiter sericeae subtus pallidiores in nervis valde exsculptae perdense sericeae vel sub- lanatae, nervis secundariis utrinque 14-18. Inflorescentiae dense ramosae in ramis pri- marlis distincte bractiiferae, bracteis foli- iformibus plerumque 3—9 cm longis et 1-3 cm latis, bracteis ramulis subnullis. Capit- ula sessilia in nodis binata vel congesta breviter campanulata 5-6 mm alta; squa- mae involucri plerumque virides interiores distaliter brunnescentes ca. 40 subimbri- catae 3—4-seriatae lanceolatae 1.5-4.0 mm longae et 0.5—1.0 mm latae plerumque extus dense pilosulae et apice subaristatae, interi- ores sparse minute puberulae apice breviter acutae saepe recurvatae. Flores ca. 30 in VOLUME 103, NUMBER 2 capitulo. Corollae albae 4.0-—4.5 mm longae extus inferne glabrae distaliter et in margi- nis lobarum dense spiculiferae, tubis infun- dibularibus ca. 2 mm longis, faucibus ca. | mm longis, lobis oblongo-lanceolatis ca. 2 mm longis et 0.6 mm latis; thecae anthera- rum ca. 2 mm longae, appendicibus api- calibus ca. 0.5 mm longis glabris; basi sty- lorum abrupte disciformes. Achaenia ca. 2 mm longae dense sericeo-setuliferae non glandulferae distincte mediocriter idioblas- tiferae; setae pappi flavae ca. 40 plerumque 3.5-4.0 mm longae superne sensim dis- tincte latiores in sereibus anguste lanceo- latae ad 0.8 mm longae. Grana pollinis in diametro ca. 45 um valde lophata, reticula in typo C. (Fig. 61). Type. — Bolivia: La Paz: Prov. Nor Yun- gas, Suapi 16 km hacia Santa Rosa, 1650 m, bosque montanoso, sub-arbusto de 2 m erecto, frutos, en el borde del camino, 25.9.1987, St. G. Beck 13640 (holotype US; isotype LPB). Paratypes.— Bolivia: La Paz: Prov. Mu- rillo, Suapi 22 km hacia Santa Rosa de Quilo Quilo, 1300 m, restos del monte en el borde del camino, arbusta trepanda alga voluble, frutos en globulos, 25.9.1987, St. G. Beck 13638 (LPB, US); Valley of the Rio Zongo, approx. 2 hour by trail which climbs from the Cahua Hydroelectric Plant on the left bank of the Rio Zongo, moist forest with scattered chacos, 16°05’S, 68°03'W, scan- dent shrub, stems up to 3 m, 23 Apr 1982, J. C. Solomon 7563 (MO; US); Prov. Nor Yungas, 5.5 km below Coroico towards Yo- losa (1.4 km above Yolosa), disturbed road- side forest, 1400 m, 16°13’S, 67°44’W, suf- frutescent, 1.5 m, corollas white, 16 May 1985, J. C. Solomon 13726 (MO, US), 21.1 km al noroeste del camino entre Yolosa y Caranavi por el camino a Suapi (ca. 2.5 km al oeste de Suapi, cerca del puente sobre el Rio Suapi), bosque humido muy tocado, 16°07’S, 67°47'W, 1200 m, corolas blancas, tallos arqueados hasta 3 m, 27 May 1988, Solomon 18400 (MO, US). The species is readily distinguished by the large, coarse leaves having rounded, nearly 483 sessile bases and veins impressed in the up- per surface. The inflorescence is also dis- tinctive in the branches with many small crowded heads that mostly lack obvious subcapitular bracts. The corollas seem to be unique in having the margins of the lobes densely spiculiferous along their whole length. Lepidaploa bolivarensis (Badillo) H. Robinson, comb. nov. Vernonia bolivarensis Badillo, Bol. Soc. Ve- nez, Ci. Nat. 23(103):291. 1963. Verno- nia glandulosa Badillo, Bol. Soc. Venez. Ci. Nat. 10:283. 1946, not V. glandulosa DC., Prodr. 5:22. 1836. Guyana, Vene- zuela. Pollen C. Lepidaploa borinquensis (Urban) H. Robinson, comb. nov. Vernonia borinquensis Urban, Symb. An- till. 3:390. 1903. Puerto Rico. Pollen C/G. Lepidaploa buchtienii (Gleason) H. Robinson, comb. nov. Vernonia buchtienii Gleason, Amer. J. Bot. 10:302. 1923. Bolivia. Pollen G. Lepidaploa canescens (H.B.K.) H. Robinson, comb. nov. Vernonia canescens H.B.K., Nov. Gen. Sp., folio ed. 4:27. 1818. V. mollis H.B.K., Nov. Gen. Sp., folio ed. 4:28. 1818. V. geminata H.B.K., Nov. Gen. Sp., folio ed. 4:28. 1818. V. bullata Benth. ex Oerst., Vidensk. Meddel. Naturhist. Foren. Kjobenhavn 1852:67. 1853. V. arbores- cens var. cuneifolia Britton, Bull. Torrey Bot. Club 18:311. 1891. V. sodiroi Hi- eron. ex Sodiro, Bot. Jahrb. Syst. 29:1. 1900. V. volubilis Hieron., Bot. Jahrb. Syst. 36:460. 1905. V. hirsutivena Glea- son, Bull. New York Bot. Gard. 4:175. 1906. V. patuliflora Rusby, Bull. New York Bot. Gard. 4:376. 1906. V. purpusil T. S. Brandegee, Univ. Calif. Publ. Bot. 484 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON HERBARIO NACIONAL DE BOLIVIA (Convenio M.N.H.N. — IE.) rnonia. Lepidaploa beckis HRosins Holotype ec. La Paz, Provincia Nor Yungas, Suapi 16 Km. hacia Santa Resa, 1650 m. 3osque montanoso. Sub-arbuste de 2m. erectc, frutas, en UNITED STATES ; : el borde del camino. 3143669 sot DaRe Be nee Ne. 19640 NATIONAL HERBARIUM Fig. 61. Holotype of Lepidaploa beckii H. Robinson, Beck 13640 (US). VOLUME 103, NUMBER 2 6:197. 1915. V. cuneifolia (Britton) Glea- son, Amer. J. Bot. 10:301. 1923, not V. cuneifolia Gardner, London J. Bot. 5:215. 1846. V. pseudomollis Gleason, Amer. J. Bot. 10:307. 1923. V. rusbyi Gleason, Amer. J. Bot. 19:753. 1932, based on V. arborescens var. cuneifolia Britton. V. polypleura Blake, J. Wash. Acad. Sci. 28: 478. 1938. V. medialis Standl. & Stey- erm., Publ. Field. Mus. Nat. Hist., Bot. Ser. 23:148. 1943. V. spiritu-sancti Cuatr., Bot. Jahrb. Syst. 77:58. 1956. V. unillen- sis Cuatr., Bot. Jahrb. Syst. 77:59. 1956. Mexico, Guatemala, Costa Rica, Pana- ma, Colombia, Venezuela, Ecuador, Peru, Bolivia. Pollen C, Figs. 7, 8. This species was placed in the synonymy of Vernonia arborescens (L.) Swartz by Kee- ley (1982), but her concept is overly broad. Typical West Indian Lepidaploa arbores- cens and continental L. canescens are some- what similar in aspect, but they are not con- sidered to be close relatives in this study. The continental species has characteristi- cally larger leaves and has smaller, more pubescent involucres. The West Indian L. arborescens characteristically has foliose bracts at the immediate bases of the heads, but the continental L. canescens essentially lacks such bracts. In the few cases where foliose bracts occur in L. canescens (Pana- ma, King 5256, US), they are attached on the opposite side of the stem from the heads. The achene surfaces in L. canescens have resiniferous idioblasts, whereas those of L. arborescens have glands instead. The syn- onymy of Keeley is redistributed between the two species. Lepidaploa canescens var. opposita (H. Robinson) H. Robinson, comb. nov. Vernonia canescens var. opposita H. Rob- inson, Phytologia 49:261. 1981. Colom- bia. 485 Lepidaploa carachensis (Badillo) H. Robinson, comb. nov. Vernonia carachensis Badillo, Ernstia 48: 41. 1988. Venezuela. Pollen C/G. Lepidaploa chalybaea (Mart. ex DC.) H. Robinson, comb. nov. Vernonia chalybaea Mart. ex DC., Prodr. 5: 54. 1836. Brazil (Bahia, Ceara). Pollen D, Figs. 43-47. Lepidaploa chamissonis (Less.) H. Robinson, comb. nov. Vernonia chamissonis Less., Linnaea 4:304. 1829. V. parodii Cabrera, Darwiniana 5: 187. 1941. Brazil (Parana, Santa Catari- na, Sao Paulo), Argentina (Chaco), Par- aguay. Pollen C. Lepidaploa chrysotricha (Alexander) H. Robinson, comb. nov. Vernonia chrysotricha Alexander, Lloydia 2:217. 1939. Guyana. Pollen C/G. Lepidaploa cleocalderonae (H. Robinson) H. Robinson, comb. nov. Vernonia cleocalderonae H. Robinson, Phytologia 46:108. 1980. Brazil (Ama- zonas). Pollen C, Figs. 17, 18. Lepidaploa commutata (Ekman) H. Robinson, comb. nov. Vernonia commutata Ekman, Ark. Bot. 13(15):77. 1914. Cuba. Pollen C. Lepidaploa complicata (Wright ex Griseb.) H. Robinson, comb. nov. Vernonia complicata Wright ex Griseb., Cat. Pl. Cub. 143. 1866. Cuba. Pollen C. 486 Lepidaploa cordiaefolia (H.B.K.) H. Robinson, comb. nov. Vernonia cordiaefolia H.B.K., Nov. Gen. Sp., folio ed. 4:29. 1818. Colombia. Pol- leng@: Lepidaploa costanensis (Badillo) H. Robinson, comb. nov. Vernonia costanensis Badillo, Ernstia 23:32. 1984. Venezuela. Pollen C. Lepidaploa costata (Rusby) H. Robinson, comb. nov. Vernonia costata Rusby, Mem. Torrey Bot. Club 6:53. 1896. Peru, Bolivia. Pollen G. Lepidaploa cotoneaster (Willd. ex Spreng.) H. Robinson, comb. nov. Conyza cotoneaster Willd. ex Spreng., Syst. Veg. 3:509. 1826. Vernonia cotoneaster (Willd. ex Spreng.) Less., Linnaea 4:298. 1829. V. axilliflora Less., Linnaea 4:297. 1829. V. debilis Mart. ex DC., Prodr. 5: 54. 1836. Brazil (Bahia, Minas Gerais). Pollen C, Figs. 9-12. Lepidaploa coulonioides (H. Robinson) H. Robinson, comb. nov. Vernonia coulonioides H. Robinson, Phy- tologia 49:263. 1981. Brazil (Rio de Ja- neiro). Pollen G. Lepidaploa crassifolia (Rusby) H. Robinson, comb. nov. Vernonia crassifolia Rusby, Bull New York Bot. Gard. 8:124. 1912. Bolivia. Pollen G. Lepidaploa cuiabensis (Baker in Mart.) H. Robinson, comb. nov. Vernonia cuiabensis Baker in Mart., Fl. Bras. 6(2):37. 1873. Brazil (Mato Grosso). Pol- len D. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lepidaploa danielis (Cuatr.) H. Robinson, comb. nov. Vernonia danielis Cuatr., Bot. Jahrb. Syst. 77:54. 1956. Colombia. Pollen C. Lepidaploa decumbens (Gardner) H. Robinson, comb. nov. Vernonia decumbens Gardner, London J. Bot. 4:115. 1845. Brazil (Espirito Santo, Rio de Janeiro, Sao Paulo). Pollen G. The species is close to Lepidaploa sub- squarrosa, which has been known as Ver- nonia geminata Less., but the stems are stouter and straighter, the leaves are more pointed, and the inflorescence is denser. Lepidaploa deflexa (Rusby) H. Robinson, comb. nov. Vernonia deflexa Rusby, Bull. New York Bot. Gard. 4:376. 1907. Bolivia. Pollen G. Lepidaploa densipaniculata (Rusby) H. Robinson, comb. nov. Vernonia densipaniculata Rusby, Bull. New York Bot. Gard. 8:126. 1912. Bolivia. Pollen G. Only the type (Cargadira, Williams 1534, NY) and one recent collection (La Paz, Mu- rillo, below Lago Zongo dam, Solomon 10740, MO, US) have been seen. Lepidaploa desiliens (Gleason) H. Robinson, comb. nov. Vernonia desiliens Gleason, Bull. Torrey Bot. Club 40:316. 1913. Cuba. Pollen C. Lepidaploa edmundoi (Barroso) H. Robinson, comb. nov. Vernonia edmundoi Barroso, Arq. Jard. Bot. Rio de Janeiro 17:21. 1959. Brazil (Ba- hia). Pollen D. VOLUME 103, NUMBER 2 Lepidaploa ehretiifolia (Benth.) H. Robinson, comb. nov. Vernonia ehretiifolia Benth., London J. Bot. 2:39. 1840. V. schomburgkiana Sch.Bip., Linnaea 20:509. 1847. Guyana, Vene- zuela. Pollen C/G. Lepidaploa ekmanii (Urban) H. Robinson, comb. nov. Vernonia ekmanii Urban, Ark. Bot. 17(7): 62. 1921. Haiti. Pollen C/G. Lepidaploa eriolepis (Gardner) H. Robinson, comb. nov. Vernonia eriolepis Gardner, London J. Bot. 5:224. 1846. V. riedelii Sch.Bip. ex Baker in Mart., Fl. Bras. 6(2):64. 1873. Brazil (Ceara, Mato Grosso, Minas Gerais, Para- na, Santa Catarina, Sao Paulo). Pollen G. Lepidaploa ferreyrae (H. Robinson) H. Robinson, comb. nov. Vernonia ferreyrae H. Robinson, Phytolo- gia 45:158. Feb. 1980. V. apurimacensis S. B. Jones, Fieldiana, Bot. N.S. 5:38. Dec 1980. Peru. Pollen C. Lepidaploa fieldiana (Gleason) H. Robinson, comb. nov. Vernonia fieldiana Gleason, Bull. Torrey Bot. Club 59:374. 1932. Peru. Pollen G. Lepidaploa fournetii (H. Robinson & B. Kahn) H. Robinson, comb. nov. Vernonia fournetii H. Robinson & B. Kahn, Phytologia 58:252. 1985. Bolivia. Pollen G. Lepidaploa frangulaefolia (H.B.K.) H. Robinson, comb. nov. Vernonia frangulaefolia H.B.K., Nov. Gen. Sp., folio ed. 4:29. 1818. Colombia. Pol- len C. 487 Lepidaploa fruticosa (L.) H. Robinson, comb. nov. Conyza fruticosa L., Sp. Pl. ed. 2. 1209. 1763. Vernonia fruticosa (L.) Swartz, FI. Ind. Occid. 3:1323. 1806. V. buchii Ur- ban, Repert. Spec. Nov. Regni Veg. 16: 146. 1919. Hispaniola. Pollen C; N = 17 (Keeley 1978). Lepidaploa glabra (Willd.) H. Robinson, comb. nov. Conyza glabra Willd., Sp. Pl. 3:1940. 1803, non Vernonia glabra Vatke, 1877. Eu- patorium obtusifolium Willd., Sp. Pl. 3: 1768. 1803, non Vernonia obtusifolia Less., 1829. Vernonia albicaulis Vahl ex Pers., Syn. Pl. 2(2):404. 1807. V. longi- folia Pers., Syn. Pl. 2(2):404. 1807. Lep- idaploa lanceolata Cass., Dict. Sci. Nat. 26:17. 1823, nom. inval., prior to vali- dation of Lepidaploa at generic level. Ver- nonia punctata Swartz ex Wikstr., Kongl. Vetensk. Acad. Handl. 1827:72. 1828. V. emarginata Wikstr., Kongl. Vetensk. Acad. Handl. 1827:73. 1828. V. vahliana Less., Linnaea 4:306. 1829. Eupatorium secundiflorum Bertero ex DC., Prodr. 5: 48. 1836. V. thomae Benth. ex Oerst, Vi- densk. Meddel. Naturhist. Foren. Kjg- benhavn 1852:66. 1853. V. longifolia var. sintenisii Urban, Symb. Antill. 1:456. 1899. V. sintenisii (Urban) Gleason, Bull. New York Bot. Gard. 4:187. 1906. Ver- nonia gleasonii Ekman, Ark. Bot. 13(15): 54. 1914. V. shaferi Gleason, Bull. Torrey Bot. Club 46:238. 1919. Puerto Rico, Lesser Antilles. Pollen C; N = 17 (Keeley 1978). In Lepidaploa, the oldest name can be used for this species, even though it (and a second older name) are blocked from use in Vernonia by being later homonyms. Ver- nonia albicaulis remains the proper name for the species in Vernonia. 488 Lepidaploa gnaphaliifolia (A. Rich.) H. Robinson, comb. nov. Vernonia gnaphalifolia A. Rich. in Sagra, Hist. Fis. Pol. Nat. Cuba, Bot. 2:33. 1850. V. membranacea Griseb., Cat. Pl. Cub. 144. 1866. V. crassinervia Wright ex Gleason, Bull. New York Bot. Gard. 4: 180. 1906. V. sublanata Gleason, Bull. New York Bot. Gard. 4:177. 1906. V. sublanata var. angustata Gleason, Bull. New York Bot. Gard. 4:177. 1906. V. an- gustata (Gleason) Gleason, Bull. Torrey Bot. Club 40:309. 1913. V. gnaphaliifolia var. platyphylla Gleason, Bull. Torrey Bot. Club. 46:238. 1919. V. platyphylla (Glea- son) Ekman ex Urban, Repert. Spec. Nov. Regni Veg. 26:100. 1929. V. nervosa Alain, Contr. Ocas. Mus. Colegio ““De la Salle.” 18:150. 1960. Cuba. Pollen C. Lepidaploa gnaphalioides (Sch.Bip. ex Mart.) H. Robinson comb. nov. Vernonia gnaphalioides Sch.Bip. ex Baker in Mart., Fl. Bras. 6(2):78. 1873. Brazil (Minas Gerais, Parana). Pollen G. Lepidaploa gracilis (H.B.K.) H. Robinson, comb. nov. Vernonia gracilis H.B.K., Nov. Gen. Sp., folio ed. 4:27. 1818. V. moritziana Sch.Bip., Linnaea 20:511. 1847. Colom- bia, Venezuela. Pollen C. The concept follows Badillo (1984). Lepidaploa grisea (Baker) H. Robinson, comb. nov. Vernonia grisea Baker in Mart., Fl. Bras. 6(2):61. 1873. Brazil (Amazonas, Ceara, Para, Piauhy). Pollen D. The species seems close to Lepidaploa ar- enaria Gardner, but the latter is basically a costal species with more succulent leaves. Vegetatively the species seems close to L. obtusifolia of southern Brazil, but the latter PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON belongs to the group with type C pollen. The latter further differs by the rather indistinct cymes in the inflorescence with little or no reduction in the size of the bracts of the inflorescence. Lepidaploa hagei (H. Robinson) H. Robinson, comb. nov. Vernonia hagei H. Robinson, Phytologia 45: 176. 1980. Brazil (Bahia). Pollen G. Lepidaploa harrisii (S. Moore) H. Robinson, comb. nov. Vernonia harrisii S. Moore, J. Bot. 66:164. 1928. Jamaica. Pollen C. Lepidaploa helophila (Mart. ex DC.) H. Robinson, comb. nov. Vernonia helophila Mart. ex DC., Prodr. 5: 50. 1836. V. subcordata Gardner, London J. Bot. 5:226. 1846. Brazil (Bahia, Mato Grosso, Minas Gerais, Sao Paulo). Pol- len G. The distribution and synonymy is taken from Baker (1873), but such a range seems unlikely for a species so poorly represented in collections. A specimen from Bahia (Har- ley 18497, US) may be this species; it has glands on the anthers with one or two glands on the apical appendages. The material shows no other important characters of the comparatively remotely related genus Chry- solaena (Robinson 1988b). Lepidaploa jenssenii (Ekman ex Urban) H. Robinson, comb. nov. Vernonia jenssenii Ekman ex Urban, Re- pert. Spec. Nov. Regni Veg. 26:98. 1929. Cuba. Pollen C. Lepidaploa karstenii (Sch.Bip.) H. Robinson, comb. nov. Vernonia karstenii Sch.Bip., Linnaea 30: 169. 1859. Colombia. Pollen C. VOLUME 103, NUMBER 2 Lepidaploa lehmannii (Hieron.) H. Robinson, comb. nov. Vernonia lehmannii Hieron., Bot. Jahrb. Syst. 19:44. 1894. Vernonia larensis Ba- dillo, Bol. Soc. Venez. Ci. Nat. 10:285. 1946. Venezuela, Colombia, Ecuador. Pollen C. Lepidaploa leptoclada (Sch.Bip.) H. Robinson, comb. nov. Vernonia leptoclada Sch.Bip., J. Bot. 1:233. 1863. Cuba. Pollen C. Lepidaploa liesneri (H. Robinson) H. Robinson, comb. nov. Vernonia liesneri H. Robinson, Phytologia 49:264. 1981. Venezuela. Pollen C. Material of this species was the basis for Badillo’s (1982) report of the Colombia Vernonia marguana Cuatr. from Venezue- la. The two species are vegetatively nearly alike and must be closely related. The Co- lombian species differs by having more ro- bust inflorescences with larger heads bear- ing more than twice as many flowers (over 20 vs. 8-9) and by bearing more numerous, shorter involucral bracts. Lepidaploa lilacina (Mart. ex DC.) H. Robinson, comb. nov. Vernonia lilacina Mart. ex DC., Prodr. 5: 48. 1836. V. adamantium Gardner, Lon- don J. Bot. 5:222. 1846. Brazil (Bahia, Minas Gerais). Pollen C. Lepidaploa luetzelburgii (Mattf.) H. Robinson, comb. nov. Vernonia luetzelburgii Mattf., Notizbl. Bot. Gart. Berlin-Dahlem 9:377. 1925. Brazil (Piauhy). Pollen D? A type photograph shows a species close in habit and pubescence to the more re- cently described Lepidaploa pinheiroi of Ba- hia, but the leaf tips are rounded and the 489 involucral bracts lack the long apical spines of the latter species. Lepidaploa macahensis (Glaziou ex Barroso) H. Robinson, comb. nov. Vernonia macahensis Glaziou ex Barroso, Arq. Jard. Bot. Rio de Janeiro 17:21. 1962. Brazil (Espirito Santo, Rio de Ja- neiro). Pollen C. I place here also material in herbaria un- der the name Vernonia tijucana Glaziou from Rio de Janeiro. The latter name was never validated by either Glaziou or by Ek- man. The lack of validation is in spite of some annotations by Ekman indicating his intention to publish the name. The Barroso publication was the first and only validation for either Glaziou name. Material under the name V. tijucana seems to have more entire leaves and more spreading cymose branches compared to typical Lepidaploa macahen- sis, but the differences do not seem worthy of species rank. Lepidaploa mandonii (Sch.Bip. ex Gleason) H. Robinson, comb. nov. Vernonia mandonii Sch.Bip. ex Gleason, Amer. J. Bot. 10:300. 1923. Bolivia. Pol- len C. Lepidaploa mapirensis (Gleason) H. Robinson, comb. nov. Vernonia mapirensis Gleason, Amer. J. Bot. 10:307. 1923. Bolivia, Peru. Pollen G. The species is redelimited to exclude Lep- idaploa trichoclada that was placed in syn- onymy by Jones (1980) and exclude the nu- merous specimens of L. sordidopapposa recently annotated by Jones as Vernonia mapirensis. The present species, as seen in the type (Buchtien 1533, US) and paratype (Buchtien 2462, US), has leaves similar to L. canescens, but the heads are somewhat larger and more densely arranged, the pap- 490 pus is dirty yellowish, the surfaces of the achenes lack resiniferous idioblasts, and the pollen seems consistently to be type G. One specimen from Paucatambo, Dept. Cuzco, Peru (Vargas 15495, US) seems to be the same species. Lepidaploa mapirensis seems closest to L. tristis, but the latter is larger, has more erect or retrorse pubescence on the stems, oblong elliptical leaves with the lateral veins widely separated, and heads that are often paired at the nodes. Lepidaploa mucronifolia (DC.) H. Robinson, comb. nov. Vernonia mucronifolia DC.., Prodr. S255 1836. Brazil (Bahia). Pollen D. The species seems very similar in aspect to Lepidaploa obtusifolia of the Rio de Ja- neiro area, but the two are apparently not closely related. The present species is a member of the group having Type D pollen and is a more pubescent plant with less ob- viously striated stems. Lepidaploa muricata (DC.) H. Robinson, comb. nov. Vernonia muricata DC., Prodr. 5:55. 1836. Brazil (Minas Gerais, Rio de Janeiro, Sao Paulo). Pollen G. Lepidaploa myriocephala (DC.) H. Robinson, comb. nov. Vernonia myriocephala DC., Prodr. 5:40. 1836. Peru. Pollen C. The species was placed in the synonymy of Lepidaploa arborescens with L. canescens by Keeley (1982), but the leaves are less pubescent and the secondary veins are less closely and less regularly arranged. Lepidaploa nitens (Gardner) H. Robinson, comb. nov. Vernonia nitens Gardner, London J. Bot. 5: 221. 1846. Brazil (Bahia, Goids). Pol- len D. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lepidaploa obtusifolia (Less.) H. Robinson, comb. nov. Vernonia obtusifolia Less., Linnaea 4:308. 1829. Brazil (Rio de Janeiro). Pollen C. The species superficially resembles some members of the genus from farther north, such as Lepidaploa grisea and L. mucroni- folia. The latter, however, have Type D pol- len, and they are not closely related. Lepidaploa orbicularis (Alain) H. Robinson, comb. nov. Vernonia orbicularis Alain, Contr. Ocas. Mus. Colegio “De la Salle” 18:15. 1960. V. leonis Alain, Contr. Ocas. Mus. Cole- gio “De la Salle” 18:14. 1960. Cuba. Pol- len C. Lepidaploa pallescens (Gleason) H. Robinson, comb. nov. Vernonia pallescens Gleason, Bull. New York Bot. Gard. 4:192. 1906. Lesser An- tilles. Pollen C. Lepidaploa pari (Badillo) H. Robinson, comb. nov. Vernonia pari Badillo, Bol. Soc. Venez. Ci. Nat. 10:284. 1946. Venezuela. Pollen C. Lepidaploa persericea (H. Robinson) H. Robinson, comb. nov. Vernonia persericea H. Robinson, Phyto- logia 44:292. 1979. V. cognata var. lun- diana Baker in Mart., Fl. Bras. 6(2):95. 1873. Brazil (Bahia, Espirito Santo, Mi- nas Gerais, Rio de Janeiro). Pollen G, Figs. 32, 33. Lepidaploa persicifolia (Desf.) H. Robinson, comb. nov. Vernonia persicifolia Desf., Cat. Pl. Hort. Par. ed. 3, 400. 1829. V. acutifolia Hook., Bot. Mag. 58:t. 3062. 1831. Brazil (Rio de Janeiro). Pollen C. VOLUME 103, NUMBER 2 During much of its taxonomic history, the species has been placed in the synonymy of Vernonia sericea L. C. Rich. The latter entity seems to be exclusively West Indian and has no particular resemblance to the present large-leaved species with its large, remote heads. Lepidaploa pineticola (Gleason) H. Robinson, comb. nov. Vernonia pineticola Gleason, Bull. New York Bot. Gard. 4:176. 1906. Cuba. Pol- len C/G. Lepidaploa pinheiroi (H. Robinson) H. Robinson, comb. nov. Vernonia pinheiroi H. Robinson, Phytolo- gia 45:179. 1980. Brazil (Bahia). Pol- len D. Lepidaploa pluvialis (Gleason) H. Robinson, comb. nov. Vernonia pluvialis Gleason, Bull. Torrey Bot. Club 40:312. 1913. V. reducta Gleason, Bull. Torrey Bot. Club 40:313. 1913. Ja- maica. Pollen C; N = 17 (Keeley 1978). Lepidaploa proctorii (Urbatsch) H. Robinson, comb. nov. Vernonia proctorii Urbatsch, Syst. Bot. 14: 589. 1989. Puerto Rico. Pollen C/G. Lepidaploa psilostachya (DC.) H. Robinson, comb. nov. Vernonia psilostachya DC., Prodr. 5:43. 1836. V. oxylepis Sch.Bip. ex Baker in Mart., 6(2):70. 1973. Brazil (Parana, Sao Paulo). Pollen B/G, Figs. 25-27. Lepidaploa purpurata (Gleason) H. Robinson, comb. nov. Vernonia purpurata Gleason, Bull. Torrey Bot. Club 40:322. 1913. V. praestans Ek- man & Urban, Repert. Spec. Nov. Regni Veg. 26:101. 1921. Cuba. Pollen C/G. 491 Lepidaploa reflexa (Gardner) H. Robinson, comb. nov. Vernonia reflexa Gardner, London J. Bot. 5:228. 1846. Brazil (Bahia, Minas Ge- rais). Pollen D. Lepidaploa remotiflora (L. C. Rich) H. Robinson, comb. nov. Vernonia remotiflora L. C. Rich., Actes Soc. Hist. Nat. Paris 1:112. 1792. V. sessili- flora Willd. ex Less., Linnaea 4:309. 1829. V. tricholepis DC., Prodr. 5:54. 1836. V. acilepis Benth. ex Oerst., Vidensk. Med- del. Naturhist. Foren, Kjobenhavn 1852: 68. 1853. Vernonia lithospermoides Ba- ker in Mart., Fl. Bras. 6(2):66. 1873. V. hirtiflora Sch.Bip. ex Baker in Mart., FI. Bras. 6(2):70. 1873. V. setososquamosa Hieron., Bot. Jahrb. Syst. 22:684. 1897. Venezuela, Guyana, Cayenne, Brazil, Paraguay, Bolivia, Argentina. Pollen G. A type photograph of Vernonia ovata Less. shows a plant from Brazil that is possibly related to Lepidaploa remotiflora, but it has broadly elliptical leaves and broader heads with recurved rather than straight outer bracts. Lepidaploa retrosetosa (H. Robinson) H. Robinson, comb. nov. Vernonia retrosetosa H. Robinson, Phyto- logia 45:159. 1980. Peru. Pollen G. Lepidaploa rigida (Swartz) H. Robinson, comb. nov. Conyza rigida Swartz, Prodr. 113. 1788. Vernonia rigida (Swartz) Swartz, Fl. Ind. Occid. 3:1322. 1806. Jamaica. Pollen C; N = 17 (Keeley 1978). Lepidaploa rimachii (H. Robinson) H. Robinson, comb. nov. Vernonia rimachii H. Robinson, Phytologia 49:266. 1981. Peru. Pollen G. 492 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lepidaploa rufogrisea (St. Hil.) H. Robinson, comb. nov. Vernonia rufo-grisea St. Hil., Voy. Distr. Diam. 2:453. 1833. V. fruticulosa Mart. ex DC., Prodr. 5:53. 1836. V. eremophila Mart. ex DC., Prodr. 5:54. 1836. V. tri- cephala Gardner, London J. Bot. 5:223. 1846. V. resinosa Gardner, London J. Bot. 6:419. 1847. V. saxicola Sch.Bip. ex Ba- ker in Mart., Fl. Bras. 6(2):81. 1873. ?V. oreophila Malme, Kongl. Svenska Veten- skapsakad Handl. N.S. 32(5):26. 1899. Brazil (Bahia, D.F., Goias, Mato Grosso, Minas Gerais). Pollen D, Figs. 48-51. A photograph of the type shows that the older St. Hilaire name applies to the same common species usually known as Vernonia fruticulosa Mart. ex DC. The leaves seem to vary in shape from broadly elliptical to linear. A type photograph of V. oreophila Malme, of Mato Grosso, seems to represent the same or a closely related species. The leaves of the latter are exclusively ovate and the inflorescences contain very few heads. Lepidaploa sagraeana (DC.) H. Robinson, comb. nov. Vernonia sagraeana DC., Prodr. 5:55. 1836. V. valenzuelana A. Rich. in Sagra, Hist. Fis. Pol. Nat. Cuba. Bot. 11:33. 1850. V. inaequiserrata Sch.Bip., J. Bot. 1:131. 1863. V. fallax Gleason, Bull. Torrey Bot. Club 40:324. 1913. V. aceratoides Glea- son, Bull. Torrey Bot. Club 40:325. 1913. V. angusticeps Ekman, Ark. Bot. 13(15): 14.1914. V. linguaefolia Ekman, Ark. Bot. 13(15):19. 1914. V. reedii Ekman & Ur- ban, Repert. Spec. Nov. Regni Veg. 26: 97. 1929, not V. reedii Daniels, 1907. V. potrerillona Ekman & Urban, Repert. Spec. Nov. Regni Veg. 26:98. 1929. Cuba. Pollen C/G. Lepidaploa salzmannii (DC.) H. Robinson, comb. nov. Vernonia salzmannii DC., Prodr. 5:55. 1836. V. poeppigiana DC., Prodr. 5:55. 1836, non V. poeppigiana DC., Prodr. 5: 20. 1836. V. argyropappa H. Buek., Ind. Gen. Sp. Syn. in DC., Prodr. 2: Praef. v. 1840. V. miersiana Gardner, London J. Bot. 4:115. 1845. V. virens Sch.Bip. ex Baker in Mart., Fl. Bras. 6(2):71. 1873. V. velutina Hieron., Bot. Jahrb. Syst. 22: 697.1897. V. herbertii Cuatr., Bot. Jahrb. Syst. 77:55. 1956. Central America, Co- lombia, Peru, Brazil. Pollen C, Figs. 13- 16. Lepidaploa sclareaefolia (Sch.Bip.) H. Robinson, comb. nov. Vernonia sclareaefolia Sch.Bip., Linnaea 30: 170. 1859. Colombia. Pollen G. Lepidaploa segregata (Gleason) H. Robinson, comb. nov. Vernonia segregata Gleason, Bull. Torrey Bot. Club 40:327. 1913. Cuba. Pollen C. Lepidaploa sericea (L. C. Rich.) H. Robinson, comb. nov. Vernonia sericea L. C. Rich., Actes Soc. Hist. Nat. Paris 1:112. 1792. Lepidaploa phyl- lostachya Cassini, Dict. Sci. Nat. 26:16. 1823, nom. inval., prior to validation of Lepidaploa at generic level. V. berteriana DC., Prodr. 5:52. 1836. Conyza portori- censis Bertero ex DC., Prodr. 5:52. 1836, nom. nud. Vernonia racemosa Delponte, Mem. Reale Acad. Sci. Torino. II, 14: 396. 1854. V. venusta Gleason, Bull. New York Bot. Gard. 4:177. 1906. V. phyllos- tachya Gleason, Bull. New York Bot. Gard. 4:181. 1906. V. angustissima Wright ex Ekman, Ark. Bot. 13(15):78. 1914. V. maestralis Ekman ex Urban, Re- pert. Spec. Nov. Regni Veg. 26:99. 1929. Greater Antilles, Virgin Isl. Pollen C, Figs. 4-6; N = 17 (Keeley 1978). Lepidaploa silvae (H. Robinson) H. Robinson, comb. nov. Vernonia silvae H. Robinson, Phytologia 46: 112, 1980. Brazil (Para). Pollen C. VOLUME 103, NUMBER 2 Lepidaploa solomonii H. Robinson, sp. nov. Plantae suffrutescentes ad 2 m altae. Caule virides striati sparse minute antrorse stri- gulosi. Folia alterna, petiolis 1.5—2.5 cm longis; laminae ovato-ellipticae 14-16 cm longae et ad 6.5 cm latae base breviter acu- tae et minime acuminatae margine integrae vel remote minime crenulatae apice brevi- ter anguste acuminatae supra et subtus subglabrae sparse minute strigulosae supra sparsius. Inflorescentiae supra folia vege- tativa laxe ramosae seriate cymosae, brac- teae foliiformes distincte minores breviter petiolatae petiolis 3-7 mm longis, laminis lanceolatis 3.0-8.5 cm longis et 0.4—2.4 cm latis in ramulis ultimis subnullis. Capitula sessilia solitaria late campanulata ca. 10 cm alta; squamae involucri in partibus atro- purpurascentes ca. 45 subimbricatae 4—5- seriatae graduatae exteriores lanceolatae 2— 5 mm longae et 0.8—1.0 mm latae apice sub- aristatae interiores oblong-lineares ad 8 mm longae apice acutae margine omnino mi- nute fimbriatae extus subglabrae vel per- minute puberulae. Flores ca. 30 in capitulo. Corollae ca. 7 mm longae extus plerumque glabrae in partibus apicalibus lobarum breviter spiculiferae, tubis cylindricis 3 mm longis, faucibus late infundibularibus 1.0- 1.3 mm longis, lobis linearibus ca. 3 mm longis et 0.6—0.7 mm latis; thecae anthera- rum ca. 2.5 mm longae, appendicibus api- calibus ca. 0.4 mm longae glabrae; basi sty- lorum abrupte disciformes. Achaenia ca. 3 mm longa dense sericeo-setulifera non glan- dulifera et non idioblastifera; setae pappi sordide ca. 30 ca. 6 mm longae subdeciduae superne vix latiores in sereivus exterioribus anguste lanceolatae 1.0-1.5 mm longae. Grana pollinis in diametro ca. 45 um valde lophata, reticulis in typo G. (Fig. 62). Type.—Bolivia: La Paz: Prov. Murillo, 44.0 km below Lago Zongo dam, vicinity of Cahua hydroelectric plant, 16°03’S, 68°01'W, 1200 m, moist forest, disturbed, alternating with Chacos and secondary for- est, shrub, 2 m, 12-15 Sep 1983, J. C. Sol- omon 10780 (holotype US; isotype MO). 493 The species resembles Lepidaploa sordi- dopapposa in the dark involucre and the brownish color of the pappus, but it does not seem closely related. The species is dis- tinctive in the subglabrous aspect of the stems and leaves, the narrowly petiolate vegetative leaves, and the lax, seriate cymes with distinct, foliose bracts. Lepidaploa sordidopapposa (Hieron.) H. Robinson, comb. nov. Vernonia sordidopapposa Hieron., Bot. Jahrb. Syst. 22:697. 1897. Bolivia, Ec- uador, Peru. Pollen G. The species is extended to include nu- merous specimens that have recently been annotated as Vernonia mapirensis by Jones for his study of the tribe in Peru (Jones 1980). The present species seems to be character- ized by the often crowded heads on short- ened inflorescence branches, by the invo- lucre remaining nearly cylindrical when dry, and by the dark pappus. The specimens show a great variation in length of the leaf pu- bescence and density of involucral pubes- cence. Lepidaploa sororia (DC.) H. Robinson, comb. nov. Vernonia sororia DC., Prodr. 5:40. 1836. V. coulonii Sch.Bip. ex Baker in Mart., FI. Bras. 6(2):93. 1873. Brazil (Espirito San- to, Rio de Janeiro). Pollen G. As indicated by Robinson (1987c), the DeCandolle species is not a synonym of Cyrtocymura scorpioides, but is an older name for the species that has been known as Vernonia coulonii. Lepidaploa stenophylla (Less.) H. Robinson, comb. nov. Vernonia stenophylla Less., Linnaea 6:667. 1831. V. corallophila Gleason, Bull. Tor- rey Bot. Club 40:309. 1913. V. nemato- phylla Ekman & Urban, Repert. Spec. Nov. Regni Veg. 26:100. 1929. Cuba, Hispaniola. Pollen C. 494 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ “Fieldwork supported by the National Science Foundation” BOLIVIA COMPOSITAE ; ip Lepidaplos selletaasaii Dil oelntaes Mololype Dept. La Pazr Prov. Murillo. 44.0 km below Lago Zongo damr vicinity of Cahua hydroelectric plant. 16°03'S, 68°01'W, elev. 1200 m. Moist forests disturbedr alternating with Chacos and secondary forest. UNITED STATES Shrub, 2 m. 3149580 12-15 Sept. 1983 NATIONAL HERBARIUM J.C. Solomon 10780 MISSOURI BOTANICAL GARDEN HERBARIUM (MO) Fig. 62. Holotype of Lepidaploa solomonii H. Robinson, Solomon 10780 (US). VOLUME 103, NUMBER 2 Lepidaploa subsquarrosa (DC.) H. Robinson, comb. nov. Chysocoma paniculata Vell., Fl. Flumin. 327. 1825, non C. paniculata [Walter] Gmel. 1792. Vernonia subsquarrosa DC.., Prodr. 5:41. 1836. V. albiflora Gardner, London J. Bot. 6:224. 1847. Brazil (Es- pirito Santo, Rio de Janeiro, Sao Paulo). Pollen G, Figs. 28-31. The species has usually been recognized under the invalid name Vernonia geminata Less. Lessing (1829) was not naming a new species but was erroneously identifying Bra- zilian material as the Colombian V. gemi- nata H.B.K. (=Lepidaploa canescens). Lat- er attempts (DeCandolle 1836, Baker 1873) to use the name sensu Lessing (exclusively in the atypical sense) were totally invalid. The oldest acceptable name for the species is here raised from synonymy. Lepidaploa tarijensis (Griseb.) H. Robinson, comb. nov. Vernonia sericea var. tarijensis Griseb., Abh. Konigl. Ges. Wiss. Gottingen 24:163. 1879. V. tarijensis (Griseb.) Hieron., Bot. Jahrb. Syst. 22:682. 1897. Argentina, Bo- livia. Pollen G. Lepidaploa tenella (D. Nash) H. Robinson, comb. nov. Vernonia tenella D. Nash, Fieldiana, Bot. 36:74. 1974. Guatemala. Pollen C/G. Lepidaploa tombadorensis (H. Robinson) H. Robinson, comb. nov. Vernonia tombadorensis H. Robinson, Phy- tologia 45:187. 1980. Brazil (Bahia). Pol- len D, Figs. 40-42. Lepidaploa tortuosa (L.) H. Robinson, comb. nov. Conyza tortuosa L., Sp. Pl. 862. 1753. C. scandens Mill., Gard. Dict. ed. 8. Conyza no. 11. 1768. Vernonia schiedeana Less., Linnaea 6:399. 1831. V. seemaniana 495 Steetz, Bot. Voy. Herald 139. 1854. V. tortuosa (L.) Blake, Proc. Biol. Soc. Wash. 39:144. 1926. Mexico (Chiapas, Oaxaca, Veracruz), Guatemala, Belize, Honduras, El Salvador, Costa Rica, Panama. Pol- len G. Lepidaploa tovarensis (Gleason) H. Robinson, comb. nov. Vernonia tovarensis Gleason, Amer. J. Bot. 19:753. 1932. Venezuela. Pollen Aynia- type, Figs. 19-24. Lepidaploa trichoclada (Gleason) H. Robinson, comb. nov. Vernonia trichoclada Gleason, Bull. Torrey Bot. Club 52:184. 1925. Peru. Pollen G. This central Peruvian species is resur- rected from the synonymy of the Bolivian and southern Peruvian Lepidaploa mapi- rensis, from which it differs in the larger size of the heads, the more erect to retrorse pu- bescence, and the more oblong leaves with widely separated veins. Lepidaploa tricho- clada is like the more recently described L. retrosetosa of southern Peru in the numer- ous outer involucral bracts that are as long as the inner bracts. But the latter species has ovate, remotely denticulate leaves with rounded bases and large, foliose bracts in the inflorescence. Lepidaploa trichoclada generally resembles the Bolivian L. tristis, but the latter has shorter outer involucral bracts and has the heads often paired at the nodes. Lepidaploa trilectorum (Gleason) H. Robinson, comb. nov. Vernonia trilectorum Gleason, Bull. Torrey Bot. Club 52:186. 1925. Colombia. Pol- len G. Lepidaploa trinitatis (Ekman) H. Robinson, comb. nov. Vernonia trinitatis Ekman, Ark. Bot. 13(15): 39. 1914. Trinidad, Venezuela. Pollen C; N = 51 ,, (Keeley 1978). 496 Lepidaploa tristis (Hieron.) H. Robinson, comb. nov. Vernonia tristis Hieron., Bot. Jahrb. Syst. 22:683. 1897. Bolivia. Pollen G. Lepidaploa uniflora (Miller) H. Robinson, comb. nov. Conyza uniflora Miller, Gard. Dict. ed. 8. Conyza no. 13. 1768. Vernonia ctenopho- ra Gleason, Bull. Torrey Bot. Club 46: 243. 1919. Mexico (Campeche), Guate- mala, Belize. Pollen C. Lepidaploa urbaniana (Ekman ex Urban) H. Robinson, comb. nov. Vernonia urbaniana Ekman ex Urban, Re- pert. Spec. Nov. Regni Veg. 26:99. 1929. Cuba. Pollen C. Lepidaploa verticillata (Proctor ex Adams) H. Robinson, comb. nov. Vernonia verticillata Proctor ex Adams, Phytologia 21:409. 1971. Jamaica. Pollen C; N = 17 (Keeley 1978). Lepidaploa viminalis (Gleason) H. Robinson, comb. nov. Vernonia viminalis Gleason, Bull. New York Bot. Gard. 4:184. 1906. Cuba. Pollen C/G. Lepidaploa violiceps (H. Robinson) H. Robinson, comb. nov. Vernonia violiceps H. Robinson, Phytologia 45:160. 1980. Ecuador. Pollen C. Lepidaploa virentiformis (Malme) H. Robinson, comb. nov. Vernonia virentiformis Malme, Ark. Bot. 24A(8):8. 1932. Bolivia (Beni), Brazil (Mato Grosso). Pollen C. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lepidaploa wrightii (Sch.Bip.) H. Robinson, comb. nov. Vernonia wrightii Sch.Bip., J. Bot. 1:234. 1863. Cuba. Pollen C/G. Lepidaploa yunquensis (Gleason) H. Robinson, comb. nov. Vernonia yunquensis Gleason, Bull. New York Bot. Gard. 4:191. 1906. Cuba. Acknowledgments The pollen specimens and photographic prints were prepared by Brian Kahn, Mary Sangrey, and Barbara Eastwood using fa- cilities of the Botany Department Palyno- logical Laboratory and darkroom. The pho- tographs were taken by Suzanne Braden and Brian Kahn of the National Museum of Natural History SEM Laboratory using a Hitachi S-570 scanning electron micro- scope. Photographs of holotypes were pre- pared by Victor E. Krantz, Staff Photogra- pher, National Museum of Natural History. Some prints were also prepared by Sherry Pittam and Suzanne Fredericq at the De- partment of Botany. The editorial efforts of Dr. David Lellinger of the Department of Botany are greatly appreciated. Literature Cited Aristeguieta, L. 1964. Compositae. Jn T. Lasser, ed., Flora de Venezuela 10:i—xiu1, 1-941, indice 1- 16. Badillo, V. M. 1982. Vernonia marguana Cuatr., nueva para Venezuela.—Ernstia 8:7. 1984. Sobre Vernonia gracilis HBK. y V. moritziana Sch.Bip.—Ernstia 23:12-13. Baker, J. B. 1873. Compositae. I. Vernoniaceae. In C. F. P. Martius, ed., Flora Brasiliensis 6(2):1— 180. Bartling, F. G. 1830. Ordines naturales plantarum. Dieterich, Gottingen, 498 pp. Bentham, G., & J. D. Hooker. 1873. Compositae. In Genera Plantarum 2(1):163-533. Cabrera, A. L. 1944. Vernonieas Argentinas (Com- positae).— Darwiniana 6:265-379. Cassini, H. 1817a. Apercu des genres nouveaux formés par M. Henri Cassini dans la famille des Syn- VOLUME 103, NUMBER 2 anthérées. Troisiéme fascicule.—Bulletin des Sciences de La Société Philomatique de Paris 1817:31-34. 1817b. Apercu des genres nouveaux formés par M. Henri Cassini dans la famille des Syn- anthérées. Quatriéme fascicule.—Bulletin des Sciences de La Société Philomatique de Paris. 1817:66-70. 1823. Lépidaple, Lepidaploa. Pp. 16-24 in G. Cuvier, ed., Dictioniare des Sciences Natu- relles 26. Levrault Strassbourg. [Reprinted in R. M. King & H. W. Dawson, eds., 1975, Cassini on Compositae, 3 volumes. Oriole Editions, New York.] . 1825. Oliganthe, Oliganthes. Pp. 18-20 in G. Cuvier, ed., Dictioniare des Sciences Naturelles 36. Levrault Strassbourg. [Reprinted in R. M. King & H. W. Dawson, eds., 1975, Cassini on Compositae, 3 volumes. Oriole Editions, New York.] 1830. Zyégée, Zoegea. Pp. 560-619 in G. Cuvier, ed., Dictioniare des Sciences Naturelles 60. Levrault Strassbourg. [Reprinted in R. M. King & H. W. Dawson, eds., 1975, Cassini on Compositae, 3 volumes. Oriole Editions, New York.] DeCandolle, A. P. 1836. Prodromus systematis na- turalis regni vegetabilis, vol. 5. Treuttel & Wirtz, Paris, 706 pp. Gleason, H. A. 1906. A revision of the North Amer- ican Vernonieae. — Bulletin New York Botanical Garden 4:144—243. 1922. Vernonieae.—North American Flora 33(1):47-110. 1923. Evolution and geographical distribu- tion of the genus Vernonia in North America. — American Journal of Botany 10:187—202. Humboldt, F. H. A. von, A. J. Bonpland, & C. S. Kunth. 1818. Compositae. In Nov. Gen. Sp., folio edition, vol. 4:1—246, pl. 301-412. Jeffrey, C. 1988. The Vernonieae in East Tropical Africa.—Kew Bulletin 43:195-277. Jones, S. B. 1977. Vernonieae: Systematic review. Pp. 503-447 in V. H. Heywood, J. B. Harborne, & B. L. Turner, eds., The biology and chemistry of the Compositae. Academic Press, London, 1189 pp. . 1979a. Chromosome numbers of Vernonieae (Compositae). — Bulletin of the Torrey Botanical Club 106:79-84. 1979b. Synopsis and pollen morphology of Vernonia (Compositae: Vernonieae) in the New World.—Rhodora 81:425—447. 1980. The Vernonieae. Jn J. F. Macbride, Flora of Peru.—Fieldiana, Botany n.s. 5:22—73. . 1981. Synoptic classification and pollen mor- 497 phology of Vernonia (Compositae: Vernonieae) in the Old World.—Rhodora 83:59-75. Keeley, S. C. 1978. A revision of the West Indian vernonias (Compositae).—Journal of the Ar- nold Arboretum 59:360-413. 1982. Morphological variation and species recognition in the Neotropical taxon Vernonia arborescens Compositae.—Systematic Botany 7:71-84. Kingham, D. L. 1976. A study of the pollen mor- phology of tropical African and certain other Vernonieae (Compositae). — Kew Bulletin 31:9- 26. Lessing, C. F. 1829. De Synanthereis herbarii regii berolinensis dissertatio prima.—Linnaea 4:240- 288, 290-356, pl. II. Reichenbach, H.G. L. 1828. Conspectus regni vege- tabilis. Cnobloch, Leipzig, 294 pp. Robinson, H. 1980. New species of the Vernonieae (Asteraceae). V. Additions to Vernonia from Brasil.— Phytologia 45:166—208. 1987a. Studies of the Lepidaploa complex (Vernonieae: Asteraceae). I. The genus Steno- cephalum Sch. Bip.— Proceedings of the Biolog- ical Society of Washington 100:578—583. 1987b. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). IJ. A new genus, Echinocoryne.—Proceedings of the Biological Society of Washington 100:584—-589. 1987c. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). III. Two new genera, Cyrtocymura and Eirmocephala. — Proceedings of the Biological Society of Washington 100: 844-855. 1988a. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). IV. The new genus, Lessingianthus.— Proceedings of the Biological Society of Washington 101:929-951. 1988b. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). V. The new genus, Chrysolaena.—Proceedings of the Biological Society of Washington 101:952-958. 1988c. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). VI. A new genus, Ay- nia. — Proceedings of the Biological Society of Washington 101:959-965. . 1989. Acilepidopsis, a new genus of Vernoni- eae from South America (Asteraceae).— Phy- tologia 67:289-292. , F. Bohlman, & R. M. King. 1980. Chemo- systematic notes on the Asteraceae. III. Natural subdivision of the Vernonieae.— Phytologia 46: 421-436. —., & V.A. Funk. 1987. A phylogenetic analysis of Leiboldia, Lepidonia, and a new genus Stra- mentopappus (Vernonieae: Asteraceae).—Bo- 498 tanische Jahrbiicher fiir Systematik, Pflanzen- geschichte und Pflanzengeographie 108:213-228. , & B. Kahn. 1986. Trinervate leaves, yellow flowers, tailed anthers, and pollen variation in Distephanus Cassini (Vernonieae: Astera- ceae). — Proceedings of the Biological Society of Washington 99:493-501. ——., & R.M. King. 1979. Mattfeldanthus muti- sioides gen. et spec. nov. (Asteraceae: Vernonie- ae) from Bahia, Brazil.— Willdenowia 9:9-12. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Spach, E. 1841. Histoire naturelle des végetaux, vol. 10. Phanaérogames. Roret, Paris, 572 pp. Stix, E. 1960. Pollenmorphologische untersuchungen an Compositen.—Grana Palynologica 2(2):41- 104, pl. 1-21. Department of Botany, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE Applications published in the Bulletin of Zoological Nomenclature The following applications were published on 19 December 1989 in Vol. 46, Part 4 of the Bulletin of Zoological Nomenclature. Comment or advice on these appli- cations is invited for publication in the Bulletin and should be sent to the Executive Secretary, I.C.Z.N., % The Natural History Museum, Cromwell Road, London SW7 SBD, U.K. Case No. 2683 Gryphaea pitcheri Morton, 1834 (currently Texigryphaea pitcheri, Mollusca, Bivalvia): proposed conservation. 2554 Myriochele Malmgren, 1867 and Myriochele oculata Zaks, 1923 (Annelida, Polychaeta): proposed conservation. 2637 Buthus vittatus (currently Centruroides vittatus,; Arachnida, Scorpionida): proposed recognition of Wood (1863) as author of the specific name and designation of a neotype, and Centrurus hentzi (currently Cen- truroides hentzi) Banks, 1904: proposed conservation of the specific name. Shoemakerella Pirlot, 1936 (Crustacea, Amphipoda): proposed designation of Lysianax cubensis Stebbing, 1897 as the type species. Corisa verticalis Fieber, 1851 (currently Trichocorixa verticalis; Insecta, Het- eroptera): proposed conservation of the specific name. Curculio viridicollis Fabricius, 1792 (currently Phyllobius viridicollis; Insecta, Coleoptera): proposed conservation of the specific name, and Rhyn- colus Germar, 1817: proposed designation of Curculio ater Lin- naeus, 1758 as the type species. Ochthebius Leach, 1815 (Insecta, Coleoptera): proposed conservation of E/o- phorus marinus Paykull, 1798 as the type species. Culex stigmatosoma Dyar, 1907 and C. thriambus Dyar, 1921 (Insecta, Diptera): proposed conservation of the specific names by the suppression of C. peus Speiser, 1904. Exoprosopa Macquardt, 1840 (Insecta, Diptera): proposed confirmation of Anthrax pandora Fabricius, 1805 as the type species. Musca heraclei Linnaeus, 1758 (currently Euleia heraclei; Insecta, Diptera): proposed conservation of heraclei as the correct spelling of the spe- cific name. Callionymus pusillus Delaroche, 1809 (Osteichthyes, Perciformes): proposed conservation of the specific name. Muraena Linnaeus, 1758 (Osteichthyes, Anguilliformes): proposed confir- mation of Muraena helena Linnaeus, 1758 as the type species, so conserving Anguilla Shaw, 1803. Haplocanthosaurus Hatcher, 1903 (Reptilia, Saurischia): proposed conser- vation. Atheris Cope, 1862 (Reptilia, Serpentes): proposed conservation, and pro- posed confirmation of Vipera chlorechis Pel, [1851] as the valid name of the type species. Bur - OE a Pat ey eptiniyt siannt - OTE oF bie ae rhin y atte “eonener a i; i palizie ine Ont 448 un un a aaa te. aouisn sein ape san ile 16 of deme ma gma en bern il INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (botany, zoology, and paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with a summary in an alternate language when appropriate. Submission of manuscripts. —Submit manuscripts to the Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History NHB Stop 108, Smithsonian Institution, Washington, D.C. 20560. 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Clarity of presentation, and requirements of taxonomic and nomenclatural procedures ne- cessitate reasonable consistency in the organization of papers. Telegraphic style is required for descriptions and diagnoses. Literature citations in the text should be in abbreviated style (author, date, page), except in botanical synonymies, with unabbreviated citations of journals and books in the Literature Cited sections. Direct quotations in the text must be accompanied by author, date, and pagination. The establishment of new taxa must conform with the requirements of the appropriate international codes of nomenclature. When appropriate, accounts of new taxa must cite a type specimen deposited in an institutional collection. Examples of journal and book citations: Eigenmann, C. H. 1915. The Cheirodontidae, a subfamily of minute characid fishes of South America.— Memoirs of the Carnegie Museum 7(1):1—99. Ridgely, R. S. 1976. A guide to the birds of Panama. Princeton, New Jersey, Princeton University Press, 354 pp. Olson, S. L. 1973. The fossil record of birds. Pp. 79-238 in D. Farner, J. King, and K. Parkes, eds., Avian biology, volume 8. Academic Press, New York. Figures and tables with their legends and headings should be self-explanatory, not requiring reference to the text. Indicate figure and table placement in pencil in the margin of the manu- script. Plan illustrations in proportions that will efficiently use space on the type bed of the Proceedings. Original illustrations should not exceed 15 x 24 inches. Figures requiring solid black backgrounds should be indicated as such when the manuscript is submitted, but should not be masked. CONTENTS Porites colonensis, new species of stony coral (Anthozoa: Scleractinia) off the Caribbean coast of Panama Vassil N. Zlatarski Solenotheca, new hyolitha (Mollusca) from the Ordovician of North America John M. Malinky A new Pyrgulopsis (Gastropoda: Hydrobiidae) from southeastern California, with a model for historical development of the Death Valley Hydrographic System Robert Hershler and William L. Pratt Some tardigrades from Colorado, with a description of a new species of Macrobiotus (Macro- biotidae: Eutardigrada) R. Deedee Kathman Pycnogonida of the Western Pacific Islands, VII. On some rare species from the Flores Sea, Indonesia K. Nakamura and C. Allan Child Pycnogonida of the Western Pacific Islands, VIII. Recent collections from islands of the Great Barrier Reef, Australia C. Allan Child Canthocamptus (Elaphoidella) striblingi, new species (Copepoda: Harpacticoida) from Costa Rica Janet W. Reid Tachidius incisipes Klie and other harpacticoids from northwestern Canada (Crustacea: Co- pepoda) Edward B. Reed The occurrence of Sphaeroma serratum (Fabricius, 1787) in the western South Atlantic (Crus- tacea: Isopoda) Ana Marta Roux and Ricardo Bastida The status of the caridean shrimp Pandalina modesta (Bate, 1888) (Crustacea: Decapoda: Pandalidae) with redescription of the species E. Macpherson Galathea coralliophilus, a new decapod crustacean (Anomura: Galatheidae) from Singapore, Gulf of Thailand, and West Irian Keiji Baba and Sang-Chul Oh Calyptraeotheres, a new genus of Pinnotheridae for the limpet crab Fabia granti Glassell, 1933 (Crustacea: Brachyura) Ernesto Campos Mesacturus dicrurus, new species, an unusual stomatopod from Micronesia (Stomatopoda: Gonodactylidae) Roy K. Kropp and Jane H. Dominguez The primary types of the Richard M. Bohart Museum of Entomology, I. Tardigrada R. O. Schuster, T. L. Tyler, J. A. Skinner, and E. A. Sugden Two new species of Mimagoniates (Teleostei: Characidae: Glandulocaudinae), their phylogeny and biogeography and a key to the glandulocaudin fishes of Brazil and Paraguay Naércio A. Menezes and Stanley H. Weitzman The tadpole of a dart-poison frog Phyllobates lugubris (Anura: Dendrobatidae) Maureen A. Donnelly, Craig Guyer, and Rafael O. de Sa Taxa of North American birds described from 1957 to 1987 M. Ralph Browning Erroneous emendations to names proposed by Hekstra (Strigidae: Otus) M. Ralph Browning Subspecies of Stenella longirostris (Mammalia: Cetacea: Delphinidae) William F. Perrin Studies in the Lepidaploa complex (Vernonieae: Asteraceae). VII. The genus Lepidaploa Harold Robinson International Commission on Zoological Nomenclature: Applications Di 265 a PRC OC FE EDINGS POPTHE = =. 7 BIOL OGICAL SOCIETY WAS H INGTON _ THE BIOLOGICAL SOCIETY OF WASHINGTON 1990-1991 Officers President: Leslie W. Knapp Secretary: G. David Johnson President-elect: Storrs L. Olson 2 Treasurer: Michael Vecchione Elected Council Roger F. Cressey, Jr. Janet W. Reid Janet R. Gomon Wayne C. Starnes Robert Hershler Jeffery T. Williams Custodian of Publications: Austin B. Williams PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: George C. Steyskal Invertebrates: Stephen D. Cairns Frank D. Ferrari Plants: David B. Lellinger Rafael Lemaitre Insects: Wayne N. Mathis Vertebrates: G. David Johnson Membership in the Society 1s open to anyone who wishes to join. There are no prerequisites. Annual dues of $15.00 ($20.00 to non-USA addresses) include subscription to the Proceedings of the Biological Society of Washington. Library subscriptions to the Proceedings are: $25.00 within the U.S.A., $30.00 elsewhere. The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued sporadically) are available. Correspondence dealing with membership and subscriptions should be sent to The Treasurer, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, National Museum of Natural History, Washington, D.C. 20560. THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 501-508 COMMENTS ON THE NOMENCLATURE OF SOME NEOTROPICAL BATS (MAMMALIA: CHIROPTERA) Alfred L. Gardner and Carolyn S. Ferrell Abstract.— We examine four nomenclatural problems concerning Neotrop- ical bats and conclude that Platyrrhinus has priority over Vampyrops; An- thorhina is a junior objective synonym of Tonatia; and Cabrera, as first reviser, selected the spelling Molossus barnesi over M. burnesi. We recommend that Wied-Neuwied, not Oken, be considered the author of Diclidurus albus, a ae af kk k& kk kkk OX @MoO * * O O OCO x*e Oo O O x® ce) % * 0 80.0 70.0 40.0 50.0 60.0 STANDARD LENGTH (mm) Fig. 3. Number of anal photophores plotted against standard length for 22 Polyipnus species. All stars = P. spinosus complex; open stars = P. fraseri; circles = P. asteroides and P. laternatus complexes combined (i.e., all other species). absence or reduced numbers in P. fraseri is a result of some process producing paedo- morphic development. The occurrence of species exhibiting pae- domorphosis has been noted for a diverse assemblage of fishes (Fink 1981, Weitzman & Vari 1988), including the deep-sea fam- ilies Gonostomatidae (Ahlstrom et al. 1984), Stomiidae (Fink 1985), and Sternoptychi- dae (Weitzman 1974). The existence of species or higher taxa with juvenile-like fea- tures in adults can be a source of difficulty in systematics. It is now generally accepted that much of observed morphological di- versity, including otherwise perplexing morphologies, may result through hetero- chronic alteration of developmental pro- grams. A framework for interpretation of such cases has been emerging over the past decade (see Alberch et al. 1979, Fink 1982, Kluge 1988). Figure 3, which compares rel- ative growth of structures among related taxa, can be interpreted as a set of ontoge- netic trajectories. Due to unavailability of a growth series of P. fraseri it is not possible to compare slopes with other species or to plot sizes at which various events take place. It is clear, however, that rate of photophore addition to the anal cluster is probably low- er than in other species. In a form that reaches a body size comparable to possible sister taxa and Polyipnus outgroups (P. as- teroides Schultz, 1938 and P. /aternatus Garman, 1899 complexes), such develop- ment is considered neotenic. Alternatively, the presence of four anal photophores could be the result of late onset of growth with the rate of addition the same as in other species (i.e., postdisplacement). The supra-anal photophore cluster is absent in the two 514 known specimens but the possibility re- mains that P. fraseri reaches a much larger body size and has unusually protracted de- velopment of AN and SAN photophores. In either case, when compared with all known possible immediate outgroups, it appears probable that some process has affected the general ontogenetic program of photophore development. Without material represent- ing the entire size range of P. fraseri it is not possible to comment on rate of appearance of the other groups of luminescent organs, or to say which of the possible heterochron- ic explanations is the more likely. Since P. fraseri is a member of the P. spinosus com- plex, then the distribution of ontogenetic photophore characters (absence of SAN, low AN number) is such that their states in P. fraseri are most parsimoniously interpreted through outgroup comparison as apo- morphic. Qualitatively, the body shape (e.g., large orbit; narrow caudal peduncle) of P. fraseri 1s very similar to juveniles of other species of the P. spinosus complex and the out- groups. The possibility remains that the known specimens of P. fraseri are juveniles ofa large species. However, this explanation requires very early development of photo- phore-scale denticles relative to the ap- pearance of typically adult photophore fea- tures. Acknowledgments I’m grateful to A. P. Andriashev, K. E. Hartel, S. L. Jewett, K. Kawaguchi, H. Ki- shimoto, K. Matsuura, M. McGrouther, N. R. Merrett, J. Nielsen, O. Okamura, N. V. Parin, J. R. Paxton, S. J. Raredon, C. B. Renaud, H. J. Walker, S. H. Weitzman, and J. Williams who provided study material. I thank H. Griffith, G. D. Johnson, R. D. Mooi, R. J. Mooi, and S. H. Weitzman whose comments led to an improved manu- script. This research was supported by NSERC (Natural Sciences and Engineering Research Council of Canada) grant #330068 to R. L. Haedrich. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Ahlstrom, E. H., W. J. Richards, & S. H. Weitzman. 1984. Families Gonostomatidae, Sternopty- chidae, and associated stomiiform groups: de- velopment and relationships. Pp. 184-198 in H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall, Jr., and S. L. Richardson, eds., Ontogeny and systematics of fishes. Special Publication Number 1, American Society of Ichthyologists and Herpetologists, 760 pp. Alberch, P., S. J. Gould, G. F. Oster, & D. B. Wake. 1979. Size and shape in ontogeny and phylog- eny.— Paleobiology 5(3):296-317. Baird, R.C. 1971. The systematics, distribution, and zoogeography of the marine hatchetfishes (fam- ily Sternoptychidae).— Bulletin of the Museum of Comparative Zoology, Harvard University 142(1):1-128. Borodulina, O.D. 1979. Composition of the “Polyip- nus spinosus complex” (Sternoptychidae, Os- teichthyes) with a description of 3 new species of the group.—Journal of Ichthyology 19(2):1— 10. Fink, W. L. 1981. Ontogeny and phylogeny of tooth attachment modes in actinopterygian fishes. — Journal of Morphology 167:167-184. 1982. The conceptual relationship between ontogeny and phylogeny.— Paleobiology 8(3): 254-264. 1985. Phylogenetic relationships of the sto- miid fishes (Teleostei: Stomiiformes). Miscel- laneous Publications of the Museum of Zoology, University of Michigan, No. 171, 127 pp. Fowler, H. G. 1934. Descriptions of new fishes ob- tained 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. — Academy of Natural Sciences of Philadelphia, Proceedings 85:233- 437. Fujii, E. 1984. Family Sternoptychidae. Pp. 47-48, pl. 49L in H. Masuda, K. Amaoka, C. Araga, T. Uyeno, and T. Yoshino, eds., Fishes of the Japanese Archipelago. Tokai University Press, Tokyo, 433 pp., 370 pl. Harold, A. S. 1989. A new species of Polyipnus (Sto- miuiformes: Sternoptychidae) from the Coral Sea, with-a revised key to the species of the P. spi- nosus complex.—Copeia 1989(4):871-876. Hubbs, C. L., & K. F. Lagler. 1958. Fishes of the Great Lakes region. The University of Michigan Press, Ann Arbor, 213 pp. Kluge, A.G. 1988. The characterization of ontogeny. Pp. 57-81 in C. J. Humphries, ed., Ontogeny and systematics. Columbia University Press, New York, 236 pp. Leviton, A. E., R. H. Gibbs, Jr., H. Heal, & C. E. Dawson. 1985. Standards in ichthyology and herpetology: Part I. Standard symbolic codes for VOLUME 103, NUMBER 3 institutional resource collections in herpetology and ichthyology.—Copeia 1985(3):802-832. Schultz, L. P. 1961. Revision of the marine silver hatchetfishes (family Sternoptychidae).—Pro- ceedings of the United States National Museum 112(3449):587-649. . 1964. Family Sternoptychidae. Pp. 241-273 in H. B. Bigelow et al., eds., Fishes of the West- ern North Atlantic, Part 4, Isospondyli. Sears Foundation for Marine Research Memoir No. 1, 599 pp. Weitzman, S. H. 1974. Osteology and evolutionary relationships of the Sternoptychidae, with a new classification of stomiatoid families. — Bulletin of the American Museum of Natural History 53(3):327-478. —,&R.P. Vari. 1988. Miniaturization in South American freshwater fishes: an overview and discussion.— Proceedings of the Biological So- ciety of Washington 101:444-465. Yamamoto, E. 1982. Sternoptychidae. P. 327 in O. Okamura, K. Amaoko, and F. Mitani, eds., Fishes of the Kyushu-Palau Ridge and Tosa Bay. Japan Fisheries Resource Conservation Asso- ciation, Tokyo, 435 pp. Department of Biology, Memorial Uni- versity of Newfoundland, St. John’s, New- foundland, Canada, A1B 3X9. Appendix Comparative material examined.—P. aquavitus Baird, 1971: USNM 298928, 1 (17.5 mm), USNM 298927, 1 (15.4 mm), USNM 298925, 10 (10.6-17.9 mm), AMS 1.19761-029, 10 (10.2-34.1 mm), AMS I.16492-008, 3 (12.0-14.9 mm), AMS I.20310-017, 3 (17.6—20.7 mm), AMS I.19762-002, 3 (21.2-34.6 mm), AMS 1.20316-005, 3 (20.2—36.1 mm); P. asteroides Schultz, 1938: MCZ 66695, 1 (41.9 mm), MCZ 66696, | (30.9 mm); P. elongatus Borodulina, 1979: AMS I.21975- Dis 007, 2 (54.6-65.0 mm); P. indicus Schultz, 1961: per- sonal collection, uncatalogued, 3 (53.6—62.3 mm); P. inermis Borodulina, 1981: personal collection, uncata- logued, 3 (48.6-49.8 mm); P. kiwiensis Baird, 1971: AMS I.15984-002, 1 (48.6 mm), AMS I.24496-001, 1 (73.6 mm); P. laternatus Schultz, 1938: USNM 298924, 3 (32.4-42.8 mm), MCZ 40575, 1 (31.1 mm); P. ma- tsubarai Schultz, 1961: ORIT 2572-2578, 2580, 2582, 2585, 2587, 11 (18.6-41.0 mm), NMC 79-0009, 1 (97.4 mm); P. meteori Kotthaus, 1967: MCZ 64694, 1 (17.9 mm), ZMUC P206931, 1 (37.6 mm), ZMUC P206928, 1 (54.0 mm), ZMUC P206929, 2 (12.2—21.3 mm), SIO 61-541-10, 1 (28.7 mm), USNM 256965, | (20.3 mm); P. nuttingi Gilbert, 1905: BPBM 24892, 3 (28.3-39.9 mm), BPBM 23779, 1 (47.1 mm); P. oluolus Baird, 1971: holotype, USNM 204390, 1 (26.9 mm); P. om- phus Baird, 1971: USNM 256967, 1 (32.2 mm); P. parini Borodulina, 1979: holotype, ZIL 43997, 1 (61.0 mm), illustration (Borodulina, 1979: Fig. 3) and x-ra- diograph; P. polli Schultz, 1961: MCZ 80400, 27 (9.7- 34.9 mm); P. polli?: MCZ 80401, 13 (5.9-11.0 mm); P. ruggeri: USNM 298920, 1 (16.9 mm), USNM 298920, 2 (8.9-12.6), ZMUC P202814, 1 (24.6 mm), ZMUC P206958, 1 (21.6 mm), ZMUC P206956, 1 (63.3 mm), AMS I.27166-004, 1 (64.7 mm), AMS 1.20312-007, 1 (21.3 mm), AMS I.20066-014, 1 (48.9 mm), AMS I.21372-006, 1 (25.5 mm); P. spinifer Bo- rodulina, 1979: USNM 289176, 3 (21.7-24.9 mm), ORIT 2552, 2555, 2556, 3 (48.1-50.5 mm), AMS 1.22808-028, 6 (32.1-51.6 mm), AMS I.22817-014, 6 (50.5—56.2 mm); P. spinosus Gunther, 1887: holotype, BMNH 1987.12.7.159, 1 (45.0 mm); P. stereope Jor- dan and Starks, 1904: ORIT 2519, 1 (47.7 mm); P. tridentifer McCulloch, 1914: AMSI.18711-014, 3 (48.4— 62.1 mm); P. triphanos Schultz, 1938: ZMUC P206963, 2 (26.5-30.4 mm), AMS I.24338-001, 1 (47.3 mm); P. unispinus Schultz, 1938: Polyipnus sp. (P. spinosus complex): USNM 298929, 1 (6.4 mm), AMS I.27171- 007, 1 (8.6 mm), AMS I.27166-003, 1 (8.4 mm); Pol- yipnus sp.: MCZ uncatalogued, field no. RHB 2056, 14 (5.5—-10.3 mm), USNM 298929, 1 (9.3 mm), USNM 298926, 1 (7.3 mm), MCZ 80402, 1 (9.2 mm). PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 516-524 A NEW, UNUSUALLY SEXUALLY DIMORPHIC SPECIES OF BRYCONAMERICUS (PISCES: OSTARIOPHYSI: CHARACIDAE) FROM THE PERUVIAN AMAZON Richard P. Vari and Darrell J. Siebert Abstract. — Bryconamericus pectinatus, a new species of characid fish, is de- scribed from the Rio Manu basin in southeastern Peru. Males of the species are characterized by unique elaborations of the distal portions of the anterior branched rays of the anal fin. They are also distinctive in the expansion of the anterior basal pterygiophores of the anal fin, and in the presence of lobulate bodies, evidently consisting of glandular tissue, along the base of the anterior portion of that fin. Resumen. — Bryconamericus pectinatus es una nueva especie de los caracidos que proviene de la cuenca del rio Manu, al sureste del Peru. Los machos de esta especie se caracterizan por la singular elaboracion de una porcion distal de los radios anteriores no ramificados de la aleta anal. Se los puede distinguir por el ensanchamiento de los pterygiophoros de la aleta anal y en la presencia de cuerpos lobulados que evidentemente consta de un tejido glandular a lo largo de la base de la parte anterior de esta aleta. As presently defined, the characid genus Bryconamericus Eigenmann (in Eigenmann et al. 1907) consists of some three dozen relatively small-sized species (Gery 1977: 386). Species of Bryconamericus occur in a variety of freshwater ecosystems at lower altitudes across a broad expanse of South and Central America to both sides of the Andean cordilleras. The genus is most di- verse, however, in the Atlantic drainages of the continent, in particular through the vast reaches of the Amazon basin. In their com- pendium of the freshwater fishes of Peru, Ortega & Vari (1986:7) listed ten species of Bryconamericus as occurring in the myriad river systems that drain the part of the coun- try to the east of the Andean Cordilleras. Those authors, following Bohlke et al. (1978), noted, however, that the entire neo- tropical freshwater fish fauna is still poorly understood, and that their tentative listing of the species known from Peru would un- doubtedly undergo numerous modifications in the future. In the course of studying a collection of fishes from Rio Manu system of the Rio Madre de Dios basin in south- eastern Peru, we discovered an undescribed species assignable to Bryconamericus that differs from the other members of the genus in several striking features. The most note- worthy of these are the unusual modifica- tions of the anterior anal-fin rays and as- sociated basal pterygiophores in adult males. Males are also characterized by a series of lobulate, evidently glandular, bodies along the lateral surfaces of the anterior anal-fin rays. Methods.— All measurements are given as proportions of standard length (SL) ex- cept for subunits of the head which are pre- sented as proportions of head length. Ver- tebral counts were taken from radiographs, and cleared and counterstained specimens. This number includes the four vertebrae in- VOLUME 103, NUMBER 3 corporated in the Weberian apparatus and considers the fused PU,+U, as a single ele- ment. In the counts of elements in the me- dian and pelvic fins, lower-case Roman nu- merals indicate unbranched rays, and Arabic numerals indicate branched rays. For me- ristic values the range for each value for the holotype and measured paratypes is pre- sented first, with the value for the holotype indicated in brackets. The following abbreviations are used for institutions: CM— Carnegie Museum, Pitts- burgh; FMNH—Field Museum of Natural History, Chicago; MHN-USM— Museo de Historia Natural de la Universidad Nacio- nal Mayor de San Marcos, Lima, Peru; and USNM-—National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. Bryconamericus pectinatus, new species Figs. 1-5, Table 1 Holotype. —MHN-USM 2057, 27.3 mm SL, female. Peru, Departamento Madre de Dios, Provincia Manu, Parque Nacional de Manu, second large quebrada along Trail 1 leading to the east from Pakitza, tributary of Rio Manu (approx. 11°50’S, 71°21’W); H. Ortega, D. Siebert, M. Rauchenberger, I. Samanez, G. Contreras, and J. Canepa; 13 Sep 1988. Paratypes.—15 specimens collected with holotype, MHN-USM 2058, 7 specimens; USNM 303442, 8 specimens including 3 males, 17.0-23.6 mm SL (1 specimen cleared and counterstained for cartilage and bone). Ten specimens collected down- stream of holotype locality by same collec- tors, 17 Sep 1988; MHN-USM 2059, 5 specimens including 2 males; USNM 303441, 5 specimens, 19.8—34.8 mm SL (1 specimen cleared and counterstained for cartilage and bone). Diagnosis. —The combination of a single row of teeth on the dentary, two rows of teeth on the premaxilla with four teeth in the inner series that are larger than those of 517 the outer row, the limited number of teeth along the anterior margin of the maxilla, the simple curve to the border of the upper jaw, the lack of scales on the caudal fin, the large third infraorbital contacting the preopercle along its posterior and ventral margins, se- tiform gill-rakers, the complete laterosen- sory canal system on the body, and the ab- sence of a glandular pouch on the caudal fin in males serve to assign the species to Bry- conamericus. Within the genus, B. pectina- tus can be distinguished by autapomorphic modifications of the anterior rays of the anal fin and of the associated basal pterygio- phores, and by the presence of lobulate, ev- idently glandular, bodies along the anterior anal-fin rays (see ““Description”’ below). The species is also distinctive within the genus on the basis of the combination of a rela- tively low number of anal-fin rays, a robust body that is relatively deep in adults, the possession of 32 or more lateral-line scales to the hypural joint, the absence of dark pigmentation on the tips of the caudal-fin lobes, and the presence of 5 or 6 moderately sized teeth on the maxilla. Description. — Body robust, slightly more compressed in smaller individuals (Figs. 1, 2). Dorsal profile of head nearly vertical along upper lip, distinctly convex from mar- gin of upper lip to vertical line through pos- terior nostril, slightly convex from that line to rear of head. Dorsal profile of body smoothly curved from rear of head to origin of dorsal fin, more convex in larger, deeper bodied specimens; straight and posteroven- trally slanted at base of dorsal fin, gently convex from base of last dorsal-fin ray to caudal peduncle, convexity more pro- nounced in larger individuals. Ventral pro- file of head notably convex anteriorly, less so posteriorly. Ventral profile of body gently curved from tip of lower jaw to caudai pe- duncle; convexity more pronounced in larg- er specimens. Greatest body depth located slightly anterior of origin of dorsal fin. Cau- dal peduncle somewhat laterally com- pressed, deeper in adult males (0.146-0.164 518 of SL) than adult females (0.121-0.139 of SL). No sexual dimorphism obvious in oth- er examined morphometric features. Head relatively large, overall profile rounded anteriorly. Jaws equal, mouth ter- minal; lips fleshy with brown tips of teeth in outer row on premaxilla visible when mouth closed; ventral margin of upper jaw concave, main axis of maxilla aligned pos- teroventrally, posterior tip of maxilla ex- tending below orbit, reaching posteriorly nearly to vertical line through center of eye. Nostrils on each side of head very close to each other; opening of anterior nares ver- tically ovoid; opening of posterior nares crescent-shaped; thin flap of skin extending laterally from strip of tissue separating two nares. Orbital rim free, no adipose eyelid present. Six well-ossified infraorbitals present, each bearing laterosensory canal segment. Third infraorbital very large, ventral and posterior margins contacting lateral surface of pre- opercle. Fourth infraorbital without poste- rior branch of laterosensory canal. Latero- sensory canal segment in sixth infraorbital (dermosphenotic) consisting of a single tube. No supraorbital present. Lower jaw with single series of 11 to 13 teeth. First three teeth largest, fourth tooth somewhat smaller, fifth tooth through end of dentary series distinctly smaller than fourth, gradually diminishing in size pos- teriorly. First five teeth typically tricuspi- date with central cusp distinctly larger. Sec- ond tooth often with additional very small cusp on lateral margin. Fourth tooth either tricuspidate or bicuspid, sometimes with two conditions in a single specimen; when bi- cuspid posterior cusp large, posterior mar- gin straight, not recurved. Sixth tooth bi- cuspid; remaining teeth on lower jaw unicuspid. Two rows of teeth on premaxilla; teeth of inner row larger. Six teeth in outer row, ori- entation of main axis of teeth alternating slightly anteriorly and posteriorly, resulting in a wavy margin of tooth edges when ex- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON amined in ventral view. Outer row teeth bicuspid, with medial cusp much larger. Four teeth in inner row of premaxilla; with three to five cusps, middle cusp distinctly larger. Maxilla with five or six small tricus- pidate teeth along upper half of exposed an- terior margin; dorsal tooth on maxilla sit- uated proximate to lateral tooth of inner row of premaxilla. Dorsal-fin rays 11,8 or 11,9 [11,9]. Margin of dorsal fin rounded, second unbranched and first branched rays subequal; depressed fin not reaching anterior margin of adipose fin in smaller specimens, overlapping an- terior of fin in largest specimens examined. Adipose fin moderately developed in all specimens. Pectoral-fin rays 10 to 12 [11]. Margin of pectoral fin rounded; tip of fin extends nearly to vertical line through in- sertion of pelvic fin in smaller specimens, just slightly short of that line in largest spec- imens examined. Pelvic-fin rays 6 to 8 [7]. Margin of pelvic fin rounded; reaches be- yond vertical line through origin of anal fin in smaller specimens, barely reaches or just short of that line in larger individuals. Cau- dal fin forked, lobes rounded; 7 or 8 pro- current rays both dorsally and ventrally. Anal-fin rays vi,12 to 14 (counts taken from radiographs; two or three anterior un- branched rays not apparent in specimens that have not been cleared and stained or radiographed). Anal fin with pronounced sexual dimor- phism in adults (sex verified by dissection). First branched anal-fin ray in females long- est, remaining branched rays becoming pro- gressively shorter posteriorly. Margin of anal fin in females somewhat emarginate (see Fig. 1). Males with posterior unbranched and an- terior branched anal-fin rays distinctively modified, longest rays proportionally short- er than in females (compare Figs. 1 and 2). Unbranched anal-fin rays of males, in par- ticular last unbranched ray, but also rays hidden by body tissue and scales, thickened basally (Fig. 4). First branched ray with VOLUME 103, NUMBER 3 519 Fig. 1. Departamento Madre de Dios, Provincia Mant, Parque Nacional de Mant. distal portion significantly expanded anteroposteriorly, division of distal ray seg- ments highly asymmetrical, with progres- sive subdivision of posterior, but not an- terior half of each branched ray segment (Fig. 3). This condition contrasts with typ- ical progressive distal subdivision of both anterior and posterior branches of anal-fin ray in females of the species and other cha- raciforms (compare Fig. 3 with figs. 38 to 43 in Weitzman & Fink 1985). Distal por- Bryconamericus pectinatus, new species, holotype, MHN-USM 2057, 27.3 mm SL, female. Peru, tions of second through fourth branched anal-fin rays also somewhat expanded anteroposteriorly; degree of distal expan- sion progressively decreasing posteriorly along fin, never as pronounced as in first branched ray (Fig. 3). Form of posterior anal- fin rays comparable to those in females of species. Proximal portions of enlarged anal-fin rays and ventral sections of associated basal pterygiophores of males thickened relative Fig. 2. Bryconamericus pectinatus, new species, paratype, USNM 303441, 34.8 mm SL, male. Peru, De- partamento Madre de Dios, Provincia Manu, Parque Nacional de Mant. 520 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Bryconamericus pectinatus, male paratype, USNM 303441; 32.0 mm SL; anterior rays of anal fin and associated basal sheath of scales; anterior to left. to condition in females (compare within Fig. 4), and in comparison to typical mor- phology of these elements in other chara- cids. First and second basal pterygiophores in males more closely associated with hemal spines of proximate vertebrae than in fe- males (Fig. 4). Large males with mass of lobulate material extending ventrally from under scales across lateral surface of anal- fin membranes (Fig. 5). Lobulate bodies across membranes of anal fin extend nearly to end of exposed unbranched fin-rays; only reaching about one half distance to tip of anterior branched fin rays. Such lobulate bodies on anal fin not known elsewhere in Bryconamericus or among other characids. Lobulate bodies appear to be glandular, but limited available material prevents histo- logical confirmation of this supposition. Scales cycloid, thin. Caudal fin not scaled. Anal fin with 1 or 2 series of scales over- lapping bases of fin rays along anterior two thirds of fin. One series of scales overlap- ping proximal portions of unbranched rays, 2 series over bases of anterior 4 or 5 branched rays. Pored lateral-line scales ex- tend from supracleithrum to hypural joint 32 to 36 [33]; all scales of lateral line pored, canals in lateral-line scales straight; 3 series of scales extend beyond hypural joint onto caudal-fin base; 6 to 6'% [64] scales in trans- verse series from origin of dorsal fin to lat- eral line; 5 to 6 [6] scales in transverse series from lateral line to origin of anal fin. Total vertebrae 35 (2), 36 (5), 37 (4). Color in alcohol. —Overall coloration of specimens fixed in formalin and preserved in alcohol light tan. No guanine remaining on scales. Scattered small chromatophores on lower lip, snout, dorsal portion of head, area posterior and ventral to orbit, and on dorsal half of opercle. Pigmentation most intense on dorsal portion of head, most no- tably in smaller specimens. Distinct, verti- cally elongate, patch of deep-lying chro- matophores forming humeral spot. Humeral spot separated anteriorly from pigmented portion of opercle by region with few or no dark chromatophores. Diffuse mid-lateral stripe of small dark chromatophores ex- VOLUME 103, NUMBER 3 tending posteriorly from slightly behind hu- meral spot to rear of caudal peduncle. Stripe more obvious posteriorly at all body sizes, overall intensity greater in larger individu- als. Portion of stripe on caudal peduncle most heavily pigmented. Margins of scale pockets on dorsal portion of body outlined by series of small dark chromatophores, in- tensity greater in adult specimens. Dorsal-fin rays outlined by series of small dark chromatophores, pigmentation most intense in large males. Membranes of dor- sal-fin rays dusky. Middle and most ventral rays of caudal fin dusky. Hyaline region on fin separates dusky dorsal caudal-fin rays from pigmented middle rays. Similar un- pigmented patch on basal portion of middle rays of lower lobe of caudal fin. Anal fin hyaline in smaller specimens, with small chromatophores giving distal portions of rays dusky appearance in larger individuals. Paired fins hyaline in specimens of all sizes. Distribution. —Known only from the type locality, the upper Rio Manu of southeast Peru. Etymology. —The species name, pectina- tus, from the Latin for rake or comb, refers to the stiffened, subdivided, comb-like an- terior rays of the anal fins in the males of the species. Ecology. —The specimens were collected in a small, moderately flowing, rainforest stream with a rock and mud bottom. Remarks. —Bryconamericus pectinatus demonstrates a number of striking features not encountered elsewhere in the genus, most notably the modifications of the anterior anal-fin rays and associated basal pteryg- iophores in males. We are consequently confronted with a choice of expanding the definition of Bryconamericus to include this very distinctive species or proposing a new monotypic genus for the species. Many au- thors publishing on characiforms have seg- regated equally distinctive species in mono- typic genera rather than expand the definition of previous genera to accommo- date the features of newly discovered species. 521 Fig. 4. Positives of radiographic plates of anal-fin regions of Bryconamericus pectinatus; top, male, USNM 303441, 34.8 mm SL; bottom, female, USNM 303441, 28.6 mm SL. Anterior to left. Fink (1976), for example, decided to de- scribe a new genus, Eretmobrycon, for E. bayano, a Bryconamericus-like species from Panama characterized by specializations of the caudal and pelvic fins. In his opinion (1976:340) it ““seems best to recognize the unique specilizations of FE. bayano and place it in a monotypic genus.” The choice between these alternatives, the expansion of the definition of a previous genus, and the proposal of a new genus, is 522 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Bryconamericus pectinatus, male, USNM 303441; 34.8 mm SL. Close-up of anal fin showing lobulate, possibly glandular tissue extending lateral to anterior fin-rays. Anterior to left. somewhat arbitrary given the general lack of information on relationships between Bryconamericus and its possible close rel- atives. The primary basis for our placement of the new species within the Bryconameri- cus 1s its possession of the combination of characters that are presently considered di- agnostic for the genus (see Diagnosis). The distinctive features of Bryconamericus pec- tinatus, in turn, are autapomorphies for the species, with no bearing at present on the question of its phyletic associations. We must admit, however, that we have little confidence that Bryconamericus in this broad sense represents a natural (monophy- letic) component of the Characidae, or that the closest relatives of B. pectinatus con- sequently occur within that genus. Problems with the naturalness of Bryconamericus have been long recognized, even by Eigenmann who first proposed the genus in 1907. Ei- genmann subsequently commented (1927: 358) ““There are three or four recognizable groups in the genus Bryconamericus and these may have been independently derived from various species of Astyanax and Hemibrycon.” Questions concerning the naturalness of many New World characi- form taxa and problems with the utility of many commonly used diagnostic features within the order, including those pertinent to Bryconamericus, also have been noted more recently by Fink (1976), Vari & Gery (1980), Weitzman & Fink (1983), and oth- ers. Given these general problems and the specific questions concerning the monophy- ly of Bryconamericus cited by Eigenmann, there is little likelihood that the inclusion of B. pectinatus in the genus will disrupt a natural assemblage. In light of this situation the provisional inclusion of B. pectinatus within Bryconamericus is, in our view, pref- erable to the creation of a new monotypic genus. One other genus in the Characidae, Car- lastyanax Géry (1972) demonstrates certain similarities with Bryconamericus pectinatus which raises the question of a possible close relationship between these two taxa. The type species of Carlastyanax, Astyanax au- rocaudatus Eigenmann (1913:26), was col- lected in the upper Rio Cauca, a tributary of the Rio Magdalena of northwestern Co- lombia, which drains into the Caribbean Sea. VOLUME 103, NUMBER 3 523 Table 1.—Morphometrics of Bryconamericus pectinatus, new species. Standard length is expressed in mm; measurements | to 11 are proportions of standard length; 12 to 16 are proportions of head length. Values for paratypes represent ranges of measured specimens. Holotype Paratypes Standard length 27.3 17.0-34.8 1. Greatest body depth 0.395 0.328—-0.420 2. Snout to dorsal-fin origin 0.623 0.561-0.631 3. Snout to pectoral-fin insertion 0.296 0.269-0.288 4. Snout to pelvic-fin insertion 0.532 0.484-0.537 5. Snout to anal-fin origin 0.691 0.636-0.689 6. Origin of dorsal fin to hypural fin 0.454 0.430-0.482 7. Length of longest dorsal-fin ray 0.234 0.220-0.244 8. Pectoral-fin length 0.227 0.206-0.237 9. Pelvic-fin legnth 0.150 0.136-0.169 10. Caudal-peduncle depth 0.139 0.121-0.164 11. Head length 0.308 0.295-0.311 12. Snout length 0.223 0.210-0.260 13. Orbital diameter 0.329 0.291-0.350 14. Postorbital length 0.447 0.440-0.500 15. Upper jaw length 0.450 0.435-0.456 16. Interorbital width 0.329 0.320-0.350 Eigenmann (1927:322) subsequently noted that the species “‘ought probably to be dis- tinguished generically,’ but did not propose a new genus for the species. Géry (1972:16— 21), however, following on that suggestion, proposed Carlastyanax for Astyanax au- rocaudatus. Carlastyanax aurocaudatus and Bryconamericus pectinatus are similar in their blunt, massive heads, heavy bodies, and in the presence of two series of scales along the base of the anal fin. These features are, however, hardly unique to these species within the Characidae. Géry (1972:18) also notes that the anterior rays of the anal fin in Carlastyanax aurocaudatus are sexually dimorphic, with those in males being thick- ened, another possible similarity with Bry- conamericus pectinatus. Examination of the holotype of Astyanax aurocaudatus (FMNH 56882, formerly CM 5162) and part of the paratype series of the species (FMNH 56883, formerly CM 5163a- d) has shown that those specimens and Bry- conamericus pectinatus differ in numerous details. The basally expanded anterior anal- fin rays in the holotype of Astyanax auro- caudatus show none of the asymmetrical distal branching and anteroposterior expan- sion characteristic of Bryconamericus pectinatus. Neither does Astyanax aurocau- datus show any indication of the modifi- cations of the anterior basal pterygiophores, or possess the lobulate, possibly glandular, bodies found along the basal portion of the anal-fin membranes in males of Brycona- mericus pectinatus. Géry (1972) listed a number of diagnostic features for Carlastyanax. Notable differ- ences between that genus and Bryconamer- icus pectinatus include the degree of devel- opment of the third infraorbital (small in aurocaudatus, very well developed in pec- tinatus), morphology of the third dentary tooth (posterior cusp greatly recurved in au- rocaudatus, tricuspid or non-recurved bi- cuspid in pectinatus), and form of the an- terior nostril (tubular in aurocaudatus, a simple opening in pectinatus). Admittedly, neither these differences, nor others we could detail would serve to refute a hypothesis that these two species are closely related. Alternatively we have been unable to iden- tify any derived feature indicative of a close relationship between the two taxa. In the 524 absence of such information we prefer to take a conservative course of action and as- sign pectinatus to Bryconamericus. Acknowledgments The material of Bryconamericus pectina- tus was collected during a survey funded by the BIOLAT program of the National Mu- seum of Natural History, Smithsonian In- stitution. The ichthyological collecting pro- gram was a collaborative effort between USNM and MHN-USM. The assistance of Prof. Hernan Ortega and his colleagues (MHN-USM) in all facets of the project is greatly appreciated. Dr. Barry Chernoff and Ms. Mary Anne Rogers (FMNH) arranged for the loan of the holotype and part of the paratype series of Astyanax aurocaudatus. Figures 1, 2, 4, 5 were prepared by Mr. Theophilus B. Griswold. The Spanish “‘Re- sumen”’ was translated by Dr. Ramiro Bar- riga of the Escuela Politécnica Nacional, Quito, Ecuador. This paper was improved by the suggestions of Dr. Stanley H. Weitz- man (USNM), Dr. Wayne C. Starnes (USNM), Dr. Scott A. Schaefer (Academy of Natural Sciences of Philadelphia), Dr. Thomas A. Munroe (National Marine Fish- eries Service, Systematics Laboratory), and two anonymous reviewers. We thank all of the above for their assistance and interest. Research associated with this study was supported in part by the Neotropical Low- land Research Program of the International Environmental Sciences Program of the Smithsonian Institution. This is contribu- tion no. 1, Biological Diversity in Latin America (BIOLAT) Project, Smithsonian Institution. Literature Cited Bohlke, J., S. H. Weitzman, & N. A. Menezes. 1978. Estado atual da sistematica dos peixes de agua PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON doce da América do Sul.— Acta Amazonica 8(4): 53-62. Eigenmann, C. H. 1913. Some results from an ich- thyological reconnaissance of Colombia, South America. Part IJ.—Indiana University Studies 18:1-30. . 1927. The American Characidae.— Memoirs of the Museum of Comparative Zoology 43(4): 311-428. —, W. L. McAtee, & D. P. Ward. 1907. On further collections of fishes from Paraguay. — Annals of the Carnegie Museum 4(2):1 10-157. Fink, W. L. 1976. A new genus and species of chara- cid fish from the Bayano River basin, Panama (Pisces: Cypriniformes).— Proceedings of the Bi- ological Society of Washington 88(30):33 1-344. Gery, J. 1972. Contribution a l’étude des poissons characoides de l’équateur.— Acta Humbold- tiana, Series Geologica, Palaentologica et Biolo- gica 2:1-110. 1977. Characoids of the world. T.F.H. Pub- lications, Neptune City, New Jersey, 672 pp. Ortega, H. & R. P. Vari. 1986. Annotated checklist of the freshwater fishes of Peru.—Smithsonian Contributions to Zoology 437:1—25. Vari, R. P., & J. Géry. 1980. Cheirodon ortegai, a new markedly sexually dimorphic cheirodon- tine (Pisces: Characoidei) from the Rio Ucayali of Peru.— Proceedings of the Biological Society of Washington 93(1):75—92. Weitzman, S. H., & W. L. Fink. 1983. Relationships of the Neon Tetras, a group of South American freshwater fishes (Teleostei, Characidae), with comments on the phylogeny of New World Characiforms.—Bulletin of the Museum of Comparative Zoology 150(6):339-395. —., & S. V. Fink. 1985. Xenurobryconin phy- logeny and putative pheromone pumps in glan- dulocaudine fishes (Teleostei: Characidae).— Smithsonian Contributions to Zoology 421:1— 121. Department of Vertebrate Zoology (Fish- es), National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Present address of Siebert: Depart- ment of Zoology, British Museum (Natural History), Cromwell Road, London SW7 SBD, England. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 525-542 MOOJENICHTHYS MIRANDA-RIBEIRO (PISCES: OSTARIOPHYSI: CHARACIDAB), A PHYLOGENETIC REAPPRAISAL AND REDESCRIPTION Ricardo M. C. Castro and Richard P. Vari Abstract. —The Neotropical characiform characid genus Moojenichthys Mi- randa-Ribeiro is hypothesized to form a monophyletic lineage with Triportheus Cope on the basis of shared derived features of the pectoral girdle and perhaps of the first infraorbital. Autapomorphies for Moojenichthys are discussed. Moo- Jenichthys myersi Miranda-Ribeiro, the only member of the genus, is rede- scribed. This species is apparently endemic to the Rio do Brago system of the state of Bahia, Brazil. Resumo. —Uma hipotese de relacoes filogenéticas reunindo em um so grupo monofilético os géneros neotropicais Moojenichthys e Triportheus, ambos per- tencentes 4 familia Characidae e 4 ordem Characiformes, é¢ formulada com base na posse em comum de caracteres derivados na cintura escapular e, pos- sivelmente, primeiro infra-orbital. Moojenichthys myersi Miranda-Ribeiro, o unico membro do género, é redescrito. Esta espécie €, aparentemente, endémica da bacia do Rio do Braco, no estado da Bahia, Brasil, e consiste na unica ocorréncia de um representante da linhagem evolutiva Triportheus-Moojenich- thys nos rios costeiros do leste do Brasil, excluindo o rio Sao Francisco. Miranda-Ribeiro (1956:546) proposed the characid genus Moojenichthys for a single species, M/. myersi, first described in that publication on the basis of two specimens collected in the Rio do Braco, near Ilhéus in the state of Bahia of eastern Brazil. Moo- jenichthys myersi has a number of external anatomical features unusual within the Characiformes. Perhaps the most striking of these are its elongate, laterally flattened body, and the pronounced, laterally com- pressed mid-ventral keel. Although the tho- racic keel is not obvious in specimens of Moojenichthys under 15 mm SL, mid- to large-sized individuals of the genus have a distinct mid-ventral ridge extending from the isthmus posteriorly to the origin of the pelvic fin. In his original description of Moojenich- thys Miranda-Ribeiro stated that the genus was “related”’ to Triportheus Cope (1872) and Clupeacharax Pearson (1924) both of which also have elongate, laterally com- pressed bodies characterized by varyingly developed mid-ventral keels. Miranda-Ri- beiro did not explicitly state which char- acters lead him to propose that these species are “related,” presumably closely. It seems reasonable, nonetheless, to assume that the overall similarities in body shapes, partic- ularly the presence of mid-ventral keels, contributed significantly to this hypothesis. Despite the distinctive external mor- phology of Moojenichthys myersi, subse- quent references to the genus and species are extremely limited. Géry (1972:55), in his key to New World characiforms, com- mented that Triportheus and Moojenichthys are probably derived from Brycon Muller and Troschel, and on the next page pro- 526 posed that C/upeacharax is close to Tripor- theus (1972:56). More recently, Géry (1977: 346) again emphasized the similarities be- tween Moojenichthys and Triportheus, not- ing, however, that the reported dentition of M. myersi is more reminiscent of the subfamily ““Tetragonopterinae” of the fam- ily Characidae. In that publication Geéry did not comment on Miranda-Ribeiro’s pro- posal of a possible relationship between Moojenichthys and Clupeacharax, but rath- er segregates Clupeacharax in the monotyp- ic subfamily Clupeacharacinae. Castro (1981:138), in turn, cited some external similarities between Clupeacharax and En- graulisoma Castro. Other than for the original description by Miranda-Ribeiro (1956) and Géry’s brief comments (1972, 1977), we know of no published citations of Moojenichthys. Sim- ilarly, the primary ichthyological literature apparently does not include records of the subsequent capture of the species. This is not surprising given that . myersi is ap- parently endemic to the Rio do Braco sys- tem, a poorly sampled river basin that drains into the Atlantic Ocean slightly north of the city of Ilhéus in the state of Bahia, Brazil. Recent collecting efforts in the coastal rivers of Bahia associated with our revi- sionary studies of the characiform families Curimatidae and Prochilodontidae, have resulted in the capture of a large series of Moojenichthys myersi with a much greater range of standard lengths than available to Miranda-Ribeiro. This additional material allows us to provide a detailed redescription of the genus and species. Those specimens also permit anatomical studies to evaluate previous suggestions about the relation- ships of Moojenichthys. Methods.— All measurements are given as proportions of standard length (SL) ex- cept for subunits of the head which are pre- sented as proportions of head length (HL). Lateral-line scale counts include all pored scales along that series, including the scales posterior of the hypural joint. Vertebral PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON counts were taken from radiographs, and specimens cleared and counterstained for bone and cartilage. The vertebral count in- cludes the four vertebrae incorporated in the Weberian apparatus, and considers the fused PU,+U, asa single element. In counts of median and pelvic fins, lower-case Ro- man numerals indicate unbranched rays, and Arabic numerals indicate branched rays. The range for each meristic value of all mea- sured specimens is presented first, with the value for the holotype indicated in square brackets. Measurements were made follow- ing the methods outlined in Fink & Weitz- man (1974:1—2). The following institutional abbreviations are used: ANSP—Academy of Natural Sci- ences of Philadelphia; MNRJ— Museu Na- cional, Rio de Janeiro; MZUSP— Museu de Zoologia, Universidad de Sao Paulo; USNM-— National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C.; and FFCLRP-USP—Faculdade de Filosofia, Ciéncias e Letras de Ribeirao Pre- to, Universidade de Sao Paulo. Phylogenetic Analysis As noted in the introductory comments, Miranda-Ribeiro (1956:546) considered Moojenichthys, Triportheus, and Clupea- charax to be related, albeit without specif- ically stating the basis for his opinion. A number of questions exist about the phy- logenetic associations of these taxa, and at least in the case of 7riportheus the species- level classification of the genus remains un- settled. Moojenichthys myersi has been collected only in the Rio do Braco system of eastern Brazil. Clupeacharax includes a single species, C. anchoveoides, a poorly known fish reported from scattered sites ranging from Argentina (Miquelarena & Casciotta 1982:333), through Bolivia (Pearson 1924: 47) and Peru (Ortega & Vari 1986:8), to Ecuador (Stewart et al. 1987:26). Triporthe- us, a much more speciose genus well rep- VOLUME 103, NUMBER 3 resented in museum collections, is found on both sides of the Andean Cordilleras across much of lowland South America. Miranda- Ribeiro (1941), who applied Chalcinus Cu- vier & Valenciennes (1849) to the members of Triportheus, recognized eleven species in- cluding Chalcinus culter Cope which Fowler (1907) segregated in the genus Coscinoxy- ron. Myers (1940:170) pointed out that Chalcinus was already occupied in the Hy- menoptera, and that Triportheus was the next available name for the members of the genus. Schultz (1944:273) and Weitzman (1960:239) agreed that Miranda-Ribeiro’s revisionary effort was unsatisfactory, a view that we share. Gery (1977:343, 654) in his key to the members of Jriportheus, recog- nized only nine of the nominal species as valid, and tentatively retained cu/ter in Triportheus. This uncertainty concerning the recognizable species of Triportheus compli- cates a determination of whether the genus is monophyletic. Those questions are be- yond the scope of this study. We wiill, rather, attempt to determine the phyletic relation- ships of Moojenichthys, and critically eval- uate suggestions that the genus is related to Triportheus and Clupeacharax. Mid-ventral keel.—One of the most ob- vious features of Moojenichthys is the dis- tinct mid-ventral keel that extends poste- riorly from the isthmus to between the origins of the pelvic fins. Whereas the pos- terior portion of this keel is formed by fleshy tissue, the anterior portion is underlain by asymmetrically expanded coracoid bones. The somewhat irregular anterior margins of the coracoids are relatively short where they meet the anteroventral margins of the cleithra (Fig. 1B). The dorsal margin of the coracoids in Moojenichthys gradually rises posteriorly to the region where it articulates with the scapula and mesocoracoid. As a consequence, the overall proportions of the plate-like ventral portion of the coracoid are distinctly asymmetrical in lateral view, with the posterior margin of the coracoid notably deeper than the anterior border of the bone. 527 Such asymmetrically expanded coracoids are relatively unusual within characiforms. A horizontally rectangular, moderately-sized coracoid with an overall horizontally rect- angular form is found in the vast majority of characiforms of all families (e.g., Charac- idae (Brycon, Fig. 1A); Prochilodontidae, see Roberts 1973:fig. 24; Hemiodontidae, see Roberts 1974:figs. 16, 53; Lebiasinidae, see Weitzman 1964:fig 10; Parodontidae, see Roberts 1974:fig. 76; and Erythrinidae, see Starks 1930:fig. 8). Indeed, only the cha- raciform families Characidae and Gaster- opelecidae include taxa in which the cora- coids are dramatically expanded to form distinct thoracic keels. Within the Charac- idae enlarged coracoids are absent among Old World members of the family, and the vast majority of Neotropical characids sim- ilarly lack significant expansions of these bones (e.g., Acestrorhynchus, see Roberts 1969:fig. 52; and Brycon (Fig. 1A), see also Weitzman 1962:figs. 18, 19). As noted in the introductory section, the species of Trip- ortheus are also characterized by a distinct thoracic keel. The coracoids in 7riportheus are similar to those of Moojenichthys in being asymmetrically and vertically ex- panded in lateral view. In Triportheus, how- ever, the dorsal margin of the plate-like ven- tral portion of the ossification is more steeply angled, and the posterior margin of the bone much more extensive vertically than that in Moojenichthys (Fig. 1C). This gives the pro- file of the bone a near equilateral triangular appearance. Given that most characids and non-characid characiforms lack enlarged coracoids, the common occurrence of ex- panded coracoids found in Moojenichthys and Triportheus is reasonably hypothesized as a derived character which, in turn, would be congruent with the hypothesis that the two genera are sister taxa. Expanded coracoids associated with a thoracic keel are, however, not unique to Moojenichthys and Triportheus among characiforms. As noted in the introductory discussion, an enlargement of that element 528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON B Fig. 1. C Coracoids of A) Brycon falcatus, USNM 226161, 75.1 mm SL; B) Moojenichthys myersi, USNM 304497, 84.8 mm SL; and C) Triportheus angulatus, USNM 270343, 76.6 mm SL; right side, medial view, anterior to left. also occurs in Clupeacharax and may have lead Miranda-Ribeiro to suggest that the ge- nus was related to Moojenichthys. Starks (1930:22—23, fig. 9) described and figured the expanded coracoid of Rhaphiodon Agassiz, a genus of large predatory Neo- tropical characiforms. Weitzman (1960:239) noted that a keeled thorax and expanded coracoids, although unusual among characi- forms (his “‘characids’’), also occur in Pia- bucus Oken, Pseudocorynopoma Perugia, Rhaphiodon, and the genera Carnegiella Ei- genmann, Gasteropelecus Pallas, and Thor- acocharax Fowler, the last three of which together constitute the Gasteropelecidae (sensu Greenwood et al. 1966:395). Ex- panded coracoids also are found in Cynodon Spix, Hydrolycus Miller and Troschel and Gnathocharax Fowler. All of the above cit- ed taxa appear to be surface feeders which, with the exception of the Gasteropelecidae, also have moderately to distinctly elongate bodies. The occurrence of expanded coracoids in a number of characid taxa other than Moo- jenichthys and Triportheus brings, into question the appropriateness of using that feature to propose a close relationship for those genera. Weitzman suggested (1954: 230-231) that the specialized expansions of the coracoids probably arose several times within the Characiformes. The present poor understanding of phylogenetic relationships within the Characiformes in general, and the Characidae in particular, restricts the degree to which we are able to critically evaluate that suggestion in all instances. Nonetheless, subsequent research by var- ious researchers has yielded data that sup- port Weitzman’s suggestion, and which in- VOLUME 103, NUMBER 3 dicates that the occurrence of expanded coracoids in characids other than Moojen- ichthys and Triportheus is homoplastic rel- ative to that feature in those genera. Vari (1977:4—6) discussed a series of dis- tinctive derived features of the posterior chamber of the gas-bladder and of the an- terior proximal pterygiophores of the anal fin that unite Piabucus with Iguanodectes Cuvier. In [guanodectes the pre-pelvic re- gion is transversely rounded. Pseudocory- nopoma is a member of the subfamily Glan- dulocaudinae, a taxon which Weitzman et al. (1985:112-113) noted may not represent a monophyletic assemblage. Nonetheless those authors tentatively suggested that Pseudocorynopoma may be part of a mono- phyletic subset of genera within the Glan- dulocaudinae. Other species in the subfam- ily lack keeled thoracic regions. Howes (1976) united Rhaphiodon, Cynodon, and Hydrolycus as a tribe, the Cynodontini, and hypothesized on the basis of a variety of characters that the lineage consisting of these three genera was most closely related to var- ious genera of the characid tribe Characini. The Characini, in turn, consists of species in which the coracoids are not dramatically enlarged. Thus the available evidence indicates that the characid taxa with keeled thoracic re- gions cited in the previous paragraph are each in turn most closely related to species or species groups without that derived mod- ification. The species with expanded cora- coids cited in the immediately preceding section also lack the derived features of the infraorbital series and lateral ethmoid com- mon to Moojenichthys and Triportheus (see discussions in following sections). Conse- quently it is most parsimonious to assume that the pre-pelvic keels of Piabucus, Pseu- docorynopoma, Rhaphiodon, Cynodon, and Hydrolycus are homoplastic relative to the expanded coracoids in Moojenichthys and Triportheus. Information concerning the phylogenetic relationships of the Gasteropelecidae, Clu- 529 peacharax, and Gnathocharax is somewhat more equivocal. The three genera of the Gasteropelecidae constitute a highly de- rived lineage presumably derived from some component of what is now recognized as the Characidae (Weitzman 1954:243). Al- though the closest relatives of the Gaster- opelecidae remain to be elucidated, we agree with Weitzman (1954) that the overall char- acters of gasteropelecids differ dramatically from those of 7riportheus and that gaster- opelecids are apparently evolved from a dif- ferent subunit within the Characiformes. Although a resolution of the phylogenetic associations of gasteropelecids would re- quire an analysis that extends far beyond the scope of this paper, one feature of the Gasteropelecidae is noteworthy relative to this question. Weitzman & Fink (1983:391) noted that the supraorbital bone is absent in all the “‘tetragonopterine”’ characids they examined. The supraorbital is widely dis- tributed among characiforms, and also among the members of the Characidae both in the New World and Africa. The absence of that ossification is thus hypothesized to be a derived condition within the Charac- idae. The supraorbital is absent in the Gas- teropelecidae (see Weitzman 1954:7), and the common absence of the supraorbital in ‘‘tetragonopterines’’ and gasteropelecids may be a derived feature indicative of com- mon ancestry of those taxa. Both 7riporthe- us and Moojenichthys, in contrast, retain a supraorbital. The relationships of Gnathocharax, a monotypic genus of the Amazon basin with an expanded coracoid, are still unresolved. Gnathocharax lacks the distinct anterior process of the lateral ethmoid common to Moojenichthys, Triportheus, Brycon, and various other characids (see discussion un- der ‘““Autapomorphies of Moojenichthys’’). Gnathocharax also does not have the dis- tinctive modification of the first infraorbital found in Moojenichthys, Triportheus, and at least some species of Brycon. Finally, Gnathocharax \acks a supraorbital and is 530 characterized by conical dentition typical of the characid tribe Characini and groups probably aligned with that tribe (see also discussions concerning the monophyly of the Characini by Menezes (in Sazima 1983), Vari (1986), and Weitzman & Vari (1987)). The cumulative data is congruent with the hypothesis that the coracoid expansion in Gnathocharax is homoplastic relative to that in Moojenichthys and Triportheus. The phylogenetic relationships of Clu- peacharax, the last characid genus in our list of genera with expanded coracoids are uncertain. Castro (1981:138) noted a series of similarities between Clupeacharax and Engraulisoma which has non-expanded coracoids. Ongoing studies by one of us (RMCC) are aimed at analyzing the signif- icance of these similarities between C/upea- charax and Engraulisoma. In the interim nonetheless, several features of C/upea- charax bring into question the hypothesis of a close relationship between that genus and Moojenichthys as first proposed by Mi- randa-Ribeiro (1956). Clupeacharax lacks the derived form of the first infraorbital common to Moojenichthys, Triportheus and some species of Brycon (see discussion un- der “‘Infraorbitals’’). Furthermore, C/upea- charax lacks the anterior process of the lat- eral ethmoid common to TJriportheus, Moojenichthys, Brycon, and various other characids (see discussion under “‘Autapo- morphies of Moojenichthys’’). Thus even in the absence of a detailed analysis of the re- lationships of Clupeacharax it is more par- simonious to hypothesize that the enlarge- ment of the coracoids in that genus are homoplastic with respect to those in Moo- Jenichthys and Triportheus. In summary, the presence of expanded coracoids in gasteropelecids and various characids besides Moojenichthys and Trip- ortheus, thus appears to have arisen inde- pendently of that feature in those genera. The possession of asymmetrically vertically enlarged coracoids is consequently pro- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON posed as a synapomorphy for Moojenich- thys and Triportheus. In our introductory discussion we note that Géry (1977:346) commented that the dentition of Moojenichthys is ““approaching [that of] the Tetragonopterinae.”’ The pres- ence of a supraorbital in Moojenichthys myersi argues, however, against a close phy- logenetic alignment of Moojenichthys with tetragonopterines in which that ossification is apparently absent (Weitzman & Fink 1983:391). Infraorbitals. —A second possible syn- apomorphy for Triportheus and Moojenich- thys is found in the infraorbital series. Moo- Jenichthys and Triportheus have the series of ossifications surrounding the orbit typical for characiforms, six infraorbitals, a su- praorbital, and an antorbital. The most no- table feature in these series of ossifications in Moojenichthys and Triportheus involves the form of the first infraorbital and its as- sociation with the second infraorbital. The posteroventral portion of the first infraor- bital in both Moojenichthys and Triportheus extends distinctly ventral of the anteroven- tral portion of the second infraorbital there- by significantly reducing the degree to which the latter element enters into the outer mar- gin of the infraorbital series (Fig. 2). This association of the two anteriormost infraor- bital elements differs from the morphology of these bones in Clupeacharax, Rhaphio- don, Piabucus, Pseudocorynopoma, Carne- giella, Gasteropelecus, Thoracocharax, Cynodon, Hydrolycus, Gnathocharax, the other characid genera known to have ex- panded coracoids. In those other taxa the first and second infraorbitals meet along a straight anteroventrally to posteroventrally aligned juncture without any invasion by the first infraorbital of the area primitively occupied by the anteroventral portion of the second infraorbital. Although the derived form of the first and second infraorbitals in Moojenichthys and Triportheus serves to distinguish those taxa VOLUME 103, NUMBER 3 Fig. 2. anterior to right. Abbreviations: ANT —antorbital; IO—infraorbitals (1 to 6); SO—supraorbital. from other characids with expanded cora- coids, we should note that the occurrence of such modifications of the infraorbitals extends beyond Moojenichthys and Tripor- theus. Howes (1982:5) illustrated a mor- phology of the first and second infraorbitals in Brycon acuminatus (Eigenmann & Nor- ris) apparently comparable to that in Moo- jJenichthys and Triportheus. At least some other species of Brycon, in contrast, have more generalized associations between the first two infraorbitals (e.g., B. meeki, see Weitzman 1962:fig. 8). Brycon has been suggested to be a likely 531 ANT Infraorbitals and supraorbital of Moojenichthys myersi, USNM 304497, 84.8 mm SL, right side, close relative to Triportheus by both Regan (1911:18) and Weitzman (1960:243). In his overview of the former genus Howes (1982: 1) questioned whether Brycon is monophy- letic, but utilized the traditional concept of the genus until future phylogenetic and re- visionary studies can be undertaken. In the absence of such phylogenetic data Howes excluded Triportheus from Brycon on the basis of the presence of the mid-ventral keel in the latter genus; noting that 7riportheus shares, however, all the other diagnostic characters of Brycon. The various similar- ities between Triportheus and Brycon noted SoZ by Regan, Weitzman, and Howes, and the derived features of the infraorbitals cited above for Moojenichthys, Triportheus, and at least some species of Brycon, raise the question of whether the lineage formed by Triportheus and Moojenichthys may be most closely related to some subunit of Brycon. Resolution of that question must await fur- ther revisionary and phylogenetic studies of both Triportheus and Brycon. Autapomorphies of Moojenichthys my- ersi.— Within the lineage formed by Moo- Jenichthys, Triportheus, and possibly Bry- con or some subunit of the latter genus (see immediately preceding discussion), one of the more notable derived features for Moo- Jenichthys myersi (hereafter referred to as Moogjenichthys) is the absence of the single symphysial tooth posterior to the main row of dentition on each dentary. Such sym- physial teeth are common to all members of Triportheus and Brycon, and also occur in Chalceus Cuvier. Chalceus shares all the defining characters of Brycon, but is ex- cluded from Brycon because it possesses a supramaxilla (Howes 1982:1—2). An inner row of teeth on the dentary developed to varying degrees also occur in various char- aciforms in both the New World (e.g., Le- biasinidae, Weitzman 1964:143) and Africa (e.g., diverse genera in the Characidae, see Poll 1957:95, and Distichodontidae, Vari 1979:275—277). This broad phyletic distri- bution of symphysial teeth both in groups proximate to Moojenichthys and other more distantly related characiforms, makes it most parsimonious to hypothesize that the ab- sence of the symphysial dentary teeth in Moojenichthys is a derived loss. The ventrally recurved form of the max- illary dentition and the large number of teeth along the anterior margin of the maxilla dis- tinguish Moojenichthys (Fig. 3) from all examined species of Triportheus. The ven- trally recurved maxillary teeth in Moojen- ichthys are unique within the assemblage formed by that genus, 7riportheus, and pos- sibly Brycon, and are hypothesized to be an autapomorphy for Moojenichthys. In con- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON trast, the large number of teeth (14 to 20) along the anterior margin of the maxilla in Mogjenichthys is more difficult to evaluate. On the one hand that dentition does serve to readily separate Moojenichthys from Triportheus which has only 2 to 4 teeth in that series. Alternatively the species of Bry- con have 10 to 30 teeth along the maxilla (Howes 1982:46), with most species over- lapping the range in tooth number of Moo- Jenichthys to some degree. This common occurrence of large numbers of maxillary teeth in Moojenichthys and Brycon raises the possibility that the relatively few max- illary teeth in Triportheus may be synapo- morphic for the members of that genus, and that the high number of maxillary teeth in Moojenichthys is primitive. A final noteworthy autapomorphy for Moojenichthys involves the anterior portion of the lateral ethmoids. In Moojenichthys, Brycon and Triportheus the anterior surface of the lateral ethmoid bears a distinct pro- cess that extends anteriorly and medially to contact the posterodorsal surface of the vomer (see Weitzman 1962:fig. 3 for an il- lustration of the condition in Brycon meeki). In Moojenichthys the anterior process of the lateral ethmoid is developed into an elon- gate anteriorly-tapering process (Fig. 4) that is significantly longer than comparable pro- cesses in 7riportheus and Brycon. Anterior- ly this elongate anterior process of the lat- eral ethmoid in Moojenichthys contacts a distinct lateral process situated on the dor- sal surface of the vomer. The space between the anterior processes of the paired lateral ethmoids and dorsal of the vomer and para- sphenoid is, in turn, filled by a large cartilage mass comparable to that in Brycon, Tripor- theus, and many other characids. Although the presence of the anterior process of the lateral ethmoid is not unique to Moojenich- thys, the degree of the anterior elongation of the structure is not equalled in other ex- amined characiforms and this modification is thus considered autapomorphic for the genus. The relationship of the anterior process VOLUME 103, NUMBER 3 PMX DEN 533 Fig. 3. Upper and lower jaws of Moojenichthys myersi, USNM 304497, 84.8 mm SL; left side, anterior to left; individual bones separated from positions in life. Abbreviations: DEN—dentary; MX—maxilla; PMX — premaxilla. of the lateral ethmoid to the parasphenoid, vomer, and associated median cartilage in Moojenichthys is also distinctive. When present, the anterior process of the lateral ethmoid in characids usually extends di- rectly along the lateral margin of the para- sphenoid and vomer (e.g., Brycon meeki, see Weitzman 1962:fig. 3). Moojenichthys, in contrast, has a distinct vertical gap be- tween the ventral margin of the anterior process of the lateral ethmoid and the dorsal surface of the vomer. This results in a broad lateral exposure of the median cartilage mass (Fig. 4). This relationship of the lateral eth- moid and proximate bones and cartilages is unknown in Triportheus, Brycon and other examined characiforms, and is consequent- ly hypothesized to represent an additional autapomorphy for Moojenichthys. Moojenichthys Miranda-Ribeiro Moojenichthys Miranda-Ribeiro 1956:546 [type Moojenichthys myersi Miranda-Ri- beiro, by original designation].—Géry 1972:55 [possible derivation from Bry- con].—Géry 1977:346 [similarities with Triportheus noted; dentition compared with that of ““Tetragonopterinae’’]. 534 Lie PARA PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Anterior portion of the neurocranium of Moojenichthys myersi, USNM 304497, 84.8 mm SL; right side, lateral view, anterior to right. Abbreviations: APLE—anterior process of lateral ethmoid; CART — cartilage; LE—main body of lateral ethmoid; PARA —parasphenoid; RH—rhinosphenoid; VO—vomer. Diagnosis. —Within the clade formed by Moojenichthys and Triportheus only the for- mer genus lacks the paired symphysial teeth posterior to the main row of dentary den- tition. Moojenichthys also differs from Trip- ortheus in the numerous ventrally recurved teeth along the anterior margin of the max- illa, and in the derived degree of develop- ment and position of the anterior process of the lateral ethmoid. Moojenichthys can also be distinguished from Triportheus on the basis of its possession of two rather than three rows of teeth on the premaxilla, and in having numerous teeth along much of the anterior margin of the maxilla rather than several teeth limited to the dorsal portions of the maxilla. Remarks. —The derived features de- scribed above are congruent with the hy- pothesis that Moojenichthys forms a mono- phyletic lineage with Triportheus, or with Triportheus and a subunit of Brycon. This conclusion raises the question of whether it iS appropriate to continue to recognize a monotypic Moojenichthys, or whether that genus should be synonymized into Tripor- theus. Mogjenichthys is characterized by a series of autapomorphic features. Within the Characiformes such phenetically distinct VOLUME 103, NUMBER 3 taxa have been traditionally segregated into monotypic genera. We reject the criterion of phenetic distinctness as an a priori basis for the continued recognition of Moojenich- thys, but nonetheless suggest that it is pre- mature to synonymize Moojenichthys into Triportheus. Our decision is a consequence of the present poor knowledge of the phy- logenetic relationships within Triportheus. Until such time as we have a rigorous hy- pothesis of the relationships within Tripor- theus it is impossible to hypothesize wheth- er Moojenichthys is the sister-group to that genus, and thus could be preserved, or sim- ply represents a subunit of 7riportheus. In the latter case the continued recognition of Moojenichthys would result in a paraphy- letic Triportheus. That problem could be re- solved in one of several ways depending on the topology of the phylogenetic tree for the clade consisting of Moojenichthys plus Trip- ortheus. Moojenichthys could be synony- mized into Triportheus along with Coscino- xyron, a genus not presently recognized by most authors. Alternatively the topology of the phylogenetic tree might be such that it would be possible to continue to recognize Moojenichthys either by expanding the def- inition of the genus, or by recognizing one or more Other genera in the clade in addition to Triportheus and perhaps Coscinoxyron. Given these diverse possibilities, we con- tinue to use Moojenichthys in this study. Moojenichthys myersi Miranda-Ribeiro Figs. 1-7, Table 1 Moojenichthys myersi Miranda-Ribeiro, 1956:546-547, fig., type locality: ““Braco River, Ilhéos [=Ihéus], state of Bahia, Brazil.—Géry, 1977:346 [citation; pos- sible relationships]. Diagnosis. —See ““Diagnosis” of Moojen- ichthys above. Description. —Morphometrics of holo- type, paratype and larger examined non-type specimens presented in Table 1. Body elon- gate, distinctly compressed laterally in all 535 specimens greater than 25 mm SL, some- what less so in smaller individuals. Greatest body depth located slightly anterior to ver- tical line through origin of pelvic fin, ap- proximately equal to one-half length of longest pelvic-fin ray in specimens over 50 mm SL; body not as deep and mid-ventral keel less developed in smaller examined specimens. Dorsal profile of head slightly convex from margin of lip to vertical line through posterior nostril, nearly straight from that line to rear of head. Dorsal profile of body slightly convex from rear of head to origin of dorsal fin, posteroventrally slanted and somewhat convex along base of dorsal fin; straight from posterior termina- tion of dorsal fin to adipose fin, and mod- erately concave along caudal peduncle. Dor- sal portion of body obtusely keeled transversely anterior to dorsal fin; trans- versely rounded posterior to fin. Ventral profile of head distinctly convex over lip, straight along anteroventral margin of jaw, and distinctly convex ventral to joint with quadrate. Ventral profile of body irregular, distinctly convex overall; very slightly con- vex from isthmus nearly to vertical line through origin of pectoral fin; convexity greater from that line to origin of pelvic fin; straight to slightly concave from origin of pelvic fin to anterior termination of anal fin; straight and posterodorsally slanted along base of anal fin; slightly concave along dor- sal peduncle. Distinct mid-ventral keel ex- tending from isthmus to between origins of pelvic fins; keel less developed in specimens under 20 mm SL; increasingly obvious in individuals over 30 mm SL, most devel- oped in specimens of over 50 mm SL. Scales along margin of keel flat, not folded over edge of keel. Head obtusely pointed in profile; mouth terminal, lower jaw longer than upper, with dentigerous portion of maxilla distinctly an- gled posteroventrally. Maxilla extending posteriorly under orbit to vertical line through anterior margin of pupil. Nostrils of each side close together; anterior opening 536 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Morphometrics of Moojenichthys myersi. Standard length is expressed in mm; measurements | to 14 are percentages of standard length; 15 to 18 are percentages of head length. Dashes indicate measurement that could not be taken due to damage to holotype. Range includes values for 24 specimens (holotype, MNRJ 4127; paratype MNRJ 4128; and 22 of the larger non-type specimens out of USNM 304497, MZUSP 40227, MNRJ 11605, and ANSP 164288), with the exception of length of the longest dorsal-fin ray which is based on 23 specimens, and length of the longest pectoral-fin ray which is based on 22 specimens. Holotype Paratype Range Mean Standard length 96.6 85.8 27.0-96.6 69.1 1. Greatest body depth 25.7 25.5 22.2-28.7 27.0 2. Snout to dorsal-fin origin 65.2 62.0 60.7-65.4 62.8 3. Length of base of dorsal fin 8.2 U5 7.1-8.7 7.8 4. Posterior terminus of dorsal fin to adipose fin 18.0 17.8 16.4-19.8 18.0 5. Posterior terminus of dorsal fin to caudal-fin base 28.9 29.6 27.9-32.4 29.8 6. Snout to origin of pelvic fin 49.6 50.5 46.7-51.3 49.2 7. Snout to origin of anal fin 68.7 67.6 62.2-68.7 65.3 8. Length of base of anal fin 31.5 28.3 28.1-32.1 30.4 9. Length of caudal peduncle 8.7 9.8 8.0-10.6 9.2 10. Length of longest dorsal-fin ray 16.8 15.3 15.3-18.6 16.8 11. Length of longest pectoral-fin ray _ 32.1 24.4—32.1 30.1 12. Length of longest pelvic-fin ray 14.7 13.6 11.1-15.0 13.5 13. Least depth of caudal peduncle 7.6 7.0 6.3-8.7 7.9 14. Head length 24.1 24.2 23.7-29.0 24.6 15. Snout length 23.6 22.6 20.0-27.0 23.5 16. Orbital diameter 36.1 36.5 31.7-38.9 36.3 17. Postorbital head length 38.2 36.5 29.3-41.8 38.1 18. Interorbital width 24.0 24.0 19.1-27.6 24.5 circular, posterior kidney-shaped. Eye rel- atively large, without adipose eyelid. Me- dian fronto-parietal fontanel well devel- oped; completely separating parietals; frontals in contact only anteromedially and at epiphyseal bar. Fontanel becoming pro- gressively wider posteriorly, extending onto dorsomedial surface of supraoccipital. Infraorbital series complete (Fig. 2), all infraorbitals with laterosensory canal seg- ments. Sixth infraorbital (dermosphenotic) with single tubular laterosensory canal seg- ment. First infraorbital expanded antero- ventrally, with distinctly convex anterior Margin, anterior portion extends over lat- eral surface of maxilla; laterosensory canal segment with three sections in larger spec- imens. Supraorbital and antorbital present. Four branchiostegal rays, first three at- tached to anterior ceratohyal, fourth to pos- terior ceratohyal. Gill-rakers relatively elongate, 17 or 18+1+40 to 42 rakers on outermost gill-arch (in 2 larger cleared and counterstained specimens). Lower jaw with one row of 12 to 14 teeth on each side on each dentary (Fig. 3); num- ber of teeth greater in largest specimens; in- ner row consisting of single symphysial tooth absent. Anterior 5 teeth on dentary notably larger than remainder, with 5 cusps, medial cusp distinctly largest. Remaining teeth usually tricuspidate, rarely unicuspidate, with largest cusp recurved somewhat pos- teriorly. Teeth on premaxilla in two rows; teeth of inner row larger (Fig. 3). Four tri- cuspidate teeth of approximately equal size in outer row. Six teeth in inner row on pre- maxilla; 2 medial teeth largest, subequal; remaining teeth gradually becoming smaller laterally; medial tooth tricuspidate, remain- ing teeth in row with 5 cusps. Lateral tooth of inner row of premaxilla approximates dorsal tooth on maxilla. Anterior margin of maxilla distinctly convex, with single row VOLUME 103, NUMBER 3 537 Fig. 5. holotype, 96.6 mm SL. of teeth (Fig. 3). Teeth on maxilla distinctly smaller than smallest tooth on premaxilla; typically unicuspidate in specimens of ap- proximately 27 mm SL, bicuspidate or usu- ally tricuspidate in specimens of about 40 mm SL and greater. Smaller specimens with 2 to 9 teeth limited to upper one-quarter to one-half of anterior margin of maxilla; larg- er specimens with 14 to 20 teeth arranged along nearly entire anterior margin of max- illa. Largest cusp of teeth on maxilla re- curved ventrally. Scales cycloid, thin, relatively large. Lat- eral line distinctly decurved ventrally, com- pletely pored from supracleithrum to base of middle rays of caudal fin. Forty to 43 [42] scales in lateral line (70% of specimens with 42 scales); 6 or 7 [7] scales in transverse series from origin of dorsal fin to lateral line; 3 or 4 [3] scales in transverse series from origin of pelvic fin to lateral line (4 scales present in only 1 specimen); 3 or 4 [3] scales in transverse series from origin of anal fin to lateral line; 17 to 21 [20] scales along mid- dorsal line between tip of supraoccipital process and origin of dorsal fin (60% of spec- imens with 19 or 20 scales); 9 or 11 [11] scales along mid-dorsal line between pos- terior termination of dorsal fin and adipose fin (91% of specimens with 10 or 11 scales); 13 to 15 [13] horizontal scale rows around caudal peduncle (86% of specimens with 14 or 15 scales). Dorsal-fin rays 11,8 or 9 or 111,9 [11,9] (41,9 Moojenichthys myersi, Brazil, Bahia, “Braco river’ ,[=Rio do Braco], Ilhéos [=Ilhéus]; MNRJ 4127, most common); anal-fin rays iv,31 to 35, or v,33 [iv,33] (iv,33 most common); pectoral- fin rays 1,9 to 12 followed by O to ii un- branched rays [1, 10,1] G, 10,1 most common); pelvic-fin rays 1,6 [1,6]; principal caudal-fin rays 10/9 [10/9]. Dorsal fin profile obtusely acute, poste- rior unbranched and first branched ray sub- equal; posterior unbranched ray typically slightly longer. Dorsal fin situated on pos- terior half of body; origin of fin located slightly posterior of vertical line through an- terior terminus of anal fin, closer to base of caudal fin than to tip of snout. Longest di- mension of adipose fin approximately equal to horizontal width of pupil; origin of adi- pose fin slightly anterior of vertical line through posterior terminus of anal fin. Pec- toral fin large, profile distinctly acute; when fin depressed, tip extends to vertical line approximately two-thirds distance along pelvic fin. Pelvic fin profile obtusely acute, origin of fin at posterior margin of mid- ventral keel, tip of depressed fin extending posteriorly slightly beyond anus, but falling short of anterior terminus of anal fin. Ax- illary pelvic scale present, its length about one-third that of longest pelvic-fin ray. Cleared and counterstained 84.8 mm SL male with 6 to 15 basally directed bony hooks along posterior margins of first 5 branched pelvic-fin rays. Ventral margin of anal fin somewhat rounded anteriorly, with last unbranched and first branched rays 538 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Moojenichthys myersi, Brazil, Bahia, Rio do Braco, 2 km SW of town of Rio do Braco, on Fazenda Luzia, USNM 304496, 33.4 mm SL. longest, subequal, following’ 10 to 12 branched rays rapidly decreasing in length, remaining anal-fin rays slowly decreasing in length. Males with 1 to 8 basally-directed bony hooks along posterior margins of dis- tal sections of 7 to 14 longest anal-fin rays. Caudal fin forked, lobes obtusely pointed. Total vertebrae 39 (2), 40 (20), 41 (2) [40]. Color in life. — Descriptions based on col- or transparencies of a series of recently pre- served specimens captured in August 1988 and February 1989. Overall coloration of specimens ranging from 15.9 to 33.4 mm SL clear to yellowish. Iris, lower jaw, in- fraorbital region, opercle, and peritoneum silvery. Tip of lower jaw, snout, dorsal por- tions of body and basal portions of caudal fin light yellow. Traces of yellow pigmen- tation apparent on dorsal, adipose, and anal fins. Other fins hyaline. Mid-lateral dark stripe on body quite obvious, but somewhat masked anteriorly by guanine. Dark stripe above anal fin, and dark pigmentation on all fins clearly visible. Specimens above approximately 50 mm SL bright silver overall, somewhat darker along dorsal portions of head and body. Dense guanine on scales completely mask- ing both dark stripes along mid-lateral sur- face of body, and those on ventrolateral sur- face of body above anal fin. Dark pigmentation on fins as in preserved spec- imens. Coloration in preservative. —Overall ground color of specimens fixed in formalin and lacking guanine on scales yellowish brown. Dense fields of small, dark chro- matophores on upper lip, snout, and dorsal surface of head (Figs. 5-7). Very intense, horizontally elongate stripe of dark pigmen- tation along dorsal portion of lower lip; less intense dark pigmentation ventral to this on lower jaw. Scattered small dark chromato- phores on lateral surface of head anterior to orbit and on opercle; pigmentation more obvious in specimens totally lacking gua- nine on head. Body with mid-lateral stripe of small, dark chromatophores extending from supra- cleithrum posteriorly to caudal peduncle; stripe gradually expanding vertically pos- teriorly; broadened into distinctly wider dif- fuse dark spot on lateral surface of caudal peduncle. Body dorsal of dark mid-lateral stripe with margins of scales outlined by dark chromatophores; pattern most ob- vious in medium sized specimens, some- what obscured by overall dusky appearance of dorsal portions of body in larger speci- mens. Stripe of dark chromatophores ex- tending posterodorsally along ventrolateral portion of body from slightly anterior of vertical through anterior terminus of fin posteriorly to posterior terminus of fin. Stripe wider and distinctly separated from base of anal fin anteriorly, becoming grad- ually narrower and approaching base of anal- fin rays posteriorly. VOLUME 103, NUMBER 3 539 Fig. 7. Moojenichthys myersi, Brazil, Bahia, Rio do Braco, 2 km SW of town of Rio do Braco, on Fazenda Luzia, MZUSP 40227, 81.6 mm SL. Dorsal fin with dense field of dark chro- matophores along distal portions of rays and membranes. Margins of adipose fin in larger specimens outlined by scattered dark chro- matophores. Anterior rays of pectoral and pelvic fins outlined by series of small dark chromatophores. Anterior margin and dis- tal portions of anal fin dusky, most rays outlined distally by dark chromatophores. Caudal fin dusky in specimens of all sizes, rays outlined by series of small dark chro- matophores. Common names. —Brazil, Bahia, Ilhéus: ““Mossarupé,” “Piaba-faca,” and “Cani- vete”’ (Miranda-Ribeiro, 1956:547). During the 1988 and 1989 expeditions the only name used by local fishermen was “‘Piaba- faca.” Ecology. —Specimens collected during the 1988 and 1989 expeditions were collected in black waters containing limited suspend- ed material. The area surrounding the river was originally a portion of the Atlantic Coastal Forest, but much of the understory vegetation has been replaced by cocoa trees. In the areas sampled for fishes the Rio do Braco was between 10 and 25 m wide, ranged from 1.5 to 3 m deep, and had mats of floating vegetation along its margins. The bottom was sandy-mud with scattered boul- ders. Other fishes captured with Moojenichthys myersi and the families to which they are presently assigned were Steindachnerina elegans (Curimatidae); Nematocharax ve- nustus, Oligoscarcus macrolepis, Astyanax sp., Characidium sp. (Characidae); Hoplias sp. (Erythrinidae); Rhamdia sp. (Pimelod- idae); Poecilia sp. (Poeciliidae); Astronotus ocellatus and Geophagus brasiliensis (Cich- lidae). The Astronotus ocellatus record rep- resents an introduction. Diet. —Examination of the stomach con- tents of the three cleared and counterstained specimens shows that the species eats mos- quito larvae and other aquatic inverte- brates. Distribution. —Known only from the Rio do Braco in the state of Bahia, Brazil. The original description of Moojenichthys my- ersi states that the type material was col- lected in the “Brago river, Ilhéos.” In ac- tuality the mouth of the “Braco river” [=Rio do Braco] is located approximately 7 km along the coast north of the city of Ilhéus (‘‘Ilhéos”’ of Miranda-Ribeiro). Limited ichthyological collecting has taken place in the Bahian coastal drainages near the Rio do Braco. Thus the lack of records of Moo- Jenichthys myersi from other neighboring river systems may be a consequence of poor sampling. Material examined. —Brazil. Bahia: “Braco river,” Ilhéos [=Ilhéus], MNRJ 540 4127, 1 specimen, holotype, 96.6 mm SL; same locality, MNRJ 4128, 1 specimen, paratype, 85.8 mm SL; Ilhéus, Fazenda Pi- rataquicé, MNRJ 5572, 1 specimen, 78.2 mm SL (locality not found in examined gaz- etteers or maps); Rio do Braco, 2 km SW of town of Rio do Braco (approx. 14°39’S, 39°16’'W), on Fazenda Luzia, USNM 304497, 9 specimens, 20.5-84.8 mm SL (1 specimen, 84.8 mm SL, cleared and coun- terstained); USNM 304496, 15 specimens, 17.8-33.4 mm SL (1 specimen, 27.0 mm SL, cleared and counterstained); MZUSP 40226, 15 specimens, 15.9-30.5 mm SL; MZUSP 40227, 9 specimens, 19:6—84.1 mm SL (1 specimen, 56.0 mm SL, cleared and counterstained); ANSP 164287, 3 speci- mens, 17.4—26.5 mm SL; ANSP 164288, 2 specimens, 63.4-76.0 mm SL; MNRJ 11604, 3 specimens, 20.1—24.4 mm SL; MNRJ 11605, 2 specimens, 63.4—74.9 mm SL. Comparative cleared and stained material examined. —Brycon falcatus, USNM 226161, 2 specimens. Carnegiella strigata, USNM 225245, 5 specimens. Clupeacha- rax anchoveoides, USNM 302245, 1 spec- imen. Cynodon gibbus, USNM 270338, 2 specimens. Engraulisoma_ taeniatum, USNM 302225, 1 specimen. Gasteropelecus sternicla, USNM 226337. Gnathocharax steindachneri, USNM 278995, 2 specimens. Rhaphiodon vulpinus, USNM 231549, 3 specimens. Triportheus angulatus, USNM 270343, 2 specimens. Triportheus sp.., USNM 280498, 4 specimens; USNM 258079, 2 specimens. Acknowledgments A number of the specimens of Moojenich- thys myersi that served as the basis for this paper was collected during a collaborative FFCLRP-USP and USNM expedition in the state of Bahia, Brazil. The success of the collecting effort was assured by the enthu- siasm of Susan L. Jewett (USNM), Hertz F. Santos, Maura H. Manfrin, Eliseu B. Dias, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and Eduardo Castadelli, Jr. (all of FFCLRP- USP) who unstintingly assisted in the often difficult fishing efforts. Hertz F. Santos made a special second trip into the region of the type-locality to collect additional material of the species, including an extensive series that included most of the larger individuals reported on in this paper. Figures 5 to 7 were prepared by Mr. Theophilus B. Gris- wold. Sandra Raredon and Jeffrey Howe as- sisted in the research efforts at USNM. Ms. India Moreira (MNRJ) arranged for the loan of the holotype and paratype of Moojenich- thys myersi, and Mr. Luis Paulo S. Portugal (MZUSP) and Dr. Naércio A. Menezes (MZUSP) facilitated our examination of those specimens. This paper was improved by the suggestions of Dr. Stanley H. Weitz- man (USNM), Dr. Wayne C. Starnes (USNM), Dr. William L. Fink (University of Michigan, Museum of Zoology), Dr. Dar- rell J. Siebert (British Museum (Natural History)), and Mrs. Marilyn Weitzman (USNM). We thank all of the above for their assistance. Extensive logistical support for the ex- pedition that collected most of the speci- mens of Moojenichthys myersi was provid- ed by FFCLRP-USP. Funding for that collecting effort was provided through the Neotropical Lowland Research Program of the International Environmental Sciences Program of the Smithsonian Institution. That program also supported the research associated with this project both at mu- seums in Brazil and at the Smithsonian In- stitution. Literature Cited Castro, R.M.C. 1981. Engraulisoma taeniatum, um novo género e espécie de Characidae da bacia do Rio Paraguai (Pisces, Ostariophysi). — Papéis Avulsos de Zoologia, Sao Paulo 34(11):135-139. Cope, E. D. 1872. On the fishes of the Ambiyacu River. — Proceedings of the Academy of Natural Sciences of Philadelphia (for 1871):257—269. Cuvier, G. & A. Valenciennes. 1849. Histoire natu- relle des poissons. Volume 22:1-395. VOLUME 103, NUMBER 3 Fink, W. L., & S. H. Weitzman. 1974. The so-called cheirodontin fishes of Central America with de- scriptions of two new species (Pisces: Charac- idae).—Smithsonian Contributions to Zoology 172:1—46. Fowler, H.W. 1906. Further knowledge of some het- erognathous fishes. Part IJ.— Proceedings of the Academy of Natural Sciences of Philadelphia (for 1906):431—488. Geéry, J. 1972. Poissons characoides des Guyanes, I. Généralitées. II. Famile des Serrasalmidae. — Zoologische Verhandelingen 122:1—250. 1977. Characoids of the world. TFH Publi- cations, Neptune City, New Jersey, 672 pp. Greenwood, P. H., D. E. Rosen, S. H. Weitzman, & G. S. Myers. 1966. Phyletic studies of teleos- tean fishes, with a provisional classification of living forms.—Bulletin of the American Mu- seum of Natural History 131(4):339-456. Howes, G. J. 1976. The cranial musculature and tax- onomy of characoid fishes of the tribes Cyno- dontini and Characini.— Bulletin of the British Museum (Natural History), Zoology 29:203-248. . 1982. Review of the genus Brycon (Teloestei: Characoidei).— Bulletin of the British Museum (Natural History), Zoology 43(1):1—47. Miquelarena, A. H., & J. R. Casciotta. 1982. Presen- cia en la Argentina de Clupeacharax anchov- eoides Pearson, 1924.—Limnobios 2(5):333- 335. Miranda-Ribeiro, P. de. 1941. Notas para o estudo dos characinidos brasileiros (Peixes—Género Chalcinus Cuv. & Val. 1849).—Papeis Avulsos do Departamento do Zoologia 1(18):159-174. 1956. On a new genus and new species of South American fishes.— Proceedings of the 14th International Congress of Zoology, Copenha- gen, 16(Vertebrates):546—-547. Myers, G.S. 1940. Suppression of Acropsis and Chal- cinus, two preoccupied generic names of South American fresh-water fishes. — Stanford Ichthy- ological Bulletin 1(5):170. Ortega, H., & R. P. Vari. 1986. Annotated checklist of the freshwater fishes of Peru.—Smithsonian Contributions to Zoology 437:1-25. Pearson, N. E. 1924. The fishes of the eastern slope of the Andes. I. The fishes of the Rio Beni basin, Bolivia, collected by the Mulford Expedition. — Indiana University Studies 11(64):1—63. Poll, M. 1957. Les genres des poissons d’eau douce de l’Afrique.—Annales du Musée Royal du Congo Belge, Series Zoologiques 54:1-191. Regan, C.T. 1911. The classification of the teleostean fishes of the order Ostariophysi.— 1. Cyprinoi- dea.— Annals and Magazine of Natural History, ser. 8, 8:14—32. Roberts, T. R. 1969. Osteology and relationships of 541 the characoid fishes, particularly the genera Hepsetus, Salminus, Hoplias, Ctenolucius, and Acestrorhynchus.— Proceedings of the Califor- nia Academy of Sciences, fourth series, 36(15): 391-500. . 1973. Osteology and relationships of the Pro- chilodontidae, a South American family of cha- racoid fishes. — Bulletin of the Museum of Com- parative Zoology 145(4):213-235. . 1974. Osteology and classification of the neo- tropical characoid fishes of the families Hemi- odontidae (including Anodontinae) and Paro- dontidae.—Bulletin of the Museum of Comparative Zoology 146(9):41 1-472. Sazima,I. 1983. Scale-eating in characoids and other fishes. Environmental Biology of Fishes 9(2): 87-101. Schultz, L. P. 1944. The fishes of the family Chara- cinidae from Venezuela, with descriptions of seventeen new forms.—Proceedings of the United States National Museum 95:235-367. Starks, E. C. 1930. The primary shoulder girdle of the bony fishes.—Stanford University Publica- tions, University Series, Biological Sciences 6(2): 149-239. Stewart, D. J., R. Barriga S., & M. Ibarra. 1987. Ic- tiofauna de la cuenca del Rio Napo, Ecuador Oriental: lista anotada de especies. — Politecni- ca, revista de Informacion Técnico-Cientifica, 12(4):9-63. Vari, R. P. 1977. Notes on the characoid subfamily Iguanodectinae, with a description of a new species.—American Museum Novitates 2612: 1-6. 1979. Anatomy, relationships and classifi- cation of the families Citharinidae and Distich- odontidae (Pisces, Characoidea).— Bulletin of the British Museum (Natural History) 36(5):261- 344. 1986. Serrabrycon magoi, a new genus and species of scale-eating characid (Pisces: Characi- formes) from the upper Rio Negro.—Proceed- ings of the Biological Society of Washington 99(2):328-334. Weitzman, S. H. 1954. The osteology and relation- ships of the South American characid fishes of the subfamily Gasteropelecinae. — Stanford Ich- thyological Bulletin 4(4):213-263. . 1960. The phylogenetic relationships of Trip- ortheus, a genus of South American characid fishes.—Stanford Ichthyological Bulletin 7(4): 239-244. 1962. The osteology of Brycon meeki, a gen- eralized characid fish, with an osteological def- inition of the family.—Stanford Ichthyological Bulletin 8(1):1-77. . 1964. Osteology and relationships of the South PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON America characid fishes of the subfamilies Le- biasininae and Erythrininae with special refer- ence to the subtribe Nannostomina.— Proceed- ings of the United States National Museum 116: 127-170. , S. V. Fink, & N. A. Menezes. 1985. Appendix 3. A critique of the monophyly of the Glandu- locaudinae. Pp. 112-121 in S. H. Weitzman and S. V. Fink, eds., Xenurobryconin phylogeny and putative pheromone pumps in glandulocaudine fishes (Teleostei: Characidae). Smithsonian Contributions to Zoology 421:1-121. —., & W. L. Fink. 1983. Relationships of the neon tetras, a group of South American fresh- water fishes (Teleostei, Characidae), with com- ments on the phylogeny of New World characi- forms. — Bulletin of the Museum of Comparative Zoology 150(6):339-395. —, & R. P. Vari. 1987. Two new species and a new genus of miniature characid fishes (Teleos- tei: Characiformes) from northern South Amer- ica.— Proceedings of the Biological Society of Washington 100:640-652. (RMCC) Departmento de Biologia, Fa- culdade de Filosofia, Ciéncias e Letras de Ribeirao Preto, Universidade de Sao Paulo, Avenida Bandeirantes 3900, 14049 Ribei- rao Preto SP, Brazil; and (RPV) Department of Vertebrate Zoology (Fishes), National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 543-549 SCORPAENODES IMMACULATUS, A NEW SPECIES OF SCORPIONFISH (OSTEICHTHYES: SCORPAENIDAE) FROM WALTERS SHOALS, MADAGASCAR RIDGE Stuart G. Poss and Bruce B. Collette Abstract.—Scorpaenodes immaculatus is described from a single specimen collected in shallow water (40-49 m) at Walters Shoals in the southwestern Indian Ocean. It differs from all known species of the genus in lacking dark markings on the head, body, and fins. The new species is most similar to Scorpaenodes smithi Eschmeyer & Rama Rao, but differs from it in the absence of a dark spot over the posterior part of the spinous dorsal fin, the absence of bars and other dark markings over the body, in having a small spine on the upper arm of the preopercle, and in having the upper unbranched rays in the pectoral fin slightly elongate. A shallow water trawl collection made in December 1988 during cruise 17 of the So- viet oceanographic vessel Vityaz in the southwestern Indian Ocean (Collette & Pa- rin 1991) resulted in the discovery of a new scorpionfish of the genus Scorpaenodes Bleeker. The new species, represented by a single specimen, was taken in shallow water (40-49 m) at Walters Shoals, an isolated submerged oceanic mountaintop that rises to within 18 m of the surface, 400 nautical miles south of Madagascar and 600 nm east of South Africa (33-35°S, 43°50—-56’E). Scorpaenodes is distinguished from other scorpaenid genera by the following combi- nation of features: a low spinous dorsal fin, usually with XIII spines (sometimes XIV), and absence of an occipital pit and palatine teeth. Like many other scorpionfish genera, it is poorly defined. The genus has been di- vided by some authors (e.g., Smith 1957), but recent work has followed Matsubara (1943) and Eschmeyer (1969a) in recogniz- ing a single genus. With the description of Scorpaenodes immaculatus, the genus now includes about 25 species. The purposes of this paper are to describe the new species so that the name is available for subsequent analysis of the Walters Shoals fish fauna, and to compare it with other species of the widely distributed genus Scor- paenodes. In addition, we take the oppor- tunity to list the described species in the genus (Table 1). Materials and methods. —Counts and head spine terminology follow those of Eschmeyer (1969b). Measurements were taken as specified in Poss (1982). Scale ter- minology and the methods of cleaning the scales follow Hughes (1981), except the con- centration of sodium hypochlorite was re- duced to 0.5%. Scales were photographed using an Olympus SZH binocular light mi- croscope. The acronym USNM designates the Di- vision of Fishes, National Museum of Nat- ural History, Smithsonian Institution, Washington, D.C., where the holotype and only specimen is deposited; CAS, the Cal- ifornia Academy of Sciences in San Fran- cisco which houses much of the comipara- tive material. Scorpaenodes immaculatus, new species Figs. 1-3 Holotype. —USNM 307748; 89.2 mm SL; western Indian Ocean, Walters Shoals, 544 Table 1.—Nominal species of Scorpaenodes and their distributions. Species Author africanus Pfaff 1933 africanus Smith 1958 albaiensis Evermann & Seale 1907 arenai Torchio 1962 asperrimus Smith 1958 brocki Schultz 1956 caribbaeus Meek & Hildebrand 1928 corallinus Smith 1957 elongatus Cadenat 1949 englerti Eschmeyer & Allen 1971 erinacea Garman 1903 floridae Hildebrand 1940 guamensis Quoy & Gaimard 1824 hirsutus Smith 1957 immaculatus Poss & Collette 1990 insularis Eschmeyer 1971 investigatoris Eschmeyer & Rama Rao 1972 keelingensis Marshall 1950 kelloggi Jenkins 1903 littoralis Tanaka 1917 minor Smith 1958 minutus Cuvier 1829 muciparus Alcock 1889 parvipinnis Garrett 1864 polylepis! Bleeker 1851 scaber Ramsay & Ogilby 1886 smithi Eschmeyer & Rama Rao 1972 steeni Allen 1977 Steinitzi Klausewitz & Froiland 1970 tredecimspinosa Metzelaar 1919 tribulosus Eschmeyer 1969 varipinnis Smith 1957 xyris PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Jordan & Gilbert 1882 Distribution eastern Atlantic = albaiensis Indo-West Pacific Mediterranean Sea = parvipinnis = minor western Atlantic Indo-West Pacific eastern Atlantic Easter Island = guamensis = tredecimspinosus Indo-West Pacific Indo-Pacific Walters Shoals St. Helena western Indian O. = ?kelloggi western Pacific Indo-West Pacific Indo-West Pacific = ?guamensis Indo-West Pacific Indo-West Pacific = guamensis = ?guamensis western Indian O. Western Australia Red Sea western Atlantic O. western Indian O. Indian Ocean eastern Pacific O. 1 Type-species of Scorpaenodes Bleeker 1857. 33°11'S, 43°52'E; 40-49 m; 29-m fish trawl; Vityaz Cruise 17, Sta. 2685; 12 December 1988; only known specimen. Diagnosis.—An entirely red species of Scorpaenodes without dark markings on body and fins. It is also separable from other species of Scorpaenodes by a combination of differences in spination, counts, and mea- surements (see below, comparisons). Description. —Dorsal-fin rays XIV, 81, (count of XIII, 912 should be expected). Anal-fin rays III, 5%. Pectoral-fin rays 19, rays 2-8 or 9 branched. Lateral line dam- aged, but with about 27 scales, with last scale extending over base of caudal fin. Gill rakers 5+ 1+ 10 (left); 6+1+10 (right). Cau- dal fin with 12 branched rays (6 ventral; 6 dorsal), 15 segmented rays (7 ventral; 8 dor- sal), and 8 unsegmented, procurrent rays (4 dorsal; 4 ventral). Vertebrae 9 precaudal + 15 caudal = 24. Head moderately large; relatively deep posteriorly. Infraorbital one with suborbital ridge, but without spine. Three lobes on in- fraorbital one extending over maxilla, one anteriorly, two posteriorly; none ending in pungent spine. Infraorbital two with strong ridge ending in distinct spine near junction VOLUME 103, NUMBER 3 Fig. 1. with third infraorbital bone. Third infraor- bital bone forming strong ridge ending in spine just anterior to preopercular spine. Nasal spine stout, slightly curved. Interor- bital broad, shallow, with weak ridges; right ridge ending in small spine. Preocular spine sharp. Supraocular spine stout. Postocular spine strong. Tympanic spine strong. Co- ronal spines absent. Sphenotic spine absent. Pterotic spine small. Parietal and nuchal spines notably stout; broad at base. Upper posttemporal spine absent. Lower posttem- poral spine small. Supracleithral spine prominent. Cleithral spine strong. Posterior margin of the preoperculum with 3 distinct spines: largest at end of suborbital ridge (stay); second ventral to but near first; third larger than second. Fourth and fifth pre- opercular spines absent. Anterior margin of dorsal arm of preopercle with small spine projecting laterally and slightly posteriorly. Supplemental preopercular spine a small point at base of largest preopercular spine. Operculum with 2 prominent strong spines, extending posteriorly beyond edge of bone. Maxilla extending almost to vertical with posterior border of orbit; without scales. In- fraorbital bones and cheek covered with 545 Holotype of Scorpaenodes immaculatus in lateral view (USNM 307748, 89.2 mm SL). Walters Shoals. scales. Operculum covered with scales, ex- cept posteriorly. Small relatively simple cir- rl present on many head spines; the longest and most elaborate cirri on supraocular spine. Body scales of moderate size, with about 50 vertical rows as counted above lateral line (damaged or missing, counted from scale pockets); scales from dorsum with about 40 ctenii in 2 alternating marginal rows and about 4 submarginal rows of ctenial bases in posterior field (Fig. 3a). Underside of head without scales. Scales on posterior part of head and on body ctenoid. Cirri absent on body. Dorsal fin origin above middle of opercle. Anterior dorsal-fin spines tipped with sim- ple cirri. Anal fin with second spine notably stronger and longer than third. Upper un- branched pectoral-fin rays slightly elongate; extending posteriorly to first segmented anal ray; ventral rays notably thicker. Caudal fin somewhat rounded, but with rays in ventral half slightly longer than those dorsally. Cau- dal skeleton with haemal spine of second preural centrum broad, weakly ankylosed to second preural centrum, not supporting caudal-fin rays; parhypural broad, free from 546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Head of holotype of Scorpaenodes immaculatus in dorsal view. hypurals supporting ventral procurrent rays, nous from compound urostylar centrum and partially ankylosed to compound urostylar hypural 3; hypural 3 supporting 3 branched centrum; hypurals | and 2 fused, supporting rays; hypural 4 supporting 2 branched and 6 branched rays and | unbranched, autoge- 1 unbranched ray; hypurals 3 and 4 anky- VOLUME 103, NUMBER 3 547 We < - ; ay) At: Via “2 oe ws sg “i Fig. 3. Comparison of scales from dorsum above lateral line and right pectoral fin in several species of Scorpaenodes. a. Holotype of Scorpaenodes immaculatus USNM 307748. b. S. varipinnis, CAS 48692. c. S. parvipinnis, CAS 31352. d. S. tribulosus, CAS 24267. Bars represent 1.0 mm. Note differences in number of ctenial bases, size and number of ctenii. losed only near compound urostylar cen- trum, distinctly separate posteriorly; hy- pural 5 autogenous; 3 broad epurals; uroneural long, autogenous; neural spine of second preural centrum broad, short. Body uniformly red immediately after capture, with small red spots over soft fin rays and no dark brown markings. No dark brown or black spot on posterior part of spinous dorsal fin. Spinous dorsal fin with more red pigment proximally. Body devoid of markings in alcohol. Measurements for the holotype in mm are as follows (percentage standard length in pa- rentheses): standard length 89.2, head length 38.2(43), snout 11.0(12), orbit 10.1(11), in- terorbit 6.0(7), upper jaw 19.9(22), postor- bit 17.8(20), body depth 31.6(35), predorsal 36.0(40), anal fin 24.6(28), caudal fin 22.6(25), pectoral fin 30.6(34), pelvic fin 22.1(25), first dorsal spine 4.4(5), second dorsal spine 7.3(8), third dorsal spine 9.9(11), fourth dorsal spine 12.2(14), fifth dorsal spine 13.6(15), penultimate dorsal spine 6.6(7), last dorsal spine 9.9(11), first anal spine 7.1(8), second anal spine 18.6(21), third anal spine 14.0(16), maximum width of interorbital ridge 2.3(3), caudal peduncle 9.6(11), snout to second dorsal spine 37.3(42), snout to third dorsal spine 39.6(44), snout to fourth dorsal spine 43.1(48), snout to fifth dorsal spine 46.6(52), first dorsal spine width at midlength 0.5(0.6), interorbital depth 0.9(1), incision of fin membrane at fourth dorsal spine 5.6(6), snout to pelvic insertion 37.7(42), opercular tip to dorsal fin 11.4(13), uppermost pre- opercular spine 4.3(5), first dorsal spine to fifth dorsal spine 10.1(11), fifth dorsal spine to pelvic insertion 32.7(37), first dorsal spine to pelvic insertion 31.7(36), fifth dorsal spine to last dorsal spine 24.1(27), last dorsal spine to last dorsal ray 16.4(18), last dorsal ray to last anal ray 12.6(14), anal-fin origin to last anal ray 11.8(13), pelvic insertion to anal- fin origin 31.1(35), first dorsal spine to anal- 548 fin origin 46.0(52), last dorsal spine to pelvic insertion 42.4(48), last dorsal spine to last anal ray 21.1(24), last dorsal ray to anal-fin origin 22.8(26), last dorsal spine to anal-fin origin 25.8(29), fifth dorsal spine to anal- fin origin 38.2(43). Comparisons. —Scorpaenodes immacu- latus most closely resembles S. smithi and S. investigatoris but differs from both in lacking any dark markings on head, fins, or body. These three species share the follow- ing traits: similar spination on the infraor- bital bones, absence or near absence of coronal spines, similar counts and body proportions. With respect to coloration, S. immaculatus resembles a specimen identi- fied as S. smithi that was photographed by Gloerfelt-Tarp & Kailola (1984:114), but does not fit their description, which noted the presence of a spot on the dorsal fin. De- tailed study of geographic variation in S. smithi is needed. Scorpaenodes immaculatus is readily dis- tinguished from its western Indian Ocean congeners by its coloration, among other features. It differs most notably from S. smithi in lacking a dark spot in the posterior part of the spinous dorsal fin and the bars and markings present on the body in S. smithi (but see below), in having a small spine on the upper arm of the preopercle, in having XIV (if normal) rather than XIII dorsal-fin spines, and in having the upper unbranched rays in the pectoral fin slightly elongate. It differs from S. investigatoris in having the upper portion of the pharynx light-colored, rather than dusky-colored. Scorpaenodes immaculatus can be separat- ed from S. tribulosus in having 3, as opposed to 4-8, spinous points on the suborbital ridge, and with less well-developed ctenti on the scales, a difference especially evident in scales from the chest, maxilla, and inter- orbital regions, which are thickly covered with ctenoid scales in S. tribulosus. It can be quickly distinguished from S. steinitzi by having one more dorsal fin spine and fewer segmented dorsal rays (81 vs. 10), in having PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 19 as opposed to 16-17 pectoral rays, and lacking coronal spines. Scorpaenodes im- maculatus has only 3 suborbital spines, un- like S. parvipinnis, which typically has from 5 to 10. The new species differs from S. hirsutus in having cirri confined to head and fin spines and not widely distributed over the body, in having a deeper body (35% vs. 30-32% SL), in having 3, rather than 4, sub- orbital spines in a row below the orbit, and in lacking a spine below those in the sub- orbital row. It can be separated from S. gua- mensis in lacking the large dark spot over the opercle and in not having coronal spines. It lacks the distinct dark spot on the sub- opercle characteristic of S. /ittoralis. Scor- paenodes immaculatus is distinguishable from S. varipinnis in having a wider inter- orbit (7% vs. 3—4% SL) and in not bearing strong dark markings over the body. It dif- fers widely from S. albaiensis and S. minor in having well-developed nasal spines, in having a much deeper body (35% vs. 27-— 30% SL), and in not having the middle pec- toral fin rays as abruptly longer as those dorsally. Scorpaenodes immaculatus differs from S. insularis, a species known only from St. Helena in the South Atlantic, in lacking a second spine below the row of 3 spines on infraorbitals 1-3, in having a wider inter- orbit (7.0% vs. 4.5% SL) and a slightly deep- er body (35% vs. 32% SL). Distribution. —Western Indian Ocean, known only from Walters Shoals. Etymology. —immaculatus, from the Lat- in meaning unstained, unspotted; in refer- ence to the diagnostic lack of pronounced markings on the body and fins typical of other species of Scorpaenodes. Discussion. — Although scales of S. im- maculatus taken from the dorsum and im- mediately above the anterior part of the lat- eral-line have a broader series of ctenial bases in the posterior field, with respect to ctenial size and number they are otherwise similar to those taken from the same region of the body of most other species assigned VOLUME 103, NUMBER 3 to Scorpaenodes (Fig. 3). The arrangement in S. tribulosus (Fig. 3d) notably contrasts with that of other Scorpaenodes in having many fewer and longer cten1i, with little trace of the development of ctenial bases. The presence of abruptly longer middle pectoral- fin rays (relative to those dorsally) was a feature used by Evermann & Seale (1907), Smith (1957), and Schultz (1966) to distin- guish species of Hypomacrus from other Scorpaenodes. However, the intermediate condition observed in S. immaculatus is somewhat more pronounced than that fig- ured by Eschmeyer & Randall 1975 for S. hirsutus and like that observed in S. gua- mensis-like specimens taken from Madang, Papua New Guinea (SGP observation). Insular endemism is evident in several other species of Scorpaenodes, being known for S. insularis (St. Helena), S. steenei (Rott- nest I., Western Australia), and S. englerti (Easter Island). Other shallow-water species, most notably S. /ittoralis, S. parvipinnis, and S. guamensis, are among the most broadly distributed of scorpionfishes. Only further collecting will determine if S. immaculatus occurs elsewhere. Walters Shoals is one of seven islands and sea mounts extending along the West Wind Drift from Gough and Tristan da Cunha in the South Atlantic to Amsterdam and St. Paul in the southern Indian Ocean. There is a high degree of endemism in the West Wind Drift Islands (Collette & Parin 1991) so S. immaculatus should be looked for around the other is- lands and sea mounts of this chain. Acknowledgments N. V. Parin kindly invited B. B. Collette to participate in the cruise of the Vityaz that resulted in collecting the unique specimen of Scorpaenodes immaculatus. Keiko Hira- tsuka Moore prepared Figs. | and 2. Robert Allen, Rebbeca Bivings, and Deneen Ben- ford assisted in preparation of the photo- micrographs. Drafts of the manuscript were read by William D. Anderson, Jr., William N. Eschmeyer, Thomas A. Munroe and 549 Clive Roberts. This work was supported, in part, through an NSF grant (BSR 8705373) to SGP. Literature Cited Collette, B. B., & N. V. Parin. 1991. Shallow-water fishes of Walters Shoals, Madagascar Ridge. — Bulletin of Marine Science (in press). Eschmeyer, W. N. 1969a. A new scorpionfish of the genus Scorpaenodes and S. muciparus (Alcock) from the Indian Ocean, with comments on the limits of the genus.— Occasional Papers, Cali- fornia Academy of Sciences 76:1-11. . 1969b. A systematic review of the scorpion- fishes of the Atlantic Ocean (Pisces: Scorpaeni- dae).— Occasional Papers, California Academy of Sciences 79: 1-130. ——., & J.E. Randall. 1975. The scorpaenid fishes of the Hawaiian Islands, including new species and new records (Pisces: Scorpaenidae). — Pro- ceedings of the California Academy of Sciences 40(1 1):265-334. Evermann, B. W., & A. Seale. 1907. Fishes of the Philippine Islands.— Bulletin of the [U.S.] Bu- reau of Fisheries 26:49-110. Gloerfelt-Tarp, T., & P. Kailola. 1984. Trawled fishes of southern Indonesia and northwestern Aus- tralia. Australian Development and Assistance Bureau, Directorate General Fisheries, Indo- nesia, and German Agency for Technical Co- operation, 406 pp. Hughes, D. R. 1981. Development and organization of the posterior field of ctenoid scales in the Platycephalidae.—Copeia 1981(3):596-606. Matsubara, K. 1943. Studies on the scorpaenoid fish- es of Japan. (II). Transactions Sigenkagaku Kenkyusyo, pp. 171-486. Poss, S. G. 1982. A new species of the aploactinid fish genus Kanekonia from Halmahera, Indo- nesia and a redescription of Kanekonia flori- da.—Japanese Journal of Ichthyology 28(4):1-6. Schultz, L. P. 1966. Fishes of the Marshall and Mari- anas Islands, vol. 3, United States National Mu- seum Bulletin 202, 176 pp. Smith, J. L. B. 1957. The fishes of the family Scor- paenidae in the Western Indian Ocean. Part I. The sub-family Scorpaeninae.—Ichthyological Bulletin, Rhodes University (4):49-69. (SGP) Gulf Coast Research Laboratory, P.O. Box 7000, Ocean Springs, Mississippi 39564; (BBC) National Marine Fisheries Service Systematics Laboratory, National Museum of Natural History, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 550-557 CYPHOCHARAX PANTOSTICTOS, A NEW SPECIES (PISCES: OSTARIOPHYSI: CHARACIFORMES: CURIMATIDAE) FROM THE WESTERN PORTIONS OF THE AMAZON BASIN Richard P. Vari and Ramiro Barriga S. Abstract. — Cyphocharax pantostictos, a species of curimatid characiform with a distinctive pattern of dark spots arranged in longitudinal series along the sides of the body, is described as new from the Rio Napo, Rio Putumayo, Rio Ucayali, and Rio Nanay in Ecuador and northern Peru. Cyphocharax multi- lineatus (Myers), the only other species in that questionably monophyletic genus with a similar pigmentation pattern has dark wavy horizontal lines, rather than discrete spots arranged in horizontal patterns. The dark body pigmentation in the two species also differs in its relative position on the scales. The pigmen- tation pattern and overall external appearance of C. pantostictos are nearly identical to that of Steindachnerina fasciata (Vari & Géry) a species endemic to the upper Rio Madeira system in Brazil. The two species can be readily distinguished on the basis of a series of meristic and morphometric features, and in differences in the portion of buccopharyngeal complex on the roof of the oral cavity. A series of polarized characters indicate furthermore that the two species are not closely related. The myriad drainage systems, range of stream gradients, and complexity of aquatic habitats found in the drainage basins of the western portions of the Amazon basin are reflected in the remarkable diversity of the fish fauna in that region (Ortega & Vari 1986, Stewart et al. 1987). This region is also one of the areas of greatest diversity for the fam- ily Curimatidae, involving both species widely distributed within the Amazon basin (e.g., Curimata aspera Gunther (Vari 1988: fig.8) and C. vittata Kner (Vari 1989b:42)), or ranging north and south of that system into the Rio Orinoco or Rio de La Plata systems (e.g., Steindachnerina guentheri (Eigenmann & Eigenmann) (Vari 1990)). Other curimatids in this area of high species diversity have much more restricted ranges (e.g., Steindachnerina quasimodoi Vari & Vari (see Vari & Vari 1989:477)) and are known only to occur in the region that Kul- lander (1986:40) termed the ‘““Western Am- azonian endemic area” based on distribu- tional data from various neotropical genera of the perciform family Cichlidae. In the course of investigations of the fish fauna of eastern Ecuador and northern Peru we in- dependently collected a distinctive species of curimatid with an unusual pigmentation pattern consisting of seven or eight longi- tudinal series of dark spots along the sides of the body. This material first appeared to represent a major extension in the known distribution of Curimata fasciata which Vari & Geéry (1985) described from the Rio Ma- deira system in Brazil, a considerable dis- tance southeast of the region from which the Ecuadorian and Peruvian specimens originated. More recently Vari (1989a:ta- bles 2, 3), in an analysis of intrafamilial phy- logenetic relationships, restricted Curimata to a single lineage within the family and reassigned fasciata to Steindachnerina Fow- ler (1906) on the basis of a series of derived VOLUME 103, NUMBER 3 characters. Further examination of our specimens surprisingly showed that they neither constitute a major range extension for Steindachnerina fasciata, nor do they even represent a species of Steindachnerina. Rather they are a species of Cyphocharax Fowler (1906) previously unknown to sci- ence. This new curimatid is described herein and is yet another fish species with a known range limited to the Western Amazonian endemic area identified by Kullander. Materials and methods. —Counts and measurements were made following meth- ods outlined in Vari (1989b, 1989c, 1990). Ranges for meristic and morphometric fea- tures include values of all examined speci- mens. The values in square brackets are those of the holotype. Subunits of the head are presented as proportions of head length (HL). Head length itself and measurements of body parts are presented as proportions of standard length (SL). The following abbreviations for institu- tions are used: Academy of Natural Sciences of Philadelphia (ANSP); British Museum (Natural History), London (BMNH); Cali- fornia Academy of Sciences, San Francisco (CAS); Stanford University, collections now deposited at CAS (CAS-SU); Instituto Na- cional de Pesquisas da Amazonia, Manaus (INPA); Indiana University, collections now deposited at various repositories (IU); Mu- seo de Biologia, Universidad Central de Venezuela, Caracas (MBUCV); Museo de Biologia de la Escuela Politécnica Nacional, Quito (MEPN); Muséum d’Histoire Natu- relle, Geneva (MHNG); Museu Nacional, Rio de Janeiro (MNRJ); Museu de Zoolo- gia, Universidade de Sao Paulo, Sao Paulo (MZUSP); Naturhistoriska Riksmuseet, Stockholm (NRM); and National Museum of Natural History, Smithsonian Institu- tion, Washington, D.C. (USNM). Cyphocharax pantostictos, new species Figs. 1, 4 Diagnosis. —The new species is assigned to Cyphocharax, a genus that Vari (1989a: 551 58-59) noted was not defined on the basis of known derived features. Cyphocharax of that classification was rather an assemblage of species lacking the derived features di- agnostic of the three other genera (Stein- dachnerina, Curimatella Eigenmann & Ei- genmann, and Pseudocurimata Fernandez- Yeépez) which together with Cyphocharax form an unresolved terminal polytomy in Vari’s hypothesis of intrafamilial relation- ships within the Curimatidae. The absence of identified synapomorphies for Cypho- charax increases the likelihood that the ge- nus may not be monophyletic. Ongoing studies by the senior author focus on the question of the monophyly of Cyphocharax and its subunits. In the interim we assign the new species to Cyphocharax given that C. pantostictos shares the synapomorphies for the clade formed by Cyphocharax, Steindachnerina, Curimatella, and Pseu- docurimata, but lacks the derived features that diagnose each of Steindachnerina, Cu- rimatella, and Pseudocurimata. The strik- ing pattern of seven or eight horizontal se- ries of prominent dark spots aligned along the center of the body scales is unique to the Cyphocharax pantostictos within the ge- nus (Fig. 1). Only one other Cyphocharax species, C. multilineatus (Myers 1927) of the Rio Negro system in Venezuela and Bra- zil, has a pattern of horizontal dark body pigmentation reminiscent of that in C. pan- tostictos. The pattern of dark body pigmen- tation in C. multilineatus (Fig. 2) differs from that in C. pantostictos in forming solid wavy horizontal lines rather than a series of dis- crete rotund spots (compare Figs. | and 2). Furthermore, the dark stripes in C. multi- lineatus are positioned along the area of overlap of horizontal rows of scales along the body, rather than being aligned along the center of the scale rows as are the spots in C. pantostictos. Thus the patterns of lon- gitudinal dark pigmentation on the bodies in the two species are apparently non-ho- mologous. Cyphocharax multilineatus also has a discrete dark band across the mid- 352 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Laguna de Jatuncocha. lateral surface of the head anterior and pos- terior to the orbit, a pigmentation pattern lacking in C. pantostictos. Cyphocharax pantostictos, in turn, is characterized by a well-developed, mid-lateral, horizontally elongate patch of dark pigmentation on the caudal peduncle that is absent in C. multi- lineatus. Itis likely that various meristic and morphometric features further distinguish C. pantostictos within Cyphocharax. Iden- tification of those characters must await the completion of revisionary studies within that speciose genus. The overall pigmentation pattern and overall external appearance of C. pantostic- tos are strikingly similar to that of one other member of the Curimatidae, Steindachne- “" ae Cm Cyphocharax pantostictos, new species, holotype, USNM 306594, 72.4 mm SL; Ecuador, Napo, rina fasciata (Fig. 3), an endemic of the Rio Madeira basin, and have lead to misiden- tifications of the two species. Cyphocharax pantostictos lacks the derived features that diagnose Steindachnerina (see Vari 1989a: 58, 1990), and lacks the intrageneric syn- apomorphies for the clades that include S. fasciata (see Vari 1990 for details). One of the most obvious differences between the two species involves the form of the buc- copharyngeal complex on the roof of the oral cavity. Cyphocharax pantostictos has three simple longitudinal fleshy folds in that region. Steindachnerina fasciata, in con- trast, has a mass of lobulate fleshy bodies that extend ventrally into the oral cavity, a hypothesized derived condition unique to a Fig. 2. Cyphocharax multilineatus, USNM 269987, 111.8 mm SL; Venezuela, Territorio Federal Amazonas, Departamento Rio Negro, Cafio Tremblador where crossed by road from San Carlos de Rio Negro to Solano. VOLUME 103, NUMBER 3 See ey ry mer padi Wd ii seasente 553 Fig. 3. Steindachnerina fasciata, MNRJ 11208, 89.6 mm SL, holotype of Curimata fasciata Vari and Géry; Brazil, Territorio de Rond6énia, Municipio de Ouro Preto do Oeste, Rio Romari (or Sao Domingo) near Nova Uniao. subunit of Steindachnerina (see Vari 1989a: 31-35, 1990, for a discussion of the buc- copharyngeal complex). The two species also differ in various me- ristic and morphometric values including the number of vertebrae (31 in Cyphocha- rax pantostictos versus 32 to 34, typically 33, in Steindachnerina fasciata), the num- ber of scales in a longitudinal series to the hypural joint (29 to 31 versus 32 to 37), the number of scale rows above the lateral line to the origin of dorsal fin (4'2 versus 514 or 61), the relative length of the pelvic fin (0.20-0.23 of SL versus 0.23-0.25), and the relative gape width (0.24—0.28 of HL versus 0.28—0.32). Description. —Body moderately elongate, somewhat compressed. Dorsal profile of head straight overall, slightly convex ante- riorly. Dorsal profile of body smoothly con- vex from posterior portion of head to origin of dorsal fin; straight or slightly convex, pos- teroventrally slanted at base of dorsal fin, greatly convex from base of last dorsal-fin ray to caudal peduncle. Dorsal surface of body with indistinct median keel anterior to dorsal fin, smoothly rounded transversely posterior to fin. Ventral profile of body gently curved from tip of lower jaw to caudal pe- duncle. Pre-pelvic region very obtusely flat- tened, scales of that area not notably en- larged relative to those on lateral surfaces of body. Median pre-pelvic scale series somewhat irregular, particularly near origin of pelvic fin. No distinct median keel pos- terior to origin of pelvic fin. Barely discern- able secondary obtuse keel on each side of post-pelvic portion of body about two scales dorsal of ventral midline. Greatest body depth at origin of dorsal fin, depth 0.35-0.40 [0.37], relatively deep- er in larger specimens; snout tip to origin of dorsal fin 0.47—0.52 [0.50]; snout tip to origin of anal fin 0.83-0.85 [0.83]; snout tip to origin of pelvic fin 0.52—-0.57 [0.56]; snout tip to anus 0.78-0.79 [0.78]; origin of dorsal fin to hypural joint 0.54—0.58 [0.56]. Margin of dorsal fin rounded posteriorly; anterior- most rays approximately two to two and one-half times length of ultimate ray. Mar- gin of pectoral fin pointed; length of pectoral fin 0.18—0.21 [0.20], extending slightly over one-half distance to vertical line through origin of pelvic fin. Margin of pelvic fin pointed, length of pelvic fin 0.20-0.23 [0.22], tip reaches to anus in holotype, falls some- what short of that point in larger specimens. Caudal fin forked. Adipose fin well devel- oped. Anal fin emarginate, anteriormost branched rays about two and one-half times length of ultimate ray. Caudal peduncle depth 0.12—0.14 [0.14]. 554 Head profile distinctly pointed anteriorly, head length 0.29-0.33 [0.31]; upper jaw somewhat longer than lower, mouth sub- terminal; snout length 0.27-0.31 [0.30]; nares of each side very close, anterior ro- tund, posterior crescent-shaped with aper- ture partially closed by thin flap of skin sep- arating nares; orbital diameter 0.27-0.32 [0.31]; adipose eyelid present, moderately developed, with rotund opening over center of eye; length of postorbital portion of head 0.42-0.46 [0.44]; gape width 0.24—0.28 [0.27]; interorbital width 0.39-0.43 [0.43]. Pored lateral-line scales from supraclei- thrum to hypural joint 29 to 31 [29]; all scales of lateral line pored, canals in scales straight; 2 or 3 pored scales extend beyond hypural joint onto caudal-fin base; 41% [414] scales in transverse series from origin of dorsal fin to lateral line; 312 to 41 [4] scales in transverse series from lateral line to or- igin of anal fin. Dorsal-fin rays ii,9 [11,9]; anal-fin rays 11,7 or iii,7 [ii,7]; pectoral-fin rays 13 to 15 [15]; pelvic-fin rays 1,8 or i,7,1 [1,7,1]. Total vertebrae 31 in 8 specimens. Color in life. —(Based on photograph of paratype (USNM 280573) from the Rio Na- nay of Peru taken shortly after capture.) Overall coloration silvery with slightly ol- ive-grey cast on dorsal portions of head and body. Series of black spots arranged in hor- izontal series along dorsal and lateral sur- faces of body. Distinct black mid-lateral stripe on caudal peduncle. Fins hyaline. Color in alcohol.—See Fig. 1 for pre- served color pattern. Available specimens largely lacking guanine on scales. Overall ground coloration yellowish-tan, darker on dorsal portions of head and body. Scales on lateral and dorsal surfaces of body with dark patch of pigmentation on each scale; size of spots largest mid-laterally; overall intensity of spots not as pronounced in smaller in- dividuals. Spots forming 7 or 8 horizontal series, dorsal most series not apparent in smaller specimens. Series of dark spots on scales less developed posteriorly on scale PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rows ventral of lateral line; very poorly de- veloped in series starting immediately dor- sal to origin of pectoral fin. Intense dark spots also progressively less pronounced in horizontal series dorsal to lateral line. Patches of dark pigmentation located on center of scale, with midpoint of spots lying medial of margin of preceding scale. Scales dorsal to lateral line with secondary area of diffuse dark pigmentation posterior to dis- crete central dark spot; secondary dark pig- mentation increasingly pronounced on dor- sal portions of body. Dark pigmentation patches on scales along lateral line merging posteriorly into distinct horizontal stripe on mid-lateral surface of caudal peduncle; stripe continuing onto base of middle caudal-fin rays. Deeper lying, dusky band extends along mid-lateral surface of body from supra- cleithrum to caudal peduncle. Caudal fin with small streak of dark pig- mentation on basal portions of middle rays; basal two-thirds of fin somewhat more dus- ky than remainder of fin. Median and paired fins somewhat dusky. Distribution. —Rio Napo, Rio Putumayo, Rio Ucayali, and Rio Nanay systems in Ec- uador and northern Peru (Fig. 4). Ecology.—Two of the specimens from Peru (USNM 280573, NRM SOK/ 1986293.5292) were collected in acidic black waters among grass and submerged vege- tation. The specimens from the holotype locality were collected in submerged vege- tation in blackwater of pH 5.5 at a depth of 1.5m. The Rio Yasuni specimens came from a slow flowing turbid stream with a pH of 6.0 lacking submerged vegetation. Etymology. —Pantostictos, from the Greek for “spotted all over’ refers to the prominent dark spots on the lateral and dor- sal surfaces of the body. Remarks.—As noted above, Cyphocha- rax pantostictos is very similar in body form and pigmentation to Steindachnerina fas- ciata which 1s apparently endemic to upper portions of the eastern drainages of the Rio Madeira basin in Brazil (see Vari 1990:fig. VOLUME 103, NUMBER 3 555 ) : : gt AM Ny, A \ a SA APPS iN Sey, ia Of aes ats \ te 3 Fig. 4. Map of Amazon basin and adjoining areas showing distribution of Cyphocharax pantostictus (dots; localities: 1 = Rio Napo above Coca; 2 = Rio Yasuni; 3 = Laguna Jatuncocha, type locality; 4 = Rio Putumayo, El Estrecho; 5 = Rio Nanay, Nanay beach), Steindachnerina fasciata (squares), and Cyphocharax multilineatus (stars) (see under ““Material examined” for additional locality information). 40). Nonetheless those two species differ in many meristic and morphometric features, and Cyphocharax pantostictos furthermore lacks the derived characters that both define Steindachnerina and clades within the ge- nus that include S. fasciata (see Vari (1990) for details). Such pronounced superficial similarities between two distantly related species would at first consideration apparently represent a case of intergeneric mimicry. Interestingly, however, there is no overlap in the distri- butions of the two species, whose known ranges are separated by over two thousand river kilometers. Thus mimicry would not appear to be involved in the remarkable external resemblance between Cyphocharax pantostictos and Steindachnerina fasciata. Material examined. — 10 specimens, 34.4— 98.2 mm SL. Holotype.—Ecuador: Napo. Laguna de Jatuncocha (01°00’S, 75°29'W), collected by R. Barriga, 29 Sep 1988, USNM 306594, 72.4 mm SL. Paratypes.—Ecuador: Napo. Laguna de Jatuncocha (1°00'S, 75°29’W) collected with holotype, MEPN 4554, | specimen, 74.9 mm SL. Estero Culebrero, tributary of Rio 556 Yasuni (0°54’45”S, 76°13’03”W) collected by R. Barriga, 9 May 1988, USNM 305617, 1 specimen, 98.2 mm SL; MEPN 4557, 1 specimen, 81.8 mm SL. Rio Napo, 2.7 km along river above the bridge at Coca (0°29.0'S, 77°04.0'W), collected by D. Stew- art, R. Barriga, and M. Ibarra, 2 Oct 1983, USNM 305616, 1 specimen, 66.7 mm SL; MEPN 4558, 1 specimen, 63.0 mm SL. Peru: Loreto. Rio Nanay, Nanay beach along river west of Iquitos (approx. 3°50’S, 073°11'W), collected by R. P. Vari, H. Or- tega, A. Gerberich, and J. A. Louton, 17 Aug 1986, USNM 280573, 1 specimen, 72.0 mm SL. Small stream approx. 65 km up- stream from mouth of Rio Nanay, collected by P. Fromm et al., 18 Aug 1989, ANSP 164981, 1 specimen, 34.4 mm SL. Rio Pu- tumayo drainage, El Estrecho, Quebrada de Las Granjas, collected by S. O. Kul- lander et al., 16 July 1986, NRM SOK/ 1986293.5292, 1 specimen, 94.0 mm SL. Along road from Genero Herrera towards Peruvian-Brazilian border, Rio Ucayali drainage, collected by P. Fromm et al., 23 Aug 1989, ANSP 164980, 1 specimen, 39.3 mm SL. Other material examined.—Cyphocha- rax multilineatus. Brazil: Amazonas. Rio Negro below Daraa, USNM 274102, 1. Rio Negro at Bucuri, CAS 58605, 1 (holotype of Curimatus multilineatus Myers, formerly IU 17672); CAS-SU 58986, 1. Rio Paduari, MZUSP 21161, 1. Venezuela: Territorio Federal Amazonas. Cano La Esmeralda, tributary of Rio Ori- noco, SE of La Esmeralda, MBUCV V-4479, 1. Rio Mawarinuma (0°55’N, 66°10'W), AMNH uncat., 5. Rio Urumi, tributary to Rio Negro upstream of Santa Lucia (1°17'N, 66°51'W), USNM 270241, 1. Cano Trem- blador where crossed by road from San Car- los de Rio Negro to Solano, USNM 269987, 7. Rio Barria (0°50’N, 66°10’W), MBUCV V-14898, 1. Steindachnerina fasciata. Brazil: Terri- torio de Rond6énia, Rio Romari (or Sao Do- mingo) near Nova Uniao, Municipio of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ouro Preto do Oeste, MNRJ 11208, 1 (ho- lotype of Curimata fasciata); USNM 270377, 4 (paratypes of Curimata fasciata); MNRJ 11271, 4 (paratypes of Curimata fasciata). Jiparana, Rio Urupa, tributary of Rio Jiparana, USNM 273306, 3. Rio Ma- chado system, 20 km upstream of Jiparana, USNM 295126, 1. Mato Grosso. Rio Ari- puana, above Cachoeira de Dardanelos (ap- prox. 10°19'42”S, 59°12'30”W), USNM 270375, 2 (paratypes of Curimata fasciata); INPA, 3 (paratypes of Curimata fasciata). Rio Aripuana, approximately 10 km above Cachoeira de Dardanelos, Cidade de Hum- boldt, USNM 270376, 2 (paratypes of Curi- mata fasciata); INPA, 2 (paratypes of Curi- mata fasciata); BMNH 1985.2.5:1-2, 2 (paratypes of Curimata fasciata); MZUSP 28724, 2 (paratypes of Curimata fasciata). Rio Aripuana, above Cachoeira das An- dorinhas, MHNG 2226.24, 6 (paratypes of Curimata fasciata). Acknowledgments Research associated with this study in Peru was supported by the Neotropical Lowland Research Program of the Inter- national Environmental Sciences Program of the Smithsonian Institution. Collecting activities in Peru were made possible by the generous assistance of Prof. Hernan Ortega (Museo de Historia Natural de la Univer- sidad Nacional Mayor de San Marcos) and the Consejo Nacional de Ciencia y Tecno- logia of Peru. Prof. Ortega and Dr. Andrés Urteaga (Universidad Nacional de la Ama- zonia Peruana) facilitated and participated in collecting efforts that resulted in the cap- ture of the new species. Mr. William G. Saul directed my attention towards material of the new species in the ANSP holdings. Dr. Jerry A. Louton (USNM) and Mr. Andrew Gerberich (USNM) contributed to the suc- cess of the expedition. Dr. Sven O. Kullan- der and Mr. E. Ahlander (NRM) arranged for the loan of a series of curimatids which included the specimen of Cyphocharax pan- VOLUME 103, NUMBER 3 tostictos from the Rio Putumayo. Figures 1 to 3 were prepared by Mr. Theophilus Britt Griswold (USNM). This paper benefitted from the comments and suggestions of Dr. Stanley H. Weitzman (USNM) and Dr. Thomas A. Munroe (National Marine Fish- eries Service, Systematics Laboratory), and two anonymous reviewers. Literature Cited Fowler, H. W. 1906. Further knowledge on some heterognathus fishes, Part. I.— Proceedings of the Academy of Natural Sciences of Philadelphia 58:293-351. Kullander, S.O. 1986. Cichlid fishes of the Amazon River drainage of Peru. Swedish Museum of Natural History, Stockholm, 431 pp. Myers, G.S. 1927. Descriptions of new South Amer- ican fresh-water fishes collected by Dr. Carl Ter- netz.— Bulletin of the Museum of Comparative Zoology 68:107-135. Ortega, H., & R. P. Vari. 1986. Annotated checklist of the freshwater fishes of Peru.—Smithsonian Contributions to Zoology 437:1-25. Stewart, D., R. Barriga S., & M. Ibarra. 1987. Ictio- fauna de la cuenca del rio Napo, Ecuador ori- ental: lista anotada de especies.— Politecnica, Revista de Informacion Técnico-Cientifica, Quito 12(4):9-63. Vari, R. P. 1988. The Curimatidae, a lowland neo- tropical family (Pisces: Characiformes); distri- bution, endemism and phylogenetic biogeog- raphy. Pp. 313-348 in P. E. Vanzolini, and W. Ronald Heyer, eds., Neotropical distribution 557 patterns: proceedings of a workshop. Academia Brasiliera de Ciéncias, Rio de Janeiro. 1989a. A phylogenetic study of the neotrop- ical Characiform family Curimatidae (Pisces: Ostariophysi).—Smithsonian Contributions to Zoology 471:iv+ 1-25. 1989b. Systematics of the neotropical Cha- raciform genus Curimata Bosc (Pisces: Characi- formes).—Smithsonian Contributions to Zool- ogy 474:111+ 1-63. 1989c. Systematics of the neotropical Cha- raciform genus Psectrogaster Eigenmann and Eigenmann (Pisces: Characiformes).—Smith- sonian Contributions to Zoology 48 1:iii+ 1-42. . 1990. Systematics of the neotropical Characi- form genus Steindachnerina Fowler (Pisces: Os- tariophysi). —Smithsonian Contributions to Zo- ology (in press). ——., & J. Géry. 1985. A new curimatid fish (Cha- raciformes: Curimatidae) from the Amazon ba- sin.— Proceedings of the Biological Society of Washington 98:1030-1034. ——., & A. W. Vari. 1989. Systematics of the Stein- dachnerina hypostoma complex (Pisces, Ostar- iophysi, Curimatidae), with the description of three new species. — Proceedings of the Biolog- ical Society of Washington 102:468-482. (RPV) Department of Vertebrate Zoology (Fishes), National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560; and (RBS) Departamento de Ciencias Biologicas, Escuela Politécnica National, Quito. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 558-572 THE GENUS AXTANASSA (CRUSTACEA: DECAPODA: THALASSINIDEA) IN THE AMERICAS Brian Kensley and Richard Heard Abstract. —The genus Axianassa Schmitt is redefined, based on external mor- phological characters. Descriptions and figures of the two previously described species A. intermedia Schmitt from Curacao and Puerto Rico, and A. mineri Boone from the Pacific Bay of Panama, and three new species, A. arenaria from the Gulf of Mexico, A. jamaicensis from Jamaica, and A. canalis from the Panama Canal, are provided. The species are distinguished primarily on differences in the structure of the antennal acicle, abdominal pleura, and first pereopods. An overview of the family Laomediidae (= senior synonym of Axianassidae) is presented. Schmitt (1924) described the genus and species Axianassa intermedia for which he established the monotypic family Axianas- sidae. Since that time only one other species, A. mineri Boone, 1931, from the Pacific coast of Panama has been described. Until the present study a total of five specimens was known for the genus, which has been alternately placed in the families Axianas- sidae or the Laomediidae (see references in synonymies for these families under Sys- tematics). Three more species of Axianassa are now described from the Gulf of Mexico, Jamaica, and the Panama Canal, but the total number of specimens is still only nine, suggesting that the species are nowhere par- ticularly abundant, that their microhabitats are not often sampled, or that their popu- lations are not readily accessible to conven- tional collecting methods. Systematics Family Laomediidae Borradaile, 1903 Laomediidae Borradaile, 1903:540.—De Man, 1928:15.—Gurney, 1938:332 [lar- val stages], 343 [key].—Chace, 1939: 529.—Balss, 1957:1578, 1580.—Wear & Yaldwyn, 1966:2, 3.—Yaldwyn & Wear, 1970:384; 1972:127.—Le Loeuff & Intes, 1974:20.—Poore & Griffin, 1979:224.— Goy & Provenzano, 1979:351.—Ngoc- Ho, 1981:247 [larval stages].— Williams, 1984:189.—Konishi, 1989:15 [larval stages]. Axianassidae Schmitt, 1924:76.—De Man, 1928:15.—Gurney, 1938:343 [key].— Balss, 1957:1580.—Wear & Yaldwyn, 1966:2.—Yaldwyn & Wear, 1972:127.— Le Loeuff & Intes, 1974:20. Diagnosis. —Carapace with linea thalas- sinica present; cervical groove distinct. Sev- eral elongate cleaning setae present on pos- terior margin of scaphognathite of maxilla 2. Epipod of maxilliped 3 with serrate mar- gin (reduced in Axianassa). Epipods present on pereopods 1-4. Pleopod 1 2 uniramous; pleopod | 6 absent; appendix interna absent from pleopods. Axianassa Schmitt, 1924 Axianassa Schmitt, 1924:76.—Balss, 1957: 1580. Type species.—By monotypy, Axianassa intermedia Schmitt, 1924:77, pl. 8, figs. 4, 5, text fig. 7. Gender: feminine. VOLUME 103, NUMBER 3 Diagnosis. — Body poorly calcified, integ- ument thin. Rostrum short, anteriorly rounded, weakly to moderately developed. Eyestalks short, cornea poorly defined, eyes poorly pigmented, generally not, or barely reaching anteriorly beyond rostral apex. Antennules with article 3 of peduncle elon- gate, slender; superior flagellum longer than inferior. Antennal scale usually well devel- oped and dagger-like; article 4 of peduncle elongate, slender. Mandible with palp of 3 articles, two basal articles incompletely fused; incisor area having several cusps; molar moderately developed, with several tubercles. Maxilla 1 with strongly setose en- dite. Maxilla 2, scaphognathite bearing 5 elongate setae extending posteriorly into branchial chamber. Exopods present on maxillipeds 1 and 2. Epipods present on maxillipeds 1-3 and pereopods 1-4. Single podobranch present on maxillipeds 2 and 3 (rudimentary on maxilliped 1) and pereo- pods 1-3; single rudimentary arthrobranch on maxilliped 1, 2 arthrobranchs on max- illipeds 2 and 3, and pereopods 1-4. Pereo- pod 1, chelae of similar size but dissimilar in chela-palm proportions and dentition of cutting edges of dactylus and propodal fin- ger. Dactyli of pereopods 3-5 flexed, with posterior margin becoming dorsal in posi- tion. Pereopod 5 base not covered by car- apace, lacking branchiae. Pleopods 2-5 bi- ramous, rami narrowly lanceolate; lacking appendices internae. Uropodal rami lacking sutures, outer ramus with variable weak spi- nation. Telson lacking spination. Distribution. —The genus is known only from the tropical and subtropical waters of the northeastern Pacific (Panama) and the northwestern Atlantic (Gulf of Mexico and Caribbean). Axianassa intermedia Schmitt, 1924 lies, I, 5.3) Axianassa intermedia Schmitt, 1924:77, pl. 8, figs. 4, 5, text fig. 7.—De Man, 1928: 17, pl. 1, fig. 2.—Gurney, 1938:332.—Goy & Provenzano, 1979:351. 559 Material. —Syntypes, National Museum of Natural History, Smithsonian Institu- tion, USNM 57512, 2 2 cl.6.8 mm, 7.8 mm, 6 cl. 6.1 mm, from muddy creek, Spanish Harbor, Curagao.—USNM 110451, 1 4, cl. 5.5 mm, Punta Arenas, Puerto Rico. Description. —Carapace strongly rounded dorsally but unarmed, with linea thalassin- ica somewhat grooved; cervical groove clearly defined; rostrum anteriorly rounded, barely surpassing cornea of eyes. Abdomen thin-walled, tergites bearing scattered setae; somite | narrowing ventrally, pleuron poor- ly defined, rounded, lacking any ventral ex- tension; somites 2 and 3 subequal in mid- dorsal length; somites 4—6 decreasing in length. Telson with greatest width slightly more than middorsal length, bearing mar- ginal plumose setae, lateral and posterior margins not clearly separated. Cornea weakly pigmented, with tiny tu- bercle distally. Antennular peduncle, arti- cles 1 and 2 together about % length of ar- ticle 3; latter slender, elongate-cylindrical; inferior flagellum of about 14 articles, reaching to distal end of antennal peduncle article 5; superior flagellum of about 28 ar- ticles, almost twice length of inferior fla- gellum. Antennal acicle slender, dagger-like, with small tooth on mesial margin; pedun- cle article 4 slender, elongate-cylindrical, al- most reaching distal margin of merus of pe- reopod 1; flagellum about 3 times carapace length. Mouthparts as illustrated. Mandible with cutting edge of 7 cusps; raised molar area on mesial face bearing 4 rounded tubercles; palp of 3 articles, articles 1 and 2 partially fused, article 3 bearing numerous stiff setae on outer surface. Maxilla 2, scaphognathite bearing 5 elongate setae on posterior mar- gin. Maxilliped 1, endopod unsegmented, paddle-shaped, distally broadly rounded; exopod bipartite, longer proximal part wid- ening distally, flagellar part about half length of proximal half, bearing plumose setae. Maxilliped 2, endopod pediform, of 5 ar- ticles, distal article inserted obliquely on 560 SO ron 9 SOs = ecpEEeStectitce aaa 2 ot Coe Sc ot SS ies eee £7 Sy Fig. 1. penultimate article; exopod bipartite, prox- imal part distally broadened, flagellar part more than half length of proximal, bearing plumose setae. Maxilliped 3, endopodal ar- ticles strongly setose on mesial margins; ba- sis with single tooth on lateral surface; is- chium having dentate crest on mesial surface bearing 15 teeth; merus with band of dense short mesiodistal fusiform setae. Pereopods 1 asymmetrical; ventral mar- gin of ischium obscurely toothed or tuber- culate; merus distally expanded, ventral margin toothed, teeth becoming larger dis- tally, suture along dorsal margin; carpus with longitudinal suture on dorsal surface, small tooth ventrodistally; both chelae carinate along dorsal margin of propodus; larger che- la broader, and fingers relatively shorter, than smaller chela, outer surface of propo- dus at base of finger finely granulate, fixed finger with 2 or 3 large teeth on cutting edge, dactylus strongly curved, with rounded ridge PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Axianassa intermedia, syntype ¢é in lateral view. scale = 5 mm. on outer surface, about 5 strong teeth on cutting edge; smaller chela lacking granu- lations on outer surface, row of small teeth on propodus at base of dactylus on both mesial and lateral surface, cutting edges of fingers more evenly serrate than in larger chela. Pereopod 2, merus, carpus, and propodus bearing elongate setae on posterior margins; dactylus with posterior margin bearing row of fine spinules. Pereopod 3, propodus with posterodistal band of dense stiff setae; dac- tylus flexed, with row of about 17 stout cor- neous spines on posterior surface, sinuous row of fine short spinules on anterior mar- gin. Pereopod 4 similar to pereopod 3, but with merus relatively shorter. Pereopod 5 flexed anteriorly, propodus bearing broad ‘pad’ of fine setae, those along posterior margin becoming more elongate distally, posterodistal margin bearing several small teeth; dactylus with posterior margin sin- VOLUME 103, NUMBER 3 561 NN Wel Ih li en aii Wil | Vi Wi LLL AI A MW Fig. 2. Axianassa intermedia, syntype 2: A, anterior carapace in dorsal view; B, antennule; C, antenna; D, E, mandible, outer and inner view; F, maxilla 1; G, maxilla 2; H, maxilliped 1; I, maxilliped 2; J, maxilliped 3, with ischial crest shown separately. 562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON oe N Cs SS aN py N\Y ‘NS Be RA WN STAN ——\ AUX YY NY Fig. 3. Axianassa intermedia, syntype 2: A, telson and right uropod in dorsal view; B, pleopod 1; C, pleopod 2; D, pereopod 2; E, pereopod 3; F, pereopod 4; G, pereopod 5, with propodus and dactylus enlarged. VOLUME 103, NUMBER 3 uous, lined with row of tiny spinules. Lat- eral uropodal ramus ovate, lacking suture, lateral margin bearing 3-5 small fixed teeth, distalmost largest and having mobile spine internal to it. Variation. —Pereopod 1, spination on ventral margin of merus: USNM 110451: 3 cl 5.5 mm-—strong row of spines on both chelipeds. USNM 57512: 2 cl 7.8 mm— spines strong on larger chela, almost obso- lete on smaller chela. 2 cl 6.8 mm-—spines fairly strong on larger chela, smaller chela missing. 6cl 6.1 mm-—spines strong on both chelae. Axianassa mineri Boone, 1931 Fig. 4 Axianassa mineri Boone, 1931:157, fig. 10.—Goy & Provenzano, 1979:351. Material. —Holotype, American Mu- seum of Natural History, AMNH 8-IllI- 1926-6403, 2 cl 6.1 mm, Pacheca Island, Pearl Islands, Bay of Panama (Pacific coast), under intertidal stones on rocky and grav- elly beach. Description. —Carapace with linea thalas- sinica somewhat grooved; cervical groove clearly defined; rostrum barely defined, an- terior margin of carapace broadly rounded, not reaching beyond cornea of eyes. Ab- domen thin-walled, bearing few setae on posteroventral region of pleura; somite | half length of 2, pleuron ventrally indistinct, rounded; somites 2—4 subequal in length, somites 5 and 6 decreasing in length. Telson about ’ longer than greatest width, poste- riorly broadly rounded. Cornea subterminal, faintly demarked from eyestalk, pigmentation lost after pro- longed preservation, with tiny tubercle dis- tally. Antennular peduncle articles 1 and 2 about 3 length of article 3, latter slender, elongate-cylindrical; inferior flagellum of about 14 articles, just reaching beyond an- tennal peduncle article 5; superior flagellum of about 25 articles, about 3 longer than inferior flagellum. Antennal acicle short, apically bifid; peduncle article 4 slender, 563 elongate-cylindrical, slightly longer than distance between rostral apex and cervical groove of carapace; flagellum 4 longer than carapace. Mouthparts typical of genus. Maxilliped 3, articles strongly setose on posterior mar- gins; ischium with mesial crest bearing about 14 teeth, crest continuing on proximal me- rus bearing 3 teeth; merus with one mar- ginal tooth in proximal half and 2 teeth in distal half of posterior margin; carpus with single distal tooth on posterior margin. Pereopod 1, left chela missing; right leg with ischium finely denticulate on posterior margin; merus inflated, with 3 small teeth proximally and single small tooth distally on posterior margin; carpus with distal rounded process on posterior margin; chela with dactylus equal in length of propodal palm, outer surface of palm smooth except for faint patch of low granulations at base of fixed finger, inner surface somewhat more granulate at base of fixed finger; latter with 4 triangular proximal teeth on cutting edge, followed by several low rounded teeth dis- tally almost to apex; dactylus with cutting edge bearing even row of rounded teeth. Pereopod 2, dactylus with row of fine spi- nules on posterior margin. Pereopod 3, pro- podus with pad of short stiff setae distally; dactylus with 37 corneous spines arranged roughly in 3 rows on flexed posterior sur- face, anteriorly with sinuous row of fine short spinules. Pereopod 4 similar to pereopod 3 but with merus relatively shorter. Pereopod 5, both legs missing. Lateral uropodal ramus ovate, with single submarginal tooth laterally, 3 marginal teeth and | mobile spine on rounded distolateral margin, | marginal and | submarginal tooth distally; inner ramus with single small tooth on dorsal surface, single tooth on distal mar- gin. Axianassa arenaria, new species Fig. 5 Material. —Holotype, USNM 211490, 6 cl 6.2 mm, Gulf of Mexico, MAFLA (Mis- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 564 FSSS88 S ji "7 77S YT APSO \\ | AQ FT / SS Z a LF K my LG ae I} \ y “i, /, \ \ ( \4 Pay // ff /, Porat INorcet —Yyi / C i HN Fig. 4. Axianassa mineri, holotype é: A, lateral view, scale = 5 mm; B, right pereopod 1; C, telson and left uropod; D, anterior carapace in dorsal view. sissippi-Alabama-Florida BLM program) reaching well beyond cornea of eyes. Ab- sta 2315, 28°33'59”"N, 84°20'09”"W, 38 m, dominal somite 1 about half middorsal fine silty sand bottom.— Paratype, USNM_ length of somite 2, pleuron tapering ven- 211491, 6 cl 7.0 mm (pleon somite 6 and _trally to somewhat calcified spiniform pro- telson missing), Gulf of Mexico, MAFLA cess. Somites 2 and 3 subequal in length, sta 2209, 27°52'30’"N, 83°33'59”"W, 34 m, somites 4, 5, and 6 shorter, subequal; pleura clayey and sandy silt bottom. of somites 2—5 with acute posteroventral Description. —Carapace with linea thalas- tooth. Telson middorsal length almost 1.5 times greatest width; tapering to broadly sinica grooved; cervical groove clearly de- fined; rostrum narrowing to rounded ante- rounded posterior margin. Cornea not clearly demarked, moderately rior margin with tiny papilla at midpoint, VOLUME 103, NUMBER 3 565 SS =~ ~y tS itn, ] Ti Fig. 5. Axianassa arenaria: A, paratype ¢ in lateral view, scale = 5 mm; B, anterior carapace in dorsal view; C, telson and left uropod in dorsal view; D, right pereopod 1, inner view; E, left pereopod 1, outer view; F, pereopod 2; G, pereopod 3; H, pereopod 4; I, pereopod 5. pigmented. Antennular peduncle articles | length of superior flagellum; latter of about and 2 together slightly less than half length 30 articles. Antennal acicle narrowly tri- of article 3; latter slender, elongate-cylin- angular, lacking tooth on mesial margin; pe- drical; inferior flagellum of 18 articles, 7% duncle article 4 slender, elongate-cylindri- 566 cal, equal in length to distance from rostral tip to cervical groove of carapace; flagellum 1%; carapace length including rostrum. Mouthparts typical of genus. Maxilliped 3, basis with strong mesiodistal tooth; is- chium with mesial crest bearing about 17 teeth; 5 distal articles heavily setose on pos- terior margins. Pereopod 1, ischium with 2 or 3 strong teeth on posterior margin; merus inflated, with posterior margin bearing single strong tooth in distal half plus several small den- ticulations; carpus with | or 2 blunt tuber- cles on posterodistal margin; left and right chelae of equal length, left chela more robust than right; left chela, fingers 74 length of pro- podal palm, latter with low granulations on proximoposterior surface, cutting edge of fixed finger with 2 strong triangular teeth and series of small rounded teeth reaching almost to apex; dactylus with strong prox- imal tooth, second strong tooth at about midlength, rest of cutting edge bearing low rounded teeth almost to apex; right chela, fingers about '4 longer than propodal palm; propodal fixed finger with 2 strong trian- gular teeth, rest of cutting edge bearing row of small rounded teeth; dactylar cutting edge bearing uniform row of small rounded teeth; tapering and curved distally. Pereopod 2, 4 distal articles strongly setose on posterior margins; dactylus about % length of pro- podus, with row of very small spinules on posterior margin. Pereopod 3, dactylus flexed, with row of 10 spines on posterior margin, spines becoming progressively more elongate distally; close-packed sinuous fringe of spinules on anterior margin. Pereopod 4 similar to pereopod 3, but merus relatively shorter. Pereopod 5, propodus with band of densely packed spinules, broadening distal- ly, on posterior margin; dactylus flexed, with close-packed row of short spinules on an- terior margin. Outer ramus of uropod with several ser- rations on outer margin; inner ramus un- armed; both rami bearing dense row of plu- mose marginal setae. Etymology.—The specific epithet ‘are- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON naria’ meaning sandy, refers to the fine sandy habitat of this species. Axianassa jamaicensis, new species Fig. 6 Material.—Holotype, USNM 155732, 6 cl 6.0 mm, St. James, Montego Bay, Ja- maica, intertidal, taken from burrow in sandy bottom, coll. C. B. Wilson, 7 Feb 1910. Description. —Carapace lightly calcified, dorsally strongly rounded, free margins of branchiostegite bearing row of setae; cer- vical groove clearly defined; linea thalassin- ica somewhat grooved; rostrum anteriorly rounded, surpassing cornea of eyes. Abdo- men thin-walled, with scattered setae es- pecially on posteroventral angles of pleura; somite | with pleuron narrowing ventrally, rounded and poorly defined, lacking any spine-like ventral extension; somite 2 long- est, somites 3—5 subequal in length, somite 6 about % longer than somite 5. Greatest telsonic width only slightly less than mid- dorsal length; posterior margin broadly rounded, bearing marginal plumose setae. Cornea not clearly demarked from eye- stalk, weakly pigmented, with poorly de- fined mesiodistal tubercle. Antennular pe- duncle with articles 1 and 2 together half length of article 3; latter slender, elongate- cylindrical; inferior flagellum of about 11 articles, slightly less than half length of su- perior flagellum; latter of 24 articles. An- tennal acicle narrowly triangular, with small tooth at about midlength of mesial margin; peduncle article 4 slender, elongate-cylin- drical, about 7 length of carapace plus ros- trum; flagellum about 3 times carapace length. Mouthparts, and disposition of exopods, epipods, and branchiae as in A. intermedia. Maxilliped 3, endopodal articles strongly setose on posterior margins; basis with sin- gle strong mesiodistal spine; mesial crest of ischium bearing 10 teeth. Pereopod 1, chelae of similar size, dissim- ilar in proportions and armature; left leg, VOLUME 103, NUMBER 3 567 nv Cc ) \ \\ \) cu do Fig. 6. Axianassa jamaicensis: A, holotype ¢ in lateral view, scale = 5 mm; B, anterior carapace in dorsal view; C, telson and left uropod in dorsal view; D, maxilliped 3. ischium with 3 strong spines on posterior margin; merus inflated, with single spine at midlength of posterior margin and few proximal denticulations; carpus with blunt distal tubercle and shoulder on posterior margin; propodus with marked patches of granulations along anterior (upper) surface, at base of fixed finger and along posterior surface, fixed finger with 4 strong rounded teeth on cutting edge; dactylus slightly more than half length of propodal palm, with proximal molar-like process and distal row of low rounded teeth. Right leg, merus, is- chium, and carpus as in left; granulations of propodus similar but not as extensive as in left, cutting edge with large triangular dis- tal and proximal tooth, row of low rounded teeth between; dactylar cutting edge bearing even row of low rounded teeth. Pereopod 2, 4 distal articles bearing numerous mar- ginal setae; dactylus with row of fine spi- nules on posterior margin. Pereopod 3, pro- podus with dense posterodistal band of setae; dactylus flexed, with single row of about 16 cormeous spines on posterior margin; row of fine spines in sinuous row on anterior margin. Pereopod 4 similar to pereopod 3, but merus relatively shorter; flexed dactylus bearing about 16 corneous spines in single row distally, double row proximally, on posterior margin. Pereopod 5, propodus bearing band of dense short setae; dactylus flexed, setose, lacking spines. Outer uropodal ramus ovate, with 2 teeth and single mobile spine on outer margin; inner and outer rami bearing dense fringe of plumose setae. Etymology.—The specific epithet refers to Jamaica, the type locality of the species. Axianassa canalis, new species Fig. 7 Material. —Holotype, USNM 125897, 6 cl 10.9 mm, Panama Canal, seaward of 568 Thatcher Ferry Bridge, from dredge flume, coll. H. O. Wright, 16 Mar 1967. Description. —Carapace with linea thalas- sinica grooved; cervical groove clearly de- fined; rostrum tapering, anteriorly narrowly rounded, reaching well beyond cornea of eyes. Abdomen thin-walled, terga and pleu- ra bearing fairly dense pile of very short setules; somite 1 somewhat more than half middorsal length of somite 2, pleuron pro- duced ventrally into strong, calcified spi- niform process, anteroventral margin bear- ing few short spinules; somite 2, pleuron ventrally broad, bearing few submarginal spinules; somites 3-6 subequal in length, pleura each bearing dense fringe of marginal and submarginal setae, and few spinules an- teroventrally, ventral margins sinuous. Tel- son length subequal to greatest width, pos- teriorly broadly rounded, bearing numerous short scattered spinules dorsally. Cornea not clearly demarked, moderately pigmented. Article 3 ofantennular peduncle slender, elongate-cylindrical, twice length of articles 1 and 2 together; inferior flagellum of at least 14 articles, less than half length of superior flagellum; latter of about 44 ar- ticles. Antennal acicle narrowly triangular, with tooth on mesial margin; peduncle ar- ticle 4 slender, elongate-cylindrical, bearing row of short spinules along ventral surface; flagellum missing. Mouthparts typical of genus. Maxilliped 3, basis with strong mesiodistal tooth; is- chium with band of fusiform setae on pos- terior margin, 9 slender teeth on mesial crest; posterior margins of 5 distal articles strong- ly setose. Pereopod 1, both legs missing. Pereopod 2, both legs missing. Pereopod 3, propodus with dense band of setae on posterodistal margin; dactylus flexed, with row of 11 slen- der spines on posterior margin, becoming more slender distally; fringe of short spi- nules on anterior margin. Pereopod 4 sim- ilar to but shorter than pereopod 3, dactylus with row of 13 slender spines on posterior margin. Pereopod 5, both legs missing. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Outer uropodal ramus ovate, bearing sin- gle marginal tooth on lateral margin, outer half of dorsal surface bearing short scattered spinules; inner ramus ovate, with outer half of dorsal surface bearing short scattered spi- nules; both rami bearing marginal plumose setae. Etymology. —The specific epithet “‘cana- lis” meaning a canal, refers to the type lo- cality of this species. Characters for the separation of the five known species of the genus Axianassa are presented in the following key. Since the combined number of specimens for all species in the genus is only nine, the follow- ing key cannot account for intraspecific variability. With more material, some fea- tures used in the key and diagnoses may prove to be of limited value in distinguish- ing the species. Key to the species of Axianassa 1. Antennal acicle long, dagger-like, apically acute — Antennal acicle short, apically bifid dal bolas Mellin Salt Ba ait da ane mineri 2. Pleura of abdominal somites 2-5 having single posteroventral tooth; antennal acicle lacking tooth on me- Sial Maree ya Cah aaa arenaria — Pleura of abdominal somites 2-5 unarmed; antennal acicle with tooth On mesial mareines 3) seen ae 3 3. Abdominal somite 1 produced ven- trally into spiniform process ... canalis — Abdominal somite 1 ventrally TOUNGEGY tc 1c, okie ta ee ee eee 4 4. Ischium of pereopod | armed with 3 strong teeth on posterior margin EPR RN ERS Ait ee hos SG Jamaicensis — Ischium of pereopod | lacking strong teeth, posterior margin finely den- ticulate Mig d coated Sees intermedia Discussion Since 1924 when Schmitt created the family Axianassidae for his new genus and VOLUME 103, NUMBER 3 569 Fig. 7. Axianassa canalis: A, holotype é in lateral view, scale = 5 mm; B, anterior carapace in dorsal view; C, telson and left uropod; D, maxilliped 3; E, pereopod 3; F, pereopod 4. species Axianassa intermedia, the position and validity of the family Axianassidae have been questioned by several authors (De Man 1928, Balss 1957; Le Loeuff & Intes 1974, Ngoc-Ho 1981), who included the genus in the earlier family Laomediidae. Others have retained the Axianassidae (Wear & Yald- wyn 1966, Poore & Griffin 1979), while Goy & Provenzano (1979:351) explicitly exclud- ed Axianassa from the Laomediidae. Gur- ney (1938) retained the Axianassidae, based on his studies of larval forms. Chace (1939), 570 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Comparison of the genera of the Laomediidae. Character Axianassa Laomedia Jaxea Naushonia Laurentiella rostrum unarmed armed armed armed ?unarmed antennular ped. elongate short elongate short short art. 3 antennal acicle spiciform reduced reduced well-dev. spiciform (reduced) antennal ped. elongate short elongate short elongate art. 4 maxilla 2 elongate 5 5 7 10 7 setae maxilliped 3 absent present present present absent exopod exopods on 0 2-5 (reduced) 0 0 0 pereopods pereopod 1 chelate > chelate chelate subchelate chelate dissimilar dissimilar similar similar dissimilar pereopod 2 simple subchelate subchelate simple simple pereopods 3 & 4 twisted twisted ?twisted unflexed twisted dactyl band of spines band of spines no band of no band of band of spines spines spines corneous no corneous no corneous no corneous corneous spines spines spines spines spines pereopod 5 subchelate subchelate subchelate simple subchelate uropodal sutures absent on both rami on bothrami onbothrami_ on outer ramus on in dealing with the species of Naushonia, divided the Laomediidae into the subfam- ilies Laomediinae and Naushoniinae, based on the structure of the uropods, antennal scale, and fifth pereopods, but did not refer to Axianassa. Le Loeuff & Intes (1974) added the new genus Laurentiella to the family Laomediidae. Five genera have now been included in this family: Jaxea Nardo, 1847, Laomedia De Haan, 1849, Naushonia Kingsley, 1897, Axianassa Schmitt, 1924, and Laurentiella Le Loeuff & Intes, 1974. Clearly, a critical reexamination of the re- lationships of these genera is necessary, along with their relationship to the rest of the tha- lassinidean families. Table 1 has been compiled for compari- son of some of the characters of the five laomediid genera. For each genus, the rel- evant column of character states may be regarded as its diagnosis. Unfortunately, information on larval morphology and development for laome- diids is limited. The larval types for species in two genera, Jaxea and Naushonia, have been definitely identified and described by Wear & Yaldwyn (1966) and Goy & Pro- venzano (1979), respectively. There have been three published reports of laomediid larvae taken in plankton samples from the western Atlantic. In one of these reports based on plankton samples from the south- western Gulf of Mexico, Ngoc-Ho (1981) tentatively identified the laomediid larvae present as “Axianassa sp.’ and discussed their similarity to the known larvae of the family Upogebiidae. Ngoc-Ho’s material appears to be similar and possibly conge- neric with the laomediid larvae reported by Gurney (1938) from Brazilian waters and by Truesdale & Andryszak (1983) from the northern Gulf of Mexico off Louisiana. However, until larvae hatched from eggs of Axianassa females can be examined and VOLUME 103, NUMBER 3 critically compared with the larvae from these previous plankton studies, Ngoc-Ho’s generic identification must be considered tentative. Sakai & Miyake (1964) reported on the first zoea of Laomedia astacina, and noted its greater similarity to larval stages of Jaxea and Naushonia than to those of Laomedia. Konishi (1989), ina compilation of the known information on thalassinidean larval development, concluded that the upogebiids are intermediate between the more primitive axiids and the more ad- vanced laomediids, and that the earlier re- ported Axianassa zoea (Ngoc-Ho 1981) had more primitive characters than the typical laomediids. While larval features are un- doubtedly of value in the question of thalas- sinidean phylogeny, conclusions about phy- logeny based on such features alone should be avoided; rather, larval characters should be considered along with morphological, bi- ological, ecological, and molecular evi- dence, to build a strong basis for phyloge- netic inference. Acknowledgments We are grateful to Dr. Harold Feinberg (American Museum of Natural History, New York) for the loan of material, and to Ms. Debbie Roney who inked the whole animal drawings of Figs. 4-7. Our sincere thanks are due to A. B. Williams, National Marine Fisheries Service, for his critical reading of the manuscript. Literature Cited Balss, H. 1957. Decapoda (Zehnfusser) Part 12.—Dr. H. G. Bronns Klassen und Ordnungen des Tier- reichs 5(1)7:1505—1672. Boone, L. 1931. A collection of anomuran and ma- cruran Crustacea from Panama and the fresh waters of the Canal Zone.—Bulletin of the American Museum of Natural History 633(2): 137-189. Borradaile, L. A. 1903. On the classification of the Thalassinidea.—Annals and Magazine of Nat- ural History (7)12:534—-551. Chace, F. A., Jr. 1939. On the systematic status of the crustacean genera Naushonia, Homoriscus, 571 and Coralliocrangon.— Annals and Magazine of Natural History (11)3:524~—530. Goy, J. W., & A. J. Provenzano, Jr. 1979. Juvenile morphology of the rare burrowing mud shrimp Naushonia crangonoides Kingsley, with a re- view of the genus Naushonia (Decapoda: Tha- lassinidea: Laomediidae).— Proceedings of the Biological Society of Washington 92(2):339-359. Gurney, R. 1938. Larvae of decapod Crustacea. Part 5. Nephropsidea and Thalassinidea.— Discov- ery Reports 17:291-344. Haan, W. de. 1833-1850. Crustacea. Jn P. F. von Siebold, Fauna Japonica sive Descriptio Ani- malium, quae in Itinere per Japoniam, Jussu et Auspiciis Superiorum, qui Summum in India Batava Imperium Tenent, Suscepto, Annis 1823- 1830 Collegit, Notis, Observationibus et Ad- umbrationibus Illustravit. Leiden, i—xxx1, 1x—xvi, 1-243, pls. A-J, L-Q, 1-55. Kingsley, J. S. 1897. Ona new genus and two new species of macrurous Crustacea. — Bulletin of the Essex Institute 27:95-99. Konishi, K. 1989. Larval development of the mud shrimp Upogebia (Upogebia) major (De Haan) (Crustacea: Thalassinidea: Upogebiidae) under laboratory conditions, with comments on larval characters of thalassinid families.— Bulletin of the National Research Institute of Aquaculture, Nansei 15:1-17. Le Loeuff, P., & A. Intes. 1974. Les Thalassinidea (Crustacea, Decapoda) du Golfe de Guinée. Sys- tématique—€cologie.— Cahiers ORSTOM 12(1): 17-69. Man, J. G. de. 1928. The Decapoda of the Siboga- Expedition. Part 7. The Thalassinidae and Cal- lianassidae collected by the Siboga—Expedition with some remarks on the Laomediidae.—Si- boga—Expeditie monographie 39a6:1-187. Nardo, G. D. 1847. Sinonimia moderna della specie registrate nell’ opera initolata: Descrizione de’ Crostacei, de’ Testacei e de’ Pesci che abitano le Laguna e Golfo Veneto, rappresentati in figure dall’ Abate S. Chiereghini Ven. Clodiense, ap- plicata per commissione governativa. Venice, xi, 128 pp. Ngoc-Ho, N. 1981. A taxonomic study of the larvae of four thalassinid species (Decapoda, Thalas- sinidea) from the Gulf of Mexico.—Bulletin British Museum of Natural History (Zoology) 40(5):237-273. Poore, G. C. B., & D. J. G. Griffin. 1979. The Thalas- sinidea (Crustacea: Decapoda) of Australia.— Records of the Australian Museum 32(6):217- Be Sakai, K., & S. Miyake. 1964. Description of the first zoea of Laomedia astacina de Haan (Decapoda Crustacea).— Science Bulletin of the Faculty of Agriculture, Kyushu University 21(1):83-87. 2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Schmitt, W. L. 1924. Bijdragen tot de kennis der fauna van Curacao. Resultaten eener reis van Dr. C. J. van der Horst in 1920. The macruran, anomuran and stomatopod Crustacea. — Bijdra- gen tot de Dierkunde 23:61-81. Truesdale, F. M., & B. L. Andryszak. 1983. Occur- rence and distribution of reptant decapod crus- tacean larvae in neritic Louisiana waters: July 1976.— Contributions in Marine Science 26:37-— 53. Wear, R. G., & J. C. Yaldwyn. 1966. Studies on thalassinid Crustacea (Decapoda, Macrura Rep- tantia) with a description of a new Jaxea from New Zealand and an account of its larval de- velopment.— Zoology Publications from Vic- toria, University of Wellington 41:1—27. Williams, A. B. 1984. Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Washington, D.C., Smithson- ian Institution Press, 550 pp. Yaldwyn, J. C., & R. G. Wear. 1970. Preliminary description of a new burrowing mud-shrimp from eastern Australia (Crustacea, Macrura, Reptantia, Laomediidae).— Australian Zoolo- gist 15(3):384-385. 5 & . 1972. The eastern Australian, bur- rowing mud-shrimp Laomedia healyi (Crusta- cea, Macrura Reptantia, Laomediidae) with notes on larvae of the genus Laomedia. — Aus- tralian Zoologist 17(2):126-141. (BK) Department of Invertebrate Zool- ogy, National Museum of Natural History, NHB-163, Smithsonian Institution, Wash- ington, D.C. 20560; (RH) Gulf Coast Re- search Lab, P.O. Box 7000, Ocean Springs, Mississippi 39564—7000. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 573-597 ON THE CRAYFISHES (DECAPODA: CAMBARIDAE) OF THE NECHES RIVER BASIN OF EASTERN TEXAS WITH THE DESCRIPTIONS OF THREE NEW SPECIES Horton H. Hobbs, Jr. Abstract.— Among the 13 crayfishes recorded here from the Neches River basin in eastern Texas are three that are previously undescribed: Procambarus (Girardiella) nigrocinctus, P. (G.) kensleyi, and P. (Ortmannicus) nechesae. The first has its closest affinities with P. (G.) tulanei Penn, the second with P. (G.) parasimulans Hobbs and Robison, and the third is a close relative of P. (O.) geminus Hobbs. Present also in the Basin is a member of the genus Cambarellus, two representatives of the genus Cambarus, two belonging to the genus Falli- cambarus, a member of the genus Faxonella, and four other species of Pro- cambarus. New locality records are cited for all of them. With the intent of stimulating an interest in the crayfish fauna of Texas, the late George Henry Penn, Jr., and I summarized (1958) all of the information available to us con- cerning the crayfishes of Texas and provid- ed a key to the species and subspecies rec- ognized at that time. Unfortunately, few contributions extending our knowledge of the fauna have appeared since. Among them is that of Rollin D. Reimer (1969) whose unpublished dissertation contains descrip- tions of several unnamed new taxa and rec- ords that are in need of further attention. One of the species (““Procambarus species E’’) included in the dissertation was cited as occurring in the Navasota River basin by Reimer & Clark (1974) in their summary of the decapod crustaceans occurring in that basin. In the following year this crayfish was described under the name of Procambarus (Girardiella) curdi by Reimer (1975). Two of the species described herein were almost certainly recognized by Reimer and were cited by him as occurring in the Neches Riv- er basin: his ““Procambarus species F’’ is almost certainly conspecific with P. (Girard- ella) nigrocinctus described below, and his Procambarus species G’’ embraces, if it is not identical with, P. (G.) kensleyi which is also described herein. As noted by Hobbs & Robison (1982:545), reluctance exists in describing crayfishes that almost certainly were recognized as undescribed by Dr. Rei- mer, but receiving no response to repeated attempts to communicate with him, de- scriptions of these two species are included here. Hobbs (1971) described Procambarus (Ortmannicus) texanus from the fish hatch- ery near Smithville, Bastrop County, but no records of its presence elsewhere have been reported. Unpublished is a detailed study of the life histories of two crayfishes occur- ring in southeastern Texas by Albaugh (1973). A new species belonging to the genus Procambarus was described by Albaugh (1975) and another of the genus Cambarel- lus by Albaugh & Black (1973). In 1986, I was greatly surprised to learn of the exis- tence of a crayfish pest in the eastern prairie section of Texas that had caused concern to farmers and to residents with lawns for a number of years, if not for generations. How Fallicambarus (F.) devastator Hobbs & Whiteman (1987) escaped earlier attention of students of crayfishes remains unan- swered to date. In attempting to obtain in- formation on the distribution and biology of this crayfish (See Hobbs & Whiteman 1990), specimens of other species of cam- 574 barids, including three that were previously undescribed, were found within and bor- dering its range. Their occurrence within the area frequented by F. (F.) devastator are re- corded herein. It would be desirable to compare the cray- fish fauna of the Neches River basin with that of other river systems along the western Gulf Slope as has been so admirably done for the freshwater fishes by Conner & Sutt- kus (1986). Unfortunately the crayfish fau- na has not been determined for a single stream between the Mississippi River and the Rio Grande. Certainly the present report does not represent a complete inventory of the crayfishes of the Neches Basin. The only survey of a Texas watershed that has pro- duced a perhaps near-exhaustive list of the crayfishes occurring within it is the study of Reimer & Clark (1974) on the Navasota watershed, a tributary of the Brazos River. They found ten species of crayfishes, six of which are reported herein to occur also in the Neches Basin: Cambarellus (Pandicam- barus) puer Hobbs, 1945, Fallicambarus (C.) hedgpethi (Hobbs, 1948) (=F. (C.) fodiens (Cottle, 1863); see Hobbs & Robison 1989), Cambarus (L.) ludovicianus Faxon, 1884, ‘“*Procambarus species A”’ (=P. (Girardiella) curdi), P. (Ortmannicus) acutus (Girard, 1852) , and P. (Scapulicambarus) clarkii (Girard, 1852). I am aware of no records for Procambarus (Capillicambarus) incilis Penn (1962), P. (G.) simulans (Faxon, 1884), P. (G.) species B, and Orconectes (Buan- nulifictus) palmeri longimanus (Faxon, 1898) in the Neches watershed. Perhaps this exposure of the comparative richness of the crayfish fauna of the Neches River basin will induce a more thorough survey by others of the crayfishes frequent- ing this and neighboring watersheds. Family Cambaridae Subfamily Cambarellinae Cambarellus (Pandicambarus) puer Hobbs Cambarellus puer Hobbs, 1945:469. Cambarellus (Pandicambarus) puer. — Fitz- patrick, 1983:268. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON New record.—Jasper Co.: Edge of creek 9.2 mi (14.7 km) NW of US Hwy 190 on St Rte 63, 3 6 II, 2 2, 17 Apr 1987, G. B. Hobbs & HHH. Subfamily Cambarinae Cambarus (Lacunicambarus) diogenes Girard Cambarus diogenes Girard 1852:88. Cambarus (Lacunicambarus) diogenes. — Hobbs, 1969:110. New records.—Newton Co.: 2.7 mi (4.3 km) NW of Newton (Courthouse) on US Hwy 190, 2j2, 11 Nov 1987, Brian F. Kens- ley & HHH. Polk Co.: 6.2 mi (9.9 km) NE of Farm Rd 3152 on Rd 350, 1 2, 13 Nov 1987, BFK & HHH. Tyler Co.: 2.7 mi (4.3 km) E of Horse Pen Creek on US Hwy 190, 2 6 II, 2 2, 13 Nov 1987, BFK & HHH. Remarks. — These specimens belonging to the subgenus Lacunicambarus are only ten- tatively assigned to this species. The ranges and limits of variation of the few described members of this species group are currently receiving a long-needed review by Ray- mond F. Jezerinac. All of the specimens cit- ed were obtained from burrows. Cambarus (Lacunicambarus) ludovicianus Faxon Cambarus diogenes var. Ludoviciana Fax- on, 1884:144. Cambarus (Lacunicambarus) ludovicia- nus. — Hobbs, 1990. New records. — Angelina Co: 5.6 mi (9 km) NW of US Hwy 59 on Farm Rd 2497, 1 9, 15 Apr 1987, GBH & HHH. Jasper Co.: 9.0 mi (14.4 km) SW of Jasper on US Hwy 190, 2 j6 1j2, 16 Apr 1987, GBH & HHH. Polk Co.: 12 mi (19.2 km) E of Livingston on US Hwy 190, 2 j2, 16 Apr 1987, GBH & HHH. 12.5 mi (20 km) E of Livingston on US Hwy 190, 1 2, 16 Apr 1987, GBH & HHH. 7.0 mi (11.2 km) W of US Hwy 287 on Farm Rd 1745, 1 61, 2 2, 12 Nov 1987, R. Ar- mentrout, J. David, BFK, HHH. San Au- gustine Co.: 19.4 mi (31 km) NE of Zavalla VOLUME 103, NUMBER 3 on US Hwy 69, 1 4 II, 1 2, 17 Apr 1987, GBH & HHH. 1.7 mi (2.7 km) NE of San Augustine on Farm Rd 353, 8 Nov 1987, 1 6 Il, BFK & HHH. Remarks. —See ““Remarks”’ under Cam- barus (L.) diogenes which are also applicable here. All of the specimens cited were ob- tained from burrows. Fallicambarus (C.) fodiens Astacus fodiens Cottle 1863:217. Cambarus hedgpethi Hobbs, 1948:224. Fallicambarus (Creaserinus) fodiens. — Hobbs, 1973:463.—Hobbs & Robison, 1989:672. New records.— Angelina Co.: southeast- ern part of Lufkin, | 2, 13 Apr 1987, Mike Whiteman & HHH. 3 mi (4.8 km) NW of US Hwy 59 on Farm Rd 2497, 1 éI, 6 8, 14 Apr 1987, GBH & HHH. 4.2 mi (6.7 km) NW of US Hwy 59 on Farm Rd 2497, 141,32, 1j6, 14 Apr 1987, GBH & HHH. 5.6 mi (9 km) NW of US Hwy 59 on Farm Rd 2497, 1 61, 16 II, 8 9, 14 Apr 1987, GBH & HHH; 3 4 Il, 5 2, 1 jd, 1 32, 15 Apr 1987, GBH & HHH. 7.7 mi (12.3 km) NW of Hwy 59 on Farm Rd 2497, 2 j4, 1 32, 15 Apr 1987, GBH & HHH; 4 jé, 2 j?, 9 Nov 1987, BFK & HHH. Jasper Co.: 0.3 mi (0.5 km) N of Angelina River on St Rte 63, 2 3 II, 1 29, 11 Nov 1987, BFK & HHH. 0.2 mi (0.3 km) SE of Angelina River on St Rte 63, 2oll, 11 Nov 1987, BFK & HHH. 3.1 mi (5 km) SE of Angelina River on St Rte 63, 131,12, 146, 4 ovig. 29, 11 Nov 1987, BFK & HHH. Polk Co.: 5.0 mi (8 km) W of Livingston on US Hwy 190, 1 ¢ II, 13 Nov 1987, BFK & HHH. Trinity Co.: 4 mi (6.4 km) SW of Farm Rd 357 on Rd 2262, 1 6 II, 16 Nov 1987, BFK & HHH. 9.0 mi (14.4 km) SW of Farm Rd 357 on Rd 2262, 1 6 II, 1 2, 16 Nov 1987, BFK & HHH. 10.6 mi(17 km) SW of Farm Rd 357 on Rd 2262, 1 2, 16 Nov 1987, BFK & HHH. Remarks. — This crayfish has been treated in considerable detail by Hobbs & Robison (1989). All of the adult specimens cited here were obtained from burrows, most of the juveniles from temporary roadside pools. 575 Fallicambarus (Fallicambarus) devastator Hobbs & Whiteman Fallicambarus (Fallicambarus) devastator Hobbs & Whiteman, 1987:403. New records.—8 localities in Angelina, Houston, Polk, Trinity, and Tyler counties (see Hobbs & Whiteman 1990). Except for a few specimens in one locality obtained near or at the mouths of burrows at night, all were removed from burrows that were excavated. Faxonella beyeri (Penn) Orconectes (Faxonella) beyeri Penn, 1950: 166. Faxonella beyeri. —Creaser, 1962:3. New record. — Angelina Co.: 4.2 mi (6.7 km) NW of US Hwy 59 on Farm Rd 2497, 3 61, 48 6 Il, 48 2, 14 Apr 1987, GBH & HHH. 5.6 mi (9 km) NW of US Hwy 59 on Farm Rd 2497, 2 61, 7 6 Il, 8 2, 15 Apr 1987, GBH & HHH. Jasper Co.: 8.7 mi (13.9 km) NW of St Rte 63 on US Hwy 190, 1 j2, 17 Apr 1987, GBH & HHH. San Au- gustine Co. 19.4 mi (31 km) NE of Zavalla on St Rte 147, 1 61, 22, 17 Apr 1987, GBH & HHH. Remarks. —Specimens of this species from the Neches River Basin are distinctly larger than any members of the species that I have examined from elsewhere. The largest first form male and female, both from the last locality cited, have carapace lengths of 18.2 and 19.6 mm, respectively. All of the spec- imens cited were taken from among vege- tation in open water. Procambarus (Girardiella) curdi Reimer Procambarus species E.—Reimer 1969:64. Procambarus species A.—Reimer & Clark, 1974:171. Procambarus (Girardiella) curdi Reimer, 1975:22. New records. — Angelina Co.: ditch at jct of farm rds 287, 1336, & 324 in SW part of Lufkin, 16 jé, 21 j2, 13 Apr 1987, MW, 576 Table 1.—Measurements (mm) of Procambarus (G.) nigrocinctus. Holotype Allotype | Morphotype Carapace: Entire length 37.5 42.2 31.2 Postorbital length 29.2 33.4 23.3 Width 2.1 Dp) 1.6 Height 17.5 20.5 13.3 Areola: Width 2.1 DP) 1.6 Length 11.1 13.5 9.3 Rostrum: Width 5.9 6.6 4.9 Length — 10.5 9.1 Right chela: Length, palm 10.4 8.7 4.8 mesial margin Palm width 11.4 9.9 Sol/ Length, lateral 31.5 26.2 15.8 margin Dactyl length 18.5 16.2 9.3 Abdomen: Width 14.7 18.1 11.9 Length 39.4 42.1 BQ) GBH, HHH. 2.4 mi (3.8 km) NW of US Hwy 59 on Farm Rd 2497, 1 4 Il, 3 2, 1 jd, 6 j2, 14 Apr 1987, GBH & HHH. 5.6 mi (9 km) NW of US Hwy 59 on Farm Rd 2497, 1 3 II, 42, 1 34, 1 32, 14 Apr 1987, GBH & HHH; 2 4 II, 2 2, 4 j4, 5 j2, 15 Apr 1987, GBH & HHH. 7.7 mi (12.3 km) NW of US Hwy 59 on Farm Rd 2497, 2 9, 4 36, 3 j9, 15 Apr 1987, GBH & HHH; 2 61,3 4 Il, 2 2, 1 j4, 5 j2, 9 Nov 1987, BFK & HHH. 0.3 mi (0.5 km) W of Farm Rd 326 on Rd 1475, 1 2, 1 js, 10 Nov 1987, BFK & HHH. An- gelina County Airport, 1 2, 10 Nov 1987, BFK & HHH. 1.3 mi (2.1 km) W of Shaw- nee Creek on Farm Rd 1818, 13 jé, 11 Jj, 11 Nov 1987, BFK & HHH. NW city limits of Zavalla on US Hwy 69, 1 2, 1 2 with young, 11 Nov 1987, BFK & HHH. Polk County: 15.1 mi (24.2 km) SE of Trinity on St Hwy 356, 1 j2, 16 Apr 1987, GBH & HHH. 2.4 mi (3.8 km) W of Tyler Co line on US Hwy 190, 1 2, 13 Nov 1987, BFK & PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON HHH. 1.8 mi (2.9 km) N of Farm Rd 350 on Rd 3152, 1 jg, 13 Nov 1987, BFK & HHH. Remarks. —This crayfish does not shun sandy soil in which the burrows, consisting of a single shaft and lacking conspicuous chambers, spiral to depths of as much as one meter. Whereas most of the specimens cited were retrieved from burrows, a few were found in roadside pools, several of which were temporary. Procambarus (Girardiella) nigrocinctus, new species Figs. i, 2a, Table 1 Procambarus species F.— Reimer, 1969:68. Diagnosis.—Body pigmented, eyes well developed. Rostrum with marginal spines, lacking median carina. Carapace with strong cervical spine. Areola 4.5 to 6.2 (mean 5.4 + 0.77) times as long as broad and consti- tuting 28.5 to 31.6 (mean 30.0 + 1.20) per- cent of total length of carapace (36.8 to 40.9, mean 39.1 + 1.42, percent of postorbital carapace length). Suborbital angle very weak and obtuse; hepatic area with few small tu- bercles; branchiostegal spine comparatively strong. Antennal scale about 2.5 times as long as broad, widest at about midlength. Mesial surface of palm of chela bearded; ventral surface tuberculate, tubercles pres- ent along proximal half of ventral surface of dactyl. Ischium of third pereiopod in first form male with simple strong hook over- reaching basioischial articulation; hook op- posed by small setiferous tubercle on cor- responding basis; coxa of fourth pereiopod lacking caudomesial boss. First pleopods of first form male reaching coxae of third pereiopods, symmetrical, bearing proxi- momesial spur and somewhat produced ce- phalic shoulder at base of terminal elements, lacking lateral subterminal setae, setae on caudoproximal ridge directed caudally, not flared; terminal elements (all sclerotized at least distally) consisting of (1) straight, ta- pering, subspiculiform, distally directed VOLUME 103, NUMBER 3 mesial process; (2) short, acute, distally di- rected cephalic process; (3) strong, acute, cephalocaudally flattened, and distally dis- posed central projection not overreaching (4) conspicuous, somewhat tapering, obliquely positioned caudal element; latter consisting of caudomesially excavate la- melliform lobe and lacking distinct digiti- form prominence. Lamelliform lobe and central projection reaching almost same level distally. Female with annulus ventralis freely movable, about 1.3 times as long as broad, and subrhomboidal (but tilted) in outline; cephalomedian trough broadening rapidly posteriorly by strong dextral diver- gence of dextral wall; sinus originating ad- jJacent to wall near midlength of annulus, and, after extending mesially to median line, following sinuous course caudally, ending on caudomedian surface of annulus; ce- phalic part of sinistral wall bordering trough tuberculate. Preannular plate poorly devel- oped; first pleopods present. Holotypic male, form I.—Cephalothorax (Fig. la, 1, 2a) subcylindrical. Second seg- ment of abdomen distinctly narrower than thorax (14.7 and 18.1 mm). Areola 5.3 times as long as broad and with 4 or 5 punctations in narrowest part. Cephalic section of car- apace approximately (rostrum broken) 2.4 times as long as areola, latter comprising about 29.5% of total length of carapace (38.0% of postorbital carapace length). Sur- face of carapace punctate dorsally, strongly granulate laterally; most tubercles in hepatic area little, if any, larger than granules on branchiostegites. Rostrum broad basally, tapering gently anteriorly from level of orbit (apical part of rostrum missing in holotype but in other specimens gradually diminish- ing in width to well developed marginal spines; slightly upturned tip reaching ulti- mate podomere of antennular peduncle); margins not thickened; dorsal surface, lack- ing median carina, concave with puncta- tions scattered between submarginal rows. Subrostral ridges weak and evident in dorsal view only slight distance anterior to pos- Sif terior margin or orbit. Postorbital ridges prominent, each ending anteriorly in cor- neous spine. Suborbital angle weak and ob- tuse. Branchiostegal spine and cervical spine well developed. Abdomen (Figs. 1k, 2a) subequal in length to carapace in all specimens. Cephalic sec- tion of telson with 3 spines in each caudo- lateral corner, lateral and mesial ones fixed. Cephalic lobe of epistome (Fig. 11) campan- ulate with slightly irregular, weakly-thick- ened anterolateral margins; main body of epistome with distinct fovea. Ventral sur- face of proximal podomere of antennular peduncle with small spine near midlength. Antennal peduncle with well developed spine on both basis and ischium; flagellum almost reaching midlength of telson. An- tennal scale (Fig. 1m) 2.5 times as long as broad, widest near midlength; greatest width of lamellar area 1.7 times that of thickened lateral part. Third maxilliped extending slightly distal to spine on ventral surface of basal podo- mere of antennule, ventral surface of pod- omeres proximal to propodus largely ob- scured by long plumose setae; lateral surface of all except distalmost part of merus hid- den in lateral aspect. Right chela (Fig. 1p) subovate in cross section, somewhat depressed; palm about 1.1 times as broad as length of mesial mar- gin; latter little more than one-third total length of chela; entire palm studded with tubercles, although those on and adjacent to mesial surface hidden by beard of plu- mose setae; 7 or 8 tubercles present in me- sialmost row, all except proximal and dis- talmost well obscured by tufts of plumose setae, row flanked by several somewhat ir- regular rows of tubercles; ventral ridge ad- jacent to base of dactyl bearing 1 tubercle larger than most others on ventral surface of palm; dorsolateral tubercles conspicu- ously smaller and more depressed than those more mesially located on dorsal surface. Both fingers with low median longitudinal ridges dorsally and ventrally, ridges flanked 578 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Procambarus (Girardiella) nigrocinctus (all from holotype except c and g which are from morphotype, and e and o from allotype): a, Lateral view of carapace (rostrum broken, reconstructed on basis of other specimens); b, c, Mesial view of first pleopod; d, Cephalic view of apical part of first pleopod; e, Annulus ventralis and adjacent sternites; f, Caudal view of apical part of first pleopod; g, h, Lateral view of first pleopod; i, Epistome; j, Basal podomeres of third, fourth, and fifth pereiopods; k, Lateral view of abdomen; |, Dorsal view of carapace (rostrum broken, reconstructed on basis of other specimens); m, Antennal scale; n, Caudal view of first pleopods; 0, p, Dorsal view of distal podomeres of cheliped. VOLUME 103, NUMBER 3 Se, expt Stormy itive 579 Fig. 2. Dorsolateral views of new crayfishes: a, Procambarus (Girardiella) nigrocinctus; b, Procambarus (Girardiella) kensleyi. by tubercles along proximal half and by se- tiferous punctations along distal half. Op- posable margin of fixed finger with row of 23 (left chela with 20) tubercles (more distal Ones too small to be included in Fig. |p), fifth from base largest, those distal to fifth decreasing in size to ultimate, which located twice length of corneous tip of finger from apex; row of 3 (2 on left) strongly cornified tubercles borne more ventrally on opposa- ble margin in distal third of finger; longi- tudinal band of minute denticles extending almost entire length, broadening in area be- tween rows of tubercles in distal third of 580 finger; lateral margin with row of tubercles almost reaching midlength of finger. Op- posable margin of dactyl with row of 17 (left with 12) tubercles, 7th (Sth on left) from base largest, along proximal two-thirds of finger, few additional contiguous small tu- bercles on dorsal side of row, and distal to 17th (12th on left) tubercle, dorsal row of 4 and ventral one of 3 small tubercles con- tinuing distally separated by band of minute denticles reaching base of corneous tip of finger; denticle band beginning at base of finger interrupted by principal tubercular row; mesial surface of finger with subserrate row of 17 (15 on left) tubercles. Carpus of cheliped longer than broad with very shallow oblique furrow flanked me- sially by squamous tubercles and laterally by scattered punctations; mesial surface with 2 acute tubercles somewhat larger than oth- ers and ventromesial triangular patch of tu- bercles with apex directed toward large, strongly acute tubercle on ventromesial dis- tal margin of podomere; another similar strong tubercle on ventrodistal margin flanking articular condyle; otherwise ventral and lateral surfaces sparsely setose punctate. Merus tuberculate dorsally with 2 strong, spiniform tubercles short distance proximal to distal margin; lateral, and most of mesial surfaces sparsely punctate, although dis- tomesial surface with few small tubercles; ventral surface of podomere with mesial row of 15 tubercles and lateral one of 8 followed by oblique row of 4 joining distal ends of lateral and mesial rows; few additional tu- bercles present between and to the sides of the longitudinal rows. Strong spine present on distolateral angle. Ischium with row of 5 small tubercles ventromesially. Hook on ischium of 3rd pereiopod (Fig. 1j) simple, heavy, overreaching basioischial articulation, and opposed by weak, setifer- ous tubercle on corresponding basis. Coxa of 4th pereiopod lacking caudomesial boss, that of 5th with small triangular one. Sternum between 3rd, 4th, and 5th pe- reiopods moderately deep; ventrolateral PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON margins with plumose setae obscuring much of lst pleopods. First pleopods (Fig. 1b, d, f, h, n) as de- scribed in “‘Diagnosis.” Uropods with both lobes of basal podo- mere bearing acute spines; mesial ramus with weak median carina but with well de- veloped, distinctly-premarginal distomedi- an spine; distolateral spine strong. Allotypic female. — Differing from holo- type, other than in secondary sexual fea- tures, in following aspects: apex of rostrum reaching slightly beyond midlength of ulti- mate podomere of antennular peduncle; in respect to total carapace length, areola pro- portionately longer (32.0% of carapace length, 40.0% of postorbital carapace length) than in any other specimens available, but apical part of rostrum probably regenerated; cephalic section of telson with 2 fixed spines in each caudolateral corner; cephalic lobe of epistome subtriangular with irregular ce- phalolateral margins; 3rd maxilliped slight- ly overreaching basal podomere of anten- nule; mesial margin of palm of right chela (Fig. lo) (left chela partly regenerated) lack- ing beard; opposable margin of fixed finger of chela with row of 11 tubercles, distalmost lying proximal to large tubercle situated ventral to level of row, denticles dispersed in single row; opposable margin of dacty! with row of 16 tubercles, 6th from base larg- est, 2nd row lacking, and denticles, for most part, arranged in single row, mesial surface with row of 15 tubercles, more distal ones not so well developed as in holotype; merus with ventromesial row of 13 tubercles, and 4 in row on ischium. Annulus ventralis (Fig. le) as described in “Diagnosis.” Preannular plate incon- spicuous, deeply embedded in sternum, and consisting of narrow calcified arch, antero- median part fusing with sternite XIII. Post- annular sclerite subtriangular 1.7 times as broad as long and wider than, but only 0.7 as long as, annulus and bearing shallow, me- dian longitudinal depression. First pleopod comparatively well developed, overreach- VOLUME 103, NUMBER 3 ing cephalic margin of annulus when ab- domen flexed. Morphotypic male, form IT.—Differing from holotype in following respects: apex of acumen reaching distal extremity of anten- nular peduncle; spines on body and on preoral appendages more strongly devel- oped than in holotype and allotype; anten- nal scale broadest slightly proximal to mid- length; beard on mesial surface of palm less well developed but clearly evident; oppos- able margin of fixed finger with row of 10 tubercles, 4th from base largest, corre- sponding margin of dactyl with row of 12 tubercles, 5th from base largest, and 2 more ventral tubercles near midlength of finger; mesial margin of dactyl with row of 13 tu- bercles; ventral surface of merus of cheliped with lateral row of 8 tubercles, mesial one of 12, and distal connecting row of 4; is- chium of cheliped with ventromesial row of 4 tubercles; hooks on ischia of 3rd pereio- pods very small, tuberculiform. First pleo- pod (Fig. lc, g) with shoulder on cephal- odistal margin much weaker than that in holotype and not produced; mesial process much heavier than that in holotype, ce- phalic process not clearly distinguishable from cephalic rim partly encircling and forming part of bulbous central projection; caudal process well developed but, like oth- er terminal elements, non-corneous. Color notes.—Holotype (Fig. 2a): Pre- dominant coloration tan with brown mark- ings, most in form of small spots. Cephalic region of carapace tan with small brown spots and few pale ones along lateral rostral and postorbital ridges; latter with dark brown line extending along ventrolateral margin. Mandibular adductor region with complex patterns of fine spots forming paired semielliptical splotches abutting cer- vical groove. Thoracic region spotted like cephalic region and with conspicuous, paired, almost black, semielliptical splotch- es at posterior dorsolateral extremity; splotches narrowing and converging me- sially into narrow transverse band. Abdo- 581 men similarly spotted but with darker pig- ments forming paired dorsolateral lines from first through basal part of 6th terga, and bases of pleura set off by scalloped sublinear series from which paler subtriangular patches extending laterally onto pleura; oth- er spots scattered between lines and patches. Telson and uropods with anastamosing maculations exhibiting same color con- trasts. Antennular and antennal peduncles and antennal scale spotted; flagella of both appendages tan. Third maxillipeds cream. Chelipeds similar in color to carapace (i.e., cream tan with brown spots) except tan re- duced almost to cream over much of carpus, propodus, and dactyl; major tubercles on merus and carpus and those on mesial mar- gin of palm and on fingers at least tipped with cream; those scattered over dorsum of merus, carpus, and palm very dark brown; both fingers fading from base distally to dis- tinctly pinkish orange distal fourth. Setal tufts on mesial surface of palm tan. Second through 5th pereiopods cream basally but with spots and splotches of grayish tan from merus through propodus. Coloration of female differing in no re- markable way from that of male. Type locality. —Jack Creek, a tributary of the Neches River, at State Route 94, about 3.0 mi (4.8 km) WSW of Lufkin Perimeter Route 287, Angelina County, Texas. There the creek was some 2 to 8 m wide, no more than 0.7 m deep, and flowed with a mod- erate current over a sandy and rocky bot- tom. The sometimes clear, weak-coffee col- ored water was slightly cloudy in November. Adult crayfish were found under the bridge among rocks and debris that had accumu- lated adjacent to the pilings. Just down- stream from the bridge a stand of Myrio- phyllum was present in a sun-drenched area, and from the debris in this clump, a few juveniles were taken. Several burrows along the bank were examined, but no crayfish was found in them. Shading the creek were trees belonging to the genera Liquidambar, Quercus, Pinus, Acer, and Salix. The pop- 582 ulation in the area of the bridge is rather small, for only five adult specimens were collected on two visits to the locality, 16 April 1987 and 9 November 1987. Juve- niles were far more common in April than in November. Disposition of types.—The holotype, al- lotype, and morphotype (4 I, ?, 6 II) are deposited in the National Museum of Nat- ural History (Smithsonian Institution), nos. 219436, 219437, and 219438, respectively, as are the paratypes consisting of 1 6 I, 2 6 II, 3 9, 22 j4, and 30 je. Size.—The largest specimen available is the allotypic female which has a carapace length of 42.2 (postorbital carapace length 33.4) mm despite the fact that the anterior part of the rostrum is apparently regener- ated perhaps shorter than it would have been had it not been injured. The smaller of the two first form males has corresponding lengths of 37.3 and 28.9 mm. Ovigerous females or ones carrying young have not been collected. Range and specimens examined. —This crayfish has been collected in only five lo- calities, all in the Neches River basin in Angelina and Jasper counties, Texas. An- gelina County: (1) Type locality, 1 61, 16 II, 1 9, 6 jd, 18 j2, 16 Apr 1987, GBH & HHH; 1 61, 1 9, 1 j2, 9 Nov 1987, BFK & HHH. (2) Jack Creek at Farm Road 2497, SW of Lufkin, 8 jé, 6 j2, 14 Apr 1987, GBH & HHH; 1 6 Il, 2 2, 1 jé, 1 je, 9 Nov 1987, BFK & HHH. (3) Moccasin Creek at Farm Rd 2497, 4.9 mi (7.8 km) SE of jct with St Rte 94, 1 4 Il, 6 jé, 3 j2, 9 Nov 1987 BFK & HHH. (4) Pool in roadside ditch on Farm Rd 2407, 1.2 mi(1.9 km) SE of intersection with St Rte 94, 1 jg. 9 Nov 1987, BFK, HHH. Jasper County: (5) Sand-bottomed creek at St Rte 63, 1.1 mi (1.8 km) SE of Angelina River, 1 jé, 11 Nov 1987, BFK & HHH. Almost certianly Reimer’s “Procam- barus species F” is conspecific with this crayfish (unfortunately, these specimens have not been available to me); he reported its presence in Anderson County : (6) 4.1 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mi E of Slocum. Nacodoches County: (6) 5 mi E of Nacogoches; (7) 4 or 5 mi NW of Nacodoches; (8) 9 mi W of Nacogoches; (10) 2.5 mi SW of Garrison, Polk County: (11) 2 mi S of Corrigan. Smith County: (12) 3.5 mi N of Troup. Shelby County: (13) 13 mi N, 2 mi W of San Augustine. Variations. —With so few adult speci- mens, and these from nearby localities, it is not surprising that the range of variation noted among them seems very small. One of the most conspicuous variations occurs in the relative development of the beard on the mesial margin of the palm of the chela: it is much the longest and densest in first form males, but it appears in the young by the time they have attained a carapace length of 10 mm. It becomes more obvious with succeeding molts in both sexes, and in at least some juvenile males approaching the molt that will take them into first form, the setal tufts are little, if any, better developed than they are in adult females. In some of the latter, the beard is reduced to a few small patches of setae that scarcely rise above the level of the adjacent tubercles, as occurs in the allotype. The chief differences noted in the rostrum seem to be associated with in- jury, primarily the loss of part or all of the acumen, and in several specimens, includ- ing both first form males, the acumen is either absent or has obviously been broken and regenerated, its tip not attaining the dis- tal extremity of the antennular peduncle. The areola in specimens with carapace lengths of at least 27.8 mm ranges from 4.5 to 6.2 times as long as broad and comprises from 28.5 to 32.0% of the total length of the carapace (37.4 to 40.9% of postorbital length). The spines on the carapace and che- lipeds are almost always well developed, al- though occasionally one or more may be shorter than the average (asymmetry in de- gree of development suggests that some re- ductions might well be associated with in- jury and regeneration). The spines in the caudolateral corners of the cephalic section of the telson vary in size and number: there VOLUME 103, NUMBER 3 may be two, three, or four in each corner, but only one, that immediately mesial to the fixed lateral spine, in each corner is movable. The differences noted in the sec- ondary sexual features deserve no com- ment. Relationships.—The color pattern, the bearded chelae, the comparatively broad areola, and the proximity of the range to that of Procambarus (Girardiella) tulanei Penn, 1953, attest to the assumed close re- lationship existing between P. (G) nigro- cinctus and that species. They are both ““speckled”’ and, although less well defined in tulanei, corresponding dark areas (line on lateral flank of postorbital ridges; large area over and adjacent to the mandibular ad- ductor region; and parts of the dark band on the caudal margin of the carapace) are comparable. The areolae of the primary types of P. tulanei, according to Penn’s mea- surements, range from 7.8 to 9.0 times as long as wide, in all of the available adults of P. nigrocinctus, from 4.5 to 6.2. The ranges of the two are contiguous, for P. tulanei has been found as far west in Louisiana as the Sabine River basin, which parallels the Neches watershed immediately to the west. Insofar as is known, however, their ranges are allopatric. The presence of marginal spines on the rostrum, well developed cervical spines, a distinct dark band at the caudal margin of the carapace, and a first pleopod of the male in which the central projection does not ov- erreach the lamelliform lobe of the caudal process will serve to distinguish P. (G.) ni- grocinctus from all other members of the subgenus Girardiella. Ecological note.— All except one of the specimens available to us were taken from streams. The single juvenile female ob- tained at locality 4, however, came from what appears to be a permanent roadside pool that appeared to be distant from a body of flowing water. Etymology. —Niger (L.) = black + Cinc- tus (L.) = girdle; alluding to the narrow black 583 Table 2.— Measurements (mm) of Procambarus (G.) kensleyi. Holotype Allotype | Morphotype Carapace: Entire length 31.8 33.5 31.3 Postorbital length 25.5 27.0 24.9 Areola: Width 2.1 2.4 1.6 Length 10.2 10.3 9.8 Rostrum: Width 5.2 6.3 5.2 Length 7.3 ES) Ve Right chela: Length, palm 8.0 6.3 S\.7) mesial margin Palm width 8.3 8.2 6.6 Length, lateral 23.3 20.0 18.6 margin Dactyl length 13.4 11.7 11.0 Abdomen: Width 13.5 14.8 12.5 Length 32.8 37.2 31.7 band across the caudal margin of the cara- pace. Associates. —Collected with this crayfish in one or more localities were: Procambarus (Ortmannicus) a. acutus (Girard, 1852), P. (Girardiella) curdi Reimer, 1975, P. (Scapu- licambarus) clarkii (Girard, 1852), P. (O.) nechesae n. sp., and Fallicambarus (C.) fodi- ens (Cottle, 1863). Procambarus (Girardiella) kensleyi, new species Figs. 2b, 3, Table 2 Procambarus species F.—Reimer, 1969:73 [in part?]. Diagnosis. —Body pigmented, eyes well developed. Rostrum without marginal spines, lacking median carina. Carapace without cervical spine. Areola 4.0 to 6.2 (mean 5.1 + 0.81) times as long as broad and constituting 30.4 to 34.7 (mean 32.2 + 1.26) percent of total length of carapace (36.9 584 to 42.9, mean 40.1 + 1.63, percent of post- orbital carapace length). Suborbital angle obtuse; hepatic area weakly tuberculate; branchiostegal spine rather weak. Antennal scale about 2.2 times as long as broad, wid- est slightly distal to midlength. Mesial sur- face of chela not bearded, ventral surface tuberculate, tubercles present along proxi- mal half of ventral surface of dactyl. Is- chium of third pereiopod in first form male with simple hook overreaching basioischial articulation; hook not opposed by tubercle on corresponding basis; coxa of fourth pe- reiopod lacking caudomesial boss. First pleopods of first form male reaching coxae of third pereiopods, symmetrical, bearing proximomesial spur, and subangular shoul- der at base of terminal elements lacking lat- eral subterminal setae; setae on caudoprox- imal ridge directed caudally, not flared; terminal elements (all sclerotized at least distally) consisting of (1) straight, tapering, subspiculiform, distally-directed mesial process; (2) very short, acute, distally-di- rected cephalic process; (3) strong, corni- fied, acute, cephalocaudally-flattened, dis- tally-disposed central projection not overreaching (4) prominent, distally-cor- nified, flattened, obliquely-positioned cau- dal element, latter consisting of caudome- sially excavate lamelliform lobe with small digitiform prominence situated in cau- domesial concavity. Lamelliform lobe and central projection reaching almost same level distally. Female with annulus ventralis freely movable, about 1.2 times as broad as long, and subrhomboidal (but tilted) in out- line; moderately deep cephalomedian trough, flanked by smooth to tuberculate ridges, broadening posteriorly, dextral wall flaring more strongly than sinistral one; si- nus originating in caudodextral part of trough, forming symmetrical loop caudo- dextrally over tongue and fossa, then turning caudosinistrally across median line before curving caudally and slightly dextrally and ending just anterior to caudal margin of an- nulus. Preannular plate poorly developed; first pleopods present. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Holotypic male, form I.—Cephalothorax (Fig. 2b, 3a, 1) subcylindrical. Second seg- ment of abdomen distinctly narrower than thorax (13.5 and 15.9 mm). Areola 4.9 times as long as broad and with room for 4 or 5 punctations across narrowest part. Cephalic section of carapace 2.1 times as long as areo- la, latter comprising 32.1% of total length of carapace (40.0% of postorbital carapace length). Surface of carapace punctate dor- sally, hepatic area mostly tuberculate and lateral and ventrolateral parts of branchios- tegites granulate, anteroventral branchio- stegal region tuberculate. Rostrum broad, gradually tapering from base to level of dis- tal extremity of proximal podomere of an- tennule, anteriorly contracting more rapidly to apex, which reaching almost midlength of ultimate podomere of antennular pedun- cle; margins not thickened and without spines or tubercles; dorsal surface, lacking median carina, concave with punctations scattered between prominent submarginal rows. Subrostral ridges weak and not evi- dent in dorsal aspect. Suborbital angle in- conspicuous and obtuse. Branchiostegal spine moderately well developed; cervical spine represented by small tubercle. Abdomen (Figs. 2b, 3k) shorter than car- apace. Cephalic section of telson with 2 spines in each caudolateral corner, laterat one fixed. Cephalic lobe of epistome (Fig. 31) subtriangular with irregularly crenulate anterolateral margins; main body of epi- stome with distinct fovea. Ventral surface of proximal podomere of antennular pe- duncle with small spine near midlength, mesial margin of peduncle with plumose se- tae, especially 2 more distal podomeres. An- tennal peduncle with small spine on both basis and ischium, distal 3 podomeres bear- ing prominent mesial fringe of plumose se- tae; flagellum reaching fourth abdominal tergum. Antennal scale (Fig. 3m) about 2.2 times as long as broad, widest slightly distal to midlength; greatest width of lamellar area about 2 times that of thickened lateral part. Third maxilliped almost reaching distal end of proximal podomere of antennule; VOLUME 103, NUMBER 3 585 Fig. 3. Procambarus (Girardiella) kensleyi (all from holotype except c and g from morphotype, e and o from allotype, and j from topotypic first form male): a, Lateral view of carapace; b, c, Mesial view of first pleopod; d, Cephalic view of distal part of first pleopod; e, Annulus ventralis and adjacent sclerites; f, Caudal view of distal part of first pleopod; g, h, Lateral view of first pleopod; i, Epistome; j. Caudal view of first pleopods; k, Lateral view of abdomen; |, Dorsal view of carapace; m, Antennal scale; n, Basal podomeres of third, fourth, and fifth pereiopods; 0, p, Dorsal view of distal podomeres of cheliped. 586 ventral surfaces of ischium and merus stud- ded with dense mat of plumose setae; lateral surface of all except distalmost part of me- rus hidden in lateral aspect. Right chela (fig. 3p) (left regenerated) sub- Ovate in cross section, moderately de- pressed; palm slightly broader than length of mesial margin; latter slightly more than one-third total length of chela; entire palm studded with tubercles; 7 present in each of 3 mesialmost rows; ventral ridge opposite base of dactyl with 1 tubercle larger than others on ventral surface. Both fingers with low median longitudinal ridges dorsally and ventrally, ridges flanked by tubercles proxi- mally and setiferous punctations distally. Opposable margin of fixed finger with row of 17 small tubercles along proximal three- fourths of finger, sixth from base largest (more distal ones very small); larger (“‘lock- ing’’) tubercle present below distalmost tu- bercle of row; band of minute denticles ex- tending from base of finger to base of corneous tip between and distal to tubercles; lateral margin with row of tubercles reach- ing midlength of finger (not visible in Fig 3p). Opposable margin of dactyl with row of 12 comparatively small tubercles, fifth from base largest, along proximal two-thirds of finger; band of minute denticles extend- ing from base of finger to base of corneous tip between and distal to tubercles; mesial surface of finger with row of 12 tubercles, distalmost just proximal to corneous tip. Carpus of cheliped longer than broad with shallow oblique furrow flanked mesially by tubercles and few punctations and laterally by widely spaced punctations; mesial sur- face with 2 tubercles larger than others, one just distal to midlength and other at dor- sodistal angle, 9 additional ones present in proximal half; ventral surface with few squamous tubercles and punctations prox- imal to 3 subacute tubercles on distal mar- gin. Merus tuberculate dorsally with 2 strong spiniform tubercles near distal margin; me- sial surface with few tubercles near distal extremity, otherwise it and lateral surface PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with scattered punctations; ventral surface with mesial row of 14 tubercles, lateral one of 10, and oblique distal row of 4; tufts of short plumose setae present between rows; short, heavy, spiniform tubercle on disto- lateral extremity. Ischium with row of 4 tu- bercles ventromesially and another of 3 dor- sally. Hook on ischium of third pereiopod (Fig. 3n) simple and overreaching basioischial ar- ticulation, not opposed by tubercles on cor- responding basis. Coxa of fourth pereiopod without caudomesial boss, that of fifth with small rounded one. Sternum between third, fourth, an fifth pereiopods moderately deep, ventral mar- ginal fringe of plumose setae concealing much of first pleopods. First pleopods (Fig. 3b, d, f, h, j) as de- scribed in “Diagnosis.” Uropods with both lobes of basal podo- mere bearing acute spines; mesial ramus with premarginal distomedian spine at end of keel and strong lateral spine. Allotypic female. —Differing from holo- type, other than in secondary sexual fea- tures, in following respects: areola 30.7% of carapace length (38.1% of postorbital car- apace length); rostrum reaching distal end of penultimate podomere of antennular pe- duncle; flagellum of antenna reaching fifth abdominal tergum; chela (Fig. 30) with 6 tubercles present in mesialmost row on palm, 5 in adjacent dorsal row and 6 in adjacent ventral row; opposable margin of fixed finger with row of 9 tubercles, fourth from base largest; corresponding margin of dactyl with 9; opposable margins of both fingers with single longitudinal row of mi- nute denticles; mesialmost of 3 tubercles on ventral surface of carpus of cheliped situ- ated more dorsally; merus with ventrome- sial row of 15 tubercles, ventrolateral one of 11, and oblique distal row of 3; only 2 tubercles present on dorsal margin of is- chium. Annulus ventralis (Fig. 3e) as described in “Diagnosis.” Morphotypic male, form II.—Dijffering VOLUME 103, NUMBER 3 from holotype in following respects: areola 6.1 times as long as broad and consisting of 31.3% of length of carapace, 39.3% of post- orbital length; abdomen slightly longer than carapace; cephalic section of telson with 3 (right) or 4 (left) spines in each caudolateral corner; anterolateral margins of cephalic lobe of epistome almost smooth; antennal flagellum reaching base of telson; setae on third maxilliped less dense and most of me- rus visible in lateral aspect; row on mesial margin of palm of chela flanked by row of 6 tubercles dorsally and one of 7 ventrally; opposable margin of fixed finger with row of 11 tubercles, fourth from base largest; corresponding margin of dactyl with row of 13 tubercles; denticles on both fingers ar- ranged in single row; carpus with tubercles situated more similar to those in allotype; ventromesial angle of merus with row of 15 tubercles, 8 in lateral row, and 4 in oblique row; ischium with ventromesial row of 5 tubercles; hook on ischium of third pereio- pod and boss on coxa of fifth markedly re- duced; setae on ventrolateral margins of sternites XII and XIII much less well de- veloped. First pleopods (Fig. 3c, g) with reduced setation; terminal elements disposed as in holotype (cephalic process slightly more posteriomesially situated), and while short- er, more stocky and none corneous. Color notes. —(Based on recently molted first form male from Trinity County, Texas; see Fig. 2b for pattern.) Basic colors dark tan with reddish brown markings. Rostrum dark tan with few small brown splotches, some of which anastomosing in basal area; postorbital ridges with brown stripe flank- ing ventrolateral border and merging with anastomosing splotches forming anterior part of irregular longitudinal stripe; lateral part of cephalic region with mosaic dark brown pattern. Suborbital angle encom- passed by cream-colored splotch extending Over antennal and mandibular regions. Thoracic region with ill-defined, irregular stripe dorsolaterally and with distinctive spotted pattern laterally (spots and back- 587 ground fading ventrally); areola mostly tan but with dark branchiocardiac grooves and submedian elongate spot anteriorly. Caudal ridge almost black; caudal flange gray. Ab- domen with tan dorsomedian stripe from first to anterior half of sixth abdominal ter- gum; stripe flanked by irregular, very nar- row brown stripe, and it, in turn, by broader dorsolateral ones continuous with irregular ones on branchiostegites; third pair of brown stripes formed by ventrally convex arcs across bases of pleura, these flanked dorsally and ventrally by mosaic of pale anasto- mosing splotches. Telson and uropods all tan with light reddish brown and cream spots; proximal podomere of uropods edged in dark brown, spines on telson and uro- podal rami also dark brown. Antennular and antennal peduncles and antennal scale with dark brown margins; flagella light brown with olive suffusion distally. Dorsum of chelipeds from merus distally tan with dark brown to black tubercles, ventral surface mostly pinkish cream to pinkish tan; tips of fingers purplish. Remaining pereiopods light pinkish tan with darker tan to brown splotches dorsally; distal part of merus and carpus darker than more proximal and dis- tal podomeres, which pinkish cream dor- sally and ventrally. Setal tufts on third max- illipeds cream. Type locality. —Roadside ditch on gentle slope 4.6 mi (7.4 km) NW of US Highway 59 on Farm Road 2497, Angelina County, Texas. There the crayfish were collected from simple burrows 0.5 to 0.8 m deep and topped by chimneys eight to 25 cm tall. The soil was a sandy clay supporting grasses and sedges. Dominant among the trees in the adjacent woods were Liriodendron tulipifera and members of the genera Pinus and Quer- CUS. Disposition of types.—The holotype, al- lotype, and morphotype (¢ I, 2, ¢ II) are deposited in the National Museum of Nat- ural History (Smithsonian Institution), nos. 219772, 219973, and 219774, respectively, as are the paratypes consisting of 461,36 II, 8 2, 1 j4, and 3 32. 588 Size. —The largest specimen available is a female from Tyler County, Texas, having a carapace length of 40.9 mm (postorbital length 32.8 mm). The largest and smallest first form males have corresponding lengths of 36.3 (28.9) mm and 30.6 (25.2) mm. Measurements are not available for females carrying eggs or young, neither of which has been collected. Range and specimens examined. —This crayfish has been found in eleven localities in eastern Texas as follows (The specimens in those collections preceded by asterisks are excluded from the type series.): Angelina County: (1) Type locality, 2 6 I, 2 6 II, 3 9, 1j9, 14 Apr 1987, GBH & HHH. *(2) Road- side ditch 5.6 mi (9.0 km) NE of US Hwy 59 on Farm Rd 2497, 1 jd, 2 j2, 14 Apr 1987, GBH & HHH. Jasper County: (3) Roadside ditch on St Rte 63, 0.7 mi (1.1 km) NW of Farm Rd 255, 1 6 I, 2 6 II (one molted to form I on 10 Feb 1988), 2 2, 11 Nov 1987, BFK & HHH. (4) Roadside ditch on US Hwy 190, 1.5 mi (2.4 km) E of Coun- ty Courthouse, 1 2, 12 Nov 1987, BFK & HHH. Newton County: *(5) Pool in road- side ditch 6.3 mi (10.1 km) NW of Newton on US Hwy 190, 13 juv, 11 Nov 1987, BFK & HHH. (6) Roadside ditch 2.7 mi (4.3 km) NW of Newton on US Hwy 190, 1 ¢ II, 1 jé, 1 j2, 11 Nov 1987, BFK & HHH. Panola County: *(7) Roadside ditch 9 mi (14.4 km) S of Carthage on US Hwy 96, 2 j6, 17 Apr 1987, GBH & HHH (tentatively assigned to this species). San Augustine County: (8) Burrow in creek bank 1.7 mi (2.7 km) NE of San Augustine on Farm Rd 353, 1 2, 8 Nov 1987, BFK & HHH. Trinity County: (9) Burrow in creek bank on Farm Rd 2262 9.0 mi (14.4 km) SW of Farm Rd 357, 1 3 II (molted to form I in laboratory), 1 j2, 14 Nov 1987, BFK & HHH. Tyler County: (10) Floodplain of Horsepen Creek on US Hwy 190, about 1.5 mi (2.4 km) E of Polk Co line, 1 2, 13 Nov 1987, BFK & HHH. (11) Floodplain of Big Cypress Creek on US Hwy 190, about 4 mi (6.4 km) E of Polk Co line, 1 2, 12 Nov 1987, BFK & HHH. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Should Reimer’s ““Pocambarus species G”’ prove to be conspecific with this crayfish, a number of localities cited by him expands the range of the species into western Loui- siana and southwestern Arkansas. The iden- ty of specimens from the localities offered by him should be confirmed. Variations. —With so few specimens available from all of the localities, there is no way to determine whether the few vari- ations noted are individual ones or typical of the populations of which they were mem- bers. No attempt is therefore made to note in which locality/ies these features were ex- hibited. The antennal scale may be broadest at midlength or slightly more distally; the third maxillipeds are all hirsute, but some decidedly more so than others; the subor- bital angle varies from being almost obso- lete to almost acute, but in none of the spec- imens is it prominent; differences noted in the first pleopod of the first form males in- clude reduction or absence of a proxi- momesial spur, a reduction in the cephalic process that renders it a virtual rudiment, and in one specimen the central projection slightly surpasses the caudal process distal- ly; the annulus ventralis exhibits much vari- ability in the nature of the ridges bordering the cephalomedian trough: they may be smooth and quite approximate so that their opposing slopes are almost vertical or, like the allotype (Fig. 3e), they may be tuber- culate with their crests more widely sepa- rated. While there are variations in the numbers of tubercles elsewhere and, to some extent, in their distribution, most fall within the range cited for the primary types. Tu- bercles on the dorsolateral margin of the ischium of the third maxilliped are absent in many crayfishes but here range in number from 2 to 5. Relationships. —Procambarus (G.) kens- leyi has its closest affinities with P. (G.) par- asimulans Hobbs & Robison (1982). In both of them the areola is broad (no more than nine times as long as wide) and short (con- stituting a maximum of 35% of the Cara- VOLUME 103, NUMBER 3 pace length), the opposable margin of the dactyl of the cheliped lacks a well defined excision in the basal third, the dorsolateral surface of the palm of the chela is tuber- culate, the central projection of the first pleopod of the first form male neither clear- ly overreaches nor projects laterally beyond the caudal process. They differ in that in P. (S.) kensleyi the areola is more densely punctate, the rostrum is almost always more tapering, the mesial margin of the anten- nular peduncle and the ventral and ventro- lateral surfaces of the third maxilliped are strongly hirsute, the cephalic process of the first pleopod does not extend beyond mid- length of the central projection which ex- tends as far distally as the caudal element, and the latter is less tapering. Ecological notes. —Even though all of the adult specimens belonging to this species that have been collected came from bur- rows, I am reluctant to state that it is a primary burrower, largely because of the simplicity of their domiciles which consist of a single subvertical, slightly sloping or loosely spiraling shaft that leads to a very slight enlargement at depths of 0.6 to 1.8 m. In most of the localities, the soil con- sisted of clay or sandy clay overlain by sand, occasionally by as much as 30 cm. Whereas no adults were found in open water, perhaps this reflects the habitats (burrows) selected by the collectors rather than the habitat dis- tribution of the crayfish. More frequently than not, a broad, short areola, such as that possessed by this crayfish, is that of a dwell- er of well areated streams. The lack of a reduction in the abdomen also suggests that it frequents open water. Some burrowing crayfishes can be enticed to the air-water interface in burrows (Hobbs, 1981:31), but not one of the specimens available could be lured from the fundus of its single shaft which lacked even one con- spicuous chamber. Etymology.—This crayfish is named in honor of my friend and colleague Brian F. Kensley whose assistance in collecting many 589 Table 3.— Measurements (mm) of Procambarus (O.) nechesae. Holotype Allotype Morphotype Carapace: Entire length 32.6 40.6 32.6 Postorbital length 25.5 20.9 24.7 Width 16.3 19.5 16.0 Height 16.0 19.0 15.1 Areola: Width 1.4 1.3 1.5 Length 10.4 12.9 9.6 Rostrum: Width 5.6 6.9 5.6 Length 8.0 11.3 8.9 Right chela: Length, palm 11.4 8.0* 7.9* mesial margin Palm width 8.2 Veda 6.3* Length, lateral 31.8 24.0* 23.8* margin Dactyl length 17.5 14.1* 13.7* Abdomen: Width 13.7 17.3 13.0 Length 33.7 43.3 33.8 * Left chela. of the crayfishes reported here was invalu- able. Associates. —Co-existing with this cray- fish in one or more localities were members of: Cambarus (L.) diogenes, C. (L.) ludovi- cianus, Fallicambarus (C.) fodiens, Procam- barus (G.) curdi, Faxonella beyeri, Procam- barus (O.) acutus acutus (Girard), and P. (Pe.) dupratzi Penn (1953). Procambarus (Ortmannicus) acutus acutus (Girard) Cambarus acutus Girard, 1852:91. Procambarus (Ortmannicus) acutus acu- tus. —Hobbs, 1972:9. New records. — Angelina Co.: 3.5 mi (5.6 km) NW of US Hwy 59 on Farm Rd 2497, 1 6 II, 1 j4, 14 Apr 1987, GBH & HHH. 4.2 mi (6.7 km) NW of US Hwy 59 on Farm Rd 2497, 1 31, 336, 2j2, 14 Apr 1987, GBH 590 & HHH. trib. Moccasin Creek at Farm Rd 2497, 4.9 mi (7.8 km) SE of St Rte 94, 1 6 II, 9 Nov 1987, BFK & HHH. 1.3 mi (2.1 km) W of Shawnee Creek on Farm Rd 1818, 336, 1j2, 11 Nov 1987, BFK & HHH. Jasper Co.: 8.7 mi (13.9 km) NW of US Hwy 190 on St Rte 63, 1 jé, 17 Apr 1987, GBH & HHH. 9.2 mi (14.7 km) NW of US Hwy 190 on St Rte 63, 3 4 II, 2 2, 17 Apr 1987, GBH & HHH. Newton Co.: 6.3 mi (10.1 km) NW of Newton on US Hwy 190, 1 jé, 2 3°, 11 Nov 1987, BFK & HHH. Polk Co.: 7 mi (11.2 km) W of US Hwy 287 off Farm Rd 1745 (David Farm), 2 ¢ II, 4 j6, 4 j9, 12 Nov 1987, J. David, RA, BFK, & HHH. Trinity Co.: 4 mi (6.4 km) SW of Farm Rd 357 on Rd 2262, 1 jd, 1 32, 14 Nov 1987, BFK & HHH. Remarks. — The specimens cited here were taken from sluggish streams, roadside pools, and from burrows consisting of single, sub- vertical shafts. Procambarus (Ortmannicus) nechesae, new species Fig. 4, Table 3 Diagnosis. —Body pigmented, eyes well developed. Rostrum of adults with or with- out marginal spines but lacking median ca- rina. Carapace with small cervical spine or tubercle. Areola 7.3 to 10.9 (mean 8.8 + 1.20) times as long as wide and constituting 29.1 to 32.4 (mean 30.9 + 1.03) percent of total length of carapace (38.0 to 42.3, mean 40.8 + 1.20 percent of postorbital length). Suborbital angle very weak and obtuse; postorbital ridges with cephalic spine or tu- bercle sometimes abraded; hepatic area weakly tuberculate; branchiostegal spine small to vestigial. Antennal scale little more than twice as long as broad, widest at about midlength. Ischia of third and fourth pe- reiopods with simple hooks, hooks of third distinctly overreaching basioischial articu- lation and lacking opposing tubercle on ba- sis, that of fourth almost reaching articu- lation and opposed by strong tubercle on basis; coxa of fourth pereiopod with strong PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON caudomesial boss, that of fifth much smaller and flattened. First pleopod of first form male reaching coxa of third pereiopod, asymmetrical, provided distolaterally with rounded prominence (caudal knob) bearing conspicuous tuft of subapical setae; termi- nal elements consisting of (1) mesial process tapering from base to acute tip, directed caudolaterally and extending distally to about tip of caudal process; (2) acute ce- phalic process directed caudally, hooding (3) beaklike central projection which also directed caudally; and (4) caudal process, arising from caudolateral extremity of shaft, bladelike with acute tip and directed cau- dodistally. Annulus ventralis about twice as broad as long, with nearly straight caudal margin and strongly arched cephalically; sinuous sinus arising near cephalic margin, progressing caudosinistrally in cephalic third of annulus before turning caudally and fi- nally caudodextrally ending on prominent protruding submedian tongue-like promi- nence. Sternum immediately anterior to an- nulus with low tubercles flanking median line. Unadorned postannular sclerite sub- triangular, almost two-thirds as broad as an- nulus. First pleopod present in female. Holotypic male, form I.—Cephalothorax (Fig. 4a, k) subcylindrical in section. Ab- domen narrower than thorax (16.7 and 20.5 mm). Greatest width of carapace slightly greater than height at caudodorsal margin of cervical groove. Areola 8.3 times longer than wide with 1 or 2 punctations in nar- rowest part. Cephalic section of carapace about 2.1 times as long as areola, length of latter 31.9% of entire length of carapace (40.8% of postorbital carapace length). Sur- face of carapace punctate dorsally, granulate laterally. Rostrum slightly deflected ven- trally with slender converging margins, its acute apex reaching slightly beyond mid- length of penultimate podomere of anten- nular peduncle; margins neither thickened nor provided with spines or tubercles, faint- ly contracted at base of short acumen; dorsal surface concave with many fine setiferous VOLUME 103, NUMBER 3 591 Fig. 4. Procambarus (Ortmannicus) nechesae (all from paratypic male, form I, except a and k from holotype, c and f from morphotype, | from allotype, and m from paratypic female): a, Lateral view of carapace, b, c, Mesial view of first pleopod; d, Caudal view of first pleopods; e, Basal podomeres of third, fourth, and fifth pereiopods; f, g, Lateral view of first pleopod; h, Epistome; i, Adductor face of mandible; j, Antennal scale; k, Dorsal view of carapace; 1, Annulus ventralis; m, n, Dorsal view of distal podomeres of cheliped. 592 punctations. Subrostral ridge evident in dorsal aspect for only short distance ante- rior to caudal margin of orbit. Postorbital ridge well developed, grooved dorsolateral- ly and bearing small tubercle at cephalic extremity. Suborbital angle very small and obtuse. Branchiostegal spine also small. Cervical spine represented by small tuber- cle. Abdomen and carapace subequal in length. Pleura of third through fifth seg- ments very broadly rounded, almost trun- cate ventrally but lacking posteroventral an- gles. Cephalic section of telson with 2 spines in each caudolateral corner, lateral ones im- movable; caudal margin of caudal section with shallow median excavation. Cephalic lobe of epistome (like Fig. 4h) broadly tri- angular with slightly elevated cephalolateral margins heavily fringed with plumose setae, central area convex; distinct anteromedian fovea present on main body. Ventral surface of proximal podomere of antennular pe- duncle with spine at midlength. Antenna with comparatively weak spiniform tuber- cles on basis and ischium; flagellum ex- tending almost to end of telson. Antennal scale (like Fig. 4j) almost 2.5 times as long as broad, widest at about midlength; great- est width of lamella about 1.8 times width of thickened lateral part. Mandible (like Fig. 41) as illustrated. Third maxilliped extending cephalically to base of penultimate podomere of antennule; is- chium not produced distolaterally, its ven- tral surface studded with plumose setae. Right chela (like Fig. 4n) subovate in cross section, not strongly depressed. Mesial sur- face of palm with row of 8 tubercles sub- tended by additional rows of more squa- mous ones dorsally and ventrally; tubercles present over all except ventrolateral part of palm and also present on basal parts of both fingers. Both fingers with low, rounded lon- gitudinal ridges dorsally and ventrally, all poorly defined except for being flanked by rows of setiferous punctations. Opposable margin of fixed finger with dorsal row of 9 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (10 on left) tubercles, third from base larg- est, on proximal half of finger, and ventral row of 6, fifth much larger than others, in middle third of finger; lateral margin with row of setiferous punctations. Opposable margin of dactyl with dorsal row of 11 tu- bercles in proximal half and ventral row of 13 (8 and 18, respectively, on left); mesial margin with row of 5 (7 on left) tubercles proximally followed by row of setiferous punctations. Mesial surface of dactyl with subserrate row of tubercles along proximal two-fifths. Carpus of cheliped longer than broad with distinct oblique furrow dorsally, tuberculate mesially and dorsomesially; mesial surface with 3 tubercles somewhat larger than others: 1 proximally, 1 near mid- length, and another distally; ventral surface with usual 2 tubercles on distal margin, oth- erwise setiferous. Merus tuberculate dorsally, distomesial- ly, and ventrally; 1 premarginal tubercle larger than others on dorsodistal surface; ventral surface with mesial row of 16 tu- bercles and lateral one of 11, 3 forming oblique distal row joining lateral and mesial rows. Ischium with ventromesial row of 3 tubercles. Hooks on ischia of third and fourth pe- reiopods (like Fig. 4e) simple, that on third overreaching basioischial articulation, that on fourth not overreaching articulation but opposed by prominent tubercle on corre- sponding basis. Coxa of fourth pereiopod with prominent, subvertically oriented cau- domesial boss; that of fifth with smaller one strongly compressed in longitudinal plane of body. Sternum between third, fourth, and fifth pereiopods comparatively deep with mat of plumose setae extending mesially from ventrolateral margins. First pleopods (like Fig. 4b, d, g) as de- scribed in “‘Diagnosis.” Uropod with both lobes of basal podomere bearing small acute spine; both rami with distolateral spines, and distomedian spine on mesial ramus sit- uated distinctly proximal to subtruncate distal margin. VOLUME 103, NUMBER 3 Allotypic female. — Differing from holo- type, except in secondary sexual characters, as follows: apex of rostrum almost reaching distal extremity of antennular peduncle; small corneous marginal spines flanking base of short, narrow acumen; suborbital angle almost obsolete; left cervical spine very small but with sharp apex; abdomen slightly long- er than carapace; chelipeds perhaps regen- erated but similar to chela of paratypic fe- male (like Fig. 4m); comparatively few plumose setae on third maxilliped; mesial surface of palm with row of 6 tubercles sub- tended by dorsal one of 6 and ventral one of 3; opposable margin of fixed finger with single row of 9 tubercles, third from base largest, on proximal half of finger with larger more ventrally located one almost at mid- length; single row of minute denticles ex- tending between tubercles and continuing to base of corneous tip of finger; opposable margin of dactyl with row of 13 tubercles, sixth from base largest, in proximal half (left with 12, fourth from base largest), minute denticles as on fixed finger; tubercles on me- sial surface of dactyl weaker than those in holotype; merus with 2 premarginal tuber- cles larger than others on dorsodistal sur- face, ventral surface with mesial row of 12 tubercles and lateral one of 11, oblique row inseparable from lateral; ischium with me- sial row of 4 (2 very small) tubercles. (See Table 3 for measurements.) Annulus ventralis (Fig. 41) as described in “Diagnosis.” Morphotypic male, form II.—Differing from holotype in following respects: rostral margins bearing minute tubercles at base of slender, short acumen, latter reaching ulti- mate podomere of antennular peduncle; ab- dominal pleura with posteroventral extrem- ities (especially of sixth segment) angular; flagella of antennae broken; third maxil- liped, probably regenerated at least in part, reaching midlength of proximal podomere of antennular peduncle, comparatively few plumose setae present; mesial surface of palm of chelae with mesialmost row of 6 593 tubercles, 6 in that on dorsal flank, and 4 in that on ventral flank; fixed finger of right chela with row of 10 tubercles (13 on left), third from base largest; opposable margin of dactyl with upper row of 19 (15 on left) and lower row of 8 (9 on left); mesial surface of carpus of cheliped with additional tu- bercle between proximalmost and that near midlength; ventral surface of merus with mesial row of 14 tubercles and lateral one of 15 (left with 12 and 14 respectively); is- chium with only 2 tubercles marking mesial row in holotype. Hooks on ischia of third and fourth pereiopods and bosses on fourth and fifth much reduced. First pleopods (Fig. 4c, f) with all terminal elements positioned as in holotype; mesial process comparatively much heavier, re- maining ones smaller and not nearly so clearly differentiated as in holotype; sub- apical setae and shoulder much less prom- inent. Juvenile oblique suture clearly de- fined on shaft. Type locality.—Semi-permanent pool in roadside ditch on Farm Road 2497, 1.2 mi (1.9 km) SE of intersection with State Route 94, southwest of Lufkin, Angelina County, Texas. The pool of grayish cloudy water, some 3 by 13 and no more than one-half m in depth, was excavated in a sandy clay soil and is situated adjacent to a wooded area in which Pinus and Quercus are the domi- nant plants; grasses and a few sedges are present in the open ditch adjacent to the pool. Both on 13 Apr and 9 Nov 1987, two other crayfishes, Procambarus (Girardiella) curdi and Fallicambarus (Creaserinus) hedgpethi shared the pool with P. (O.) ne- chesae. Disposition of types.—The holotype, al- lotype, and morphotype (4 I, 2, ¢ II) are deposited in the National Museum of Nat- ural History (Smithsonian Institution), numbers 219733, 219735, and 219734, re- spectively, as are the paratypes consisting of 261,96 II, 8 2, 6 j4, and 1032. Size.—The largest specimen available is a first form male having a carapace length 594 of 40.8 mm (postorbital carapace length 31.2 mm); the smallest has corresponding lengths of 30.8 and 24.2 mm. Length of females carrying eggs or young are not available be- cause of lack of such specimens. Range and specimens examined. —This crayfish has been found at the following lo- calities in the Neches River basin of An- gelina and Trinity counties, Texas. Angelina County: (1) Type locality, 1 61, 1 6 I], 1 9, 15 Apr 1987, GBH & HHH; 1 4 II, 9 Nov 1987, BFK & HHH. (2) Burrows in flooded roadside ditch in southeastern Lufkin, | ¢ I, 1 2, 13 Apr 1987, Mike Whiteman & HHH. (3) Drainage ditch and pool at An- gelina County Airport, about 5 mi (8.0 km) S of Lufkin, 1 41, 4 4 II, 4 2, 6 jd, 10 j9, 13 Apr 1987, Harold Brockman, MW, & HHH. (4) Roadside pool at junction of Farm Road and road into airport, about 3.5 mi (5.6 km) S of Lufkin, 13 Apr 1987, MW & HHH; 1 éII, 1210 Nov 1987, BFK & HHH. Trinity County: (5) Caney Creek, 12 mi (19.2 km) NE of Trinity on St Rte 94, 1 6 II, 16 Apr 1987, BFK & HHH. Variations. —Perhaps the most conspic- uous of the variations noted is in the ros- trum, which in some of the juvenile speci- mens is not so strongly contracted anteriorly and bears prominent marginal spines; in most of the specimens there is at least a trace of these spines, but in a few of the larger individuals there is hardly an indentation along the gently contracting margins. In most specimens the antennal scale is broadest at about midlength, but occasionally the great- est width is clearly more proximal, and as for the length, the apices reach to or beyond the tip of the acumen. Cervical spines, like the marginal spines on the rostrum, may be strong in juveniles, but may be reduced to tubercles or even become, at most, rudi- mentary in the adults. The epistome is usu- ally little different from that illustrated for a paratype in Fig. 4h, but in the small male from locality 5 its shape approaches that of an isosceles triangle. The ventrolateral sur- face of the ischium of the third maxilliped may be almost hidden by the dense mat of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON plumose setae borne on it, or the setae may be comparatively smail, sparse, or so in- conspicuous that the podomere appears al- most naked. The telson bears 2 to 4 spines in the caudolateral corner of the cephalic segment. Variations in the secondary sexual characters are almost all so slight that they are not noteworthy, but the proximomesial angle of the first pleopod of the male does show a conspicuous variation which in some 1s produced into a conical prominence (as in right member of Fig. 4d) that is reminis- cent of, if not homologous to, the proxi- momesial spur typical of several species groups, but not of most members of Ort- mannicus. Relationships. —Procambarus (O.) ne- chesae has its closest relationships with Pro- cambarus (O.) geminus Hobbs (1975), P. (O.) lecontei (Hagen, 1870), and P. (O.) tex- anus Hobbs (1971). This is most clearly re- vealed in the short, caudally bent terminal elements of the first pleopod of first form males and in the configuration of the an- nulus ventralis. Procambarus (O.) nechesae resembles P. (O.) geminus more closely than it does the other two, but it may be distin- guished from this crayfish and P. (O.) le- contei by the less strongly reflexed (40 to 50 degrees instead of 80 to 90 degrees) terminal elements of the first pleopod of the male, by the strong tubercle on the basis of the fourth pereiopod which opposes the hook on the corresponding ischium, and usually by the more strongly tapered margins of the rostrum. (Some of the females of P. (O.) nechesae and P. (O.) geminus are virtually impossible to distinguish.) The new crayfish differs from P. (O.) texanus in several strik- ing respects: the areola is usually broader (7.3 to 10.9, avg. 8.8, as opposed to 8.5 to 16.4, avg. 12.8, times longer than broad); the first pleopods of the male are strongly asymmetrically situated with their proxi- momesial extremities overlapping as op- posed to being almost symmetrically dis- posed; there is a strong tubercle on the basis of the fourth pereiopod that opposes the hook on the corresponding ischium in P. VOLUME 103, NUMBER 3 nechesae that is lacking in P. texanus; the sternum in the female of the former is much more weakly tuberculate than that of the latter, and the tongue-like prominence, so conspicuous on the caudomesial part of the annulus ventralis of P. nechesae, is repre- sented at most by a rudiment in P. texanus. These three species constitute a closely al- lied group and may well represent geograph- ic races of a single species. Until their ranges are more clearly determined and evidence of gene exchange between them is found, assigning them specific rank seems prefer- able to me. Ecological notes. —All of the known members of this species, except the single small male collected in Caney Creek (lo- cality 5), were collected from burrows con- sisting of a single subvertical shaft or from temporary or semi-permanent pools in roadside ditches. Caney Creek, flowing over a bed-rock bottom, is a clear stream 3 to 10 m wide and with depths exceeding 1 m. The single specimen, the only crayfish obtained at that locality, was found under a rock ad- jacent to the shore. It seems likely to me that it had wandered or perhaps was washed into this unlikely habitat during a period of high water. Etymology.—The name is derived from the Neches drainage basin, the only wa- tershed in which it has been collected. Crayfish associates. —Taken from the same pool, or dug from burrows adjacent to those occupied by P. (O.) nechesae were members of the following species: Fallicam- barus (F.) devastator, F. (Creaserinus) fod- iens, Procambarus (Scapulicambarus) clar- kii, P. (Girardiella) curdi, and P. (G.) nigrocinctus. Procambarus (Pennides) dupratzi Penn Procambarus dupratzi Penn, 1953:1. Procambarus (Pennides) dupratzi. — Hobbs, 1972:10. New records. —Jasper Co.: Small creek 0.3 mi (0.5 km) E of St Rte 63, 16 mi (25.6 km) N of jct with US Hwy 90, 1 é1, 1 3 Il, 3 jd, 595 4 j2, 17 Apr 1987, GBH & HHH. Polk Co.: Menard Creek 16 mi (25.6 km) W of Tyler Co line on US Hwy 190, 2 4 II, 1 2, 10 36, 9 j2, 12 Nov 1987, BFK & HHH. Tyler Co.: Russell Creek at US Hwy 287, 7 ¢ II, 5 2, 8 j6, 3 j2, 12 Nov 1987, BFK & HHH. Remarks. —The specimens cited here were taken from lotic habitats in which the water was translucent but coffee-colored. Procambarus (Scapulicambarus) clarkii (Girard) Cambarus Clarkii Girard, 1852:91. Procambarus (Scapulicambarus) clarkii.— Hobbs, 1972:12. New records.— Angelina Co.: SE part of Lufkin, 2 6 I, 1 2, 13 Apr 1987, MW & HHH. Trib to Moccasin Creek at Farm Rd 2497, 1 2, 4 j6, 3 j2, 4.9 mi (7.8 km) SE of St Rte 96, 9 Nov 1987, BFK & HHH. Polk Co.: Menard Creek on US Hwy 190, 16 mi (25.6 km) W of Tyler Co line, 1 j2, 13 Nov 1987, BFK & HHH. Creek 5 mi (8 km) W of Livingston on US Hwy 190, 1 jé, 13 Nov 1987, BFK & HHH. Trinity Co.: 4.0 mi (6.4 km) SW of Farm Rd 357 on Rd 2262, | 9, 1 34, 1 32, 16 Nov 1987, BFK & HHH. Remarks. —Except for the adult speci- mens from the first locality listed, which were dug from shallow, flooded, burrows, all were found either in creeks or in a road- side pool. Acknowledgments Thanks are extended to the following per- sons for their assistance in collecting the specimens on which this report is based: Bob Armentrout, Extension Agent for Tyler County, Texas; Hal Brockman of the Soil Conservation Service in Lufkin, Texas; my wife, Georgia B. Hobbs; Brian F. Kensley of the National Museum of Natural History; and Mike Whiteman, Extension Agent for Angelina County, Texas. For their criti- cisms of the manuscript I am indebted to J. F. Fitzpatrick, Jr., University of South Alabama, H. H. Hobbs III, Wittenberg Uni- versity; and Brian F. Kensley. 596 Literature Cited Albaugh, Douglas W. 1973. Life histories of the cray- fishes Procambarus acutus and Procambarus hi- nei in Texas. Unpublished Ph.D. dissertation, Texas A&M University, xiii + 135 pp. 1975. A new crawfish of the genus Procam- barus, subgenus Capillicambarus, from Texas, with notes on the distribution of the subgenus. — Tulane Studies in Zoology and Botany 19(1 & 2):1-7. , & Joe B. Black. 1973. A new crawfish of the genus Cambarellus from Texas, with new Texas distributional records for the Genus (Decapoda, Astacidae). —Southwestern Naturalist 18(2):177- 185. Conner, John V. & Royal D. Suttkus. 1986. Zooge- ography of freshwater fishes of the western Gulf Slope of North America. Chapter 12. Pp. 413- 456 in Charles H. Hocutt and E. O. Wiley, eds., The zoogeography of North American fresh- water fishes. John Wiley and Sons, New York. Cottle, T. J. 1863. Two species of Astacus found in upper Canada.— Canadian Journal of Industry, Science, and Arts (n.s.) 45:216-219. Creaser, Edwin P. 1962. Notes on homologies and genetic relationships in the Cambarinae cray- fishes. 7 pp. [Privately printed]. Faxon, Walter. 1884. Descriptions of new species of Cambarus, to which is added a synonymical list of the known species of Cambarus and Asta- cus. — Proceedings of the American Academy of Arts and Sciences 20:107-158. 1898. Observations on the Astacidae in the United States National Museum and in the Mu- seum of Comparative Zoology, with descrip- tions of new species. — Proceedings of the United States National Museum 20(1136):643-694. Fitzpatrick, J. F., Jr. 1983. A revision of the dwarf crawfishes (Cambaridae, Cambarellinae). — Journal of Crustacean Biology 3(2):266—277. Girard, Charles. 1852. A revision of the North Amer- ican Astaci, with observations on their habits and geograpical distribution.— Proceedings of the Academy of Natural Sciences of Philadel- phia 6:87-91. Hagen, Herman A. 1870. Monograph of the North American Astacidae.—Illustrated Catalogue of the Museum of Comparative Zoology at Har- vard College 3:viii + 109 pp. Hobbs, Horton H., Jr. 1945. Two new species of crayfishes of the genus Cambarellus from the Gulf coastal states, with a key to the species of the genus (Decapoda, Astacidae).— American Midland Naturalist 34(2):466-474. . 1948. A new crayfish of the genus Cambarus from Texas, with notes on the distribution of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cambarus fodiens (Cottle).— Proceedings of the United States National Museum 98(3230):223- 231. 1969. On the distribution and phylogeny of the crayfish genus Cambarus. In Perry C. Holt, Richard L. Hoffman, and C. Willard Hart, Jr., eds., The distributional history of the biota of the Southern Appalachians, Part I: Inverte- brates. — Virginia Polytechnic Institute, Re- search Division Monograph 1:93-178. 1971. New crayfishes of the genus Procam- barus from Alabama and Texas (Decapoda: As- tacidae). — Proceedings of the Biological Society of Washington 84(1 1):81-94. 1972. The subgenera of the crayfish genus Procambarus (Decapoda: Astacidae).—Smith- sonian Contributions to Zoology 117:1-22. 1973. New species and relationships of the members of the genus Fallicambarus. —Pro- ceedings of the Biological Society of Washington 86(40):461-481. 1975. New crayfishes (Decapoda: Cambari- dae) from the southern United States and Mex- ico.—Smithsonian Contributions to Zoology 201:1-34. . 1981. The crayfishes of Georgia. —Smithson- ian Contributions to Zoology 318:viii + 549 pp. 1990. An illustrated checklist of the Amer- ican crayfishes (Decapoda: Astacidae, Cambar- idae, and Parastacidae). —Smithsonian Contri- butions to Zoology 480:iii + 236 pp. — ., & Henry W. Robison. 1982. A new crayfish of the genus Procambarus from southwestern Arkansas.— Proceedings of the Biological Soci- ety of Washington 95(3):545-553. ——., & 1989. On the crayfish genus Fal- licambarus (Decapoda: Cambaridae) in Arkan- sas, with notes on the fodiens complex and de- scriptions of two new species.— Proceedings of the Biological Society of Washington 102(3):65 1— 697. —, & Mike Whiteman. 1987. A new, econom- ically important crayfish (Decapoda: Cambari- dae) from the Neches River Basin, Texas, with a key to the subgenus Fallicambarus. —Pro- ceedings of the Biological Society of Washington 100(2):403-411. —_, & 1990. Notes on the burrows, be- havior, and color of the crayfish Fallicambarus (F.) devastator (Decapoda: Cambaridae).— Southwestern Naturalist (in press). Penn, George H., Jr. 1950. A new crawfish of the genus Orconectes from Louisiana (Decapoda: Astacidae).— Journal of the Washington Acad- emy of Sciences 40(5):166-169. 1953. Two new crawfishes of the genus Pro- cambarus from Texas, Louisiana, and Arkansas VOLUME 103, NUMBER 3 (Decapoda, Astacidae).— American Museum Novitates 1636:1-10. 1962. A new crawfish of the Hinei Section of the genus Procambarus (Decapoda, Astaci- dae).—Crustaceana 3(3):222-226. Penn, George H., Jr., & Horton H. Hobbs, Jr. 1958. A contribution toward a knowledge of the cray- fishes of Texas (Decapoda, Astacidae).— Texas Journal of Science 10(4):452-483. Reimer, Rollin D. 1969. A taxonomic study of the Gracilis Section of the Genus Procambarus. Un- published Ph.D. Dissertation, Tulane Univer- sity, New Orleans, Louisiana, 190 pp. 597 . 1975. Procambarus (Girardiella) curdi, a new crawfish from Arkansas, Oklahoma, and Texas (Decapoda, Astacidae).— Tulane Studies in Zo- ology and Botany 19(1, 2):22—25. —., & William J. Clark. 1974. Decapod crusta- ceans of the Navasota River system in central Texas. —Southwestern Naturalist 19(2):167-178. Department of Invertebrate Zoology Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 598-601 PANOPEUS MARGENTUS, A NEW CRAB FROM THE ARGENTINE WARM TEMPERATE SUBREGION (DECAPODA: XANTHIDAE) Austin B. Williams and E. E. Boschi Abstract. — Panopeus margentus new species, a xanthid crab from warm, temperate, marine Argentine waters is described, illustrated, and compared with the recently described Panopeus mirafloresensis from the Pacific side of the Panama Canal. The first male pleopods and carapace of these species are similar and resemble these features in P. bermudensis. Notes on associated xanthids are given. An unidentified Argentine species of crab belonging to the brachyuran family Xan- thidae (family Panopeidae Ortmann, 1893, sensu Guinot 1978, and subfamily Pano- peinae Ortmann, 1893, sensu Martin & Abele 1986) has been known to the junior author for a number of years. Boschi (1964) described and illustrated brachyurans from Argentine marine waters, including xanthid species belonging to the genera Platyxan- thus, Pilumnoides, and Pilumnus, and he (1979) added Panopeus sp. from the warm temperate region of Buenos Aires and northern Patagonia to this array in an an- notated checklist of decapods from the re- gion. Williams (1984) treated species of the Panopeus “‘herbstii complex,” which has representatives in Brazil and Uruguay, but at that time were not known from Argen- tina. Description of the species listed by Boschi (1979) as Panopeus sp. is the central subject of this paper. Type materials are deposited in the crus- tacean collection of the United States Na- tional Museum of Natural History, Smith- sonian Institution (USNM), Washington, D.C. Panopeus margentus, new species Fig. la-e Material.— Argentina: USNM 239191. Holotype 6 (damaged carapace and right chela), Escollera Norte, Mar del Plata, Prov- ince of Buenos Aires, E. Boschi, 18 Jan. 1964.—239192, Paratypes, 7 6, 15 2 ovig., same.— 239193. Paratypes, 1 6, 1 2 ovig., Escollera Norte, Mar del Plata Puerto, Ris- so, Jan. 1964.—239194, Allotype 2 ovig., Mar del Plata, E. Boschi, 18 Jan. 1964.— 239195, Paratypes, 3 6, 4 2 ovig., same. Specimens in all of these collections are very brittle, and legs are detached from most of them. Description. —Carapace (Fig. la) wider than long, surface smooth, regions faintly indicated, margins finely granular; few lightly granular transverse lines, sinuous epibranchial line more or less evident, less evident arcuate line leading toward but not continued on 4th anterolateral tooth, faint oblique lines on mesogastric lobes still less evident. Faint shallow and sinuously trans- verse depression near posterior '4 passing between meso- and metabranchial, meso- gastric and cardiac regions, another faint depression curving posteromesially from juncture of 3rd and 4th anterolateral teeth, and still another similar groove coursing from between 4th and 5th anterolateral teeth to join transverse groove; slight submargin- al swelling on metabranchial region near coxa of Sth leg. Front with V-shaped me- dian notch, each transverse half shallowly concave; orbital regions broad, moderately VOLUME 103, NUMBER 3 raised and cut by 2 nearly equal fissures; coalesced Ist and 2nd anterolateral teeth separated by well defined but variable notch (progressively developed with increasing age); 1st tooth narrower than 2nd and ex- ceeding it, posterior slope straight to slightly concave; 2nd tooth bluntly triangular, pos- terior margin arcuate; 3rd and 4th teeth broader, each with anterior margin shorter than posterior, tip rounded, rectilinear, or occasionally subacuminate; 5th tooth much smaller, acute or rounded (sometimes on either side of same specimen) and directed laterally; posterodorsal margin slightly and unevenly granular. Abdomen of mature 6 with segments 3 to 5 fused and narrowing distally, segment 6 free; telson narrowly subtriangular. Abdo- men of mature 2 ovate in outline, 6 seg- ments free, 4th broadest, 6th longest in mid- line; telson broadly subtriangular. Chelipeds (Fig. 1a—c) not markedly asym- metrical; microscopically granular except for smooth fingers, carpus unevenly granular and sometimes faintly rugose, with mesial lobelike tooth stout and rounded; hand with outer surface variably marked by broad shallow groove between dorsal crest and swollen palm, lower margin sinuous; curved dactyl of each chela longer and more slender than stout wedge shaped fixed finger, occlu- sive edge of each rather sharp; major dactyl with strong blunt basal tooth and 1 or more less prominent triangular teeth distal to it; triangular tooth on fixed finger, sometimes compound, closing against distal edge of basal tooth on dactyl, other smaller teeth variable; fingers of minor chela each with tooth row somewhat uneven, not strongly developed. Male first pleopod (Fig. 1d—e) obscurely trilobed at tip; accessory process much re- duced, not exceeding membranous collar nearly surrounding it, being perhaps a mod- ification of median process, subterminal lat- eral tooth present; terminal tract of short spinules on lateral rim of collar, subterminal row of about 8 spinules on folded edge of 599 shaft just proximal to collar, grading prox- imad from long to short, and scattered shorter spinules on shaft proximal to these. Measurements in mm.— Holotype 4, car- apace length 7.7, width 10.6; Allotype 9, same, 6.7, 9.5; Paratypes (239192), same, 6 Vals NOD Oth 4s Ue Color.—Color of fingers extending onto distal part of palm, color boundary sharply defined but both fingers and palm faded in specimens available for study. Known range.—Limited to the type lo- cality. Remarks.—The holotype and allotype, although damaged, are the most complete specimens in the material examined. The carapace of the holotype ¢ has been punc- tured in two places, the carpus of the major chela has been broken, and the minor chela has been detached from the body, but no essential features are lost through these damages. The allotype 2 lacks the left fifth leg. The first pleopod of the holotype has the accessory process blunted at the tip, and spines on the membranous collar surround- ing it are swollen, presumably because of poor preservation. Pleopods on ¢ paratypes show both the accessory process and fea- tures of the collar in a better state of pres- ervation. The first pleopods of Panopeus margentus bear some resemblance to those of Pano- peus mirafloresensis Abele & Kim, 1989 from the Pacific side of the Panama Canal (see also Martin & Abele 1986: fig. 2A), as do features of the carapace. Otherwise, the species is comparable in these regards with Panopeus bermudensis Benedict & Rath- bun, 1891 (see Martin & Abele 1986). Associated with the material studied are three other xanthid species. Panopeus me- ridionalis Williams, 1984, is represented by 1 2, cl 9.7, cw 13.2 mm, a specimen with characters less definitely expressed than those of larger individuals of the species known previously only from the Montevi- deo region of Uruguay. Pi/umnoides hassleri A. Milne Edwards, 1880 (see Guinot & 600 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Panopeus margentus new species, holotype é: a, dorsal view showing carapace and chelipeds, broken exoskeleton indicated; b, minor; and c, major chelae in frontal view; first pleopod, d, abdominal; and e, sternal views. Scales: 1 = 1 mm, 2 = 0.1 mm. VOLUME 103, NUMBER 3 Macpherson 1987) is represented by | 6, cl 9.7, cw 13.2 mm. Pilumnus reticulatus Stimpson, 1860 is represented by 1 frag- mentary 6, cl 5.9, cw 7.9 mm, which is seem- ingly closest to the variant forma tessellata discussed by Rathbun (1930) and Boschi (1964). Etymology. —The specific name is a con- struct from the Latin equivalent of Mar del Plata, ““mare,” sea, and “‘argentum,”’ silver, treated as a masculine noun. Acknowledgments The manuscript has been critically re- viewed by F. A. Chace, Jr., R. B. Manning, and M. Vecchione. The illustrations were rendered by Keiko Hiratsuka Moore. Literature Cited Abele, L. G., & W. Kim. 1989. The decapod crus- taceans of the Panama Canal.—Smithsonian Contributions to Zoology 482:1—50. Boschi, E. E. 1964. Los crustaceos decapodos Brach- yura del litoral bonaerense (R. Argentina).— Boletin del Instituto de Biologia Marina 6:1—76. 1979. Geographic distribution of Argenti- nian marine decapod crustaceans. Jn A. B. Wil- liams, ed., Symposium on the composition and evolution of crustaceans in the cold and tem- perate waters of the World Ocean.— Bulletin of the Biological Society of Washington 3:134-143. Guinot, D. 1978. Principes d’une classification évo- lutive des Crustacés Décapodes Brachyoures. — Bulletin Biologique de la France et de la Bel- gique, n.s. 112(3):211-292. , & E. Macpherson. 1987. Révision du genre 601 Pilumnoides Lucas, 1844, avec description de quatre espécies nouvelles et création de Pilum- noidinae subfam. nov. (Crustacea Decapoda Brachyura).— Bulletin du Muséum National d’Histoire Naturella, Paris, s. 4, 9 (sect. A, 1): 211-247. Martin, J. W., & L. G. Abele. 1986. Notes on male pleopod morphology in the brachyuran crab family Panopeidae Ortmann, 1893, sensu Gui- not (1978) (Decapoda). — Crustaceana 50(2):182- 198. Ortmann, A. 1893. Die Decapoden-Krebse des Strassburger Museums, mit besonderer Bertick- sichtigung der von Herrn Dr. Déderlein bei Ja- pan und bei den Liu-Kiu-Inseln gesammelten und zur Zeit im Strassburger Museum aufbe- wahrten Formen. VII Theil. Abtheilung: Brach- yura (Brachyura genuina Boas). II. Unterabtei- lung: Cancroidea, 2 Section: Cancrinea, 1. Gruppe: Cyclometopa. —Zoologische Jahrbiich- er. Abtheilung fiir Systematik, Geographie und Biologie der Thiere 7:41 1-495, pl. 17. Rathbun, M. J. 1930. The cancroid crabs of America of the families Euryalidae, Portunidae, Atele- cyclidae, Cancridae and Xanthidae.— United States National Museum Bulletin 152:i—xvi, 1- 609, 230 pl. Williams, A. B. 1984. The mud crab Panopeus herb- Stii, s.1. Partition into six species (Decapoda: Xanthidae).— Fishery Bulletin, U.S. 81(4, for 1983):863-882. (ABW) National Marine Fisheries Ser- vice-NOAA Systematics Laboratory, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560; (EEB) Instituto Nacional de Investigacion y Desarrollo Pesquero, Casilla de Correo 175, 7600 Mar del Plata, Argentina. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 602-607 A NEW SPECIES OF CHACEON FROM NEW ZEALAND (CRUSTACEA: DECAPODA: GERYONIDAE) Raymond B. Manning, Elliot W. Dawson and W. Richard Webber Abstract. —Chaceon yaldwyni, new species, is described from material col- lected off New Zealand in 1040-1045 and 1228-1276 m. It is the second species from the central Pacific characterized by laterally compressed dactyli on the walking legs. It differs from the other species known from the area, Chaceon bicolor Manning & Holthuis, 1989, by the slenderness of its legs in combination with characters of leg spinulation. Among material of the commercially im- portant crab genus Chaceon taken in New Zealand waters are two specimens, one from northeast of the Chatham Islands and the second from deeper water east of the North Island, that represent an undescribed species, characterized below. The holotype and paratype have been deposited in the Na- tional Museum of New Zealand (NMNZ). Abbreviations used in the account below include cb for carapace width, including lat- eral spines, cl for carapace length on the midline, P5 for fifth pereopod, and FV for Fishering Vessel. Chaceon yaldwyni, new species Figs. 1-3 ““second species.” — Webber et al., 1990:10. Material.—New Zealand: Northeast of Chatham Islands, 43°40.4’S, 174°09.6'W to 43°26.6'S, 174°05.5'W, 1040-1045 m, leg. Henry Kavale, FV Oyang 86, Sta. 33, 14 Apr 1988: 1 6, holotype (NMNZ Cr. 6048).—Off Portland Island, Mahia Pen- insula, 39°51.9'S, 177°55.3'E to 39°48.9'S, 177°55.6'E, 1228-1276 m, leg. Alan Hart, FV Willwatch, Sta. WIL/163/89, 19 Oct 1989: 1 2, paratype (NMNZ Cr. 6468). Diagnosis. —A moderately large Cha- ceon, cl to 105 mm, cb to 128 mm, with well-developed anterolateral teeth on the carapace in adults and with laterally com- pressed dactyli on the walking legs. Cara- pace 1.2—1.3 times broader than long, mod- erately inflated, distinctly convex from front to back, surface appearing smooth, pitted posterolaterally, not tuberculate. Median pair of frontal teeth short, rounded, sepa- rated by U-shaped emargination, medians extending further forward than laterals. An- terolateral teeth well-developed but not spi- niform, second and fourth smaller than re- mainder. Distance from first to second tooth slightly less than distance from third to fourth tooth, distance from first to third less than distance from third to fifth. Suborbital tooth strong, visible in dorsal view, extend- ing about to level of lateral frontal tooth; suborbital margin evenly curved, with some low, rounded tubercles. Cheliped: merus with sharp spine subdistally and with distal dorsal spine; carpus rough dorsally, with distal outer spine, denticulate anterior mar- gin, and with strong, distinct inner spine; propodus roughened on dorsal and outer surfaces, with trace of distal dorsal spine on larger chela. Meri of posterior 3 walking legs with distal dorsal spine. Dactyli of walking legs laterally compressed, height at mid- length greater than width. PS: merus 5.1 1- 5.25 times longer than high and 0.5-0.6 times cb, with distal dorsal spine; carpus with erect spinules dorsally; propodus dis- tinctly longer than dactylus, 4.9 times longer than high. VOLUME 103, NUMBER 3 603 Fig. 1. Chaceon yaldwyni, male holotype, cb 128 mm: a, dorsal view; b, carapace; c, P5. 604 Cia A aaae PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Chaceon yaldwyni, male holotype, cb 128 mm: a, ventral view of orbit; b, merocarpal articulation of P5; c, dactylus of P5, dorsal view; d, dactylus of P5, posterior view. Size. — Male holotype, cl 105 mm, cb 128 mm; female paratype, cl 72 mm, cb 92 mm. Color. —The carapace of both specimens, when fresh, was a fairly uniform yellowish- red, in contrast to C. bicolor, in which, despite its name, the color ranges from a uniform tan to varying shades of purple forming a distinctive pattern on the cara- pace (see color figures of C. bicolor in Sakai (1978:pl. 2, fig. D). Remarks.—Manning & Holthuis (1989: 75) listed five species of Chaceon known from the Indo-West Pacific region, two from the Pacific Ocean, three from the Indian Ocean. Three of these species resemble C. yaldwyni in having laterally compressed dactyli on the walking legs: C. bicolor Man- ning & Holthuis (1989) from the central and southwestern Pacific, C. crosnieri Manning & Holthuis (1989) from Madagascar, and C. paulensis (Chun) (1903) from Amster- dam Island, southern Indian Ocean. Chaceon yaldwyni resembles both C. crosnieri and C. paulensis in having distinct meral spines on the walking legs. It further resembles C. crosnieri in having the cara- pace, especially the protogastric regions, distinctly inflated, so that the carapace 1s VOLUME 103, NUMBER 3 arched in lateral view; the carapace is much more inflated dorsally in C. crosnieri than in C. yaldwyni. Chaceon yaldwyni differs from C. cros- nieri in having more strongly developed an- terolateral spines on the carapace and longer walking legs, with a line of erect spinules on the carpus. This new species differs from C. paulensis in having the carapace much more inflated and in having much shorter anterolateral spines on the carapace. In C. paulensis the second and fourth anterolateral spines are well developed in adults, whereas in C. yaldwyni they are reduced in relation to the other anterolateral spines. In the New Zealand region, C. bicolor is the more commonly found species of Cha- ceon. It is known from east and northeast of the North Island (Webber et al. 1990), whereas C. yaldwyni is only known at pres- ent from two areas, in one of which C. bi- color also occurs, although at shallower depths. Chaceon yaldwyni differs from C. bicolor principally in having quite strikingly slender legs in relation to body size and with strong carapace spines (although such spines have their counterparts in some specimens of C. bicolor, also, particularly in very young specimens tentatively identified with C. bi- color). Measurements of the proportions of the propodus of the walking legs give an index of this slenderness which can serve as a basis of comparison of the two species of Chaceon known from New Zealand waters. A linear regression analysis of the length/ height ratio of the propodus of PS on car- apace length for a total sample of 25 spec- imens of Chaceon available to us gave a low positive correlation coefficient (r) of 0.174. In contrast, omitting the two specimens of C. yaldwyni with the distinctly higher length/ height ratios, the analysis yielded a high positive correlation coefficient of 0.775 (Fig. 3). The analysis includes all adult specimens previously determined as C. bicolor. A comparison of the means of the P5 length/height ratios gave a t value of 3.879. 605 6.0 S15) 5 L/H 3.0 hae tale tae eee 70 80 90 100 110 120 130 140 150 160 170 180 CL Fig. 3. Length/height ratio (L/H) of propodus of fifth leg (PS) in relation to carapace length (CL) in Chaceon bicolor (+) and C. yaldwyni (@); regression line applies to C. bicolor only. For 23 degrees of freedom, the probability of obtaining a value of ¢ greater than 2.807 is 0.01. Hence, the difference between the means is judged to be significant at the 1% level. In other words, the observed differ- ence in propodus slenderness reflects, 99% of the time, a real difference between C. bicolor (n = 23) and C. yaldwyni (n = 2) so far as statistical discrimination based on the small sample sizes allows. Chaceon yaldwyni also appears to differ from C. bicolor in having much more strongly developed anterolateral spines on the carapace, a longer, sharper suborbital tooth, and in having distinct distal dorsal spines on the meri of the walking legs in the adult. Meral spines are present in juveniles from New Caledonia identified with C. bi- color by Manning & Holthuis (1989); these juveniles, however, might well belong to C. yaldwyni or even another species rather than to C. bicolor, as they were not taken together with C. bicolor but in deeper water near areas where C. bicolor was collected. How- 606 ever, C. bicolor shows variability in its PS distal meral spines, which, so far as New Zealand specimens are concerned, can range from mere protuberances to distinct low spines. Similarly, the dorsal margin of the carpus of P5 may vary from being quite smooth to bearing low spinules. These fea- tures certainly are much more pronounced in both of our specimens of C. yaldwyni. It would be useful to be able to distinguish smaller C. bicolor from C. yaldwyni of the same size, but only two of the 23 specimens of C. bicolor available to us have carapace lengths of less than 100 mm. With only two specimens of C. yaldwyni so far known, it is hardly valid to attempt to establish any limits for its population mean in compari- son with the mean of the sample. However, the limits for the population mean of the length/height ratios of the PS propodus, the measure of leg slenderness, derived from a sample of 23 C. bicolor at the 95% confi- dence level (P = 0.05) is 4.206 to 4.467 (mean 4.337). For C. yaldwyni such a limit is 4.293 to 6.07. The specimens of C. bicolor closest in size to the known C. yaldwyni have cl and length/height ratio of the PS propodus of 83 mm and 3.6, 95 mm and 3.7, and 110 mm and 4.0 whereas in C. yaldwyni these values are 72 mm and 5.25 and 105 mm and 5.11. It might be expected, then, that smaller C. bicolor will be distin- guishable by having proportionately stouter legs, in addition to a variable degree of con- spicuousness of the carpal spinules, the dis- tal meral spine, and the compression of the dactylus on the walking legs. So far as taxonomic discrimination is concerned, the two populations of Chaceon found in New Zealand waters can be distin- guished at the species level, even if some subjectivity may be needed in the deter- mination of smaller individuals of C. bi- color in New Zealand using the range of morphological diagnostic criteria proposed by us. The genus Chaceon now comprises 22 species. Manning & Holthuis (1989) listed PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 19 nominal species, and subsequently two other species have been named, Chaceon chilensis Chirino-Galvez & Manning (1989), from Chile, and C. ramosae Manning, Ta- vares, & Albuquerque (1989), from Brazil. The female paratype of C. yaldwyni, cl 72 mm, cb 92 mm, has enlarged, open vulvae with slightly blackened edges and longitu- dinal markings posteriorly consistent with the abrasive action of male pleopods. Size at sexual maturity might possibly be further evidence for the distinctness of C. yaldwyni, although, in the case of the West African C. maritae (Manning & Holthuis) (1981), Mel- ville-Smith (1989) has suggested that ma- ture females may moult more than once and continue to grow extremely slowly. Name.—This species is named for John C. Yaldwyn, recently retired Director of the National Museum of New Zealand. Distribution.—Known only from New Zealand. Acknowledgments We thank John Yaldwyn for his com- ments on the manuscript. Henry Kavale and Alan Hart collected the specimens, and we thank them for their interest. The line draw- ings were done by Lilly K. Manning, who also prepared the figures for publication. Si- mon Dawson prepared Fig. 3. Literature Cited Chirino-Galvez, L. A., & R. B. Manning. 1989. A new deep-sea crab of the genus Chaceon from Chile (Crustacea, Decapoda, Geryonidae).— Proceedings of the Biological Society of Wash- ington 102:401—404. Chun, C. 1903. Aus den Tiefen der Weltmeeres, 2nd edition. Gustav Fischer, 592 pp. Manning, R. B., & L. B. Holthuis. 1981. West African brachyuran crabs.— Smithsonian Contributions to Zoology 306, 379 pp. ——, & 1989. Two new genera and nine new species of geryonid crabs (Crustacea, De- capoda, Geryonidae).— Proceedings of the Bi- ological Society of Washington 102:50-77. ——.,, M. S. Tavares, & E. F. Albuquerque. 1989. Chaceon ramosae, a new deep-water crab from Brazil (Crustacea: Decapoda: Geryonidae).— VOLUME 103, NUMBER 3 Proceedings of the Biological Society of Wash- ington 102:646-650. Melville-Smith, R. 1989. A growth model for the deep-sea red crab (Geryon maritae) off South West Africa/Namibia.—Crustaceana 56(3):279- 292. Sakai, T. 1978. Decapod Crustacea from the Emperor Seamount Chain.— Researches on Crustacea 8 (supplement):1-—39, pls. 1-4. Webber, R., E. Dawson, & B. Stephenson. 1990. The deep-sea red crab—a new resource?.— New Zealand Professional Fisherman 3(6):10—11. Note added in proof: A reference to Cha- ceon yaldwyni (as Chaceon sp.) taken north- east of the Chatham Rise in 962 meters came to our attention while this manuscript was in proof. The citation of this species was in: 1990. Fishes collected by the R/V Shinkai Maru around New Zealand. Japan Marine Fishery Resource Center, Tokyo, 410 pp. 607 The account of the crustaceans in this vol- ume apparently is by M. Takeda, but the pagination of Takeda’s article is unknown to us. (RBM) Department of Invertebrate Zo- ology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560; (EW) Research Associate, Nat- ural History Unit, National Museum of New Zealand, P.O. Box 467, Wellington, New Zealand (formerly New Zealand Oceano- graphic Institute, DSIR, Wellington); (WRW) Crustacea Department, Natural History Unit, National Museum of New Zealand, P.O. Box 467, Wellington, New Zealand. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 608-613 A NEW CRAYFISH (DECAPODA: CAMBARIDAE) FROM SOUTHEASTERN TEXAS Horton H. Hobbs, Jr. and H. H. Hobbs III Abstract. — Procambarus (Ortmannicus) zonangulus is described from Jeffer- son, Hardin, and Orange counties Texas. It may be distinguished from its closest relatives: P. (O.) texanus Hobbs, P. (O.) lecontei (Hagen), and P. (O.) geminus Hobbs by the attenuated distal part of the first pleopod of the male and a strong cephalomesial shoulder situated proximal to the caudodistally disposed ter- minal elements. Some years ago one of us (HHH, Jr) be- came aware of the variation that exists in populations of crayfishes that were being identified as members of Procambarus (Ort- mannicus) acutus acutus (Girard, 1852), and that opinion was stated in the most recent checklist of the American crayfishes (Hobbs 1989). With the accumulation of collections of these crayfishes from Maine to Mexico and northward to Minnesota, the two of us began a study of the cluster of species cur- rently included in the “complex.” The description of this crayfish, while premature in terms of the scope of our proj- ect, is offered to enable David L. Bechler, of Lamar University, Beaumont, Texas, and the late Xuehuai Deng of East China Nor- mal University, Peoples’ Republic of China, to associate the results of their studies with a taxon that will be employed in our revi- sion. A review of the literature involving this crayfish, a summary of its range, and geographic variations are anticipated to be included in our study of the complex. Procambarus (Ortmannicus) zonangulus, new species Fig. 1, Table 1 Diagnosis.—Body pigmented, eyes well developed. Rostrum of adults with or with- out minute marginal tubercles but lacking median carina. Carapace with cervical tu- bercle scarcely larger than others in row on caudoventral flank of cervical groove. Ar- eola 13.1 to 29.0 (mean 19.15 + 3.89; n = 42) as long as wide, constituting 32.0 to 37.1 (mean 33.86 + 1.17; nm = 42) percent of total length of carapace and 41.0 to 46.8 (mean 42.79 + 1.22; n= 42) percent of postorbital length. Suborbital angle very weak and ob- tuse; postorbital ridges with or without in- conspicuous cephalic tubercle; hepatic area tuberculate; branchiostegal spine small and acute. Antennal scale approximately twice as long as broad, widest slightly distal to midlength. Ischia of third and fourth pe- reiopods with simple hooks, hooks of third distinctly overreaching basioischial articu- lation and lacking opposing tubercle on ba- sis, that of fourth almost attaining articu- lation and not opposed by tubercle on basis; coxa of fourth pereiopod with strong cau- domesial boss, that of fifth much smaller and flattened. First pleopods of first form male reaching coxae of third pereiopods, symmetrical, and conspicuously tapering distally; cephalomesial margin with well de- veloped, rounded hump projecting cephalo- mesially. Terminal elements consisting of: (1) tapering acute mesial process directed caudodistally and inclined laterally; (2) ce- phalic process, obscuring central projection in cephalic aspect, corneous, acute, tapering from broad base, inclined mesially, and di- rected caudally, its apex lying mesial to cen- tral projection; (3) caudal element consist- ing of corneous, tapering, acute caudal VOLUME 103, NUMBER 3 process lying almost against caudal surface of central projection and comparatively in- conspicuous, non-corneous, setiferous cau- dal knob at lateral base of cephalic process; and (4) corneous central projection, largest of corneous terminals, inclined mesially and tapering to subacute apex which directed caudodistally and slightly laterally. Annulus ventralis more than twice as broad as long, dextral half elevated little more than sinis- tral; sinus originating on median line, dis- appearing beneath dextral wall and emerg- ing on caudal flank of dextrally oriented tongue near midlength where crossing me- dian line and turning almost caudally before forming arc and extending caudomesially onto conspicuous posteromedian promi- nence and terminating before reaching cau- dal extremity of latter. Sternum immedi- ately anterior to annulus strongly cleft and multituberculate. Unadorned postannular sclerite approximately half width of annu- lus, subtriangular. First pleopods present in female. Holotypic male, form I.—Cephalothorax (Fig. la, m) ovate in section, taller than broad. Abdomen narrower than thorax (17.1 and 20.5 mm). Greatest width of carapace slightly less than height at caudodorsal mar- gin of cervical groove. Areola 20.4 times as long as wide with no more than | punctation in narrowest part. Cephalic section of car- apace 1.9 times as long as areola, length of latter 34.0% of entire length of carapace (42.6% of postorbital carapace length). Sur- face of carapace punctate dorsally, granulate to tuberculate laterally. Rostrum slightly deflected ventrally with converging slender margins, acute apex of short acumen reach- ing midlength of ultimate segment of an- tennular peduncle; minute marginal tuber- cles marking base of acumen; dorsal surface concave with many fine setiferous puncta- tions. Subrostral ridges evident in dorsal as- pect for short distance anterior to caudal margin of orbit. Postorbital ridges well de- veloped, grooved dorsolaterally and bearing small acute tubercle at cephalic extremities. 609 Table 1.—Measurements (mm) of Procambarus (O.) zonangulus. Holotype Allotype | Morphotype Carapace: Entire length 42.0 49.8 39.0 Postorbital length 33.6 40.0 30.2 Width 20.5 24.5 19.5 Height 21.0 24.1 18.3 Areola: Width 0.7 1.0 1.0 Length 14.3 17.6 13.1 Rostrum: Width 7.0 9.2 6.9 Length 10.5 11.6 10.6 Right chela* Length, palm 16.0 12.5 12.1 mesial margin Palm width 12.5 13.5 8.4 Length, lateral 48.7 40.5 35.0 margin Dactyl length 29.2 25.0 20.2 Abdomen: Width 17.1 21.9 15.8 Length 42.1 48.2 40.0 * Left chela in morphotype. Suborbital angle very small and obtuse. Branchiostegal spines small. Cervical spines represented by small acute tubercles scarce- ly larger than neighboring ones on caudal flank of cervical groove. Abdomen (Fig. 1j) subequal in length to carapace. Pleura of third through fifth seg- ments subtruncate to rounded with caudo- ventral extremities subangular on third and fourth. Cephalic section of telson with 3 spines in each caudolateral corner, middle one in both clusters movable; caudal margin of caudal section with shallow median ex- cavation. Cephalic lobe of epistome (Fig. 11) ovate with elevated free margins; central area subplane and sparsely punctate; dis- tinct anteromedian fovea present on main body. Ventral surface of proximal podo- mere of antennular peduncle with spine slightly proximal to midlength. Antenna with small spiniform tubercles on basis and 610 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON QS E ue Fig. 1. _Procambarus (Ortmannicus) zonangulus (all from holotype except c and g from morphotype, e and p from allotype, and o from paratypic first form male): a, Lateral view of carapace; b, c, Mesial view of first pleopod; d, Caudal view of distal part of first pleopod; e, Annulus ventralis and adjacent sternites; f, Cephalic view of distal part of first pleopod; g, h, Lateral view of first pleopod; i, Caudolateral view of distal part of first pleopod; j, Lateral view of abdomen; k, Proximal podomeres of third, fourth and fifth pereiopods; 1, Epistome; m, Dorsal view of carapace; n, Antennal scale; 0, Caudal view of first pleopods; p, q, Dorsal view of distal podomeres of cheliped. VOLUME 103, NUMBER 3 ischium; flagella overreaching caudal mar- gin of telson. Antennal scale (Fig. In) 2.2 times as long as broad, widest slightly distal to midlength; greatest width of lamella al- most twice width of thickened lateral part. Third maxilliped extending cephalically to level of ultimate podomere of antennule; ischium not produced distolaterally, its ventral surface densely studded with plu- mose setae. Right chela (Fig. 1q) subovate in cross section, not strongly depressed. Mesial sur- face of palm with row of 8 tubercles sub- tended dorsally by additional row and ven- trally by only few tubercles. Entire palm moderately tuberculate, but ventrolateral tubercles much reduced and some replaced by punctations. Dorsal and ventral longi- tudinal ridges scarcely evident on either fin- ger. Opposable surface of fixed finger with dorsally situated row of 24 (32 on left) tu- bercles along proximal three-fifths and low- er row of 14 (15 on left), one of which much larger than other tubercles on finger; broad band of minute denticles situated between rows of tubercles and extending to base of corneous tip of finger; except for few tuber- cles proximoventrally, finger otherwise smooth except for longitudinal rows of se- tiferous punctations. Opposable margin of dactyl with upper row of 32 (29 on left) tubercles on proximal three-fifths and lower row of 17 (15 on left); broad band of minute denticles present between rows and extend- ing distally to base of corneous tip of finger; dorsal and ventral surfaces with few tuber- cles in basal portion, otherwise with few longitudinal rows of setiferous punctations; mesial surface of finger with row of 6 (4 on left) tubercles along proximal fourth fol- lowed distally by row of setiferous puncta- tions reaching base of corneous tip. Carpus of cheliped longer than broad with promi- nent oblique furrow dorsally, tuberculate mesially and dorsomesially; mesial surface with 3 (2 on left) tubercles larger than oth- ers, two near midlength and one distome- sially; ventral surface with usual 2 tubercles on distal margin. Merus tuberculate dor- 611 sally, distomesially, and ventrally; 2 pre- marginal tubercles larger than others on dorsodistal surface; ventral surface with mesial row of 16 (15 on left) tubercles and lateral one of 14 (13 on left). Ischium with ventromesial row of 4 tubercles. Hooks on ischia of third and fourth pe- reiopods (Fig. 1k) simple, that on third ov- erreaching basioischial articulation, that on fourth almost reaching articulation and un- opposed by prominent tubercle on corre- sponding basis. Coxa of fourth pereiopod with prominent subvertically oriented cau- domesial boss; that of fifth with smaller one strongly compressed in longitudinal plane of body (when legs positioned at right angles to longitudinal axis). Sternum between third, fourth, and fifth pereiopods comparatively deep with mat of plumose setae extending mesially from ventrolateral margins. First pleopods (Fig. 1b, d, f, h, 1, 0) as described in “Diagnosis.”’ Uropod with both lobes of basal podomere bearing small acute spine; both rami with distolateral spines, and distomedian spine on mesial ramus sit- uated distinctly proximal to subtruncate distal margin. Allotypic female. — Differing from holo- type, except in secondary sexual characters, as follows: acumen of rostrum, marked ba- sally by sudden but shallow contraction of rostral margins reaching base of ultimate podomere of antennular peduncle; tubercles at cephalic extremities of postorbital ridges rounded; suborbital angle vestigial; bran- chiostegal spine tuberculiform; pleura of third through fifth abdominal segments an- gular posteroventrally; cephalic lobe of epi- stome more nearly subtriangular with an- teromedian prominence; mesial surface of palm of chela (Fig. 1p) with row of 6 (7 on left) tubercles; opposable surface of fixed finger with single row of 13 (left with 10 tubercles, latter with additional large one at base of distal fourth; opposable margin of dactyl with single row of 14 (13 on left) tubercles and mesial margin with row of 5; ventral surface of merus of cheliped with mesial row of 14 (13 on left) tubercles and 612 lateral row with 15 (13 on left). Ventrome- sial surface of ischium with 4 (3 on left) tubercles. Annulus ventralis (Fig. le) as described in “Diagnosis.” Morphotypic male, form II.—Differing from holotype in following respects: greatest width of carapace slightly more than height at caudodorsal margin of cervical groove; narrowest part of areola with 2 punctations; acumen reaching base of ultimate segment of antennular peduncle; marginal tubercles at base of acumen much more strongly de- veloped; cephalic section of telson with 2 spines in each caudolateral corner, mesial pair movable; antennal scale with acute dis- tolateral spine and broadest slightly proxi- mal to midlength; longitudinal ridges on fixed finger of chela more distinct, oppos- able margin with dorsal row of 14 tubercles and ventral row of only 3; opposable margin of dactyl with corresponding rows of 22 and 11 tubercles, mesial surface with row of 4 tubercles along proximal fourth; ventral surface of merus of cheliped with mesial row of 17 tubercles and lateral one of 14; is- chium of cheliped with ventromesial row of 5 tubercles; hooks on ischia of third and fourth pereiopods much smaller, neither overreaching articulation with correspond- ing ischium; bosses on coxae of fourth and fifth pereiopods weaker. First pleopod (Fig. lc, g) with apical section much more stocky and while all terminal elements described for holotype present, all reduced and none corneous. Type locality.—Dishman Road at Tram Road north of Meeker, Jefferson County, Texas 30°7'N, 94°15'W). Roadside ditch, occasionally drying, with sedges and grass- es, bordered by Salix niger. Disposition of types. —The holotype, al- lotype, and morphotype (USNM 220297, 220298, and 220299, respectively) are de- posited in the National Museum of Natural History, Smithsonian Institution, as are the paratypes, consisting of 24 6 I, 2 4 Il, 8 2, 2 2s PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Specimens examined. —(1) Type locality: 16 3 I, 7 2, 27 May 1989, coll. Edmund Farmer and David L. Bechler. (2) Massey Lake Slough and unnamed slough, Hardin County, Texas (Lat. 30°7’, Long. 94°12’), 3 61, 1 2, 252, 12 May 1989, coll. DLB, Lynn Sadler, Roy King. (3) The Crawdad Farm, Mauriceville, Orange County, Texas: 5 4 I, 27 May 1989, coll. Boyce and Sharon Ward. (4) Pond adjacent to Neches River at Beau- mont, Orange County, Texas: | 6 I, 3 4 II, 1 2, 26 Sept 1953, coll. R. J. Baldauf. Relationships. —Procambarus (Ortman- nicus) zonangulus has its closest affinities with P. (O.) texanus Hobbs (1971), which has been reported from a single locality in Bastrop County, Texas. More distant affin- ities exist with P. (O.) lecontei (Hagen, 1870) and P. (O.) geminus Hobbs (1975). Among the features in which it differs from the latter two are a much narrower, less densely punc- tate areola, a rostrum with more strongly convergent margins and weaker marginal spines; the first pleopods are symmetrical and, except for the caudal process, the ter- minal elements of the first pleopod of the first form male are directed caudally at ap- proximately right angles to the main shaft of the appendage. This crayfish differs from P. (O.) texanus chiefly in features of the first pleopod, especially in the first form male: the distal part of the pleopod is strongly tapering; the cephalomesial shoulder prox- imal to the terminal elements is much more strongly developed; the central projection and caudal process are proportionally long- er, and more tapering. Etymology. — Zona (L.) = belt + Angulus (L.) = bay or gulf, noting its range in the gulf coastal area of the United States. Acknowledgments Our thanks are extended to David L. Bechler, Lamar University, for providing us with most of the specimens selected to be included in the type series of this crayfish and to Raymond B. Manning, Smithsonian VOLUME 103, NUMBER 3 Institution, for his criticisms of the manu- script. Literature Cited Girard, Charles. 1852. A revision of the North Amer- ican Astaci, with observations on their habits and geographical distribution.— Proceedings of the Academy of Natural Sciences of Philadel- phia 6:87-91. Hagen, Hermann A. 1870. Monograph of the North American Astacidae.—Illustrated Catalogue of the Museum of Comparative Zoology at Har- vard College 3:viiit+ 109 pp. Hobbs, Horton H., Jr. 1971. New crayfishes of the genus Procambarus from Alabama and Texas (Decapoda, Astacidae).— Proceedings of the Bi- ological Society of Washington 84:8 1-94. 613 1975. New crayfishes (Decapoda: Cambari- dae) from the southern United States and Mex- ico.—Smithsonian Contributions to Zoology 201:1-34. 1989. An illustrated checklist of the Amer- ican crayfishes (Decapoda: Astacidae, Cambar- idae and Parastacidae).—Smithsonian Contri- butions to Zoology 480:i11 + 236 pp. (HHH, Jr) Department of Invertebrate Zoology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560; (HHH III) Department of Bi- ology, P.O. Box 720, Wittenberg Univer- sity, Springfield, Ohio 45501. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 614-616 POMATOGEBIA, A NEW GENUS OF THALASSINIDEAN SHRIMPS FROM WESTERN HEMISPHERE TROPICS (CRUSTACEA: UPOGEBIIDAE) Austin B. Williams and Nguyen Ngoc-Ho Abstract.—A new genus, Pomatogebia, is proposed for 3 species of thalas- sinidean shrimps from the tropical western Atlantic and eastern Pacific that are specialized for burrowing in massive stony corals. The caudal part of the shrimp’s abdomen is shaped as an operculum that can be inserted from within the burrow system into its entrance, the exoskeletal surface mimicking that of the coral. Pomatogebia operculata (Schmitt, 1924), placed in Upogebia until now, becomes the type species of the new genus. A tropical western Atlantic thalassini- dean shrimp, identified until now as Upo- gebia operculata Schmitt, 1924, is special- ized for commensal existence in several species of massive stony corals (Kleemann 1984, Scott et al. 1988). The shrimps at an early stage of development commence ex- cavation of burrows in host corals and es- tablish themselves eventually as male-—fe- male pairs that grow into maturity while becoming confined in the burrows. The cau- dal part of the shrimp’s abdomen is shaped as an operculum or plug, with a surface mimicking that of the host coral. In this specialization and in morphology of the car- apace and appendages, this shrimp and two other species in the tropical eastern Pacific differ from related shrimps in the region. Indeed, there are no other upogebiids in the world known to be specialized in this man- ner, and we regard this structural novelty as worthy of generic rank. Pomatogebia, new genus Fig. la—c Diagnosis. —Carapace anierior to cervi- cal groove more or less flattened dorsally and armed on its gastric *4 with field of spines grading from strong anteriorly to weak or obsolescent posteriorly; spines irregularly distributed but tending to arrangement in rows that diverge posteriorly, many with tufts of setae emerging anterior to base. Gas- tric region projected into broadly subtrian- gular rostrum barely exceeding short eye- stalks and bearing pair of subterminal spines, similar spine at each posterolateral corner where rostrum merges with gastric field. Lateral margin of spine field flanked on each side by poorly developed furrow, and that in turn by imperfectly developed lateral ridge bearing crest of about 8-11 spines grading from strong anteriorly to obsolescence pos- teriorly. Incomplete orbital margin concave in dorsal portion; postorbital margin spine- less. Abdomen broadly and smoothly arched dorsally on segments 1—4, segment 4 with dense fringe of setae on posterior margin and transverse band across anterior half; pleura of segment | narrowly rounded pos- terolaterally, those of 2—5 broadly rounded, margins unspined; dense fine setae in tracts on pleura of segments 3—4, tuft on postero- lateral corner of 2 and anterolateral corner of 5; segment 6 irregularly rectangular, broader than long, its lateral margin scal- loped anteriorly and adapted posteriorly for articulation with base of uropod; dorsal sur- VOLUME 103, NUMBER 3 615 Fig. 1. Pomatogebia operculata (Schmitt): a, Cephalic region, lateral; b, Anterior carapace, dorsal; c, Caudal opercular complex, abdominal segments 5-6, telson and uropods; Paratype 2, USNM 57952, Barbados, Lesser Antilles, West Indies. Upogebia affinis (Say): d, Telson, left uropods and posterior margin of abdominal segment 6; 6, USNM 31289, near Bluffton, South Carolina, USA. Scale = 1 mm. face of segments 5 and 6 ornamented with symmetrical pattern of meandering rugae. Tail fan with exposed aspect generally concave. Telson with sides diverging pos- teriorly and posterior margin convex, stiff- ened with radiating longitudinal ribs; exo- pod and endopod of uropods bearing similar radiating ribs. Entire tail fan with dense fringe of setae on distal margin, forming to- gether with segments 5 and 6 an almost cir- cular operculum when fully extended. Maxilliped 1 with an epipod, maxilliped 3 lacking even rudimentary epipod. Chelipeds equal, rather slender, more slender in females than in males; articles spineless; fixed finger nearly as long as dac- tyl, toothed proximally, rather stout and gently curved; dactyl curved, setose, stouter than fixed finger, abruptly tapered to tip and hooking beyond tip of opposed finger. Pe- reopods 2-5 spineless. Type species. —Upogebia operculata Schmitt, 1924. Etymology.—From the Greek “‘poma- tos,’ operculum, for the operculate caudal complex, and the stem “‘gebia,”’ from Upo- gebia, underground digger. The gender is feminine. Remarks. — The operculate abdomen best distinguishes Pomatogebia operculata and 616 its two sister species in the eastern Pacific, P. rugosa (Lockington, 1878) and P. cocosia (Williams, 1986), from members of the ge- nus Upogebia Leach, 1814, distributed worldwide in shallow temperate and trop- ical seas and currently containing 78 rec- ognized species. Pomatogebia has a multi- ribbed telson with convex posterior margin and divergent lateral margins, and the rami of the uropods are also ribbed. Upogebia does not have an operculate abdomen; the telson (Fig. 1d) is basically rectangular, with straight or slightly concave posterior margin and lateral margins straight, somewhat sin- uous, or slightly convergent distally, never divergent, and although reinforced by a var- iously developed transverse proximal ridge confluent with low submarginal lateral ridges, never displays multiple longitudinal ribbing (see Williams 1986). In addition to the two above mentioned genera, three other genera are currently rec- ognized in the family Upogebiidae: Tuer- kayogebia Sakai, 1982 (Japan, | species), Wolffogebia Sakai, 1982 (Malay Peninsula- northwestern Australia, 3 species), and Ge- biacantha Ngoc-Ho, 1989 (Réunion, In- donesia, and New Caledonia, 11 species). Acknowledgments We thank Fenner A. Chace, Jr., Bruce B. Collette, and Raymond B. Manning for crit- ical review of the manuscript, and Keiko Hiratsuka Moore for rendering the draw- ings. Literature Cited Kleemann, K. 1984. Lebensspuren von Upogebia operculata (Crustacea, Decapoda) in karibisch- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON en Steinkorallen (Madreporaria, Anthozoa).— Beitrage zur Palaontologie von Osterreich, In- stitut fir Palaontologie der Universitat Wien, No. 11:325-49. Leach, W. E. [1814.]. Crustaceology. 7n Edinburgh encyclopaedia. Edinburgh, 76:383—437, pl. 221. Lockington, W. N. 1878. Remarks upon the Thalas- sinidea and Astacidea of the Pacific coast of North America, with description of a new species. — Annals and Magazine of Natural His- tory 5(2):299-304. Ngoc-Ho, N. 1989. Sur le genre Gebiacantha gen. nov., avec la description de cing espéces nou- velles (Crustacea, Thalassinidea, Upogebi- idae).—Bulletin du Muséum National d’His- toire Naturelle, Section A, Série 4, 11(1):117— 145. Sakai, K. 1982. Revision of Upogebiidae (Decapoda, Thalassinidea) in the Indo-West Pacific region. Researches on Crustacea, The Carcinological Society of Japan, Special No. 1:1—106. Schmitt, W. L. 1924. Report on the Macrura, Ano- mura and Stomatopoda collected by the Bar- bados-Antigua Expedition from the University of Iowa in 1918.—University of Iowa Studies in Natural History 10(4):65—99, pls. 1-5. Scott, P. J. B., H. M. Reiswig, & B. M. Marcotte. 1988. Ecology, functional morphology, behaviour, and feeding in coral- and sponge-boring species of Upogebia (Crustacea: Decapoda: Thalassini- dea). — Canadian Journal of Zoology 66(2):483- 495. Williams, A. B. 1986. Mud shrimps, Upogebia, from the eastern Pacific (Thalassinoidea: Upogebi- idae).—San Diego Society of Natural History, Memoir 14:1-60. (ABW) National Marine Fisheries Ser- vice-NOAA Systematics Laboratory, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560; (NN-H) Muséum National d’Histoire Na- turelle, Laboratoire de Zoologie (Arthro- podes), 61 Rue de Buffon, 75231 Paris, CEDEX 05, France. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 617-623 GITANA DOMINICA, A NEW SPECIES FROM THE CARIBBEAN SEA (AMPHIPODA: AMPHILOCHIDAE) James Darwin Thomas and J. L. Barnard Abstract. —The new species Gitana dominica is described from Dominica in the Lesser Antilles; it is the first record of the genus from the western Atlantic Ocean. The new species differs from its closest relative, Gitana calitemplado, from California, in the shorter and posteroventrally excavate coxa 1. Gitana is reviewed. Gitana is reviewed and followed by the description of the new species. The first name after each species in the review is the author of the species with date of publica- tion; additional dates or information in pa- rentheses refer to more modern compre- hensive references to the several species; numbers in brackets refer to geographic dis- tribution codes described by Barnard & Bar- nard (1983); those codes represent a uni- versal geographic descriptive system but codes for these species are not included therein and are modernized and presented herein for the first time. Gitana Boeck Gitana Boeck, 1871:132 (Gitana sarsi Boeck, 1871, designated by Sars, 1895: 229, “‘Remarks.’’).—Stebbing, 1906: 155.—Lincoln, 1979:162. Diagnosis. —Mandibular molar large, cushion-shaped, triturative. Lower lip or- dinary. Palp of maxilla 1 uniarticulate. Maxilla 2 ordinary. Outer plate of maxil- liped weakly excavate or not, palp articles 1-2 equal or not strongly different in length. Gnathopods 1-2 small, more or less car- pochelate, poorly subchelate or almost sim- ple, palms very oblique, dactyls lacking large inner nodiform process. Urosomite 3 poor- ly alate. Variables. —Inner plate of maxilla 2 rath- er thin (/ongicarpa) but broader than outer; gnathopods simple (rostrata, longicarpa, etc.), gnathopod 2 scarcely carpochelate (longicarpa, rostrata; etc.). Relationship. —Like Gitanopsis but palp of maxilla 1 uniarticulate. Like Amphilo- chopsis but gnathopod 2 feeble. Species. —See important notes on bio- geography and taxonomy in Chevreux (1911), Chevreux & Fage (1925), Gurja- nova (1951), Krapp-Schickel (1982), Le- doyer (1973), Schellenberg (1942), Stephen- sen (1938). Gitana abyssicola Sars, 1895 (Ledoyer, 1973), eastern Atlantic, warm-temperate to boreal [355]; Gitana bilobata Myers, 1985, Fiji [576]; Gitana calitemplado J. L. Barnard, 1962, 1964, northeastern Pacific warm-temper- ate [370]; Gitana dominica Thomas & Barnard, herein, Caribbean, Leeward Islands [491]; Gitana gracilis Myers, 1985, Fiji [576]; Gitana liliuokalaniae J. L. Barnard, 1970, Hawaiian Islands [381]; Gitana longicarpa Ledoyer, 1977, north- eastern Mediterranean, bathyal [348B]; Gitana rostrata Boeck, 1871 (Sars, 1895) boreal east Atlantic, bathyal [240B]; Gitana sarsi Boeck, 1871 (Sars, 1895) (= Gitana sabrinae Stebbing, 1878) (Lin- coln, 1979) amphi-Atlantic, Mediterra- nean, warm-temperate to arctic ocean [SSSI 618 Biogeography. —Marine, cold and warm northeast Atlantic, warm mid to east Pa- cific, 0-575 m, 9 species. Gitana dominica, new species Figs. 1-3 Etymology. —dominica, noun in apposi- tion, from the type locality. Diagnosis. — Rostrum large (see Fig. 1B) and downturned; lateral cephalic lobes broadly rounded; eyes large and pigmented deep orange in life. Articles 1-2 of antenna 1 of equal length, article 3 shorter, accessory flagellum forming small articulate scale. Labrum almost symmetrically and deeply lobed. Right lacinia mobilis slender and bi- fid, left broad and multitoothed. Outer lobes of lower lip widely spread, unnotched, api- ces each with articulate “salivary spout’; faint inner lobes present. Coxa 1 spout-shaped, with weakly exca- vate posteroventral margin and weakly bifid apically. Carpi of gnathopods 1-2 medium- short, propodi much longer; carpus of gnathopod 1 with short posterodistal lobe, of gnathopod 2 with medium lobe extending along posterior fifth of propodus but not appressed to it; palms obsolescent but bare- ly distinct and armed with sparse spinules. Pereopodal dactyls simple, locking spines paired and unspecialized. Article 2 of pereo- pods 5-7 of diverse sizes, of 5—6 setose but not serrate posteriorly, of pereopod 7 se- tulose and serrate posteriorly. Epimera 1 and 3 with rounded-quadrate posteroventral corner, epimeron 2 with small posteroventral tooth. Outer ramus of uropod | barely shorter than inner ramus, of uropod 2 one third shorter. Telson about Fig. 1. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON two-thirds as long as peduncle of uropod 3, sharply tapering and pointed. Description. —Apical 3-4 articles of fla- gellum on antenna | each with 3—4 aesthet- ascs. Incisors multitoothed. Inner plate of maxilla 1 with 1 apical setule, outer plate with 5 medial serrate spine-setae, apico- medial serrate hump and 8 apical spines; uniarticulate palp with 4 apical spines. In- ner plate of maxilla 2 with 9 medial setae and small stiff setule; outer plate very slen- der, with apical spout and 4 setae. Inner plate of maxilliped with subsidiary apical hump bearing apicolateral spine-tooth, in- cipient apicomedial spine-tooth, and lateral setule inserted below facial pocket, primary truncate margin with 2 apicomedial and one apicolateral spines; outer plate with 3 me- dial spines, no combs on these plates; dactyl with barely visible nail and 2 minute ac- cessory setules. Coxae 2 and 3 with 2 and 3 weak pos- teroventral serrations. Oostegites 2—4 me- dium-broad, oostegite 5 small, all sparsely setose. Gills sac-like, on coxae 2-6, gills 2— 3 smaller than gills 4-6. Epimera 1-2 with lateral facial ridge. Color.—Colors cited are from USA Na- tional Bureau of Standards ISCC-NBS Col- or-Name Charts. In life, eyes very orange (48) to deep orange (51); somatic chro- matophores dark olive brown (96) but when fully expanded yielding overall cast from 40 power observation of dark gray yellow brown (81); during maximum color rendi- tion anterior part of body from head to pos- terior margin of segment 6, including coxae, very dark but splotchy; chromatophores not fully joined nor interspersed, thus rendering densely dotted appearance; dense chro- = Gitana dominica, figures without lower case letter on left of label = holotype, female “‘n,” 2.01 mm; o = female “‘o” 1.98 mm; p = male “‘p” 2.08 mm; q = female ‘“‘q” 2.10 mm. (Legend: Capital letters in figures refer to parts; lower case letters to left of capital letters refer to specimens and to the right refer to adjectives as described: B, body; C, coxa; D, dactyl; G, gnathopod; H, head; I, inner plate or ramus; L, labium; M, mandible; N, molar, or incisor and lacinia mobilis; P, pereopod; R, uropod; S, maxilliped; T, telson; U, upper lip; W, pleon; Y, gill; Z, oostegite; h, half; t, left.) VOLUME 103, NUMBER 3 619 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 620 XM = Q \\ ji yes Wi VOLUME 103, NUMBER 3 621 ( i oC2 Y5 7a Fig. 3. Gitana dominica, figures without lower case letter on left of label = holotype, female “‘n,” 2.01 mm; o = female “‘o” 1.98 mm. —_— Fig. 2. Gitana dominica, figures without lower case letter on left of label = holotype, female “‘n,”’ 2.01 mm; o = female “‘o” 1.98 mm; p = male “‘p” 2.08 mm; q = female “‘q” 2.10 mm; r = female “’r” 2.70 mm. 622 matophores also in articles 2—3 of pereo- pods 5-6, article 1 of antenna | and articles 1-4 of antenna 2; medium density chro- matophores present in urosomites 2-3 and posterior end of urosomite 1, midlateral face of pleonite 3 and peduncles of uropods which, when expanded, provide dark cast; very scattered chromatophores present in some apical articles of pereopods 3-6, pe- reonite 7, pleonites 1—2, article 2 of antenna 1, article 5 of antenna 2, and telson. Faint pigmentary cast not well organized into giant chromatophores present generally through- out gnathopods, article 2 of pereopods 3-4, coxa 3, article 2 of pereopods 5—6. Antero- ventral corner of coxa 4 and pereopod 7 not pigmented. Drawing of holotype body showing principal chromatophores and dif- fuse casts based on field notes and male “‘p,”’ the pigment of which was best preserved. Pigment of chromatophores concentrated on death and to some extent broken and contents released to form pigmentary cast. Microhabitat unusual for amphilochids which are generally thought to be slime lap- pers on sedentary organisms, especially coelenterates. Holotype.—USNM 242015, female ‘“‘n” 2.01 mm (Figs. 1-3). Type locality. -—JDT-DOM 2A, Pte. Guignard, Dominica, 2 km south of Ro- seau, sediments amongst reef outcrop, 19 May 1987; coarse quartzose sand with dark mineral inclusions, 6.3 meters, sediment appearing greyish-black; large amount of unicellular algal strands; J. D. Thomas and J. Clark, collectors. Material.—All paratypes from type lo- cality, female “‘o”’ 2.48 mm; female “‘p” 2.58 mm; female “‘q’’ 2.60 mm; female “‘r’’ 2.70 mm; male “s” 1.80 mm. Six other speci- mens from the type locality provisionally labeled as this species are deposited in USNM collections but are not discretely identified and are left undamaged for future examiners. Relationship. —Our species appears most similar to Gitana calitemplado. These two PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON species differ from other species of Gitana in the short carpi and relatively long pro- podi of the gnathopods. This may have ge- neric significance and is a condition so far restricted to species in the western hemi- sphere. Our species differs from Gitana cali- templado in the short, posteroventrally ex- cavate coxa |. Differing from Gitana sarsi in the shorter carpi of the gnathopods, spout-like shape of coxa 1, and lack of inner notches on the lower lip. Differing from Gitana longicarpa in the downturned rostrum, short carpi of the gnathopods, poorly lobed article 2 of per- eopods 5-7, shorter peduncle of antenna 2, unpointed coxa 2, short article 1 of man- dibular palp, less extended palp of the max- illiped and shorter broader telson. Differing from Gitana abyssicola in the short carpi of the gnathopods, spout-like coxa 1, slightly larger rostrum and greater difference in relative sizes of article 2 on pereopods 5-7. Differing from Gitana rostrata in the downturned rostrum, short carpi of the gnathopods, thicker peduncle of antenna 2, relatively greater differences in article 2 of pereopods 5-7. Differing from Gitana liliuokalaniae in the relatively long propodi and short carpi of the gnathopods, distinct tooth of epi- meron 2, bifid right lacinia mobilis, more elongate peduncle of antenna 1, and spout- like coxa 1. Differing from Gitana bilobata and Gi- tana gracilis in the short carpi of the gnatho- pods, the much longer antenna 1, and much longer (but normal) coxa 2. Distribution. —Dominica, 6 m. Acknowledgments Fieldwork for this report was supported by Smithsonian’s CCRE committee; this is CCRE Contribution No. 260; we thank Dr. Klaus Ruetzler, head of this program, and Mike Carpenter for their assistance. The VOLUME 103, NUMBER 3 second author was supported by NSF Grant 8515186. Linda Lutz of Vicksburg, Missis- sippi, inked our drawings. Literature Cited Barnard, J. L. 1962. Benthic marine Amphipoda of southern California: families Amphilochidae, Leucothoidae, Stenothoidae, Argissidae, Hyali- dae.— Pacific Naturalist 3:116-163. 1964. Los anfipodos bentonicos marinos de la costa occidental de Baja California. — Revista de la Sociedad Mexicana de Historia Natural 24:205-274, 11 figs., 5 tabs. . 1970. Sublittoral Gammaridea (Amphipoda) of the Hawaiian Islands. —Smithsonian Contri- butions to Zoology 34:1-286. , & C. M. Barnard. 1983. The freshwater Am- phipoda of the world.—Mt. Vernon, Virginia, Hayfield Associates, 2 volumes, xix and 849 pp. Boeck, A. 1871. Crustacea Amphipoda Borealia et Arctica.—Forhandlinger i Videnskabs-Selska- bet i Christiania 1870:83-280. Chevreux, E. 1911. Campagnes de la Melita. Les am- phipodes d’Algerie et de Tunisie. — Memoire de la Societe Zoologique de France 23:145-285, plates 6-20. —., & L. Fage. 1925. Amphipodes.—Faune de France 9:1-488, 438 figs. Gurjanova, E. 1951. Bokoplavy morej SSSR 1 so- predel’nykh vod (Amphipoda-Gammaridea). — Akademiia Nauk SSSR, Opredeliteli po Faune SSSR 41:1-1029, 705 figs. Krapp-Schickel, Gitana. 1982. Family Amphilochi- dae. In S. Ruffo, ed., The Amphipoda of the Mediterranean, Part 1, Gammaridea (Acan- thonotozomatidae to Gammaridae).— Mem- oires de I’Institut Oceanographique 13:70-93, figs. 48-63. Ledoyer, M. 1973. Amphipodes gammariens nou- 623 veaux Ou peu connus de la region de Marseille. — Tethys 4:881-898, 13 pls. . 1977. Contribution a l’etude de l’ecologie de la faune vagile profonde de la Mediterranee nord occidentale 1. Les gammariens (Crustacea, Am- phipoda).— Bollettino del Museo Civico di Sto- ria Naturale, Verona 4:321-—421. Lincoln, R. 1979. British marine Amphipoda: Gam- maridea. London, British Museum (Natural History), vi + 658 pp., 280 figs., 3 pls. Myers, A. S. 1985. Shallow-water, coral reef and mangrove Amphipoda (Gammaridea) of Fiji. — Records of the Australian Museum supplement 5:1-144, 109 figs. Sars, G. O. 1895. An account of the Crustacea of Norway with short descriptions and figures of all the species. Christiania: Alb. Cammer-Mey- ers 1:1-711. Schellenberg, A. 1942. Krebstiere oder Crustacea IV: Flohkrebse oder Amphipoda.—Die Tierwelt Deutschlands, Jena 40:1—252, 204 figs. Stebbing, T. R. R. 1878. On two new species of am- phipodous crustaceans.—-Annals and Maga- zine of Natural History, series 5, 2:364—370, pl. 15. 1906. Amphipoda I. Gammaridea.—Das Tierreich 21:1-806, 127 figs. Stephensen, K. 1938. The Amphipoda of N. Norway and Spitsbergen with adjacent waters. — Tromso Museums Skrifter 3:141-278, figs. 20-31. USA National Bureau of Standards. ISCC-NBS Cen- troid Color Charts Standard Sample No. 2106. Supplement to NBS Circular 553. (Version with 257 colors.) (JDT) Reef Foundation, P.O. Box 569, Big Pine Key, Florida 33043; (JLB) NHB- 163, Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 624-632 THE JAPANESE AMPHIPOD GENUS EONIPHARGUS, REDISCOVERED IN A SOUTH KOREAN CAVE Jan H. Stock and Young Won Jo Abstract. —Eoniphargus, a monotypic genus, known only from the Tokyo area, Japan, has been rediscovered in cave waters of South Korea. The Korean animals are a new species, E. glandulatus, which bridges to a certain extent the gap between Eoniphargus and the Indian genus Indoniphargus. Their mor- phology indicates that these genera are more closely related to the Gammaridae than to the Crangonyctidae. Among a number of stygobiont amphi- pods collected by the junior author in cave waters of South Korea, a representative of a new species was discovered belonging to a small group of two monotypic genera, Eoniphargus Ueno, 1955 (Eoniphargus was originally described as a subgenus of Neoni- phargus, but was elevated to generic rank by Straskraba 1964), and IJndoniphargus Straskraba, 1967. The former is known from a single locality near Tokyo (Japan), where- as the latter is known from the states of West Bengal and Orissa (India). The Korean tax- on bridges to a certain extent the gap be- tween Eoniphargus and Indoniphargus: of the ten discriminating characters, three agree better with Indoniphargus, whereas seven correspond with Eoniphargus (Table 1). The resemblance to Eoniphargus bears amongst others on the shape of the gnathopods 1 and 2, and the armature of the inner and outer lobes of maxilla 1, both considered of prime importance in amphipod taxonomy. The resemblance to Indoniphargus bears on “‘weak’’ characters (urosome spination, number of segments in the accessory fla- gellum, absence of calceoli), known to be variable, sometimes even at infraspecific level. Therefore, the Korean form has been attributed to Eoniphargus rather than to Jn- doniphargus. Eoniphargus and Indoniphargus have often been thought related to the crango- nyctids or crangonyctoids (e.g., Bousfield 1977:table VI, Bousfield 1982:262, Stras- kraba 1964:table I, StraSkraba 1967). Bar- nard & Barnard (1983) classify Eoniphargus near the gammarids, but Indoniphargus near the melitids. Clearly enough, and contrary to what the generic names suggest, these genera are not closely related to the Ni- phargidae. In our opinion, several points in their morphology prohibit inclusion in the crangonyctids s.l. (= superfamily Crango- nyctoidea Bousfield), e.g., the asymmetrical palps of left and right maxillae, the occur- rence of spines on the urosome, the ‘eulim- nogammarid’ shape of the gnathopods of Eoniphargus, the absence of sternal gills, the absence of a coxal gill on pereiopod 7, the absence of a double row of distally-notched spine teeth on the palm of the two gnatho- pods, the rather elongate third uropod, the occasional presence of calceoli described as shoe-shaped by Uéno (1955:fig. 1) on an- tenna 2 of the male. None of these char- acters alone is sufficient proof of a non-cran- gonyctid relationship, but in combination they probably show that Eoniphargus forms a subgroup of the gammarids (or gamma- roids, if one believes in superfamilies in this group). Eoniphargus glandulatus, new species Material. —1 2 (holotype), 1 6 (allotype), 4 paratypes. South Korea, Ondal-gul (gul = cave), Prov. Choongbuk, Danyang-gun, VOLUME 103, NUMBER 3 625 Table 1.—Comparison of some salient characters of the new Korean taxon (K), Indoniphargus indicus (1), and Eoniphargus kojimai (E). K spines + setules 1-segmented absent long broad, with 11 setae Armature urosomites Al, acc. flag. A2 4, calceoli Distal segm. Md. palp Mx.1, inner lobe Mx.1, outer lobe 10 spines Mx.2, inner lobe, oblique present row of setae Gn.1 & 2, carpus non-lobate Gn.1 & 2, propodus elongate Peduncle uropod 1, absent ventral spine(s) Youngchoon-myeon, Ha-ri (128°30’E, 37°04'N); pools of 10-20 cm deep in lime- stone cave, 50-120 m from entrance (dim light to complete darkness); water temper- atunes|2:5-C491 Oct 1986 leg Yn W. Jo and H. J. Lee, ZMA Amph. 108.633. Description. —Body length of 2 up to 6 mm, ¢ 4.5 mm. Blind, colourless in alcohol. Body shape as in Fig. la. Lateral lobe of head rounded (Fig. 1b); antennal sinus very shallow. Coxal plates 1 to 4 deep, 5 to 7 shallow. Dorsum of metasome with a few setules. Urosomite 1 with variable arma- ture: 1 lateral spine + 1 dorsal setule or 1+1 setules; urosomite 2 with 2 lateral spines, sometimes 1 additional lateral setule, and 1 dorsal setule; urosomite 3 unarmed (Fig. Lh). Antenna | (Fig. 1c): Peduncle segment 1 longest; segment 2 slightly longer than 3. Flagellum with up to 25 segments; short aesthetascs on segments 5 to 22. Accessory flagellum slightly longer than first flagellum segment, 1-segmented. Antenna 2 (Fig. 1d) with very long, ta- pering gland cone. Peduncle segments 4 and 5 thin and slender. Flagellum 7-segmented. Calceoli absent (6, @). Upper lip (Fig. 3c) rounded. Lower lip (Fig. 3d) without inner lobes. I E spines + setules setules 2-segmented 4-segmented absent present short long finger-shaped, with broad, with 8 setae 2 setae 8 spines 10 spines absent present lobate non-lobate mittenform elongate present absent Mandible (Fig. 2a): Molar setae present both on right and left appendage; left la- cinia mobilis 5-dentate; right lacinia bifid; 3 spines + 3 plumose setae between molar and incisor of left mandible, 2+ 2 right. Palp (Fig. 2b) strong; segment 1 unarmed; seg- ment 2 with row of about 10 ventrodistal setae; segment 3 with regular row of c. 15 D-setae, 1 B-setule, 4 E-setae, but without A- or C-setae. Maxilla 1 (Fig. 2c, d): Palps asymmetrical (left more slender, with 5 slender spines and 1 seta; right more robust, with 6 robust spines and | seta). Outer lobe with 10 distal spines, each spine with 4 to 10 medial den- ticles. Inner lobe rounded-triangular, with 11 plumose setae on medial margin. Maxilla 2 (Fig. 2e) with oblique row of 14 strong setae on inner lobe. Maxilliped (Fig. 2f): Inner lobe with 3 short, robust distal spines. Outer lobe with 5 mediodistal spines; medial margin setose. Gnathopods | and 2 with very slight sex- ual dimorphism (propodus in ¢ slightly larg- er in size than in 2; palmar margin in 6 with more setules than in 9; largest palmar angle spine of Gn.2 longer in ¢ than in 9). Gnathopod 1 (Fig. 3a): Basis with 3 an- terior and 5 posterior setae. Carpus un- lobed. Propodus of similar shape in both 626 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON aces ee SS a a a Fig. 1. Eoniphargus glandulatus: a, female, from the right, pleopods omitted (scale 1); b, head, 2 (2); c, antenna 1, 2 (2); d, antenna 2, ¢ (2); e, telson, left half, 2 (3); f, telson, 6 (3); g, epimeral plates, 6 (2); h, contour of urosome, ¢ (4). sexes, almost rectangular; 4 palmar angle spines (2 short, 1 medium, | very long); palmar margin with setules only (c. 6 in 8, cf. Fig. 4a; c. 15 in 6, Fig. 4b); claw slender; dactylus with 2 inner, | outer, and 3 distal setae; unguis shorter than dactylus. Gnathopod 2 (Fig. 3b) slightly longer than Gn.1, but propodus “‘feebler’’ in appearance than that of Gn.1. Basis with 2 anterior and 7-9 posterior setae. Carpus more elongate than that of Gn.1. Propodus very narrow and slender in both sexes. Palmar angle with VOLUME 103, NUMBER 3 627 Fig. 2. Eoniphargus glandulatus °: a, left mandible; b, right mandibular palp; c, right maxilla 1; d, palp of left maxilla 1; e, maxilla 2; f, maxilliped. All scale 5. 4 spines (1 very long); palmar margin with setules only (c. 10 in 2, Fig. 4c; c. 14 in 4, Fig. 4d). Armature of claw as in Gn.1. Pereiopod 3 (Fig. 5a) with long and nar- row coxal plate. Distal segment poorly se- tose/spinose. Claw thin and slender. Coxal plate 4 (Fig. 5b) with shallow posterior emargination; remaining segments of P4 as in P3. Pereiopods 5 and 6 (Fig. 5c, e) broken in all specimens examined (reconstructed from the fragments in Fig. la). P5 shorter than P6, P7 longer than P6. Basis of PS to P7 with strong posteroventral lobe. Dactylus 628 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 6 2.2mm ( ae Fig. 3. Eoniphargus glandulatus: a, gnathopod 1, é (scale 6); b, gnathopod 2, 4 (6); c, upper lip, 2 (5); d, lower lip, 2 (6). thin and slender, unguis small (Figs. 5c, 6d). No sternal gills. Oostegites linear, non-se- Basis practically without sexual dimor- _ tiferous (diapause stage), but with scars in- phism (Fig. 5d). dicating insertion of c. 8 setae on margins Coxal gills large, ovate, with long basal _ of distal part (Fig. 5a); present on Gn.2 and stalk, present on Gn.2 and P3 through P6. P3 through PS. VOLUME 103, NUMBER 3 629 Fig.4. Eoniphargus glandulatus: a, palma of gnathopod 1, ?; b, palma of gnathopod 1, 4; c, palma of gnathopod 2, 2; d, palma of gnathopod 2, 6. All scale 7. 630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Eoniphargus glandulatus: a, pereiopod 3, ¢ (scale 8); b, coxal plate of pereiopod 4, 2 (8); c, pereilopod 5, 2 (9); d, basal segments of pereiopod 5, 6 (9); e, basal segments of pereiopod 6, 2 (9). Epimeral plates with angular posteroven- tral corners (Fig. 1g), unarmed ventral mar- gins and | to 3 setules on posterior margin. Pleopod peduncle with 2 retinacula (Fig. 6c), shorter than rami. Pleopod 1: both rami 11-segmented. Pleopods 2 and 3 with 11- segmented exopodite and 10-segmented en- dopodite. All setae of pleopod rami feath- ered, no clothes-peg spines. Uropod 1 (Fig. 6a): Peduncle without proximoventral spine; | distomedial and 2 distolateral spines; 2 rows of 3 to 6 dorsal VOLUME 103, NUMBER 3 631 Fig. 6. Eoniphargus glandulatus 2: a, uropod 1 (scale 8); b, uropod 2 (8); c, pleopod 2 (8); d, pereiopod 7 (9); e, uropod 3 (6). spines. Exopodite slightly shorter than en- dopodite, both rami with dorsal, lateral and distal spines. Uropod 2 (Fig. 6b) much shorter than U1, with proximal peduncular spine, but oth- erwise rather similar. Uropod 3 (Fig. 6e) not sexually dimor- phous, reaching far beyond tip of uropod 1 (Fig. la). Exopodite 1-segmented, terminal margin truncate with 4 spines; 4 groups of lateral spines and 3 groups of medial spines; no setae. Endopodite scale-like, small, with 1 distal setule and a minute laterodistal notch. Telson (Fig. le, f) deeply incised (over about 70% of its length); cleft V-shaped, rather wide. Armature variable: a lateral spine may be present or absent; distal ar- mature of each telson lobe consisting of 2 or 3 spines and | setule; 2 long, plumose (“sensorial”) setae distolaterally on each lobe. 632 Discussion.—The Korean taxon differs from Indoniphargus indicus (Chilton 1923) in the characters listed in Table 1 (see also Straskraba 1967 and Stephensen 1931). From the only species known of Eoniphar- gus, E. kojimai Uéno, 1955, it differs in (1) the monomerous accessory flagellum (4- segmented in E. kojimai); (2) the absence of calceoli on the male second antenna; (3) the shape of the exopodite of uropod 3 (with acuminate, unarmed distal end in E. koji- mati; with truncate, spinose distal end in the new species); (4) the very strongly elongated antennal gland cone; (5) a slightly less elon- gate carpus of gnathopod 2; (6) the presence of spines on urosomites 1 and 2 (Setules only in E.. kojimai); (7) shorter aesthetascs on the flagellum of antenna 1. Character states (1) and (2) are frustrating since they are often considered of some taxonomic value. However, the presence or absence of calceoli is a variable character in the genus Gammarus (e.g., G. insensibilis is permanently devoid of calceoli, cf. Stock 1967, whereas in G. fossarum certain pop- ulations lack calceoli, cf. Goedmakers 1972). Apparently, the new species is apomor- phous in character states (1) and (2), in com- parison with Eoniphargus kojimai. Etymology. —The specific name is based on glandula (Latin = gland), alluding to the large size of the antennal gland. Literature Cited Barnard, J. L., & C. M. Barnard. 1983. Freshwater Amphipoda of the world. Hayfield Associates, Mt. Vernon, Virginia, 1, 2:830 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bousfield, E. L. 1977. A new look at the systematics of gammaroidean amphipods of the world.— Crustaceana, Suppl. 4:282-316. . 1982. Amphipoda. Pp. 254—293 in S. P. Par- ker, ed., Synopsis and classification of living or- ganisms. McGraw-Hill, New York. Chilton, C. 1923. A blind amphipod from a mine in Bengal. — Records of the Indian Museum 25(2): 195-196. Goedmakers, A. 1972. Gammarus fossarum Koch, 1835: redescription based on neotype material and notes on its local variation (Crustacea, Am- phipoda).— Bijdragen tot de Dierkunde 42:124— 138. Stephensen, K. 1931. Neoniphargus indicus (Chil- ton), an Indian fresh-water amphipod.—Rec- ords of the Indian Museum 33:13-19. Stock, J. H. 1967. A revision of the European species of the Gammarus locusta-group (Crustacea, Amphipoda).—Zo6logische Verhandelingen Leiden 90:3-56. Straskraba, M. 1964. Perthia n. g. (Amphipoda, Gammaridae) from fresh water of western Aus- tralia, with remarks on the genera Neoniphargus and Uroctena.—Crustaceana 7(3):125-139. . 1967. Re-examination of the taxonomic sta- tus of Neoniphargus indicus Chilton (Amphipo- da, Gammaridae) and its zoogeographical re- lations.— Proceedings of the Symposium on Crustacea, Ernakulam 1:126-132. Uéno, M. 1955. Occurrence of a freshwater gam- marid (Amphipoda) of the Niphargus group in Japan. — Bulletin of the Biogeographical Society of Japan 16:146-152. Institute of Taxonomic Zoology, Univer- sity of Amsterdam, P.O. Box 4766, 1009 AT Amsterdam, The Netherlands. Markham (1985) proposed the genus PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 633-642 TAXONOMIC REMARKS ON SCHIZOBOPYRINA MARKHAM, 1985, WITH THE DESCRIPTION OF S. BRUSCAI (CRUSTACEA: ISOPODA: BOPYRIDAE) Ernesto Campos and Alma Rosa de Campos Abstract.—Schizobopyrina striata (Nierstrasz & Brender a Brandis, 1929) that infests Thor algicola Wicksten, 1987 is reported from the Golfo de Cali- fornia, México. The morphology of the female examined agrees well with that of the lectotype female from San Diego Bay, California. Schizobopyrina bruscai is described from a single adult female collected in Bahia Concepcion, Baja California Sur, México. This species is a parasite of a new species of Thor. It differs from other species of Schizobopyrina principally in the pleomeres and pleotelson morphology. In addition, we suggest Bopyrina platylobata Bourdon, 1983, from Seram, Seleman Bay and Queensland, should be included within Schizobopyrina, and S. lobata Bourdon & Bruce, 1983, from Queensland, should be redescribed in order to determine whether this species is correctly placed within this genus. Resumen. —El isopodo bopirido Schizobopyrina striata (Nierstrasz & Brender a Brandis, 1929) se registra como parasito del camaron carideo Thor algicola Wicksten, 1987 en el Golfo de California. La morfologia de la hembra corres- ponde a la del lectotipo colectado en San Diego Bay, California, EEUU. Schizo- bopyrina bruscai se describe en base a un espécimen hembra adulto que se colecto en Bahia Concepcion, Baja California Sur, México. Esta especie es parasita de una especie indescrita de Thor. Difiere de otras especies del género Schizobopyrina principalmente en morfologia de los pleomeros y pleotelson. Adicionalmente se sugiere que Bopyrina platylobata Bourdon, 1983, de Seram, Bahia de Seleman y Queensland, debe ser asignada al género Schizobopyrina, y que S. Jobata Bourdon & Bruce, 1983, de Queensland, necesita redescribirse a fin de establecer si su inclusion a este género es correcta. (Richardson, 1904), was collected from Schizobopyrina to include 10 species that had been assigned formerly to the closely allied genus Bopyrina Kossman, 1881. Schizobopyrina has a maxilliped palp, elon- gate second through fifth oostegites, and at least lateral separation of the six pleomeres. These features contrast with the lack of maxilliped palp, tiny oostegites 2—5 and fu- sion of pleomeres on the short side in Bo- pyrina. Two species of Schizobopyrina have been recorded from American waters only. The type species, Schizobopyrina urocaridis North Carolina, western Florida, and Be- lize, while S. striata (Nierstrasz & Brender a Brandis, 1929) has been recorded only from San Diego Bay, California (Nierstrasz & Brender a Brandis 1929, Markham 1985). In the present note we extend the known distribution, report a new host and give ad- ditional morphological information for S. striata, a species that apparently has not been collected since its description. Con- comitantly, a new northeastern Pacific species of Schizobopyrina is described. 634 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON These species records increase from 12 to 14 the number of bopyrid species reported from the Mexican Pacific (Campos & Cam- pos 1989). In addition, some comments on taxonomy of the Australian species S. /ob- ata Bourdon & Bruce, 1983 and Bopyrina platylobata Bourdon, 1983 are included. Specimens of S. striata and Schizobopyrina bruscai described herein have been depos- ited in the Collection of Invertebrates, Es- cuela Superior de Ciencias, Universidad Autonoma de Baja California. The terms posteroventral border of head and spur de- scribed by Markham (1985) are respectively herein referred as barbula and plectron in agreement with Markham (1988). Infor- mation recorded in Table | was obtained from the below-noted descriptions and pa- pers of Chopra (1923); Nierstrasz & Brender a Brandis (1923); Shiino (1933, 1939a, 1939b, 1942); Bourdon (1983); Bourdon & Bruce (1983); and Markham (1985). Schizobopyrina striata (Nierstrasz & Brender a Brandis, 1929) Figs. 1, 2 Bopyrina striata Nierstrasz & Brender a Brandis, 1929:40—42, figs. 51-53. Schizobopyrina striata.—Markham, 1985: 46. Distribution. —San Diego Bay, California (type locality); upper Golfo de California (new record, herein). Hosts recorded. —Caridea-Hippolytidae: Hippolyte californiensis Holmes, 1895. Material examined.—One female, Puer- tecitos, Golfo de California, Baja California, km 72 road San Felipe-San Luis Gonzaga 30°30’N, 114°40'W, 23 May 1986, lower midlittoral, E. Campos coll. New host.—Rubén Rios (pers. comm.) noted that the host of S. striata agrees well with description of 7. algicola Wicksten, 1987; however, he suggested that Wick- sten’s species should be considered as a ju- nior synonym of 7. manningi Chace, 1972. A discussion of this topic will be published by R. Rios (in prep.). Description. — Female (Fig. 1A, B). Length 4.0 mm, maximum width 2.0 mm, head length 0.8 mm, pleon length 0.9 mm, body axis distortion 126°. Head roughly triangular, separated from pereon by a deep groove. Large and indis- tinctly set-off frontal lamina extending into sub-triangular points and notched on short- er side of body; frontal lamina with medial extensions. Antennae tiny, obscure. Max- illiped (Fig. 2A) with setose palp well de- veloped, articulating with margin (Fig. 2B). Barbula medially produced, with two blunt projections on each side (Fig. 2C). Seven well defined pereomeres dorsally and lat- erally, inconspicuous ventrally. Coxal plates on pereomeres 1-4 of both sides; dorsolat- eral bosses on pereomeres 1-4 of long side. Pereopods subequal, increasing slightly in length posteriorly, seventh pereopod with bilobed and pseudoarticulated merus (Fig. 2D). First pair of oostegites asymmetrical (Fig. 2E); right one with falcate posterolater- al point, left one with larger and sickle- shaped posterolateral point; internal ridges of both first oostegites unadorned. Pleon with 5 medially fused pleomeres and pleotelson, latter deeply arcuate and embedded in the fifth pleomere. Four pairs of lanceolate uniramous pleopods; ventral surface of lateral plates of long side bearing tuberculiform projections decreasing pos- teriorly, conical on first four pleomeres, rounded on fifth one. On short side such projections are not evident. No uropods. Remarks. —Schizobopyrina striata is the only species of this genus recorded from the East Pacific (Markham 1985). It was de- scribed by Nierstrasz & Brender a Brandis (1929) from specimens collected in San Di- ego Bay, California. There are no additional published records to our knowledge. Mor- phologically our female specimen (Fig. 1A, B) is almost identical to that described and figured in 1929 by Nierstrasz & Brender a VOLUME 103, NUMBER 3 635 Ue Fig. 1. Schizobopyrina striata (Nierstrasz & Brender 4 Brandis, 1929): A, C, Female ventral view; B, E, Female dorsal view; D, Male. (C—E redrawn from Nierstrasz & Brender a Brandis 1929). 636 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON See | Fig. 2. Schizobopyrina striata (Nierstrasz & Brender a Brandis, 1929), Female: A, Maxilliped; B, Maxilliped palp; C, Barbula; D, Pereopod 7; E, Left oostegite 1. Brandis (fig. 1C, D). Differences occur in the body distortion (126° in our specimen, 85° in the lectotype); the festoons of the frontal lamina (lacking in the lectotype); and the degree of pleonal fusion (more complete in our female). Since we did not collect a male specimen, the original description and figure given by Nierstrasz & Brender a Bran- dis (1929) is reproduced here (Fig. 1E). The features that they recorded are the follow- ing: ““mit Augen; Pigmentierung wie auf def Abbildung; Grenze zwischen Cephalon und VOLUME 103, NUMBER 3 Fig. 3. D, Male dorsal view (all redrawn from Bourdon 1983). Thoracomer II nicht deutlich; Pleon mit kleinem caudalen Fortsatz; Pleomere und Pleotelson verwachsen, I und II aber seitlich angedeutet” (= with eyes; pigmentation as in the illustration. Border between cephalon and first pereomere not clear; pleon with a small caudal process; pleomeres and pleo- telson fused, however I and II indicated lat- erally). Schizobopyrina platylobata (Bourdon, 1983), new combination Fig. 3 Restricted synonymy: Bopyrina platylobata Bourdon, 1983:867-— 869, fig. 13a—-d.—Bourdon & Bruce, 1983: 99. Schizobopyrina platylobata (Bourdon, 1983): A, Female dorsal view; B, Right oostegite 1; C, Barbula; Remarks. — Based on the original descrip- tions and figures recorded in Bourdon (1983) we suggest this species should be included within Schizobopyrina. The female (Fig. 5A— C) has (1) the frontal lamina evident, (2) the maxilliped with palp, (3) the barbula with two lanceolate projections, (4) dorsolateral bosses and coxal plates on pleomeres 1-4, (5) elongated oostegites 2—5, and (6) five pairs of laterally indicated pleomeres and pleo- telson. The exclusion of S. platylobata from Bopyrina Kossmann 1881 is because fe- males in this genus have a maxilliped with- out palp, tiny oostegites 2-5 and complete pleomeral fusion on the short side of body. These features are lacking in S. platylobata. The male of this species (Fig. 3D) is almost identical to those described within Schizo- bopyrina. 638 Schizobopyrina(?) lobata Bourdon & Bruce, 1983 Restricted synonymy: Bopyrina lobata Bourdon & Bruce, 1983: 100-101, fig. 3a-d.— Markham, 1985:46. Remarks. —Bourdon & Bruce (1983) de- scribed this bopyrid and pointed out that the generic position of this species will re- main uncertain. Subsequently Markham (1985) excluded this species from Bopyrina and included it within Schizobopyrina. In agreement with Markham’s first action we considered that this isopod is not a species of the genus Bopyrina; however, we found in the original description given by Bourdon & Bruce (1983) a partial support to Mark- ham’s second action. According to Bourdon & Bruce (1983) S.(?) /obata has a maxilliped apparently without palp (close to Bopyrina), the barbula with single pair of poorly de- veloped projections (like Bopyrina) and lack of dorsolateral bosses and coxal plates (sim- ilar to Bopyrina). In contrast, Schizobopyr- ina possesses a maxilliped palp well defined, the barbula with two pairs of projections, and with dorsolateral bosses or coxal plates or both on pereomeres 1—4. However, the species /obata agree well with Schizobopyr- ina in other features, e.g., oostegites 2—5 elongate, pleomeres laterally indicated, frontal lamina visible. The above-men- tioned similarities and dissimilarities may suggest this species is intermediate between -Schizobopyrina and Bopyrina, but a rede- scription is necessary to resolve the system- atics of this enigmatic species. Schizobopyrina bruscai, new species Figs. 4, 5 Material examined. —One female, holo- type, Bahia Concepcion, Golfo de Califor- nia, south side of El Coyote beach, Baja California Sur, México, 26°44’N, 111°55’W, 2 m depth, 6 Oct 1981, Rubén Rios coll. Host. According to R. Rios (pers. comm.) this host is conspecific with Thor spinosus PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON recorded by Wicksten (1983) [not Boone 1935]. Rios (pers. comm.) has found this cariden is an undescribed species of Thor. Description. — Female (Fig. 4A, B). Length 2.2 mm, maximal width 1.1 mm, head length 0.5 mm, pleon length 0.7 mm, body axis distortion 85°. Head distinct, moderately set into per- eon; frontal lamina large, well differentiated from head, extended into lateral lobules, margin on short side of body slightly con- cave and notched. Antennae inconspicuous; barbula (Fig. 5A) medially produced, with 2 blunt projections. Maxilliped palp large, setose and obscurely articulated (Fig. 5B), plectron projecting, large and slender (Fig. 4C). Pereomeres distinctly separated dor- sally and laterally, much less so ventrally. Coxal plates on pereomeres 1—4 of both sides. Dorsolateral bosses on pereomeres 1- 4 of long side. Pereopods subequal, increas- ing a little posteriorly, each of the 7 articles well defined. First oostegites asymmetrical, left one with posterolateral point short, slender and slightly hooked (Fig. 4D); right one large, broad and hook-shaped (Fig. 5C) internal ridge of both first oostegites slightly orna- mented, with setae on distomedial margin. Oostegites 2—5 slender, fifth one larger. Pleon (Fig. 5D) of 4 distinct pleomeres and pleotelson that are obscurely separated medially; pleotelson notched in middle; 4 pairs of leaf-like and uniramous pleopods diminishing in length posteriorly, first pair suboblong, last three pairs subovate. No uropods. Distribution. —Known only from the type locality, El Coyote beach, Bahia Concep- cion, Baja California Sur, México. Etymology.—This species is named in honor of Richard C. Brusca, for his scientific contributions to the knowledge of the ma- rine fauna of the Golfo de California. Remarks. —Markham’s (1985) diagnosis of the genus Schizobopyrina was based on the type species and took into account all 10 species that he included within this ge- VOLUME 103, NUMBER 3 639 iD Fig. 4. Schizobopyrina bruscai, n. sp., Female holotype: A, Dorsal view; B, Ventral view; C, Maxilliped; D, Left oostegite 1. ee sa J | D, Pleon, ventral view. nus. Schizobopyrina bruscai is similar to another congener in its frontal lamina, max- illiped palp, dorsolateral bosses, shape and number of oostegites and number of pleo- pods. Comparison of the Schizobopyrina species, including S. bruscai, is recorded in Table 1. Schizobopyrina bruscai principally differs from S. amakusaensis (Shino, 1939) [from Amakusa, Kyusyu, Japan], S. an- damica (Chopra, 1923) [from India and Ja- pan], S. brachytelson (Nierstrasz & Brender a Brandis, 1923) [from Dangar, Besar, Saleh Bay], S. gracilis Chopra, 1923 [from India and Japan], and S. /obata (Bourdon & Bruce, 1983) [from off Caloundra, Queensland], in that the first species has the cephalon sep- arated from the first thoracic somite (Fig. 4A), instead of fused as recorded in the last six species (Table 1). Furthermore, S. brus- cai possesses 4 pleomeres and pleotelson, while these species, except S. amakusaensis, have 5 pleomeres and pleotelson. This last pleomeral trait is also observed in S. coch- inensis (Chopra, 1923) [from Cochin, In- dia], S. platylobata (Bourdon, 1983) [from PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Schizobopyrina bruscai, n. sp., Female holotype: A, Barbula; B, Maxilliped palp; C, Right oostegite; Seram, Seleman Bay], S. striata (Nierstrasz & Brender a Brandis, 1929) [from California and Golfo de California, Mexico) and S. urocaridis (Richardson, 1904) [from Atlan- tic coast of U.S. and Belize], and represents the most evident difference between these last four species and S. bruscai. Additional differences are recorded in Table 1. Schizobopyrina bruscai differs from S. miyakei Shiino, 1939, from Palao and Aus- tralia, in the margin of pleotelson, bilobed- symmetrical in S. bruscai, and from obliquely-continuous to bilobed-asymmet- rical in S. miyakei. Additionally, S. bruscai has the cephalon and first thoracic somite well separated by a deep groove, instead of fused with suture line indicated by a shallow groove as in S. miyakei. With regard to the diagnosis of Schizobopyrina, Markham (1985) pointed out that this genus possesses six distinct pleomeres (= 5 pleomeres and pleotelson). S. amakusaensis, S. bruscai, and S. miyakei have 4 pleomeres and pleotel- son. This pleonal variation is analogous to that recorded in other genera of Bopyridae VOLUME 103, NUMBER 3 641 Table 1.—Comparison of selected diagnostic features among species of Schizobopyrina. Species marked with (*) have the cephalon and pereomere 1 fused. Abdominal pleomeres and Dorsolateral pleotelson bosses Coxal plates (PT) S. amakusaensis (*) present present 4+ PT S. andamica (*) present present 5 + PT S. brachytelson (*) present present 5 + PT S. bruscai present present 4+ PT S. cochinensis indistinct present 5+ PT S. gracilis (*) absent present 5) ar Jee S. kossmanni (*) present present Saati S. lobata (*) absent absent 5 + PT S. miyakei present present 3+ PT 4+ PT S. platylobata present present 5 + PT S. striata present present 5 + PT S. urocaridis present present 5 + PT Pleomere V No indicated Margin of pleomere V No indicated Pleotelson margin Rounded con- tinuous Extending far Angular Rounded con- beyond telson tinuous margin Extending far Angular Straight contin- beyond telson margin uous No indicated No indicated Bilobed No extending Rounded Rounded con- far beyond tinuous telson margin Extending far Rounded Straight contin- beyond telson uous margin Slightly extend- Angular Rounded sin- ing far be- uous yond telson margin No extending Rounded Asymmetrical far beyond bilobed telson margin No indicated No indicated Obliquely con- tinuous or asymmetrical bilobed Extending far Angular Bilobed beyond telson margin No extending Rounded Concave con- far beyond tinuous telson margin Slightly extend- Rounded Widely rounded ing far be- yond telson margin infesting caridean shrimps (e.g., Bopyrina, Markham 1985; Bopyrione, Bourdon and Markham 1980) and should be included in the diagnosis of Schizobopyrina. Acknowledgments We are grateful to Rubén Rios (Centro de Investigacion Cientifica y de Educacion Su- perior de Ensenada) who allowed us to ex- amine and describe S. bruscai, and gave us information about taxonomy of the species of Thor from the Golfo de California. We are indebted and deeply grateful to Roland Bourdon and John Markham for their sup- port of our bopyrid studies. Our great ap- preciation is due to Roger Seapy for his crit- icism of the first draft of this note and to Lon McLanahan and Lady Janie Yeo Kirk for their support during our stay in Cali- fornia State University, Fullerton. This work 642 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON was sponsored by project ““Sistematica y bioecologia de los Arthropoda dominantes en areas selectas del municipio de Ensena- da, Baja California” of the Escuela Superior de Ciencias, Universidad Autonoma de Baja California and by the agreement SEP-UABC 089-01-0352. Literature Cited Boone, L. 1935. Crustacea: Anomura, Macrura, Eu- phausiacea, Isopoda, Amphipoda and Echino- dermata: Asteroidea and Echinoidea. Jn Sci- entific results of the world cruise of the yacht “Alva,” 1931 William K. Vanderbilt, com- manding.—-Bulletin of the Vanderbilt Marine Museum 6:1-264. : Bourdon, R. 1983. Expedition Rumphius II (1975) Crustacés parasites, commensaux, etc. (Th. Monod Ed.). VIII. Crustacés Isopodes (3e par- tie: Epicarides Bopyridae).—Bulletin du Mu- seum d’Histoire Naturelle, Paris, Serie 4, 5, Sec- tion A, 3:845-869. —., & A. J. Bruce. 1983. Six bopyrid shrimps (Isopoda, Epicaridea) new to the Australian fau- na.—Crustaceana 45:96-106. —, & J.C. Markham. 1980. A new genus and species of bopyrid isopod infesting alpheid shrimps in the genus Synalpheus in the Eastern Atlantic Ocean. —Zoologische Mededelingen (Leiden) 55:221—230. Campos, E., & A. R. de Campos. 1989. Epicarideos de Baja California: distribucion y notas ecoldé- gicas de Probopyrus pandalicola (Packard, 1879) en el Pacifico oriental.—Revista de Biologia Tropical 37(1):29-36. Chopra, B. 1923. Bopyrid isopod parasitic on Indian Decapoda Macrura.—Records of the Indian Museum 25:411-550. Markham, J. C. 1985. A review of the bopyrid iso- pods infesting caridean shrimps in the north- western Atlantic Ocean, with special reference to those collected during the Hourglass cruises in the Gulf of Mexico.—Memoirs of the Hour- glass Cruises 7(3):1—-156. 1988. Description and revision of some species of Isopoda Bopyridae of the north West- em Atlantic Ocean. Zoologische Verhandelin- gen, Leiden 246:1-63. Nierstrasz, H. F., & G. A. Brender a Brandis. 1923. Die Isopoden der Siboga-Expedition. II. Isopo- da Genuina. I. Epicaridea. Siboga-Expeditie Monography 32°: 4 unnum + 1-65. ——,, & 1929. Papers from Dr. Th. Mor- tensen’s Pacific Expedition 1914-16. XLVIII. Epicaridea I.— Videnskabelgie Meddelelser fra Dansk Naturrhistorisk Forening i Kobenhavn 87:1-44. Shiino, S. M. 1933. Bopyrid from Tanabe Bay.— Memoirs of the College of Science, Kyoto Im- perial University, Series B, 7:249-300. . 1939a. A bopyrid from Palao.—Palao Trop- ical Biological Station Studies 24:598-601. . 1939b. Bopyrid from Kyusyu and Ryukyu. — Amakusa Marine Biological Laboratory 70:79- 99. 1942. Bopyrids from the South Sea Islands with description of a hyperparasitic cryptonis- cid.—Palao Tropical Biological Station Studies 2:437-458. Wicksten, M. K. 1983. A monograph on the shallow water caridean shrimps of the Gulf of California, Mexico.—Allan Hancock Monographs in Ma- rine Biology 134:1-59. (Postal address) Escuela Superior de Ciencias, Universidad Autonoma de Baja California, Apartado Postal 2300, Ensena- da, Baja California, México; Department of Biological Sciences, California State Uni- versity—Fullerton, Fullerton, California 92634. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 643-648 TRIDENTELLA WILLIAMST, A NEW SPECIES OF ISOPOD CRUSTACEAN FROM THE BRITISH VIRGIN ISLANDS, WESTERN ATLANTIC (FLABELLIFERA: TRIDENTELLIDAE) Paul M. Delaney Abstract. —Tridentella williamsi, n. sp., described from the British Virgin Islands, is distinguished from other Atlantic species by its distinctively tuber- culate pleotelson. A species list for the genus is given. Previously only two Tridentella species have been described from the Atlantic, 7. virginiana (Richardson, 1900) and 7. re- cava Bowman, 1986. Herein a third Atlan- tic species, 7. williamsi, is described from the tropical waters of the Virgin Islands, based on examination of specimens from the United States National Museum of Nat- ural History (USNM). Family Tridentellidae Bruce, 1984 Genus Tridentella Richardson, 1905 Tridentella williamsi, new species Figs. 1-3 Material. —Caribbean Sea, British Virgin Islands, “‘fish parasites,’ two males, 8.5 mm holotype (from Epinephelus mystacinus), USNM 239198 and 4.8 mm paratype (from E. flavolimbatus), USNM 239199. Description of male.—Length up to 8.5 mm, about 2.3 x as long as wide. Cephalon without tubercles, with small pointed ros- trum meeting frontal lamina and slightly separating antennule bases. Eyes large, fac- ets well-developed. Pereonites 1—3 without dorsal ornamentation, 4—7 with small tu- bercles along posterior margins, 7 with short, incomplete tubercle rows extending medi- ally on lateral margins. Pereon widest at pereonite 3. Coxae 6—7 visible in dorsal view; coxae 6 extending to midlength of pereonite 7, coxae 7 extending to pleonite 2. All coxae without carinae, tubercles, spines or setae (Fig. 1). Pleonites 1-5 with row of tubercles on posterior margins; pleonites 3—4 with ad- ditional, short median tubercle row, pleo- nite 5 with 2 additional tubercle rows. Pleon widest at pleonite 1. Pleonite | laterally en- compassed by pereonite 7 coxae; pleonite 5 narrow, encompassed laterally by pleonite 4. Pleotelson subtriangular, apex rounded, crenulate, with 6 setae; lateral margins with very large tubercles, dorsum with tubercle row on either side of medial depression, and 2 short tubercle rows near pleotelson base (Fig. 1). Antennule extending slightly past poste- rior margin of cephalon (Fig. 1); peduncle article 3 longer than 1 and 2 combined, fla- gellum of 13-14 articles (Fig. 2A). Antenna extending beyond midlength of pereonite 3 (Fig. 1), peduncle article 5 longest, flagellum of 15 articles (Fig. 2B). Frontal lamina pen- tagonal, lateral margins slightly concave; clypeus short, very broad; labrum small, not encompassed by clypeus (Fig. 2C). Left mandible with narrow, unicuspid incisor process and triangular, setose molar pro- cess; middle palp article longest, with 14 simple setae and | plumose marginal seta (hereafter PMS), apical article with 20 sim- ple setae and 1 PMS (Fig. 2D). Maxillule lateral lobe with 5 large apical spines, 5 smaller subapical hooklike spines; medial lobe slender, with simple rounded apex and 1 apical seta (Fig. 2E). Maxilla 2-segmented, 644 apex with many denticles (Fig. 2F). Max- illiped palp articles 1-3 with simple medial marginal setae, distal 3 articles with comb- like setae; endite extending to penultimate palp article, without coupling hooks, with single apical seta (Fig. 2G). Pereopods 1-3 subsimilar, subprehensile. Pereopod 1 stouter than 2-3, ischium with 2 simple spines on distomedial and disto- lateral angles; merus medial margin with 3 stout bifid spines, | bifid and 2 simple setae, distolateral angle with 2 long spines; carpus medial margin with | bifid, 2 simple spines; propodus medial margin with | bifid, 1 sim- ple seta, and | short spine distally, lateral margin with 3 simple setae; dactylus with 1 simple seta at base of unguis on medial mar- gin, lateral margin with 3 simple setae prox- imal to unguis (Fig. 3A). Pereopods 4—7 slender, ambulatory. Pereopod 4 basis with simple spine on distomedial angle; ischium medial margin with 2 simple setae and 2 spines, distolateral angle with 1 large bifid spine, | simple spine; merus medial margin with | simple and 9 bifid spines, distolateral angle with | simple, 2 bifid spines; carpus distomedial angle with 2 bifid, 2 simple spines, | seta; propodus medial margin with 1 bifid, 2 simple spines and 2 simple setae, distolateral margins with 3 simple setae; dactylus with | seta at base of unguis on medial margin (Fig. 3B). Pereopod 7 basis with 2 spines on distomedial angle, lateral margin with 8 short setae; ischium medial margin with 4 bifid and 4 simple spines, distolateral angle with 2 plumose setae, 2 bifid spines; merus medial margin with 4 simple and 3 bifid spines, distolateral angle with 3 bifid spines, | simple seta; carpus medial margin with | simple and 4 bifid spines, distolateral angle with 1 spine, 2 simple and 10 plumose setae; propodus me- dial margin with 5 bifid spines, distolateral angle with | seta; dactylus medial margin with single seta at base of unguis, lateral margin with 3 setae proximal to unguis (Fig. 3C). Penes set distinctly apart on sternite 7. Pleopods 1—5 rami with PMS as figured, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON endopod of pleopod 5 naked; exopods of 3- 5 with complete or partial transverse su- tures. Peduncles of pleopods 1—5 with 1 spine on lateral margins. Peduncles of pleopods 1—5 with 1 spine on lateral margins. Pleopod 1 peduncle with 3 coupling spines, 1 PMS on medial margin; pleopods 2-4 peduncles with 4 coupling spines and 2 PMS on medial margins. Appendix masculina of pleopod 2 rodlike, with pointed apex, arising from proximal medial margin of endopod (Fig. 3D-H). Uropods extending well beyond pleotel- son apex; exopod one-half width of endo- pod, both rami with scalloped margins inset with PMS, lacking spines. Uropod peduncle with 3 simple setae, 2 PMS on lateral mar- gin, distomedial angle with 3 PMS (Fig. 3]). Etymology.—This species is named for Ernest H. Williams, Jr. in recognition of his contributions to Caribbean isopod biology. Remarks. — Tridentella williamsi is easily distinguished from the other two Atlantic species by its strongly tuberculate pleotelson and lack of cephalic tubercles. 7. virginiana (Richardson) has four cephalic tubercles and an unornamented, smoothly rounded pleo- telson. 7. recava Bowman lacks cephalic tu- bercles, has a pleotelson apex with a dis- tinctive U-shaped excavation, and the pleotelson dorsum is unornamented. Tridentella williamsi is the 14th species known in the genus, and has a highly or- namented or sculpted pleotelson as do 9 of the other 13 species. T. williamsi also lacks the marginal uropodal spines found in five other 7ridentella species. Species list.—The currently-known Tri- dentella species and their localities are listed below. See Delaney & Brusca (1985) for syn- onymies. 1. T. acheronae Bruce, 1988; New Zea- land. 2. T. cornuta Kussakin, 1979; northwest Pacific. 3. T. glutacantha Delaney & Brusca, 1985; California. VOLUME 103, NUMBER 3 645 Fig. 1. Tridentella williamsi, n. sp., dorsal view of holotype male, USNM 239198, British Virgin Islands, 646 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Tridentella williamsi, holotype: A, Antennule; B, Antenna; C, Frontal lamina, clypeus and labrum; D, Left mandible; E, Maxillule; F, Maxilla; G, Maxilliped. 647 VOLUME 103, NUMBER 3 Pleopod 1; E, > D Tridentella williamsi, holotype: A, Pereopod 1; B, Pereopod 4; C, Pereopod 7; Fig. 3. Pleopod 2; F, Pleopod 3; G, Pleopod 4; , Pleopod 5; I, Ventral view of uropod. H >) 648 4. T. japonica Thielemann, 1910; Japan. 5. T. laevicephalax Menzies, 1962; south- ern Chile. 6. T. ornamenta (Menzies & George, 1972); Peru-Chile Trench. 7. T. quinicornis Delaney & Brusca, 1985; California. 8. T. recava Bowman, 1986; New York Bight. 9. T. saxicola (Hale, 1925); Australia. 10. T. sculpturata Kussakin, 1955; north- west Pacific. 11. 7. tangaroae Bruce, 1988; New Zea- land. 12. T. virginiana (Richardson, 1900); Vir- ginia to Nova Scotia. 13. T. vitae Bruce, 1984; Fiji. 14. T. williamsi, n. sp.; British Virgin Is- lands, Caribbean. Acknowledgments Specimens were loaned to the author by Dr. Thomas E. Bowman (USNM), and Frances Runyan drew the figures. Institu- tional support provided by the University of the Virgin Islands is gratefully acknowl- edged. I thank Dr. Ernest H. Williams, Jr. for his correspondence and specimen loan which aided evaluation of this Tridentella species. This paper benefited from reviews by several anonymous reviewers. Literature Cited Bowman, T. E. 1986. Tridentella recava, a new iso- pod from tilefish burrows in the New York Bight (Flabellifera: Tridentellidae).—Proceedings of the Biological Society of Washington 99:269- 273. Bruce, N. L. 1984. A new family for the isopod genus Tridentella Richardson, 1905, with description of a new species from Fiji.— Zoological Journal of the Linnean Society, London 80(4):447-455. . 1988. Two new species of Tridentella (Crus- tacea, Isopoda, Tridentellidae) from New Zea- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON land.— National Museum of New Zealand Rec- ords 3(7):71-79. Delaney, P. M., & R. C. Brusca. 1985. Two new species of Tridentella Richardson, 1905 (Isopo- da: Flabellifera: Tridentellidae) from California, with a rediagnosis and comments on the family, and a key to the genera of Tridentellidae and Corallanidae.—Journal of Crustacean Biology 5(4):728-742. Hale, H. M. 1925. Review of the Australian isopods of the cymothoid group. Part I.—Transactions of the Royal Society of South Australia 49:128- 185. Kussakin, O. 1955. New for far-eastern waters U.S.S.R.—the warm water families of Isopoda. — Transactions of the Institute of Zoology and Academy of Sciences U.S.S.R. 18:228-234 (text in Russian). . 1979. Marine and brackish-water Isopoda of cold and temperate waters of the Northern Hemisphere. I. Flabellifera.—Opredeliteli po Faune SSSR 12:1-470, Academy of Sciences, Leningrad, U.S.S.R. (text in Russian). Menzies, R. J. 1962. The zoogeography, ecology and systematics of the Chilean marine isopods.— Reports of the Lund University Chile Expedi- tion 1948-49, No. 42, Lunds Universitets Ar- skriffter, Avd. 2, Bund 57:1-162. —, & R. Y. George. 1972. Isopod Crustacea of the Peru-Chile Trench.—Anton Bruun Report Number 9:124 pp., Texas A&M Press. Richardson, H. R. 1900. Synopses of North Amer- ican invertebrates. VII. The Isopoda.—Ameri- can Naturalist 34:207-230, 295-309. 1905. Isopods of North America.— United States National Museum Bulletin 54:1-727. Thielemann, M. 1910. Beitrage zur Kenntnis der Iso- podenfauna Ostasiens.—Abhandlungen der Mathematisch-Naturwissenschaftlichen Klasse der Koeniglich-Bayerische Akademie der Wis- senschaften, Supplementary Volume 2, Abhan- dling 3:1-109. Marine Science Program, University of the Virgin Islands, St. Thomas, U.S. Virgin Islands 00802. (Current address) Life Science Division, Los Angeles County Natural History Mu- seum, 900 Exposition Blvd., Los Angeles, California 90007. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 649-654 DESCRIPTION OF THE CRYPTONISCIUM LARVA OF ENTOPHILUS OMNITECTUS RICHARDSON, 1903 (CRUSTACEA: ISOPODA: EPICARIDEA) AND RECORDS FROM THE GULF OF MEXICO Daniel L. Adkison and Sneed B. Collard Abstract.—Entophilus omnitectus is reported from two new host species, Munida microphthalma and M. valida, and its range in the Atlantic Ocean is extended from the Azores to the northern Gulf of Mexico. The cryptoniscium larva is described and comments on its maturation to the adult male form are presented. Entophilus omnitectus is an internal parasite enclosed by a host sheath that is similar to that surrounding the Entoniscidae. Seven of the eleven host specimens were also parasitized by Aporobopyrina anomala Markham, 1973 and/or an unidentified rhizocephalan. A specimen of Munida valida Smith, 1883 collected on a University of West Florida training cruise was parasitized by Entophi- lus omnitectus Richardson, 1903, Aporo- bopyrina anomala Markham, 1973 and an unidentified rhizocephalan. The presence of E. omnitectus was indicated by a pore near the anterolateral margin of the host’s car- apace and by swelling and discoloring of the hepatic region of the carapace. This pore is herein referred to as the “larval exit pore” for its presumed function. Galatheoids from subsequent cruises were examined for ex- ternal signs of infestation noted above and additional Entophilus omnitectus speci- mens were found. The material examined has been depos- ited in the collections of the Smithsonian Institution, Washington, D.C. (USNM) and Marine Research Laboratory, Florida De- partment of Natural Resources, St. Peters- burg (FSBC). Entophilus omnitectus Richardson, 1903 Figs. 1, 2 Entophilus omnitectus Richardson, 1903: 53-54, figs. 6-8. — Danforth, 1963:17-18, 40 pls. 3, 7; 1970:13, 72-73, fig. 18.— Bourdon 1976:385-391, figs. 20-23; 1979:511. Material examined. —Gulf of Mexico. In- festing Munida valida Smith, 1883. R/V Bellows B8403 sta 4; 22 Mar 1984, coll. S. B. Collard, 15 ft flat trawl, 185-370 m, 29°16.53'N, 29°12.06’W to 29°12.06'N, 87°47.97'W, Aporobopyrina anomala | 2, 1 6 USNM 240129, Entophilus omnitectus 1 2 (no 6) USNM 240128, host with rhizo- cephalan USNM 240127.—R/V Oregon IT sta 42230; 13 Nov 1984, coll. T. H. Hans- knecht, 55 ft shrimp trawl, 640-685 m, 29°09'N, 87°59’W, 2 hosts each with a pair of branchial and internal bopyrids, Aporo- bopyrina anomala 2 2, 26 USNM 240138, Entophilus omnitectus 2 2, 1 6, 1 cryptonis- cium larva USNM 240137.—R/V Gyre 86- G-2 sta 42; 28 Feb 1986, coll. D. L. Adki- son, 30 ft trawl, 620-850 m, 27°47.0'N, 90°16.9'W to 27°46.5’N, 90°12.5'W, En- tophilus omnitectus 1 ° (gravid), 1 6, FSBC I-33079, host FSBC I-33080.—R/V Gyre 86-G-2 sta 70; 6 Mar 1986, coll. D. L. Ad- kison, 30 ft trawl, 600-770 m, 27°43.2'N, 91°15.7’W to 27°43.2'N, 91°19.4'W, Apo- robopyrina anomala | 2 (gravid), 1 6 FSBC I-33081, Entophilus omnitectus 1 2 (no 4) 650 FSBC I-33082, host FSBC I-33083.—R/V Gyre 86-G-2 sta 76; 7 Mar 1986, coll. D. L. Adkison, 30 ft trawl, 600-750 m, 27-22 AIN 9323 dE Warton 27224, 95N¢ 93°27.8'W, 2 hosts each with a female bran- chial and internal bopyrid, Aporobopyrina anomala 2 2 (gravid), 2 6 USNM 240132, Entophilus omnitectus 2 2 (1 gravid), 1 6 USNM 240131, host with rhizocephalan USNM 240130.—R/V Gyre 87-G-2 sta 26; 5 Mar 1987, coll. D. L. Adkison, 30 ft trawl, 660-1010 m, 27°42.9’N, 91°16.6’W to 27°43.0'N, 91°19.6'W, Entophilus omnitec- tus 1 2 (gravid), 1 6 USNM 240134, host with rhizocephalan USNM 240133.—M/V H.O.S. Citation sample 4502 sta E2A; 13 May 1985, coll. LGL, trawl, 625 m, 28°35.61'N, 86°46.31'W to 28°35.40'N, 86°46.24’W, Entophilus omnitectus 1 2, 1 6 USNM 240126.—M/V H.O.S. Citation sample 4508 sta E2D; 16 May 1985, coll. LGL, trawl, 624-631 m, 28°07.44'N, 85°52.31'W to 28°06.78'N, 85°54.09’W, Entophilus omnitectus 1 2, 1 6 USNM 240124. Infesting Munida microphthalma A. Milne Edwards, 1880. R/V Gyre 86-G-2, sta 47; 2 Mar 1986, coll. D. L. Adkison, 30 ft trawl, 430-690 m, 28°06.1'N, 89°58.9’'W to 28°03.7'N, 89°58.9'W, Entophilus om- nitectus 1 29, 1 6 USNM 240136, host USNM 240135. Description. — Bourdon (1976) gave ex- cellent descriptions and illustrations of E. omnitectus adults. The present material agrees in most details with descriptions of Richardson (1903) and Bourdon (1976) though several differences for the male are noted. The antenna is elongate and more similar to that illustrated by Richardson (1903) than that by Bourdon (1976). All pe- reopods have five segments. Several males appear to be intermediate between adult and cryptoniscium larva with the pleopodal en- dopod being a distinct article. Several spec- imens have a vestigial pleopodal exopod but pereopods are as in mature males. Dis- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cussed in greater detail under maturation of the male. Cryptoniscium larva (Fig. 2): Eyes pres- ent. Antennule, biramous; peduncle of 2 ar- ticles, distally with a tuft of long setae; basal segment without posterior teeth. Antenna of 9 articles, reaching pereomere 4. Oral cone without terminal sucker. Dorsal cuticular striation not seen, presumed absent. Coxal plates without teeth on posterior and ven- tral margins; posterior margin weakly scal- loped; ventral margin with minute serra- tion; posteroventral corner blunt on pereomere |, corner becoming elongate on posterior pereomeres. Pereopods increasing in length posteriorly, dactylus with 1 short seta; propodus palm with 2-3 stout spines; carpus distally with 2 long setae and 0-2 stout spines; merus with 1 large spine on posterior margin; ischium and basis grooved to receive distal segments. Pleopods bira- mous, pleopods 1-3 with 2 setae on disto- medial corner of peduncle; exopod quadrate with | seta on distolateral corner and 5 long feathered setae; endopod broad, rounded laterally, distally with 5 long feathered setae and distolateral corner with 4 crenations. Pleopod 4 with 2 setae on distomedial cor- ner of peduncle; exopod quadrate with 1 seta on distolateral corner and 4 long feath- ered setae; endopod quadrate with 4 long feathered setae and distolateral corner with 3 crenations. Pleopod 5, peduncle without setae on distomedial corner; exopod with 1 seta and 4 long feathered setae on distola- teral corner; endopod elongate, distally with 4 long feathered setae. Uropod peduncle long, approximately twice length of endo- pod, distomedial margin with dense band of setae, distomedial corner with 2 strong teeth, distolateral corner with | seta; exopod approximately 74 length of endopod; en- dopod with 1 seta on ventral surface. Pos- terior margin of pleomere 6 entire. Maturation of the male. —The cryptonis- cium larva is larger than the mature male. The larva appears to attain adult form over VOLUME 103, NUMBER 3 651 Fig. 1. Host carapace and male of Entophilus omnitectus: a, Carapace of Munida microphthalma showing exit pore (arrow) of the female Entophilus omnitectus, anterolateral view; b—d, Antennae; e, Pleopod 1; f, Pleopod 4; g, Pleopod 5; g, Pleopod 5; h, Pleopod 4; i, Pleopod 5; j-1, Uropod. Scale bar A is 1 cm, for Fig. a; scale bar B is 0.3 mm, for Fig. b to 1. Figures from USNM 240136 (host Munida microphthalma) a, b, h, i, k. Figures from USNM 240137 c, j. Figures from USNM 240134 d-g, 1. several molts. While pereopods of ali male specimens except cryptoniscium larva are of adult form, antennae show variable seg- mentation. In general, the basal segment of antennule becomes bilobed and then loses its distal articles. Antenna becomes reduced and then loses its flagellar articles, resulting in an appendage of one or two articles. In the adult form, antenna is largely covered by antennule, thus appearing to be the distal articles of antennule. After the cryptoniscium stage, pleopod setae are lost early except for one long seta on endopod of pleopod 5. This seta reaches nearly to the distal end of the uropod pe- duncle. Between the cryptoniscium and ma- ture male, two forms of pleopods were found. The first has a free endopod and a vestigial exopod. The later form has a uni- ramous pleopod with fused endopod. Changes in uropods are similar to those of pleopods. Distal spines, setae and exopod are lost first, followed by the loss of the dense setal band on the peduncle. In one 652 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Cryptoniscium larva: a, Antennae; b, Pereopod 1; c, Pereopod 3; d, Pereopod 5; e, Pereopod 7; f, Pleopod 1; g, Pleopod 4; h, Pleopod 5; i, Uropod, dorsal view; j, Uropod, distal ventrolateral view. Scale bars are 0.3 mm; scale A for Fig. a; scale B for Fig. b to j. specimen (from Munida microphthalma) a deep invagination is present at the location of the dense setal band. Next, the endopod fuses with the peduncle although its location is indicated by the asymmetrical shape of the distal end of the uropod. Last, all in- dication of the endopod is lost, and the dis- tal end of the uropod is symmetrical. Distribution. —Entophilus omnitectus has a sporadic distribution from off the Hawai- VOLUME 103, NUMBER 3 ian Islands on Munida normani Henderson (type locality; Richardson 1903), off Aus- tralia and the Philippines (Bourdon 1976 as unpublished records), off Madagascar Is- land on M. incerta Henderson (Bourdon 1976), off the Azores on M. sanctipauli Henderson (Bourdon 1979) and in the Gulf of Mexico on M. microphthalma and M. valida (reported here). Position of parasite.—Danforth (1963, 1970) described the position of female En- tophilus omnitectus as mid-dorsal under the host’s carapace. In the present material and those reported by Bourdon (1976), the fe- male E. omnitectus is located anterolater- ally under the host’s carapace, anterior to branchial chamber. The female parasite is oriented with the posterior end of its pleon at the larval exit pore. The long axis of the female is directed obliquely across the host towards the lateral wall on the other side of the host’s carapace. The dorsal surface of female parasite is directed toward the ven- tral surface of host and the brood chamber faces the dorsal surface of the host’s cara- pace. Asin Bopyridae and Dajidae, the male is usually found on ventral surface of the female pleon. As previously noted, presence of an adult female parasite is indicated by a swelling on the dorsal surface of the host’s carapace and a small pore anterior to bran- chiostegite region of the carapace. Female Entophilus omnitectus are en- closed in a host-derived sheath that appears similar to but thicker than the sheath sur- rounding female entoniscid isopods. The host derived-sheath around E. omnitectus is in less intimate contact with the female parasite than around female entoniscids (Shiino 1942, Kuris et al. 1980) where the sheath is very difficult to remove without damaging the specimen. Like the entonis- cids (Kuris et al. 1980), Entophilus omni- tectus is clearly an internal parasite. Discussion. —For the nine hosts with an adult female E. omnitectus, female parasite length is 39% (range 33 to 45%) of host carapace length. The length of the two im- mature female parasites are 19% of host car- 653 apace length. In both cases, the host with an immature female had a partly closed lar- val exit pore and no dorsal swelling of the host carapace. The remains of the previous adult female E. omnitectus were found at the larval exit pore in specimen, FSBC I-33082, and the immature female parasite was found in a swelling on the lateral margin of the host carapace dorsal to the larval exit pore. In the other case (USNM 240131), the immature parasite was found in a “normal” position, with its pleon at the partly closed larval exit pore. It is unknown whether the immature females (both without males) were the transformed males of the previous adult female or the result of subsequent infesta- tion. Entophilus omnitectus has low numbers of large eggs per clutch. The number of eggs in three broods are 435 (USNM 240134), 563 (USNM 240124), and 1524 (FSBC- 133079). One female carried 2228 embryos (USNM 240126). The average egg diame- ters for four broods were 403 um (USNM 240124), 438 um (FSBC-I33079), 548 um (USNM 240131), and 561 wm (USNM 240134). The number of eggs and clutch volume (calculated by treating eggs as spheres and multiplying egg volume times number of eggs) were not found to be related to length of either female parasite or host. Eggs of Entophilus omnitectus are the larg- est reported for any epicaridean and are within the size range of eggs of other isopods (Stromberg 1971). Of the 11 hosts infested by Entophilus omnitectus, only four were not parasitized by females of other parasites. Six hosts were infested by Aporobopyrina anomala, three were infested by an unidentified rhizoceph- alan, and two were infested by females of all three parasites. One rhizocephalan ap- pears parasitized by a cryptoniscid isopod. Markham (1973) reported the simultaneous infestation of Munida valida by a rhizo- cephalan and A. anomala. The known distribution of Entophilus omnitectus is sporadic, no doubt in part due to the relatively subtle inflation of the host 654 carapace when compared to that caused by the branchial bopyrids. The examination of galatheoids using the signs of infestation noted above will turn up more material and should help address the question of whether E. omnitectus is a single species or a species complex. Acknowledgments We wish to thank Thomas H. Hansknecht for allowing us to use material collected by him on the R/V Oregon II and Randall Howard of LGL Ecological Research As- sociates Inc., Bryan, Texas for allowing the use of material under his care. Material from LGL was collected in the Northern Gulf of Mexico Continental Slope Study with fund- ing by the U.S. Department of Interior, Minerals Management Service, Gulf of Mexico Regional Office, under contract numbers 14-12-0001-30046 and 14-12- 0001-30212. We also wish to thank Dr. James M. Brooks, Geochemical and Envi- ronmental Research Group, Texas A&M University, College Station, Texas, for al- lowing one of us (DLA) to make several cruises on the R/V Gyre and use material collected on these cruises. Support for R/V Gyre time was partially provided by the Ma- rine Chemistry Program of the National Science Foundation (grant OCE83-01538 to Brooks). We wish to thank David Camp of Marine Research Laboratory, Florida De- partment of Natural Resources, St. Peters- burg, Florida, for confirmation of the iden- tification of the host specimens. We wish to thank Drs. Thomas E. Bowman and John C. Markham for comments on an early draft of this manuscript. The assistance of Dr. Linda Adkison in the preparation of this paper is gratefully acknowledged. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Bourdon, R. 1976. Epicarides de Madagascar. I.— Bulletin du Muséum National d’Histoire Na- turelle, Paris, Ser. 3, No. 371, Zoologie 259:353- 392. 1979. Bopyridae de la campagne Biacores (Isopoda: Epicaridea).—Bulletin du Muséum National d’Histoire Naturelle, Paris, Ser. 4, No. 1, Sec. A, 2:507-512. Danforth, C.G. 1963. Bopyridian (Crustacea, Isopo- da) parasites found in the eastern Pacific of the United States. Unpublished Ph.D. Dissertation, Oregon State University, Corvallis, 110 pp. 1970. Epicaridea (Crustacea: Isopoda) of North America.—Ann Arbor, Michigan, Uni- versity Microfilms, 191 pp. Kuris, A. M., G. O. Poinar, & R. T. Hess. 1980. Post- larval mortality of the endoparasitic isopod cas- trator Portunion conformis (Epicaridea; Ento- niscidae) in the shore crab, Hemigrapsus ore- gonensis, with a description of the host response. — Parasitology 80:21 1-232. Markham, J. C. 1973. Six new species of bopyrid isopods parasitic on galatheid crabs of the genus Munida in the western Atlantic.—Bulletin of Marine Science 23:613-648. Richardson, H. 1903. Isopods collected at the Hawai- ian Islands by the United States Fish Commis- sion Steamer Albatross.—United States Fish Commission Bulletin for 1903:47-54. Shiino, S. M. 1942. On the parasitic isopods of the family Entoniscidae, especially those found in the vicinity of Seto.—Memoirs of the College of Sciences, Kyoto Imperial University, Series B 17:37-76. Stromberg, J.-O. 1971. Contribution to the embryol- ogy of bopyrid isopods with special reference to Bopyroides, Hemiarthrus and Pseudione (Isopo- da, Epicaridea).—Sarsia 47:1—46. (DLA) 1699 Wesleyan Bowman Road, Macon, Georgia 31210; (SBC) Biology De- partment, University of West Florida, Pen- sacola, Florida 32514. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 655-662 A NEW SPECIES OF ISOPOD, AEGA (RHAMPHION) FRANCOISAE (FLABELLIFERA: AEGIDAE), FROM THE CLOACA OF AN ASCIDIAN FROM THE GALAPAGOS ISLANDS Regina Wetzer Abstract.—Aega (Rhamphion) francoisae, a new species of Aegidae, is re- ported from a depth of 316 m in the Galapagos Islands. All 10 specimens of the type series were found in the cloaca ofa single ascidian, Halocynthia hispida (Pyuridae: Stolidobranchiata). This is the first species of aegid isopod to be reported inhabiting a urochordate. The nature of the relationship is not known. In November and December 1986 the Harbor Branch Oceanographic Institute and SeaPharm Inc. undertook a collecting ex- pedition in the Galapagos Islands. Chief sci- entists for the expedition were Drs. Shirley Pomponi and Kenneth Rinehart, while Drs. Richard Brusca and Francoise Monniot un- dertook studies of the Crustacea and Uro- chordata, respectively. During this expedi- tion, a single deep-water specimen of the stolidobranchiate ascidian Halocynthia his- pida (Herdman, 1881) was collected by the Johnson-Sea- Link research submersible; the ascidian had 10 specimens of an unde- scribed aegid isopod in its cloacal chamber. Members of the family Aegidae are typ- ically predators/parasites on marine fishes and to my knowledge no aegid or any other isopod species have been reported as com- mensals or parasites in ascidians. In the eastern Pacific this family has been most recently treated by Brusca (1983) and Brusca & Iverson (1985). Knowledge of the Aus- tralian aegid fauna has been brought current by Bruce (1983, 1988). Abbreviations used in this paper are: PMS = plumose marginal setae; LACM = Los Angeles County Museum of Natural His- tory, Los Angeles, California, SDNHM = San Diego Natural History Museum, San Diego, California, USNM = National Mu- seum of Natural History, Washington, D.C.; BMNH = British Museum (Natural His- tory), London. Aega (Rhamphion) francoisae, new species Material examined. —Ten individuals, from one specimen of the solitary ascidian Halocynthia hispida (Herdman, 1881) (Family Pyuridae: Order Stolidobranchia- ta), SeaPharm no. 24-XI-86-3-9V. Ecuador, Galapagos Islands, near Fernandina Island, ca. 0.23°S, 91.32°W, 316 m depth, 24 No- vember 1986. Holotype (SDNHM 2220), female 9.6 mm long (with oostegites). Five paratypes (SDNHM 2221) (2 females 10.7 mm, 10.3 mm both with oostegites; 2 males 9.2 mm, 9.2 mm; | manca 4.3 mm). One male paratype each sent to the USNM (250192) (7.4 mm), LACM (86-446.1) (7.5 mm), BMNH (1990:31) (7.8 mm) and the Australian Museum, Sydney (P40180) (8.3 mm). Diagnosis. — Body unornamented; lateral body margins strongly convex, cephalon and pleon considerably narrower than pereonite 4. With a single frontal plate of uncertain homology (frontal lamina wanting; clypeus and labrum fused?). Antennule article 3 at least 3.5 times longer than article 2. Max- illule with 3 apical spines and 3 subapical spines. Distal article of maxillipedal palp in males with ctenate setae. Pleotelson evenly 656 rounded, without marginal notches. Uropo- dal endopod without a deep cleft or notch on lateral margin; apex of inner peduncular angle with 1 long circumplumose seta; ex- opod oblong with prominent, stout, apical spine; endopod subtriangular. Description.—Body smooth and com- pact, slightly more than twice as long as wide; pleon narrower than pereon; pleonites progressively narrower posteriorly (Fig. 1A, B). Cephalon wider than long, subtriangu- lar. Eyes large, well pigmented, with distinct facets, covering 73 of cephalon but not con- tiguous. Cephalon produced anteriorly into a small triangular rostrum extending ven- trally to separate basal articles of antennules (Fig. 2A). Frontal plate, of uncertain ho- mology (fused clypeus and labrum?), shield- like, elevated anteriorly on a broad pedicel, with median transverse ridge, sloping downward posteriorly, and separating an- tennae. Antennules extending to middle of second pereonite; basal articles 1 and 2 somewhat expanded, article 3 3.5—4 times as long as article 2; peduncular articles 2 and 3 with simple and palmate setae; 8—12 flagellar articles, each with 3—4 aesthetascs (not all figured); terminal flagellar article usually with 4 long simple setae and | seta bearing fine setules over most of its length (Fig. 2B). Antennae extending to posterior margin of second pereonite; with 5 pedun- cular articles and 9-10 flagellar articles; pe- duncular articles 2—S with simple setae, ar- ticle 5 also with 3 setae bearing fine setules over most of their length; simple setae on all flagellar articles (Fig. 2C). Both right and left mandibles with a prominent incisor process; with a rounded, fleshy molar process lacking setation; palp of 3 articles subequal in length, proximal (first) article shorter than middle article; middle article with 5 plumose setae with short setules and 3 simple setae on margin; distal article with row of about 18 simple setae on margin (Fig. 3A). Maxillule with 3 stout apical and 3 subapical spines (Fig. 3B). Maxilla with 2 unequal lobes, each with 3 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON stout, slightly curved spines (Fig. 3C). Max- illipedal palp of 5 articles; terminal article very short, with 5 curved ctenate setae; ar- ticle 4 with 5 small recurved marginal spines; article 3 with 4 marginal spines and 1 sub- marginal spine; article 2 with 2 distolateral spines; article | without spines or setae (Fig. 3D). In oostegite-bearing females, proximal maxillipedal palp articles somewhat indis- tinct and basis expanded into a large plate with PMS as figured (Fig. 3E). Pereonites 4—6 considerably longer than pereonites 1-3. Pereonite 7 as long as pereo- nite 3. Coxal plates of pereonites 5—7 visible in dorsal aspect. Pereopods 4—7 consider- ably longer than 1-3 and increasing in length posteriorly (Fig. 4A—G). Pereopods 1-3 with very few setae. Pereopods 4—7 more spinose than pereopods 1-3, as figured. Pereopods 1-3 with large curved dactyl, as long as pro- podus, without expanded distal lobe on pro- podus. Penes small, on sternite 7; fused ba- sally. Pleon with 5 free pleonites plus pleotel- son; pleonite 1 almost entirely covered by pereonite 7; pleonites 2—4 equilength. Pleo- telson with 8 short spines on posterior mar- gin and PMS as figured (Fig. 1C). Uropodal exopod oval; endopod subtriangular, with- out a deep cleft or notch on lateral margin; both with spines and PMS as figured; en- dopods slightly longer than exopods. Inner angle of peduncle strongly produced, about half length of endopod and with 1 long ter- minal plumose seta; outer distal angle of peduncle with one dorsal and one ventral spine (Fig. 1D). Peduncles of pleopods not divided; pleo- podal peduncles 1-4 with 3-6 basally plu- mose coupling spines and 3—9 PMS on inner margin; peduncle of pleopod 5 without cou- pling spines or PMS; peduncles of pleopods 2-5 with | large spine on outer margin (Fig. 5A-F); spine on pleopod 2 often missing (probably broken off). Pleopodal rami un- divided; all pleopods with PMS as figured; density of PMS more-or-less equal on all exopods, but decreasing posteriorly on en- VOLUME 103, NUMBER 3 Fig. 1. C, pleotelson of holotype; D, left uropod of female paratype (10.7 mm). dopods and absent on pleopod 5 endopod; endopod of pleopod 5 with proximal lobe. Pleopods of males similar to those of fe- males. Appendix masculina on male pleo- pod 2 arising basally, long and slender, ta- pering evenly, extending beyond apex of exopod. Remarks. — Aega francoisae fits Brusca’s (1983) diagnosis of the subgenus Rham- phion: antennular articles 1 and 2 are not 657 Aega francoisae n. sp. A, dorsal view of holotype (female); B, lateral view of male paratype (9.2 mm); inflated or dilated, the antennae extend to the posterior margin of pereonite 2, pereo- pods 1-3 lack distal lobes on the propodi, and the uropodal endopods are subtrian- gular in outline. Aega francoisae differs slightly from Brusca’s diagnosis of Rham- phion in that the distal maxillipedal palp article bears ctenate rather than simple setae in males. Brusca (1983) noted that oostegite-bear- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Aega francoisae n. sp. A, frontal lamina, clypeus, labrum of female paratype (10.7 mm); B, left antennule of male paratype (9.2 mm); C, left antenna of male paratype (9.2 mm). ing females of the families Aegidae, Coral- lanidae and Cymothoidae tend to replace the maxillipedal spines with PMS; Aega francoisae has both PMS and spines. Bruce (1988) stated that in Aegidae the mandible possesses a molar process and a 3- or 4-articulate palp. Aega francoisae has a mo- lar process and a 3-articulate palp. A review of the figures in the published literature on Aega, and examination of var- 10us species during this study, indicates that the morphology of the frontal lamina, clyp- eus and labrum can be variable and ex- tremely valuable in distinguishing among closely related species of Aega. Unfortu- nately these structures have often not been illustrated by previous workers. In A. fran- coisae the clypeus and labrum appear to be fused into a single frontal “‘plate.’’ The ap- parent shape of this plate varies depending upon the angle at which the specimen is viewed. Prior to this description only three species of Aega had been recorded from the Gala- pagos Islands (Brusca 1987). Aega acumi- nata Hansen, 1897 is Panamic in distri- bution, occurring at depths greater than 1000 m, and Brusca (1983) assigned this species to the subgenus Aega. Aega plebia Hansen, 1897 and A. longicornis Hansen, 1897, like A. francoisae belong to Brusca’s subgenus Rhamphion. Aega plebia is widespread VOLUME 103, NUMBER 3 659 7 Uf, Fig. 3. Aega francoisae n. sp. A, left mandible of holotype; B, left maxillule of male paratype (9.2 mm); C, left maxilla of male paratype (9.2 mm); D, right maxilliped of male paratype (9.2 mm); E, right maxilliped of holotype. 660 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Aega francoisae n. sp. Left pereopods of holotype (female): A, pereopod 1; B, pereopod 2; C, pereopod 3; D, pereopod 4; E, pereopod 5; F, pereopod 6; G, pereopod 7. throughout the Panamic and northwest Pa- cific temperate regions at depths exceeding 500 m. Aega longicornis, collected at Al- batross Station 3402 (eastern Pacific Ex- pedition) in 1891 at 0°57'30”S, 89°3'30”W at 842 m, is only known from the nonovig- erous female holotype. Of these three species, A. Jongicornis most closely resem- bles A. francoisae. Aega francoisae can be distinguished from VOLUME 103, NUMBER 3 \ \ | 0 \ fj y Ve Oe \ . ¥ D posterior (P), distal tip (DT), and secondary tooth (ST), plus the interior clear streak (S), crescent (C), and triangle (T). The lines numbered 1, 2, 3, indicate where the three adjacent Transmission Electron photomicrographs were taken. The upper angle formed between X and Y is used when referring to tip angle. B. Light microscope view. C. Scanning electron photomicrograph of two rings of hooks. Scale lines = 20 um. Tissue for TEM embedded in Spurr’s medium, sectioned 2000 A thick, and photographed on a Zeiss 902. (Fig. 1A). Exceptions to this include P. pa- cificum (90-125 um), P. meteori and P. sa- prophagicum (15-30 wm). Three species (P. arcuatum, P. nigrescens, and P. stephen- soni) are reported to have hooks in the 50- 80 um range, but we have measured hooks up to 120 um in the latter. Small worms tend to have smaller hooks than larger members of the same population (see Ap- pendix). Therefore, while this character state is not diagnostic for most species it can be helpful for a few at the two ends of this continuum. D. Shape (secondary tooth and tip): In- VOLUME 103, NUMBER 3 trovert hooks differ in shape but how to describe the differences has often proved difficult. Consequently the same hook is not always consistently described. A “‘second- ary” or “accessory” tooth or “keel” is com- monly present on the posterior edge (con- cave side) of most species of Phascolosoma (Figs. 1, 2). It ranges from being small, frag- ile, and sharp to being large, hump-like and blunt. The tooth is constant within a species (except for P. nigrescens) and has taxonomic importance. The distal tip may also vary from an acute point to a rounded, blunt apex. While there is a generally consistent pattern within a species, one does find worn and rounded hooks on an individual with normally pointed hooks, consequently more than a few hooks per worm must be examined. The tip of the hook bends at an angle to the perpendicular and this angle varies within (and among) populations. We have mea- sured this angle as shown in Fig. 1A. Line X is drawn perpendicular to the base through the most anterior part of the concave side and Y is drawn from the tip to intersect X in the middle of the point. Some species have angles less than 90°, and others in the 90-115° range. This second group includes those species commonly described as hav- ing hooks bent at a right angle (e.g., P. al- bolineatum and P. glabrum). The variation in angle within a deme shows no correlation to hook size or whether the hook came from a proximal or distal ring. E. Internal clear streak (apical canal) and triangle: When viewed by transmitted light one sees zones of differing opacity. Trans- mission electron micrographs show the pal- er, lighter regions are where there is little or no organic material (Fig. 1A). The hooks are open basally so that a hollow space (clear streak) extends along the basal edge. This may not always be apparent when the hooks are removed from the tissue. The clear streak (hollow space) extends upward from the base to near the apex. There may also be a tri- angular clear area in the anterio-basal cor- 695 ner that may or may not be separated from the slender clear streak. Additionally, P. ste- phensoni has a pale “crescent” posterior to the clear streak (Fig. 1A, B). The width of the streak, whether it has a sharp or gradual bend near the midpoint, and the presence of swellings along its length have been used in species diagnoses. Using TEM, SEM, and light microscopy together with a comparison of the internal architec- ture of newly formed hooks (from very small worms or from the first two rings) to older, more mature hooks (Fig. 7G, H), it becomes clear that Phascolosoma hooks begin with an open internal space into which partitions of reinforcing material are secreted even- tually. The resulting subdivisions into tri- angles, streaks, or crescents must be at least partially genetically determined. Our ex- amination of the literature and many hooks from within demes suggests that there are some species with distinctive morphs (e.g., P. nigrescens and P. stephensoni), but the degree of variation in many populations is great. One must accept the fact that within some species-complexes (e.g., P. scolops/ agassizil) there is overlap and a degree of variation that can be confusing. Therefore, one must use caution when looking at in- ternal hook-structure despite its potential usefulness. F. Posterior basal elaborations: Separate, small units of diverse shape are associated with the posterior basal edge of most hooks in Phascolosoma (only P. arcuatum has no basal ornamentation). The units are sepa- rated from the main body of the hook and each other, but they really form a single structure which is considered part of the hook. Scanning electron micrographs illus- trate three variations: like P. stephensoni most species are warty (Fig. 3A). In P. sa- prophagicum and P. turnerae these basal processes (rootlets) are taller and thinner (Rice 1985, fig. le; Gibbs 1987, fig. 1) while in P. glabrum an elaborate series of toes to the side of the hook and differing in the two subspecies (Fig. 3B, C) is present. Therefore, 696 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. SEM photographs of hooks from several Phascolosoma: A. P. granulatum; B. P. perlucens; C. P. scolops; D. P. albolineatum; E. P. glabrum glabrum,; F. P. glabrum multiannulatum. Scale lines equal 10 wm. VOLUME 103, NUMBER 3 Fig. 3. SEM photographs of basal elaborations: A. Warts of P. stephensoni; B. Toes of P. glabrum glabrum; C. Toes of P. glabrum multiannulatum. Scale lines equal 10 um. for these latter species basal elaborations are taxonomically useful. 2. Pigmented introvert bands/stripes. — The dorsal side of the introvert is darker than the ventral side in many species. When present, this reddish-brown color is distrib- uted in patches of varying size, from large and almost continuous in P. perlucens, to the more common pattern of irregular, nar- row bands as in P. scolops and P. agassizii. With the exception of P. nigrescens the pres- ence or absence of pigmented bands is con- sistent within a species and, therefore, of systematic value. 3. Introvert length.—The difficulty in us- ing the relative length of the introvert lies in the elastic nature of this body region, easily observed in living worms. Addition- ally, as demonstrated in other genera, the trunk grows faster than the introvert so that the introvert becomes a smaller part of the whole in larger worms (allometric growth). The slope of the linear regression is always negative (Fig. 4). Depending on whether or not the introvert was extended and, if so, how well narcotized the worm was when preserved, the same introvert may appear to be from 75% to 125% of the trunk length. Within one population such a range is com- mon in preserved material (see Appendix). Despite the fact that some populations have introverts averaging less than 90% of the trunk and others average over 125%, the overlapping ranges around these means pre- clude use of these data to identify an indi- vidual worm (Fig. 4 insert). 4. Number of longitudinal muscle bands. — The longitudinal muscle layer on the inside of the body wall is divided into a variable number of bundles or bands. In almost every species there are 18—24, occasionally as few as 15 in smaller or as many as 30 in larger worms (P. pacificum has 30-40). The bands anastomose occasionally and generally are more numerous towards the posterior end (see Appendix). This anterior/posterior dif- ference can range from 0 to 10, and the order of magnitude of this difference is larger for some species than others. The number of bands is not as absolute or constant as stated in Stephen & Edmonds (1972:291), and ap- pears to have no systematic value in the genus. 5. Nephridiopore/anus relationship. — These openings almost always occur at the same anterior/posterior level. If there is a 698 220 190 160 100 Introvert Length as % of Trunk oe) (o} N jo) 40 10 O 5C Trunk Length (inmm) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON BB Ooex%Ke A 70 Fig. 4. Relationship between trunk length and introvert length of four Phascolosoma species; lines are linear regressions (y = a + bx): Squares = P. agassizii, b = —1.01; Circles = P. nigrescens, b = —2.06; X’s = P. perlucens, b = —1.04; Triangles = P. albolineatum, b = —0.47. Insert shows introvert length of same four species to illustrate overlapping ranges despite significantly different means. Each dot represents one extended worm; the horizontal lines are the mean length for each species. difference, the nephridiopores shift anterior (occasionally) or posterior (rarely) to the anus. When a shift occurs, the distance is on the order of 0.5 mm. To use this char- acter in a taxonomic context for this genus would be a mistake. 6. Nephridia length and attachment. — The nephridia are a pair of tubular organs ex- tending towards the posterior end of the trunk. In a few species (e.g., P. pacificum and P. glabrum) they are quite long, 95-— 100% of the trunk, and in a few species (e.g., P. perlucens) they are short (25-45%). In most species they are 40-65% of the trunk length (see Appendix). Between the body wall and the nephridia there are thin connective tissue strands and most authors have noted the extent of their VOLUME 103, NUMBER 3 attachment. While in most species the value averages around 50%, the range within any population can be 30-70% of the nephridia length with no clear correlation to size of nephridia or trunk. Because of the delicate nature of this attachment it can be easily torn during dissection. In a few species this organ is attached along almost its entire length (e.g., P. pacificum). These nephridial features are taxonomically helpful for only a few special cases. Most species are similar, 1.e., in the broad mid-range and variable. 7. Rectal caecum. —Located near the be- ginning of the rectum there may be a small digitiform or bulbous caecum. The presence or absence of this structure has been as- sumed to be consistent within a species. In Stephen & Edmonds’ table (1972:291) they list 10 species with a caecum, 12 without, and 14 unknown. In our analysis (see Ap- pendix) we find from 11-57% of the indi- viduals in a population have a caecum. Sometimes it is clearly visible on the straight part of the gut, but at other times it is cov- ered by the coiled portion and more difficult to find. It is hard to understand how this degree of polymorphism can exist if this structure might have some physiological function. Nevertheless, based on these data, we conclude that this character cannot be used for taxonomic purposes. 8. Papillae (shape/platelets/size). —De- scriptions of the glandular epidermal struc- tures known as papillae are varied, but not always precise or helpful. The careful draw- ings and measurements provided by many early workers are an expression of a typo- logical species concept that overlooks the variations present within demes. These structures do differ but in less precise ways. Three general shapes occur: domelike, mammillate, and conical. Many worms ex- hibit at least two types in various regions of the same animal. The papillae around the base of the introvert (pre-anal) are of par- ticular interest because they help to differ- entiate species (Fig. 5). When observed by transmitted light the papillae appear to have granular platelets of 699 different sizes arranged in diverse patterns around a central pore. The pattern, how- ever, seems to be consistent and unique only in P. noduliferum. When viewed with the SEM the papillae surface appears smooth. This observation suggests that the platelets are sub-cuticular. Additionally, the cuticle over the pre-anal papillae of a few species (e.g., P. stephensoni and P. perlucens) has a smooth hardened appearance (not granular in transmitted light). Size measurements (height, basal diam- eter) have been presented, but the possible range of the measurements over the surface of a single worm is great, usually being smaller in the mid-trunk and larger towards the ends. In a general manner one can speak of large or small papillae, but this is a sub- jective, overall impression and larger worms tend to have larger papillae. 9. Contractile vessel villi. — Along the dor- sal side of the esophagus runs a tubular con- tractile vessel. In some genera the vessel possesses digitiform extensions or villi, but Phascolosoma species do not. Stephen & Edmonds (1972:315) repeat ten Broeke’s (1925) assertion that P. nigrescens has villi, but our examination of her material shows only a bulbous vessel with vesicular pouch- es, not villi. 10. Retractor muscle origins.—In most species the pair of dorsal retractor muscles has its origins from the body wall about 45% of the distance towards the posterior end of the trunk (the range may be as wide as 30- 60% in a given population). The ventral pair originates at about 65% but may range from 50-75% within a population (see Appen- dix). Only P. saprophagicum has the ventral retractors in the posterior quarter of the trunk. In this species and P. arcuatum, both pairs of muscles originate at about the same distance from the ventral nerve cord so that the terms dorsal and ventral lose their meaning. Except for these two species, re- tractor origins cannot be used as an aid in identifying specimens. 11. Karyotypes.—While karyology may eventually be useful to systematists, our PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. (dome); B. P. scolops (mammilate); C. P. albolineatum (dome); D. P. perlucens (cone); E. P. stephensoni (cone). Scale lines = 0.2 mm. present data base is too small to be of value at the species level. Through the efforts of J. Silverstein (pers. comm.) we have infor- mation on the chromosomal morphology of Phascolosoma scolops, P. pacificum, P. per- lucens, and P. agassizii. All have 2N = 20 with a common tendency toward asym- metry of chromosome arm length, e.g., 70- 100% of the chromosomes are subtelocen- tric or telocentric. They all show a gradual reduction in chromosome length from pair one to pair 10 rather than a bimodal pattern of distinctly large and small chromosomes as is seen in other sipunculan genera. Summary. —Four character states are de- termined to be consistent and broadly use- ful to the taxonomist at the species level. These are the number of hook rings, nature of the secondary tooth of the hook, pigment bands on introvert, and shape/size/texture of the pre-anal papillae. Eight characters can be used in a more restricted manner for smaller subsets or special cases. These in- SEM photographs of different shaped pre-anal papillae from several Phascolosoma: A. P. granulatum clude hook size, tip angle, internal clear streak/triangle/crescent, basal elaborations (warts/toes), nephridial length, trunk pa- pillae shape, arrangement of papillae plate- lets, and the position of the retractor muscle origins. Six previously used attributes are of no value to the taxonomist, because of the within-deme variability. These are the presence/absence of hooks, introvert length, the number of longitudinal muscle bands, nephridiopore/anus relationship, the pres- ence of a rectal caecum, and the nature of the contractile vessel. Systematic Section Genus Phascolosoma Leuckart, 1828 Phascolosoma Leuckart, 1828:22, fig. 5.— Keferstein, 1863:39 (part); 1865a:422 (part).—Baird, 1868:91.—Fisher, 1950: 551; 1952:422.—Stephen & Edmonds, 1972:270-271.—Gibbs & Cutler, 1987: 54. VOLUME 103, NUMBER 3 Phascolosomum Diesing, 1851:63 (part); 1859:758 (part). Phymosomum de Quatrefages, 1865:621. Phymosoma Selenka et al., 1883:54 (emen- dation of Phymosomum). Physcosoma Selenka, 1897:460.—Spengel, 1898:50. Diagnosis. —Introvert of variable length often equal to trunk with numerous rings of recurved hooks. Body wall with longitudi- nal muscle layer gathered into bands. Ten- tacles (less than 30) in crescent around nu- chal organ (peripheral tentacles lacking). Contractile vessel without true villi. Four introvert retractor muscles, lateral pairs sometimes partially, rarely completely, fused. Spindle muscle may or may not be attached posteriorly. Two nephridia. Type species.—Phascolosoma granula- tum Leuckart, 1828. Subgenus Phascolosoma Leuckart, 1828 Phascolosoma (Phascolosoma) Stephen & Edmonds, 1972:289-291.—Gibbs & Cutler, 1987:54. Diagnosis. —Spindle muscle attached posteriorly. Introvert hooks without acces- sory spinelets. The following three taxa are not consid- ered valid members of this genus and are discussed first. After the key, the remaining species are alphabetically arranged. Phascolosoma corallicola (ten Broeke, 1925) Physcosoma corallicola ten Broeke, 19235: 90, text fig. 12 (not Sipunculus coralli- colus Pourtalés, 1851:41). Phascolosoma (Phascolosoma) corallico- lum.—Stephen & Edmonds, 1972:298- 299. Discussion. —In the bottle at ZMUA la- beled as type (V.Si. 96) is one piece of coral and three anthozoans but no sipunculans. There are unresolvable peculiarities in ten 701 Broeke’s description of a damaged, hookless worm of unspecified size. She did say it had 12 tentacles despite Stephen & Edmonds (1972) statement that they were absent. Ex- tensive collections in the type locality in recent decades by ourselves and others have failed to yield any worms like this. For these reasons it seems most prudent to place this name on the list of incertae sedis. Phascolosoma longicolle Ruppell & Leuckart, 1828 Phascolosoma longicolle Ruppell & Leuck- art, 1828:6, fig. 1. Grube, 1840:47; 1868: 644.—Diesing, 1851:64; 1859:762.— Baird, 1868:95.—Stephen & Edmonds, 1972:339.—Saiz Salinas, 1989:208. Material examined.—MNHU, Griibe’s Red Sea specimen (3171). Discussion. —The recent analysis by Saiz Salinas shows this name (considered incer- tae sedis by Stephen & Edmonds, 1972) to be associated with worms that belong to the species Golfingia vulgaris. Griibe’s speci- men in Berlin is a P. scolops. Phascolosoma nigritorquatum (Sluiter, 1881) Phymosoma nigritorquatum Sluiter, 1881: 151-152, pl. 1, figs. 3, 8, 11; 1891:117.— Selenka et al., 1883:68-69. Physcosoma nigritorquatum. —Sluiter, 1902: 13.—Fischer, 1919:280; 1921:4—5; 1927: 416. Phascolosoma (Satonus) nigritorquatum. — Stephen & Edmonds, 1972:286-287.— Edmonds, 1980:61-62.—Cutler & Cut- ler, 1983:186. Material examined. —ZMUA, type (V.Si. 80); ZMUH, Fischer’s 1921 material (V8916). Discussion. —We reaffirm our position taken in Cutler & Cutler (1983) that these worms belong in this subgenus, but the quality of the material precludes a mean- ingful definition of this species and the sta- 702 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tus of incertae sedis is appropriate including Fischer’s material. Phascolosoma abyssorum (Southern, 1913) Physcosoma abyssorum Southern, 1913:12- 14S pls lee ties pl 2h. le —ischer 1916:6; 1922b:6; (not Phascolosoma abyssorum Koren & Danielssen, 1877 = Golfingia abyssora). Phascolosoma (Phascolosoma) abysso- rum. —Stephen & Edmonds, 1972:292. Discussion. —We concur with Gibbs (1986), who showed Southern’s species to be conspecific with, and a junior synonym to, Apionsoma capitatum. Key to Phascolosoma Species This is based on the characters of mature hooks taken from complete rings (not distal 2 or 3 or posterior worn and scattered ones). Ik More than 50 rings (complete and/ or incomplete) of hooks ........ 2 Less than 50 rings (complete and/ or incomplete) of hooks ........ 9 . Most hooks over 100 um tall, ne- phridia as long as trunk (Fig. 6A) Fo) as RA MONG Nice hey came an aN P. pacificum Hooks less than 100 um tall .... 3 . Concave side of hook with large rounded hump, toes present but not warts (2 subspecies, Fig. 6B) .... Lhd aia 5 Rte apa are tes oe nates Ea rt P. glabrum Concave side of hook is smoothly tapered or with small secondary TOOU ET: Ace ee peice 4 . Hooks with long, thin basal pro- cessesi(iges ©) earl ee P. turnerae Basal processes, if present, as nor- Tal WartStez-tsee eres ae Ae eee 5 . Hooks without basal warts (Fig. OD) i hore Ere ae Baers Hooks with basal warts P. arcuatum . Clear streak of hook with swelling 13. 14. US. in middle of vertical and horizon- tal portion (Fig. 6E) ... P. nigrescens Clear streak without abrupt swell- TBS! 0 eee nae ts ae op 7 . Hooks with posterior crescent, many over 75 um tall, pre-anal pa- pillae smooth cones, pigment bands on introvert (Fig. 1) ..... SDS ANIR TN uh en a Sa P. stephensoni Hooks without crescents, most less than 75 um tall, no pigment bands On INtTOVert 4/2. 4... eee 8 . Hooks with granular triangle (Fig. 6F), close-set, randomly distribut- ed papillae platelets ...P. granulatum Hooks without triangle, posterior hooks more triangular shape and narrower clear streak (Fig. 7D, E), papillae platelets around pore small and close-set then abruptly more wide-spread P. noduliferum . Hooks less than 25 um tall ..... 10 Hooks over 25 um tall ......... 11 . Fewer than 10 rings of inconspic- uous hooks. (Fig. 6G) .... P. meteori More than 15 hook rings (Fig. 6H) P. saprophagicum . Angle of hook tip greater than 90° (Fig. 61) P. albolineatum Angle of hook tip 90° or less .... 12 Large rounded hump on concave side of hook, pre-anal papillae smooth, posteriorly directed, cone- shaped (Fig. 7A) ....... P. perlucens Concave side smooth or with small toothe.8 ).. Lee ee ee 13 Hooks with separate anterior basal triangles) see ae ee eee 14 Hooks without triangles (Fig. 7B) ee TA ene thee Lata P. maculatum Trunk papillae platelets extending onto inter-papillae surfaces (Fig. 7C) P. annulatum Papillae platelets restricted to pa- pillae surfaces Hook with distinct triangle, nar- row band of red cone-shaped pre- er Cy VOLUME 103, NUMBER 3 703 Fig. 6. Light microscope photographs of Phascolosoma introvert hooks arranged as in the Key. Scale lines = 20 um: A. P. pacificum; B. P. glabrum; C. P. turnerae; D. P. arcuatum; E. P. nigrescens; F. P. granulatum; G. P. meteori; H. P. saprophagicum; I. P. albolineatum. anal papillae (Fig. 7F) ...P. scolops — Hook triangle indistinct or ab- sent, pre-anal papillae not dis- tinct from dome-shaped trunk Phascolosoma agassizii Keferstein, 1866: papillae (Fig. 7G, H) .. P. agassizii 218-219; 1867:46.—Baird, 1868:92; Phascolosoma agassizii Keferstein, 1866 704 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. As for Fig. 6: A. P. perlucens; B. P. maculatum; C. P. annulatum; D & E. P. noduliferum (anterior & posterior); F. P. scolops,; G & H. P. agassizii from most distal ring, not completely erupted (G) & middle ring (H). Fisher, 1952:424—430.— Wesenberg- 1895:10; 1914a:6; 1914b:67-68; 1923:23; Lund, 1954a:8-9; 1957a:3; 1959b:210; 1927:200.—Shipley, 1891:123; 1899a: 1963:126.—Stephen, 1960:516-517; 155; 1902:133; 1903:174.—Chamberlin, 1966:147-148.—Rice, 1967:143-171; 1920a:30; 1920b:5D.—Leroy, 1936:424 1973:1-51.—Stephen & Cutler, 1969:115; (not Fischer, 1919:280; 1922a:7-9). Johnson, 1971:599-600. Phascolosoma (Phascolosoma) agassiZii. — Phymosoma agassizii. —Selenka et al., 1883: Stephen & Edmonds, 1972:292-293.— 78-79. Physcosoma agassizii. — Fischer, Dybas, 1976:67—75.—Cutler & Cutler, VOLUME 103, NUMBER 3 1979a:982-983.—Rice, 1980:492.— Frank, 1983:23-24.—Saiz Salinas, 1986: 52-55. Phymosoma lordi Baird, 1868:92.—Rice & Stephen, 1970:62. Phascolosoma japonicum Grube, 1877: 73.—Fisher, 1952:429.—Wesenberg- Lund, 1963:116-119.—Stephen & Cut- ler, 1969:115.—Cutler 1977b:154—-155.— Cutler et al., 1984:293-296. Phymosoma japonicum.—Selenka et al., 1883:76-78.—Ikeda, 1904:22—23. Physcosoma japonicum. —Selenka, 1888: 220.—Shipley, 1891:122.—Fischer, 1895: 12; 1914a:5-6; 1916:15; 1922a:13-14; 1923:3.—Chamberlin, 1920b:5D.—Os- troumov, 1909:321-322.—Sato, 1930:9- 11; 1937b:149-150; 1939:383-386.— Stephen, 1942:247-248; 1948:220 (vide P. noduliferum).—Leroy, 1936:424 (not Selenka, 1885:21.—Fischer, 1922b:7-8). Phascolosoma (Phascolosoma) japoni- cum. —Stephen & Edmonds, 1972:309- 310. Physcosoma yezoense \keda, 1924:32-34, pl. 1, figs. 3-6. Phascolosoma (Phascolosoma) yezoense. — Stephen & Edmonds, 1972:328-329.— Cutler & Cutler, 1981:91-92. Physcosoma glaucum Sato, 1930:15—-17, pl. 1, fig. 6, fl. 2, figs. 7-8, text fig. 4 (not Phascolosoma glaucum Lanchester, 1905a:32 = Golfingia glauca). Phascolosoma (Phascolosoma) glaucum. — Stephen & Edmonds, 1972:306.—Cutler & Cutler, 1981:89-90. Physcosoma formosense Sato, 1939:398- 401, pl. 20, fig. 15, text figs. 36-41. Phascolosoma (Phascolosoma) formo- sense. —Stephen & Edmonds, 1972:304.— Cutler & Cutler, 1981:89. Phascolosoma (Rueppellisoma) golikovi Murina, 1975:54—55, fig. 1.—Cutler & Cutler, 1983:180. Material examined.—MCZH, type ma- terial (186, 439); our recent California col- lections; P. japonicum, MNHU, type (1024) plus 979 & 1025, and Fischer’s 1922 Port 705 Elizabeth specimens (6087); NHRS, Fi- scher’s 1922 Eugenie material (162, 164); recent Japanese collections (Cutler et al. 1984). Discussion. —Considerable lack of clarity about this taxon has existed. Our decision to combine these names comes after pro- tracted analysis of numerous specimens. As presently defined P. agassizii has fewer than 30 rings of hooks that are 30-70 um tall, each having a variable clear streak, a tri- angle that is usually indistinct, and if a sec- ondary tooth is present, it is small (Fig. 7G, H, and Fisher 1952, plates 37-38 for vari- ety). When others have reported this species as having more than 30 rings of hooks they were probably looking at a different species. The introvert has irregular pigment bands and is about as long as the trunk. The trunk is covered with papillae that exhibit much variety in color (generally darker than the skin). Papillae platelets are variable in size (3-9 um) and with a random distribution. Published illustrations overlook the varia- tion present, even on a single worm. Previous decisions to reduce some taxa to junior synonyms of P. japonicum are re- affirmed here with the subsequent move into synonymy with P. agassizii. These are P. glaucum and P. yezoense (Cutler & Cutler 1981), plus P. golikovi (Cutler & Cutler 1983). In the description of P. formosense, Satd (1939) seems inconsistent. He stated that the hooks are scattered but later wrote about the number of rings. His figure shows three hooks in a line as if part of a ring. As sug- gested in Cutler & Cutler (1981) Sato prob- ably had two worms with few hook rings that were partially covered by a fold of skin. The hook and papillae shape fit within our concept of P. agassizil. Distribution. —Common on both sides of the north Pacific Ocean (Mexico to Alaska on the eastern side and Japanese waters in the west), plus scattered records from cooler Indian Ocean waters. Several records from south and west Africa exist, and while these may include some misidentified P. granu- 706 latum, others are correct. Australian and other warm water Indian and Pacific Ocean records (including Keferstein’s Panama ma- terial) are considered to be other Phasco- losoma species. Phascolosoma agassizii kurilense (Satd6, 1937) Physcosoma kurilense Sato, 1937a:117-120, text fig. 14. Phascolosoma (Phascolosoma) kurilense. — Stephen & Edmonds, 1972:310-311.— Cutler & Cutler, 1981:91. Material examined. —None. Discussion. —The earlier decision (Cutler & Cutler 1981) to reduce the northern P. kurilense to a subspecies of P. japonicum is reafhrmed here. However, P. japonicum 1s now a junior synynym of P. agassizii. The one morphological difference is the presence of a small secondary lobe on the nephridia of mature worms. Distribution. —Kurile Islands. Phascolosoma albolineatum (Baird, 1868) Phascolosoma albolineatum Baird, 1868:91- 92.—Wesenberg-Lund, 1963:128.—Ste- phen, 1967:45.—Rice & Stephen, 1970: 59.—Edmonds, 1980:56—-57.— Murina, 1981:13.—Cutler et al., 1984:292-293. Phymosoma albolineatum. —Selenka et al., 1883:71-72.—Augener, 1903:301-302. Physcosoma albolineatum. — Fischer, 1913: 99: 1914a:6; 1922a:9.—Ikeda, 1924:32.— Leroy, 1942:6-9.—Sato, 1935:312; 1939: 395-396.—Tokioka, 1953:140. Phascolosoma (Phascolosoma) albolinea- tum.—Stephen & Edmonds, 1972:293- 295.—Cutler & Cutler, 1979a:983. Phymosoma microdontoton Sluiter, 1886: 506, pl. 4, fig. 9; 1891:118; 1902:13. Phascolosoma (Phascolosoma) microdon- toton. —Stephen & Edmonds, 1972:312- 313. Phascolosoma multiannulata Wesenberg- Lund, 1954b:378-383, text figs. 2-6 (part). Phascolosoma andamanensis Johnson, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1971:603-604, pl. 1, figs. 1-7.— Haldar, 1976:3—-4. Material examined.—BMNH, type; In- dian Ocean and Japanese material (Cutler & Cutler 1979a, Cutler et al. 1984); P. mul- tiannulatum, UZMK, type material, (cata- loged as Golfingia multiannulata); P. mi- crodontoton, ZMUA, type (V.Si. 77). Discussion. —This taxon is characterized by the tip of the hooks being bent at an angle of more than 90°. Superficially it strongly resembles pale specimens of P. scolops, and unless hooks are examined the two species could be misidentified. On the concave side the hooks have a large bulge but no sec- ondary tooth, giving the base a less trian- gular form. The clear streak often does not extend beyond the mid-point (Fig. 61). The hooks are from 25-65 um tall and arranged in less than 40 rings. The introvert is shorter than the trunk, and the nephridia are usually less than one-half the trunk length. In some specimens only two or three retractor mus- cles are present owing to some degree of fusion (Cutler & Cutler 1979a:983, Cutler et al. 1984:292). Phascolosoma microdontoton (Sluiter, 1886) is considered a junior synonym. The original illustration of a hook is misleading. The internal structure and the external shape match P. albolineatum except that the tips of some hooks are worn down. The band of pre-anal papillae are also comparable. When Wesenberg-Lund (1954b) de- scribed P. multiannulatum she reported 500 specimens, of which only 20 can now be located. Of these, 16 are clearly P. alboli- neatum, with pale trunks 8-25 mm long. Her description was a combination of char- acters and we have redefined her taxon be- low. The name P. andamanensis was intro- duced by Johnson (1971) and used by Hal- der in 1976 for a population that was like P. albolineatum except that the hooks had basal bar and warts (see discussion in Mor- phological Section). Haldar (1988), after collecting many specimens in the Andaman VOLUME 103, NUMBER 3 Islands, placed the name in synonymy with P. albolineatum and we concur. Distribution. —Wide spread but not com- mon in Indo-West Pacific tropical shallow waters. Phascolosoma annulatum (Hutton, 1879) Phascolosoma annulatum Hutton, 1879: 278.—Benham, 1903:174.—Edmonds, 1960:160-162. Phascolosoma (Phascolosoma) annula- tum.—Stephen & Edmonds, 1972:296- 297.—Cutler, 1977a:151-152.—Ed- monds, 1980:57—58. Physcosoma _ scolops var. ciense. —Augener, 1903:339. Physcosoma scolops tasmaniense Fischer, 19i4a:3-4. Physcosoma scolops.— Wheeler, 1938:346. Phascolosoma tasmaniense Edmonds, 1956: 285-286. mossambi- Material examined.—BMNH, type; two worms identified by Edmonds from New Zealand and Australia. Discussion.—This species has been re- corded up to 50 mm long with introverts 1-2 times the trunk bearing up to 30 ten- tacles. The hooks in this species are very similar to P. scolops, 45-60 um tall, in up to 25 complete rings. The separate triangle is less distinct (Fig. 7C). Proximal, incom- plete, dorsal rings of smaller (30-35 um), more triangular hooks are partially ob- scured by the papillae and pigment. The bases of the trunk papillae appear polygonal (not oval) and are covered by dark polyg- onal platelets. These platelets spread out over the skin in the interpapillae spaces in a distinctive manner. Distribution. —Southern Australia, New Zealand, and Campbell Island in cooler water. Phascolosoma arcuatum (Gray, 1828) Siphunculus arcuatus Gray, 1828:8.— Baird, 1868:88.—Rice & Stephen, 1970:50-51. Phascolosoma (Phascolosoma) arcuatum 707 arcuatum. —Stephen & Edmonds, 1972: 297-298.—Edmonds, 1980:58—59. Phymosoma lurco Selenka et al., 1883:61-— 63. Physcosoma lurco. —Fischer, 1895:12; 1914a:4-5; 1922a:15.—Sluiter, 1902: 12.—Lanchester, 1905b:37.—Leroy, 1936:424. Phascolosoma lurco.—Edmonds, 1956:290— 291. Phymosoma lurco malaccensis Selenka et al., 1883:63. Physcosoma lurco malaccensis. —Sluiter, 1902:12. Phascolosoma arcuatum malaccense. —Ste- phen & Edmonds, 1972:298. Phascolosoma rhizophora Sluiter, 1891:119- ERIS O23» Physcosoma ambonense Fischer, 1896:337— 338, text figs. 1-3. Phascolosoma (Phascolosoma) ambo- nense. —Stephen & Edmonds, 1972:295-— 296. Phymosoma deani Ikeda, 1905:171-172, pl. 8, figs. 5-8. Phascolosoma (Phascolosoma) deani. —Ste- phen & Edmonds, 1972:299.—Cutler & Cutler, 1981:88-89. Physcosoma esculenta Chen & Yeh, 1958: 273-274, text figs. 1-2. Phascolosoma esculenta.—Murina, 1964: 263. Phascolosoma (Phascolosoma) esculen- tum. —Stephen & Edmonds, 1972:301. Material examined.—P. deani, ZMUT, type; P. esculenta, material identified by Murina; P. /urco, MNHU, type (488, 971): UZMC, Lanchester’s material; P. rhizoph- ora, ZMUA, type (V.Si. 84). Discussion. —This species has very dark, large papillae with sharp borders against a light yellow-brown skin. The introvert may be twice the trunk length and bears over 100 complete and incomplete rings of hooks. The hooks are 40—70 um tall, without sec- ondary teeth, warts, or toes. Internally the hooks are simple, with the clear streak great- ly expanded basally. The retractor muscles 708 are unique in that the origins of the broad posterior pair have shifted dorsally (to lon- gitudinal muscle bands 2-3) and the origins of the thinner anterior pair have shifted ventrally (to muscle band 1). Also, the entire retractor complex may appear as a single fused unit with four roots when contracted. The circular muscle layer is divided into anastomosing bands thereby creating “‘coe- lomic sacs”’ where gas exchange between the environment and the coelomic fluid can oc- cur more readily as in many Sipunculidae. In 1970 Rice & Stephen concluded that P. arcuatum and P. lurco were conspecific and in 1972 Stephen & Edmonds added P. rhizophora. In 1981 Cutler & Cutler con- tinued this process by adding P. deani and P. esculenta to the list of junior synonyms. The justification for retaining P. arcu- atum malaccense as a valid subspecies is weak. ““The papillae on the middle of the trunk are formed of many concentric plates instead of being arranged irregularly”’ (Ste- phen & Edmonds 1972:298). Later they state “*. . figure of. . .nominate form shows only a partial concentric arrangement of the plates.”’ Selenka et al. (1883) did not pro- vide any illustrations of their subspecies, and we conclude that this taxon does not merit separate status. The type material of P. ambonense Fi- scher, 1896, is not located in those mu- seums where Fischer deposited other spec- imens, and there are no other records of this taxon. The similarities of this to P. arcu- atum, a species common in Indonesia, have been noted by earlier writers and include: the retractor muscles (ventral pair anterior to dorsal), the weakly developed and few longitudinal muscle bands (only 16 in a 140 mm worm (total length and not 14 mm as in Stephen & Edmonds 1972:285)), and hook shape. Our inability to verify this in- formation is not helpful. Rather than add this name on the list of species inquiren- dum, pending future clarification, we place it in the present synonymy. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Distribution. —Northeast India, Anda- man Islands, southern China, Vietnam, Philippines, Malaysia, Indonesia, and northern Australia. Tolerates extended pe- riods out of the water and in brackish water, e.g., In Mangrove estuaries. Phascolosoma glabrum glabrum (Sluiter, 1902) Physcosoma glabrum Sluiter, 1902:14-15, pl. 1, figs. 7-8. Phascolosoma (Phascolosoma) glabrum. — Stephen & Edmonds, 1972:305. Phymosoma_ microdontoton. —Shipley, 1898:471; 1899b:56. Physcosoma funafutiense Fischer, 1914a:6— 8, pl. 1, fig. 8; 1922a:11-13. Phascolosoma (Phascolosoma) funafu- tiense. —Stephen & Edmonds, 1972:304— 305; Cutler & Cutler, 1979a:983-984. Material examined. —ZMUA, type (V.Si. 128/5); P. funafutiense, ZMUH, 4 worms collected in West Java by Beneden and de- termined by Augener but never published (V10983); NHRS, Fischer’s 1922 material; Pacific and Indian Ocean specimens (Cutler & Cutler 1979a). P. microdontoton, UZMC, Shipley’s 1898 worms. Discussion. —In several ways (nephridia 75-100% of trunk, trunk papillae evenly distributed and of uniform size all over, a large number of hook rings) P. glabrum is similar to P. pacificum, but is distinguished by having somewhat smaller hooks (60-85 um) with a different shape and internal structure (Fig. 6B). The hooks have a large hump on the posterior edge, the clear streak has an apical expansion, and they are the only hooks that have basal toes (Fig. 3B). In addition, the skin has a smoother texture because of smaller dome-shaped papillae (vs. tall cones). The differences between the two subspecies are described in the following section. When Fischer (1914a) described P. funa- futiense he made no reference to P. glabrum. VOLUME 103, NUMBER 3 Sluiter’s illustration does not show any bas- al toes while Fischer’s material has them. Sluiter overlooked them, although they are present on the type. Cutler & Cutler (1979a) focused on the hook structure and used Fischer’s name. It is now clear that these two species are conspecific. Fischer (1914a) maintained that Shipley’s (1898, 1899b) P. microdontoton actually was synonymous with his new species. Our anal- ysis confirms that conclusion, and, thus, Shipley’s reports are included here. Distribution. —Scattered records from Indo-West Pacific (Diego Garcia, Indone- sia, Eniwetok, Funafuti, Rotuma, and Christmas Island). Phascolosoma glabrum multiannulatum (Wesenberg-Lund, 1954) Phascolosoma multiannulata Wesenberg- Lund, 1954b:378-383, text figs. 2-6 (part). Phascolosoma (Phascolosoma) multiannu- latum. —Stephen & Edmonds, 1972:313- 314. Material examined. —UZMK, type ma- terial, (cataloged as Golfingia multiannu- lata). Discussion. —In the museum collections there are 20 specimens from Wesenberg- Lund’s type material; only four of which have hooks that fit her description. The oth- er 16 worms are P. albolineatum. We do not know what happened to the other 480 worms in the original collection and suspect that Wesenberg-Lund’s description is an amal- gamation of features of both populations. Her description and drawings of the hooks (her figs. 3-5) are of P. g. multiannulatum. Her comments about the papillae and her drawings of these and the internal anatomy (her figs. 2 and 6) fit the P. albolineatum morph. The differences between these four spec- imens and the nominate form are subtle. Our present action allows for future work- ers, having access to a larger sample size, to 709 treat this population as conspecific or re- establish it as a species. The hooks are smaller (up to 60 wm), the clear streak is without an apical expansion, and there are 12 toes on the left instead of 10 toes on the right side (Fig. 3C). The hook rings are far- ther apart, the trunk papillae are larger domes, and sometimes lie in rectangles formed by folds of skin. Distribution. — Type locality at Hikueru, Tahiti. This is at the eastern edge of the species’ range. Phascolosoma granulatum Leuckart, 1828 Phascolosoma granulatum Leuckart, 1828: 22, text fig. 5. (For references between 1828 and 1969 see Stephen & Edmonds, 1972:306-307 and Saiz Salinas, 1984b: 203-205.) Phascolosoma (Phascolosoma) granula- tum.—Stephen & Edmonds, 1972:306—- 309.— Walter, 1973:487.— Haldar, 1975: 59.—Zavodnik, 1975:99.—Zavodnik & Murina, 1975:127; 1976:83.—Gibbs, 1977:28-29.—Saiz et al., 1979:209- 210.—Saiz Salinas & Rallo Gruss, 1980: 114.—Saiz Salinas, 1984a:185-186: 1984b:203-205; 1986:55-62: 1988b: 11.—Ocharan, 1980:115—-116.—Murina, 1977:160-162; 1981:13-14 (not Cutler & Cutler, 1979b:109; 1987b:71). Physcosoma lanzarotae Harms, 1921:307 (part). Physcosoma japonicum. —Fischer, 1922b: 7-8. Material examined. -UZMK, Wesen- berg-Lund’s 1959a material from Cape Verde and Canary Islands; Saiz Salinas’ 1986 material from Spain; Brazil and Cape Verde material (Cutler & Cutler 1979b); P. japon- icum, MNHU, Fischer’s 1922b Port Eliz- abeth material (6087); P. laeve, MNHU, Keferstein’s 1865a worms (6990); P. loveni, UZMK, Wesenberg-Lund’s 1930 material. Discussion.—There has been a lack of 710 clarity about the distinction between this species and P. stephensoni. Some of the ear- lier reports of P. granulatum are really P. stephensoni. Their ranges overlap. It is also probable that some descriptions have been based on a mixture of the two species. The papillae of P. granulatum are granular and dome-shaped, occasionally darker than the body wall, without the taller, smoother, cone-shaped pre-anal papillae that charac- terize P. stephensoni. The introvert is with- out any dark pigmented bands. The hooks are arranged in over 50 rings, some incom- plete. The hooks are 35-70 wm tall, have a narrow clear streak with an indistinct granular triangle, and ill-defined or no sec- ondary tooth (Fig. 6F). The number of lon- gitudinal muscle bands varies considerably between the anterior and posterior in a giv- en worm, averaging 18 and 26 respectively. In general, there is a lack of diagnostic apo- morphic characters suggesting that this may represent an evolutionarily significant mor- phology—closest to the ancestral stock. The material described by Wesenberg- Lund (1930) as P. loveni is clearly Apion- soma capitatum. Sat6’s two reports (1935, 1939) cannot be verified, but based on his comments, and upon zoogeographical con- siderations, we propose that his worms were probably P. scolops. Collin (1892), Wesenberg-Lund (1959c), and Murina (1981) have reported P. gran- ulatum from the Indian Ocean. We have not been able to examine these collections, the descriptions are brief with no figures, and we suggest that the worms belong to one of the tropical Phascolosoma species. Reexamination of the material from Brazil and Cape Verde (Cutler & Cutler 1979b), as well as Wesenberg-Lund’s (1959a) from the Canary Islands, show them to be what we now consider P. stephensoni. Distribution. —Common in the north- eastern Atlantic Ocean from southern Nor- way along the coasts of Europe and the Brit- ish Isles to northern Africa out to the Azores PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and Cape Verde Islands. It extends into the Mediterranean and Adriatic Seas. The Bra- zilian record was an error and we suspect that the Indian Ocean records are not this species either. Phascolosoma maculatum (Sluiter, 1886) Phymosoma maculatum Sluiter, 1886:511- 512, pl. 4, fig. 4; 1891:118-119.—Auge- ner, 1903:308-310. Physcosoma maculatum.—Sluiter, 1902:11. Phascolosoma (Satonus) maculatum. —Ste- phen & Edmonds, 1972:285.—Cutler & Cutler, 1983:185. Material examined. —ZMUA, type (V.Si. 76). Discussion. — Although based on only three individuals whose differences from other species are not clear, P. maculatum is retained as a species. The hooks are dis- tinctive: 90-95 um tall, slender, with the clear streak reaching one-half to two-thirds the distance towards the tip and usually ex- panding into a broad basal triangle (Fig. 7B). Only some of the hooks have a secondary tooth and up to 12 basal warts are present. The posterior trunk papillae are tall cone- shaped while the anterior papillae are mam- miform. The decision by Cutler & Cutler (1983) to move this from the subgenus Satonus was based on our observation that a posteriorly attached spindle muscle was present. Distribution. — Indonesia. Phascolosoma meteori (Hérubel, 1904) Phymosoma meteori Hérubel, 1904a:477- 478, figs. 2-3; 1904b:563; 1907:123-128. Physcosoma meteori.—Stephen, 1941:405. Phascolosoma meteori. —Wesenberg-Lund, 1957b:12. Phascolosoma (Phascolosoma) meteori.— Stephen & Edmonds, 1972:312. Material examined. -MNHN, type ma- terial (V-22). VOLUME 103, NUMBER 3 Discussion.—These are grayish worms with introvert shorter than the trunk. The trunk papillae are small domes and the pre- anal papillae are not randomly arranged but in clusters of 2-5. We found up to ten in- complete rings of pale hooks (15-30 um) with a bent clear streak and anterior triangle only on the ventral surface. These were overlooked by Hérubel. This species is well founded on well pre- served specimens and, except for an as- sumed lack of hooks, is very similar to the other members of this genus. Hérbuel’s three papers are all based on the same material, the second 1904 paper is simply a listing of the stations while in 1907 he gave a more detailed morphological description. How- ever, not all of the museum material is this species (the six Djibouti worms are a mix- ture of P. nigrescens and P. scolops). Some of the animals were surrounded by a mud coating that must have been formed by mucus secretions much like that of Phas- colion lutense. One had a commensal poly- chaete sharing the shelter as do some Phas- colion species. The contractile vessel is large and folded with vesicular swellings, and not villi (even in those specimens with the in- trovert completely extended.) The many tentacles and enlarged contractile vessel are probably adaptations to increase the surface area for gas exchange, a feature seen in other sipunculans from this region (Sipunculus, Phascolion). Distribution. —Red Sea and Gulf of Aden, 18-38 m. Phascolosoma nigrescens (Keferstein, 1865) Phascolosoma nigrescens Keferstein, 1865a: 424, pl. 31, fig. 2, pl. 32, figs. 14-15; 1865b:198—-199. (For references between 1865 and 1965 see Stephen & Edmonds, 1972:315.).—Murina, 1964:263-265; 1967:43-44; 1968:422; 1970:68; 1971: 83.—Stephen, 1965:82; 1967:46.—Cut- 711 ler & Kirsteuer, 1968:353.—Edmonds, 1971:148-149.—Amor, 1975:22-23.— Haldar, 1975:59-60; 1976:4—5.—Saiz Sa- linas, 1988a:165; 1988b:12. Phascolosoma (Phascolosoma) nigres- cens.—Stephen & Edmonds, 1972:315- 316.—Cutler, 1977a:152.—Cutler & Cut- ler, 1979a:984—-985:; 1979b:108.—Ed- monds, 1980:59-61.—Murina, 1981:14; Cutler et al., 1984:296.—Saiz Salinas, 1984b:208-210. Phascolosoma puntarenae Griibe & Oecer- sted, 1858:13.—Diesing, 1859:761.—Ke- ferstein, 1863:40.—Fisher, 1952:430-432 (not Wesenberg-Lund, 1959a:191-193). Sipunculus (Phymosomum) puntarenae. — de Quatrefages, 1865:624. Phascolosoma (Phascolosoma) puntaren- ae. —Stephen & Edmonds, 1972:319-320. Phascolosoma varians Keferstein, 1865a: 424-426, pl. 32, fig. 22; 1865b:199-200; 1867:48-49.—de Quatrefages, 1865: 623.—Wesenberg-Lund, 1954a:7-8.— Rice, 1975:35—49. Phymosoma varians. —Selenka et al., 1883: 69-70.—Shipley, 1890:1—24.—Augener, 1903:340. Physcosoma varians.—Shipley, 1898:468- 473.—Gerould, 1913:419-420.—Fischer, 1922a:16.—ten Broeke, 1925:5.—Sat6, 1939:391-394.—Leroy, 1936:424. Phascolosoma (Phascolosoma) varians.— Stephen & Edmonds, 1972:327-328.— Cutler, 1977a:153.—Rice & MacIntyre, 1979:314. Phascolosoma agassizii Keferstein, 1867:46 (part). Physcosoma agassizii var. puntarenae. — Selenka et al., 1883:79. Phascolosoma planispinosum Baird, 1868: 93.—Rice & Stephen, 1970:65. Phymosoma spengeli Sluiter, 1886:498—-499, oll Sy 1s, By poll, Gh antes 7S INET Physcosoma spengeli.—Shipley, 1899a:156. Phascolosoma (Phascolosoma) spengeli. — Stephen & Edmonds, 1972:325. Phymosoma duplicigranulatum Sluiter, 712 1886:501-502; 1903:307-308. Physcosoma_ duplicigranulatum. —Shipley, 1899a:155.—Sluiter, 1902:13. Phascolosoma (Satonus) duplicigranula- tum.—Stephen & Edmonds, 1972:283- 284.—Cutler & Cutler, 1983:185. Phymosoma lacteum Sluiter, 1886:507-508, pie 4a figs le Os 12-2 aSo Tels. Physcosoma lacteum. —Sluiter, 1902:13.— Shipley, 1899a:155; 1902:134. Phascolosoma (Phascolosoma) lacteum.— Stephen & Edmonds, 1972:311-312. Phymosoma diaphanes Sluiter, 1886:509- 510, pl. 4, figs. 2, 11; 1891:118. Phascolosoma (Phascolosoma) diaph- anes. —Stephen & Edmonds, 1972:299- 300. Physcosoma extortum Sluiter, 1902:15-16, pl. 1, figs. 9-10. Phascolosoma (Phascolosoma) extortum. — Stephen & Edmonds, 1972:303-304. Physcosoma evisceratum Lanchester, 1905a: 31, pl. 1, fig. 1.—Stephen & Robertson, 1952:437. Phascolosoma (Phascolosoma) eviscera- tum.—Stephen & Edmonds, 1972:301- 303. Physcosoma minutum ten Broeke, 1925:87- 88, text figs. 6-7 (not Phascolosoma mi- nutum Keferstein, 1863:40 = Golfingia minuta (Kef.)). Phascolosoma (Antillesoma) minutum.— Stephen & Edmonds, 1972:281.—Cutler & Cutler, 1983:179. Physcosoma horsti ten Broeke, 1925:89, text tres uloles Phascolosoma (Antillesoma) horsti.—Ste- phen & Edmonds, 1972:280.—Cutler & Cutler, 1983:178. 1891:118.—Augener, Material examined. -MNHU, type (6976) plus Keferstein’s from Nordwachter (6998); UWMP, listed as type; RSME, Ste- phen’s from Senegal (1960.48.10); Indian Ocean material (Cutler & Cutler 1979a); Japanese material (Cutler et al. 1984); our 1988 Caribbean material; P. diaphanes, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ZMUA, type (V.Si. 101); P. duplicigranu- latum, BMNH, ZMUA (V.S1. 102), co-types; UZMC, Shipley’s 1899a material; P. evis- ceratum, BMNH, type (1924.3.1.163); RSME, Stephen & Robertson’s 1952 ma- terial (1958.23.66 & 67); P. extortum, ZMUA, type (V.Si. 128/6); P. horsti, ZMUA, type (V.Si. 108); P. lacteum, ZMUA, type (V.Si. 75); P. minutum, ZMUA, type (V.Si. 117); P. puntarenae, MNHU, type (1023); UZMK, from Gribe’s original collections; USNM, identified by W. K. Fisher from Bay of Panama (21463) and from Mexico (24732); P. spengeli, ZMUA, type (V.Si. 128); UZMC, Shipley’s material; P. varians, MCZH, type material (423, 428, 1604). Discussion. —The character that most easily distinguishes this species is the clear streak in the hook that has an expansion near the mid-point of both the vertical and the horizontal portion (Fig. 6E). The angle of the point is usually less (as low as 65°) but may be slightly more than 90°, and with- in one deme the secondary tooth may be large or absent, but is usually small. Hooks are arranged in over 100 mostly incomplete rings and measure 35-90 ym tall. The trunk and the uniform dome-shaped papillae are commonly brown. The introvert is longer than the trunk and may have pigmented bands. The contractile vessel may be en- larged with vesicular swellings but not true villi. As earlier authors, including Edmonds (1980:60) have pointed out, “It is difficult to find a satisfactory character which can be used to distinguish P. nigrescens from P. puntarenae Gribe. If they are the same, 12, puntarenae Griibe, 1859 [sic] has priority.” We agree with this but have been reluctant to submerge the more familiar name. The type material of P. puntarenae is in poor condition, lacks the introvert, and is of no value as a voucher specimen. In Copenha- gen there are two specimens that appear to be part of Griibe’s original material. These had never been dissected nor had any hooks been removed. Our examination showed VOLUME 103, NUMBER 3 them to be Nephasoma and probably N. pel- lucidum. So while there is a very poor foun- dation for the name P. puntarenae, P. ni- grescens 1s represented by the type plus 13 other bottles of worms in Berlin. The single P. nigrescens in Poland, labeled as type, is completely dried out and had not been dis- sected. The older name has been used only twice in the past 125 years. Fisher (1952) used it for five worms from Panama and Mexico but commented on its similarity to P. nigrescens, the difference stated as the bend in the hook. Wesenberg-Lund (1959a) used this name for four worms collected off the Cape Verde Islands. Her comments about the hooks (distinct triangle, absence of swelling on clear streak in many), ne- phridia (longer than the trunk), and the cool water habitat cast doubt on her record. Saiz Salinas (pers. comm.), having looked at her material, concludes that it is P. thomense that we consider conspecific with P. perlu- cens. The name P. puntarenae, based on a shaky foundation, has been avoided by most bi- ologists for over a century, while P. nigres- cens has been used over 60 times during that same period. Therefore, while we con- sider these taxa to be conspecific, we shall continue the past practice of using the fa- miliar junior synonym to preserve nomen- clatural stability and avoid confusion. We reafirm Cutler & Cutler’s (1983) con- clusion that three names, previously in oth- er subgenera, are conspecific with this species: P. duplicigranulatum, P. horsti, and P. minutum. The name P. varians (Keferstein, 1865a) has been used by several biologists for Ca- ribbean worms (3 from the Pacific) with hooks having the internal structure of P. nigrescens, but with the tip being more bent. Most of these hooks also have a blunt sec- ondary tooth on the concave border. Ship- ley (1890) made a detailed micro and macro anatomical study of P. varians. Many fea- tures he studied are common to all Phas- colosoma, and they cannot be used to dif- ae ferentiate P. varians from P. nigrescens. We could find no consistent differences between the two species. When one compares hooks from different populations, a continuum of curvature and secondary tooth develop- ment can be observed (Fig. 8). One possible hypothesis is that hook morphology is de- termined by more than one pair of genes and that allelic frequencies vary from place to place. The alleles for sharp angle and large secondary tooth occur at a high frequency in the Caribbean and a low frequency in the Indo-West Pacific. It is our conclusion that the species are conspecific and that P. var- ians is the junior name because it was de- scribed later on the page. The reason Sluiter (1886) named one worm P. diaphanes seems to be the peculiar arrangement of the retractor muscles. The two ventral muscles are partially split into several strands (not uncommon in this ge- nus), the right dorsal retractor is absent, and the left dorsal retractor has its origin atyp- ically close to the ventral nerve cord. Such fusion and splitting of retractor muscles has been observed elsewhere (Gibbs 1973). This species (individual) with anomalous retrac- tors is a junior synonym of P. nigrescens. In the same paper Sluiter (1886) erected P. lacteum on the basis of one worm, with no differential diagnosis. The distal hooks are up to 90 um tall with long points like some P. nigrescens from Madagascar (Fig. 8). Sluiter evidently measured the smaller (55 um) proximal hooks. In all ways this worm fits the P. nigrescens morph, and we find no reason to retain it as a separate tax- on. When Sluiter (1886) erected P. spengeli, on the basis of two worms, he offered no differential diagnosis so it is not clear how he thought it differed from P. nigrescens. He counted only 22 rings of hooks, and we pre- sume he found differences in hook and pa- pillae morphology. Part of the introvert of the dissected worm is missing and some rings have only a few remaining hooks. A large part of each hook is hidden by folds in the 714 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON {JIL ae [x ee Fig. 8. Composite drawing of P. nigrescens hooks from different populations (internal structure same for all but only shown in first): Row one from single Japanese deme, last three from a single worm. Row two from left to right: Keferstein’s type (first two), P. puntarenae of Fisher, type of P. extortum (same as Cutlers’ Hawaii P. nigrescens), type of P. varians, type of P. lacteum (same as Cutler’s Madagascar P. nigrescens). Row 3; types of P. horsti, P. spengeli, P. minutum. (Scale line = 50 um.) tissue of the retracted introvert and conse- quently easily overlooked. The second worm has at least 50 rings of hooks. The papillae in the mid-trunk are smaller and less dense than in most worms, but overall these an- imals fall within the range of variation de- scribed above and P. spengeli becomes a junior synonym. Of the five worms Shipley (1899a) thus described, two are a species of Themiste, and the remaining three are P. nigrescens. The putative differences used by Sluiter (1902) to differentiate P. extortum from P. nigrescens were the loosely wound gut coil and the hook morphology. Sluiter’s figure of the hook matches that of P. nigrescens very well. The looseness of the gut coil in this single worm is not significant, and there is no reason to retain P. extortum as a valid specific name. Lanchester (1905a) named P. evisceratum on the basis of hooks and papillae of one specimen that consisted of the epidermis only. The hook that is illustrated is clearly one from the most posterior part of the in- trovert. Stephen & Robertson (1952) used this name for two complete worms but with- out removing any hooks. Our examination of their material convinced us that it is P. nigrescens. Distribution. —The most wide spread, cir- cum-tropical species in the genus; generally between 30°N and S in shallow water of all oceans. Phascolosoma noduliferum Stimpson, 1855 Phascolosoma noduliferum Stimpson, 1855: 390.—Keferstein, 1865a:423; 1865b: 198.—Baird, 1868:92.—Edmonds, 1956: 286-288; 1966:176. Phascolosoma (Phascolosoma) nodulifer- um.—Stephen & Edmonds, 1972:316- 317.—Cutler, 1977a:152.—Murina, 1978: 121.—Edmonds, 1980:62. Siphunculus tuberculatus Gray, 1828:8 (part).— Rice & Stephen, 1970:52-53. Sipunculus (Phymosomum) nodulosus de Quatrefages, 1865:621-622. Sipunculus (Phymosomum) javanensis de Quatrefages, 1865:622 (part); Stephen & VOLUME 103, NUMBER 3 Edmonds, 1972:339 (incertae sedis).— Saiz Salinas, 1984b:144—-148. Phascolosoma javenense.—Baird, 1968:94 (part). Sipunculus (Phymosomum) noduliferus de Quatrefages, 1865:624. Phascolosoma grayi Baird, 1868:88.—Rice & Stephen, 1970:52. Phymosoma japonicum Selenkaetal., 1883: 76-78 (part). Physcosoma japonicum.—Stephen, 1948: 220. Material examined. —MNHU, specimen listed as type but appears to be one of Ke- ferstein’s worms; S. Edmonds’ specimen from Victoria, Australia. Discussion. —The similarities to P. agas- sizil are marked, but the differences seem real and lie in the papillae and hooks. The introvert lacks pigment bands, and the pa- pillae are of uniform size, more crowded posteriorly. The uniqueness of the papillae lies in the arrangement of platelets, i.e., around the central pore there is a narrow ring of closely set units that abruptly be- come much more dispersed, but do not ex- tend onto the interpapillae skin. In P. agas- sizil the platelets are more uniformly distributed over the papillae surface. Illus- trations of the hooks in the literature appear to be of the posterior hooks (from incom- plete rings or scattered). The more anterior hooks are much more like P. agassizii (Fig. 7D, E). In general, these hooks have a more narrow Clear streak, lack a triangle, are 60— 90 um tall, usually in more than 50 rings, and have no secondary tooth. Distribution. —Southern Australia and Tasmania plus deep water records from Philippines, New Guinea, and New Zealand (Cutler 1977a). Phascolosoma pacificum Keferstein, 1866 Phascolosoma pacificum Keferstein, 1866: 8-9; 1867:49-50.— Baird, 1868:96.—Ed- monds, 1956:291-—292; 1971:146-148.— 715 Wesenberg-Lund, 1957b:6—7; 1959c:62- 63.—Stephen, 1967:46.— Haldar, 1976:5- 6.—Cutler et al., 1984:296-297. Phymosoma pacificum. —Selenka et al., 1883:63—-65.—Fischer, 1895:12; 1896: 337.—Augener, 1903:310-311. Physcosoma pacificum. —Shipley, 1898:470; 1899a:156; 1902:134.—Sluiter, 1902: 11.—Ikeda, 1904:25-26; 1905:169.— Fischer, 1914a:6; 1922b:8; 1926:108.— Leroy, 1942:23.—Monro, 1931:34.— Sato, 1935:310; 1939:390-391.—Ste- phen & Robertson, 1952:436.—Stephen, 1952:182. Phascolosoma (Phascolosoma) pacifi- cum.—Stephen & Edmonds, 1972:317- 318.—Cutler & Cutler, 1979a:965-986. — Edmonds, 1980:62-63.—Saiz Salinas, 1984b:210-211. Sipunculus (Phymosomum) javanensis de Quatrefages, 1865:622 (part).—Stephen & Edmonds, 1972:339 (incertae sedis). — Saiz Salinas, 1984b:148-155. Phascolosoma javanense. — Baird, 1868:94. Phascolosoma asperum Grube, 1868:642-— 643. Phymosoma asperum. —Selenka et al., 1883: 61. Material examined. —MNHU, labeled type (6969) but only contains eggs; seven additional bottles in this collection, at least two named by Keferstein; MCZH, dried out but appear to be part of the original collec- tion and co-types (500). Discussion.—The uniformly distributed, tall, cone-shaped papillae give this species a rough, sand-paper texture. It is one of the largest species, commonly being up to 80 um (occasionally 125 mm), and is uniform- ly colored. The hooks resemble those of P. perlucens, have a broad base, are 70-125 mm tall, arranged in 80-200 rings, the sec- ondary tooth is hump-like, and an irregular clear streak with a separate triangle is pres- ent (Fig. 6A). The nephridia are long (range from 75-125% of the trunk length) and at- tached for most of their length. Up to 40 716 anastomosing longitudinal muscle bands have been recorded. Distribution. —From the Red Sea, throughout the Indian Ocean, and the west- ern Pacific Ocean from southern Japan to northern Australia including Indonesia and numerous tropical islands at depths less than 3} ile Phascolosoma perlucens Baird, 1868 Phascolosoma perlucens Baird, 1868:90-91, pl. 10, fig. 2.—Rice & Stephen, 1970:63- 64.—Rice, 1975:35-48.—Rice & Mac- Intyre, 1979:314.—Edmonds, 1980:63- 64.—Cutler et al., 1984:297.- Phascolosoma (Phascolosoma) perlucens. — Stephen & Edmonds, 1972:318-319.— Cutler, 1977a:151.—Cutler & Cutler, 1979a:987. Sipunculus (Phascolosomum) vermiculus de Quatrefages, 1865:621. Phascolosoma vermiculum.—Baird, 1868: 85. Sipunculus vermiculus. —Stephen & Ed- monds, 1972:339 (incertae sedis). Phascolosoma (Phascolosoma) vermicu- lus. —Saiz Salinas, 1984b:90-97, 203. Phymosoma dentigerum Selenka et al., 1883: 67—68.—Sluiter, 1886:500. Physcosoma dentigerum. —Sluiter, 1891: 118; 1902:11-12.—Shipley, 1898:474; 1902:134.—Augener, 1903:304-305.— Fischer, 1922a:10-11; 1922b:7.—Mon- ro, 1931:34. Phascolosoma dentigerum.—Fisher, 1952: 432-434.—Murina, 1964:262.—Cutler, 1965:58.—Cutler & Kirsteuer, 1968: 354.—Murina, 1972:306-307. Physcosoma thomense Augener, 1903:343— 344, text fig. 19. Phascolosoma thomense.—Murina, 1967: 44-45; 1968:422. Phascolosoma (Phascolosoma) thomense. — Stephen & Edmonds, 1972:327. Aspidosiphon insularis Lanchester, 1905b: 40, pl. 2, fig. 4.—Gibbs & Cutler, 1987: 56. Paraspidosiphon insularis. —Stephen & Ed- monds, 1972:247. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Physcosoma microdentigerum ten Broeke, 1925:88-89, text figs. 8—10.—Stephen, 1960:517-518. Phascolosoma (Antillesoma) microdentiger- um.—Stephen & Edmonds, 1972:280- 281.—Cutler & Cutler, 1983:178. Physcosoma scolops. —ten Broeke, 1925:86. Phascolosoma puntarenae. —Wesenberg- Lund, 1959a:191-193. Phascolosoma spinosum Johnson, 601-602, pl. 2, figs. 1-9. Material examined. —BMNH, 3 type specimens; our 1988 Caribbean collections; P. dentigerum, MNHU, 2 type specimens (976); P. microdentigerum, ZMUA, type (V.Si. 111) plus additional specimens; A. insularis, BMNH, type (1924.3.1.80); P. thomense, MNHU, type (7010); specimen from Cuba identified by Murina. Discussion. —In 1970 Rice & Stephen re- described Baird’s material and reduced the more familiar P. dentigerum to a junior syn- onym. The assertion that this is the most common rock boring sipunculan in the Ca- ribbean (Rice & MacIntyre 1979) is con- firmed by our recent observations. The red- dish, conical, posteriorly directed, pre-anal papillae on the dorsal base of the introvert characterize this species. The hooks are 30- 60 um tall, in 15-25 rings (8-10 in a 5 mm worm), and have a large rounded secondary tooth. The internal triangle is separate from the clear streak similar to that in P. scolops. The trunk is commonly up to 35 mm long (a few at 45-50 mm), and the introvert is not longer than the trunk, usually with patches of reddish pigment on the dorsal surface. In 1984b Saiz Salinas reintroduced the name P. vermiculus de Quatrefages, 1865. He considered this a senior synonym of P. thomense. While we do not disagree with this, we conclude that both names are syn- onymous with P. perlucens. While strict ap- plication of the ICZN rules would allow us- ing this older name (not used for over 120 years) we will continue applying the more familiar name to avoid confusion and pre- serve nomenclatural stability. 72 VOLUME 103, NUMBER 3 The name proposed by Augener (1903) for two worms from St. Thomas, P. tho- mense, was used twice by Murina (1967, 1968) for some Cuban material. Neither au- thor used the name P. perlucens and our analysis of their specimens, especially the hook morphology, makes it clear that these two names are conspecific. In 1983 Cutler & Cutler examined ten Broeke’s P. microdentigerum and conclud- ed that it is conspecific with P. perlucens. We reaffirm that decision. The contractile vessel exhibits only vesicular swellings, not true villi. The only record of this taxon from the eastern Atlantic is Stephen (1960) with no description or illustrations. Many spec- imens of P. stephensoni (a species also hav- ing enlarged, reddish, pre-anal papillae) have been collected there. The latter may be what Stephen saw, but we cannot confirm this since his specimens cannot be located. Gibbs & Cutler (1987) observed that As- pidosiphon insularis was a Phascolosoma species and probably P. perlucens. This in- formation was repeated in Cutler & Cutler (1989), and is reaffirmed here. Haldar (1988) compared his Indian ma- terial to Johnson’s (1971) description of P. spinosum and concluded that they were con- specific, and we agree. Distribution. —Common in the Caribbe- an (Venezuela to southern Florida), and the western Pacific (Queensland to central Ja- pan). Also recorded from several Indian Ocean locations and in the eastern Pacific off Panama and northern Mexico. The two eastern Atlantic records complete this cir- cum-tropical but patchy distribution. Phascolosoma saprophagicum Gibbs, 1987 Phascolosoma saprophagicum Gibbs, 1987: 135-137, fig. 1. Material examined. —Several worms identified by P. Gibbs from type collection. Discussion.—The single known popula- tion was collected from a decaying whale skull. The worms have small (20-25 um) bluntly rounded hooks with a simple nar- Ui row internal clear streak (Fig. 6H) and small inconspicuous papillae. The nephridiopores are anterior to the anus by up to 10% of the trunk length, and the ventral retractor mus- cles originate in the posterior quarter of the trunk. Distribution. —Chatham Island, New Zealand, 880 m. Phascolosoma scolops (Selenka, de Man & Bulow, 1883) Phymosoma scolops Selenka et al., 1883: 75-76, pl. 2, fig. 17, pl. 10, figs. 138-144. (For synonymy between 1883 and 1965 see Stephen & Edmonds, 1972:321 and Saiz Salinas, 1984b:206—208.) (Not Phys- cosoma scolops ten Broeke, 1925:86.) Phascolosoma scolops. —Stephen, 1965:83; 1966:148; 1967:46-47.—Cutler, 1965: 57.—Stephen & Cutler, 1969:116.—Mu- rina, 1971:83.—Haldar, 1975:61; 1976: 6—7.—Edmonds, 1980:55-60. Phascolosoma (Phascolosoma) scolops.— Cutler, 1977a:152—153.—Murina, 1978: 121.—Cutler & Cutler, 1979a:987-988; 1979b:108-109.— Cutler et al., 1984:298- 299.—Saiz Salinas, 1984b:206—208; 1986: 62-63. Phascolosoma (Phascolosoma) scolops scol- ops.—Stephen & Edmonds, 1972:321- 323) Phymosoma scolops var. mossambiciense Selenka et al., 1883:76, pl. 10, fig. 144. Physcosoma_ scolops var. mossambi- ciense. —Sluiter, 1898:444.—Leroy, 1936: 424.—Stephen, 1942:249.—Edmonds, 1956:285 (not Augener, 1903:339). Phascolosoma (Phascolosoma) scolops mossambiciense. —Stephen & Edmonds, 1972:324. Phascolosoma carneum Riuppell & Leuck- art, 1828:7.—Diesing, 1859:764.—Baird, 1868:85.—Stephen & Edmonds, 1972: 321.—Saiz Salinas, 1989:210-211. Phascolosoma longicolle.—Grube, 1840:47. Phymosoma psaron Sluiter, 1886:505; 1891: 118. Physcosoma psaron. —Sluiter, 1902:13. 718 Phascolosoma (Phascolosoma) psaron. — Stephen & Edmonds, 1972:319. Physcosoma spongicola Sluiter, 1902:16—17, pl. 1, figs. 11-12. Phascolosoma (Phascolosoma) spongico- lum.—Stephen & Edmonds, 1972:325- 326, Physcosoma scolops var. adenticulatum Herubel, 1904b:563. Phascolosoma (Phascolosoma) scolops ad- enticulatum. —Stephen & Edmonds, 1972: 323-324. Phymosoma nahaense \keda, 1904:29-31, figs. 8, 59-62. Phascolosoma (Rueppellisoma) naha- ense. —Stephen & Edmonds, 1972:274.— Cutler & Cutler, 1981:85-86; 1983:180. Physcosoma socium Lanchester, 1905b:37-— 38, pl. 2, fig. 1 (not Phascolosoma socium Lanchester, 1908:1 = Golfingia socia (Lanchester) = Golfingia margaritacea (Sars)). Phascolosoma (Phascolosoma) socium.— Stephen & Edmonds, 1972:324—325. Physcosoma agassizii. — Fischer, 1919:280; 1922a:7-9 (part). Phascolosoma_ rottnesti Edmonds, 282-284, text figs. 1-4. Phascolosoma (Phascolosoma) rottnesti. — Stephen & Edmonds, 1972:320-321.— Edmonds, 1980:64—65. Phascolosoma dunwichi Edmonds, 1956: 292-293, text figs. 12-13; 1980:65-66. Phascolosoma (Phascolosoma) dunwichi. — Stephen & Edmonds, 1972:300-301. Phascolosoma riukiuensis Murina, 1975:55— Sse ys Phascolosoma (Antillesoma) pelmum. — Cutler, 1977a:150. 1956: Material examined. —MNHU, Selenka’s material (960, 977, 978); ZMUA, ten Broeke’s 1925 material (V.Si. 119); our In- dian, Pacific, Atlantic Ocean material (Cu- tler 1965, 1977a, Cutler & Cutler 1979a, 1979b); P. scolops mossambiciense, MNHU, type material (956, 958); P. scolops tas- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON maniense, ZMUH, type (V5407); P. psaron, ZMUA (V.Si. 83); P. rottnesti, AMSS, type (W3598); P. soclum, BMNH, type; P. spon- gicolum, ZMUA, type (V.Si. 128-7). Discussion. —These are usually pale worms with reddish-brown mammiform to dome-shaped pre-anal papillae. While the mid-trunk papillae are small and widely scattered, their size, color, and density in- creases towards the posterior end. The in- trovert is commonly shorter than the trunk and exhibits pigmented bands. The hooks are arrayed in less than 25 rings, are 20-60 um tall, and if a secondary tooth is present it is small (Fig. 7F). The clear streak is sep- arate from the distinct triangle. Apart from hook morphology there are two minor internal differences from the similar P. albolineatum: the rectal caecum is present in more than half the worms (vs. 10%), and the retractor origins are closer to the posterior end of the trunk by about 10% (ventral retractors around 60% vs. 50%). Previous action by Cutler & Cutler (1981) reduced P. nahaense to a junior synonym of P. scolops, and we reaffirm that decision. The two subspecies in Stephen & Ed- monds (1972) have characters well within the range we observe in the nominate form and are here considered to have no distinct status. In 1980 Edmonds reduced his P. dunwichi to a junior synonym of P. scolops and we concur. When Sluiter (1886) described P. psaron he asserted that it lacked hooks and had only fine spines. Our examination revealed rows of normal hooks and papillae of the P. scolops type. Thus we place this name in synonymy. The single 25 mm worm Sluiter (1902) described as P. spongicolum had unusually puffy, thickened skin, but over time this must have changed because it now appears rather normal. The papillae are mammi- form to cone-shaped and while large, not outside the P. scolops range. Figure 33Q in Stephen & Edmonds (1972), presented as VOLUME 103, NUMBER 3 Sluiter’s P. spongicolum hook, is really Slui- ter’s fig. 14 of an Aspidosiphon cylindricus conical hook. Sluiter did illustrate these hooks as fig. 11, which do not show a sep- arate clear steak and triangle. Our obser- vations show the hooks to have the P. scol- ops attributes, and these are considered conspecific taxa. Lanchester (1905b) described P. socium as lacking hooks. Of the three specimens, two lack introverts while the third has rings of P. scolops hooks. He compared this to P. psaron, but we can find nothing to distin- guish this from P. scolops. Examination of P. rottnesti material con- vinced us that Edmonds’ (1956) distinc- tions (papillae platelets and presence of cae- cum) overlooks the variation within P. scolops. While some authors contend that P. scolops lacks a caecum, we observed one in 57% of those we examined. Another hookless worm was given the name P. riukiuensis by Murina (1975). The distal end of the introvert is hookless, but it is also white and gives the impression of a regenerating body part. On the basis of other attributes, we conclude that this is an anomalous individual P. scolops and not a distinct species. Saiz Salinas (1989) reexamined P. car- neum that Stephen & Edmonds (1972) placed in incertae sedis. He concluded that it is conspecific with P. scolops and is the most senior available name. However, be- cause the name has been unused for over 120 years, we would again appeal to the logic of ICZN article 79b to preserve no- menclatural stability and conserve the very often used junior synonym in this case. Distribution.—Common throughout the Indo-West Pacific area including the Red Sea, up to northern Japan, down to northern Australia, and out to Hawaii. The several records from west Africa (Gulf of Guinea and south) cannot all be verified, but we suspect that some of these are P. stephensoni as well as the one Bermuda record (Rice 719 1986). The single Caribbean worm (ten Broeke 1925) is P. perlucens. Phascolosoma stephensoni (Stephen, 1942) Physcosoma stephensoni Stephen, 1942:250, pl. 11, figs. 3-5. Phascolosoma stephensoni. —Wesenberg- Lund, 1963:121-126. Phascolosoma (Phascolosoma) stephen- soni.—Stephen & Edmonds, 1972:326- 327.—Cutler, 1977a:153.—Edmonds, 1980:67-68; 1987:197.—Saiz Salinas, 1982:197-199; 1984b:205—206; 1986:63-— 70; 1988a:166.— Haldar, 1988:127-130. Phascolosoma laeve. —Keferstein, 1863:38-— 39 (part). Sipunculus (Phymosomum) spinicauda de Quatrefages, 1865:621. Phascolosoma spinicauda, Baird, 1868:93. Physcosoma lanzarotae Harms, 1921:307 (part). Physcosoma agassizil. — Fischer, 1922a:7 (part). Physcosoma scolops.— Monro, 1931:35. Phascolosoma heronis Edmonds, 1956:293- 295, text fig. 14. Phascolosoma (Phascolosoma) heronis.— Stephen & Edmonds, 1972:309. Phascolosoma granulatum. —Wesenberg- Lund, 1959a:193-194 (part); Cutler & Cutler, 1979b:108. Material examined.—RSME, type ma- terial (1958:23:17); recent east Atlantic ma- terial (Saiz Salinas 1982 and as P. granu- latum, Cutler & Cutler 1987a); P. laeve, MNHU, Keferstein’s 1863 worms (6991). Discussion. — The large smooth cone-like pre-anal and posterior papillae distinguish P. stephensoni from the similar P. granu- latum that shares part of its range. When viewed under the microscope these papillae show none of the platelets typical of this genus. The smaller papillae in the mid-trunk exhibit only very small, uniform size gran- ules, not platelets. The introvert has irreg- 720 ular pigmented bands and over 40 rings of hooks, 60-110 um tall. There may be only 10-20 complete rings, the remainder rep- resented by dorsal patches only. Internally, the clear streak is smooth, the triangular space is clear, and posterior to the clear streak is a distinct crescentic clear area (Fig. 1). This figure shows that the clear areas are hollow spaces within a more solid matrix. The shape of the hooks varies from distal to proximal, the latter are more scattered, blunter, smaller (30-45 wm), more trian- gular, and the crescentic clear area is not often present. In most hooks the secondary tooth is present and distinct. Edmonds (1980) synonymized P. heronis and Fischer’s (1922a) Port Jackson P. agas- sizii under P. stephensoni, and we concur. For many years P. /aeve Keferstein, and P. spinicauda de Quatrefages, have been con- sidered synonyms of P. granulatum. Kefer- stein’s (1863) material is a mixture, some from Sicily is P. stephensoni while another portion from the Adriatic is P. granulatum. Saiz Salinas has reexamined de Quatrefages single worm and transfers it here, and we concur. As noted above, other biologists have confused these two species (see also Saiz Salinas 1986). An example of this is Harms (1921). Based on his figures P. lan- zarotae 1s clearly a mixture of these. Distribution. — Mediterranean Sea (Sicily and southern Spain), east Atlantic (Azores, Canaries, Gulf of Guinea to South Africa), Indian Ocean (Durban, Mozambique, southwest India), and western Pacific (northern Australia, Solomon Islands, and Hawaii). Phascolosoma turnerae Rice, 1985 Phascolosoma turnerae Rice, 1985:54-60, figs. 1-4. Phascolosoma (Phascolosoma) kapalum Edmonds, 1985:43—44, 2 figs. Material examined. -USNM, type (96687). Six worms from Gyre cruise 87- G-2, sta. EJ-87-127, in cluster of vestimen- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tiferans, 600 m, Bush Hill, off Louisiana, two deposited at USNM (118769). Discussion. — The sharply bent hooks (45-— 80 um tall) have up to 10 long, posterior, basal processes/rootlets (Rice 1985, fig. 2). In light microscope preparations these ap- pear as warts (Fig. 6C). The anterior base is drawn out into a thin prong-like extension. The clear streak is close to the anterior side and often narrows towards the base. The papillae are large and mammillate with ““prominent apical protuberances.” The Bush Hill material differed from Rice’s in minor ways: the papillae in the middle of the trunk are almost as large as those at the extremities, and the nephridia are slightly anterior to the anus (not at the same level) and only 50% attached vs. 75%. The introvert is 1.7 (vs. up to 1.4) times the trunk length in a 19 mm worm with an ex- tended introvert. Edmonds’ (1985) P. kapalum while geo- graphically separated is also from deep water and the putative differences can be ex- plained by the larger size (hook size) and retracted introverts (shorter than trunk). Edmonds (pers. comm.), after looking at specimens of P. turnerae, agrees with this synonymy. Distribution.—Two of the records are from the Gulf of Mexico; off Florida and Alabama at 366-1184 m in wood, and off Louisiana at 600 m near a cold water seep. In both cases these were in close association with Pogonophora. The third record, from 710 m off New South Wales does not in- clude comparable ecological data. Zoogeographical Summary Four of the 16 Phascolosoma species ap- pear to have very restricted ranges, three in specialized habitats: P. meteori from the Red and Arabian Seas (warm, high salinity, low oxygen tension), P. turnerae from the Gulf of Mexico and off Australia (deep cold water in wood or near cold-water seeps), and P. saprophagicum from New Zealand (deep VOLUME 103, NUMBER 3 cold water from rotting whale skull). Phas- colosoma maculatum is from more typical habitat off Indonesia. The most striking feature of the remain- ing 12 species is that 11 of these occur in the waters between the Indian and Pacific Oceans (Australia/Indonesia/Philippines). Adding the two local endemic species and P. turnerae means that 88% live here. This species richness is unparalleled by other si- punculan genera. Eight of these (50% of species in the genus) are widespread throughout the Indo-West Pacific (IWP). Only P. granulatum (from cooler waters of the northeastern Atlantic Ocean and the Mediterranean Sea) and P. meteori are not found in this part of the world. A second noteworthy feature is that only three species live in the Caribbean basin (the circum-tropical P. perlucens, and P. nigres- cens, plus the restricted P. turnerae). This situation contrasts sharply with that found in Aspidosiphon, the other burrowers in hard substrata, and this suggests a different evo- lutionary history. In the eastern Pacific one finds the two shallow water Caribbean species just men- tioned, plus P. agassizii (cool water), and P. scolops (warm water); these four also are found throughout the IWP. Phascolosoma stephensoni extends eastward only as far as Hawaii, but also westward to the Mediter- ranean and the eastern Atlantic. Four species do not extend outside the IWP (P. alboli- neatum, P. arcuatum, P. glabrum, and P. pacificum), while P. noduliferum and P. an- nulatum are restricted to the boundary be- tween the Indian and western Pacific Oceans. Acknowledgments An open exchange of information, ideas, and specimens with J. Saiz Salinas, Bilbao, were most important to several stages of this work, especially in resolving the P. granu- latum/stephensoni complex. The continued encouragement and exchange of specimens with S. J. Edmonds, Adelaide, has helped 721 us complete this work. We are indebted to B. P. Haldar, Calcutta, for the opportunity to read an unpublished manuscript. The field work in Cura¢gao was dependent on the co- operation of W. Bakhuis, Carmabi Foun- dation. In Cumana J. Perez at the Univer- sidad de Oriente assisted us in many ways. The visit to the Fundacion Cientifica Los Roques was made possible by the cooper- ation of B. Alverez and her associates. Lynn Cutler, Mountain View, and J. Silverstein, Seattle, provided invaluable assistance in the field and lab, the former also provided the transmission electron micrographs. Correct spelling of species’ names was kind- ly verified by G. Steyskal, Washington, and K. Bart, Clinton, New York, assisted with the scanning electron microscope. Partial fi- nancial support was provided by the Na- tional Science Foundation (BSR 86-15315) and part by our respective institutions. The cooperation of the following persons and institutions in the loan of reference ma- terial and/or providing access to their col- lections was essential to the completion of this project and greatly appreciated: P. Ber- ents (AMSS); E. Easton (BMNH); H. Levi (MCZH); J-L. Justine, (MNHN); G. Hart- wich (MNHU); R. Olerod (NHRS); S. Chambers (RSME); M. Rice (USNM); J. Wiktor (UWMP); R. C. Preece (UZMC); J. Kirkegaard (UZMK); A. Pierrot-Bults (ZMUA); M. Dzwillo (ZMUH); University of Tokyo (ZMUT). Literature Cited Amor, A. 1975. 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Geselschaft der Wissenschaften und der Georg-Augusts-Uni- versitat zu Gottingen, Gottingen 1865:189-209. 1866. Untersuchungen tiber einige ameri- kanische Sipunculiden.— Nachrichten von der K6nigl. Geselschaft der Wissenschaften und der Georg-Augusts-Universitat zu Gottingen, G6t- tingen 1866(14):215-228. 1867. Untersuchungen iiber einige ameri- kanische Sipunculiden. — Zeitschrift flir Wissen- schaftliche Zoologie 17:44—55. Koren, J., & D. C. Danielssen. 1877. The natural history of the Norwegian gephyreae. Pp. 111- 156 in M. Sars, ed., Fauna Littoralis Norvegiae, Druck und Verlag von Johann Dahl, Christi- ania. Lanchester, W. F. 1905a. Gephyrea in the marine fauna of Zanzibar and British East Africa from the collections made by C. Crossland in 1901- 1902.— Proceedings of the Zoological Society of London 1:28-35. . 1905b. On the sipunculids and echiurids col- lected during the “‘Skeat” Expedition to the Ma- lay Peninsula.— Proceedings of the Zoological Society of London 1:35—-41. . 1908. Sipunculoidea of the National Antarc- tic Expedition 1901—1904.— Natural History 4: 1-6. Leroy, P. 1936. Les sipunculiens du Muséum d’His- toire Naturelle de Paris.— Bulletin du Muséum National d’Histoire Naturelle, Paris Serie 2, 8: 423-426. 1942. Sipunculiens d’Indochine.—40e Note Institut Océanographique de |’Indochine 40:1- 51. Leuckart, F.S. 1828. Breves animalium quorundam maxima ex parte marinorum descriptiones. Heidelbergae, A. Osswaldi, 24 pp. Monro, C. A. 1931. Polychaeta, Oligochaeta, Echiu- roidea and Sipunculoidea. — Scientific Report of the Barrier Reef Expedition 4:1037. Murina, V. V. 1964. Report on the sipunculid worms from the coast of South Chinese Sea.—Trudy Instituta Okeanologii Akademii Nauk SSSR 69: 254-270. 1967. On the sipunculid fauna of the littoral of Cuba.—Zoologicheskii Zhurnal 46:35—47. 1968. On the distribution of new sipunculid fauna from Cuba and Gulf of Mexico.—Revue Roumaine de Biologie-Zoologie 13(6):421—423. 1970. Contributions to the fauna of sipun- culid worms from the Aden Gulf.—Vestnik Zoologii 2:65-71. 1971. On the species composition and ecol- ogy of sipunculids of the Red Sea. Pp. 76-88 in VOLUME 103, NUMBER 3 A. O. Kovalevski, ed., Shelf benthos of the Red Sea. Academii Nauk SSSR, Moscow. 1972. Contribution of the sipunculid fauna of the Southern Hemisphere.—Zoologii Insti- tuta Akademii Nauk SSSR 11(19):294-314. 1975. New species of genus Phascolosoma (Sipuncula).— Vestnik Zoologii 197:54—59. . 1977. Marine worms of the Arctic and boreal waters of Europe. Works on the fauna of the USSR. Academy of Science USSR, Leningrad, 283 pp. . 1978. On the sipunculids and priapulids fau- na of the southern Pacific.—Trudy Instituta Okeanologii Academii Nauk SSSR 113:120-131. . 1981. First report of the sea worms Sipuncula and Echiura from littoral Somali.—Vestnik Zoologii 1:11-18. Ocharan, F. J. 1980. Primeros datos sobre los sipun- culides del litoral N.Y. N.W. de Espana (Santan- der, Asturias y Galicia).—Boletin de Ciencias de la Naturaleza I 26:111-119. Ostroumov, A. 1909. Sur les gephyriens du nord de la mer du Japan. —Ezhegodnik Zoologischeskii Muzei 14:319-321. Pourtalés, L. F. de. 1851. On the Gephyrea of the Atlantic coast of the United States.— Proceed- ings of the American Association for the Ad- vancement of Science, 5th meeting 5:39—42. Quatrefages, M. A. de. 1865. Géphyriens Inermes. — Histoire naturelle des Annelés marins et d’eau douce, Paris 2:599-632. Rice, M. E. 1967. A comparative study of the de- velopment of Phascolosoma agassizii, Golfingia pugettensis, and Themiste pyroides with a dis- cussion of developmental patterns in the Sipun- cula.—Ophelia 4:143-171. 1973. Morphology, behavior and histogene- sis of the pelagosphera larva of Phascolosoma agassizii (Sipuncula).—Smithsonian Contribu- tions to Zoology 132:1-51. 1975. Survey of the Sipuncula of the coral and beach rock communities of the Caribbean Sea. Pp. 35—49 in M. E. Rice and M. Todorovic, eds., Proceedings of the International Sympo- sium of Sipuncula and Echiura. Nau¢no Delo Press, Belgrade. 1980. Sipuncula and Echiura. Pp. 490-498 inR. H. Morris, D. P. Abbott, & E. C. Haderlie, eds., Intertidal invertebrates of California, chap- ter 19. Stanford University Press, Palo Alto. . 1985. Description of a wood dwelling sipun- culan, Phascolosoma turnerae, new species.— Proceedings of the Biological Society of Wash- ington 98:54-60. . 1986. Phylum Sipuncula (Peanut worms). Pp. 224-228 in W. Sterrer, ed., Marine fauna and flora of Bermuda. John Wiley & Sons, Inc., New York. 725 ——,, & I. G. MacIntyre. 1979. Distribution of Si- puncula in the coral reef community, Carrie Bow Cay, Belize.—Smithsonian Contributions to Marine Sciences 12:311-320. ——, & A.C. Stephen. 1970. The type specimens of Sipuncula and Echiura described by J. E. Gray and W. Baird in the collections of the British Museum of Natural History.—Bulletin of the British Museum of Natural History 20(2):49- 72. Rippell, E., & F.S. Leuckart. 1828. Neue wirbellose Thiere des rothen Meers. P. 50 in E. Riippell, ed. Atlas zu der Reise im nordlichen Afrika; 1. Zoologie. Senckenbergische Naturforschung Gesellschaft, Frankfurt. Saiz, J. I., A. Rallo, F. Rodriguez, & E. Gutiérrez. 1979. Sipuncula y Echiura de la costa Via- caina.— Actas del Primer Simposio Ibérico de Estudios del Bentos Marino 1:207-221. Saiz Salinas, J. I. 1982. Nuevos datos sobre los si- punculidos de costas espanolas y de mares ad- yacentes.— Actas del Primer Simposio Ibérico de Estudios del Bentos Marino 3:193-201. 1984a. 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Ona new species of Phymosoma, with a synopsis of the genus and some account of its geographical distribution.—Quarterly Journal of Microscopical Science 32:111-126. 1898. Report on the Gephyrean worms col- lected by Mr. J. Stanley Gardiner at Rotuma and Funafuti.—Journal of Zoology 1898:468- 473. 1899a. A report on the Sipunculoidea col- lected by Dr. Willey at the Loyalty Islands and in New Britain. Pp. 151-160 in A. Willey, ed., Zoological results based on material from New Britain, New Guinea, Loyalty Islands and else- where collected 1885-1897 by A. Willey. Uni- versity Press, Cambridge. 1899b. On Gephyrean worms from Christ- mas Island.— Proceedings of the Zoological So- ciety of London 22:54—-57. 1902. Sipunculoidea, with an account of a new genus Lithacrosiphon. Pp. 131-140 in J. S. Gardiner, ed., Fauna and geography of the Mal- dive and Laccadive Archipelagoes. University Press, Cambridge. 1903. Report on the Gephyrea collected by PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Prof. Herdman at Ceylon in 1902.—Ceylon Pearl Oyster Fisheries Supplementary Reports 3:169-— 176. Sluiter, C. P. 1881. Beitrage zu der Kenntnis der Gephyreén aus dem Malayischen Archipel. Zweite Mittheilung. — Natuurkundig Tijdschrift voor Nederlandsich-Indie 41:148-171. 1886. Beitrage zu der Kenntnis der Gephy- reén aus dem Malayischen Archipel.—Natuur- kundig Tijdschrift voor Nederlandisch-Indie 45: 472-517. 1891. Die Evertebraten aus der Sammlung des KG6niglichen naturwissenschaftlichen Ver- eins in Nederlandisch-Indien in Batavia. —Na- tuurkundig Tijdschrift voor Nederlandisch-In- die 50:102-123. 1898. Gephyreen von Sid Africa.—Zoolo- gische Jahrbiicher Abteilung ftir Systematik 11: 442-450. 1902. Die Sipunculiden und Echiuriden der ““Siboga” Expedition; nebst Zusammenstellung der Uberdies aus den indischen Archipel be- kannten Arten.—Siboga-Expeditie, Monogra- phie 25:1-53. Southern, R. 1913. Gephyrea of the coasts of Ire- land.—Scientific Investigations of the Fisheries Branch, Department of Agriculture for Ireland 3:1-46. Spengel, J. W. 1898. Der Name Physcosoma. —Zool- ogischer Anzeiger 21:50. Stephen, A. C. 1941. Sipunculids and Echiurids of the John Murray Expedition to the Red Sea and Indian Ocean 1933-34.—Scientific Reports of the John Murray Expedition 7:401-409. . 1942. The South African intertidal zone and its relation to ocean currents. Notes on the in- tertidal sipunculids of Cape Province and Na- tal.—Annals of the Natal Museum 10(2):245- 256. 1948. Sipunculids.—B.A.N.Z.A.R.E. Re- ports, Series B 5(4):213-220. 1952. Gephyrea. The ““Manihine” Expedi- tion to the Gulf of Aqaba. — Bulletin of the Brit- ish Museum (Natural History) 1:181-182. 1960. Echiuroidea and Sipunculoidea from Senegal, West Africa.—Bulletin de I’Institut Francais d’Afrique Noire 22a(2):512-520. . 1965. Echiura and Sipuncula from the Israel South Red Sea Expedition.—Sea Fisheries Re- search Station Haifa Bulletin 40:79-83. . 1966. A collection of Sipuncula taken on the summit of the Vema Sea-mount, South Atlantic Ocean.— Annals and Magazine of Natural His- tory 9(13):145-148. . 1967. Sipuncula and Echiura from Nouvelle- Caledonie.—Cahiers du Pacifique 10:44—S0. —, & E. B. Cutler. 1969. Ona collection of Si- VOLUME 103, NUMBER 3 puncula, Echiura, and Priapulida from South African waters. — Transactions of the Royal So- ciety of South Africa 38(2):111-121. —., & S.J. Edmonds. 1972. The phyla Sipuncula and Echiura. Trustees British Museum (Natural History), London, 528 pp. —., & J. D. Robertson. 1952. A preliminary re- port on the Echiuridae and Sipunculidae of Zan- zibar.— Proceedings of the Royal Society of Edinburgh 64(4):426-444. Stimpson, W. 1855. Description of some new marine invertebrates. — Proceedings of the Academy of Natural Sciences of Philadelphia 7:390-391. ten Broeke, A. 1925. Westindische Sipunculiden und Echuiriden. — Bijdragen tot de Dierkunde 24:8 1— 96. Tokioka, T. 1953. Invertebrate fauna of the intertidal zone of the Tokara Islands. 111. Echiuroidea and Sipunculoidea.— Publications of the Seto Ma- rine Biological Laboratory 3(2):140. Walter, M. D. 1973. Fressverhalten und Darmin- haltsuntersuchungen bei Sipunculiden.—Hel- golander Wissenschaftliche Meeresuntersu- chungen 25:486-494. Wesenberg-Lund, E. 1930. Priapulidae and Sipun- culidae.— The Danish “‘Ingolf’ Expedition 4(7). 1954a. Priapuloidea, Sipunculoidea and Echiuroidea.— Bulletin Institut royal des Sci- ences naturelles de Belgique 30(16):1-18. 1954b. Sipunculids and Echiurids collected by Mr. G. Ranson in Oceania in 1952.— Bulletin du Muséum National d’Histoire Naturelle, Paris 26:376-384. 1957a. Sipunculoidea and Echiuroidea from West Africa together with a bibliography on Ge- phyrea after 1920.—Bulletin Institut royal des Sciences naturelles de Belgique 33(42):1-24. . 1957b. Sipunculoidea and Echiuroidea from the Red Sea.— Contributions to the Knowledge of the Red Sea No. 3, The Sea Fisheries Re- search Station, Haifa Bulletin 14:1-15. 727 . 1959a. Sipunculoidea and Echiuroidea from tropical West Africa. — Atlantide Report 5:177- 210. . 1959b. Campagne de la ““Calypso”’: Golfe de Guinée 7. Sipunculoidea and Echiuroidea. — Ré- sultats Scientifiques des Campagnes de la ““Ca- lypso” 4:207-217. 1959c. Sipunculoidea and Echiuroidea from Mauritius. — Videnskabelige Meddelelser Dansk fra Naturhistorisk Forening 121:53-73. 1963. South African sipunculids and echiu- roids from coastal waters. — Videnskabelige Meddelelser Dansk fra Naturhistorisk Forening 125:101-146. Wheeler, M. B. 1938. The Sir Joseph Banks Islands. Reports of the expedition of the McCoy Society for field investigations and research. — Proceed- ings of the Royal Society, Victoria 50(1):345. Zavodnik, D. 1975. Contribution to the ecology and zoogeography of North Adriatic sipunculans. Pp. 93-102 in M.E. Rice and M. Todorovic, eds., Proceedings of the International Symposium of Sipuncula and Echiura, volume 1. Naucno Delo Press, Belgrade. —., & V.G. Murina. 1975. Contribution to Si- puncula of North Adriatic insular region. — Rap- ports et procés-verbaux des Reunions Commis- sion Internationale pour 1l’Exploration Scientifique de la Mer-Méditerranée 23(2):127- 128. ——, & 1976. Sipuncula of the region of Rovinj (North Adriatic Sea).—Biosistematika 2(1):79-89. (NJC) Biology Department, Hamilton College, Clinton, New York 13323; (EBC) Biology Department, Utica College of Syr- acuse University, Utica, New York 13502. 728 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Appendix.—Comparative morphological data. Trunk Introvert length length as Number of Longitudinal muscle bands __ Retractor origins as % of Nephnidia length & % Rectal in mm. % trunk hook rings anter. post. trunk dorsal : ventral attached caecum P. agassizii— California 8 188 20 14 20 50 62 75 83 N 10 300 18 17 22 50 70 40 75 N 10 90 14 14 18 50 60 50 60 YC 13 154 14 99 21 46 69 46 50 Y 13 108 16 22 24 54 69 53 57 N 15 107 23 18 21 40 60 40 67 N 18 78 18 17 20 44 67 44 50 Y 19 142 24 26 30 47 68 37 43 N 20 60 18 23 24 55 75 50 50 N 23 109 19 21 25 52 65 57 a We 26 100 19 17 18 50 58 69 72 , IN 26 65 12 22 22 54 69 46 67 *Y; AY 78 24 Dy 22 44 63 37 70 NG 28 71 20 20 21 54 71 48 4 NC 32 56 DD) 22, 21 47 69 47 73 N 34 103 24 25 28 59 74 47 63 N 35 80 20 20 23 43 57 37 ? N 40 125 2B 23 24 45 62 45 72 N 43 70 20 19 22 53 70 47 70 N 45 100 18 18 24 49 60 67 67 N 50 V2 22 21 19 52 68 44 77 N 63 63 23 20 25 48 63 51 69 N 65 69 20 22 23 63 78 57 59 N 69 67 D2 20 25 54 68 41 71 Y P. agassizii— Japan 5 140 16 20 21 50 70 100 80 N 7 143 10 16 18 57 64 71 60 N 7 71 12 20 20 57 71 71 40 N 11 136 37 24 26 55 73 45 40 WC 12 100 16 21 24 42 58 75 50 Y 17 118 15 18 26 47 71 41 57 N 18 117 54 21 24 50 V2, 44 50 N 19 95 16 24 26 53 63 37 86 N 21 129 16 20 25 62 71 33 57 N 22 73 56 Dy Dp 45 64 45 % N 23 87 18 20 25 43 57 57 38 N 24 88 27 26 28 46 67 29 71 N 30 73 20 17 99 53 67 50 67 Y P. albolineatum—West Pacific 7 114 17 Mp) 22 43 57 57 50 N 9 56 18 16 16 56 67 33 67 N 10 50 16 18 20 50 60 40 75 N 11 91 18 16 18 24) 45 36 50 N 12 58 16 16 18 42 50 33 75 N 14 43 18 20 18 50 64 29 75 N 14 50 18 18 20 43 57 50 71 N 1S 67 18 20 14 33 47 20 67 N 17 82 20 20 22 29 41 41 43 Y SS VOLUME 103, NUMBER 3 Appendix. — Continued. 729 Trunk Introvert length length as Number of | Longitudinal muscle bands __ Retractor origins as % of Nephridia length & % Rectal in mm. % trunk hook rings anter. post. trunk dorsal : ventral attached caecum P. granulatum—Spain 18 122 35 18 23 61 V2 56 50 N 24 133 74 20 28 46 67 62 40 N 28 86 44 15 24 43 61 46 54 Y 31 113 68 19 30 48 58 39 67 Y P. nigrescens—1.W.P. 12 267 150 20 26 58 75 67 50 N 14 71 v gy 26 36 5)1/ 50 71 N 15 167 125 24 30 47 60 40 50 Y 21 129 100 22 26 33 52 48 ¥ 2 23 187 200 26 28 39 52 % uf N 26 131 100 28 24 42 54 54 57 N Di, 159 125 20 28 30 44 44 50 N 27 93 100 24 30 33 56 33 ? N 30 117 125 26 28 40 63 60 56 N 30 83 70 30 32 37 53 40 42 N 33 148 200 20 26 30 45 67 55 Y 40 112 150 26 32 40 5)5) 3S) 68 N P. nigrescens— Caribbean 5 160 45 ? ? 40 60 30 67 N 6 217 60 20 28 33 67 50 33 N 7 171 150 18 ? 29 43 71 ? N 10 140 150 16 22 50 70 70 57 N 11 127 70 ? 24 45 64 45 60 N 13 169 133 26 36 31 46 46 50 N 15 133 50 16 24 47 67 47 5)// N 19 142 150 22 32 37 74 47 56 N P. perlucens—Caribbean 7 114 14 20 18 43 a7 Si) 75 N 10 90 17 18 18 40 50 40 50 ays 12 58 16 19 21 42 58 25 67 N 13 62 18 18 23 46 62 38 40 N 14 93 14 18 20 50 57 43 50 N 16 56 18 20 18 44 56 38 83 N 16 88 19 18 20 44 50 44 29 N 17 88 18 18 23 35 41 35 50 N 19 74 18 16 18 37 53 26 60 N 22 82 DD, 18 20 41 50 36 8 N 24 54 20 18 22 38 50 25 67 N 25 96 16 17 23 36 48 40 70 N 26 65 24 19 23 35 42 23 33 YA 29 66 20 16 19 41 Sy 38 73 N 54 43 16 20 20 44 56 37 55 bY? P. scolops—1I.W.P. 5 100 10 16 20 60 80 40 50 Y4 6 67 10 16 18 50 67 50 67 N 7 57 14 16 18 57 86 29 50 We 7 57 10 18 18 29 43 29 50 N 730 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Appendix. — Continued. Trunk Introvert length length as Number of Longitudinal muscle bands __ Retractor origins as % of Nephridia length & % Rectal in mm. % trunk hook rings anter. post. trunk dorsal : ventral attached caecum 8 88 10 16 18 50 62 75 67 Y 9 89 12 20 20 56 67 56 80 Y 10 100 12 18 18 50 60 50 60 N 11 82 12 18 20 55 64 27 67 Y 24 67 15 19 yp) 38 50 33 88 Y 27 93 14 19 21 52 59 37 60 Y P. stephensoni— Hawaii 11 91 16 20 24 36 45 45 80 N 19 105 20 20 D2 63 74 53 70 N 21 62 20 Bee 26 57 71 57 83 N Oy; 114 18 18 26 59 41 64 71 Y BY 96 18 22 20 48 63 48 54 N PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 731-745 CORRECTION OF IJIMA’S (1927) LIST OF RECENT HEXACTINELLID SPONGES (PORIFERA) Henry M. Reiswig Abstract. —Errors in the final species list of Ijima’s posthumously published Siboga Monograph on the Hexactinellida (Porifera) have been the cause of misallocations of species in later publications and disorder of museum collec- tions. The text of the entire monograph has been reviewed in detail in an attempt to provide a faithful reflection of Ijima’s opinions and a corrected list of species as understood by him at the time of his death. A total of 421 forms are recognized, consisting of 381 species and 40 additional subspecies. The taxonomic actions attributable to Ijima in this monograph are summarized in an action list. Isao Tjima’s (1927) monograph, “The Hexactinellida of the Siboga Expedition,” is still regarded by specialists as the most important authority for present taxonomic arrangement of this class of Porifera. It does, however, contain significant, unrecognized hazards for both casual and serious research users. Its prominence is attributable both to its major reorganization of higher taxa, which has served as the basis for later re- views (de Laubenfels 1936) and modifica- tions (e.g., Reid 1957, 1963), and to its in- clusion of a revised list of all extant hexactinellids—the last such attempt. The monograph’s appended “List of Recogniz- ably Known Recent Hexactinellids Ar- ranged Systematically,” pp. 364-377 (here- after referred to as the “‘Final List” or ““FL’’), has served as reference for subsequent hex- actinellid collection reports (e.g., Burton 1954) and as the basic guide for systematic arrangement of specimen collections of most major museums throughout the world. Un- fortunately, the Final List is extensively flawed at lower taxonomic levels due to ed- itorial actions in the preparation of the final publication. As noted in Kaburaki’s preface and the introduction to the monograph, Jjima in- herited responsibility for the Siboga hex- actinellid collection in 1907 after ill health incapacitated F. E. Schulze. Ijima spent the final ten years of his life preparing the re- port, but died in 1921 before completing his revision of the second half of the material — basically the Lyssacinosa. Responsibility for final preparation fell upon Y. Okada, Ji- ma’s assistant, and Max Weber, the editor of the Siboga Series. While the main text is accepted here as a faithful reflection of Iji- ma’s taxonomic opinions, the Final List, presumably assembled by Okada and/or Weber, contradicts Ijima’s text conclusions on many points. The Final List, with its unappreciated errors, continues to be the most widely used portion of the monograph due to omission of an index of scientific names in this work. Even Burton (1928), in his careful taxonom- ic review for the Zoological Record, failed to detect several new genera erected by IJji- ma. These errors continue to be perpetuated in modern literature (e.g., Hartman 1982, Koltun 1967, Reiswig 1979, Tuzet 1973), hence correction of the list is urgently need- ed. Ijima, a major figure in hexactinellid systematics, ranking second only to F. E. Schulze in terms of total contributions to the field, deserves better recognition for his last major accomplishment than appearing 732 to be the source of taxonomic confusion. Museum collection managers likewise need an accurate scheme for arranging specimens without the necessity of scrutinizing the fragmented literature of this class. A cor- rected version of the Final List is also re- quired for eventual development of a cur- rent list of the recent hexactinellids and, ultimately, inclusion of fossil groupings into a single classification. Because of the numerous contradictions to Ijima’s text, the details presented in the Final List of the Siboga Monograph cannot be attributed to Ijima. The transfers and synonomies implied by that list should be ignored where these conflict with state- ments or implications of Ijima’s text. In preparation of the corrected list, my pri- mary aim has been to reflect Ijima’s con- cepts as faithfully as possible. Simple errors of spelling and statements of authority as presented in the text have been corrected where these have been detected. Names for typical subspecies have been added where Ijima’s recognition of atypic subspecies re- quires formal recognition of the reference taxon. Clear statements of uncertainty by Ijima on allocation of specific taxa are in- dicated by symbol. Text contradictions or absence of clear statement of allocation by Ijima were resolved by reviewing his pre- vious publications and, occasionally, by ed- ucated guess. These points are clearly iden- tified in the annotations. Species which Ijima either did not mention, or mentioned only in trivial context (lacking any implication of recognition), are identified by symbol. All transfers and synonomies attributed to IJji- ma in this work are included in an Action List, with the cautionary note that several of these may have been previously suggest- ed, but without conviction of formal intent. A complete index of generic, specific and subspecific names contained in the Siboga Monograph is available from the author upon request. Publication date of the Siboga Monograph is often erroneously cited as PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1926; the publisher, E. J. Brill, Leiden, has advised me that their booklet advertising the Siboga Series clearly states the date of availability as January, 1927 (H. Reerink, pers. comm.). Corrected version of Ijima’s 1927 “LIST OF RECOGNIZABLY KNOWN RE- CENT HEXACTINELLIDS ARRANGED SYSTEMATICALLY.” Ijima’s overall systematic arrangement is retained, but in the absence of data on phy- logenetic relationships at lower levels, species are listed alphabetically under gen- era and subgenera. All page numbers refer to Ijima, 1927. An asterisk (*) before the species number identifies species or sub- species either not mentioned or, if men- tioned, not clearly allocated in the text; Iji- ma’s concept of the status of these names, at the time of his death in 1921, cannot be assessed. Taxa for which Ijima explicitly or implicitly expressed uncertainty in recog- nition or allocation are indicated by a ques- tion mark after the species number. Brack- ets identify nominate subspecies which are implied but not formed in the text. They are required by recognition of non-nomi- nate subspecies by ljima, and, although formed here by HMR, their status is implied by Ijima. Annotations following the list are indicated by parentheses. Suborder I AMPHIDISCOPHORA Schulze, 1899 Fam. I PHERONEMATIDAE Gray, 1870 (emended) Gen. 1 Pheronema Leidy, 1868; p. 7, 9 *1 annae Leidy, 1868 *2 _carpenteri (Thomson, 1869) 3 echinatum Yjima, 1927; p. 9, 21 4 giganteum Schulze, 1886; p. 9, 10 5 ?gigas (Schulze, 1886); p. 9, (a) 6 globosum Schulze, 1886; p. 21 *7 grayi Kent, 1869 *8 hemisphaericum Gray, 1873; p. 8 9 raphanus Schulze, 1894; p. 21 10 weberi Ijima, 1927; p. 9, 17 VOLUME 103, NUMBER 3 733 Gen. 2 Poliopogon Thomson, 1873; p. 7,9 Subgen. 3 Coscinonema \jima, 1927; p. 50 1 amadou Thomson 1873; p. 9 Gen. 3. Sericolophus Yjima, (1894) 1901; p. 9, 25, (b) 1 reflexus (jima, 1894); p. 25, 26 Gen. 4 Semperella Gray, 1868; p. 9, 28 1 cucumis Schulze, 1895; p. 28 2 schultzei (Semper, 1868); p. 28 3 similis Yjima, 1927; p. 28 4 spicifera Schulze, 1904; p. 28 5 stomata Yjima, 1896; p. 28 Gen. 5 Platylistrum Schulze, 1904; p. 7, 9 1 platessa Schulze, 1904; p. 25 Fam. II MONORHAPHIDIDAE Jjima, 1927 Gen. 1 Monorhaphis Schulze, 1904; p. 36, 37 1 chuni Schulze, 1904; p. 37 2 dives Schulze, 1904; p. 37 3 ?fruticosa (Schulze, 1893); p. 37, (c) Fam. III HYALONEMATIDAE Gray, 1857, emend. Gen. 1 Chalaronema Ijima, 1927; p. 41, 43 1 sibogae \jima, 1927; p. 43 Gen. 2 Hyalonema Gray, 1832; p. 43, 49 Subgen. | Euhyalonema Ijima, 1927; p. 50 intermedium Ijima, 1927; p. 50, 55 keianum \jima, 1927; p. 50, 58 pellucidum (Ijima, 1894); p. 50, 52 proximum (Schulze, 1904); p. 50 sieboldi (Gray, 1835); p. 50 aA BWN Subgen. 2. Pteronema Yjima, 1927; p. 50 1 aculeatum (Schulze, 1894); p. 50 cebuense (Higgin, 1875); p. 50 globus (Schulze, 1886); p. 50 heideri (Schulze, 1894); p. 50 polycaulum (Lendenfeld, 1915); p. 50, (c) 6 topsenti Ijima, 1927; p. 50, 61 nA BW bd 1 conus (Schulze, 1886); p. 51 2 elegans (Schulze, 1886); p. 51 3. ?gracile (Schulze, 1886); p. 51 heymonsi Schulze, 1895; p. 51, (d) indicum andamanense (Schulze, 1895); p. 51, (e) 6 indicum laccadivense (Schulze, 1895); p. 51, (e) 7 kenti (Schmidt, 1880); p. 51 8 kirkpatricki Yjima, 1927; p. 51, 68 9 lamella (Schulze, 1900); p. 51 10 pateriferum (Wilson, 1904); p. 51, 52 11 schmidti (Schulze, 1899); p. 51 12 toxeres (Thomson, 1877); p. 51, (c) nN & Subgen. 4 Cyliconema ljima, 1927; p. 50, Sl 1 apertum apertum (Schulze, 1886); p. 51, (f) 2 apertum maehrenthali (Schulze, 1895); p. 51, 72, (b, g) 3. coniforme (Schulze, 1904); p. 51 4 drygalskii (Schulze, 1910); p. 51 5 globiferum (Schulze, 1904); p. 51 6 infundibulum (Topsent, 1896); p. 51 7 keiense Yjima, 1927; p. 51, 90 8 martabanense (Schulze, 1900); p. 51, (h) 9 molle (Schulze, 1904); p. 51 10 nicobaricum (Schulze, 1904); p. 51, (i) 11 ovatum (jima, 1895); p. 51 12 pirum (Schulze, 1895); p. 51 13. rapa (Schulze, 1900); p. 51, (b) 14 simile (Schulze, 1904); p. 51 15 somalicum (Schulze, 1904); p. 51, (i) 16 tenerum (Schulze, 1886); p. 51 17. thomsonis (Marshall, 1875); p. 51, (c) 18 timorense Ijima, 1927; p. 51, 88 19 tulipa (Schulze, 1904); p. 51 20 valdiviae (Schulze, 1904); p. 51 Subgen. 5 Paradisconema Ijima, 1927; p. 30), SZ 1 alcocki (Schulze, 1895); p. 52 734 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2 investigatoris (Schulze, 1900); p. 52 3 vosmaeri Yjima, 1927; p. 52, 93 Subgen. 6 Leptonema Lendenfeld, 1915; p. S50; SZ acuferum (Schulze, 1893); p. 52 campanula Lendenfeld, 1915; p. 52 divergens (Schulze, 1886); p. 52 flagelliferum Yjima, 1927; p. 52, 97 lusitanicum (B. d. Bocage, 1864); p. Sy 6 ovuliferum (Schulze, 1899); p. 52 7 solutum (Schulze, 1904); p. 52 8 urna (Schulze, 1904); p. 52 aABWN Subgen. 7 Thamnonema Ijima, 1927; p. 50, 52 1 thamnophorum \jima, 1927; p. 52, 99 Subgen. 8 Phialonema Lendenfeld, 1915; p. 50, 52 1 brevancora Lendenfeld, 1915; p. 52 Subgen. 9 Prionema Lendenfeld, 1915; p. 50, 52 1 agujanum Lendenfeld, 1915; p. 52, 53 2 azuerone Lendenfeld, 1915; p. 53 3 crassum Lendenfeld, 1915; p. 53 4 fimbriatum Lendenfeld, 1915; p. 53 5 pinulifusum Lendenfeld, 1915; p. 53 6 poculum (Schulze, 1886); p. 53 7 spinosum Lendenfeld, 1915; p. 53 8 validum (Schulze, 1904); p. 53 Subgen. 10 Corynonema ljima, 1927; p. 50, 53 calix (Schulze, 1904); p. 53 clathratum (Ijima, 1895); p. 53 clavigerum (Schulze, 1886); p. 53 ?cupressiferum (Schulze, 1893); p. 53, (h) 5 ?depressum (Schulze, 1886); p. 53 6 grandancora (Lendenfeld, 1915); p. 53 7 hercules (Schulze, 1899); p. 53 8 masoni (Schulze, 1895); p. 53 BRWNre 9 owstoni (ljima, 1894); p. 53 10 placuna (Lendenfeld, 1915); p. 53 11 populiferum (Schulze, 1899); p. 53 12 rotundum \jima, 1927; p. 53, 102 13 tenuifusum (Lendenfeld, 1915); p. 53 14 tylostylum (Lendenfeld, 1915); p. 53 15 weltneri (Schulze, 1895); p. 53 Subgen. 11 Onconema Ijima, 1927; p. 50, 53 1 agassizi (Lendenfeld, 1915); p. 53 2 obtusum gracile (Lendenfeld, 1915); p. 53, (h,i) 3 obtusum robustum (Lendenfeld, 1915); p. 53, (h,1) Subgen. 12 Oonema Lendenfeld, 1915 (emended); p. 50, 53 1 aequatoriale (Lendenfeld, 1915); p. 54, (k) 2 bianchoratum (Wilson, 1904); p. 53, 54 3 crassipinulum Lendenfeld, 1915; p. 54 4 densum Lendenfeld, 1915; p. 54 5 geminatum (Lendenfeld, 1915); p. 54, (k) 6 henshawi Lendenfeld, 1915; p. 54 7 pedunculatum (Wilson, 1904); p. 54 8 robustum (Schulze, 1886); p. 54 9 sequoia Lendenfeld, 1915; p. 54 QO umbraculum (Lendenfeld, 1915); p. 54, (k) Gen. 3. Compsocalyx Schulze, 1904; p. 41, 43 *1 gibberosa Schulze, 1904 Gen. 4 Lophophysema Schulze, 1900; p. 41, 43 *] inflatum Schulze, 1900 Suborder II HEXASTEROPHORA Schulze, 1899 Tribe A HEXACTINOSA Schrammen, 1910-1912 Subtribe a CLAVULARIA Schulze, 1886 Fam. I FARREIDAE Schulze, 1886 Gen. | Farrea Bowerbank, 1862; p. 130 VOLUME 103, NUMBER 3 1 aculeata Schulze, 1899; p. 131 2 convolvulus Schulze, 1899; p. 131, 158 hanitschi Yjima, 1927; p. 151 lendenfeldi Ijima, 1927; p. 159 mexicana Wilson, 1904; p. 160 nodulosa Vjima, 1927; p. 154 occa claviformis Wilson, 1904; p. 132, 140 8 occa clavigera (Schulze, 1887); p. 131, 37 9 occa cuspidata Ijima, 1927; p. 145 10 occa erecta Wjima, 1927; p. 132 11 occa foliascens Topsent, 1906; p. 162 *12 occa laminaris Topsent, 1904 13. occa mammillata ljima, 1927; p. 142 14 occa occa Bowerbank, 1862, Carter, 1885; p. 131, (b) 15. occa ouwensi Ijima, 1927; p. 148 *16 occa scutella Lendenfeld, 1915 17 occa subclavigera jima, 1927; p. 140 18 sollasi Schulze, 1886; p. 153 19 spirifera Ijima, 1927; p. 156 20 vosmaeri Schulze, 1886; p. 131, 154 21 weltneri Topsent, 1901; p. 153 Gen. 2 Lonchiphora \jima, 1927; p. 130, (1) 1 inversa Yjima, 1927; p. 162, (1) NDA BB W Gen. 3 Claviscopulia Schulze, 1899; p. 126, 130, (h) *1 intermedia Schulze, 1899 Gen. 4 Sarostegia Topsent, 1904; p. 130 *1 oculata Topsent, 1904; p. 124 Subtribe b. SCOPULARIA Schulze, 1886 Fam. I EURETIDAE Schulze, 1886 Gen. 1 Eurete Semper, 1868; p. 165, 166, (b, m) 1 bowerbanki Schulze, 1886; p. 165, 166 freelandi \jima, 1927; p. 167, 176 marshalli Schulze, 1886; p. 166, 167 schmidti kampeni Yjima, 1927; p. 173 schmidti schmidti Schulze, 1886, Iji- ma, 1927; p. 166, 171, (f) 6 schmidti treubi lima, 1927; p. 170 On BWN 735 7 simplicissimum Semper, 1868; p. 166 8 spinosum Lendenfeld, 1915; p. 166 9 trachydocus Yjima, 1927; p. 168 Gen. 2 Pararete Ijima, 1927; p. 165, 178 1 baliense Yjima, 1927; p. 178, 187, (n) 2 carteri (Schulze, 1886); p. 166, 178 3 farreopsis farreopsis (Carter, 1877) Tjima, 1927; p. 166, 178 4 farreopsis fragiferum Ijima, 1927; p. 182 5 farreopsis jakosalemi Ijima, 1927; p. 178, 185 6 farreopsis subglobosum Ijima, 1927: p. 178, 183 7 freeri \jima, 1927; p. 178, 190 8 gerlachei(Topsent, 1901); p. 166, 178 9 kangeanganum Yjima, 1927; p. 178, 189, (n) 10 semperi (Schulze, 1886); p. 178, 193 Gen. 3 Pleurochorium Schrammen, 1910- 1912; p. 165, 195 1 annandalei (Kirkpatrick, 1908); p. 166, 196, (h) 2 cornutum Yjima, 1927; p. 196, 197 Gen. 4 Periphragella Marshall, 1875; p. 165, 203 1 challengeri Yjima, 1927; p. 204 2 elisae [elisae| Marshall, 1875; p. 165, 204, (0) 3. elisae japonica Yjima, 1927; p. 204, (p) 4 irregularis Ijima, 1927; p. 205, 208 5 lusitanica Topsent, 1890; p. 205 6 parva Tjima, 1927; p. 205 Gen. 5 Lefroyella Thomson, 1877; p. 165, 210 1 ceramensis ljima, 1927; p. 210 2 decora Thomson, 1877; p. 165, 210 Gen. 6 Myliusia Gray, 1859; p. 165, 213 callocyathus Gray, 1859; p. 165, 215 conica (Schmidt, 1880); p. 214 ?subglobosa (Gray, 1867); p. 214, (b) verrucosa Tjima, 1927; p. 214, 217 BRWNr- 736 Gen. 7 Margaritella Schmidt, 1880; p. 113, 165, (h) 1 coeloptychioides Schmidt, 1880; p. 165 Gen. 8 Chonelasma Schulze, 1886; p. 118, 165, (q) *1 ijimai Topsent, 1901 lamella {lamella Schulze, 1886; p. 165, (0) *3 lamella choanoides Schulze & Kirk- patrick, 1910 Gen. 9 Ptychodesia Schrammen, 1910- 1912; p. 165 1 doederleini (Schulze, 1886); p. 165 Gen. 10 Bathyxiphus Schulze, 1899; p. 165 1 subtilis Schulze, 1899; p. 165 Gen. 11 Heterorete Dendy, 1916; p. 165 1 pulchrum Dendy, 1916; p. 165, (h) Gen. 12 Dactylocalyx Stutchbury, 1841; p. 114, 166, (r) 1 ingalli (Bowerbank, 1869); p. 214 ?patella Schulze, 1887; p. 231 *3 potatorum Schmidt, 1880 4 pumiceus Stutchbury, 1841; p. 166, 214 Gen. 13 Iphiteon Bowerbank, 1869; p. 166, (h) 1 panicea Bowerbank, 1869; p. 166 Gen. 14 Conorete Ijima, 1927; p. 165, 166, (f, m) 1 erectum erectum (Schulze, 1899); p. 165, 166 *2 erectum tubuliferum (Wilson, 1904) *3 erectum gracile (Wilson, 1904) *4 erectum forma D (Lendenfeld, 1915) *5 mucronatum (Wilson, 1904) Gen. 15 Gymnorete Ijima, 1927; p. 165, 166, (f, m, s) 1 alicei (Topsent, 1901); p. 165, 166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fam. II TRETODICTYIDAE Schulze, 1886 1886 Gen. 1 Tretodictyum (Schulze, 1886); p. 219, 220, (u) 1 2abyrinthicum (Wilson, 1904); p. 231, (t) 2 pumicosum Tjima, 1927; p. 220, 229 3 schrammeni \jima, 1927; p. 220, 224 4 tubulosum (Schulze, 1886); p. 220, DI Gen. 2 Hexactinella Carter, 1885; p. 219, 231 grimaldii Topsent, 1890; p. 231, 234 lata (Schulze, 1886); p. 231, 235 lingua Yjima, 1927; p. 231, 242 rugosa ljima, 1927; p. 231, 237 ?spongiosa Ijima, 1927; p. 231, 246, (u) 6 ventilabrum Carter, 1885; p. 231, 232 7 vermiculosa Ijima, 1927; p. 231, 239 Gen. 3 Auloplax Schulze, 1904; p. 219, 231 1 auricularis Schulze, 1904; p. 219, 231 ?filholi (Topsent, 1904); p. 231, 244, (v) Gen. 4 Cyrtaulon Schulze, 1887; p. 219 1 sigsbeei (Schmidt, 1880); p. 6 2 solutus Schulze, 1886; p. 219 Gen. 5 Tretocalyx Schulze, 1900; p. 219 aA BWN N 1 monticularis (Lendenfeld, 1915); p. 231 2 polae Schulze, 1900; p. 219 Gen. 6 Sclerothamnopsis Wilson,1904; p. 219, 248 1 compressa Wilson, 1904, p. 219, 248 2 schulzei Yjima, 1927; p. 248, 249 Gen. 7 Sclerothamnus Marshall, 1875; p. 219, 253 1 clausi Marshall, 1875; p. 219, 253 Gen. 8 Psilocalyx Yjima, 1927; p. 219, 265 1 wilsoni Yjima, 1927; p. 219, 265 VOLUME 103, NUMBER 3 Gen. 9 Anomochone Tjima, 1927; p. 219, 268 1 expansa \jima, 1927; p. 219, 269 2 globosa \jima, 1927; p. 272 Fam. IIIT AULOCALYCIDAE IJjima, 1927 Gen. | Aulocalyx Schulze, 1886; p. 274, 276 1 irregularis Schulze, 1886; p. 274 2 serialis Dendy, 1916; p. 274 Gen. 2 Rhabdodictyum (Schmidt, 1880); p. 274, 276 1 delicatum (Schmidt, 1880); p. 274, Dad) 2 kurense \jima, 1927; p. 274, 277 Gen. 3 Euryplegma Schulze, 1886; p. 274, 276 1 auriculare Schulze, 1886; p. 274 Gen. 4 Fieldingia Kent, 1870; p. 274, 276 1 Jagettoides Kent, 1870; p. 274 Gen. 5 Tretopleura Ijima, 1927; p. 276, 280 1 candelabrum \jima, 1927; p. 274, 280 Fam. IV CRATICULARIDAE Rauff, 1893 Gen. 1 Tretorete Ijima, 1927; p. 298 1 incertum Yjima, 1927; p. 298 Fam. V APHROCALLISTIDAE Gray, 1867 Gen. 1 Aphrocallistes Gray, 1858; p. 284, 286, (w) 1 beatrix beatrix Gray, 1858, Tjima, 1916; p. 286, 297 2 beatrix orientalis Ijima, 1916; p. 287, 288 3 vastus Schulze, 1886; p. 286, 287 Gen. 2. Heterochone Ijima, 1927; p. 284, 285, (f, a) 1 calyx (Schulze, 1886); p. 118, 284 2 ?hamata (Schulze, 1886); p. 284, (h) 3 tenera (Schulze, 1899); p. 284, (h) 737 Tribe BLYCHNISCOSA Schrammen, 1903 Fam. I AULOCYSTIDAE Schulze, 1886, (b) Gen. 1 Aulocystis Schulze, 1886; p. 302 1 grayi grayi (Bowerbank, 1869); p. 302, 304 2 grayi polae \jima, 1927; p. 303, 304 3 zitteli sibogae \jima, 1927; p. 304, 305 4 zitteli zitteli (Marshall & Meyer, 1877) Ijima, 1927; p. 302, 304 Incertae Sedis (Aulocystidae) 1 ?superstes Schmidt, 1880, (Cystis- pongia); p. 302, (f) 2 ?mixtum Schmidt, 1880, (Diplacod- ium); p. 302, (f) 3 anterna Schmidt, 1880, (Sclero- plegma); p. 214, 302, (f) Fam. If DIAPLEURIDAE Jjima, 1927 Gen. | Diapleura Ijima, 1927; p. 314 1 maasi \jima, 1927; p. 314 Tribe C LYSSACINOSA Ijima, 1927 Fam. I LEUCOPSACADIDAE Jjima, 1903, (h) Gen. 1 Leucopsacas Ijima, (1898) 1903; p. 320 1 orthodocus Tjima, 1898; p. 321 scoliodocus {scoliodocus] Yjima, 1898; p. 320, 321, (0) 3 scoliodocus retroscissa Topsent, 1904; p. 321, (h Gen. 2 Placopegma Schulze, 1896; p. 320 *1 solutum Schulze, 1896; p. 105, 319 Gen. 3 Caulocalyx Schulze, 1886; p. 320 *1 tenera Schulze, 1886, (h) Gen. 4 Chaunangium Schulze, 1904; p. 320 *1 crater Schulze, 1904 Gen. 5 Chaunoplectella Ijima, (1896) 1903; p. 320, 323 1 cavernosa \jima, 1896; p. 323, 324 738 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2 spinifera \jima, 1903; p. 323 3 stelleta Ijima, 1927; p. 320, 323 Fam. II EUPLECTELLIDAE (Gray, 1867) Ijima, 1903, (b) Subfam. a Euplectellinae Ijima, 1903 Gen. 1 Euplectella Owen, 1841; p. 327 *1 aspera Schulze, 1895 2 aspergillum Owen, 1841; p. 106, 329 *3 crassistellata Schulze, 1886 *4 cucumer Owen, 1857 *5 curvistellata ljima, 1901 6 imperialis ljima, 1894; p. 105, 327 *7 jovis Schmidt, 1880 8 marshalli Yjima, 1895; p. 328, 334 *Q nobilis Schulze, 1904 *10 nodosa Schulze, 1886 *11 oweni Herklots & Marshall, 1868 12 regalis Schulze, 1900; p. 327, 334 *13 simplex Schulze, 1895 *14 suberea Thomson, 1876 15 timorensis Yjima, 1927; p. 327 Gen. 2 Holascus Schulze, 1886; p. 327 *1 edwardsi Lendenfeld, 1915 *2 fibulatus Schulze, 1886 *3 obesus Schulze, 1904 *4 polejaevi Schulze, 1886 *5 ridleyi Schulze, 1886 *6 robustus Schulze, 1895 *7 — stellatus Schulze, 1887 *8 tener Schulze, 1895 *Q tenuis Schulze, 1904 *10 undulatus Schulze, 1899 Gen. 3. Malacosaccus Schulze, 1886; p. 327 *1 coatsi Topsent, 1910; p. 105 *2 floricomatus Topsent, 1901 *3 pedunculatus Topsent, 1910 *4 unguiculatus Schulze, 1886 *5 vastus Schulze, 1886 Gen. 4 Docosaccus Topsent, 1910; p. 327 *1 ancoratus Topsent, 1910; p. 105 Gen. 5. Acoelocalyx Topsent, 1910; p. 327 *1 brucei Topsent, 1910; p. 105 Gen. 6 Holascella Lendenfeld, 1915; p. 327 *1 ancorata Lendenfeld, 1915 *2 euonyx Lendenfeld, 1915 *3 taraxacum Lendenfeld, 1915 Subfam. b Corbitellinae Ijima, 1903 Gen. 1 Regadrella Schmidt, 1880; p. 334, 335 1 cylindrica Yjima, 1927; p. 335 2 decora Schulze, 1900; p. 335 3. delicata Wilson, 1904; p. 337 4 komeyamai Ijima, 1901; p. 337 5 okinoseana \jima, 1896; p. 335 6 phoenix Schmidt, 1880; p. 337 Gen. 2 Corbitella Gray, 1867; p. 344 *1 elegans (Marshall, 1875) *2 pulchra (Schulze, 1887) *3 speciosa (Quoy & Gaimard, 1833); p. 106 Gen. 3 Heterotella Gray, 1867; p. 344 *1_ corbicula (Bowerbank, 1858) Gen. 4 Walteria Schulze, 1886; p. 343 1 flemmingi Schulze, 1886; p. 343 leuckarti Yjima, 1896; p. 343 Gen. 5 Dictyaulus Schulze, 1895 *1 elegans Schulze, 1895 Gen. 6 Dictyocalyx Schulze, 1886 *1 gracilis Schulze, 1886; p. 277 Gen. 7 Hertwigia Schmidt, 1880 *1 falcifera Schmidt, 1880 Gen. 8 Trachycaulus Schulze, 1886 *1 gurlitti Schulze, 1886 Gen. 9 Saccocalyx Schulze, 1895; p. 276 *1 pedunculata Schulze, 1895; p. 279, (h) Gen. 10 Rhabdopectella Schmidt, 1880; p. 276 *] tintinnus Schmidt, 1880; p. 279 VOLUME 103, NUMBER 3 Gen. 11 Hyalostylus Schulze, 1886 *1 dives Schulze, 1886 Gen. 12 Bolosoma (ljima, 1903) Schulze, 1904; p. 334, 340, (b) 1 cavum \jima, 1927; p. 340 2 paradictyum (Ijima, 1903); p. 341 Fam. IIT CAULOPHACIDAE Ijima, 1903 Gen. | Caulophacus Schulze, 1886, p. 345, 346 *1 agassizi Schulze, 1899 *2 antarcticus Schulze & Kirkpatrick, 1910 *3 arcticus (Hansen, 1885) *4 elegans Schulze, 1886 *5 instabilis Topsent, 1910 *6 Jatus Schulze, 1886 7 oviformis (Schulze, 1886); p. 345 *8 pipetta (Schulze, 1886) *Q schulzei Wilson, 1904 *10 scotiae Topsent, 1910 *11 valdiviae Schulze, 1904; p. 106 Gen. 2. Caulophacella Lendenfeld, 1915; p. 345, 346 *1 tenuis Lendenfeld, 1915 Gen. 3 Sympagella Schmidt, 1870; p. 345, 346, (h) *1 anomala Tyima, 1903 *2 cantharellus helix Lendenfeld, 1915; p. 345, (x) *3 cantharellus megonychia Lenden- feld, 1915; p. 345, (x) *4 cantharellus simplex Lendenfeld, 1915; p. 345, (x) 5 gracilis (Schulze, 1903); p. 345 *6 johnstoni (Schulze, 1903) 7 nux Schmidt, 1870; p. 106 Gen. 4 Caulodiscus \jima, 1927; p. 345, 346, (f) 1 /otifolium (jima, 1903); p. 345, (fp Fam. IV ROSSELLIDAE (Schulze, 1886), Tjima, 1903; (b) Subfam. a Lanuginellinae Schulze, 1897 739 Gen. 1 Lanuginella Schmidt, 1869; p. 352 1 pupa Schmidt, 1870; p. 352 Gen. 2 Lanugonychia Lendenfeld, 1915; p. 347 1 flabellum Lendenfeld, 1915; p. 347 Gen. 3 Calycosoma Schulze, 1899; p. 345 1 validum Schulze, 1896; p. 345 Gen. 4 Mellonympha Schulze, 1897; p. 105, 319 *1 yvelata (Thomson, 1873) Gen. 5 Lophocalyx Schulze, 1887; p. 348 1 philippinensis (Gray, 1872); p. 348, S52) 2 spinosa Schulze, 1900; p. 352 3. suluana Yjima, 1927; p. 348, 352, (h) Subfam. b Rossellinae Schulze, 1897 Gen. 1 Rossella Carter, 1872; p. 105, 319 *1 antarctica antarctica Carter, 1872; p. 105 *2 antarctica gaussi Schulze & Kirkpat- rick, 1910 *3 anarctica solida Kirkpatrick, 1907 *4 dubia (Schulze, 1886) *5 fibulata Schulze & Kirkpatrick, 1910 *6 gaussi Schulze & Kirkpatrick, 1910 *7 lychnophora Schulze & Kirkpatrick, 1910 *8 mixta Schulze & Kirkpatrick, 1910 *9 nodastrella Topsent, 1915 *10 podagrosa Kirkpatrick, 1907 *11 racovitzae hexactinophila Kirkpat- rick, 1910 racovitzae microdiscina Topsent, 1916 racovitzae minuta Schulze & Kirk- patrick, 1910 racovitzae [racovitzae] Topsent, 1901; (0, y) racovitzae racovitzae Schulze & Kirkpatrick, 1910; (b, y) Gen. 2 Aulorossella Kirkpatrick, 1907; p. 355) ale #13 *14 *15 740 *1 aperta Topsent, 1916 *2 gaini Topsent, 1916 *3 Jaevis Kirkpatrick, 1907 *4 longstaffi Kirkpatrick, 1907 *5 pilosa Kirkpatrick, 1907 *6 vanhoeffeni armata Schulze & Kirk- patrick, 1910 *7 yanhoeffeni [vanhoeffeni] Schulze & Kirkpatrick, 1910; (0) Gen. 3 Gymnorossella Topsent, 1916; p. 353 *1 inermis Topsent, 1916 *2 nuda (Topsent, 1901) Gen. 4 Scyphidium Schulze, 1900 *1 chilense Yjima, 1904, 1927; (b) *2 longispina (Ijima, 1896) *3 namiyel (Ijima, 1898) *4 septentrionale Schulze, 1900 Gen. 5 Vitrollula Ijima, 1898 *1 fertilis Ijima, 1898 Gen. 6 Schaudinnia Schulze, 1900; p. 105, 319 *1 arctica Schulze, 1900; p. 105 Gen. 7 Crateromorpha Gray, 1872 *1 corrugata Ijima, 1898 *2 meyeri [meyeri] Gray, 1872 *3 meyeri rugosa ljima, 1898 *4 meyeri tuberosa Yjima, 1898 *S pachyactina jima, 1898 *6 thierfelderi Schulze, 1886 *7 tumida Schulze, 1886 Gen. 8 Hyalascus Ijima, 1896 *1 giganteus Ijima, 1898 *2 hodgsoni Kirkpatrick, 1907 *3 sagamiensis Ijima, 1896 *4 similis Yjima, 1904 Gen. 9 Anaulosoma Kirkpatrick, 1907; p. 353 *1 schulzei Kirkpatrick, 1907 Gen. 10 Aulochone Schulze, 1886 *1 cylindrica Schulze, 1886 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON *2 lankesteri (Kirkpatrick, 1902) *3 lilium Schulze, 1886 . 11 Aulosaccus Yjima, 1896 *1 imae (Schulze, 1899) *2 misukurii ima, 1898 *3 schulzei Ijima, 1896 . 12 Asconema Kent, 1870 *] setubalense Kent, 1870 Gen. 13 Trichasterina Schulze, 1900; p. 105, 319 1 borealis Schulze, 1900; p. 105 *2 sagittaria Topsent, 1913 Gen. 14 Aphorme Schulze, 1899 *1 horrida Schulze, 1899 Gen. 15 Bathydorus Schulze, 1886; p. 353 baculifer Schulze, 1886; p. 355 fimbriatus Schulze, 1886; p. 355 laevis [laevis] Schulze, 1895; p. 355 laevis ciliatus Topsent, 1910 laevis spinosus Wilson, 1904 pedunculatus Yjima, 1927; p. 353 spinosissimus Lendenfeld, 1915 spinosus Schulze, 1886; p. 355 stellatus Schulze, 1886; p. 355 uncifer Schulze, 1899; p. 355 Ge Gp * OoOoWoO NDNA BWNYN eH — Gen. 353 *1 ijimai Kirkpatrick, 1907 16 Anoxycalyx Kirkpatrick, 1907; p. Gen. 17 Scolymastra Topsent, 1916; p. 353 *1 joubini Topsent, 1916 Subfam. c. Acanthascinae Schulze, 1897; (b) Gen. 1 Acanthascus Schulze, 1886; p. 355 *1 alani Yjima, 1898 *2 cactus Schulze, 1886 *3 platei Schulze, 1899 Gen. 2 Staurocalyptus Tjima, 1897; p. 355 1 affinis Yjima, 1904; p. 357 2 celebesianus ljima 1927; p. 355, (h) 3 dowlingi (Lambe, 1893); p. 357 VOLUME 103, NUMBER 3 *4 entacanthus Ijima, 1904 *5 fasciculatus Schulze, 1899 6 glaber \jima, 1897; p. 357 Ay) 8 heteractinus Ijima, 1897; p. 357 9 microchetus jima, 1898; p. 357 hamatus Lendenfeld, 1915 *10 pleorhaphides Yjima, 1897 *11 roeperi (Schulze, 1886) *12 solidus Schulze, 1899 *13 tubulosus Yjima, 1904 Gen. 3 Rhabdocalyptus Schulze, 1886; p. 355 *1 asper Schulze, 1899 *2 australis Topsent, 1901 *3 baculifer Schulze, 1904 *4 capillatus Ijima, 1897 *5 dawsoni (Lambe, 1892); (h) 6 mirabilis Schulze, 1899; p. 107, 108 *7 mollis Schulze, 1886 *8 nodulosus Schulze, 1899 *9 plumodigitatus Kirkpatrick, 1901 *10 tener Schulze, 1899 *11 tenuis (Schulze, 1899) *12 wunguiculatus \jima, 1904 13 victor Ijima, 1897; p. 325 Annotations (a) Ijima suggested transfer of Poliopogon (b) (c) (d — (e) gigas to Pheronema or to a new un- named genus. Authority corrected—name(s), date or punctuation. Suffix differs from FL or text; form cho- sen to conform either to text usage or original spelling where recent ICZN rule ammendment permits. Erroneously treated in FL as a synonym of H. indicum and misspelled as “‘hey- mousi.” Ijima makes no mention of the two sub- species H. (C.) indicum andamanense (Schulze, 1895) and H. (C.) indicum lac- cadivense (Schulze, 1895); the species and the two subspecies are erroneously given separate (duplicate) entries in the FL. Distinction of the two subspecies is retained as common for the period. (f) (g) (h) (i) Q) (k) (I) 741 Name (genus, species or subspecies) was erroneously omitted from FL. Indication as n. ssp. on p. 55 is erro- neous. Misspelling of FL or text is corrected. FL entry of subspecies nicobaricum and somalicum does not agree with text; Iji- ma may have been “doubtful” that these are specifically distinct (p. 51) but he listed them as recognized species. FL also contradicts Schulze (1904) who, with doubt, recognized somalicum as a species but did not mention nicobari- cum. Both are included here in con- formity with Ijima’s text list. FL erroneously contains separate entries for H. (Onconema) obtusum as well as the two constituent varieties: gracilis and robusta; since Ijima’s concept of the va- rieties (Subspecies here) 1s not stated, they are accepted as for the period. Ijima clearly dropped the subgenera Skianema Lendenfeld, 1915, and Thal- lonema Lendenfeld, 1915, to synonomy with Oonema Lendenfeld, 1915, in the text. Their retention in FL, and the po- sition of their contained species, are er- roneous. Ijima did not intend his terse descrip tions of Lonchiphora and L. inversa pro- vided here to constitute formal diag- noses; strict reflection of his opinion would require deletion of these from the corrected FL. However, his intent to provide a more detailed report at a later date was never realized. The informa- tion given by Ijima has been accepted by Reid, 1958 and 1963, and by the author as adequate indications for both. The relation of Ijima’s “L. sp.” to L. inversa needs reevaluation. (m) Species and subspecies entries of FL un- der Eurete are particularly troublesome. Text statements clearly indicate ljima’s movement of E. erectum erectum, E. erectum tubuliferum and E. erectum gracile to Conorete. E. alicei Topsent, 1901, is also clearly transferred to Gym- 742 norete alicei (Topsent, 1901). The al- location of E. erectum forma D Lenden- feld, 1915, and E. mucronatum Wilson, 1904, are not indicated in the text, but symbols on FL are interpreted to reflect Ijima’s text implication to transfer these to Conorete. (n) Placement in FL as a subspecies is er- roneous; it is a newly described species. (o) The required nominate subspecies name is inserted by HMR. (p) FL entry as a full species is erroneous; japonica is clearly described as a sub- species of P. elisae in the text. There is no earlier use of P. japonica. Ijima’s transfer of C. calyx Schulze, 1886, and C. tenerum Schulze, 1899, and suggested transfer of H. hamatum Schulze, 1886, to Heterochone are ac- cepted. He made no mention of H. /a- mella subspecies, but they are included with indication of their uncertain status. (r) Ijima’s (p. 210) recognition of D. crispus Schmidt, 1870, as a questionable syn- onym of Lefroyella decora Thomson, 1877, is accepted as intent; retention of D. crispus in FL is considered erro- neous. The undescribed species, Gymnorete variolosum mentioned on p. 120, 166, was a nomen nudem as of 1927. Ref- erence to the ms form Gymnodictyum variolosum (p. 164) is taken as a mis- taken reference to that species. (t) Ijima’s intent to transfer Hexactinella labyrinthica to Tretodictyum is ambig- uous but accepted. Ijima’s occasional use of “Tretodictyum spongiosum” in the text, e.g., p. 209, 236, is nowhere stated to imply taxo- nomic validity of that combination; these are interpreted to be errors in ref- erence to Hexactinella spongiosa \jima. Text statements concerning distinction between A. auricularis and filholi are contradictory; the final text remark is adopted. (w) Here (p. 297) and in his last (1916) pub- — (q (s) — (u (v — PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lication, Ijima clearly rejects formal rec- ognition of many historic names for va- rieties and subspecies of A. beatrix (e.g., bocagei, ramosus and azoricus) and forms of A. vastus (vastus and whiteaves- ianus). He does, however, maintain dis- tinction of A. beatrix orientalis which necessitates acceptance of a named nominate subspecies, A. beatrix beatrix Gray, but it is understood to include all of the other poorly known forms not included in A. beatrix orientalis. The species status accorded to 4. bocagei, ramosus, and whiteavesianus in FL is contrary to Ijima’s concepts. FL erroneously retains Calycosilva Len- denfeld, 1915, and the single species consisting of 3 varieties (here subspe- cies); the move to Sympagella is clearly indicated on p. 345. Schulze and Kirkpatrick, 1910, em- ployed the nominate trinomial, Rossella racovitzae racovitzae, for a form clearly not assignable to Topsent’s type; it re- quires a new subspecific name. (x) (y) Action List Taxonomic actions, at and below generic level, which are attributable to Ijima, 1927, are arranged alphabetically within subclass. Newly erected taxa may be indentified by authority on the preceeding Corrected List. Transfer is represented by “‘origin — final’’; synonymy by “junior = senior’; “?” indi- cates that Ijima expressed lack of strong conviction for the action or inconsistency in his statements. AMPHIDISCOPHORA: aculeatum (Schulze, 1894), p. 50 (Hyalo- nema) — (Pteronema). aequatoriale Lendenfeld, 1915, p. 54 (Ski- anema) ~ (Oonema). agassizi (Lendenfeld, 1915), p. 53 (Hyalo- nema) — (Onconema). alcocki (Shulze, 1895), p. 52 (Hyalonema) — (Paradisconema). VOLUME 103, NUMBER 3 apertum apertum (Schulze, 1886), p. 51 (Hyalonema) — (Cyliconema). apertum maehrenthali (Schulze, 1895), p. 51 (Hyalonema) reinstated and > (Cyli- conema). calix (Schulze, 1904), p. 53 (Hyalonema) > (Corynonema). cebuense (Higgin, 1875), p. 50 (Hyalonema) — (Pteronema). clathratum (jima, 1895), p. 53 (Hyalone- ma) — (Corynonema). clavigerum (Schulze, 1886), p. 53 (Hyalo- nema) — (Corynonema). coniforme (Schulze, 1904), p. 51 (Hyalo- nema) — (Cyliconema). conus (Schulze, 1886), p. 51 (Hyalonema) — (Coscinonema). cupresiferum (Schulze, 1893), p. 53 (Pri- onema) ?— (Corynonema). depressum (Schulze, 1886), p. 53 (Lepto- nema) ?— (Corynonema). drygalskii (Schulze, 1910), p. 51 (Hyalo- nema) — (Cyliconema). elegans (Schulze, 1886), p. 51 (Hyalonema) 2 (Coscinonema). fruticosum Schulze, 1893, p. 37 Hyalonema 2?— Monorhaphis. geminatum Lendenfeld, 1915, p. 54 (Thal- lonema) > (Oonema). gigas Schulze, 1902, p. 9, Poliopogon ?-> Pheronema. globiferum (Schulze, 1904), p. 51 (Oonema) — (Cyliconema). globus (Schulze, 1886), p. 50 (Hyalonema) — (Pteronema). gracile (Schulze, 1886), p. 51 (Hyalonema) ?— (Coscinonema). grandancora (Lendenfeld, 1915), p. 53 (Hy- alonema) — (Corynonema). heideri (Schulze, 1894), p. 50 (Hyalonema) — (Pteronema). hercules (Schulze, 1899), p. 53 (Hyalonema) — (Corynonema). heymonsi Schulze, 1895, p. 51 (Hyalonema) — (Coscinonema). (Hyalonema) s. restr. Lendenfeld, 1915, p. 49, 50 subdivided to 7 new subgenera: (Euhyalonema), (Pteronema), (Coscino- 743 nema), (Cyliconema), (Paradisconema), (Corynonema), (Onconema). indicum (Schulze, 1895) p. 51 (Hyalonema) with 2 ssp. ~ (Coscinonema). infundibulum (Topsent, 1896), p. 51 (Hy- alonema) — (Cyliconema). investigatoris (Schulze, 1900), p. 52 (Hyalo- nema) — (Paradisconema). kenti (Schmidt, 1880), p. 51 (Hyalonema) — (Coscinonema). lamella (Schulze, 1900), p. 51 (Hyalonema) — (Coscinonema). lusitanicum (B. d. Bocage, 1864), p. 52 (Pri- onema) — (Leptonema). martabanense (Schulze, 1900), p. 51 (Hy- alonema) — (Cyliconema). masoni (Schulze, 1895), p. 53 (Hyalonema) — (Corynonema). molle (Schulze, 1904), p. 51 (Hyalonema) — (Cyliconema). nicobaricum (Schulze, 1904), p. 51 (Hyalo- nema) — (Cyliconema). obtusum (Lendenfeld, 1915), p. 53 (Hyalo- nema) with 2 ssp. ~ (Onconema). ovatum (Ijima, 1895), p. 51 (Hyalonema) — (Cyliconema). ovuliferum (Schulze, 1899), p. 52 (OQonema) — (Leptonema). owstoni (Ijima, 1894), p. 53 (Hyalonema) — (Corynonema). pateriferum (Wilson, 1904), p. 51 (Phialo- nema) — (Coscinonema). pellucidum (Ijima, 1894), p. 50 (Phialone- ma) — (Euhyalonema). pirum (Schulze, 1895), p. 51 (Hyalonema) > (Cyliconema). placuna (Lendenfeld, 1915), p. 53 (Hyalo- nema) — (Corynonema). polycaulum (Lendenfeld, 1915), p. 50 (H)y- alonema) — (Pteronema). populiferum (Schulze, 1899), p. 53 (Hyalo- nema) — (Corynonema). proximum (Schulze, 1904), p. 50 (Hyalo- nema) — (Euhyalonema). rapa (Schulze, 1900), p. 51 (Hyalonema) > (Cyliconema). schmidti (Schulze, 1899), p. 51 (Hyalone- ma) — (Coscinonema). 744 sieboldi (Gray, 1835), p. 28, 50 (Hyalone- ma) — (Euhyalonema). simile (Schulze, 1904), p. 51 (Hyalonema) — (Cyliconema). (Skianema) Lendenfeld, 1915, p. 49 = (Oonema) Lendenfeld, 1915. somalicum (Schulze, 1904), p. 51 (Hyalo- nema) — (Cyliconema). tenerum (Schulze, 1886), p. 51 (Qonema) — (Cyliconema). tenuifusum (Lendenfeld, 1915), p. 53 (Hy- alonema) — (Corynonema). (Thallonema) Lendenfeld, 1915, p. 49 => (Oonema) Lendenfeld, 1915. thomsonis (Marshall, 1875), p. 51 (Hyalo- nema) — (Cyliconema). toxeres (Thomson, 1877), p. 51 (Hyalone- ma) — (Coscinonema). tulipa (Schulze, 1904), p. 51 (Hyalonema) — (Cyliconema). tylostylum (Lendenfeld, 1915), p. 53 (Ay- alonema) > (Corynonema). umbraculum Lendenfeld, 1915, p. 54 (Ski- anema) ~ (Oonema). valdiviae (Schulze, 1904), p. 51 (Hyalone- ma) — (Cyliconema). weltneri (Schulze, 1895), p. 53 (Hyalonema) > (Corynonema). HEXASTEROPHORA: alicei Topsent, 1901, p. 166 Eurete > Gym- norete. annandalei Kirkpatrick, 1908, p. 166 Eu- rete > Pleurochorium. Calycosilva Lendenfeld, 1915, p. 345 => Sympagella Schmidt, 1870. calyx Schulze, 1886, p. 118, 284 Chonelas- ma ~— Heterochone. cantharellus Lendenfeld, 1915, p. 345 Ca- lycosilva with 3 ssp. implied > Sympa- gella. carteri Schulze, 1886, p. 166, 178 Eurete > Pararete. Chonelasma Schulze, 1886, p. 118, 284 part ~ Heterochone Ijima, 1927. clavigera (Schulze, 1887) Topsent, 1904, p. 137 Claviscopulia > Farrea occa clavi- gera (Schulze, 1887). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON conicum Schmidt, 1880, p. 214 Scleropleg- ma — Myliusia. doederleini Schulze, 1886, p. 165 Chone- lasma — Ptychodesia. erectum Schulze, 1899, p. 166 Eurete — Conorete. farreopsis Carter, 1877, p. 166, 178 Eurete — Pararete farreopsis farreopsis. filholi Topsent, 1904, p. 112, 231, 244 Hex- actinella ?— Auloplax. gerlachei Topsent, 1901, p. 166 Eurete > Pararete. hamatum Schulze, 1886, p. 284 Chonelas- ma ?— Heterochone. ingalli Bowerbank, 1869, p. 214 Iphiteon > Dactylocalyx. labyrinthica Wilson, 1904, p. 231 Hexac- tinella ?— Tretodictyum. latum Schulze, 1886, p. 220, 231 Tretodic- tyum — Hexactinella. Leucopsacus ljima, 1898, p. 320 — Leu- copsacas (spelling correction). lotifolium Tjima, 1903, p. 345 Caulophacus > Caulodiscus. monticularis (Lendenfeld, 1915), p. 231 Hexactinella > Tretocalyx. occa Bowerbank, 1862, Carter, 1885, p. 131 — Farrea occa occa Bowerbank, 1862, Carter, 1885. oviformis (Schulze, 1886), p. 345 Pleorhab- dus implied — Caulophacus. Pleorhabdus (Schulze, 1887), p. 345 = Cau- lophacus Schulze, 1886. Ramella Schulze, 1904, p. 130 = Saroste- gia Topsent, 1904. schmidti Schulze, 1886, p. 171 Eurete > Eurete schmidti schmidti. schulzei Topsent, 1890, 1892, 1901, 1904, p. 205 Chonelasma => Periphragella lu- sitanica Topsent. semperi (Schulze, 1886), p. 166 Eurete — Pararete. subglobosus Gray, 1867, p. 214 Dactyloca- lyx 2? Myliusia. subglobosus of Schmidt, 1880 & Schulze, 1887, p. 214 Dactylocalyx ? = Dactylo- calyx ingalli (Bowerbank). Syringidium Schmidt, 1880, p. 210 = Lef- royella Thomson, 1877. VOLUME 103, NUMBER 3 tenerum Schulze, 1899, p. 284 Chonelasma — Heterochone. tubulosa Schulze, 1904, p. 130 Ramella ? = Sarostegia oculata Topsent, 1904. zitteli Schmidt, 1880, p. 210 Syringidium = Lefroyella decora Thomson, 1877. Acknowledgments This work was supported by an operating grant from the Natural Sciences and Engi- neering Research Council of Canada. Literature Cited Burton, M. 1928. Porifera or Spongida.—The Zoo- logical Record [1927] 64(II1):1-11. 1954. Sponges (Rosaura Expedition).—Bul- letin of the British Museum (Natural History), Zoology 2:215—239, pl. 9. Hartman, W. D. 1982. Porifera. Pp. 640-666 in S. P. Parker, ed., Synopsis and classification of liv- ing organisms, volume 1. McGraw Hill, New York. Ijima, I. 1916. Notes on Aphrocallistes beatrix Gray, particularly with reference to the form occurring in East Asiatic Seas.— Annotationes Zoologicae Japonenses [X:173-183. 1927. The Hexactinellida of the Siboga Ex- pedition. Siboga Expedition reports, volume 6. E. J. Brill, Leiden, i—vii, 1-383, pls. 1-26. Koltun, V. M. 1967. (Vitreous sponges of the north- ern and far-eastern seas of the USSR.) (In Rus- sian).—Opredeliteli po Faune SSSR, Izdavaye- myye Zoologicheskik Institutom Akademii Nauk SSSR 94:1-124. 745 Laubenfels, M. W.de. 1936. A discussion of the sponge fauna of the Dry Tortugas, in particular, with material for a revision of the families and orders of the Porifera.— Carnegie Institution of Wash- ington Publication 467:1—225, pls. 1-22. Reid, R. E. H. 1957. Notes on the hexactinellid sponges. II. Dactylocalyx Stutchbury and the family Dactylocalycidae Gray.—Annals and Magazine of Natural History (12)10:821-826. . 1958. A monograph of the Upper Cretaceous Hexactinellida of Great Britain and Northern Ireland. Part II.—Paleontographical Society of London CXII:xlvii—xlviii, 1-26, pls. I-IV. 1963. Notes on a classification of the Hex- actinosa.— Journal of Paleontology 37:218-231. Reiswig, H. M. 1979. Histology of Hexactinellida (Porifera). Pp. 173-180 in C. Levi & N. Boury- Esnault, eds., Biologie des spongiaires. Centre National de la Recherche Scientifique, Paris. Schulze, F.E. 1904. Hexactinellida. Wissenschafliche Ergebnisse der deutschen Tiefsee-Expedition auf dem Dampfer “ Valdivia’ 1898-1899, Band IV. Gustav Fisher, Jena, i—viii, 1-266, 52 Taf. Schulze, F. E. & R. Kirkpatrick. 1910. Die Hexac- tinelliden der deutschen Siidpolar-Expedition 1901-1903.—Deutsche Siidpolar-Expedition 1901-1903, Band 12, Heft 1, G. Reimer, Berlin, 1-62, 10 Taf. Tuzet, O. 1973. Hexactinellides ou Hyalosponges. Pp. 633-690 in P.-P. Grasse, ed., Traite de Zoo- logie, III Spongiaires. Masson et Cie, Paris. Redpath Museum and Biology Depart- ment, McGill University, 859 Sherbrooke St. West, Montreal, Quebec, Canada H3A 2K6. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 746-768 TWO NEW SPECIES OF CENTROLENELLA (ANURA: CENTROLENIDAE) FROM NORTHWESTERN PERU John E. Cadle and Roy W. McDiarmid Abstract.—Two new species of Centrolenella are described from the Pacific versant of the Andes in northern Peru (Rio Zana, Cajamarca Department). These constitute the first records for the genus on the western slopes of the Andes in Peru. Centrolenella euhystrix, a large species (males to 31.3 mm, females to 33.6 mm snout-—vent length), has a distinctive coloration (dark greenish black when active at night), broad digits, and large hands. Males of this species have very spinose dorsal skin and lack humeral spines. Although males of C. euhystrix call from rock ledges in or along cascading streams, an unusual calling site for Centrolenella, neither egg masses nor reproduction were observed in this species. Centrolenella hesperia is a smaller species (males to 27.3 mm, females to 28.8 mm snout—vent length) which lacks a typanum, is bright green when active, and has a distinctive lateral white stripe from the snout to the groin. Males of this species have spinose dorsal skin and humeral spines. The reproductive behavior, vocalizations, and larval morphology of C. hesperia are similar to those known for other species of Centrolenella except that females have an unusual behavior at the clutch and egg masses contain empty capsules. The intercalary elements in these species are mineralized; the nature of this element in the Centrolenidae and other frogs is discussed. Resumen. —Se describen dos nuevas especies de Centrolenella de la vertiente pacifica de los Andes en el norte del Peru (Rio Zaha, Departamento de Caja- marca). Estas especies se componen el primer registro del género en Peru occidental. Centrolenella euhystrix es una especie grande (los machos alcanzan a 31.3 mm de longitud corporal y las hembras a 33.6 mm) que tiene una coloracion distintiva (negro verdoso oscuro durante la actividad nocturnal), los dedos anchos, y las manos grandes. Los machos de esta especie son muy espinosos y carecen de espinas humerales. Aunque los machos de C. euhystrix cantan desde anaqueles rocosos en torrentes, 0 a lo largo de los mismos, un sitio de canto poco frecuente para Centrolenella, ni masas de huevos ni la reproduccion fueron observadas en esta especie. Centrolenella hesperia es una especie mas pequena (machos hasta 27.3 mm de longitud corporal, hembras hasta 28.8 mm) que carece de un timpano, es verde brillante cuando esta activa, y tiene una raya lateral blanca desde el hocico hasta la ingle. Los machos de esta especie son también espinosos y tienen espinas humerales. El comporta- miento reproductivo, las vocalizaciones, y la morfologia larval de C. hesperia son similares a aquellas conocidas de otras especies de Centrolenella, menos que las hembras muestran un comportamiento extrana a la nidada, y que las masas de huevos contienen capsulas vacias. Los elementos intercalares de estas especies estan mineralizados; se discute la forma de este elemento en las Cen- trolenidae y otras ranas. VOLUME 103, NUMBER 3 Fig. 1. Map of the central Andes of southern Ec- uador and northern Peru showing the Monte Seco area (star), the type locality for Centrolenella euhystrix and C. hesperia, and all other localities (dots) at which spe- cies of Centrolenella are known to occur. The light hatching indicates areas between 1000 m and 3000 m; darker areas indicate those regions above 3000 m. The Huancabamba Depression of northern Peru and south- ern Ecuador has only a few, disjunct areas above 3000 m. Many frogs of the genus Centrolenella (Centrolenidae) are known from the eastern Andean slopes of Colombia, Ecuador, and Peru (Frost 1985), and new species are being discovered at an astonishing rate (Duellman & Burrowes 1989; Flores 1985, 1987; Flores & McDiarmid 1989). Cannatella & Duell- man (1982) noted 10 species in Peru (nine discovered since the mid-1970s), and in- dicated five undescribed species from the Amazonian slopes. Although Centrolenella is known from the western foothills and slopes of the Andes in Colombia and Ec- uador, none has been reported from Peru’s western slopes. Herein we describe two spe- cies from the Pacific versant of northern 747 Peru about 350 km south of the nearest re- ported localities on the Pacific versant in Ecuador, and about 175 km southwest of the nearest reported locality (Cannatella & Duellman 1982) on the Amazonian versant of Peru (Fig. 1). Cadle collected during the periods 1 May to 25 June, 1987 and 13 to 31 January, 1989, near Monte Seco, a coffee cooperative at 1200 m in the valley of the Rio Zana, Department of Cajamarca, Peru (Fig. 2). Al- though the coast and western slopes of the Peruvian Andes are generally arid and sup- port primarily desert or dry scrub forest, local conditions sometimes permit more mesic environments, particularly in north- ern Peru (Koepcke 1961). The slopes north and east of Monte Seco above about 1500 m support a wet forest that receives heavy rains from January to April, and that is en- veloped by dense clouds for at least the early part of the dry season (May to December). Frogs were collected near streams and wa- terfalls whose headwaters originate on a ridge north and east of Monte Seco. Locally, the highest mountains extend to about 3000 m elevation. Methods and Materials Using dial calipers, we measured to the nearest 0.1 mm as follows: snout—vent length (SVL), straight line from tip of head to vent; head length (HL), angle of the jaw to tip of the head; head width (HW), width of the head at the angle of the jaw; snout length (SL), anterior border of eye to tip of head; eye diameter (ED), measured in the hori- zontal plane; eye to nostril (EN), anterior border of eye to middle of the nostril; tym- panum width (TW), measured in horizontal plane; tibia length (TL), measured with the limb flexed; hand length (HnL), from the proximal border of the outer palmar tuber- cle to tip of digit III. Webbing formula no- tations follow Savage & Heyer (1967), as modified by Myers & Duellman (1982). In determining webbing formulae, we used the point of intersection of the web base with a 748 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ) ¢ El Chorro t 7 } F \ 4 ! -77 Monte Seco, 77°09 Fig. 2. Map of the Monte Seco region showing type localities and local place-names. Numbered sites refer to the following: (1) Type locality for Centrolenella euhystrix; (2) Type locality for Centrolenella hesperia; (3) ‘*‘Chorro Blanco”’—a waterfall and local landmark. Cadle’s field camps were near site 2. Contours are in meters. perpendicular drawn to the digit at that point. Digit lengths were determined as the relative extent of digit protrusion from the hand or foot. Nuptial pads were classified according to Flores (1985). Developmental stages follow Gosner (1960), and tadpole terminology follows Altig (1970). We recorded calls of one species with a SONY ECM-929LT microphone and Walkman Professional® cassette tape re- corder. A copy of the tape has been depos- ited in the tape archive, Division of Am- phibians and Reptiles, National Museum of Natural History. These calls were analyzed using a Kay Digital Sona-Graph 7800 and a Multigon Uniscan II real-time analyzer. Pulse rates and call lengths were measured from waveform analyses or from wide-band audiospectrograms. Call rates were mea- sured from the real-time analyzer screen. Two specimens cleared and double- stained for cartilage and bone using the method of Dingerkus & Uhler (1977) were used for osteological descriptions. Museum depositories for specimens are abbreviated as in Leviton et al. (1985), except for the museum formerly known as the Museo de Historia Natural “Javier Prado” (MHNJP) in Lima, Peru. This museum, presently known as the Museo de Historia Natural de San Marcos, is abbreviated as MHNSM. Descriptions Centrolenella euhystrix, new species Figs. 3-5 Holotype. —Field Museum of Natural History (FMNH) 232510 (field number J. E. Cadle 7628), an adult male, taken from the ridge above basecamp (near Chorro VOLUME 103, NUMBER 3 Blanco), about 4—4.5 km (airline) NE of Monte Seco, Rio Zana, Department of Ca- jamarca, Peru, 2610 m. (Fig. 2). Collected 16 May 1987 by J. E. Cadle. Paratypes. — Fifteen specimens with the following data: FMNH 232509 (adult male), 232511 (adult male, cleared and stained), 232513-14 (adult male and female respec- tively), and MHNSM 3501 (adult female), from about 4.5 km (airline) NE of Monte Seco, 2630 m, 2 June 1987; USNM 292588 (adult female) and ANSP 31574—75 (adult female and male respectively), from 4 km (airline) NE of Monte Seco, 2550-2650 m, 23-24 January 1989; FMNH 232512 (adult male collected 7 May 1987), 231763 (sub- adult collected 24 June 1987), 231770 (adult male collected 24 June 1987), MHNSM 3502 (adult male), 3503 (subadult male), 3504 (adult female), collected 6 May 1987, and USNM 292587 (adult male), collected 28 January 1989, all from basecamp on trail between Monte Seco and Chorro Blanco, about 2.5 km (airline) NE of Monte Seco, 1800 m. Definition. —The following diagnostic features (Lynch & Duellman 1973, Flores 1985) distinguish C. euhystrix from other Centrolenella species: (1) vomerine teeth absent; (2) bones green; (3) parietal perito- neum white, visceral peritoneum clear; (4) color in life: when active at night, dark greenish black with lighter green spicules; when concealed during day, nearly black; when active during day, dark green (brown- ish in sunlight) with light green spicules; col- or in preservative: dark gray to medium gray with white spicules; (5) finger webbing (I- II(2-)-(3.5)I1I(2.5)(2*)1V; (6) toe webbing I(1IH2 Jd )H2*-2.5) TI )H42.5)1V(3>)- (1)V; (7) head round in dorsal outline; snout truncate from above and truncate (slightly rounded in some females) in profile; (8) dor- sal skin texture finely spinose with large pointed spicules in males, smoother in fe- males; (9) ulnar and tarsal ridges absent; (10) no humeral spine; (11) tympanum two- thirds to completely visible; (12) prepollex 749 well developed, no externally visible pre- pollical spine; (13) nuptial excrescences white, forming large Type I pad; (14) no lateral glands; one pair of tubercles ventral to cloacal opening. Description. —Head distinct, wider than body, wider than long, round in outline (most easily seen from below). Snout pro- truding, forming anterior outline of head viewed from above, truncate (occasionally with weak point) from above and truncate to slightly rounded (females) in profile, SL about 23% of HL; canthus rostralis straight, distinct, rounded in section; loreal region shallowly concave; lips not or only slightly flared. Nostril small, slit-like to elliptical, directed laterally on protuberance; inter- narial area flat to slightly concave. Eye mod- erate, directed slightly anterior of antero- laterally (>135°). Tympanum distinct, heavily pigmented, two-thirds to complete- ly visible, dorsomedially inclined, postero- laterally directed; usually smooth or with low spicule; annulus tympanicus indicated as elevated and unpigmented border along anteroventral third to lower two-thirds of tympanum. Supratympanic fold heavy, es- pecially in males, sometimes covering up to one-third of tympanum. Tongue round to weakly cordate. Vomerine teeth absent. Choanae round to slightly elongate, occa- sionally slightly rectangular, about size of subarticular tubercle on finger III, separated by distance 5.5 times their diameter. Vocal slits paired, elongate, posterolateral to tongue. Dorsal skin texture of males finely spi- nose with large, white (in preservative), pointed (often more rounded medially) spicules; spicules sparsely distributed on snout and dorsum of head and best devel- oped dorsolaterally, above tympanum, and on upper arms (Fig. 3); dorsal surfaces of limbs spiculate; spicules present on sides of head and legs, extending onto lateral sides of foot to basal part of digit V; lateral parts of body and dorsal surface of hands smooth. Dorsal skin of females much smoother, 750 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON by FS a . ss De. - oe es ee rs Bt i ._ se Fig. 3. Adult male Centrolenella euhystrix (SVL 31.3 mm, FMNH 232513), collected on 2 June 1987 from about 4.5 km (airline) NE of Monte Seco, 2630 m, Department of Cajamarca, Peru. spicules absent (usually) or sparsely distrib- uted on sides of head, dorsolateral surface of body, and on limbs. Skin on belly vari- able (possible preservation artifact), nearly smooth or weakly to coarsely granulose, sometimes areolate; pectoral area and throat smooth; ventral surfaces of thighs weakly granulose. Moderately to poorly developed pair of cloacal (ventral to pubis) tubercles. Cloacal opening high between thighs; cov- ered by moderately large flap with straight or slightly incised edge; flap margin smooth to crenulate; cluster of 0-9 spicules on each side below the opening. Hands large (HnL/HL, 1.22); digits very wide, lateral ridges along all fingers. Digit lengths of hand I < II < IV < III; webbing absent between I and II, basal between II and III, moderate between III and IV; web- bing formula I-II(2-)—-(3.5)III(2.5){2*)IV. Thumb tip expanded, disc I 60% width of discs III or IV; discs on fingers II-IV ex- panded, broader than disc I, those of III and IV about equal in width. Prepollical area enlarged, no external prepollical spine. Nuptial excrescences large, whitish, Type I. Subarticular tubercles present, that on finger IV largest. Inner metacarpal tubercle elon- gate; outer tubercle more elliptical, equal to or slightly smaller than inner. Small palmar tubercles at base of fingers. No fringes or tubercles along outer edge of forearm. Fore- arm larger than upper arm; no obvious sex- ual dimorphism. No humeral spine. Digit lengths on foot I < II < III <=> V < IV. Webbing extensive, toe webbing formula I(1)-(2-)II(1-)—(2*—2.5)MI(1~)— (2.5)IV(3-)-(1)V. All toe discs expanded; tip on toe I 60% of that on toe IV; discs on toes IIJ-V about equal in size. No tarsal fold. Subarticular tubercles moderate, rounded, about of equal size. Distinct inner metatar- VOLUME 103, NUMBER 3 ° se: o 0 0° 0 oo ego eo OP ga%g oO ° oS o Fig. 4. Dorsal and lateral views of the head of the holotype of Centrolenella euhystrix (male, SVL 31.1, FMNH 232510). sal tubercle, width about equal to tip of digit I; outer tubercle small, poorly developed. Snout—vent lengths of adult males 28.5- 31.3 mm; of adult females 31.1—-33.6 mm. Coloration. —In life the dorsum is dark greenish black in active individuals, and dark green in inactive ones. The head (es- pecially eyelids and lips), back, and limbs of males are densely covered with large, light green spicules, lighter than dorsal ground color and giving a finely spotted appearance to dorsum. Females generally have smooth skin with few spicules on forelimbs, upper lip, and shank, and generally lack the spot- ted appearance. The venter is greenish with yellow wash on anterior belly region. The parietal peritoneum is white over anterior half of belly. The feet and hands are clear or greenish yellow with dull yellowish wash; Fig. 5. Diagrammatic representations of the right hand and foot of the holotype of Centrolenella euhys- trix (male, SVL 31.1, FMNH 232510). the webbing is transparent yellowish. The bones are green. The irises of female (MHNSM 3504) and subadult male (MHNSN 3503) were “‘gray with fine black irregular reticulations”’ (Cadle, field notes); that of adult male (FMNH 232512) deep medium brown. In bright sunlight the dor- sum changes to light greenish gray. Centrolenella euhystrix is capable of rapid color changes from a uniform black to dull dark green. When collected, the holotype was totally black dorsally, with some yellow on the palmar and plantar surfaces, match- ing the black log under which it was found. After a short time in the collecting bag, this frog turned a dull dark green. In preservative, dorsal surfaces of head, body, and limbs of males dark gray to me- dium gray, tubercles white; dorsal surfaces of hands, feet, and webbing pale gray; flanks gray, gradually lightening ventrally; venter creamish gray, palest on proximal parts of thighs and pectoral region; some males dark gray on chest and throat; nuptial excres- cences creamish. Females pale to dark gray dorsally; creamish to gray ventrally. Nicti- tating membrane covered with gray mela- nophores except in central portion of upper half, which lacks pigment. Measurements of holotype. —SVL—31.1; 752 HL—9.7; HW—11.0; ED—3.1; EN—2.2; SL—4.7; TW—0.65; TL—15.9; HnL—11.8. Osteology.—A cleared and stained male specimen of C. euhystrix (SVL 31.0; FMNH 232511) has the prepollex completely en- closed within thumb base; the prepollex ex- tends about 80% the length of metacarpal I. The tip of the prepollex is distinct, car- tilaginous, with some mineralization (see Discussion for use of this term). Metacarpal III bears a medially projecting bony flange near the midpoint. Digit tips are T- to slight- ly Y-shaped; the intercalary elements are mineralized. The proximal flange (deltoid crest) of the humerus is about 45% the length of the bone; it lacks a projecting spine. The vomers are widely separated medially, em- bedded in the large cartilaginous floor of the nasal capsule, and lack teeth. The fronto- parietal fontanelle is large; the anterior pro- jections of the frontoparietals on the skull table overlap the sphenethmoid. The para- sphenoid does not reach the level of the palatines. On the lateral surface of the brain- case, the sphenethmoid is separated by a small cartilaginous gap from the posterior portions of the cranium. Quadratojugals are present but do not articulate with the max- illa. A cartilaginous annulus tympanicus and bony columella are present. The hyoid of Centrolenella euhystrix dif- fers from that illustrated by Eaton (1958, fig. 5) for C. prosoblepon. Eaton indicated only the bony posteromedial processes of the hyoid plate (Duellman & Trueb 1986, fig. 13.21), whereas the hyoid of C. ewhystrix bears cartilaginous anterolateral and pos- terolateral processes as well. David C. Can- natella (pers. comm.) informed us that these processes are typical of Centrolenella hyoids and that Eaton’s figure is erroneous in these details. In general form the hyoid of C. eu- hystrix is similar to that illustrated for Lep- todactylus ocellatus by Duellman & Trueb (1986, fig. 13.21D). The hyoid plate is broad, about 1.5 times as wide as long. The tips of the anterolateral processes are flared distal- ly. The posterolateral processes are slightly PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON longer than the anterolateral processes, and tapered distally. The only bony elements of the hyoid are the posteromedial processes. All digits have intercalary elements be- tween the penultimate and ultimate pha- langes, and in all cases these elements are mineralized (i.e., they stain red with aliza- rin; see Discussion for further consideration of terminology). In the hand the intercalary elements are as wide as the distal ends of the penultimate phalanges. They are prox- imo-distally compressed with the proximal surface slightly convex and the distal surface concave. The intercalary elements in the feet are of the same general form as on the fore- limb. The epiphyses of all phalanges are well- mineralized. Natural history notes. — At night, individ- uals of Centrolenella euhystrix were active mostly on vertical rockfaces in the splash/ spray zone of waterfalls, on rock ledges (up- per or lower surfaces) in or around water- falls, and on liverwort- and moss-covered wet boulders in midstream. Most were lo- cated by eye-shine. Many individuals were observed on rock ledges in waterfalls 6-8 m high, but were too high to collect. At night their dark greenish bodies were difficult to see against the mosses and liverworts that usually cover their perches. A juvenile male (MHNSM 3503; 28.3 SVL) clung to a twig hanging from a vertical rock face 0.8 m above the water in a waterfall spray zone. Another individual was perched on a leaf over water at night. The holotype, the only individual found by day, was under a log in a pile of debris in the middle of a stream. In general, C. euhystrix seems to prefer mi- crohabitats along streams close to water splashing or pouring over rocks. Observations of the behavior of adult C. euhystrix are similar to those reported for Centrolene geckoideum in Colombia (Lynch et al. 1983). These authors reported female and calling male Centrolene on vertical or overhanging rock faces in the spray zones of waterfalls, and egg masses attached to the same rock faces. Given that virtually all in- VOLUME 103, NUMBER 3 dividuals of C. euhystrix were associated with rock faces near or in waterfalls and that no egg masses attributable to this species were found attached to vegetation, we con- sider it possible that eggs of this species are attached to rocks as in Centrolene. Such be- havior has not been reported for any species of Centrolenella. Vocalization and reproductive behavy- ior.— Males called from rock ledges or ver- tical faces at two sites around waterfalls at about 2600 m on 2 June. A lone male called from a large boulder 20 cm above a stream at 1800 m on 6 May. The call sites often were within the splash zone of small water- falls and on dripping rock faces covered with mosses and liverworts. The call is a short “click” or “‘chirp”’ usually given as two notes in rapid succession; occasionally a call con- sisted of only a single note. Although no individuals were heard calling in January 1989, the noise from the torrential streams which are excessively swollen during the rainy season, may have precluded hearing the calls. One female (MHNSM 3504) was collected 0.3 m above the rushing stream and was difficult to see on the bryophyte- covered rock. Two frogs were found in close proximity on a vertical rock face in a spray zone of a waterfall on 23 January. The male (ANSP 31575) was above the female (ANSP 31574) but mostly lateral to her body, and the pair was not observed in amplexus. Pos- sibly, the pair had been disturbed by the observer before they were noticed. The male rapidly ascended the rockface when a light was shone on them. No egg masses that we could attribute to this species were found. Distribution. —Centrolenella euhystrix is known only from two streams at the type locality and from about 2.5 km (airline) NE of Monte Seco at 1800 m along a trail to Chorro Blanco (Fig. 2). Monte Seco is a cof- fee cooperative located approximately 80 km ESE of Chiclayo. The type locality is along an abandoned logging road that par- allels the ridge to the northeast of Monte Seco. Chorro Blanco is a waterfall near the 753 site of Cadle’s field camp. The recorded ele- vational range is 1800 m to 2630 m. Etymology.—The specific epithet ‘‘eu- hystrix” is derived from the Greek word “hystrix,” meaning porcupine, and the pre- fix “‘eu,”’ meaning very. We use it as a noun in apposition to refer to the very spiny, por- cupine-like dorsum of males of euvhystrix, which for a centrolenid is quite unusual. Centrolenella hesperia, new species Figs. 6-8 Holotype. —Field Museum of Natural History (FMNH) 236200 (field number J.E. Cadle 9260), an adult male, taken near the basecamp on trail between Monte Seco and Chorro Blanco, about 2.5 km (airline) NE of Monte Seco, Rio Zana, Department of Cajamarca, Peru, 1800 m (Fig. 2). Collected 28 Jan 1989 by J. E. Cadle. Paratypes. —Sixty-seven specimens with the following data: FMNH 232495, 232503 (adult females), 232496-232501, 232504— 232508 (adult males), 232502 (adult male, cleared and stained), and MHNSM 3505- 3507 (adult males), 3508-3509 (adult fe- males), collected between 6 and 17 May 1987; USNM 292582, 292584 (adult males), 292583 (adult female), and ANSP 31576, 31586, 31588 (adult females), 31577-85, 31587 (adult males) collected 14 Jan 1989; USNM 292585 (adult male) and ANSP 31601-06, 31608 (adult males), 31607 (adult female) collected 18 Jan 1989; USNM 292586 (adult male) and ANSP 31609-13, 31615-18 (adult males) and 31614 (adult female) collected 28 Jan 1989; all from same locality as holotype. ANSP 31589-90, 31592-94 (adult males), 31591 (adult fe- male) from 1.5 km (airline) NE of Monte Seco on trail to Chorro Blanco, 1530 m, and ANSP 3159598 (adult males), 31599-600 (adult females), from the same locality at 1630 m, 15 Jan 1989. Definition. —The following diagnostic features (Lynch & Duellman 1973, Flores 1985) distinguish C. hesperia from other 754 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Adult male Centrolenella hesperia (SVL 26.0, FMNH 232501), collected on 8 May 1987 from 2.5 km (airline) NE of Monte Seco, 1800 m, Department of Cajamarca, Peru. Centrolenella species: (1) vomerine teeth absent; (2) bones green; (3) parietal perito- neum white, visceral peritoneum clear; (4) color in life leaf green with green spicules, light lateral stripe, and white cloacal patch; in preservative pale lavender with irregu- larly placed, pigmentless spots surrounding large, white spicules, giving the appearance of light-colored dots; (5) webbing formula on fingers I-II(2*)-(> 3.5)III(3-)-(2.5)IV; (6) webbing on toes I(27)-(2.5)II(1*) —(2°-2.5)III(1 *-1.5)-(3 -—3.5)IV(3*)-(1.5- 2-)V; (7) head round in dorsal outline; snout weakly truncate from above, obtuse to mod- erately sloped in profile; (8) dorsal skin tex- ture in males shagreen with minute spicules and scattered larger spicules, smoother in females; (9) ulnar and tarsal ridges present; (10) humeral spine present in males; (11) tympanum not visible; (12) prepollex slight- ly enlarged, lacking an external prepollical spine; (13) creamish nuptial excrescences, Type I pad; (14) no lateral glands; one pair of tubercles ventral to cloacal opening; skin beneath cloacal opening thickened, appear- ing glandular. Description. —Head wide, 1.2 times length, only slightly wider than body, round in outline. Snout short (SL about 18% of HL), not or only slightly protruding, weakly truncate in dorsal outline, obtuse to mod- erately sloped in profile; canthus rostralis indistinct; loreal region weakly concave to flat, slightly obtuse; lips slightly flared. Nos- trils oval to slitlike, slightly recessed, di- rected posterolaterally; internarial area con- cave to nearly flat. Eyes small, directed slightly anterior of 135°. Tympanum not dif- ferentiated externally; annulus tympanicus not obvious, tympanic region usually in- dicated by raised area which, when present, is most distinct anteroventrally. Supratym- VOLUME 103, NUMBER 3 Fig. 7. holotype of Centrolenella hesperia (male, SVL 25.5, FMNH 236200). Dorsal and lateral views of the head of the panic fold indistinct. Tongue round to oval, slightly broader anteriorly, occasionally slightly notched anteriorly and posteriorly. Vocal slits large, posterolaterally situated. No vomerine teeth. Choanae round, about 30-40% width of pad on finger I; separated by distance about four times their diameter. Dorsally, skin in males shagreen with minute spicules and larger, scattered spic- ules with white tips; spicules smaller and densest on lip below eye and below tym- panic area on head, larger and more evenly distributed on back, somewhat sparser on snout and upper surfaces of limbs; skin smooth or with few scattered spicules in fe- DS males; lateral (below stripe) skin smoother, without spicules; ventral skin coarsely and uniformly granulose on belly, smooth on throat and chest, less granulose on ventral surfaces of thighs. Moderately developed pair of ventral tubercles below cloacal open- ing. Cloacal opening on upper quarter be- tween limbs, subterminal (visible from above), covered by straight-edged (rarely weakly scalloped), shallow flap; area below cloacal area appears glandular. Hands moderately large (HnL/HL, 1.19); order of finger lengths I < II < IV < III. Finger webbing absent between digits I and II, basal between fingers II and III, and slight between III and IV; formula I-II(2*)- (>3.5)I1I(3-)-(2.5)1V. Ridges along lateral edge of all digits, especially well developed on lateral margins of II and III, and on me- dial margin of IV. Tip of thumb slightly expanded, about 55% of disc III; disc III widest, discs II and IV about equal. Pre- pollical area moderately expanded, no ex- ternal prepollical spine. Nuptial excres- cences cream-colored but not greatly enlarged, Type I pad, relatively large cells on ventral surface of thumb base. Subartic- ular tubercles low, round, largest on finger IV. Metacarpal tubercles indistinct, inner elongate to kidney-shaped, outer more oval; palm tubercular. Ulnar fringe distinct, ex- tending onto hand. Humeral spine present, not protruding externally. Forearm larger than upper arm, no obvious sexual dimor- phism in arm size. Toe lengths I < II < III < V < IV. Toe webbing smooth to granulose, of moderate extent; webbing formula I(2>)—(2.5)II(1*)- (2--2.5)III(1 *-1.5)-(3 —3.5)1V(3*)-(1.5- 2-)V. Thumb disc slightly expanded, 60% of width of disc IV; disc IV widest, III and V about equal. Lateral tarsal ridge extending onto base of digit V. Subarticular tubercles moderate, rounded. Inner metatarsal tuber- cle small, elongate, slightly larger than low outer tubercle, both weakly developed. Snout—vent lengths of adult males 23.0- 27.3 mm; of adult females 24.7—28.8 mm. 756 Coloration. —In life, the dorsum of C. hesperia is light leaf green with pale green spicules on upper surfaces of limbs, head, and body (Fig. 6). A pale cream upper labial stripe continues laterally along the body to the groin, and separates the dorsal color from the yellowish cream ventral color. The irises are grayish white with black reticulations. Ventral surfaces of throat and shank are clear green. Feet, webbing, and ventral surfaces of thighs have a yellow wash. Anterior belly region is cream to yellowish cream. Parietal peritoneum is white over most of belly. A white rump patch is present. A white line runs along the outer border (with frog in sitting position) of the limbs. The bones are green. In preservative males are pale lavender with some scattered, irregular pigmentless spots and white spines giving the appear- ance of pale dots; females are uniform lav- ender with a few irregular pale areas. The upper lip is pale, without pigment. Eyelids also lack pigment except for a few mela- nophores along the upper margins and more at the bases. Under 20 x magnification me- lanophores on some specimens are arranged in a circular pattern in the ear region. A sharply demarcated line occurs laterally be- tween the dorsal pigmentation and the light unpigmented sides. Venter is creamy white; palmar and plantar surfaces and webbing are white. White labial, lateral, ulnar, and tarsal lines, and a white rump patch are vis- ible in specimens preserved less than eight months but not visible in the 1987 sample. (This may reflect some difference in pres- ervation between samples rather than fad- ing.) Measurements of holotype. —SVL—25.5; HL—7.8; HW—9.2; ED—2.6; EN—1.4; SL—3.4; TL—14.5; HnL—9.3 Osteology.—A cleared and stained male specimen (22.2 SVL; FMNH 232502) has a prepollex that extends about 75% of length of metacarpal I and is completely enclosed within the thumb base; a non-mineralized cartilaginous tip is distinct. Metacarpal III PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. Diagrammatic representations of the right hand and foot of the holotype of Centrolenella hesperia (male, SVL 25.5, FMNH 236200). has a medial bony flange with cartilaginous anterior and posterior borders. The tips of the digits are T- or shallowly Y-shaped; in- tercalary elements are mineralized. The proximal flange (deltoid crest) of the hu- merus bears a distal projecting spine that extends about half the length of the flange. The vomers lack teeth and are embedded in the cartilaginous matrix of the floor of the nasal capsule. The frontoparietal fon- tanelle is large; anterior projections of the frontoparietals on the skull table narrowly overlap the sphenethmoid. The anterior tip of the parasphenoid nearly reaches the level of the palatines. On the lateral surface of the braincase, the sphenethmoid is separat- ed by a large cartilaginous gap from the pos- terior cranial elements. The quadratojugals are present but do not articulate with the maxilla. A cartilaginous annulus tympani- cus and bony columella are present. The hyoid structure of Centrolenella hes- peria is similar to that described previously for C. euhystrix with two exceptions. The distal tips of the anterolateral processes are irregularly-shaped and not expanded as in C. euhystrix. In C. hesperia the posterolater- al process on the right side is interrupted by a small non-cartilaginous gap and is less ro- bust than on the left side, which bears a complete process. Undoubtedly, this is an VOLUME 103, NUMBER 3 individual aberration or artifact of prepa- ration, but it may reflect a developmental source of variation in this structure for this or other species of Centrolenella. Intercalary elements are present between the penultimate and ultimate phalanges of all digits. These elements are small (narrow- er than the distal end of the penultimate phalanx) and of the same form but not as well mineralized as those in C. euhystrix. In the hand only a trace of mineralization is evident in the intercalary element of digit I. The intercalary elements in the foot show a progressive increase in mineralization from that in digit I, in which no mineralization is detectable, to that in digits IV and V, which are well mineralized. A similar pat- tern of mineralization was observed in a preparation of C. orientalis (USNM 257182). The epiphyses of all phalanges are entirely cartilaginous. Natural history notes. — At night nearly all individuals were perched on the upper sur- faces of leaves over streams. One female (FMNH 232503) was found by day, appar- ently having been disturbed during survey work. She was hopping on the litter in low vegetation >30 m from the nearest stream. Several others were observed during the day as they rested vertically on upper leaf sur- faces over streams where breeding occurred. Breeding in this species apparently begins with the onset of rains. Males call from the upper surfaces of leaves 0.5-3 m above streams. In 1989, regular daily rains did not begin until 20 January (as inferred from de- gree of soil moisture and interviews with locals), and this coincided with males call- ing regularly each night. A search for eggs along two streams on 14 January revealed only a single mass, despite many calling in- dividuals on that date. Within two weeks, however, egg masses along these streams were abundant. During the dry season males were calling between 6 May and 15 June, but none thereafter (Cadle left the field site on 25 June). Although no males were individually 757 marked, it is our impression that males are active at the same site for several nights and may attract more than one female. Multiple egg masses in different developmental stages often were seen on the same or adjacent leaves near a calling male. As many as four masses, three with eggs in different devel- opmental stages ranging from early neurula to near-hatching, and the jelly remains of another clutch already hatched, were ob- served in the same bush near one male on 7 May. Males (presumably the same indi- viduals) were observed calling from the same sites over the span of several days. Male densities often were very high, as was breed- ing activity (based on the number of ob- served egg masses in some stream sections) compared to our experience with other spe- cies of the genus. In two frequently-worked sections of stream, male densities some- times exceeded 30 individuals along ap- proximately 10 m. Densities apparently de- pend primarily on the amount of foliage available for perches. Although only one kind of vocalization was noted, aggressive interactions occurred among males. On 7 May at 2204 hr, two males (MHNSM 3505, 23.0 mm SVL; 3506, 24.3 mm SVL) were observed fighting belly- to-belly on the petiole of an elephant-ear leaf (Araceae) in a head-to-vent position. They released their grappling holds when disturbed by the light. Grappling or similar behavior between males has been observed in other species of Centrolenella: C. fleisch- manni in Mexico and C. valerioi in Costa Rica (McDiarmid & Adler 1974), C. grif- fithsi in Ecuador (Duellman & Savitzky 1976), C. fleischmanni and C. prosoblepon in Costa Rica (Jacobson 1985), C. prosob- lepon in Panama (Jungfer 1988), and in Ec- uador (McDiarmid, unpublished field notes), and is not unexpected in high-density sit- uations especially among territorial males. This head-to-vent position has not been ob- served previously, but we are not surprised and view it as another variant of the several grappling positions reported. Males of many 758 centrolenid species equipped with special- ized morphological structures (e.g., humeral hooks or spines, prepollical spines) are ter- ritorial (McDiarmid, unpublished notes) and use them in an agressive fashion against oth- er males during fights over calling and egg- laying sites. The outcome of the fight, not the positions assumed by the combatants during the grappling, is the more significant parameter. Taylor’s suggestion (1949) that the prepollical spines in Centrolenella spi- nosa serve as a grasping organ during mat- ing has not been supported by field obser- vations and seems highly unlikely to us. Likewise, suggestions in the literature that the humeral spines in Centrolene gecko- ideum may have some scansorial function, perhaps enabling an individual to maintain its hold on a tree (Noble 1920), or in other species (e.g., Centrolenella prosoblepon) ap- parently functioning to insure amplexus (Noble 1924) or involved mechanically in amplexus (Eaton 1958) also are unverified and lack merit. We have seen mating pairs of several species (including C. prosoblepon) whose males have humeral spines, and in no instance did the males insure their grips on the females or otherwise use their spines during amplexus. We contend that humeral spines are secondary sexual traits used by males to defend their territories from other males and in this sense are analogous to horns in certain territorial mammals. Centrolenella hesperia lays light greenish- white eggs in a single, loosely arranged layer on the upper surfaces of leaves above water, usually on the distal quarter of the leaf near its tip. Egg masses (FMNH 232713-15; ANSP 31619-21; USNM 292591) were found on ferns (Polystichum), a melastome, leaflets of a small palm, elephant ears (Ara- ceae), and other understory plants along the stream margins. In two or three instances masses were attached to aerial roots of epi- phytes situated on tree limbs overhanging the water. An egg mass (FMNH 232715) taken from a leaf 1.5 m above the water at 2114 hr on PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 7 May contained 26 light green eggs in Gos- ner stage 1. These eggs measured about 2.2 mm diameter and may have been laid ear- lier that evening. A calling male (FMNH 232508, SVL 25.7) was collected on the same bush about 1.0 m above the water. Another egg mass (USNM 292591) collected during the day of 29 January contained 38 eggs in stages 4 and 5 that measured 2.5 to 2.7 mm diameter. A sleeping male was found on top of a leaf near this clutch. This clutch of 38 eggs was attached to the upper surface of a fern frond (Polystichum sp.) and had 16 empty capsules on its proximal (top) edge (Fig. 9). All egg masses examined had empty capsules on the side closest to the leaf pet- iole. Empty capsules have been observed in egg masses of certain other Centrolenella species (McDiarmid, unpublished field ob- servations) but not with the apparent con- sistency of this species. Because the capsules are always positioned along the upper edge of the clutches of Centrolenella hesperia, we suspect that they may function to increase the adhesion of the egg mass to the leaf or, by way of their hygroscopic tendencies, re- tain water and serve as an additional water source for the eggs during their several days of development when they are exposed to the air. The size of 12 “dry season’ clutches (2— 9 May) range from 16 to 30, X = 20.7 + 4.4. This is significantly smaller than the size of 15 clutches laid at the beginning of the “‘wet season” (29 Jan), range 19-42, X = 33 + 6.9 (t-test for difference between the means, t = 4.096, P <0.001). Some observations suggest that loss of egg masses due to desiccation was a problem in 1987 for those laid after mid-May. Breeding activity for this species, as indicated by the number and regularity of calling males and recent egg masses, declined rapidly after mid-May. By this time the regular rains had ceased, fogs characteristic of the site were less intense and enduring, and the humidity was lower. The last viable clutches were ob- served on 25 May, when three masses with VOLUME 103, NUMBER 3 Fig. 9. Drawing of an egg mass (USNM 292591) of Centrolenella hesperia attached to the upper surface of a fern (Polystichum sp.) frond showing the position of the 38 eggs and 16 capsules. Line equals 20 mm. advanced tadpoles were found along the stream where most of the previous breeding activity had occurred. Two desiccated clutches and no fresh egg masses were ob- served here on 15 June. On 26 May along another stream two desiccated clutches were found on the same palm frond where viable clutches had been seen earlier in the month. At 2055 hr on 7 May, a male was calling about 40 cm from a female that was alone 759 on top of an egg mass. When checked at 2248 hr the male was still calling but the female was gone; 17 eggs were in the clutch. At the time Cadle pondered the meaning of this behavior and wondered whether the fe- male was performing some egg-laying ac- tivity in the absence of the male. More de- tailed observations of what appears to be the same behavior were made later at a dif- ferent site on the same stream. On 10 May a female (FMNH 232495; 28.8 SVL) was found at 2116 hr next to an egg mass. When first observed, the female was flattened against the leaf in a normal sitting position with her cloacal opening oriented above the upper (proximal for the leaf) edge of the mass containing 16 recently laid eggs. After maintaining this position for several min- utes, she raised the rear part of her body, and at the same time positioned her thighs perpendicular to the body axis. She held this new position for several minutes. During the next hour, the female alternated several times between this behavior and the more normal sitting position, at which time the frog was collected. We do not understand the significance of these females’ behavior. We do not think that the females were depositing eggs in- dependently of the males, and no issuances from the cloaca were observed. Dissection of female FMNH 232495 revealed small Ova in the right ovary, and small, as well as somewhat larger ova in the left ovary. No eggs were present in the body cavity or ovi- ducts (which were enlarged). These obser- vations suggest that she was not ready to lay eggs when collected, but the enlarged Oviducts indicate that she may have re- cently laid eggs. We speculate that the fe- males were: (1) wetting the egg mass (likely their own and presumably with water stored in their bladder), a behavior that has been reported for males of C. fleischmanni in Costa Rica (Jacobson 1985, Mark Hayes, pers. comm.); (2) depositing empty capsules in the mass presumably just after depositing the eggs (see previous discussion); or (3) 760 pressing the eggs against the leaf surface to increase adhesion. Aichinger (1987) report- ed a similar behavior by female Hyla brevi- frons that he described as brushing the eggs into a monolayer. Because rain is more like- ly to detach an egg mass placed on the upper surface ofa leaf than one placed on the lower surface of a leaf, and because the choice of an egg deposition site apparently is species- specific, those species that place their eggs on the upper surface of leaves might be ex- pected to have behaviors that increase egg mass adhesion. We suspect (but did not ob- serve) that the females’ behavior was as- sociated with depositing empty egg cap- sules, but cannot rule out the other two explanations. Whatever the functional sig- nificance of this peculiar behavior in fe- males might be, to our knowledge it has not been reported previously in species of Cen- trolenella. Larvae.—A few tadpoles (FMNH 232710-232712) were reared to stage 25_ (Gosner 1960) from clutches taken from leaves in the field. The largest of these mea- sured 14.9 mm total length with a tail 10.2 mm long. The tadpole is typical of centro- lenids and has a fusiform body, rounded snout, and long, narrow tail. The eyes are small, cresent-shaped and dorsally located. The nostrils are located slightly closer to the eye than to the tip of the snout. Near the midpoint of the tail, the ventral fin is slightly deeper than the dorsal fin, and each is about 20% of the tail height. The tail has a rounded tip. The spiracle is small, oval, and opens posterolaterally about three-fourths of the way back on the left side of the body below the midline. The vent tube is medial. The oral disc is anteroventral and has a uniserial row of about 22 large marginal papillae lat- erally and posteriorly (the anterior edge is bare). Jaw sheaths are narrow, weakly kera- tinized, and serrate; they appear striated. The labial tooth rows are barely visible and not completely keratinized; A—2 has a wide medial gap; the short sections are located nearly lateral to the anterior jaw sheath. The labial tooth row formula is 2(2)/3. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Vocalizations. —Centrolenella hesperia has a short, two-note, high-frequency call (Fig. 10). A calling male (FMNH 232499, SVL 25.4) was recorded on 8 May 1987. The only air temperature data available are maximum (26°C) and minimum (10°C) val- ues recorded for that day. An analysis of seven calls shows the following call char- acteristics (means are given with ranges in parentheses). The call consists of two notes with an overall duration of 0.12 sec (0.10—- 0.13). The first note has either two distinct pulses (two calls) or three distinct pulses (five calls) with an overall duration of 0.04 sec (0.03—0.05). The second note, consisting of one (occasionally two) pulses, lasts 0.02 sec (0.02—0.03) and is separated from the first note by 0.05 sec (0.04—0.06). In both notes the dominant frequency of the call is in the 3300-4000 hz range (mean about 3630 hz). Calls (Fig. 10) were given at a rate of about two per second. The pulse rate cal- culated for the first note was 71.4 notes per second (62.5—76.9). The rise time for these pulses is very fast, averaging 0.0014 sec (0.0008-0.0025). Distribution. —Centrolenella hesperia 1s known only from the vicinity of the type locality (Fig. 2). The recorded elevational range is 1500 m-1800 m. Although stream inaccessibility made searching extensively above 1800 m difficult, this species does not appear to occur above 2500 m where its detection would have been likely. Etymology.—The specific epithet “hes- peria”’ is a Latinized word derived from the Greek “‘hesperos”” meaning western or of the evening. We use it as an adjective in reference to its being one of two species of Centrolenella first recorded from western Peru, and in refrence to its evening activi- ties. Discussion Comparisons. — We do not understand the relationships among centrolenid frogs. In part this is due to the incredible rate at which new species have been discovered and de- VOLUME 103, NUMBER 3 761 aN = = Pe eT nN FREQUENCY (khz) TIME (sec) Fig. 10. Audiospectrograms of five advertisement calls of a male Centrolenella hesperia (SVL 25.4, FANH 232499) with a wide-band filter (300 hz). Calls were recorded on 8 May 1987 at the type locality. Waveforms of the first and third calls are shown above the audiospectrogram. Line between waveforms represents 0.034 sec. scribed. By our calculation about 80% of the recognized species (Frost 1985 and publications since) in the family have been described in the past 30 years and nearly 30 species (> 40% of the total described species diversity) in the past 10 years. The number of known but undescribed species also is quite high. In addition to the two we de- scribe here, we know of three others from Peru, four from Ecuador, and four or more from Venezuela that are undescribed; Pedro Ruiz-Carranza (pers. comm.) informed one of us of a large number of undescribed forms from Colombia. Moreover, the lack of detailed compar- ative morphological descriptions has pre- cluded a comprehensive phylogenetic treat- ment of the family. Consequently, the presently recognized genera and species groups do not easily accommodate many of the species being described. This problem stems at least partly from the relatively re- cent recognition of the family (Taylor 1951) and partly because two of the genera pro- posed in that paper were inadequately de- fined and generally have not been accepted (Goin 1964, Savage 1967). When species groups have been proposed (Savage 1967, Savage & Starrett 1967, Starrett & Savage 1973), coverage has been restricted to geo- graphic regions (i.e., Costa Rica or lower Central America), and the groups were sub- sequently claimed not to accomodate easily some species from other geographic regions (Lynch & Duellman 1973). These latter au- thors recognized the problems associated with species-group definitions based on geo- graphic subsets (they reported species from Ecuador) of the family and preferred not to alter previous groupings or propose new ones. We do not attempt to evaluate critically the various groupings that have been pro- posed within the family. We also recognize that our groups are ones of convenience and may not be monophyletic lineages, but for purposes of comparison we use the follow- ing five groups of centrolenids. (1) The ge- nus Centrolene Jiménez de la Espada (see Ruiz-Carranza et al. 1986 for diagnostic traits), which includes two described forms (large species, presumably with exposed hu- 762 meral spines in males) and probably some undescribed ones. (2) Species within the currently recognized genus Centrolenella Noble (type C. antioquiensis Noble) made up of smaller species, the males of which have humeral spines (part of Savage’s [1967] prosoblepon group). (3) A second group within Centrolenella that includes species in which males lack humeral spines. This cluster of species also is part of Savage’s prosoblepon group; and if shown to be tax- onomically distinct, they would be placed in Taylor’s (1951) genus Cochranella (type C. granulosa Taylor). (4) Another group currently within the genus Centrolenella in- cludes several small species with distin- guishing characteristics (clear parietal peri- toneum, no humeral spines, white color in preservative). This group, originally defined by Savage (1967) and Starrett & Savage (1973), has found general acceptance and has been referred to in the literature (Lynch & Duellman 1973, Cannatella 1980, Can- natella & Lamar 1986) as the fleishmanni group. As currently understood, this group has no available taxonomic name. (5) A fi- nal group currently placed within Centro- lenella includes those species with prepolli- cal spines. This trait was recognized by Taylor (1951) in his description of the genus Teratohyla (type C. spinosa Taylor). Two species (C. gemmata and C. lynchi) with prepollical spines also have humeral spines (Flores 1985) while the third, C. spinosa, does not. For this discussion, we include C. gemmata and C. lynchi in group 5. Centrolenella euhystrix lacks humeral spines and therefore is associated with spe- cies in our group 3. The absence of humeral spines will distinguish it from all species of Centrolene (group 1) and those species of Centrolenella in group 2. Centrolenella eu- hystrix also lacks prepollical spines and thus differs from species in group 5. Though sim- ilar to members of the fleischmanni group (group 4 above) in that they all lack humeral spines and vomerine teeth, C. euhystrix is easily distinguished from all species in group PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 4 (characteristics in parentheses) by the fol- lowing: larger size (usually less than 25 mm SVL), spinose skin (smoother skin), color- ation in life dark green to nearly black with- out spots (pale green or yellow green with diffuse yellow spots or with pale reticula- tion) and in preservative dark gray (creamy white), truncate snout (truncate or round- ed), distinct canthus (often flattened), and white parietal peritoneum (clear parietal peritoneum). Centrolenella euhystrix lacks vomerine teeth and so differs from many species in group 3 which also lack humeral spines but have vomerine teeth. The following com- bination of characters distinguishes C. eu- hystrix from other group 3 species that lack vomerine teeth: green bones, large hands with wide digits and expanded toe tips, ex- tensive toe webbing, color in life dark green- ish black without light or dark flecks, spots, or ocelli, dark gray in preservative, and prominent (in males) pale green (life) or white (in preservative) spines on dorsum. In comparison to the eleven currently rec- ognized species of Centrolenella from Peru (Cannatella & Duellman 1982, Flores & McDiarmid 1989), Centrolenella euhystrix can be distinguished from all by its larger size, dark coloration in life and in preser- vative, and the extremely spinose dorsum of males. Other differences from Peruvian species include the following (charcteristics of euhystrix in parentheses): C. azulae, C. spiculata, and C. midas have vomerine teeth (lacking); C. bergeriand C. munozorum have white bones and concealed tympana (green bones and visible tympanum); and C. phe- nax has a clear parietal peritoneum (white). The only species on the western versant of northern South America that approaches C. euhystrix in size, webbing, and coloration is C. orejuela (Duellman & Burrowes 1989), but this species is reported to have vomer- ine teeth, smooth skin, and white visceral peritoneum. We note that the holotype of C. orejuela, reported as an adult female by Duellman & Burrowes (1989:5), in fact is VOLUME 103, NUMBER 3 an adult male with vocal slits and large nup- tial pads. Also, the paratype (IND-AN 1520) illustrated in Duellman & Burrowes’ paper (1989, fig. 1) is listed as a female in the caption but as an adult male in the text. In many respects (size, coloration, spi- nose dorsum, absence of vomerine teeth, webbing) C. euhystrix is similar to C. joh- nelsi (Cochran & Goin 1970) from northern Colombia. However, C. johnelsi has hu- meral spines and smaller finger discs. The closest relative of C. euhystrix may be found among several undescribed species recently collected from the Amazonian side of the Andes in northern Peru, or perhaps 1n other areas of northern Peru or southern Ecuador (see below). Centrolenella hesperia (placed in our group 2) has humeral spines and thus 1s dis- tinct from members of groups 3, 4, and C. spinosa in group 5. It also is distinct from C. gemmata and C. lynchi, two species with both humeral spines and prepollical spines (prepollical spines are lacking in C. hespe- ria). This species is easily distinguished from the described species of Centrolene (group 1) by the following traits: smaller size (<30 mm vs. >45 mm in Centrolene), concealed humeral spine (exposed in C. geckoideum), proportionally broader head (HW/SVL about 36% vs. <28% in Centrolene), con- cealed tympanum (visible), lack of vomer- ine teeth (present or absent), and light eggs (dark eggs). Among species in group 2, C. hesperia differs from those with vomerine teeth by lacking vomerine teeth, and from many oth- ers by having a uniform green dorsum with small spines. It can be distinguished from C. johnelsi and C. pipilata, two species with spines, by its lavender color in preservative (gray or purplish gray) and concealed (dis- tinct) tympanum. The combination of hu- meral spines, spiculate and spinose dorsum, concealed tympanum, and moderately sloped snout distinguishes C. hesperia from nearly all other species of Centrolenella. It is the only described species with a distinct 763 white lateral line, white cloacal patch, and ulnar and tarsal ridges. Of the described Peruvian species of Cen- trolenella, C. hesperia can be distinguished from all as follows (characters of C. hesperia in parentheses): C. bejaranoi, C. midas, C. ocellata, C. phenax, C. pluvialis, C. spicu- lata, and C. truebae lack humeral spines in males, have visible tympana, and lack lat- eral stripes (present, absent, and present, respectively). In addition, C. midas and C. spiculata have vomerine teeth (absent). Centrolenella bergeri and C. munozorum have white bones and lack humeral spines and lateral stipes (green bones, spines and stripes present). Centrolenella azulae has a visible tympanum and vomerine teeth (tympanum and vomerine teeth absent). Centrolenella mariae has a visible tympa- num and no lateral stripes (undifferentiated tympanum and lateral stripes). Males of C. mariae remain unreported so that the con- dition of the humeral spine is unknown. As with C. euhystrix from this locality, the closest relative of C. hesperia may be found among several undescribed species from Peru’s Amazonian slopes. However, some species at Monte Seco are distributed through the Huancabamba Depression area of northern Peru (Amazonian versant), in other areas of northwestern Peru or western Ecuador (Pacific versant), or have their clos- est relatives in these areas (Fig. 1). Examples include Dendrophidion brunneum, which has a narrow distribution along the Pacific versant of Ecuador and northern Peru (Lieb 1988); Eleutherodactylus lymani, widely distributed throughout the Huancabamba Depression and southern Ecuador (Lynch 1969); and Coniophanes longinquus, which is closely related to Coniophanes dromici- formis of southwestern Ecuador (Cadle 1989). Centrolenella euhystrix, Centrole- nella hesperia, and their close relatives pos- sibly have distributions similar to these, but resolution of this problem must await ad- equate delineation of species groups within the genus. 764 Intercalary elements and their descrip- tors. — Because the family Centrolenidae has been characterized by the presence of car- tilaginous intercalary elements between the penultimate and ultimate phalanges (Duell- man 1975, Duellman & Trueb 1986, Duell- man 1988), our discovery that these ele- ments are mineralized in the two species of Centrolenella described herein was, to us, somewhat surprising, although David C. Cannatella (pers. comm.) informed us that this condition is common in centrolenids. Certain descriptors (e.g., ““ossified,” “‘min- eralized,” and “‘calcified’’?) have been used indiscriminately in the literature to describe the nature of intercalary elements in some frogs. Such use can lead to misinterpreta- tions and subsequent confusion. For ex- ample, in their description of Scarthyla os- tinodactyla Duellman & de Sa (1988) used all three terms to describe the intercalary elements of hylid frogs but did not distin- guish clearly among their use. The confu- sion arises because several types of tissues, including cartilage, can receive deposits of calcium salts by a variety of processes. These tissues then become red with alizarin stain in the standard cleared and stained whole- mount preparations used for anuran oste- ological studies. Drewes (1984) noted that in most adult hyperoliids, the intercalary “cartilages” are partly or wholly mineral- ized (1.e., stain red with alizarin), and used the term “intercalary elements” and the de- scriptor “mineralized” to refer to these structures. We prefer this terminology for the following reasons. The terms ‘“‘calcified”’ (as used with reference to cartilage) and ‘“‘os- sified’ (used specifically with reference to bone) have precise meanings in the histo- logical and anatomical literature that imply specific processes by which a tissue becomes impregnated with calcium deposits. Calci- fied cartilage and bone are distinguished, among other things, by differences in vas- cularity, growth properties, extracellular matrix components, and cell types involved (see Ham & Cormack 1979, Poole et al. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1989). Because these features rarely are studied in evaluating the nature of inter- calary elements in anurans, and in any case cannot generally be distinguished in cleared and stained preparations, a need for precise usage of these terms exists. We argue that the term “‘mineralized”’ is most appropriate to describe those intercalary elements that have calcium deposits and for which the mechanism of deposition has not been de- termined histologically; this term does not imply the specific process or histological structure involved. Noble & Jaeckle (1928: 271) reported that the intercalary element in Polypedates leucomystax occurs as “‘.. . fully ossified bone with a marrow cavi- ty....’? Drewes (1984) examined histolog- ical sections of the intercalary element of Kassina senegalensis, a species with a sim- ilarly appearing and identically shaped in- tercalary element to that of P. leucomystax, and found only calcified cartilage. This con- dition appears to be characteristic of nearly all mineralized intercalary elements that have been studied histologically in anurans (David C. Cannatella, pers. comm.). We suggest that ““mineralized”’ be used as a de- scriptor when only a gross characterization is available (as in cleared and stained prep- arations). Our reading of the literature suggests that most workers use the terms “‘intercalary car- tilages” or “cartilaginous intercalary ele- ments” to refer to non-mineralized inter- calary cartilages, whereas both “‘ossified”’ and “calcified” have commonly referred to intercalary elements composed of calcified cartilage (for which we prefer the term ““mineralized”’). According to Duellman & de Sa (1988), the only neotropical frogs hav- ing “ossified” intercalary elements are the hylines Sphaenorhynchus carneus, Aplas- todiscus perviridis, and Scarthyla ostinodac- tyla, and species in the pseudid genera Ly- sapsus and Pseudis. We now add to this list some species of Centrolenella. Both C. eu- hystrix and C. hesperia, as well as one prep- aration of C. orientalis (USNM 257182), VOLUME 103, NUMBER 3 have mineralized intercalary elements. Two preparations of C. prosoblepon (USNM 252617-18) did not show mineralized in- tercalary elements, but these preparations were only lightly stained with alizarin. We suggest that the lack of evidence for min- eralization of these intercalary elements may be an artifact. The histology of the intercalary elements within the family Centrolenidae has not been clarified. Noble (1920:443) stated that Cen- trolene possessed “. . . an intercalary bone (or partly ossified cartilage). . .”’ but was less clear about the nature of the element in later papers. Although Noble (1925) defined the Hylidae, into which he placed Centrolene and Centrolenella, as a group having char- acters of the Bufonidae but with an inter- calary cartilage or bone between the ulti- mate and penultimate phalanges of each digit, he did not state whether the element in the two genera of concern was cartilage or bone. In his monumental work on the biology of the Amphibia, Noble (1931) re- ferred to the Hylidae as bufonids with in- tercalary cartilages; no mention was made of bony elements. In defining the family Centrolenidae, Taylor (1951) listed the trait as intercalary cartilages. All subsequent ref- erences to the trait in the Centrolenidae have called the structures intercalary cartilages (e.g., Eaton 1958, Duellman 1975, Dowling & Duellman 1978, Duellman 1988) or car- tilaginous intercalary elements (Duellman & Trueb 1986). Peters (1964:166) called the structure “‘intercalary cartilage’’ and de- scribed it as “a phalanx-like cartilaginous (occasionally bony?) element....” Even though one of the other synonyms (inter- calary phalanx) listed by Peters (1964) re- ferred to both cartilage or bone intercalated between the normal components of the dig- it, the use of a question mark suggests to us that Peters was unsure of the nature of the element or confused by previous usage. Given the prevalence of mineralized in- tercalary elements among species of centro- lenids as evidenced by our observations and 765 those of David C. Cannatella (pers. comm.), we recommend that “‘mineralized interca- lary elements” be recognized as a diagnostic feature of the family Centrolenidae (contra Duellman & Trueb 1986, Duellman 1988). The Monte Seco area. —The western slopes of the Peruvian Andes generally are arid, through local conditions sometimes permit more humid areas, as in the vicinity of Monte Seco. Koepcke (1961) summa- rized general features of the vegetation and climate of the western Andean slopes of Peru, including the Rio Zana valley. In gen- eral, the western slopes of northern Peru receive more moisture than those in central and southern Peru. The Rio Zana lies at the southern edge of a transition along the An- dean slope where the vegetation changes to a more mesic type; humidity increases as one moves north from this area (Weber- bauer 1936). Cadle (1989) briefly detailed some aspects of climate in the Monte Seco area. Additional observations from the be- ginning of the rainy season (January 1989) are noted here. When Cadle arrived in the Monte Seco area on 13 January, little rain had fallen as evidenced by the dry soil and lack of recent herbaceous and epiphytic growth. No rain fell on several days between 13 and 20 Jan- uary. Thereafter, each day was character- ized by steady rains and much cloud cover. Precipitation fell either as relatively heavy rains lasting for one to several hours during the day, usually in the afternoon, or as steady daytime drizzles of varying intensity. Rain occurring at night was usually of limited intensity and duration, with most falling be- fore midnight. Dense daytime fogs as had happened during the beginning of the dry season (Cadle 1989) did not occur during this period. The slopes above Monte Seco are steep with relatively few permanent streams and many waterfalls. Because of the topography, nocturnal work along certain portions of streams is difficult and even dangerous. The highest waterfall, Chorro Blanco, drops 766 about 200 m. Many others occur in the study streams (Fig. 2) and vary in height from <1 m to >18 m. During the rainy season, many small temporary streams flow from the hills, and in 1987 some of these lasted into the beginning of the dry season (mid-June) but with greatly reduced flow. In addition to natural streams, local inhabitants have con- structed a series of drainage ditches (ace- quias) along the hillside to channel water for domestic use. Long portions of these ditches extend through forested areas, are well-shaded, and have overhanging vege- tation that seemingly could have provided suitable egg-laying sites for Centrolenella but none were found. Centrolenella hesperia bred along both permanent and temporary streams between 1500 m and 1800 m from the vicinity of the type locality; but C. eu- hystrix was observed only along permanent streams near Cadle’s field camp (site 2) and at higher elevations (site 1 and one other nearby site, Fig. 2). Apparently, neither spe- cies bred along the drainage ditches. This may reflect differences in substrate quality of the ditches (generally sandy) compared to natural streams (gravelly to rocky), gra- dient (gradually sloping versus precipitous), or some other unknown variable (e.g., dif- ferences in oxygen tension between the ditches and streams). Tadpoles of Coloste- thus sp. were found in the drainage ditches, as well as elsewhere in relatively oxygen- poor, standing water with sandy substrates. Centrolenella euhystrix and C. hesperia were found sympatrically only at the type locality of the latter (Fig. 2). This repre- sented the lowest and highest elevations (1800 m), respectively, at which the two species were observed. Greater elevational overlap in the ranges of the two species probably occurs, but the precipitous nature of the steams precluded adequate explora- tion of this possibility. Owing to its dis- tinctive call and overt behavior, C. hesperia was easily detected and thus appeared to be absent from the sites at 2500 m and above where C. euvhystrix was observed. Although PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON individuals of C. euhystrix were more cryp- tic, none were observed during a careful search of several waterfalls below 1800 m and at waterfalls along one other permanent stream at about 1800 m. The only other frog which lays eggs along streams in the Monte Seco area is Telmatobius sp., but it most likely lays its eggs in the water. Other frogs in the area use terrestrial sites (Colostethus and Eleutherodactylus) or carry their eggs (Gastrotheca). Tadpoles of Colostethus sp. were found in the streams (among other sites) at all elevations where adults and eggs of Centrolenella were observed. Tadpoles of Telmatobius sp. were found only in the streams above 2500 m where Centrolenella euhystrix adults (but no tadpoles) were ob- served; Gastrotheca sp. tadpoles were found in ponds or small pools away from streams but only at higher elevations. Acknowledgments JEC thanks Robert F. Inger, Michael O. Dillon, and Abundio Sagastegui for en- couraging his work at Monte Seco and for discussions and logistic support. Field work was supported by the Field Museum of Nat- ural History and the Academy of Natural Sciences, and was ably assisted by Raul Qui- roz, José Santisteban, José Guevara B., He- lena Siesniegas, and Segundo Leiva. Local inhabitants of the Monte Seco area, espe- cially Selzo Gil, Ing. Aldo Maquina, and the family of Francisco Quiroz, made the field work possible. JEC was supported by the National Science Foundation through grant BSR 84-00116 during the 1987 field work. We also thank Ted Kahn for preparing the drawings in Figs. 4, 5, 7, 8, 9; Rex Cocroft for discussions about Centrolenella vocali- zations and review of the manuscript; Ronn Altig for comments on Centrolenella larvae and review of the manuscript; George Stey- skal for providing advise on derivations of the species names; Ron Crombie for notable suggestions for the specific epithets (though their meanings did not allow their use in VOLUME 103, NUMBER 3 print); David Lellinger for identification of a fern; Cristian Ruiz for help with the Span- ish resumen; and Aaron Bauer, James Han- ken, John Harshbarger, Ron Heyer, Dave Wake, and George Zug for discussing inter- calary elements with us. We extend special thanks to David C. Cannatella for a detailed critique of the manuscript, for sharing with us and allowing us to cite some of his un- published observations, and for examining specimens for us in the LSUMZ collections. Literature Cited Aichinger, M. 1987. Freilandbeobachtungen zum Fortpflanzungsverhalten von Hyla brevifrons Duellman & Crump, 1974.—Salamandra 23: 269-276. Altig, R. 1970. A key to the tadpoles of the conti- nental United States and Canada.— Herpetolo- gica 26:180-207. Cadle, J. E. 1989. A new species of Coniophanes (Serpentes: Colubridae) from northwestern Peru.— Herpetologica 45:41 1-424. Cannatella, D. A. 1980. Two new species of Centro- lenella from Bolivia (Anura: Centrolenidae).— Proceedings of the Biological Society of Wash- ington 93:714—724. ——.,, & W. E. Duellman. 1982. Two new species of Centrolenella, with a brief review of the genus in Peru and Bolivia.—Herpetologica 38:380- 388. , & W. W. Lamar. 1986. Synonymy and dis- tribution of Centrolenella orientalis with notes on its life history (Anura: Centrolenidae).— Journal of Herpetology 20:307-317. Cochran, D. M., & C. J. Goin. 1970. Frogs of Co- lombia.— United States National Museum Bul- letin 288:1-605. Dingerkus, G., & L.D. Uhler. 1977. Enzyme clearing of alcian blue stained whole small vertebrates for demonstration of cartilage. — Stain Technol- ogy 52:229-232. Dowling, H. G., & W.E. Duellman. 1978. Systematic herpetology: a synopsis of families and higher categories. HISS Publications, New York. Drewes, R. C. 1984. A phylogenetic analysis of the Hyperoliidae (Anura): treefrogs of Africa, Mad- agascar, and the Seychelles Islands. — Occasion- al Papers of the California Academy of Sciences. 139:1-70. Duellman, W.E. 1975. On the classification of frogs. — Occasional Papers on the Museum of Natural History, The University of Kansas 42:1-14. 767 1988. Evolutionary relationships of the Am- phibia. Pp 1-34 in B. Fritzsch, M. J. Ryan, W. Wilczynski, T. E. Hetherington, & W. Walko- wiak, eds., The evolution of the amphibian au- ditory system. John Wiley & Sons, New York. —., & P. A. Burrowes. 1989. New species of frogs, Centrolenella, from the Pacific versant of Ec- uador and southern Colombia. — Occasional Pa- pers of the Museum of Natural History, The University of Kansas 132:1-14. —.,,&R.O.deSa. 1988. Anew genus and species of South American hylid frog with a highly mod- ified tadpole.— Tropical Zoology 1:117-136. —, & A. H. Savitzky. 1976. Aggressive behavior in a centrolenid frog, with comments on terri- toriality in anurans. — Herpetologica 32:401—404. —., & L. Trueb. 1986. Biology of amphibians. McGraw-Hill Book Company, New York, 670 pp. Eaton, T. H. 1958. An anatomical study of a neo- tropical tree frog, Centrolene prosoblepon (Sali- entia: Centrolenidae).— The University of Kan- sas Science Bulletin 39:459-472. Flores, G. 1985. A new Centrolenella (Anura) from Ecuador, with comments on nuptial pads and prepollical spines in Centrolenella.—Journal of Herpetology 19:313-320. 1987. A new Centrolenella from the Ama- zonian lowlands of Colombia. — Journal of Her- petology 21:185-—190. —.,& R. W. McDiarmid. 1989. Two new species of South American Centrolenella (Anura: Cen- trolenidae) related to C. mariae.—Herpetolo- gica 45:401-411. Frost, D. R. (ed.). 1985. Amphibian species of the world, a taxonomic and geographical reference. Association of Systematics Collections, Law- rence, Kansas, 732 pp. Goin, C.J. 1964. Distribution and synonymy of Cen- trolenella fleischmanniin northern South Amer- ica. — Herpetologica 20:1-8. Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on iden- tification. — Herpetologica 16:183-190. Ham, A. W., & D. H. Cormack. 1979. Histophysi- ology of cartilage, bone, and joints. J. B. Lip- pincott Company, Philadelphia. Jacobson, S. K. 1985. Reproductive behavior and male mating success in two species of glass frogs (Centrolenidae).— Herpetologica 41:396—-404. Jungfer, K.-H. 1988. Froschlurche von Fortuna, Pan- ama II. Hylidae (2), Centrolenidae, Dendroba- tidae.— Herpetofauna 10(56):6—-12. Koepcke, H. W. 1961. Syn6dkologische studien an der westseite der peruanischen Anden.— Bonner Geographische Abhandlungen 29:1-—320. Leviton, A. E., R. H. Gibbs, Jr., E. Heal, & C. E. 768 Dawson. 1985. Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology.—Copeia 1985:802-832. Lieb, C.S. 1988. Systematic status of the neotropical snakes Dendrophidion dendrophis and D. nu- chalis (Colubridae).— Herpetologica 44:162—175. Lynch, J. D. 1969. Taxonomic notes on Ecuadorian frogs (Leptodactylidae: Eleutherodactylus).— Herpetologica 25:262-274. —., & W. E. Duellman. 1973. A review of the centrolenid frogs of Ecuador, with descriptions of new species.— Occasional Papers of the Mu- seum of Natural History, The University of Kansas 16:1-66. , P. M. Ruiz, & J. V. Rueda. 1983. Notes on the distribution and reproductive biology of Centrolene geckoideum Jiménez de la Espada in Colombia and Ecuador (Amphibia: Centroleni- dae).—Studies on Neotropical Fauna and En- vironment 18:239-243. McDiarmid, R. W., & K. Adler. 1974. Notes on ter- ritorial and vocal behavior of neotropical frogs of the genus Centrolenella. —Herpetologica 30: 75-78. Myers, C. W., & W. E. Duellman. 1982. A new spe- cies of Hyla from Cerro Colorado, and other tree frog records and geographical notes from western Panama.—American Museum Novi- tates 2752:1-32. Noble, G. K. 1920. Two new batrachians from Co- lombia.— Bulletin of the American Museum of Natural History 42:441-446. 1924. Some neotropical batrachians pre- served in the United States National Museum with a note on the secondary sexual characters of these and other amphibians. — Proceedings of the Biological Society of Washington 37:65-72. 1925. An outline of the relation of ontogeny to phylogeny within the Amphibia. I.—Amer- ican Museum Novitates 165:1-17. 1931. The Biology of the Amphibia. Mc- Graw-Hill Book Company, New York. 577 pp. ——.,, & M. E. Jaeckle. 1928. The digital pads of the tree frogs. A study of the phylogenesis of an adaptive structure.— Journal of Morphology and Physiology 45:259-292. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Peters, J. A. 1964. Dictionary of Herpetology. Hafner Publishing Company, New York. 392 pp. Poole, A. R., Y. Matsui, A. Hinek, & E. R. Lee. 1989. Cartilage macromolecules and the calcification of cartilage matrix.—The Anatomical Record 224:167-179. Ruiz-Carranza, P. M., J. Hernandez-Camacho, & M. C. Ardila-Robayo. 1986. Una nueva especie Colombiana del genero Centrolene Jiménez de la Espada 1872 (Amphibia: Anura) y redefini- cion del genero.—Caldasia 15(71-—75):43 1-444. Savage, J. M. 1967. A new tree-frog (Centrolenidae) from Costa Rica.—Copeia 1967:325-331. ——, & W.R. Heyer. 1967. Variation and distri- bution in the tree-frog genus Phyllomedusa in Costa Rica, Central America.—Beitrége zur Neotropischen Fauna 5(2):111-131. —, & P. H. Starrett. 1967. A new fringe-limbed tree-frog (Family Centrolenidae) from lower Central America.—Copeia 1967:604—609. Starrett, P. H., & J. M. Savage. 1973. The systematic status and distribution of Costa Rican glass-frogs, genus Centrolenella (Family Centrolenidae), with description of a new species.—Bulletin of the Southern California Academy of Sciences 72: 57-78. Taylor, E.H. 1949. Costa Rican frogs of the genera Centrolene and Centrolenella.—The University of Kansas Science Bulletin 33:257—270. 1951. Two new genera and a new family of tropical American frogs.—Proceedings of the Biological Society of Washington 64:33—40. Weberbauer, A. 1936. Phytogeography of the Peru- vian Andes.—Field Museum of Natural His- tory, Botanical Series, 13. Flora of Peru, Part 1: 13-81. (JEC) Department of Herpetology, Acad- emy of Natural Sciences, 1 9th and the Park- way, Philadelphia, Pennsylvania 19103; and (RWM), United States Fish and Wildlife Service, Biological Survey, National Mu- seum of Natural History, Washington, D.C. 20560. VOLUME 103, NUMBER 1 769 INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE % BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON, SW7 5BD Applications published in the Bulletin of Zoological Nomenclature The following applications were published on 27 March 1990 in Vol. 47, Part 1 of the Bulletin of Zoological Nomenclature. Comment or advice on these applica- tions is invited for publication in the Bulletin and should be sent to the Executive Secretary, ICZN, % The Natural History Museum, Cromwell Road, London SW7 SBD. Case No. 2714 Pleuractis Verrill, 1864 (Cnidaria, Anthozoa): proposed designation of Fun- gia paumotensis Stutchbury, 1833 as the type species, with conser- vation of Lobactis Verrill, 1864. 2547 CYMATIINAE Iredale, 1913 (1854) (Mollusca, Gastropoda) and CYMA- TIINAE Walton in Hutchinson, 1940 (Insecta, Heteroptera): pro- posal to remove the homonymy. 2641 Limax fibratus Martyn, 1784 et Nerita hebraea Martyn, 1786 (actuellement Placostylus fibratus et Natica hebraea; Mollusca, Gastropoda): con- servation proposée pour les noms spécifiques; et Placostylus Beck, 1837: désignation proposée de L. fibratus comme espéce-type. Proptera Rafinesque, 1819 (Mollusca, Bivalvia): proposed conservation. Mirochernes Beier, 1930 (Arachnida, Pseudoscorpionida): proposed confir- mation of Chelanops dentatus Banks, 1895 as the type species. Holostaspis subbadius var. robustulus Berlese, 1904 (currently Macrocheles robustulus; Arachnida, Acarina): proposed conservation as the cor- rect spelling of the specific name. Bathynomus A. Milne Edwards, 1879 (Crustacea, Isopoda): proposed pre- cedence over Palaega Woodward, 1870. Carcinochelis Fieber, 1861 (Insecta, Heteroptera): proposed designation of Carcinochelis alutaceus Handlirsch, 1897 as the type species. Steno attenuatus Gray, 1846 (currently Stenella attenuata; Mammalia, Ce- tacea): proposed conservation of the specific name. Mammuthus Brookes, 1828 (Mammalia, Proboscidea): proposed conserva- tion, and Elephas primigenius Blumenbach, 1799 (currently Mam- muthus primigenius): proposed designation as the type species of Mammuthus, and designation of a neotype. 770 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE Opinions published in the Bulletin of Zoological Nomenclature The following Opinions were published on 27 March 1990 in Vol. 47, Part 1 of the Bulletin of Zoological Nomenclature. Opinion No. 1567 Nonion de Montfort, 1808 (Foraminiferida): Nautilus faba Fichtel & Moll, 1798 designated as the type species. 1568 Hanzawaia Asano, 1944 (Foraminiferida): conserved. 1569 Calcarina d’Orbigny, 1826 (Foraminiferida): conserved. 1570 Dendritina d’Orbigny, 1826 (Foraminiferida): conserved. 1571 Planularia Defrance, 1826 (Foraminiferida): conserved. 1572 Nautilus repandus Fichtel & Moll, 1798 (currently Eponides repandus; Fo- raminiferida): neotype replaced by rediscovered holotype. 1573 Madrepora limax Esper, 1797 (currently Herpolitha limax) and Fungia tal- pina Lamarck, 1801 (currently Polyphyllia talpina; both Cnidaria, Anthozoa): specific names conserved. 1574 Sphaeroma hookeri Leach, 1814 (currently Lekanesphaera hookeri; Crus- tacea, Isopoda): specific name conserved. 1575 Coenobita Latreille, 1829 (Crustacea, Decapoda): conserved. 1576 Palaemon longirostris H. Milne Edwards, 1837 (Crustacea, Decapoda): spe- cific name conserved. 1577 Hydrobius Leach, 1815 (Insecta, Coleoptera): Dytiscus fuscipes Linnaeus, 1758 conserved as type species, and Berosus Leach, 1817 (Insecta, Coleoptera): conserved. 1578 Vespa triangulum Fabricius, 1775 (currently Philanthus triangulum; Insecta, Hymenoptera): specific name conserved. 1579 Pycinaster magnificus Spencer, 1913 (Echinodermata, Asteroidea): specific name conserved. 1580 Cordylodus? dubius Rhodes, 1953 (currently Distomodus dubius; Conodonta): specific name conserved. 1581 HAydrolycus Muller & Troschel, 1844 (Osteichthyes, Cypriniformes): Hydroc- yon scomberoides Cuvier, 1819 confirmed as the type species. 1582 Ictiobus Rafinesque, 1820 (Osteichthyes, Cypriniformes): conserved. 1583 Scorpaenichthys marmoratus (Osteichthyes, Scorpaeniformes): Ayres, 1854 to be taken as the author of the specific name. 1584 Ameiurus Rafinesque, 1820 (Osteichthyes, Siluriformes): Si/urus lividus Ra- finesque, 1820 designated as the type species. 1585 Ascalabotes gigas Bocage, 1875 (currently Tarentola gigas; Reptilia, Squa- mata): specific name conserved. 1586 Euryotis brantsii A. Smith, 1834 (currently Parotomys brantsii; Mammalia, Rodentia): specific name conserved. PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 771 BIOLOGICAL SOCIETY OF WASHINGTON 117TH Annual Meeting, 8 May 1990 The meeting was called to order by Kris- tian Fauchald, President, at 12:30 p.m. in the Waldo Schmitt Room, National Mu- seum of Natural History. Kristian announced the election results. The new elected officers and council mem- bers are listed on the inside front cover of this issue. Mike Vecchione, Treasurer, presented the report of past Treasurer, Don Wilson. In- come from dues in 1989 was slightly higher than that of 1988, and income from sub- scriptions, sales of past issues of the Pro- ceedings and Bulletins, and page charges was substantially lower. Total income for 1989 was $83,661 and total expenditures were $81,673. The proposed budget for 1990 es- timates income of $91,000 and expendi- tures of $97,000, predicting a potential def- icit of $6000. Brian Robbins, Editor, then presented his report. The four issues of Volume 102 of the Proceedings were published on 29 March, 28 June, 18 October, and 19 De- cember 1989, containing a total of 104 pa- pers and 1085 pages. A copy of the new *““Guidelines for Manuscripts” was sent to all members as a Supplement to the first issue of Volume 103. The One-Hundred- Year Index is nearing completion and will be published camera-ready as a Supplement to Volume 100. Rafael Lemaitre replaced Ray Manning as one of the Associated Ed- itors for Invertebrate Zoology. Kristian noted that Austin Williams re- placed Dave Pawson as Back-Issues Cus- todian and announced that he has appoint- ed an ad-hoc committee to review problems associated with storage and disposition of back issues. A motion was made and seconded that the meeting be adjourned; the new Presi- dent, Leslie W. Knapp, adjourned the meet- ing at 1:15 p.m. Respectfully submitted, G. 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Literature citations in the text should be in abbreviated style (author, date, page), except in botanical synonymies, with unabbreviated citations of journals and books in the Literature Cited sections. Direct quotations in the text must be accompanied by author, date, and pagination. The establishment of new taxa must conform with the requirements of the appropriate international codes of nomenclature. When appropriate, accounts of new taxa must cite a type specimen deposited in an institutional collection. Examples of journal and book citations: Eigenmann, C. H. 1915. The Cheirodontidae, a subfamily of minute characid fishes of South America.— Memoirs of the Carnegie Museum 7(1):1—99. Ridgely, R. S. 1976. A guide to the birds of Panama. Princeton, New Jersey, Princeton University Press, 354 pp. Olson, S. L. 1973. The fossil record of birds. Pp. 79-238 in D. Farner, J. King, and K. Parkes, eds., Avian biology, volume 8. Academic Press, New York. Figures and tables with their legends and headings should be self-explanatory, not requiring reference to the text. Indicate figure and table placement in pencil in the margin of the manu- script. Plan illustrations in proportions that will efficiently use space on the type bed of the Proceedings. Original illustrations should not exceed 15 x 24 inches. Figures requiring solid black backgrounds should be indicated as such when the manuscript is submitted, but should not be masked. CONTENTS Comments on the nomenclature of some neotropical bats (Mammalia: Chiroptera) Alfred L. Gardner and Carolyn S. Ferrell Redescription of Polyipnus fraseri Fowler, 1934 (Teleostei: Stomiiformes: Sternoptychidae), with remarks on paedomorphosis Antony S. Harold A new, unusually sexually dimorphic species of Bryconamericus (Pisces: Ostariophysi: Charac- idae) from the Peruvian Amazon Richard P. Vari and Darrell J. Siebert Moojenichthys Miranda-Ribeiro (Pisces: Ostariophysi: Characidae), a phylogenetic reappraisal and redescription Ricardo M. C. Castro and Richard P. Vari Scorpaenodes immaculatus, a new species of scropionfish (Osteichthyes: Scorpaenidae) from Walters Shoals, Madagascar Ridge Stuart G. Poss and Bruce B. Collette Cyphocharax pantostictos, a new species (Pisces: Ostariophysi: Characiformes: Curimatidae) from the western portions of the Amazon basin __ Richard P. Vari and Ramiro Barriga S. The genus Axianassa (Crustacea: Decapoda: Thalassinidea) in the Americas Brian Kensley and Richard Heard On the crayfishes (Decapoda: Cambaridae) of the Neches River basin of eastern Texas with the descriptions of three new species Horton H. Hobbs, Jr. Panopeus margentus, a new crab from the Argentine warm temperate subregion (Decapoda: Xanthidae) Austin B. Williams and E. E. Boschi A new species of Chaceon from New Zealand (Crustacea: Decapoda: Geryonidae) Raymond B. Manning, Elliot W. Dawson, and W. Richard Webber A new crayfish (Decapoda: Cambaridae) from southeastern Texas Horton H. Hobbs, Jr., and H. H. Hobbs III Pomatogebia, a new genus of thalassinidean shrimps from western hemisphere tropics (Crus- tacea: Upogebiidae) Austin B. Williams and Nguyen Ngoc-Ho Gitana dominica, a new species from the Caribbean Sea (Amphipoda: Amphilochidae) James Darwin Thomas and J. L. Barnard The Japanese amphipod genus Eoniphargus, rediscovered in a South Korean cave Jan H. Stock and Young Won Jo Taxonomic remarks on Schizobopyrina Markham, 1985, with the description of S. bruscai (Crustacea: Isopoda: Bopyridae) Ernesto Campos and Alma Rosa de Campos Tridentella williamsi, a new species of isopod crustacean from the British Virgin Islands, western Atlantic (Flabellifera: Tridentellidae) Paul M. Delaney Description of the cryptoniscium larva of E ntophilus omnitectus Richardson, 1903 (Crustacea: Isopoda: Epicaridea) and records from the Gulf of Mexico Daniel L. Adkison and Sneed B. Collard A new species of isopod, Aega (Rhamphion) francoisae (Flabellifera: Aegidae), from the cloaca of an ascidian from the Galapagos Islands Regina Wetzer Supplementary description and phylogenetic significance of Arcoscalpellum conradi (Gabb) (Cirripedia: Scalpellidae) from the Paleocene Vincentown formation of New Jersey Victor A. Zullo Metacyclops leptopus totaensis, new subspecies (Crustacea, Copepoda) from Lago de Tota, Colombia Janet W. Reid, Javier A. Molina Arevalo, and Manuel M. Fukushima A new genus and species of Polychaeta commensal with a deep-sea thyasirid clam James A. Blake Exogone acerata (Exogoninae: Syllidae: Polychaeta), a new species without antennae from the Mediterranean Sea Guillermo San Martin and Julio Parapar A revision of the subgenus Phascolosoma (Sipuncula: Phascolosoma) Norma J. Cutler and Edward B. Cutler Correction of Ijima’s (1927) list of Recent hexactinellid sponges (Porifera) Henry W. Reiswig Two new species of Centrolenella (Anura: Centrolenidae) from northwestern Peru John E. Cadle and Roy W. McDiarmid International Commission on Zoological Nomenclature: Applications and Opinions Biological Society of Washington: 117th Annual Meeting 501 509 516 525 543 550 558 SS 598 602 608 614 617 624 633 643 649 655 663 674 “681 687 691 Ta 746 769 771 7 "PROCEEDINGS 4 _ OF THE : WA SHING TON VOLUME 103 NUMBER 4 19 DECEMBER 1990 oa ee THE BIOLOGICAL SOCIETY OF WASHINGTON 1990-1991 Officers President: Leslie W. Knapp Secretary: G. David Johnson President-elect: Storrs L. Olson Treasurer: Michael Vecchione Elected Council Roger F. Cressey, Jr. Janet W. Reid Janet R. Gomon Wayne C. Starnes Robert Hershler Jeffery T. Williams Custodian of Publications: Austin B. Williams PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: George C. Steyskal Invertebrates: Stephen D. Cairns Frank D. Ferrari Plants: David B. Lellinger Rafael Lemaitre Insects: Wayne N. 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Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, National Museum of Natural History, Washington, D.C. 20560. THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 773-783 THE IDENTITY OF FANNYELLA ROSSI J. E. GRAY (COELENTERATA: OCTOCORALLIA) Frederick M. Bayer Abstract.—The conspecificity of Ascolepis splendens Thomson & Rennet, 1931, with Fannyella rossii J. E. Gray, 1873, is established. A syntype of Ascolepis splendens, register number G-13237 in the Australian Museum, Syd- ney, is selected as lectotype of the species and simultaneously established as neotype of Fannyella rossii. A revised description of the species is provided, illustrated by scanning electron micrographs, in comparison with the original illustrations of Fannyella rossii and Ascolepis splendens. During the Antarctic voyage of HMS Er- ebus and Terror 1839-1843 under the com- mand of Captain Sir James Clark Ross, a dredge haul made on 19 January 1841 ata depth of 270 fathoms (=494 m) at 72°31’S, 173°39’E, in the vicinity of Cape Hallett, Ross Sea, obtained a “‘new species of Prim- noa, which I name P. Rossii,”’ along with another gorgonian, a scleractinian coral, and some bryozoans (Stokes 1847:258—259). Ross entrusted these, together with some other invertebrates from the expedition, to Charles Stokes for investigation. In 1846, Stokes sent drawings and notes on the spec- imens to Ross (Stokes 1847:259), who, in turn, gave the drawings to J. E. Gray. They were not published in the volume on the zoology of the expedition, remaining in Gray’s custody until 1872, when he pre- sented them together with descriptive text of his own at the meeting of the Zoological Society of London June 18, 1872. Accord- ing to Gray (1873:744), the specimens by that time had been “lost to science.” Although Stokes (1847) specifically dis- cussed his new Primnoa rossii, he referred only to the then unpublished drawings with- out giving any characters to make the name nomenclaturally available. Gray (1873), in publishing the drawings, established a new genus, Fannyella, and used the name rossii proposed by Stokes for the species. He seems to have been unaware of the paper pub- lished by Stokes. Good though the drawings are, they have not been associated with any gorgonians obtained by later Antarctic ex- peditions. Numerous specimens provisionally iden- tified as Ascolepis splendens Thomson & Rennet, 1931, aspecies first obtained at four localities between 92°10’E and 145°21’E by the Australasian Antarctic Expedition, have been taken at various localities around the Antarctic continent by expeditions of the U.S. Antarctic Research Program. This ma- terial agrees with the published drawing of Fannyella rossii, leaving no doubt that they represent the same species, and comparison with a syntype of Ascolepis splendens con- firms that the two species also are identical. Although the generic name Ascolepis has been used in keys (see below), the species A. splendens Thomson & Rennet, 1931, has not been reported again in the literature since 1931, so no justification exists for retaining it as the valid name for the genus and species in preference over the senior synonym Fan- nyella rossii Gray, 1873, also used only once. In order to place this undoubted but sub- jective synonymy on an objective basis, the illustrated syntype of A. splendens Thomson & Rennet, 1931 (Australian Museum, Syd- ney, register no. G-13237), is hereby si- multaneously selected as lectotype of As- 774 colepis splendens and designated as neotype of Fannyella rossii Gray, 1873. Fannyella J. E. Gray, 1873 Primnoa (part). —Stokes, 1847:260 Fannyella Gray, 1873:744. (Type species, Fannyella rossii Gray, 1873, by mono- typy.) Ascolepis Thomson & Rennet, 1931:20.— Bayer, 1981:936 (in key only); 1982:120- 122 (passim). — Bayer & Stefani, 1989:454 (in key only). (Type species, Ascolepis splendens Thomson & Rennet, 1931, here designated.) Diagnosis. —Dichotomously branched Primnoidae with verticillate polyps pro- tected by sclerites arranged in distinct lon- gitudinal rows on abaxial and lateral aspects of body; adaxial surface with fewer sclerites, in some cases unarmed except for a few small scales immediately below opercular and cir- cumopercular scales. Body sclerites verti- cally overlapping, exposed part smooth and distinctly differentiated from strongly tu- berculate covered part. Circumopercular (i.e., marginal) scales folding over bases of opercular scales. Outer layer of coenen- chyme with thick polygonal, rounded, or irregular plates and many smaller, rounded scales; inner layer with irregularly tuber- culate discoidal forms and spindles, which occasionally are triradiate or branched. Fannyella rossii Gray, 1873 Figs. 1-6 Primnoa Rossii Stokes, 1847:260, 261 (no- men nudum). Fannyella rossii Gray, 1873:745, pl. 62, figs. 1-3 (Antarctic Ocean). ?Caligorgia antarctica Kiikenthal, 1912:321, figs. 27-35, pl. 21, fig. 10 (Gauss-Station, 385 m). ?Caligorgia ventilabrum.—Gravier, 1914: 85, figs. 109-119; pl. 6, fig. 30 (Marguerite Bay, Antarctic Peninsula, 176-230 m).— PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Molander, 1929:60 (Seymour I., Graham Land, 150 m; South Georgia, 75 m). Not Caligorgia ventilabrum Studer, 1879: 647, pl. 2, fig. 12 (North of New Zealand, 165 m).—Versluys, 1906:74, figs. 83, 84 (illustrates polyp from Studer’s type spec- imen). Ascolepis splendens Thomson & Rennet, 1931:20, pl. 9, figs. 6, 7; pl. 10, figs. 1, 2; pl. 11, fig. 6 (Off Wilkes Land, from Ade- lie Coast west to Davis Sea, 46-582 m). Gray (1873:745) gave no description of the species, but a combined description of genus and species (1873:744), which runs as follows: “Coral slightly furcately branched; branches club-shaped, enlarging up- wards, and then rapidly contracting at the tip; polypiferous cells many, in numerous close concentric rings, forming regular whorls round the branches, the cells ob- long, cylindrical, contracted at the base, and each covered with six longitudinal series of transverse oblong hexangular scales, truncated at top and closed with elongated more or less acute scales, con- verging to a point when the animal is withdrawn; axis covered with small scales.” The club-shaped branches are related to the small size of the very young colony with only two bifurcations (Fig. 2d), and are not characteristic of fully developed colonies. As the adaxial surface of the polyps is naked save for a few small scales just below the operculum, Gray’s description of “‘six lon- gitudinal series” of scales, rather than the more usual eight, is basically correct. The more exhaustive description of As- colepis splendens provided by Thomson & Rennet (1931:20) is more nearly definitive. The specimen from Aurora station 2, 66°55'S, 145°21’E, 318 fathoms (=582 m), 28 Dec 1913 (Australian Museum no. G-13237) conforms with the specifications VOLUME 103, NUMBER 4 cited by Thomson & Rennet (1931:20) and clearly is the specimen upon which the de- scription of Ascolepis splendens is based. A photograph of this specimen is given here- with (Fig. 1). The drawing reproduced on Thomson & Rennet’s Plate IX, Fig. 6, ap- pears to be a somewhat simplified repre- sentation either of the detached branch ac- companying the complete specimen, or of a branch of the complete colony as isolated by the artist. The original description of A. splendens is accurate except in regard to the form of the polyp sclerites: ““The striking peculiarity is that the main sclerites of the vertical rows have a distinct ascus-like or chalice-like form. That is to say, the basal portion is the substantial very warty support of a delicate cup whose cavity is open to the exterior! The delicate edges of the cups are weakly notched, sometimes almost entire. The cup of the sclerite is broader than the substantial knobbed support, so that the appearance is somewhat like a short-stalked chalice or fruit-basket.’’ The authors go on to say that “We must emphasise the point that the specimen is on the whole like a Caligorgia, but its hollow ascus-like sclerites are very far from the ctenoid-scale type” (Thomson & Rennet 1931:20-21). As is immediately apparent from scan- ning electron micrographs (Figs. 3-5), the “delicate” cups are an optical illusion. As the calcite of which the sclerites are com- posed is almost glassy clear, only the edges of the smooth outer portion of the body sclerites are visible as a bright line under the light microscope; the fact that it is solid is not easily detected in a bulk preparation of sclerites. It is, however, readily perceived in intact polyps viewed with a binocular Fig. 1. 775 dissecting microscope, and it is a mystery why this was not noticed during preparation of the drawing reproduced by Thomson & Rennet as fig. 7 of Plate [IX (Fig. 2b). When sclerites seen by Thomson & Rennet as transparent objects are viewed by SEM as Opaque objects, it can be seen that the sur- face of the exposed part of the body sclerites is sometimes very weakly concave, but as often it is flat or slightly convex and orna- mented by rows of low, smooth, elongate granules (Figs. 4, 5). The abaxial and lateral opercular scales are broad and have a marginal point located more or less off center, but the adaxial oper- culars are narrow and toothlike. The outer lateral rows of body sclerites are well de- veloped and composed of nearly as many scales as the abaxial rows, but these are more rounded in outline and not nearly so “‘ascus- like’”’ as the abaxial sclerites (Fig. 5). The inner lateral rows are present but short, comprised of only 2 or 3 scales below the operculars, and the adaxial rows are reduced usually to a single small marginal below each opercular (Fig. 4). In order to show the true form of the scler- ites illustrated by Thomson & Rennet, the various types included in their fig. 6 of Plate XI (Fig. 2c) have been photographed under the SEM and arranged so as to reconstruct their illustration (Fig. 6), which appears to have been drawn free-hand. Distribution. —Circum-Antarctic, 46-852 m. The southernmost locality reported thus far is from 77°13.7'S, 40°03.8'W, off Filch- ner Ice Shelf (Polarstern ‘“‘Antarktis I’ sta. 180); the northernmost record is from 54°22.1'S, 03°38.2’'E, off Bouvet Is. (Marion Dufresne sta. 17, haul CM 30), which also is the deepest record. — Fannyella rossii J. E. Gray, 1873. Neotype colony, natural size. Lectotype of Ascolepis splendens Thomson & Rennet, 1931. Australasian Antarctic Expedition, HMS Aurora sta. 2. Australian Museum register no. G-13237. — eee ig Ap Se ; Nees wR “a4 Pees; = Z © val ©) a ja 7) < S ay O > aed Sl Oo fe) N é o) fe) = 2 A eal rr oa fy O DN O Ss al ea aa Oo fe) 4 ay) a » o eg ——— a eae Pa . 7 —ee ‘a ge Mle See 777 VOLUME 103, NUMBER 4 TORS ry, ,, og KECESERE SEER 3 on? neler on ecETSEE Ep, = ai PEED oy eeeeeseeeeit riers SPAT: B37. ory 2 EEE 2 sere, Be peih3333103932 032d ree” a—c.—Ascolepis splendens Thomson & Rennet, 1931. Original illustrations. a, Copy of pl. IX, fig. 6; Fig. 2. b, Copy of pl. IX, fig. 7; c, Copy of pl. XI of pl. XLII, figs. 1-3. fig. 6. —d-f, Fannyella rossii Gray, 1873. Original illustrations. Copies > 778 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON +N. 1790 = «10K¥ SS ee an 1798 = 18KY Fig. 3. Fannyella rossii J. E. Gray, 1873, neotype =Ascolepis splendens Thomson & Rennet, 1931, lectotype. Stereomicrographs of whorls of polyps. VOLUME 103, NUMBER 4 68mm 1798 soar x y op REE aod ~7 see oe ene ewes »oomm Fig. 4. Fannyella rossii J. E. Gray, 1873, neotype =Ascolepis splendens Thomson & Rennet, 1931, lectotype. Top, Whorl of polyps. Bottom, Adaxial surface of isolated polyp. Stereomicrographs. Fannyella rossii is one of the most abun- U.S. Antarctic Research Program. Spec- dant and widely distributed gorgonians of imens in the U.S. National Museum of Nat- the Antarctic and sub-Antarctic fauna. It ural History: has been taken by various expeditions at the Ross Sea, off Cape Hallett: 72°32'S, following locations. 171°26’E, 337-329 m, USNS Eltanin sta. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON »oomm 1791 1akKy Fig. 5. Fannyella rossii J. E. Gray, 1873, neotype =Ascolepis splendens Thomson & Rennet, 1931, lectotype. Stereomicrographs of isolated polyp. Top, Lateral view. Bottom, Abaxial view. 1875, 15 Jan 1967. Five large more or less Ross Sea, off Cape Washington: 75°01’S, complete colonies without holdfasts, and 168°23’E, 334-335 m, USNS Eltanin sta. detached branches, USNM 82949 (SEM _ 2036, 18 Jan 1968. One large colony lacking 1417, 1427). holdfast, USNM 82078 (SEM 1471-1473). VOLUME 103, NUMBER 4 781 Fig. 6. Fannyella rossii J. E. Gray, 1873, neotype =Ascolepis splendens Thomson & Rennet, 1931, lectotype. Sclerites. Reconstruction of Thomson & Rennet’s plate XI, fig. 6. Scanning electron micrographs. Ross Sea: 76°00’S, 176°48’W, 566-569 m, USNS Eltanin sta. 2045, 20 Jan 1968. One colony and detached branches, USNM 82079 (SEM 1407). Ross Sea: 76°25’S, 170°24'W, 568 m, USNS Eltanin sta. 2075, 30 Jan 1968. One colony without holdfast, and 1 detached branch, USNM 82109. Ross Sea, Bay of Whales: 76°08’S, 165°04'W, 494-498 m, USNS Eltanin sta. 2097, 4 Feb 1968. 2 small incomplete spec- imens, USNM 82950. South Shetland Islands: 61°18’S, 56°09’'W, 220-240 m, USNS Eltanin sta. 410, 31 Dec 1962. One colony lacking holdfast, USNM 82072 (SEM 1404, 1405). South Shetland Islands: 61°25’S, 56°30'W, 164 fathoms (=300 m), USNS Eltanin sta. 993, 13 Mar 1964. One complete colony with holdfast attached to coral, USNM 60342. South Shetland Islands: 63°26'S, 62°15'W, 119-124 m, R/V Hero cruise 691 sta. 26, 10 Feb 1969. One large colony nearly com- plete but lacking holdfast, and 1 detached branch, USNM 82952 (SEM 1448, 1449). 782 Antarctic Peninsula, vicinity of Brabant Island: 64°21'24”S, 61°28'12”W, 110-155 m, R/V Hero sta. 21-1, 23 Mar 1982, coll. G. Hendler. One large colony complete with holdfast, USNM 77360 (SEM 1798, 1799). Antarctic Peninsula: 64°46'28’S, 63°26.5'W, 100-150 m, R/V Hero cruise 731 sta. 1944, 11 Mar 1973. Three colonies lacking holdfasts, USNM 82071. Antarctic Peninsula: 64°50’S, 63°12’W, 155 fath. (=283 m); Eastwind sta. 66-006, haul 3, coll. D. F. Squires and D. L. Pawson, 29 Jan 1966. Two colonies, USNM 58151 (SEM 303). Antarctic Peninsula: 65°03.71'S, 63°57.05'W, 360-375 m, R/V Hero cruise 833 sta. 8-2, 10 March 1983. One colony lacking holdfast, USNM 82082. Antarctic Peninsula: 65°06.7’S, 65°00.7’W, 100-180 m, R/V Hero cruise © 731 sta. 1884. One colony extensively over- grown by hydroids, USNM 82118. Antarctic Peninsula: 65°54.5'S, 65°15.5'W, 246-270 m, R/V Hero cruise 824 sta. 5-1, 16 Mar 1982. Two incomplete colonies, USNM 82119 (SEM 1440-1442). Weddell Sea off Filchner Ice Shelf: 76°50'S, 40°55’W, 513 m, IWSOE Univer- sity of Connecticut, USS Glacier cruise 2 sta. 0006, 1 Mar 1969. One small branch, USNM 82951. R/V Polarstern. Specimens in Sencken- berg Museum Frankfurt: Off Atka Iceport, Princess Martha Coast: 70°30.3’S, 08°04.0’W, 261-263 m, cruise “‘Antarktis I’ sta. 220, 1 Mar 1983. One large, slender colony without holdfast. Weddell Sea, off Nantucket Inlet, Lassiter Coast, Antarctic Peninsula: 74°49.9’S, 61°08.3'W, 637 m, cruise ‘‘Antarktis II/4”’ sta. 386, 31 Jan 1984. Two small colonies without holdfasts. Weddell Sea, off Nantucket Inlet, Lassiter Coast, Antarctic Peninsula: 74°57.3’S, 60°31.4'W, 662 m, cruise “‘Antarktis II/4” sta. 378, 31 Jan 1984. Six colonies without holdfasts. Weddell Sea, off Cape Adams, Antarctic PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Peninsula: 75°00.1’S, 59°38.0’'W, 621 m, cruise ““Antarktis II/4”’ sta. 372, 30 Jan 1984. Three colonies without holdfasts and 6 frag- ments. Weddell Sea, off Gardner Inlet, Antarctic Peninsula: 75°08.5'S, 59°38.1’W, 627 m, cruise ““Antarktis II/4” sta. 369, 30 Jan 1984. Two large and 2 smaller colonies without holdfasts, and many fragments. Weddell Sea: 75°49.2'S, 56°15.1'W, 456 m, cruise ““Antarktis II/4”’ sta. 450, 8 Feb 1984. Two small colonies with holdfast, 8 small colonies without holdfasts, and 10 fragments. Weddell Sea, north of Luitpold Coast, Coats Land: 76°08.4’S, 32°37.6’W, 788 m, cruise ““Antarktis II/4”’ sta. 510, 22 Feb 1984. One large incomplete colony. Weddell Sea: 76°09.7'S, 52°21.4'W, 416 m, cruise ““Antarktis II/4”’ sta. 438, 7 Feb 1984. Two incomplete colonies. Weddell Sea: 76°38.6’S, 52°10.3'W, 304 m, cruise ““Antarktis II/4”’ sta. 341, 26 Jan 1984. One small colony without holdfast. Weddell Sea: 76°52.5’S, 50°40.4’W, 259 m, cruise ““Antarktis II/4”’ sta. 308, 20 Jan 1984. One large incomplete colony. Weddell Sea, off Filchner Ice Shelf: 77°13.7'S, 40°03.8’W, 673-717 m, cruise ‘“‘Antarktis I’? sta. 180, 18 Feb 1983. One large detached branch. R/V Marion Dufresne. Specimen in Mu- seum National d’Histoire Naturelle, Paris: East of Bouvet Is.: 54°22.1'S, 03°38.2’E, 965-852 m, sta. 17, Marinovich trawl 30, 22 Aug 1980. One large colony almost com- plete, with holdfast, and 2 detached branch- es. The original material of Ascolepis splen- dens taken by HMS Aurora of the Austral- asian Antarctic Expedition, now preserved in the Australian Museum Sydney, was ob- tained at station 2, 66°55’S, 145°21’E, 318 fathoms (=582 m), 28 Dec 1913 (AMS no. G-13245 and G-13237 [lectotype]; station 7, 65°42'S, 92°10’E, 60 fathoms (=110 m), 21 Jan 1914 (AMS no. G-13221); station 8, 66°08'S, 94°17'E, 120 fathoms (=219 m), VOLUME 103, NUMBER 4 27 Jan 1914; and at Commonwealth Bay, Adelie Land, in 25-318 fathoms (=47-582 m), 3 Sep 1912 (AMS no. G-13273). Acknowledgments In the preparation of this paper I am in- debted to Mr. Simon Moore (Natural His- tory Museum, London) and Ms. Penny Ber- ents (Australian Museum, Sydney), for the loan of pertinent material. The records of Fannyella rossii from stations of R/V Po- larstern and R/V Marion Dufresne were supplied by Dr. Manfred Grasshoff (Natur- Museum Senckenberg, Frankfurt). As al- ways, I am grateful to Dr. Grasshoff for his perceptive comments and advice in regard to the taxonomic and nomenclatural prob- lems considered in this paper. Dr. Curtis W. Sabrosky, former president of the Interna- tional Commission on Zoological Nomen- clature, provided helpful critical comments on the manuscript, for which I express sin- cere thanks. The scanning electron micro- graphs given here were made by Mr. Walter R. Brown, chief of the Scanning Electron Microscope Laboratory, U.S. National Mu- seum of Natural History, Smithsonian In- stitution. Literature Cited Bayer, F.M. 1981. Key to the genera of Octocorallia exclusive of Pennatulacea (Coelenterata: An- thozoa), with diagnoses of new taxa. —Proceed- ings of the Biological Society of Washington 94: 902-947, figs. 1-80. , 1982. Some new and old species of the prim- noid genus Callogorgia Gray, with a revalida- tion of the related genus Fanellia Gray (Coelen- terata: Anthozoa).— Proceedings of the Biological Society of Washington 95:116-160, figs. 1-29. , & J. Stefani. 1989. Primnoidae (Gorgonacea) de Nouvelle-Calédonie.— Bulletin du Muséum National d’Histoire Naturelle, Paris (4)10(A no. 3):449-5 18, including pls. 1-42. Duncan, F.M. 1937. On the dates of publication of the Society’s ‘Proceedings,’ 1859-1926. With an appendix containing the dates of publication of 783 ‘Proceedings,’ 1830-1858, compiled by the late F. H. Waterhouse, and of the “Transactions,” 1833-1869, by the late Henry Peavot, originally published in P.Z.S. 1893, 1913.—Proceedings of the Zoological Society of London (A)107:71- 84. Gravier,C. 1914. Alcyonaires. Deuxiéme Expédition Antarctique Francaise (1908-1910). Sciences Naturelles: Documents scientifiques. Pp. 1-118, pls. 1-10. Gray, J.E. 1873. Notes on corals from the South and Antarctic seas.—Proceedings of the Scientific Meetings of the Zoological Society of London for the year 1872(part 47):744-747, pls. 62-64. [According to Duncan (1937:72), the date of publication of the Proceedings pp. 737-927 is April 1873; Neave (1939:398) records 1873 as the date of Fannyella.] Kikenthal, W. 1912. Die Alcyonarien der deutschen Siudpolar-Expedition 1901-1903. Deutsche Sudpolar-Expedition 1901-1903, 13 Band.— Zoologie 5(3):289-349, pls. 20-23. Molander, A.R. 1929. Die Octactiniarien.— Further Zoological Results of the Swedish Antarctic Ex- pedition Expedition 1901-1903, 2(2):i-iv + 1- 86, pls. 1-5. Neave, S. A. (ed.) 1939. Nomenclator zoologicus. Zoological Society, London 2:1-1025. Stokes, C. 1847. Remarks on some corals obtained from great depths in the Antarctic Ocean, in a letter from Charles Stokes, Esq., F.R.S., F.G:S., &c., to Captain Sir James C. Ross, R.N.—The Edinburgh New Philosophical Journal, exhib- iting a view of the Progressive Discoveries and Improvements in the Sciences and the Arts 43: 258-262. Studer, Th. 1879. Ubersicht der Anthozoa Alcyon- aria, welche wahrend der Reise S.M.S. Gazelle um die Erde gesammelt wurden.— Monatsber- icht der K6niglich Preussischen Akademie der Wissenschaften zu Berlin 1878:632-688, pls. 1-5. Thomson, J.A., & N.I. Rennet. 1931. Alcyonaria, Madreporaria, and Antipatharia.— Australasian Antarctic Expedition 1911-14. Scientific Re- ports. (Series C)9(3):1—46, pls. 8-14. Versluys, J. 1906. Die Gorgoniden der Siboga Ex- pedition. II. Die Primnoidae.—Siboga-Expedi- tie Monographie 13a. Pp. 1-187, 178 figs. 10 pls. chart. Department of Invertebrate Zoology, Na- tional Museum of Natural History, NHB- 163, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 784-788 MEIOPRIAPULUS FIJIENSTS MORSE (PRIAPULIDA) FROM SOUTH ANDAMAN, ANOTHER EXAMPLE OF LARGE-SCALE GEOGRAPHIC DISTRIBUTION OF INTERSTITIAL MARINE MEIOFAUNA TAXA Wilfried Westheide Abstract.—The interstitial priapulid Meiopriapulus fijiensis Morse, 1981, is described for the first time outside of its type locality on Fiji from a coral reef on the Andaman Islands. The species identification is based on SEM investi- gations of cuticular structures. Priapulids are among the more recent dis- coveries in the marine meiofauna. Remane (1963) saw the first interstitial representa- tive of this taxon during his investigations in the Red Sea. Van der Land (1968) gave the first description of a meiobenthic priap- ulid, Tubiluchus corallicola, which he found in shallow coral sand from Curagao. Since that time, four additional Tubiluchus spe- cies have been described and two genera erected. Maccabeus Por, 1973, with at least one species is obviously identical with Sal- vini-Plawen’s (1974) Chaetostephanus (Cal- loway 1988). Meiopriapulus Morse, 1981, the most recently discovered genus, is rep- resented by the species fijiensis known from a single beach on Viti Levu, Fiji. Details of its integument and anatomy were published by Storch et al. (1989a, 1989b). Neverthe- less, priapulids are extremely rare represen- tatives of the interstitial meiofauna. Present records (reviewed by Calloway 1988) ap- pear to indicate their endemic distribution in more or less insular habitats, except for Tubiluchus corallicola which is recorded from Curacao, the Carribean Sea, Bonaire, Barbados and Bermuda (Van der Land 1970). All the more surprising and of zoo- geographic interest was the discovery of the Pacific M. fijiensis in the Andaman Sea dur- ing a short collecting trip by the author to the Andaman Islands in February 1988. Meiopriapulus fijiensis Morse, 1981 Methods and Materials Extraction took place in a room of the Hotel Aasiana, Port Blair. The sediment was mixed with a solution of about 8% MgCl, in tap water to relax any meiofauna present, and sieved through a 63 wm screen. The fraction was placed in fresh sea water and the animals were observed and sorted live using a transportable dissecting microscope. Oocytes were observed in at least two of the specimens. Following narcotization with the MgCl,-solution, the specimens were fixed for several weeks in either 10% formalin or in Bouin’s fluid and then transferred into 70% ethanol. The fixed animals exhibited all stages of contraction, invertion, and evertion of the introvert. Four specimens were postfixed with 1% OsO, in aqua dest., critical point dried in CO,, coated with gold and examined with a Cambridge Stereo Scan 250. One specimen was whole-mounted in W 15 mounting medium and examined us- ing interference contrast microscopy. Results Material. —Twelve specimens of different size were found. Five of them were used for identification, two were deposited in the National Museum of Natural History, VOLUME 103, NUMBER 4 Smithsonian Institution, Washington, D.C.: USNM 128299, and five remain in the col- lection of the author. Locality.—Bay of Bengal, Andaman Is- lands, South Andaman. Coral reef in front of a small village about 10 km south of Port Blair. In water covered patches of clean cor- al sand between coral clumps in the surf zone of the reef edge; during low tide, 19 Jan 1988. Measurements. —Length between 1.2 and 1.7 mm; width of the abdomen about 300 pm. External morphology. —The terminology used is based on the species description of Morse (1981) and also incorporates the findings of Storch et al. (1989a, 1989b). Only cuticular details are described. The overall external morphology totally agrees with the description of Morse (1981): a cylindrical body consisting of a bulbous anterior intro- vert and an elongate abdomen, the posterior end of which is conical in shape and bears a circle of protruding hooks (Fig. 1A). The introvert sensory scalids are arranged in three circlets. The eight unornamented elongate scalids of the anterior circlet (length about 70 wm) consist of a basal half with two scale-like cover elements (Fig. 2B) and tubular protrusions at the tip. The upper cover element is triangular (t), and the lower one is comb-like with fine fimbriations (c). Fimbriae in groups or singly may occur proximally to the lower cover element (s). Two small flosculi with circularly arranged papillae (“‘petals’’) (f) occur at the base of the scalids. The second and third circlets of alternately-positioned ornamented scalids each consist of eight elongate structures (length about 50 um) bearing overlapping fimbriated cover elements (Fig. 2A). There are usually fewer cover elements in the pos- terior third circlet (three or four elements) than in the anterior second circlet (four to six). Each scalid bears three distal tubular protrusions; the middle protrusion is longer than the two lateral ones. 785 The small introvert locomotory scalids of ““semi-palm-tree-like shape” are arranged in rows oblique to the longitudinal body axis. Thirteen to 15 scalids occur in each row. The spoon-shaped scalid hood has about 40 fimbriations (Figs. 1E, 2A). The pharynx is everted in the form of a buccal cone and bears 16 longitudinal rows of fringed teeth with eight (nine?) teeth in each row (Fig. 1A—D). Teeth of adjacent rows alternate, so that 16 (18?) alternating rings of teeth surround the buccal cone. Three different types of teeth are present. The anterior teeth surround the pharynx opening with comb-like arranged fimbriae of nearly equal length (a); the teeth in the following two rings are scale-like with tri- angularly arranged fimbriations (m), and have a covering row of small fimbriae of equal length on the upper side and fine fim- briations on the inner side. The teeth at the base of the everted pharynx are tripartite, with a central comb-like structure and two lateral bundles of long fimbriae (b) that ex- tend beyond the central comb. The cuticle of the abdomen is regularly structured with small spherical protrusions, wrinkled by an irregular pattern of folds due to fixation (Fig. 1A) and characteristic tu- bercles (less than 1 wm diameter) that ap- pear to be equidistant from each other. Be- hind the introvert the anterior portion of the abdomen possesses triangular scalids that have fimbriated apical edges and a base width of about 20 um. The posterior end of the abdomen bears a ring of robust hooks each consisting of two strong prongs and a central tubular projection that Storch et al. (1989b) found to be a sensillum. There are numerous flosculi on the abdomen, which increase in number posterior to the ring of hooks. Often, one or two flosculi are asso- ciated with a larger cone-shaped structure with a prominent apical tubular projection and a subdistal ring of short petals (““mod- ified tubulus” in Morse (1981); “setae” in Storch et al. (19895). The flosculi have a 786 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. cone with anterior comb-like teeth (a), middle triangular teeth (m), and basal tripartite teeth (b); C, Close up of middle triangular pharyngeal teeth (m) and basal tripartite teeth (b); D, Close up of anterior comb-like pharyngeal teeth (a) and middle triangular teeth (b); E, Locomotory scalids of introvert. palisade-like ring of about 14 longer petals, and several smaller inner projections sur- rounding a central opening. Discussion The agreement between the Andaman an- imals and those from the type-locality (Fiji) Meiopriapulus fijiensis: A, Whole specimen with pharynx and introvert everted; B, Everted pharynx (Morse 1981) is extremely high. A few dif- ferences, however, exist. The so-called unornamented first circlet scalids of the in- trovert possess two fimbriate ornamenta- tions, whereas Morse states that they “gen- erally are without ornamentations.” Careful examination of additional SEM micro- graphs of Fijian specimens kindly provided VOLUME 103, NUMBER 4 787 Fig. 2. Meiopriapulus fijiensis: A, Locomotory scalids of introvert and sensory “ornamented” scalids of second and third circlet; B, Sensory “unornamented” scalid of first circlet with two flosculi at the base (f). t = triangular cover element, c = comb-like cover element, s = small group of fimbriations. by Dr. Morse shows them to be present there as well, although being more delicate. The number of scale-like cover-elements on or- namented scalids of the second circlet is probably higher (8) than in the Fiji speci- mens and may be related to age and to moulting (pers. comm. of Morse). Moulting has also been assumed by Storch et al. (1989b). Storch et al. (1989a) also counted 16 rows of pharyngeal teeth but mention only seven teeth in each row, whereas the Andaman specimens have eight (nine?) teeth per row. The three types of pharyngeal teeth are not described by Morse (1981), but do exist in the Fijian animals (see fig. 5 in Storch et al. 1989a, and fig. 2 in Storch et al. 1989b). Slight quantitative differences also seem to exist in the number of fimbriae of the lo- comotory scalids, the triangular abdominal scalids and in the number of petals of trunk flosculi, but this may well be due to general variability of these details or because mi- crographs are unfavorable. The generally less delicate appearance of the fimbriae in the Andaman species may be the result of in- adequate fixation. The absence of distinct differences within the structure-rich details of the body surface suggests that the Andaman and Fiji popu- lations are conspecific. Meiopriapulus fijien- sis joins the series of intertidal sand inhab- iting meiofauna species that appear to exhibit an extremely wide geographic range (Sterrer 1973, Westheide 1977, Ax & Ar- monies 1987). However, little is known about the taxonomic significance of the ex- ternal cuticular characters within the Priap- ulida. Van der Land (1982:333) could find “not a single useful differential character in the adult”? between Tubiluchus corallicola (from the Carribean Sea) and 7. remanei (from the Red Sea) unless he examined the peculiar cuticular organs associated with the male genital pores. No such organs appear to exist for comparison of the two Meio- 788 priapulus populations. Further investiga- tions (e.g., by TEM) may elucidate features which may otherwise differentiate between these externally nearly identical members of two widely separated populations, as in the case of the Microphthalmus listensis spe- cies-complex found on both sides of the At- lantic (Westheide & Rieger 1987). Acknowledgments I thank Dr. G. C. Rao for providing trans- portation on the Andaman Islands. I greatly acknowledge the help of Dr. M. P. Morse, who sent me her SEM micrographs for comparison. Dr. R. P. Higgins kindly read the manuscript. W. Mangerich skillfully performed the SEM investigations. Literature Cited Ax, P., & W. Armonies. 1987. Amphiatlantic iden- tities in the composition of the boreal brackish water community of Plathelminthes. A com- parison between the Canadian and European Atlantic Coast.—Microfauna Marina 3:7-80. Calloway, C. B. 1988. Priapulida. Pp. 322-327 inR. P. Higgins and H. Thiel, Introduction to the study of meiofauna. Smithsonian Institution Press, Washington. Morse, M. P. 1981. Meiopriapulus fyiensis n. gen., n. sp.: an interstitial priapulid from coarse sand in Fiji.—Transactions of the American Micro- scopical Society 100:239-252. Remane, A. 1963. The systematic position and phy- logeny of the pseudocoelomates. Pp. 247-255 in E. C. Dougherty et al., eds., The lower Meta- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON zoa. Comparative biology and phylogeny. Uni- versity of California Press, Berkeley. Salvini-Plawen, L. O. 1974. Zur Morphologie und Systematik der Priapulida: Chaetostephanus praeposteriens, der Vertreter einer neuen Ord- nung Seticoronaria.— Zeitschrift fiir Zoolo- gische Systematik und Evolutionsforschung 12: 31-54. Storch, O., R. P. Higgins, & M. P. Morse. 1989a. Internal anatomy of Meiopriapulus fijiensis (Priapulida).—Transactions of the American Microscopical Society 108:245-261. 1989b. The ultrastructure of the integument of Meiopriapulus fijiensis (Priapulida).—Trans- actions of the American Microscopical Society 108:319-331. Sterrer, W. 1973. Plate tectonics as a mechanism for dispersal and speciation in interstitial sand fau- na.— Netherlands Journal of Sea Research 7:200- 222. Van der Land, J. 1968. A new aschelminth, probably related to the Priapulida.— Zoologische Mede- delingen 42:237-250. 1970. Systematics, zoogeography, and ecol- ogy of the Priapulida.—Zoologische Verhan- delingen 112:1-118. 1982. A new species of Tubiluchus (Priapu- lida) from the Red Sea.— Netherlands Journal of Zoology 32:324—-335. Westheide, W. 1977. The geographical distribution of interstitial Polychaeta. — Mikrofauna des Meeresbodens 61:297-302. —, & R.M. Rieger. 1987. Sytematics of the M- crophthalmus-listensis-species group (Polychae- ta: Hesionidae): facts and concepts for recon- struction of phylogeny and speciation. — Zeitschrift fiir Zoologische Systematik und Evo- lutionsforschung 25:12-39. Spezielle Zoologie, Fachbereich Biologie/ Chemie, Universitat Osnabrtick. D-4500 Osnabrtick, Federal Republic of Germany. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 789-802 A CHECKLIST OF THE BRYOZOA OF THE GALAPAGOS William C. Banta and John C. Redden Abstract. —The 184 records of Bryozoa known from the Galapagos archipel- ago (Ecuador) are listed, including 4 new records and 3 unidentified species from Academy Bay, Santa Cruz. Thirty- four species (18%) of Galapagos bryozo- ans are endemic. Proportionally, there are more endemic stenolaemates (42%) than there are anascans or ascophorans. The Galapagos belongs in the Panamic Province, but is represented disproportionately by colder water species of the Northern Hemisphere with weedier, r-selected tendencies. There is evidence of difficulty in arriving at the Islands and of genetic isolation and local adap- tation, especially among stenolaemates. -__eoeoo This report lists all records of Bryozoa we know of from the Galapagos archipelago, Ecuador. The broader zoogeographical im- plications, discussed in detail by Banta (1991), are summarized herein. Collections Previous collections.—There are seven published works describing bryozoan col- lections from the Galapagos. Hastings (1930) had 37 taxa from 11 shallow-water sites made by C. Crossland (1927). Canu & Bass- ler (1930) studied 50 species from four sta- tions collected in 1888 and 1891 from the Albatross collections of the U.S. Fisheries Commission. One of these stations was from relatively deep water (684 fathoms = 125] m). Osburn (1950, 1952, 1953) and Osburn and Soule (1953) had extensive collections, from at least 25 stations, some of great depth, made in the 1930s by the Velero JI expe- ditions of the Allan Hancock Foundation (see Fraser 1943). Soule ( 1963b) adds 3 ad- ditional records from the Hancock Gala- pagos collections in his monograph on Bryozoa of the Gulf of California. New data.—In July, 1980 the senior au- thor collected about 30 species at snorkel depth near boat docks at Academy Bay, Santa Cruz Island. Specimens were dried and wrapped in paper for transport to the laboratory. Where necessary, calcareous specimens were cleaned in dilute KOCI. Specimens are for deposit in the National Museum of Ecuador, Quito. Four of the specimens collected are new records for the Galapagos: Lichenopora fim- briata, Membranipora arborescens, Ar- thropoma circinata and Parasmittina cross- landi. Another three are undetermined species, probably new, unnamed taxa: Hip- poporella sp., with affinities in H. gorgo- nensis (Hastings 1930); Lagenicella sp., with affinities in L. Jacunosa (Bassler 1934): and Parasmittina sp., with affinities in P. has- tingsae Soule & Soule 1973. Known records in the Galapagos. —The appendix includes every record we were able to find where bryozoan species or subspecies were identified from the Galapagos Islands. Bryozoan subspecies in the Galapagos are based entirely on morphological features: no good criteria are known to distinguish subspecies from species in the area. Au- thorities for Galapagos records are listed. These are followed by the family of each species and the islands from which each rec- ord was taken. The order of the Islands is arbitrary. Where there is more than one name for an Island we use the name in most common use by the inhabitants of the Ga- 790 lapagos (Bowman 1966). The distributions listed are culled from scattered literature. We place each species into one of seven gen- eral distribution categories: (1) Cosmopol- itan: recorded worldwide from the equator into Arctic and Antarctic waters; (2) Cir- cumglobal: worldwide, but absent from high latitudes above about 60°N and S; (3) Pan American: recorded in both the eastern Pa- cific and the Caribbean or Gulf of Mexico; present or absent elsewhere in the Atlantic and the Mediterranean; (4) Pacific: known from both the eastern and western Pacific; (5) Eastern Pacific: found exclusively along the western coasts of the Americas, includ- ing areas outside the Panamic; (6) Panamic: coastal areas of the Americas from about the U.S. Mexican border to Guayaquil in Ecuador; (7) Endemic: not recorded outside the Galapagos. Summary of the Zoogeography of Galapagos Bryozoa One hundred eighty-four species of bryo- zoans are known from the Galapagos Is- lands, a number comparable to that of the most diverse bryozoan faunas known. This number is probably an underestimate, es- pecially for the soft-bodied ctenostomes. The sparse data provide no evident patterns of differential distribution of species among the islands, so at least for the purposes of this report, the Galapagos can be treated as a single zoogeographical unit. Although the relative numbers of anascan and ascopho- ran cheilostomes are near the average for the rest of the world, the proportion of sten- olaemates (18%) is about twice as high as that in the rest of the Panamic. Comparable proportions of stenolaemates are found in the Magellanic and Chilean waters off South America, but otherwise there is zoogeo- graphic affinity between these areas and the Galapagos. Relatively low water tempera- tures and elevated productivities associated with upwelling do not fully explain the high proportion of stenolaemates in Galapagos waters. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON At the family level, few absences are con- spicuous, except for the Steginoporellidae, characteristic of coral communities, rare in the Galapagos. Zoogeographically, the Galapagos Islands belong to the Northern Hemisphere, se- curely in the Panamic Province, with 46% of Galapagos species shared with the rest of the Panamic. The distribution of Panamic bryozoans seems to divide the Panamic into two subprovinces near 10°N. Within the Panamic, strong affinities also are seen among the Galapagos and Cocos Island and the Gulf of California. More distant affini- ties are seen with the Gulf of Mexico and Indo-Pacific. The fauna seems unrelated to that of the Chilean and Magellanic Prov- inces of South America. The few shared spe- cies are mostly circumglobal or cosmopol- itan in distribution. Thirty-four of 184 species (18%) of Ga- lapagos bryozoans are endemic to the Is- lands. This proportion is comparable to that of other major groups of shallow-water an- imals, but smaller than that of the molluscs, especially gastropods. There are 2 endemic genera among 53 in the islands, about 4%. Proportionally, there are more endemic stenolaemates (14 of 33 species, 42%) than endemics in anascans or ascophorans. This difference does not seem to be a simple case of more lumping in the Cheilostomata than among stenolaemates; on the contrary, lumping probably has been more severe among stenolaemates. Affinities with neighboring zoogeograph- ical provinces are stronger for cheilostomes than for stenolaemate Bryozoa. This, taken together with the higher rate of endemism for stenolaemates, suggests that the dispers- al abilities of stenolaemates are inferior to those of cheilostomes. The difference can- not be accounted for by the presence of planktotrophic larvae among some cheilo- stomes, because Panamic species with planktotrophic larvae are no better repre- sented in the Islands than are those with lecithotrophic larvae. VOLUME 103, NUMBER 4 In general, the Galapagos bryozoan fauna seems to be proportionately more repre- sented by species which range into latitudes higher than about 30 degrees. There also are proportionately more species with geo- graphical ranges wider than those indige- nous to waters nearer the Galapagos. The bryozoan distribution data taken as a whole convey a picture of the Galapagos as a biological isolate belonging securely in the Panamic Province, but represented dis- proportionately by colder water species of the Northern Hemisphere with weedier, r-selected tendencies. There is evidence of difficulty in arriving at the Islands and of genetic isolation and local adaptation, es- pecially among stenolaemates. The rarity of some habitats, notably estuaries and coral reefs, has contributed to a unique species composition. The stenolaemates of the Galapagos in particular are disproportionately abundant relative to the rest of the Panamic. There is evidence that the Galapagos stenolaemate fauna may be more isolated and/or more rapidly evolving than the cheilostome fau- na. Acknowledgments Michael Temkin, Donald Bonyai and Re- becca Beerman helped collect specimens in the Galapagos. Joann Sanner at the US. National Museum of Natural History helped with the collections and literature and iden- tified some Parasmittina specimens. Alan H. Cheetham, also of the Museum, dis- cussed the subject with me and was a great help with both details and broad ideas in- cluded in this paper. Steven Cairns made a number of suggestions and corrected several Latin and Greek names. Thanks also are due the Government of Ecuador, the Charles Darwin Foundation and the Smithsonian Institution for assistance in collecting and studying specimens. Supported in part by an American University Research Award. Contribution 18-89, The American Uni- versity Biology Department. 791 Literature Cited Banta, William C. 1969a. Systematic list of Bryozoa identified from Farnsworth Bank, Santa Cata- lina Island. Pp. 419-422 in O. Hartman, Quan- titative survey of the benthos of San Pedro Ba- sin, southern California. Part II, Final results and conclusions.—Allan Hancock Pacific Ex- peditions 19(2):1-456. . 1969b. Watersipora arcuata, a new species in the subovoidea-cucullata-nigra complex (Bryo- zoa, Cheilostomata).— Bulletin Southern Cali- fornia Academy of Sciences 68(2):96-102. . 1991. Bryozoa of the Galapagos. in M. James, ed., Galapagos marine invertebrates: taxonomy, biogeography and evolution in Darwin’s is- lands. Plenum Publishing Company, New York (in press). , & Rennate J. M. Carson. 1977. Bryozoa from Costa Rica.— Pacific Science 31(4):38 1-424. Borg, F. 1933. Ona supposed species of Heteropora (Bryozoa).—Annals and Magazine of Natural History (10)12:583-587. Bowman, R.I. 1966. Comments on the names of the Islands of the Galapagos. P. vxii in R. I. Bow- man, ed., The Galapagos. Proceedings of the symposium of the Galapagos International Sci- entific Project. University of California, Berke- ley. Busk, George. 1854. Catalogue of marine Polyzoa in the collection of the British Museum, Part II, Cheilostomata (Part). British Museum pp. i—viii + 55+120. Reprinted 1966, Johnston Reprint Company, New York. Canu, Ferdinand, & Raymond S. Bassler. 1928a. Fos- sil and Recent Bryozoa of the Gulf of Mexico region.— Proceedings of the U.S. National Mu- seum 72(14):1-199. , & 1928b. Les Bryozoaires du Maroc et de Mauritanie.—Mémories de la Société des Sciences Naturelles du Maroc 18:1-85. ——.,,& . 1929. Contributions to the biology of the Philippine Archipelago and adjacent regions. — Bulletin of the U.S. National Museum 100(9): 1-683. 5, & 1930. The bryozoan fauna of the Galapagos Islands.—Proceedings of the U.S. National Museum 76(13):1-78. Cheetham, Alan H., & Philip A. Sandberg. 1964. Quaternary Bryozoa from Louisiana mud- lumps.—Journal of Paleontology 38(6):1013- 1046. Crossland, C. 1927. The expedition to the south Pa- cific of the S.Y. ‘St. George’.— Transactions of the Royal Society of Edinborough 55(2):531- 554. Dick, Matthew H., & June R. P. Ross. 1988. Inter- 792 tidal Bryozoa (Cheilostomata) of the Kodiak vi- cinity, Alaska.— Occasional Papers of the Cen- ter for Pacific Northwest Studies (Western Washington University, Bellingham, Washing- ton) 23:1-133. ——.,& . 1985. Intertidal cheilostome bryo- zoans in rock-pile habitat at Narrow Strait, Ko- diak, Alaska. Pp. 87-93 in Claus Nielsen & Gil- bert P. Larwood, eds., Bryozoa: Ordovician to Recent. Olsen and Olsen, Fredensborg, Den- mark. Fraser, C. McLean. 1943. General account of the scientific work of the Velero III in the eastern Pacific, 1931—41. Part III. A ten-year list of the Velero III collecting stations.— Allan Hancock Pacific Expeditions 1(3):259-431. Gordon, Dennis P. 1982. The genera of the Chap- erlidae (Bryozoa). — New Zealand Journal of Zo- ology 9:1—24. 1984. The marine fauna of New Zealand: Bryozoa: Gymnolaemata from the Kermadec Ridge.— Memoirs of the New Zealand Ocean- ographic Institution 91:1-198. Harmer, Sidney F. 1926. The Polyzoa of the Siboga Expedition. Part II. Cheilostomata Anasca.— Siboga-Expedite Monograph 28b:181-501. . 1934. Polyzoa of the Siboga Expedition. Part III. Cheilostomata Ascophora. I. Family Rete- poridae. —Siboga-Expedite Monograph 28c: vii + pp. 503-640. 1957. Polyzoa of the Siboga Expedition. Part IV. Cheilostomata Ascophora. II (Ascophora, except Reteporidae, with additions to part II, Anasca).—Siboga-Expedite Monograph 28d: xv + pp. 641-1147. Hastings, Anna B. 1930. Cheilostomatous Bryozoa from the vicinity of the Panama Canal collected by Dr. C. Crossland on the cruise of the S.Y. “St. George’’.— Proceedings of the Zoological Society of London 1929(4):1-17. ——, & 1979. The South African Mu- seum’s Meiring Naude cruises. Part 13. Bryozoa II.—Annals of the South African Museum 91(1): 1-161. Hayward, Peter J., & John S. Ryland. 1979. British ascophoran bryozoans. Synopsis of the British Fauna (new series).— Linnean Society, London 14:1-312. Lagaaij, Robert. 1963. New additions to the bryo- zoan fauna of the Gulf of Mexico. — Publications of the Texas University Institute of Marine Sci- ence 9:162—236. MacGillivray, Paul H. 1889. On some South Aus- tralian Polyzoa.— Transactions and Proceedings of the Royal Society of South Australia 12:24— 30. Marcus, Ernst. 1938. Bryozoarios Brasileiros - II.— Boletins da Faculidade de Philosophia, Scien- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cias e Letras, Universidad de Sao Paulo 4, Zoo- logia 2:1-137. Maturo, Frank J. S. 1968. The distributional pattern of Bryozoa of the east coast of the United States exclusive of New England.— Atti della Societa Italiana di Scienze Naturali 108(36):261—284 , & Thomas J. M. Schopf. 1968. Ectoproct and entoproct type material: examination of species from New England and Bermuda named by A. E. Verrill, J. W. Dawson, and E. Desor.—Pos- tilla 120:1-95. Morris, Penny A. 1980. The bryozoan family Hip- pothoidae (Cheilostomata-Ascophora), with emphasis on the genus Hippothoa. — Allan Han- cock Monographs in Marine Biology 10:1—145. Moyano, Hugo. I. 1983. Southern Pacific Bryozoa: a general view with emphasis on Chilean spe- cies. —Gayana 46:3-45. Neviani, Antonio. 1895. Nuove genere e nuove spe- cie di Briozoi fossili.—Rivue Italiana Paleon- tologica Bologna 1:82-84. Osburn, Raymond C. 1950. Bryozoa of the Pacific coast of North America. Part I, Cheilostomata- Anasca.—Allan Hancock Pacific Expeditions 14(1):1-269. 1952. Bryozoa of the Pacific coast of North America. Part II, Cheilostomata-Ascophora. — Allan Hancock Pacific Expeditions 14(2):271- 611. 1953. Bryozoa of the Pacific coast of Amer- ica. Part III. Cyclostomata, Ctenostomata, En- toprocta and Addenda.— Allan Hancock Pacific Expeditions 14(3):613-841. —., & John D. Soule. 1953. Order Ectoprocta, Suborder Ctenostomata. Pp. 726-758 in R. C. Osburn, Bryozoa of the Pacific coast of America. Part II. Cyclostomata, Ctenostomata, Ento- procta and Addenda.— Allan Hancock Pacific Expeditions 14(3). Powell, Neil A. 1967. Polyzoa (Bryozoa) Ascophora from New Zealand. — Discovery Reports 34:199- 394. . 1971. The marine Bryozoa near the Panama Canal.— Bulletin of Marine Science 21(3):766— 778. Prenant, Marcel, & Genevieve Bobin. 1966. Bry- ozoaires. Deuxiéme partie: Chilostomes Anas- ca.— Fauna de France 68:1-647. Robertson, Alice. 1908. The incrusting cheilostom- atous Bryozoa of the west coast of North Amer- ica.— University of California Publications in Zoology 4(5):253-344. Ryland, John S., & Dennis P. Gordon. 1977. Some New Zealand and British species of Hippothoa (Bryozoa: Cheilostomata).— Journal of the Roy- al Society of New Zealand 7(1):17-49. ——., & Peter J. Hayward. 1977. British anascan VOLUME 103, NUMBER 4 bryozoans. — Synopsis of the British Fauna (new series) 10:1-188. Soule, Dorothy F., & John D. Soule. 1964. The Ec- toprocta (Bryozoa) of Scammon’s Lagoon, Baja California.— American Museum Novitates 2199: 1-56. ——,, & 1973. Morphology and speciation of Hawaiian and eastern Pacific Smittinidae (Bryozoa, Ectoprocta).— Bulletin of the Ameri- can Museum of Natural History 152(6):365—440. , & 1975. Species groups in Watersi- poridae.—Documments Laboratorie Geologie Faculte des Sciences de Lyon 3(2):299-309. , & 1985. Effects of oceanographic phenomena such as ‘El Nifio’ on the zoogeog- raphy and endemism of tropical/subtropical Pa- cific Watersiporidae. Pp. 293-300 in C. Nielsen and G. P. Larwood, eds., Bryozoa: Ordovician to Recent. Olsen & Olsen, Fredensborg, Den- mark. Soule, John D. 1959. Results of the Puritan-Ameri- can Museum of Natural History expedition to western Mexico. 6. Anascan Cheilostomata (Bryozoa) of the Gulf of California.— American Museum Novitates 1969:1-54. 1963a. Phylum Bryozoa. Pp. 232-234 in L. G. Hertline, ed., Contribution to the biogeog- raphy of Cocos Island, including a bibliogra- phy.— Proceedings of the California Academy of Sciences 32(8):232—234. . 1963b. Results of the Puritan-American Mu- seum of Natural History expedition to western Mexico. 18. Cyclostomata, Ctenostomata (Ec- toprocta) and Entoprocta of the Gulf of Cali- fornia.— American Museum Novitates 2144:1- 34. Winston, Judith E. 1982. Marine bryozoans (Ecto- procta) of the Indian River area (Florida). — Bul- letin of the American Museum of Natural His- tory 173(2):100-176. , & Beverly F. Heimberg. 1986. Bryozoans from Bali, Lombok, and Komodo.— American Museum Novitates 2841:1-106. Department of Biology, The American University, Washington, D.C. 20016. Appendix Bryozoan species known from the Galapagos Ar- chipelago, arranged alphabetically within Orders (there are no Phylactolaemata known). STENOLAEMATA Borgiola rugosa (Borg, 1933).—Osburn, 1953:697. Heteroporidae. Islands: Marchena. Distribution: Ja- pan and the Galapagos (Pacific). 793 Borgiola sp. (Canu & Bassler, 1930).—Heteropora sp.— Canu & Bassler, 1930:60. Heteroporidae. Islands: Espanola. Distribution: Endemic. Apertures mea- sure 0.10 mm in diameter and lunaria are present; we assign it tentatively to Borgiola, distinct from B. rugosa (Borg, 1933). Probably a new endemic spe- cies. Cavaria praesens Canu & Bassler, 1930.—Canu & Bassler, 1930:58. Lichenoporidae. Islands: Floreana. Distribution: Endemic. Crisia maxima Robertson, 1910.—Osburn, 1953:682. Crisiidae. Islands: Espanola, San Crist6bal, Santa Fé. Distribution: British Columbia to the Gulf of Cali- fornia and the Galapagos (Eastern Pacific). Crisia occidentalis Trask, 1857.—Osburn, 1953:680. Crisiidae. Islands: Seymour. Distribution: British Columbia to Costa Rica and the Galapagos (Eastern Pacific). Crisia serrulata Osburn, 1953.—Osburn, 1953:679. Crisiidae. Islands: Isabela, Floreana, San Cristdbal, Santa Cruz, Onslow. Distribution: British Columbia, California and the Galapagos (Eastern Pacific). Crisulipora occidentalis Robertson, 1910.—Osburn, 1953:686. Crisiidae. Islands: Espanola. Distribution: California, Peru, Japan, Brazil, Panama, Gulf of Cal- ifornia, Baja California (Pan American). Diaperoecia flabellata (Canu & Bassler, 1923).—Canu & Bassler, 1930:52. Diaperoeciidae. Islands: Espa- nola. Distribution: Pleistocene of California; Gala- pagos (Endemic). Disporella astraea Osburn, 1953.—Osburn, 1953:719. Lichenoporidae. Islands: Floreana, Isabela. Distri- bution: Endemic. Disporella octoradiata (Waters, 1904.).—?Disporella octoradiata.—Osburn, 1953:718. Lichenoporidae. Islands: Isabela. Distribution: Antarctic, Chile (Moyano 1983:15) (Eastern Pacific). Osburn’s query mark reflected his uncertainty about the generic placement, not his confidence in the species deter- mination. Disporella ovoidea Osburn, 1953.—Osburn, 1953: 713.—Lichenopora radiata. —Canu & Bassler, 1928. Lichenoporidae. Islands: Isabela, Espanola. Distri- bution: California, Colombia and the Galapagos (Eastern Pacific). Entalophora sp. Osburn, 1953.—Entalophora sp.— Osburn, 1953:669. Entalophoridae. Islands: Isabela. Distribution: Endemic. The animal appears to be distinct; Osburn did not describe it because ovicells were lacking. Entalophora symmetrica Osburn, 1953.—Osburn, 1953:667. Entalophoridae. Islands: San Cristobal. Distribution: Endemic. Heteropora sp. Osburn, 1953.—Osburn, 1953:695. Heteroporidae. Islands: Wenman, San Cristobal. Distribution: Unknown. Similar to Heteropora pa- cifica Borg, 1933:317 in its measurements, but avail- able specimens are in poor condition. 794 Lichenopora (Radiopora) pacifica (Osburn, 1953.—De- francia stellata.—Canu & Bassler, 1930:57.—Dis- porella stellata var. pacifica.—Osburn, 1953:716. Lichenoporidae. Islands: Wenman, Isabela, Espano- la, Gardner-by-Hood. Distribution: Endemic. This ‘variety’ surely deserves specific status. Lichenopora fimbriata Busk, 1875.—Present study. Lichenoporidae. Islands: Santa Cruz. Distribution: British Columbia, California, Chile, Falklands, Tris- tan de Cunha, Azores, Cape Verde Islands, Australia, New Zealand, Kerguelens, Juan Fernandez (Circum- global). First record in the Galapagos. For descrip- tion see Osburn 1952:709. Lichenopora novae-zelandiae (Busk, 1875).—Osburn, 1953:705. Lichenoporidae. Islands: Floreana. Dis- tribution: New Zealand, Australia, Ceylon, Japan, British Columbia, California (Pacific). Plagioecia lactea Calvet, 1903.—Canu & Bassler, 1930: 48.—?Plagioecia lactea.—Osburn, 1953:653. Dia- stoporidae. Islands: Wenman, Espanola. Distribu- tion: Atlantic (France), Galapagos (Pan American). The identity with Atlantic specimens is question- able. Plagioecia meandrina (Canu & Bassler, 1930).—Os- burn, 1953:635.— Diaperoecia meandrina. —Canu & Bassler, 1930:51. Diastoporidae. Islands: Wenman, San Cristobal, Espanola, Isabela, Floreana, Gardner- by-Hood. Distribution: California, Guadalupe Is- land (off Mexico), Galapagos (Eastern Pacific). Plagioecia patina (Lamarck, 1816).—Osburn, 1952:631. Diastoporidae. Islands: Wenman, Isabela, San Cris- tobal. Distribution: British Columbia to Baja Cali- fornia and the Galapagos; Atlantic. (Pan American). Plagioecia sarniensis (Norman, 1864).—Osburn, 1953: 632. Plagioeciidae. Islands: Santa Fé. Distribution: Colombia, California, Washington; ““Cosmopoli- tan” (Osburn 1953:633) (Circumglobal). Plagioecia striatula Canu & Bassler, 1930.—Diaper- oecia striatula.—Canu & Bassler, 1930:49. Diaper- oeciidae. Islands: Espanola. Distribution: Endemic. See Osburn (1953:641) regarding generic placement of this species. Plagioecia subpapyracea Canu & Bassler, 1930.— Dia- peroecia subpapyracea.—Canu & Bassler, 1930:50. Diaperoeciidae. Islands: Espanola. Distribution: En- demic. See Osburn (1953:641) regarding the generic placement of this species. Plagioecia tubiabortiva Canu & Bassler, 1930.—Os- burn, 1953:636.—Microecia tubiabortiva.—Canu & Bassler, 1930:48. Diastoporidae. Islands: Wenman, Espanola. Distribution: Endemic. Platonea expansa Osburn, 1953.—Osburn, 1953:663. Tubuliporidae. Islands: Isabela, Espanola, Gardner- by-Hood. Distribution: Endemic. Platonea veleronis Osburn, 1953.—Osburn, 1953:662. Tubuliporidae. Islands: Isabela, Espafiola, Duncan, Floreana, Gardner-by-Hood, Santa Fé. Distribution: Endemic. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Proboscina lamellifera Canu & Bassler, 1930.—Canu & Bassler, 1930:46.—Osburn, 1953:623. Oncousoe- ciidae. Islands: Wenman, Espanola, Isabela, Gard- ner-by-Hood. Distribution: Endemic. Proboscina major (Johnston, 1847).—Osburn, 1953: 621.—Present study.— Proboscina (Oncousoecia) major Canu & Bassler, 1930:46. Oncousoeciidae. Is- lands: Espanola, Isabela, Santiago, Santa Fé, Santa Cruz. Distribution: British Columbia to Costa Rica, Cocos Island, Galapagos; Norway to the Cape Verde Islands; Mediterranean (Osburn 1953:621); Chile (Moyano 1983:15) (Pan American). Tubulipora flexuosa (Pourtalés, 1867).—Osburn, 1953: 653. Tubuliporidae. Islands: San Salvador. Distri- bution: California, Cocos Island, the Galapagos, Cuba, Porto Rico and the southern shore of the Ca- ribbean Sea. (Pan American). Tubulipora liliacea var. tenuis Canu & Bassler, 1930.— Canu & Bassler, 1930:55. Tubuliporidae. Islands: Floreana. Distribution: Endemic. Tubulipora pulchra MacGillivray, 1885.—Osburn, 1953: 653. Tubuliporidae. Islands: San Salvador. Distri- bution: Australia, California, British Columbia, Mexico, Costa Rica (Pacific). Tubulipora sp. Canu & Bassler, 1930.—Canu & Bass- ler, 1930:54. Tubuliporidae. Islands: Espanola. Dis- tribution: Unknown. Tubulipora sp. Canu & Bassler, 1930.—Canu & Bas- sler, 1930:52. Tubuliporidae. Islands: Floreana. Dis- tribution: Not given (Unknown). CTENOSTOMATA Amathia vidovici (Heller, 1867).—Osburn & Soule, 1953:741. Vesiculariidae. Islands: Not specified. Distribution: California to Ecuador and Galapagos; Massachusetts to Colombia; France to the Mediter- ranean and the Cape Verde Islands; eastern; Indian Ocean (Prenant & Bobin 1966:283( (Pan American). Buskia seriata Soule, 1953.—Soule in Osburn & Soule, 1953:747. Buskiidae. Islands: Seymour. Distribu- tion: California and the Galapagos (Eastern Pacific). ANASCA Aetea anguina (Linnaeus, 1758).—Hastings, 1930: 702.—Osburn, 1950:11. Aeteidae. Islands: Santiago, Santa Cruz, Isabela; Osburn’s stations unspecified. Distribution: Arctic, Great Britain, Maine to Brazil, Mediterranean, Indian Ocean, Malay, Australia, Ja- pan, Chile, Patagonia, Juan Fernandez, California, British Columbia, Antarctic (Cosmopolitan). Pren- ant and Bobin (1966:80) present an extensive syn- onymy. Aetea ligulata (Busk, 1852).—Osburn, 1950:13. Ae- teidae. Islands: Isabela, San Cristobal, Bartolomé. Distribution: British Columbia to Chile, Caribbean to Argentina, Red Sea, Baja California, Panama, Co- lombia, New Zealand, possibly the Mediterranean (Gordon 1984:39); South Atlantic. (Circumglobal). VOLUME 103, NUMBER 4 Aetea recta Hincks, 1861.—Osburn, 1950:12. Aetei- dae. Islands: Isabela, Santiago. Distribution: Europe south, Caribbean, Brazil, Gulf of California, Wash- ington, Galapagos, Chile (Pan American). Velero III Station converted from Fraser (1943). Aetea truncata (Landsborough, 1852).—Osburn, 1950: 13. Aeteidae. Islands: Isabela. Distribution: Europe, Indian Ocean, Japan to Australia, Nova Scotia to Brazil, British Columbia, California, Colombia, Peru and the Galapagos; Chile (Cosmopolitan). Synony- my, Prenant and Bobin 1966:86. Alderina smitti Osburn, 1950.—Osburn, 1950:60. Al- derinidae. Islands: Isabela, San Cristébal. Distri- bution: Gulf of Mexico, Caribbean, California, Gulf of California, Baja California, Panama (Pan Amer- ican). Antropora claustracrassa (Canu & Bassler, 1930).—Os- burn, 1950:53.—Membrendoecium claustracras- sum.—Canu & Bassler, 1930:7. Hincksinidae. Is- lands: Espanola, Floreana and unspecified stations of Osburn 1950:53. Distribution: Guaymas (Mexico) to Ecuador and the Galapagos (Panamic). Antropora tincta (Hastings, 1930).—Osburn, 1950:54.— Fresent study.—Crassimarginatella tincta. —Has- tings, 1930:708.—Osburn, 1950:54. Hincksinidae. Islands: Isabela, Santa Cruz; Osburn’s collection sites not specified. Distribution: California to the Gala- pagos (Eastern Pacific). There is much variation in avicularium shape among Pacific specimens. Aplousina filum (Jullien, 1903).—Canu & Bassler, 1930: 7.—Osburn, 1950:47. Hincksinidae. Islands: Espa- nola, Daphne Major. Distribution: Eastern Atlantic, Gulf of Mexico, Gulf of California, Cocos Island (Pan American). Aplousina major Osburn, 1950.—Osburn, 1950:48. Alderinidae. Islands: Santa Fé. Distribution: Cali- fornia, Mexico, Galapagos, Chile (Eastern Pacific). Beania mirabilis Johnston, 1840.—Osburn, 1950:170. Bugulidae. Islands: Isabela. Distribution: Scandi- navia to the Mediterranean; California to Panama and the Galapagos, Australia, Indian Ocean (Prenant & Bobin 1966:555) (Circumglobal). Bugula californica Robertson, 1905.—Osburn, 1950: 157. Bugulidae. Islands: San Cristobal. Distribution: Brazil, British Columbia, California, Gulf of Cali- fornia, Galapagos (Pan American). Bugula longirostrata Robertson, 1905.—Osburn, 1950: 156. Bugulidae. Islands: Isabela (Tagus Cove), San- tiago. Distribution: California, Gulf of California, Galapagos (Eastern Pacific). Bugula minima (Waters, 1909).— Bugula neritina var. minima. —Hastings, 1930:704. Bugulidae. Islands: Isabela. Distribution: Gulf of Mexico, Caribbean, Red Sea, Indian Ocean, Malay, Australia; Cocos Is- land, Costa Rica, Panama and the Galapagos (Os- burn 1950:155) (Circumglobal). Bugula mollis Harmer, 1926.—Hastings, 1930:704.— Osburn, 1950:158. Bugulidae. Islands: Santiago, Is- 795 abela. Distribution: New Guinea, Java Sea, Califor- nia to Panama and the Galapagos (Pacific). Bugula neritina (Linnaeus, 1758).—Hastings, 1930: 704.—Osburn, 1950:155.—Present study. Buguli- dae. Islands: Santiago, Isabela, Floreana, Santa Cruz. Distribution: Worldwide in warm ports (Circum- global). Bugula uniserialis Hincks, 1884.— Hastings, 1930:705. Bugulidae. Islands: Isabela. Distribution: Western Australia, California (Pacific). Callopora horrida (Hincks, 1880).—Osburn, 1950:70. Alderinidae. Islands: Isabela (Tagus Cove), Daphne Major. Distribution: California, Puget Sound, Brit- ish Columbia, Gulf of California. (Eastern Pacific). Callopora verrucosa Canu & Bassler, 1929.—Canu & Bassler, 1930:10. Alderinidae. Islands: Espanola. Distribution: Endemic. Osburn 1950:71 questions the generic assignment. Cauloramphus brunea Canu & Bassler, 1930.—Canu & Bassler, 1930:10.—Osburn, 1950:56.—Present study. Alderinidae. Islands: Santiago, Isabela, Flo- reana, Santa Cruz. Distribution: Alaska to Colombia and the Galapagos. (Eastern Pacific). The report of ‘Caulorhamphus brunnea’ in Alaska (Dick & Ross 1985:89) is of C. variegatus (Dick & Ross 1988:41; California: Banta 1969a:419 (as ‘C. brunnea’). Cauloramphus spiniferum (Hincks, 1838).— Hastings, 1930:713. Alderinidae. Islands: Santiago. Distribu- tion: Britain, Chile, California, British Columbia, Alaska; South Atlantic (see Moyano 1983:18) (East- ern Pacific). Cellaria diffusa Robertson, 1905.—Osburn, 1950:118. Cellariidae. Islands: Santiago. Distribution: Southern California, Baja California, Galapagos (Eastern Pa- cific). Cellaria veleronis Osburn, 1950.—Osburn, 1950:118. Cellariidae. Islands: Isabela, Daphne Major, Mar- chena, Santiago. Distribution: Clipperton Island, Cocos Island and the Galapagos (Soule 1963a:232) (Panamic). Chaperiopsis condylata (Canu & Bassler, 1930).— Chapperia condylata.—Canu & Bassler, 1930:44.— Osburn, 1950:91. Chaperiidae. Islands: Floreana. Distribution: Southern California to Colombia and the Galapagos; Post-Pleistocene of Louisiana (Cheetham & Sandberg 1964:1020) (Eastern Pacific). =Chaperiella condylata: Soule 1963b, Cocos Island. Gordon (1982) discusses the Chaperiidae. Chaperiopsis frontalis Osburn, 1950.— Chapperia fron- talis. —Osburn, 1950:93. Chaperiidae. Islands: Es- panola, Isabela, Santa Cruz. Distribution: Endemic. Copidozoum planum (Hincks, 1880).— Hastings, 1930: 713. Alderinidae. Islands: Santiago. Distribution: Australia; Mediterranean, Japan, California, Colom- bia, Panama (Osburn 1952:73) (Circumglobal). This record may belong to C. tenuirostre (Hincks); see Osburn 1952:72. Copidozoum tenuirostre (Hincks, 1880).—Osburn, 796 1952:72.—Callopora tenuirostris.—Canu & Bassler, 1929:9. Alderinidae. Islands: Espanola, Floreana; Osburn’s stations are unspecified. Distribution: Western Atlantic, Gulf of Mexico, Caribbean, Brazil; Mediterranean; Indian Ocean, Western Pacific, Brit- ish Columbia to Peru and the Galapagos (Prenant & Bobin 1966:257) (Circumglobal). Cupuladria canariensis (Busk, 1885).—Osburn, 1950: 33. Cupuladriidae. Islands: Not given. Distribution: Gulf of California to Ecuador and the Galapagos: Gulf of Mexico; Eastern Atlantic, Mediterranean (Cheetham & Sandberg 1964:1021) (Pan American). Discoporella umbellata (Defrance, 1823).— Hastings, 1930:719.—Osburn, 1950:113.—Cupularia umbel- lata.—Canu & Bassler, 1930:12. Cupuladriidae. Is- lands: Santiago, Espafiola; Osburn’s stations not specified. Distribution: Gulf of Mexico, Eastern and Western Atlantic; Mediterranean; South Africa, Cal- ifornia, Cocos Island and the Galapagos (Cheetham and Sandberg 1964:1022) (Pan American). Electra bellula var. bicornis (Hincks, 1881).— Hastings, 1930:706. Electrinidae. Islands: Floreana. Distri- bution: Pacific (Pacific). Membranipora aragoi var. pacifica Osburn, 1950.— Osburn, 1950:175. Membraniporidae. Islands: Wenman, Isabela. Distribution: Gulf of California, Baja California, Galapagos (Panamic). Membranipora arborescens (Canu & Bassler, 1928b) (Morocco).—Present study. Membraniporidae. Is- lands: Santa Cruz. Distribution: Morocco, West Af- rica, Panama, Galapagos (Pan American). Closely related and possibly identical to M. savarti and M. tenuis. If so, it is a circumglobal subtropical to trop- ical species (see Banta & Carson 1977:387). Membranipora savarti (Audouin, 1826).—Osburn, 1950.—27.—Acanthodesia savartii (Canu & Bassler, 1930:4). Membraniporidae. Islands: Santiago, Es- panola, Santa Cruz, San Cristobal. Distribution: Cal- ifornia, Baja California, Costa Rica, Panama, Gulf of Mexico, Caribbean, Philippines, Australia. (Cir- cumglobal.) Membranipora tenuis Desor, 1848.—Acanthodesia denticulata.—Hastings, 1930:707. Membranipori- dae. Islands: Isabela. Distribution: Cape Cod to Bra- zil; Baja California, Panama, Galapagos, Peru, Gulf of California (Osburn 1950:27). (Pan American.) See Osburn (1950:27) about the identity of Acanthodesia dentidulata with M. tenuis. Membranipora tuberculata (Bosc, 1803).—Osburn, 1950:23.—Nichtina tuberculata.—Hastings, 1930: 706. Membraniporidae. Islands: Isabela; Osburn’s stations not listed. Distribution: British Columbia to Peru, Massachusetts to Patagonia; Europe to An- gola; Japan to Australia and New Zealand; Indian Ocean (Gordon 1984:24); Chile (Circumglobal). Micropora coriacea (Esper, 1794).— Hastings, 1930.— Canu & Bassler, 1930:11.—Osburn, 1950:105. Mi- croporidae. Islands: Espanola, Santiago, Isabela, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Floreana; Osburn’s stations not specified. Distribu- tion: British Columbia to Chile, the Juan Fernandez Islands and the Galapagos; Britain to the Mediter- ranean, Antarctic, South Atlantic, Tasman Sea (Cir- cumglobal). Micropora coriacea var. inarmata Soule, 1959.—Soule, 1959:29. Microporidae. Islands: Marchena, Santa Cruz. Distribution: Baja California, Soccoro Island, Cocos Island, New Zealand (Gordon 1984:52) (Pa- cific). Parellisina curvirostris (Hincks, 1862).— Osburn, 1950: 75.—Present study.—Ellisina curvirostris. —Has- tings, 1930:711.—Callopora curvirostris Canu & Bassler 1930:9. Alderinidae. Islands: Isabela, Flo- reana, Espanola, Santa Cruz. Distribution: Gulf of Mexico, Caribbean, Mexico to the Cocos Island and the Galapagos, Britain to Morocco, Hawaii, Western Pacific, Indian Ocean, Indonesia (Winston & Heim- berg 1986:6 (Circumglobal). Retevirgula areolata (Canu & Bassler, 1923).—Soule, 1963a:233. Alderinidae. Islands: Not given. Distri- bution: California to the Galapagos Islands, Cocos; (Soule 1963a); Chile (Moyano 1983:21) (Eastern Pa- cific). “Common at the Galapagos Islands” (Soule 1963a:233). Retevirgula lata Osburn, 1950.—Osburn, 1950:87. Calloporidae. Islands: Santiago, off Santa Cruz. Dis- tribution: Endemic. Retevirgula tubulata (Hastings, 1930).—Osburn, 1952: 86.—Pyrulella tubulata. —Hastings, 1930:709. Cal- loporidae. Islands: Floreana, Marchena, San Cris- tobal, Isabela. Distribution: Baja California and the Gulf of California to Panama and the Galapagos (Panamic). Scruparia ambigua d’Orbigny, 1841.—Scruparia che- lata. —Hastings, 1930:702. Scrupariidae. Islands: Is- abela (Tagus Cove). Distribution: “‘Distributed throughout the world except in polar waters” (Ry- land & Hayward 1977:50) (Circumglobal). Confused with S. chelata until about 1941. Scrupocellaria bertholleti (Audouin, 1826).— Hastings, 1930:733.—S. bertholetti (sic). —Osburn, 1950:133. Scrupocellariidae. Islands: Santiago, Floreana, Isa- bela. Osburn reports it from Galapagos at stations not specified. Distribution: Southern California to Panama and the Galapagos; Atlantic, Mediterra- nean, Red Sea (Circumglobal). Scrupocellaria ferox Busk, 1852.—Osburn, 1950:137. Scrupocellariidae. Islands: San Cristobal, South of Santa Cruz, Isabela (Tagus Cove). Distribution: Widely distributed in the Indian and Pacific oceans, Zanzibar, the eastern Pacific (Circumglobal). Scrupocellaria harmeri Osburn, 1947.—Osburn, 1950: 138. Scrupocellariidae. Islands: Isabela. Distribu- tion: Caribbean, California, Galapagos (Pan Amer- ican). Scrupocellaria panamensis Osburn, 1950.—Osburn, VOLUME 103, NUMBER 4 1950:142. Scrupocellariidae. Islands: Floreana. Dis- tribution: Panama and the Galapagos (Panamic). Scrupocellaria pugnax Osburn, 1950.—Osburn, 1950: 143. Scrupocellariidae. Islands: Floreana. Distribu- tion: Endemic. Scrupocellaria scruposa (Linnaeus, 1758).— Hastings, 1930:703. Scrupocellariidae. Islands: Santiago. Dis- tribution: Europe, Madeira, New Zealand. “Widely distributed in warm and temperate seas” (Ryland & Hayward 1977:140) (Circumglobal). Scrupocellaria unguiculata Osburn, 1950.—Osburn, 1950:149. Scrupocellariidae. Islands: Santiago; south of Santa Cruz; Floreana. Distribution: Endemic. Sessibugula translucens Osburn, 1950.—Osburn, 1950: 164.—Present study. Bugulidae. Islands: Isabela, Santa Cruz. Distribution: Gulf of California, Costa Rica, Galapagos (Panamic). Smittipora americana (Canu & Bassler, 1930).— Vel- umella americana. —Osburn, 1950:103. Onychocel- lidae. Islands: San Crist6bal. Distribution: West Indes, Panama, Galapagos (Pan American). Thalamoporeila californica (Levinsen, 1909). — Has- tings, 1930:716.—Osburn, 1950:112. Thalamopo- rcllidae. Islands: Santiago, Santa Cruz, Isabela; Os- burn’s stations not given. Distribution: California to Colombia and the Galapagos (Eastern Pacific). ASCOPHORA Aimulosia palliolata (Canu & Bassler, 1928).—Osburn, 1952:353. Hippoporinidae. Islands: Wenman. Dis- tribution: Florida, Gulf of California and Galapagos (Pan American). Aimulosia uvulifera (Osburn, 1914).—Osburn, 1952: 352. Hippoporinidae. Islands: South Seymour. Dis- tribution: Florida, Caribbean, Baja California, Costa Rica, Ecuador, Cocos Island (Pan American). Arthropoma cecili (Audouin, 1826).—Canu & Bassler, 1930:16. Schizoporellidae. Islands: Espafiola. Dis- tribution: Western Pacific, Indian Ocean, Red Sea, Mediterranean, Britain to South Africa, Brazil, Brit- ish Columbia to the Galapagos (Gordon 1984:85) (Circumglobal). Arthropoma circinatum (MacGillivray, 1868).—Pres- ent study. Schizoporellidae. Islands: Santa Cruz. Distribution: Australia, New Zealand, Tristan da Cunha, Southern California to Colombia and the Galapagos (Osburn 1952:334) (Circumglobal). First record in the Galapagos. Bellulopora bellula (Osburn, 1950).—Colletosia bel- lula.—Osburn, 1950:189. Cribrilinidae. Islands: Is- abela. Distribution: Cape Hatteras to Florida, Gulf of Mexico, Gulf of California, Baja California, Pleis- tocene of California, Galapagos (Winston 1982:1 34); Chile (Moyano 1983:17) (Pan American). See Lagaaij (1963:183) regarding generic placement of this spe- cies. Celleporaria albirostris (Hincks, 1884).—Holoporella 797 albirostris.—Osburn, 1952:498. Celleporariidae. Is- lands: Daphne Major. Distribution: Florida, Gulf of Mexico and Caribbean, Indian Ocean, Australia, New Zealand (Circumglobal). Celleporaria brunnea (Hincks, 1884).—Present study. — Holoporella brunnea. —Hastings, 1930:731.—Os- burn, 1952:497. Celleporariidae. Islands: Santiago, Santa Cruz, Isabela; Osburn records it from “‘21 sta- tions in the Galapagos.” Distribution: British Co- lumbia to Ecuador, Cocos Island and the Galapagos. (Eastern Pacific.) Celleporaria hancocki (Osburn, 1952).—Holoporella hancocki. —Osburn, 1952:500. Celleporidae. Is- lands: “Between Seymour and Daphne” and at Daphne Major. Distribution: Endemic. Celleporaria peristomata (Osburn, 1952).—H. oloporel- la peristomata.—Osburn, 1952:501. Celleporidae. Islands: “Between South Seymour and Daphne,” Santiago, Isabela. Distribution: Endemic. Celleporaria quadrispinosa Canu & Bassler, 1930.— Holoporella quadrispinosa.—Canu & Bassler, 1930: 38. Celleporidae. Islands: Espafiola, Floreana. Dis- tribution: Baja California and the Galapagos (Pan- amic). Celleporella hyalina (Linnaeus, 1767).—Hippothoah 'y- alina. —Hastings, 1930:720.—Osburn, 1953:277. Hippothoidae. Islands: Santiago, Isabela; Osburn’s stations not specified. Distribution: Cosmopolitan. Chorizopora brogniarti (Audouin, 1826).—Canu & Bassler, 1930:14. Chorizoporidae. Islands: Espano- la. Distribution: Australia, New Zealand, Philip- pines, China, Burma, Sri Lanka, Red Sea, Mediter- ranean, Britain, South Africa, Galapagos: Gordon 1984:113. (Circumglobal.) Cigclisula hexagonalis (Canu & Bassler, 1929).—Holo- porella hexagonalis.—Canu & Bassler, 1930:39.— Trematooecia hexagonalis.—Osburn, 1952:503. Stomachetosellidae. Islands: Espanola, San Crist6- bal, Isabela, Floreana, Santa Fé. Distribution: Gulf of California, Mexico, Galapagos (Panamic). Cigclisula porosa (Canu & Bassler, 1929).—Present study. — Holoporella porosa. —Canu & Bassler, 1930: 39.—Trematooecia porosa.—Osburn, 1952:503. Stomachetosellidae. Islands: Floreana, Santa Cruz. Distribution: Gulf of California and the Galapagos (Panamic). Cigclisula tridenticulata (Busk, 1884).—Holoporella tridenticulata.—Canu & Bassler, 1930:41.—Osburn, 1952:499. Celleporidae. Islands: Wenman, Flo- reana, Isabela, San Cristobal, Santiago. Distribution: Indian Ocean, Australia, New Zealand; Galapagos (Pacific). Cigclisula turrita (Smitt, 1873).—Holoporella turri- ta. —Hastings, 1930:732. Celleporidae. Islands: Is- abela. Distribution: Indian Ocean, China, Philip- pines, Florida, Pleistocene of Panama (Circumglobal). Powell 1971:773 gives a synonomy. American spe- cies of Cigclisula are often difficult to distinguish and 798 several species may be confused under this name (Banta & Carson 1977:400). Cleidochasma contractum (Waters, 1899).— Perigas- trella contracta.—Hastings, 1930:722.— Hippopori- na contracta.—Osburn, 1952:347. Cleidochasmati- dae. Islands: Santiago, Isabela; Osburn’s Galapagos stations not given. Distribution: Madeira; Gulf of Mexico, Caribbean, Massachusetts to Brazil; Gulf of California to Ecuador and the Galapagos (Cheetham & Sandberg 1964:1032 (Pan American). Cleidochasma porcellanum (Busk, 1860).— Present study.—Hippoporina porcellana.— Hastings, 1930: 722.—Osburn, 1952:345.—Hippoporina cleidosto- ma.—Canu & Bassler 1929:19. Cleidochasmatidae. Islands: Santiago, Isabela, Santa Cruz. Distribution: Gulf of Mexico, Caribbean; Gulf of California, Bra- zil, Pliocene; British Columbia to Peru; Mediterra- nean, Eastern Central Atlantic, Japan, Indonesia, New Zealand (Gordon 1984:124) (Circumglobal). Cleidochasma tuberculata Osburn, 1952.—Present study.—Hippoporina tuberculata.—Osburn, 1952: 346. Cleidochasmatidae. Islands: San Cristobal, Santa Cruz. Distribution: Endemic. Absence of spines is a useful character to distinguish C. tuberculata from C. porcellana. Codonellina anatina (Canu & Bassler, 1927).—Os- burn, 1952:42.—Codonella granulata.—Canu & Bassler, 1930:29.—Hastings, 1930:725. Hippopor- inidae. Islands: Wenman, Isabela, Santiago, Flo- reana, Onslow, Espanola, San Cristobal, ‘Albany.’ Distribution: Gulf of California, Galapagos, Hawaii, Gulf of Mexico (Osburn 1952:422) (Pan American). Osburn synonymized C. granulata and C. anatina, but suggested there may be ‘varietal’ differences. Ga- lapagos specimens identified as C. anatina may be of a distinct species (Soule & Soule 1973:431). Crepidacantha poissoni (Audouin, 1826).— Present study.— Hippoporina porcellana.—Hastings, 1930: 722.—Osburn, 1952:345.—Hippoporina cleidosto- ma Canu & Bassler 1929:19. Crepidacanthidae. Is- lands: Espanola, Floreana; Osburn’s stations not specified. Distribution: Southern California, Gulf of California, Panama, Cocos Island, Ecuador, Gala- pagos, Hawaii, Tahiti, New Zealand, Madeira, In- donesia, Red Sea; Florida, Brazil (Gordon 1984:124) (Circumglobal). Crepidacantha setigera (Smitt, 1873).—Osburn, 1952: 479. Crepidacanthidae. Islands: Wenman, Isabela. Distribution: Florida, Cocos Island, Galapagos (Soule 1963b) (Panamic). Cyclicopora longipora (MacGillivray, 1883).— Osburn, 1952:285. Cyclicoporidae. Islands: Southwest of Santa Cruz. Distribution: Gulf of California, Gala- pagos, San Bonito Ids.; Australia (Pacific). Cycloperiella rosacea Osburn, 1947.—Osburn, 1952: 297. Petraliidae. Islands: Isabela, Santa Cruz. Dis- tribution: Caribbean; Mexico, Panama, Galapagos (Pan American). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Dakaria biserialis (Hincks, 1885).—Osburn, 1952:330. Schizoporellidae. Islands: San Cristobal. Distribu- tion: New Zealand, Cocos Island, Galapagos, Chile (Pacific). Ascribed to Arthropoma by Moyano 1983: 17. Dakaria sertata Canu & Bassler, 1930.—Canu & Bass- ler, 1930:57.—Osburn, 1952:329.—Present study. Schizoporellidae. Islands: Floreana, Espanola, Isa- bela, Gardner-by-Hood, Santa Cruz. Distribution: Southern California, Gulf of California, Galapagos (Eastern Pacific). Diplonotos costulatus Canu & Bassler, 1930.—Canu & Bassler, 1930:31 (as D. costulatum). Sertellidae. Is- lands: Espanola, Marchena. Distribution: Endemic. S. Cairns (pers. comm.) points out Diplonotos must be masculine; it could be neuter only if it ended in -on or -um. The correct name, therefore, appears to be D. costulatus. Enantiosula manica Canu & Bassler, 1929.—Canu & Bassler, 1929.—Osburn, 1952:469.— Present study. Cheiloporinidae. Islands: Isabela, San Cristobal, Duncan, Gardner, Santiago, Onslow, South Sey- mour, Floreana. Distribution: Mexico, Gulf of Cal- ifornia, Cocos Island, Galapagos (Panamic). Among the most common and characteristic of the Gala- pagos bryozoans. Escharella major (Hincks, 1884).—Mucronella ma- jor.—Osburn, 1952:439. Smittinidae. Islands: Isa- bela, Santiago, Floreana. Distribution: Southern Cal- ifornia, Gulf of California (Eastern Pacific). Escharina pesanseris Smitt, 1973.— Mastigophora pe- sanseris. —Osburn, 1952:480. Crepidacanthidae. Is- lands: Wenman, Daphne Major, Isabela. Distribu- tion: Florida, Gulf of Mexico, Caribbean, Brazil; Gulf of California, Panama, Colombia (Pan American). Escharoides praestans (Hincks, 1882).—Osburn, 1952: 373. Exochellidae. Islands: Wenman, Isabela, Daph- ne Major. Distribution: Australia, New Zealand, Baja California, Galapagos (Pacific). Fenestrulina malusi (Audouin, 1826).—Osburn, 1952: 388. Microporellidae. Islands: Wenman, Isabela, Floreana. Distribution: British Columbia to the Ga- lapagos; Gulf of Mexico, Caribbean, Bermuda; north Atlantic, Mediterranean; Western Pacific; Indian Ocean (Cheetham & Sandberg 1964:103); Australia, Chile, southern Atlantic (Moyano 1983:19) (Cir- cumglobal). Possibly a cosmopolitan species, but morphological differences exist between Alaskan specimens and more southern material (Dick & Ross 1988:77). Hippaliosina rostigera (Smitt, 1873).—Osburn, 1952: 475. Cheiloporinidae. Islands: Isabela. Distribution: Caribbean, Gulf of Mexico, Gulf of California to Colombia and the Galapagos (Cheetham & Sand- berg 1964:1044) (Pan American). Hippomonavella parvicapitata (Canu & Bassler, 1930).—Osburn, 1952:367.—Hippomonella parvi- capitata.—Canu & Bassler, 1930:19. Hippoporini- VOLUME 103, NUMBER 4 dae. Islands: Wenman, Espafiola. Distribution: Gulf of California, Ecuador, Colombia (Panamic). Hippopleurifera mucronata (Smitt, 1873).—Osburn, 1952:301. Umbonulidae. Islands: North Seymour, Isabela, Espafiola. Distribution: Cape Hatteras south (Maturo 1968:278), Gulf of Mexico; Gulf of Cali- fornia to Colombia and the Galapagos (Pan Amer- ican). Hippopodinella adpressa (Busk, 1854).— Hastings, 1930: 729.—Osburn, 1952:467.—Present study. Cheilo- porinidae. Islands: Santa Cruz, Isabela. “A common species” at unnamed Galapagos stations (Osburn 1952:467). Distribution: Gulf of California to Costa Rica, Panama, Ecuador, Chile, and the Galapagos; Falklands, Antarctic; Britain, Mediterranean (Pan American). Hippopodinella turrita Osburn, 1952.—Osburn, 1952: 468. Cheiloporinidae. Islands: Floreana, San Chris- tobal. Distribution: Endemic. Hippoporella gorgonensis Hastings, 1930.— Hastings, 1930:723.—Osburn, 1952:349.— Present study. Cleidochasmatidae. Islands: Wenman, Santiago, Santa Cruz, Santiago, Isabela, Espafiola. Distribu- tion: Southern California to the Galapagos (Eastern Pacific). Hippoporella rimata Osburn, 1952.—Osburn, 1952: 352. Cleidochasmatidae. Islands: Isabela, San Cris- tobal. Distribution: Ecuador, Mexico, Galapagos (Panamic). Hippoporella species indetermined.— Present study. Hippoporellidae. Islands: Santa Cruz. Distribution: Endemic. Resembles H. gorgonensis, but there are six oral spines and the colony is persistently unilam- inar. Hippoporidra granulosa Canu & Bassler, 1930.—Canu & Bassler, 1930:43.—Osburn, 1952:357. Cleido- chasmatidae. Islands: San Cristobal. Distribution: Gulf of California and the Galapagos (Panamic). Hippoporidra spiculifera (Canu & Bassler, 1930).— Present study.—Hippotrema spiculifera.—Canu & Bassler, 1930:43. Cleidochasmatidae. Islands: Es- panola, Floreana, Santa Cruz. Distribution: Mexico, Clarion Island, Panama, Galapagos (Panamic). Hippoporina ampla Osburn, 1952.—Osburn, 1952:348. Hippoporinidae. Islands: San Cristdbal, Floreana, Santiago. Distribution: Endemic. Hippoporina pertusa (Esper, 1796).— Hippodiplosia pertusa.—Hastings, 1930:724. Hippoporinidae. Is- lands: Santiago, Isabela. Distribution: Gulf of Mex- ico, Caribbean, Massachusetts to Brazil; Spitzbergen to the Mediterranean; southern California to Colom- bia and the Galapagos (Pan American). Cheetham & Sandberg (1964:1041) discuss the generic place- ment. Hippoporina verilli Maturo & Schopf, 1968:56.—Hip- podiplosia americana.—Osburn, 1952:339. Hippo- porinidae. Islands: Isabela, Santiago. Distribution: Massachusetts to Brazil, Gulf of Mexico; Gulf of 799 California, to Panama, Costa Rica, and the Gala- pagos (Winston 1982:139) (Pan American). Hippothoa divaricata (Lamouroux, 1821).—Osburn, 1952:278. Hippothoidae. Islands: Not listed. Dis- tribution: Widely distributed (Cosmopolitan). De- tailed synonymy and distribution in Ryland & Gor- don (1977:20). “Typical H. divaricata seems to be Atlantic in distribution” Gordon (1984:111). A sub- species occurs in New Zealand. Hippothoa flagellum Manzoni, 1870.—Osburn, 1952: 278. Hippothoidae. Islands: Not given. Distribution: Europe, Southeastern U.S., Caribbean, Japan, Ha- wali, California, Mexico, Galapagos (Morris 1980); Chile, south Atlantic (Moyano 1983:20) (Circum- global). The world wide synonymy is complex. Ry- land & Gordon (1977:25) consider many eastern Pa- cific specimens to belong to a distinct species. Lagenicella hippocrepis (Busk, 1856).—Costazia hip- pocrepis.—Hastings, 1930:731.—Lagenipora hip- pocrepis (Busk, 1856).—Osburn, 1952:489. Teucho- poridae. Islands: Wenman, Floreana. Distribution: Southern California to Panama and the Galapagos (Eastern Pacific). Lagenicella lacunosa Bassler, 1934.—Lagenipora ver- rucosa.—Canu & Bassler, 1930:35.—Lagenipora la- cunosa. —Osburn, 1952:491. Teuchoporidae. Is- lands: Wenman, Floreana, Isabela, Santiago, Espanola, Fernandina. Distribution: southern Cali- fornia to Peru and the Galapagos (Eastern Pacific). The family Teuchoporidae Neviani, 1895 has pri- ority over the Phylactellidae Harmer, 1957:896 (Gordon 1984:80). Lagenicella marginata Canu & Bassler, 1929.—La- genipora marginata.—Osburn, 1952:490. Teucho- poridae. Islands: Isabela, San Cristdbal, Espanola, Santa Fé. Distribution: Panama, Cocos Island, Gulf of California, Galapagos; east coast of Florida, Gulf of Mexico (see Winston 1982:144) (Pan American). Lagenicella punctulata (Gabb & Horn, 1862).—La- genipora punctulata.—Osburn, 1952:486. Teucho- poridae. Islands: Not given. Distribution: Alaska to the Galapagos (Eastern Pacific). Lagenicella species undetermined. —Present study. Teuchoporidae. Islands: Santa Cruz. Distribution: Endemic. Oligoserial like L. /acunosa Bassler, but the zoids are minute for this genus (0.35 x 0.22 mm); peristome low, much shorter than zoids: inner ends of pseudopores circular. No complete ovicells ob- served. Lagenicella spinulosa Hincks, 1884.—Lagenipora spi- nulosa.—Hastings, 1930:730. Teuchoporidae. Is- lands: Santiago. Distribution: Alaska, British Co- lumbia, California, Gulf of California (Osburn 1952: 487) (Eastern Pacific). Mamillopora cupula Smitt, 1873.—Osburn, 1952: 517.—Canu & Bassler, 1928:45. Mamilloporidae. Is- lands: Espafiola (Canu & Bassler 1928a:45) ‘Abun- dant about the Galapagos Islands’ (Osburn 1952: 800 518). Distribution: Gulf of California to Panama and the Galapagos; Gulf of Mexico, Caribbean. (Pan American.) Microporella californica (Busk, 1856).-Osburn, 1952: 382. Microporellidae. Islands: Isabela, Santiago. Distribution: British Columbia to the Galapagos (Eastern Pacific). Microporella ciliata (Pallas, 1766).—Hastings, 1930: 727.—Osburn, 1952:377. Microporellidae. Islands: Santiago, Isabela; Osburn’s stations not given. Dis- tribution: Australia, New Zealand; Philippines, Or- egon to the Galapagos; Chile, Falklands, Juan Fer- nandez, Tristan de Cunha; Britain, Mediterranean (Gordon 1984:101) (Circumglobal). Microporella gibbosula Canu & Bassler, 1930.—Canu & Bassler, 1930:20.—Osburn, 1952:386.— Present study. Microporellidae. Islands: Espanola, Santa Cruz; Osburn recorded it from 8 unspecified stations in the Islands. Distribution: Gulf of California, Pan- ama, Costa Rica, Colombia and the Galapagos (Os- burn 1952:386) (Panamic). Microporella marsupiata (Busk, 1860).—Osburn, 1952: 383. Microporellidae. Islands: Isabela, Santiago, Santa Fé, San Cristobal. Distribution: Gulf of Cal- ifornia, Cocos Island and the Galapagos (Pan Amer- ican). Synonym, M. coronata Marcus 1938:235: “Mediterranean, Madeira, Cape Verde Islands, Brit- ain, Gulf of Guinnea, Saint Helena.” The synonymy of American species is in doubt (Gordon 1984:102). Microporella pontifica Osburn, 1952.—Osburn, 1952: 383. Microporellidae. Islands: Isabela. Distribution: Gulf of California to the Galapagos; Post-Pleistocene of Louisiana (Pan American). Microporella tractabilis Canu & Bassler, 1930.—Canu & Bassler, 1930:22.—Present study. Microporelli- dae. Islands: Espanola, Floreana, Santa Cruz. Dis- tribution: Colombia, Panama, Galapagos; Post- Pleistocene of Louisiana (Pan American). Cheetham & Sandberg (1964:1036) found Osburn’s descrip- tions differed in important respects from Canu & Bassler’s Galapagos specimens. Pachycleithonia nigra Canu & Bassler, 1930.—Canu & Bassler, 1930:27. Watersiporidae. Islands: Espa- Nola. Distribution: Endemic. Osburn (1952:472) considered P. nigraa subspecies of W. cucullata (Busk 1854). At least some of his P. nigra specimens were of W. arcuata Banta 1968. See also Soule & Soule (1975:308). Parasmittina californica (Robertson, 1908).—Osbum, 1952:416. Smittinidae. Islands: Wenman. Distri- bution: Southern California, Baja California, Gala- pagos (eastern Pacific). The identity of Galapagos material with that of Mucronella californica Rob- ertson 1908:308 is questionable. Confused with and possibly identical to P. trispinosa of authors (Soule & Soule 1973:424). Parasmittina crosslandi (Hastings, 1930).—Present PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON study. Smittinidae. Islands: Santa Cruz. Distribu- tion: Gulf of California, Baja California, Panama and the Galapagos (Panamic). The first record in the Ga- lapagos, if Soule & Soule (1973:382) are correct in regarding all of Canu & Bassler’s and Osburn’s spec- imens of P. crosslandi as P. dolobrata. Parasmittina dolobrata Soule & Soule, 1973.—Smit- tina crosslandi.—Hastings, 1930:726.—Parasmit- tina crosslandi.—Osburn, 1952:418 (part).—Para- smittina trispinosa: Canu & Bassler 1930:27; Osburn 1952:418 (part); P. spathulata: Osburn 1952:415 (part); Parasmittina dolobrata: Soule & Soule 1973: 421 and 393. See Soule & Soule (1973:421). Smit- tinidae. Islands: Wenman, Santiago, Santa Cruz, San Cristobal, Marchena, Floreana, Baltra, Isabela, Es- panola. Distribution: Colombia, Panama, Costa Rica and the Galapagos (Soule & Soule 1973:423) (Pan- amic). Parasmittina frazeri Osburn, 1952.—Osburn, 1952:420. Smittinidae. Islands: Isabela. Distribution: Mexico, Ecuador, Galapagos (Panamic). Parasmittina species undetermined.—Present study. Smittinidae. Islands: Santa Cruz. Distribution: En- demic. Like P. hastingsae Soule & Soule 1973:417, but smaller, and the aperture is broader than long. Phidolipora pacifica (Robertson, 1908).—Osburn, 1952: 449. Sertellidae. Islands: Wenman, Isabela, Santia- go. Distribution: Oregon, Gulf of California, to Peru and the Galapagos (Eastern Pacific). Phylactella aperta Osburn, 1952.—Osburn, 1952:482. Phylactellidae. Islands: Isabela. Distribution: En- demic. Porella columbiana O’Donoghue, 1923.—Osburn, 1952:398. Smittinidae. Islands: Isabela and at a sta- tion between Floreana and Santa Fé. Distribution: British Columbia, Puget Sound, Southern California (Eastern Pacific). Porella patens Osburn, 1952.—Osburn, 1952:397. Smittinidae. Islands: Wenman. Distribution: Southern California, Baja California, Cocos Island, Galapagos (Eastern Pacific). Puellina innominata (Couch, 1844).—Canu & Bassler, 1930:13.—Present study. Cribrilinidae. Islands: Es- paola. Distribution: Philippines, India, Mediter- ranean, Britain, Madeira, Gulf of Mexico, Califor- nia, Australia, New Zealand (Gordon 1984:64); Peru, Chile (Moyano 1983:19) (Circumglobal). Osburn (1950:187) considered P. innominata to be synon- ymous with Cribrillaria radiata; some of his records of C. radiata in the Galapagos may be of P. innom- inata. Puellina radiata (Moll, 1803).—Canu & Bassler, 1930: 13.—Present study.—Colletosia radiata.—Osburn, 1950:188. Cribrilinidae. Islands: Floreana, Santa Cruz. Distribution: British Columbia to Peru, Cocos Island and the Galapagos; Gulf of Mexico, Carib- bean, North Carolina to Brazil; Western Pacific, In- VOLUME 103, NUMBER 4 dian Oceans (Circumglobal). The Cribrilina radiata of MacGillivray (1889:317) and Powell 1967:223 is probably C. innominata (Gordon 1984:64). See Gor- don for generic placement. Puellina setosa (Waters, 1889).—Osburn, 1950:186. Cribrilinidae. Islands: Isabela. Distribution: British Columbia, Puget Sound, Oregon, California, Ma- deira, Naples (Pan American). Reptadeonella tubulifera (Canu & Bassler, 1930).— Adeona tubulifera.—Canu & Bassler, 1930:34.—Os- burn, 1952:442. Adeonidae. Islands: Wenman, Is- abela, San Cristobal, Santa Fé, Santiago, Floreana, Espanola. Distribution: Ecuador and the Galapagos Islands (Osburn 1952:442) (Eastern Pacific). Reptadeonella violaecea (Johnston, 1847).—Adeona violaecea.—Osburn, 1952:441. Adeonidae. Islands: Not given. Distribution: Britain, Mediterranean, Cape Verde Islands, Western Africa, Gulf of Mexico, Ca- ribbean; Baja California to Colombia and the Ga- lapagos (Hayward & Ryland 1979:82) (Pan Ameri- can). Reteporella striata Canu & Bassler, 1930.—Diplonotos striatum. —Canu & Bassler, 1930:32. Sertellidae. Is- lands: Marchena. Distribution: Endemic. See Harm- er (1934:624) regarding the tentative generic place- ment of this species. Reteporellina denticulata var. gracilis Osburn, 1952.— Osburn, 1952:447. Sertellidae. Islands: Wenman, Is- abela, South Seymour, Daphne Major, Santiago. Distribution: Costa Rica, Cocos Island, Galapagos (Panamic). Rhynchozoon rostratum (Busk, 1856).— Hastings, 1930: 728.—Osburn, 1952:456.—Present study. Sertelli- dae. Islands: Santa Cruz; unspecified stations in the Galapagos (Osburn 1952:458). Distribution: Carib- bean, Gulf of Mexico, Southern California to Co- lombia, Panama and the Galapagos; Indonesia (see Winston & Heimberg 1986:38) (Circumtropical). If R. rostratum is synonymous with R. tumulosum (Soule & Soule 1964:33), the range is Arctic to trop- ical eastern Pacific (Dick & Ross 1988:84). Robertsonidra oligopus Osburn, 1952.—Osburn, 1952: 295. Petraliidae. Islands: about 30 km northeast of Pinta. Distribution: California, Baja California, Ga- lapagos (eastern Pacific). Savignyella lafontii (Audouin, 1826).—Savignyella la- fonti.—Osburn, 1952:288. Savignyellidae. Islands: Isabela. Distribution: Circumglobal in warmer wa- ters, including in America, the Gulf of Mexico, Bra- zil, California, Colombia, Galapagos (Circumglob- al). Schizmopora anatina Canu & Bassler, 1930.—Osburn, 1952:493.—Osthimosia anatina.—Canu & Bassler, 1930:42. Celleporidae. Islands: Isabela, Floreana, South Seymour, Santiago, Espafiola, Marchena, San- ta Fé, Gardner, Duncan. Distribution: Gulf of Cal- ifornia, Mexico, Panama (Panamic). 801 Schizmopora margaritacea (Pourtalés, 1867).—Os- burn, 1952:495. Celleporidae. Islands: Santiago. Distribution: North Carolina to the Caribbean; Ga- lapagos (Pan American). Schizoporella cornuta (Gabb & Horn, 1862).—Has- tings, 1930:721.—Osburn, 1952:321.—Present study. —Schizopodrella biaperta.—Canu & Bassler, 1930:1. Schizoporellidae. Islands: Espafiola, Santi- ago, Isabela, Santa Cruz; Osburn’s stations not listed. Distribution: Alaska to the Galapagos; Gulf of Mex- ico, Caribbean, Massachusetts to Florida (Cheetham & Sandberg 1964:1030) (Circumglobal). Schizoporella dissimilis Osburn, 1952.—Osburn, 1952: 322. Schizoporellidae. Islands: Isabela, Santiago, Santa Fé. Distribution: Gulf of California and the Galapagos (Panamic). Schizoporella trichotoma (Waters, 1918).—Hastings, 1930:721.—Osburn, 1952:318. Schizoporellidae. Is- lands: Wenman, Floreana, San Cristobal, Santa Cruz, Isabela. Distribution: Cape Verde Islands; Gulf of California, Galapagos (Pan American). Schizoporella unicornis (Johnston, 1847).—Osburn, 1952:317. Schizoporellidae. Islands: Santiago. Dis- tribution: Reported from almost every ocean, but lumping makes many of these records suspect (Cos- mopolitan). Schizotheca umbonata Osburn, 1952.—Osburn, 1952: 452. Sertellidae. Islands: Isabela. Distribution: En- demic. Semihalswellia sulcosa Canu & Bassler, 1930.—Canu & Bassler, 1930:15.—Osburn, 1952:304. Giganto- poridae. Islands: Marchena, Isabela. Distribution: Endemic. Smittina landsborovi (Johnston, 1847).—Osburn, 1952: 401. Smittinidae. Islands: Not listed. Distribution: Western Norway to the western Mediterranean; du- biously recorded world wide (Hayward & Ryland 1979:99) (Cosmopolitan). The identity with Euro- pean material is questionable. Smittina smittiella Osburn, 1947.— Osburn, 1952:405. Smittinidae. Islands: Wenman, Santa Cruz, Isabela. Distribution: Gulf of Mexico, Caribbean, Brazil; Ec- uador and the Galapagos (Pan American). Smittoidea pacifica Soule & Soule, 1973.—Osburn, 1952:410.—Soule & Soule, 1973:381.—Smittina re- ticulata.—Canu & Bassler, 1930:27.—Smittoidea pacifica Soule & Soule 1973:380. Smittinidae. Is- lands: Wenman, Isabela, San Cristobal, Duncan, Santiago (Osburn 1952:410); off Floreana (Canu & Bassler 1930:27). Distribution: Hawaii, Galapagos (Soule & Soule 1973:380) (eastern Pacific). Stephanosella vitrea Osburn, 1952.—Osburn, 1952:370. Hippoporinidae. Islands: San Cristébal. Distribu- tion: Puget Sound to the Galapagos (eastern Pacific). Stylopoma informatum (Lonsdale, 1845).—Osburn, 1952:337. Schizoporellidae. Islands: Floreana, San- tiago. Distribution: West Indies, Brazil, Galapagos. 802 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON I have a specimen from the Gulf of California. (Pan American). Stylopoma spongites (Pallas, 1766).— Hastings, 1930: 721. Schizoporellidae. Islands: Santiago, Isabela. Distribution: Florida, Gulf of Mexico, Caribbean; Bermuda, Brazil, Pliocene of Panama, Galapagos (Pan American). The taxonomy is discussed by Cheetham & Sandberg (1964:1030). Tetraplaria veleroae Osburn, 1952.—Osburn, 1952:467. Cheiloporinidae. Islands: About 10 km SSW of Flo- reana; Isabela. Distribution: Endemic. Trypostega venusta (Norman, 1864).—Canu & Bassler, 1930:15.—Osburn, 1952:281.— Present study. Hip- pothoidae. Islands: Espanola, Floreana, Santa Cruz; Osburn’s stations not listed. Distribution: Britain to the Caribbean, Gulf of Mexico; Indian Ocean; Cal- ifornia to Cocos Island and Ecuador, the Philippines, Australia, Loyalty Islands, and elsewhere. (Circum- global). : Watersipora arcuata Banta, 1968.—Wéatersipora cu- cullata.—Hastings, 1930:729.— Watersipora cucul- lata var. nigra. —Osburn, 1952:472. Watersiporidae. Islands: Santiago, Isabela; Osburn’s stations in the Galapagos are unspecified. Distribution: California, Baja California, Gulf of California and the Gala- pagos; Australia and New Zealand (Pacific). Hast- ing’s Pl. 15, figs. 99 and 101, are probably of W. arcuata. Some or all of Osburn’s records of this spe- cies in the Galapagos probably belong here. See Soule & Soule 1975:308. Watersipora cucullata (Busk, 1854).— Hastings, 1930: 729.—Osburn, 1950:472. Watersiporidae. Islands: Santiago, Isabela; Osburn’s stations unspecified. Distribution: Atlantic, Mediterranean, Red Sea, In- dian Ocean, Amoy (Circumglobal). This species has not yet been recorded with certainty in the Gala- pagos; it may be present but lumped with W. arcuata and Pachycleithonia nigra; see Soule & Soule (1975: 308, 1985:297). PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 803-814 PYRGULOPSIS BRUNEAUENSIS, A NEW SPRINGSNAIL (GASTROPODA: HYDROBIIDAE) FROM THE SNAKE RIVER PLAIN, SOUTHERN IDAHO Robert Hershler Abstract. — Pyrgulopsis bruneauensis, new species, from thermal springs along Hot Creek and Bruneau River in Owyhee County, Idaho (upper Snake River drainage), is described. Distinguishing features include a combination of small size (<2.8 mm shell height); globose-low conic shell; and penis with an elongate, muscular filament and a small lobe bearing a single distal glandular ridge. Pyrgulopsis bruneauensis appears closest morphologically to P. amargosae Hershler, 1989, from the Death Valley System to the south, but this opinion may require modification after other congeners from the Snake River region receive anatomical study. The diverse molluscan fauna of the Snake River drainage includes several gastropods that lack formal description, despite their having been known to malacologists for a number of years. One of these is a spring- snail from the Hot Creek area of upper Snake River drainage in northern Owyhee County, Idaho, that was first collected by Borys Malkin in 1952 (letter, W. F. Barr to J. P. E. Morrison, 14 July 1953). Malkin’s col- lection, plus one made by Barr in 1954, were sent to Morrison, who recognized the dis- tinctiveness of the snail and used new manuscript genus and species names for it in the National Museum of Natural History (USNM) collection. Taylor (1982) later pur- sued additional field and laboratory study of this snail. Springflows in the Hot Creek area have declined dramatically in recent years be- cause of groundwater mining, and conser- vation of the Bruneau Hot Springsnail has become an important issue. This species has been proposed for listing under the Endan- gered Species Act (USDI 1985), and mea- sures currently are being taken to protect remaining populations. The author visited the Hot Creek area in June 1989 and made collections of the snail, which were used to prepare the following description. Family Hydrobiidae Troschel, 1857 Genus Pyrgulopsis Call and Pilsbry, 1886 Pyrgulopsis bruneauensis, new species Bruneau Hot Springsnail Figs. 1-6, Table 1 Bruneau Hot Spring snail.—USDI 1984: 21673.—USDI 1985:33803. Warm Springs Snail.— Armantrout 1985:35. “undescribed genus.’ — Taylor 1985:291. Material examined.—Spring along west side of Bruneau River, ca. 100 m downflow from Hot Creek’s confluence with the river, Owyhee County, Idaho, T 7S, R 6E, SW 4 sec. 34, USNM 860507 (holotype), USNM 860508 (paratypes; 18 dry shells and a large alcohol series), ANSP 376090 (paratypes; six dry shells), UF 161474 (paratypes; six dry shells).—Seep along south-east side of Hot Creek, ca. 50 m upflow from confluence with Bruneau River, T 8S, R 6E, NE 4 sec. 3, USNM 860509.— Hot Creek Falls, T 8S, R 6E, NW 4 sec. 3, USNM 791466 (Malkin coll.), USNM 860510 (Barr coll.). Diagnosis. —A moderate-sized species with globose to low-conic shell. Penis with PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. SEM micrographs of shells of Pyrgulopsis bruneauensis Hershler, n. sp. a, Holotype, USNM 860507; b-d, Paratypes, USNM 860508, lot as above; e, f, USNM 791466; g-i, USNM 860509. The holotype is 2.48 mm tall (other micrographs printed to the same scale). VOLUME 103, NUMBER 4 small lobe and elongate, muscular filament. Distal edge of lobe bearing a single glandular ridge. Description. — Shell (Figs. 1, 2a, b): mor- phometric data were obtained as described in Hershler (1989) and are in Table 1. Thin, transparent, white-clear, but appearing black because of animal pigmentation. Slightly taller than wide; body whorl large (79-91% of shell height). Whorls, 3.75—4.25, round- ed, with pronounced adapical shoulders. Aperture about 55% of shell height, ovate, slightly taller than wide, rounded below, very slightly angled above. Outer lip thin, slightly prosocline (Fig. 1d); inner lip thickened but not reflected, narrowly adnate above or slightly separated from body whorl. Um- bilicus broadly open. Protoconch slightly protruding; surface usually whitened and slightly eroded, generally smooth although small area of wrinkled mMicrosculpture sometimes visible (Fig. 2b). Teleoconch growth lines moderately pronounced; nu- merous faint spiral lines also present. Operculum (Fig. 3a, b): amber, thin, nar- rowly ovate, paucispiral, with eccentric nu- cleus. End farthest from nucleus angled. Radula (Fig. 4): radular formula, (4-6)-1- (4-6)/1-1, 3-1-3(4), 21-26, 22-26. Width of central tooth, 28-34 um. Basal cusps of cen- tral teeth small, originating from lateral an- gles; lateral angles well expanded; basal pro- cess moderately excavated. Central cusps of central and lateral teeth slightly enlarged. Anatomy (from relaxed, preserved ma- terial; Figs. 2c-f, 3b, d, 5, 6): head-foot and dorsal visceral coil covered with dark, grey- 805 black epithelial pigment. Pigment some- what lighter around the eyes, on cephalic tentacles (Fig. 5a), along broad strips along the sides of the head-foot, and on dorsal prostate gland (of male). Edge of mantle col- lar, sole of foot, distal lips of snout, and majority of dorsal operculigerous lobe un- pigmented. Cephalic tentacles without hypertrophied ciliary tufts (Fig. 2c). Ctenidial filaments, ca. 20, triangular, plicate, moderately broad relative to width of pallial cavity. Frontal edges of filaments with two narrow stripes of grey pigment. Osphradium about 25% of ctenidium length, slightly posterior to cen- ter of ctenidial axis. Kidney scarcely bulging into pallial cavity; renal opening small, ringed with fleshy lips. Hypobranchial gland not obvious in dissection or section. Stom- ach and style sac sub-equal in length; small triangular caecal chamber protruding from posterior edge of stomach. Testis ofnumerous simple lobes (draining to vas efferens), large (1.5 whorls), filling about 50% of body length and extending from near apical tip of animal to posterior edge of prostate gland (covering stomach). Seminal vesicle of a few thickened coils pressed against posterior edge of stomach. Prostate gland small, bean-shaped, with about 50% of length in pallial roof. Anterior vas deferens exiting proximal to tip of gland. Penis (Figs. 2d-f, 3d, 6) moderate-large, usually uncoiled and protruding beyond edge of mantle collar. Black sub-epithelial pig- ment dense in virtual entirety of filament. somewhat lighter in lobe and scattered in 2 Fig. 2. SEM micrographs of shell protoconchs and critical point dried bodies of P. bruneauensis. a, b, Protoconch, paratypes, USNM 860508 (scale bars = 136 um, 150 um); c, Dorsal head, USNM 860509 (200 um); d, Dorsal aspect of penis, USNM 860508 (250 um); e, f, Dorsal aspects of head-foot and penis, USNM 860509 (0.43 mm). Pf = penial filament; Pl = penial lobe. Fig. 3. Operculae and histological sections of P. bruneauensis, paratypes, USNM 860508. a, Dorsal operculum (0.38 mm); b, Cross section of mid capsule gland (0.1 mm). Arrow indicates ventral channel of gland; c, Ventral operculum (0.38 mm); d, Cross section of male just posterior to nerve ring (0.25 mm). Note elongate, muscular penis (to left). Z 0 Ee cS a rm NH < ca fy ° > fH =| Oo eo N J < 2S © ° — © faa] ea rc B fu ©) N © Z A sal ea Q © pe Au VOLUME 103, NUMBER 4 807 808 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pn aS Fig. 4. SEM micrographs of radulae of P. bruneauensis. a, Central teeth, paratypes, USNM 860508 (15.0 um); b, Centrals, USNM 860510 (13.6 um); c, Laterals, USNM 860508 (12.0 um); d, Outer marginals, lot as above (7.5 um). e, Laterals, USNM 860510 (10.0 um); f, Inner marginal, USNM 860508 (7.5 um). VOLUME 103, NUMBER 4 remaining penis. Dorsal penis without hy- pertrophied ciliary patches. Filament nar- row, equal to or longer than proximal penis, much longer than lobe. Filament tapering little along most of length, near-circular in cross section. Thick layer of circular mus- cles of filament obvious in section (Fig. 3d). Lobe stout, slightly tapered distally. Single glandular ridge confined to lobe, usually along distal edge. Vas deferens uncoiled in penis, positioned near outer edge. Ovary about one whorl, of simple lobes, covering posterior stomach chamber. Al- bumen gland (Ag) slightly longer than cap- sule gland (Cg). Renal oviduct (Ov) with large, complex coil on posterior albumen gland. Seminal receptacle (Sr) a small sac with short duct into posterior loop of ovi- duct. Bursa copulatrix (Bu) moderate-sized, ovate, partly posterior to albumen gland. Duct from bursa copulatrix broad and elon- gate, traversing much of albumen gland be- fore joining oviduct. Ventral channel (Vc) moderately wide, separated from capsule gland lumen by pronounced fold (Fig. 3b). Capsule gland opening simple, sub-termi- nal. Distal to opening, edge of gland (base of ventral channel) continued distally (as gutter) to near edge of mantle collar (not illustrated). Variation. —Shells from the Hot Creek spring and those along the Bruneau River differed significantly in shell measurements (9 of 12 pairwise comparisons [by sex] sig- nificant, P = 0.05, Tukey HSD Test) and translation rate (0.4 > P > 0.3), reflecting the larger size and squatter shells from the type locality. These populations also are dif- ferentiated by penial form, with males from the type locality (Fig. 6a—d) having smaller penes (scarcely protruding beyond mantle collar) with shorter filaments than those from the Hot Creek spring (Fig. 6f-i). These differences do not, however, appear to jus- tify taxonomic distinction of the popula- tions. Etymology. —The species name refers to the snail’s occurrence in Bruneau River drainage. 809 0.5 mm b Sr Bu Ag Ov Cg 2 0.5 mm Fig. 5. Anatomy of P. bruneauensis, paratypes, USNM 860508. a, Dorsal aspect of head showing typ- ically near-uniform dark pigment; b, Left lateral aspect of pallial oviduct and associated structures. Ag = al- bumen gland; Bu = bursa copulatrix; Cg = capsule gland; Cga = opening of capsule gland; Dbu = duct from the bursa copulatrix; Ov = renal oviduct; Sr = seminal receptacle; Vc = ventral channel of capsule gland. Remarks. —Pyrgulopsis bruneauensis is readily separable from all but one hydrobiid of the Snake River and adjacent areas by its small size and squat shell (““Fluminicola”’ minutissima Pilsbry, 1907, from Snake River basin near the Idaho-Oregon border is even smaller and more globose than the 810 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON b VOLUME 103, NUMBER 4 811 Table 1.—Shell parameters for Pyrgulopsis bruneauensis. WH = number of whorls, SH = shell height, SW = shell width, LBW = length of body whorl, WBW = width of body whorl, AL = aperture length, AW = aperture width, W = whorl expansion rate, D = distance of generating curve from coiling axis, T = translation rate, SA = aperture shape. Measurements are in mm. USNM 560507 (holotype) 4.00 2.48 2.03 2.12 USNM 560508 (paratypes) Nessie 234 6 196) 22011 (6, n = 8) s 0.12 0.10 0.12 0.12 (paratypes) x BS) DRE NAD) 139 (2, n = 7) s 0.17 0.15 0.04 0.10 USNM 560509 (6, n = 10) MBO DAI DEG YL s 0.18 0.16 0.20 0.17 (2, n = 10) X 3.88 2.35 1.96 2.02 s 0.21 0.16 0.08 0.12 1G ORR IES Saeeele? ale Am O160) 83555). Lett Ae) West} es AOS) OB SES I O/ O07 O07 10108) F0s7- 0:04 .0'53' 0104 LES ele POS see alG O54" = 93:96) 1512 OOS OLO7T O03 0237005 0:41 0104 1S 2a eye le Ss Se OD OFS 908 83229) 109 OS OLN Ol O19 O08 O42 O07 oo) esky} AAG OLS Ses 0.06 0.08 0.05 0.32 0.04 0.43 0.05 above). The penis of P. bruneauensis is sim- ilar to that of P. amargosae from Death Valley System, California, in that the latter also has an elongate filament darkly pig- mented with sub-epithelial granules (Her- shler 1989:figs. 17b, 18b-—e). A close historic relationship between the upper Snake River (prior to its integration with the lower river) and Lahontan drainage to the south (which then was integrated with Death Valley Sys- tem) was postulated by Minckley et al. (1986: 534, fig. 15.4). A detailed discussion of the relationships of P. bruneauensis must, how- ever, await anatomical study of other re- gional Pyrgulopsis. Distribution and habitat.—The snail oc- curs in small thermal springs (to 35°C) along Hot Creek and along the Bruneau River in the immediate vicinity of the creek’s con- fluence (Figs. 7, 8).1 Hot Creek, fed by nu- merous small springs, flows about one km before entering Bruneau River near the mouth of Bruneau Canyon. Discharge of the creek historically was at least 4.0 cfs (War- ing 1965:30). At the upper end of the creek springflow historically drained via Hot Creek Falls into a large pool (Indian Bath- tub), which now is virtually dry (compare Fig. 8b and 8c) as a result of water table decline. During a trip to the area on 23 June 1989, the following snail-positive localities were visited: ' Local endemism of this species appears likely. There are no historic records for the snail from extra-limital areas, and a helicopter survey of thermal springs in southwest Idaho and southeastern Oregon (Bruneau, Jarbridge, South Owyhee River basins) conducted dur- ing January 1987 did not reveal additional populations (S. M. Chambers, unpublished report, February 3, 1987; Hershler, personal observations on materials collected on this survey). — Fig. 6. Whole mounted penes of P. bruneauensis. a-e, Paratypes, USNM 860508 (a-c, dorsal, d, e, ventral aspects). f-i, USNM 860509 (dorsal aspects). The scale bar in ‘‘a” equals 1.0 mm. All other photographs are printed to the same scale except “‘e”’ (0.25 mm). Note the darkly pigmented filament and small area of glandular ridges in the distal penial lobe (particularly obvious in ‘‘e’’). 812 Fig. 7. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 4 Gaging / Map showing collecting localities in Hot Creek area, Owyhee County, Idaho. Modified from USGS 1947 Hot Spring quadrangle (7.5 minute series [topographic]). (a) Small (ca. 10 cm?) seep on rock wall just above floor of Indian Bathtub. About four snails were observed (none collected) on the algal-covered drip wall (none found in small pool below). (b) Small seeps emerging along base of low outcrop on the east side of lower Hot Creek. Snails were common just below spring orifices, on moistened ground cov- ered by grass. (c) A large (3-4 m high, 3 m across) drip wall on western side of Bruneau River downflow from Hot Creek confluence. Snails were extremely common on the wet, algal- Fig. 8. Photographs of the Hot Creek area. a, Confluence of Hot Creek (to right) and Bruneau River, 9-20- 89; b, Indian Bathtub under historic flow conditions, 11-73; c, Indian Bathtub and upper Hot Creek under recent flow conditions, 4-4-89. 814 covered walls and in a pool below. Snails have also been collected from small thermal spring vents along the east side of the river just upflow from the creek’s confluence (P. Olmstead, pers. comm., June 1989). Several basommatophoran pulmonates were found in association with P. bru- neauensis in the spring brooks, and an un- related hydrobiid, Fluminicola cf. hindsi (Baird), was collected at the confluence of Hot Creek and the Bruneau River. Acknowledgments Fieldwork was supported by funds pro- vided by the United States Fish and Wildlife Service (contract to W. Minshall). Assis- tance in the field was provided by J. Gore (USFWS), P. Olmstead (BLM), G. W. Min- shall, and C. T. Robinson. Scanning elec- tron micrographs were taken and developed by W. Brown and S. Braden (NMNH Scan- ning Electron Microscopy Laboratory). Beth Fricano prepared histological sections and whole mounts. Paul Greenhall and M. Ryan (NMNH, Invertebrate Zoology) measured shells, and prepared line drawings and maps respectively. Peter Bowler and P. Olmstead provided habitat photographs. Useful crit- icism of the manuscript was provided by P. Bowler, J. Gore, G. W. Minshall, G. Mla- denka, and P. Olmstead. Literature Cited Armantrout, N. B. 1985. Agency Report. Bureau of Land Management. — Proceedings of the Desert Fishes Council 13:34—37. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hershler, R. 1989. Springsnails (Gastropoda: Hydro- biidae) of Owens and Amargosa (exclusive of Ash Meadows) River drainages, Death Valley System, California-Nevada.— Proceedings of the Biological Society of Washington 102:176-248. Minckley, W. L., D. A. Hendrickson, & C. E. Bond. 1986. Geography of western North American freshwater fishes: description and relationships to intracontinental tectonism. Pp. 519-613 (+ bibliography) in C. H. Hocutt & E. O. Wiley, eds., The zoogeography of North American freshwater fishes. John Wiley & Sons, New York, New York. Taylor, D. W. 1982. Status report on Bruneau Hot Spring Snail.— Unpublished report submitted to United States Fish and Wildlife Service, Boise Endangered Species Field Office, Idaho, 14 pp. . 1985. Evolution of freshwater drainages and molluscs in western North America. Pp. 265- 321 in C. J. Hocutt & A. B. Leviton, eds., Ce- nozoic history of the Pacific Northwest. AAAS & California Academy of Sciences, San Fran- cisco, California. United States Department of the Interior. 1984. En- dangered and threatened wildlife and plants; re- view of invertebrate wildlife for listing as en- dangered or threatened species.— Federal Register 49:21664-21675. . 1985. Endangered and threatened wildlife and plants; proposed endangered status for the Bru- neau Hot Spring Snail.—Federal Register 50: 33803-33805. Waring, G. A. 1965. Thermal springs of the United States and other countries of the world—a sum- mary.—United States Geological Survey Pro- fessional Paper 492:1-383. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 815-824 REDISCOVERY OF TULOTOMA MAGNIFICA (CONRAD) (GASTROPODA: VIVIPARIDAE) Robert Hershler, J. Malcolm Pierson, and R. Stephen Krotzer Abstract. — Recent reports have concluded that the monotypic viviparid snail genus Tulotoma Haldeman, 1840, once relatively widespread in lotic waters of the Coosa-Alabama River System, is extinct or reduced to one or two small colonies at most. The authors conducted an extensive survey for the snail during 1988-1989 and discovered five populations in shoal habitat of the Coosa River and its larger tributaries. These populations are separated by unsuitable snail habitat consisting of reaches of impounded river and probably are not in genetic contact with one another. The genus Tulotoma (type species, Pal- udina magnifica Conrad, 1834) is one of many aquatic mollusks endemic to the Coo- sa-Alabama River System. Tulotoma mag- nifica (commonly known as the tulotoma) is a particularly impressive snail, with a large (>25 mm tall) shell usually ornamented with prominent, spirally arranged knobs (Fig. 1). The genus was erected by Haldeman in 1840 and includes four species described (as Pal- udina) between 1834-1841. While early workers usually recognized at least two of these species (see Wetherby 1877), most re- cent authors (e.g., Clench 1962, Burch 1982) considered the genus to be monotypic. Note, however, that Patterson (1965) documented differences in chromosome numbers be- tween two tulotoma populations and sug- gested that several species may be involved. The historic distribution of the tulotoma, based on examination of various museum collections, is shown in Fig. 2. In the Coosa River System of Alabama, the snail ranged widely from Big Canoe Creek south to We- tumpka, just above the confluence with the Tallapoosa River. Localities included nu- merous sites on the Coosa River as well as lower reaches of several large tributaries. The snail has been recorded from only two localities in the Alabama River System: the main river in the vicinity of Claiborne (type locality for 7. magnifica), and Chilatchee Creek southwest of Selma. Records from waters apart from the above are doubtful, in some cases because the material was from Indian middens and probably was trans- ported from original localities (e.g., Black Warrior River records; Clench 1962), and in other cases because the localities are well separated from the general range of the snail and have not been verified by later collect- ing (e.g., Bridgeport, Jackson Co.; Patsaliga Creek, Crenshaw Co.). The biology of the tulotoma is virtually unknown, apart from the fact that it broods young and filter-feeds (as do other members of the Viviparidae). Early descriptions in- dicated that the tulotoma inhabited riffles and shoals, clinging tightly to the undersides of large rocks. Local abundance was com- mon: Hinkley (1904:43) noted that in the Coosa River, ““They were generally in col- onies; it was not uncommon to find 20 or 30 under a single stone a foot square or more.” Modifications of the Coosa-Alabama River System have been extensive since the description of the tulotoma in 1834. Six ma- jor dams were completed along the Coosa River in Alabama between 1914 and 1966, resulting in impoundment of much of the river and lower portions of tributaries. Pol- 816 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Shells of Tulotoma magnifica (Conrad): to left, USNM 858098, Kelley Creek, St. Clair County, Alabama (27.2 mm tall); to right, USNM 858097, Coosa River above Wetumpka, Elmore County, Alabama (22.2 mm). lution of this drainage also has been on the increase, due to introduction of waste wa- ters of various municipalities (Hurd 1974). Dredging of the Alabama River channel be- gan in 1878 and continues to the present day. Locks and dams on that river were completed in the 1960s. Decline of the tulotoma has been evident for at least 50 years. The snail no longer could be found at Claiborne by the mid- 1930s (Goodrich 1944, Clench 1962), nor has it been found elsewhere in Alabama River drainage in the past 50 years. Reduc- tion of numbers (of all prosobranch snails) in the Coosa River was obvious by 1944, as summarized by Goodrich (1944: 1-2): “‘For the waters have been backed up be- hind great dams, miles of reefs are cov- ered and formerly quiet reaches between rapids have been expanded into silt-ac- cumulating lakes. At the foot of the low- ermost dam are remains of the old We- tumpka rapids, but I have been told that ten to fifteen feet of water [actually 1-2 m depending on number of turbines op- erating; Pierson, personal observation] rush over them in the hours that the dy- namos are operating. Moreover, up- stream sections which once ran clear, Rome to Cedar Bluff, for example, are turbid with field wash, even in a dry Au- gust, and one gets specimens, if any, by feeling for them.” The last live collections of the tulotoma were those of Herbert Athearn (in Stein 1976), who located three populations in up- per Coosa River drainage between 1955-— 1963 (before completion of Logan Martin Dam), and U.S. Army Corps of Engineers (1981), which found a single live individual in Lay Reservoir below Kelley Creek. Re- cent reports concluded that the snail is now either extinct or consists of only one or two remaining colonies (Heard 1970, Davis 1974, Stein 1976). The snail currently is a candidate for possible addition to the List — Fig. 2. Historic distribution of Tulotoma magnifica. Map modified from United States Geological Survey 1:500,000 State of Alabama sheet (1970 edition). Filled circles indicate a single or two or more closely spaced localities. VOLUME 103, NUMBER 4 817 Birmingham gy C Ness \ TS 0 10 20 30 40 50 Kilometers Cet ee J Birmingham jj ) 1) 20 30 40 50 Kilometers Fig. 3. Localities surveyed (descriptions in Appendix). VOLUME 103, NUMBER 4 of Endangered and Threatened Wildlife (USDI 1989). During 1988-1989 the authors conducted a field survey for the tulotoma, principally in the Coosa basin, and discovered several live populations, which are reported below. A redescription of this remarkable snail will be published elsewhere. Methods Sites visited included those where the tu- lotoma had been found previously as well as other locales within the historic range of the species. Large tributary creeks and rivers were visited, usually at road crossings. In a number of instances, sites were considered unsuitable for the snail based on scrutiny from bridges and search was not conducted. At potentially suitable sites (e.g., ones in which riffles were present), search was con- ducted by two or three people, who exam- ined surfaces (particularly undersides) of rocks for time periods of 15 minutes to 3 hours. Boats occasionally were used to reach sampling sites and diving was necessary to gather and/or examine stones from deep water. Results Seventy-eight sites (Appendix 1, Fig. 3) were visited, of which fifty-four were inten- sively searched. The tulotoma was found living in five areas (Fig. 4), which are de- scribed below: 1) Ohatchee Creek, Calhoun County: Snails were found in a small shoal several kilometers upstream from the creek’s mouth. Substrate consisted of slabs, ledges, and sharp, angular boulders. Snails were rare (six individuals collected) on medium-sized boulders in moderate current. The species was not found at two sites upstream from the above. 2) Kelley Creek, St. Clair County: Snails were collected at various points along a two km section of the moderate-sized stream (generally <1.5 m deep and 8 m wide) be- 819 ginning at the boat landing about a hundred meters above the mouth (and upstream to SW % sec. 29, T 18S, R 3E). The creek bottom was muddy, and stones were un- common except in riffles. Snails were found throughout this area but were particularly common in a long riffle zone above the first bridge crossing (border of secs. 29, 32), where up to 20-30 snails were found on single large stones. The stream was subject to a rapid rise in water level when Logan Martin Dam was generating, during which time snails had to be collected by diving for stones. Snails were not found in the Coosa River at or near the mouth of the creek (but note the 1981 Corps of Engineers record cited above). 3) Weogufka Creek, Coosa County: Snails were found commonly in the first set of shoals (rocky bottom, no silt) above Mitch- ell Reservoir.! The tulotoma was much less abundant at two sites 3-4 km above the mouth of creek where fewer than 20 indi- viduals were found after almost 2 hours of searching. At the latter sites the stream was fairly small (3-5 m across, <1 m deep) and predominantly riffle, with the bottom densely littered with small rounded stones covered with green algae. Snails were absent at one site upstream from the above. 4) Hatchet Creek, Coosa County: Snails were found commonly (50-60 individuals on a single large rock) in fast shoals at the lower end of the creek E-NE of Kelly’s Crossroads.' Snails also were found in the creek ca. 3 km upstream from the conflu- ence with Swamp Creek (ca. 6 km upstream from Mitchell Lake). At this site the creek was about 7 m across and had both short riffles and large pooled areas. Snails were rare: only 6 individuals were collected in 30 minutes. Snails were absent at a site up- stream from the above. 'R. G. Bowker, United States Fish and Wildlife Ser- vice (Jackson Field Office) reported finding the tulot- oma at the beginning of reservoir influence on both Weogufka and Hatchet Creeks in October, 1989 (letter to RH, 10-23-89). Birmingham Montgomery S/ 0 10 20 30 40 50 Kilometers L 1 = St Nl ! Fig. 4. Current distribution of Tulotoma magnifica. VOLUME 103, NUMBER 4 821 Fig. 5. Photographs of tulotoma site on Hatchet Creek, Coosa County, Alabama: a, snail-bearing shoals; b, underside of large stone showing clustered snails. 5) Coosa River, between Jordan Dam and Wetumpka, Elmore County: The river is free-flowing for at least the upper portion of this area and contains a series of im- pressive shoals. Snails were collected from two such areas separated by 2-3 km. In both sites the river was constricted by a complex of small islands through which the water rushes. The bottom was gravel-sand and was littered with stones of various sizes. Snails were very common on rocks in narrow riffle areas (up to 40-50 specimens/large stone) and less numerous in pooled areas. During normal daily generation from one turbine (4400 cfs for 2.25 hours) the water level would rise a maximum of | m at Moccasin Shoal. Maximum turbine discharge at full gate (21,000 cfs) would raise the river level about 2 m at this location (a very rare event). Snails were absent both from a shoal about 822 1 km upstream from the above, and down- stream at Wetumpka. Associated molluscan fauna at the above sites included diverse pleurocerid snails and unionid clams, and Corbicula. Discussion A synthesis of results of this survey and available literature allows a general inter- pretation of the habitat requirements of the tulotoma. The snail is restricted to cool, well- oxygenated, clean water in free-flowing riv- er and lower portions (ca. lowermost 4—5 km) of large tributary streams (Fig. 5a). The tulotoma never has been recorded from up- per reaches of tributaries: perhaps such hab- itat is unsuitable owing to its small size and/ or adverse physiochemical conditions (e.g., softer water compared to downstream). The animal clings to undersides of submerged large stones (which usually rest on hard bot- toms), with individuals densely clustered rather than uniformly distributed (Fig. 5b). Densities are highest in riffles or shoals, al- though snails also occur in pooled areas. Although the tulotoma can tolerate diurnal variation in hydrologic variables (as seen in the Kelley Creek population), continued persistence of the snail in waters where such variation is extreme (as in the river im- mediately below a dam) appears unlikely. Perusal of the historic and current distri- bution maps indicates that a drastic reduc- tion of the tulotoma’s range has occurred over the past 150 years (compare Figs. 2 and 4). The snail now is apparently extinct in the Alabama River system and restricted to several km of free-flowing Coosa River below Jordan Dam and short sections of four large creeks tributary to the river. These populations are isolated by reaches of un- suitable tulotoma habitat (impounded riv- er) that are quite long, with the exception of Weogufka and Hatchet Creeks (which discharge into Mitchell Lake at points sep- arated only by a few km), and probably are not in genetic contact with one another. The PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON largest population appears to be that of the Coosa River below Jordan Dam where some tens of thousands of individuals occur. Much of the range reduction can be attributed to unequivocal habitat destruction associated with impoundment of a large portion of the Coosa River in Alabama. Acknowledgments Hershler’s fieldwork was funded by a con- tract from the United States Fish and Wild- life Service (Cooperative Agreement No. 14- 16-004-87-931). Assistance in the field was provided by H. Athearn, A. Gerberich, P. Greenhall, J. Lochamy, C. Norquist, J. Stewart, and E. Tyberghein. Victor Krantz prepared shell photographs, and M. Ryan and K. Flamer drafted the map. Literature Cited Burch, J. B. 1982. North American freshwater snails. Identification keys, generic synonymy, supple- mental notes, glossary, references, index. — Walkerana 4:217-365. Clench, W. J. 1962. A catalogue of the Viviparidae of North America with notes on the distribution of Viviparus georgianus Lea.—Occasional Pa- pers on Mollusks, Museum of Comparative Zo- ology 2:261-287. Davis, G. M. 1974. Report on the rare and endan- gered status of a selected number of freshwater gastropods from southeastern U.S.A.—Report to U.S. Department of Interior Fish and Wildlife Service, Contract 14-16-0008-766. Goodrich, C. 1944. Certain operculates of the Coosa River. — Nautilus 58:1—10. Heard, W. H. 1970. 3. Eastern freshwater mollusks (II). The South Atlantic and Gulf drainages. — Malacologia 10:23-27. Hinkley, A.A. 1904. List of Alabama shells collected in October and November, 1903.— Nautilus 18: 37-45. Hurd, J. 1974. Systematics and zoogeography of the unionacean mollusks of the Coosa River drain- age of Alabama, Georgia, and Tennessee. — Un- published Ph.D. dissertation, Auburn Univer- sity. 240 pp. Patterson, C. M. 1965. The chromosomes of Tu/ot- oma angulata (Streptoneura: Viviparidae).— Malacologia 2:259-265. VOLUME 103, NUMBER 4 Stein, C. B. 1976. Gastropods. Pp. 1-41 in H. Bos- chung, ed., Endangered and threatened plants and animals of Alabama. Bulletin Alabama Mu- seum of Natural History 2. United States Army Corps of Engineers. 1981. Mont- gomery to Gadsden Coosa River channel, Al- abama. Design memorandum 1. Volume 5. Ap- pendix V—Environmental Systems (EIS Appendix C—Part 2). United States Department of the Interior. 1989. En- dangered and threatened wildlife and plants; an- imal notice of review. — Federal Register 54:554— 579. Wetherby, A. G. 1877. Review of the genus Tulot- oma, with remarks on the geographical distri- bution of the North American Viviparidae— Quarterly Journal of Conchology 11:207-215. (RH) Department of Invertebrate Zool- ogy, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (JMP, RSK) G.S.C. #8, Alabama Power Company, Post Office Box 2641, Bir- mingham, Alabama 35291. Appendix Localities surveyed. Data include topographic co- ordinates and date of visitation. Positive localities are indicated by an asterisk. ALABAMA. Autauga County: Shoal Creek, State HW 143 (crossing), T 20N, R 16E, SW % sec. 27, 6-2-88. Calhoun County: Coosa River below Neely Henry Dam, 3.2 km SW of Ohatchee, T 14S, R 6E, SE % sec. 31, 10-11-88.—Ohatchee Creek: N of State HW 62, T 14S, R 6E, SW % sec. 27, 9-20-89; State HW 62, T 14S, R 6E, SW % sec. 27, 6-6-88; *1.9 km S of Ohatch- ee, T 14S, R 6E, SE % sec. 33, 10-11-88.—Cane Creek: State HW 77, T 15S, R 6E, SW % sec. 18, 9-20-89; County Road 73, T 15S, R 5E, SW % sec. 13, 9-20- 89. Cherokee County: Little River: State HW 35, T 7S R 10E, SE % sec. 30, 6-4-88; State HW 273, 8.8 km NW of Cedar Bluff, T 9S, R 9E, NE % sec. 3, 10-13- 88.—Terrapin Creek: 0.6 km S of Ellisville, T 11S, R 9E, SW % sec. 20, 10-13-88; NW of Providence Church, T 11S, R 9E, SE % sec. 34, 9-20-89. Chilton County: Coosa River, Lay Dam tailrace, T 23N, R 15SE, NW % sec. 24, 10-11-88.—Yellowleaf Creek: Crossing S of Page Creek confluence, T 23N, R 15E, SW ‘4 sec. 22, 9-13-88; Crossing N of Lime Springs Church, T 23N, R 15E, NE % sec. 20, 9-13-88: 1.6 km 823 SW of Lay Dam, T 23N, R 15E, SW 4 sec. 25, 10-5- 88.— Walnut Creek: County Road 325 1522. ReISE: SE 4 sec. 14, 9-18-89; 10.9 km E-NE of Clanton, T 22N, R 16E, NE sec. 19, 10-5-88.—Chestnut Creek: State HW 3 at Verbena, T 21N, R 15E, SE 4 sec. 36, 6-2-88; 8.0 km E-NE of Verbena, T 21N, R 16E, NE Ys sec. 35, 10-6-88. Coosa County: Paint Creek, 3.2 km S-SE of Marble Valley, T 24N, R 16E, SE % sec. 35, 10-11-88.—Weo- gufka Creek: NW of Moriah, T 23N, R 17E, NE “sec. 34, 6-3-88, 10-3-88; *Horse Stomp Campground, T 22N, R 17E, SE % sec. 6, 9-18-89: *Crossing SW of above, T 22N, R 17E, NW % sec. 7, 9-18-89; *First shoals above Mitchell Reservoir, T 22N, R 16E, SE 4 sec. 13, 10-6-88.— Hatchet Creek, 9-18-89: US 231, T 23N, R 19E, SE % sec. 30, 9-18-89; *SE of Lyle, T 22N R 18E, NW % sec. 19, 9-18-89: *3.7 km N-NE of Kelly’s Crossroads, T 22N, R 17E, NW ‘4 sec. 25, 10- 7-88.—Swamp Creek: State HW 22, T 22N, R 18E, SE % sec. 30, 9-18-89; County Road 29, 2.4 km N of Kelly’s Crossroads, T 22N, R 17E, SW % sec. 25, 10- 27-88. Dallas County: Soapstone Creek, US 80, T 16N, R 12E, NW % sec. 31, 6-3-88.—Cedar Creek, State HW 41, T 14N, R 10E, NE % sec. 14, 6-3-88.—Oak Creek. State HW 41, T 13N, R 10E, SW 4 sec. 8, 6-3-88. Elmore County: Coosa River: Just below Jordan Dam, T 19N, R 18E, NE % sec. 22, 6-2-88: Shoal ca. 1 km below dam, T 19N, R 18E, SE 4 sec. 22, 9-14- 88; *Moccasin Shoal, ca. 5 km upstream from We- tumpka, T 18N, R 18E, NE % sec. 2, 9-18-89: *Ca. 2.5 km upstream from Wetumpka, T 18N, R 19E, NE “4 sec. 7, 9-14-88; Vicinity of Wetumpka boat ramp, T 18N, R 19E, NE % sec. 24, 9-14-88. Monroe County: Tallahatchee Creek, State HW 41. T 10N, R 7E, NE % sec. 20, 6-3-88.—Beaver Creek. State HW 41, T 9N, R 6E, NE % sec. 24, 6-3-88.—Big Flat Creek, State HW 41, T 7N, R 6E, NE 4 sec. 1. 6-3-88.—Alabama River, 9-14-88: Base of limestone bluff on west side of river, ca. 3.0 km below Claiborne Lock and Dam, T 7N, R 5E, SE 4 sec. 11: ca. 1.6 km below Claiborne-Murphy Bridge, T 7N, R SE, NE 4 sec. 26.—Limestone Creek: State HW 41, T 7N, R 7E, NE '% sec. 22, 6-3-88; from mouth to ca. 0.5 km up- stream, T 7N, R 6E, SE % sec. 19, 9-14-88. St. Clair County: Big Canoe Creek, US 231, T 14S, R 4E, SE % sec. 6, 6-4-88.— Beaver Creek, US 231, T 14S, R 3E, NE ‘4 sec. 36, 6-4-88.—Shoal Creek, US 231, T 15S, R 3E, SE % sec. 1, 6-4-88.—Coosa River, ca. 0.8 km below Logan Martin Dam, T 18S, R 3E, SW ‘4 sec. 33, 9-12-88.— Kelley Creek: *E-NE of Logan Martin Dam substation (upstream from crossing), T 18S, R 3E, SW 4 sec. 29, 9-21-88, 9-13-89; *Kelley Creek Landing, T 19S, R 3E, NE '% sec. 6, 9-12-89; Mouth, T 19S, R 3E, NW ‘4 sec. 5, 9-12-89. Shelby County: Stream tributary to Kelley Creek, W-SW of Macedonia Church, T 18S, R 2E, SE ' sec. 824 23, 6-1-88.—Spring Creek N of Vincent, T 19S, R 2E, SW 4 sec. 11, 6-1-88.—Morgan Creek NW of Klein, T 20S, R 2E, SE % sec. 17, 6-1-88.— Yellowleaf Creek: State HW 25, T 20S, R 2E, SE % sec. 29, 6-1-88; 6.1 km S-SW of Harpersville, T 20S, R 2E, NE % sec. 18, 10-4-88.—Fourmile Creek, County Road 441, T 20S, R 2E, SE % sec. 30, 6-1-88.— Waxahatchee Creek: 4.8 km S-SW of Shelby, T 24N, R 15E, SE % sec. 7, 10- 11-88; State HW 145 S of Shelby, T 24N, R 15E, SE % sec. 20, 6-1-88. Talladega County: Blue Eye Creek, State HW 77, T 16S, R 5E, SW % sec. 22, 6-6-88.—Eastaboga Creek, in Old Eastaboga, T 16S, R 6E, SW % sec. 33, 6-6- 88.—Choccolocco Creek: County Road 399, T 17S, R 6E, NW sec. 15, 9-19-89; 2.9 km S of Old Eastaboga (near Brick Store Church), T 17S, R 6E, SW % sec. 9, 10-27-88; County Road 5, T 17S, R 6E, SW % sec. 17, 9-19-89; Lower end of Jackson Shoals, T 17S, R 5E, SW % sec. 14, 6-1,4-88, 9-21-88.—Cheaha Creek, County Road 5, T 17S, R 6E, SW % sec. 20, 10-27- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 88, 9-19-89.—Coosa River, east bank below Logan Martin Dam, T 18S, R 3E, NW % sec. 33, 9-28-88.— Talladega Creek: 5.6 km N-NE of Childersburg, T 20S, R 3E, NE % sec. 3, 9-28-88; At Kymulga, T 19S, R 3E, center of sec. 35, 9-19-89.—Tallasseehatchee Creek: County Road 105, T 20S, R 4E, SE % sec. 30, 9-19- 89: N-NW of Friendship Church, T 20S, R 3E, SE 4 sec. 14, 9-19-89; 3.8 km E-NE of Childersburg, T 20S, R 3E, sec. 22, 9-28-88.— Kahatchee Creek, 6.6 km SW of Childersburg, T 21S, R 2E, NW % sec. 2, 10-4-88. Tallapoosa County: Tallapoosa River, 10.7 km S-SW of Daviston, T 23N, R 24E, SE % sec. 19, 9-27- 88. Wilcox County: Pine Barren Creek, State HW 41, T 13N, R 9E, SW % sec. 28, 6-3-88.—Pursley Creek, State HW 41, T 11N, R 7E, NE % sec. 2, 6-3-88.— Gravel Creek, State HW 41, T 11N, R 7E, NW % sec. 22, 6-3-88.—Chilatchee Creek, Alberta, T 15N, R 7E, SE % sec. 30, 9-15-88. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 825-838 NEW SPECIES AND NEW RECORDS OF SCALED POLYCHAETES (POLYCHAETA: POLYNOIDAE) FROM THE AXIAL SEAMOUNT CALDERA OF THE JUAN DE FUCA RIDGE IN THE NORTHEAST PACIFIC AND THE EAST PACIFIC OCEAN OFF NORTHERN CALIFORNIA Marian H. Pettibone Abstract.—A new species of Harmothoe globosa from the Axial Seamount Caldera on the Juan de Fuca Ridge is described. Polynoids from the Gorda Ridge off northern California include Harmothoe gordae, new species, in Harmothoinae, from Wood Island; Parabathynoe brisinga, new genus, new species, in Arctonoinae, commensal with a brisingid asteroid; Lepidonotopo- dium piscesae, new record, in Lepidonotopodinae; Levensteiniella kincaidi, new record, and L. intermedia, new species, in Macellicephalinae; and Branchinoto- gluma grasslei and Opisthotrochopodus tunnicliffeae, new records, in Bran- chinotogluminae. A new species of Harmothoe (Subfamily Harmothoinae) has been discovered from fresh basalts up to 100 meters from the hy- drothermal vents in the Axial Seamount Caldera of the Juan de Fuca Ridge, collected by DSRV Alvin. An additional seven species of polynoids were collected on the Gorda Ridge off northern California by DSRV A/- vin and in the northern Escanaba Trough of the Gorda Ridge by DSRV Sea Cliff: The seven species from the Gorda Ridge include a new species of Harmothoe; a new genus and new species (Subfamily Arctoninae) commensal with a brisingid asteroid; new records of Lepidonotopodium piscesae (Sub- family Lepidonotopodiinae); a new species of Levensteiniella and new records of L. kin- caidi (Subfamily Macellicephalinae); and new records of Branchinotogluma grasslei and Opisthotrochopodus tunnicliffeae (Sub- family Branchinotogluminae). The specimens are deposited in the Na- tional Museum of Natural History, Smith- sonian Institution (USNM) and the Scripps Institution of Oceanography (SIO). Subfamily Harmothoinae Willey, 1902 Genus Harmothoe Kinberg, 1856 Harmothoe globosa, new species Figs. 1, 2 Material examined.—Northeast Pacific, Axial Seamount Caldera, Juan de Fuca Ridge, 45°58’N, 130°03’W, 1570 m, DSRV Alvin dive 2087, 15 Aug 1988, from fresh basalts, up to 100 m from hydrothermal vents, holotype (USNM 123368). Description. —Length of holotype 32 mm, width with setae 12 mm, segments 41, last one minute. Body flattened ventrally, arched dorsally, tapering slightly anteriorly and more so posteriorly, with long parapodia nearly as long as body width. Middorsum brownish with 2 light transverse ciliated bands per segment between bases of elytro- phores and dorsal tubercles (Fig. 1A, C). Elytra 15 pairs, on segments 2, 4, 5, 7, al- ternate segments to 23, 26, 29, and 32. El- ytra round, subreniform to oval, large, over- lapping, covering dorsum, rather thick stiff, with most of surface covered with conical 826 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. antenna, dorsal and ventral tentacular cirri missing; B, Same, ventral view, right side and palps only partially shown; C, Dorsal view of right half of segments 5 and 6, style of dorsal cirrus on segment 6 missing; D, Right 1st elytron from segment 2, with detail of microtubercles and papillae; E, Right 2nd elytron from segment 4; F, right middle elytron, with detail of macrotubercle, microtubercles and papillae. Scales = 1.0 mm for A-C; 1.0 mm for D-F. microtubercles, bare region near antero- medial side; additional baloonlike, bulbous macrotubercles with nipple-like tips emerg- ing from wide bases near posterior and lat- eral borders plus smaller oval macrotuber- cles on middle part, and some scattered delicate papillae on border and surface (Fig. 1 D-F). Elytrophores large, bulbous, reddish glandular area on anterior side (Figs. 1A, C, 2A). Dorsal cirri on segments without ely- tra; cirrophores long, cylindrical, on pos- terior sides of notopodia, wider basally and enclosing anterior and posterior reddish glandular areas; styles long, tapering distal- ly, extending beyond setae, with delicate clavate papillae; dorsal tubercles nodular, enclosing reddish glandular areas (Figs. 1C, 2B) smaller from segment 24, indistinct on 9 posterior cirrigerous segments. Harmothoe globosa, holotype, USNM 123368: A, Dorsal view of anterior end, styles of median Prostomium bilobed, wider than long, with distinct cephalic peaks including del- icate tips; median antenna with large cylin- drical ceratophore in anterior notch, style missing (probably long); ceratophores of lat- eral antennae small, inserted ventrally and converging midventrally, with short, su- bulate, papillate styles; eyes large, anterior pair of anterolateral just anterior to widest part of prostomium, posterior pair postero- dorsal; palps stout, long, tapered, minutely papillate (Fig. 1A, B). First segment not dis- tinct dorsally; tentaculophores lateral to prostomium, each with small acicular lobe, 2 setae on inner side, and pair of dorsal and ventral tentacular cirri with styles missing (probably long, similar to buccal cirri); dis- tinct ventrally forming anterior and lateral lips of ventral mouth, with facial ridge but VOLUME 103, NUMBER 4 B — —————e Fig. 2. Harmothoe globosa, holotype USNM 123368: A, Right elytrigerous parapodium, anterior view, acicula dotted, with detail of neuropodial supraacicular process; B, Right cirrigerous parapodium, posterior view; C, Long and short notosetae; D, Lower and upper subacicular and supraacicular neurosetae, with detail of tips. Scales = 1.0 mm for A, B; 0.1 mm for C, D. without distinct facial tubercle (Fig. 1A, B). Second or buccal segment with small nuchal lobe, first pair of large elytrophores, well- developed, biramous parapodia, and long, papillate buccal cirri, and forming posterior lip of ventral mouth (Fig. 1A, B). Pharynx not extended and not examined; dark color of pharynx visible through delicate integu- ment. Biramous parapodia with both rami well developed, with acicular processes sharply pointed and tips of acicula projecting; no- topodium shorter than neuropodium, rounded, with acicular process on lower side; neuropodium with conical presetal acicular lobe, long digitiform supraacicular process and shorter rounded postsetal lobe with red- dish glandular area on distal part (Fig. 2A, B). Notosetae very numerous, forming ra- diating bundle, stouter than neurosetae, of 3 lengths, shorter, curved and longer nearly Straight, with numerous rows and tapered blunt bare tips, some of longest ones show- ing faint longitudinal groove (Fig. 2C). Neu- rosetae very numerous, forming fan-shaped bundle, with numerous spinous rows; su- praacicular neurosetae more slender than subacicular ones; mostly showing delicate secondary tooth or indication of one; lower ones with entire tips (Fig. 2D). Ventral cirri short, tapering, with small clavate papillae (Fig. 2A, B). Pygidium small, with dorsal anus medial to last pair of parapodia, with pair of anal cirri (styles missing). Nephridial papillae beginning on segment 6, small, oval, on lat- eral side of rounded, inflated area. Etymology.—Named globosa, referring to the globular macrotubercles on the elytra. Remarks. —The unusual baloonlike mac- 828 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Comparison of Harmothoe globosa, new species and H. macnabi Pettibone. Harmothoe globosa Harmothoe macnabi Elytral macrotubercles face (Fig. 1D-F) Prostomium large cephalic peaks; 2 pairs of large eyes (Fig. 1A) Tentaculophore with 2 setae (Fig. 1A) Neuropodial presetal acic- ular lobe process (Fig. 2A) balloonlike, near border and on sur- with well-developed supraacicular extensions on posterior border (Petti- bone 1985c, Fig. 6C) small cephalic peaks; without eyes (Pettibone 1985c, Fig. 6A) with 6-8 setae (Pettibone 1985c, Fig. 6A, B) without supraacicular process (Petti- bone 1985c, Fig. 7B) rotubercles found on the elytra of Har- mothoe globosa suggest a greater develop- ment of the macrotubercles than those found on H. macnabi Pettibone (1985c:749, figs. 6, 7), described from the Galapagos Rift in the East Central Pacific. The two species differ as shown in Table 1. Harmothoe gordae, new species Fig. 3 Material. —East Pacific Ocean off north- ern California, Gorda Ridge, Alvin dive 2034, 4 Jun 1988, 41°00’N, 127°29'W, 3362 m, Wood Island, holotype (USNM 123367). Description.—Length of holotype (male with sperm) 17 mm, width with setae 9 mm, segments 36, last one minute. Body flat- tened dorsoventrally, tapering slightly an- teriorly and posteriorly, with long parapo- dia nearly as long as body width, without color. Elytrophores 15 pairs, on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, and 32, large, bulbous (Fig. 3A, C). Elytra (all missing except left 14th on segment 29) large, oval, soft, opaque, nearly covered with conical microtubercles, low, oval on ante- rior part, bare region near anteromedial side; scattered cylindrical papillae on surface and posterior and lateral borders (Fig. 3B). Dor- sal cirri on segments without elytra, with long cylindrical cirrophores on posterior sides of notopodia, wider basally; styles long, extending beyond setae, with slender tips and scattered short papillae; dorsal tuber- cles prominent, nodular (Fig. 3D). Prostomium bilobed, with prominent ce- phalic peaks, without eyes; median antenna with large bulbous ceratophore in anterior notch, with style about twice as long as pro- stomium, with scattered short papillae; cer- atophores of lateral antennae rather large, inserted ventrally, not meeting midventral- ly, with short, subulate papillate styles; palps stout, tapered, longer than median antenna (Fig. 3A). First segment not distinct dor- sally; tentaculophores lateral to prostomi- um, each with small acicular lobe, 3 setae on inner side and pair of dorsal and ventral tentacular cirri similar to median antenna; ventral surface with facial ridge but without distinct facial tubercle (Fig. 3A). Second or buccal segment with first pair of large elytro- phores, well-developed biramous parapo- dia, and long papillate ventral buccal cirri (Fig. 3A). Pharynx not extended and not examined. Biramous parapodia with both rami well developed, with projecting acicular lobes and tips of acicula projecting; notopodium shorter than neuropodium, rounded, with projecting acicular process on lower side; neuropodium with conical presetal acicular lobe, with slight indication of rounded su- praacicular process; postsetal lobe shorter, rounded (Fig. 3C, D). Notosetae numerous, forming radiating bundle, stouter than neu- rosetae, short and slightly curved to long VOLUME 103, NUMBER 4 829 Fig. 3. Harmothoe gordae, holotype, USNM 123367: A, Dorsal view of anterior end, right dorsal and ventral tentacular cirri missing; B, Left 14th elytron from segment 29, with detail of microtubercles and papillae; C, Right elytrigerous parapodium, anterior view, acicula dotted; D, Right cirrigerous parapodium, posterior view; E, Long and short notosetae; F, Lower, middle and upper neurosetae. Scales = 1.0 mm for A; 1.0 mm for B; 0.5 mm for C, D; 0.1 mm for E, F. and straight, with spinous rows and rather long, bare, tapering tips (Fig. 3E). Neuro- setae numerous, forming fan-shaped bun- dle; upper ones more slender, with longer spinous regions, lower ones stouter, with shorter spinous regions; all with bifid tips, with long slender secondary tooth (Fig. 3F). Ventral cirri short, tapering (Fig. 3C, D). Pygidium small, rectangular, with dorsal anus medial to last pair of small parapodia; anal cirri missing. Nephridial papillae be- ginning on segment 6, small, on lateral side, projecting dorsally between parapodia. Etymology.—Named for the collecting site, the Gorda Ridge. Remarks. —Harmothoe gordae, collected on Wood Island on the Gorda Ridge, is close to H. vagabunda Pettibone (1985a:146, fig. 830 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Comparison of Harmothoe gordae, new species and H. vagabunda Pettibone. Harmothoe gordae Harmothoe vagabunda Prostomium (Fig. 3A) Neuropodial presetal acic- ular lobe Neurosetae process (Fig. 3C) (Fig. 3F) prominent distinct cephalic peaks with small rounded supraacicular with long slender secondary tooth very small cephalic peaks (Pettibone 1985a, Fig. 6A) without supraacicular process (Petti- bone 1985a, Fig. 6B) with shorter secondary tooth (Petti- bone 1985a, Fig. 6E) 6), described from the North Atlantic in the Tongue of the Ocean, Bahamas, and off St. Croix, Virgin Islands, associated with wood panels. The two species differ as indicated in Table 2. Subfamily Arctonoinae Hanley, 1989 Parabathynoe, new genus Type species. —Parabathynoe brisinga, new species. Diagnosis.—Body short, up to 46 seg- ments. Elytra and prominent elytrophores 18 pairs, on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, 32, 35, 40, and 43. Elytra thickly covered with conical micro- tubercles and papillae. Dorsal cirri on seg- ments lacking elytra, with cirrophores and distal styles; dorsal tubercles in line with elytrophores. Prostomium bilobed, round- ed, with 2 pairs of large eyes, 2 long palps, and 3 antennae: median antenna with cer- atophore and long style inserted in anterior notch; lateral antennae inserted subtermi- nally, ventral to median antenna, with cer- atophores and short styles only slightly set off from prostomium. First or tentacular segment not visible dorsally; tentaculo- phores lateral to prostomium, achaetous, with 2 pairs of tentacular cirri; ventral sur- face with bulbous upper lip. Second or buc- cal segment with bulbous elytrophores of first pair of elytra, small subbiramous par- apodia, ventral buccal cirri, and bulbous lat- eral and wrinkled lower lips. Parapodia sub- biramous; notopodia with small conical acicular lobe and bundle of short acicular notosetae; neuropodia larger, with rounded presetal acicular lobe; postsetal lobe hook- shaped, of two parts: upper part short, rounded; lower part curved dorsally, with thick bushy bundle of long papillae on outer curved border; neurosetae numerous, long, nearly smooth, with tapered, mostly bifid tips; ventral surface of neuropodia thickly papillate. Ventral cirri lacking, except on segment 2. Extended pharynx with 9 pairs of papillae and 2 pairs of chitinous jaws. Commensal with brisingid asteroids. Etymology. —From the Greek para, plus the genus Bathynoe Ditlevsen, in the sub- family Arctonoinae; gender feminine. Remarks. —In comparison with the gen- era included in the subfamily Arctonoinae by Hanley (1989), Parabathynoe is closest to Bathynoe Ditlevsen, 1917. Both are rel- atively short-bodied (less than 50 seg- ments), ventral cirri are present only on seg- ment 2, and lower parts of neuropodial postsetal lobes are curved dorsally and pa- pillated on the curved border. Both are commensal with asteroids. Bathynoe differs from Parbathynoe in a number of features. In the former, the body has prominent middorsal tubercles, lacking in the latter; elytra have a different arrange- ment and have nodular tubercles and glob- ular micropapillae on the surface, instead of microtubercles and cylindrical papillae; prostomium has short, stumpy palps and has no eyes, instead of long palps and two pairs of eyes; notosetae are absent on sub- biramous parapodia, instead of present; and neurosetae are few, short, stout, and strong- ly hooked, instead of numerous, long, and slender, with minute bifid tips. VOLUME 103, NUMBER 4 Parabathynoe brisinga, new species Figs. 4, 5 Material. —East Pacific Ocean off north- erm California, Gorda Ridge, Alvin dive 2033, 41°00'N, 127°29’W, 3 Jun 1988, 3356 m, commensal with brisingid asteroid, ho- lotype (USNM 123371), paratype (USNM 123372). Description. —Length of holotype (female with large yolky eggs in body cavity) 18 mm, width 6 mm with setae, segments 47, last one minute. Paratype incomplete posteri- orly, with 32 segments, 11 mm long and 4.5 mm wide. Body flattened ventrally, slightly arched dorsally, tapering slightly anteriorly, and more so posteriorly. Ventral side of body with 2 pairs of large, rounded papillae per segment and numerous small papillae on ventral side of parapodia (Fig. 4E). Elytra 18 pairs, large, covering dorsum, on seg- ments 2, 4, 5, 7, alternate segments to 23, 26, 29, 32, 35, 40, 43. Elytra oval to sub- reniform, thickly covered with conical mi- crotubercles and papillae; thicker, slightly raised area across posterior fifth or sixth of surface (Fig. 5 E—G); elytrophores large and prominent (Figs. 4A, C, 5A). Dorsal cirri on segments lacking elytra, with short cy- lindrical cirrophores and styles extending beyond neuropodia, styles with proximal 7 cylindrical and slightly bulbous, distal ' fil- amentous, with long papillae on lower side of cylindrical part; dorsal tubercles nodular, in line with elytrophores (Figs. 4A, B, D, 5B). Prostomium oval, bilobed, wider than long, with 2 pairs of eyes on lateral sides, anterior pair very large, twice as large as posterior pair; median antenna with bul- bous ceratophore in anterior notch, style about twice as long as prostomium, with filamentous tip; lateral antennae with cer- atophores indistinctly set off from prosto- mium and inserted ventral to median an- tenna; short styles with clavate enlargements and filamentous tips; palps stout, cylindri- cal, with filamentous tips; tentaculophores 831 lateral to prostomium, with small acicular lobe on inner side, achaetous; dorsal and ventral tentacular cirri bulbous subdistally with filamentous tips; facial ridge on bul- bous anterior lip of ventral mouth (Fig. 4A, B). Second or buccal segment without nuchal lobe, with first pair of large elytrophores, subbiramous parapodia much smaller than following parapodia, ventral buccal cirri, similar to tentacular cirri, and bulbous lat- eral and posterior lips enclosing ventral mouth (Fig. 4A-C). Parapodia from segment 3 on larger, sub- biramous, with small conical notopodial acicular lobe on anterodorsal side of larger neuropodium with longer, rounded presetal acicular lobe with low rounded upper part; shorter, hook-shaped postsetal lobe con- sisting of short rounded upper half and lon- ger lower half curving dorsally, with bushy bundle of long papillae on outer curved bor- der of lower part (Figs. 4B, 5A, B). Without ventral cirri. Notosetae extending to tip of neuropodium, in small bundle, short, acic- ular, slightly stouter than neurosetae, nearly smooth, with faint close-set spinous rows and entire tips (Fig. 5C). Neurosetae very numerous, forming fan-shaped bundle, long, enlarged subdistally, tapering to slightly hooked tips, mostly with very small sec- ondary tooth, others entire; upper few (5 or so) with split tips (Fig. 5D). Pharynx (cut open) with 9 pairs of marginal papillae and 2 pairs of amber-colored jaws. Pygidium short, rectancular, between posterior, mi- nute parapodia, with anal cirri (missing). Etymology. —The species is named for its commensal relationship with brisingid as- teroids. Subfamily Lepidonotopodiinae Pettibone, 1983 Genus Lepidonotopodium Pettibone, 1983 Lepidonotopodium piscesae Pettibone, 1988 Lepidonotopodium piscesae Pettibone 1988: 193, figs. 1-4. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Parabathynoe brisinga, A-D, holotype USNM 123371: A, Dorsal view of anterior end, left ventral tentacular cirrus missing; B, Same, ventral view; C, Left elytrigerous parapodium from segment 2, anterior view, acicula dotted; D, Left cirrigerous parapodium from segment 3, anterior view, acicula dotted; E, Ventral view of left side of two middle segments lateral to midventral neural groove. Scales = 1.0 mm for A, B, E; 0.5 mm for C, D. Lepidonotopodium pettibonae Detinova, 1988:858, fig. la—h. Material. —East Pacific Ocean off north- ern California, Gorda Ridge, in northern Escanaba Trough, Sea Cliff dive 764, 3 Sep 1988, 41°00’N, 127°30'W, 3500 m, 11 spec- imens (USNM 123369, SIO). Remarks. — The specimens from the Gor- da Ridge agree with the specimens from the hydrothermal vents of the Northeast Pacific Explorer and Juan de Fuca Ridges, de- scribed earlier as Lepidonotopodium pisces- ae by Pettibone (1988, 29 April) and as L. pettibonae by Detinova (1988, June). Un- fortunately, the former name has priority. The eleven specimens from the Gorda Ridge were collected in the northern Esca- naba Trough in a heavily sedimented area with a large clump of vestimentiferans, in- cluding Rigia spp., and long filamentous bacteria. The elytra and setae of the poly- noids were covered with foreign material, including sediment and bacteria. The larger specimens measured 20-22 mm in length, 10-11 mm in width, with 25-26 segments. Distribution. —Hydrothermal vents of Northeast Pacific Explorer and Juan de Fuca Ridges, in 1500-2208 m and northern Es- canaba Trough of Gorda Ridge, in 3500 m. Subfamily Macellicephalinae Hartmann-Schroder, 1971; emended Pettibone, 1976 Genus Levensteiniella Pettibone, 1985 The genus includes L. kincaidi, described from the Galapagos Rift and the East Pacific Rise at 21°N by Pettibone (1985c), with ad- ditional records from the Northeast Pacific Explorer and Juan de Fuca Ridges by Pet- tibone (1988), and L. raisae, described from the Western Pacific in the Mariana Back- VOLUME 103, NUMBER 4 | 833 Fig. 5. Parabathynoe brisinga, holotype, USNM 123371: A, Right elytrigerous parapodium from segment 15, anterior view, acicula dotted; B, Right cirrigerous parapodium from segment 16, posterior view (few eggs in body cavity dotted); C, Notosetae, with detail of faint spinous rows; D, Middle and upper neurosetae, with detail of tips; E, Left 1st elytron from segment 2, with detail of microtubercles and papillae; F, Left 2nd elytron from segment 4, with detail of microtubercles and papillae; G, Left middle elytron, with detail of microtubercles and papillae. Scales = 0.5 mm for A, B; 0.1 mm for C, D. Arc Basin by Pettibone (1989b). From the Gorda Ridge, additional records of L. kin- caidi showing some varietal differences are added and a new species is described below. Levensteiniella kincaidi Pettibone, 1985 Fig. 6 Levensteiniella kincaidi Pettibone, 1985c: 741, figs. 1-3; 1988:199, fig. 5; 1989b: 159. Material. —East Pacific Ocean off north- ern California, Gorda Ridge, Alvin dive 2036, 6 Jun 1988, 41°00’N, 127°29'W, 3240 m, 1 specimen (USNM 123366). Gorda Ridge in northern Escanaba Trough, Sea Cliff dive 764, 3 Sep 1988, 41°00’N, 127°30’W, 3500 m, 5 specimens (USNM 123365; SIO). Description of specimen from Alvin dive 2036.—Length 19 mm, width with setae 10 mm, with 26 segments and 11 pairs of el- ytra. Some elytra with papillae on posterior and lateral borders enlarged basally; papil- lae on border and surface variable in shape (Fig. 6G-I). Prostomium, tentacular and first buccal segments as described earlier (Fig. 6A). Dorsal cirri and dorsal tubercles, bi- ramous parapodia, notosetae and neurose- tae, and ventral cirri, as described earlier (Fig. 6C—F). Body without 2 pairs of long ventral papillae on segments 11 and 12 but with small rounded paired papillae near basal sides of parapodia on segments | 1-19 834 (Fig. 6B); as reported earlier, long ventral papillae found on only some adults (Petti- bone, 1985c, fig. 1D; 1988). Remarks.—The setae of the specimens from the Escanaba Trough, collected on heavily sedimented fresh basalts, were cov- ered with numerous round and long fila- mentous bacteria; one was covered with stalked peritrich protozoa. The elytra of the stem form of L. kincaidi, described from the Galapagos Rift and 21°N, have only cylindrical micropapillae near the border and on the surface (Pettibone, 1985c, fig. 1E). The elytra of the varietal form, de- scribed on a small specimen from the Juan de Fuca Ridge (Pettibone, 1988, fig. 5A, B) and described here from the Gorda Ridge (Fig. 6G-I) have some additional border pa- pillae enlarged basally. Distribution. —Hydrothermal vents of tropical East Pacific, Galapagos Rift and East Pacific Rise at 21°N, in 2450-2633 m; Northeast Pacific Explorer Ridge, Endeav- our Segment and Axial Seamount, Juan de Fuca Ridge, in 1546-2213 m; and off north- ern California Gorda Ridge, in 3240-3500 m. Levensteiniella intermedia, new species Figs 7, 8 Material. —East Pacific Ocean off north- ern California, Gorda Ridge, Alvin dive 2042, 12 Jun 1988, 41°00’N, 127°29'W, 3271 m, “clam washings,”’ holotype (USNM 123370). Description.—Body flattened ventrally, arched dorsally, slightly tapered anteriorly and more so posteriorly, with parapodia about as long as body width. Eleven pairs of elytra, oval to subreniform, thick, stiff, opaque, with thick oval raised areas or pro- jections on posterior border and some scat- tered oval micropapillae on surface (Fig. 7D— G). Prostomium wider than long, deeply bi- lobed with subtriangular lobes projecting anteriorly, with terminal filaments; without PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON eyes; ceratophore of median antenna large, bulbous, in anterior notch; style missing; palps long, stout, tapered; tentaculophores lateral to prostomium, achaetous, with small acicular lobe on inner side; dorsal and ven- tral tentacular cirri about same length as palps (Fig. 7A, B) Segment 2 with first pair of large elytrophores, biramous parapodia, and ventral buccal cirri attached basally lat- eral to ventral mouth, longer than following ventral cirri (Fig. 7A, B). Biramous parapodia with notopodium shorter than neuropodium, notopodium rounded with projecting acicular lobe on lower side; neuropodium with conical pro- jecting presetal acicular lobe; postsetal lobe shorter, rounded, deeply cut on dorsal side (Fig. 8A, B). Notosetae very numerous, forming radiating bundle, much stouter than neurosetae, of several lengths, stout, taper- ing to bare, blunt tips, with smooth or with widely spaced spines along one side (Fig. 8C). Neurosetae very numerous, forming fan-shaped bundle; supraacicular neurose- tae with longer spinous regions, with 2 rows of widely spaced spines along borders, with shorter close-set spines on tapered tips (Fig. 8E); subacicular neurosetae with slightly hooked bare tips and close-set minute spines along one border (Fig. 8D). Dorsal cirri with cylindrical cirrophores on anterior side of notopodia; styles with long filamentous tips, extending to about tips of neurosetae; dorsal tubercles nodular to truncate; ventral cirri short, tapering, and extending to distal end of ventral side of neuropodia (Fig. 8B). Single pair of long ventral papillae on seg- ment 11, much wider basally and extending almost to tip of neuropodium and ending in distal knob and subdistal filament (Fig. 7C). Pygidium small rounded lobe between bases of posterior small parapodia; anal cirri missing. Etymology. —The species is named based on its intermediate position between the other two species of Levensteiniella. Comparisons. —Important characters separating L. intermedia and the other two VOLUME 103, NUMBER 4 835 Fig. 6. Levensteiniella kincaidi, variety, USNM 123366: A, Dorsal view of anterior end; B, Ventral view of left side of segments 10-12, showing rounded papillae on segments 11 and 12; C, Right elytrigerous parapodium, anterior view, acicula dotted; D, Right cirrigerous parapodium, posterior view; E, Short and long notosetae; F, Tip of supraacicular and subacicular neurosetae; G, Right 2nd elytron from segment 4, with detail of papillae; H, Right 5th elytron from Segment 9, with detail of surface and border papillae; I, Right 10th elytron from segment 19, with detail of papillae. Scales = 1.0 mm for A, B; 1.0 mm for C, D; 0.1 mm for E, F; 2.0 mm for G-I. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Levensteiniella intermedia, holotype, USNM 123370: A, Dorsal view of anterior end, styles of median antenna and left ventral tentacular cirrus missing; B, Ventral view of same; C, Ventral view of left side of segments 10-12, showing long ventral papilla on segment 11; D, Right Ist elytron from segment 2; E, Right 2nd elytron from segment 4; F, Right middle elytron; G, Left posterior elytron. Scales = 1.0 mm for A-C; 2.0 mm for D-G. species of Levensteiniella are indicated in Table 3. Subfamily Branchinotogluminae Pettibone, 1985 Genus Branchinotogluma Pettibone, 1985 Branchinotogluma grasslei Pettibone, 1985 Branchinotogluma _ grasslei Pettibone, 1985b:457, figs. 5, 6; 1988:199: 1989b: LSS. Material. —East Pacific Ocean off north- ern California, Gorda Ridge, Alvin dive 2036, 6 Jun 1988, 41°00'N, 127°29’W, 3240 m, 9 specimens (USNM 123363); Alvin dive 2037, 7 Jun 1988, 3261 m, washings from vestimentiferans and alvinellids, 2 speci- mens (USNM 123362). Gorda Ridge, in northern Escanaba Trough, Sea Cliff dive 764, 3 Sep 1988, 41°00’N, 127°30'W, 3500 m, heavily sedimented area with vestimen- tiferans and long filamentous bacteria, 4 specimens (USNM 123364; SIO). Remarks.—The 15 specimens from the Gorda Ridge, in 3240-3500 m, agree with the specimens previously described from the VOLUME 103, NUMBER 4 837 \ Fig. 8. Levensteiniella intermedia, holotype, USNM 123370: A, Right elytrigerous parapodium, anterior view, acicula dotted; B, Right cirrigerous parapodium, posterior view; C, Long and short notosetae; D, Lower and upper supraacicular neurosetae; E, Supraacicular neuroseta. Scales = 1.0 mm for A, B; 0.1 mm for C-E. Table 3.—Comparison of three species of Levensteiniella. L. kincaidi Pettibone, 1985c L. raisae Pettibone, 1989 L. intermedia n. sp. Elytra with filiform micropapil- | with macro- and microtu- with thickened bulbous lae on border & sur- bercle-papillae on sur- projections on posterior face; some with en- face and posterior bor- border and oval micro- larged bases on der (Pettibone 1989a, papillae on surface (Fig. posterior border (Fig. Fig. 1G) 7D-F) 6G-I) Notosetae with spinous rows (Fig. with widely spaced spines smooth or with widely 6E) along one side (Petti- spaced spines along one bone 1989a, Fig. 2E) side (Fig. 8C) Long paired ventral 2 pairs on segments 11 & 2 pairs on segments 11 & single large pair on seg- papillae 12 (Pettibone 1985c, 12 (Pettibone 1989a, ment |1 (Fig. 7C) Fig. 1D) Fig. 1E) 838 Galapagos Rift and 21°N vent sites (Petti- bone 1985b) and reported from the Ex- plorer and Juan de Fuca Ridges (Pettibone 1988) and the Guaymas Basin (Pettibone, 1989b), in 1495-2633 m, associated with vestimentiferans. The specimens measured 9-22 mm in length, 5—11 mm in width, with 21 segments. Genus Opisthotrochopodus Pettibone, 1985b Opisthotrochopodus tunnicliffeae Pettibone, 1988 Opisthotrochopodus tunnicliffeae Pettibone, 1988:203, figs. 6-9 Material. —East Pacific Ocean off north- ern California, Gorda Ridge, Alvin dive 2036, 6 Jun 1988, 41°00’N, 127°29’W, 3240 m, 5 specimens (USNM 123375); Alvin dive 2037, 7 Jun 1988, 3261 m, washings from vestimentiferans and alvinellids, 1 speci- men (USNM 123376). Gorda Ridge, in northern Escanaba Trough, Sea Cliff dive 764, 3 Sep 1988, 41°00’N, 127°30'W, 3500 m, heavily sedimented area with vestimen- tiferans and long filamentous bacteria, 7 specimens (USNM 123374; SIO). Remarks.—The 13 specimens from the Gorda Ridge, in 3240-3500 m, agree with the specimens previously described from the Explorer and Juan de Fuca Ridges, in 1818- 1533 m. The larger specimens from the Gorda Ridge measured 19-22 mm in length, 9-12 mm in width, with 21 segments. One specimen from the Sea Cliff dive had a long coiled parasitic copepod in the body cavity and extending to the outside. Acknowledgments I wish to thank Verena Tunnicliffe of the University of Victoria for the specimen from the Juan de Fuca Ridge, J. Frederick Grassle and Rosemarie Petrecca of the Woods Hole Oceanographic Institution for the DSRV Alvin specimens, and Michael A. Boudrias of the Scripps Institution of Oceanography for the DSRV Sea Cliff specimens from the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Gorda Ridge. The manuscript benefited from the reviews of Thomas H. Perkins and James A. Blake. Literature Cited Detinova, N. N. 1988. New species of polychaetous annelids from hydrothermal vents of the Juan de Fuca Ridge (Pacific Ocean). — Zoologichesky Zhurnal 67(6):858-864. (In Russian, English summary.) Hanley, J.R. 1989. Revision of the scaleworm genera Arctonoe Chamberlin and Gastrolepidia Schmarda (Polychaeta: Polynoidae) with the erection of a new subfamily Arctonoinae. —The Beagle, Records of the Northern Territory Mu- seum of Arts and Sciences 6:1-34. Pettibone, M. H. 1985a. Polychaete worms from a cave in the Bahamas and from experimental wood panels in deep water of the North Atlantic (Polynoidae: Macellicephalinae, Harmothoi- nae). —Proceedings of the Biological Society of Washington 98:127-149. 1985b. Additional branchiate scale-worms (Polychaeta: Polynoidae) from Galapagos hy- drothermal vent and rift-area off Western Mex- ico at 21°N.—Proceedings of the Biological So- ciety of Washington 98:447-469. 1985c. New genera and species of deep-sea Macellicephalinae and Harmothoinae (Poly- chaeta: Polynoidae) from the hydrothermal rift areas off the Galapagos and Western Mexico at 21°N and from the Catalina Channel.—Pro- ceedings of the Biological Society of Washington 98:740-756. . 1988. New species and new records of scaled polychaetes (Polychaeta: Polynoidae) from hy- drothermal vents of the Northeast Pacific Ex- plorer and Juan de Fuca Ridges. — Proceedings of the Biological Society of Washington 101: 192-208. i 1989a. New. species of scale-worms (Poly- chaeta: Polynoidae) from the hydrothermal rift- area of the Mariana Back-arc Basin in the west- ern central Pacific.— Proceedings of the Biolog- ical Society of Washington 102:137-153. 1989b. Polynoidae and Sigalionidae (Poly- chaeta) from the Guaymas Basin, with descrip- tions of two new species, and additional records from hydrothermal vents of the Galapagos rift, 21°N, and seep-sites in the Gulf of Mexico (Flor- ida and Louisiana).— Proceedings of the Bio- logical Society of Washington 102:154—168. Department of Invertebrate Zoology, National Mu- seum of Natural History, Washington, D.C. 20560 PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 839-846 REDEFINITION OF TENERIDRILUS HOLMQUIST (OLIGOCHAETA: TUBIFICIDAE), WITH DESCRIPTION OF TWO NEW SPECIES FROM NORTH AMERICA Christer Erséus, Jarl K. Hiltunen, Ralph O. Brinkhurst, and Don W. Schloesser Abstract.—The tubificid genus Texeridrilus Holmquist, 1985 (subfamily Tubificinae) is redefined. The genus was originally monotypic for the type species, formerly classified as I/yodrilus mastix Brinkhurst, 1978. The genus now includes TJeneridrilus columbiensis (Brinkhurst & Diaz, 1985), a new com- bination for /sochaetides columbiensis, and two new species. The first of these, Teneridrilus calvus, is described by Erséus & Brinkhurst, and the second, Te- neridrilus flexus, by Erséus & Hiltunen. Synapomorphies for the genus are strongly modified chaetae in II and an enlarged eversible pharynx. The genus is distributed from China to British Columbia, Washington, and California in freshwater near the mouths of large rivers, and in the St. Marys River, which connects Lake Superior with Lakes Michigan and Huron. An aberrant freshwater tubificid with modified chaetae in segment II and greatly enlarged mouth and pharynx was described as Ilyodrilus mastix by Brinkhurst (1978) from the Fraser River in British Columbia, and later recorded by Erséus and Qi (1985) from the Pearl River in China. Holmquist (1985:343), who re-examined the type se- ries, challenged the original generic posi- tion: ““The species mastix with its quite dif- ferently built male apparatus, its entire lack of spermathecae, and the peculiarly built head region is not compatible with any of the above species, nor with any of the other members of the ‘J/yodrilus’ complex scru- tinized, but holds its own position.’ She accordingly established a monotypic genus, Teneridrilus, for I. mastix, an action that was supported by the original author (Brinkhurst, pers. comm., in Erséus & Qi 1985:194). The male efferent duct of 7. mastix con- sists of a narrow vas deferens, a stout atrium bearing a small prostate gland, and a small cone-shaped penis without thickened cuti- cle (see Holmquist 1985 for details and di- mensions). All atrial epithelial cells are of one type, unlike those of the tubificine genus Tubificoides. Although clearly a member of the Tubificinae, the species has a simple male apparatus atypical of most species in the subfamily. There are large ovaries but no spermathecae—the latter feature usually attributed to self-fertilizing or parthenoge- netic species in the Tubificidae (see Brink- hurst 1986a). Species with these character- istics are commonly classified together with species that have normal reproductive sys- tems in genera defined by phylogenetic prin- ciples. These involve recognizing shared apomorphies (synapomorphies) that indi- cate descent from a common ancestor de- spite more recent adaptations. Teneridrilus mastix was originally found in the tidal freshwater part of the Fraser River, British Columbia. Tubificoides fraseri Brinkhurst was described from more seaward sites in the same estuary in more saline water (Brinkhurst 1986b). It, too, is a suspected parthenogen with a simplified reproductive 840 system (spermathecae lacking), but is rec- ognizable as a true Tubificoides. Because such an adaptation is not seen as a generic character, we recognize three additional members of the genus Teneridrilus that share the peculiar modified chaetae of II and the enlarged pharynx of the type species but which have more elaborate male reproduc- tive systems than that of T. mastix. Tener- idrilus columbiensis and T. calvus, n. sp. are both found in tidal freshwaters in the Pacific Rim, but T. flexus, n. sp. is known only from the exit channel from Lake Superior. Material and Methods Specimens of Teneridrilus mastix and T. calyus, n. sp. were found during a study in the Sacramento-San Joaquin Delta in Cal- ifornia, and were placed at our disposal by Dr. W. C. Fields, Jr. (Newcastle, California). Specimens of T. flexus, n. sp. were collected in St. Marys River, the exit channel from Lake Superior, by one of us (D.W.S.). All specimens were mounted whole in Canada balsam and studied with a light microscope. The type series of the new species are de- posited in the National Museum of Natural History (USNM), Smithsonian Institution, Washington, D.C. Family Tubificidae Subfamily Tubificinae Teneridrilus Holmquist, 1985 Definition (emended).—Small tubificids. Prostomium small. Anterior end of worm wide and rather bluntly rounded; pharynx large, eversible, forming a folded divertic- ulum when withdrawn inside animal. All chaetae of II bifid, but modified: stouter, and with distal tooth thinner and shorter than bifid chaetae of other segments. Hair chaetae present in dorsal bundles (in 7. mastix) or absent. Modified (grooved) sper- mathecal chaetae present (in T. columbien- sis) or absent. Modified penial chaetae ab- sent. Male pores paired in XI. Spermathecal PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pores absent (in 7. mastix) or paired in X. Coelomocytes of the “‘rhyacodriline-type”’ absent. Male ducts paired. Vasa deferentia narrow, confined to XI. Atria cylindrical, small, each bearing one small, solid prostate gland (not yet confirmed for 7. flexus, n. sp.). Penes present, sometimes with thick- ened cuticular sheath (T. columbiensis and T. calvus, n. sp.), generally enclosed in mus- cular sacs. Spermathecae (absent in 7. mas- tix) with roundish ampullae and narrow ducts. Sperm as short and broad sperma- tozeugmata (in 7. columbiensis), or ran- dom, in spermathecae. Type species.—Ilyodrilus mastix Brink- hurst, 1978. Other species.—Teneridrilus columbien- sis (Brinkhurst & Diaz, 1985), n. comb., T. calvus Erséus & Brinkhurst n. sp., and T. flexus Erséus & Hiltunen n. sp. Remarks. —Monophyly of Teneridrilus is indicated by two synapomorphies in the four species of the genus: the large, modified feeding apparatus, and the modified bifid chaetae in segment II. All other features ap- pear homologous to states known also from other Tubificinae. Most of these traits are thus likely to prove to be plesiomorphic and should not be used as a basis for a phylo- genetic definition of the genus. The four species of the genus are all fresh- water forms, mostly occurring in riverine habitats. Teneridrilus mastix (Brinkhurst, 1978) Ilyodrilus mastix Brinkhurst, 1978:2171- 2173, fig. 3.—Erséus & Qi 1985:193-194, fig. 1. Teneridrilus mastix. —Holmquist 1985:332- 334, 336-341, 357-360, figs. 18, 21D-E, 22E, 23K—-L, 24P-R, 31A. New material (studied by, and in collec- tion of, R. O. Brinkhurst).—Several speci- mens from various sites in the Sacramento- San Joaquin Delta, in fine sediments with slow water flows (collected on several dates since April 1984; W. C. Fields, Jr.). VOLUME 103, NUMBER 4 Remarks.—This species, the type of the genus, is in fact the most atypical member of Teneridrilus. Unlike the other three spe- cies, it bears both hairs and bifid chaetae in the dorsal bundles. It has very simple penes in penial sacs that are not particularly mus- cular, and lacks spermathecae. Teneridrilus mastix is likely to be a re- productive opportunist, not reproducing by normal cross-fertilization. Distribution and habitat.—British Co- lumbia and California (new record), south- erm China. Fine sediments of rivers; in reshwater. Teneridrilus columbiensis (Brinkhurst and Diaz, 1985), new combination Tsochaetides columbiensis Brinkhurst & Diaz, 1985:949-952, figs. 1-2. Remarks. — Originally placed in Isochae- tides Hrabé, the monophyletic state of which is highly questionable (Brinkhurst 1986a), this species is here transferred to Teneri- drilus because it has the synapomorphies now recognized for the genus. Teneridrilus columbiensis is further char- acterized by lack of hair chaetae, possession of grooved spermathecal chaetae in segment X, and atria appearing as “simple widenings of vasa deferentia, with prostates attached near midpoints”’ (Brinkhurst and Diaz 1985: 950). Short ejaculatory ducts connect the atria with muscular penial bulbs that bear short, rounded to blunt-ended penis sheaths. The spermathecal ampullae are large and spherical and have short and broad sper- matozeugmata. Distribution and habitat.—Known only from Columbia River, Oregon. Tidal fresh- water marsh, muddy sediment. Teneridrilus calvus Erséus & Brinkhurst, new species Fig. 1 Type material.—USNM 123377, holo- type, whole-mounted specimen from the 841 Sacramento-San Joaquin Delta, Sacramen- to, California, muddy silt and clay (26 Feb- ruary 1987, W. C. Fields, Jr.). USNM 123378-123380, paratypes, 3 whole- mounted specimens from type locality. Other material. —Several specimens from the type locality, in the collections of R. O. Brinkhurst and C. Erséus. Etymology. —The epithet “‘calvus”’ is Lat- in for “bald, without hair,” and refers to the lack of hair chaetae in this species. Description. —Holotype 6.9 mm long, but not complete, consisting of the first 29 an- terior segments. Paratypes 4.6—5.3 mm long, with 22-33 segments. Width at XI (com- pressed specimens) 0.21—0.31 mm. Prosto- mium (Fig. 1A, pro) triangular, small. Body wall with scattered particles on surface. Cli- tellum extending over XI—XII. Chaetae all bifid, those of II (Fig. 1B) stout, 30-35 um long, about 2.5 wm thick, 2—3 per bundle; with distal tooth distinctly thinner and shorter than proximal. Chaetae of other an- teclitellar segments (Fig. 1C) about 50 wm long, 2.0—2.5 wm thick, up to 5(6) per bun- dle, with distal tooth almost as thick as, but longer than, proximal. Postclitellar chaetae up to about 40 um long, about 2 um thick, 2 per bundle, with teeth about equal in length and thickness. Chaetae absent ventrally in XI. Male pores paired in line with ventral chaetae in XI. Spermathecal pores imme- diately anterolateral to ventral chaetae in X. Mouth and pharynx (Fig. 1A) enlarged and complexly folded (semi-schematical in drawing), apparently forming a massive, eversible feeding apparatus. In all speci- mens studied, esophagus and gut filled (and distended) with sediment and detritus along most of worm, indicating that large amounts are engulfed. Pharyngeal glands not ob- served. Male genitalia (Fig. 1D) paired. Vas deferens 6—9 um wide, much longer than atrium, entering apical end of latter. Atrium 45-60 um long, 20—28 wm wide, somewhat comma-shaped or spindle-shaped, with api- cal end tilted over to posterior. Atrium with ventral bulge bearing (somewhat broadly at- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. semi-schematically drawn; B, Chaeta of segment II; C, Chaeta of segment IV; D, Male genitalia; E, Spermatheca from one specimen; F—G, Spermathecae from another specimen. Abbreviations: a atrium; br brain; m muscular bulb; mo mouth; p penis; ph pharynx; pr prostate gland; pro prostomium; vd vas deferens. tached?) discrete prostate gland. Atrium en- tering heavily muscular bulb, interior to which is a hollow sac with a cylindrical or somewhat tapering penis. Penis 60-95 um long, 18—30 wm wide, with thin but distinct cuticular lining (= a cylindrical sheath) along most of its length, but with ‘soft’? and some- what pointed tip, protruding from opening of sheath. Spermathecae (Fig. 1E-—G) with wide and distinct ducts, and spherical or oval ampullae; body wall thickened and somewhat folded at pore, often with a cir- cular groove surrounding pore (but details not clear in available material); ampullae 30-60 um long, 25—70 um wide; sperm ran- dom or as loose bundles in ampullae. Remarks. —This species is distinguished from 7. mastix and T. columbiensis by its characteristic, slender penes (Fig. 1D), which, however, are not as long as those of T. flexus, n. sp. (see below). The complex spermathecal pores of 7. calvus (Fig. 1G) are unique within the genus. The corresponding pores are inconspicuous in T. columbiensis and T. flexus, and absent in T. mastix. Distribution and habitat.—Known only from the Sacramento-San Joaquin Delta in Teneridrilus calvus Erséus & Brinkhurst, n. sp.: A, Anterior end of worm, with folding of pharynx California, where it is commonly associated with the following invertebrates (W. C. Fields, Jr., pers. comm.): Corbicula flumi- nea, Corophium stimpsoni, Anisogamma- rus ramellus, Limnodrilus angustipenis, L. hoffmeisteri, Bothrioneurum vejdovskya- num, Aulodrilus limnobius, Nereis limni- cola, and Prostoma graecense. Freshwater muddy sediments. Teneridrilus flexus Erséus & Hiltunen, new species Fig. 2 Type material.—USNM 123381, holo- type, whole-mounted specimen from St. Marys River, exit channel from Lake Su- perior, Chippewa Co., Michigan, U.S.A. (June 1985, D. W. Schloesser). USNM 123382-123383, paratypes, 2 whole- mounted specimens (one represented by fragments only) from type locality. Etymology. — The epithet ‘‘flexus”’ is Lat- in for “winding” and refers to the flexible shape of the penes in this species. Description. —Length of holotype 4.1 mm, about 32 segments (worm coiled and dis- torted), paratypes not complete. Width at VOLUME 103, NUMBER 4 843 Fig. 2. Teneridrilus flexus Erséus & Hiltunen, n. sp.: A, Chaeta from segment II; B, Bundle of chaetae from segment V; C, Chaeta from postclitellar segment; D, Male genitalia from one side of much distorted specimen (paratype), with prostate gland (if present at all?) and vas deferens detached from atrium; E, Copulatory organ of other paratype, with penis protruded; F, Spermatheca. Abbreviations: a atrium; m muscular bulb; p penis; ?pr possible attachment of prostate gland; vd vas deferens. XI (compressed specimens) 0.24—0.25 mm. Prostomium somewhat triangular, small. Body wall without adhering particles. Cli- tellum extending over XI—XII. Chaetae all bifid, those of II (Fig. 2A) stout, about 30 um long, about 2 wm thick, 2—3 per bundle, with distal tooth distinctly thinner and shorter than proximal. Chaetae of segments III through VII or thereabouts (Fig. 2B) clearly longer than those of II, 40-47 um long, 2.0—2.5 um thick, 4—6 per bundle, with distal tooth about as thick as the proximal, but clearly longer. Chaetae of mid-body and posterior part of worm (Fig. 2C) about 35 um long, about 1.5 um thick, 1—3 per bundle (bundles sometimes missing?), somewhat similar in shape to those of II. Chaetae ab- sent ventrally in XI. Male and spermathecal pores paired, ventrally in XI and X, re- spectively, but exact position in relation to chaetal lines difficult to see in available ma- terial. Mouth and pharynx modified as in T. cal- vus (see Fig. 1A); esophagus and gut filled with sediment along most of worm. Pha- ryngeal glands not observed. Male genitalia (Fig. 2D-E) paired. Vas deferens about 5-— 7 wm wide, longer than atrium, but detached from atrium in the paratype, where it is best visible (Fig. 2D). Atrium inconspicuous (visible only in one paratype; Fig. 2D), slen- der, about 40 um long, 9-11 um wide, with non-muscular outer lining and virtually non- granulated inner epithelium. Prostate gland not observed, but a small structure may be remains of a prostate stalk in (much dis- torted) paratype (Fig. 2D:?pr). Atrium en- tering heavily muscular bulb, interior of which is a hollow sac with a very long, ta- pering, flexible penis. Penis about 150-200 um long, 15—20 um wide at base, only about 5 um wide at tip, either coiled and retained within sac (Fig. 2D) or protruding through male pore, reaching far out from worm (Fig. 844 2E). Penis appearing somewhat cuticular- ized, but distinct sheath not formed. Sper- mathecae (Fig. 2F) small, with very slender ducts, 80-90 um long, 12—15 wm wide, and round or oval ampullae, 35—40 um long, 30- 35 um wide. Sperm as loose bundle or ran- dom mass in ampullae. Remarks.—This species is easily distin- guished from the closely related T. calvus by its clearly longer penes and more slender spermathecal ducts (cf. Figs. 1-2). Distribution. —Known only from type lo- cality (St. Marys River). Discussion Aulodrilus paucichaeta Brinkhurst & Bar- bour, 1985, known from freshwater marsh- es in Maryland (and possibly North Caro- lina), is a very large form, up to 50 mm long, which has some features in common with these smaller Teneridrilus species. The small atria, intimately associated with large, spherical, muscular penial bulbs, the wedge to spherical shaped penes, and the bundled sperm in the spermathecae make A. pau- cichaeta similar to the taxa treated here. However, in A. paucichaeta the chaetae of segment II are not much different from those of the following segments, and the pharynx does not appear to be modified in the TJe- neridrilus manner (observations on some of the original specimens of A. paucichaeta in Brinkhurst’s collection). Therefore, the spe- cies is excluded from Teneridrilus. The European Peipsidrilus, established by Timm (1977) for P. pusillus Timm, 1977, later revised by Finogenova (1983) to in- clude Umbadrilus saamicus Timm, 1978 (described in Timm & Popchenko 1978), and further revised by Giani et al. (1984) to include Neoaulodrilus libanus Giani et al., 1982, is probably closely related to Te- neridrilus. The three species of Peipsidrilus are also small freshwater forms (length 6-8 mm), with simple atria and small penes en- closed in muscular sacs. In P. pusillus, the chaetae of segment II, as well as those of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the postclitellar segments, have teeth that are equally long, whereas the chaetae of oth- er segments have distal teeth that are longer than the proximal ones (Finogenova 1983). This is noteworthy, as it foreshadows the state in Teneridrilus, in which the distal tooth of the segment II chaetae is shorter than the proximal. Similarly, although none of the descriptions of P. pusillus (Timm 1977, Finogenova 1983) mentions any modifica- tion of the feeding apparatus, the original drawing by Timm (1977:fig. 1) gives the im- pression that the pharynx is slightly wider than in other tubificids, and thus appears transitional toward the modification seen in Teneridrilus. These circumstances suggest that P. pusillus in fact may be the plesiomor- phic sister taxon to Teneridrilus, but wheth- er the two genera should be united cannot be determined until the states of these char- acters (chaetae of II and feeding apparatus) in all species of Peipsidrilus have been care- fully reexamined. The taxa discussed so far are only a part of a whole complex within the Tubificinae characterized by rather simple atria and pe- nes, and by a tendency toward bearing ran- dom sperm instead of spermatozeugmata in the spermathecae. Some loose sperm were observed in the spermathecae of 7. co- lumbiensis by one of us (C.E.) but the sig- nificance of this is unclear until tubificines with the same feature are examined. Genera involved in this complex are Aulodrilus Bretscher (Giani et al. 1984), Isochaetides Hrabé emend., Brinkhurst (Brinkhurst 1984, 1986a), Peipsidrilus Timm (1977), Krened- rilus Dumnicka (Gianiet al. 1990), and Ske- todrilus Karaman, 1976. The morphologi- cal features of these often appear to be plesiomorphic and the species in question are thus likely to be less derived Tubificinae. However, some of the species may be ad- vanced members of the subfamily; i.e., their lack of genital complexity is due to regres- sion linked with reduction in body size or modification in reproductive strategy (see comments on 7. mastix in Introduction). VOLUME 103, NUMBER 4 For instance, some species of Aulodrilus combine relatively simple genitalia with presumed advanced traits such as modified and very numerous chaetae, tube-dwelling, and asexual reproduction. Phylogenetic analysis to determine such questions should be attempted after some of the generic def- initions have been clarified. The scarcity of unequivocal apomorphic states has made it difficult to resolve the phylogenetic relationships within the lower Tubificinae, and the generic classification has to a great extent represented phenetic similarity and unique combinations of char- acters rather than monophyletic groups based on synapomorphies—a statement that unfortunately is true also for many other tubificids. In the present paper, we do not attempt to rectify this difficulty for other genera, but we claim that there is evidence that the four Teneridrilus species constitute a true monophyly. Acknowledgments We are grateful to Dr. W. C. Fields, Jr. for placing oligochaete material at our dis- posal, and to Ms. Barbro Lofnertz and Ms. Aino Falck-Wahlstrom (University of Go- teborg) for technical assistance. This paper is Contribution 742 of the National Fish- eries Research Center—Great Lakes, U.S. Fish and Wildlife Service, Ann Arbor, Michigan 48105. Literature Cited Brinkhurst, R. O. 1978. Freshwater Oligochaeta in Canada.—Canadian Journal of Zoology 56: 2166-2175. 1984. A revision of the Tubificidae and Ly- codrilidae (Annelida, Oligochaeta) known from Lake Baikal. — Canadian Journal of Zoology 62: 494-509. 1986a. Guide to the freshwater aquatic mi- crodrile oligochaetes of North America.—Ca- nadian Special Publication of Fisheries and Aquatic Sciences 84:1-259. 1986b. Taxonomy of the genus Tubificoides Lastochin (Oligochaeta, Tubificidae): Species 845 with bifid setae.—-Canadian Journal of Zoology 64:1270-1279. , & M. T. Barbour. 1985. A new species of Aulodrilus Bretscher (Oligochaeta:Tubificidae) from North America.—Proceedings of the Bi- ological Society of Washington 98:93 1-934. ——, & R. J. Diaz. 1985. Isochaetides columbien- sis, new species (Oligochaeta:Tubificidae) from the Columbia River, Oregon.— Proceedings of the Biological Society of Washington 98:949- 953. Erséus, C., & Qi Sang. 1985. Two aberrant Tubifici- dae (Oligochaeta) from Pearl River in the Peo- ple’s Republic of China.—Hydrobiologia 127: 193-196. Finogenova, N. P. 1983. A revision of the genera Peipsidrilus Timm, 1977 and Umbadrilus Timm, 1978 (Tubificidae, Oligochaeta). Pp. 199-203 in Limnologicheskie issledovaniya na zalive Onezhskogo ozera Bol’shoye Onego:Sbornik nauchnykh trudov. Academy of Sciences U.S.S.R., Zoological Institute, Leningrad. [In Russian. |] Giani, N., C. Erséus, & E. Martinez-Ansemil. 1990. Redefinition of the subterranean genus Krened- rilus Dumnicka with a redescription of K. papil- latus Dumnicka, 1983 and the description of two new species. —Stygologia 5:55-65. , E. Martinez-Ansemil, & R. O. Brinkhurst. 1984. Revision du statut taxonomique des Aulodrilinae (Tubificidae, Oligochaeta).—Bul- letin de Ja Societe d’Histoire Naturelle de Tou- louse 120:17-22. , E. Martinez-Ansemil, Z. Moubayed, & A. Dia. 1982. Les Oligochetes aquatiques du Liban. I. Neoaulodrilus libanus n. g.,n. sp. et Nais iorensis Pataridze.— Annales de Limnologie 18:179-190. Holmquist, C. 1985. A revision of the genera Tubifex Lamarck, J/yodrilus Eisen, and Potamothrix Vejdovsky & Mrazek (Oligochaeta, Tubifici- dae), with extensions to some connected gen- era.— Zoologische Jahrbiicher (Systematik) | 12: 311-366. Karaman, S. 1976. The second contribution to the knowledge of the freshwater Oligochaeta of Slo- venia.— Bioloski Vestnik 24:201-—207. Timm, T. 1977. Peipsidrilus pusillus gen. n. sp. n. (Oligochaeta, Tubificidae).—Eesti NSV Tea- duste Akadeemia Toimetised 26 (Biologia):279-— 283. — , & V. Popchenko. 1978. The aquatic Oligo- chaeta of the Murmansk region. Pp. 71—132 in Hydrobiological researches VII. Seasonal phe- nomena in fresh-water biology. Academy of Sci- ences of the Estonian S.S.R., Institute of Zool- ogy and Botany, Tartu. [In Russian, with English summary.] 846 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (CE) Zoo-tax, Swedish Museum of Nat- ural History, Stockholm, and (postal ad- dress) Department of Zoology, University of Goteborg, Box 25059, S-400 31 Gote- borg, Sweden; (JKH and DWS) National Fisheries Research Center—Great Lakes, 1451 Green Road, Ann Arbor, Michigan 48105; (ROB) Institute of Ocean Sciences, P.O. Box 6000, Sidney, British Columbia V8L 4B2, Canada. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 847-853 ADDITIONAL RECORDS OF STOMATOPOD CRUSTACEANS FROM ISLA DEL COCO AND GOLFO DE PAPAGAYO, EAST PACIFIC OCEAN David K. Camp and Hans G. Kuck Abstract.—Four species of stomatopod crustaceans are reported from Isla del Coco, East Pacific Ocean: Gonodactylus zacae Manning, 1972; Pseudosquilla adiastalta Manning, 1964; Crenatosquilla oculinova (Glassell, 1942); and Het- erosquilloides mccullochae (Schmitt, 1940). Three of the species, G. zacae, C. oculinova, and H. mccullochae, are reported from Isla del Coco for the first time. The only other stomatopod known from there is Neocoronida cocosiana (Manning, 1972). A monodactyla postlarva, tentatively assigned to P. adiastal- ta, is briefly characterized and illustrated. Records from the Golfo de Papagayo, Costa Rica, of Squilla panamensis Bigelow, 1891, and the poorly known S. biformis Bigelow, 1891, are also presented. The original definition of Crena- tosquilla Manning, 1984, contained errors that are corrected here. Heterosquil- loides mccullochae is shown to differ from the current definition of every family in the superfamily to which the species is assigned. Isla del Coco is an isolated oceanic island located at 5°32'57”N, 86°59'17”W, approx- imately 500 km southwest of Costa Rica and 630 km northeast of the Galapagos Ar- chipelago (see Bakus 1975, Hogue & Miller 1981, and Abele & Kim 1984 for general information, biogeography, and ecology of the island; see Hertlein 1963 for faunal checklist and bibliography). Although the molluscan fauna of Isla del Coco has been extensively sampled (e.g., Hanna & Hertlein 1938, Hertlein 1963, Shasky 1983), little effort has been directed toward surveying the Stomatopoda until now. Collections made during 1932-1933 and 1938 by the R/V Velero III of the Allan Hancock Foun- dation, University of Southern California (AHF), proved that a rich crustacean fauna existed on and around the island, but no stomatopods were collected (John S. Garth and Janet Haig, AHF, pers. comm.). Hert- lein (1963) reported fifty species of Crus- tacea from the island, but none were sto- matopods. Few additional species of crustaceans have been reported from Isla del Coco since that time (Manning 1972, Abele & Kim 1984). Forty-two species of stomatopods are known to occur in the tropical eastern Pa- cific (Salgado-Barragan & Illescas-Monter- roso 1987:159). Reaka & Manning (1980) reviewed the stomatopod fauna of Pacific Costa Rica and nearby offshore islands. They listed only two species known from Isla del Coco: Neocoronida cocosiana (Manning, 1972), the only stomatopod endemic to the island, and Pseudosquilla adiastalta Man- ning, 1964, the only species of Pseudosquilla known from the East Pacific Region (Reaka & Manning 1987:15). In 1988 one of us (HGK) participated in a general collecting expedition to Parque Nacional Isla del Coco (Fig. 1) aboard the schooner M/S Victoria af Carlstad. The present report is principally based on small collections of stomatopods taken during that trip by personnel from the Natural History Museum of Los Angeles County (LACM) using SCUBA (3-43 m depths). Also in- cluded in this note are specimens collected 848 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 5°30’ | Fig. 1. Isla del Coco, showing sites at which stomatopods were collected. Closed circle = Gonodactylus zacae; Open square = Pseudosquilla adiastalta; Cross = Crenatosquilla oculinova; Star = Heterosquilloides mccullochae. at Isla del Coco by Kirstie Kaiser, Michel Montoya, and Douglas von Kriegelstein us- ing trawl, dredge, and tangle net (18—46 m depths) during March 1989 and donated to LACM. Finally, specimens captured by trawling (66-276 m depths) in the Golfo de Papagayo, Costa Rica (Fig. 2), by William Bussing aboard the Japanese vessel R/V Nisshin Maru during 1987-1988 are listed. The waters at Isla del Coco abound with suitable habitats for stomatopods. During the 1988 trip, inclement weather prevented collecting anywhere but in and around the protected bays of the north side of the is- land. Specimens reported here were found in several habitats. Some specimens were collected in sand and rubble beneath dead patches of the coral Porites lobata Dana, 1846. Others were collected from deep within the branches of living colonies of the much less abundant Pocillopora capitata Verrill, 1864. Many specimens were found in the medium-grained, light brown to white sand under basaltic rocks in the wide talus zones at the bases of the many islets sur- rounding Isla del Coco. The sheer rock walls of some islets and seamounts have many crevices and holes with stomatopods, but VOLUME 103, NUMBER 4 Bahia Playa Blanca _): e+ 849 PROVINCIA DE GUANACASTE Golfo de Papagayo Fig. 2. Golfo de Papagayo, Pacific coast of Costa Rica, showing sites at which stomatopods were collected. Cross = Squilla biformis; Closed circle = Squilla panamensis. specimens are difficult to collect because of the depth to which the stomatopods recede within the crevices and because of the strong currents in those habitats, which make div- ing difficult. Carapace length (cl) is measured on the midline and does not include the rostral plate, but total length does include the plate. All specimens have been deposited at LACM and at the Florida Marine Research Insti- tute, Marine Invertebrate Collection (cata- log number prefix FSBC I). Family Gonodactylidae Giesbrecht, 1910 Gonodactylus zacae Manning, 1972 Material.—Isla del Coco: Northeast of Isla Manuelita, seamount, 5°33.88'N, 87°02.78'W, depth 18-43 m; leg. H. G. Kuck and A. Jahn, 29 Apr 1988; 1 male, cl = 2.5 mm; LACM 88-38.1.—Bahia Wafer, Pori- tes reef, east side of Isla Gissler, 5°32.8’N, 87°03.82’W, depth 3-6 m; leg. H. G. Kuck and R. W. Peck, 25 Apr 1988; 1 female, cl = 4.3 mm; FSBC I 34187 (LACM 88-12).— Bahia de Chatham, depth 46 m; leg. K. Kai- ser, 26 Mar 1989; 1 male, cl = 2.7 mm; LACM 89-28.1. Remarks. —The specimens reported here agree well with the illustrations of G. zacae presented by Manning (1974:fig. 1), except that the lateral margins of the ocular scales are not as concave as those he depicted. The species has not been reported before from Isla del Coco, but its occurrence there is not surprising. Gonodactylus zacae is thought to be the most widely distributed gonodac- tylid and the most common Gonodactylus 850 in the East Pacific Region (Manning 1974: 104, Reaka & Manning 1980:9). Family Pseudosquillidae Manning, 1977 Pseudosquilla adiastalta Manning, 1964 Material.—Isla del Coco: Bahia Wafer, Porites reef, east side of Isla Gissler, 5°32.8'N, 87°03.82'’W, depth 3-6 m; leg. H. G. Kuck and R. W. Peck, 25 Apr 1988; 1 male, 2 females, cl = 4.6-6.7 mm; LACM 88-12.1.—Bahia Weston, Porites reef, southwest side of Isla Pajara, 5°33.28'N, 87°03.45'W, depth 9-15 m; leg. R. W. Peck and H. G. Kuck, 27 Apr 1988; 1 male, cl = 14.8 mm; FSBC I 34188 (LACM 88-23).— Approx. 8 km north of Bahia de Chatham, 5°38.0'N, 87°02.55'W, from stomach of yel- lowfin tuna Thunnus albacares (Bonnaterre, 1788) (approx. length 1 m, approx. weight 165 kg, hook and line); leg. M. Dell’ Aquila, 30 Apr 1988; 1 monodactyla postlarva, cl = 4.4 mm; LACM 88-41.1 Remarks. —The subadult male with car- apace length 4.6 mm (LACM 88-12.1) lacks the two large patches of dark chromato- phores on the carapace that are usually char- acteristic of adults (see Manning 1964:fig. 1). The white spots on that specimen are relatively larger than similar spots on adults and are positioned evenly over the dorsal surface of the entire body. We have assigned to this species the post- larva (LACM 88-41.i) found in the stom- ach contents of a tuna caught north of Isla del Coco, even though the specimen is not in good condition and the postlarval stage of P. adiastalta has never been described. Manning (1969) reviewed the morphologi- cal characteristics of postlarvae of the At- lantic species P. ciliata (Fabricius, 1787) and P. oculata (Brullé, 1837). The specimen from off Isla del Coco differs from P. ciliata in the same way that P. oculata does (see Man- ning 1969:270, 277). The specimen is more similar to the postlarva of P. oculata, which shares the following characteristics. Total PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON length of the specimen is 25 mm. The eyes are large, with broadened cornea, and the rostral plate is cordiform (Fig. 3a). The sixth abdominal somite has submedian spines but lacks intermediate spines (Fig. 3b). The me- dian cleft of the telson is pronounced. The outer margin of the uropod is armed with ten movable spines, and the proximal seg- ment of the outer ramus of the uropod is more than three times the length of the dis- tal segment (Fig. 3b, d). The outer spine of the basal prolongation of the uropod is lon- ger than the inner, and the inner margin of the outer spine of the prolongation is angled. There are posterolateral spines on the fifth and sixth abdominal somites; however, un- like the postlarva of P. oculata, the fourth somite is acutely angled posterolaterally and lacks a spine (Fig. 3c). A complete descrip- tion of the postlarva of P. adiastalta must await collection of more suitable material from the area. Family Squillidae Latreille, 1803 Crenatosquilla oculinova (Glassell, 1942) Material. —Isla del Coco: East side of Isla Manuelita, depth 18 m; leg. M. Montoya and K. Kaiser, 21 Mar 1989; 1 female, cl = 5.1 mm; LACM 89-29.1. Remarks. — The specimen agrees well with Glassell’s (1942) description of the species. Manning (1984) erected the monotypic ge- nus Crenatosquilla for Squilla oculinova. His definition of Crenatosquilla contained two errors: 1) specimens of C. oculinova have three epipods on the maxillipeds, not four as Manning stated; and 2) the dorsal ridge of the carpus of the raptorial claw, rather than being indistinct, is prominent and ends in a distal tooth (see Glassell 1942:54). Crenatosquilla oculinova has not been re- ported before from Isla del Coco. Reaka & Manning (1980:16) reported the species to be the most common stomatopod collected in their study of Pacific Costa Rica. VOLUME 103, NUMBER 4 C Fig. 3. Pseudosquilla adiastalta postlarva, LACM 88-41.1: a, Anterior portion of body, dorsal view, setae and antennular flagella omitted; b, Sixth abdominal somite, telson and left uropod, dorsal view, setae omitted: c, Abdominal somites 3-6, lateral view of right side; d, Left uropod, ventral view, setae omitted. (Scale line = 2 mm.) Squilla biformis Bigelow, 1891 Material. —Costa Rica, Provincia de Guanacaste: Golfo de Papagayo, 10°45.0’N, 86°12.0’'W-10°46.2'’N, 86°13.0’W, depth 206-208 m; leg. W. Bussing, R/V Nisshin Maru, 7 Dec 1987; 1 female, cl = 28.3 mm; FSBC I 34185 (LACM 87-217).—Golfo de Papagayo, 10°30.5'’N, 86°13.6'’W- 10°32.0'N, 86°14.6’'W, depth 268-276 m; leg. W. Bussing, R/V Nisshin Maru, 10 Dec 1987; 2 females, 1 male, cl = 30.5—33.2 mm; LACM 87-216.1.—Bahia Playa Blanca, northwest of Golfo de Papagayo, 10°58.4’N, 86°07.5'W-10°00.1'N, 86°08.1'W, depth 131-139 m; leg. W. Bussing, R/V Nisshin Maru, 9 Jan 1988; 1 female, cl = 30.8 mm; LACM 88-108.1. Remarks.—Squilla biformis is captured infrequently despite its rather wide distri- bution in the tropical East Pacific. It has been reported only from the Golfo de Cal- ifornia, the Golfo de Panama, and off Peru (Manning 1974:108). Squilla panamensis Bigelow, 1891 Material. —Costa Rica, Provincia de Guanacaste: Golfo de Papagayo, 10°45.0'N, 86°12.0’'W-10°46.2'N, 86°13.0'W, depth 852 206-208 m; leg. W. Bussing, R/V Nisshin Maru, 7 Dec 1987; 1 male, cl = 24.2 mm; FSBC I 34186 (LACM 87-217).—Golfo de Papagayo, 10°40.7’N, 85°46.3’W- 10°42.1'N, 85°47.2'W, depth 66-67 m; leg. W. Bussing, R/V Nisshin Maru, 13 Nov 1987; 2 females, cl = 14.9-approx. 22 mm (latter damaged), 1 female abdomen and tel- son only, 1 male, cl = 22.7 mm; LACM 87- 214.1.—Bahia Playa Blanca, northwest of Golfo de Papagayo, 10°58.4'N, 86°07.5’W— 10°00.1’N, 86°08.1’W, depth 131-139 m; leg. W. Bussing, R/V Nisshin Maru, 9 Jan 1988; 1 female, 1 male (abdomens and tel- sons only); LACM 88-108.1 _ Remarks.—This species is known from Bahia Petatlan, México, to Tumbes, Peru (Manning 1974:108). Family Lysiosquillidae Giesbrecht, 1910 Heterosquilloides mccullochae (Schmitt, 1940) Material.—Isla del Coco: Porites reef, northeast side of Isla Manuelita, 5°33.87’'N, 87°02.12'W, depth 12-20 m; leg. R. W. Peck and H. G. Kuck, 26 Apr 1988; 2 females, cl = 10.6—14.8 mm; LACM 88-19.1.—Pori- tes reef, east side of Isla Manuelita, 5°33.8'N, 87°02.07'W, depth 9-12 m; leg. H. G. Kuck and R. W. Peck, 26 Apr 1988; 1 male, cl = 10.3 mm; LACM 88-20.1.—Bahia Weston, west side of Isla Pajara, 5°33.48'N, 87°03.40'W, depth 21 m; leg. K. Burke, 27 Apr 1988; 1 male, cl = 9.8 mm; LACM 88- 24.1.—Bahia de Chatham, boulder talus zone at base of wall, west side of Isla Ulloa, 5°33.20'N, 87°02.07'W, depth 8-26 m; leg. R. W. Peck, H. G. Kuck, and M. Dell’ Aq- uila, 29 Apr 1988; 1 female, cl = 13.8 mm; FSBC I 34189 (LACM 88-36). Remarks. —The specimens agree well with descriptions of the species published by Schmitt (1940:197) and Manning (1969:55); all have the distinctive chromatophore pat- tern illustrated by Schmitt. Heterosquil- loides mccullochae, although relatively un- common, has the widest distribution of any PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON stomatopod (Reaka & Manning 1987:15). It is a circumtropical species, known from the East Pacific, West Atlantic, Central At- lantic (Ascension Island), and Indian oceans (Reaka & Manning 1980:11, 1987:15). A new family might have to be erected to receive this species. Manning (1980:367— 368) revised the superfamilies, families, and genera of Recent stomatopods. Characters of Heterosquilloides mccullochae do not fit the definition of any of the three families (Lysiosquillidae, Coronididae, and Nan- nosquillidae) that he placed within the su- perfamily Lysiosquilloidea, nor do the char- acters fit the more recently discovered Erythrosquillidae, also tentatively placed in the Lysiosquilloidea (see Manning & Bruce 1984:332). Heterosquilloides mccullochae differs from members of the Lysiosquillidae and the Erythrosquillidae by having the en- dopods of the first two walking legs subcir- cular rather than slender and strap-like, from members of the Nannosquillidae by lacking a proximal fold on the outer margin of the uropodal endopod, and from members of the Coronididae by lacking the basal infla- tion of the dactylus of the raptorial claw. Because of the morphological characteris- tics and unusually broad distributional pat- tern of Heterosquilloides mccullochae, the species is certainly unique within the Ly- siosquilloidea. Acknowledgments The 1988 collecting trip in which HGK participated was sponsored by the Ichthy- ology Section of the Natural History Mu- seum of Los Angeles County, Los Angeles, California, and funded by the LACM Life Science Research Fund and the Taylor Re- search Fund through a grant to Robert J. Lavenberg (Curator of Fishes, LACM). We extend our gratitude to Robert Lavenberg for providing financial support for HGK to participate in the 1988 expedition; Manuel Murillo, Michel Montoya, and William Bussing of the Universidad de Costa Rica, VOLUME 103, NUMBER 4 Museo de Zoologia for providing lab space, specimens, logistical support, and hospital- ity during the 1988 stay of HGK in Costa Rica; Robert W. Peck, formerly of LACM, for his keen eye in spotting stomatopods during many dives off Isla del Coco; the crew of the M/S Victoria af Carlstad for excellent logistical support and good com- panionship during the expedition; John S. Garth and Janet Haig (AHF) for informa- tion and data on early AHF expeditions to Isla del Coco; Kirstie Kaiser and Douglas von Kriegelstein for the donation of speci- mens to LACM; Raymond B. Manning (Smithsonian Institution) for discussion of the familial placement of Heterosquilloides mecullochae and for reading an early draft of the manuscript; Joel W. Martin (LACM) for his motivation, guidance to HGK, and editorial comments; Judy Leiby, William G. Lyons, Thomas H. Perkins and James F. Quinn, Jr. (Florida Marine Research In- stitute, FMRI) for advice about the manu- script; and Llyn French (FMRI) for assis- tance with the figures. Literature Cited Abele, L. G., & W. Kim. 1984. Notes on the fresh- water shrimps of Isla del Coco with the descrip- tion of Macrobrachium cocoense, new species. — Proceedings of the Biological Society of Wash- ington 97:95 1-960. Bakus, G. J. 1975. Marine zonation and ecology of Cocos Island, off Central America.— Atoll Re- search Bulletin 179:1-11. Glassell, S. A. 1942. A new stomatopod crustacean from the west coast of Mexico. — Proceedings of the United States National Museum 92(3138): 53-56. Hanna, G. D., & L. G. Hertlein. 1938. Land and brackish water Mollusca of Cocos Island.— Al- lan Hancock Pacific Expeditions 2(8):123-135. Hertlein, L. G. 1963. Contribution to the biogeog- raphy of Cocos Island, including a bibliogra- phy.— Proceedings of the California Academy of Sciences 32(8):219-289. Hogue, C. L., & S. E. Miller. 1981. Entomofauna of Cocos Island, Costa Rica.— Atoll Research Bul- letin 250:1-29. Manning, R. B. 1964. A new West American species 853 of Pseudosquilla (Stomatopoda).—Crustaceana 6(4):303-308. 1969. Stomatopod Crustacea of the western Atlantic.—Studies in Tropical Oceanography, Miami, 8:viii + 380 pp. 1972. Three new stomatopod crustaceans of the family Lysiosquillidae from the eastern Pa- cific region.— Proceedings of the Biological So- ciety of Washington 85(21):271-278. 1974. Stomatopods collected by Th. Mor- tensen in the eastern Pacific region (Crustacea, Stomatopoda).—Steenstrupia 3(11):101-109. . 1980. The superfamilies, families, and genera of Recent stomatopod Crustacea, with diagno- ses of six new families. — Proceedings of the Bi- ological Society of Washington 93:362-372. 1984. Crenatosquilla, a new genus of sto- matopod crustacean from the East Pacific. — Proceedings of the Biological Society of Wash- ington 97:191-193. »&A.J. Bruce. 1984. Erythrosquilla megalops, a remarkable new stomatopod from the western Indian Ocean.—Journal of Crustacean Biology 4(2):329-332. Reaka, M. L., & R. B. Manning. 1980. The distri- butional ecology and zoogeographical relation- ships of stomatopod Crustacea from Pacific Costa Rica.—Smithsonian Contributions to the Marine Sciences 7:iii + 29 pp. ——.. 1987. The significance of body size, dispersal potential, and habitat for rates of morphological evolution in stomatopod Crustacea.—Smith- sonian Contributions to Zoology 448:iii + 46 pp. Salgado-Barragan, J., & C. Illescas-Monterroso. 1987. First record of Lysiosquilla panamica Manning, 1977 (Crustacea: Stomatopoda) in the Pacific waters of Mexico.—Revista de Biologia Tropi- cal 35(1):159-160. Schmitt, W. L. 1940. The stomatopods of the west coast of America, based on collections made by the Allan Hancock Expeditions, 1933-38.—Al- lan Hancock Pacific Expeditions 5(4):129-225. Shasky, D. R. 1983. New records of Indo-Pacific Mollusca from Cocos Island, Costa Rica. —The Nautilus 97(4):144-145. (DKC) Florida Marine Research Insti- tute, 100 Eighth Avenue, S.E., Saint Pe- tersburg, Florida 33701-5095; (HGK) Sec- tion of Invertebrate Zoology, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 854-860 A NEW SPECIES OF CHIROSTYLID CRUSTACEAN (DECAPODA: ANOMURA) FROM OFF THE WEST COAST OF NORTH AMERICA Keiji Baba and Janet Haig Abstract. —A new chirostylid, Gastroptychus iaspis, is described from depths of 600-1189 m off Mexico, California and Oregon. Its affinities to Gastroptychus defensa (Benedict, 1902) from the Galapagos Islands are discussed based on examination of two syntypes of that species. This is the fifth species of the genus in the eastern Pacific. In the course of studying benthic com- munities of deep hard bottom seamounts Dr. Amatzia Genin of the Hebrew Univer- sity of Jerusalem, then of the Scripps Insti- tution of Oceanography, has observed spec- imens of an anomuran crustacean belonging to the chirostylid genus Gastroptychus on Jasper Seamount off Baja California. Ac- cording to him, this chirostylid was a very important member of the seamount com- munity at the 600 to 1100 m depth interval, usually seen on gorgonians and antipathar- ians (A. Genin, pers. comm.). Two speci- mens of this chirostylid were collected; one is deposited in the Smithsonian Institution (USNM) and the other in the Allan Hancock Foundation (AHF). The specimens were made available for study by the courtesy of Dr. Genin and Dr. Austin B. Williams of the National Marine Fisheries Service. In the meantime one of us (JH) found that the following specimens were identical with those from the Jasper Seamount: four AHF specimens taken in 950-1189 m off Cah- fornia, and reported by Wicksten (1982:245) as Chirostylus sp.; 12 USNM specimens on loan to AHF, collected off Oregon in 914 m by the Commando, U.S. Bureau of Com- mercial Fisheries-AEC Project; and mate- rial from British Columbia and Washington described and illustrated as Chirostylus sp. by Hart (1982:166, fig. 65). Their identity was confirmed also by K. Baba during a visit to the Allan Hancock Foundation in 1989 and they represent an undescribed species. The measurements of the specimens ex- amined are given in parentheses under ‘*Material,”’ indicating postorbital carapace lengths. Gastroptychus iaspis, new species Figs. 1, 2 Chirostylus sp. Wicksten, 1982: 245 (Gin Ta- ble 1). Chirostylus sp. Hart, 1982: 166, fig. 65. Type material. —MEXICO: 1 ovig. 2 (17.7 mm), holotype, AHF 861, Jasper seamount off Baja California (30°25.6'N, 122°43.7'W to 30°25.5'N, 122°44.3’W), 950-840 m, rock dredge, 1 Nov 1986, Seatomado Expedition Sta 12; 1 6 (26.5 mm), USNM 234412, Jas- per Seamount (30°25’N, 122°45’W), 600- 800 m, dredge, 20 Oct 1984. CALIFOR- NIA: 1 6 (30.0 mm), 2 ovig. 2 (24.0, 25.5 mm), AHF 787, W of Tanner Bank (32°46'41’"N, 119°59'23”W), 950 m, baited fish traps, 1978, coll. commercial fishing boat; 1 ovig. 2? (25.0 mm), AHF 7720, off (inside) San Clemente Island (approx. 33°N, 118°W), 1189 m, sablefish trap, 25 July 1977, coll. boat Pete Boy. OREGON: 7 6 (11.9-18.5 mm), 3 ovig. 2 (19.8—21.0 mm), 2 2 (15.0, 15.4 mm), USNM 243911, SW of Columbia River mouth (46°02.7'N, 124°57.3'W), 914 m, 72-foot shrimp trawl, VOLUME 103, NUMBER 4 30 May 1964, coll. Commando, U.S. Bu- reau of Commercial Fisheries—AEC Project. Description. —Rostrum nearly one-third to barely one-fourth as long as postorbital carapace length, rostral base laterally ridged, rostral spine curving dorsad. Outer orbital angle rounded. Carapace, excluding ros- trum, |.i-1.2 times as long as its greatest width. Lateral margins strongly convex in posterior two-thirds of length, with distinct constriction at one-third from anterior end, bearing spines as figured (Fig. la, b), an- terolateral spine much pronounced. Gastric region convex, distinctly separated from cardiac region by wide depression (cervical groove), indistinctly from anterior branchi- al region, bearing 6 prominent spines in hexagonal arrangement with another spine in center (left posterior spine of hexagon set well back from level of right posterior one in holotype), occasionally accompanied by few additional smaller spines inside, beside or behind hexagon. Mid-cervical groove slightly anterior to midlength of carapace. Anterior branchial region with 2 convexi- ties bordered by deep groove, each with 1 or 2 pronounced central spines, occasion- ally with small accompanying spine, ante- rior convexity weakly elevated and indis- tinctly separated from gastric region, posterior convexity well elevated and bor- dered by deep posterior groove. Cardiac re- gion indistinctly defined, with pair of prom- inent anterior spines, usually followed by few small or somewhat pronounced spines. Branchial and intestinal spines as figured (Fig. la); elevation along posterior margin of carapace with several erect spines. Pter- ygostomian flap spinulose, anterior end with prominent spine, occasionally accompanied by second smaller spine behind it. First segment of abdomen bearing pos- terior transverse elevation with 5—11 spines (usually 5, 6 or 7, rarely 8, 9 or 11), lacking pleura. Second segment having tergum with low hump-like elevations in transverse line, each elevation with few simple or pointed granules, occasionally lacking granules, 855 pleuron separated by deep groove from ter- gum, tapering, transversely hollowed, bear- ing elevation with 1—5 small dorsal spines along anterolateral margin. Third and fourth segments similar, terga feebly elevated, pleura more weakly tapering than preced- ing; pleuron of third segment with 1 or 2, or, rarely 3 or 4 small posterior marginal spines in addition to terminal one, rarely spineless; that of fourth segment usually ta- pering, occasionally rounded, with 2 or 3 (rarely 1 or 4) posterior marginal spines or without spines. Fifth segment having pleu- ron ending in rounded margin, bearing 1-7 (usually 2 or 3) small spines on posterolat- eral margin, occasionally with 1 or 2 small spines on surface; tergum unarmed. Tergum of sixth segment usually with 3 prominent posterior marginal and 6 dorsal spines, rare- ly with few more spines; pleuron usually spineless, rarely with few (on surface) to 7 or 8 spines (on posterolateral margin). Tel- son divided into anterior and posterior lobes, posterior lobe slightly wider than, and bare- ly twice as long as anterior lobe, fully twice as wide as long. Eyes barely reaching end of rostral spine, cornea dilated and distinctly wider than re- maining eyestalk. Basal segment of antennule simple, spine- less. Antennal peduncle having second seg- ment with small distolateral spine, ultimate segment about 3 times as long as penulti- mate, bearing slender distoventral spine, penultimate segment with tiny distoventral spine; antennal scale rudimentary. Third maxilliped having coxa with slen- der but prominent ventral spine; ischium half as long as merus, with 12-22 (average 16) denticles on mesial ridge; merus with dorsodistal spine; carpus with distinct dis- tolateral spine; propodus shorter than mer- us, with ventral margin somewhat expand- ed on distal portion. Sternite at base of third maxilliped blunt- ly produced on anteromedian margin, usu- ally with pair of small, low spines somewhat posterior to anterior end, rarely with 3 ad- 856 ditional spines: 2 on anterior margin and 1 somewhat posterior to left anterior ventral spine. Next sternite at base of cheliped with 2 pairs of strong spines on anterolateral margins, pair of very tiny, tubercle-like spines between first lateral pair, and another pair of larger spines between posterior limits of posterolateral margins. Chelipeds 5 times as long as carapace, subcylindrical, sparsely provided with long coarse setae. Coxa ventrally bearing well de- veloped distomesial spine and few tuber- cular processes. Merus with 6 regular rows of spines. Carpus equalling length of palm, with lines of spines continued from merus. Paim, exclusive of spines, 7 times longer than wide, 1.3 times as long as movable finger, bearing 7 regular rows of spines: 2 dorsal, 1 mesial, 1 lateral, 1 ventromesial, and 2 ventral. Fingers largely gaping, slen- der, subcylindrical; opposable margins with line of tubercular teeth, prominent truncate basal tooth on movable finger opposed to 2 somewhat smaller ones on fixed finger; me- sial margin of movable finger with few tiny spines proximally. Walking legs slender, laterally com- pressed distally, sparsely provided with stiff setae. First walking leg falling short of end of carpus of cheliped when extended for- ward; coxa with 2 prominent mesial mar- ginal spines near ventral surface; merus about twice as long as carpus, 1.5 times as long as propodus, bearing 6 rows of spines continued onto carpus; propodus with dor- sal, mesial and dorsolateral spines in rows, all small and fixed, dorsolateral ones less numerous; ventral margin with more than 20 movable slender, closely set spinules in- cluding distal pair; dactylus distinctly less than one-third length of propodus, termi- nating in acute curved corneous claw, ven- trally bearing 11 or 12 slender spinules. Sec- ond walking leg similar to first, third walking leg with longer propodus. Diameters of ova measuring 1.7 mm in holotype, 1.1-1.2 mm in paratypes. Male with 2 pairs of gonopods similar to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON those illustrated for G. hawaiiensis Baba, 1977 (see Baba 1977a:fig. 2f, g). Etymology. —The specific name is a noun in apposition from the Greek “‘iaspis,”’ jas- per, alluding to the type locality. Remarks. —From the eastern Pacific Ocean four species of Gastroptychus are known (Baba 1977b:205). The new species resembles G. defensa (Benedict, 1902) from the Galapagos Islands in the arrangement of the gastric spines and the less spinose terga of the abdominal segments. Although the original description of the latter species is brief, the following characters seem unique to G. defensa: 1) the carapace has more prominent spines than in G. iaspis and fewer interspersed small ones on the posterior dorsal surface; 2) the second abdominal seg- ment bears prominent spines on the tergum, and the next three segments bear distinct spines on the pleura; 3) the telson has a longer posterior lobe (if Benedict’s figure 44 is correctly depicted); and 4) the branchial region of the carapace bears less numerous but prominent dorsal spines. In addition, Chace’s notes on G. defensa (see Haig 1968: 276) concerning the sternite at the base of the cheliped that lacks tubercles between the lateral spines, and the chelipeds that have the spination widely spaced and irregular, also seem helpful to separate it from the new species. Examination of two syntypes (one male and one female, USNM 20563) of Gastrop- tychus defensa taken from Albatross Station 2818, northeast of Indefatigable Island, Ga- lapagos Islands, in 717 m (392 fms), showed that most of the above characteristics are true in the male (12.9 mm in postorbital carapace length) upon which Benedict’s de- scription is based. In the female (8.7 mm), however, the carapace, and its posterior half in particular, bears fewer small dorsal spines interspersed among prominent ones; spines on the chelipeds are fewer and widely spaced between spines on rows; the spinules on the abdominal pleura are very reduced in both size and number so as to be discernible only VOLUME 103, NUMBER 4 857 Fig. |. dorsal view; b: same, lateral view. under high magnification; the terga of the third to fifth segments are elevated from levels of pleura, bearing a distinct spine on the lateral extremity; the sixth abdominal tergum is less spinose; width-length ratio of the posterior lobe of the telson is 1.87 in the female syntype, 1.84 in the male syntype, instead of 2.25 in the new species. In both syntypes, the spination of the carapace is much more pronounced than in the new species; in particular, prominent spines on the posterior half of the carapace are ar- ranged in four longitudinal rows (Fig. 3a, b); chelipeds are comparatively slender, and the spines are arranged more irregularly than in the new species; the carapace is relatively long, the length-width ratios being 1.32 in the male syntype, 1.28 in the female syn- type, while 1.18 (average) in the males and 1.13 (average) in the females of the new species. Gastroptychus iaspis, new species, ovigerous female holotype (AHF 861): a, carapace and abdomen, The hexagonal arrangement of the gastric spines is also known in the Indo-West Pa- cific G. hendersoni (Alcock & Anderson, 1899), though a few additional accompa- nying spinules are occasionally present within or directly behind the hexagon (AlI- cock & Anderson 1899: pl. 45: figs. 2, 2a; Baba 1988:14). That species differs from the new species in the very spinose pleura of the abdominal segments. Color. —Hart (1982:166) noted in detail the color of living specimens from British Columbia and Washington. The San Cle- mente specimen (AHF 7720) when fresh, was dark carrot-orange in color (M. K. Wicksten, pers. comm.). Key to the eastern Pacific species of Gastroptychus 1. Abdominal segments very spinose 2 858 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A —S 1mm = (hee 3S Set Cc va = OAT tS A i Yypy OX O MW Fig. 2. Gastroptychus iaspis, new species, ovigerous female holotype (AHF 861): a, sixth abdominal segment and telson, extended, dorsal view; b, right antennal peduncle, ventral view; c, endopod of left third maxilliped, ischium omitted, lateral view; d, anterior part of sternum; e, distal portion of right chela, dorsal view; f, distal segments of right first walking leg, lateral view. — Atleast third and fourth abdominal — Propodus of third maxilliped un- segments nearly spineless ........ 3 armed G. cavimurus Baba, 1977b:202 2. Propodus of third maxilliped with (Off northern Peru and Ecuador, distinct dorsal spines ............ 388-500 m) CUPS CM Sen SRN Act BS G. milneedwardsi 3. Carapace with many small spines (Henderson, 1885:418) interspersed among prominent ones; (Southern Chile, 732 m) mid-cervical groove distinctly an- VOLUME 103, NUMBER 4 859 Fig. 3. Gastroptychus defensa (Benedict, 1902), syntypes (USNM 20563): a, male carapace, dorsal view; b, female carapace, dorsal view; c, sixth abdominal segment and telson of same, extended, dorsal view; d, distal part of right cheliped of same, dorsal view. terior to midlength of carapace ... with prominent spines arranged in eae G. perarmatus (Haig, 1968:272) hexagon with central spine; mid- (Off California, 229 m) cervical groove about at midlength — Carapace with prominent spines and OMCArAPACe 7) chi acme es 4 few interspersed ones, gastric region 4. Terga of third to fifth abdominal 860 segments lacking spines near pleura; sternite at base of cheliped with 2 anterior spines between first lateral marginal spines. G. iaspis, new species (Off northern Mexico to British Columbia, 600-1189 m) — Terga of third to fifth abdominal segments provided with spines at least on lateral extremities near pleura; sternite at base of cheliped lacking spine between first lateral margimalispines..)..68 4.84 oho oe eae G. defensa (Benedict, 1902:299) (Galapagos Islands, 717 m) Acknowledgments We thank A. Genin and A. B. Williams for the opportunity of studying the Jasper Seamount specimens. We also thank Ray- mond B. Manning for loaning type material of Ptychogaster defensa Benedict, and A. B. Williams and two anonymous reviewers for critical reading of the manuscript. Literature Cited Alcock, A., & A. R. S. Anderson. 1899. Illustrations of the Zoology of the Royal Indian Marine Sur- veying Steamer “Investigator.” Crustacea, part 7, pls. 36—45. Calcutta. Baba, K. 1977a. Five new species of chirostylid crus- taceans (Decapoda, Anomura) from off Midway Island.— Bulletin of the National Science Mu- seum, Tokyo, series A (Zoology) 3(3):141-156. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1977b. Gastroptychus cavimurus sp. nov., a new chirostylid (Crustacea, Decapoda, Ano- mura) from off the western coast of South Amer- ica.—Zoologische Mededelingen Uitgegeven door het Rijksmuseum van Natuurlijke Historie te Leiden 52(16):201-207. 1988. Chirostylid and galatheid crustaceans (Decapoda: Anomura) of the “Albatross” Phil- ippine Expedition, 1907—1910.— Researches on Crustacea, The Carcinological Society of Japan, Special Number 2:v + 203 pp. Benedict, J.E. 1902. Descriptions ofa new genus and forty-six new species of crustaceans of the family Galatheidae, with a list of the known marine species. — Proceedings of the United States Na- tional Museum 26:243-334. Haig, J. 1968. First report of the crab family Chi- rostylidae off California, and description of a new species of Chirostylus.—California Fish and Game 54(4):270-277. Hart, J. F.L. 1982. Crabs and their relatives of British Columbia.—British Columbia Provincial Mu- seum Handbook 40:11 + 266 pp., 12 pls. Henderson, J.R. 1885. Diagnoses of the new species of Galatheidea collected during the “Challeng- er’ Expedition.— Annals and Magazine of Nat- ural History (5)16:407-421. Wicksten, M. K. 1982. Crustaceans from baited traps and gill nets offsouthern California. — California Fish and Game 68:244-248. (KB) Faculty of Education, Kumamoto University, Kumamoto, 860 Japan; (JH) Allan Hancock Foundation, University of Southern California, Los Angeles, Califor- nia 90089-0371. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 861-890 NEW SPECIES, CLARIFICATIONS, AND CHANGES IN STATUS WITHIN EOSENTOMON BERLESE (HEXAPODA: PROTURA: EOSENTOMIDAE) FROM THE UNITED STATES Ernest C. Bernard Abstract. —Nine new species of Eosentomon Berlese are described from the United States: seven from South Carolina (E. crypticum, E. hargrovei, E. nu- dilabratum, E. renateae, E. richardi, E. savannahense, E. xenomystax; one from Arkansas (E. osageorum); and one from Michigan (E. snideri). Eosen- tomon brassicae Bernard is synonymized with E. pusillum Ewing, which is redescribed from the holotype. Eosentomon pusillum ewingi Bonet is rede- scribed and raised to specific rank as E. ewingi Bonet. The genus Eosentomon Berlese (Eosen- tomidae), the largest genus among the Pro- tura, now contains more than 180 species. Recently, Dr. R. J. Snider, Michigan State University, sent to me several thousand specimens of Protura collected from the U.S. Department of Energy’s Savannah River Site (SRS) in South Carolina. In this collection were 13 Eosentomon species, of which sev- en are undescribed. In addition, specimens in collections from Arkansas and Michigan were determined to represent other unde- scribed species of Eosentomon. Also, type specimens of E. pusillum Ewing and E. pu- sillum ewingi Bonet were redescribed as a step in the evaluation of the South Carolina material. Materials and Methods Protura were extracted from soil and litter in Tullgren funnels and preserved in 95% ethanol. Specimens were cleared, expanded, and mounted in a polyvinyl alcohol-lacto- phenol medium. Slides were dried over- night in an oven at 60°C. Most drawings were made with the use of a drawing tube on an interference-contrast microscope. Fe- male genital structures were interpreted with brightfield and interference-contrast optics, and relative sclerotization of the parts was estimated with a phase-contrast micro- scope. Head and labral lengths were gen- erally measured in dorsal view. For most species, only females were used for descrip- tions because of difficulties in identifying males and juveniles. All holotypes and some paratypes are deposited in the National Mu- seum of Natural History (NMNH). Re- maining paratypes, except as designated, are in the Apterygote Section of the Uni- versity of Tennessee Entomology Museum (UTEM). Unless otherwise noted, all spec- imens were collected by W. W. Hargrove. Terminology Terms and designations, except as noted below, are used as given in Tuxen (1964). Szeptycki (1984) has been followed for des- ignations of anterior additional head seta (aa), posterior additional head seta (pa), an- terior sensillum (as), posterior sensillum (ps), median subposterior seta (sp), and median posterior seta (p). In this paper, the other paired, median head setae are designated for convention as follows, from anterior to sp: rs (median rostral seta) and cs (clypeal seta) (Copeland 1964), mJ, m2, m3, m4, and m5; rs, cs, m2, m3, m5, sp, and p are considered 862 the innermost setae of transverse rows of setae (Fig. 8). Identification of these rows greatly facilitates the location of aa and as. Seta m4 is present in all the species de- scribed in this paper, but is absent in E. vermiforme Ewing. The form of the galea is similar among Eosentomon spp., with an external spine and three apical projections (Fig. 3). The shape of the projections, here termed digits (outer, median, inner), varies among species and is a very useful supplementary taxonomic character. The mesonotum and metanotum each have certain characters not hitherto used in eosentomid systematics. On each segment, seta p3' may be setiform, lanceolate, clav- iform, or ovate. Mesonotal seta p2' is al- most always setiform but occasionally mimics a modified p3’. On the metanotum, p5 is associated with two minute micro- chaetae near the anterolateral margin (Fig. 1), which for convention may be designated p5' and p5". The bases of seta p5 and p5’ are usually contiguous, although occasion- ally there is discernible space between them, useful as a supplementary character. Ima- date (1989) observed and sketched the lo- cation of these microchaetae in Eosentomon woroae Imadate. Bernard (1975a) noted that, in addition to the 5+5 posterior setae usually reported on abdominal tergite I, there was an addi- tional microchaeta (p3”) near each posterior corner (Fig. 2). The socket of p3” is much less distinct than that of p3’, making it more difficult to detect. All Eosentomon spp. I have examined possess p3”, but Imadate (1989) recently reported that E. woroae Imadate does not possess it. The tergite I setal formula introduced by Copeland (1964) is used in this paper. A formula of 3:1:2 means three normal setae, one delicate seta, and two microchaetae. The microchaetae, sensilla, and setae of eosentomids display a wide variety of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON shapes. Terms that define these shapes are given (Fig. 5); some of these terms have been in use for many years. A “delicate” seta (Tuxen 1964) is one found in the pos- terior rows of the terga, less sclerotized than the normal setae and without a pronounced socket. Inflated and indented setae are un- usual forms of the median rostral setae; in- flated setae are swollen in the basal half, while indented setae suddenly narrow in the middle third. Copeland (1964) has referred to the inflated type as “‘alate.’’ Sensilla are those specialized sensory setae found on the forelegs. Microchaetae are the small sensory setae found on the head, terga, middle legs, and hind legs. Among the more specialized forms, “‘aristate”’ refers to a lanceolate mi- crochaeta with a terminal filament; “‘capit- ulate’’ (small-headed) applies to minutely capitate microchaetae on tergum VIII, and ‘““semmate”’ (bud-like) refers to the minute microchaetae of the metanotum and some- times tergum VIII. The metatarsus possesses, at least in North American species, a constant number and arrangement of setae, some of which are variable in thickness and length. A num- bering system for the metatarsal setae is pro- posed and illustrated in Fig. 4. Six dorsal setae (D1-6) are arranged in an irregular zigzag row; the most variable of these is D2, referred to as the basal seta (bs) by Szeptycki (1984). Since BS is already used for a fore- tarsal ratio, it is desirable to use a different term for this metatarsal seta. D2 varies from a thin seta to a thick spine. D5 is always a distinct spine but its thickness varies. D4 is variably spiniform, while D1, D3, and D6 are setiform. On the exterior surface, El and E2 are setiform, while E3 is spiniform. All nine ventral setae (V1—V9) are rather heavy spines, whose length may vary among spe- cies; V4 is the shortest of the ventral setae. The interior side is considered to be free of setae. In this paper, usually only the view that best shows D2 is illustrated. VOLUME 103, NUMBER 4 Figs. 1-5. 863 Eosentomon vermiforme: 1, metanotum, left side; 2, abdominal tergum I, left side; 3, galea (S: exterior spine; O: outer digit; M: median digit; I: inner digit); 4, metatarsus, dorsal and ventral views (D: dorsal setae; E: exterior setae; V: ventral setae). Eosentomon spp.: 5, Setal types (A—D, setae: A, normal: B, delicate: C, inflated; D, indented. E-J, sensilla: E, linear; F, rounded spatulate; G, pointed spatulate; H, claviform: I, oval; J, oblong. K-Q, microchaetae: K, linear; L, lanceolate; M, aristate; N, oblong; O, capitulate; P, falcate; Q, gemmate). Eosentomon crypticum, new species Figs. 6-17; Table 1 Color and dimensions. —Body weakly sclerotized, yellow-amber posteriorly. Length 795 um; length of head 99 um; length of foretarsus without claw 64 wm. LR = 12.3; PR = 7.0; TR unknown (foretarsal claw broken). Morphology.—Pseudoculus oval with a few longitudinal striae (Fig. 8). Clypeal apo- deme small but robust, with thick anterior bar and a central keel-like appendage ex- tended posteriorly (Fig. 10). Labral margins coarsely scalloped anteriorly, apices round- ed to a very narrow U-shaped notch; labral setae present, extended well past labrum apex (Fig. 10). Rostral setae greatly inflated. Mandible slender with two apical teeth; ga- lea with cylindrical, apically rounded outer 864 digit and slender-conoid middle and inner digits, the middle digit shorter than the in- ner; outer lobe of lacinia slender, slightly curved at apex, not serrate; inner lacinial lobe slender and curved anteriorly, not hooked (Fig. 9). Empodia of middle and hind tarsi short. Central lobe of praecosta sinuate (Fig. 15). Female squama genitalis (Fig. 17) with very thin caput processus bent sharply toward inner stylus margins; corpus processus ex- tending as a narrow lobe anteriorly past the caput, and without prominent alae; median sclerotizations present, rod-like; proximo- lateral sclerotizations very heavy and well- developed. Filum processes short, stylus apex bluntly pointed. Chaetotaxy. —Cephalic seta aa absent, pa present, anterior sensillum present (Fig. 8). Setae sp and p about equal in length. On mesonotum and metanotum (Figs. 12, 13), lengths of p/ and pl’ equal, p2' two- thirds the length of p2, p3 setiform and shorter than p2’; on mesonotum, p5 very short, its base contiguous with that of p5’. On foretarsus (Figs. 6, 7), all sensilla pres- ent; b, a’, b’2, and t narrowly spatulate; b’/ very large and ovate, nearly level with b'2; e and g rounded-spatulate and large; f/ lin- ear, long; d broken near base; t3 robust; sen- silla a, c, f2, and c’ short and slender. Sen- sillum ¢/ closer to a3 than to a3’; BS = 0.81. Metatarsal seta D2 of moderate thickness (Fig. 11). Abdominal setal composition given in Table 1. Tergite I formula 3:1:2, pl’ longer than p/. On tergites II-VI, p/' and p2’ twice the length of p/ and p2; p4' delicate and shorter than p4 (Fig. 14) but becoming pro- gressively longer and stronger on tergites V— VII. On tergite VII, p/’ short and weakly capitulate, p2’ longer than p2. On tergite VIII, pe, pl’ and p!” aristate, the filament short; p2’ clearly capitulate (Fig. 16). Collection data.—Holotype female col- lected from wet sand and liverworts along Road 5, SRS, Barnwell County, South Car- olina, 9 Sep 1983; deposited in NMNH. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Derivation of name. —The latinized Greek word crypticum means “hidden,” and refers both to the small size of the holotype and my failure to find it initially in the thousands of SRS proturans. Diagnosis.—Eosentomon crypticum is distinguished from all other Eosentomon spp. by the large, anteriorly displaced fore- tarsal sensillum b'/. The structure of the corpus processus is reminiscent of the ““transitorium” species group (Tuxen 1964), but the very thin caput processus is dis- tinctive. Eosentomon ewingi Bonet, new status Figs. 18-29; Table 2 Eosentomon pusillum f. ewingi Bonet 1950: 113; Tuxen 1964:112, fig. 131. Ewing (1940), in his description of E. pu- sillum, included three specimens from North Carolina in the type series. Bonet (1950) considered these specimens a distinct form of E. pusillum because they lacked anterior setae on sternite VIII, and designated them as forma ewingi. Tuxen (1964:112) noted the very shrivelled state of the specimens and considered that “It seems advisable. . . to omit this form from the catalogue.” This action, apparently a recommendation to de- clare E. pusillum f. ewingi a forma inqui- renda, has no validity since specimens and a published, albeit very inadequate, descrip- tion exist. With the collection of many undescribed species from South Carolina, it became nec- essary to reassess E. pusillum f. ewingi. A male proturan collected from Great Smoky Mountains National Park is nearly identical to Ewing’s specimens and is used to com- plete the following description, which is based on the specimen designated as lec- totype (Type #65881, USNM) by Tuxen. Characters determined from the male are described in parentheses, following the des- ignation GRSM. Color and dimensions. —Body small and VOLUME 103, NUMBER 4 865 Figs. 6-17. Eosentomon crypticum: 6, Foretarsus, dorsal view; 7, Foretarsus, ventral view; 8, Right side of head, dorsal view; 9, Lacinia, galea, mandible; 10, Labrum and rostral region; 11, Metatarsus, exterior view; 12, Mesonotal margin; 13, Metanotal margin; 14, Posterior corner of tergum IV; 15, Praecosta of tergum VI; 16, Posterior margin of tergum VIII; 17, Squama genitalis. (20 um scale applies to Figs. 6-7, 9-11, 15-17, 25 um scale to all others.) 866 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Abdominal chaetotaxy of Eosentomon crypticum, n. sp. I II-III IV V-VI VII VIII IX-X XI XII 4 10 10 g! 6 6 ae ae ees Se ees = 8 8 9 Bers iia 16 16 16 16 9 4 6 6 6 6 0 2 = o a a Ms 6 8 12 Visa! 4 4 10 10 10 7 1 Seta a3 absent. 2 Setae al, a3 absent. slender, weakly sclerotized, light amber. Length 603 um (GRSM: 868 um); head length about 70 um (GRSM: 82 um); PR approximately 3.6 (GRSM: 4.5); (GRSM: foretarsus length 65 um). LR: (GRSM 9.9); TR: (GRSM 4.4). Morphology. —Pseudoculus extremely large, appearing round or broadly oval with at least three prominent longitudinal striae (Figs. 20, 21). Labrum, mouthparts, and clypeal apodeme all unobservable on lec- totype. (GRSM: Labrum rounded-truncate anteriorly, the lobes angled inward to a small V-shaped notch, labral setae present [Fig. 22]. Median rostral setae inflated. Mandible tridentate with small subapical and two api- cal teeth; on galea, outer digit the longest, inner digit the shortest [Fig. 23]; lacinia not clearly observable. Clypeal apodeme stout, side arms short and heavy.) Empodium of middle leg almost a third the length of the unguis, EUII = 0.29 (GRSM: 0.31); empodium of hind leg long, EU = 0.69 (GRSM: 0.70) (Fig. 24). Central lobe of praecosta slightly incised (Fig. 27). Styli of female squama genitalis presenting slightly different appearances (Fig. 29): caput processus semicircular; thickened rods in corpus processus united to form a Y-shape, the Y spread to the ex- terior edge on the left stylus but in the usual position on the right stylus. Median scler- otizations linear; proximo-lateral scleroti- zations distinct. Filum processus very long and slender; stylus apices narrowly round- ed. Chaetotaxy.—Cephalic seta aa not ob- served on lectotype, pa present (Fig. 20); (GRSM: aa and pa present, anterior sensil- lum present [Fig. 21]). Seta sp 1.6x the length of seta p (GRSM: 1.5). Mesonotal seta p2'’ and p3’ short; on metanotum, seta p3’ longer than p2’, p5 very short and contiguous with p5' (Fig. 25). Foretarsi absent on lectotype. (GRSM: foretarsus [Figs. 18, 19] broad, all sensilla present. Sensilla a’ and b narrowly spatu- late; b'2, fl, and t2 long and slender; /2 ovate; sensillum ¢/ closer to a3 than to a3’, BS = 0.94.) Metatarsal seta D2 a spine of moderate thickness; D5 relatively slender (Fig. 24). Abdominal setal composition given in Table 2. Tergite I formula 2:2:2, p3 delicate (Fig. 26). (GRSM: on tergites II-VI, p/’ and p2' longer than p/ and p2; p4 delicate and slightly longer than p/ on tergite II, delicate and equal to p4 on tergites IIJ—-VII. On ter- gite VII, pl’ short and capitulate, p2 and p2' of equal lengths; on tergite VIII [Fig. 28], pl’ and p!” aristate, p2' capitulate). Material examined.—Lectotype female (USNM Type #65881) collected in peach orchard soil by W. F. Turner, Moore Coun- ty, North Carolina, 8 May 1937; one male collected by E. Bernard from Great Smoky Mountains National Park, North Carolina, Swain County, southern face of Mine Ridge, elev. 817 m, in mixed hemlock-oak forest, deposited in UTEM. Diagnosis. —Eosentomon ewingiis distin- guished from most Eosentomon spp. by the enormous pseudoculus, long metatarsal em- podium, and semicircular caput processus. VOLUME 103, NUMBER 4 867 = a5 -====---02y G re i ™ 27 = Figs. 18-29. Eosentomon ewingi: 18, Foretarsus, exterior view (GRSM); 19, Interior view (GRSM); 20, Left side of head (lectotype); 21, Right side of head (GRSM); 22, Labrum and rostral region (GRSM); 23, Galea and mandible (GRSM); 24, Metatarsus (lectotype); 25, Metanotal margin (lectotype); 26, Posterior setae of tergum I (lectotype); 27, Clypeal apodeme (lectotype); 28, Posterior margin of tergum VIII (GRSM); 29, Squama genitalis (lectotype). (25 wm scale refers to Figs. 20-21, 20 um scale to all others.) 868 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Abdominal chaetotaxy of Eosentomon ewingi Bonet. I T-I IV-VI VII Vill IX-X XI XII 4! 10 10 6 el oe ae 2 8 8 9 Dorsal aD iG ie 9 (9) (4) (6) 6 0 — — — — 6 8 12 Ventral a) (A) iG 7 (12) 1 Setal formulas in parentheses determined from male collected in Great Smoky Mountains National Park. 2 Setae al, a3 absent. It shares these characters with E. brevicor- pusculum Yin (1965, 1982), but lacks an- terior setae on sternite VIII (two setae in E. brevicorpusculum) and has a much longer foretarsal sensillum fl. Eosentomon ewingi does not bear any special relationship to E. pusillum; on the basis of the squama geni- talis, E. ewingi is related to E. yosemitense Ewing, while E. pusillum is allied to E. uda- gawal. Eosentomon hargrovei, new species Figs. 30-40; Table 3 Color and Dimensions. — Body moderate- ly sclerotized, yellowish. Length of fully ex- panded female 1097 um. Length of foretar- sus without claw 64 wm. Length of head without labrum 98 wm. LR = 14.2; PR = 10.6; TR = 5.1. Morphology. —Pseudoculus broadly oval, with obscure linear and circular depressions in the posterior half (Fig. 34). Clypeal apo- deme not visible. Labrum in lateral view short, with labral setae extending past an- terior edge of labrum (Fig. 32). Central pair of rostral setae inflated basally. Mandible (Fig. 32) with two apical and one subapical teeth. Galea of maxilla with slender outer spine; median apical lobe slightly clavate and rounded, two inner apical lobes adja- cent, fused on galeal blade; outer lobe of lacinia short, almost straight, broadly rounded apically, without inner teeth, inner lobe thin and slightly curved, not hooked at apex (Fig. 33). Empodium of foretarsus almost as long as claw, EU = 0.96. Empodia of middle and hind legs very short, less than one-fifth the length of the claws. Central lobe of praecosta incised (Fig. 38). Squama genitalis of female (Fig. 40) short and broad, basal apodeme thick and heavy. Median sclerotization of each stylus weakly developed; caput processus strong, curving and tapering to a narrowly rounded point; corpus processus weakly sclerotized, ex- tending as a lobe to the inner edge of the stylus and with distinct ala; lateral sclero- tizations of perigynium and stylus present; filum short, less than half the length of the stylus; apex of stylus rounded. Chaetotaxy.—Cephalic setae aa and pa absent, anterior sensillum present (Fig. 34). Seta sp about 1.4 the length of seta p. On mesonotum and metanotum, seta pl’ about equal in length to p/; p2’ much short- er than p2; seta p3’ normal, length equal to spiracle width (Figs. 35, 36). On metano- tum, seta p5 as long as p3’, its base close to but not contiguous to p5' (Fig. 36). On foretarsus (Figs. 30, 31), sensilla b’/ and c’ present; sensillum a short and slen- der, b reaching the base of G6, c reaching 3; d reaching a6; fl slender, f2 shorter and broader than f7. Sensilla a’, b'2, and ¢2 nar- rowly spatulate, e and g broadly spatulate. Sensillum ¢t/ closer to a3 than to a3', BS = 0.81; ¢3 long and slender. Metatarsal seta D2 a long, slender spine (Fig. 37). Abdominal setal composition given in Table 3. Tergum I setal formula 3:1:2. Seta p'1 longer than p/ on terga I-VI, very short VOLUME 103, NUMBER 4 869 Figs. 30-40. Eosentomon hargrovei: 30, Foretarsus, exterior view; 31, Foretarsus, interior view; 32, Labral region and mandible; 33, Lacinia and galea; 34, Right side of head, dorsal view; 35, Mesonotal margin; 36, Metanotal margin; 37, Metatarsus; 38, Praecosta of tergum VI; 39, Posterior margin of tergum VIII; 40, Squama genitalis. (20 um scale applies to Figs. 34-36, 10 um scale to all others.) 870 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.—Abdominal chaetotaxy of Eosentomon hargrovei, n. sp. I II-III IV-VI VII VIII IX-X XI XII 4 10 8! q2 6 od eas oe a © 8 8 9 Dorsal 12 16 16 16 9 4 6 6 6 2 = 2 a sue 2 4 8 12 Ventral 4 4 10 10 7 ' Seta a3 absent. 2 Setae a4 absent on right side of holotype. and weakly claviform on tergum VII. Seta p'2 longer than p2 on terga II-VI, equal to p2 on tergum VII. Accessory seta p’4 similar in appearance to but slightly shorter than p4. On tergum VIII, p/” aristate (Fig. 39). Collection data.—Holotype female ex- tracted from deciduous litter along Road F past Road 4, SRS, Aiken County, South Carolina, 18 May 1984, deposited in NMNH. Derivation of name. —This species is named with gratitude after Mr. William W. Hargrove, who collected most of the pro- turan material described in this paper. Diagnosis.—Eosentomon hargrovei can be separated from other members of the genus by the very strong caput processus and well developed corpus process with prominent lateral alae. The squama geni- talis is reminiscent of that in some ‘“‘whee- leri’’ group species (Copeland 1964, Tuxen 1964), but otherwise FE. hargrovei bears no resemblance to them. Eosentomon nudilabratum, new species Figs. 41-53; Table 4 Color and dimensions. —Body moderate- ly sclerotized, yellowish. Mean length of partially expanded adults 792 um (699-892, n = 4). Length of foretarsus without claw 64 um (61-67, n = 4). Length of head with- out labrum 96 um (91-100, nm = 4). LR = 0.17 (0.15-0.20, n = 4), PR = 8.8 (7.59.7, n = 6); TR = 6.1 (5.6-7.1, n = 7). Morphology. —Pseudoculus broadly oval, with a small circular, central depression and several longitudinal striae (Fig. 45). Clypeal apodeme (Fig. 43) stout, the side arms heavy and slightly enlarged posteriorly. Labrum prominent, tapering anteriorly to small, rounded apices and a V-shaped notch; la- bral setae absent (Fig. 43). Median pair of rostral setae not inflated or otherwise mod- ified. Mandible (Fig. 43) with three to five minute teeth along the inner edge. Galea of maxilla with outer spine, tapering outer dig- it and minute median and inner digits, all three digits appearing bluntly pointed; outer lobe of lacinia smoothly curved, not serrate; inner lobe tapering and straight (Fig. 44). Empodium of foretarsus slightly longer than the claw, EU = 1.08 (0.95-1.15, n = 7). Empodia of middle and hind legs very short, less than one-fifth the length of the claws. Central lobe of praecosta trapezoidal, not incised (Fig. 51). Squama genitalis of female (Fig. 53) small, each stylus tapering to a point; median sclerotization strong, capitate proximally; caput processus thin and smoothly curved, shaped like a question mark; corpus processus weakly developed, bifurcate; filum short, about half the length of the stylus. Chaetotaxy. —Cephalic seta aa absent, pa present; anterior sensillum absent (Fig. 45). Seta sp 1-1.3 x the length of seta p. Seta pl’ on mesonotum and metanotum slightly shorter than p/’; seta p3’ short and clavate; on metanotum, seta p5 nearly as long as p2’, its base close to that of p5' (Figs. 47, 48). VOLUME 103, NUMBER 4 871 Figs. 41-53. Eosentomon nudilabratum: 41, Foretarsus, dorsal view; 42, Foretarsus, ventral view; 43, Labrum and rostral region; 44, Galea and lacinia; 45, Right side of head, dorsal view; 46, Metatarsus, subdorsal view; 47, Mesonotal margin; 48, Metanotal margin; 49, Posterior margin of tergum III; 50, Posterior margin of tergum IV; 51, Praecosta of tergum VI; 52, Posterior margin of tergum VIII; 53, Squama genitalis. (25 um scale applies to Figs. 45, 47-50, 20 um scale to all others.) 872 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 4.—Abdominal chaetotaxy of Eosentomon nudilabratum, n. sp. I II-Ill IV-VII nae 4 8! g! ake 12 16 16 Venta x g Or eae 4 4 10 1 Seta a3 absent. On foretarsus (Figs. 41, 42), sensilla b'/ and c’ absent; sensillum a not reaching y2, b long and nearly reaching 87, c reaching 73, d extending to f2; f1 short and linear, f2 oblong; e and g spatulate. Sensilla t2 and b'2 narrowly spatulate; t3 reaching the base of a7; a' long and slender. Sensillum ¢/ close to a3, BS = 0.94 (0.87-1.00, n = 7). Abdominal setal composition given in Table 4. Tergum I setal formula 3:1:2; pl’ almost twice the length of p/. On tergites II-VI, pl' and p2' longer than p/ and p2; p4’ tergites II-III setiform, short (Fig. 49), but on tergites IV-VII, p4’ longer than p4 and delicate (Fig. 50). Seta pl’ short and slightly capitulate on tergum VII. On tergite VIII, pl” minute, oblong (Fig. 52); p2’ pointed, not clubbed. Collection data.—Holotype female and two paratype females extracted from wet oak and Spanish moss litter, SRS near Dun- barton, Barnwell County, South Carolina, 13 Apr 1984; one paratype female, same locality, 18 May 1984. Holotype and one paratype in NMNH, two paratypes in UTEM. Derivation of name. —The specific epithet is a compound Latin word meaning “‘pro- vided with a naked lip,”’ referring to the lack of labral setae. Diagnosis.—Eosentomon nudilabratum is distinguished from all other Eosentomon spp. by the lack of labral setae, mandibles with three to five inner teeth, foretarsal sen- sillum t/ very close to a3, and claviform seta p3’ on the mesonotum and metanotum. The squama genitalis is somewhat similar to that of E. pomari Bernard, which also VIII IX-X XI XII g 8 8 9 9 g 6 8 12 7 lacks labral setae, but possesses median sclerotizations of the styli (absent in E. po- mari). In addition, E. nudilabratum lacks foretarsal sensillum c’ (c’ present in E. po- mari) and has eight anterior setae on ab- dominal terga II-VI (ten in E. pomari). Eosentomon osageorum, new species Figs. 54-66; Table 5 Color and dimensions. — Body well scler- otized, amber brown. Mean length of fully expanded individuals 1332 um (1217-1446, n = 6). Length of foretarsus without claw 103 wm (101-106, n = 9). Length of head without labrum 124 wm (122-127, n = 6). LR = 11.2 (8.9-13.5, n = 6); PR = 10.0 (9.1-12.0, n = 9); TR = 6.4 (5.9-6.6, n = 9). Morphology. —Pseudoculus oval, with several weak longitudinal striae in the an- terior half (Fig. 48). Clypeal apodeme in- distinct. Labrum with truncated apices sloping inward to a U-shaped notch, inner corners with fine, curved striae (Fig. 56); labral setae present and reaching just past labral apices. Median rostral setae slightly inflated basally. Mandible tridentate, with two apical and one subapical teeth (Fig. 56). Digits of galea well-developed and rounded apically, the median and inner digits of equal length; outer lobe of lacinia smoothly curved and tapering, distinctly serrate on the inner edge; margin of inner lobe bent at tip to form finger-like process supporting a thin inner lamina (Fig. 57). Foretarsal empodium shorter than the VOLUME 103, NUMBER 4 claw, EU = 0.87 (0.79-0.84, n = 7). Em- podia of both middle and hind legs long, EU of middle leg 0.59 (0.52-0.67, n = 10), EU of hind leg 0.66 (0.61-0.74, n = 11) (Figs. 59-61). Central lobe of praecosta weakly incised on tergites IV-V, very weak and linear on tergites VI-VII (Figs. 64, 65). Styli of female squama genitalis (Fig. 63) elongate, median sclerotizations present and with small median spurs. Caput processus thin, smoothly curved, and reaching stylus edge, not heavily sclerotized; outer and in- ner edges of corpus process heavily sclero- tized, the outer edge thickened and clavate proximally. Filum processus long and slen- der; tips of styli bluntly pointed. Chaetotaxy.—Cephalic setae aa and pa present, anterior sensillum present (Fig. 58); seta sp 1.2-1.3 x the length of seta p. Seta p/' on meso- and metanotum slight- ly shorter than p/; p3' normal, longer than the width of the spiracles; on metanotum, p5 remote from p5’ (Fig. 62). On foretarsus (Figs. 54, 55), c’ present, b'I absent. Most sensilla long and slender; c, d, fl, and b'2 all about of equal length; a’ nearly reaching d; sensillum e long and slen- der, not spatulate; g weakly spatulate. Sen- sillum t] much closer to a3 than to a3’, BS = 1.09 (1.02-1.12, n = 9). Metatarsal seta D2 a very stout spine, nearly as large as D5 (Fig. 61). Abdominal setal composition given in Table 5. Tergum I setal formula 3:1:2; p/’ 1.5 X the length of p/. Accessory setae p/’ and p2’ longer than p/ and p2 on terga II- VI, p#’ short and delicate but becoming stronger on each succeeding segment. On tergum VII, p/'’ much shorter than p/, rounded but not capitulate at its apex. On tergum VIII, p/’ and pl” aristate, p2’ slen- der, very slightly capitulate apically (Fig. 66). Collection data.—Holotype female and three male, three female, and two maturus junior paratypes collected from swampy soil near Pine Bluff, Jefferson County, Arkansas, 8 Jul 1988, N. Elkassabany and M. A. Muegge, coll. Holotype and four paratypes 873 deposited in NMNH, four paratypes in UTEM. Derivation of name. —The specific epithet is derived from the name ofa Native Amer- ican tribe, the Osage, who inhabited parts of what is now Arkansas. Diagnosis.—Eosentomon osageorum, in its possession of long mesotarsal and meta- tarsal empodia, differs from all Eosentomon spp. except E. saharense Conde and E. adakense Bernard, but differs radically from both in the structure of the corpus proces- sus, which appears similar to that of the Japanese E. asahi Imadate. The combina- tion of linear foretarsal sensillum e and spat- ulate g appears to be unique within the Eos- entomidae; Madagascarentomon Nosek has both sensilla long and setiform. Eosentomon pusillum Ewing Figs. 67-75; Table 6 Eosentomon pusillum Ewing, 1940:527, fig. 13.—Bonet, 1950:111.—Bonet and Tux- en, 1960:281, figs. 37-42.—Tuxen, 1964: 110-112, figs. 128-130.—Tuxen, 1976: 436, fig. 19. Eosentomon brassicae Bernard, 1975b:167— 169, figs. 31-40. New synonymy. This species, known originally from a sin- gle specimen collected in Florida, has been examined several times and until 1976 was considered to have a short hind empodium (Bonet & Tuxen 1960, Tuxen 1964). Tuxen (1976) studied the holotype once again and found it possessed foretarsal sensillum b'/ and a long metatarsal empodium. However, many characters are still not described, ne- cessitating yet another restudy of the ho- lotype. After this reexamination, it is ob- vious that FE. brassicae Bernard from Michigan is a junior synonym of E. pusil- lum; references to E. brassicae below are in parentheses and designated MS (Michigan specimens). Color and dimensions. — Body lightly sclerotized, amber. Length 570 um (MS: 918). Length of head without labrum 87 hm 874 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 54-63. Eosentomon osageorum: 54, Foretarsus, dorsal view; 55, Foretarsus, ventral view; 56, Labrum and rostral region; 57, Galea and lacinia; 58, Right side of head, dorsal view; 59, Apical half of mesotarsus; 60, Apical half of metatarsus, lateral view; 61, Metatarsus, dorsal view; 62, Postero-lateral margin of metanotum; 63, Squama genitalis. (10 um scale refers to Figs. 56, 57, 63; 40 um scale to Fig. 58; 20 um scale to all others.) VOLUME 103, NUMBER 4 875 Table 5.—Abdominal chaetotaxy of Eosentomon osageorum, 0. sp. I I-III IV-VI VII VII IX-X xI x 4 10 10 8! 6 Dorsal — —_— — — = 12 16 16 16 9 : : 2 4 6 6 6 0 Ventral - = — — = 4 4 10 10 7 5 3 i ' Seta a3 absent. (MS:85). Length of foretarsus without claw 55 um (MS:62). LR = 17.6; PR = 6.1 (MS: 5.6). TR = 5.6 (MS:6.4). Morphology. —Pseudoculus very large but obscure in holotype, (MS: broadly oval, three longitudinal striae). Clypeal apodeme stout, side arms hooked inward (Fig. 70). Labrum short, apices broadly rounded, medially with a deep V-shaped notch; labral setae present, extending past labral tip (Fig. 70). Rostral setae not inflated. Mandible with two apical teeth and a minute subapical denticle; galea with slender outer digit, rudimentary mid- dle digit, and short, slender inner digit; out- er lobe of lacinia short, stout, and slightly curved, inner lobe curved inward at apex but not hooked (Fig. 71). Empodium of foretarsus about equal to claw, EU = 1.0 (Tuxen [1964]: 0.9, MS 1.0). Empodium of middle leg less than one- fourth the length of its unguis, EU = 0.22; empodium of hind leg nearly two-thirds the length of its unguis, EU = 0.63 (MS:0.64) (Fig. 72). Central lobe of praecosta very weakly in- cised (Fig. 73). Female squama genitalis (Fig. 75) distorted, the basal apodeme lying in the horizontal plane but the remainder turned sharply upward. Proximo-lateral sclerotizations present; caput processus shaped somewhat like a duck’s head, the apex slightly recurved anteriorly; caput and ala processus appearing to be combined into an S-shape. Median sclerotizations present but thin; filum processus of medium length; stylus apex narrowly rounded. Chaetotaxy.—Cephalic setae aa and pa present, anterior sensillum present (Fig. 69). Seta sp about equal to seta p. Setae p/ and pl' of equal length on mesonotum and metanotum; seta p3’ the same length as the spiracle width; on metanotum, bases of p5 and p5’ contiguous. On foretarsus (Figs. 67, 68) sensillum c’ absent; b, a’, and t2 narrowly spatulate; e and g rounded spatulate, f7 spatulate but turned upward and foreshortened; sensil- lum b/' present; f2 not clearly visible but appearing oblong; t/ very close to a3, BS = 0.75; t3 thin, curving, reaching the level of 66. Metatarsal seta D2 a slender spine, slightly thicker than D4 (Fig. 72). Abdominal setal composition given in Table 6. Setal formula of tergum I 3:1:2; p/’ longer than p/. On terga II-VI, p/’ and p2' longer than p/ and p2; p4’ delicate and shorter than p4. Seta p/” on tergite VIII oblong, minute, p/' setiform but very short; p2' not capitulate (Fig. 74). Tergum XI with only six setae (p2 absent), the middle pair minute (Fig. 75). Diagnosis.—Eosentomon pusillum is similar to the “kumei’’ group of Tuxen (1964) and Imadate (1974), and to E. uda- gawai Imadate in the S-shaped form of the caput processus + ala processus. It differs from all but E. udagawai in the presence of a long metatarsal empodium. Eosentomon pusillum differs from E. udagawai in the presence of labral setae and lack of foretarsal sensillum c’ (in E. udagawai, labral setae absent, sensillum c’ present). In Bernard (1985), E. pusillum will key to brassicae, a species here synonymized with E. pusillum. 876 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 64-75. Eosentomon osageorum: 64, Praecosta, tergum V; 65, Praecosta, tergum VI; 66, Posterior margin of tergum VIII. Eosentomon pusillum, holotype: 67, Foretarsus, interior view; 68, Foretarsus, exterior view; 69, Right side of head, dorsal view; 70, Labrum and rostral region; 71, Mandible, galea, and lacinia; 72, Metatarsus; 73, Praecosta of tergum VI; 74, Posterior margin of tergum VIII; 75, Posterior abdominal segments with extruded squama genitalis (dotted circles: setal bases of sternite XI; setae of abdominal segment XII not shown). (20 um scale refers to Figs. 66, 69; 10 um scale to all others.) VOLUME 103, NUMBER 4 877 Table 6.—Abdominal chaetotaxy of Eosentomon pusillum Ewing. I -II IV-VI Dorsal ae LO uv 12 16 16 Ventral Bs 2 & 4 4 10 ' Setae al, a3 absent. ? Seta p2 absent. Eosentomon renateae, new species Figs 76-86; Table 7 Color and dimensions. —Body moderate- ly sclerotized, yellowish. Mean length of fe- males with various degrees of expansion 884 um (675-1024, n = 3). Length of head with- out labrum 106 wm (97-110, n = 3). Length of foretarsus without claw 67 um (66-68, n = 5). LR = 12.2 (10.7-14.0, n = 3); PR = 10.2 (9.6-10.7, n = 2); TR = 5.1 (4.8-5.9), n= 5). Morphology. —Pseudoculus round, with several very obscure circular depressions in the center (Fig. 80). Clypeal apodeme mod- erately developed, side arms slightly clavate (Fig. 78). Labrum with rounded apices and deep U-shaped notch; labral setae present, extending past anterior margin (Fig. 78). Central pair of rostral setae indented, apical three-fifths of each seta much narrower than the basal part. Mandible with two apical and one subapical teeth; digits of galea rounded, the outer digit curving outward and longer than the other two digits; outer lobe of la- cinia curved and tapering distally and not serrate, inner lobe hooked at apex (Fig. 79). Empodium of foretarsus shorter than un- guis, EU = 0.91 (0.86-0.96, n = 5). Em- podia of middle and hind legs less than one- sixth the lengths of their ungues. Central lobe of praecosta incised (Fig. 84). Female squama genitalis (Fig. 86) with short, wide styli; weak proximo-lateral scleroti- zations present; caput processus thin and shaped like a boomerang, angled toward in- ner stylus margin; corpus processus extend- VII VIII IX-X XI XII 6 16 9 8 67 9 2 = 2 7 6 8 12 ing nearly to inner stylus margin; median sclerotizations very well-developed: filum processus almost as long as stylus; stylus apices tapering to finely rounded tips. Chaetotaxy.—Cephalic setae aa and pa present, anterior sensillum also present (Fig. 80). Seta sp 1.5 the length of seta p. On mesonotum, p2’ and p3’ lanceolate (Fig. 82), p/ and p/' of equal length: meta- notal chaetotaxy similar except p2’ seti- form; bases of p5 and p5’ contiguous (Fig. 83). All sensilla present on foretarsus (Figs. 76, 77), sensilla 6, e, g, t2, a’, and b’2 spat- ulate; a, c, d, fl, b'l, c', and ¢3 linear, but t3 slightly broader than the others; f2 ob- long. Sensillum ¢/ closer to a3 than to a3’, BS = 0.84 Metatarsal seta D2 spiniform, stouter than D4 (Fig. 81). Abdominal setal composition given in Table 7. Tergum I setal formula 3:1:2, p/’ longer than p/. On terga II-VI, p/' and p2’ longer than p/ and p2, p4' about half the length of p4 and less robust. On tergum VII, p/’ short and capitulate, p4' nearly the length of p4 and equally ro- bust. On tergum VIII, pc, pl’, and p/” all aristate, p2’ clearly capitulate (Fig. 85). Collection data.—Holotype female and one paratype female extracted from decay- ing wood, Upper Three Runs Creek and Road F, SRS, Aiken County, South Caro- lina, 17 Aug 1983; one paratype female from mixed pine-deciduous litter, SRS, Barnwell County, Road F near the guard rail, 4 Jan 1984. Holotype and one paratype in NMNH; one paratype in UTEM. Derivation of name.—It is a pleasure to 878 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 76-86. Eosentomon renateae: 76, Foretarsus, dorsal view; 77, Foretarsus, ventral view; 78, Labrum and rostral region; 79, Lacinia, galea, and mandible; 80, right side of head, dorsal view, and pseudoculus; 81, Metatarsus; 82, Posterior margin of mesonotum; 83, Posterior margin of metanotum; 84, Praecosta of tergum VI; 85, Posterior margin of tergum VIII; 86, Squama genitalis. (25 wm scale refers to Figs. 80, 82, 83, 85; 20 um scale refers to all others.) VOLUME 103, NUMBER 4 879 Table 7.—Abdominal chaetotaxy of Eosentomon renateae, n. sp. I Il-Ill IV V-VI Vil Vill IX-X XI XII Dorsal ei NY ae S a 2 8 8 9 12 16 16 16 16 9 4 6 6 6 6 0 Ventral — = — — — = 2 4 4 10 10 10 7 e 5 ie ' Seta a3 absent. 2 Setae al, a2, a3 absent. name this species for Dr. Renate M. Snider, Michigan State University, who has made significant contributions to our knowledge of soil arthropod ecology. Diagnosis.—Eosentomon renateae is similar to E. richardi (this paper) in most chaetotaxic and foretarsal characteristics, but differs primarily in the squama genitalis. In E. renateae, the caput processus is boo- merang-shaped and the median sclerotiza- tions are strong and prominent, whereas in E. richardi the caput processus is shaped like the head of a duck and bent sharply, and the median sclerotizations are thin and rod-like. Also, E. renateae has indented me- dian rostral setae, while those of E. richardi are thin and not modified. The squama gen- italis of E. renateae is similar to that of E. asakawaense Imadate, but lacks the distinct accessory sclerotization described for that species, and possesses prominent median sclerotizations (very thin in E. asaka- waense). Eosentomon richardi, new species Figs. 87-95; Table 8 Color and dimensions. —Body moderate- ly sclerotized, yellow-orange. Mean length 782 wm (699-892, n = 9). Length of fore- tarsus 71 um (69-74, n = 13). Length of head without labrum 103 wm (93-115, n = 9). LR = 10.7 (9.0-11.3, n = 9); PR = 11.0 (9.1=13.3, n = 110; tr = 4.5 (4.1-5.3, n = 13). Morphology. —Pseudoculus round, with an Oval median depression and a few faint, longitudinal striae in the anterior half (Fig. 91). Clypeal apodeme thin, side arms di- vergent. Labrum with truncate apices an- gled inward and terminating medially as a U-shaped notch; labral setae present, ex- tending past tip of labrum (Fig. 89). Central pair of rostral setae not inflated. Mandible with three apical teeth, the middle teeth slightly larger than the others; digits of galea cylindrical and rounded apically, the outer digit about twice the length of the other two; outer lacinial lobe tapering, slightly curved, and without serrations; inner lacinial lobe sharply bent at apex (Fig. 90). Empodium of foretarsus about the same length as the unguis, EU = 0.97 (0.89-1.07, n = 10). Empodia of middle and hind legs less than one-fifth the lengths of their un- gues. Central lobe of praecosta deeply sinuate (Fig. 93). Female squama genitalis (Fig. 95) with caput processus in the shape ofa duck’s head; corpus processus weakly developed, appearing striate in the proximal part, with outlines of the alae processus distinct; me- dian sclerotizations rod-like, filum proces- sus of medium length; stylus apices bluntly pointed. Chaetotaxy. —Cephalic setae aa and pa present, anterior sensillum present; seta sp 1.2-1.5 the length of seta p (Fig. 91). Seta p/' on mesonotum much longer than, and on metanotum slightly longer than, p/; p3' setiform, slightly longer than spiracle width; sockets of p5 and p5’ on metanotum contiguous. 880 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 87-95. Eosentomon richardi: 87, Foretarsus, ventral view; 88, Foretarsus, dorsal view; 89, labrum and rostral region; 90, Mandible, galea, lacinia; 91, Right side of head and pseudoculus; 92, Metatarsus; 93, Praecosta of tergum VI; 94, Posterior margin of tergum VIII; 95, Squama genitalis. (30 um scale applies to Figs. 91, 94; 10 um scale to all others.) VOLUME 103, NUMBER 4 881 Table 8.—Abdominal chaetotaxy of Eosentomon richardi, n. sp. _——— ee ee eee I I-III IV-VI VII VIII IX-X XI XI ——— re aa A ee oD ee ee ee 4 10 8! 4? 6 Dorsal — — — — = 12 16 16 16 9 3 3 A 4 Ventral — 2 ack — Y 6 8 12 4 4 10 10 7 ' Seta a3 absent. 2 Setae al, a2, a3 absent. All sensilla present on foretarsus (Fig. 97, 88); b, c, a’, b'2, t2 and t3 very weakly spat- ulate, e and g broadly spatulate; 2 oblong, unusually large; all other sensilla except t/ linear; sensilla d, a’, b'2, t2, and t3 all about equal in length. Sensillum t¢/ closer to a3 than to a3’, BS = 0.83 (0.76-0.91, n = 13). Metatarsal seta D2 a slender spine, weaker than D4 (Fig. 92). Abdominal setal composition given in Table 8. Tergum I formula 3:1:2, p/’ longer than p/. On terga II-VI, p/’ and p2’ longer than p/ and p2; p4' shorter than p4 but robust. On tergum VII, p/' short and weakly capitulate, p2' shorter than p2. On tergum VIII, pc, pl’, and pl” aristate and rather slender, p2' capitulate (Fig. 94). Collection data.—Holotype female, two male paratypes, and two female paratypes collected from deciduous litter near the Road F guard rail SRS, Aiken County, South Car- olina, 26 Mar 1984; one male and two fe- male paratypes from soil under a log in de- ciduous forest along Road F, SRS, Barnwell County, South Carolina, 18 Jul 1983; one female paratype, decaying wood from Up- per Three Runs Creek at Road F, SRS, Ai- ken County, South Carolina, 17 Aug 1983. Holotype and three paratypes deposited in NMNBH, five paratypes in UTEM. Derivation of name. —This species is named for Dr. Richard J. Snider, Michigan State University, who generously allowed me to examine the many proturans collect- ed from the SRS. Diagnosis.—Eosentomon richardi is one of the few known North American repre- sentatives of the mostly European “transi- torium” group (Tuxen, 1964), species of which are distinguished by the sharply bent and recurved caput processus, resembling a duck’s head. The presence of the two ce- phalic setae, aa and pa, places it among the subset named ‘“‘delicatum” group by Szep- tycki (1985) where it will key to E. zodion Szeptycki. It differs from this and the other “delicatum” group species, and from the North American species E. vermontense Nosek and E. bernardi Nosek (Nosek & Kevan 1984), in the presence of foretarsal sensillum b’1, the presence of eight, rather than ten, anterior setae on abdominal ter- gum IV, and the presence of six, not four, setae on sternites [X—X. Eosentomon savannahense, new species Figs. 96-108; Table 9 Color and dimensions. —Body moderate- ly sclerotized, yellowish. Mean length of fe- males with various degrees of expansion 759 um (603-1036, n = 9). Length of head with- out labrum 82 wm (70-95, n = 9). Length of foretarsus without claw 57 um (48-63, 7 = 17). LR = 15.3 (11.8-18.5, n = 9); PR 6.1 (5.6-6.5, n = 6); TR = 5.8 (5.3-6.4, 7 = 16). Morphology. —Pseudoculus broadly oval, with several faint longitudinal striae in the anterior half and a small, circular central depression (Fig. 101). Clypeal apodeme with thin connecting bar and bulbous side arms (Fig. 98). Labrum small, much wider than ~ ~ 882 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 96-108. Eosentomon savannahense: 96, Foretarsus, dorsal view; 97, Foretarsus, ventral view; 98, Labrum and rostral region; 99, Mandibles; 100, Galea and lacinia; 101, Right side of head, dorsal view; 102, Posterior margin of mesonotum; 103, Posterior margin of metanotum; 104, Metatarsus; 105, Praecosta of tergum VI; 106, Posterior margin of tergum VIII; 107, Variation in squama genitalis. (20 um scale applies to Figs. 101— 103, 15 um scale to all others.) VOLUME 103, NUMBER 4 883 Table 9.—Abdominal chaetotaxy of Eosentomon savannahense, n. sp. I II-II IV-VII Vill Keone XI XI 4 8! 8} 6 Dorsal — — — = 12 16 16 9 : : Z 4 6 6 2 Ventral — = — = 2 4 4 10 7 2 8 i ' Seta a3 absent. long, apices truncated to form a shallow V (Fig. 98); labral setae present. Central pair of rostral setae not inflated. Mandible (Fig. 99) slender, rounded terminally or with an obscure, rounded subapical protuberance. Digits of galea spine-like, the outer digit the longest; outer lacinial lobe slightly curved, not serrate, inner lacinial lobe slender and curved but not bent near apex (Fig. 100). Empodium of foretarsus usually shorter than the unguis, EU = 0.90 (0.85-1.0, n = 14). Empodia of middle and hind tarsi less than one-sixth the lengths of their ungues. Central lobe of praecosta slightly incised (Fig. 105). Female squama genitalis (Figs. 107, 108) with well developed, clubbed or hooked median sclerotizations and weakly sclerotized, semi-circular caput processus; corpus processus reduced except for well- developed alae processus; filum of variable length; stylus apices broadly rounded with a weak to well-developed subterminal spur. Basal apodeme rather thin; proximo-lateral sclerotizations clearly present. Chaetotaxy.—Cephalic setae aa and pa and anterior sensillum all absent (Fig. 101). Seta sp 1.1-1.4 the length of seta p; pos- terior sensillum longer than the posterior setae. On mesonotum and metanotum, setae p/ and p/' of equal lengths; p3’ lanceolate and as long as width of spiracle (Fig. 102). On mesonotum, p2’ similar in shape to p3’; on metanotum p2’ setiform; p5 about equal to p2' and contiguous with p5’ (Fig. 103). On foretarsus (Figs. 96, 97), sensilla b'/ and c’ absent; b, a’, b'2 and ¢2 all pointed- spatulate; lengths of b'2 and ¢2 variable; sen- sillum d long and broad, reaching or by- passing the base of f2; t3 broad, half the length of d; f7 linear. Sensillum ft/ closer to a3 than to a3’, BS = 0.91 (0.83-0.96, n = 17). Metatarsal seta D2 a slender spine, much thinner than D5 (Fig. 104). Abdominal setal composition given in Table 9. Tergum I setal formula 3:1:2; seta pl’ twice the length of p/. On terga II-VI, pl’ and p2’ longer than their primary setae; p4' delicate and longer than p4 on terga II- V, robust and shorter than p4 on terga VI and VII. On tergum VII, p/' two-thirds the length of p/ and capitulate, p2’ longer than p2. On tergum VIII, seta p/’ oblong, p/” gemmate, p2’ short, not noticeably capitu- late (Fig. 106). Collection data.—Holotype female and two paratype females collected from oak lit- ter, Road F near Dunbarton, SRS, Barnwell County, South Carolina, 18 May 1984; one paratype female, rotted wood and decidu- ous litter, same general locality, junction of Road 6 and Road C, 10 Mar 1982; one paratype female, soil cores near Road C, SRS, Barnwell County, 26 Sep 1983; one paratype female, soil cores 1n live oak stand, near Road F— Road B junction, SRS, Barn- well County, 10 Oct 1983; two paratype fe- males, deciduous litter near Road 6 and guard rail, SRS, Barnwell County, 4 Jan 1984; one paratype female, pitfall trap in decid- uous swamp near Road 4 and guard rail, SRS, Aiken County, 2 Mar 1984; one para- type female, wet oak and Spanish moss litter near Dunbarton, SRS, Barnwell County, 13 Apr 1984; one paratype female, oak litter in U.S. Forest Service pine plantation, SRS, 884 Aiken County, 15 May 1984; one paratype female, deciduous litter near Road F guard rail, SRS, Aiken County, 23 Jul 1984. Ho- lotype and four paratypes deposited in NMNBH, the remaining paratypes in UTEM. Derivation of name. —This species is named after the Savannah River, which flows near the type locality. Diagnosis.—Eosentomon savannahense is very similar to E. ewingi, E. pseudoyo- semitense Copeland and White, and E. yo- semitense Ewing in the form of the squama genitalis. The following key will serve to separate the four species: 1. Sternum VII with two anterior se- tae; terga II-VII each with eight an- LCEIOR SCtace st Ree Fes ieee eee ar, 2 — Sternum VIII without anterior se- tae; tergum VII with four or six an- feTlor Setae teas fies AAMC ey Bee 3 2. Foretarsal sensillum 5’/ present; sterna IX—X each with six setae; me- dian rostral setae indented ee gales E. pseudoyosemitense — Foretarsal sensillum b’1 absent; sterna IX—X each with four setae; median rostral setae thin ot ERE IN ATL CARTE My «aes E. savannahense 3. Empodium of metatarsus short; ter- gum VII with four anterior setae . RIGA Lo aE, APE Ed Mane E. yosemitense — Empodium of metatarsus long; ter- gum VII with six anterior setae SSPE ae aa ge eget eapee sense ct Soe E. ewingi Eosentomon snideri, new species Figs. 109-118; Table 10 Eosentomon australicum Womersley of Bernard, 1975b:187, figs. 1-5. The specimens of this new species from southern Michigan were identified origi- nally as E. australicum Wormersley 1939 by Bernard (1975b). After reexamination of the specimens with the aid of interference contrast microscopy, and especially with a reinterpretation of the female genitalia, it is PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON clear they represent a species allied to east Asian taxa such as EF. udagawai Imadate. Color and dimensions. —Body moderate- ly sclerotized, amber-yellow. Mean length of adults 1183 um (940-1362, n= 6). Length of head without labrum 134 um (121-148, n = 6). Length of foretarsus without claw 88 um (83-90, n = 11). LR = 11.0 (10.1- 12.1, n= 6); PR = 11.9 (10.2-12.1, n = 12); TR = 4.8 (4.2-5.2, n = 11). Morphology. —Pseudoculus broadly oval with two small, contiguous, oval median depressions (Fig. 113). Labrum robust, api- ces rounded-truncate, with several fine, lon- gitudinal lines distally; median notch a rounded U-shape; labral setae present, ex- tending past labral apex (Fig. 111). Clypeal apodeme with large, clavate side arms. Ros- tral setae inflated. Mandible with two apical and one minute inner teeth; digits of galea cylindrical and apically rounded, the me- dian and inner digits about two-thirds the length of the outer digit; outer lobe of lacinia tapering and curved inward slightly, not ser- rate on the inner margin; inner lacinial lobe hooked at apex (Fig. 112). Empodium of foretarsus slightly shorter than the unguis, EU = 0.92 (0.85—1.03, n = 11). Empodia of middle and hind legs less than one-fifth the lengths of their ungues. Central lobe of praecosta sinuate (Fig. 116). Processus sternalis of female squama genitalis (Fig. 118) formed into an S-shape by the caput processus and an ala processus, the caput processus shaped like a duck’s head; median sclerotization very broad but weakly sclerotized; filum processus nearly as long as stylus; stylus pointed apically. Chaetotaxy.—Cephalic seta aa and pa present, anterior sensillum present (Fig. 113). Seta sp 1.3—-1.5 x the length of seta p. On mesonotum and metanotum, p3’ se- tiform, slightly longer than spiracle width; pl’ longer than p/; on metanotum, p5 near- ly as long as p3’, bases of p5 and p5' nearly contiguous (Fig. 114). On foretarsus (Figs. 109-110), sensillum b'l absent; sensilla b, f1, and a’ slender and VOLUME 103, NUMBER 4 spatulate, e and g rounded-spatulate; /2 clearly claviform; except for t/, all other sensilla linear. Sensillum d nearly reaching f2; t3 and c' of equal length; a reaching ¥2. Sensillum ¢/ closer to a3’ than to a3, BS = 1.00 (0.93-1.05, nm = 11). Metatarsal seta D2 a robust spine; most of the ventral spi- niform setae very long and slender (Fig. 115). Abdominal setal composition given in Table 10. Tergum I formula 3:1:2, p/’ lon- ger than p/. On terga II-VI, pi’ and p2’ longer than p/ and p2; p4’ a normal seta, two-thirds of the length of p4. On tergum VII, pl’ capitulate, very short, less than one- fifth the length of p2’. Setae pl” and p2’ on tergum VIII aristate, base of pc far from margin (Fig. 117). Collection data.—Holotype female and two paratype females extracted from moss near Monahan Lake, Livingston County, Michigan, 19 May 1974, E. Bernard, coll.; three paratype females, pine forest soil, Kel- logg Forest, Kalamazoo County, Michigan, 23 Jun 1963. Holotype and one paratype deposited in NMNH, three paratypes in UTEM, and one paratype in the Michigan State University Entomology Museum. Derivation of name. —This species is named with pleasure for Dr. Richard J. Snider, world authority on the Collembola and my first mentor for studies on aptery- gote insects. Diagnosis.—Eosentomon snideri is simi- lar to E. kumei Imadate and Yosii from Japan and E. udagawai Imadate from Japan and China (Imadate 1974) but differs in ter- gal chaetotaxy, length of metatarsal empo- dium, and details of the squama genitalis. Eosentomon snideri has a short metatarsal empodium (long in EF. udagawai), ten a-setae on terga IV—V, four a-setae on ter- gum VII, no a-setae on sternite VIII, and four setae on sternites IX—X (in E. kumei, eight a-setae on terga IV—V; in E. udagawai, six a-setae on terga IV—V, two a-setae on sternite VIII, and six setae on sternites [X-— X). The squama genitalis of FE. snideri has a compact “‘S’”’ formed from the caput pro- 885 cessus and ala processus well anterior to the stylus apex, as in FE. udagawai (“‘S” large in E. kumeiand close to stylus apex), possesses median sclerotizations (absent in E. kumei), and has sublateral alae (absent in E. kumei and EF. udagawai). Eosentomon xenomystax, new species Figs. 119-129; Table 11 Color and dimensions. —Body lightly sclerotized except posteriorly, yellowish amber. Mean length of expanded adults 759 um (627-831, n = 4). Length of foretarsus without claw 51 um (49-54, n = 6). Length of head without labrum 81 wm (75-87, n = 4). LR = 8.9 (7.2-10.3, n = 4); PR = 7.4 (6.8-8.0, n = 8); TR = 5.7 (5.4-6.3, n = 6). Morphology. —Pseudoculus nearly round, with median line, a few obscure lines an- teriorly, and two small circular depressions near its center (Fig. 123). Clypeal apodeme thin, side arms slightly swollen. Labrum of a peculiar shape, the sides concave in the distal half and tapering to small, truncated apices and a minute U-shaped cleft; labral setae present but minute, much shorter than the labrum (Fig. 121). Central pair of rostral setae not inflated. Mandible small and bi- dentate, with an apical tooth and an inner retrorse tooth; galea of maxilla with the usu- al outer spine and thin, delicate digits, the outer digit the longest; all three digits fused on the galeal blade; each lacinial lobe curved and evenly tapering, serrations not ob- served on the outer lobe (Fig. 122). Empodium of foretarsus generally slightly shorter than the unguis, EU = 0.9 (0.8-1.0, n = 7). Empodia of middle and hind legs less than one-sixth the lengths of their un- gues. Central lobe of praecosta trapezoidal, not incised (Fig. 126). Female squama genitalis (Fig. 129) with well-developed sclerotiza- tions of the perigynium and stylus; caput process thin but distinct, smoothly curved to the inner stylus edge; corpus processus PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 886 Figs. 109-118. Eosentomon snideri: 109, Foretarsus, dorsal view; 110, Foretarsus, ventral view; 111, Labrum and rostral region; 112, Mandible and head of maxilla; 113, Right side of head, dorsal view; 114, Posterior margin of metanotum; 115, Metatarsus; 116, Praecosta of tergum VI; 117, Posterior margin of tergum VIII; 118, Squama genitalis. (10 um scale applies to Figs. 111, 112, 118; 15 um scale applies to Figs. 109, 110, 115- 117; 20 um scale applies to Fig. 114; 25 wm scale applies to Fig. 113.) VOLUME 103, NUMBER 4 Table 10.—Abdominal chaetotaxy of Eosentomon Snideri, n. sp. I -Ill IV-V VI Dorsal as a Wo oe 12 16 16 16 Ventral 2 © o Bod 4 4 10 10 887 Vil VIII IX-X XI XI 4 6 16 9 : ‘ ‘ 6 0 — = 4 10 7 : os "Seta a3 absent in holotype and four adult paratypes, present in one adult paratype. 2 Setae al, a2, a3 absent. well developed and variably thickened, scalloped on anterior edge, and extending as ala processus to the outer stylus edge; median sclerotizations long and thin, dou- bled in the proximal fourth. Filum proces- sus short, stylus tips rounded. Chaetotaxy. —Cephalic seta aa absent, pa present; anterior sensillum absent (Fig. 123). Seta sp 0.9-1.4x the length of seta p. Seta pl’ on mesonotum and metanotum similar to pJ and of the same length; seta p3' short and asymmetrically claviform with flattened, oblique tip appearing minutely crenulate (Fig. 124). On metanotum, bases of p5 and p5’ close but not contiguous. On foretarsus (Figs. 119, 120), sensillum c’ absent; b'/ present, setiform. Sensillum a nearly reaching 72, a’ reaching 12, b long and thinly spatulate, c reaching a3; t/ closer to a3 than to a3’; BS = 0.94 (0.91-0.96, n = 6); t2 and b’2 of moderate length and pointed-spatulate; d and t3 each long, t3 particularly robust and reaching nearly to the base of the claw. Sensilla e and g spat- ulate; f7 linear; f2 shorter than f/7 and ovate. Metatarsal setae D2, D4, and DS very stout (Fig. 125). Abdominal setal composition given in Table 11. Tergum I setal formula 3:1:2; pl’ almost twice as long as p/. On terga II-VI, pl' and p2' nearly twice the length of p/ and p2; p4' on terga II-III setiform and much shorter than p4 but delicate and longer than p4 on terga IV—VI; on tergum VII p4#’ equal in length to p4, p/' more than half the length of pl and not noticeably capitulate (Fig. 127). On tergum VIII, p/” minute and bluntly pointed, p2' very weakly capitulate (Fig. 128). Collection data.—Holotype female and two paratype females collected from wet oak and Spanish moss litter near Dunbarton, SRS, Barnwell County, South Carolina, 13 Apr 1984; one paratype female, same lo- cation, from oak litter, 18 May 1984. Ho- lotype and one paratype deposited in NMNH; two paratypes deposited in UTEM. Derivation of name. —The specific epithet is a compound Greek word meaning ““strange upper lip,” in reference to the un- usual labrum with very short setae. Diagnosis.—Eosentomon xenomystax possesses many unusual characteristics that separate it from other Eosentomon spp.: la- bral setae very short; foretarsal sensillum 3 long and strong, equal in length to d; seta p3’ on the mesonotum and metanotum clavi- form-truncate; caput processus shaped like a barbless fishhook, median sclerotizations doubled proximally. The Minorcan species E. coiffaiti Conde has very short labral setae but is otherwise dissimilar. The squama genitalis is reminiscent of that of E. pseu- doyosemitense Copeland & White but dif- fers from that species in many characters, including those listed above. Remarks In addition to the seven new species de- scribed herein from the SRS, six other eo- sentomid species were collected from the same area: Eosentomon pallidum Ewing, E. pseudoyosemitense Copeland & White, E. 888 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 119-129. Eosentomon xenomystax: 119, Foretarsus, interior view; 120, Foretarsus, exterior view; 121, Labrum and rostral region; 122, Lacinia, galea, and mandible; 123, Right side of head, dorsal view; 124, Postero- lateral margins of mesonotum and metanotum; 125, Metatarsus; 126, Praecosta of tergum VI; 127, Posterior margin of tergum VII; 128, Posterior margin of tergum VIII; 129, Squama genitalis. (20 um scale applies to Figs. 123-124, 15 um scale to all others.) VOLUME 103, NUMBER 4 ‘ 889 Table 11.—Abdominal chaetotaxy of Eosentomon xenomystax, n. sp. I I-III IV-VII VIII IX-X XI XI 4 8! 8! 6 Dorsal = = = = 12 16 16 9 : 8 z 4 6 6 2 Ventral — =— — = 2 4 4 10 7 j : y ' Seta a3 absent. turneri Bonet, E. vermiforme Ewing, E. wheeleri Silvestri, and Styletoentomon ros- tratum (Ewing). This total of 13 species from the SRS represents a more diverse fauna than hitherto observed in North America, and is a richer assemblage of species for the size of the area than reported either in Eu- rope or Japan, two well-studied regions. Imadate (1974) reported 13 species from all of Japan, and Nakamura (1983a, 1983b) has since decribed two more. Nosek (1973) list- ed 14 species from all of Europe; since then 26 additional species have been described, mostly by Szeptycki (e.g., 1986); the ma- jority of these are very similar forms sep- arated by small setal and sensilla differ- ences. The greater perceived diversity in the SRS may be due partly to very intensive collecting and the discrimination of isolated specimens of very small species (e.g., E. crypticum, E. renateae, E. xenomystax), which relates to the method of extraction. Proturans usually are extracted with Tull- gren-type funnels, which favor the collec- tion of larger, more heavily sclerotized spe- cies (> 1 mm length). All of the new species from the SRS described in this paper have lengths usually much less than one mm, whereas all of the previously described SRS species, except E. pseudoyosemitense, are longer than one mm. Small proturan species tend to inhabit soil, rather than leaf litter and fermentation layers. Since there is no reason to assume that small species are rarer than large species, it is likely that the choice of extraction method is critical in the quan- titative study of Protura. The species described in this paper help clarify geographical relationships among various Eosentomon species groups, which Tuxen (1964) differentiated on the basis of the squama genitalis. The primarily Euro- pean “‘transitorum” group is represented in North America by three species: E. bernardi (Quebec), E. vermontense (Vermont), and E. richardi (South Carolina). The east Asian “kumei” group is represented in North America by E. pusillum (Michigan, Florida) and E. snideri (Michigan). Conversely, the “‘wheeleri’’ group is almost exclusively North American, with two species in western Eu- rope and one in Japan. The ““maya” group, which includes E. ewingi (North Carolina), E. pseudoyosemitense (North Carolina, South Carolina, Tennessee), E. savannah- ense (South Carolina), and E. yosemitense (California), appears restricted to North America, except for E. brevicorpusculum in China. Acknowledgments This research was partially supported by the U.S. Department of Energy’s National Environmental Research Park Program un- der contract DE-AC09-76SROO-819_ be- tween the U.S. Department of Energy and the University of Georgia’s Savannah River Ecology Laboratory. Literature Cited Bernard, E. C. and records of Protura from Michigan.—Great Lakes Entomologist 8:157-181. 1975a. A new genus, six new species, 890 1975b. New species and additional records of Protura from Michigan.—Great Lakes En- tomologist 8:187-195. 1985. Two new species of Protura (Insecta) from North America.—Proceedings of the Bi- ological Society of Washington 98:72-80. Bonet, F. 1950. Descripcion preliminar de especies nuevas del genero Eosentomon (Protura). II. El E. pallidum Ewing y sus especies afines.— Anal- es de la Escuela Nacional de Ciencias Biologicas 6:109-130. —, & S. L. Tuxen. 1960. Reexamination of spe- cies of Protura described by H. E. Ewing. — Pro- ceedings of the United States National Museum 112:265-305. Copeland, T. P. 1964. New species of Protura from Tennessee. — Journal of the Tennessee Academy of Science 39:17-29. : Ewing, H.E. 1940. The Protura of North America. — Annals of the Entomological Society of America 33:495-551. Imadate, G. 1974. Fauna Japonica. Protura (Insecta). Tokyo, Keigaku Publishing Company, Ltd., 351 pp. . 1989. Proturans from Java.—Azao 1:91-118. Nakamura, O. 1983a. A new species of the genus Eosentomon (Protura, Eosentomidae) from Hokkaido, northern Japan.—Kontyu 51:140- 143. 1983b. Eosentomon nupri sp. nov. from Hokkaido (Protura, Eosentomidae).—Kontyu 51:596-600. Nosek, J. 1973. The European Protura. Geneva, Mu- seum L’Histoire Naturelle, 345 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON , & D. K. McE. Kevan. 1984. New species of Eosentomon Berlese, 1909 (Protura: Eosento- midae) from Quebec and Vermont.—Canadian Entomologist 116:19-26. Szeptycki, A. 1984. Three new species of Eosento- mon Berlese, 1909, from Poland with redescrip- tion of Eosentomon germanicum Prell, 1912.— Polskie Pismo Entomologiczne 54:195-213. 1985. Polish Protura. II]. Eosentomon deli- catum Gisin, 1945, and related species.—Pol- skie Pismo Entomologiczne 55:139-186. . 1986. Polish Protura. IV. Eosentomon “tran- sitorium” group.—Polskie Pismo Entomolo- giczne 56:48 1-530. Tuxen, S.L. 1964. The Protura. Paris, Hermann, 360 pp 1976. The Protura (Insecta) of Brazil, espe- cially Amazonas.— Amazoniana 5:417-463. Yin, Wen-ying. 1965. Studies on Chinese Protura I. Ten species of the genus Eosentomon from Nanking-Shanghai regions.—Acta Entomolo- gica Sinica 14:71-92 (in Chinese with English summary). 1982. Studies on Chinese Protura: Twelve species of the genus Eosentomon from Yunnan Province.— Zoological Research 3:11-30 (in Chinese with English summary). Department of Entomology and Plant Pa- thology, University of Tennessee, Knox- ville, Tennessee 37901-1071. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 891-906 TAXONOMIC NOTES ON EPHYDRIDAE (DIPTERA) Wayne N. Mathis and Tadeusz Zatwarnicki Abstract. —Several zoological and nomenclatural items concerning Nearctic Ephydridae, which will be modified in a forthcoming checklist of North Amer- ican Diptera, are presented and explained. Usage of the family-group names Gymnomyzinae and Gymnomyzini is discussed; Athyroglossa (Parathyroglos- sa) dinorata, a new species, is described (= A. ordinata of American authors, not Becker); the synonymy of the genus Gymnomyza Fallén with Mosillus Latreille is explained; the tribe Ochtherini is transferred to the subfamily Gym- nomyzinae where character evidence allies it with the tribe Gymnomyzini; the tribe Lipochaetini is recognized and Nearctic genera assigned; Pelignellus is synonymized with Atissa; Pelignus salinus Cresson is transferred to Schema Becker; Hecamedoides glaucellus unispinosus is given species status (= H. glaucellus of American authors, not Stenhammar); Nesopsilopa Mathis & Wirth is relegated to subgeneric status under Guttipsilopa Wirth; the subfamily Hy- adininae Phillips et al. is given precedence over Gastropinae Cresson by our present action as first revisors; three subspecies in the genus Pelina (bispinosa, latiforma, and prospinosa) are given species status; possible relationships of the tribe Parydrini are discussed; Parydra parasocia Clausen is synonymized with Ephydra fossarum Haliday; Parydra halteralis joaquinensis Clausen is given species status; the tribe Philygriini is diagnosed for the first time; Philygris opposita Loew, 1861 is synonymized with Notiphila puanctatonervosa 1813; Ephydra lata Walker, 1858 is synonymized with Ephydra ripara Fallén, 1813; Ephydra oscitans Walker is discussed; and Ephydra gracilis Packard, 1871 is given precedence over Ephydra cinerea Jones, 1906. While preparing material for a catalog on the dipterous family Ephydridae, more commonly known as shore flies, we noted several zoological and nomenclatural changes that should be explained in greater detail than the format of a catalog normally permits. The changes that pertain to the Ne- arctic fauna are explained here, as that fauna will soon be treated in a checklist of Diptera to be published by the United States De- partment of Agriculture. Some of the prob- lems concern only nomenclature; others en- tail both zoological and nomenclatural matters. Methods. —As a framework for this paper and to be explicit about the classification that results from changes made herein, we have arranged each taxon being treated by its subfamily. If taxa being treated within a subfamily belong to different tribes, then tribes are also cited. The general method- ology used in this study was explained pre- viously (Mathis 1986). The descriptive ter- minology, with the exceptions noted (Mathis 1986), follows that published in the recent Manual of Nearctic Diptera, Vol. 1 (Mc- Alpine 1981). One head and two venational ratios are used commonly in the descrip- tions and are defined here for the conve- nience of the user (all ratios are averages of three specimens, the largest and smallest available and one other). Eye-to-cheek ratio: genal height (imme- diately below eye)/eye height. 892 Costal vein ratio is the straight line dis- tance between the apices of R,,, and R,,;/ distance between the apices of R, and Rj, 3. M vein ratio: straight line distance along vein M between crossveins r-m and dm-cu/ distance apical of crossvein dm-cu. The illustrations of the male terminalia were first drawn in pencil by the second author (TZ) and then inked by Elaine R. S. Hodges. Most specimens we examined are in the National Museum of Natural History, Smithsonian Institution (USNM). Numer- ous others were borrowed from the follow- ing collections: Academy of Natural Sci- ences of Philadelphia (ANSP); Museum of Comparative Zoology, Harvard (MCZ); National Museum of Ireland (NMI); Na- turhistoriska Riksmuseet, Stockholm, Swe- den (NRS); Hope Entomological Collec- tion, Oxford University, England. Subfamily Gymnomyzinae Gymnomyzides Latreille, 1829:535. Type genus: Gymnomyza Fallén, 1810 (= Mo- sillus Latreille, 1804). Tribe Gymnomyzini Gymnomyzides Latreille, 1829:535. See subfamilial listing. Gymnopini Cresson, 1922:326. Type ge- nus: Gymnopa Fallén, 1820 (= Mosillus Latreille, 1804). Remarks. —The family-group names Gymnomyzinae and Gymnomyzini are both based on the genus Gymnomyza Fallen (see treatment of this genus under Mosillus). At the subfamilial and tribal levels, these names are nearly 100 years older and clearly have precedence over Psilopinae (Cresson, 1925) or Gymnopini (Cresson, 1922), the family- group names used at the subfamilial and tribal levels in recent catalogs (Wirth 1965, 1968; Cogan & Wirth 1977; Cogan 1980, 1984; Mathis 1989). We are advocating use of Gymnomyzinae and Gymnomyzini rath- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON er than requesting their suppression because priority will promote stability at these cat- egorical levels. For example, two other fam- ily-group names, Lipochaetinae Becker (1896) and Discocerinini Cresson (1925), are also older than Psilopinae, and they would also have precedence over the latter. Thus to promote stability, we are using Gymnomyzinae and Gymnomyzini, as both are the oldest family-group names in Ephydridae and are unlikely to be replaced with still older names. The priority of Gymnomyzidae does pose a problem, however. As a family-group name, Gymnomyzidae is also older than Ephydridae Zetterstedt (1837). Because Ephydridae has and remains the commonly used name for the family, we have requested that the ICZN use its plenary powers to give precedence to Ephydridae over Gymno- myzidae when the two are considered to be synonyms (Mathis & Zatwarnicki, 1990b). Athyroglossa (Parathyroglossa) dinorata, new species Figs. 1-3 Athyroglossa ordinata of various American authors, not Becker, 1896.— Wirth, 1965: 735 [Nearctic catalog]. Diagnosis. —Small to moderately small shore flies, length 1.65 to 2.50 mm. Head: Two proclinate fronto-orbital se- tae, anterior seta subequal in size to recli- nate seta, posterior seta about '4 length of anterior seta; frons mostly uniformly smooth, shiny; pseudopostocellar setae well developed, about '2 length of ocellar setae; ocelli arranged to form an isosceles triangle, distance between posterior pair greater; an- tenna blackish; arista with 5—8 dorsal rays, length of longer dorsal rays about 73 width of Ist flagellomere; antennal groove bare ventrally to invested with distinctly whitish microtomentum dorsally; face pointed, transversely grooved; parafacials bare of microtomentum; eye-to-cheek ratio 0.45. VOLUME 103, NUMBER 4 893 Figs. 1-3. Male terminalia of Athyroglossa dinorata: 1, Epandrium, cerci, and surstyli, posterior view; 2, Epandrium, cercus, and surstylus, lateral view; 3, Internal male genitalia, lateral view. Thorax: Mesonotum mostly bare, shiny, microtomentum only along posterior mar- gin of scutum and anterior margin of scu- tellum; acrostichal setae conspicuous in 4 rows. Halter blackish. Wing unicolorous, nearly hyaline; venation unicolorous, brownish black; costal vein ratio 0.33; M vein ratio 0.48. Legs concolorous, each with coxae, femora, tibiae, and apical 2-3 tar- someres black, basitarsomeres yellowish to whitish; fore femur unarmed. Abdomen: Dorsum of tergum 1, most of 2, anterior margin of 3, and most of 5 mi- crotomentose, otherwise bare and shiny. Male terminalia (Figs. 1-3) as follows: width of epandrium in posterior view about as wide as high (Fig. 1); surstylus in lateral view roughly conical, ventral apex broadly rounded, bearing a few setulae apicad (Figs. 1, 2); gonite (postgonite) in lateral view cla- vate (Fig. 3); hypandrium in lateral view broadly Y-shaped (Fig. 3). Type material. —The holotype male is la- beled “[USA] COLORADO.Rio GrandeCo. [,]8000’[,]South Fork[,] 20 June 1972[,], W. W. Wirth[,] Malaise trap.” The allotype fe- male and one male paratype bear the same locality label as the holotype. The holotype is double mounted (minute nadel in poly- porus block), is in excellent condition, and is deposited in the USNM. Other specimens examined. -CANADA. ALBERTA. Banff, 20 Aug 1925, O. Bryant (1 6; USNM). Okotoks, Sheep River, 27 Jun 1968, W. W. Wirth (4 4, 22; USNM). NOVA SCOTIA. Truro, 16 Aug 1913, R. Matheson (1 2; USNM). ONTARIO. Hearst (75 mi W), 5 Jul 1954, A. H. Sturtevant (1 2; USNM). UNITED STATES. ARIZONA. Mari- copa Co., Wickenburg, 16-18 May 1950, A. H. Sturtevant (3 6; USNM). CALIFOR- NIA. Monterey Co., Lucia, 28 Jul 1940, A. L. Melander (1 6; USNM). “Toumey Park,” 19 Jun 1935, A. L. Melander (1 ¢; USNM). COLORADO. Gunnison Co., Crested Butte (8300 ft), 9 Jul—-S Aug, 1957, A. H. Stur- tevant (1 6, 1 2; USNM). Rio Grande Co., South Fork (8000 ft, Malaise trap), 20 Jun 1972, W. W. Wirth (2 6, 1 2; USNM). CON- NECTICUT. Litchfield Co., Woodbury, 8 Jun 1931, A. L. Melander (1 2; USNM). IDAHO. Bonner Co., Priest Lake, 1-22 Aug LOGE M920 WAL EL. Melanders (1 ices; 894 USNM); Priest Lake (4-mile camp), Aug 1920, A. L. Melander (1 6; USNM); Priest Lake (Tule Bay), 19 Aug 1919, A. L. Me- lander (2 6; USNM). Latah Co., Moscow, J. M. Aldrich (1 2; USNM); Potlatch (1 4; USNM); Viola (1 6; USNM). INDIANA. Tippecanoe Co., LaFayette, 25 May 1915 (1 ?; USNM). MICHIGAN. Cheboygan Co., 2 Aug 1935, H. B. Hungerford (1 9; USNM). Midland Co., 22 Jun 1953, R. R. Dreisbach (1 2; USNM). MONTANA. Glacier Co., St. Mary River, 2 Aug 1935, A. L. Melander (1 2; USNM). Lake Co., Swan Lake (2.5 mi S), 9 Aug 1972, W. N. Mathis (1 9; USNM). Mineral Co., Upper St. Regis River, 28 Jul 1918, A. L. Melander (2 6; USNM). Sher- idan Co., Medicine Lake, 9 Jun 1969, W. W. Wirth (2 9; USNM). NEW YORK. Gen- esee Co., Portageville, Genesee River, 13 Jun 1963, W. W. Wirth (1 6; USNM). Tompkins Co., Ithaca, 15 Aug 1928, A. L. Melander (1 6; USNM). Wyoming Co., Warsaw, Oatka Creek, 11 Jun 1963, W. W. Wirth (1 6; USNM). Fish Creek Pond, 13 Aug 1941, A. L. Melander (1 °; USNM). OHIO. Ashtabula Co., Pymatuning Lake State Park, 13 Sep 1976, B. A. Steinly (7 °; USNM). Erie Co., Huron River, 17 Jul 1976, B. A. Steinly (1 2; USNM). Lorain Co., Am- herst, Beaver Creek, 24 Aug 1977, B. A. Steinly (2 2; USNM); Mill Hollow C. P., Vermillion River, 22 Sep 1976, B. A. Stein- ly (1 6; USNM). Mercer Co., Grand Lake, Montezuma, 26 May 1977, B. A. Steinly (1 6, 1 2; USNM); St. Marys, Windy Point, Grand Lake, near Montezuma, 11 Oct 1976, B. A. Steinly (1 2; USNM). OREGON. Ben- ton Co., Cary’s Grove, 2 Sep 1974, W. N. Mathis (1 6; USNM); Rock Creek (4 mi SW Philomath), 29 May 1972, W. N. Mathis (1 2; USNM). Curry Co., Gold Beach (30 mi E), 2 Sep 1965, K. Goeden (1 2; USNM). Linn Co., Waterloo, 24 Jul 1974, W. N. Mathis (1 2; USNM). Polk Co., Helmick State Park, 20 Mar 1972, W. N. Mathis (1 2; USNM). UTAH. Duchesne Co., Moun- tain Home (20 mi N), 7 Jul 1968, W. N. Mathis (1 6; USNM). VERMONT. Cale- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON donia Co., Lyndon, 13 Jun 1914 (1 2; USNM). WASHINGTON. Benton Co., Prosser, 4 May 1911 (1 2; USNM). Chelan Co., Lake Chelan, Stehekin, 30 Jul 1919, A. L. Melander (1 6; USNM). Clallam Co., Lake Crescent, Fairholm, 26 Jul 1917, A. L. Me- lander (1 2; USNM). Ferry Co., Keller, 4 Jul 1935, A. L. Melander (1 2; USNM). Grant Co., O’Sullivan Dam, 13 Jul 1968, W. N. Mathis (1 6; USNM). Klickitat Co., Glen- wood, Klickitat River, 27 Jun 1917, A. L. Melander (2 6, 1 2; USNM). Mason Co., Lake Cushman, 22 Jul 1917, A. L. Melander (1 6; USNM); Lilliwaup, 23 Jul 1917, A. L. Melander (1 6; USNM); Potlatch, Hood Ca- nal, 28 Jul 1917, A. L. Melander (1 4; USNM). Spokane Co., Spokane, 24 Jun, J. M. Aldrich (1 6; USNM). Walla Walla Co., Walla Walla, Mill Creek, 2-6 Jul 1922, A. L. Melander (1 6; USNM). Whitman Co., Wawawai, 7 Apr 1954, M. T. James, J. Quist (1 9; USNM). WYOMING. Park Co., Yel- lowstone National Park, Old Faithful, 14 Jul 1923, A. L. Melander (1 2; USNM); Yel- lowstone Lake, 9 Aug 1916, A. L. Melander (1 6; USNM). State unknown: “Pipestone Pass,” 3 Jul 1923, A. L. Melander (1 34; USNM;; this could be a pass associated with Pipestone Canyon, Okenagan Co., Wash- ington). Distribution. —Nearctic: British Colum- bia to Nova Scotia, south to Arizona and Maryland. Remarks.—This species was confused with and usually misidentified as Athyro- glossa ordinata Becker, a species from the Old World. We are revising the species of Athyroglossa from the western Palearctic Region (Mathis & Zatwarnicki, 1990a), and as part of that study, we compared Euro- pean material of A. ordinata with those that were labeled as such from North America. Although very similar, the North American species differs as follows (for comparisons, characters of A. ordinata are cited in paren- thesis): aristal rays comparatively short, length up to 1.5 x width of aristal base (3 x basal aristal width); three to four proclinate VOLUME 103, NUMBER 4 fronto-orbital setae (one proclinate fronto- orbital seta); face in lateral view prominent (face in lateral view shallowly prominent, nearly flat); and structures of male termin- alia (Figs. 1-3): ventral apex of the surstylus rounded in lateral view (somewhat point- ed); gonite with narrow process extended from near middle (extended process not as narrow and arising from posterior end). Genus Mosillus Latreille Mosillus Latreille, 1804:196. Type species: Mosillus arcuatus Latreille, 1805 (= Syr- phus subsultans Fabricius, 1794), subse- quent monotypy, Latreille, 1805:390. Gymnomyza Fallén, 1810:19. Type species: Syrphus subsultans Fabricius, 1794), by present designation. Remarks. —With designation of Syrphus subsultans as the type species of Gymno- myza, this generic name becomes an objec- tive, junior synonym of Mosillus. Gymno- myza is an available name and is the type genus for the oldest family-group name in the family (see ““Remarks” section under our treatment of Gymnomyzinae and Gym- nomyzin1). Tribe Ochtherini Ochtherinae Dahl, 1959:105. Type genus: Ochthera Latreille, 1802. Genus Ochthera Latreille Ochthera Latreille, [1802]:462. Type spe- cies: Musca manicata Fabricius, 1794, subsequent designation, Latreille, 1810: 444. Remarks.—Members of this genus and monobasic tribe are among the most easily recognized shore flies, largely because of their raptorial forelegs and triangular-shaped head. Although easily recognized, the phy- logenetic status of the genus and tribe re- mains unresolved. Most recent authors 895 (Wirth 1965, 1968; Cogan & Wirth 1977; Cogan 1980, 1984) have preferred place- ment of Ochthera in the tribe Hyadinini of the subfamily Parydrinae (= Hyadininae); others (Dahl 1959, Miyagi 1977, Mathis 1989) as a separate subfamily, Ochtherinae, or as a tribe, Ochtherini, within the subfam- ily Parydrinae (= Hyadininae). Our studies indicate that Ochthera is better placed in the subfamily Gymnomyzinae, close to the tribe Gymnomyzini. For the present, we are also recognizing the tribe Ochtherini. The character evidence we have found to sup- port this relationship is as follows: 1. Oral margin deeply emarginate anteri- orly. 2. Clypeus button-shaped and exposed through deeply emarginate oral margin. 3. Arista and branching rays typical of gymnomyzine type (3-4 rays inserted to- ward base). 4. Mesonotum lacking well developed se- tae anterior of transverse suture. 5. Surstyli well developed, posteroventral margin extended under anterior margin of epandrium. Apomorphies that indicate the monophy- ly of the genus and tribe are: 1. Head triangular-shaped from an anterior view. 2. Face in profile with distinct protuber- ance near middle. 3. Eyes relatively large, characteristic of predators. 4. Foreleg raptorial, with coxa and femur greatly enlarged and tibial apex projected as a ventroapical spinelike process. 5. Fronto-orbital setae greatly reduced or lacking. 6. Surstyli undulate ventrally, apices re- curved medially. 7. Gonal arch fused distally with gonites. Tribe Lipochaetini Lipochaetini Becker, 1896:275. Type genus: Lipochaeta Coquillett, 1896. 896 Remarks. —In the most recent catalog of Nearctic Diptera (Wirth 1965), the tribe Li- pochaetini was placed in the subfamily Par- ydrinae (= Hyadininae). We suggest, how- ever, that the character evidence of this tribe indicates a closer relationship to members of the subfamily Gymnomyzinae, especially those of the tribe Atissini, as was indicated by Mathis (1984). We are tentatively leav- ing Lipochaetini and Atissini as distinct tribes, although other studies now in pro- gress may alter this status. Genera of this tribe that occur in the Nearctic Region are the following: Glenanthe Haliday and Li- pochaeta Coquillett. Tribe Atissini Atissini Cresson, 1942:102. Type genus: Atissa Haliday, 1837. Genus Atissa Haliday Atissa Haliday in Curtis, 1837:281. (Pub- lished in synonymy, first used for a taxon by Haliday, 1839:401.) Type species: Ephydra pygmaea Haliday, 1833, mono- typy. Pelignellus Sturtevant & Wheeler, 1954:252. Type species: Pelignellus subnudus Stur- tevant & Wheeler, 1954, original desig- nation. New synonym. Remarks. —Sturtevant & Wheeler (1954: 252) noted that Pelignellus resembled Atissa and Pelignus Cresson (= Schema Becker), and further, that J. E. Collin, to whom they submitted a paratype for examination, rec- ommended that P. subnudus would be“. . . best considered an aberrant Afissa.” To resolve whether Pelignellus should be recognized as separate or a part of Afissa, we re-examined characters, external as well as those of the male terminalia. We found that the position of the interfrontal setae (anterior in Pelignellus, and posterior to me- dian ocellus in Atissa) is of trivial value at the generic level. This observation, more- over, is corroborated by the close similarity PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON in the shape of the male terminalia. Al- though the exact shape differs slightly (see Figs. 4—5 of A. subnuda), the differences are but variations on the theme of Afissa, hence our synonymy. Atissa subnuda (Sturtevant & Wheeler), new combination Figs. 4—5 Pelignellus subnudus Sturtevant & Wheeler, 1954:252.—Wirth, 1965:737 [Nearctic catalog]. Distribution. —California. Remarks.—The structures of the male terminalia are as illustrated in Figs. 4—5. Also see our remarks under the generic syn- onymy. Genus Schema Becker Schema Becker, 1907:302. Type species: Schema minutum Becker, 1907, mono- typy. Pelignus Cresson, 1926:254. Type species: Atissa durrenbergensis Loew, 1864, orig- inal designation. —Cogan, 1984:130 [syn- onymy]. Discussion. —In the recent catalog of Pa- learctic Ephydridae, Cogan (1984) listed Pe- lignus as ajunior synonym of Schema. Until then, Schema and its type species, S. minu- tum, had been obscure and for the most part forgotten names. Now that the status of these taxa and their names have been clarified, we are adhering to that precedent (Cogan 1984) and are here transferring P. salinus Cresson to the genus Schema, which is neu- ter in gender. Schema salinum (Cresson), new combination Pelignus durrenbergensis of Cresson (in part), 1926:254, not Loew. Pelignus salinus Cresson, 1942:109.— Wirth, 1965:737 [Nearctic catalog]. VOLUME 103, NUMBER 4 Figs. 4, 5. Male terminalia of Atissa subnuda: 4, Epandrium, cerci, surstyli, and aedeagus, posterior view; 5, Epandrium, cercus, surstylus, aedeagus, and hypandrium, lateral view. Tribe Discocerinini Discocerini Cresson, 1925:228. Type genus: Discocerina Macquart, 1835. Hecamedoides unispinosus (Collin), new status Figs. 6-8 Discocerina (Hecamedoides) glaucella var. unispinosa Collin, 1943:148. Hecamedoides glaucella var. unispinosa. — Cogan, 1976:83 [generic combination]. Hecamedoides glaucellus of American au- thors, not Stenhammar, 1844. Diagnosis (for comparison, external char- acters of H. glaucellus are cited in paren- thesis).—Length 2.05-2.20 mm (2.75-3.0 mm); eye-to-cheek ratio 3.0—3.2 (2.1-2.4); fore femur with | posteroventral, spinelike seta (5 setae); general coloration brownish gray (light gray); costal vein ratio 1.8—2.1 (2.43.0); aedeagus elongate in dorsal view, slightly narrower anteriorly, rounded api- cally and with cleft in lateral view, generally conical in shape (divided into 2 elongate parts, these united basally); aedeagal apo- deme semicircular in lateral view with dor- sal margin straight (spoonlike, rounded ba- 898 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 6-8. Male terminalia of Hecamedoides unispinosus: 6, Epandrium and cerci, posterior view; 7, Internal genitalia (gonites, aedeagus, and hypandrium), dorsal view; 8, Internal genitalia (gonite, aedeagus, aedeagal apodeme, and hypandrium), lateral view. sally). The following characters of the male terminalia (Figs. 6-8) also distinguish H. unispinosus: anterior margin of hypan- drium with medial cleft (Fig. 7); gonite with 1 ventromedial seta (Figs. 7-8); gonite wide at base, thereafter tapered and elbowed to moderately pointed apex (Figs. 7—8). Type material.—Lectotype male, here designated, is labeled ““Monnow Valley, 10. 7. 09 [10 Jul 1909]/LECTOTYPE Disco- cerina (Hecamedoides) glaucella var. uni- spinosa Collin.” There is also a paralecto- type female that is labeled ““Monnow Valley, 15.8. 34 [15 Aug 1934].”’ The lectotype and paralectotype are preserved in the Hope En- tomological Collection (Oxford). Distribution. —Holarctic. Palearctic: Great Britain. Nearctic: Alberta to Ontario, south VOLUME 103, NUMBER 4 to California, Colorado, and New York; Mexico (Baja California). Remarks. —Study of the appropriate type material reveals that H. glaucellus and H. unispinosus are distinct species and that only the latter occurs in North America. North American specimens were usually misiden- tified as H. glaucellus. Although Collin published this species as a variety of H. glaucellus, it is clear from his paper that his “‘variety” is of subspecific status, and being published before 1961, his varietal name is available at the species lev- ele Tribe Psilopini Psilopini Cresson, 1925:241. Type genus: Psilopa Fallén, 1823. Genus Guttipsilopa Wirth Guttipsilopa Wirth, 1956:9. Type species: Guttispsilopa haydeni Wirth, 1956, orig- inal designation. Nesopsilopa Mathis & Wirth, 1977:64. Type species: Psilopa umbrosa Loew, 1862, original designation. New synonym. Discussion.— Although these two taxa represent monophyletic lineages, they are sister groups and are sufficiently similar that recognition of them as separate genera is unwarranted. Nesopsilopa will be used at the subgeneric level, however. In accor- dance with this change, the following spe- cies names are transferred from Nesopsilopa to Guttipsilopa as new combinations: Gut- tipsilopa (Nesopsilopa) bahamaensis (Math- is & Wirth), G. (N.) caeruleiventris (Loew), G. (N.) stonei (Mathis & Wirth), G. (N.) umbrosa (Loew), and G. (N.) wirthi (Mathis & Freidberg). Subfamily Hyadininae Phillips et al., revised status Hyadinini Phillips et al., in Cresson, 1949: 251. Type genus: Hyadina Haliday, 1837. 899 See “Discussion” under tribe Hyadinini for comments. Tribe Hyadinini Phillips et al. Hyadinini Phillips et al., in Cresson, 1949: 251. Type genus: Hyadina Haliday, 1837. Discussion. —Cresson’s last publication (1949) was assembled and published post- humously by a “Publication Committee” of the American Entomolgoical Society (1949: 225). The committee was headed by Mau- rice E. Phillips, who served as editor for the Society’s publications. Most of the paper was taken directly from Cresson’s notes and a nearly completed manuscript that he was preparing at the time of his death. Where necessary for clarification, the publication committee inserted brief annotations that are clearly indicated by being included with- in brackets. One such annotation, this one rather lengthy, contains the family-group name Hyadinini and an explanation of the name, including a brief but adequate diag- nosis (adequate in the sense of complying with the rules of nomenclature). The name is available, having been published correct- ly, but is attributable to the members of the ‘Publication Committee,” not Cresson. The oldest available family-group names for this subfamily, Gastropinae and Hyadi- ninae, were both published in the same pa- per (Cresson 1949). As first revisors, we elect to use Hyadininae for the subfamilial name because its type genus, Hyadina, is a better known taxon that is also more widespread. Gastrops Williston, the type genus for Gas- tropinae, is known only from the New World. The relationships among the genera and tribes of this subfamily are being studied by Edmiston & Mathis, and it would be pre- mature to cite their results, which are still inconclusive. Pelina bispinosa Clausen, new status Pelina truncatula bispinosa Clausen, 1973: 139. 900 Pelina latiforma Clausen, new status Pelina truncatula latiforma Clausen, 1973: 141. Pelina prospinosa Clausen, new status Pelina truncatula prospinosa Clausen, 1973: 143. Remarks. —Clausen (1973:137) de- scribed these three taxa as subspecies of P. truncatula Loew because of an “. . . appar- ent tendency toward intergradation.” He noted, however, that (p. 137) “All groups show overlaps in distribution” and further, that “. . . two groups are often found in the same collections but in these collections no intergradation can be found.” After publi- cation of Clausen’s revision, more field work, especially in western US, was conducted, and in more than one locality, two or more of these taxa were found occurring together, often collected in the same sweep of an ae- rial net. The available evidence now sug- gests that these taxa represent independent populations that can be consistently distin- guished by characters of the male terminalia and that ought to be recognized as full spe- cies. Tribe Parydrini Parydrini Wirth & Stone, 1956:464. Type genus: Parydra Stenhammar, 1844. Remarks. —This tribe may be the sister group to the subfamily Ephydrinae. If evi- dence corroborates this relationship, the concept of Ephydrinae would need to be broadened to include most genera that were previously placed under Parydrini. Ephyd- rinae would thus include four tribes, viz., Ephydrini, Dagini, Parydrini, and Scatelli- ni. At present, we have not opted to rec- ognize Ephydrinae in this sense (Ephydri- nae + Parydrini), preferring to accumulate additional and more convincing evidence. As a preliminary step toward better reso- lution of these relationships, we have iden- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tified the following characters that may be synapomorphies and that would confirm the monophyly of this lineage and the sister- group relationship: 1. Fronto-orbital setae lateroclinate. These setae are well developed in most but not all species of the subfamily, but even when weakly developed, they are latero- clinate. Elsewhere in the family, these setae are proclinate and/or reclinate. 2. Face broadly projected anteriorly, shieldlike. This condition may have evolved more than once within the Ephydridae, where a projected, shield- like face appears in the Hydrelliinae, 1.e., Paralimna and related genera, and in the Ephydrinae. In both cases, it is probably a synapomorphy. Other shore flies usu- ally have the face slightly convex or flat. 3. Oral opening large, gaping. See number 2. 4. Clypeus broadly developed, commen- surate with the gaping oral opening. 5. Anterior spiracle of larvae protrusile, mostly with 3-7 digitiform papillae (in a few cases with 2-18). For the Nearctic Region the tribe Pary- drini includes: Callinapaea Sturtevant & Wheeler, Eutaenionotum Oldenberg, Par- ydra Stenhammar, and Rhinonapaea Wirth. Parydra (Chaetoapnaea) fossarum Haliday Ephydra fossarum Haliday, 1833:175. Parydra fossarum. — Loew, 1860:32 [gener- ic combination]. Parydra (Chaetoapnaea) parasocia Clau- sen, in Clausen & Cook, 1971:83, new synonym. Type material. —The lectotype female of Ephydra fossarum, here designated, is la- beled “Ireland [Northern Ireland, Down- shire, Holywood; green]/Haliday 20. 2. °82 [NMI’s registration number ]/fossarum [handwritten, apparently by J. E. Collin]/ Named by J. E. Collin/LECTOTYPE 2 Ephydra fossarum Haliday by Mathis & VOLUME 103, NUMBER 4 Zatwarnicki [gender symbol, name and au- thor, designators all handwrittten].”’ The lectotype is in poor condition (pointed to paper triangle), and is deposited in NMI. There are two additional female syntypes (NMI) that bear the same label data as the lectotype except they lack the handwritten, determination label; these two specimens are designated as paralectotypes. The holotype male of Parydra parasocia was collected in Sidney, Fremont County, Iowa, and is deposited in the collection at Iowa State University. Distribution. —Holarctic. Nearctic: Brit- ish Columbia and Northern Territories to Nova Scotia, south to California, Arkansas, and Michigan. Palearctic: Austria, Great Britain, Hungary, Italy, Finland, Germany (FRG), Netherlands, Poland, Sweden. Remarks. —We compared European and North American specimens, especially characters of the male genitalia, and con- sider the specimens to be conspecific. For figures of the male terminalia of P. fossa- rum, please refer to those (fig. 100) in Clau- sen & Cook (1971:132). Parydra (Chaetoapnaea) joaquinensis Clausen, new status Parydra halteralis joaquinensis Clausen, in Clausen & Cook, 1971:79. Remarks. —This taxon was described as a subspecies of P. halteralis (Cresson). Al- though the two “‘subspecies”’ are similar, es- pecially externally, they can be distin- guished by the shape of the aedeagus, and there is no apparent intergradation. The populations of the two taxa are allopatric, and as the only available evidence suggests that they are independent, both are given species status. Philygriini, new tribe Philygriini of authors (Nomen nudum, lack- ing a diagnosis).—Wirth, 1965:745; Wirth, 1968:16.—Cole, 1969:400.—Co- gan & Wirth, 1977:335.—Cogan, 1980: 666; Cogan, 1984:149.—Ferrar, 1987: 169. 901 Diagnosis. —A tribe of Hyadininae that is distinguished from others of this subfamily by the following characters: Head: Ocellar setae weakly developed or lacking; face nar- row, slightly to distinctly convex, most prominent at mid facial height and with re- ceding lower facial margin; eye microsetu- lose; subcranial cavity small to moderately large; clypeus not prominent. Thorax: Pos- terior notopleural seta inserted distinctly dorsad from level of anterior seta; katepi- sternal seta lacking or greatly reduced. Costa extended to vein M. Abdomen: Tergum 4 at most twice length of 5. Male terminalia: Cerci connected anterodorsally with epan- drium; gonites united with hypandrium and bearing long setae anterodorsally. Remarks. — Although Philygriini has been widely used in the literature for the last few decades, it is a nomen nudum, lacking a diagnosis. The North American genera that are included in this tribe are Philygria Sten- hammar and Nostima Coquillett. Lemna- phila Cresson, which was included in Phily- griini (Wirth 1965, and elsewhere, see synonymy listed above), is better placed in the tribe Hydrelliini (subfamily Hydrelli- inae) near the genus Hydrellia Robineau- Desvoidy based on characters of the male terminalia and larvae (Johannsen 1935, Hennig 1943, Lizarralde de Grosso 1978, and Edmiston & Foote, pers. comm.). Philygria punctatonervosa (Fallén) Figs. 9-13 Notiphila punctatonervosa Fallén, 1813:254. Philygria punctatonervosa. —Loew, 1860:25 [generic combination]. Hydrina punctatonervosa. — Becker, 1926:57 [generic combination]. Philygria opposita Loew, 1861:356, new synonym. Diagnosis. —Specimens with short stump veins on veins R,,3;, R4,;, and M. Male terminalia (Figs. 9-13): Cerci semicircular and broad, fused anteroventrally with nar- row epandrium (Figs. 9, 10); anteroventral 902 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 9-13. drium and cercus, lateral view; 11, Gonite, hypandrium, aedeagal apodeme, and aedeagus, lateral view; 12, Aedeagal apodeme and aedeagus, dorsal view; 13, Hypandrium and gonite, ventral view. margin of epandrium with 6-7 long setae (Figs. 9, 10); gonite with 1 anterodorsal long seta and 2 short apical spinulae (Fig. 11); aedeagal apodeme triangular in lateral view (Fig. 11); aedeagus curved upwards, C-shaped, basal part with 2 laterodorsal processes (Figs. 11, 12); hypandrium flat, U-shaped, closely associated with gonites (Fig. 13). Type material.—The lectotype é of No- tiphila punctatonervosa, here designated, is labeled ““N. puncta-to-nervosa 6 [handwrit- ten, apparently by Fallén; species name un- derlined]/355 90[pink; “90” handwritten]/ Riksmuseum Stockholm [green]/LECTO- TYPE ¢ Notiphila punctatonervosa Fallen By Mathis & Zatwarnicki [black subborder; Male terminalia of Philygria punctatonervosa: 9, Epandrium and cerci, posterior view; 10, Epan- gender, name, author and designators hand- written].”’ Six other specimens (16, 22, 37) from the Fallén collection, all in poor con- dition (mouldy, missing structures), are des- ignated as paralectotypes. The lectotype is in poor condition (mouldy, left wing stuck to pin), is pinned directly, and is deposited in the NRS. The lectotype male of Philygria opposita, here designated, is labeled ““Penn./Loew Coll./opposita/Type 11161 [red]/Philygria opposita Lw det. W. Wirth °61.”’ Two par- alectotype males bear labels as follows ““Penn./Loew Coll./Type 11161.” The lec- totype and paralectotypes are deposited in the MCZ. Distribution. —Holarctic. Palearctic: Bel- VOLUME 103, NUMBER 4 gium, Denmark, Germany (GDR), USSR. Nearctic: British Columbia to Quebec, south to California, Illinois, and Maryland. Remarks. — Loew (1861) characterized P. opposita as being very similar to P. punc- tatonervosa, differing mainly by the darker coloration and fewer number of wing spots. The number of wing spots is unstable in populations occurring in the Holarctic Re- gion, and in specimens from both the Pa- learctic and Nearctic Regions there is only very slight variation in the shape of the male terminalia, hence our synonymy. The shape of the male terminalia appears to provide good characters for recognition of species in Philygria. Subfamily Ephydrinae Ephydrinae Zetterstedt, 1837:48. Type ge- nus: Ephydra Fallén, 1810. Ephydra (Ephydra) riparia Fallén Ephydra riparia Fallén, 1813:246. Ephydra lata Walker, 1858:233, new syn- onym. Remarks.—The primary type of EF. lata is apparently lost, but the description is ad- equate to identify this species as being con- specific with E. riparia. Ephydra oscitans Walker Ephydra oscitans Walker, 1858:253, pre- occupied, Walker, 1849:1106. Remarks. —Walker’s 1858 name is a homonym and, therefore, is unavailable, being preoccupied by FE. oscitans Walker, 1849:1106. As the primary type of the spe- cies described in 1858 is apparently lost and unrecognizable from the description, the species will not be renamed. Ephydra (Halephydra) gracilis Packard, revised status Ephydra gracilis Packard, 1871:105.—A\l- drich, 1912:78 [biology, description]. 903 Ephydra cinerea Jones, 1906:159.—Stur- tevant & Wheeler, 1954:168 [distribu- tion, notes, synonymy].— Wirth, 1956:19 [distribution in Bahamas]; 1968:22 [Neo- tropical catalog]. Ephydra (Halephydra) cinerea. —Wirth, 1971:371 [revision].—Simpson, 1976:264 [description and figures of larva and pu- parium]. Remarks. —Our usage of Ephydra gracilis as the senior synonym for this species re- verses the precedent of Sturtevant & Wheel- er (1954:168-169) who cited Ephydra ci- nerea Jones as the valid name for this species and listed Ephydra gracilis as a questionable synonym. Sturtevant & Wheeler’s (1954) argument, in part, is a misleading para- phrase of Aldrich (1912), i.e., that Packard’s description of the “larva [sic, Packard de- scribed the puparium] does not agree with the present species—a point that we can confirm.’’ We reexamined the characters and have concluded that Ephydra gracilis can indeed be recognized. The evidence is as follows. Packard’s description of the pu- parium, although brief, does mention a few salient characters that are unique to this spe- cies (1871:78). The respiratory tube is much longer than that of any other species, “being as long as the body,”’ the body is generally smaller and more slender, and the “feet” (prolegs) are more prominent. In addition, as Packard also noted, this species occurs abundantly around the Great Salt Lake. This combination of characters is unique to and clearly identifies E. gracilis, which is the most abundant species around Great Salt Lake. Aldrich (1912:79), contrary to Sturtevant & Wheeler’s (1954) incorrect paraphrase, did not say that Packard’s description “does not agree with the present species.” Instead, Al- drich (1912) noted the brevity of Packard’s description, which, according to Aldrich, was “scarcely” recognizable and unsatisfac- tory by the omission ofa striking distinction (the basal filaments of the “‘anal’ tube). 904 Nevertheless, Aldrich (1912) went on to conclude that “*. . . it is certain that Packard was describing a strikingly small Ephydra common in Great Salt Lake, and there is but one species [E. gracilis], whether he [Packard] described it well or not.” Acknowledgments Several individuals contributed to this paper by loaning specimens, answering let- ters, and providing space, equipment, and general assistance while visiting museums and studying collections. D. Azuma (ANSP), Charles Vogt (MCZ), James P. O’Connor (NMI), Per Inge Persson (NRS), G. Mc- Gavin (Hope Entomological Collection, Oxford). This draft was also reviewed by F. C. Thompson, W. W. Wirth, James Ed- miston, Philip J. Clausen, and Amnon Freidberg and made substantially better as a result. We wish to acknowledge the special interest and contribution of Curtis W. Sa- brosky, who provided much time, knowl- edge, and many discussions to this paper. Donald Azuma kindly provided informa- tion on the “Publication Committee” of Cresson’s last paper (1949), and Elaine R. S. Hodges skillfully inked the illustrations of the male terminalia. Literature Cited Aldrich, J. M. 1912. The biology of some western species of dipterous genus Ephydra. — Journal of the New York Entomological Society 20:77-99. Becker, T. 1896. Dipterologische Studien IV: Ephyd- ridae.— Berliner Entomologische Zeitschrift 41: 67-195. 1907. Zur Kenntnis der Dipteren von Cen- tral-Asien. I. Cyclorrhapha schizophora holo- metopa und Orthorrhapha brachycera.—An- nuaire du Musée Zoologique de L’Academie Impériale des Sciences de St.-Petersbourg 12: 253-317. . 1926. Ephydridae und Canaceidae: Families 56a, 56b. In E. Lindner, ed., Die Fliegen der Palaarktischen Region. 6(1):1—115, figures 66- 134. Stuttgart. Clausen, P.J. 1973. A revision of the Nearctic species of the genus Pelina (Diptera: Ephydridae).— PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Transactions of the American Entomological Society 99:119-156. Clausen, P. J., & E. F. Cook. 1971. A revision of the Nearctic species of the tribe Parydrini (Diptera: Ephydridae).—Memoirs of the American En- tomological Society 27:1-150. Cogan, B. H. 1976. 67. Ephydridae. Pp. 83-85 in G. S. Kloet and W. D. Hicks, eds., A check list of British insects, 2nd edition. 5:1-139 + ix pp. . 1980. Family Ephydridae. Pp. 655-669 inR. W. Crosskey, ed., Catalogue of the Diptera of the Afrotropical Region. British Museum (Nat- ural History), London, 1437 pp. . 1984. Family Ephydridae. Pp. 126-176 inA. Soos and L. Papp, eds., Catalogue of Palaearctic Diptera. Vol. 10. Elsevier Press, Budapest, 402 pp. Cogan, B. H., & W. W. Wirth. 1977. Family Ephyd- ridae. In M. D. Delfinado and D. E. Hardy, eds., A catalog of the Diptera of the Oriental Region. Vol. III. Suborder Cyclorrhapha (excluding Di- vision Aschiza). University Press Hawaii, Ho- nolulu, 854 pp. Cole, F. R. (with the collaboration of E. I. Schlinger). 1969. The flies of western North America. Univ. Calif. Press, Berkeley and Los Angeles, 673 pp. Collin, J. E. 1943. The British species of Psilopa Fin. and Discocerina Mcq. (Dip., Ephydridae).— The Entomologist’s Monthly Magazine 79:147-152. Cresson, E. T., Jr. 1922. Studies in American Ephyd- ridae (Diptera). III. A revision of the species of Gymnopa and allied genera constituting the sub- family Gymnopinae.— Transactions of the American Entomological Society 47:325—343. . 1925. 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Studies on Scandinavian Ephyd- VOLUME 103, NUMBER 4 ridae (Diptera Brachycera).—Opuscula Ento- mologica, Supplementum 15:1—224. Fallén, C.F. 1810. Specimen Entomologicum novam Diptera disponendi methodum exhibens. Lund, 26 pp. . 1813. Beskrifning Ofver nagra 1 Sverige funna Vattenflugor (Hydromyzides).—Kongliga Ve- tenskaps-Academiens Handlingar, series 3 1813: 240-257. Ferrar, P. 1987. A guide to the breeding habits and immature stages of Diptera Cyclorrhapha.—En- tomolograph 8:1—907. Haliday, A.H. 1833. Catalogue of Diptera occurring about Holywood in Downshire.—The Ento- mological Magazine (London) 1:147-180. 1839. Remarks on the generic distribution of the British Hydromyzidae (Diptera).—An- nals of Natural History 3:217-224, 401-411. Hennig, W. 1943. Ubersicht iiber die bisher bekann- ten Metamorphosestadien der Ephydriden, mit Neubeschreibungen nach dem Material der Deutschen Limnologischen Sundaexpedition. (Diptera: Ephydridae).—Arbeiten iiber mor- phologische und taxonomische Entomologie 10(2-3):105-136. Johannsen, O. A. 1935. Aquatic Diptera. Part II. Orthorrhapha-Brachycera and Cyclorrhapha. — Cornell University Agricultural Experiment Station. Memoir 177, 62 pp., 12 plts. Jones, B. J. 1906. Catalogue of the Ephydridae with bibliography and description of new species. — University of California, Publications in Ento- mology 1:153-198. Latreille, P. A. [1802]. Histoire naturelle, générale et particuliére, des Crustacés et des Insectes. Tome troisiéme. Jn C. S. Sonnini, ed., Histoire natu- relle par Buffon. Paris, 468 pp. 1804. Tableau methodique des Insectes. Jn Tableaux methodiques d’histoire naturelle, pp. 129-200, 5 plts. Paris, 238 pp. 1805. Histoire naturelle, générale et parti- culiére, des Crustacés et des Insectes. Tome qua- torziéme. Jn C. S. Sonnini, ed., Histoire natu- relle par Buffon. Paris, 432 pp. 1810. Considérations générales sur l’ordre naturel des animaux composant les classes des crustacés, des arachnides, et des insectes; avec un tableau méthodique de leurs genres, disposés en familes. Paris, 444 pp. 1829. Suite et fin des Insectes. Jn [Cuvier, G.C.L.F.D.] Le régne animal distribué d’aprés son organisation, pour servir de base a l’histoire naturelle des animaux et d’introduction a l’an- atomie comparée. Ed. 2, Volume 5. Paris, 556 pp. Lizarralde de Grosso, M. 1978. Nuevos aportes al conocimiento del genero Lemnaphila Cresson 905 (Diptera-Ephydridae).— Neotropica 24(71):13- 20. Loew, H. 1860. Neue Beitrage zur Kenntniss der Dip- teren. Siebenter Beitrag.—Programm K. Real- schule zu Meseritz 1860:1-46. 1861. Diptera Americae Septentrionalis In- digena. Centuria prima.—Berliner Entomolo- gische Zeitschrift 5:307-359. Mathis, W. N. 1984. A revision of the genus Asmer- inga Becker (Diptera: Ephydridae).—Israel Journal of Entomology 17[1983]:67-79. . 1986. Studies of Psilopinae (Diptera: Ephyd- ridae), I: A revision of the shore fly genus Pla- copsidella Kertész.—Smithsonian Contribu- tions to Zoology 430, 30 pp. 1989. 66. Family Ephydridae. Pp. 639-649 in N. L. Evenhuis, ed., Catalog of the Diptera of the Australasian and Oceanian regions. E. J. Brill and B. P. Bishop Museum special publi- cation 86, 1155 pages. , & W. W. Wirth. 1977. A new genus of psi- lopine flies (Diptera: Ephydridae) with notes on its relationships. — Proceedings of the Entomo- logical Society of Washington 79:63-74. —, & T. Zatwarnicki. 1990a. A revision of the palearctic species of Athyroglossa (Diptera: Ephydridae).—Transactions of the American Entomological Society 116(1):103-133. —-, & 1990b. Ephydridae Zetterstedt, 1837 (Insecta, Diptera): proposed conserva- tion.—Bulletin of Zoological Nomenclature (in press). McAlpine, J. F. 1981. Morphology and terminolo- gy—adults [chapter]. Pp. 9-63 in J. F. McAlpine etal., eds., Manual of Nearctic Diptera, Volume 1. Agriculture Canada, Research Branch, Mono- graph 27, Hull, Quebec, 674 pp. Miyagi, I. 1977. Ephydridae (Insecta: Diptera). Jn Fauna Japonica. Keigaku Publishing, Tokyo, 113 pp. Packard, A. S., Jr. 1871. On insects inhabiting salt water. No. 2. American Journal of Science and Arts (3)l[whole series 101]:100-110. Simpson, K. W. 1976. The mature larvae and puparia of Ephydra (Halephydra) cinerea Jones and Ephydra (Hydropyrus) hians Say (Diptera: Ephydridae).— Proceedings of the Entomologi- cal Society of Washington 78:263-269. Sturtevant, A. H., & M. R. Wheeler. 1954. Synopses of Nearctic Ephydridae (Diptera).—Transac- tions of the American Entomological Society 79: 151-257. Walker, F. 1849. List of the specimens of dipterous insects in the collection of the British Museum 4:689-1172. 1858. Characters of undescribed Diptera in the collection of W. W. Saunders.—Transac- 906 tions of the Entomological Society of London, new series [= series 2] 4:190-235. Wirth, W. W. 1956. The Ephydridae (Diptera) of the Bahama Islands.—American Museum Novi- tates 1817, 20 pp. . 1965. Family Ephydridae. In A. Stone et al., eds., A catalog of the Diptera of America north of Mexico. U.S. Department of Agriculture, Ag- riculture Handbook, 276, iv + 1696 pp. 1968. 77. Family Ephydridae. In N. Papa- vero, ed., A catalogue of the Diptera of the Americas south of the United States. Departa- mento de Zoologia, Secretaria de Agricultura, Sao Paulo, 43 pp. 1971. The brine flies of the genus Ephydra in North America (Diptera: Ephydridae).— An- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nals of the Entomological Society of America 64:357-377. ——, & A.Stone. 1956. Aquatic Diptera. Pp. 372- 482 in R. L. Usinger, ed., Aquatic insects of California. Univ. Calif. Press, Berkeley, 508 pp. Zetterstedt, J. W. 1837. Conspectus familiarum, ge- nerum et specierum dipterorum, in fauna in- sectorum Lapponica descriptorum.—Isis (Oken’s), 1837:28-67. (WNM) Department of Entomology, NHB 169, Smithsonian Institution, Wash- ington, D.C. 20560; (TZ) Department of Zoology, Academy of Agriculture, ul. Cy- bulskiego 20, Wroclaw, 50-205, Poland. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 907-912 A NEW SPECIES OF PARASPADELLA (CHAETOGNATHA) FROM THE COASTAL WATERS OF JAPAN Jean-Paul Casanova Abstract.— A new species of the benthic chaetognath genus Paraspadella, P. gotoi, is described from Amakusa Island, Japan. It differs from the hitherto known species mainly by the shape of adhesive organs and seminal vesicles and the ventral position of the female genital orifices. During the first workshop on chaeto- gnaths held in the University of Surrey (Great Britain) in September 1988, Dr. Tai- chiro Goto (Mie University, Japan) gave me several specimens of a Paraspadella that looked like P. schizoptera Conant, 1895 but which appeared to him to be different from this species. Indeed, it is a new species and thus named P. gotoi. Paraspadella gotoi, new species Figs. 1-2, Table 1 The holotype and four paratypes are de- posited in the National Science Museum Tokyo (NSMT-Ch. 006 and 007-010 re- spectively). Other paratypes are presented to the Muséum national d’Histoire Natu- relle, Paris (UC 96) and to the National Mu- seum of Natural History, Washington, D.C. (USNM 128300). All were collected in April 1987 and 1988. Description. —Eighteen specimens stud- ied. Body stumpy (Fig. 1), reaching up to 5.9 mm without tail fin. Caudal segment represents 46.5 to 51% of body length. Head bears 8 to 10 very recurved hooks. Anterior teeth only, 4—6 on each side, thin and long, the second innermost being the longer (Fig. 2d). Eyes large, with a large four or three branched pigmented cell (Fig. 2e). Corona ciliata unique and easier to describe with a drawing and photographs (Figs. La, 2f, g, h): outline of the three lengthened parts very irregular, one extending towards the eyes and the two others on both sides of neck. Thick collarette stretching from the neck to the end of caudal segment, covered with numerous sensory tufts. Numerous colored pits (dark grey or brownish yellow) arranged in symmetrical areas on the dorsal and lateral sides from head to tail. Gut sin- uous, of a brick reddish color. Well-marked intestinal diverticula at the level of neck, clearly visible since the body is rather trans- parent. Longitudinal muscles not strongly developed as in other spadellids and lateral fields rather wide. Transverse musculature extends from neck to end of trunk. Ventral ganglion large, occupying about one third of trunk length. Lateral fins divided into two parts, a small one (length ~ 0.7 mm) before the female genital orifice and a considerably larger one, extending from this orifice to seminal ves- icle. Different directions of rays in the small part and in the beginning of the larger one (Fig. 21) seem to indicate that there are in fact two pairs of lateral fins as in Paraspa- della schizoptera. Caudal fin spatulate; all fins completely provided with rays. Two adhesive organs on each side meet and unite on half their length at level of anterior part of the seminal vesicle; finger- like processes which constitute each adhesive organ themselves united so that adhesive organs look like a cockscomb sur- rounding ventrally each seminal vesicle (Fig. 1b). Ovaries reaching the level of neck when 908 Fig. 1. Or A eee Ole: CY Bay OO PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VY ES C8eyi [a Shea 0.5mm 0.5mm 2) Paraspadella gotoi, n. sp.: a, Dorsal view; b, Ventral view; c, Right lateral view, with pigmented areas. In a, sensory tufts on fins are stained with methylene blue. fully mature and filled with about twenty large ova of 0.20—0.22 mm in diameter ar- ranged into two rows (only one row when small). Ovaries connected near their base by a duct surrounding ventrally the intestine as already described in three other species of Paraspadella (Conant 1895, Mawson 1944) and in Heterokrohnia and Archetero- krohnia (Casanova 1985, 1986). Female genital orifices ventral with regard to lateral fins (Fig. 1b, c). Seminal vesicles crooked, their orifice at the top of a stout tube per- pendicular to the axis of tail (Fig. 2c). Comparisons with other species. —Species ~ Fig. 2. Paraspadella gotoi, n. sp.: a, Ventral view of caudal segment showing adhesive organs and seminal vesicles. Arrows indicate the thickenings on the tail fin analogous to the finger-like processes of the adhesive organs; b, Enlargement of the right part of the tail fin showing that the thickened edge is an extension of the adhesive organ; c, Seminal vesicle (rays of the anterior part of the adhesive organ laying above the vesicle are visible); d, Teeth and tips of hooks; e, Right eye; f, Dorsal view of the anterior part of a mature specimen (black VOLUME 103, NUMBER 4 909 area in the trunk = ovaries) showing the corona ciliata (arrows); g, Enlargement of the corona ciliata; h, Central part of the corona ciliata of another specimen; i, Anterior and posterior fins with rays in different directions. AF = anterior fins, AO = adhesive organs, E = eyes, PF = posterior fins, SV = seminal vesicles. Scale bars: 0.25 mm (a, f, i), 0.05 mm (b, c, d, g, h) and 0.02 mm (e). 910 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.— Diagnostic characters of the two species of Paraspadella with two pairs of adhesive organs. Characteristics Maximal length 6.5 mm Tail segment 44-45% Intestinal diverticula absent Anterior teeth 3 Posterior teeth 0 Hooks 11 Eyes small Corona ciliata line Lateral fins angular, well separated Position of female genital orifices Seminal vesicles Adhesive organs fins small, crescent shaped of the genus Spadella s. 1. have been sepa- rated into two groups by Tokioka & Pa- thansali (1964), and into three other ones by Alvarino (1981a). The latter three were raised to the rank of genus by Salvini-Plaw- en (1986): Spadella s. s. for the species of the cephaloptera group, Gephyrospadella for those of the schizoptera group with one pair of lateral fins, and Paraspadella for those of the schizoptera group with two pairs of lat- eral fins. Finally, Bowman & Bieri (1989) combined the last two groups because they did not recognize the existence of two sep- arate pairs of lateral fins, thus minimizing the importance of that characteristic. This proposal agrees with Tokioka and Pathan- sali’s groups, making two genera: Spadella and Paraspadella, which differ by the ab- sence or presence of adhesive organs. There are ten known species of Paraspa- della if, as it is very probable, P. humme- lincki Alvarino, 1970 is a junior synonym of P. pulchella Owre, 1963 (Owre 1973). The following species have only one pair of adhesive organs with finger-like processes: P. schizoptera Conant, 1895; P. johnstoni Mawson, 1944; P. pulchella and P. nana Owre, 1963; P. legazpichessi Alvarino, 1981b; P. pimukatharos Alvarino, 1987; P. Paraspadella sheardi Mawson, 1944 three cornered shape with regular out- anterior and posterior fins almost rect- lateral, between anterior and posterior anterior and posterior separated, each with 10-11 free processes Paraspadella gotoi, n. sp. 5.9 mm 46.5-51% well developed 4-6 0 8-10 large three lengthened, irregularly shaped pro- cesses anterior fins triangular and posterior ones roundish, in close contact ventral, below the junction of anterior and posterior fins very large, crooked anterior and posterior more or less unit- ed, cockscomb shaped coecafera Salvini-Plawen, 1986; and P. an- ops Bowman & Bieri, 1989. Only one spe- cies, P. sheardi Mawson, 1944, has also two pairs of adhesive organs, but they differ from those of P. gotoi since they are well sepa- rated and consist of numerous finger-like processes. The principal characteristics of these spe- cies of Paraspadella with two pairs of ad- hesive organs are summarized in Table 1. Remarks.—P. gotoi is the most highly colored chaetognath. The numerous colored pits on the body are reminiscent of Archete- rokrohnia rubra Casanova, 1986 in which they are localized dorsally in the transverse septum region (unpublished data). The col- or of its gut also is similar to the bathyal A. rubra that is rather unusual in a superficial form, but it must be noted also that orange- brown spots of pigment have been described on the body of Spadella angulata (TYokioka & Bieri, 1966) and clusters of reddish cells in the walls of the intestine of Spadella (= Paraspadella) pulchella (Owre 1963), which are both neritic species. A detailed observation of the adhesive organs of P. gotoi shows that they probably have the same origin as fins, the former be- ing modified parts of the latter. Indeed, in VOLUME 103, NUMBER 4 911 Reihoku AMAKUSA Fig. 3. Paraspadella gotoi, n. sp.: Sampling station in Japan (type locality). one specimen, one finger-like process of the right posterior adhesive organ is fused with the tail fin (Fig. 2a, b). In many others, the posterior edges of both lateral and caudal fins bear tiny papillae as those observed on the extremity of adhesive organs. Lastly, when the collarette tissue is stripped off, these organs appear to be rayed as fins. This is contrary to Mawson’s (1944:330) opinion when she wrote: “‘They [finger-like process- es] are therefore not to be regarded as part of the fin,” but agrees with previous opin- ions (Conant 1895, Yosii & Tokioka 1939). Some morphological features of P. gotoi are unique and can be explained by the mat- ing behavior described by Goto & Yoshida (1985). There is cross-fertilization and, when mating, the two partners stand face-to-face. Then, the one that acts as a male jumps and deposits a sperm cluster on the genital or- ifice of the other, which acts as a female. Thus this orifice is ventral and cannot be dorsal as in all the other chaetognaths but one, P. sheardi, where it is exactly lateral, and the particular shape of the seminal ves- icles can help to precisely deposit the sperm. Likewise, the two adhesive organs on each side (one directed forward and the second backwards) allow perhaps the complete erection of the body when mating. In P. sheardi, the other species of the genus with two pairs of adhesive organs, the outspread adhesive processes, according to Mawson (1944), serve as “‘props”’ to support the body in an almost vertical position. This fact, as with also the lateral opening of oviducts, seems to indicate that the mating behavior is the same as in P. gotoi. Distribution.—Two species of Paraspa- della are known from the coasts of Japan: Paraspadella coecafera was described as Spadella schizoptera by Yosii & Tokioka (1939) from a single specimen caught near Misaki, not far from Tokyo, but differs from Conant’s description of Spadella (= Para- spadella) schizoptera by the absence of an- terior fins and the presence of intestinal diverticula. These reasons recently led Sal- 912 vini-Plawen (1986) to recognize its specific status. Paraspadella gotoi, known as S. schi- zoptera in papers dealing with ultrastruc- tural studies, has been collected in tide pools (type locality) near the Amakusa Marine Biological Laboratory, Amakusa Island, Reihoku, in Kytisht, southwestern Japan (Fig. 3). Acknowledgments I wish to express my sincere thanks to Dr. Taichiro Goto of the Mie University in Ja- pan who gave me the specimens to describe this new species. Literature Cited Alvarino, A. 1970. Anew species of Spadella (benthic Chaetognatha).—Studies on the Fauna of Cu- racao and Other Caribbean Islands 34:73-89. 1981la. Los quetognatos benticos, estudios y distribucion.—Mem VIIIe Latino-American Congress of Zoology, Merida (Venezuela, Nov. 1980) 2:1109-1128. . 1981b. Spadella legazpichessi, a new benthic chaetognath from Enewetak, Marshall Is- lands. — Proceedings of the Biological Society of Washington 94:107-121. . 1987. Spadella pimukatharos, a new benthic chaetognath from Santa Catalina Island, Cali- fornia.— Proceedings of the Biological Society of Washington 100:125-133. Bowman, T.E., & R. Bieri. 1989. Paraspadella anops, new species, from Sagittarius cave, Grand Ba- hama Island, the second troglobitic chaeto- gnath. — Proceedings of the Biological Society of Washington 102:586-589. Casanova, J.-P. 1985. Description de l’appareil gén- ital primitif du genre Heterokrohnia et nouvelle classification des chaetognathes.—Compte-ren- dus hebdomadaires des Séances de l’Académie des Sciences, Paris t.301, III(8):397—402. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1986. Archeterokrohnia rubra n. gen., n. sp., nouveau chaetognathe abyssal de 1|’Atlantique nord-africain: description et position systéma- tique, hypothése phylogénétique.— Bulletin du Muséum national d’Histoire naturelle, Paris 4é série, 8, section A, (1):185-194. Conant, F.S. 1895. Description of two new chaeto- gnaths (Spadella schizoptera and Sagitta hispi- da).— Annals and Magazine of Natural History, Series 6, 16:288—292. Goto, T., & M. Yoshida. 1985. The mating sequence of the benthic arrow-worm Spadella schizop- tera. — Biological Bulletin 169:328—333. Mawson, P. M. 1944. Some species of the chaeto- gnath genus Spadella from New South Wales. — Transactions of the Royal Society of South Aus- tralia 68:327-333. Owre, H. B. 1963. The genus Spadella (Chaetogna- tha) in the western North Atlantic Ocean, with descriptions of two new species.—Bulletin of Marine Science of the Gulf and Caribbean 13: 378-390. . 1973. A new chaetognath genus and species, with remarks on the taxonomy and distribution of others. — Bulletin of Marine Science 23(4):948— 963. Salvini-Plawen, L. Von. 1986. Systematic notes on Spadella and on the Chaetognatha in general. — Zeitschrift fiir Zoologische Systematik und Evo- lutionsforschung 24(2):122-128. Tokioka, T., & D. Pathansali. 1964. Spadella cepha- loptera forma angulata raised to the rank of spe- cies.— Publication of the Seto Marine Biological Laboratory 12:145-148. —, & R. Bieri. 1966. The colour pattern of Spa- della angulata Tokioka.— Publication of the Seto Marine Biological Laboratory 14:323-326. Yosii, N., & T. Tokioka. 1939. Notes on Japanese Spadella (Chaetognatha).— Annotationes Zoo- logicae Japonenses 18:267-273. Laboratoire de Biologie animale (Planc- ton), Université de Provence, 13331 Mar- seille Cedex 3, France. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 913-921 A NEW COMBINATION AND SYNONYMY FOR TWO SUBSPECIES OF CUCUMARIA FISHERI WELLS (ECHINODERMATA: HOLOTHUROIDEA) Philip Lambert Abstract.—Cucumaria fisheri fisheri Wells is a junior subjective synonym of Cucumaria piperata (Stimpson). Cucumaria fisheri astigmata Wells is referred to the genus Pseudocnus Panning. The new combination Pseudocnus astigmatus (Wells) is redescribed here. The taxonomy of Cucumariidae from the west coast of North America needs revision. This paper addresses one taxonomic prob- lem as part of a larger study into the sys- tematics of a group of brooding holothu- roids, to be reported on later. Wells (1924) described Cucumaria fisheri (later in the same paper he referred to it as C. f, forma fisheri) as yellow with brown spots, and with podia in double rows. C. f. astigmata was described as orange yellow, without spots, and having scattered podia. The obvious differences in podia arrangement led me to question whether the two forms were con- specific. In this paper I shall assume “‘for- ma” to be equivalent to subspecies (Int. Trust Zool. Nomen., 1985, Section 45(g)). My examination of the holotypes (Figs. 1, 2) confirmed that the external characters are as Wells originally described. He stated that the ossicles of C. f, astigmata were“... identical with those of Cucumaria fisheri, forma fisheri” (Wells, 1924:118). I sampled skin ossicles from the holotypes and found them to be substantially different. Skin os- sicles from the holotype of C. f astigmata (USNM E01196) are thick, knobbed, per- forated buttons or plates (Fig. 3). A few are pine-cone shaped with a spiny handle-like extension at one end. They appear to match the ossicles illustrated by Wells (1924, fig. 1). Ossicles from the holotype of C. f. fisheri Wells (USNM E01198) are, on the other hand, relatively thin, perforated plates with serrated edges. Most are oval with a handle- like extension and pointed bumps on both surfaces (Fig. 4). The ossicles from these two forms are undoubtedly from distinct species rather than from two conspecifics as sug- gested by Wells. The holotype of C. fisheri fisheri should be referred to Cucumaria piperata (Stimp- son, 1864). Cucumaria piperata was de- scribed from specimens collected in Puget Sound. Unfortunately, the type specimen has been lost. There is little doubt, however, that Stimpson was describing the white, speckled sea cucumber commonly found in the sheltered waters of Washington and British Columbia (Fig. 5). Deichmann (1937: 169) confused the identification of this spe- cies by referring a specimen from off San Jose Point, west of Lower California, to C. piperata (Stimpson); however, that speci- men had “... knobbed perforated plates with dentate handle and small four-holed swollen or knobbed buttons.” C. piperata does not have the latter ““four-holed. . . but- tons.”’ Deichmann probably had either C. californica Semper or C. f. astigmata. Pan- ning (1962) illustrated the ossicles of C. pi- perata for the first time, and also stated that the four-holed plates, mentioned by Deich- mann, were absent. As a result of this confusion in the liter- ature, the identification of white, spotted sea cucumbers on the west coast of North America has been unclear. I have collected 914 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Holotype of Cucumaria fisheri fisheri Wells (USNM E01198); length 41 mm. VOLUME 103, NUMBER 4 plain white and spotted individuals with os- sicles that match those described for C. f. astigmata (Fig. 6). It appears that Wells combined the external features of C. piper- ata and the ossicles of the spotted form of C. f. astigmata in describing C. f. fisheri. Based on my re-examination of the two holotypes, I consider C. f fisheri Wells, 1924 to be a junior subjective synonym of C. pi- perata (Stimpson). The subspecies astig- mata should be raised to species status as Cucumaria astigmata Wells. In his revision of the Family Cucumari- idae, Panning (1949) placed fisheri Wells and fisheri forma astigmata Wells in the genus Stereoderma. He also placed piperata (Stimpson) into Pseudocnus. In two later pa- pers (Panning 1962, 1964) he reversed his decision, returned piperata to Cucumaria, and then removed fisheri from Stereoder- ma. Panning (1962:58) also redefined the ossicles of Pseudocnus. His description translates from German as “plates in the shape of pine cones, tightly layered. Below those, more deeply within the skin, round plates can be found.” C. astigmata has pine- cone ossicles in variable numbers usually clustered around the bases of the podia, and numerous thick buttons; thus, I believe as- tigmata should be placed in the genus Pseu- docnus. Genus Pseudocnus Panning, 1949 Pseudocnus astigmatus (Wells, 1924) (Figs. 2, 3, 6) Cucumaria fisheri forma astigmata Wells, 1924:117, fig. 2. Stereoderma fisheri forma astigmata Pan- ning, 1949:422. ONS Stereoderma fisheri Cherbonnier, 1951:42, plates 11 and 12. Material examined.—Collections by P. Lambert unless otherwise indicated. Ossi- cles were examined from | to 5 specimens in each lot. California: Holotype USNM E01196, length 33 mm, H. Wells, 1923, off Monterey and Cabrillo Point, Pacific Grove, depth 22 m; CASIZ 021555, 2 specimens, length 20, 25 mm, 10 May 1904, Albatross Station 4441, Point Pinos, Monterey Bay, depth 51- 64 m; CASIZ 15222, 5 specimens, length 20, 14, 22, 20, 8 mm, M. H. Spaulding, Pacific Grove, Monterey Bay, no depth; CASIZ 021554, 3 specimens, length 45, 40, 20 mm, summer 1908, China Point to Del Monte, depth 31 m; CASIZ 1200, | speci- men, length 8 mm, 8 Apr 1973, Ano Nuevo Cove, San Mateo Co., intertidal. British Columbia: RBCM 982-237-2, 4 specimens, 30 Sep 1982, Ogden Point, Vic- toria, Juan de Fuca Strait, 15 m; RBCM 988-751, 32 specimens, 2 Jun 1988, Whiffin Spit, Sooke, Juan de Fuca Strait, low inter- tidal; RBCM 973-154-5, 5 specimens, 28 Jun 1973, Dicebox I., Barkley Sound, depth 21 m; RBCM 973-183-30, 100 specimens, 25 Jul 1973, Cree Island, Barkley Sound, depth 26 m; RBCM 973-152-32, 2 speci- mens, 27 Jun 1973, Gilbert Island, Barkley Sound, depth 12 m; RBCM 974-595-2, 2 specimens, D. B. Quayle, 20 Jul 1959, Louie Creek, Esperanza Inlet, intertidal; RBCM 980-343, 1 specimen, 6 Jul 1980, Rugged Point, Kyuquot Sound, depth < 21 m; RBCM 985-384-7, 6 specimens, | Jun 1985, Sobry Island, Kyuquot Sound, intertidal; NMC 2221, 2 specimens, R. O’Clair, 29 Jun ———) Fig. 2. Holotype of Cucumaria fisheri astigmata Wells (USNM E01196) herein called Pseudocnus astigmatus (Wells); length 33 mm. VOLUME 103, NUMBER 4 Fig. 3. Ossicles from the mid-dorsal skin of the holotype of C. f astigmata Wells (= Pseudocnus astigmatus (Wells)); scale bar = 100 um. 918 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Ossicles from mid-dorsal skin of the holotype of C. f. fisheri Wells (= Cucumaria piperata Stimpson); scale bar = 100 wm. 1976, San Josef Bay, Vancouver Island, in- tertidal; RBCM 980-338, 2 specimens, 1 Jul 1980, Cliffe Point, Quatsino Sound, depth 18 m; RBCM 980-329-7, 3 speci- mens, 28 Jun 1980, Hunt Islets, Queen Charlotte Strait, depth < 21 m; RBCM 977- 444-5, 3 specimens, 12 Jan 1963, D. B. Quayle, FRB 63-2, Cormorant Channel, Queen Charlotte Strait, depth 26 m; RBCM 976-1037-7, 1 specimen, 27 Mar 1976, Juan Perez Sound, Queen Charlotte Islands, depth < 18 m; RBCM 102-19, 1 specimen, 3 Jun 1969, G. C. Carl, Frederick Island, Queen Charlotte Islands, intertidal; RBCM 108- 18, 7 specimens, 4 Jun 1969, G. C. Carl, Louis Point, Graham Island, Queen Char- lotte Islands, intertidal; RBCM 984-219-1, 1 specimen, 29 Apr 1961, D. B. Quayle, FRB S-4 H-9.7, Langara Island, Dixon En- trance, depth 82 m. Diagnosis. —Small form with robust ven- tral podia in rows as well as between the ambulacra. Dorsal podia visible as scattered dimples. Ten tentacles of equal size. Cal- VOLUME 103, NUMBER 4 919 careous ring without posterior prolonga- tions. Majority of ossicles are knobbed but- tons with from 4 to 10 perforations, some with a spiny, handle-like extension and more than 20 perforations; three-armed support- ing rods in podia. Color varies from yellow- ish-white to white with fine brown spots primarily on the dorsum and ends of ani- mal. Holotype. —USNM E01196. Type locality. —Monterey, California, 22 m. Range. —British Columbia to California; intertidal to 82 m. Remarks. —Pseudocnus astigmatus (Wells) appears to be closely related to Cu- cumaria curata Cowles and Cucumaria lu- brica Clark. Like these latter two species, P. astigmatus, collected at Whiffin Spit, Juan de Fuca Strait, brood large yolky eggs and juveniles between the ventral surface and the substrate. P. astigmatus also has similar tentacles, podia, body shape and has certain ossicle shapes in common with C. curata and C. /ubrica. Further studies are planned to investigate the systematics of these close- ly related forms. Acknowledgments I wish to thank Dave Pawson, National Museum of Natural History (USNM); Ailsa Clark, British Museum (BM); Peter Frank, Canadian Museum of Nature (CMN) (for- mally National Museums of Canada (NMC); and Elizabeth Kools, California Academy of Sciences (CASIZ), for loan of specimens. Translations of relevant German papers were kindly provided by Dave Pawson, Mary Bergen and Maria Abbott. I especially thank my colleagues Alex Peden and Rob Cannings for their constructive suggestions to improve the manuscript and to Rich Mooi Pe Fig. 5. Cucumaria piperata (Stimpson) collected from Nasparti Inlet, west coast Vancouver Island; depth 12 m; length 71 mm (RBCM 985-409-3). Z OS sal 0 Sg oe nN < 3 ee © > on = O fo) Nn | < O © ° — g ~Q eal rm onl fy ©) Nn © Z A ea sal QO © pe AY VOLUME 103, NUMBER 4 and an anonymous reviewer for their useful comments. Thank you also to Barbara Ku- luah for typing it. Funding was provided by the Royal British Columbia Museum (RBCM) and the Friends of the Royal B.C. Museum. Literature Cited Cherbonnier,G. 1951. Holothuries de L’Institut Royal des Sciences Naturelles de Belgique. —Me- moires de L’Institut Royal des Sciences Natu- relles de Belgique, 2 serie, fasc 41. Deichmann, E. 1937. 10. The Templeton Crocker Expedition. IX. Holothurians from the Gulf of California, the West Coast of Lower California and Clarion Island.— Zoologica 22:161-176. International Trust for Zoological Nomenclature. 1985. International Code of Nomenclature. Third edition. British Museum (Natural His- tory), 338 pp. Panning, A. 1949. Versuch einer Neuordnung der Familie Cucumariidae (Holothurioidea, Den- drochirota).—Zoologische Jahrbiicher Abteil- ung fiir Systematik, Okologie und Geographie der Tiere 78:404—-470. 921 1962. Bemerkungen iiber die Holothurien- Familie Cucumariidae (Ordnung Dendrochi- rota). 3. Tiel. Die Gattung Pseudocnus Panning, 1949.—Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 60:57-80. 1964. Bemerkungen uber die Holothurien— Familie Cucumariidae (Ordnung Dendrochi- rota). 3 Teil. Die Gattungen Stereoderma, Stau- rothyone und Trachythyone. — Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut. Kosswig-Festschrift 61:159-174. Stimpson, W. 1864. Descriptions of new species of marine invertebrata from Puget Sound, collect- ed by the naturalists of the Northwest Boundary Commission.— Proceedings of the Academy of Natural Sciences of Philadelphia 16:153-161. Wells, H. 1924. IV. New species of Cucumaria from Monterey Bay, California.—Annals and Maga- zine of Natural History Series 9, 14:113-121. Invertebrate Unit, Royal British Colum- bia Museum, 675 Belleville Street, Victoria, British Columbia, Canada, V8V 1X4. en eS E Uy EEE SEIS SISSSSSSSSSSSSSSSSSSS SS — Fig. 6. Plain and spotted forms of Pseudocnus astigmatus (Wells) (= C. f- astigmata Wells) collected at Whifhin Spit, Sooke, British Columbia; low intertidal amongst holdfasts of Hedophyllum; length of spotted specimen, 42 mm; plain specimen, 37 mm (RBCM 988-751). PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 922-930 A NEW GENUS AND SPECIES OF ANTHIINE FISH (PISCES: SERRANIDAE) FROM THE EASTERN SOUTH PACIFIC WITH COMMENTS ON ANTHIINE RELATIONSHIPS William D. Anderson, Jr., N. V. Parin, and John E. Randall Abstract. —Anatolanthias apiomycter, a new genus and species of anthiine fish, is described from two specimens collected in the eastern South Pacific, about 1500 km west of Chile at almost 26°S, near the southwest end of the Nazca Ridge. It is distinguished from all other serranids in having the following combination of characters: maxilla abruptly expanded distally, anterior naris somewhat remote from posterior naris, vomer edentate, vertebrae 26 (10 pre- caudal + 16 caudal), dorsal fin continuous—not incised between spinous and soft portions, opercle with three well-developed spinous processes, pleural ribs on vertebrae 3 through 11, epipleural ribs associated with first 11 vertebrae, epihaemal ribs on vertebrae 12 through 19, predorsal bones 2, principal caudal- fin rays 15, scales ctenoid with only marginal cteni, maxilla with scales, dorsal and anal fins naked (but each with low scaly sheath as its base), supramaxilla absent, and preopercle without antrorse spines. Characters that may be useful in clarifying relationships within the Anthiinae are discussed. During a recent cruise in the eastern South Pacific, personnel aboard the Soviet re- search vessel Prof. Shtokman collected two specimens of a new species of anthiine ser- ranid fish. These specimens are so distinc- tive that they also warrant description as a new genus. The main purpose of this paper is to describe the new genus and species. In addition, because of the inadequately un- derstood relationships of the serranid sub- families and the chaotic generic classifica- tion within the Anthiinae, we discuss characters that appear to be useful in elu- cidating serranid intrarelationships—par- ticularly those within the Anthiinae. The Anthiinae include a plethora of brightly colored species of small to medium size that inhabit tropical to temperate seas worldwide at shallow to moderate depths, usually on rocky bottoms or coral reefs which provide shelter. Most species feed on zooplankton a short distance above the bot- tom to which individuals rapidly retreat when predators approach. These fishes of- ten occur in aggregations, and, as far as known, are protogynous hermaphrodites. The sexes are often colored differently and may exhibit morphological differences, par- ticularly in fin structure; typically males at- tend large harems. There are about 165 valid described spe- cies of Anthiinae, variously classified in more than 20 genera. A remarkable 73 spe- cies of the subfamily were first described between 1975 and 1987—46 of them in the years 1979-1982. At least 14 undescribed species (in addition to the one described herein) are in museum collections, and still more must remain to be discovered. This explosive increase in the number of anthiine species known is due to more intensive col- lecting. Of particular importance has been the relatively recent advent of SCUBA, which has allowed collecting at previously inaccessible depths. Because only one genus of anthiines has VOLUME 103, NUMBER 4 been revised since 1975 (Plectranthias, Randall 1980), it is not surprising that the generic classification is inadequate. We hope that our comments on relationships will contribute to a better understanding of ge- neric limits and relationships within the Anthiinae. Materials and Methods The holotype is deposited in the Division of Fishes, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. (USNM); the paratype, in the Zoolog- ical Institute, Academy of Sciences USSR, Leningrad (ZIL). Counts and measurements follow Ander- son & Heemstra (1980), except as indicated below. The first vertebra with a haemal spine was considered the first caudal vertebra; the urostylar vertebra, the last. Mabee (1988) interpreted the predorsal bones of teleosts as the phylogenetic homologues of supra- neurals, and because supraneural is the more widely used senior synonym, she recom- mended its use for those bones in teleosts. We prefer to use the term “‘predorsal bone”’ because it is firmly entrenched in the liter- ature of percoid fishes. Gillrakers and pseu- dobranchial filaments were counted on the right side. Internarial distance was the dis- tance between the posterior border of the anterior naris and anterior border of the posterior naris. Body depth was measured at the dorsal-fin origin and at greatest depth; body width, just posterior to gill opening; and prepelvic length, from premaxillary symphysis to origin of pelvic fin. Caudal concavity was the horizontal distance be- tween verticals at distal tips of longest and shortest caudal-fin rays. The symbol > pre- ceding the measurement of a fin ray indi- cates that the element measured was slightly damaged. Most measurements are present- ed as percentages of the standard length (SL), but some are given as quotients of SL, head length, snout length, or orbital diameter. These quotients are rounded to the nearest 0032 Anatolanthias, new genus Diagnosis. —A genus of Anthiinae distin- guishable from all other genera of Serrani- dae by the following combination of char- acters. Maxilla abruptly expanded distally, particularly on labial border, where a shelf or rostrally directed hook is present at point of expansion. Anterior naris located rather far anteriorly on snout, somewhat remote from posterior naris. Vomer edentate. Ver- tebrae 26 (10 precaudal + 16 caudal). Dor- sal fin single, not incised between spinous and soft portions. Posterior margin of bony opercle with three well-developed spinous processes, middle one largest. Pleural ribs on vertebrae 3 through 11. Epipleural ribs associated with first 11 vertebrae. Epihae- mal ribs (bones in the same series as the epipleural ribs which appear to be modified intermuscular bones; see Stiassny & Jensen 1987:300) on vertebrae 12 through 19. For- mula for configuration of predorsal bones, anterior neural spines, and anterior dorsal pterygiophores 0/0/2/1+1/1/ (using sym- bolization of Ahlstrom et al. 1976). Caudal fin forked; principal rays 15 (8 + 7); branched rays 13 (7 + 6). Scales ctenoid, with only marginal cteni (i.e., no ctenial bas- es present proximal to marginal cteni; see Hughes 1981); no secondary squamation. Scales present on maxilla and in interorbital region. Well-developed axillary process of modified scales at pelvic-fin base. Dorsal and anal fins without scales, but each with low scaly sheath at its base. No supramax- illa. Preopercle without antrorse spines. Description. —Characters included in the generic diagnosis form part of the generic description and are not repeated. Mouth terminal; upper and lower jaws about equal. Premaxillae protrusile. Posterodorsal bor- der of maxilla not covered by elements of circumorbital series when mouth closed. Palatine with teeth. Pterygoids and tongue edentate. Fleshy papillae on posterior half of orbital border. Posterior margin of pre- opercle serrate; ventral margin of preopercle essentially smooth. Branchiostegals 7. Pseu- 924 dobranch present. Gill arches 4, with slit behind fourth. Longest gillrakers longer than longest gill filaments. Lateral line complete, extending to base of caudal fin (running par- allel to dorsal body contour below dorsal fin, curving gently to near mid-lateral axis of body on caudal peduncle). Squamation well developed on bases of pectoral, pelvic, and caudal fins and continuing onto fins. Pelvic-fin rays I, 5; pelvic fin inserted slight- ly posterior to vertical through pectoral-fin base. Procurrent spur (Johnson 1975) ab- sent; penultimate ventral procurrent cau- dal-fin ray not shortened basally. Parhy- pural with well-developed hypurapophysis. Autogenous hypurals 5. Epurals 3. Uro- neurals 1 pair (posterior pair absent). No trisegmental pterygiophores associated with dorsal and anal fins. Other characters are those of the single species. Etymology. — Anatolanthias (anatole, east; anthias, a seafish) is from the Greek, refer- ring to the occurrence of the genus in the eastern Pacific. The gender is masculine. Type species.—Anatolanthias apiomyc- ter, new species. Anatolanthias apiomycter, new species Figs. 1, 2 Holotype: USNM 309202, 93.9 mm SL; 25°41.7'S, 85°23.7'W; 160-168 m; R/V Prof. Shtokman cruise 18, station 1922; 26 April 1987; bottom otter trawl. Paratype: ZIL 49471, 89.0 mm SL; same data as for holotype. Description. —Characters presented in the generic diagnosis and description form part of the species description and are not re- peated unless necessary for clarification. Most of the scales have been lost from the paratype; as a consequence it is impossible to enumerate or accurately estimate most of its scale counts. Data for the holotype are followed, in parentheses, by those for the paratype, when different. Dorsal-fin rays X,16. Anal-fin rays III, 7. Pectoral-fin rays PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 21, both sides (22 both sides); dorsalmost pectoral-fin ray unbranched on both sides (right ventralmost unbranched), other rays branched. Procurrent caudal-fin rays 14, both dorsally and ventrally. Gillrakers on first arch 10 + 27 (11 + 26), no rudimentary rakers. Pseudobranchial filaments 18 (16). Fleshy papillae on posterior half of orbital border 22 left, ca. 20 right (20 left, 21 right). Tubed lateral-line scales 63 left, ca. 62 right (estimated ca. 62 left). Rows of cheek scales ca. 8 left, ca. 9 right. Scale rows between lateral line and mid-base of spinous dorsal fin 3. Scales from dorsal-fin origin to lateral line 5. Scales from anal-fin origin to lateral line ca. 17. Serrae on posterior margin of preopercle ca. 19 left, ca. 20 right (16 left, ca. 14 right); serrae enlarged into bifurcate spinous process at angle on right side (blunt process at angle), not enlarged on left side; ventral margin of preopercle essentially smooth. Body depth (at dorsal-fin origin) 3.95 (4.15), head length 3.75 (3.60) in SL. Bony orbital diameter 3.05 (3.40) in head length. Bony interorbital width 3.75 (4.50) in head length, 1.20 (1.35) in bony orbital diameter. Snout length 1.60 (1.50) in bony orbital di- ameter. Internarial distance 3.05 (2.80) in snout length. The following morphometric data are in percentages of SL. Head length 26.7 (27.9). Snout length 5.5 (5.4). Bony orbital diameter 8.7 (8.2). Postorbital head length 12.9 (13.7). Upper jaw length 12.5. Maxilla width 4.2 (4.3). Internarial distance 1.8 (1.9). Least bony interorbital width 7.1 (6.2). Body depth at dorsal-fin origin 25.3 (24.0). Greatest body depth 26.9 (24.0). Body width 16.4 (14.4). Predorsal length 28.8 (28.5). Prepelvic length 35.0 (32.0). Preanal length 64.9 (63.9). Caudal-peduncle length 21.7. Least caudal-peduncle depth 11.1 (10.6). Pectoral-fin length 29.8 (28.4). Pelvic-fin length 23.6 (21.9). Pelvic spine 13.0 (13.1). Dorsal-fin base 56.7 (58.2). First dorsal spine >3.7 (4.4). Second dorsal spine 8.1 (8.4). Third dorsal spine 10.1 (>9.1). Fourth dorsal spine 10.8 (11.3). Longest VOLUME 103, NUMBER 4 dorsal spine 11.8, fifth (11.6, sixth). Tenth dorsal spine >7.8 (9.3). Longest dorsal soft ray ca. 14.0, tenth (broken). Anal-fin base 17.5 (17.8). Depressed anal-fin length 28.6 (27.9). First anal spine 4.3 (5.1). Second anal spine >9.2 (10.2). Third anal spine >8.3 (9.2). Longest anal soft ray ca. 14.1, second (ca. 15.2, fourth). Upper caudal-fin lobe ca. 33.8 (broken). Lower caudal-fin lobe ca. 31.9 (broken). Caudal concavity ca. 20.7 (fin damaged). Premaxilla with lateral row of conical teeth and medial series of much smaller conical teeth; one or two small canines at anterior end of lateral row; medial series with one to a few small posteriorly directed canines adjacent to symphysis; no teeth at symphy- sis. Dentary with row of conical teeth, teeth smaller near symphysis; one to a few teeth enlarged into small canines about 30 to 35 percent of distance from anterior end of jaw to posterior end of row of teeth; exserted canine at anterior end of jaw; no teeth at symphysis. Vomer edentate, but with sev- eral fleshy papillae. Palatine with band of small conical teeth. Maxilla reaching just posterior to vertical through middle of eye. Posterior border of anterior naris produced into flap which falls well short of posterior naris when reflected. Free margins of intercpercle and subopercle somewhat irregular, but not serrate. Max- illa, interorbital region, lachrymal, cheek, preopercle, interopercle, opercle, and sub- opercle densely covered with scales; scales on dorsum of snout not reaching anterior end of snout—leaving considerable area scaleless (anteriormost scales on dorsum of snout very small); most of lateral aspect of snout naked; no scales on lower jaw, gular region, branchiostegals, and branchiostegal membranes. Modified scales (interpelvic process) overlapping pelvic fin bases along mid-ventral line. Tubes in lateral-line scales simple. Distal margin of anal fin rounded. Second anal spine more robust than first or third, about twice as long as first, slightly longer 925 0) iff ¥ a 39 +3 we we! o oe 3 — =) ou wv S & > 8 ru) AYA D\ Ae AYRAWN th \ x . ; - . ~" at SBS N . ra _ NY NE > oa 2 NY 3 7 A\ \ N ” AY eu \ ~ he ~ . SI ae + 4 “el IS eee > J ‘ , “ x = L—d . P re ne fi . i | \ ie wat . , C \ aK ee Fig. 1. Holotype of Anatolanthias apiomycter, USNM 309202, 93.9 mm SL; eastern South Pacific. 926 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON T.[,1990 Fig. 2. Head of holotype of Anatolanthias apiomycter, USNM 309202, 93.9 mm SL; eastern South Pacific. than third. Pectoral fin symmetrical, middle rays longest. Pectoral fin reaching vertical between bases of first and second dorsal soft rays (vertical through base of first dorsal soft ray), falling short of vertical through vent. Pelvic fin reaching vertical through base of ninth dorsal spine (vertical between bases of eighth and ninth dorsal spines), falling short of vent. Coloration.—In alcohol the types are straw-colored with no distinctive pigmen- tation. A color transparency of the holotype, taken shortly after capture, shows: body uniformly red, iris red, dorsal and caudal fins red, anal fin paler, paired fins not clearly visible. Distribution. —Anatolanthias apiomycter is known only from the type-locality in the eastern South Pacific, about 1500 km off the coast of Chile, near the southwest end of the Nazca Ridge. Etymology.—The name apiomycter (apios, far away; mykter, nostril) is from the Greek and is an allusion to the unusual po- sition of the anterior nostril. The new name is a masculine noun in apposition to Ana- tolanthias. Relationships Johnson (1983) defined the Serranidae with respect to the Percichthyidae (sensu Gosline 1966) on the basis of three reduc- tive specializations (absence of the posterior uroneural, procurrent spur, and third pre- ural radial cartilages), and showed that VOLUME 103, NUMBER 4 members of the Serranidae share at least one innovative specialization (the presence of three spines on the opercle)— thus support- ing the hypothesis of monophyly for the family. Following Gosline (1966), Johnson (1983, 1988) recognized three subfamilies in the Serranidae: the Serraninae, Epine- phelinae, and Anthiinae, but identified an autapomorphy for only the Epinephelinae. Anderson & Heemstra (1989) mentioned two characters (one reductive, the other in- novative) that may prove useful in defining the Anthiinae. The reductive character, ab- sence of a tooth plate on the second epi- branchial, was discussed by Baldwin (1990) and interpreted as an autapomorphy of the Anthiinae. As noted by Johnson (1983), it is difficult to evaluate the significance of vertebral number in determining relationships among the Percoidei; nevertheless, this character may be of value in delimiting the bound- aries of the Anthiinae. Species of Serraninae and Epinephelinae almost always have 24 vertebrae, but species of Anthiinae have 25 to 28, usually 26. (Acanthistius, a serranine, has 26 vertebrae, and Niphon, a primitive epinepheline, has 30 [Johnson 1983]. Pseu- dogramma has 26 vertebrae; Suttonia has 26 or 27; and Aporops has 27 or 28 [Leis & Rennis 1983; Carole C. Baldwin, pers. comm.]. These last three genera are highly derived grammistin epinephelines, whose progenitors presumably acquired additional vertebrae subsequent to the divergence of the grammistins from the main line of epi- nepheline evolution.) Anderson & Heem- stra (1989) presented an analysis of verte- bral number as a character in the Serranidae; they considered 24 or 25 as the most prim- itive state in the Serranidae and 26, 27, and 28 as progressively more derived states. Realizing that additional study is needed before the Anthiinae can be definitively de- fined, we accept, at present, the absence of the second epibranchial tooth plate and high vertebral number (26 to 28, usually 26) as autapomorphies delimiting this subfamily. 927 Recognition of the Anthiinae as a distinct taxon is of considerable practical value be- cause the concept anthiine unites a large number of look-alike species that share uniquely derived characters at some level within the Serranidae. According to Johnson (1984), the prim- itive and most common number of princi- pal caudal-fin rays (branched rays + 2) in percoids is 17 (9 + 8), and the most com- mon and presumably primitive number of predorsal bones is three. Johnson (1984) noted that most percoids (59 groups) have ctenoid scales in which the cteni are discrete bony plates that are added continually to the posterior field as the scale grows, and that in most of these the posterior field is filled with the remains of old cteni (ctenial bases of Hughes 1981). The retention of ctenial bases in the posterior field is pre- sumably more primitive in the Percoidei than the condition in which only primary and secondary rows of marginal cteni are present (i.e., no ctenial bases remaining in the posterior field) because, according to Johnson (1984), only a few groups (includ- ing the Anthiinae) have species lacking cte- nial bases in the posterior field. If, in the Serranidae, 15 principal caudal-fin rays, one or two predorsal bones, and absence of cte- nial bases in the posterior field are derived states, as they appear to be, they may be helpful in clarifying the generic classifica- tion of the Anthiinae.! Baldwin (1990) found that all of the spe- cies of Anthiinae that she examined that have 17 principal caudal-fin rays also have three predorsal bones, whereas those with 15 principal rays have either two or three predorsal bones (all Atlantic and eastern Pa- cific species with 15 principal rays have only two predorsal bones). Our data corroborate ' We realize that the argument common equals ple- siomorphous is logically flawed, but, because the sister group of the Serranidae has not been identified and because there are no other pertinent data, it is the only argument available. 928 Baldwin’s with the exceptions of Giganthias immaculatus (if it is an anthiine) from Jap- anese waters and Plectranthias vexillarius from the Gulfof Oman, which have 17 prin- cipal rays and two predorsal bones, and Plectranthias japonicus from the western Pacific, which has 16 or 17 principal rays and two predorsal bones (also see Randall & Heemstra 1978, Randall 1980). Randall (1980) reported five other species of Plec- tranthias that have variable or unusual numbers of principal caudal-fin rays (one species with 15 to 17, two with 16, one with 15 or 16, and one with 14 to 16), but he did not give counts of predorsal bones. Randall & Lubbock (1981) reported that four species of Pseudanthias, of the Indo-Pacific sub- genus Mirolabrichthys, have 15 principal rays but only one predorsal bone and that one other Mirolabrichthys has 15 principal rays and either one or two predorsal bones. Among anthiines there is a strong corre- lation in number of principal caudal-fin rays, number of predorsal bones, and type of ctenoid scale. Species with 17 principal rays and three predorsal bones usually have scales in which ctenial bases have been retained in the posterior field, but among those with 15 principle rays apparently all lack ctenial bases in the posterior field (Anderson, un- published data). Although all three of the presumed derived states (15 principal cau- dal-fin rays, one or two predorsal bones, and absence of ctenial bases in the posterior field) are reductive, the shared possession of all three may be indicative of propinquity of descent. (Based on our incomplete data we speculate that the sequence of appearance of these derived characters in the main line of anthiine evolution was: loss of ctenial bases in the posterior field, reduction in number of principal caudal-fin rays, and re- duction in number of predorsal bones.) An- atolanthias belongs to a large group of an- thiines (which includes, e.g., species of Anthias, Hemanthias, Holanthias, Luzon- ichthys, Pronotogrammus, and Rabaulich- thys) that displays all three derived traits. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Anatolanthias shares the following ap- parently derived traits with species of the Indo-Pacific genera Luzonichthys (six nom- inal species) and Rabaulichthys (two spe- cies): anterior naris rather remote from pos- terior naris (nares usually close together in anthiines; in the above genera internarial distance 2.45—4.35 times in snout length, other anthiines for which data are avail- able—with the exception of Anthias tenu- is—5.10—14.00 [usually 6.00-1 1.00], A. ten- uis 4.30—4.95), vomerine dentition absent or extremely reduced (vomerine dentition usually well developed in anthiines, Pseud- anthias fucinus being the only other anthiine known to lack vomerine teeth; see Randall 1981, Allen 1984, Randall & Ralston 1984, Randall & Pyle 1989), and the sum of num- bers of pairs of epipleural and epihaemal ribs 16 to 19 (specimens of about 60 species of anthiines representing 14 other genera with 8-15 [usually 9-13], Nemanthias car- berryi with 16 and one species of Pseud- anthias with 17). In addition, another char- acter state that may be a synapomorphy for Anatolanthias, Luzonichthys, and Rabaul- ichthys is the number of pleural ribs; these genera have nine pairs, but anthiines usually have eight (about three-fourths of the spe- cies for which data are available with eight). Based on these characters we hypothesize that the genera Luzonichthys, Rabaulich- thys, and Anatolanthias constitute a mono- phyletic assemblage. We recognize three genera for this assemblage because all three possess characteristics that can be inter- preted as autapomorphies and no two of the genera bear identified synapomorphies. Luzonichthys is quite distinctive (Randall 1981); species of this genus have two dorsal fins (other anthiines have a single dorsal fin) and have the third or ventralmost opercular spine greatly reduced—apparently absent in some specimens (other anthiines have three well developed opercular spines). Also Lu- zonichthys has 11 precaudal and 15 caudal vertebrae; among anthiines with 26 verte- brae the vast majority (including Anatolan- VOLUME 103, NUMBER 4 thias and Rabaulichthys) have 10 and 16, respectively. Rabaulichthys (Allen 1984, Randall & Pyle 1989) is characterized by the possession of a high sail-like spinous dorsal fin in males (no other anthiines have a dorsal fin with this shape) and the absence of palatine dentition (other anthiines have teeth on the palatine). In Anatolanthias the maxilla is abruptly expanded distally, par- ticularly so on the labial border where a shelf or a rostrally directed hook is present at the point of expansion (the shelf or hook is absent in Luzonichthys and Rabaulich- thys and the distal expansion of the maxilla is not particularly abrupt). This configura- tion of the maxilla has, to our knowledge, not been reported in the literature for any anthiine, although Phillip C. Heemstra in- formed us (pers. comm.) that he has ob- served it in several species, including An- tnias nicholsi, A. salmopunctatus, A. tenuis, Hemanthias vivanus, and Sacura parva; we have observed it in a number of other spe- cies, e.g., Anthias anthias, A. asperilinguis, Hemanthias peruanus, and Holanthias martinicensis. The abrupt expansion of (or shelf or hook on) the maxilla is easily over- looked because it is usually hidden by the lip. Despite the fact that other anthiine spe- cies (apparently not closely related to An- atolanthias) have the abruptly expanded maxilla, this trait may have been indepen- dently derived in Anatolanthias and thus be an autapomorphy for this genus. Acknowledgments P. C. Heemstra called the senior author’s attention to the hook on the maxilla of the holotype of Anatolanthias apiomycter, W. F. Hoffman and P. P. Maier made the ra- diographs used in this study; J.-E. Trecartin prepared the illustrations; M. Jones typed the manuscript; and C. C. Baldwin, P. C. Heemstra, V. G. Springer, and C. C. Swift reviewed the manuscript. This research was supported by a grant from the Faculty Re- search and Development Committee of the 929 College of Charleston to the senior author. To all who contributed to the completion of this research we express our appreciation. This is contribution number 87 of the Grice Marine Biological Laboratory, College of Charleston. Literature Cited Ahlstrom, E. H., J. L. Butler, & B. Y. Sumida. 1976. Pelagic stromateoid fishes (Pisces, Perciformes) of the eastern Pacific: kinds, distributions, and early life histories and observations on five of these from the northwest Atlantic.— Bulletin of Marine Science 26:285-402. Allen,G.R. 1984. Anew genus and species of anthiid fish from Papua New Guinea.— Revue francaise d’Aquariologie 11:47—50. Anderson, W. D., Jr., & P. C. Heemstra. 1980. Two new species of western Atlantic Anthias (Pisces: Serranidae), redescription of A. asperilinguis and review of Holanthias martinicensis. —Copeia 1980:72-87. —., & 1989. Ellerkeldia, a junior syn- onym of Hypoplectrodes, with redescriptions of the type species of the genera (Pisces: Serrani- dae: Anthiinae).— Proceedings of the Biological Society of Washington 102:1001-1017. Baldwin, C. C. 1990. Morphology of the larvae of American Anthiinae (Pisces: Serranidae) with comments on relationships within the subfam- ily.—Copeia (in press). Gosline, W. A. 1966. The limits of the fish family Serranidae, with notes on other lower per- coids.— Proceedings of the California Academy of Sciences, 4th Ser., 33:91-111. Hughes, D. R. 1981. Development and organization of the posterior field of ctenoid scales in the Platycephalidae.—Copeia 1981:596-606. Johnson, G. D. 1975. The procurrent spur: an un- described perciform caudal character and its phylogenetic implications. — Occasional Papers of the California Academy of Sciences 121:1- 23. 1983. Niphon spinosus: a primitive epinephe- line serranid, with comments on the monophy- ly and intrarelationships of the Serranidae.— Copeia 1983:777-787. 1984. Percoidei: development and relation- ships. Pp. 464-498 in H. G. Moser et al., eds., Ontogeny and systematics of fishes. American Society of Ichthyologists and Herpetologists, Special Publication No. 1. 1988. Niphon spinosus, a primitive epinephe- line serranid: corroborative evidence from the 930 larvae. —Japanese Journal of Ichthyology 35:7- 18. Leis, J. M., & D.S. Rennis. 1983. The larvae of Indo- Pacific coral reef fishes. New South Wales Uni- versity Press, Sydney, Australia, 269 pp. Mabee, P.M. 1988. Supraneural and predorsal bones in fishes: development and homologies. — Cope- ia 1988:827-838. Randall, J. E. 1980. Revision of the fish genus Plec- tranthias (Serranidae: Anthiinae) with descrip- tions of 13 new species.— Micronesica 16:101— 187. 1981. Luzonichthys earlei a new species of anthiine fish from the Hawaiian Islands.— Freshwater and Marine Aquarium 4(9):13-18. —, & P. C. Heemstra. 1978. Reclassification of the Japanese cirrhitid fishes Serranocirrhitus la- tus and Isobuna japonica to the Anthiinae.— Japanese Journal of Ichthyology 25:165-172. —, & R. Lubbock. 1981. A revision of the ser- ranid fishes of the subgenus Mirolabrichthys (Anthiinae: Anthias), with descriptions of five new species. — Contributions in Science, Natural History Museum of Los Angeles County, No. 333:1-27. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON —,&R.M.Pyle. 1989. Anewspecies ofanthiine fish of the genus Rabaulichthys (Perciformes: Serranidae) from the Maldive Islands. — Special Publication, J. L. B. Smith Institute of Ichthy- ology, No. 47:1-7. ——,, &S. Ralston. 1984. A new species of serranid fish of the genus Anthias from the Hawaiian Islands and Johnston Island.— Pacific Science 38:220-227. Stiassny, M. L. J., & J. S. Jensen. 1987. Labroid intrarelationships revisited: morphological complexity, key innovations, and the study of comparative diversity.—Bulletin of the Muse- um of Comparative Zoology 151:269-319. (WDA) Grice Marine Biological Labo- ratory, College of Charleston, 205 Fort Johnson, Charleston, South Carolina 29412; (NVP) P. P. Shirshov Institute of Ocean- ology, Academy of Sciences USSR, Krasi- kova 23, Moscow 117218, USSR; (JER) Bernice Pauahi Bishop Museum, Box 19000-A, Honolulu, Hawaii 96817. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 931-954 THREE NEW SPECIES OF SYMPHURINE TONGUEFISHES FROM TROPICAL AND WARM TEMPERATE WATERS OF THE EASTERN PACIFIC (SYMPHURUS: CYNOGLOSSIDAE: PLEURONECTIFORMES) Madhu N. Mahadeva and Thomas A. Munroe Abstract.—Three new species of Symphurus are described based on speci- mens collected from the Gulf of California southward along Central America and northern South America to northern Peru. Symphurus oligomerus, n. sp., occurs from the southern Gulf of California to northern Peru and is a relatively deep-water (84-481 m) species characterized by: a 1-3-2 pattern of interdigi- tation of dorsal pterygiophores and neural spines (ID pattern), 85-97 dorsal- fin rays, 71-83 anal-fin rays, 12 caudal-fin rays, 48-52 total vertebrae, 5 hy- purals, black peritoneum, an alternating series of dark rectilinear pigment blotches and unpigmented areas on the posterior portion of the dorsal fin and throughout the anal fin, a dark pigment blotch on the proximal portion of the caudal fin, with distal portions distinctly unpigmented, and the absence of a pupillary operculum. Symphurus chabanaudi, n. sp., occurs from the northern Gulf of California to northern Peru and is among the largest species in the genus, reaching lengths of at least 233 mm SL. This shallow-water species (2- 59 m) is characterized by: a 1-5-3 ID pattern, 12 caudal-fin rays, 4 hypurals, small scales on blind-side dorsal- and anal-fin rays, a prominent black spot on the ocular-side opercle, and absence of a pupillary operculum. Symphurus diabolicus, n. sp., known only from a single specimen collected at 501 m off Chatham Island in the Galapagos Archipelago, is readily distinguished by the combination of: a 1-3-2 ID pattern, 12 caudal-fin rays, 5 hypurals, 109 dorsal- fin rays, 94 anal-fin rays, 58 total vertebrae, small scales (ca. 135 in longitudinal series), spotted peritoneum, and large eyes (18.6% of head length). Symphurus is the only genus of tongue- fishes (Cynoglossidae) occurring in the New World, with over 40 nominal species re- corded from marine waters on both sides of the Americas (Ginsburg 1951, Mahadeva 1956, Menezes & Benvegnt 1976, Munroe 1987). In the eastern Pacific, at least 14 spe- cies inhabit coastal and deep-waters (Jordan & Evermann 1898, Mahadeva 1956, Mun- roe & Mahadeva 1989, Munroe & Nizinski 1990) from southern Oregon (Eschmeyer et al. 1983) to northern Peru (Hildebrand 1946, Mahadeva 1956, Chirichigno 1974). The majority of species occur in tropical waters. Recent collecting in inshore and deeper areas off the coasts of Mexico, Central, and South America have resulted in the capture of many tonguefishes, including specimens of the new species described herein. This paper provides formal descriptions of three of these new species. Methods. —Specimens examined are list- ed by collection acronym following Leviton et al. (1985). Counts and measurements fol- low Mahadeva (1956), Munroe & Maha- deva (1989), and Munroe (1990a). Standard length (SL) is used throughout. Where pos- sible, measurements to 150 mm were taken to the nearest 0.1 mm with dial calipers or ocular micrometer; measurements larger 932 than 150 mm were made to the nearest mm with a steel ruler. Measurements are ex- pressed either as thousandths of standard length or thousandths of head length. Ab- breviations in text and tables are: head length (HL), head width (HW), eye diameter (ED), snout length (SNL). Unless stated other- wise, observations and descriptions of pig- ment patterns are based on fishes preserved in formalin and stored in ethyl] or isopropyl alcohol. Interdigitation pattern (ID pattern).— Patterns of interdigitation of proximal dor- sal pterygiophores and neural spines were counted and recorded for the first three, or in unusual cases, the first four interneural spaces. The number of dorsal pterygio- phores inserted into interneural spaces 1-3 was found to be diagnostic for species or groups of species of Symphurus (Munroe 1987). ID patterns are indicated by a pteryg- iophore formula such as 1-3-2. The 1-3-2 ID pattern indicates one pterygiophore in- serts in interneural space one, three in in- terneural space two, and two in interneural space three. The first neural spine abuts di- rectly against the cranium so there is no obvious space between it and the cranium. Therefore, the first interneural space reflect- ed in the formula is that between the first and second neural spines. Variation in dorsal- and anal-fin rays and total vertebrae were examined using SYS- TAT programs (Wilkinson 1988) for one- way ANOVA and Tukey HSD multiple comparison test on log-transformed vari- ables of specimens divided into groups based on capture location. Symphurus oligomerus, new species Figs. la, b, 2, Tables 1-3 Symphurus atramentatus (not of Jordan & Bollman).—Garman 1899:229 (counts; measurements; color description; Pana- ma and Colombia). Symphurus sp.—Lavenberg & Fitch 1966: 108 (Gulf of California; photograph). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Holotype. —SIO 84-70, (female, 85.8 mm), Mexico, Gulf of California, Bahia La Paz, 24°26.7'-24°24.8'N, 110°36.0’— 110°37.5'W (Toadhop I, Station 4), 240 m, collected with 25’ otter trawl by R. Rosen- blatt and party aboard RV E. B. Scripps, 14 Jul 1984. Paratypes. —(171 specimens, 38.0-145 mm SL): Gulf of California: SIO 68-94, (15 of 40, 38-121), 29°19.9'—29°20.4'N, 113°10.4’— 113°12.0’'W, 273 m, 20 Jan 1968. LACM 21719, (1, 64.2), ca. 23°N, 109°W, 6.5 mi from Punta Gorda, outer Gorda Bank, ca. 130 m, 12 Mar 1949. LACM 20407, (19, 49-76), outer Gorda Bank, 12 Mar 1949. UCLA W56-79, (10, 54-71), outer Gorda Bank, 12 Mar 1949. LACM 20261, (3, 44— 55.5), 29°33’45"N, 113°30'35”W, Puerto Refugio, Isla Angel de la Guarda, 94 m, 29 Jan 1940. Nicaragua: SIO 73-280, (25 of 174, 66.7— 119.4), 11°07.4-07.8’N, 86°35.0-35.5'W, 159 m, 18 Apr 1973. Costa Rica: UF 33932, (3, 97.4—109.6), Gulf of Nicoya. LACM 33827-10, (6, 87.4— 120.6), Gulf of Nicoya, central Puntarenas Province, 29 Jun 1973. SIO 73-281, (24, 86.2—110), 10°50.2—53.2'N, 86°20—24.3'W, 196 m, 18 Apr 1973. UCR 425-18, (1, 113.9), Puntarenas Province, between Cabo Blanco and Punta Herradura, 310 m. UCR 494-6, (1, 91.6), Puntarenas Province, off Parrita, 259 m. Panama: MCZ 28540, (14, 74.2—105.1), 7°40.0'N, 79°17.9'W, 235 m, 8 Mar 1891. USNM 57882, (2, 110.5—113.8), 7°33'40’N, 79°43'20"W, 283 m, 9 Mar 1891. UMML 26051, (12, 91.7-113.1), Bay of Panama, 7°30.5'-7°31.0’N, 79°41.5'-79°43.3'W, 210 m, 4 May 1967. MCZ 28537, (1, 145), 7°12.3'N, 80°55'W, 337 m, 23 Feb 1891. UMML 31935, (20, 80.1-111.5), 6°31.2- 32.1'N, 77°32.2-34.4'W, 205 m, 16 Jan 1972. UMML 31947, (7, 70.7-116.2), 6°28.8-29.3'N, 77°29.2-30'W, 216 m, 16 Jan 1972. Ecuador: CAS 57858, (1, 126.3), 2°14’S, 81°11’30”W, 481 m, 31 Aug 1968. VOLUME 103, NUMBER 4 Peru: CAS 24201, (6, 93.1-111.9), 5°02’S, 81°24'W, 254 m, 3 Jun 1966. Diagnosis.—A Symphurus with a 1-3-2 ID pattern; 12 caudal-fin rays; a black peri- toneum usually showing through abdomi- nal wall on both sides of body; 8—13 black, irregularly rectilinear, blotches alternating with unpigmented areas on posterior two- thirds of dorsal fin and entire length of anal fin; similar black blotch across base of cau- dal fin with distal three-fourths of caudal fin distinctly unpigmented; 85—97 dorsal-fin rays; 71-83 anal-fin rays; 48—52 (usually 49— 51) total vertebrae; 5 hypurals; 86—96 scales in a longitudinal series; 39-46 scales in transverse row; no pupillary operculum; rel- atively large eyes (ED/SNL 1.3 to 1.5); up- per jaw not extending beyond vertical line through posterior margin of pupil of lower eye; well-developed dentition on ocular-side jaws; dorsal-fin origin at vertical line through middle of pupil of upper eye; and body color yellowish tan to dark brown with 3-7, most- ly incomplete, crossbands. Description. —Frequency distributions of meristic data are provided in Table 1. ID pattern typically 1-3-2 (182/205), infre- quently 1-3-3 (8/205), 1-2-3 (5/205), or 1-4-2 (4/205), rarely otherwise. Caudal-fin rays 12 (215 of 228 specimens), less fre- quently 11 (9 individuals) or 13 (two spec- imens), rarely 10 or 14 (one each). Dorsal- fin rays 85-97, usually 88-94, X¥ = 91.3. Anal-fin rays 71-83, usually 74-80, X¥ = 77.2. Pelvic-fin rays 4. Total vertebrae 48- 52, usually 49-51, rarely 48 or 52, ¥ = 50.3; abdominal vertebrae 9 (3+6). Hypurals 5 (180/202), less frequently 4 (22/202). Lon- gitudinal scale rows 86-96, X = 90.5. Scale rows on head posterior to lower orbit 20-— 23. Transverse scales 39-46, X = 43.3. Proportional measurements are provided in Table 2. Medium-sized Symphurus with relatively deep body, 259-307 SL, X = 281; greatest depth in anterior third of body; gradual posterior taper to body beginning approximately at anterior third of standard length. Trunk length 709-750 SL, X = 729. Head relatively long (250-291 SL, X = 271), 933 usually slightly shorter than body depth. Snout length 40-69 SL, ¥ = 58; covered with small ctenoid scales. Dermal papillae well developed on blind-side snout. Mouth moderately sized, upper jaw length 58-94 SL, X = 78. Posterior margin of jaws reach- ing verticals between mid-eye and posterior margin of pupil of lower eye. Lower eye moderately sized, 32-38 SL, ¥ = 34: eyes usually slightly subequal in position with upper in advance of lower eye. Anterior and medial surfaces of eyes with 2 or 3 rows of 3 or 4 scales lying posterior to posterior nos- tril, and in narrow interorbital region; no scales posteriorly between eyes or on pos- terior surface of eyes. Pupillary operculum absent. Dorsal-fin origin at vertical line through mid-point of upper eye; predorsal length 61-115 SL, X¥ = 100. Longest dor- salfin ray 85-115 SL (X = 100). Dorsal- and anal-fin rays without scales on blind side. Pelvic fin relatively long, 64-99 SL, ¥ = 86. Anal-fin origin at vertical line approxi- mately through bases of dorsal-fin rays 12- 13. Caudal fin with three or four rows of scales on base; scales diminishing in size posteriorly. Teeth well developed on blind- and oc- ular-side jaws. Blind-side dentary with 3-4 rows of teeth in middle of crescentic tooth band. Crescent tapers smoothly laterally with number of tooth rows decreasing dis- tally, ending in single tooth. Tooth band on blind-side premaxilla with 3+4 rows of teeth posteriorly, tapering anteriorly to single tooth. Ocular-side premaxilla with one ir- regular row of prominent conical teeth ex- tending from anterior tip to below anterior margin of lower eye; one irregular row of prominent conical teeth on lower jaw ex- tending to vertical through anterior margin of pupil of lower eye. Pigmentation. —Ground color tan to dark brown, with variable number (usually 3-7) of darker brown, mostly incomplete cross- bands (6-9 scales wide) extending from behind nape to base of caudal fin; 8-11 well- defined, rectilinear, black blotches alternat- ing with unpigmented areas on median fins. 934 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Symphurus oligomerus. a. SIO 84-70, holotype; female, 85.8 mm SL; Gulf of California, Bahia La Paz, 24°25.45'N, 110°36.75'W. b. SIO 73-280, paratype; male, 87.2 mm SL; Nicaragua, 1 1°07.4—07.8'N, 86°35.0— 35.5'W. Specimens from southern Mexico, Costa Rica, Nicaragua, and Panama generally darker than those from other locations. Blind side usually unpigmented, suggesting creamy whiteness in life. Some specimens with lon- gitudinal series of small melanophores along body midline on blind side. Peritoneum black, showing through abdominal wall on both sides of body. Head usually uniformly pigmented; oc- casionally with diffuse, faint crossband, 4— 5 scales wide, extending from dorsal margin to ventral border at posterior margin of operculum. Ocular-side outer opercle oc- casionally with diffuse, irregularly-shaped spot on ventralmost surface. Inner linings of both ocular- and blind-side opercles speckled with melanophores. Isthmus un- pigmented on both sides of body. Ocular- side lips with dark band of pigment. Posterior two-thirds of dorsal fin and en- tire anal fin with clearly defined, rectilinear blotches, not extending to tips of fin rays. Blotches (8-13 in dorsal fin and 8-12 in anal fin) covering proximal three-fourths of fin rays, often continuous at fin-ray bases with body crossbands. Dark blotches on dorsal fin increase in intensity and size posteriorly; blotches to about middle of body lighter and smaller (covering only 2—4 rays) compared with those on posterior half of fin (black and better defined, covering 5-10 rays). Blotch- es in anal fin roughly parallel those in dorsal fin in position and intensity of pigmenta- VOLUME 103, NUMBER 4 935 Table 1.—Frequency distributions of meristic features for Symphurus oligomerus. (Asterisks indicate counts for holotype.) nN Dorsal-fin rays 85 86 87 88* 89 90 91 92 93 94 95 96 97 3% Frequency 1 1 1 13 28 37 41 50 44 21 2 l 1 91.3 Anal-fin rays 71 72 73 We TS 716 77 78 79 80 81 82 83 X Frequency 1 2 4 13 22 39 38 59 39 17 2 l 1 W122 Caudal-fin rays 10 11 12* 13 14 Frequency 1 925 2 1 Hypurals 4 5* Frequency 22 180 Total vertebrae 48 49* 50 51 52 X Frequency 3 35 75 84 5 50.3 Longitudinal scale rows 86 87 88 89 90 91 92* 93 94 95 96 xX Frequency 2) — 7 1 19 2 11 — 1 1 Ay SOLS) Transverse scale count 39 40 41 42 43 44 45 46* X Frequency 2 D, 2 7 4 23 2 4 43.3 Pattern Frequency % Interdigitation Pattern 1-3-2-2* 182 88.8 1-3-3-2 8 3.9 1-2-3-2 5 2.4 1-4-2-2 4 2.0 1-200 3 NSS) 1-3-3-1 2 1.0 2-3-2-2 ] 0.5 tion. Posteriormost blotch in dorsal and anal fins often coalescing with blotch at base of caudal-fin rays. Distal three-fourths of caudal-fin rays hy- aline, contrasting sharply with strongly pig- mented posterior dorsal and anal fins, and base of caudal fin. Specimens occasionally with heavy pigmentation on proximal fourth of caudal fin on blind side of body. Etymology. —‘‘Oligomerus”’ from the Greek “oligos” meaning few and “meros” meaning part of segment, in reference to the relatively low number of vertebrae and dor- sal- and anal-fin rays of this species com- pared with most other eastern Pacific Syym- phurus. Distribution. —Offshore habitats on the continental shelf (Fig. 2) from the Gulf of California at Puerto Refugio, off Isla Angel de la Guarda (29°33'45”N, 113°30'35’W), including Gorda Banks at the entrance to the Gulf, and continuing southwards to the @ S. chabanaudi B® S. oligomerus A 5S. diabolicus PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Geographical distribution of Symphurus oligomerus, S. chabanaudi, and S. diabolicus. Symbols may represent more than one locality and/or more than a single lot from each locality. continental shelf off Peru (6°21’'S, 80°59’W). There were no specimens collected between ca. 20°N and 10°N available for our study. At this time, the occurrence or abundance of this species between these latitudes is un- known, however, there is no a priori reason to assume that S. oligomerus does not occur in this region. The disjunct distribution in- dicated in Fig. 2 is presumed to be an ar- tifact of collecting intensity. Symphurus oligomerus has been collected at depths ranging from approximately 80 to 481 m. Ninety-three percent of 356 speci- mens for which reliable depth information is available were collected between 111 and 300 m. Only five specimens have been col- lected shallower (one each at 80 and 84 m; three at 94 m) while approximately 6% were taken deeper than 300 m (17 specimens be- tween 301-350 m, 3 specimens at 389 m, and one specimen at 481 m). Available information for ca. 10% of the VOLUME 103, NUMBER 4 Table 2.—Summary of morphometrics for the ho- lotype (SIO 84-70) and 33 paratypes of Symphurus oligomerus. Proportional measurements, except stan- dard length in mm, in thousandths of standard length. aS Paratypes Holotype Range Mean EE ee eee Standard length 85.8 44.4-76.2 _ Body depth 295 259-307 281 Predorsal length 61 85-115 100 Pelvic-fin length 64 75-99 86 Head length 256 250-291 271 Snout length 40 49-69 58 Upper jaw length 68 58-94 78 Eye diameter 35 32-38 34 specimens indicates this species is frequent- ly collected on substrates containing a large sand component, including muddy sand (1 collection, 7 specimens), sand (4 collections, 5 specimens), sand and gravel (1 collection, 3 individuals), rock (1 collection, 1 speci- men), and a sand, shell, and rock mixture (1 collection, 30 individuals). Geographical variation.—Comparisons of selected meristics (Table 3) revealed that specimens collected from the Gulf of Cali- fornia and Mexico had significantly fewer dorsal- (f = 43.84; P < 0.001) and anal-fin rays (f = 36.60; P < 0.001), and total ver- tebrae (f = 66.78; P < 0.001) than those fishes collected from Central America (El Salvador, Costa Rica, and Panama) and northern South America (Colombia to northern Peru). Fin-ray and vertebral counts of S. oligomerus taken between El Salvador and Panama, although slightly greater, were not significantly different from those of specimens collected off northern South America. Comparisons. —Symphurus oligomerus is the third species described (of four known species) from eastern Pacific waters char- acterized by a 1-3-2 ID pattern, 12 caudal- fin rays, and black peritoneum. Other east- ern Pacific species with these characteristics are S. microlepis Garman, S. gorgonae Cha- banaud and S. diabolicus (described below). 9377 Table 3.—One-way ANOVA for selected meristic features of Symphurus oligomerus collected from dif- ferent portions of the species range. (Asterisk indicates difference at P < 0.001.) n Range Mean SD Dorsal-fin rays (f= 43.84; P < 0.001) *Gulf of California 46 85-93 90.1 1.35 Central America 146 89-97 92.3 1.37 South America 11 90-94 91.8 1.47 Anal-fin rays (f= 36.60; P < 0.001) *Gulf of California 46 71-79 76.1 1.42 Central America 146 74-83 78.1 1.39 South America 11 76-80 77.8 1.47 Total vertebrae (f= 66.78; P < 0.001) *Gulf of California 46 48-50 49.3 0.59 Central America 145 49-52 50.6 0.67 South America 11 50-51 50.4 0.50 These other species are quite different from S. oligomerus and easily distinguished ei- ther by their lack of alternating rectilinear pigmented blotches and unpigmented areas on the dorsal and anal fins characteristic of S. oligomerus (fin pigmentation uniformly pigmented in S. diabolicus and S. gorgonae) or by differences in meristic values. Sym- phurus oligomerus has lower vertebral and fin-ray counts than S. microlepis and S. dia- bolicus (total vertebrae 48-52 versus 57 in S. microlepis and 58 in S. diabolicus; dorsal- fin rays 85-97 versus 106 in S. microlepis and 109 in S. diabolicus; anal-fin rays 71- 83 versus 94 in S. microlepis and 92 in S. diabolicus). Furthermore, the body of S. oligomerus is deepest in the anterior third of the standard length with a relatively rapid posterior taper compared with that of S. diabolicus in which the body is relatively elongate with almost uniform depth occur- ring throughout much of the mid-body re- gion and with only a gradual posterior taper. Symphurus oligomerus is readily distin- guished from S. gorgonae in that the blind side is uniformly creamy-white compared with that of S. gorgonae in which the blind side has a pepper-dot pattern of melano- phores, especially well developed along the 938 basal pterygiophore regions of the dorsal and anal fins. These two species differ further in that there is little overlap for several me- ristic features (total vertebrae 48-52 in S. oligomerus versus 46-49 in S. gorgonae; dorsal-fin rays 85—97 versus 80-89; anal-fin rays 71-83 versus 63-74; hypural numbers 5 versus 4). Other eastern Pacific Symphurus pos- sessing pigmented spots or blotches on the dorsal and anal fins that may be confused with S. oligomerus are S. callopterus Mun- roe & Mahadeva (sometimes collected with S. oligomerus) and S. atramentatus Jordan & Bollman. However, both S. callopterus and S. atramentatus have an unpigmented peritoneum (versus black in S. oligomerus) and a well-developed pupillary operculum (absent in S. oligomerus). Additionally, S. oligomerus has 5 hypurals (4 in S. callop- terus and S. atramentatus) and a 1-3-2 ID pattern (usually 1-3-4 in S. callopterus; 1-3-3 in S. atramentatus). Symphurus atramen- tatus also has well-defined oval black spots on the fins (versus rectilinear black blotches in S. oligomerus). Symphurus oligomerus differs further from S. callopterus in several counts (total vertebrae 48—52 versus 57-61 in S. callopterus; dorsal-fin rays 85—97 ver- sus 105-114; anal-fin rays 71-83 versus 91- 98; longitudinal scale rows 86-96 versus 97— 114), and the distal half of the caudal fin is unpigmented in S. oligomerus, whereas in S. callopterus, the distal half of the caudal fin either has an ill-defined blotch or the entire fin is uniformly darkly pigmented. Some meristic features of seven other eastern Pacific species, S. varius Garman, S. williamsi Jordan & Culver, S. fasciolaris Gilbert, S. /eei Jordan & Bollman, S. atri- caudus (Jordan & Gilbert), S. melanurus Clark, and Symphurus sp. (the species de- scribed by Munroe & Nizinski (1990) with the lower meristic values; hereafter referred to as Symphurus sp.) overlap those of S. oligomerus. Symphurus oligomerus, how- ever, differs from all of these species in hav- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ing alternating blotches in the dorsal and anal fins (versus uniformly pigmented fins without alternating blotches, or dorsal and anal fins with posterior intensification of pigment, or with dorsal and anal fins speck- led). Symphurus oligomerus also differs from these species in ID pattern (1-3-2 versus 1-3-3 in S. varius; 1-4-3 in S. fasciolaris and S. leei; 1-5-3 in S. atricaudus, S. melanurus, and Symphurus sp.; and 1-5-3 or 1-4-3 in S. williamsi) and in the number of hypurals (5 in S. oligomerus versus 4 in all others, except S. varius). Symphurus oligomerus has larger scales (86—96 in a longitudinal series) than does S. varius (120-124). Certain me- ristic values of S. oligomerus completely overlap those of S. williamsi, but the two species are easily distinguished. Symphurus oligomerus has a black peritoneum (unpig- mented in S. williamsi), lacks a pupillary operculum (present in S. williamsi), and lacks scales on the blind side dorsal- and anal-fin rays (present in S. williamsi). Sym- phurus fasciolaris and Symphurus sp. differ further from S. oligomerus in caudal-fin ray counts (12 in S. oligomerus versus 10 in S. fasciolaris and 11 in Symphurus sp.). Among Symphurus species occurring out- side the eastern Pacific region, some counts for S. oligomerus overlap those of S. piger (Goode & Bean), S. pusillus (Goode & Bean), S. plagiusa (Linnaeus), S. plagusia (Schnei- der, in Bloch & Schneider), S. diomedeanus (Goode & Bean), and S. civitatium Ginsburg from the western Atlantic, and S. trifascia- tus (Alcock) from the Indian Ocean. All of these species, except S. diomedeanus, lack the highly pigmented fins found in S. oli- gomerus. Although S. diomedeanus has counts and pigmented dorsal and anal fins reminiscent of those of S. oligomerus, these two species are distinct in caudal-fin ray counts (12 versus 10 in S. diomedeanus), hypural numbers (5 versus 4), and ID pat- terns (1-3-2 versus 1-4-3). Symphurus di- omedeanus also has a pupillary operculum (absent in S. oligomerus). VOLUME 103, NUMBER 4 Additional Material Examined Counts were taken from the following 150 non-type specimens (28.5—117.5 mm): Gulf of California: LACM 8818-10, (15, 67-109), 28°55'N, 112°50.5'W, midway be- tween southern tip of Isla Tiburon and Isla Angel de la Guarda. SIO 60-97, (7, 54.1- 81.1), 28°13.8-15.0'N, 111°46.7-48.0'W, off Estero de Tastiuta, Costa de Hermosillo, Sonora. SIO 60-110, (1, 82.2), 28°12.9’N, 112°3.2'W, off Costa de Hermosillo, So- nora. SIO 60-98, (25, 79.4—110.1), 28°02’— 28°06’N, 111°47.2'-111°53.2'W, 162 m, 21 Mar 1960. LACM 20259, (2, 42.0—52.5), 27°58'40’N, 111°24'10”W, Isla San Pedro Nolasco, 111 m, 12 Mar 1936. LACM 20262, (1, 86.5), 27°58'35’N, 110°22'40’W, Isla San Pedro Nolasco, 6 Feb 1940. LACM 8842-5, (1, 65.5), 26°57'N, 111°49.8'W, 3- 5 mi north of Punta Concepcion. SIO 65- 293, (7, 28.5-58.9), 25°40.6-44.3'N, 111°05.3-—5.6'W, northwest of Isla Monser- rate, Baja California Sur. SIO 84-70, (1, 86.5), Bahia La Paz, 24°25.45'N, 110°36.75'W, 240 m, 14 Jul 1984. SIO 84- 80, (3, 91.6-100.7), Baja California Sur, 23°31.5-32.5'N, 110°28-30'W, 143 m, 17 Jul 1984. SIO 84-81, (7, 76.1-117.0), north- west of Lobos Point, 23°29.95'N, 110°27.1'W, 17 Jul 1984. LACM 20260, (1, 54.9), 23°02’N, 109°0'15”W, Inner Gorda Bank, Baja California Sur, 20 Jan 1940. Costa Rica: CAS 43872, (1, 97.9), Cabo Blanco, 268 m, 9 Mar 1974. CAS 44104, (1, 100.4), Cabo Blanco, 268 m, 9 Mar 1974. Panama: UMML 27031, (73 of 101, 54.1- 137.7), Bay of Panama, 7°39.5'—7°40.9'N, 79°40.7'-79°42.7'W, 117 m, 4 May 1967. USNM 57883, (1, 99.7), 7°16'45’N, 79°56'30"W, 389 m, 9 Mar 1891. Peru: CAS 24200, (1, 118.5), 4°53’S, 81°20-23'’W, 84 m, 2 Jun 1966. IMARPE 4, (1, 94.0), 5°0.5’S, 83°24.5'W. CAS 24979, (1, 100.1), 6°21’S, 80°59’W, 142 m, 4 Jun 1966. Additional material examined but not counted or measured (277 specimens): 939 Gulf of California: SIO 68-94, (25 of 40, 38-121), 29°19.9’-29°20.4'’N, 113°10.4’— 113°12.0'W, 273 m, 20 Jan 1968. SIO 60- 98, (37, 79.4-110.1), 28°2-6'’N, 111°47.2- 53.2'W, off San Juan Bautista Flats, Sonora, 162 m, 21 Mar 1960. SIO 68-103, (15, 73.2- LOUES Pie 282195372 8°90 SNe. Al 22870— 112°10.5'W, 303 m, 21 Jan 1968. Nicaragua: SIO 73-280, (149, 64-120), 11°07.407.8'N, 86°35.0—35.5'W, 159 m, 18 Apr 1973. Costa Rica: SIO 73-281, (12 of 24, 86.2- 110), 10°50.2—53.2'N, 86°20-24.3’W, 196 m, 18 Apr 1973. LACM 33827-56, (1, 106.9), Gulf of Nicoya, central Puntarenas Province, 29 Jun 1973. UCR 682-6, (1, 79.7), Puntarenas Province, Isla del Cano, 80 m. UCR 2190-1, (2, 70.8-81.9), off Cabo Matapalo, tip of Osa Peninsula, 23 Jun 1973. Panama: UMML 27031, (28 of 101, 54.1- 137.7), Bay of Panama, 7°39.5’-7°40.9’N, 79°40.7'-79°42.7'W, 117 m, 4 May 1967. MCZ 28539, (5, 107.0-121.2), 7°33.7'N, 79°43.3'W, 283 m, 9 Mar 1891. MCZ 70978, (2, 89.4—100.8), 7°16.8'N, 79°56.5’W, 389 m, 9 Mar 1891. Symphurus chabanaudi, new species Figs. 2, 3a, b, Tables 4-7 Aphoristia elongata (not of Gunther). —Jor- dan & Gilbert, 1883:24 (in part) (listed, Panama). Symphurus elongatus (not of Gunther).— Jordan & Goss, 1889:323 (in part) (after Jordan & Gilbert; Panama).—Jordan & Evermann, 1898:2707 (in part) (after Jor- dan & Gilbert; Panama).—Gilbert & Starks, 1904:203 (in part) (fish market, Panama City).—Meek and Hildebrand, 1928:1006 (counts; measurements; color description; in key; Panama).—Breder, 1936:5 (in part) (Gulf of California).— Seale, 1940:14 (listed, Tenacatita Bay, Mexico and Colombia).—?Fowler, 1944: 495 (listed, Panama).— Phillips, 1981:54 (in part) (Jiquilisco Bay, El Salvador). 940 Symphurus atricaudus (not of Jordan & Gil- bert).—Breder, 1936:6 (in part) (Gulf of California).— Phillips, 1981:54 (in part) (Jiquilisco Bay, El Salvador). Symphurus sechurae Hildebrand, 1946:476 (in part) (Gulf of California). Holotype. —USNM 305717, (male, 130.8 mm), El Salvador, El Potrero, Jiquilisco Bay, Station Number 6, 13°16’N, 88°38'’W, on mud bottom canal in mangroves, 5 m try- net, collected 14 Jul 1976, P. Phillips and party. Paratypes. —(115 specimens, 47.5—233 mim): Gulf of California: Near San Felipe, 31°18—22'N, 114°47-50’W, ca. 5 m, 6—9 Apr 1947, 50 specimens distributed to the fol- lowing collections: USNM 164493, (1, 139), USNM 164494, (10, 47.5-152), BMNH 1956.3.1:6-14, (9, 90.5-159), CAS 20696, (10, 59.5—156), UCLA W53-196, (19, 93- 153), SIO 47-53, (1, 109). YPM 630, (10, 124-157), north of San Felipe, 26 m, 20 May 1926. UCLA W52-46, (4, 72-127), 10 mi off Santa Clara, Sonora. USNM 119742, (1, 184), south of Guaymas, 1940. El Salvador: USNM 236606, (2, 185- 188), Saite, 26 m, 28 Oct 1975. USNM 291339, (1, 188), La Libertad, 26 m, 29 Oct 1975. USNM 220701, (5, 110.1—131.0), El Potrero, Jiquilisco Bay, 14 Jul 1976. Costa Rica: LACM 30716-11, (4, 182- 211), Puntarenas Province, Gulf of Nicoya, Isla Negritos, 13 Feb 1968. LACM 30714- 15, (3, 178-185), Puntarenas Province, Isla Chira, 1968. UCR 1122-3, (2, 147.8-149.5), Puntarenas and Guanacaste Provinces, Golfo de Nicoya and Playa Hermosa, 18-— 28 m. Panama: UCLA W53-275, (11, 137-217), Panama Bay. SIO 80-23, (5, 128-184), Pan- ama Bay, Isla Verde, 4 m, 9 Apr 1980. Colombia: USNM 305718, (5, 138-200), 4°20-18'N, 77°28-29'W, off Rio Togorama. USNM 305719, (7, 164-178), 2°57'N, 77°48'W, Punta Coco, south of Buenaven- tura. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ecuador: CAS 24199, (5, 161-233), Gulf of Guayaquil, 3°16’S, 80°25’W. Diagnosis. —A Symphurus with 1-5-3 ID pattern; 12 caudal-fin rays; 98-109 dorsal- fin rays; 82-92 anal-fin rays; 52-57 total vertebrae; 91-104 scales in longitudinal se- ries; 32-42 transverse scales; no pupillary operculum; relatively small eye (ED/SNL 2.4 to 2.7); 2—5 small, ctenoid scales on blind-side dorsal- and anal-fin rays; dis- tinct, large black blotch on ocular-side outer opercle (not to be confused with black bran- chial cavity lining showing through oper- culum in many species of Symphurus); up- per jaw usually reaching to or just posterior to vertical line through middle of pupil of lower eye (rarely extending posteriorly to vertical through rear margin of lower eye); dorsal-fin origin usually equal with or just posterior to vertical line through middle of pupil of upper eye (rarely originating at ver- tical line through anterior margin of upper eye); posterior dorsal and anal fins and cau- dal fin dusky to dark black; body tan to dark brown usually with 6-8, sharply contrasting crossbands 6-10 scales wide; and unpig- mented peritoneum. Description. — Frequency distributions of meristic data are provided in Table 4. ID pattern usually 1-5-3 (47 of 95 individuals), less frequently 1-4-3 (28/95) or 1-4-4 (13/ 95), rarely 1-4-3-3, 1-5-2, or 1-5-3-3. Cau- dal-fin rays 12 (103/108), rarely 11 (5/108). Dorsal-fin rays 98-109, usually 100-107, X = 103.4. Anal-fin rays 82-92, usually 84— 89, X = 86.6. Pelvic-fin rays 4. Total ver- tebrae 52-57, usually 53-56, rarely 57 (1 of 125), X = 54.1; abdominal vertebrae 9 (3 + 6). Hypurals 4 (90/90). Longitudinal scale rows 91-104, X¥ = 97.9. Scale rows on head posterior to lower orbit 21—23. Transverse scales 32-42, ¥ = 39.0. Proportional measurements appear in Table 5. Large-sized (to at least 233 mm SL) species with relatively deep body (239- 293 SL, ¥ = 265); greatest depth in anterior third of body with gradual posterior taper starting near middle of body. Trunk length VOLUME 103, NUMBER 4 94] Table 4.—Frequency distribution of meristic features for Symphurus chabanaudi. (Asterisks indicate counts for holotype.) Dorsal-fin rays SuSE ee ee eee 98 99 100 101 102 103 104 105* 106 107 108 109 ¥ Frequency 1 1 OA 4 2 0) 6 ee 4 8 — Anal-fin rays Frequency 1 3 1@ 20 28 AZ 15 13 3 2 1 86.6 Caudal-fin rays 11 12* Frequency 5 103 Hypurals 4* Frequency 90 Total vertebrae 52 53 54 55 S6emnST x Frequency 8 22 DAG peDO ole: 1 54.1 Longitudinal scale rows 91 92 93* 94 95 96 97 98 99 100 101 102 103 104 ¥ Frequency 1 2) 1 4 3 @ 5) 16 9 5) ] 4 4 Lee g7e9 Frequency D, 1 = 7) 3 3 10 14 17 8 6 39.0 Pattern Frequency % a Interdigitation Pattern 1-5-3-2* 47 49.5 1-4-3-2 28 29.5 1-4-4-2 13 NS 7/ 1-5-3-3 1 1.0 1-5-2-2 2 2a 1-4-3-3 2 251 1-5-4-2 1 1.0 1-4-2-2 ] 1.0 770-816 SL, X = 799. Head relatively short, 185-213 SL, X¥ = 201: somewhat shorter than body depth. Snout length 42-57 SL, V = 47; covered with small ctenoid scales. Dermal papillae well developed on blind- side snout, chin, and head region just ven- tral to anterior portion of dorsal fin. Mouth moderately sized, upper jaw length 49-69 SL, X¥ = 64; usually reaching to or just pos- terior to vertical line through middle of pu- pil of lower eye (rarely extending posteriorly to vertical through rear margin of lower eye). Lower eye relatively small, 15-21 SL, ¥ = 19; eyes usually slightly subequal in position with upper in advance of lower eye. Surface of head from region posterior to posterior nostril and including narrow interorbital re- gion to posterior margin of eyes with 13- 14 rows of scales, each row with 6-8 scales (difficult to discern in juveniles). Pupillary operculum absent. Dorsal-fin origin at or just posterior to a vertical line through mid- 942 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Symphurus chabanaudi. a. USNM 305717, holotype; male, 130.8 mm SL; El Salvador, Jiquilisco Bay, El Potrero. b. USNM 291339, paratype; male, 188 mm SL; El Salvador, La Libertad. dle of upper eye, occasionally at a vertical line through anterior margin of upper eye; predorsal length, 50-88 SL (XY = 80); longest dorsal-fin ray, 62-86 SL (X = 71). Blind sides of dorsal- and anal-fin rays with 3—5 very thin, ctenoid scales (difficult to discern in juveniles). Anal-fin origin approximately at vertical line through base of dorsal-fin rays 14-16. Pelvic fin relatively long, 60- 83 (X = 73); reaches first or occasionally second anal-fin ray when depressed poste- riorly. Caudal fin with approximately 9-10 relatively large scales on basal half of fin; distal half of caudal fin with numerous mi- nute scales diminishing in size distally. Teeth well developed on blind-side jaws. Lower jaw on blind side with crescentic tooth band about 5—8 rows wide. Arms of crescent tapering smoothly to a single tooth at ex- tremities. Upper jaw on blind side with 5- 8 rows of teeth posteriorly; band tapering to single tooth anteriorly. Juveniles with only 4—5 visible rows of teeth in bands on blind- side jaws. Teeth on ocular-side jaws in ir- regular rows. Ocular-side premaxilla with one irregular row of feeble teeth extending from anterior tip to point equal with vertical line through anterior base of anterior nos- tril; ocular-side dentary with irregular row of very feeble teeth extending posteriorly to vertical line through anterior base of ante- rior nostril. Pigmentation. —Ground color of live specimens tannish brown with pinkish hue. Median fins pale lavender. Blind side pink- ish to creamy white. Ground color in preserved specimens tannish to dark brown, usually with variable number (6-10) of sharply contrasting black- ish-brown crossbands 6—10 scale rows wide extending from behind nape to base of cau- dal fin. Color of specimens from the Gulf VOLUME 103, NUMBER 4 of California typically lighter than those col- lected on muddy bottoms along coastal El Salvador, Costa Rica, and Panama. Blind side uniformly creamy white. Occasionally, individuals of both sexes have dark pigment patches on caudal region and scattered along bases of dorsal and anal fins on blind side. Crossbands on body varying in intensity, usually somewhat darker than ground color. Peritoneum unpigmented. Head with one, or occasionally two, faint crossbands. Anterior band diffuse, about 3- 4 scales wide, located immediately posterior to eyes and barely reaching ventral margin of head. Posterior band crossing opercular region and extending ventrally slightly onto blind side of head; often incomplete in dor- sal area of head, usually fading, disappear- ing completely, or occasionally forming faintly pigmented crossband, about 6 scales wide, at dorsal margin of operculum; ven- tralmost portion of band always expanded into distinct black blotch about 6-10 scale rows wide composed of large epidermal me- lanophores; clearly visible even in very young specimens; covering more than half the outer surface of ocular-side operculum. Inner linings of opercles on both sides of body heavily pigmented; inner lining on oc- ular-side opercle more heavily pigmented than that on blind side, often visible exter- nally on outer opercular surface as poorly- defined, dusky blotch. Isthmus heavily spotted on both sides of body. Basal portions of dorsal and anal fins to about a fifth or sixth the height of fin paler than remainder of fin. Dorsal fin anterior to a vertical line through anal-fin origin usually unpigmented; remainder of dorsal fin and entire anal fin light to dark brown or black without discrete spots or blotches; progres- sively darker posteriorly, with the posterior- most fourth or fifth of fins almost black in mature fish (fins usually darker in mature males). Posterior portions of dorsal, anal, and caudal fins on blind side often with small patches of melanophores. Caudal fin dark, 943 Table 5.—Summary of morphometrics for holotype (USNM 305717) and 35 paratypes of Symphurus cha- banaudi. Measurements, except standard length in mm, in thousandths of standard length. Paratypes Holotype Range Mean Standard length 130.8 70.8-226 _ Body depth 263 239-293 265 Predorsal length 50 70-88 80 Pelvic-fin length 60 63-83 73 Head length 193 185-213 201 Snout length 42 43-57 47 Upper jaw length 49 58-69 64 Eye diameter 17 15-21 19 similar to posterior portions of dorsal and anal fins. Median fins of juveniles usually only lightly pigmented or appearing almost unpigmented to naked eye, but with mela- nophores discernible under magnification. Etymology.—Named in honor of Paul Chabanaud who contributed greatly to our knowledge of flatfishes, especially of the ge- nus Symphurus. Distribution.—A widespread and com- monly collected species (Fig. 2) in shallow warmer waters ranging throughout the Gulf of California and extending coastally south- ward to approximately Callao, Peru (12°06'S, 76°55'W). There were no speci- mens collected between ca. 20°N and 10°N available for our study. At this time, the occurrence or abundance of this species be- tween these latitudes is unknown, however, there is no a priori reason to believe that S. chabanaudi does not occur in this region. The disjunct distribution indicated in Fig. 2 is presumed to be an artifact of collecting intensity. Symphurus chabanaudi has been collect- ed at depths ranging from approximately 2 to 59 m; however, the majority of speci- mens (97%) were taken at depths shallower than 28 m. The deepest captures of 277 specimens for which depth information is available are for seven fish taken between 944 Table 6.—One-way ANOVA for selected meristic features of Symphurus chabanaudi collected from dif- ferent portions of the species range. (Asterisk indicates difference at P < 0.001.) n Range Mean SD Dorsal-fin rays (f = 30.68; P < 0.001) *Gulf of California 33 99-103 101.2 1.14 *Central America 46 98-109 103.8 2.00 *South America 5 103-109 105.7 1.49 Anal-fin rays (f = 39.77; P < 0.001) *Gulf of California 33 82-87 84.8 1.18 *Central America 51 84-91 86.8 1.56 *South America 5 86-92 88.4 1.50 Total vertebrae (f= 50.44; P < 0.001) *Gulf of California 33 52-54 . 53.0 0.71 *Central America 53 53-57 54.6 0.92 *South America 5 55-56 55.8 0.45 30 and 37 m and for solitary specimens taken at 39 m and 59 m. Symphurus chabanaudi has most often been collected on muddy or sand-mud sub- strates. Available substrate data list mud for 13 collections (123 specimens), mud and sand (11 collections, 76 specimens), mud and shell (2 collections, 9 specimens), mud and rocks (1 collection, 2 specimens), sand (5 collections, 13 specimens), clay and bits of shell (1 collection, 50 specimens), and rock (1 collection, 1 specimen). Geographical variation. —Numbers of dorsal- (f = 30.68; P < 0.001) and anal-fin rays (f = 39.77; P < 0.001), and total ver- tebrae (f = 50.44; P < 0.001) differed sig- nificantly (Table 6) in specimens collected in each of three different portions (Mexico, Central America, and northern South America) of the species range. Specimens collected in the Gulf of California and coast- al waters off northern Mexico had the lowest numbers of meristic elements, whereas specimens taken off Colombia and Peru had the highest numbers of finrays and verte- brae for any group examined. Remarks. —Earlier investigators fre- quently misidentified Symphurus chaba- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON naudi as S. elongatus, undoubtedly due to similarities in meristic features of these spe- cies. Symphurus elongatus occurs sympat- rically with S. chabanaudi, especially in southern Central America and in coastal waters of northern South America, and the two species are collected syntopically. How- ever, these species are quite distinct (see comparisons below). Jordan & Gilbert (1883) and later, Jordan & Goss (1889) and Jordan & Evermann (1898) included specimens of S. chabanaudi in their accounts of S. elongatus. These ac- counts were based on three specimens of S. chabanaudi (CAS-SU 6900), misidentified as S. elongatus by Jordan and co-workers, collected from Albatross station 2804 off Colombia, together with numerous speci- mens of S. /eei and 10 specimens of S. elon- gatus. At least one specimen (USNM 50333) identified as S. elongatus by Gilbert & Starks (1904) is S. chabanaudi, but their account probably included other specimens. Mor- phometric proportions reported (table on p. 204) more closely match those of S. chaba- naudi than S. elongatus, especially body depth measurements. These authors re- ported body depths equalling 24.5-—28.0% SL, which are larger values than those (20.4— 25.5% SL) noted for specimens of S. elon- gatus examined by Mahadeva (1956). Also, the large sizes (up to 255 mm) reported for specimens referred to as S. elongatus by Gil- bert & Starks more nearly reflect sizes at- tained by S. chabanaudi; S. elongatus is a much smaller species usually not exceeding 150 mm SL. It is apparent that Meek & Hildebrand (1928) included specimens of S. chabanaudi in their account of S. elongatus from Pan- ama, given the pigment characters listed in their key, especially references to dark crossbars, a dark blotch on the opercle, and posterior darkening of dorsal and anal fins (pigment characters not found in S. elon- gatus). Two lots (USNM 81032 and 81674) examined in the present study, cited as S. VOLUME 103, NUMBER 4 elongatus by Meek & Hildebrand, are S. chabanaudi. It is possible that Meek & Hil- debrand did not examine any S. elongatus in their study, as they noted that the maxilla in their specimens reached a point only equal with the middle of the lower eye, whereas, in S. elongatus, the maxilla usually extends to a point well beyond the vertical line through the posterior margin of the lower eye. Specimens (YPM 630-633a) collected in the Gulf of California and reported as S. elongatus and S. atricaudus by Breder (1936) are S. chabanaudi. Hildebrand (1946) ten- tatively identified specimens of S. chaba- naudi (USNM 126741 and USNM 50333) collected in the Gulf of California and Pan- ama Bay as S. sechurae, a nominal species now placed in the synonymy of S. melanu- rus Clark (Munroe 1990b). Hildebrand (1946) also incorrectly identified as S. elon- gatus several lots (USNM 144788-144791) collected in the Miraflores Locks, Panama Canal, in which were actually represented four different species, including juvenile S. chabanaudi, S. elongatus, S. melanurus and S. williamsi. Comparisons. —Eastern Pacific Symphu- rus with counts comparable to those of S. chabanaudi include S. elongatus (Gunther), S. atricaudus, S. fasciclaris, S. leei, S. mela- nurus, S. diabolicus, and S. microlepis. The pigmentation of S. chabanaudi (well-de- fined crossbands and a black blotch on the ocular-side opercle) differs from that of S. elongatus, S. melanurus, S. microlepis, and S. diabolicus, which usually lack crossbands and an opercular spot. Symphurus leei and S. atricaudus both have crossbands in a pat- tern different from S. chabanaudi. Symphu- rus leei has four or fewer wide bands (when bands are present) and lacks an opercular spot. The crossbands of S. atricaudus, in turn, are disrupted and incomplete. Sym- phurus leei and S. atricaudus also lack an opercular spot. In S. fasciolaris, the ocular- side of the body often has a number of spherical spots accompanying crossbands 945 (when present), an ocellated spot in the cau- dal fin (absent in S. chabanaudi), and this species lacks the opercular spot character- istic of S. chabanaudi. Symphurus chaba- naudi and S. fasciolaris also differ in caudal- fin ray counts (12 versus 10 in S. fasciolaris). Symphurus chabanaudi and S. elongatus are further distinguished in the length of the upper jaw, which in SS. chabanaudi does not extend beyond a vertical line through the posterior margin of the pupil of the lower eye, whereas in S. e/ongatus it extends to a vertical line through or beyond the posterior margin of the lower eye. Symphurus chaba- naudi also has small ctenoid scales on the blind-side dorsal- and anal-fin rays (absent in S. elongatus) and the posterior taper of the body starts anterior to the body mid- point rather than near the middle of the standard length as in S. elongatus. Addi- tionally, S. chabanaudi has a much larger eye ranging from 1.5-2.1% (¥ = 1.9) SL versus only 0.9-1.5% (X = 1.2) SL in S. elongatus. Symphurus chabanaudi can be further distinguished from S. atricaudus in lacking both a pupillary operculum and small cte- noid scales on the ocular-side dorsal- and anal-fin rays that are present in S. atricau- dus. Symphurus chabanaudi differs most no- tably from S. /eei in having the head length (185-213 SL, ¥ = 201) considerably less than the body depth (239-293 SL, ¥ = 265), in contrast to S. /eei, in which the head length (235-256 SL, ¥ = 245) nearly equals body depth (237-269 SL, X = 253). Symphurus chabanaudi is easily distin- guished from S. me/anurus in the possession of small ctenoid scales extending onto distal portions of the blind-side dorsal- and anal- fin rays and in lacking a pronounced fleshy ridge on the posterior extent of the ocular- side lower jaw. In contrast, in S. me/anurus, scales are either absent or there are 1—2 scales limited to the bases of the fin rays and the posterior extent of the ocular-side lower jaw has a distinct fleshy ridge. The dorsal-fin 946 origin is always posterior to a vertical line through the anterior margin of the upper eye in S. chabanaudi, whereas in S. melanu- rus, the dorsal-fin originates more anteriorly with the first dorsal-fin ray located anterior to a vertical line through the anterior margin of the upper eye. In S. chabanaudi, the pos- terior margin of the jaw does not extend posterior to a vertical line through the pos- terior margin of the lower eye, whereas in S. melanurus, the jaw always reaches a ver- tical line through, or posterior to, the pos- terior margin of the lower eye. Symphurus chabanaudi is readily distin- guished from S. microlepis and S. diabolicus in: having an unpigmented peritoneum (versus black in S. microlepis and spotted in S. diabolicus); in ID pattern (1-5-3 or 1-4-3 versus 1-3-2); hypural number (4 ver- sus 5); and by the number of scales in a longitudinal series (91-104 versus 126 and 135, respectively, in S. microlepis and S. diabolicus). Species of Symphurus possessing 1-5-3 or 1-4-3 ID patterns (patterns occurring in S. chabanaudi) are found only in the New World (Munroe 1987). Western Atlantic Symphurus with similar ID patterns, fin-ray counts, or pigment patterns to those ob- served in S. chabanaudi include S. tessel- latus (Quoy & Gaimard), an undescribed species (species D of Munroe 1987), and S. plagusia (Schneider, in Bloch & Schneider). Symphurus chabanaudi is easily distin- guished from S. plagusia by the large black opercular spot and scales on blind-side dor- sal- and anal-fin rays (both absent in S. pla- gusia); the absence of a fleshy ridge on the ocular-side lower jaw (present in S. plagu- sia); the dorsal-fin origin placed at a vertical line through the middle or anterior margin of the upper eye (versus dorsal-fin origin at a vertical line anterior to anterior margin of upper eye); and meristic values (dorsal-fin rays 98-109 versus 89-98 in S. plagusia; anal-fin rays 82—92 versus 73-81; total ver- tebrae 52-57 versus 47-51; and scales in a longitudinal series 91-104 versus 79-89). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Of all species in the genus, S. chabanaudi is most similar in form, size, and pigmen- tation pattern to S. tessellatus. Symphurus chabanaudi, however, differs from S. tes- sellatus in its modally higher counts of dor- sal- and anal-fin rays and total vertebrae (Table 7). The most useful character to dis- tinguish S. chabanaudi from S. tessellatus is the number of dorsal-fin rays, which range from 98-109 in S. chabanaudi compared with only 91-102 in S. tessellatus. Com- parison of frequency distributions for dor- sal-fin ray counts in these species (Table 7) reveals that about 12% of the specimens overlap with respect to dorsal-fin ray num- ber. However, S. chabanaudi usually has 101 or more dorsal-fin rays (95 of 107 spec- imens) while S. tessellatus usually has 100 or fewer dorsal-fin rays (224 of 233 with 100 or fewer dorsal-fin rays). Corresponding modal differences, although not as great as those noted for dorsal-fin rays, also occur in number of anal-fin rays (82-92 versus 74-86 in S. tessellatus) and total vertebrae (52-57, usually 53-56 in S. chabanaudi ver- sus 48-54, but usually 50-53 in S. tessel- latus). There are also differences in the rel- ative frequencies of occurrence of particular ID patterns in the two species. In S. chaba- naudi, 50% (47/95) of the individuals had a 1-5-3 ID pattern and only 30% (28 spec- imens) featured a 1-4-3 pattern. In contrast, 173 of 233 (74%) S. tessellatus possessed a 1-4-3 ID pattern, while only 6% (13 speci- mens) had a 1-5-3 pattern. Many meristic features of S. chabanaudi completely overlap those of undescribed species D from the Caribbean Sea. Sym- phurus chabanaudi differs from undescribed species D, however, in having 4—8 small, but well-developed scales on the blind-side dorsal- and anal-fin rays (especially prom- inent in specimens larger than 60 mm); a somewhat larger eye (1.5-2.1, ¥ = 1.9 SL versus 1.2-1.9, ¥ = 1.5 SL); and S. chaba- naudi lacks a fleshy ridge on the ocular-side lower jaw (usually present and well developed in undescribed species D). The VOLUME 103, NUMBER 4 947 Table 7.—Comparison of selected meristic features for the eastern Pacific Symphurus chabanaudi and the western Atlantic S. tessellatus. 91 92 93 94 95 96 97 98 S. chabanaudi 1 S. tessellatus 2 1 5 16 S. chabanaudi S. tessellatus be. — Ss Dil Bs} ay AIS Total vertebrae 48 49 50 51 52 53 54 55 S. chabanaudi DD i DP S. tessellatus 1 AD aS Bil Wl By 2) posterior extension of the jaws is slightly more anterior in S. chabanaudi, reaching only to a vertical line through the rear mar- gin of the pupil or rear margin of the lower eye. In undescribed species D, the jaws ex- tend further backwards reaching a vertical line through the posterior margin of the eye and in many specimens, the jaws actually extend slightly beyond the eyes. Symphurus chabanaudi also differs from undescribed species D in the relative frequencies of spec- imens possessing 1-5-3 and 1-4-3 ID pat- terns. Symphurus chabanaudi has a much higher frequency of occurrence of the 1-5- 3 ID pattern (50% of individuals examined) compared with only 30% with a 1-4-3 pat- tern. In contrast, 40 of 45 (89%) of the un- described species D examined had a 1-4-3 pattern and only one specimen possessed a 1-5-3 pattern. Symphurus chabanaudi also differs from undescribed species D in subtle features of its pigmentation. Symphurus chabanaudi generally has about nine wide, dark-brown crossbands; undescribed species D has 10- 14 (usually 10-12) narrower bands. Addi- tionally, the posterior third of the dorsal and anal fins, and the caudal fin in S. chabanaudi are usually uniformly dark brown or black without alternating blotches and unpig- Dorsal-fin rays 99 100 101 102 103 104 105 106 107 «#108 = 109 OR W414 20 6217 AS er Psy MNS) 3 1 Anal-fin rays 82 83 84 85 86 87 88 89 90 91 92 on LO 20235 iy US 3) 33 Sv 268 183 1 No _ 56 37 mented areas. In undescribed species D, the posterior two-thirds of the dorsal and anal fins usually have an alternating series of blotches and unpigmented areas. Additional material examined. — The fol- lowing non-type specimens were also ex- amined (393 specimens, 19-226 mm): Gulf of California western shore: UCLA W54-367, (52, 106-145), about 5 mi north of San Felipe. CAS 24062, (5, 120-133), between Punta Majoro and Punta Ensenada Blanca, about 3 mi north of San Felipe. UCLA W49-427, (2, 152-161), off Ensena- da Blanca, 31°03'30’N, 114°50-51'W. CAS- SU 47384, (1, 101). UCLA W55-26, (1, 160). CAS 24060, (1, 162) and CAS 24071, (11, 119-164), near San Felipe. UCLA W55-2, (13, 99-168), about 9 mi east of San Felipe. YPM 632 (1, 111), San Felipe Bay, 6 m, 19 May 1926. YPM 631 and 633a, (2, 75-122), San Felipe Bay, 6 m, 14 May 1926. CAS 24065, (2, 138-149), 2-3 mi north of Punta Estrella (Diggs), about 7 mi south of San Felipe. CAS 24066, (2, 144-161), about 9 mi south of San Felipe. CAS 24064, (1, 138), about 10 mi south of San Felipe. UCLA W62-61, (22, 66-154), off Punta Estrella (Diggs). UCLA W52-45, (6, 90-182), about 2-3 mi off Punta San Fermin. CAS 24063, (3, 100-150), about 15 mi south of San Fe- 948 lipe. UCLA W59-17, (1, 191), off Punta Willard, Bahia San Luis Gonzaga. USNM 126741, (2, 122-132) and CAS-SU 5571, (2, 126-128), 30°50'45’N, 114°29'45’W, off Isla San Luis. Gulf of California eastern shore: UCLA W49-55, (20, 114-178), 20 mi southwest of El Golfo, Sonora. UCLA W56-28, (10, 156—- 166), north of Punta Lobos, Sonora. UCLA W58-233, (1, 149), north of Punta Lobos, Sonora. UCLA WS50-27, (1, 166), near mouth of Rio Muerto, in tidal waters west of Mapoli, Sonora. CAS 24059, (2, 70-101), Bahia Guaymas, Sonora. CAS 24061, (2, 96-97), just south of channel and east of Punta Baja, Guaymas Harbor, Sonora. UCLA W52-40, (7, 71-105), Guaymas Harbor, Sonora. UCLA WS50-42, (10, 111- 180), south of Boca del Rio Mayo, Sonora. CAS 24068, (1, 122) and UCLA W5S0-43, (2, 19-22) (larvae with eyes recently mi- grated), 26°40'N, 109°47’W, in vicinity of Boca del Rio Mayo, Sonora. UCLA W56- 113, (2, 150-168), south of mouth of Bahia Topolobampo, Sinaloa. SIO 60-95, (1, 218), 24°32.6—33.4'N, 108°03-04.5’W, off Altata, Sinaloa. UCLA W51-36, (2, 98-103), near Astillero at Mazatlan, Sinaloa. Pacific coast of southern Mexico: CAS 4670, (1, 168), near Isla Isabela. SIO 60-87, (2, 173-200), 21°46.4—-50.5'N, 105°25.2- 44.9'W, near mouth of Rio San Pedro, Na- yarit. UCLA W58-18, (12, 43-97), Boca del Asodero, Nayarit. UCLA W58-3, (3, 161- 197), 1-3 mi north of Ensenada Chacala, Nayarit. El Salvador: CAS-SU 46259, (2, 144— 159), 13°20'03”N, 87°48'57’W, off La Union, Gulf of Fonseca. USNM 220804, (7, 122.0-133.7, five specimens cleared and stained), Rio Chaguantique, Jiquilisco Bay, 14 Sep 1976. USNM 291340, (19, 66.7— 123.3), Jiquilisco Bay, Sept 1975—Mar 1976. Nicaragua: CAS-SU 46258, (1, 160), 13°02'30’N, 87°29'30"W, off Punta Mony- penny, Gulf of Fonseca. Costa Rica: SIO 64-465, (1, 179), vicinity of Cabo Blanco. LACM 9754-1, (1, 198), PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON between Isla San Lucas and Isla Negritas, Gulf of Nicoya. UCLA W54-35, (4, 67-101), Erdman Cove, Isla Caballo, Gulf of Nicoya. CAS 24067, (2, 64-106), off Isla Chira, Gulf of Nicoya. UCR 297-16 and 297-17, (7, 150-193), Puntarenas Province, Gulf of Ni- coya. UCLA W54-434, (23, 116-165), exact locality unknown. UMMZ 194670, (2, 199— 206), Gulf of Nicoya, off Puntarenas, Pun- tarenas Province, | Jul 1973. Panama: YPM 4369, (8, 149-201), Canal Zone. ANSP 123572, (2, 174-181), Canal Zone, Oct 1953. CAS 24965, (3, 123-136), 8°58’15”N, 79°25'15”"W. CAS 24962, (7, 146-191), 8°43'15”N, 79°41'W. CAS 24967, (8, 174-204), 8°43'10”N, 79°15’W. SIO 52- 193, (1, 126), 8°40’N, 79°45’W, off Punta Chame. CAS 24963, (14, 152-214), 8°38’N, 78°40'W. CAS 24961, (1, 116), 8°27'15’N, 78°49'50’W, Isla del Ray. UMML 34330, (2, 167-186), 8°19’N, 78°36’ W, 29 m, 7 May 1967. CAS 24964, (1, 162), 8°15'N, 78°26'W. CAS 24968, (2, 156-183), 8°08'15”N, 80°20’W. CAS 24966, (4, 133- 176), San Miguel, Bay of Garachine. UCLA W54-325, (1, 136). UCLA W54-345, (1, 107.5). UCLA W58-278, (1, 208). UCLA W58-304, (20, 82.5—158). UCLA W58-305, (17, 84-170). CAS 24070, (1, 202). LACM 6509-26, (2, 189-193), Bahia de Panama. CAS-SU 6900, (3, 172-226), 8°16'30’N, 79°37'45”"W. USNM 50333, (1, 220). USNM 81674, (1, 43.5) and USNM 81032, (1, 189), from Panama City Market. CAS 24069, (2, 208-212), off Chiman, Gulf of Panama. ANSP 123579, (1, 194), off Chi- man Province, Gulf of Panama, 9 m, 10 Sep 1953. USNM 291357, (1, 161), Canal dredge effluent, Fort Amador, 16 Mar 1967. Colombia: USNM 305725, (3, 160-180), 3°37-39'N, 77°20.5-21'W, Tortugas grounds south of Buenaventura, 18 m, 22 Oct 1970. USNM 305722, (1, 172), 3°39’N, 77°18'W, Tortugas grounds south of Bue- naventura, 9 m, 19 Sep 1969. USNM 305724, (1, 178), 3°31-33'N, 77°22-22.5'W, Tortugas grounds south of Buenaventura, 9 m, 22 Oct 1970. USNM 305720, (1, 202), VOLUME 103, NUMBER 4 3°18-16'’N, 77°33.5-34'W, 9 m, 23 Oct LOTOSTWSNM (3057.23.41, 197), 2°43'N, 77°51'W, Timbiqui, 7 m, 25 Jan 1969. USNM 305721, (1, 205), 1°42'N, 79°00'W, Punta Manglares, 11 m, 28 Jan 1969. Renucn UVIAISPE 2 (1 204). Sols: 81°12'W, Paita. IMARPE 13, (1, 142), 12°06'S, 76°55'W, Lima. Symphurus diabolicus, new species Figs. 2, 4a, Table 8 Holotype.—USNM 135653 (male, 112.6 mm), collected by the U. S. Fish Commis- sion Steamer Albatross, Sta. 2817, 15 Apr 1888, west of Isla San Cristobal (Chatham Island), Galapagos Islands (0°46’S, 89°42'’W), Ecuador, 501 m. Diagnosis.—A Symphurus with a 1-3-2 ID pattern, 109 dorsal-fin rays; 94 anal-fin rays; 12 caudal-fin rays; 58 total vertebrae; 5 hypurals; extremely small scales, 135 in longitudinal series and 58 in transverse row; no pupillary operculum; large, prominent eyes (ED/SNL 0.96), with narrow interor- bital space; upper jaw reaching vertical line just posterior to anterior margin of lower eye; well-developed dentition on ocular-side jaws; origin of dorsal fin at vertical between anterior margin and mid-point of pupil of upper eye; head length slightly larger (1.04 times) than body depth, spotted peritoneum (possibly black in life); and uniform body color without crossbands. Description. —ID pattern 1-3-2. Caudal- fin rays 12 (caudal fin broken and nearly completely severed from body). Dorsal-fin rays 109. Anal-fin rays 94. Pelvic-fin rays 4. Total vertebrae 58; abdominal vertebrae 9 (3 + 6). Hypurals 5. Longitudinal scale rows approximately 135. Scales on head posterior to lower orbit missing. Transverse scales approximately 58. Body medium-sized, relatively elongate, depth at anus 203 SL; greatest body depth 212 SL, beginning about at base of anal-fin ray 5 and continuing almost uniformly over 949 Table 8.—Comparison of meristic and morphomet- ric features for holotypes and only known specimens of Symphurus diabolicus (USNM 135653, male, 112.6 mm) and S. microlepis (MCZ 28535, male, 99.5 mm). S. diabolicus S. microlepis ID pattern 1-3-2 |-3-2 Caudal-fin rays 12 12 Dorsal-fin rays 109 106 Anal-fin rays 94 92 Total vertebrae 58 57 Hypurals 5 S Longitudinal scales 135 126 Postorbital scale rows _ 30 Transverse scale count ca. 58 55 Morphometrics (thousandths of SL) Body depth at anus 203 279 Greatest body depth 212 279 Preanal length 248 271 Dorsal-fin base 938 967 Anal-fin base 756 746 Head length 220 248 Head width 194 PTS Postorbital length 131 176 Upper head lobe 103 110 Lower head lobe 97 170 Predorsal length 62 35 Morphometrics (thousandths of HL, except HW/HL) HW/HL 0.88 Lee Postorbital length 597 708 Snout length 194 154 Upper jaw length 230 231 Eye diameter 186 130 Chin depth 157 109 large area in middle of body (to approxi- mately anal-fin rays 60-70); body taper an- terior and posterior of that region smooth and gradual. Anterior curvature of body not pronounced. Preanal length 248 SL; some- what longer than body depth. Head rela- tively long, 220 SL, greater than body depth. Head length greater than head width (194 SL). Postorbital length 131 SL. Lower head lobe (97 SL) slightly smaller than upper head lobe (103 SL). Snout length 194 HL, with small number of small ctenoid scales. Der- mal papillae scarcely evident on blind-side snout. Anterior nostril relatively long, but not reaching lower eye when depressed pos- 950 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. (00°46'S, 89°42'W). b. Symphurus microlepis, MCZ 28535, holotype; male, 99.5 mm SL; off Panama (7°32.6'N, 79°16'W). teriorly. Evidently 6-8 rows of 2-4 tiny and flimsy scales behind posterior nostril and in narrow interorbital region to posterior mar- gin of eyes (most scales lost; counts made from scale pockets). Mouth relatively large, upper jaw 230 HL, reaching to a vertical line through anterior margin of lower eye. Chin depth 157 HL, shorter than snout length. Lower eye large, 186 HL; eyes slight- ly subequal with upper in advance of lower; eyeballs almost touching. Pupillary oper- culum absent. Length of dorsal-fin base 938 SL. Dorsal-fin origin reaching vertical line through point between anterior margin and mid-point of pupil of upper eye; predorsal length relatively long, 62 SL. Length of anal- fin base 756 SL; blind sides of dorsal and anal fins apparently without scales. Anal-fin origin approximately at vertical line be- a. Symphurus diabolicus, USNM 135653, holotype; male, 112.6 mm SL; Ecuador, Galapagos Islands tween bases of fourteenth and fifteenth dor- sal-fin rays. Pelvic fin with 4 rays; distal half of fin rays broken. Caudal fin damaged pos- teriorly; with four or five rows of tiny scales to point where rays broken. Teeth well developed on both jaws. Blind- side dentary with 5-6 rows of strong teeth across middle of crescentic tooth band. Arms of crescent tapering to fewer rows, ending in single tooth. Premaxilla on blind side with tooth band as broad posteriorly as middle of lower jaw crescent, but narrowing ante- riorly to fewer rows, terminating in single tooth. Ocular-side premaxilla with single row of prominent teeth extending from tip to about mid-point of bone (at point equal with vertical midway between anterior mar- gin of lower eye and posterior margin of anterior nostril). Ocular-side dentary with VOLUME 103, NUMBER 4 row of teeth extending to point below an- terior margin of lower eye. Scales small (smallest among eastern Pa- cific tonguefishes), numerous, ctenoid on both sides of body. Scales on blind side with fewer cteni than those on ocular side (a fea- ture not observed in other eastern Pacific Symphurus). Pigmentation. —Specimen uniformly yel- lowish throughout without visible signs of any other form of pigmentation. Whether yellowish color is natural pigmentation of the species or results from bleaching of nat- ural pigment during fixation and long-term storage is unknown. Etymology. — From the Latin “diabolus” meaning devil, in reference to the large, gro- tesque eyes; an apparent adaptation to the deep-water habitat occupied by this species. Distribution. —Known only from the ho- lotype collected at 501 m on white sand substrate west of Isla San Cristobal (Chat- ham Island) in the Galapagos Islands (Fig. 2). Comparisons. —Symphurus diabolicus is only the fourth known eastern Pacific Sym- phurus with the following combination of characters: a 1-3-2 ID pattern, 12 caudal- fin rays, and a black peritoneum (although the peritoneum in the holotype is spotted and not now solid black, it is thought to have had a black peritoneum and that most natural pigment has been bleached from this specimen during fixation and storage. Four- teen nominal species in addition to S. dia- bolicus are characterized by a 1-3-2 ID pat- tern and 12 caudal-fin rays (Munroe 1987). Of these, ten species, all occurring in deep water (>50 m), have a black peritoneum. Only four species inhabiting relatively shal- low-waters (<45m) have an unpigmented peritoneum.) Other eastern Pacific species characterized by this combination of char- acters include S. microlepis, S. gorgonae, and S. oligomerus. Symphurus diabolicus differs strikingly from these other Symphu- rus with the exception of S. microlepis (see below). Symphurus diabolicus differs from 951 S. gorgonae and S. oligomerus in vertebral counts (total vertebrae 58 versus 46—49 in S. gorgonae and 48-52 in S. oligomerus); numbers of dorsal-fin rays (109 versus 80- 89 in S. gorgonae and 85-97 in S. oligo- merus); numbers of anal-fin rays (94 versus 63-74 in S. gorgonae and 71-83 in S. oli- gomerus);, and body shape (relatively elon- gate with gradual posterior taper in S. dia- bolicus versus much deeper body with greatest body depth in anterior third of body and rapid posterior taper in S. gorgonae and S. oligomerus). Symphurus diabolicus dif- fers further from S. gorgonae in having 5 hypurals (versus 4) and in its much larger size (112.6 mm versus adults smaller than 70 mm). Among all other congeners, S. diabolicus is most similar in morphology and meristic features to S. microlepis, a second eastern Pacific species known only from the holo- type collected in deep-water off the coast of Panama. Although only the types are avail- able for comparison, the differences be- tween these specimens are, nevertheless, substantial and beyond the range of intra- specific variation normally encountered in other species of Symphurus. These differ- ences are particularly noteworthy in light of the fact that both specimens are males and of nearly the same size (112.6 versus 99.5 mm SL), which reduces or eliminates con- founding factors attributable to size or sex- ually related variations in morphology. The two species have similar meristic features, especially ID pattern (1-3-2), total vertebrae (58 versus 57 in S. microlepis), numerous small scales in a longitudinal series (ca. 135 and 126), and caudal-fin rays (12); however, they are easily distinguished by a number of characters (compare Figs. 4a and 4b and see Table 8 for detailed comparison of mor- phological characteristics of both species). Notable differences between these species are body shape and relative size and shape of the head. Symphurus diabolicus has a more elongate body featuring a smooth, gradual taper with the greatest depth (21.2% 52 SL) occurring slightly posterior to the anus and diminishing posteriorly only slightly throughout the middle of the body. In con- trast, S. microlepis has a much deeper body (27.9% SL) with the greatest depth occurring at the anus and the body tapers posteriorly much more rapidly. In S. diabolicus, the head is shorter (HL = 22.0% SL versus 24.8% SL in S. microlepis) and longer than wide (HW/HL = 0.88 versus 1.11) com- pared with that of S. microlepis. Differences in head shape between these species are par- ticularly evident in the relative size of the lower head lobe, which is only 0.94 the width of upper head lobe in S. diabolicus, in con- trast to being 1.6 times larger in S. micro- lepis. Although both species have relatively large eyes, that of S. diabolicus is larger, more elliptically-shaped, and equal in length to about one-fifth of the head length while that of S. microlepis is only about one-eighth of head length and almost spherical. None of the Atlantic Symphurus possess- ing 1-3-2 ID pattern, 12 caudal-fin rays, and black peritoneum have vertebrae or fin-ray counts approaching those of S. diabolicus. Other species in the genus with meristic features similar to S. diabolicus include two rarely collected, Indian Ocean, deep-water species, S. macrophthalmus Norman and S. fuscus Brauer. Symphurus macrophthalmus is a large-eyed, deep-water species known only from the holotype and a single paratype collected in the Gulf of Oman near the Per- sian Gulf (Norman 1939). Large eye size, black peritoneum, and generally slender body are the only similarities this other- wise distinctive species shares with S. dia- bolicus. Symphurus diabolicus has a differ- ent ID pattern than that observed in S. macrophthalmus (1-3-2-2-2 versus 1-2-2-1- 2); fewer caudal-fin rays (12 versus 14); and some higher counts (total vertebrae 58 ver- sus 48; 109 versus 87 dorsal-fin rays and 94 versus 75 anal-fin rays). Certain similarities exist in meristic fea- tures of S. diabolicus and those observed for S. fuscus known only from the holotype col- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lected off the east coast of equatorial Africa (Brauer 1906). Both species have 58 total vertebrae and similar numbers of dorsal- (109 in S. diabolicus versus 105 in S. fuscus) and anal-fin rays (94 versus 93). However, S. diabolicus differs from S. fuscus primarily in ID pattern (1-3-2-2-2 versus 1-2-2-1-2) and caudal-fin ray counts (12 versus 14). Discussion Interestingly, southward increases in sev- eral meristic features occurred in S. oligo- merus and S. chabanaudi (Tables 3, 6). Sim- ilar increases in meristic features occur in other eastern Pacific tonguefishes that range from the Gulf of California to northern Peru, including S. atramentatus, S. fasciolaris, S. leei, S. melanurus, and S. gorgonae (Ma- hadeva 1956). Increased meristic elements in tonguefishes occurring in the southern- most extent of the species range in the Northern Hemisphere contradicts the well- known Jordan Rule (Jordan 1891), in which specimens of Northern Hemisphere species inhabiting the northernmost portions of the species range usually have the highest num- ber of meristic elements. Hubbs (1924, 1926, 1934), in reviewing probable causes of lat- itudinal variation in meristic elements, not- ed strong correlations between the number of elements formed during development and ambient water temperatures, but he recog- nized that other factors such as salinity could also influence the complement of meristic elements formed during ontogeny. One or more of those factors may account for the cline we find in these Symphurus species; however, because virtually nothing is known concerning the seasonality and location of spawning and rates of larval development of these tonguefishes, it is premature to speculate which factors contribute to this trend in these species. Acknowledgments The following curators and support staff generously provided space and arranged for VOLUME 103, NUMBER 4 loan of specimens: W. Smith-Vaniz and W. Saul (ANSP); A. Wheeler, M. Halloway, and J. Chambers BM(NH); W. Eschmeyer, M. E. Anderson, W. Follett, L. Dempster, and D. Catania (CAS); N. Chirichigno (IM- ARPE); R. Lavenberg, J. Seigel, and R. Fee- ney (LACM); K. Hartel (MCZ); C. Hubbs, R. Rosenblatt, D. Gibbons, and H. J. Walk- er (SIO); L. Knapp (SOSC); B. Walker and D. Buth (UCLA); W. Bussing (UCR); C. R. Robins (UMML); R. R. Miller, D. W. Nel- son, and A. S. Creighton (UMMZ); V. G. Springer, R. P. Vari, and S. L. Jewett (USNM); L. P. Schultz (formerly USNM); and K. Thompson (YPM). We thank R. Gibbons and R. Wisner for assistance with photography and radiography. M. Nizinski provided assistance reading radiographs and prepared Fig. 2. The manuscript was im- proved through critical reviews by B. Col- lette, R. Vari, and R. Lavenberg. Portions of this study comprised a partial require- ment for the Ph.D. degree from UCLA to the first author. Portions of this study were independently initiated while the second author was a postdoctoral fellow in the Di- vision of Fishes, USNM. Travel funds en- abling Munroe to work at UMML in 1982 were provided by the Department of Fish- eries, VIMS. Literature Cited Brauer, A. 1906. Wissentschaftliche ergebnisse der deutchen tiefsee expedition auf dem Dampfer ‘Valdivia’, 1898-1899.—Jena (G. Fischer) 15: 1-432. Breder, C. M., Jr. 1936. Heterosomata to Pediculati from Panama to Lower California. Scientific re- sults of the Second Oceanographic Expedition of the “Pawnee”, 1926.—Bulletin of the Bing- ham Oceanographic Collection 2(3):1-56. Chirichigno, F. N. 1974. Clave para identificar los peces marinos del Peru.—Informes Instituto del Mar del Peru 44:1-387. Eschmeyer, W.N.,E.S. Herald, & H. Hammann. 1983.A field guide to Pacific coast fishes from the Gulf of Alaska to Baja California. Boston, Houghton Mifflin Company, 336 pp. Fowler, H. W. 1944. Results of the fifth George Van- 953 derbilt Expedition (1941) (Bahamas, Caribbean Sea, Panama, Galapagos Archipelago and Mex- ican Pacific Islands).—Monograph of the Acad- emy of Natural Sciences of Philadelphia 6:57— 583. Garman, S. 1899. The fishes. Reports on an explo- ration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U. S. Fish Commission Steamer “‘Albatross”’, during 1891, Lieut. Commander Z. L. Tanner, U. S. N., commanding.— Memoir of the Museum of Comparative Zoology, Harvard 24:1-431. Gilbert, C.H., & E.C. Starks. 1904. Fishes of Panama Bay.—Memoir of the California Academy of Sciences 4:3-304. Ginsburg, I. 1951. Western Atlantic tonguefishes with descriptions of six new species. — Zoologica, N.Y. 36:185-201. Hildebrand, S. F. 1946. A descriptive catalog of the shore fishes of Peru.—Bulletin of the United States National Museum 189:1—530. Hubbs, C.L. 1924. Seasonal variations in the number of vertebrae.— Occasional Papers of the Mich- igan Academy of Sciences and Arts 1 1:207-214. 1926. The structural consequences of mod- ifications of the developmental rate in fishes, considered in reference to certain problems of evolution.—American Naturalist 60:57-8 1. 1934. Racial and individual variation in an- imals, especially fishes.—American Naturalist 68:115-128. Jordan, D.S. 1891. Relations of temperature to ver- tebrae among fishes.— Proceedings of the Unit- ed States National Museum 14:107-120. ——, & B. W.Evermann. 1898. The fishes of North and Middle America.—Bulletin of the United States National Museum 47(3):2183-3136. ——., & C.H. Gilbert. 1883. List of fishes collected at Panama by Charles Henry Gilbert. — Bulletin of the United States Fish Commission 2:109- ite ——.,&D.K.Goss. 1889. Areview of the flounders and soles (Pleuronectidae) of America and Eu- rope.— Report of the United States Fish Com- mission for 1886 14:225-342. Lavenberg, R. J., & J. E. Fitch. 1966. Annotated list of fishes collected by mid-water trawl in the Gulf of California, March-April 1964.—California Fish and Game 52:92-110. Leviton, A. E., R. H. Gibbs, Jr., E. Heal, & C. E. Dawson. 1985. Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology.—Copeia 1985:802-832. Mahadeva, M.N. 1956. A review of the tonguefishes of the eastern Pacific, with descriptions of six 954 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON new species. Unpublished Ph.D. dissertation, University of California at Los Angeles, 272 pp. Meek, S. E., & S. F. Hildebrand. 1928. The marine fishes of Panama.—Field Museum of Natural History Publication 249, Zoological Series 15(Part 3):709-1045. Menezes, N. A., & G. De Q. Benvegnu. 1976. On the species of the genus Symphurus from the Bra- zilian coast, with descriptions of two new species (Osteichthyes, Pleuronectiformes, Cynoglossi- dae).— Papeis Avulsos de Zoologia, Sao Paulo 30:137-170. Munroe, T. A. 1987. A systematic revision of Atlan- tic tonguefishes (Symphurus: Cynoglossidae: Pleuronectiformes), with a preliminary hypoth- esis of species group relationships. Unpublished Ph.D. dissertation, College of William and Mary, Williamsburg, Virginia, 598 pp. 1990a. Eastern Atlantic tonguefishes (Sym- phurus: Cynoglossidae; Pleuronectiformes), with descriptions of two new species.— Bulletin of Marine Science 47(3) (in press). 1990b. Symphurus melanurus Clark, 1936, a senior synonym for the eastern Pacific tongue- fishes, S. seychellensis Chabanaud, 1955 and S. sechurae Hildebrand, 1946.—Copeia 1990:229- DED, ——, & M.N. Mahadeva. 1989. Symphurus cal- lopterus (Cynoglossidae, Pleuronectiformes), a new deepwater tonguefish from the eastern Pa- cific.— Proceedings of the Biological Society of Washington 102:458-467. —, & M.S. Nizinski. 1990. Symphurus and S. (Cynoglossidae, Pleuronecti- formes), two new eastern Pacific tonguefishes with 11 caudal-fin rays.—Copeia 1990(4) Gn press). Norman, J. R. 1939. Fishes. John Murray Expedi- tion, 1933-34.—Scientific Report of the British Museum (Natural History) 7:1-116. Phillips, P. C. 1981. Annotated checklist of fishes at Jiquilisco Bay, El Salvador.— Revista de Biolo- gia Tropical 29:45—58. Seale, A. 1940. Report on fishes from Alan Hancock Expeditions in the California Academy of Sci- ences.— Alan Hancock Pacific Expeditions 9(1): 1-46. Wilkinson, L. 1988. SYSTAT: the system for statis- tics. SYSTAT Inc., Evanston, Illinois, 822 pp. (MNM) Department of Biology, Univer- sity of Wisconsin—Oshkosh, Oshkosh, Wis- consin 54901; (TAM) National Marine Fisheries Service Systematics Laboratory, National Museum of Natural History, Washington, D.C. 20560. (All correspon- dence should be addressed to the second author.) _ OF sae aoe PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 955-961 CROCODYLUS RANINUS S. MULLER AND SCHLEGEL, A VALID SPECIES OF CROCODILE (REPTILIA: CROCODYLIDAE) FROM BORNEO Charles A. Ross Abstract.—Crocodylus raninus S. Miller & Schlegel 1844, described on the basis of two syntypes from Borneo, is resurrected and diagnosed. Squamation and cranial morphology of C. raninus are compared with other sympatric and neighboring Indopacific species. The systematics and distribution of In- dopacific Crocodylus are inadequately known. Many museum specimens identi- fied as the widely distributed Crocodylus po- rosus Schneider, 1801, actually represent unrelated and poorly known insular species. This is particularly true in the Indonesian Archipelago, where specimens of unde- scribed Crocodylus have been examined from Banka Island and Sulawesi (Ross 1986). Unfortunately, diagnoses of many of these populations are precluded by insufficient material and complicated by a long history of misidentifications in the literature. One exception is the palustrine crocodile of Bor- neo which, although equally confused in the older literature and known from very few specimens, can be diagnosed on the basis of limited museum material and as an une- quivocal name is available, its resurrection is uncomplicated. Methods Scale counts used in this study were made on 481 individual crocodiles from the In- dopacific region. Specimens were examined from museums, private collections, and from the wild. Some specimens critical to this study in European museums were not examined autoptically but photographs of them were examined and scale counts made from the photographs. Scale counts follow Ross & Roberts (1979), King & Brazaitis (1971), and Bra- zaitis (1973, 1974). Dorsal and neck armor terminology of F. Ross & Mayer (1983) is given in parentheses where pertinent. Museum abreviations follow Leviton et al. (1985). Historical Discussion S. Miller & Schlegel (1844) described Crocodylus raninus (as a race of C. bipor- catus Cuvier = C. porosus) on the basis of an adult skull and a whole juvenile from Borneo. They differentiated C. raninus from C. porosus on the basis of snout shape and neck squamation but the name was not for- mally proposed and specific locality data and disposition of the syntypes were not included. Many crocodile specimens col- lected by M. S. Miller from Borneo in 1836 were in the Muséum d’Histoire Naturelle des Pays-Bas and listed by Lidth de Jeude (1898) in the “Catalogue Ostéologique.” These specimens are individually identified by letter of the alphabet and are now in the Rijksmuseum van Natuurlijke Historie, Leiden. The syntypes of C. raninus were not specifically noted by Lidth de Jeude (1898) and have not been located. They were, how- ever, clearly figured (S. Miller & Schlegel 1844, pl. 3, figs. 7, 8), and show an adult skull (fig. 7) and the neck and head of a juvenile (fig. 8). There is no indication that any subsequent worker examined the syn- 956 types as all later comments refer directly to S. Miiller & Schlegel’s (1844) figures. Gray was clearly confused by the status of S. Miller & Schlegel’s C. b. raninus. In 1844 he synonymized it (in part, the juve- nile syntype) with C. porosus Schneider (1844:58) which he correctly characterized as having reduced nuchal (= postoccipital) squamation. He (1844:62) referred the adult skull to Crocodylus palustris Lesson. Later (1862), he synonymized the adult skull with both C. porosus and C. trigonops Gray (= C. palustris) but failed to mention the ju- venile. He later concluded that the adult skull was C. porosus, and suggested that the juvenile syntype may be a species similar to C. slamensis (1869, 1872). Strauch (1866) identified the juvenile syntype with C. palustris and the adult skull with C. porosus. Bartlett (1895), without re- ferring to earlier authors, presumed that the second species of crocodile occurring in Borneo (that is, other than C. porosus) must be C. palustris but did not examine any specimens of it. More recently several authors, often with- out access to specimens or data, have spec- ulated on the identity of the palustrine, or ‘freshwater,’ crocodile of Borneo. Schmidt (1935) suggested that the name C. raninus may be referable to it. But later authors (Pe- ters 1967; Neill 1971, 1973; Hooijer 1972; Steel 1973; Fuchs 1974; Wermuth & Mer- tens 1977; Groombridge 1982, 1987; Whi- taker & Whitaker 1989) either refer to the Bornean palustrine crocodile as C. siamen- sis or C. palustris (which is restricted to the Indian subcontinent) or ignore it complete- ly. The name Crocodylus raninus has re- mained in the synonomy of C. porosus (Boulenger 1889; Mertens & Wermuth 1955; Wermuth 1953; Wermuth & Mertens 1961, 1977; Werner 1933). Results While examining a large number of In- dopacific crocodiles I reidentified three PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON specimens as a “‘palustrine”’ species possibly referable to C. raninus, following Schmidt (1935). These were a 36 cm long skull lack- ing mandibles ostensibly from Borneo (AMNH 24294) and two preserved juve- niles from Borneo but without further lo- cality data (MCZ 6727 and RMNH 3219) that had previously been identified as C. porosus. The skull (AMNH 24294, Fig. 1) has a transverse palatine-pterygoid suture. In C. porosus this suture is directed posterolater- ally and deeply indents the anterior margin of each pterygoid (Schmidt 1928, 1932, fig. 28 & pl. 7). The lacrymal ridges are less developed and angled from the orbit di- rectly to the posterolateral margin of the frontals (60-—65°), instead of running from the orbit parallel to the frontals as in C. porosus. The skull is robust with heavy sur- face pitting. The maxillaries at the level of the 5th maxillary tooth are distinctly flared outward, accentuating the constriction of the snout at the level of the crocodylid notch at the lateral edges of the maxillary-premax- illary suture. Crocodylus porosus skulls of similar size are more attenuate, lack the ob- vious lateral sculpturing and have less sur- face pitting. The two juveniles have four well-devel- oped postoccipital scales (PC 24—26) (ab- sent or poorly developed in C. porosus; O- 2 scales, n = 60, ¥ = 0.5, +0.83 SD, Fig. 2) and 25 transverse ventral scale rows (po- rosus has 29-35 scale rows n = 67, X = 31.6, +1.29 SD, Fig. 3). The skull (AMNH 24294) differs from another southeast Asian palustrine croco- dile, C. siamensis, which has a broad and rounded snout with weak Sth maxillary tooth bulges, flat or raised nasals forming a promenade Gin AMNH 24294 they are sunken), lacrymal ridges that are poorly de- veloped or absent (in AMNH 24294 they are well developed) and more similar in ori- entation to those of C. porosus in being gen- tly angled inward from the orbits along the frontals (60°). This skull (AMNH 24294) VOLUME 103, NUMBER 4 Fig. 1. Skull of Crocodylus raninus, AMNH 24294 (from top to bottom), dorsal, ventral, and lateral views. 958 NOV MIN RAN SIA PAL POR nN 52 46 NUMBER OF SCALES 80 Fig. 2. Mean number of postoccipital scales (PC 25-26) of Indopacific crocodiles. Numbers at the base of each vertical line indicate number of specimens ex- amined. The vertical line represents the range, the rect- angle is one standard deviation, and the horizontal line represents the mean. Species are identified by the first three letters of their specific names. completely lacks the distinctive interorbital ridge characteristic of C. siamensis (n = 4), and the cranial table is distinctly trapezoidal in shape, being much narrower anteriorly. In C. siamensis the cranial table has parallel sides. The two juveniles differ from C. siamen- sis by having fewer transverse ventral scale rows, 25 (versus 29-33, n = 14, X¥ = 31.3, +1.14 SD, Fig. 3). The anterior throat scales are flat and relatively large and not small and almost granular in appearance as in C. siamensis (Fig. 4). There are 38-39 trans- verse throat scale rows (49-53 in C. Sia- mensis, n = 15, X = 50.3, £1.49 SD, Fig. 5). Both juveniles are in poor condition and it is not possible to detect whether the an- terior ventral caudal scale irregularity of C. siamensis (Brazaitis 1973) is present. On the basis of external morphology, the three Borneo specimens are most similar to Crocodylus novaeguineae Schmidt of New Guinea (Ross 1986), Crocodylus mindor- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON JOH NOV MIN RAN SIA PAL POR 384 364 344 324 3074 ae ue 67 26 | : NUMBER OF SCALES 24-7 224 Fig. 3. Mean number of transverse ventral scale rows of Indopacific crocodiles. Numbers at the base of each vertical line indicate number of specimens ex- amined. The vertical line represents the range, the rect- angle is one standard deviation, and the horizontal line represents the mean. Species are identified by the first three letters of their specific names. ensis Schmidt of the Philippine Islands (Ross & Alcala 1983), and Crocodylus johnsoni Krefft, of northern Australia. They agree with these taxa in having, among other char- acters, similarly well-developed postoccip- ital squamation (Fig. 2), a reduced number of transverse ventral scale rows (Fig. 3), rel- atively symmetrical dorsal armor with weak keels, poorly developed lacrymal ridges, and a transverse palatine-pterygoid suture. In combination these characters separate these crocodiles from all other Crocodylus. The skull (AMNH 24294) can easily be distinguished from C. johnsoni by its robust snout with the dorsal median suture of the premaxillaries nearly separated by the na- sals and barely in contact. In C. johnsoni the snout is attenuate and the dorsal median suture of the premaxillaries is in contact and not separated by the anterior edge of the nasals. Crocodylus mindorensis and AMNH 24294 both have relatively robust snouts es VOLUME 103, NUMBER 4 Fig. 4. Throat squamation of Crocodylus siamensis USNM 76089 (left), and C. porosus BMNH 71.9.1.46 (right), showing the small granular anterior throat scales of C. siamensis and typical Indopacific crocodile throat squamation. with well-developed Sth maxillary tooth bulges and similar lacrymal ridge develop- ment and shape. In both species the squa- mosals are slightly elevated and the cranial table is slightly concave. They differ in the shape of the cranial table, which is nearly parallel sided in C. mindorensis but ante- riorly narrowed in AMNH 24294; the rel- ative width of the dorsal bridge between the supratemporal fenestrae, which is narrow in C. mindorensis; the interorbital bridge, which is concave in C. mindorensis but flat in the other; and the flat nasals of C. min- 959 =x > z = fo ro) ° = < < < ro) =) z = « 77) a a 32 36 40 82 NUMBER OF SCALES 44 33 60 43 52 Fig. 5. Mean number of throat scales of Indopacific crocodiles. Numbers at the base of each vertical line indicate number of specimens examined. The vertical line represents the range, the rectangle represents one standard deviation, and the horizontal line represents the mean. Species are identified by the first three letters of their specific names. dorensis at the level of the 8th maxillary tooth, which are concave in AMNH 24294. The skull is most similar to C. novaegui- neae from the north coast of New Guinea but differs in palatal suture lengths and in the shape of the infraorbital fenestrae, which are more attenuate in C. novaeguineae re- flecting its generally longer and narrower skull shape. The squamosal elevations of similarly-sized C. novaeguineae are better developed than those of AMNH 24294. Conclusions S. Miller & Schlegel’s (1844) pl. 3, fig. 8, clearly shows a crocodile with four well- developed postoccipital scales, a condition I have not observed in C. porosus but agree- ing with the material from Borneo that | 960 examined. The figured skull (S. Muller & Schlegel 1844, pl. 3, fig. 7) likewise agrees in general with the single skull, ostensibly from Borneo, currently available to me. Consequently I conclude the “palustrine”’ crocodile of Borneo is distinct at the species level and that the name Crocodylus raninus S. Muller & Schlegel, 1844, should be res- urrected for this poorly-known population of Indopacific crocodile. There is no evi- dence that more than two species of recent Crocodylus, C. porosus and C. raninus, oc- cur on Borneo, and, as both of the figured syntypes of C. raninus are conspecific and distinguishable from C. porosus, there ap- pears no need to designate a lectotype or neotype for C. raninus. Acknowledgments I would like to thank Marinus S. Hoog- moed (RMNH); Konrad Klemmer (SNM); Roger Bour (MNHN); Jose Rosado and Franklin D. Ross (MCZ); Richard G. Zwei- fel and George Foley (AMNH); George R. Zug and W. Ronald Heyer (USNM); Wil- liam E. Duellman (KU); and, Andrew F. Stimson (BMNH), for permitting the ex- amination of C. siamensis and/or C. rani- nus material used in this study. Franklin D. Ross also provided literature critical to this study. Chip Clark provided photographs used in Fig. 1 and Edward Phillips illus- trated comparative throat squamation (Fig. 4). Innumerable individuals facilitated and assisted with field examination of wild and live crocodiles. In particular I would like to thank Angel C. Alcala, the late Howard W. Campbell, Pedro C. Gonzales, F. Wayne King, and John Lever. I wish to thank Ron- ald I. Crombie and Leslie K. Overstreet for providing photographs of, and scale counts on, critical material in European collections which were not available on loan. Ronald I. Crombie, Storrs L. Olson, George R. Zug, and Gregory C. Mayer read drafts of this manuscript. Peter Brazaitis and James M. Clark reviewed this article for the editor and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON I wish to thank them for their valuable com- ments. Literature Cited Bartlett, Edward. 1895. The crocodiles and lizards of Borneo in the Sarawak Museum with descrip- tions of supposed new species, and the variation of colours in the several species during life.— Journal of the Royal Asiatic Society, Straits Branch 28:73-96. Boulenger, G. A. 1889. Catalogue of the Chelonians, Rhynchocephaliens, and Crocodiles in the Brit- ish Museum (Natural History). Taylor and Francis, London, x + 311 pp. Brazaitis, Peter. 1973. The identification of Crocody- lus siamensis Schneider.— Zoologica 58:43-45. 1974. The identification of living crocodili- ans.— Zoologica 58:(3—4):59-101. Fuchs, Karlheinz. 1974. Die Krokodilhaut. Eduard Roether, Darmstadt, 183 pp. Gray, John Edward. 1844. Catalogue of the Tor- toises, Crocodiles, and Amphisbaenians, in the collection of the British Museum. Edward New- man, London, viii + 80 pp. 1862. A synopsis of the species of croco- diles.— Annals and Magazine of Natural History Ser. 3, 10(58):265-274. . 1869. Synopsis of the species of recent Croc- odilians or Emydosaurians, chiefly founded on the specimens in the British Museum and the Royal College of Surgeons. — Transactions of the Zoological Society of London 6(4):125-169, pls. 31-34. 1872. Catalogue of the shield reptiles in the collection of the British Museum. Part 2. Emy- dosaurians, Rhynchocephalia, and Amphisbae- nians. Taylor and Francis, London, vi + 41 pp. Groombridge, B. 1982. The IUCN Amphibia—Rep- tilia Red Data Book. Pt. 1, Testudines, Crocody- lia, Rhynchocephalia. Unwin Brothers Limited, Old Woking. xliii + 426 pp. 1987. The distribution and status of world crocodilians. Pp. 9-21 in G. J. W. Webb et al., eds., Wildlife management: crocodiles and al- ligators. Surrey Beatty and Sons Pty Limited, Chipping Norton, xiv + 552 pp. Hooyer, D. A. 1972. Pleistocene vertebrates from Celebes. XIV. Additions to the Archidiskodon— Celebochoerus fauna. —Zoologische Mededelin- gen 46(1):1-16, pls. 3-4. King, F. Wayne, & Peter Brazaitis. 1971. Species identification of commercial crocodilian skins. — Zoologica 56(2):15—70. Leviton, Alan E., Robert H. Gibbs, Jr., Elizabeth Heal, & C. E. Dawson. 1985. Standards in herpe- tology and ichthyology: Part 1. Standard sym- VOLUME 103, NUMBER 4 bolic codes for institutional resource collections in herpetology and ichthyology.—Copeia 1985(3):802—-832. Lidth de Jeude, T. W. van. 1898. Catalogue ostéo- logique des poissons, reptiles et amphibies.— Muséum d’Histoire Naturelle des Pays-Bas 10(2): 1-16. Mertens, Robert, & Heinz Wermuth. 1955. Die re- zenten Schildkréten, Krokodile und Briickene- chsen.—Zoologische Jahrbucher (Systematik) 83(5):323-440. Miller, S., & H. Schlegel. 1844. Over de Krokodillen van den Indischen Archipel. 28 pp., pls. 1-3 in C. J. Temminck 1839-1844. Verhandelingen over de natuurlijke geschiedenis der Neder- landsche overzeesche bezittingen, door de leden der Natuurkundige Commissie in Indié en an- dere Schrijvers. Leiden, 259 pp., pls. 1-70. Neill, W. T. 1971. The last of the ruling reptiles, alligators, crocodiles, and their kin. Columbia University Press, New York and London, xvii + 486 pp. 1973. Twentieth-century Indonesia. Colum- bia University Press, New York and London, 391 pp. Peters, Gunther. 1967. Klasse Reptilia—Kriechtiere. Urania Tierreich. Band: Fische, Lurche, Kriech- tiere: Klasse Reptilia—Kriechtiere, pp. 355-507. Ross, Charles A. 1986. Comments on Indopacific crocodile distributions. Pp. 349-354 in Croco- diles. Proceedings of the 7th working meeting of the Crocodile Specialists Group convened at Caracas, Venezuela, 21—28 October 1984. IUCN Publ. NS. —, & Angel C. Alcala. 1983. Distribution and status of the Philippine Crocodile (Crocodylus mindorensis).—Kalikasan (Philippine Journal of Biology) 12(1—2):169-173. —., & Charles D. Roberts. 1979. Scalation of the American Alligator. — Fish and Wildlife Service, Special Scientific Report— Wildlife 225:1-8. 961 Ross, Franklin D., & Gregory C. Mayer. 1983. On the dorsal armor of the Crocodilia. Pp. 305-331 in Anders G. J. Rhodin and Kenneth Miyata, eds., Advances in herpetology and evolutionary biology. Museum of Comparative Zoology, Cambridge. Schmidt, Karl P. 1928. A new crocodile from New Guinea.—Field Museum of Natural History, Publication 247, Zoological Series 12(14):177- 181. 1935. A new crocodile from the Philippine Islands.— Zoological Series of the Field Muse- um of Natural History 20(8):67—70. Steel, R. 1973. Crocodylia.—Encyclopedia of Pa- leoherpetologie 16:vii—116. Strauch, Alexander. 1866. Synopsis der gegenwartig lebenden Crocodiliden.— Mémoires de L’Aca- démie Impériale des Sciences de St.-Pétersbourg (Series 7) 10(13):1-120. Wermuth, Heinz. 1953. Systematik der rezenten kro- kodile. Mitteilungen aus dem Zoologischen Mu- seum, Berlin 29:375-514. — ,&R. Mertens. 1961. Schildkréten, Krokodile Briickenechsen. Gustav Fischer, Jena, xxvi + 422 pp. ——,, & 1977. Das Tierreich. Liste der re- zenten Amphibien und Reptilien. Testudines, Crocodylia, Rhynchocephalia. Walter de Gruy- ter, Berlin, xxvii + 174 pp. Werner, Franz. 1933. Das Tierreich. Loricata. Walter de Gruyter, Berlin and Leipzig, xiv + 40 pp. Whitaker, Rom, & Zai Whitaker. 1989. Status and conservation of the Asian crocodilians. Pp. 297- 308 in Crocodiles. Their ecology, management, and conservation. IUCN Special Publication, NS. Department of Vertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 962-965 A NEW SUBSPECIES OF DIGLOSSA GLORIOSTSSIMA (AVES: THRAUPINAE) FROM THE WESTERN ANDES OF COLOMBIA Gary R. Graves Abstract. —A new subspecies of flower-piercer, Diglossa gloriosissima boylei, is described from two localities at the northern end of the Western Cordillera of the Colombian Andes in the Department of Antioquia. The nominate race is apparently restricted to the highest part of the Western Cordillera west of Popayan, Department of Cauca. The Chestnut-bellied Flower-Piercer, Diglossa gloriosissima (Chapman 1912), an allospecies of the Diglossa lafresnayii su- perspecies (Vuilleumier 1969), is known from only three isolated paramos in the Western Cordillera of the Colombian Andes (Hilty & Brown 1986) (Fig. 1). Specimens are scarce in collections and those reported in the literature were collected on two ex- peditions sponsored by the American Mu- seum of Natural History (AMNH) in the early part of this century. Leo E. Miller and W. B. Richardson collected the type series of ten specimens (six adults) in July 1911 on the crest of the coast range west of Po- payan (Chapman 1912), and Miller and Howarth Boyle collected a series some 500 km north of that locale on Cerro Paramillo in January 1915 (Chapman 1917, Zimmer 1929). The species was discovered at a third locality in August 1951 by Melbourne A. Carriker, Jr., who collected a series of six adults on Paramo Frontino for the National Museum of Natural History (USNM), Smithsonian Institution. In a systematic review of the D. lafres- nayil superspecies, Zimmer (1929:30) noted that, “specimens of gloriosissima from the western Andes near Popayan show consid- erable black along the flanks, and in this respect approach closer to /afresnayii than do birds from Paramillo. ...’’ Contrary to his penchant for attaching subspecific names to the most subtly marked populations, Zimmer declined to designate the speci- mens from Cerro Paramillo as a new sub- species. With the exception of the type, the adult specimens from Popayan were dispersed to other museums soon after Chapman’s de- scription of the species. I was able to ex- amine three of these, most of the adults from Paramo Paramillo, and all of Carriker’s specimens from Paramo Frontino. My anal- yses confirm Zimmer’s observations and in- dicate that the northern populations from Paramo Frontino and Cerro Paramillo rep- resent an undescribed subspecies. Materials and methods. —Measurements of wing chord, tail from point of insertion of central rectrices to tip of longest rectrix, tarsus, and bill from anterior edge of nostril, were taken with digital calipers (Table 1). Wing and tail measurements were rounded to the nearest millimeter, those of tarsus and bill to the nearest 0.1 mm. T-tests were per- formed on untransformed data with SYS- TAT software. Diglossa gloriosissima boylei, new subspecies Holotype. — National Museum of Natural History, Smithsonian Institution (USNM) No. 436792; male in adult plumage from Paramo Frontino, 11,850 ft (ca. 3613 m), VOLUME 103, NUMBER 4 Department of Antioquia, Colombia, col- lected 25 Aug 1951 by Melbourne A. Car- riker, Jr. (original number 21013). Diagnosis. —Diglossa gloriosissima boylei differs from D. g. gloriosissima in having uniformly chestnut flanks, sides, and un- dertail coverts, these regions in D. g. glo- riosissima being sooty black or chestnut with black markings and spots. Measurements of Holotype (mm).— Wing (chord) 68; tail 60; tarsus 22.9; bill 10.5. Geographic range. — As far as known, re- stricted to elfin forest at timberline and patches of shrubbery in paramo on Paramo Frontino and Cerro Paramillo, Department of Antioquia, Colombia. Specimens examined.—Diglossa g. glo- riosissima: Coast range west of Popayan, Department of Cauca (AMNH Holotype, é; USNM 6; FMNH 2; MCZ 8). D. g. boylei: Paramo Frontino, Department of Antio- quia (USNM including Holotype, 6 38); Cer- ro Paramillo (AMNH 4 4é, 2 92; USNM 1 6; MCZ 34). Specimens of D. gloriosissima were compared directly with series of all named taxa of the D. lafresnayii (Vuilleu- mier 1969) and D. carbonaria superspecies (Graves 1982). Etymology.—I am pleased to name this flower-piercer for Howarth S. Boyle, who with Leo Miller, made the first collections of D. glorisissima boylei. 963 Fig. 1. Distribution of specimens of D. gloriosis- sima examined in this study from the Western Cor- dillera of the Colombian Andes. D. g. gloriosissima: W. Popayan (circle). D. g. boylei: Paramo Frontino (diamond); Cerro Paramillo (triangle). Degrees of north latitude are labeled along the y-axis. Dotted lines out- line the 1000 m contour interval. Hatching indicates areas above 3000 m elevation. Table 1.—Ranges and means of measurements (mm) of Diglossa gloriosissima. Subspecies Sex n Wing chord Tail Tarsus Bill D. g. gloriosissima W Popayan foto} 4 74-77 64-67 24.0-24.9 9.2-9.8 es TSS Na—16005 aa X¥ = 9.4 D. g. boylei Paramo Frontino rele} 6 68-71 59-63 22.9-24.3 9.5-10.5 X¥ = 69.7 X¥ = 60.2 X = 23.7 X = 9.8 Cerro Paramillo rok} 6 TA—=T15 64-65 23.5—24.6 9.9-10.7 VCS MPe af X = 64.6 X = 242 ¥ = 10.1 99 2 69-70 63 22.5-23.9 10.3-10.4 964 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—One-tailed t-tests of character means of male population samples of Diglossa gloriosissima (see Table 1). Significant t values, adjusted for the number of simultaneous tests (P = 0.05/12 = 0.004), are indicated by *. Population contrasts Wing chord Tail Tarsus Bill Popayan versus Paramo Frontino 5.00* 4.84* DSP 1.70 Popayan versus Cerro Paramillo 2.35 1.06 1.10 3.58 Paramo Frontino versus Cerro Paramillo 3.80* 6.04* 1.83 1.45 Results There appear to be significant size differ- ences among populations of D. gloriosissi- ma, with specimens from Paramo Frontino, geographically interposed between Popayan and Cerro Paramillo, having shorter wings and tails than those from the other localities (Table 2). Because intraspecific size varia- tion in birds may reflect environmental in- duction as well as genetic differentiation (James 1983), it is deemed unwise to rec- ognize these differences nomenclaturally. Moreover, significant geographic size vari- ation occurs within several subspecies of Diglossa carbonaria that exhibit little or no variation in plumage color and pattern. In these cases, I consider size as a taxonomi- cally unreliable character. On the other hand, qualitative plumage differences, such as those expressed among populations of D. gloriosissima, are thought to be genetically determined. The avifauna of the Western Cordillera is poorly known relative to other parts of the Colombian Andes. As an example, Hilty & Brown (1983) documented 63 major range extensions of species in the Western Cor- dillera based on previously unreported ma- terial collected by Carriker in the 1940’s and 1950’s. Although populations of D. glorio- sissima and other taxa now restricted to Cerro Paramillo and Paramo Frontino (e.g., Coeligena orina, Metallura williami recisa) may occur on several of the ornithologically unexplored peaks between Popayan and Paramo Frontino, the insular nature of hab- itat above 3000 m elevation suggests that gene flow among populations of D. glorio- sissima has been drastically reduced since the Wisconsinan glacial (Graves 1980). Acknowledgments I thank Richard Banks, Scott Lanyon, and Storrs Olson for comments on the manu- script and the curators of the American Mu- seum of Natural History (AMNH), Field Museum of Natural History (FMNH), and Museum of Comparative Zoology, Harvard University (MCZ) for permission to ex- amine specimens. Museum work was par- tially funded by the Frank M. Chapman Memorial Fund of the American Museum. Literature Cited Chapman, F.M. 1912. Diagnoses of apparently new Colombian birds.—Bulletin of the American Museum of American History 31:139-166. . 1917. The distribution of bird-life in Colom- bia: a contribution to a biological survey of South America.— Bulletin of the American Museum of Natural History 36:1-729. Graves, G. R. 1980. A new subspecies of Diglossa (carbonaria) brunneiventris.—Bulletin of the British Ornithologist’s Club 100:230-232. 1982. Speciation in the Carbonated Flower- piercer (Diglossa carbonaria) complex of the Andes.— Condor 84:1-14. Hilty, S. L., & W. L. Brown. 1983. Range extensions of Colombian birds as indicated by the M. A. Carriker Jr. collection at the National Museum of Natural History, Smithsonian Institution. — Bulletin of the British Ornithologist’s Club 103: 5-17. ——.,, & . 1986. A guide to the birds of Co- lombia. Princeton, New Jersey, Princeton Uni- versity Press, 717 pp. James, F. C. 1983. Environmental component of morphological variation in birds.—Science 221: 184-186. VOLUME 103, NUMBER 4 Vuilleumier, F. 1969. Systematics and evolution in Diglossa (Aves, Coerebidae).—American Mu- seum Novitates 2381:1-44. Zimmer, J. T. 1929. Variation and distribution in two species of Diglossa. — Auk 46:21-37. 965 Department of Vertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 103(4), 1990, pp. 966-972 THE HOLOTYPE OF NATALUS STRAMINEUS GRAY (MAMMALIA: CHIROPTERA: NATALIDAE) Charles O. Handley, Jr. and Alfred L. Gardner Abstract.—-The description of Natalus stramineus Gray was based on an unspecified number of specimens of unknown provenance. We review the critical specimens and their history in the mammal collections of the British Museum (Naturai History). We identify the holotype of NV. stramineus, and we believe that Goodwin (1959) was correct in claiming that it originated in the West Indies rather than in South America. In the original description of Natalus stra- mineus, Gray (1838) did not specify the number of specimens examined and indi- cated that he did not know the origin of his material. Later Gray (1843) said that he had a fluid-preserved specimen from South America and another (presumedly dry) from St. Blas, North America; this information was repeated by Tomes (1856). Dobson (1878) listed a male in alcohol from Brazil, an unsexed skin from South America, and a male in alcohol from Duenas, Guatemala. Authors subsequent to Dobson gave Brazil as the type locality of N. stramineus. Good- win (1959), however, suggested with good reason that the type came from the West Indies, not Brazil as previously supposed. When he shifted the type locality from Bra- zil to the West Indies, Goodwin synony- mized Shamel’s (1928) West Indian Natalus dominicensis with N. stramineus, and re- named the Brazilian population JN. stra- mineus natalensis. Goodwin (1959) argued correctly that the specimen in the British Museum (Natural History) he believed to be the holotype of N. stramineus was the same as that listed by Gray (1843) and Tomes (1856) from South America and by Dobson (1878) from Brazil. Following Goodwin (1959), the nomenclature of N. stramineus appeared stable, but some au- thors (Carter & Dolan 1978, Honacki et al. 1982) have continued to list Brazil as the type locality. We intend to set the record straight in this report. The Evidence In 1971, Handley examined the putative holotype of N. stramineus in the British Mu- seum, reviewed its history in collections of the Mammal Section, and attempted to de- termine its provenance. However, there are alternative possible types or syntypes of N. stramineus in the British Museum that rep- resent other populations, and the situation is complicated by apparent multiple rela- beling of specimens. Because the clues lead- ing to recognition of the holotype and a sec- ond specimen reported by Gray in 1843 come from a number of sources, we list the evidence in considerable detail. Gray 1838.—Gray (1838:496) described Natalus as a new genus and N. stramineus as a new species. He gave no morphological measurements, but indicated that he actu- ally had a specimen by his statement, “‘in- habits ? British Museum.” By impli- cation he had at least one specimen of unknown origin. Gray’s mss. catalogue.—Soon Gray had another specimen, as his undated hand- written catalog of Primates and Chiroptera in the British Museum (referred to in lit- erature as ‘““Gray’s Mss. Catalogue’’) shows the following entry: “‘69./ Vespertilio lon- VOLUME 103, NUMBER 4 gicaudatus, Gray Mss/ Natalus stramineus Gray Mag Zool & Bot. 1837 [sic]/ a. S. America/ b. St. Blass/ Purchased of Mr. Gould/ 42.8.17.10.” Museum register 1842.—The museum register has the following information for number 42.8.17.10 (in the registry system of the Mammal Section this was the tenth specimen registered on 17 August 1842): ““Vespertilio/ St. Blas/ Purchased of Mr. Gould,/ Cheirop. 69b.” Gray 1843.—The data from Gray’s mss. catalogue and the museum register were re- peated in Gray’s published “Catalogue” (1843:28) with important additions: “The Natale. Natalus stramineus, Gray, Mag. Zool. and Bot. I. 14. Vespertilio longicaudatus, Gray, Brit. Mus. a. In spirits. South America. b. N. America, St. Blas.” In the British Museum copy of his 1843 “Catalogue,” Gray inked the figure “69” in the margin beside specimen “‘a,”’ referring back to his ‘““Mss. Catalogue”” number. Rec- ognition of handwriting in books, papers, and specimen labels in the Mammal Section of the British Museum is a relatively simple task because the Section has preserved sam- ples of handwriting of all who have worked there. Evidently, by August 1842, there were only two specimens of N. stramineus in the British Museum. The facts that specimen ““69’°a was from an indefinite locality and that it was preserved in spirits are two im- portant bits of information. Clearly the ho- lotype of N. stramineus must be “69” a, be- cause b was from a definite locality, and Gray’s original reference (1838) specified ‘inhabits ? British Museum.” Tomes 1856.—Tomes (1856:176-178, pl. 43) redescribed the genus Natal/us and the species N. stramineus in great detail. He concluded with the statement, “The whole of the above [description] has been taken, by the kind permission of Dr. Gray, from the two examples mentioned in his Cata- 967 logue [1843], and the following are their di- mensions. The first column [No. 1.] refers to the specimen in spirits from South Amer- ica, and the second [No. 2.] to the one from St. Blas, North America.”’ A table of mea- surements of the two specimens followed. Plate 43 is colored, shows great detail, has accurate proportions, and seems to have been drawn life size. Although its source was not specified, measurements taken from the figure on Plate 43 coincide closely with those listed by Tomes for his specimen No. 2, the one from “‘St. Blas.’’ Thus, at the time of Tomes’ writing there continued to be only two specimens of N. stramineus in the Brit- ish Museum. The specimen labeled “‘South America” and preserved in spirits must be the holotype Gray described in 1838. It is the larger of the two. The other, evidently dry, labeled “St. Blas,” and figured by Tomes (1856:pl. 43), is smaller. Dobson 1878. — Dobson (1878:343) listed three specimens of N. stramineus in his cat- alogue of Chiroptera in the British Museum and gave the distribution of the species as “Brazil; Central America.”” The first two specimens were listed as: Brazil. South America. a. 6 ad., al. b. ad. sk. The third was listed on the same page under Var a. as: Duenas, Guatemala. O. Salvin, Esq. [C.]. Dobson (1878:344) gave a table of exter- nal measurements of two of the specimens. The first column is headed “NV. stra-/mi- neus” and the second, “‘Var a.”> The mea- surements under N. stramineus must have been of either Dobson’s specimen “a” or “*h.*? Some time later, Thomas wrote “type” in the margin beside specimen “a. é ad., al. Brazil” in the Mammal Section’s copy of Dobson’s catalogue. Cabrera 1958.—Cabrera (1958:73) re- stricted the type locality of N. stramineus to Lagoa Santa, Minas Gerais, Brazil, on a. 6 ad., al. 968 the basis of Winge’s (1892) report of the first definite locality for the species from the country where Dobson (1878) said the type had originated. Goodwin 1959. —Goodwin (1959:4—5, 16), on the basis of measurements and cra- nial morphology, stated that the holotype of N. stramineus matched representatives of populations inhabiting the Lesser Antil- les. As is evident from material in the mam- mal collections of the National Museum of Natural History, Washington, D.C., the Brazilian population differs sufficiently from that of the Lesser Antilles to make confu- sion of specimens between the two popu- lations unlikely. On the same basis, Good- win (1959) ruled out Venezuela, Trinidad and Tobago, Central America, and México as possible origins. He said that Gray’s type could not have come from Brazil or any- where else in South America and proposed restricting the type locality to the island of Antigua, British West Indies. Carter & Dolan 1978 and subsequent re- ports. —Carter & Dolan (1978) said that of the two specimens of Natalus stramineus reported by Gray (1843:23), the one in fluid labeled Brazil was the only identifiable one remaining. They said it was an unnumbered adult male in alcohol (with skull removed), and gave the type locality as Lagoa Santa, Minas Gerais, as restricted by Cabrera (1958: 95). Because they reported this specimen as unnumbered, their information must have come from Carter’s visit to the British Mu- seum in 1966. In 1970 this specimen was registered as No. 70.2324. Presumably, it was during 1966 and not when they revis- ited the British Museum in 1976 that Carter & Dolan (1978:11) were unable to find the second specimen listed by Gray (1843) from ““N. America, St. Blas.”? Apparently misled by Gray’s (1843) mention of two specimens, Carter & Dolan (1978) presumed them to be syntypes. They also commented that Goodwin (1959), in disagreement with Ca- brera (1958), designated the type locality as PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Antigua, Lesser Antilles. They probably fol- lowed Cabrera’s restriction of the type lo- cality to Lagoa Santa, Minas Gerais, be- cause of Dobson’s jar label, which said Brazil. Hall (1981) followed Goodwin’s des- ignation of Antigua; but Honacki et al. (1982) followed Cabrera’s (1958) restric- tion. The specimens. — All three specimens list- ed by Dobson were still in the British Mu- seum in February 1971 when Handley ex- amined them, and in October 1987 when Gardner examined them. Numerous other specimens of Natalus stramineus now in the British Museum bear dates later than 1878, have definite locality data, and are not rel- evant in this discussion. Presuming that it still exists, the holotype of N. stramineus must be one of the three specimens listed by Dobson (1878:343). The oldest label at- tached to each of these three specimens was written by Dobson himself. Presumably, he was responsible for changes where the label data differed from those published by Gray (1838, 1843) and Tomes (1856). The male in alcohol with skull removed and cleaned is labeled ““Natalus stramineus Gray/ (G. E. Dobson).” This specimen was finally registered in 1970, and at that time the jar label was emended to read: ““Type/ 6/ Natalus stramineus, Gray/ 70.2324/ Brazil.”’ Unfortunately and unaccountably, 15 of the 20 finger bones and both tibiae of this specimen have been broken. Otherwise its condition was reasonably good in 1971. The second specimen is an unsexed, dry skin with skull inside that lacked a registry number in 1971. It is labeled ‘“Natalus stra- mineus, Gray/ (G. E. Dobson) ? (type) South America,” all in Dobson’s handwriting. The third specimen, number 75.2.27.72, a male in alcohol with skull removed and cleaned, was labeled “‘Natalus stramineus/ (G. E. Dobson)/ Duenas, Guatemala/ O. Salvin, Esq.” It agrees morphologically with other specimens of Natalus from Guate- mala, and because there is no reason to doubt VOLUME 103, NUMBER 4 the accuracy of its label information, it can be eliminated from the search for the ho- lotype of N. stramineus. Discussion The specimens. — The situation thus posed is of a single specimen of N. stramineus de- scribed by Gray in 1838 without data, two specimens in the period 1842-1856 with data (Gray 1843, Tomes 1856), and two specimens in 1878 (Dobson) with different data. Correlation of the measurements (Ta- ble 1) from Tomes’ (1856) table and plate, Dobson’s (1878) table, and the two speci- mens in the British Museum in 1971 with the information published by Gray (1838, 1843), Tomes (1856), and Dobson (1878), and with the present label data of the spec- imens leads to the conclusion that only two specimens have been involved throughout, that the label data for the holotype changed at least twice, and that neither specimen was properly labeled in 1971 (Table 2). The coincidence of measurements is too great (see Table 1) to suggest other than that the “b” of Gray (1843), specimen BM 42.8.17.10 from St. Blas, ““No. 2” and plate 43 of Tomes (1856), ““b” of Dobson (1878), and the unregistered specimen labeled “South America” in 1971 are all the same. This specimen, which agrees morphologi- cally with Natalus from western México, is the other implied “‘syntype”’ Carter & Dolan (1978) were unable to recognize because it was labeled ““South America”? when Carter examined it in 1966. Evidently Dobson, or someone else between 1856 and 1878, con- fused the data and transferred ‘‘South America” from the holotype (which origi- nally had been without locality data) to the San Blas specimen. As of 1971, no bat in the British Museum was labeled “St. Blas” Of 42.6. li alO The possibility exists that specimen “‘a”’ of Dobson (1878) from “Brazil” (presently no. 70.2324) was a new specimen obtained 969 Table 1.— Measurements (in millimeters) of the ho- lotype (BM 70.2324) and contemporaneous specimen (BM 42.8.17.10) of Natalus stramineus from a table in Tomes (1856:178), plate XLIII in Tomes (1856: mea- sured by Handley), a table in Dobson (1878:344), and the specimens in the British Museum (Nat. Hist.) mea- sured by Handley in 1971. Measurements in brackets indicate approximations. Tomes (1856) Dobson Measurement (1878) Specimens 1971 Specimen Table Plate Table (left/right) Forearm 70.2324 331/55) 37.9 38.3/38.1 42.8.17.10 35:45 36t5 [35.9]/ Tibia 70.2324 20.1 19.1 [19.0]/ 42.8.17.10 GEO A? 18.0/ Third finger 70.2324 76.2 71.7/72.4 42.8.17.10 69.9 66.0 67.3/ Fourth finger 70.2324 55.0 54.0/53.5 42.8.17.10 48.9 50.5 51.6/ sometime between 1856 and 1878, and had nothing to do with Gray’s type of N. stra- mineus. However, again based on the co- incidence of measurements given by Tomes (1856) and Dobson (1878), and those of specimen BM 70.2324 (Table 1), as well as the mode of preservation, the evidence is compelling that BM 70.2324 is the speci- men Gray had at hand in 1838 when he described N. stramineus. The labels. —Having established that Gray (1838) had only one specimen before him when he described N. stramineus, and hav- ing identified that specimen from among others now in the British Museum, we at- tempt to account for the several changes in localities assigned to it (see Table 2). We start with Gray’s original statement (1838) that the provenance of the holotype was un- known. One can see how Gray later could have come to label it “South America.” Re- ceipt of the second specimen from a definite locality, ““St. Blas’ (= San Blas, Nayarit, 970 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—An outline and chronology of the information available on specimens of Natalus stramineus reported by Gray in 1838 and 1843. Reference Specimen Specimen Gray (1838) “Inhabits ___?”” (holotype of N. stramineus) Gray (ca. 1842, S. America St. Blass mss. catalogue) “‘Natalus stramineus Gray “purchased of Mr. Gould” Mag. Zool. & Bot. 42.8.17.10 1837 [sic] 69.a 69.b Gray (1843) South America North America St. Blas In spirits [Dry?] a. b. Tomes (1856) South America North America St. Blas In spirits [Dry?] No. 1 in table of measurements No. 2 in table of measurements Dobson (1878) Brazil South America In alcohol [Dry?] N. stramineus in table of measurements a. b. Specimen labels Brazil South America (1971) In alcohol Dry BM 70.2324 Unregistered Goodwin (1959) Antigua, Lesser Antilles Conclusions Origin unknown San Blas, herein [Lesser Antilles] Nayarit, México BM 70.2324 42.8.17.10 Mexico), must have prompted him to as- sume that the unlabeled holotype also came from somewhere in Latin America and led him to label it, in the broad sense, as from “South America” (a common euphemism for Latin America). If these events are prop- erly reconstructed, this was the first of other erroneous relabelings of this specimen. Emending labels or relabeling, practices at the British Museum that began with Gray and were continued by Tomes, Dobson, and Thomas, have produced some label data that are unreliable or suspect. Another potential problem with Mammal Section spirit spec- imens is that, in the 19th Century, usually the jars rather than the specimens them- selves were labeled, leading to possible con- fusion between jars or within jars when they contained more than one specimen (e.g., Smith 1971:80-81). Although we can understand how the ho- lotype of N. stramineus could have been erroneously labeled “‘South America,” it is a mystery how Dobson (1878) came up with the label “‘Brazil’’ for it. Possibly, when re- labeling specimens, Dobson’s selection of “Brazil” as the origin of the holotype was influenced by the similarity between the name Natalus and Natal, the capital of the Brazilian State of Rio Grande do Norte (eventually to be the type locality of N. stra- mineus natalensis Goodwin, 1959). It seems unlikely that he could have mistaken the handwritten “St. Blas” in Gray’s “Mss. Cat- alogue’’ for Brazil. Equally unlikely is the possibility that he got new information di- VOLUME 103, NUMBER 4 rectly from Gray. Dobson and Gray over- lapped only briefly at the British Museum. Dobson, an army surgeon who came to Lon- don in 1873, took up the project of listing the Chiroptera in the British Museum while he continued medical work at Nethley Hos- pital. Gray suffered a stroke in 1869 and, although paralyzed on the right side, he re- turned to the Museum after a few months and continued to work there until the latter part of 1874. He died in 1875 (Ginther 1912). Besides, if Gray had had new infor- mation, surely it would have been used by Tomes. There is nothing to suggest that Dobson’s “Brazil” and “South America” localities for two of the three specimens he reported were based on actual information on their origins. We are not aware of any South American populations clearly assignable to N. stra- mineus except for the widespread Brazilian population Goodwin (1959) named N. stra- mineus natalensis. Cuervo D. et al. (1986) listed N. stramineus for Colombia, but in- dicated neither specimens nor localities. According to Goodwin (1959) the two spec- imens Sanborn (1941) listed as N. strami- neus from Trinidad proved to be N. tumi- dirostris. Goodwin (1959) also pointed out that the Guyanan (British Guiana) speci- men identified as N. stramineus by Jentink (1893:79), Young (1896:44), and Beebe (1919:219), is a Furipterus horrens. Conclusions Gray (1838) had only a single specimen before him when he described Natalus stra- mineus aS a new genus and species. It was an adult male in spirits. Its origin was not known, but we believe as did Goodwin (1959) that it came from the Lesser Antilles (restricted type locality, Antigua). It is BM 70.2324, erroneously labeled “Brazil,” and preserved in fluid with skull removed. The second specimen is a dry skin with skull inside from San Blas, Nayarit, Mexico (= St. Blas, North America), first mentioned in Gray’s mss. catalogue, probably in 1842. It is BM 42.8.17.10; but is erroneously la- 971 beled “South America’”’ without registry number. It is an example of N. stramineus saturatus Dalquest & Hall, 1949. Acknowledgments We express our appreciation to the staff of the Mammal Section of the British Mu- seum, and particularly to Mr. J. E. Hill, for the many courtesies and assistance gra- ciously accorded us during our work in the mammal collections. Darelyn Handley was indispensable in the search for information in London in 1971 and in assembly of early drafts of this manuscript. Literature Cited Beebe, W. 1919. High vertebrates of British Guia- na.— Zoologica 2:205—227. Cabrera, A. 1958. Catalogo de los mamiferos de América del Sur.— Revista del Museo Argenti- no de Ciencias Naturales ““Bernardino Riva- davia,” Ciencias Zoologicas 4(1):iv + 308 pp. Carter, D. C.,& P.G. Dolan. 1978. Catalogue of type specimens of Neotropical bats in selected Eu- ropean museums.—Special Publications the Museum, Texas Tech University 15:1-136. Cuervo D., A., J. Hernandez C., & A. CadenaG. 1986. Lista actualizada de los mamiferos de Colombia anotaciones sobre su distribucion. — Caldasia 15: 471-502. Dalquest, W. W., & E. R. Hall. 1949. A new sub- species of funnel-eared bat (Natalus mexicanus) from eastern Mexico.—Proceedings of the Bi- ological Society of Washington 62:153-154. Dobson, G. E. 1878. Catalogue of the Chiroptera in the collection of the British Museum. British Museum (Natural History), London, xlii + 567 pp., 30 pls. Goodwin, G. G. 1959. Bats of the subgenus Natal- us. —American Museum Novitates 1977:1-—22. Gray, J. E. 1838. A revision of the genera of bats (Vespertilionidae), and the description of some new genera and species. — Magazine of Zoology and Botany 2:482-S0S. 1843. List of the specimens of Mammalia in the collection of the British Museum. British Museum (Natural History), London, xxviii + 216 pp. Giinther, A. 1912. General history of the Department of Zoology from 1856-1895. Pp. i-ix + 1-109 in The history of the collections contained in the natural history departments of the British Museum. British Museum (Natural History), London, Vol. 2, Appendix. 972 Hall, E.R. 1981. The mammals of North America. Second edition. J. Wiley & Sons, New York, 1: xv + 1-600 + 90. Honacki, J. H., K. E. Kinman, & J. W. Koeppl. 1982. Mammal species of the world. Allen Press, In- corporated, and The Association of Systematics Collections, Lawrence, Kansas, 694 pp. Jentink, F. A. 1893. On a collection of bats from West-Indies.— Notes Leyden Museum 15:278- 283. Sanborn, C. C. 1941. Descriptions and records of Neotropical bats.—Field Museum of Natural History, Zoological Series 27:371-387. Shamel, H. H. 1928. A new bat from Dominica.— Proceedings of the Biological Society of Wash- ington 41:67-68. Smith, J. D. 1971. Review of the Mormoopidae.— Miscellaneous Publication Museum of Natural History, University of Kansas 56:1--132. Tomes, R. E. 1856. On three genera of Vespertilion- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON idae, Furipterus, Natalus and Hyonycteris, with descriptions of two new species.— Proceedings of the Zoological Society of London 1856:172- 181, pls. 42 and 43. Winge, H. 1892. Jordfundne og nulevende Flagermus (Chiroptera) fra Lagoa Santa, Minas Geraes, Brasilien.—E Museo Lundii, Kjobenhavn 2(part 1):1-65, 2 pls. Young, C.G. 1896. Notes on Berbice bats. —Timehri, New Series 10:44—46. (COH) Division of Mammals, National Museum of Natural History, Washington, D.C. 20560; (ALG) Biological Survey Field Station, National Ecology Research Center, U.S. Fish and Wildlife Service, National Museum of Natural History, Washington, D.C. 20560. OO INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE Applications published in the Bulletin of Zoological Nomenclature The following applications were published on 29 June 1990 in Vol. 47, Part 2 of the Bulletin of Zoological Nomenclature. Comment or advice on these applications is invited for publication in the Bulletin and should be sent to the Executive Secretary, I.C.Z.N., % The Natural History Museum, Cromwell Road, London SW7 5BD. Case No. 2630 2699 1643 2762 2687 2738 Helix (Helicigona) barbata Férussac, 1832 (currently Lindholmiola barbata; Mollusca, Gastropoda): proposed confirmation of lectotype desig- nation. RISSOOIDEA (Or RISSOACEA) Gray, 1847 (Mollusca, Gastropoda): proposed precedence over TRUNCATELLOIDEA (Or TRUNCATELLACEA) Gray, 1840. Mytilus anatinus Linnaeus, 1758 (currently Anodonta anatina; Mollusca, Bivalvia): proposed designation of a neotype. Griffithides Portlock, 1843 (Trilobita): proposed confirmation of Griffithides longiceps Portlock, 1843 as the type species, so conserving Bollandia Reed, 1943. Longitarsus symphyti Heikertinger, 1912 (Insecta, Coleoptera): proposed conservation of the specific name. Acanthophthalmus van Hasselt in Temminck, 1824 (Osteichthyes, Cyprin- iformes): proposed conservation, and proposed designation of Cobi- tis kuhlii Valenciennes in Cuvier & Valenciennes, 1846 as the type species. Trionyx sinensis Wiegmann, 1834 (Reptilia, Testudines): proposed conser- vation of the specific name. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE Opinions published in the Bulletin of Zoological Nomenclature The following Opinions were published on 29 June 1990 in Vol. 47, Part 2 of the Bulletin of Zoological Nomenclature. Opinion No. 1587 1588 1589 1590 1591 1592 ISDS 1594 1595 1596 Ly 1598 S99 1600 1601 1602 1603 1604 1605 1606 1607 Orbitolina d’Orbigny, 1850 (Foraminiferida): Orbulites concava Lamarck, 1816 confirmed as the type species. Hapalorhynchus beadlei Goodman, 1987 (Trematoda, Digenea): holotype replaced by a lectotype. Phyllodoce (Carobia) rubiginosa Saint-Joseph, 1888 (currently also Nerei- phylla rubiginosa; Annelida, Polychaeta): specific name conserved. Pleuromma princeps Scott, 1894 (currently Gaussia princeps; Crustacea, Co- pepoda): specific name conserved. Fizesereneia Takeda & Tamura, 1980 (Crustacea, Decapoda): Troglocarcinus heimi Fize & Seréne, 1956 confirmed as the type species. Bodotria Goodsir, 1843 (Crustacea, Cumacea): conserved. Iphinoe Bate, 1856 (Crustacea, Cumacea): conserved. Leucon Kroyer, 1846 (Crustacea, Cumacea): conserved. Aleuropteryx Low, 1885 (Insecta, Neuroptera): Aleuropteryx loewii Klapalak, 1894 designated as the type species. Semblis Fabricius, 1775 (Insecta, Trichoptera): Phryganea phalaenoides Lin- naeus, 1758 conserved as the type species, thus conserving Sialis Latreille, 1802 (Insecta, Megaloptera). Coryphium angusticolle Stephens, 1834 (Insecta, Coleoptera): generic and specific names conserved. Ophonus Dejean, 1821 and Tachys Dejean, 1821 (Insecta, Coleoptera): Car- abus sabulicola Panzer, 1796 and Tachys scutellaris Stephens, 1828 designated as the respective type species. Papilio carthami Hubner, [1813] and Syrichthus serratulae major Staudinger, 1879 (currently both in Pyrgus; Insecta, Lepidoptera): the specific names carthami and major conserved. Tachina orbata Wiedemann, 1830 (currently Peribaea orbata; Insecta, Dip- tera): neotype designation confirmed. Rapport sur les Myodaires du Docteur Robineau Desvoidy (1826): suppressed for nomenclatural purposes. Tenthredo zonula Klug, 1817 (Insecta, Hymenoptera): specific name con- served. Saccopharynx Mitchill, 1824 (Osteichthyes, Saccopharyngiformes): con- served. ICHTHYOPHIIDAE Taylor, 1968 (Amphibia, Gymnophiona): conserved. Thorius pennatulus Cope, 1869 (Amphibia, Caudata): specific name con- served. Semioptera wallacii Gray, 1859 (Aves, PARADISAEIDAE): conserved as the correct spelling of the generic and specific names. Mus musculus domesticus Schwarz & Schwarz, 1943 (Mammalia, Rodentia): specific name conserved. INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (botany, zoology, and paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. 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Authors are encour- aged to consult this article before manuscript preparation. Copies of the article are available from the editor or any associate editor. The establishment of new taxa must conform with the requirements of appropriate inter- national codes of nomenclature. Decisions of the editor about style also are guided by the General Recommendations (Apendix E) of the International Code of Zoological Nomenclature. When appropriate, accounts of new taxa must cite a type specimen deposited in an institutional collection. Review.— One of the Society’s aims is to give its members an opportunity for prompt pub- lication of their shorter contributions. Manuscripts are reviewed by a board of Associate Editors and appropriate referees. Proofs. — Authors will receive first proofs and original manuscript for correction and approval. Both must be returned within 48 hours to the Editor. Reprint orders are taken with returned proofs. Publication charges.— Authors are required to pay full costs of figures, tables, changes in proofs ($3.00 per change or revision), and reprints. Authors are also asked to assume costs of page-charges. The Society, on request, will subsidize a limited number of contributions per volume. If subsidized manuscripts result in more than 12 printed pages, the additional pages must be financed by the author(s). Multiple authorship will not alter the 12 page limit (each author will be viewed as having used his/her 12 subsidized pages). Payment of full costs will facilitate speedy publication. Costs. —Printed pages @ $60.00, figures @ $10.00, tabular material @ $3.00 per printed inch. One ms. page = approximately 0.4 printed page. CONTENTS The identity of Fannyella rossii J. E. Gray (Coelenterata: Octocorallia) | Frederick M. Bayer Meiopriapulus fyiensis Morse (Priapulida) from South Andaman, another example of large- scale geographic distribution of interstitial marine meiofauna taxa Wilfried Westheide A checklist of the Bryozoa of the Galapagos William C. Banta and John C. Redden Pyrgulopsis bruneauensis, a new springsnail (Gastropoda: Hydrobiidae) from the Snake River Plain, southern Idaho Robert Hershler Rediscovery of Tulotoma magnifica (Conrad) (Gastropoda: Viviparidae) Robert Hershler, J. Malcolm Pierson and R. Stephen Krotzer New species and new records of scaled polychaetes (Polychaeta: Polynoidae) from the Axial Seamount Caldera of the Jaun de Fuca ridge in the northeast Pacific and the east Pacific Ocean off northern California Marian H. Pettibone Redefinition of Teneridrilus Holmquist (Oligochaeta: Tubificidae), with description of two new species from North America Christer Erséus, Jar! K. Hiltunen, Ralph O. Brinkhurst, and Don W. Schloesser Additional records of stomatopod crustaceans from Isla del Coco and Golfo de Papagayo, east Pacific Ocean David K. Camp and Hans G. Kuck A new species of chirostylid crustacean (Decapoda: Anomura) from off the west coast of North America Keiji Baba and Janet Haig New species, clarifications, and changes in status within Eosentomon Berlese (Hexapoda: Pro- tura: Eosentomidae) from the United States Ernest C. Bernard Taxonomic notes on Ephydridae (Diptera) Wayne N. Mathis and Tadeusz Zatwarnicki A new species of Paraspadella (Chaetognatha) from the coastal waters of Japan Jean-Paul Casanova A new combination and synonymy for two subspecies of Cucumaria fisheri Wells (Echino- dermata: Holothuroidea) Philip Lambert A new genus and species of anthiine fish (Pisces: Serranidae) from the eastern South Pacific with comments on anthiine relationships William D. Anderson, Jr., N. V. Parin, and John E. Randall Three new species of symphurine tonguefishes from tropical and warm temperate waters of the eastern Pacific (Symphurus: Cynoglossidae: Pleuronectiformes) Madhu N. Mahadeva and Thomas A. Munroe Crocodylus raninus S. Miiller and Schlegel, a valid species of crocodile (Reptilia: Crocodylidae) from Borneo Charles A. Ross A new subspecies of Diglossa gloriosissima (Aves: Thraupinae) from the western Andes of Colombia Gary R. Graves The holotype of Natalus stramineus Gray (Mammalia: Chiroptera: Natalidae) Charles O. Handley, Jr. and Alfred L. Gardner International Commission on Zoological Nomenclature Table of Contents, Volume 103 Index to New Taxa, Volume 103 WIS 784 789 803 815 825 839 847 854 861 891 907 913 922 931 255 962 966 973 978 982 ‘4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON GUIDELINES FOR MANUSCRIPTS For Publications of the BIOLOGICAL SOCIETY OF WASHINGTON Ir Supplement to Volume 103, No. March 1990 Proceedings of the Biological Society of Washington VOLUME 103 NUMBER 1 MARCH 1990 PE —~ = P aALTHSOn: SMIIhoON AA E60, TMUIOV ‘BREL HOSAM 4 snes Fett eee 8 | ea “Bee RARE 1a cine ck reds vif Send Proof To; Dr. C. Brian Robbins Division of Mammals National Museum of Natural History Smithsonian Institution Washington, D.C. 20560 GUIDELINES FOR MANUSCRIPTS FOR PUBLICATIONS OF THE BIOLOGICAL SOCIETY OF WASHINGTON C. Brian Robbins and David B. Lellinger Abstract.--The abstract should be <3% of the length of the text of the article. It should be a concise statement of findings, rather than a listing of subjects covered, and should be written as a single paragraph and double spaced. Except under unusual circumstances, references should not be cited in the abstract. All newly described taxa must be given by name in the abstract. This paper provides authors with guidelines and examples to aid in preparing manuscripts for the Biological Society of Washington. The required format for manuscripts for the Proceedings and Bulletin is presented and followed herein as a model of format and style. However, except for the title, author(s), abstract, and figure and table captions, the final printed papers appear in double-column format rather than occupying a complete page as does this manuscript. It is neither possible nor desirable to provide detailed examples of every format or style item, but we have attempted to address most of the potential questions facing an author preparing a manuscript. As a general guide, we recommend the fifth edition of the CBE Style Manual (CBE Style Manual Committee 1983), available from the Council for Biology Editors, 9650 Rockville Pike, Bethesda, Maryland 20814; cost is $24.00, including postage. The latest edition is a more comprehensive guide for scientific writers than were previous 2 editions. A shorter but useful guide is Day's (1988) How to Write & Publish a Scientific Paper, Third Edition; softcover copies of this guide are available from Oryx Press, 2214 North Central at Encanto, Phoenix, Arizona 85004; $14.95 for copies mailed to North America, $17.95 mailed elsewhere. Methods Manuscript preparation.--Type manuscripts on letter size (215- by 280-mm, 8 1/2= by ll-in.) or A4 (210- by 297-mm, 8 1/4- by 12 1/8-in.) paper. All parts of the manuscript should be double-spaced, including text, literature cited, tables, table headings, and figure legends, so that reviewers, editors, printers, and proofreaders can make essential notations. Do not justify the right margin. Allow >3-cm (1 1/2 in) margins on all sides. Do not use a title page. Type name, address, and telephone number of the author to whom all correspondence and proof is to be sent at the upper left of the first page. (Note to foreign authors: this is the place to indicate the name and address of the person in the U.S.A. who will read proof.) Place page numbers at upper right of all pages except the first. Titles should be brief and include key words that will be useful for indexing and information retrieval. Include phylum, class, order, and family names (separated by a colons and a space [e.g., Crustacea: Copepoda: Caligidae]) for species likely to be unfamiliar to many readers. In subsequent use of scientific names, abbreviate generic names where possible (e.g., Studies on Botrychium meridionale and B. virginianum); do not abbreviate scientific names in titles or subheadings if they have not been previously spelled out in the same title or subheading. Center the title and type it in uppercase letters, including underlined scientific names. Three spaces below the title, type the name(s) of the author(s) centered on the page. Triple-space after the author(s) name(s) and begin the abstract, using a normal paragraph indentation as shown on page 1 (secondary heading - see below) and write the entire abstract as a single paragraph. Also, indent five spaces at the beginning of each paragraph throughout the text. Double-space after the abstract and insert a 5-cm horizontal line. Double space after the line (as on page 1) and begin the introductory material. Do not use "Introduction" as a heading. In general, use no more than three levels of organization: primary headings (centered); secondary headings (indented from left-hand margin, underlined, followed by a period and two dashes); and tertiary headings (indented from left-hand margin, ending with a colon). New taxon names and associated citation of their taxonomic placement (phylun, class, order, family, etc.), and figure and table references are primary headings, for example: (Note: unless the higher taxonomic categories are the primary topic, author and date citations are not necessary.) Systematic Paleontology Class Mammalia Linnaeus, 1758 Order Chiroptera Blumenbach, 1779 Family Desmodontidae Gill, 1884 Genus Desmodus Maximilian, 1824 Desmodus archaeodaptes, new species Fig. 1 Long lists or complex material that is related only indirectly to the primary subject should be placed in an appendix at the end of the manuscript. Items that are appropriate for appendices include: lists of specimens examined; lists of morphometric characters, or ecologic characters; derivations of complex mathematical or statistical formulas; and algorithms for computer programs. Appendices may be arranged as paragraphs, in tables, or in other formats as appropriate. Short lists, single formulas, and material related more directly to the subject should be placed in the text. Material not essential to the article should be omjtted. Never break words at the right-hand margin anywhere in the manuscript for any reason. This includes hyphenated words as well as words divided between syllables. Spell out the entire scientific name the first time a plant or animal is mentioned; thereafter, abbreviate generic names (first letter and period), except at the beginning of a paragraph or sentence. Include author(s) and date where such is essential to identification of a taxon (such as ina synonymy). If subgeneric names are used, they should be placed between the generic and specific names, and be enclosed in parentheses; abbreviate them after their initial use, using the first letter of their name. Use subspecies or infraspecific names only when they are essential to the understanding of the article. New descriptions in zoology should be spelled out, as: "new genus," "new species," "new subspecies", and "new combination," or for botanical papers uses igen.) nova spemnoviet,. USubSpe nov. wy, Om ='Combin nove Each of these should be preceded by a comma. Descriptions must be in English, except for Latin descriptions or diagnoses in botanical papers (fossils excepted). Provide the scientific name (generic and specific names) of all organisms at first mention of the vernacular name both in abstract and text (even if the name appears in the title); use the scientific name of the appropriate taxon if a vernacular name is inclusive (e.g., crickets [Gryllidae}). Tables and Figures.--Because of the cost of publication, tables should be used only when large quantities of data must be summarized or trends in data illustrated to support inferences. Pertinent data in most small tables (two-row or two column tables) usually can be presented in the text in less space and without loss of intelligibility. Consider carefully the presentation of non-tabular data in the two- 5 column format of the Proceedings. In many instances, a table would be easier to read and type-set. Figures and tables should be numbered in the sequence in which they are referred to in text. Reference in text should be in the style: (Fig. 1) and (Table 1). Reference to several tables or figures in text should be in the style: (fables 1, 2) and (Figs. 1, 2). Parts of composite figures should be labeled with uppercase or lowercase letters, or numbers, and should be used to refer to parts of figures both ini legendsiand) wimetexct) ((eng/,/! (Hagia! vp airy Figshi-1—3), 5:!,0r Figs. 1B-D, 2D-F). Then note the desired location for each figure and table in pencil in the left margin of the manuscript. Because many readers prefer to scan data before reading text, each table or figure with its title or legend should be understandable without reference to the text. Be sure to include species, localities, and dates of study, if appropriate, in addition to a description of the content. Abbreviate words in column heads or data identifiers (stub columns) only when necessary. Spell out names of months except where space does not permit. When abbreviated, and in the Materials section, use three-letter abbreviations without periods (e.g., Jan, Jun, Oct). Reference in the text to tabular and graphic presentations of data should be in support of inferences, not simply to direct the reader to specific tables or graphs. For example, use sentences such as "Litter size did not seem related to month of onset or duration of the reproductive season (Table 1)," but avoid sentences such as "Data on litter size, and month of onset and duration of the reproductive season in some Spermophilus are presented in Table 1." Do not describe the content of tables and figures in text; the need for such description indicates that the tables and figures are not understandable by themselves. Data should be presented either in graphic (figures) or tabular form, whichever is more understandable 6 or economical of space. Figures should be constructed to be fully legible and not wasteful of space when reduced to column width or page width. Orient multiple graphics having two or three illustrations one above the other if reduction to a single column is desired. Illustrations that are intended for a full-page presentation, should be mounted accordingly. Type each table on a separate sheet(s), give ita complete, intelligible title, and refer to it by number in text. In titles of tables, italicized and Roman type is presented asian the text -))\Capitalize) only) the first) wordvor column heads and items in the secondary or data identifier column heads; do not use all uppercase letters in headings. Do not use a dash or minus sign in a table to indicate lack of observations or tests if the table contains plus and minus signs; otherwise its use is clear (= missing data). Footnotes to tables should be kept to a minimum. Lowercase letters should be used to denote footnotes except those limited to probability; one, two, or three asterisks (*) should be used for P < 0.05, 0.01, and 0.001, respectively. Provide a legend for each figure and refer to the figures in the text. Type all figure legends, double-spaced, on a separate sheet at the end of the manuscript; the typescript format for figure legends is illustrated in Appendix I. In legends, underline only those words or terms (i.e., genus and species names) that will be set in italics (as listed elsewhere in this guide). Place any necessary identifications (e.g., symbols, cover types, scale bars, regression formulas) directly on the figure rather than in the figure legend. Do not submit figures larger than 360 by 620 mm (15 by 24 in.); figures that are 215 by 280 mm (8 1/2 by 11 in.) are preferred. Figures exceeding this size are difficult to mail, handle, and store. Photographically reduce figures larger than 15 in. wide to page width (5.5 in.) or larger than 24 in. long 4 to page length (8 in.). Prepare line drawings with lines of sufficient width and letters of sufficient size to remain legible when the figures are reduced to final page or column width (CBE Style Manual Committee 1983). Graphics produced by coarse dot-matrix printers are not satisfactory for reproduction. Legibly mark all photographs and figures on the back with the author(s) name(s), figure number, and indicate "TOP." Use soft pencils (blue preferred) on the backs of photographs and drawings, because markings and pencil or pen indentations may show through. Leave a margin at the top of each figure (including photographs) of at least 5 cm (1/2 in.). Provide photocopies of figures to be sent with the manuscript to reviewers. Use press-on letters very carefully, and only if the illustration is mounted on a board. Pressed-on letters on a large oversized illustration often flake and break. The use of a clear spray fixative and a cover sheet is suggested. The best solution, when pressed- on letters are used, is to keep the original illustration approximately the size of the desired final reproduction. This will reduce handling problems and damage. Submission of photographs (either those produced by standard processes or by diffusion-transfer imaging [PMT]) of graphics is recommended. Submitted photographs must not be produced by making a photograph through a halftone screen (pre-screened). Mount original graphics or photographs on cardstock or other heavy material; use the same material when several small graphics are submitted for a composite figure. In the case of composite graphics, they must be mounted as the author desires. If more than one photo is mounted on a plate, it is best to cut square edges and mount photos neatly together so that no white space appears between the photos. Also, try to place the identifying letters or numbers inside each individual photograph. The editors will not mount photographs. For clarity of presentation, the Biological Society of Washington prefers to print photographs in their 8 original submitted size, or a size reduction to fit a journal page; consideration of size should be made before preparing a composite figure. Results and Discussion Style and Usage Punctuation.--Use quotation marks sparingly for emphasis or special use of a word or term, thus preserving them mainly for actual quotations. Use the slant line or solidus (/) only to indicate "divided by" or in dental, vertebral, or chaetotaxy formulas. Underline generic and specific names in the text (underlined words will become italic in print). Do not use italic or bold type in manuscripts even if your typewriter or word processor has that capability. Underlined words (italics) are discouraged for use other than scientific names and headings. However, all terms and symbols in mathematical equations and those used to denote statistical tests should be underlined. Hyphenate compounds used as adjectives (e.g., 3-year-old male, 77-day gestation period, 10- by 10-trap grid, home- range area, life-history strategy, 0.5-m plot). The same holds for adverbs (e.g., well-developed dentition) except those ending in "ly." However, to emphasize the true subject, use prepositions to avoid strings of modifiers (e.g., estimates of home-range size, not home-range-size estimates). Hyphenation in formation of compounds is complex (e.g., "semi-independent" but "semiarboreal" and "mid-July" but "midday"); use of Webster's Third New International Dictionary, Unabridged is advised. Always use serial commas, including that preceding the conjunction (mice, voles, and shrews). Use no punctuation between state and zip code in addresses (address line of these guidelines is an example). Do not use "back-to-back" parentheses () (), as for citing a reference and referring to a table in text; use (Smith 1984, Table 1) not (Smith 1984) (Table 1). To separate a parenthetical statement from a reference in the text, use a 1-em dash (two dashes on typewriter) if the authority is cited in support of statement as in (. . . as commonly seen in Peromyscus--Smith 1984); otherwise, separate with a semicolon as in (. . . as commonly seen in deer mice [Peromyscus leucopus; Wolff 1988]). Abbreviations, spelling, and miscellaneous items.-- Paragraphs and sentences should not begin with an abbreviation (such as - ha sitaattindaarea rg Walsh talkers ss: 9s) he DO not use contrived acronyms or mnemonics for names of localities, study areas, morphological characteristics, governmental agencies, physiological parameters, statistical tests, or most other items. Acronyms for museums, standard abbreviations for protein and enzyme loci, and symbols used in mathematical equations are acceptable if referenced or defined at first use (words should be used for the latter when not used in a mathematical sense; e.g., "The area is mr’, but the radius is shorter than the diameter"). Spelling and use of words should be in accordance with Webster's Third New International Dictionary, Unabridged. Use of words not included therein should be avoided, but newly coined words and foreign words may be used sparingly if precisely defined at first use in manuscripts. Scientific terms should follow recent specialized dictionaries and glossaries. Refrain from using words in other than their standard meanings. | Use only "male" and "female" to distinguish the two sexes. Avoid terms such as rams, ewes, bucks, or other similar names. Numbers and mathematics.--Use decimals rather than fractions except in equations. Decimals not preceded by a whole number always should be preceded by a zero (0.75) in text, tables, and figures. In the text (introduction, discussion, conclusions, etc.) and in titles, use numerals for numbers greater than 10 nine except when starting a sentence (associated units should then not be abbreviated: "Eleven minutes," but "About 11 min"; "Fifteen percent," but "More than 15%"). For numbers one through nine, use words except when used with units of measure (6 mm), time (3 days, 3 summers, 4 years), but not enumeration (five dugongs, two crabs, seven observations). Also, use numerals for all items in a series that includes at least one number greater than nine (1 dik-dik, 7 numbats, and 19 slow lorises). Treat ordinal numbers in the same manner as cardinal numbers (first panda, 14th deer mouse, 1st month, 6th) min) foursthiseraials)e. In the specimens examined and description or diagnosis sections, use numerals throughout. Avoid the use of numbered sentences or phrases in text. Reference to specific items by number in text is acceptable but names of items should be lowercase (e.g., day 1, experiment 4, setiger 5, grid 6, site 1, coxa 3, segment 7). Do not refer to individual animals either by name or field- catalog number; reference to individual animals tends to indicate that the material being presented is anecdotal. Use commas in numbers of five digits or more (e.g., 10,000 and 100,000). The exceptions are field and catalog numbers of museum specimens, and pagination in references. Use a colon (not a solidus [{/]) to express ratios (e.g., 1:3.2, males:females). Do not present the numbers of males and females in a total sample (e.g., 15:48) as sex ratios but calculate the true ratio and give the sample size. For example, ". . .1:3.2 (n = 63) in favor of females." Use < and > for "less than" and "greater than" with numerals (< 5 g, not "less than 5 g"; > 20 captures, not" more than 20 captures"). Use < for "equal to or less than" or "not more than" and 2 for "equal to or greater than" or "at least" with numerals. Alot When giving ranges, use "from 10.1 to 31.0 mm," but "the range is 10.1-31.4 mm." In other words, do not use "to" unless it is preceded by "from." When presenting equations and formulas, use the solidus (/) for simple fractions and give the meanings of all symbols and variables in the text. When presenting values with respect to another factor such as time or space, use the solidus (/) if only two measurements are involved (g/ha); if three or more measurements are involved use the exponent -1 asiamyit).). 43" mg g} h?." Do not present equations published elsewhere unless they have been modified; a simple reference will do. In reporting measures of central tendency (means, modes, etc.) or dispersion (standard deviations, standard errors), units of measure should not be more precise than the original measurements. For example, if mammals are weighed to the nearest 0.1 9g, then means such as 54.56 should be reduced to 54 jon ande xn Sb Shouldiber 54.16) Oh not, 547.16. (0.109), both in tables and in text. Dental formulas should be presented as i 1/1, c 0/0, p 1/0, m 3/3, total 18. Vertebral formulas should be presented ASMA, Sie wOumls a2 Seo mande2io—Ssia Cayuatotalgs55—60)— Uppers teeth should be referred to by capital letters and lower teeth by lowercase letters (e.g., P4 is the fourth upper premolar and ml is the first lower molar); do not use superscript and subscript numerals to designate upper and lower teeth. Time and dates.--Write dates as 24 April 1989, with no internal punctuation; an exception is in the specimen examined section where the three-letter month code should be used. Indicate time of day on the 24-h system with four digits. Midnight is written as 0000 h, 8:30 a.m. as 0830 h, and 11:15 p.m. as 2315 h. Also, indicate time similarly as "4-h intervals," and "3 h/day." To express the age of a stratigraphic unit or the time of a particular geologic 12 event, and where a specific dating technique has been used, employ "Ma" for mega-annum (time greater than one million years), or "B.P." (years before present) for radiocarbon dated material, which goes up to 40,000 yrs. To express other time relations, use "m.y." (million years) or m.y.a. (million years ago). For example, from 50 to 25 Ma, 25 m.y. elapsed and it is possible that these organisms lived at least 40 m.y.a. The ratio of light (L) hours to dark (D) hours under laboratory conditions is to be shown in this form: 141L:10D. Units of measure:--Use of the metric system and the international system of units (Systéme International d'Unités) is advocated. Exceptions to the use of the metric system and the international system of units are English units in localities from specimen labels and in quotations from other publications. Also, the following variables may be reported in other units: Temperature in degrees Celsius (°C) instead of degrees Kedivame (okie Time in minutes (min) and hours (h) instead of seconds (sec). Sound intensity (relative terms) in decibels (dB) instead of watts/meter square (W/m’) . Volume in liters (1) instead of cubic decimeters (am?) . Area in hectares (ha) instead of TO mice Accepted abbreviations that can be used in text, tables, and figures of manuscripts, without explanation or punctuation, and some words that should be spelled out follow: Units of measure (use abbreviations only with numerical values, otherwise spell out; e.g.,". . . FN = 72-76, but populations with low fundamental numbers .. ."): re degrees Celsius day 2N (not ppm gal 2n) diploid number (N = chromosome number) fundamental number gravity second minute hour day week month year million years gram (not gm) milligram kilogram hertz kilohertz megahertz kilometer (s) meter (s) millimeter(s) centimeter (s) fathom(s) mile(s) foot (feet) inch (note period) diameter hectare molar parts per million watt (or west) magnification microgram micrometer (micron) gallon 2:3 14 m1 milliliter and cubic centimeter liter* should be spelled out when confusion can exist; mainly when it is used alone. When in Prine) COM Looks leukeya poorly spaced 101. Confusion does not exist in a list of ingredients [(i.e., 14 kg sawdust, 500 g potassium nitrate, 1.5 1 water)] or when in combination (ml). Miscellaneous: Standard abbreviations used in text, their proper punctuation, and some terms that should be spelled out. AC DC alternating current and direct current Bev (BS Marae UMS cap Mics Cru ye hieiDr. college degrees Co. county counties spell out (no abbreviation) elev. elevation (note period) ISSKS {Gong se ako fc figure maximum no abbreviation minimum no abbreviation pers. comm. personal (oral) communication P.O. post office R/V Searcher name of research vessel is underlined SEM scanning electron microscope States spell out to avoid confusion Table write out completely TEM transmission electron microscope UK United Kingdom (no spaces) U.S.A. United States of America USNM National Museum of Natural History Wis SoSoixs Union of Soviet Socialist Republics weight no abbreviation 15 Directions and coordinates: N E S W ENE SE SSW _ (no periods) T14N, R10OW, SW Sec. 2 legal description for localities 10°06'N, 25 °07'W latitude and longitude Latin and other foreign language terms (the following are accepted as written, without underlining; definitions follow terms). ewebe a affinis, having affinity with but not identical with a posteriori known from experience a priori intuitively, independent of experience adminis ad libitum, freely available auct. auctoris, of the author auctt. auctorum, or authors ca. circa, about (he 6 conferre, compare e.g., (note comma) exempli gratia, for example emend. emended, emendation en masse in a body, as a whole et al. et alia, and others etc. et cetera, and so on BOG (note comma) such as nly ake. in a letter (Note: data or information in recent unpublished manuscripts should not be cited in any form; however, quotations or use of data extracted from old manuscripts, as exist in archives, libraries, and occasionally herbaria, may be used) in situ in place in utero in the uterus 16 in vitro outside the living organism in vivo in the living organism leg. collector or collected by nec not nom. dub. nomen dubium, name of doubtful application nom. nud. nomen nudum (plural, nomina nuda), name without validation part. partim, part per se by itself, as such p-p pro parte, in part sensu as defined by sic thus (to signal exact transcription) So dl sensu lato, in the broad sense s.s sensu stricto, in the strict sense via by way of, by means of vis-a-vis WAleze VS. Or versus Statistical terms: in relation to, as compared with videlicet, namely against, in contrast to ANOVA analysis of variance (AY coefficient of variation aft degrees of freedom n sample size or number in sample Pp probability rorR correlation coefficient SD standard deviation SE standard error SEM standard error of the mean statistical tests mean Symbols.--Male (dc) and female (9) symbols should not be used in the text, but may be used in figures and lists of specimens examined (8 o or 14 99 - note space between number and symbol). Write percent as one word in the text, but use iL the percent sign after numerals (1%, 99%) and in bodies of tables. Because of possible confusion with similar symbols, do not use X as a symbol for the word "by." Write, for example, Wawretyoys) Wieyerel (bes oS Jon Wo lone AAs} Cl 5 9G 6 oP ibe Fl multiplication sign is used in mathematical formulae or for indications of magnification, identify it as such in the margin of the manuscript. Documentation and Literature Citation The function of literature citation is to assist readers in locating material referenced by the author, a process that permits an orderly growth of knowledge through continued testing and reassessment. Documents written primarily to fill administrative requirements are not catalogued in most libraries and do not enter the body of knowledge that supports research. Therefore, such documents are not to be included in the Literature Cited section; the quarterly reports of U.S. Fish and Wildlife Service Cooperative Wildlife Research Units and job completion reports for Pittman-Robertson Federal Aid in Wildlife Restoration projects are examples of this kind of material. Certain other state, provincial, and federal reports also are excluded from lists of citations. Abstracts of oral presentations delivered at professional meetings and printed separately are excluded, but references to abstracts in Dissertation Abstracts and Masters Abstracts and abstracts published in journals (e.g., American Zoologist, American Journal of Botany) are permitted. Style of documentation.--When citing informational references in text, use the form "Jones (1983)" if the author's name is part of the sentence and "(Jones 1983)" if it is not. Two articles by one author cited at one time should be written "(Jones 1975, 1982)"; two articles published by the same author in the same year, "(Jones 1981la, 1981b)." Other examples follow: 18 (Cameron 1977:507) or Smith (1957:23, table 3) [Note: only cite pagination in text references for direct quotations, in a synonymy, or in reference to a specific table or figure in another publication.] In text, citations of figures or tables other than those in the present manuscript should be in lowercase letters. Citations of figures or tables in the text should begin with a capitol letter, as: Fig. 1 or Table 2. "In press" citations in text should be avoided. Use the year of expected publication (current or subsequent year) and end the citation listing in the Literature Cited section with "(in press)" - see Appendix II. This allows for less costly corrections when the citation is updated at the page proof stage. Citations with more than two authors are cited in the text as: Lidicker et al. (1976). Do not underline the "et al.". Multiple citations in the text should be ordered chronologically and then alphabetically if in the same year, for example (Jones 1961, 1963; Hennings & Hoffmann 1977; Phillips 1978; Jones & Smith 1981; Jones & Baker 1983; Mares et al. 1983). Use an ampersand (&) between surnames of authors rather than "and". For in text multiple citations of three or more authors, in the same year and same first author, cite all authors, for example (Jones, Smith, & Williams 1988; Jones, Williams, & Smith 1988; Jones, Williams, Smith, & Baker 1988). In the text, unpublished material can be referenced as follows: (pers. comm.) denotes information obtained orally; (in litt.) denotes information obtained in a letter. Names of persons providing unpublished information should include initials when referenced in text (e.g., R. H. Tamarin, pers. comm.; D. P. Christian, in litt.). In the Literature Cited section, do not cite or use information from unpublished recent manuscripts (except theses and dissertations) or tS papers in preparation. Unpublished data may not be referenced in any context. Citations in the text that indicate the author(s) of zoological scientific names should be as follows: Family Caryophyllidae Dana, 1846; Didelphys microtarsus Wagner, 1842:359; and, Themiste dyscrita (Fisher, 1952). Note the use of a comma after the author(s) name(s). In botanical scientific names, dates may be used in the case of homonyms. The proper form for new zoological descriptions, plus their associated synonymy, is as follows: Chaceon bicolor, new species Figs. 1-3 Geryon affinis.--Griffin & Brown, 1976:256, figs. 7-9.-- Sakai, SO 7e8ci9pmergs.) e—19), pili) 2), fag. Di i(collox)).. [Not Geryon affinis A. Milne Edwards & Bouvier, 1894] The proper form for new botanical descriptions, plus their associated synonymy, is as follows: Pityrogramma mortonii Jackson, nom. nov. Pityrogramma mutabilis Gleas., Proc. Biol. Soc. Wash. AGA Sici LOSS) wnon, Liy 7D Siee LY. Pes) MU). SiaAnseyaAre zc Onasy, Cochase Co.: 7 mi NW of Wigwam, 4800 ft elev, Brooks 482 (US; isotype UGE All citations or author and date, whether informational, taxon describers, or essential references in the synonyny, except for botanical synonymies, must be listed in Literature Cited. Literature in botanical synonymies should be abbreviated according to the Botanico-Periodicum-Huntianum or Taxonomic Literature, ed. 2. The list of references at the end of the manuscript should have the primary heading "Literature Cited." Sample literature citations are listed in Appendix II. Like all other parts of the manuscript, this section must be double- spaced. Only papers referred to in the text may be listed. The list must be alphabetical by authors' last names. Therefore, within this structure, papers with one author 20 would be listed first, then those with two authors (alphabetical by second author), then three authors, and so on. Use first author et al. (e.g., Patton, J. L., et al.) for papers having seven or more authors. Where the author line is identical, the listing should be chronological by publication date. If two or more papers by the same author or sequence of authors are listed, the name(s) are not repeated but are replaced by a 3-em dash (six dashes in typescript) and a period. Use a 3-em dash in subsequent entries for all repeated authors. For example: Cockrum, Ee ole er LOWAar RRE2E> oy) LLY Do DSSS S5 pS Wo io VOMeS sr wies ~ doze SO OSS , so----, & J. D. Smith. 1980. (Note: Throughout the text and in the Literature Cited, insert a space between the initials of a persons name, as above.) The volume number of a journal or other serial publication should be cited. However, do not include the issue number in journals having continuous pagination throughout the year or volume. When citing publications that have only a number and no volume, treat the number as a volume (Occasional Papers of the Museum of Natural History, University of Kansas 25:1-39). Underline scientific names or other words only if italicized in the original title. In citing books, omit Roman pagination (e.g., "J. Wiley and Sons, New York, 432 pp." not "xxii + 432 pp.™") except in instances that those pages contain the first reference to a taxon under consideration. Also, provide only the first city listed for publishers (e.g., John Wiley and Sons, New York" not John Wiley and Sons, New York, Chichester, Brisbane, and Toronto"). Plates (or figures), if not included in paginated materials, should be referenced after pagination (e.g., 286 Ppt plisiiai=24 “for Eigse 1=24)))) J. hE oplates), VEilguiscesi) tor tables are paginated, do not cite them again in the 21 reference. Journal names and book titles are set in Roman type, so do not underline. Abbreviations used in Literature Cited.-- Abbreviations must not be used in the Literature Cited, except for those used in the title of the cited paper. Include and spell out all words that comprise the reference, including articles, conjunctions, and prepositions. Manuscripts will be returned to the author(s) if journal names are not completely spelled out. Conclusions A summary in French, German, Russian, or Spanish is acceptable. This summary, which may be a translation of the abstract, should be placed immediately after the English abstract. Do not include a additional summary or summary paragraphs in English at the end. Before submitting your manuscript, carefully cross-check all citations in the text, including figure and table captions, against listings under Literature Cited, and check each entry in the Literature Cited section against its original source to verify title, year of publication, names, quotations, and page numbers. Include all citations, when appropriate, in synonymy listings. The editors scan the Literature Cited and make spot checks for accuracy, but cannot assume responsibility for verifying all citations. The sequence of material in each copy should be: Title, Author(s), Abstract, Text, Acknowledgments, Literature Cited, Author's(s) Address(es), Appendix, Figure Legends, Figure copies (each numbered and identified), Tables (each table numbered with an Arabic numeral and with heading provided). Number every page sent with the manuscript, including Literature Cited, figure legends, and tables. Any appendices should appear after Literature Cited. Figure legends and then copies of the figures follow. Next, the tables, each on a separate sheet. Glossy prints of each figure (or original artwork if high-contrast photographs of graphics cannot be 22 obtained), should complete the parcel of manuscript materials. Three copies of all materials, including figures (legible photocopies acceptable), should be submitted with a cover letter stating the title, and full name(s) of author(s), and availability of publication funds. Do not submit manuscripts under consideration for publication elsewhere. Acknowledgments Parts of this manuscript were extracted intact and other parts paraphrased from a similar manuscript prepared by D. E. Wilson; Al.) ‘LeGardner,.iandB. )di Vents tom Lhe Tounnaly or Mammalogy, and from "Standard style for technical publications" prepared for customers of Allen Press, Inc., by G. Dresser. Helpful suggestions for this publication were provided by 1. E)..;Bowman, |S). D2 iCasicns/ 4h )D.herrarcl Ga DE Johnson, R. B. Manning, W. N. Mathis, and D. E. Wilson. The acknowledgments section should contain a few brief statements in a single paragraph to recognize the contribution of others and support from patrons or agencies. Use only initials for names of persons acknowledged, but spell out names of agencies (e.g., National Science Foundation not NSF). Literature Cited CBE Style Manual Committee. 1983. CBE style manual. Fifth edition. Council of Biology editors, Bethesda, Maryland, 324 pp. Costain, D. C. 1978. Dynamics of a population of Belding's ground squirrels in Oregon. Unpublished M.S. Thesis, Oregon State University, Corvalis, 66 pp. Gouchy, “Lie iKe 1932. Breeding notes on a few Washington mammals.--The Murrelet 13:25. Day, R. A. 1988. How to write & publish a scientific paper. Third edition. Oryx Press, Phoenix, Arizona, 211 pp. 2s Junge, R., & D. F. Hoffmeister. 1980. Age determination in raccoons from cranial suture obliteration.--The Journal of Wildlife Management 44:725-729. Linsdale, J. M. 1946. The California ground squirrel: a record of observations made on the Hastings Natural History Reservation. The University of California Press, Berkeley, 475 pp. McKeever, S. 1966. Reproduction in Citellus beldingi and Citellus lateralis in northeastern California. Pp. 365- 385 in I. W. Rolands, ed., Symposia of the Zoological Society of London, Academic Press, London 15:1-559. Scheffer, T. H. 1941. Ground squirrel studies in the Four- Rivers country, Washington.--Journal of Mammalogy 228 2 OSB) 6 Tomich, P. Q. 1962. The annual cycle of the California ground squirrel.--University of California Publications in Zoology 65:213-281. WielSOnN De whe) Ais Gardner a Be dis Vents. 11989). Guidelines for manuscripts for publications of the American society of mammalogists.--Journal of Mammalogy (supplement) 70(4):1-17 + 21 unnumbered. (CBR) Department of Vertebrate Zoology (Mammals), National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (DBL) Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. 24 Appendix I Examples of typescript for page having Figure legends Fig. 1. Localities at which Sorex trowbridgii (open circles) and S. bendirii (closed circles) were collected in Benton Co., Oregon, 1975-1986. Fig. 2. Relationship between body mass and body length for (a) 12 Sorex trowbridgii, and (b) five S. bendirii collected in Benton Co., Oregon, 1975-1986. Imsleps Be Syringonomus dactylatus. A. Photomicrograph of the cervical region of the holotype, USNM 77172; unlabeled arrows point to amphidial gland; scale equals 100 pm. B. Photomicrograph of the posterior body region and tail of the paratype USNM 77175; scale equals 100 um. Abbreviations. -- AV, anal vent; CG, caudal gland; NR, nerve ring. 25 Appendix II Examples of Literature Citations Journals.-- Bader, R. S. 1965. A partition of variance in dental traits of the house mouse.--Journal of Mammalogy 46:384-388. Blair-West, J. R., et al. 1968. Physiological, morphological and behavioural adaptation to a deficient environment by wild native Australian and introduced species of animals.--Nature 217:922-928. [use of "et al." for papers with seven or more authors] Carleton, M. D., & C. B. Robbins. 1985. On the status and affinities of Hybomys planifrons (Miller, 1900) (Rodentia: Muridae) .--Proceedings of the Biological Society of Washington 98:956-1003. [Underlined scientific names will be set in italics; note use of an ampersand (&) instead of "and" between author's names] Fauvel, P. 1936. Sur quelques annélides polychétes de l'ile de Paques.--Bulletin du Muséum National d'Histoire Naturelle, Série 2, 8:257-259. [Include all diacritical marks] OGniaz, Ds Iu, P.yCostal &) Be wi. LeBoeut. 1990.5 Water, and energy flux in elephant seal pups fasting under natural conditions.--Physiological Zoology (in press). [Note use of probable year of publication after authors names and the use of "in press," in parentheses, after the journal name} Whitaker, J. 0., Jr., & R. Ej. Wrigly. 1972. Napaeozapus insignis.--Mammalian Species 14:1-6. [underline scientific names only if italicized in original title] Wicksten, M. K. 1984. Alpheopsis harperi (Decapoda: Alpheidae): A new species of snapping shrimp from Texas.--Northeast Gulf Science 7(1):97-100. Proceedings and transactions.-- Chitty, D. 1952. Mortality among voles (Microtus agrestis) at Lake Vyrnwy, Montgomeryshire in 1936-9.-- 26 Philosophical Transactions of the Royal Society of London, Series B, 236:505-552. Kirsch) be Mey & 7A.) Dis )Druse.| |) 1973/4) ¢Pradrile) fines sand wildlife.--Proceedings of the Annual Tall Timbers Fire Ecology Conference 12:289-304. Peek) di Ma, Mien IScote,) le di Nelson” De di) PLercey 71k) viaemeiae Irwin. 1982. Role of cover in habitat management for big game in northwestern United States.--Transactions of the North American Wildlife and Natural Resources Conference 47:363-373. Books. -- Baird, S. F. 1857 [1858]. Mammals. in Reports of explorations and surveys for a railroad route from the Mississippi River to the Pacific Ocean. Beverly Tucker, Printer, Washington, D. C., 8(part 1):1-757 + pls. 17- 28, 30-60. [give correct publication date in brackets] Barbour, R. W., & W. H. Davis. 1969. Bats of America. The University of Kentucky, Lexington, 286 pp. [City is not followed by state when name of state is part of the name of the press] Eisenberg, J. F. 1981. The mammalian radiations: an analysis of trends in evolution, adaptation, and behavior. The University of Chicago Press, Chicago, 610 pp. [Names of large, universally recognized cities are not followed by state] Honacka: di Hy, Ki) SE) Kanman\9&) di Wa Koepple(edse i. lose Mammal species of the world: a taxonomic and geographic reference. Allen Press, Inc. and The Association of Systematics Collections, Lawrence, Kansas, 694 pp. Hsu, T. C., & K. Benirschke. 1969. Microtus oregoni (creeping vole) 2n = 17, 18. An atlas of mammalian chromosomes.” Vol?)'3), Folio) 11217 Springer-Verlag, New York, unpaged. [Pages are not numbered] Linnaeus, C. 1758. Systema nature per regna tria naturae, secundum classes, ordines, genera, species, cum 27 characteribus, differentii, synonymis, locis. Tenth edition. Laurentius Salvius, Stockholm 1:1-824. [give complete title, do not use ellipsis] Nowak, R. M., & J. L. Paradiso. 1983. Walker's mammals of the world. Fourth edition. Johns Hopkins University Press, Baltimore, 1 and 2:1-1362. Part of book.-- Bourliere, F. 1955. Ordre des Fissipedes. Systématique. Pp. 215-291 in P. =P. Grasse, ed., Traité de Zoologie. Masson et Cie., Paris 17(1):1-1170. [period after title of volume] Calder, W. A., III. 1974. Consequences of body size for avian energetics. Pp. 86-151 in R. A. Paynter, Jr., ed., Avian energetics. Publication of the Nutall Ornithological Club, Cambridge, Massachusetts 15:1-334. Carleton, M. D., & G. G. Musser. 1984. Muroid rodents. Pp. 289-379 in S. Anderson & J. K. Jones, Jr., eds., Orders and families of Recent mammals of the world. John Wiley and Sons, New York, 686 pp. [give pagination for entire volume] Leach, W. E. 1821. Galatéadées. Pp. 49-56 in F. G. Levrault, ed., Dictionnaire des Sciences Naturelles, 18(1820), Strasbourg. Spezharsky. T. N. 1939. Order Ostracoda. Pp. 193-196, pl. 46 in B. Likharev, ed., Atlas rokovodyashchikh form iskopaemykh faun SSSR. Volume 6, Permskaya Systema. Tsentral'nyi Nauchno-Issledova-tel'skiy Geologo- Razvedochnyi Institut (TsNIGRI). (The atlas of the leading forms of the fossil fauna USSR. Volume 6, Permian System. Central Geological and Prospecting Institute) (in Russian). Theses and dissertations.-- Lackey, J. A. 1973. Reproduction, growth, and development in high-latitude and low-altitude populations of Peromyscus leucopus (Rodentia). Unpublished Ph.D. dissertation, University of Michigan, Ann Arbor, 128 pp. 28 Wrazen, J. 1976. Feeding ecology of a population of eastern chipmunks (Tamias striatus) in southeast Ohio. Unpublished M.S. thesis, Ohio University, Athens, 26 pp. 29 Table 1.--Some reproductive patterns in four species of Spermophilus in western United States. a EEE EEE EEE EEE ee Species and Liter size® Reproductive state I bad Range season” Authority S. washingtoni Washington 26 8.0 Sa01 February-March Scheffer 1941 S. beldingi Oregon 110 522 5-8 April-June Costain 1978 California a7 Pak 3=10 April-July McKeever 1966 S. columbianus Washington 21 5.8 2-7 March-May Couch 1932 S. beecheyi California 40 Lys 5-12 February-April Linsdale 1946 California 34 61 1-9 March-June Tomich 1962 * Based on counts of embryos. > From first evidence of reproduction behavior to first appearance of young. Table 2.--Means (+SE) and ranges for mass and length of 514 bacula from raccoons (Procyon lotor) among five age classes, northwestern Oregon, 1982-1983 and 1983-1984 furbearer seasons. 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