ORDO Pati pe , ak tte ened ; ES ae atraAb tied ay ef fete , : Pitas Mele el ai ne ‘ ‘ reaieg bedbae 7 ‘ Aaattatcha! . teat an? a Ske 4 PURSE eno See bet it By, Veda cpt ytethele te vat oValietamy, eo 7 “ * , Mervin Tih tet 7 Dede bat gi sete ad tet ehh Jape dnet ote Mom ge vata iy hee Phe RE eo we et tere On A cty eben? ot Ne a ae Hae Va Od oD oat nth af Nine sit git2 ted Wy Nei mt ately meet es i fe tee Meir a t ee 7 * 4 ve Pe ; 7 ab ares Ste an ae . ‘ pt reg eagt z Py cL meet a we eat gels re ate fees bw teu, hed Pave Ue% OE ene tie? . Pree . s PAUL Nee are ede Me an Se cred ’. site Pe ey hs 4 . Aah Pathe’ Sete ayy Ee Fete te tei Fak ein hinted sinntas te be } : ua ‘ : AV eMC BNI De Ate Oat MMe He oe c OID TOING GDN ie At dy aphid ee h : i ‘ . J emel dy ae bifle ewe de Me shee bet tie ted fe . . Gd " : edi Mn ane Teer Me Me a, BIN Gein ties ind ane tte ten ae ‘ , og 4 ? a é ' hotgertonine 4 see! 90 Los Mate Biger : ot tauren scab és . carn : gah” 4 a oe veres fedinel Hon (aM ane é HUAN Comte fy : wit a tnd Labsa s SAAD omy rites F rey os ‘ ° J 2 5 MMV 8 Re ote ered s . : r ta 12 S ieth a thyme Baars Te agra ain wire FE Teh he ee ee R ety Bs cet wg whe he ete Viterbo Neate A Te Mower t Hed al eSy grein! | 20% apt ediy php win tt as > d ee ony De Se Pa OU ce et ae . s 2 « ta tielbe lan CGR AM an OA Se ph AS Heo ty Die te ee eee y ore ier Tae : 4 , 5 OPEP IH, a , ae ‘ 4 eRe amt tare yao ett Wenge ess ‘ ‘ . Shee SU hee eg | . ; . 4 tt . Y “ i 7 aS er twieetive whe we OR stiwt Sunes AWE ae ae * NebemeSeis Da me Pi he Reema Hel dg Be PS ona tate # ey SEI SIRE at Beto? = ty Ay gy i : Beg tig ae Pe Ndtnelle n 3 He ay hy tote eile Mae wee tre ghee Me PM outty Leake te Nyt fee eee Pe Bie? este Ny tet Ad MME Fal <4 yp Neate Naha te Me Bt shen etn Gwent ete te aig ain PONS EV eed ge! Sedat cite eed at fpenath date at y® aoe Ne Pan ate Me Ae they oo eh ee ea oS wer ite: St Tha Datta ome te aret St eee nA he Pao sPebsd pa, Hee Aee ees Ae ee ene de® a Mees ste rte ta he A Derrenetdare ep Aeoyedle La @iny efit Aaa Pine ee hag st PER Fp od NINES Pee aera cnt ns ee Lae ee tla na aoe eee Heri aoe ® wser ew Abe Sate eB Gy. eee Tate hE MP Oe mw eT hae Ge an tetete Neon ete weil Vet Oe eta tate Seka ey ot Pfu Lele amate Ww ane te iter « eet i eid “yi A] cH ad mh) Le fae ty tae ate 4a a 7 : ; oy “ 4 Yh ie " ‘ 4 a iy \hy ’ . ne x 4), | 1 ¥ Ne Jou q | { | i é > ne me ee .: : a if A tan a +f ha = é i. Uae a a PROCEEDINGS of the Biological Society of Washington VOLUME 110 ey) Vol. 110(1) published 15 April 1997 Vol. 110(3) published 10 October 1997 Vol. 110(2) published 9 July 1997 Vol. 110(4) published 12 December 1997 WASHINGTON PRINTED POR THE SOCIBTY EDITOR C. BRIAN ROBBINS ASSOCIATE EDITORS Classical Languages Invertebrates FREDERICK M. BAYER JON L. NORENBURG FRANK D. FERRARI RAFAEL LEMAITRE Plants Vertebrates DAvID B. LELLINGER GARY R. GRAVES 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 1996-1997 OFFICERS President STEPHEN D. CAIRNS President-Elect RICHARD P. VARI Secretary CAROLE C. BALDWIN Treasurer T. CHAD WALTER COUNCIL Elected Members JOHN FORNSHELL RAFAEL LEMAITEE ALFRED L. GARDNER DIANA LIPSCOMB SUSAN L. JEWETT JAMES N. NORRIS + i] 71 Te Civin Ae iT ir) “ati te _ ACTOVRIZAW FO YTaIbO? JAIWOIONE : : 7 ; re ‘tata j % ae re Fa ‘i. ? ’ 7 - 4 S- { “9 } CMJ) eee F viele) A Vor? =) Spe T A way 4 CIMACT ' 4 Yi > 4 tJ Al Wittbsin HH * H I< nA 4 , ; ‘4 XA Nalert CIT TAM ARE } wee . iy ’ ‘ y A - rl ri i mae -- ee iy wistensth Wwatiad A er BEA RE Lone viame Wa" k, Bay ETE Aa) a MEAD 2OEMAN > et ne tee Wren, PR 5 TABLE OF CONTENTS Volume 110 Adams, Dean C. and James F. Jackson. A phylogenetic analysis of the southern pines (Pinus subsect. australes Loudon): biogeographical and ecological implications ______ Alvarez, Fernando and José Luis Villalobos. Pseudothelphusa ayutlaensis, a new species of freshwater crab (Crustacea: Brachyura: Pseudothelphusidae) from Mexico ________ Amaral, A. Cecilia Z. and Eloisa Helena Morgado. Stratiodrilus (Annelida: Polychaeta: Histriobdellidae) associated with a freshwater decapod, with the description of a new So see ae aN a eee eel De hk TE Be Se ee Ae Baeza, Juan A., David Véliz, Luis M. Pardo, Karin Lohrmann, and Chita Guisado. A new polyclad flatworm, Tytthosoceros inca (Platyhelminthes: Polycladida: Cotylea: Pscumoccrotigde) Mom mlean) coastal Waters. oie ieee eee ee be Sy Baldwin, Carole C., G. David Johnson, and John R. Paxton. Protoblepharon rosenblatti, a new genus and species of flashlight fish (Beryciformes: Anomalopidae) from the tropical South Pacific, with comments on anomalopid phylogeny —__-__> Bayer, Frederick M. Narella nuttingi, a new gorgonacean octocoral of the family Prim- netdae \(Aitberoa)iiiom theveastem Pacificws 221 6 ive be Ne time Berner, Dorothy B. The cladoceran collection of the National Museum of Natural His- fory, Smithsonian’ Institutions Washimegton, D:@€°i 4 ee Bravo, Manuel Rafael and Masaaki Murano. New records of the genus Hansenomysis in Japan with description of a new species (Crustacea: Mysidacea: Peep baciaGaC). SSRI ara ee) aie) )) eas De eign Ts bene» Brown, Walter C., Angel C. Alcala, and Arvin C. Diesmos. A new species of the genus Platymantis (Amphibia: Ranidae) from Luzon Island, Philippines —---_-_>>>SSE Burukovsky, Rudolf N. Selection of a type species for Farfantepenaeus Burukovsky SebenIe ACCA MCCADGGA PCR ACIG AC) (att tte et Campbell, Jonathan A. and Eric N. Smith. A new species of Tantilla (Serpentes: Co- tebridac) tent netiheastem: tiratemala Mss. 2) 0 eh Campos, Ernesto and Mary K. Wicksten. A new genus for the Central American crab Pinnixa costaricana Wicksten, 1982 (Crustacea: Brachyura: Pinnotheridae) ______ Carleton, Michael D. and Erik Van der Straeten. Morphological differentiation among Subsaharan and North African populations of the Lemniscomys barbarus complex ET ae ETE gee ee er oR SL te ee eee 5 i de a IC, eee Child, C. Allan. Some deep-sea Pycnogonida from the Argentine Slope and Basin ____- Ciros-Pérez, J. and M. Elias-Gutiérrez. Macrothrix smirnovi, a new species (Crustacea: Anomopoda: Macrothricidae) from Mexico, a member of the M. triserialis-group — Cooper, John E. and Martha Riser Cooper. A new species of troglobitic crayfish of the genus Cambarus, subgenus Aviticambarus (Decapoda: Cambaridae), endemic to White SL SUD Ee 22 202 th dh come pelt ec Sal ha ete Ane AS ee ate eA ENS Tete lett TOE DeBiase, Adrienne E. and Barbara E. Taylor. Aglaodiaptomus atomicus, a new species (Crustacea: Copepoda: Calanoida: Diaptomidae) from freshwater wetland ponds in South Carolina, U.S.A., and a redescription of A. saskatchewanensis (Wilson, wpe s ) ieskieh 2 SS hat up oh hire ti raat arty Sn a ee er aR eae ee nae rites ART aC 8 eee Fitzpatrick, Jr., J. F. and Arnold G. Eversole. A new crayfish of the genus Distocam- barus, subgenus Fitzcambarus (Crustacea: Decapoda: Cambaridae) from South (SEED NT SE est RDU Sp ee SES Peete Pee nents Neo a eee ee ee Se ee Frey, Jennifer K., Robert D. Fisher, and Luis A. Ruedas. Identification and restriction of the type locality of the Manzano Mountains cottontail, Sylvilagus cognatus Nelson, NERS eee Beso irvine) a ole) 0 ts) See ee ere Fukuoka, Kouki, Monica S. Hoffmeyer, and Maria D. Vinas. Lophogaster muranoi, a new species of mysid from the coastal waters of Argentina (Crustacea: Mysidacea: en Loe) |e cae ee eee Oe a ee ae el eer eee eee eee ee Gee, J. Michael and Robert Burgess. Triathrix montagni and T. kalki, a new genus and two new species of Cletodidae (Crustacea: Copepoda: Harpacticoida) from California Pee Gaee TUeeRLT 9 Meyer, C. and W. Westheide. Boguea panwaensis, a new species from Thailand: the first member of the Bogueinae (Polychaeta: Maldanidae) to be found outside northeast PULLS) € 1 Cy: ee IR ee ihe Bs M8 Te eles he Ree rela Prete orig py aly ly te Miura, Tomoyuki, Junzo Tsukahara, and Jun Hashimoto. Lamellibrachia satsuma, a new species of vestimentiferan worms (Annelida: Pogonophora) from a shallow hydro- thermal venti’ Kasoshitnia’ Bay, Sap 20 ee eee ee ee Modlin, Richard F. and James J. Orsi. Acanthomysis bowmani, a new species, and A. aspera li, Mysidacea newly reported from the Sacramento-San Joaquin Estuary, Cal- iformia (Cristaceat Miysidae yee ORY oh BESO U8 oO eee eee Murano, Masaaki. Nanomysis philippinensis, a new species (Crustacea: Mysidacea) from brackish watersof-the Philippines sc 02-0 nce pen se 8 eed ee a ar Naiyanetr, Phaibul and Peter K. L. Ng. Esanpotamon namsom, a new genus and species of potamid crab (Crustacea: Decapoda: Brachyura) from a waterfall in northeastern Thailand 457-470 314-319 320-325 65-68 393-398 39-48 338-365 629-639 558-559 422-425 74-98 384-387 107-114 198—202 256-262 280-284 399-411 366-372 203-209 447-456 439-446 236-241 417-421 Olson, Storrs L. and Cyril A. Walker. A trans-Atlantic record of the fossil tropicbird Heliadornis ashbyi (Aves: Phaethontidae) from the Miocene of Belgium —... Opresko, Dennis M. Review of the genus Schizopathes (Cnidaria: Antipatharia: Schi- zopathidae) with a description of a new species from the Indian Ocean _... Osawa, Masayuki. A new species of the genus Petrolisthes Stimpson, 1858 (Crustacea: Decapoda: Anomura: Porcellanidae) from Yonaguni Island, the Ryukyu Islands _____ Pettibone, Marian H. Revision of the scaleworm genus Eulagisca McIntosh (Poly- chaeta: Polynoidae) with the erection of the subfamily Eulagiscinae and the new DEES ACERS IT Se os Es oe MSE 2 0 5s ee ee Pecs Menor de Oliveiza. Cnidaec ot Scletactinia Price, Roger D. and Robert M. Timm. A new subgenus and four new species of Gliricola (Phthiraptera: Gyropidae) from Caribbean hutias (Rodentia: Capromyidae) _...____ Reid, Janet W. Argyrodiaptomus nhumirim, a new species, and Austrinodiaptomus kle- erekoperi, a new genus and species, with redescription of Argyrodiaptomus macro- chaetus Brehm, new rank, from Brazil (Crustacea: Copepoda: Diaptomidae) ______ Riser, Nathan W. Syllides eburneus, a new species, with notes on other members of the genus (Polychaeta: Syllidae) from the coast of New England and New Brunswick __. Riser, Nathan W. Protodrilus gelderi, a new species of infralittoral, interstitial poly- SNE Ete Tal BO NAAU IN ASSIA CURA SEES 8 Eu Roman-Contreras, Ramiro and Ingo Wehrtmann. A new species of bopyrid isopod, Pseu- dione chiloensis, a parasite of Nauticaris magellanica (A. Milne-Edwards, 1891) (GCrsstaces: ecapous Elippolylidac) 2.0 ee ee Sankarankutty, C. and Raymond B. Manning. Observations on Hexapanopeus schmitti Rathbun from Brazil (Crustacea: Decapoda: Xanthidae) Soest, Rob W. M. van and Helmut Lehnert. The genus Julavis de Laubenfels (Porifera: eG AT CRE C1 2 eee ee ie ee ee eee Solis-Marin, Francisco A., Alfredo Laguarda-Figueras, and Antonio Leija-Tristan. Mor- phology, systematics, and distribution of Meoma ventricosa grandis and M. ventricosa ventricosa (Echinodermata: Echinoidea: Brissidae) along Mexican coasts ___________ Sterrer, Wolfgang. Luriculus minos (Platyhelminthes: Rhabdocoela: Luridae) from the ESE LG ETE GS ae ae a ps SS pc ec ee Sterrer, Wolfgang. Gnathostomulida from the Canary Islands —_----_-__________ Suarez-Morales, E. and N. Riccardi. Redescription and first record of Cymbasoma tenue (Isaac, 1975) (Copepoda: Monstrilloida) in the Mediterranean Sea _____---_-_________ Talbot, M. S. Doxomysis acanthina, a new leptomysinid (Crustacea: Mysidacea) from the northern Great Barrier Reef, Australia, with extensions to the known distributions of D. australiensis W. M. Tattersall, 1940 and D. spinata Murano, 1990, and a key Vi) DEVE SE EE ESE cE OTC eh Se a So ee ve Taylor, Christopher A. and Mark H. Sabaj. A new crayfish of the genus Orconectes fom western. ennessee (Decapoda: Cambaridac) . Thomé, José W., Patricia H. dos Santos, and Luciana Pedott. Annotated list of Veron- icellidae from the collections of the Academy of Natural Sciences of Philadelphia and the National Museum of Natural History, Smithsonian Institution, Washington, D.C., Dee (i ioluses: Gustropoda: Soleolifera) 00 Tyler, Diane M. and James C. Tyler. A new species of chaenopsid fish, Emblemariopsis ruetzleri, from the western Caribbean off Belize (Blennioidei), with notes on its life SSE Ng oe NN i Se ne mr enn eT Ucar EE Tyler, James C. New species of Paratriacanthodes spikefish (Triacanthodidae: Tetra- ooueutionmiecs) trom: the South China Seay. Volkmer-Ribeiro, Cecilia and Klaus Riitzler. Pachyrotula, a new genus of freshwater sponges from New Caledonia (Porifera: spongillidae) —-__________ Weems, Robert E. and Jon M. Bachman. Cretaceous anuran and dinosaur footprints fram tue Paiumemt Pompaton ol Vitpiaia. 22 Wehrtmann, Ingo S. and Alberto Carvacho. New records and distribution ranges of shrimps (Crustacea: Decapoda: Penaecoida and Caridea) in Chilean waters __________ Williams, Austin B. Occurrence of three species of mud shrimps in aquiculture ponds on Caribbean coasts of Venezuela and Colombia, with a redescription of Upogebia onussagoe Williams, 4993 (Decapoda: Upogebiidae) Williams, Austin B. Two new species and a range extension of mud shrimps, Upogebia, from Pacific Costa Rica and Mexico (Decapoda: Thalassinidea: Upogebiidae) —______ 624-628 157-166 58-64 937-551 167-185 285-300 581-600 143-149 552-557 242-248 249-255 502-510 301-309 150-153 186—197 99-106 426-438 263-271 520-536 24-38 310-313 489-501 412-416 617-623 wy Ran ditegees sateen te eaten B46 ‘ons (ei atate. NE en wrt anak A nabaty esata, ee a, Ta i Spee h Tag: ik Vyas 2) SPURNS cart ag Olay en, YER Maines . caleecensG)} siatinnnT caaleaane ST Vs, WP a OE ees fade |dow peta bof (ile E At eM ie Ree ee ae dauitegpe To ee A ay ‘tsi orton M4 tho ype eget Sacbeat i AP) mvgnviegyy cumin tien graih baw a seeds einges SS ae od : Lb aad OSS Or sabes Gomi oll) : : ARIE ST, (Ro abigeart wets “ty RTD het of, eT) AD pee See aM nats iW? 2 mi) “eho ee Pann sevicecaod 5 urbe we os. genetic * enlghesie al) cure — NEC, fj ‘ x win J haben Se re sal nas tate ul its aires shots % daddies) Weer tyra untied wD eopnak”, ae ’ Cerage. | ; | ie uct une’ ted malt ete) Lwae i oy > ia 4 Ss ee ST ritws sd iad oT Panini, wut Senha ee wrannadeoh buiac) whl del yw rf flaPenlndes . TER. Spares: 1 tad eege one NeerE) cil belie pares Sat acne Mies. ale ity bioy- moch eeoiaiioc coda at m3 gency ae a es a | mS Nelle Large emma aig se eae ir, + eran ene adinviatin. Siew? stevcrarhenaid paeoaunne’ Dy a: bs a. Diese Aeon. eee aslel) 0: 4 sean: boda avhorensae 5 A te Belen Saale my sesithsuitergi Tp iby Padet kwatee. Sas eae mth ! rat ee leaded in, diated heal A ine male biad . as ‘risa ol sd harmerets raphe Need, oar et crane cone ray. Sayty, hse Ag anry INDEX TO NEW TAXA Volume 110 (New taxa are indicated in italics; new combinations designated n.c.) PORIFERA Heterorotula caledonensis © NREL ASCE E ITS Sie i a ty A Ae Se RP Oncosclera diahoti n.c. IESG PRE oe ea nT ee ce eR dS raceki n.c. CNIDARIA Btn tr LE 1h) amen elie we ht ce NEE ser Ne ere eh Schizopathes amplispina Tytthosoceros inca Austrognathis clavigera Sa eC 7 ame eee eee DN ee lel ROBE TORI a IU ee Se ee Re ee eee eee ee ANNELIDA Polychaeta Boguea panwaensis TS SUSE S OS) TENCE ONT Oc Re ea I ee SS IL Pa ETE CTCL a eee a ec a BOE AGIS GINA Ee wes biainnily 25a ee lO) a Se IPL tae ce ee Ca Ne ll a ere T TEAS LUO Comet eae ted ase reed a I ne en Sac we tes So ee ee Pe HEU yD G2) AUN TTS) SCY 1 CoS ee ae EPEE YEU, GoPhone ee Sp ee Le one CARON MR CIRI PATER NER se 8 HERE SS Py a | LN Si rn A En cE OPEC Se (2 OUTS TES Z SyZCTe 271 ha ea) I CD eae aba a? NNN ae a MEO A COPE WIG PE LEL TOE Ge ey PAT 1 el a eRe ae OASIS" 2 ¢_ JEe ARTHROPODA Crustacea Bees Ad RMB TEAIOTE UPPER ISFEIN ed theese oe te ts a Ee le Ne ea Ea a Ea SSR MMR TIT P EN wo oa el ee. Se ret st EVEE PSUS SR CUVEE) EAST tie eae NA ecto ne 7 Sn CR ee MEE SORE AEE OREO IZISFIVUIDINY 2 ssc Pas 3 te OS oe TRACE IPRS 0 IGE SAN SB I el Deh ra 20S eae de Le nn ene Pie SS SPI be 0 2 ee Kleerekonert. 2222 Se ee 594 ALSULRU Nn a a 393 Were yi en ee ee ee 395 Cambarus (Aviticambarus) veltchorum —f23:% Setanta 608 Cyathura (Cyathura) esquivel: — 2s Sy ee 74 mistocatbams (CPitzcambarws) (Rantert 5s ee PH i MPD Oa COUEN SEG CRERFUT TRATES a 428 Poymamenelila see vibes: ae ee ee 84 EES CRERTOOHLEEIREOAT Ee a ee 417 TES | AY2 || alaed eae ee Peele ia ee Pe Ne et iar A BT oe en Laren mer eee eee msneNU RNR) Saltod yd ee eres 418 PE TRtTCSNE CONS CUCCHIUELIS OE 0s ee 256 PTITILCDENRGGUS, THEW SUDR CIS cs EE a ee 154 LESS aS a RE 2 A Rt ee a Pe ee ee ee 69 COS ERETe Ame TMC) ees rs ese ie A a ee 70 EAARIS ORIGIN Y SUS fEAIOTELCGL, he nek SE a ee i ec ee 228 FIC LETOMVMIPNONCGECLOC IIIT ea aes ee aan ge ee 130 OES IS HEV OMELE So ep 8 Sed ey hah ee el he ee ae ee 82 PELTED) 90 (2 ila aR cee SSRI, ANGE Fad ORR cls, Fee OS eine eae tee ace AA OB eve ee ek 84 pa ASCE MALICE OU a Ee ee ee 601 IBVPArSRSE CM LUMEN CSUDEE FUE OVID a hl a he ld to a 115 STS ARUN UMMA fA LELO CS ta l h e k wel peecte dN 78 INAOUINSIS DRUID DIROIISES: (oc. 8 2 aN ee ee a Ee A A ee eer 236 INEOPONOGUGCLVIUS COMID! nc. J Ey Jem Sane nl SE em URI A wo RY Re aes 4 is a ee 280 VATA PONLCMR CLEA LELID cy MPs ok 2 oS ON Se ee ee ee ee 132 LEU PUIG cn i hh a nels Heer De Na ore a a iers ss pana puis. 134 LC CERID. ces eo Soa te ly Ek es 8 i ee a 139 RCONECIGS PASE! tne Se Bato ae ON ee ee ee ee Ne ae Se la 263 Pacnycnel escaitanragas: e228 see A ae ee ee ee 65 AUALIRN@ IIS 2 PZ OL OC 3 ca err ee Pe ls De i es et ae eth no ee tee. Se 89 DERI IRURU AES ee asl an seen Doren dale, abe in ie I pr Aan ad So Si i es es dai IN Visser Ah I AR ot tte A 93 Penichimenes Melellangy. noose ena a hes ae ae eal ge Ae Rye en eel nile Leer VE 39 PEtrOlistheESIGOHGNCISES 2.0 2. toes Den ee ee hie et oy ee Se 58 IPSEUGIONSHCHTOISES: 25 a5 a 2 pt ait ee meh Sg IN ge ee ee 243 Psevdothelphusaayutlaensis, |<: 2 8th, seals AM SK 88 eS ee 388 ERUEIIC REVS WCEEAI COILED CCE i ese Nn Sate ced PN i a a co ec aD ane 400 STEMS MLS AM COTIN ACA NEN CS mee 5 es I lah 1 De ll 422 BUR ISECEU AICTE x OS VOU IP Naot Mots Wr ABs I ig WN lM tag tac iN ENT a WO MI Mei aie a 211 TeBUNES Meet acke Mag re NI SNe eo Wa AT oe ees Ast, ig LOAN Ue PO SRE ee eR eee a 219 WIORIGGRE, os ere Ne A ee Se ee en ee 212 Melipnca ebb, Dada aera ec aid ot Nh ae nS A a se 617 WAT OAS LG ice Sa Na ARE mel Sede HOR ance Sep I a ne on 620 CHORDATA Amphibia Boltoclossa decora: x32 28a et a a er ee a 367 Mobeveenerioona Ti erate ena cc a eh a oN ce ee 369 Ph styriasitis: renee: os a 2 ey A Sr ee Se ce 8 a a 19 Reptilia Cyclemiys @tripons: 255 as ee es eg ee, 632 ie Ee, (/ ac aaa alee NPI OR RRA CE, ow) See CON ene oo Sh eR a ie as 333 Pisces Einblemariopsis-ruetghert FREQUENCY (KHZ) 1 TIME (sec) 2 Fig. 2. Audiospectrograms of advertisement calls of (A) Platymantis dorsalis (CAS 201561) and (B) Pla- tymantis mimulus (CAS 201191) both from Mt. Maquiling, Luzon Island. er species similarly small in size for mature specimens (most notably subterrestris, montanus, and polilloensis) have large fin- ger disks and broad “T’’-shaped, terminal phalanges. In terms of the finger and toe disks, finger length, subarticular tubercles, and ornamentation of the dorsum and limbs, mimulus and dorsalis are very sim- ilar. Adults of mimulus have heretofore been confused with juveniles of dorsalis (Taylor 1922b). Platymantis mimulus differs morpholog- ically from dorsalis primarily in its much smaller size at maturity and minor differ- ences in ornamentation of the dorsum. Their very similar general appearance con- tributed to the confusion of mimulus with juveniles of dorsalis. This is supported by the fact that the only significant, propor- tional difference, among those examined, is a slight difference in head shape (SnL/HW for mimulus: mean = 33.300, SD = 2.849, n = 20; for dorsalis: mean = 35.850, n = 20; t = 3.010, df = 38, P = 0.005). These two species also have very differ- ent advertisement calls. The call of Platy- mantis dorsalis sounds like whet-whet (pro- duced by whistling). The first part starts at about 1400 to 1500 Hz but is the dominent frequency between 2200 and 3600 Hz (Fig. 2A). The duration of each call ranges from about 0.056 to 0.075 seconds, and the in- tervals between calls from about 0.22 to 0.80 seconds for those recorded. The call of Platymantis mimulus sounds like osek-sek-sek. The dominant frequency ae PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON is 3300 or 3400 to 4500 or 4600 Hz], sweeping up at the beginning from about 2000 Hz and at the end dropping back to the same level (Fig. 2B). The duration of each call is about 0.13 to 0.15 seconds. The intervals between calls, for those recorded, ranged from 0.20 to about 1.5 seconds. Habitat.—Platymantis mimulus is re- corded from original lowland forest and man-made forest (mostly exotic Mahogany species) on Mt. Maquiling at 200 to 650 m. It is a ground dwelling, forest species, as is P. dorsalis. Taylor (1922b) notes that young frogs of dorsalis (including his ex- amples of mimulus) are usually found along streams, whereas adults of dorsalis are usu- ally further away in the forest. Our recent samples of mimulus are mostly from the forest floor and thick grasses at the forest edge, one is from under a rotting log and one from inside a curled leaf on a shrub about 30—40 cm above ground. Range.—Known thus far only from Mt. Maquiling in southern Luzon Island. Discussion The discovery of this diminutive Platy- mantis on Mt. Maquiling, Luzon Island, that has long been identified with juveniles of P. dorsalis indicates the need to reex- amine other populations from throughout the islands that have been assigned to P. dorsalis. Field studies as well as reexami- nation of existing samples in numerous mu- seums may be necessary in the effort to identify other mimics of dorsalis. Other examples possibly already exist based on our evidence of populations of di- minutive, dorsalis-like frogs on Mt. San Cristobal and Mt. Banahao at Lucban, Lu- zon. However, because of the small size of these samples (six males and two females and eight males respectively) and the need for verification of some field observations, we hold in abeyance our taxonomic assign- ment of these populations. Their calls ap- pear different from the calls of either P. mi- mulus or P. dorsalis. However, we still need a recording of the voices for analysis and comparison and larger samples for infor- mation on intra-population variation. Acknowledgments We thank R. E Inger (FMNH), A. Dubois (MNHN), E. N. Arnold and B. Clarke (BMNH) for providing specimens cited in this paper. We are indebted to A. E. Levi- ton, R. Drewes, J. Vindum, and the Orni- thology Department, California Academy of Sciences. We also thank Dr. Joselito Jara, President of Southern Luzon Polytechnic College, Lucban, Luzon Island for courte- sies extended to field parties of Alcala and Diesmos. Literature Cited Brown, W. C., & A. C. Alcala. 1974. New frogs of the genus Platymantis (Ranidae) from the Phil- ippines.—Occasional Papers of the California Academy of Sciences No. 113:1—12. , & 1982. A new cave Platymantis (Amphibia: Ranidae) from the Philippine Is- lands.—Proceedings of the Biological Society of Washington 95:386-391. , R. M. Brown, & A. C. Alcala. 1997. Species of the hazelae Group of Platymantis (Amphibia: Ranidae) from the Philippines, with Descrip- tions of Two New Species.—Proceedings of the California Academy of Sciences (in press). Duméril, A. 1853. Mémoire sur les Batraciens An- oures, de la famille des Hylae-formes ou Rai- nettes, conprenant la description d’un genre nouveau et de onze espéces nouvelles.—Anna- les des Sciences Naturelles (3)19:135—179. Guenther, A. 1873. Notes on some reptiles and batra- chians obtained by Dr. Adolph Bernhard Meyer in Celebes and the Philippine Islands.—Pro- ceedings of the Zoological Society, London 1873:165-172. Inger, R. EF 1954. Systematics and zoogeography of the Philippine amphibia.—Fieldiana: Zoology 33:183-531. Taylor, E. H. 1922a. Additions to the herpetofauna of the Philippine Islands.—I. Philippine Journal of Science 21:161—206. . 1922b. The herpetological fauna of Mt. Ma- quiling.—Philippine Agriculturist 11:127—139. VOLUME 110, NUMBER 1 Appendix A Specimens Examined Cornufer dorsalis A. Duméril, 1853. Philippines (Java is an error): MNHN 4880 (holotype) and Mt. Isarog, Camarines Sur Province, Luzon Island: FANH 251648, 251656, 251653, 251660, 251662-—63, 251665, CAS 200414-—416; Mt. Maguiling, Laguna Province, Luzon Island: CAS 61005, 61009, 61011, 61013, 61015, 61027—28, 61030-31, 61033, 61227, 61231, 61233—-34, 61239—40, 61242—43, 61264-65, 196358, 196365-—67, 200417-18, 200420-—27; Mt. Banahao, Quezon Province, Luzon Island: CAS 61180, 201013—14, 201186—-88. Hylodes corrugatus A. Duméril, 1853. Philippines (Java is an error.): MNHN 4884 (holotype). 23 Platymantis meyeri Giinther, 1873. Laguna del Bay, Luzon Island, Philippines: BMNH 1947.2.5—5 (ho- lotype). Platymantis levigatus Brown and Alcala, 1974. Tablas Island, Philippines: CAS 136097 (holotype) and CAS 136098 (same locality). Platymantis spelaeus Brown and Alcala, 1982. south- ern Negros Oriental, Philippines: CAS 153469 (ho- lotype) and CAS 153470-83, (same locality). Platymantis mimulus new species. Mt. Maquiling, Lu- zon Island, Philippines: (See account of this spe- cies.) Platymantis sp. Mt. San Cristobal and Mt. Banahao at Lucban (Mt Banahao massive), Luzon Island, Philippines: CAS 201009-11, 201178-85, 201 197-200. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):24—38. 1997. A new species of chaenopsid fish, Emblemariopsis ruetzleri, from the western Caribbean off Belize (Blennioidei), with notes on its life history Diane M. Tyler and James C. Tyler (DMT) Smithsonian Institution Press (MRC-950), Washington, D.C. 20560, U.S.A. (JCT) National Museum of Natural History, Smithsonian Institution (MRC-106), Washington, D.C. 20560, U.S.A. Abstract.—A new species of Emblemariopsis is described from the shallow waters around Carrie Bow Cay, Belize. Emblemariopsis ruetzleri differs from other species of the genus in having a combination of 14 pectoral-fin rays and a low supraorbital cirrus, whereas the other two species with 14 pectoral-fin rays (E. pricei and E. randalli) have no supraorbital cirri. Sexually mature males of the new species are dark headed and live in holes in coral (usually dead coral), whereas females and immature males are pale and surface dwell- ing. The new species is diminutive, with an average size of 14.5 mm SL for the 35 dark-headed and sexually mature territorial males collected from holes (largest specimen 19.4 mm SL). While studying the life history of the sea fan blenny, Emblemariopsis pricei Green- field (1975), at the Smithsonian Institution’s marine laboratory at Carrie Bow Cay (Ellen Cay) on the Belize Barrier Reef, we rou- tinely collected dark-headed territorial male specimens of Emblemariopsis that we found living in holes in coral. Nearly all of the dark-headed males collected from holes in living coral proved to be the territorial, sexually mature males of E. pricei; females, nonbreeding males, and immatures of E. pricei are found on sea fans and coral sur- faces (Tyler & Tyler 1997). Nearly all of the dark-headed males col- lected from holes in dead coral differed from E. pricei by being somewhat less darkly pigmented in preservation (but not in life as seen with only the front of the head protruding from the hole), by having a low, simple supraorbital cirrus, and by be- ing smaller in average size. This smaller species, which has 14 pectoral-fin rays like E. pricei (and E. randalli Cervigon, 1965), is an undescribed species. Although dark- headed males of the new species usually are found in holes in dead coral (rarely in live coral), the two females captured to date are pale and inhabit the algal turf in the same dead coral habitat where holes containing mature males are found. The only known pale, immature male was found on the sur- face of a live coral, near dead coral con- taining a hole with a dark-headed male. We describe below this diminutive spe- cies whose life history and shelter specific- ity we continue to investigate. Methods Abbreviations: ANSP = Academy of Natural Sciences of Philadelphia; CBC = Carrie Bow Cay, Belize; SL = standard length; USNM = United States National Museum specimens at the National Muse- um of Natural History, Smithsonian Insti- tution. All specimens were collected using a weak solution of quinaldine sulfate (1 gram per 500 ml water). Those from holes were trapped in clear plastic tubes placed upright over the opening of the holes immediately VOLUME 110, NUMBER 1 after the solution was squirted into them, driving out the partially narcotized and dis- oriented specimens. Those collected on the surface had been partially narcotized by the solution dispensed around them before be- ing caught in plastic tubes or plastic bags. Emblemariopsis ruetzleri, new species Diagnosis.—The new species differs from all other species of Emblemariopsis by having a combination of 14 pectoral-fin rays and a supraorbital cirrus (low and sim- ple). The two other species of Emblemariopsis with 14 pectoral-fin rays (E. randalli and E. pricei; see Tyler & Tyler 1997 for meristics and other differences between these two species) do not have supraorbital cirri. Of the seven species of Emblemariopsis with 13 pectoral-fin rays, a supraorbital cirrus is either absent (E. bahamensis Stephens, E. bottomei Stephens, E. diaphana Longley) or present (E. leptocirris Stephens, E. oc- cidentalis Stephens, E. signifera (Gins- berg), E. tayrona (Acero)) (see Stephens 1963, 1970, for meristics of the first six species and Acero 1987, for the last). Type locality—Al|l 38 specimens are from the reefs around Carrie Bow Cay (or Ellen Cay on some hydrographic charts), Belize (see below for details), on the Belize Barrier Reef, 16°48.15’N, 88°04.91'W, 18 km off the nearest point on the mainland (Sittee Point, south of Dangriga), 1-8 m depth. Type specimens.—The 35 dark-headed, sexually mature, hole-dwelling males range from 11.9-19.4 mm SL, average 14.5 mm SL; the single pale, immature, surface- dwelling male is 14.6 mm SL; and the two females are 12.3—13.4 mm SL. Holotype: USNM 337496, 14.4 mm SL, dark-headed male from hole in dead coral, spur and groove formation just east of CBC, 5 m, 1 Mar 1995, J. and D. Tyler, A. Sundberg. Paratypes: Dark-headed males from holes in coral: USNM 337497, 1, 11.9 mm 25 SL, from hole in dead coral, same data as holotype (notes on color in life). USNM 337509, 1, 19.4 mm SL (cleared and stained), from hole in dead coral, patch reef about 800 m southwest of CBC, 8 m, 12 Mar 1994, J. and D. Tyler, A. Sundberg. USNM 337510, 2, 13.5-14.3 mm SL (cleared and stained), from holes in dead coral, spur and groove formation just east of CBC, 7 m, 7 Mar 1995, J. and D. Tyler, A. Sundberg. USNM 325490, 1, 13.6 mm SL, from hole in living Porites astreoides, surge channel in forereef along east side of CBC, 2.5) m,; 19 Mar 1991, R:; and -K. Clarke (the only hole-dwelling male from living coral). USNM 325491, 2, 13.4—16.0 mm SL, from holes in dead Acropora pal- mata, back of reef crest along east side of CBC, 1 m, 21 Mar 1991, R. and K. Clarke. USNM 337499, 5, 12.9-15.7 mm SL, from holes in different but close-by large heads of dead coral (four from the sides of the head and one from the top) marked with ribbons for subsequent observations after removal of original occupants, patch reef just northwest of CBC, 1.5 m, 3 Mar 1995, J. and D. Tyler. USNM 337498, 1, 15.2 mm SL, replacement from one of the holes from which individuals were removed on 3 Mar 1995 (see preceding entry), patch reef just northwest of CBC, 1.5 m, 7 Mar 1995, J. and D. Tyler. USNM 337500, 1, 12.6 mm SL, from hole in dead coral marked with ribbon for subsequent observations after re- moval of original occupant (no replace- ments seen during next seven days of ob- servations), patch reef just northwest of CBC, 1 m, 28 Feb 1995, J. and D. Tyler (notes on color in life). USNM 337501, 1, 13.6 mm SL, from hole in dead coral rub- ble, patch reef at Curlew Cay, southeast of CBC, 5 m, 2 Mar 1995, J. and D. Tyler, A. Sundberg (notes on color in life). USNM 337502, 1, 12.1 mm SL, from hole in dead part of otherwise living Montastrea sp., patch reef about 800 m southwest of CBC, 8 m, 3 Mar 1995, J. and D. Tyler, A. Sund- berg. USNM 337503, 1, 17.6 mm SL, from hole in dead part of otherwise living Diplo- 26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ria sp., spur and groove formation just east of CBC, 5 m, 15 Mar 1995, J. and D. Tyler, A. Sundberg. USNM 337504, 2, 13.6—15.0 mm SL, from holes in dead coral (openings of 2.7 and 3.0 mm diameter; burrow with the 3.0 mm opening collected), patch reef just northwest of CBC, 1 m, 13 Mar 1994, J. and D. Tyler. USNM 337505, 1, 12.5 mm SL, from hole in dead part of otherwise liv- ing Montastrea sp., patch reef about 1000 m southwest of CBC, 8 m, 6 Mar 1994, J. and D. Tyler, A. Sundberg. USNM 337506, 4, 14.2-17.0 mm SL, from holes in dead coral, patch reef just northwest of CBC, 1 m, 12 Mar 1994, J. and D. Tyler. USNM 337507, 2, 12.9-13.0 mm SL, from holes in dead coral (both openings 2.5 mm di- ameter), spur and groove formation just east of CBC, 8 m, 13 Mar 1994, J. and D. Tyler, A. Sundberg. USNM 337508, 1, 15.3 mm SL, from hole in dead Acropora palmata (opening of 5.3 mm diameter; burrow col- lected), patch reef just northwest of CBC, 1 m, 6 Mar 1994, J. and D. Tyler. USNM 341534, 1, 16.6 mm SL, from hole in dead part of otherwise living Montastrea sp., patch reef just northwest of CBC, 1 m, 16 Mar 1996, J. and D. Tyler (see below for a pale male collected nearby on the surface, USNM 341535). USNM 341536, 1, 15.2 mm SL, from hole in dead part of otherwise living Montastrea sp., cut in back reef just east of south end of CBC, 1 m, 21 Mar 1996, J. and D. Tyler (notes on color in life). USNM 341537, 1, 13.8 mm SL, from hole in dead part of otherwise living Mon- tastrea sp., patch reef just northwest of CBC, 1 m, 24 Mar 1996, J. and D. Tyler (see below for a female collected at the Same time nearby on the surface, with the same catalogue number). ANSP 173572, 1, 15.6 mm SL, from hole in dead part of oth- erwise living Montastrea sp., patch reef just off south end of CBC, 1 m, 6 Mar 1994, J. and D. Tyler, R.E. Clark (notes and photo- graph in laboratory of color in life). ANSP 173573, 1, 13.1 mm SL, from hole in dead coral rubble, spur and groove formation just east of CBC, 3 m, 12 Mar 1994, R. and K. Clarke. ANSP 173574, 2, 13.4-15.3 mm SL, from holes in dead Acropora palmata, spur and groove formation just east of CBC, 5 m, 15 Mar 1994, J. and D. Tyler. Pale male from surface of coral: USNM 341535, 1, 14.6 mm SL, from surface of living part of Montastrea sp., within 20 cm of hole in dead part of coral from which a dark-headed male was collected just mo- ments before (see USNM 341534, above), patch reef just northwest of CBC, 1 m, 16 Mar 1996, J. and D. Tyler. Females: USNM 325489, 1, 12.3 mm SL, from surface of algal-turf covered coral boulder, back of reef crest along east side of CBC, 1 m, 18 Mar 1991, R. and K. Clarke. USNM 341537, 1, 13.4 mm SL, from surface of algal-turf area in dead re- gion of a mostly living Montastrea sp., within 10 cm of hole in dead part of coral from which a dark-headed male was col- lected just moments before (see above un- der same catalogue number), patch reef just northwest of CBC, 1 m, 24 Mar 1996, J. and D. Tyler. Description.—See Table 1 for meristics and Table 2 for measurements. Snout short, slightly less than diameter of orbit. Supraorbital cirrus short, simple; cir- rus length usually less than one-half pupil diameter (and about equal to length of nasal cirrus), sometimes about one-half pupil di- ameter, and about equal to pupil in largest specimen. Anterior nasal opening tubular, with a simple cirrus projecting from rear edge; cirrus length about equal to length of tubular part. Depth of head about 6—7 times in SL. Depth of body at anal-fin origin about 7-8 times in SL. Anterior two or three dorsal-fin spines moderately elongate, second spine usually longest (but often the first longest), greatest spine-length about 2 times in head length; interradial membrane more deeply incised in the two females and the pale surface-dwelling male than in the dark-headed hole-dwelling males. Pelvic fins shorter than pectoral fins; pelvic rays extending beyond incised membranes, third ray about one-third to two-thirds (usually VOLUME 110, NUMBER 1 97 Table 1.—Meristics of Emblemariopsis ruetzleri new species. Dorsal-fin spines XX XXI 10 il 12 13 19 20 21 zl 7. ] 23 5 1 fi 23 2 Dorsal-fin rays Anal-fin rays* Pectoral-fin rayst Caudal vertebraet ‘5 14 tS 25 26 7 56 3 1 16 A | * All specimens with two anal-fin spines. + Two specimens with 13 rays on both sides; three specimens with 13 rays on one side and 14 on the other; one specimen with 15 rays on both sides; one specimen with 14 rays on one side and 15 on the other. + The number of abdominal vertebrae was not clearly visible on radiographs, but all three cleared and stained specimens have 11 abdominal vertebrae, and we presume that there are usually a total of 37 veterbrae. one-half) length of second ray; pelvic spine short and only visible in cleared and stained specimens. Dorsal fin usually XX,11; anal fin usu- ally 11,20; pectoral fin usually 14 on both sides; pelvic fin I,3 (spine inconspicuous); caudal fin with 13 segmented unbranched rays and 2-4 procurrent rays above and be- low. About 15-20 teeth in upper jaw, with fifth, sixth, and sometimes seventh from front enlarged on both sides. About 17—19 teeth in lower jaw, the fifth to eighth or ninth from front enlarged on both sides. About 7-8 large palatine teeth, in a cres- cent. About 7—8 moderate-size vomerine teeth. Gill rakers two above and six below the angle (from a cleared and stained spec- imen). There being no standardized terminology for the cephalic sensory pore system widely accepted for blennies, we prefer to illustrate the major features in the diagnostically im- portant orbital and temporal regions (Fig. 1) rather than to verbally describe them. We follow the terminology used for Emblema- riopsis and related genera by Stephens (1970) in simply noting that there is a sin- Table 2.—Measurements of Emblemariopsis ruetzleri new species, in percent of standard length (SL). Dark-headed males from holes in coral Pale male from Pale female from Number of Range in values Average Value for coral surface algal turf Character specimens in percent of SL value holotype (one specimen) (one specimen)* Head length 197 23.3—27.8 25.8 258 24.7 26.1 Head depth 18 13.7—16.3 14.7 13.9 13.7, 14.9 Snout length 16 4.6-6.1 SE 5.9 3 6.0 Orbit diameter 15 6.1-7.3 6.6 6.9 6.8 7.1 Interorbital width 15 2.6—3.6 a9 2.8 a 3.0 Body depth 10.4—15.0 12.8 11.8 11.6 mt Pelvic-fin spine length ay 1.5—2.1 1.8 — —_— — First pelvic-fin ray length 12 9.0-11.3 10.5 10.4 9.2 9.0 Second pelvic-fin ray length 12 12.8-15.4 14.3 14.6 13,7 13.4 Third pelvic-fin ray length 11 7.2-9.2 8.4 8.7 i ta Pectoral-fin length 12 18.4—21.6 19.8 20.1 pie 26.1 Spiny dorsal-fin height 17 10.8—14.0 12.6 125 14.4 15.7 * A second female specimen is too poorly preserved and twisted for most measurements to be meaningful, but it has a relatively high spiny dorsal fin at 15.4% SL and a long pectoral fin of 26.0% SL like the female in the table. + RANGE 12.1—19.4 mm SL. + Cleared and stained specimens, 13.5—19.4 mm SL. 28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON d.f. Fig. 1. Cephalic sensory pore system in Emblemariopsis ruetzleri, based mostly on USNM 337505, 12.5 mm SL: above, pores of the infraorbital and the preopercular-temporal series; below, pores in the interorbital area and the more posterodorsal ones of the infraorbital series and those of the preopercular-temporal series evident in dorsal view. Scale line applies to both dorsal and lateral views. Abbreviations: a.n. = anterior nostril; p.n. = posterior nostril; d.f. = dorsal fin; t.p. = temporal pore; s.c. = supraorbital cirrus. There are no postorbital pores. VOLUME 110, NUMBER 1 Fig. 2. Living dark-headed male of Emblemariopsis ruetzleri from a hole in a dead part of an otherwise living head of Montastrea sp., ANSP 173572, 15.6 mm SL, 35 mm photograph of specimen under anesthesia in the lab, showing a typical amount of melanophore development for mature males; patch reef just off south end of Carrie Bow Cay, Belize, 1 m. gle temporal pore (Fig. 1, lateral view) and that postorbital pores are absent. Coloration of dark-headed, hole-dwelling males in life (Fig. 2): The following de- scription is based on notes from five males (see list of type specimens) collected on separate occasions. Each specimen was kept alive under light anesthesia with quinaldine for several hours in the laboratory before preservation. These males have low anal mounds, prominent genital papillae, and elongate, relatively plump testes. The head of living males is uniformly jet black when observed protruding from holes in coral. However, immediately upon emer- gence from the hole after application of quinaldine and capture in a clear plastic test tube, the head notably lightens and small dark spots predominate over a yellowish green background. The rest of the body is relatively transparent except for dark spot- ting on the spinous dorsal fin, scattered black and white markings on the bases of the dorsal and anal fins, yellowish green suffusions over parts of the body, and a prominent internal pattern of dark and light bars on the upper abdomen and along the vertebral column and pterygiophores. The entire head is covered with numer- ous small black spots; those on the preo- percular region of the cheek tend to be larg- er than those on the snout and the opercle, and two larger black spots are especially prominent on the lower jaw. All of these black spots (whose expansion in undis- turbed specimens presumably leads to the jet black head seen protruding from the hole of the burrow) are underlain by a yellowish to gold-green coloration, with some patchy areas of chalky white. There are also a few discrete chalky white markings in the fol- lowing locations: one is along the lower rear of the orbit; one is just above and be- hind this mark; and several other smaller ones are scattered around the cheek region, including one or two that seem to outline cephalic lateral-line pores. The black pupil is surrounded by a narrow ring of golden yellow, outside of which the rest of the eye is creamy with gold-brown suffusions, blu- ish to purplish to yellow-green iridescence, and irregular streakings of black. The su- praorbital cirrus is creamy with black spots or streakings; the nasal cirrus is sometimes dark spotted. As seen dorsally on the head, there is a large, roundish area in the occip- ital region from the rear of the eyes to the origin of the spinous dorsal fin that has a creamy basal color overlain by golden yel- low, with numerous small spots of grayish blue to greenish iridescence and even smaller and fewer black dots. The black spotting of the head broadly continues posteriorly onto the lateral side of the body to the level of the pectoral-fin base; dorsal to the pectoral-fin base the re- gion of spotting tapers posteriorly variously to the base of about the eighth to the last dorsal-fin spine. There is a relatively unpig- mented region between this tapered region 30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of spotting and the upper edge of the rear half of the abdominal cavity. The fleshy base of the pectoral fin has black spots about as large as those on the preopercular region. The basal one-fifth to one-third of the pectoral-fin rays has chalky white markings overlain with much smaller black spots; sometimes there is one large irregular white mark and/or sometimes sev- eral more rounded ones more distal regions of the pectoral-fin rays are pale. The rest of the thoracic area toward the isth- mus and the lower half of the abdomen be- hind the pectoral-fin base are covered by small black spots. The dorsal half of the abdominal cavity from behind the pectoral-fin base has a dis- tinctive pattern of internal coloration. There is a chalky white band that is transected in the middle by a vertical saddle-shaped bar of blackish to dark yellow-green pigment (color partially dependent on both the con- dition of the living specimen and the angle of observation), with the dark saddle being wider dorsally than ventrally. Ventral to this chalky, saddled, upper region of the abdom- inal cavity, the lower abdomen is slightly yellowish with small black external spots. There is a dark spot at the front of the anal region and a white spot just behind it. The membranes between the first three dorsal-fin spines are chalky white with black spots and, especially basally, black reticulations. There are scattered black spots covering most or parts of the other- wise pale interradial membranes posterior to the third dorsal-fin spine variously to the eighth to last dorsal-fin spine. The density of the spotting and its posterior extent is a function of the degree of overall body dark- ness. Posterior to the darkened part of the spinous dorsal fin, the interradial mem- branes are usually without many melano- phores and become more uniformly pale. In the paler parts of the dorsal fin, each spine or soft ray has specks or slightly larger dots of black along much of the length of the element or immediately alongside it but not broadly on the interradial membrane; there also are a few white spots along some of these fin elements. At the base of each dor- sal-fin element is a discrete spot of either black or white, with the two colors usually alternating with one another on the bases of successive elements. The anal fin has essentially no pigmen- tation on the interradial membranes, but there are black specks along most of the fin rays. At the base of each anal-fin spine and ray is a discrete spot of either black or white, alternating on successive elements just as in the dorsal fin (with the base of the first ray following the two spines usu- ally with the first black spot). The caudal fin is pale except for two white elongate marks, one along the bases of several rays in the upper lobe and one along the bases of several rays in the lower lobe, and several smaller white spots of variable occurrence more distally on the fin. Besides the external pigmentation pat- terns described above, there is little other surface pigmentation on the body. The transparent body is, however, distinctly pat- terned by internal markings associated with the vertebrae and pterygiophores, as well as by those of the abdominal cavity. There are rusty pink patches of internal color on the top of the head and the top of the iris. The vertebral column has prominent dark markings that alternate with shorter white marks to form a continuous series along the top of the centra. The dark markings range in color from blackish to dark golden green (depending on specimen condition and an- gle of observation), and the white markings are creamy. The white markings span the tops of two centra and have no ventral ex- tensions. The dark markings usually span three or four centra, and, at their anterior and posterior ends, they have ventral exten- sions along the front of the centra that con- tinue down the haemal spines. These dark markings extend only partially down the length of the haemal spines in the region just behind the level of the anus, but they extend most of the way down the lengths of more posterior haemal spines. The net VOLUME 110, NUMBER 1 31 Fig. 3. Preserved holotype of Emblemariopsis ruetzleri from a hole in a dead coral of unidentified species, USNM 337496, 14.4 mm SL; spur and groove formation just east of Carrie Bow Cay, Belize, 5 m. effect is of short white horizontal markings separating longer dark horizontal markings along the top of the vertebral column, with dark ventral extensions onto every other or every third or fourth haemal spine. Coloration of preserved dark-headed males (Figs. 3, 4): With preservation, the transparent body becomes increasingly Opaque, and nearly all of the internal col- oration of the vertebral column, pterygio- phores, and abdominal cavity is obscured; the surface melanophores form nearly all that remains of the pattern seen in life. Of the internal coloration, only rusty or pink Fig. 4. Preserved specimens of a 13.8 mm SL dark-headed male (above) and a 13.4 mm SL female of Emblemariopsis ruetzleri collected within moments of one another, the male from a hole in a dead part of an otherwise living head of Montastrea sp. and the female from the surface of the algal turf within 10 cm of that hole, both specimens USNM 341537; patch reef just northwest of Carrie Bow Cay, Belize, | m. 32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Preserved pale male of Emblemariopsis ruetzleri from the surface of the living part of a mostly living head of Montastrea sp. (collected within moments of a dark-headed male from a hole 20 cm away in a dead part of the coral head), USNM 341535, 14.6 mm SL; patch reef just northwest of Carrie Bow Cay, Belize, 1 m. patches on the top of the head and the top half of the iris usually are evident. All of the chalky white and greenish yellow to grayish blue iridescent undertones of live specimens are lost after months in preser- vative, except that sometimes there are rem- nants of the chalky markings on the cheek and pectoral-fin base. The pattern of dark melanophores on the surface remains as de- scribed above in living specimens, with the darkest males having closely spaced spots of dark pigment over most of the head and anterior half of the body, on the entire course of the spinous dorsal fin, and along the bases of every few dorsal- and anal-fin rays. Coloration of live and preserved females and an immature male (Figs. 4, 5): When alive, females (two specimens collected, with high anal mounds and no genital pa- pillae) and immature males (one specimen collected, with low anal mound, genital pa- pilla low and not protruding, and testes thin) have transparent bodies that show all of the internal markings described above for dark-headed males. However, the only external coloration evident in life comprises some small black and white alternating spots along the bases of the dorsal-/and anal-fin elements (as above for dark-headed males) and some coloration between the first two or three dorsal-fin spines (details not specified in our notes). In preservative, the opaque body obscures all of the internal markings on the vertebral column, ptery- giophores, and abdominal cavity. Some rusty pink coloration is found on the head in the roundish occipital area, top half of the eye, and sometimes in the opercular re- gion. External melanophores (sometimes rusty in color rather than black) evident in preserved specimens are as follows: dark spots along the bases of some of the dorsal- fin and anal-fin elements; lightly scattered spots along the course of the dorsal- and anal-fin elements laterally but not on the in- terradial membranes; irregular bands of speckling in the interradial membrane be- tween the first two or three dorsal-fin spines (mostly as an irregular band just internal to the distal edge of the membrane, with some chalky color also present between the sec- ond and third spines in the larger female); and a few spots along the upper jaw and on the underside of the lower jaw. Sexual dimorphism.—In addition to ma- ture dark-headed males of E. ruetzleri dif- fering from pale females and immature males in coloration, there is dimorphism in the length of the pectoral fin and in the height of the elevated anterior part of the spinous dorsal fin. In both female speci- mens, the pectoral fin (26.0—26.1% SL) is longer and the anterior spinous dorsal-fin rays (15.4-15.7% SL) are higher than in mature males (18.4—21.6% SL for the pec- toral fin and 10.8—14.0% SL for the dorsal fin). In the single immature pale male spec- imen, the pectoral-fin length (21.2% SL) is similar to the high end of the range of ma- ture males, and the elevation of the anterior VOLUME 110, NUMBER 1 elements of the spinous dorsal fin is only slightly greater than that of mature males. In mature dark-headed males the inter- radial membrane of the anterior region of the dorsal fin is not greatly indented be- tween the spines, but in both of the female specimens and in the immature male spec- imen the interradial membrane is deeply in- cised (to about one-half of its greatest depth) between the third and fourth dorsal spines. This difference between mature males and females in the depth of the in- dentation between the third and fourth spines also is evident in Emblemariopsis occidentalis (see figures in BOhlke & Chap- lin 1968:548, incorrectly identified as Pseu- demblemaria signifera, as shown by Ste- vens 1970). Characters supporting generic alloca- tion.—Although Acero (1987) recommend- ed synonymizing Emblemariopsis with Coralliozetus, we follow the ongoing cla- distic-based revisions of various chaenop- sids by Hastings (e.g., 1992, 1997) and Hastings & Springer (1994) in recognizing the distinctiveness of Emblemariopsis with- in a Coralliozetus-like clade. The most recent revision of the taxono- my of the species of Emblemariopsis and related genera is that of Stephens (1970). Emblemariopsis ruetzleri has all of the di- agnostic features given by Stephens for that genus (Stephens 1970:283; based on seven species, E. bahamensis, E. bottomei, E. dia- phana, E. leptocirris, E. occidentalis, E. signifera, E. randalli, i.e., all but E. pricei, E. ruetzleri, and E. tayrona). These features are as follows (condition of E. ruetzleri giv- en in parenthesis if there is variation in the generic character): no spinous projections on the frontal or parietal; nasals separate; snout short; pelvics shorter than pectorals; pelvic fin with membranes deeply incised between first two rays and last pelvic ray short; one or two temporal pores (one in E. ruetzleri); no postorbital pores; dorsal fin not sail-like; dorsal fin XIX—XXII, 10-14 (usually XX, 11 in E. rvetzleri), anal fin II, 19-23 (usually II, 20 in E. ruetzleri); pec- 33 toral rays 12—14 (usually 14 in E. ruetzleri); no anterior flap on first dorsal spine; first three dorsal spines closely spaced; 5-9 moderate- to large-size teeth on palatine (7— 8 large-size teeth in E. ruetzleri); 6—12 small- to moderate-size teeth on vomer (7— 8 moderate-size teeth in E. ruetzleri); su- praorbital cirrus either present and short or absent (short in E. rvetzleri); nasal cirrus simple or absent (simple in E. ruetzleri); sexual dimorphism well developed or ab- sent (dimorphism in coloration, pectoral-fin length, and height of spinous dorsal fin in E. ruetzleri). In conjunction with the description and phylogenetic relationships of a new genus of Chaenopsidae, Hastings (1992: table 1), mentioned five derived characteristics that in combination are unique to the three spe- cies of Emblemariopsis that he studied (E. diaphana, E. signifera, E. tayrona). Emble- mariopsis ruetzleri has all five of these de- rived features: females with pelvic-fin rays prolonged, the membranes not extending to the tips; males same as preceding; meso- pterygoid absent; nasals with ridges; and third pelvic-fin ray reduced, its length about one-half that of second ray but longer than pelvic spine. Hastings (1992) also listed another four derived features, each of which is found in one or two but not all three of the species he examined: supraorbital cirrus present (primitive, the condition in E. ruetzleri) or absent; fleshy flap anteriorly on first dorsal- fin spine present (primitive) or absent (as in E. ruetzleri); dorsal fin of males relatively low (primitive, the condition in E. ruetzleri) or elevated; and third basibranchial ossified (primitive) or unossified (unossified in the two smaller cleared and stained specimens of E. ruetzleri, 13.5 and 14.3 mm SL, but ossified in the largest specimen, 19.4 mm SL). In a manuscript nearing completion, P. Hastings (pers. comm.) describes a syna- pomorphy of the four species of Emble- mariopsis he has examined (the three spe- cies given in Hastings 1992, plus E. lepto- 34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cirris): the neural spine of the penultimate vertebra is truncate and much shorter than the preceding neural spines and the single epural is expanded proximally and occupies the position of the neural spine. Emblema- riopsis ruetzleri has a shortened penulti- mate neural spine and expanded epural (as does E. pricei; Fig. 6), which gives us fur- ther assurance of the proper generic allo- cation of the new species. Habitat and natural history notes.—Ma- ture males are dark-headed and inhabit holes in coral, usually dead coral, whereas females are pale and are found on the sur- face of the algal turf overlying dead coral and near the hole-dwelling males; the single immature, pale male collected during the study was on the surface of a live part of a coral that also had a dead section containing a dark-headed male in a hole. Two burrows of E. ruetzleri were col- lected along with their male residents. The internal surface of both burrows was smooth and brownish black, and each was the empty coiled shell of a vermetid gastro- pod. M. G. Harasewych (pers. comm.) ten- tatively identified these two shells, on the basis of their structure and locality, as those of Dendropoma, perhaps D. irregularis. We have not collected any other chaenopsid at Carrie Bow Cay inside a vermetid shell. Both burrows contained both eyed and eyeless eggs that were being incubated by the resident male. Moreover, both burrows had at least two different developmental stages of eyed and eyeless eggs, indicating several different clutches. One of the bur- rOWS was So intact that almost the full com- plement of eggs could be counted; there were about 32 eyeless and 58 eyed eggs, a total of about 90 (+3). This burrow had an opening diameter of 5.3 mm and a similar diameter deep inside the coil where the eggs (both eyed and eyeless) were attached. The other burrow was broken in the region where eggs were attached, and a partial count of those remaining yielded about 60 eggs. This burrow had an opening diameter of 3.0 mm and was about 3.7 mm diameter lower inside the coil in the region of the eggs. The eggs in both burrows were be- tween 0.6—0.7 mm in diameter. The smaller of the two female specimens collected during this study, 12.3 mm SL, is not fully ripe. The right side ovary has 13 developing eggs of about 0.25—0.3 mm di- ameter and many much smaller egg pri- mordia. The larger female, 13.4 mm SL, is fully ripe, with nine eggs of 0.55—0.6 mm diameter on the right side and only a few much smaller eggs. We presume, therefore, that there are a total of about 18 eggs avail- able for laying by this mature female. This is further confirmation that burrows with about 90 incubating eggs must be the result of multiple deposits. To determine whether vacant holes in dead coral were as keenly competed for by E. ruetzleri as are those in live coral by E. pricei (Tyler & Tyler 1997), we removed a dark-headed male (USNM 337500) of E. ruetzleri from a hole in a shallow patch reef just off the northwest side of Carrie Bow Cay on 28 Feb 1995, marked the location with a ribbon, and returned to it daily to see if a replacement had entered the empty hole. No replacement individual was pres- ent during the seven days of subsequent monitoring. Additionally, on 3 Mar 1995 we collect- ed five dark-headed males of E. ruetzleri (USNM 337499, mature males, 12.9—15.7 mm SL) on a shallow patch reef on the northwest side of Carrie Bow Cay, and marked their holes with ribbons. The five collected specimens were kept alive in the laboratory. They had blanched only mod- erately when collected and thereafter in the aquaria, and had good retention of darkness in preservative. We next observed the marked holes on 7 Mar and found that four of them were empty whereas one had a dark-headed replacement that we removed. The replacement was a 15.2 mm male (USNM 337498) that blanched consider- ably when collected. It had a minimal speckling of melanophores on the head and body, and it is now one of the least dark VOLUME 110, NUMBER 1 35 Fig. 6. Caudal skeletons in two specimens of, above, Emblemariopsis ruetzleri, USNM 337510, 13.5 mm SL to left and USNM 337507, 19.4 mm SL to right, and, below, two specimens of E. pricei, both USNM 325446, 13.6 mm SL to left and 16.5 mm SL to right, to show the derived condition in both species of the short neural spine of the penultimate centrum, typical of most species of the genus. The caudal skeleton has a single epural, hypural 5 free, and hypurals 1—4 consolidated. Abbreviations: h5 = fifth hypural; e = epural. males in preservative. We conclude that, in marked contrast to E. pricei, holes in coral appropriate for habitation of mature males may not be in short supply for E. rvetzleri. We presume that E. ruetzleri feeds on mi- croplankton in the water column based on finding the remains of small or larval crus- taceans in the stomach contents of several specimens. We found the holes occupied by E. ruet- zleri on both the relatively horizontal (top) and vertical (side) surfaces of dead corals, but we did not find a predominance of one orientation versus the other. 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Etymology.—We take pleasure in naming this new species in honor of Dr. Klaus Ruetzler, Curator of Invertebrate Zoology at the National Museum of Natural History, who has so effectively directed the Smith- sonian’s Caribbean Coral Reef Ecosystem (CCRE) Program and its marine laboratory at Carrie Bow Cay, Belize, since its incep- tion in 1972; he has given generously of his time and talents for all those years in pro- viding research opportunities for many Smithsonian scientists and their colleagues throughout the world. Discussion Emblemariopsis ruetzleri and E. pricei are the only species of chaenopsids with strikingly dark-headed males living in holes in large coral mounds that are relatively common around Carrie Bow Cay. The hole-dwelling males of both of these species are found in similar habitats around the reef crest, on patch reefs in the lagoon, and in the spur and groove zone around Carrie Bow Cay (to at least 8 m depth for E. ruetzleri, and deeper for E. pricei). Ma- ture males of E. ruetzleri, however, are al- most always found in holes in dead coral (only one of 35 dark-headed males in a hole in living coral), whereas those of E. pricei are almost always found in holes in live coral. Most of the burrows that have been col- lected of breeding males of E. pricei are in the empty, partially tubular, and partially ir- regularly eroded tubes of serpulid worms (Spirobranchus) embedded in coral, but the two burrows that have been collected of breeding males of E. ruetzleri were both in empty, coiled, smooth-surfaced tubes of vermetid gastropods embedded in coral. Another chaenopsid present around Car- rie Bow Cay, the sailfin blenny Emblemaria pandionis Evermann & Marsh, is found in irregular cavities formed by many different kinds of invertebrates (worms, bivalves, Snapping shrimps) and by erosion, in a great variety of sizes and shapes of dead coral rubble. In a few cases these cavities are probably the old corroded remains of vermetid gastropods (Smith & Tyler, in litt.). However, the cavities occupied by E. pandionis are of various shapes and are rough surfaced in loose rubble, in compar- ison to those of the complete coiled smooth tubes in large coral heads or boulders oc- cupied by Emblemariopsis ruetzleri. Some chaenopsids in the eastern Pacific are known to have their shelter burrows in va- cated mollusk tubes, including Emblemaria hypacanthus (Jenkins & Evermann) in Tur- bo gastropod shells (Hastings 1991) and Acanthemblemaria crockeri Beebe & Tee- Van in, among other kinds, Lithophaga bor- ing bivalve cavities (Hastings 1988). Another ecological difference between the two common species of Emblemariopsis at Carrie Bow Cay is the habitat of females, nonbreeding males, and immatures. Those of E. pricei are found mostly on the surface of soft-coral sea fans (Gorgonia ventalina Linnaeus) and clumps of live elkhorn coral (Acropora palmata (Lamarck)), whereas fe- males of E. ruetzleri are on the surface of the algal turf on the same kinds of coral heads and boulders as the holes of the breeding males, and the immature males are also surface dwelling (on live coral near dead sections of coral for the single im- mature specimen collected). The new species is one of the many chaenopsids living at least some of its life history in holes in dead coral around Carrie Bow Cay. All five species of Acanthemble- maria found there have both males and fe- males in holes in dead coral (Clarke 1994), in contrast to E. ruetzleri having only dark- headed breeding males in such holes. Hole- dwelling males of the two common species of Emblemariopsis around Carrie Bow Cay are easily distinguished from the five spe- cies of Acanthemblemaria there because the heads of the species of the latter genus have Spiny or rugose upper surfaces and are nev- er as darkly and uniformly black as those of the two species of Emblemariopsis. There are several other chaenopsids be- VOLUME 110, NUMBER 1 sides Emblemariopsis ruetzleri and E. pri- cei in which mature males utilize different habitats than females and immature males. In the sailfin blenny, Emblemaria pandion- is, at Carrie Bow Cay, mature males are res- ident in holes in coral rubble, but females and immature males move about much more freely along the sandy bottom and fre- quently change their rubble shelter sites (Smith & Tyler, in litt.). In the Gulf of Cal- ifornia sailfin blenny, Emblemaria hypa- canthus, Hastings (1991) has shown that in areas of low predation risk, males are close- ly tied to their shelters whereas females move about more freely in the open, but that in areas of high predation both sexes spend most of their time in shelters. Has- tings (1986) also has shown that most males of Coralliozetus angelica (Bohlke & Mead) in the Gulf of California are resident in bar- nacle shells, but females spend much time foraging in the open. With sexually active males ranging in size from 11.9 to 19.4 mm SL (X¥ = 14.5 mm SL for 35 individuals), E. ruetzleri is a relatively small species. A few other spe- cies of Emblemariopsis apparently are also relatively small, such as E. leptocirris and E. occidentalis, which have maximum known sizes of about 21 mm SL. Others are only slightly larger, such as E. bahamensis, E. diaphana, E. signifera, and E. tayrona, which reach between 23—25 mm SL (Ste- phens 1963, 1970; Acero 1987). The largest members of the genus are E. randalli, E. bottomei, and E. pricei, which have maxi- mum sizes of 28-30 mm SL (Cervigon 1965, Stephens 1963, Tyler & Tyler 1997). Emblemariopsis ruetzleri is about as di- minutive as the smallest species of Acan- themblemaria at Carrie Bow Cay, A. paula Johnson & Brothers, which has a maximum size of about 18 mm SL and is sexually active as small as 11 mm SL (Johnson & Brothers 1989). Acknowledgments We greatly appreciate the help we re- ceived in our field work at Carrie Bow Cay 37 from the following individuals: Raymond and Karen Clarke, Sarah Lawrence College; Arthur Sundberg, University of Delaware; and Roy Clark, Jr., Smithsonian Institution. Much information on chaenopsid system- atics and behavior was generously given to us by Philip Hastings, University of Ari- zona, and Raymond Clarke. Photographic services were provided by Roy Clark, Jr., and Carl Hansen, Smithsonian Institution. We are indebted to M. G. Harasewych, Smithsonian Institution, for his time and ex- pertise in identifying the gastropod source of the burrows of the new species. We thank Sandra Raredon, Lisa Palmer, and Kris Murphy, Smithsonian Institution, for radiographs, loans, and cataloguing related to this work, and Eugenia Bohlke and Wil- liam Saul for the loan of materials at the Academy of Natural Sciences of Philadel- phia. The manuscript benefited from the suggestions by Carole Baldwin and Jeffrey Williams, Smithsonian Institution. This is Contribution 491 from the Smith- sonian Institution’s Caribbean Coral Reef Ecosystems Program. Literature Cited Acero, A. 1987. The chaenopsine blennies of the southwestern Caribbean (Pisces, Clinidae, Chaenopsinae). III. The genera Chaenopsis and Coralliozetus.—Boletin Ecotropica, Fundacion Universidad de Bogota Jorge Tedeo Lozano 16: 1-21. Bohlke, J. E., & C. ‘C. G. Chaplin. 1968. Fishes of the Bahamas and adjacent tropical waters. Academy of Natural Sciences of Philadelphia, 771 pp. Cervigon, F 1965. Emblemariopsis randalli nov. sp., una nueva especie de Chaenopsidae de las cos- tas de Venezuela.—Novedades Cientificas, Contribuciones Ocasionales del Museo do His- toria Natural La Salle, Serie Zoolégica 33:1—4. Clarke, R. D. 1994. Habitat partitioning by chaenop- sid blennies in Belize and the Virgin Islands.— Copeia 1994(2):398—405. Greenfield, D. W. 1975. Emblemariopsis pricei, a new species of chaenopsid blenny from Belize.— Copeia 1975(4):713-715. Hastings, P. A. 1986. Habitat selection, sex ratio and sexual selection in Coralliozetus angelica (Blennioidea: Chaenopsidae). Pp. 785-793 in T. 38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Uyeno, R. Arai, T. Taniuchi, and K. Matsuura, eds., Indo-Pacific fish biology: Proceedings of the Second International Conference on Indo- Pacific Fishes. Ichthyological Society of Japan, 985 pp. 1988. Correlates of male reproductive suc- cess in the browncheek blenny, Acanthemble- maria crockeri (Blennioidea: Chaenopsidae).— Behavioral Ecology and Sociobiology 22:95-— 102. 1991. Flexible responses to predators in a marine fish.—Ethology Ecology & Evolution 3: 177-184. . 1992. Phylogenetic relationships of Tanyem- blemaria alleni, a new genus and species of chaenopsid (Pisces: Blennioidei) from the Gulf of Panama.—Bulletin of Marine Science 51(2): 147-160. . 1997. Phylogenetic relationships of the Cor- alliozetus clade of chaenopsid blennies, with description of a new genus (Teleostei, Blen- nioidei).—Bulletin of Marine Science (in press). , & V. G. Springer. 1994. Review of Stath- monotus, with redefinition and phylogenetic analysis of the Chaenopsidae (Teleostei: Blen- nioidei).—Smithsonian Contributions to Zool- ogy 558:1—48. Johnson, G. D., & E. B. Brothers. 1989. Acanthem- blemaria paula, a new diminutive chaenopsid (Pisces: Blennioidei) from Belize, with com- ments on life history—Proceedings of the Bi- ological Society of Washington 102:1018—1030. Stephens, J. S. 1963. A revised classification of the blennioid fishes of the American family Chaen- opsidae.—University of California Publications in Zoology 68:1—165. . 1970. Seven new chaenopsid blennies from the western Atlantic.—Copeia 1970(2):280-— 309. Tyler, J. C., & D. M. Tyler. 1997. Natural history of the sea fan blenny, Emblemariopsis pricei (Pis- ces, Chaenopsidae), in the western Caribbe- an.—Smithsonian Contributions to Zoology (in press). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):39—48. 1997. Pontoniine shrimps (Decapoda: Caridea: Palaemonidae) of the northwest Atlantic. V. Periclimenes mclellandi, a new species, a gorgonian associate from Pine Cay, Turks and Caicos Islands, British West Indies Richard W. Heard and Stephen Spotte (RWH) Invertebrate Zoology Section, Institute of Marine Sciences, University of Southern Mississippi, P.O. Box 7000, Ocean Springs, Mississippi 39566, U.S.A.;(SS) Marine Sciences & Technology Center, The University of Connecticut at Avery Point, Groton, Connecticut 06340, U.S.A. Abstract.—Specimens of Periclimenes mclellandi, new species, were col- lected from gorgonians, primarily Pseudopterogorgia americana Gmelin, at depths of 14 to 43 m off Pine Cay, Turks and Caicos Islands, British West Indies. The new species belongs to the “‘iridescens’’ complex and is similar to P. patae Heard & Spotte, 1991, also associated with gorgonians at Pine Cay and the Florida Keys, but at shallower depths. Periclimenes mclellandi is dis- tinguished from P. patae by a bilobate endite on the second maxillae, two rows of combsetae on the propodus of fifth pereopods, shorter and deeper rostrum with fewer rostral teeth, and an overall greenish coloration in life. Both species are distinguished from other described members of the ‘‘iridescens’’ complex by the chelae of the second pereopods, which are similar and equal or nearly equal in size, and by the dactyls of pereopods 3-5, which are simple instead of biunguiculate. Some specimens of P. mclellandi were parasitized by an unidentified aceolid trematode metacercaria and an undescribed bopyrid isopod. This report is fifth in a series devoted to the taxonomy, distribution, and ecology of shrimps belonging to the palaemonid sub- family Pontoniinae. Previous contributions to this series are Heard (1986), Heard & Spotte (1991), Heard et al. (1993), and Spotte et al. (1994). An undescribed species of Periclimenes Costa, 1844, was discovered on colonies of gorgonians, primarily Pseudopterogorgia americana (Gmelin, 1791), along the fore- reef slope west of Pine Cay in the Turks and Caicos Islands, British West Indies, near 21°53'N, 72°05'W. The new species resembles P. patae Heard & Spotte, 1991, another gorgonian associate from Pine Cay and the Florida Keys. Carapace length (CL) is defined here as the distance from the tip of the rostrum to the posterodorsal margin of the carapace. Type material has been de- posited in the collections of the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM), and Gulf Coast Research Laboratory Museum (GCRL). Periclimenes mclellandi, new species Figs. 1-5 Periclimenes sp. A.—Spotte & Bubucis 1996:229 Material examined.—Holotype, Oviger- ous female, CL = 2.40 mm, (USNM 276042), forereef slope west of Pine Cay, Turks and Caicos Islands, British West In- dies, on gorgonian Pseudopterogorgia americana, 29 m, 14 Nov 1989, colls. S. Spotte and P. M. Bubucis. Paratypes from P. americana, same general locality as ho- lotype, colls. S. Spotte and P. M. Bubucis. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 40 ‘Ul CT ‘O66I Alng ¢ ‘spue]s] soored pue syiny ‘AeD ould 1 ns ul poydeisojoyg ‘Spueq ISISASUBT] OM} 9} 9ION ‘UBIUOSIOS PoyMUspiuN ue UO J[VUISJ SNOIOSIAO UL JO MIIA [eIO}e] ‘sotoeds MoU ‘1IPUD]JaJIU SauamMyouad “| ‘SIyq a VOLUME 110, NUMBER 1 4] Fig. 2. Periclimenes mclellandi, new species, lateral view of ovigerous female. Scale = 2.0 mm. Male, CL = 1.92 mm, (USNM 276043), 21 m, 8 Nov 1989; male, CL = 1.67 mm, (USNM 276044), 20 m, 9 Nov 1989; 1 ovi- gerous female, CL = 2.88 mm, 1 male, CL = 2.05 mm, (USNM 276045), 26—29 m, 14 Nov 1989; 2 females, CL = 2.40, 2.43 mm, (USNM 276046), 17-19 m, 10 May 1990; 1 male, CL = 1.92 mm, 1 female, CL = 2.46, (GCRL 1342), 26-32 m, 8 May 1990. Diagnosis.—Small (CL less than 3 mm), greenish in life. Rostrum deep, unarmed ventrally, nearly straight or curved upward at tip, in females extending anteriorly be- yond midsection of first segment of anten- nular peduncle, in males not extending to midsection of first segment of antennular peduncle; both sexes with as many as 5 dor- sal rostral teeth, rarely more than 4 in males. Carapace with well developed anten- nal and hepatic spines, lacking other spines or ornamentation. Abdominal pleura round- ed, abdominal somite 6 twice length of so- mite 5. Mesial spines on posterior margin of telson *% length of intermediate spines. Cornea slightly narrower than eye-stalk, constricted at junction with eye-stalk; ac- cessory pigment spot and associated om- matidia present on dorsoproximal margin of cornea. Maxilla 2, endite bilobate. Pereo- pods 2 equal or nearly equal, undifferen- tiated, chelae lacking distinct teeth (vestiges of teeth sometimes present). Pereopods 3— 5 with dactyls simple, propodi with 1 spine- seta on distoflexor margin; propodus of per- eopods 5 with 2 rows of comb-setae on dis- tal inner margin, (1 or both rows can be absent in small specimens). Description of adult female (Figs. 1, 2, 4a-f, h, i, 5, 6).—Carapace with rostrum curved slightly upward at tip, not reaching to distal margin of first article of antennular peduncle; 4, or occasionally 5, dorsal teeth interspersed with small setae; first tooth of rostrum usually even with hepatic spine, second tooth usually anterior to posterior margin of orbit; ventral surface without 42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figen 3: maxilliped 2 (setae omitted); f, maxilliped 3; g, eye; h, left antennule (ventral aspect); i, right antennule (ventral aspect). Scale = 0.4 mm for a—e, 1.0 mm for g-—i. teeth, posterior % with setae. Antennal spine well developed. Hepatic spine well developed, slightly larger, more robust, and less attenuated than antennal spine. No oth- er spines or ornamentation on carapace. Abdominal somites (Fig. 2) with pleura rounded, somite 6 twice length of somite 5 and same length as telson when measured along dorsal margin. Telson (Fig. 41) with 2 pairs of dorsolat- eral spines, anterior pair slightly anterior to mid-length, posterior pair closer to end of telson than to anterior pair. Three pairs of apical spine-setae; lateral pair short; inter- mediate pair longest, 4 times length of lat- eral pair; mesial pair plumose, approxi- mately %4 length of intermediate pair. Mi- nute, acute apical process present between mesial spine-setae. Antennular peduncle (Fig. 3h) with Periclimenes mclellandi, new species: a, mandible; b, maxilla 1; c, maxilla 2; d, maxilliped 1; e, sharp, slender stylocerite extending nearly to midline of basal article; distolateral mar- gin of basal article with 1 spine; combined length of second and third articles % that of basal article. Lateral antennular flagellum with 2 branches fused for approximately 5 articles; shortest unfused part of branch consisting of 3 articles, each bearing aesth- etascs (3 on terminal article, 2 on proximal unfused articles), shorter than fused portion, distal articles of which bear 2—4 aesthe- tascs. Antennal scale (Fig. 31) with just over- reaching antennular peduncle, length 3 times width, lateral margin nearly straight, distolateral spine well short of anteromesial angle of blade. Antennal peduncle extend- ing to mid-length of blade. Eye: (Fig. 3g) with cornea diameter dis- tinctly less than proximile diameter of eye- VOLUME 110, NUMBER 1 43 Fig. 4. Periclimenes mclellandi, new species (ovigerous female except for g): a, pereopod 1; b, pereopod 2 (dorsolateral aspect); c, same (lateral aspect of propodus and dactyl, setae omitted); d, same (distal aspect of fingers); e-g, pereopods 3—5; h, appendix interna and appendix masculina of adult male (CL 2.1 mm); i, telson and left uropod (fringing plumose setae omitted). Scale 1: a, d-g = 1.0 mm, b = 0.4 mm. Scale 2: c, h = 0.2 mm, i= 1.0 mm. 44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ° Carapace length, mm 1 2 3 4 5 6 Total appendix masculina spines , Carapace length, mm Dorsal rostral teeth Carapace length, mm 1 2 3 4 5 6 Dorsal rostral teeth Fig. 5. a, Males of Periclimenes mclellandi, new species, compared with males of P. patae. Standard deviation plot of carapace length against total number of spines on the appendix masculina (both apical and lateral). Bars are 1 SD with n shrimp shown above. Correlation between variables is significant for both species: P. mclellandi (Spearman’s r = 0.74, t,_, = 11.99, P <0.001); P. patae (Spearman’s r = 0.55, t,_5 = 3.24, P <0.01). b, Females of Periclimenes mclel- landi, new species, compared with females of P. patae. Standard deviation plot of carapace length against the number of dorsal rostral teeth. Bars are 1 SD with n shrimp shown above. Correlation between the vari- ables is significant for both species: P. mclellandi (Spearman’s r = 0.51, t,_, = 6.36, P <0.001); P. patae (Spearman’s r = 0.69, t,_, = 5.00, P <0.001). c, Males of Periclimenes mclellandi, new species, compared with males of P. patae. Standard deviation plot of car- apace length against the number of dorsal rostral teeth. Bars are 1 SD with n shrimp shown above. Correlation between the variables is significant for P. mclellandi stalk, barely constricted at junction with eye-stalk; accessory pigment spot and as- sociated ommatidia present on dorsoproxi- mal margin of cornea. Mandible (Fig. 3a) with lacking palp; in- cisor process terminating in 4 distinct teeth, distal tooth of incisor distinctly larger than others; molar process dentate with numer- ous small spine setae. Maxilla 1 as illus- trated (Fig. 3b). Maxilla 2 (Fig. 3c) with bilobate endite; palp slender, lacking setae. Maxilliped 1 (Fig. 3d) having well-devel- oped exopodal flagellum (lash) bearing 3 or 4 terminal plumose setae; epipod present. Maxilliped 2 (Fig. 3e) with well developed exopod bearing 2 terminal setae; epipod present. Maxilliped 3 (Fig. 3f) with well de- veloped exopod bearing 4 terminal plumose setae; exopod (excluding setae), not extend- ing to distal end of antepenultimate article of endopod. Pereopods 1 (Fig. 4a) of equal size, not extending to distal end of antennal scale; dactyls of chelae weakly bifid, same length as palm; carpus % longer than propodus; merus and carpus of equal length; ischium % length of merus. Pereopods 2 (Fig. 4b— d) equal or subequal in size, undifferentiat- ed, weakly developed, both extending past antennal scale by less than length of dac- tyls; fingers same length as palm, lacking distinct teeth (vestiges of teeth sometimes present; Fig. 4c, d); carpus less than % length of propodus; merus 7, length of car- pus, slightly shorter than ischium. Pereo- pods 3-5 (Fig. 4e—-g) of nearly equal size and shape, dactyls simple, propodi each with 1 pair of spine-setae on distal flexor margin. Pereopods 3 and 4 nearly equal in size and shape, not extending past anten- nular peduncle; pereopods 5 longer than 3 or 4, morphologically distinct. Pereopods 3 (Fig. 4e) with propodus 4 times length of 0.05). VOLUME 110, NUMBER 1 Table 1.—Periclimenes mclellandi vs. P. patae, size comparison based on carapace length (CL). Species Mean CL (+SD), mm CL range, mm P. mclellandi (males) 1.73 (0.33), n = 119 0.73-2.91 P. mclellandi (females) 1.95 (0.63), n = 172 0.72-3.17 P. patae (males) 2.40 (0.30), n = 26 1.82—3.04 P. patae (females) 2.80 (0.97), n = 29 1.11—5.02 dactyl, carpus % length of propodus, merus slightly less than twice length of carpus, is- chium slightly more than % length of me- rus. Pereopods 4 (Fig. 4f) with propodus nearly 5 times length of dactyl, carpus 7 length of propodus, merus slightly more than twice length of carpus, ischium slight- ly less than % length of merus. Pereopods 5 (Fig. 4g) longer than pereopods 3 or 4, extending to or beyond proximal end of second article of antennular peduncle; prop- odus nearly 5 times longer than dactyl, 2 rows of comb-setae on inner distal surface (Fig. 4g) (not always present on small spec- imens); carpus slightly more than 7, length of propodus; merus slightly less than twice length of carpus; ischium slightly more than ¥% length of merus. Uropods (Fig. 41) with exopods extend- ing well beyond telson; strong, moveable spine-seta between distolateral spine and blade; moveable spine distinctly longer than distolateral spine-seta. Endopods extending beyond telson, shorter than exopod. Description of male.—Adult usually smaller than adult female (Table 1), less ro- bust. Rostrum deep, relatively short, not ex- tending to cornea or beyond mid-region of basal peduncular article of antennule; short- er branch (fused and unfused parts) of an- tennule of lateral flagellum bearing 6 to 8 groups of aesthetascs; appendix masculina of second pleopod (Fig. 4h) armed with 1 to 3 weakly serrate apical spine-setae and 0 to 4 simple lateral spine-setae along inner margin. Similar to female in other respects. 45 Color pattern.—The overall appearance in life is iridescent green shading toward aquamarine; two transverse greenish-gold bands are often evident on large specimens (Fig. 1). Subtle markings vary and probably are not diagnostic. Carapace clear (trans- parent), speckled faintly with tiny gold and russet spots. Cornea gold, sometimes with an aquamarine ring around base. Eye-stalks with aquamarine spots and smaller spots of russet and ringed distally in deep aquama- rine. Antennules and antennular peduncle clear with russet and aquamarine spots; al- ternatively, antennules clear and tinged with aquamarine, containing spots of gold and russet; antennal peduncle clear with red, russet, and gold spots. Antennal scale, fla- gella, and rostrum clear; alternatively, an- tennal scale rimmed with gold. Antennae clear with tiny russet spots. Maxillipeds clear. Abdominal somites edged with faint gold spots, dorsal surfaces marked strongly by two transverse bands of greenish gold. The first band crosses the posterior section of abdominal somite 2; a second band tra- verses somite 3 directly across the hump. Both bands widen ventrolaterally, becom- ing narrowest on dorsal surface, sometimes with an underlying band of aquamarine; gold band sometimes contains russet spots; there is occasionally a fainter third band of gold and russet on posterior dorsal surface of abdominal somite 5; somites 4 and 5 of- ten are rimmed at dorsoposterior surface with aquamarine. Pereopods clear. Uropods clear with small gold spots distributed evenly; alternatively, telson and uropods ir- idescent aquamarine, pigmented most strongly near edges; base of telson some- times aquamarine tinged with greenish-gold and having a greenish-gold band across base. Pleopods with gold and russet spots. Protopodites with irregular longitudinal markings of aquamarine speckled with gold and russet but mainly clear; exopods and endopods clear. Early ova gold, later (eyed) ova pale aquamarine often tinged with gold. Habitat.—Associated with the slimy sea plume (Pseudopterogorgia americana) and 46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON other species of gorgonians along the fring- ing reefs of Pine Cay in depths from 14 to at least 43 m. Distribution.—Presently known only from the type locality off Pine Cay, Turks and Caicos Islands, British West Indies. Etymology.—Named for Jerry A. Mc- Lelland, in recognition of his contributions to the investigations of the marine inverte- brate fauna of the West Indies. Remarks.—Lebour (1949) described Per- iclimenes iridescens from an adult female netted off Bermuda. The “‘iridescens’’ com- plex comprises several species of close morphological affinity with P. iridescens (Heard & Spotte 1991, Heard et al. 1993, Spotte et al. 1994), and we include P. mclellandi in this complex. Periclimenes mclellandi is similar to P. patae, a species also collected on gorgonians at Pine Cay and the Florida Keys (Heard & Spotte 1991) and tentatively reported from the British Virgin Islands as Periclimenes cf. patae (see Spotte et al. 1995). Several characters of Periclimenes mclel- landi appear to be transitional between larg- er, less highly derived members of the “‘ir- idescens’’ complex (e.g., P. iridescens Le- bour, 1949; P. iridescens sensu Holthuis 1951; P. infraspinis (Rathbun, 1902); P. antipathophilus Spotte, Heard, & Bubucis, 1994), none of which are known to asso- ciate with gorgonians, and the apparently more highly derived P. patae. The affinity of P. mclellandi to the larger members of the “‘iridenscens’’ complex is indicated by the presence of a bilobate endite on the sec- ond maxillae and more than one row of comb-setae on the propodi of pereopods 5. Periclimenes mclellandi and P. patae are distinguished from other described mem- bers of the “‘iridescens’’ complex by: their smaller size, the chelae of pereopod 2 being equal or nearly equal; and having simple dactyls on pereopods 3-5. In these charac- teristics they resemble juveniles of the larg- er members of the complex, raising the pos- sibility that a neotenic condition has evolved in P. mclellandi and P. patae dur- ing their close association with gorgonians. Based on adult specimens, Periclimenes mclellandi differs from P. patae by its max- illa 2 having a bilobate endite; propodus of pereopods 5 with two rows of comb-setae, rostrum shorter, deeper, and generally with fewer rostral teeth; smaller adult body size (Table 1); smaller corneal diameter relative to eye-stalk; overall greenish coloration in life accentuated by two transverse, green- ish-gold bands on the first and second ab- dominal somites (Fig. 1); and occurrence at a greater mean depth. To highlight some of the size and meris- tic differences between Periclimenes mclel- landi and P. patae, we plotted the standard deviation of carapace length against total number of spines on the appendix mascu- lina (both apical and lateral) for males of the species (Fig. 5a) and graphed the stan- dard deviation of carapace length against the number of dorsal rostral teeth in the sexes of both species (Fig. 5 b, c). Corre- lations for the variables were significant for P. mclellandi in all comparisons (see Fig. 5). Mean carapace lengths for adults of both sexes are displayed in Table 1, indicating that adults of P. patae are larger than those of P. mclellandi. At Pine Cay, Periclimenes patae is found at a mean depth of 9.6 m (range 2.7—22.2 m); in contrast, P. mclellandi was never collected at depths shallower than 14 m, and its range extended to 43 m (Spotte & Bubucis 1996). We emphasize that 43 m marked our deepest descents. Pseudopter- ogorgia americana occurs even deeper off Pine Cay, and P. mclellandi might also. Associates and symbionts.—Other cari- dean shrimps, most commonly Hippolyte nicholsoni Chace, 1972 and Pseudocouti- erea antillensis Chace, 1972, co-occurred with P. mclellandi on Pseudopterogorgia americana and other gorgonians at Pine Cay. Co-inhabiting less frequently the same gorgonian with P. mclellandi were three other species, P. patae, Periclimenes sp. B, and Tozeuma sp. (Spotte & Bubucis 1996). VOLUME 110, NUMBER 1 Periclimenes mclellandi serves as the second intermediate host of an opaceolid digenean parasite. When microscopically examining living specimens of P. mclellan- di that are infected with opaceolid larvae, the oval metacercarial cysts can be easily observed in the shrimp host’s transparent abdominal muscles. An unknown gastropod mollusk serves as the first intermediate host. The adult stages of all known mem- bers of family Opecoelidae are intestinal parasites of fishes (Yamaguti 1971). At some collection sites, ordinarily between 18 and 26 m, up to 20% of the adult P. mclel- landi studied were infected with the meta- cercarial cysts of this larval digenean. A bopyrid isopod occasionally occurred underneath the abdomen of P. mclellandi. This parasite was found most often on shrimp collected in depths between 25 and 30 m. It appears to be an undescribed spe- cies of Hyperphrixus Niertrasz & Brender 4 Brandis, 1931, a genus known to parasit- ize other shrimps of the subfamily Ponto- niinae (see Markham 1985). Acknowledgments This research was sponsored by the Oak- leigh L. Thorne Foundation through a grant to Stephen Spotte. We thank members and staff of The Meridian Club and the Turks and Caicos Islands government for support and encouragement. Patricia M. Bubucis, Cherie S. Heard, Jerry A. McLelland, and Roy Manstan provided technical assistance. Patricia M. Bubucis, Sara LeCroy, and Jer- ry A. McLelland kindly reviewed earlier drafts of the manuscript. This is contribu- tion No. 289 of the Marine Sciences & Technology Center. Literature Cited Costa, O. G. 1844. Su due nuovi generi di Crostacei Decapodi Macrouri nota.—Annali della Acca- demia degli Aspiranti Naturalista, Napoli 2: 285—(?). Chace, E A., Jr. 1972. The shrimps of the Smithson- ian-Bredin Caribbean Expeditions with a sum- mary of the West Indian shallow-water species 47 (Crustacea: Decapoda: Natantia).—Smithsonian Contributions to Zoology 98:1—179. Gmelin, J. EF 1791. Caroli a Linné Systema Naturae per regna tria naturae, secundum classes, ordi- nes, genera, species, cum characteribus, differ- entiis, synomymis, locis. Editio decima tertia, aucta, reformata. Cura Jo. Frid. Gmelin, Lip- siae, Geog, Emanuel, Beer.—1(6):3021—4120, 3 pls. Heard, R. W. 1986. Pontoniine shrimps (Decapoda: Caridea: Palaemonidae) of the northwest Atlan- tic. I. The genus Neopontonides Holthuis, 1951, with the description of N. chacei, new species, and the erection of Pseudopontonides, new ge- nus, to receive N. principis Criales, 1980.— Journal of Crustacean Biology 6:471—484. , & S. Spotte. 1991. Pontoniine shrimps (De- capoda: Caridea: Palaemonidae) of the north- west Atlantic. II. Periclimenes patae, new spe- cies, a gorgonian associate from shallow reef areas off the Turks and Caicos Islands and Flor- ida Keys.—Proceedings of the Biological So- ciety of Washington 104:40—48. , S. Spotte, & P.M. Bubucis. 1993. Pontoniine shrimps (Decapoda: Caridea: Palaemonidae) of the northwest Atlantic. III. Neopericlimenes thornei, new genus, new species, from Pine Cay, Turks and Caicos Islands, British West In- dies.—Journal of Crustacean Biology 13:793- 800. Holthuis, L. B. 1951. A general revision of the Pa- laemonidae (Crustacea Decapoda Natantia) of the Americas. I. The subfamilies Euryrhynchi- nae and Pontoniinae.—Occasional Papers, Al- lan Hancock Foundation Publications No. 11, 332 pp. Lebour, M. V. 1949. Some new decapod Crustacea from Bermuda.—Proceedings of the Zoological Society of London 11(Part IV):1107—1117. 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 Hoyr- glass Cruises 7(3):1—156. Niertrasz, H. EF, & G. A. Brender 4 Brandis. 1931. Papers from Dr. Th. Morstensen’s Pacific Ex- pedition 1914-16. LVII. Epicaridea II. Vidensk Med Dansk naturhist Foren Khobenhavn 91: 147-226. Rathbun, M. J. 1902. Brachyura and Macrura. Papers from the Hopkins Stanford Galapagos Expedi- tion, 1898-1899. VIII.—Proceedings of the Washington Academy of Sciences 4:275-291. Spotte, S., & P M. Bubucis. 1996. Diversity and abundance of caridean shrimps associated with the slimy sea plume (Pseudopterogorgia amer- icana) at Pine Cay, Turks and Caicos Islands, 48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON British West Indies.—Marine Ecology Progress idae) of the northwest Atlantic. IV. Periclimenes Series 133:229-—232. antipathophilus new species, a black coral as- , P M. Bubucis, & R. M. Overstreet. 1995. sociate from the Turks and Caicos Islands and Caridean shrimps associated with the slimy sea Eastern Honduras.—Bulletin of Marine Science plume (Pseudopterogorgia americana) in mid- 95°212=227. summer at Guana Island, British Virgin Islands, Yamaguti, S. 1971. Synopsis of the digenetic trema- West Indies.—Journal of Crustacean Biology todes of vertebrates.—Tokyo, Keigaku Publish- 15:291—300. ing Company. Volume 1: 1074 pp; Volume 2: , R. W. Heard, & P. M. Bubucis. 1994. Pon- 349 plates. toniine shrimps (Decapoda: Caridea: Palaemon- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):49-57. 1997. New records and distribution ranges of shrimps (Crustacea: Decapoda: Penaeoidea and Caridea) in Chilean waters Ingo S. Wehrtmann and Alberto Carvacho (ISW) Instituto de Zoologia “Ernst F Kilian,’’ Universidad Austral de Chile, Casilla 567, Valdivia, Chile; present address: Alfred-Wegener-Institut fiir Polar- und Meeresforschung, Am Handelshafen 12, D-27570 Bremerhaven, Germany; (AC) SINERGOS, Santa Luisa de Marillac 1081, La Reina, Santiago, Chile; present address: Museo Nacional de Historia Natural, Casilla 787, Santiago, Chile Abstract.—Extensions of the geographical distribution are provided for eight species of shrimps. Ranges were mostly extended toward the south (Halipo- roides diomedeae, Pasiphaea magna, Betaeus emarginatus, Alpheus inca, Hip- polyte williamsi and Lysmata intermedia), while the boundary of the distri- bution of Eualus dozei was shifted to the north. Two species (Pasiphaea magna and Lysmata intermedia) are reported for the first time from Chilean waters. An additional species, Ogyrides sp., was collected from a stomach content of the fish Eleginops maclovinus in central-southern Chile, representing the first record of a representative of the family Ogyrididae from Chilean waters. This record is remarkable since species of Ogyrides are considered to inhabit warm- temperate and tropical waters. Chile has an extraordinary long, diver- sified coastline that lies in two distinct zoo- geographic regions (cold-/and warm-tem- perate region) connected by a transitional zone (Brattstr6m & Johanssen 1983). Sev- eral national and international expeditions (see Retamal 1981, Brattstrom & Johanssen 1983, Arntz et al. 1996, Wehrtmann & Lar- dies 1996) have carried out intensive col- lections along the Chilean coast, such as The Lund University Chile Expedition 1948—49 (LUCEB), of special importance to the knowledge of the Chilean decapod fau- na. Holthuis (1952), in his account of the ““‘macruran’’ decapods collected during LUCE, noted the relative small number of Decapoda Macrura known from Chile, which included a total of 37 species, 5 of which inhabit freshwater, 13 live in the deep sea or are pelagic, and 19 littoral. Sub- sequently, Retamal (1981) reported a total of 44 penaeoid and caridean shrimp species from Chilean waters, but excluded Hippol- yte williamsi Schmitt, 1924, previously re- ported by Zufiga et al. (1978). Recently, Retamal (1993), and Retamal & Soto (1995), examined samples collected from deep waters off northern Chile, and report- ed four additional species from Chile: Ben- thesicymus tanneri (Faxon, 1893), Sergia Phorca (Faxon, 1893), Psathyrocaris fra- gilis Wood-Mason, 1893 and Plesionika santaecatalinae Wicksten, 1983a. This study updates and summarizes available information concerning species composition and distribution of the pen- aeoid and caridean shrimp fauna of Chile. We include results from our samplings car- ried out during the last years as well as from examination of collections of Museo Nacional de Historia Natural (MNHNC), Santiago, Pontificia Universidad Catélica de Chile (“‘Sala de Sistematica’’: SS-UC), Santiago, and National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C. (USNM). The specimens that we collected are deposited in the collections of ‘“‘Sinergos”’ (SIN), and the Institute of Zo- 50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ology (IZUA), Universidad Austral de Chile, Valdivia. Carapace length (CL) was measured from the posterior edge of the or- bital arch to the mid-dorsal margin of the carapace. Total length (TL) refers to the dis- tance between the posterior margin of the orbital arch and the distal margin of the tel- son, excluding setae. Infraorder Penaeidea Rafinesque, 1815 Family Solenoceridae Wood-Mason, 1891 Haliporoides diomedeae (Faxon, 1893) Previous known distribution——Panama (06°30’N) to central Chile (37°40'S); from 240 to 1886 m (Faxon 1895, Del Solar & Mistakidis 1971, Del Solar & Flores 1972, Noziglia & Arana 1976, Pérez-Farfante 1977, Holthuis 1980, Méndez 1981, Reta- mal R 1993). Material examined.—1 female (CL 41.0 mm), Gulf of Corcovado, off Morro Yeli (43°32'S), bottom trawl, 50-95 m, May 1996, MNHNC 660025. Remarks.—The specimen collected ex- tends considerably (approximately 660 km) to the south the known geographical distri- bution of this species. The depth at which the specimen was collected is the shallow- est recorded for this species. Infraorder Caridea Dana, 1852 Family Pasiphaeidae Dana, 1852 Pasiphaea magna Faxon, 1893 Previous known distribution.—Califor- nia, U.S.A. (off Point Arena to off Ponit Loma) to Pert (17°05'S); from 509 to 1019 m (Faxon 1893, Schmitt 1921, Méndez 1981). Material examined.—1 ovig. female (CL 47.9 mm, TL 113.0 mm), off northern Chile (22°13’S, 70°23'W), 815 m, Feb 1981, MNHNC 81085. Remarks.—This is the first record of this species from Chile, and extends its distri- bution to the south by approximately 560 km. Family Oplophoridae Dana, 1852 Acanthephyra pelagica (Risso, 1816) Previous known distribution.—Mediter- ranean, North and South Atlantic, Indo Pa- cific, and, perhaps Panantarctic (Zariquiey 1968, Chace 1986, Tiefenbacher 1994), Ar- gentina between 38°S and 54°S (Boschi et al. 1992), South Africa (without indication of latitude) (Kensley 1981), New Zealand (without indication of latitude) (Richardson & Yaldwyn 1958). Material examined.—1 male (CL 15 mm), Valle de La Mocha (38°04’'S, 73°52'W), 400-548 m, Jun 1965, MNHNC 660011. Remarks.—The systematic position of the species seems to be unclear (Chace 1986). The specimen studied by us coin- cides perfectly with Boschi et al.’s (1992: 25) description and illustration of A. pelag- ica from southern Argentina. To our knowl- edge, this is the first report of this species from central-southern Chile and, therefore, the most northern record for the Pacific Ocean. Vinuesa (1977:14, table 1) reported A. pelagica occurring between Magellan Strait and Chiloé Island (approximately 42°S), but he did not indicate where he ob- tained information on the distribution of this species. Superfamily Alpheoidea Rafinesque, 1815 Family Alpheidae Rafinesque, 1815 Betaeus emarginatus (H. Milne Edwards, 1837) Previous known distribution.—Pert (be- tween 06°55'40"S and 14°14’S) and Chile (between 20°16’S and 41°51’'S) (Holthuis 1952, Wicksten & Méndez 1983). Material examined.—2 males (CL 9.84, 15.96 mm; TL 29.26, 47.61 mm) and 1 fe- male (CL 10.10 mm; TL 30.05 mm), west coast of Chiloé Island, Quiutil (42°30’S), Apr 1993, SIN CR(97)023. Remarks.—This record extends the range of this species slightly to the south by ap- proximately 90 km, and now includes Chi- loé Island. VOLUME 110, NUMBER 1 Alpheus inca Wicksten & Méndez, 1981 Previous known distribution.—Galapa- gos Islands (Wicksten 1991), Peri (between 06°55’S and 13°50’S) and Chile (22°08’S) (Wicksten & Méndez 1981). Material examined.—1 female (CL 4.66 mm; TL 14.35 mm), Tocopilla (22°S), Jan 1965, SS-UC 2822; 1 male (CL 5.89 mm; TL 17.48 mm) and 1 female (CL 5.32 mm; TL 18.34 mm), Peninsula of Meyjillones (23°S), Jan 1965, SS-UC 2759; 2 females (CL 4.73, 6.46 mm; TL 15.66, 21.00 mm), Mejillones Bay (23°S), Jan 1965, SS-UC 2823 and 2802; 4 males (CL 8.97-—17.02 mm; TL 25.84—42.28 mm) and 3 females (CL 6.23-8.97 mm; TL 19.61—24.93 mm), Totoralillo-Panul (30°05’S), Jan 1963, SS-UC 764; 1 male (CL 10.64 mm; TL 34.35 mm) and 2 ovig. females (CL 14.29 and 16.26 mm; TL 44.23 and 49.25 mm), Los Molles (32°14'S), Jan 1963, SS-UC 451; 1 ovig. female (CL 16.72 mm; TL 50.77 mm), Los Molles (32°14’S), Jan 1963, SS-UC 452; 1 male (CL 9.42 mm; TL 34.05 mm), El Tabo (33°28’S), Feb 1960, SS-UC 33; 1 female (CL 5.99 mm; TL 17.67 mm), Las Cruces (33°35’S), Oct 1966, SS-UC 3769; 1 male (CL 4.94 mm; TL 17.58 mm), Duao (35°S), Jan 1963, SS-UC 615. Remarks.—The present report extends the southern limit of this species from Me- jillones (22°08’S) to Duao (35°S), a dis- tance of approximately 1430 km. Family Hippolytidae Dana, 1852 Hippolyte williamsi Schmitt, 1924 Previous known distribution—México, Gulf of California to Chile (between 20°14'30"S, 70°10'50”W to Isla Santa Maria (23°24'30"S, 70°36'10"W) (Zufiga et al. 1978, Wicksten & Hendrickx 1992). Material examined.—7 females (CL 3.7, 3.4, 2.5, 3.2, 3.7, 3.6 and 2.7 mm; TL 22.2, 17.2, 12.3, 16.3, 20.0, 19.3 and 13.9 mm, respectively) and 9 males (CL 2.0, 2.1, 2.1, 2.4, 2.9, 3.5, 3.2, 3.0 and 3.1 mm; TL 10.2, 10.9, 10.6, 13.3, 14.7, 17.1, 15.9, 14.9 and a 14.4 mm, respectively) associated with Ulva sp., Bahia de Coquimbo (29°50’S, 71°15'W), 2 m, 15 Mar 1995, IZUA 540; 1 female (CL 4.3 mm; TL 24.0 mm) and 3 males (CL 2.8, 2.4 and 2.8 mm; TL 16.3, 13.7 and 15.6 mm, respectively) associated with Ulva sp., Calbuco (51°50'S, 73°05’W), 2m, 5 Nov 1994, IZUA 541; 3 females (CL 6.23, 6.00 and 5.62 mm; TL 36.01, 35.11 and 32.83 mm, respectively) and 2 males (CL 4.56 and 4.03 mm; TL 25.80 and 21.81 mm, respectively), Lechagua, near Ancud, Chiloé Island (41°52’S, 73°52'W), 6 Feb 1958, MNHNC 940002; 3 females (CL 6.23, 4.15 and 6.08 mm; TL 34.50, 35.12 and 34.35 mm, respectively), Lechagua, near Ancud, Chiloé Island (41°52’S, 73°52'W), 6 Feb 1958, MNHNC 940005; 4 males (CL 4.26, 4.56 and 4.86 mm; TL 25.84, 24.48 and 26.76 mm, respectively; 1 individual incomplete), Lechagua, near An- cud, Chiloé Island (41°52’S, 73°52'W), 6 Feb 1958, MNHNC 940008; 1 ovig. female (CL 7.3 mm; TL 38.0 mm) associated with mussel raft culture, Putemtin, Chiloé Island (42°25'S, 73°43'W), approximately 5 m, 12 May 1994, IZUA 542. Remarks.—The present material extends considerably the southern limit of this spe- cies from northern (Isla Santa Maria, 23°S) to southern Chile (Isla Chiloé, 42°S), a dis- tance of approximately 2030 km. Thus, the range of this species now covers the area between the Gulf of California and southern Chile (see Wicksten 1990). Hippolyte wil- liamsi has been confused with H. califor- niensis in the Gulf of California, but the two can be separated by the number of spines on the distal end of the first anten- nular segment (Wicksten 1983b). H. cali- forniensis has one or rarely two spines, whereas H. williamsi has three. Zufiga et al. (1978) pointed out some differences be- tween the original description (Schmitt 1924) and the specimens collected in north- ern Chile. Additionally, we have observed the following variations: (1) The first seg- ment of antennular peduncle bears 3—4 and (2) the merus of the fourth pereiopod has 2 52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON or 3 spines. Our measurements indicate a latitudinal cline in adult size (TL, CL), with considerably larger individuals in Chiloé Is- land (largest female from Putemutn: 38.0 mm) compared to specimens from northern Chile (Coquimbo: mean female TL of 17.3 mm). A similar tendency has been reported by Boschi et al. (1969) for the brachyuran crab Halicarcinus planatus (Fabricius) from the Argentinean coast. Eualus dozei (A. Milne-Edwards, 1891) Previous known distribution.—Southern Chile (between 55°36’S, 67°40’W and 42°20'S, 72°22'W), from 8 to 300 m (A. Milne-Edwards 1891, Holthuis 1952, Ara- cena & Lépez 1973). Material examined.—1 ovig. female col- lected from sand and shell, Galapagos Is- lands (Tagus Cove, Isla Isabela), 15 Jan 1934, 18-33 m, Velero III station 157-34; collection of the USNM 276061; 4 females (CL 2.2, 2.2, 2.4 and 2.0 mm; TL 12.3, 12.1, 12.1 and 12.9 mm, respectively) as- sociated with suspended scallop culture, Bahia de Guanaqueros (30°08’S, 71°25’W), approximately 5 m, 13 Nov 1993, IZUA 538; 2 females (CL 2.8 and 2.6 mm; TL 12.9 and 12.4 mm, respectively) associated with suspended scallop culture, Bahia de Tongoy (30°13’S, 71°32’W), approximately 5 m, 27 Jul 1992, IZUA 539. Remarks.—Our material represents the northernmost record of this species, and the first from Galapagos Islands. So far, the species has not been reported from Perdé (see Méndez 1981) or Argentina (Boschi et al. 1992). Zarenkov (1970) reported a sin- gle specimen from 53°S 52’'S-64°10’E, but was unable to determine it with certainty due to the poor condition. It is of interest to note that our material as well as those of E. dozei collected during the LUCE (see Holthuis 1952) and the Soviet Antarctic Ex- pedition (see Zarenkov 1970), are all fe- males. As indicated by our material, this Species occurs in relatively deep and shal- low waters (approximately 5 to 300 m). Lysmata intermedia (Kingsley, 1878) Previous known distribution.—Florida Keys to Tobago and Curagao; Azores; Ga- lapagos Islands (Wicksten 1990); eastern Pacific: from México to Peri (03°30’S— 03°44'S) (Méndez 1981, Wicksten & Mén- dez 1983, Hendrickx & Wicksten 1987); up to 36 m (Méndez 1981). Material examined.—1 male collected from suspended scallop cultures (CL 8.2 mm, TL 33.8 mm), Bahia de Guanaqueros (30°08’S; 71°25'W), approximately 8 m, 31 Jul 1993, IZUA 537. Remarks.—This is the first report of a species of Lysmata from Chile. The speci- men collected extends the known geograph- ical distribution of the species by almost 27° of latitude (approximately 2960 km) to the south. Monthly sampling in the Bahia de Guanaqueros did not produce additional specimens of this species. Genus Ogyrides Stebbing, 1914 Ogyrides sp. Material examined.—1 female (CL 5.8 mm, TL 21.1 mm) obtained from a stomach content of Eleginops maclovinus (Pisces, Nototheniidae), Mehuin, estuary of river Lingue (39°25’S, 73°13'W), Feb 1991, IZUA 543. Remarks.—Species of Ogyrides are known from warm-temperate and tropical waters (Wicksten & Méndez 1988); how- ever, our sampling site (Mehuin) lies in the cold-temperature region (see Brattstro6m & Johanssen 1983). The specimen collected is a noteworthy record, representing the first account of a representative of the family Ogyrididae from Chile. Two species of Ogyrides have been reported from the east- ern Pacific Ocean (Carvacho & Olson 1984, Wicksten & Méndez 1988): O. alphaeros- tris (Kingsley 1880) and O. tarazonai Wicksten & Méndez, 1988. Due to the in- complete condition of our specimen, we were unable to identify it to the species lev- el. Our specimen clearly has more than one Spine on the dorsal midline of the carapace VOLUME 110, NUMBER 1 Pasiphaea magna Lysmata intermedia Alpheus inca Alpheopsis chilensis Rhynchocinetes typus Lysmata porteri Latreutes antiborealis Hippolyte williamsi Alpheus chilensis Betaeus emarginatus Haliporoides diomedeae Synalpheus spinifrons Argentina and Falkland Islands gentina Argentina; Brazil Acanthephyra carinata Pasiphaea dofleini Argentina circumpolar 200 m) species are excluded. The distributions of the species considered are summarized in Fig. 1. Of the species reported from Chile, a rel- atively small group occur in the subantarc- tic or cold-temperate regions; these are Acanthephyra carinata Bate 1888, Pasi- phaea dofleini Schmitt 1932, Campylonotus semistriatus Bate 1888, C. vagans Bate 1888 and Chorismus antarcticus (Pfeffer 1887). Members of this group generally in- habit waters around 50°S and 55°S of lati- tude, although the range of one species (A. carinata) extends its northernmost distri- bution up to Chiloé Island (42°S). Another group of species, Pasiphaea acutifrons Bate 1888, Nauticaris magellan- ica (A. Milne Edwards 1891), Betaeus trun- catus Dana 1852 and Austropandalus grayi (Cunningham 1871), seems to be fairly eu- rythermic with distributions that include warm-temperate as well as cold-temperate 54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON regions. The most distinctive feature of this group is the capacity of its members to cross the Chiloé boundary (see Brattstr6m & Johanssen 1983). The majority of the species considered show clear boreal affinities, with their northern distribution limit [except Lysmata porteri (Rathbun 1907)] extending to the Peruvian Province. Chiloé Island is the southern limit of the distribution of five species, which confirms that the area around 42°S serves as a separation zone be- tween the cold-/and warm-temperate regions described by Brattstr6m & Johans- sen (1983) for the Chilean coast. The south- ern distribution limit of the remaining spe- cies of this group is between 30° (Coquim- bo) and 42°S (Chiloé Island), which ac- cording to Brattstrom & Johanssen (1983) corresponds to a transitional area between the Magellanic and Peruvian Provinces. As is typical for transitional zones, this area does not contain an endemic fauna, al- though one species (Lysmata porteri) is found exclusively in this area. The bound- ary between this transitional area and the Peruvian Province is not clear, but probably is located around Coquimbo (30°S). Addi- tional information on the shrimp fauna of northern Chile is needed to properly eval- uate the various zoogeographic boundaries. The influence of the Peruvian Province on the caridean fauna of Chile becomes ap- parent from Coquimbo (30°S) on north- ward. A distributional boundary apparently is located around Iquique (20°S), where the northern limit of four eurythermic species (Pasiphaea acutifrons, Nauticaris magel- lanica, Betaeus truncatus and Austropan- dalus grayi) is located. However, this ob- servation could be attributed to the much more intense sampling programs in this area (Holthuis 1952, Retamal & Soto 1993). The northern limit of carideans from the Peruvian Province seems to be located around the equator. The several distribution patterns of cari- deans from Chile are influenced by the pre- vailing current system (see Brattstro6m & Johanssen 1983, and references cited there- in). The Chiloé boundary corresponds to the area where the Westwind Drift reaches the continent and splits into the Peri Coast- al Current (northward) and the Cape Horn Current (southward). The Peri Coastal Cur- rent gradually joins the South Equatorial Current, which may explain, in part, the presence of some species [Alpheus chilensis Coutiére (in Lenz 1902), Hippolyte william- si, Eualus dozei, Lysmata intermedia and Glyphocrangon loricata Faxon 1895] that occur both in Chile and the Galapagos Is- lands (see Chavez & Brusca 1991, Wick- sten 1991). Acknowledgments Financial support was provided to I. S. Wehrtmann by the “Deutscher Akadem- ischer Austauschdienst’? (DAAD), “‘Ge- sellschaft fiir Technische Zusammenarbeit”’ (GTZ) and Universidad Austral de Chile (Project DID N° E-91-1 and S-94-53), and to A. Carvacho by NOVIB-Netherlands, as part of the project ““Medio Ambiente en la Xa Region de Chile,’”” SINERGOS. Our thanks are due to all institutions who made their collections available for us. Nibaldo Bahamonde, Lipke B. Holthuis, Carlos Jara, and Mary K. Wicksten revised the manu- Script, and their suggestions improved con- siderably the quality of the manuscript. Ingo S. Wehrtmann is greatly indebted to Mary K. Wicksten for providing specimens of Hippolyte williamsi to verify the identi- fication of the Chilean material, and who also allowed us to include the data con- cerning Eualus dozei from the Galapagos Islands. Literature Cited Aracena P.,, O., & M. T. Lépez. 1973. Observaciones biol6gicas en organismos encontrados en sub- Stratos artificiales. Caleta Leandro, Talcahuano, Chile. I. Crustacea Decapoda Macrura.—Tra- bajo de V Congreso Latinoamericano de Zool- ogia 1:40—48. Arntz, W. E., M. Gorny, I. S. Wehrtmann, M. Lardies, & R. Soto. 1996. Crustacea Decapoda: sum- VOLUME 110, NUMBER 1 mary report. Pp. 57-59 in W. Arntz & M. Gor- ny, eds., Cruise Report of the Joint Chilean- German-Italian Magellan ‘“‘Victor Hensen”’ Campaign 1994. Berichte zur Polarforschung 190. Bate, C. S. 1888. Report on the Crustacea Macrura collected by H. M. S. Challenger during the years 1873—76.—Report on the Scientific Re- sults of the Voyage of H. M. S. “Challenger” during the Years 1873-76, 24:1—942. Boschi, E. E., C. E. Fischbach, & M. I. Iorio. 1992. Catalogo ilustrado de los crustaceos estomat6- podos y decapodos marinos de Argentina.— Frente Maritimo, Montevideo 10:7—94. , M. A. Scelzo, & B. Goldstein. 1969. Desar- rollo larval del cangrejo, Halicarcinus planatus (Fabricius) (Crustacea, Decapoda, Hymenosom- idae), en el laboratorio, con observaciones sobre la distribucion de la especies.—Bulletin of Ma- rine Science 19:225—242. Brattstrom, H., & A. Johanssen. 1983. Ecological and regional zoogeography of the marine benthic fauna of Chile.—Sarsia 68:289-—339. Carvacho, A., & R. Olson. 1984. Nuevos registros para la fauna carcinoldgica del noreste de Méx- ico y descripcién de una nueva especia: Eualus subtilis n.sp. (Crustacea: Decapoda: Natan- tia).—The Southwestern Naturalist 29:59-71. Chace, E A. 1986. The Caridean Shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedi- tion, 1907-1910, Part 4: Families Oplophoridae and Nematocarcinidae.—Smithsonian Contri- bution to Zoology 432:1—182. Chavez, FE P, & R. C. Brusca. 1991. The Galapagos Islands and their relation to oceanographic pro- cesses in the Tropical Pacific. Pp. 9-33 in M. J. James, ed., Galapagos marine invertebrates. Plenum Publishing Cooperation, New York, 474 pp. Cunningham, R. O. 1871. Notes on the Reptiles, Am- phibia, Fishes, Mollusca, and Crustacea ob- tained during the voyage of H. M. S. “‘Nassau”’ in the years 1866—69.—Transactions of the Lin- nean Society of London 27:465—502. Dahl, E., 1960. The cold temperate zone in Chilean seas.—Proceedings of the Royal Society, B 152:63 1-633. Dana, J. D. 1852. Conspectus Crustaceorum quae in Orbis Terrarum Circumnavigatione, Carolo Wilkes e Classe Reipublicae Foederatae Duce, lexit et descripsit—Proceedings of the Acade- my of Natural Sciences of Philadelphia 1852: 10-28. Del Solar, E., & M. Mistakidis. 1971. Informe del crucero SNP-1 7105. Exploracién de Crusta- ceos.—Instituto del Mar del Pert, Serie de In- formes Especiales IM-89:1-—10. , & L.A. Flores. 1972. Exploraciones de Crus- 35 taceos (Zona Sur), Crucero SNP-1 7201.—In- stituto del Mar del Pert, Serie de Informes Es- peciales IM-107:1-8. Fabricius, J. C. 1798. Supplementum Entomologiae Systematicae. Hafniae, Copenhagen, 572 pp. Faxon, W. 1893. Reports on the dredging operations off the west coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commis- sion Steamer “Albatross” during 1891, Lieut. Commander Z. L. 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Brewster, ed., Edinburgh Encyclopedia, Volume 7. 56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lenz, H. 1902. Die Crustaceen der Sammlung Plate (Decapoda und Stomatopoda).—Fauna chilen- sis 2(3). Zoologische Jahrbiicher, supplement 5: 731-772. Méndez, G. M. 1981. Claves de identificacion y dis- tribucién de los langostinos y camarones (Crus- tacea: Decapoda) del Mar y rios de la costa del Peri.—Boletin del Instituto del Mar de Pert- Callao 5:1—170. Milne-Edwards, A. 1881. Description de quelques Crustacés Macroures provenant des grandes profondeurs de la mer des Antilles—Annales Sciences Naturelles, Paris, Zool., ser. 6, vol. 11: 1-16. 1891. Crustacés.—Mission scientifique du Cap Horn. 1882-1883, 6, Zool., 1—54.. Milne Edwards, H. 1837. Histoire Naturelle des Crus- tacés, comprenant |’ anatomie, la physiologie et la classification de ces animaux. Volume 2, Ror- et, Paris, 532 pp. Noziglia, L., & P Arana. 1976. Observaciones biol- 6gico-pesqueras de la gamba, Hymenopenaeus diomedeae Faxon, 1893, en el Pacifico sur ori- ental.—Revista de la Comisi6n Permanente del Pacifico Sur 5:187—195. Farfante, I. 1977. American solenocerid shrimps of the genera Hymenopenaeus, Hali- poroides, Pleoticus, Hadropenaeus new species and Mesopenaeus new genus.—Fishery Bulle- tin 75:261—346. Pfeffer, G. 1887. Die Krebse von Stid-Georgien nach der Ausbeute der Deutschen Station 1882-— 83.—1. Teil. Jahrbuch der Hamburger wissen- schaftlichen Anstalt 4:41—150. Rafinesque, C. S. 1815. Precis des decouvertes et tra- vaux somiologiques. Rouale Typographie Mili- tarie, Palerme, 55 pp. Rathbun, M. J. 1907. South American Crustacea.— Revista Chilena de Historia Natural 11:45—50. Retamal, M. A. 1981. Catalogo ilustrado de los crus- taceos decapodos de Chile.—Gayana Zoologia 44:7-110. , M. A. 1993. Crustaceos decapodos abisales de la zona Iquique-Arica.—Estudios Oceanol- Ogicos 12:1-8. Richardson, L. R., & J. C. 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A new species of the genus Petrolisthes Stimpson, 1858 (Crustacea: Decapoda: Anomura: Porcellanidae) from Yonaguni Island, the Ryukyu Islands Masayuki Osawa Department of Aquatic Biosciences, Tokyo University of Fisheries, 4-5-7, Konan, Minato-Ku, Tokyo 108, Japan Abstract.—A new species of the genus Petrolisthes Stimpson, 1858, P. don- anensis, is described and illustrated on the basis of two specimens collected from Yonaguni Island, the Ryukyu Islands. This species closely resembles P. decacanthus Ortmann, 1897, but is distinguishable from the latter species by the following characters: sinuously triangular rostrum; absence of hepatic spines; antennular peduncles with basal segment unarmed on anterior margin except for a strong spine on lateral corner; chelipeds with palm bearing a small spine at subdistal corner of dorsoflexor margin; and ambulatory legs with slen- derer propodi, and dactyli bearing four flexor spines. The porcellanid crab genus Petrolisthes Stimpson, 1858, includes numerous inter- tidal or subtidal species. Eighteen species have been reported from the Ryukyu Is- lands, southwestern Japan (Miyake 1982, Kropp 1986, Osawa 1996), some of which are conspicuous on the intertidal boulder beaches of the islands (Asakura 1991). Through the courtesy of Mr. K. Nomura of the Kushimoto Marine Park and Dr. T. Komai of the Natural History Museum and Institute, Chiba, I had the opportunity to ex- amine two porcellanid crabs collected from the intertidal zone of Yonaguni Island (24°27'N, 122°56’E), the westernmost of the Ryukyus chain. The examination re- vealed that the two specimens belong to an undescribed species of the genus Petrolis- thes. This new species is herein described and illustrated. The carapace length (CL) and width (CW), as indications of specimen size, were measured, CL from the anterior tip of the rostrum to the median posterior end of the carapace, and CW at the broadest part. Measurements of ambulatory legs were made as follows: lengths of carpus and propodus along extensor margin, and height of propodus at midline. The type specimens are deposited in the Natural History Mu- seum and Institute, Chiba (CBM). Petrolisthes donanensis, new species Figs. 1, 2 Type material.—Holotype: ovig. female (CL 8.6 mm, CW 8.9 mm), Yonaguni Is- land, Ryukyu Islands, intertidal, 15 Apr 1995; CBM-ZC 2147. Paratype: 1 female (CL 5.9 mm, CW 5.9 mm), same data as holotype; CBM-ZC 2148. Description.—Carapace (Fig. 1A, B) weakly convex dorsally, slightly broader than long or as long as broad, broadest on posterior branchial regions. Branchial mar- gins strongly convex, with longitudinal tu- berculated edges, posterior branchial mar- gins with strong transverse rugae. Rostrum (Fig. 1C) moderately broad, slightly bent ventrally, sinuously triangular; median lobe strongly produced; dorsal surface with nu- merous small, flattened tubercles. Protogas- tric ridges distinct, divided into 2 lobes by median groove extending to tip of rostrum. Orbits (Fig. 1G) deep; orbital margin armed VOLUME 110, NUMBER 1 59 YA ag git ENON Fig. 1. Petrolisthes donanensis, new species. Holotype, ovigerous female (CL 8.6 mm, CW 8.9 mm; CBM-ZC 2147). A, carapace and proximal somites of abdomen, dorsal; B, carapace and pterygostomian flap, lateral; C, rostrum, frontal; D, thoracic sternites, ventral; E, telson, exposed; F left basal segment of antennular peduncle, ventral; G, left eye and antennal peduncle, dorsal; H, left antennal peduncle, ventral. Scales equal 1.0 mm. 60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ZNO > Fig. 2. Petrolisthes donanensis, new species. Holotype, ovigerous female (CL 8.6 mm, CW 8.9 mm; CBM-ZC 2147). A, left third maxilliped, long setae omitted on flexor margin, ventral; B, right cheliped, dorsal; C, same, ventral; D, left first ambulatory leg, lateral; E, same, distal part of merus, mesial; FE left second ambulatory leg, lateral; G, left third ambulatory leg, lateral. Scales equal 1.0 mm. VOLUME 110, NUMBER 1 with well developed supraocular spine; out- er orbital angle produced into strong, acute spine. Gastric region with numerous flat- tened tubercles and short, transverse rugae; metagastric region with rugae longer and more distinct. Hepatic margins tuberculate, lacking spines. Cervical grooves moderate- ly marked. Two epibranchial spines well developed on each side. Anterior branchial regions with weakly developed tubercles. Posterior branchial and cardiac regions cov- ered with short plumose or simple setae. Rugae and flattened tubercles on carapace fringed with short plumose or simple setae anteriorly. Pterygostomian flaps (Fig. 1B) provided with longitudinal ridges, anterodorsal mar- gin fringed with small tubercules. Third thoracic sternite (Fig. 1D) strongly depressed, trilobate anteriorly; median lobe not exceeding laterals, with broadly round- ed apex. Fourth sternite provided with se- ries of short rugae along concave, anterior Margin. Telson (Fig. 1E) as illustrated. Ocular peduncles (Fig. 1G) moderately large, short, with several short striae on dor- sal surface; dorsal extension onto cornea rounded; anterodistal margin fringed with short plumose setae. Basal segment of antennular peduncles (Fig. 1F) transversely rugose on anterior re- gion of ventral surface; anterior margin weakly produced, tuberculate, with strong, acute spine laterally. Antennal peduncles (Fig. 1G, H) 4-seg- mented; first segment immovable, follow- ing segments movable. First segment not strongly produced forward in lateral view, with small projection anterodistally. Second segment armed anteriorly with triangular crest bearing small tuberculate spine at proximal end, dorsal and ventral surfaces relatively smooth. Third segment rectan- gular, anterior margin minutely crenulate, dorsal and ventral surfaces with oblique ru- gae. Fourth segment smooth. Third maxillipeds (Fig. 2A) robust. Is- chium broad, ovate, transversely rugose on 61 ventral surface. Merus provided with lami- nate and subrectangular lobe with small tu- berculate spine on ventroflexor margin, transversely rugose on ventral surface. Car- pus with distinct projection on subproximal region of flexor margin, and longitudinal rugae on ventral surface, one of those rugae along ventroextensor margin forming ridge. Propodus with scattered short setae on ven- tral surface, and longitudinal striae along extensor margin. Dacty] subtriangular, ven- tral surface smooth. Exopod laminate, slen- der, inflated proximally, with distal flagel- lum. Rugae on ventral surface of ischium to carpus fringed with short plumose setae on anterior sides. Chelipeds (Fig. 2B, C) subequal (left cheliped missing in holotype). Ischium armed with strong, acute spine near distal end of ventroflexor margin. Merus short; dorsal surface transversely rugose, with dis- tinct transverse ridge submedially, and 1 median spine near extensor margin; dorso- flexor margin provided with crenulate lobe with 1 or 2 small spines at distal end; dor- sodistal margin armed with 2 spines; ven- tral surface rugose, distoflexor margin with 2 spines. Carpus (excluding flexor marginal teeth) 2.3 times as long as broad; dorsal sur- face with numerous small, flattened or weakly developed tubercles, median, lon- gitudinal, weak ridge (composed of series of squamae in paratype) and shallow sulcus along extensor margin; dorsoflexor margin armed with 4 strong acute teeth serrated or crenulate along edges; dorsoextensor mar- gin provided with 8—10 spines along entire length, distal spine with double-pointed api- ces; ventral surface transversely rugose, flexor margin crenulate. Palm with extensor margin thin, weakly arched and serrated; dorsal surface with distinct, median, longi- tudinal ridge extending from proximal end of palm to base of dactyl, and composed of series of flattened, squamose tubercles; sur- face between flexor margin and longitudinal ridge with numerous small, flattened tuber- cles; surface between extensor margin and longitudinal ridge with numerous small, 62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON flattened or weakly to moderately devel- oped tubercles; dorsoflexor margin with longitudinal rugose ridge and 1 small but distinct spine at subdistal corner; ventral surface with numerous short, oblique rugae. Fixed finger with numerous small, weakly developed tubercles on dorsal surface; ven- tral surface rugose as in palm. Dactyl with dorsal surface armed with longitudinal ridge of flattened and imbricated tubercles along midline, and rugose ridge along flex- or margin; ventral surface with small, flat- tened tubercles. Rugae and tubercles on dorsal surface of merus to dactyl fringed with short plumose setae arising from distal sides. Ambulatory legs (Fig. 2D—G) relatively slender. Merus elongate, narrowing distally; extensor margin serrate, number of spines 5-7 in first, 7—9 in second, and 5-7 in third legs, and furnished with short, plumose or simple setae; lateral surface with numerous short, transverse rugae; distoflexor margin armed with 1 acute spine (lacking on third leg of paratype); flexor margin on mesial surface provided with 1 strong spine sub- distally. Carpus relatively long, with short, longitudinal rugae furnished with simple, short and long setae on lateral surface; sub- distoextensor angles of first and second legs armed with 1 spine (lacking on second leg of paratype), unarmed on third leg. Propo- dus slender, long, approximately 1.4 times as long as carpus and 6.6—7.1 times as long as high; lateral surface and extensor margin with scattering, simple, short and long se- tae; flexor margin armed with 2 or 3 mov- able spines in addition to distal pair. Dactyl terminating in slightly curved claw, flexor margin with 4 movable spines. Color (preserved in ethanol).—Carapace, chelipeds, and ambulatory legs dark or light reddish except for following parts of am- bulatory legs: distal part of merus, proximal and distal parts of carpus and propodus (whitish), and dactyl (whitish but dark brown in distal claw). Distribution.—So far known only from the type locality, Yonaguni Island, the west- ernmost of the Ryukyus; intertidal. Etymology.—This specific name is de- rived from “‘Donan,’”’ which means Yona- guni Island in the Ryukyu dialect. Remarks.—Haig & Kropp (1988) de- scribed Petrolisthes eldredgei, and rede- scribed two poorly known species, P. de- cacanthus Ortmann, 1897 and P. bispinosus Borradaile, 1900, all from the Indo-West Pacific. They mentioned that the three spe- cies stand out as a group quite distinct from other Indo-West Pacific congeners by the sharing of the following characters: trans- versely rugose carapace; presence of two pairs of the epibranchial spines; unarmed mesobranchial margins; and merus of the first ambulatory leg with a strong subdistal spine on the mesial flexor margin. These four characteristics are also found in the present new species, therefore, P. donanen- sis can be considered the fourth member of this group. Petrolisthes donanensis most closely re- sembles P. decacanthus. These two species share the following characters: orbital an- gles produced into a strong spine; presence of a pair of supraocular spines; and cheli- peds with carpus and palm covered with nu- merous imbricated tubercles and granules on the dorsal surface. The second feature also distinguishes the two species from P. bispinosus and P. eldredgei. In addition to these characters, the spine on the ventro- flexor margin of the ischium of the cheli- peds found in P. donanensis, which has not been described previously for any species of Petrolisthes, was also confirmed in spec- imens of P. decacanthus from Guam (R. K. Kropp, pers. comm.), and in the figure of this species by Haig & Kropp (1988: fig. 3e), although the spine was rather weak. Petrolisthes donanensis, however, differs from P. decacanthus in: the sinuously tri- angular rostrum (trilobate in P. decacan- thus); absence of hepatic spines; antennular peduncles with basal segment unarmed on the anterior margin except for a strong spine at the lateral corner (with several VOLUME 110, NUMBER 1 strong spines and denticles in P. decacan- thus); chelipeds with palm bearing a small spine at the subdistal corner of the dorso- flexor margin; and ambulatory legs with slenderer propodi and dactyli bearing four flexor spines (three spines in P. decacan- thus). The presence of a pair of supraocular spines and two pairs of epibranchial spines in P. donanensis and P. decacanthus, also links these to several Indo-West Pacific spe- cies such as P. scabriculus (Dana, 1852), P. militaris (Heller, 1862), P. perdecorus Haig, 1981, and P. heterochrous Kropp, 1986. Based on the examination of material of P. militaris from the Ryukyus and In- donesia, the species has been found to pos- sess a strong subdistal spine on the mesial flexor margin of the merus of the first am- bulatory leg, as well as P. bispinosus, P. eldredgei, P. decacanthus, and P. donanen- SiS. A key to the Indo-west Pacific species with a pair of supraocular spines and two pairs of epibranchial spines is provided be- low. 1. Branchial margins of carapace armed ULLAL Sa a ae 2 — Branchial margins of carapace unarmed 2. Orbits deep. Dorsal surface of carapace with dense, short pubescence and scat- fetem. tone Selae .......... P. perdecorus — Orbits rather shallow 3. Inner orbital angles produced into small UL nee Se ee ee P. scabriculus Inner orbital angles unarmed ......... + . Rostrum trilobate ........ P. heterochrous — Rostrum sinuously triangular .... P. militaris Rostrum trilobate ........ P. decacanthus — Rostrum sinuously triangular P. donanensis, new species - | 2 “ee © © we “ee © © © © © we we ew Acknowledgments I express my sincere appreciation to Mr. K. Nomura of the Kushimoto Marine Park and Dr. T. Komai of the Natural History Museum and Institute, Chiba, for giving me the opportunity to examine this interesting 63 material. My deep gratitude is due to Dr. M. Murano of the Shin-Nippon Meteoro- logical and Oceanographical Consultant Co. Ltd., Dr. R. K. Kropp of the Battelle Ocean Sciences, Dr. V. Wadley of the CSIRO Di- vision of Fisheries, and Dr. R. Lemaitre of the National Museum of Natural History, Smithsonian Institution, for critical review- ing the manuscript and offering many help- ful comments. Literature Cited Asakura, A. 1991. Differences in the patterns of dis- tribution of the porcelain crab, Petrolisthes ja- ponicus on an oceanic island (Chichijima, Oga- sawara) and on a continental island (Zamami, Okinawa).—Researches on Crustacea 20:23- 28. Borradaile, L. A. 1900. On the Stomatopoda and Ma- crura brought by Dr. Willey from the South Seas. Pp. 395-428 in A. Willey, ed., Zoological results based on material from New Britain, New Guinea, Loyalty Islands and elsewhere, collected during the years 1895, 1896 and 1897. Part IV, University Press, Cambridge. Dana, J. D. 1852. Crustacea, Part 1.—United States Exploring Expedition during the years 1838, 1839, 1840, 1841, 1842, under the command of Charles Wikes, U. S. N. 13:1—685. (Sherman, Philadelphia. ) Haig, J. 1981. Three new species of Petrolisthes (De- capoda, Anomura, Porcellanidae) from the Indo-West Pacific.—Journal of Crustacean Bi- ology 1:265-—271. , & R. K. Kropp. 1988. Petrolisthes eldredgei, a new porcellanid crab from the Indo-West Pa- cific, with redescription of two related spe- cies.—Micronesia 20:171—186. [Issue dated De- cember 1987.] Heller, C. 1862. Neue Crustaceen gesammelt wahrend der Weltumseglung der K. K. Fregatte “‘Nova- ra.’ Zweiter vorlaufiger Bericht—Verhandlun- gen der kaiserlich-k6niglichen Zoologisch-Bo- tanischen Gesellschaft in Wien 28:519-528. Kropp, R. K. 1986. A neotype designation for Pe- trolisthes tomentosus (Dana), and description of Petrolisthes heterochrous, new species, from the Mariana Islands (Anomura: Porcellani- dae).—Proceedings of the Biological Society of Washington 99:452—463. Miyake, S. 1982. Japanese crustacean decapods and stomatopods in color. I. Macrura, Anomura and Stomatopoda. Hoikusha Publishing Co., Osaka, 261 pp. 64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ortmann, A. E. 1897. Carcinologische Studien.— Zoologische Jahrbiicher. Abtheilung fiir Syste- matik, Geographie und Biologie der Thiere 10: 258-372. Osawa, M. 1996. Two new species of the genus Pe- trolisthes (Decapoda: Anomura: Porcellanidae) from the Indo-West Pacific.—Journal of Crus- tacean Biology 16:602—612. Stimpson, W. 1858. Prodromus descriptionis animal- ium evertebratorum, quae in expeditione ad oceanum pacificum septentrionalem, a republica federata missa, Cadwaladaro Ringgold et Johan- ne Rodgers ducibus, observavit et descripsit. Pars VII. Crustacea Anomoura.—Proceedings of the Academy of Natural Sciences of Phila- delphia 10:225—252. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):65—68. 1997. A new species of Pachycheles from the Hawaiian Islands (Crustacea: Decapoda: Porcellanidae) Alan W. Harvey and Elizabeth M. De Santo Department of Invertebrates, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024, U.S.A. Abstract.—Pachycheles attaragos, new species, is described from two males from the Hawaiian Islands. The species is characterized by a relatively elon- gated, tufted carapace; carapace sidewalls with a large anterior piece but only a single vestigial posterior fragment; subequal chelae with a distinctive carpus margin; and third maxillipeds with an acutely triangular meral lobe. Although porcelain crabs are well repre- sented in tropical waters of both Indo-Pa- cific and eastern Pacific waters, only two species, Pachycheles pisoides (Heller, 1865) and Petrolisthes coccineus (Owen, 1839), are known from the Hawaiian Is- lands. Here we describe a distinctive new species, which we discovered during ex- amination of the porcellanid collections of the Museum of Comparative Zoology (MCZ). These specimens are part of a large collection of Hawaiian marine invertebrates and fishes made in the mid-1800’s by the conchologist Andrew Garrett for the MCZ through the patronage of a wealthy Boston merchant named James M. Barnard (Tho- mas 1954). Carapace length (CL) is provided as an indicator of specimen size. Illustrations were created with the approach used by Harvey & De Santo (1996): specimen im- ages were first captured on a Macintosh™ computer with a digital camera connected to a Wild M8 dissecting microscope, then prepared for publication using the programs Adobe Photoshop® and Adobe [IIlustra- tor™. Pachycheles attaragos, new species Fig. 1 Holotype.—Male (CL 3.85 mm), Sand- wich Islands (= Hawaiian Islands), coll. A. Garrett, donated to MCZ by J. M. Barnard, MCZ 1185la. Paratype.—Male (CL 2.90 mm), Sand- wich Islands (= Hawaiian Islands), coll. A. Garrett, donated to MCZ by J. M. Barnard, MCZ 11851b. Diagnosis.—Carapace longer than broad, with lateral margins convex, regions poorly defined. Front triangular in dorsal view, with tuft of setae. Sidewall of carapace con- sisting of 1 large anterior plate and 1 very small posterior fragment. Basal segment of antennule armed with 2 blunt spines on an- teromesial margin and smaller lateral spine on anterior surface. Merus of third maxil- liped with pronounced medial lobe, acutely triangular in shape. Chelipeds subequal. Carpus of cheliped with strongly projecting, angular lobe occupying proximal half of an- terior margin, and joining distal portion of margin in broad, smooth curve. Manus of cheliped covered with small flattened gran- ules. Walking legs with scattered marginal setae. Telson 5-plated. Second pleopods present in males. Females unknown. Description.—Carapace (Fig. 1A) longer than broad; regions faintly defined; dorsal surface with posterolateral regions plicate, otherwise punctate; anterolateral regions with scattered short setae, front with tuft of short plumose setae; dorsolateral ridges pronounced; posterolateral margins convex; posterior margin straight. Front triangular 66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cc A, B, G, H (4mm); |, J (2 mm) = C,D,E, F (1 mm) Fig. 1. Pachycheles attaragos, new species; holotype male, MCZ 11851a. A, carapace; B, sidewall of car- apace; C, basal segment of left antennule, ventral view; D, sternite of outer maxillipeds, ventral view; E, merus of left outer maxilliped, ventral view; FE ischium of left outer maxilliped, ventral view; G, major cheliped; H, minor cheliped; I, third right pereopod, lateral view; J, third right pereopod, mesial view. in dorsal view, trilobate in frontal view, me- eyes large. Sidewall of carapace (Fig. 1B) dian lobe projecting farther than lateral consisting of 1 large anterior plate and 1 lobes. Outer orbital angle produced into very small posterior fragment. acute tooth, inner orbital angle approxi- Basal segment of antennule (Fig. 1C) mately right-angled. Orbits deep and broad, armed with 2 large, blunt spines on anter- VOLUME 110, NUMBER 1 omesial margin (visible from dorsal view); with 1 somewhat smaller lateral spine on anterior surface; ventral surface with 2 transverse granular lines. Second segment of antenna with low crest on anterior mar- gin; third segment with proximal tubercle, low medial crest and distal tubercle along anterior margin. Flagella with short setae. Outer maxillipeds with trilobate sternite (Fig. 1D), median lobe exceeding lateral lobes; merus (Fig. 1E) with pronounced medial lobe, acutely triangular in shape, with anterior margin entire; ischium (Fig. 1F) with medioproximal angle obtuse. Chelipeds (Fig. 1G, H) subequal in length, major manus somewhat wider than minor manus. Merus with granular tooth on anterior margin, not projecting as far as car- pus tooth; ventral margin of merus distinct, ventrodistal angle granular. Carpus and ma- nus with very short, often vestigial, plu- mose setae arising in scattered groups from distal side of larger granules; less apparent on major chela. Carpus about as broad as long, with strongly projecting angular lobe occupying proximal half of anterior margin, and joining distal portion of margin in broad, smooth curve; dorsal surface rugose; carpus of major chela with medial and me- diolateral longitudinal ridges, each topped with row of enlarged granules. Manus cov- ered with small granules, more pronounced near base of dactyl; posterior margin of fixed finger with 2 parallel rows of gran- ules; dorsal surface of fingers with smaller, flattened granules. Major manus with single medial tubercle on cutting edge of pollex; dactyl with basal tubercle on cutting edge; fingers gaping, crossing at tips, with trace of setae in gape. Minor cheliped with fin- gers meeting entire length of cutting edge, crossing at tips. Walking legs (Fig. 11, J) with scattered setae on anterior margins of merus, carpus and propodus. Carpus with medial longitu- dinal ridge on lateral surface; with 1 or 2 granules at anterodistal angle. Propodus with 2 distal, 1 subdistal and 1 medial mov- 67 able spines ventrally. Dactyl with 3 corne- ous spines on ventral margin. Abdomen smooth; telson with 5 plates in males (females unknown). Second pleopods present in males. Distribution.—At present, known only from the type locality; bathymetric and hab- itat distributions unknown. Etymology.—From the Greek attaragos, meaning small piece or bit, and referring to the unusually small posterior fragment of the sidewall of the carapace. Used as a noun in apposition. Remarks.—Pachycheles attaragos pos- sesses several features unusual within the genus. The shape of the anterior lobe of the carpus of the cheliped is unlike any other species of Pachycheles, and strongly resem- bles that of porcelain crabs in the genus AI- lopetrolisthes (Haig, 1960). The elongate, acutely triangular meral lobe of the outer maxilliped is more similar to certain species of Petrolisthes than to other species of Pa- chycheles. Perhaps the most distinctive fea- ture of the genus Pachycheles, the fragmen- tation of the sidewall of the carapace, is barely evident in P. attaragos, which has only a single very small fragment posterior to the large anterior plate. Likewise, the dif- ference in size between the major and mi- nor chelipeds of P. attaragos is among the smallest in the genus (A. Harvey and E. M. De Santo, pers. obs.). Pachycheles attaragos is easily distin- guished from P. pisoides, the only other species in the genus reported from Hawaii. In P. pisoides, the carapace is broader than long; the front lacks a tuft of setae and is nearly straight; the posterior plate of the sidewall is quite large (approximately half the size of the anterior plate); and the an- terior margin of the carpus of the chelipeds possesses three or four acute, forwardly- pointing teeth. Pachycheles attaragos appears to be most closely related to three species, P. pectinicarpus Stimpson, 1858, currently known only from Hong Kong; the eastern Pacific P. grossimanus (Guérin-Méneville, 68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1835); and the western Atlantic P. laevi- dactylus Ortmann, 1892. Characters shared by these species include the tuft of setae on the carapace front; relatively elongated car- apace; lack of teeth on the anterior margin of the carpus of the chelipeds; anterior spines on the basal antennular segment; granules along the ventrodistal margin of the merus of the cheliped; and the pattern and degree of setation of the major and mi- nor chelae (Harvey & De Santo 1996). Acknowledgments We are indebted to Ardis Johnson (MCZ) for making these specimens available for study. Lu Eldredge provided valuable in- formation about Andrew Garrett. Lara Tol- chin and Portia Rollings assisted greatly with the illustrations. This work was sup- ported by a NSF Research Experience for Undergraduates fellowship and the AMNH Crustacean Fund (E. M. De Santo). Literature Cited Guérin-Méneville, E E. 1835. Observations sur les Porcellanes.—Bulletin de la Societé Naturelle de France:115—116. Haig, J. 1960. The Porcellanidae (Crustacea Anomu- ra) of the eastern Pacific.—Allan Hancock Pa- cific Expedition 24:1—440. Harvey, A. W., & E. M. De Santo. 1996. On the status of Pachycheles laevidactylus Ortmann, 1892 (Crustacea: Decapoda: Porcellanidae).—Pro- ceedings of the Biological Society of Washing- ton 109:707-—714. Heller, C. 1865. Crustaceen.—Reise der oesterrei- chischen Fregatte ‘“‘Novarra’”’ um die Erde, in den Jahren 1857, 1858, 1859, unter den Befeh- len des Commodore B. von Wiillerstorf-Ur- bair.—Zoologischer Theil 2(3)(1):1—280. Ortmann, A. E. 1892. Die Decapoden-Krebse des Strassburger Museums IV. Thiel. Die Abthei- lungen Galatheidea und Paguridea.—Zoologis- che Jahrbticher, Abtheilung ftir Systematik, Geographie und Biologie der Thiere 6:241—326. Owen, R. 1839. Crustacea. Pp. 77-88 in The Zoology of Captain Beechey’s Voyage; Compiled from the Collections and Notes Made by Captain Beechey, the Officers and Naturalist of the Ex- pedition, during a Voyage to the Pacific and Behring’s Straits Performed in His Majesty’s Ship Blossom, under the Command of EW. Beechey, R.N., ER.S., &c. &c. in the Years 1825, 26, 27, and 28. London, Henry G. Bohn. Stimpson, W. 1858. Prodromus descriptions animal- ium evertebratorum. Pars VII. Crustacea Ano- mura.—Proceedings of the Academy of Natural Sciences, Philadelphia 10:225-—252. Thomas, W. S. 1954. King of shell collectors—Pacific Ocean style. Hawaiian Shell News 2:69—71. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):69-—73. 1997. A new genus for the Central American crab Pinnixa costaricana Wicksten, 1982 (Crustacea: Brachyura: Pinnotheridae) Ernesto Campos and Mary K. Wicksten (EC) Facultad de Ciencias, Universidad Aut6noma de Baja California, Apartado Postal 2300, Ensenada, Baja California 22800, México (MKW) Department of Biology, Texas A&M University, College Station, Texas 77843-3258 Abstract.—A new monotypic genus, Glassella, is recognized from the trop- ical East Pacific for Pinnixa costaricana Wicksten, 1982. This genus shares with Alarconia Glassell, 1938, Indopinnixa Manning & Morton, 1987, Scler- oplax Rathbun, 1893 and Pinnixa White, 1846, a carapace wider than long, and third pair of walking legs the longest. Glassella is distinguished from other genera by: MXP3 with ischium-merus pyriform, carpus larger than the conical propodus, and small digitiform dactylus inserted sub-distally on the inner face of propodus. The type species is redescribed and illustrated. Pinnixa costaricana was originally de- scribed by Wicksten (1982) and placed in the genus Pinnixa White, 1846, because of its carapace shape and relative length and shape of the walking legs. During a recent revision of the Pinnixa-complex from the eastern Pacific, P. costaricana was com- pared to other Pinnixa species and to spe- cies of other genera in the Pinnotheridae with a Pinnixa-like morphology: Alarconia Glassell, 1938, Indopinnixa Manning & Morton, 1987, and Scleroplax Rathbun, 1893. We concluded that P. costaricana should be removed from the genus Pinnixa and placed in a new genus herein diag- nosed. The morphological analysis of P. costaricana was based upon the study of the female holotype (AHF 806) deposited in Los Angeles County Museum of Natural History. Other species studied were: the type species of Alarconia, A. seaholmi, the type species of Scleroplax, S. granulata, and the species of Pinnixa reported by Bon- fil et al. (1992), Zmarzly (1992), Martin & Zmarzly (1994), and Hendrickx (1995), all deposited in the San Diego Natural History Museum and at the Invertebrates Labora- tory, Facultad de Ciencias, Universidad Au- tonoma de Baja California. The information used on the type species of Indopinnixa, I. sipunculana, was obtained from published description and figures, although critical features were confirmed by Dr. Raymond B. Manning from types deposited in the Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. (USNM). Comparison of these genera is provided in a dichotomous key based on adult characters. The third maxilliped is abbreviated to MXP3 and the walking legs are indicated as WL1-WL4. AHF is an acronym for Al- lan Hancock Foundation. Family Pinnotheridae Glassella, new genus (Figs. 1, 2) Diagnosis.—Carapace suboblong, dorsal surface pockmarked, wider than long, in- tegument firm, regions not defined; cardiac ridge lacking; front truncated, with shallow median sulcus. MXP3 with ischium-merus pyriform, fused, separated by faint line and distal margin truncated; palp as long as is- chium-merus, 3-segmented, dactylus small, 70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. holotype AHF 806. Dorsal view. Carapace length 1.8 mm, width 4.2 mm. Glassella costaricana (Wicksten, 1982), digitiform, inserted sub-distally on inner face of conical propodus; carpus stout, lon- ger than combined length of propodus and dactylus; exopod with median lobe on outer margin, flagellum 2-segmented. WL1-4 pockmarked, relative length 3 > 2 > 1 > 4, WL3 considerably the longest. Abdomen of female with 6 somites and telson free, widest at third somite; tapering from fourth somite to triangular telson. Male unknown. Etymology.—Named in honor of Steve A. Glassell, who studied the pinnotherid crabs of the eastern Pacific and made in- valuable contributions on this group. Gen- der feminine. Type species.—Pinnixa costaricana Wicksten, 1982, by present designation and monotypy. Glassella costaricana (Wicksten, 1982), new combination (Figs. 1, 2) Pinnixa costaricana Wicksten, 1982:579— 582. fig. 1, 2A—D; Hendrickx, 1995:148. Material examined.—Female (holotype AHF 806). Redescription.—(Modified from Wick- sten 1982). Carapace suboblong, cylindri- cal, dorsal surface pockmarked, regions not defined; anterolateral margins diverging posteriorly, forming shoulders from which side walls drop vertically; lacking antero- lateral crest or cardiac ridge. Posterior mar- gin straight. Front not advanced, truncate; with shallow median sulcus. Carapace mar- gin with setae, especially on ventral sur- face. Orbits small, slightly inclined down- ward, filled by eyes when retracted. Anten- nule plicate in small fossettes. Antenna large, multi-articulated. Cheliped slender, setose. Margins of che- la subparallel; ventral margin of propodus with small tubercles. Manus with group and lines of tiny tubercules. Fingers slender; tips pointed, curved, and leaving no gape when closed; dorsal margin of dactylus with blunt and acute tubercles. WLI1-2 slender; meri trigonal; dactyli sharp, nearly straight. WL1 twisted, some- what smaller than second, nearly reaching end of propodus of WL2. WL2 reaching carpus of WL3. WL3 very wide; merus VOLUME 110, NUMBER 1 71 Fig. 2. Glassella costaricana (Wicksten, 1982), holotype AHF 806. A, third maxilliped (inner view); B, female abdomen; C, left chela (outer view); D, front, anterior view. 1.25X as long as wide, with stout tooth and small teeth and tubercles on ventral margin; carpus without teeth; propodus 1.5X as long as wide, with granules along flexor margin; dactyl sharp, slightly curved. WL4 short, reaching almost to end of WL3; dac- tyl stout, and triangular. All legs pilose and pockmarked. Distribution and habitat—Known only from the type locality, Playa de Coco, Prov- incia de Guanacaste, Costa Rica (about 10°5'N, 85°45’W); low intertidal zone, sand and rock. Remarks.—As noted in the key given be- low, the genus Glassella most resembles Pinnixa, Alarconia, Indopinnixa and Scler- oplax, all of which have a firm carapace that is wider than long and a third pair of WL considerably longer than the rest. Dif- ferences among these genera include: shape, degree of development of regions and hardness of the carapace; shape, degree of fusion, articulation point and relative length of the MXP3 articles; relative length of WL; and, width and degree of fusion of the abdominal somites. Glassella costaricana can be distin- guished from all other pinnotherids by the following presumed autapomorphies: MXP3 with pyriform ischium-merus; f2. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON shape, relative length and insertion point of the articles of the palp; and shape of the abdomen. Key to Pinnixa-like genera of the world (Carapace wider than long, firm or hard; third pair of walking legs longest.) 1. Palp of MXP3 with dactylus shorter than and inserted sub-distally on inner face of propodus; carpus larger than propodus RR rien: oie, ema Glassella, new genus (Pacific coast of Costa Rica; type species Pinnixa costaricana Wicksten, 1982; host unknown). — Palp of MXP3 with dactylus as long as or longer than and inserted proximally on ventral margin of propodus; carpus shorter than propodus 2. Ischium and merus of MXP3 not fused, subequal in length; carapace regions well defined; gonopods protruding from ster- nal trench and reaching to buccal cavity A ee Alarconia Glassell, 1938 (Mexican Pacific; type species Alarconia seaholmi Glassell, 1938; host unknown). — Ischium shorter than merus or both arti- cles fused; carapace regions not well de- fined, gonopods not protruding from sternal trench and not reaching the buc- cal cavity 3. Carapace subpentagonal, hard, dorsally very convex; WL1-4 of similar shape, third pair slightly longer, fourth not notice- ably reduced .... Scleroplax Rathbun, 1893 (Eastern Pacific, British Columbia, Canada to Baja California, Mexico; type species Scleroplax granulata Rathbun, 1893; hosts: in burrows of Crustacea-Decapoda (Callianassidae) and Echiurida. — Carapace oblong, firm but not hard, flat or slightly convex; WL1-4 very dissim- ilar in shape, third pair stout and longer, and fourth noticeably shorter 4. Propodus of MXP3 elongated, distal end expanded far beyond mid-length of dac- tylus, both articles of similar shape; male abdomen of 6 free somites and telson PAs SES AS 8 Re Pinnixa White, 1846 (Western Atlantic [Massachusetts, U.S.A. to Argentina]; Eastern Pacific [Alaska, U.S.A. to Chile]; Indo West Pa- cific [Japan, East Africa]; type species Pinnotheres cylindricum Say, 1818; host: Polychaeta, Enteropneusta, Echiur- ida, Sipunculida, Holothuroidea, Mollus- ca-Bivalvia, Crustacea-Decapoda [Calli- anassidae], Tunicata). — Propodus of MXP3 short and stout, dis- tal end not reaching far beyond middle length of dactylus, both articles very dis- similar in shape; male abdomen with fifth and sixth somites fused ....... ... Indopinnixa Manning & Morton, 1987 (Indo West Pacific [Hong Kong]; type species Indopinnixa sipunculana Man- ning & Morton, 1987; in burrows of Si- punculida). Acknowledgments We are deeply grateful to Raymond B. Manning (USNM) for provide valuable in- formation on Indopinnixa sipunculana; to R. B. Manning, Rafael Lemaitre and Mar- cos Tavares for reviewing our manuscript with great care; to Joel W. Martin and George E. Davis (LACMNH) for the loan of the holotype of Glassella costaricana; and to Alma Rosa de Campos for her very fine artistic work. This work was partially supported by program 0134 “Crustaceos Simbiontes del Pacifico Mexicano (former- ly de Baja California)”’ of the Facultad de Ciencias, Universidad Autonoma de Baja California (UABC) and by agreement UABC-CONACYyT 3587-N9311. EC is a fellow of the “‘Programa de Estimulo al Personal Académico 96/97”’ of the UABC. Literature Cited Bonfil, R., A. Carvacho, & E. Campos. 1992. Los cangrejos de la Bahia de Todos Santos, Baja California. Parte II. Grapsidae, Pinnotheridae y Ocypodidae (Crustacea; Decapoda: Brachy- ura).—Ciencias Marinas (México) 18(3):37—5S6. Glassell, S. A. 1938. New and obscure decapod Crus- tacea from the west American coasts.—Trans- actions of the San Diego Society of Natural His- tory 8(33):411—454. Hendrickx, M. E. 1995. Checklist of brachyuran crabs (Crustacea: Decapoda) from the eastern tropical Pacific.—Bulletin de L’'Institut Royal des Sci- ences Naturelles de Belgique (Biologie) 65: 125-150. VOLUME 110, NUMBER 1 Manning, R. B., & B. Morton. 1987. Pinnotherids (Crustacea: Decapoda) and Leptonaceans (Mol- lusca: Bivalvia) associated with sipunculan worms in Hong Kong.—Proceedings of the Bi- ological Society of Washington 100:543-—551. Martin, J., & D. L. Zmarzly. 1994. Pinnixa scamit, a new species of Pinnotherid crab (Decapoda: Brachyura) from the continental slope off Cal- ifornia.—Proceedings of the Biological Society of Washington 107:354-—359. Rathbun, M. J. 1893. Scientific results of explorations by the U.S. Fish commission steamer Albatross, XXIV. Description of new genera and species of crabs from the west coast of North America and the Sandwich Islands.—Proceedings of the United States National Museum 16:233—260. Say, T. 1817-1818. An account of the Crustacea of ~) ee) the United States.—Journal of the Academy of Natural Sciences of Philadelphia 1(1—2):57-63, 65-80, 97-101, 155-160, 161-169 [all 1817], 235-253, 313-316, 317-319, 374-380, 381- 401, 423-441 [all 1818]. Wicksten, M. K. 1982. Pinnixa costaricana, a new species of crab from Central America (Brachy- ura: Pinnotheridae).—Proceedings of the Bio- logical Society of Washington 95:579-582. White, A. 1846. Notes on four new genera of Crus- tacea.—Annals and Magazine of Natural His- tory 18(118):176—-178. Zmarzly, D. L. 1992. Taxonomic review of pea crabs in the genus Pinnixa (Decapoda: Brachyura: Pinnotheridae) occurring on the California shelf, with description of*two new species.— Journal of Crustacean Biology 12:677-713. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):74—98. 1997. New records of marine Isopoda from Cuba (Crustacea: Peracarida) Brian Kensley, Manuel Ortiz, and Marilyn Schotte (BK & MS) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (MO) Centro de Investigaciones Marinas, Universidad de la Habana, La Habana, Cuba Abstract.—Seven new species are described from localities on both the north and south coasts of C.:ba: Cyathura (Cyathura) esquivel, Mesanthura frances, Joeropsis juvenilis, Joeropsis unidentata, Dynamenella nuevitas, Paraimene ibarzabalae, Paraimene tumulus. Several new records, including Carpias har- rietae, Caecijaera horvathi, and Sphaeromopsis mourei are noted. The material was collected from a range of habitats, including shallow coral reefs, man- groves, seagrass beds, coastal lagoons, river mouths, rocky and sandy shore inter-/and shallow infratidal areas. Knowledge of the marine, freshwater, and cave isopod fauna of Cuba has grown slow- ly, from the earliest records of two species of Aega by Schioedte & Meinert in 1879. Numerous short papers, often describing a single species, have accumulated over the years. The first cave isopod was described by Hay in 1903; since then several addition- al species especially in the genus Cyathura have been added to the list. Coineau & Bo- tosaneanu (1973) produced the only report on interstitial isopods from Cuba. Ortiz et al. (1987) provided an updated list of Cuban isopods along with a bibliography. From the relatively small number of species in this list, it is obvious that many more await dis- covery, as many regions of the island’s shal- low and deep waters have not been collect- ed. In an attempt to fill some marine distri- butional gaps, and to document the diversity of several groups of shallow water marine organisms, two collecting trips, in April 1994, and May/June 1995, were carried out jointly by Cuban and Smithsonian Institution scientists. This work was sponsored chiefly by the Center for Marine Conservation, Washington, D.C. The 1994 trip was based aboard the R/V Ulises, which travelled along the north coast through the Archipielago de Camagiiey as far as Bahia de Nuevitas, stop- ping at several localities to carry out inten- sive sampling. The 1995 trip was to the south-western part of the island, mainly in the region around the Isla de la Juventud. The material reported in this paper was collected primarily by the authors, although several other individuals assisted. K-CUBA station numbers refer to field notes for the two trips. Holotypes have been deposited in the Centro Colecciones Naturales Marinas, Instituto de Oceanologia (IO), Havana, Cuba; paratypes and additional materials are deposited in both the CCNM and the Na- tional Museum of Natural History (USNM), Smithsonian Institution. An annotated checklist of the marine isopod fauna of Cuba is being compiled, based on a variety of sources of material. Dimensions in millimeters are always to- tal length measured along the dorsal midline. Systematic Section Suborder ANTHURIDEA Leach, 1814 Family Anthuridae Leach, 1814 Cyathura (Cyathura) esquivel, new species Figs. 1, 2 Material.—Holotype, IO-12.055, ¢ 2.5 mm, Allotype, IO-12.057, ovigerous 2 4.0 VOLUME 110, NUMBER 1 75 A e ak SS on cart << SSS — SE = See = ss —~ age ise BS ws VY Fig. 1. Cyathura esquivel, new species. A, Ovigerous female in dorsal view, scale = 1 mm; B, Antenna; C, 3 Antennule; D, 2 Antennule; E, Mandible; EK Maxilla; G, Maxilliped; H, 2 Pleopod 1; I, ¢ Pleopod 1; J, 3 Pleopod 2. 76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Cyathura esquivel, new species. A, Pleotelson; B, Uropodal endopod and exopod; C, d Pereopod 1, mesial surface; D, Pereopod 2; E, Pereopod 7; EK 2 Pereopod 1, lateral surface; G, 2 Pereopod 1, mesial surface. VOLUME 110, NUMBER 1 mm, Paratypes, IO-12.056, 4 6, 4 oviger- ous 2, 5 non-ovigerous ¢; Paratypes, USNM 253270, 5 6, 4 ovigerous 2, 9 non- ovigerous 2, 4 manca, sta K-CUBA-22, Cayo Esquivel, north coast of Cuba, coral rubble from exposed side of cay, 2—2.5 m, 12 Apr 1994.—4 ¢ 2.5 mm, 2 non-oviger- ous ¢, 12 manca 1.8—2.2-mm, sta K-CU- BA-35, Cayo Coco, north coast of Cuba, coral rubble from shallow reef area, 1.5—2 m, 14 Apr 1994. Description.—Male: Antennule with pe- duncle article 3 shorter than in female, car- rying distal band of fine aesthetascs; flagel- lum of 2 articles, basal article also carrying band of fine aesthetascs. Pereopod 1, mesial surface of propodus having row of about 14 setae. Pleopod 1, endopod shorter than, and about % width of exopod. Pleopod 2, en- dopod having slender, parallel-sided copu- latory stylet articulating in proximal half of mesial margin, with 3 distal plumose mar- ginal setae; exopod with transverse suture in distal half, 5 plumose marginal setae dis- tally. Pigment pattern similar in male and fe- male, with large red-brown chromatophores in band between eyes and forming 2 pos- terior lobes on dorsal cephalon. Pereonites each with fine dorsal squiggles, those on pereonite 2 forming 2 contiguous rings be- ing most characteristic. Pleon with pair of irregular lateral rings often with intersecting line. Ovigerous female: Body proportions: C<1>2>3=4=5=6>7>P. Cephalon wider than middorsal length, with low rounded rostrum. Eyes well pigmented. Body bent between pereonites 1 and 2. Anterior fused segments of pleon short. Pleotelson with broad transparent marginal band, having 6 pairs of submarginal setae in posterior half, slightly bilobed posterior margin with elon- gate pair of setae submesially. Antennule having 3 relatively large arti- cles; flagellum of 3 short articles, together shorter than peduncle article 3; terminal ar- ticle bearing 2 aesthetascs. Antennal pedun- cle with article 2 stout, grooved to accom- 77 modate antennule; articles 3 and 4 sub- equal; article 5 longer than 4; flagellum of 2 very short setose articles. Mandibular in- cisor of 3 sclerotized cusps; lacinia dentata having 8 teeth; molar thin-walled, distally rounded; palp with article 2 twice length of article 1; article 3 short, bearing 5 stout se- tae. Maxilla having single strong spine and several slender subsidiary spines. Maxilli- ped of 3 articles, article 2 slightly longer than 1, with few setae submesially; article 3 semicircular, with 5 mesial setae; endite lacking. Pereopod 1, carpus short, lacking free anterior margin, with 2 setae distally; propodus expanded, with transparent pal- mar flange bearing 6 setae on lateral sur- face, with low rounded proximal protuber- ance, with row of 5 setae on mesial surface; unguis % length of rest of dactylus. Pereo- pods 2 and 3 with short triangular carpus lacking free anterior margin; propodus roughly rectangular, with stout serrate pos- terodistal spine; unguis % length of rest of dactylus, with small accessory spine at its base. Pereopods 4-7 similar, carpus trian- gular, with short free anterior margin; prop- odus with several posterodistal scales and stout serrate posterodistal spine. Pleopod 1 protopod with 2 coupling hooks; exopod opercular; endopod subequal to exopod in length, between % and % width of exopod. Uropodal endopod roughly ovate, with broad transparent border and numerous marginal setae; exopod with broad trans- parent border, almost 3 times longer than wide, lateral margin crenulate with plumose setae set in each gap. Remarks.—The only species of the sub- genus Cyathura previously recorded from Cuba (and from the general Caribbean re- gion), is C. cubana Negoescu, 1979, an es- tuarine species also known from the main- land of Belize (Kensley & Schotte 1989). Cyathura cubana differs from C. esquivel, which occupies fully marine habitats, in having a more heavily pigmented and se- tose body, in having an evenly rounded posterior margin of the pleotelson, a much shorter uropodal exopod, a much shorter 78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON endopod of pleopod 1, and a more elongate mandibular palp. Cyathura tridentata Wag- ner, 1990, from the interstitial intertidal of the Dominican Republic, is a blind and un- pigmented species, differing in many ap- pendage characters from the present spe- cies. Etymology.—The specific name derives from the type locality, Cayo Esquivel. Mesanthura frances, new species Fig. 3 Material.—Holotype, IO-12.058, non- ovigerous ? 8.2 mm, Paratypes, IO-12.059, 2 non-ovigerous ¢, Paratypes, USNM 253271, 3 non-ovigerous 2, sta K-CUBA- 27, Cayo Francés, from clumps of Hali- meda in Thalassia flat, shallow infratidal, 13 Apr 1994.—Non-ovigerous 2 4.1 mm, sta K-CUBA-35, Cayo Coco, coral rubble from shallow reef area, 1.5—2.m, 14 Apr 1994. Description.—Non-ovigerous female: Body proportions: C<1=2<3<4=5>6>7 Ss, Sa we) SSS PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. A. cubensis holotype, adult female: left 6th limb, lv. Infold: Typical for genus (Fig. 1b). Central adductor muscle attachments (Fig. la, c): Comprising about 25 elongate attachments. Carapace size: Holotype, length 3.2 mm, height 2.5 mm; height 78.1% of length. First antenna (Fig. 2a, b): 1st joint with groups of very short spines near dorsal mar- gin and groups of longer spines near ventral margin. 2nd joint: ventral and dorsal mar- gins spinous; dorsal margin with 2 long dis- tal bristles; lateral surface with 6 short dis- tal bristles (these are dashed in Fig. 2a). 3rd joint with 12 long dorsal bristles and 1 short ventral bristle. 4th joint with 4 ventral bris- tles (3 short, 1 long (about 3X length of Sth joint)) and 1 long terminal dorsal bristle. 5th joint; dorsal margin with 7 or 8 nodes; ventral sensory bristle with 7 short proxi- mal filaments separated by short space from 2 filaments about 3X length of proximal fil- aments and then 7 long terminal filaments (Fig. 2a, b). 6th joint with long terminal medial bristle. 7th joint: a-claw bare, longer than 5th joint: b-/and c-bristles each with 9 marginal filaments (distal 2 longer than oth- ers). 8th joint: d-/and e-bristles about % length of c-bristle, both bare with blunt tips; f-bristle bent dorsally, tip missing on ho- lotype, with 9 filaments on remaining part; g-bristle about same length as c-bristle, with about 9 filaments. Second antenna (Fig. 2c): Protopodite with short distal medial bristle. Endopodite 3-jointed: 1st joint with 6 short bristles; 2nd joint shorter than Ist, with 2 distal bristles; 3rd joint short with long terminal filament. Exopodite: 1st joint with minute medial ter- minal bristle; bristles of joints 3—8 with na- tatory hairs and stout ventral spines; joints 2-8 with stout basal spines; 9th joint with stout lateral spine and 4—5 terminal bristles (2 with stout ventral spines). Mandible (Fig. 3a): Coxale endite lost. Basale: endite with about 11 long bristles (triaenid and end types) and 5 ventral dwarf bristles; ventral margin with 8 or 9 triaenid bristles (ventral margin folded over in Fig. 3a); dorsal margin with about 16 short bris- tles and 2 long terminal bristles. Exopodite reaching just past distal end of dorsal mar- gin of Ist endopodial joint, with 2 subter- VOLUME 110, NUMBER 1 4 ye , \S pr a“ Tas rBeaccmabll / | va / i gitd_ a => / Lad Yj | tas is iy! VA ia | by Teh 3 iy Nur ts mn / Ry * L | Ns | Os / Fig. 6. 113 a, A. cubensis holotype, adult female, posterior of body from left side (note small thumb-like process (pr)). b, A. americana (Miiller), USNM 157148, adult female from Pacific, posterior of body from left side. minal bristles and hirsute tip. 1st endopo- dial joint with 6 ventral bristles. 2nd en- dopodial joint: ventral margin with 2 sub- terminal and 2 terminal bristles; dorsal margin and medial surface near dorsal mar- gin with numerous long and short bristles. 3rd endopodial joint broken off. Maxilla (Fig. 4a): Epipodite triangular. Endites I and II fused, with 11 bristles (6 long, 5 shorter). Basale: medial surface with 7 proximal bristles and 6 distal bris- tles; lateral side with short proximal bristle; dorsal margin with proximal spines and about 9 distal bristles (2 long, remainder short); ventral margin with 18 short bristles followed by 3 longer bristles and a very long terminal bristle. Exopodite small with 3 bristles (2 short, 1 long). Endopodite % length of basale: 1st joint with short ante- rior alpha-bristle and long beta-bristle; 2nd joint with 5 terminal bristles (2 short, 3 long). Fifth limb (Fig. 4b): Dorsal margin of comb with 5 small proximal bristles; ante- rior edge of comb smoothly rounded and hirsute; exopodial bristles consisting of long stout spinous bristle, 4 minute bristles ventral to base of stout bristle, and 6 small bristles (with several long hairs near tip) close to ventral edge of comb near mid- length. Sixth limb (Fig. 5): With 1 epipodial 114 bristle; anterior margin with 2 distinct su- tures; anterior edge of stem and ventral margin of skirt with numerous bristles; pos- terior end of skirt with 4 plumose bristles (all hairs not shown); posterior extension of skirt with numerous spines; tip of lateral flap with 2 bristles. Seventh limb (Fig. 1d): Each limb with about 64 bristles, about same number on each side; each bristle with 5—7 bells; ter- minus with opposing combs, each with 18— 20 teeth. Furca (Fig. 3b): Each lamella with 3 stout claws followed by 1 slender bristle, a fourth stout claw, and then 7 slender bris- tles; ventral margin of lamellae with many small spines. Bellonci organ (Fig. 3c): Typical for ge- nus, with striae near midlength and narrow tip. Posterior of body (Fig. 6a): Hirsute, with small thumb-like posterodorsal process. Number of eggs: Holotype with 57 eggs in marsupium. Length of typical egg 0.32 mm. Comparisons.—The new species A. cub- ensis differs from A. americana (Miiller, 1890) in having a thumb-like node on the posterodorsal corner of the body (compare Fig. 6a and 6b). The dorsal margin of the Sth joint of the Ist antenna of the unique female A. cubensis bears 7—8 nodes com- pared to 4—6 for A. americana, but vari- ability of former is not known. The dorsal margin of the mandibular basale of A. cub- ensis bears about 16 short bristles compared to 7 or 8 for A. americana. The Ist antenna of A. cubensis differs from that of A. antyx Kornicker, 1981, in having nodes along the dorsal margin of the 5th joint, and in not having teeth on the a-bristle of the 7th joint. Acknowledgments The senior author wishes to thank Eliz- abeth Harrison-Nelson (Smithsonian Insti- tution) for general assistance during a stay at the U.S. National Museum of Natural PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON History. The unique specimen described herein was collected with a net by Marilyn Schotte (Smithsonian Institution). We thank the personnel aboard the R/V Ulises for their participation. The illustrations were drawn by the senior author using a camera- lucida. Literature Cited Hartmann, G. 1974. Die Ostracoden des Untersu- chungsgebiets, Part 3: Zur Kenntnis des Euli- torals der afrikanischen Westkiiste zwischen Angola und Kap der Guten Hoffnung und der afrikanischen Ostktiste von Stidafrika und Mo- ¢ambique unter besonderer Beriticksichtigung der Polychaeten und Ostracoden.—Mitteilun- gen aus dem Hamburgischen Zoologischen Mu- seum und Institut 69:229—520. Kornicker, L. S. 1958. Ecology and taxonomy of Re- cent marine ostracodes in the Bimini area, Great Bahama Bank.—Publications of the Institute of Marine Science (The University of Texas) 5: 194-300. 1981. Revision, distribution, ecology, and ontogeny of the ostracode subfamily Cyclaster- opinae (Myodocopina: Cylindroleberididae).— Smithsonian Contributions to Zoology 319:1— 548. 1985. Sexual dimorphism, ontogeny, and functional morphology of Rutiderma hartmanni Poulsen, 1965 (Crustacea: Ostracoda).—Smith- sonian Contributions to Zoology 408:1—28. 1986. Cylindroleberididae of the western North Atlantic and northern Gulf of Mexico, and zoogeography of the Myodocopina (Ostra- coda).—Smithsonian Contributions to Zoology 425:1-139. 1994. Ostracoda (Myodocopina) of the SE Australian Continental Slope, part 1.—Smith- sonian Contributions to Zoology 553:1—200. Miiller, G. W. 1890. Neue Cypridiniden.—Zoologis- che Jahrbiicher 5:211—252. . 1906. Die Ostracoden der Siboga-Expedition. Leiden: E.J. Brill, 40 pp. Poulsen, E. M. 1965. Ostracoda-Myodocopa, 2: Cy- pridiformes—Rutidermatidae, Sarsiellidae, and Asteropidae.—Dana Report 65:1—484. Sars, G. O. 1865. Oversigt af Norges marine Ostra- codes.—Fordhandlinger i Videnskabs-Selskabet I Christiania 7:1—130. [Preprint: Serial pub- lished 1866.] Skogsberg, T. 1920. Studies on marine ostracods, I: Cypridinids, Halocyprids, and polycopids.— Zoologiska Bidrag fran Uppsala, (supplement) 1:1—784. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):115-127. 1997. Macrothrix smirnovi, a new species (Crustacea: Anomopoda: Macrothricidae) from Mexico, a member of the M. triserialis-group J. Ciros-Pérez and M. Elias-Gutiérrez Laboratorio de Zoologia, Universidad Nacional Aut6noma de México Campus Iztacala, A.P. 314, C.P. 54000. Los Reyes Iztacala, Tlalnepantla, Edo. Mexico, Mexico Abstract.—Macrothrix smirnovi, new species, is described from a small res- ervoir located at a transitional zone between neartic and neotropic regions in Mexico. It resembles taxa related to M. triserialis, mainly the South-American M. superaculeata, but parthenogenic females are characterized by differences in spinulation of the antennae. Trunk limb II has a unique lobe carrying a stout conical seta densely ciliated at its distal portion located at the external surface of the endopod, near to the insertion of scrapers 4 and 5. The postabdomen is also distinct. The ephippium has a structure similar to M. rosea. Adult males have a postabdomen fairly similar to females and a copulatory hook with one crescentic ridge at the tip of an irregular margin which gives a spoon-like appearance to this portion. Recently, new concepts about diversity in tropical freshwater zooplankton indicate that at minimum the same number of spe- cies of cladocerans (ca. 50 per lake) are found in tropical as in temperate systems (Dumont 1994). In addition, the statements of Frey (1982a, 1988a) about non-cosmo- politanism in cladoceran species were con- firmed (Frey 1988b), mainly on chydorids. On the other hand, if temperate regions were the most surveyed for their freshwater fauna while tropical zones generally were ignored until the last decade, many new species will remain to be described from tropical zones. Dumont (1994), based on data published by different authors, esti- mates a future increase at about 25% in the number of known cladoceran taxa. Most new descriptions will be from the tropics. This paper mainly deals with a new ma- crothricid cladoceran from central Mexico which is located at a transitional zone be- tween neartic and neotropic regions. Samples were collected from the littoral zone of water bodies, with a 50 wm-mesh plankton net and were fixed with sugar- formaldehyde (Haney & Hall 1973). The paratype material of M. superaculeata (Smirnov) (deposited at the Instituto Na- cional de Pesquisas de Amazonia, Brazil INPA-CR 3035) and mature parthenogenet- ic females of M. triserialis s.1. from Malay- sia, Nepal, Nigeria, Nicaragua and Mexico, M. paulensis (Sars) from Brazil, M. rosea (Jurine) from Russia, M. capensis (Sars) from South Africa and Australia and M. odiosa (Gurney) from Malaysia also were studied. All measurements were made according to Smirnov (1971). Structures were dissect- ed with tungsten needles and mounted in a mixture of glycerin-formaldehyde for the fine analysis. Drawings were made with a camera lucida attached to a microscope Ni- kon Labophot-2. Macrothrix smirnovi, new species Figs. 1—35 Material.—Holotype: One adult oviger- ous female (total length 0.8 mm, height 0.5 mm), The Natural History Museum (BMNH), London, England, 1995.1290, 16 Aug 1995. Allotype: One adult male (total 116 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 200 ym x eens \ ne Mt 11n \ SNih > Sa e &S : Ze ; oo ~ Dp: B I) \SSSSa ‘o ‘SZ SoS ) ‘Sy i ss ¢ » — / \ b Y fe S 4 ~ ~) \ si u 400 um ri} s Aw» AY NY AN ws TNS Figs. 14-23. Macrothrix smirnovi, new species from kilometer 28 on the highway Jilotepec-Ixtlahuaca, State of Mexico, Mexico. All are parthenogenetic female except Figs. 22 and 23 which are ephippial females. 14, left trunk limb II, lateral; 15, right trunk limb III, medial; 16, right trunk limb IV, lateral; 17, left trunk limb V, medial; 18, lateral view of postabdomen; 19, terminal claw of postabdomen, lateral; 20, termina! claw of pos- tabdomen, medial; 21, seta natatoria of postabdomen; 22, frontal view of ephippial female; 23, lateral view of ephippial region of a sexual female, showing the slough line. Scale bar “‘b” is for Figs. 14, 15, 16 and 17. Abreviations: EX, exopod; EN, endopod; GN, gnathobase; DCS, densely ciliated seta; IA, inner armature; EA, external armature; EP, epipodite. 118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 24—35. Macrothrix smirnovi, new species from kilometer 28 on the highway Jilotepec-Ixtlahuaca, State of Mexico, Mexico. Males. 24, lateral view of instar-I male; 25, lateral view of instar-II male; 26, lateral view of mature male; 27, AI of instar-I male, medial (aesthetascs are omitted); 28, AI of instar-II male, medial (aesthetascs are omitted) arrow indicates point of insertion of external, medial seta; 29, AI of instar-II male, medial (aesthetascs are omitted), arrow indicates point of insertion of external medial seta; 30, AI of instar-II male, lateral; 31, part of trunk limb I of instar-I male; 32, part of trunk limb I of instar-II male, showing the VOLUME 110, NUMBER 1 length 0.48 mm, height 0.32 mm) BMNH 1995.1291, 16 Aug 1995. Paratypes: One ephippial female, BMNH 1995.1292, 16 Aug 1995. All mounted on slides in gly- cerine jelly sealed with depict .mounting medium. One adult ovigerous female, mounted on a slide in glycerine jelly sealed with DePeX mounting medium and five parthenogenetic females in 4% formalde- hyde solution, with a drop of glycerol add- ed, National Museum of Natural History, Washington, D.C., U.S.A., USNM 274176, 29 Aug 1995. Five parthenogenetic females in 4% formaldehyde solution, with a drop of glycerol added, Zoological Museum of Moscow University, ZMMU 4010, 14 Aug 1995; one adult ovigerous female mounted on a slide in glycerine jelly sealed with DePeX mounting medium and five parthe- nogenetic females in 4% formaldehyde so- lution, with a drop of glycerol added, Insti- tuto Nacional de Pesquisas da Amaz6nia, Manaos, Amazonia, Brazil, INPA-CR 569 and INPA-CR 570. One slide each of in- stars I, II and III of males and one ephippial female mounted in glycerine jelly sealed with DePeX mounting medium; 10 parthen- ogenic females in 4% formaldehyde solu- tion, with a drop of glycerol added, Museo de Zoologia, Universidad Nacional Autén- oma de México, Campus Iztacala, Mexico. UNAM-CI 1047 to UNAM-C1 1051. All remaining specimens and ephippia of both populations, including two dissected Ovigerous females and one dissected male of each instar, mounted on slides in glycerol sealed with DePeX mounting medium, are deposited at the Museo de Zoologia, Univ- ersidad Nacional Aut6énoma de México, Campus Iztacala, Mexico. Type locality and habitat——A small res- ervoir located at kilometer 28 on the high- <_ 119 way Jilotopec-Ixtlahuaca, State of Mexico. The geographical coordinates are 19°49'14’N, 99°41'50”W at 2740 m above sea level. At the time of sampling, we recorded a water temperature of 14°C (air temperature 19°C), conductivity 135 mS/cm and pH 7.0. No other macrothricid species were found at the time of collection. The date of collec- tion was 3 Dec. 1993. Second locality: A temporary pond lo- cated at kilometer 44 on the highway To- luca-Atlacomulco, State of Mexico. The ap- proximate coordinates are 19°38'54’N, 99°47'24"W at 2540 m above sea level. The date of collection was 3 Dec 1993. Etymology.—The species is named for Dr. Nikolai N. Smirnov, from the Russian Academy of Sciences, as a tribute to his work in Cladocera. Diagnosis.—This species is_ character- ized by the arrangement of antennal arma- ture, which is one spine on the distal edge of segments 2 and 4 from the endopod branch, plus an accessory spine on seg- ments 2 and 3, about a half length of the other spines. A distinctive lobe carrying a stout, conical seta is found on endopod ex- ternal surface of trunk limb II. The exopod of trunk limb V has only one seta. The pre- anal postabdominal spinules subequal in size are arranged in a stripe of several rows. The distal segment of postabdominal seta natatoria is three to four times shorter than the proximal segment. Males are character- ized by an ontogenetic development over the three instars; the last instar is mature. Copulatory hook on trunk limb I is strong with a crescentic ridge at the tip. The pos- tabdomen is slightly shorter and its shape is as in female. The antennula has a charac- teristic pattern of setae and spines proxi- mally on the medial surface. It has two ba- enlarging copulatory hook; 33, part of trunk limb I of mature male, showing the highly modificated setae on ODL and IDL; 34, detail of tip copulatory hook on trunk limb I of mature male; 35, lateral view of postabdomen of mature male. Scale bars: ‘“‘c’’ is for Figs. 24, 25 and 26; “‘d”’ is for Figs. 27, 28, 29 and 30; ‘‘e”’ is for Figs. 31 and 32. Abreviations: ODL, Outer distal lobe; IDL, inner distal lobe; CH, copulatory hook; GP, genital pore. 120 sal sensory setae and a long seta attached at the middle region of the external surface. Ephippial female with two ephippial eggs; about the same shape in lateral view as par- thenogenetic female. The ephippial surface has a mesh-like pattern and a dorsal margin with a row of sclerotized rounded papillae. Measurements.—Mature parthenogenetic females: Total length of the body from 0.70 to 1.05 mm, height from 0.48 to 0.75 mm (n = 31). Males: Instar I, length from 0.37 to 0.4, height from 0.24 to 0.26 (n = 3); instar II, length from 0.45 to 0.52, height from 0.27 to 0.32 (n = 25); mature, length from 0.45 to 0.55, height from 0.29 to 0.45 (7 = 13). All measurements are in mm. Description of parthenogenetic female.— Shape and shell: Body ovoid (length : max- imum height = 1.45—-1.60 mm). Dorsal margin is curved from the supraocular re- gion to the posterior-dorsal angle, with a slight depression in the vicinity of the head pore (Fig. 1); a pointed keel at the top of the shell is evident, clearly shown from a frontal view of the animal (Fig. 2). The dor- sal margin of the shell is without serration. The ventral margin is deeply serrated with two rows of movable spines which are in- serted at the submarginal edge. The external row of sparse needle-like spines increases in length distally and are outwardly direct- ed. The inner row of bilaterally spinulated spines is inwardly directed, with every two of these followed by one spine of the ex- ternal series; close to the middle region, the proximal member of the spinulated spines Starts becoming stout and longer, while the second decreases in size toward the poste- rior angle. The entire surface of the shell has dots and is striated with a polygonal pattern. Head: Evenly rounded with a slight su- praocular bulge, tapering to the rostral re- gion. The frontal part of the rostrum is tri- lobed with a medial large lobe and two lat- eral, blunt tips (Fig. 2), widening from the tip of rostrum onwards. The lateral ridges begin from the rostral apex running above PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the ocellus and the eye until it reaches the dorsal margin of the headshield. The head pore is large and subcircular. The ventral margin of the head is even, slightly convex with a transversal squamose pattern. The la- brum is cuneiform and continuous with the ventral margin of the head (Fig. 3). The la- bral apex is blunt. Compound eye is located close to the middle distance between the dorsal and the ventral edges. The ocellus is close to the apex of rostrum, from 4.75 to 5.6 times smaller than compound eye. First antenna (Figs. 4, 5) rod-shape (body length: antennule length = 3.4—3.8) insert- ed subapically, barely dilated distally, reaching over half-way to the labrum tip. One basal sensory seta is located ventrola- terally. Distally with a row of relatively long spinules near the lateral side of aesth- etascs insertion. The medial surface has 7 to 9 transverse rows of small spines on the two distal thirds. Lateral and anterior sur- face armed with several rows of minute scale-like spinules plus a subapical group of spines on the posterior side. Nine aesthe- tascs, unequal in length (longest member: shortest member = 2.0—2.2) with a bifur- cated tip, the longest one is 2.6—3.0 times shorter than the antennular length. Second antenna: Stout and long (Fig. 6). Coxa massive, the basal region appears an- nulated with several folds and provided ~ with a row of tiny denticles at the ventral © side; the middle region presents two ven- trolateral, soft sensory setae and the distal one is made up of ventrolateral margin armed with some rows of sclerotized spi- nules. The external surface of the distal por- tion has a spine used for burrowing and a longer soft seta at the medial face extended well beyond the end of the second exopod segment. Swimming setae O-0-1-3/1-1-3, spines 0-2-1-1/0-0-1. All exopod setae car- ry spines at the distal septum except for the innermost apical member. The exopod is characterized by one spine on the distal edge of segments 2 and 4, as in all the Ma- crothricidae taxa (Smirnov 1992), plus an accessory, internal curved spine on seg- VOLUME 110, NUMBER 1 ments 2 and 3, both subequal in length (Fig. 7). The spine is attached to the second seg- ment (2nd segment: spine = 2.2—3.2) which is about 1.8 to 3.0 times longer than the other one. The surface of all antennal seg- ments includes rows of fine scale-like spi- nules. The longest antennal seta (Figs. 8, 9) is on the first endopod segment, and is bi- segmented (distal segment: proximal seg- ment = 1.2—1.6) stout, sclerotized and long (body length:seta length = 1.2—1.4). Its convex margin is provided with a series of fine setules along the three proximal quar- ters of the first segment, followed by a row of stout spines, two or three of which are subequal in length and are attached to the proximal joint (Fig. 8). The remaining row of spines runs along the second segment, decreasing in size gradually toward the tip. Beyond the middle third of the seta, these spines form groups of different number. There is a row of sparse spinules that con- tinues along the next segment on the exter- nal surface of distal quarter of the first seg- ment (Fig. 9). Trunk limb I (Fig. 10): The exopod (ODL) is slender, bearing a long apical seta unilaterally having fine setules along its dis- tal portion, and a short lateral seta (Fig. 11). Endite 4 (IDL) has three setae unequal in length, the longest one with a pattern sim- ilar to seta of ODL, the other two setae pos- ses a row of stout setules, increasing mod- erately in size distally (Fig. 11). The pos- terior surface of IDL is provided with groups of strong setae. There are four setae different in length on endite 3, two of them are more sclerotized and bisegmented. En- dite 2 has three bisegmented setae, setulated along the distal portion and with a row of sparse stout setules through the proximal joint. Endite 1 presents two fine plumose setae (Fig. 10). The inner spines on endite 1 and 2 have two and one teeth respectively on a pointed apex and one lateral spinule (Figs. 12, 13). Trunk limb II (Figs. 14): The exopod is reduced, with an apical soft seta bilaterally 121 setulated. The endopod has a row of eight spine-scrapers, six similar in structure, with an enlarged base, and distally with a pecten of strong, sclerotized teeth. Scraper 4 has the largest and strongest pecten. Scrapers 7 and 8 are long with an armature similar to the seta of ODL from trunk limb I. There is a lobe carrying a stout conical seta dense- ly ciliated on its distal portion, located at the external surface of the endopod, near the insertion of scrapers 4 and 5. The gna- thobase presents a filtering comb composed of four short setae which are different in structure and with four long plumose setae. Trunk limb III (Fig. 15): The exopod car- ries four setae. The two outermost are fine and bilaterally setulated, the third one is densely setulated in the distal portion. The last one is strong and provided with two rows of spiniform setules. The lateral sur- face of exopod is covered with some groups of needle-like setae. Endopod with a distal- most seta stout, sclerotized, with a hook- like shape and proximally followed by five setae, all of them different in structure. Se- tae of the external armature are straight, finely ciliated in the distal portion. The gna- thobase is rounded and it has a setose out- growth on the external surface and two blunt papillae. Trunk limb IV (Fig. 16): There are two apical setae on the exopod, the outermost is bilaterally setulated, the other one is dense- ly setulated at the distal portion. The medial armature consists of five equal setae; the lateral row bears four setae, the distal one is stout and spiniform, the remaining ones are strongly hairy. There is also a tubular sensillum between the gnathobase and the proximal endopod. The gnathobase is re- duced, composed of a setose furry-like seta and a finger-like lobe. Trunk limb V (Fig. 17): Exopod subob- long, reduced to a small flap, with one fine setulose seta. The endopod has a rounded hairy flap and three setae remarkably dif- ferent in size, the two outermost are re- duced and strongly ciliated; the third one is the longest, bisegmented, unilaterally setu- 122 lated along its distal portion. The gnatho- base is composed of three well defined lobes, each one is armed with a series of long setae. The epipodite is large and su- bovoid. Postabdomen (Fig. 18): Large, subovoid in lateral view (body length: postabdomen length = 2.8—3.4). The ventral margin is straight, somewhat convex with one or two groups of minute spinules. The dorsal mar- gin is asymmetrically convex, non-bilobed, however, the preanal region is well differ- entiated from the rest of postabdomen (pre- anal portion: anal-postanal portion = 1.65— 1.90). The preanal region has a dorsal stripe of needle-like spinules arranged in several transversal rows (ca. 25—30) which are sub- equal in length. The anus is bordered by 5 to 7 groups of larger spinules on each side with lateral surface covered with crescent rows of fine setae. The postanal portion has no setae. The claw is heavily sclerotized, evenly curved with a lateral face armed by two rows of small spinules on both concave and convex margins (Fig. 19). The medial side is provided with a row of three contin- uous pectens, obliquely arranged (Fig. 20) The two most proximal are composed of fine setae distributed along two thirds of the claw length, with a large and sclerotized member between them. The distal pecten is provided with a series of short, stout spi- nules, hardly sclerotized, the proximal member is stouter and longer than its com- panions. The setae natatoria arise from a sclerotized heel located at the proximal por- tion of the postabdomen (Fig. 21). The dis- tal segment is shorter than the proximal one (proximal segment: distal segment = 3.0— 4.0), having long, bristle-like setae. Description of ephippial female and ephippium.—Female with two ephippial eggs is about the same shape in lateral view as parthenogenetic female (Fig. 22). The major difference is in cross section (Fig. 23) in which the pointed dorsal median keel of the parthenogenetic female is replaced by a broadly rounded median ridge. Dorsal pro- file of shell is higher than in parthenoge- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON netic female, the maximum height is near the anterior slough line; the egg locule is well developed on each side. The shell sur- face of ephippial region is finely granulated with mesh-like pattern, weak and diffuse at egg locule and dorsal region. Dorsal margin with one row of chitinous rounded papillae. Slough line evidenced by a simple line that follows closely below the ephippium, so the entire anterior ventral portions of shell are lost after molting except for a small pos- tero-ventral portion left in the ephippium. Portion of shell that sloughs has no punctae, meshes or pigmentation. Most of ephippial region is yellowish brown, mainly around the dorsal and the egg locule regions. Description of males.—Two different prereproductive instars (Figs. 24, 25) and only one mature instar (Fig. 26). The func- tional male has a height somewhat lower than in female (length: height = 1.50-1.80). There are some appreciable ontogenetic changes over the three instars in shape of head. The rostrum of adults is directed more anteriorly than in immature instars, so the general outline of body has an attenu- ated ovoid aspect in lateral view. The two immature instars have a general shape sim- ilar to females (length: height of instar I and II = 1.45-1.70). Antennula in instar I has a basal sensory seta and the setulation of internal surface is about as in female (Fig. 27). In instar II the antennula increases in size, and it has an additional sensory seta near its base, which arises from a rod-like projection, proximal to a characteristic group of long setae on the proximal internal portion (Fig. 28). In the adult male, the antennula is highly mod- ified and is curved inwardly (Figs. 29, 30). The two basal sensory setae are well de- veloped. One originates at the anterior mar- gin and the other is located behind the for- mer and close to the posterior margin. The latter is long and thin while the first is thicker and arises from a rod-like projec- tion. On the external surface near the mid- dle of antennular length (Fig. 30), there is a long soft seta (antennular length: seta VOLUME 110, NUMBER 1 length = 2.0-2.4). It is evident from instar II. Proximally, on the medial surface, close to the rod-like projection, there is a group of long hair-like setae and three groups of spinules, the distal and proximal ones are made up of sparse minute members, while the medial one has long and stronger spi- nules (Fig. 29). Nine aesthetascs in all in- stars; instars I and II aesthetascs with bi- furcated tips as that in female, but in instar III this character is lost or barely visible. There are two aesthetascs 2.25 to 3.3 times longer than the others, these latter ones are subequal in length (Fig. 30). Trunk limb I with typical instar sequence of development of copulatory hook (Figs. 31-33); adults with a long, stout free por- tion, with one crescentic ridge at tip of ir- regular margin that gives a spoon-like ap- pearance to this portion (Fig. 34). IDL of adult male with three setae shorter than in female, the shortest one modificated as a spine-like seta, naked and hardly sclero- tized. ODL with a large seta provided with small tubercle-like setules along its distal portion (Fig. 33). Postabdomen (Fig. 35) roughly similar in shape to that of female although smaller; ventral margin tends to be irregular, in part because of the presence of genital pores, which in mature male open near ventral midline at a notch, located about two basal postabdominal claw lengths from tip. The postabdominal claw is the same as in the female, except that it is relatively longer and somewhat stouter. Differential diagnosis and relation- ship.—Macrothrix smirnovi is a member of the M. triserialis-group. It shares with this Species-group the general shape of the body, structure of the antennulae, the post- abdomen, and the largest seta of the anten- na. M. triserialis Brady was described from Sri Lanka. Fryer (1974) and Smirnov (1992) mentioned that M. triserialis is a species with wide and complex distribution, but their conclusions were based on only general morphological aspects. Sometimes, finer scaled morphology has been used F233 (e.g., Fryer 1974, Dumont & Van de Velde 1977, Korinek 1984), but the differences observed were considered as variations within a single species rather than differ- ences between species. Through detailed morphological comparisons Smirnov (1976, 1992) and Brandorf et al. (1982) distin- guished three separated geographically iso- lated species from this taxa-group: M. gau- thieri from Africa, M. superaculeata from Brazil and M. flabelligera from Australia. In addition M. rosea (Jurine) is mentioned by Dumont & Van de Velde (1977) as a possible synonymy of M. triserialis. This species differs from M. triserialis s. Str. mainly because it lacks the two stout, hook-like spines on IDL of limb I, clearly shown by Fryer (1974: fig. 69) and pointed out by Korinek (1984); instead of hooks, these spines are not curved and they seem to be less strong, both armed with rows of setules (Fig. 11). The combination of char- acters suggests a closer relationship of this new species to M. superaculeata (Smirnov) described from Brazil and M. rosea (Jurine) from Central Europe than to the other spe- cies of the group. Parthenogenetic females can be easily distinguished from both relat- ed taxa, mainly because of the antennal for- mula, the structure and armature of postab- domen and some fine details of trunk limbs (Table 1). M. smirnovi, M. capensis (Sars), M. paulensis (Sars) and M. gauthieri Smir- nov have accessory spines on the antennal exopod, but M. capensis can be distin- guished by a hump-like protuberance on the ventral part of the head and the preanal re- gion with very small spinules. M. paulensis and M. gauthieri as well as M. odiosa Gur- ney and M. sioli (Smirnov) differ from M. smirnovi because of the large spines along the antennulae and the structure of postab- domen (Brandorf et al. 1982, Smirnov 1992). Detailed morphology of mature males also shows evident differences among ™M. triserialis-group species. There are no de- scriptions of M. triserialis males on popu- lations from type locality (Sri Lanka), the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 124 ee eee nen en ee ‘JUDUIZIS [eISIP SB SUC] Se SOWT) JOIN], ‘Ios1e] pue JosuoNs AJPAeJOI ore SMOI TeUITXOId jsouI ay) ATUO ouy puv [jeuIs ore sajnuids Jo sMOI yons "ae19S OM} YUM podox| ‘soynurds jjeuis Jo 33m) e Aq pamoyTjoj 4100} Josie] yeordeqns ev YIM ¢ pue p JodeIOS ‘eJOS pue aqo] YONs ON “AJOAN -oadsai sojnuids [eloyey suo puke OM} ‘xode pajutod uo 4300} suo yyIM sourds 1duUI YIOg ‘A][eISIP ozZIs ut ATAATIOUTISTIp SUISVIIOUI SO[NJOS JO MOI & YIIM 9ej}OS dso], ‘sourds [[euls [e -1aA0s Aq poMoyjo} AT][eISIp ‘s}uoUIsas [eISIp pue jeurxold sjt uaeMjoq juIOf ay) Jo vole oY} UI souIds IdSIe] OM} YIM BjoS JsOSIeT 1-0-0 /T-0-1-0 seurds ‘sotnuids Suoms [eordeqns on ‘uloned ssouenbs esioasuey APYSIS & WIM UISIeEW [eIUDA ‘uoTWBLIAas OU Sey o3po [esI0q ‘JUSS [e}SIP SB SUC] SB SOW) C'O[-N' [|] JWOUIZeS [eWITxOIg “A][ewtxoid ozIs ul SUISPOIOUI SUOIS O1e sopnuIds Jo smo YONS ‘UOT}BULIOJUI ON ‘so[nulds |[euls Jo 1n} Be Aq poMmoTo} 4100} Josie] [eoideqns & YIM ¢ pue p JadeIDS "eJOS pue oqo] Yons ON ‘aInuids [e1oye] suo pue xode pojutod uo 490} OM} YIM soulds IoUUT POG —+Aypeasip ozis ut Ajaanounstp BSUISESIOUI So[NIdS JO MOI B YIM JeIOS BSOUT, ‘yurof s}t Jo Aj[eIstp sourds [ews [ero -AdS pue oS1e] & pue ‘juIof sjI 0} [euTxoId duids os1e] & pur [Tes B Sey LOS JSOBIeT 1-0-0 /I-I-Z-1 soutds sornu -Ids poziunolejos pue suonNs ¢—7 jo dnois B UJIM SopIs JOLIojue pue JolIa\}sod yIog ‘quids & YIM poulre ouO Yyove sospi oyI][-dojs [eSIOASURI] YIM UISIeU [eIUO A ‘uOTeLIOs ou AIOA sey d3po [esIog ‘(IZ “Btq) uourses [eysIp Sv SUC] SB SOUT) O'p—-O'E JUSWIZES [eUITXOIg ‘(QI ‘SI4) OzIs ul yenboqns so[nuids [Jews JO SMOI YA UOTZoI [eueoIg ‘(LI ‘814) 818s 9UO YIM podox| ‘(pI “314) YISUs] OUIeS JO 4190) YIM ¢ puke p SIodeIDS ‘(pI “S3I4) ¢ pue p siodeios reou po -JB90| ‘UOIZ9I [eISIP S}I UO Payer]IO ATosuop *eJOS [BOIUOD jNOoJs & SUTALIVD oqo] SATIOUTISIG ‘(EL ‘ZI ‘S8t4q) curds [e1aye] suo pue xode pajutod ay} uo AjoATjoedsa1 yI90) UO pue OM} SUIARY Z pue | SopuS UO souTds JOUUT “(IT Sta) Aj[e3Stp ozIs ur AToyeIopour SursvoroUI ‘soyn) -9S JO MOI B YIM OM} JsopIOYsS OY} ‘yIZUST jUSIOJJIP JO BIOS Som) YIM (TqO) pv oupug (6 “3tq) JUSUISOS [eISIP oy) UO dy oy) preMO} OZIS UI SUISvoIDOp souTds [BIOADS JO SOLIOS B pue ‘(g ‘31,) quIOf sz 0} [ewTxoid yZu9] UT Jenbagns soutds 9014) 0} OM) Sey BIOS JSOSIeT ‘(L ‘9 S814) 1-0-0 /I-1-Z-0 seutds "(¢ “BIq) soqnurds €-z jo dnois [eordeqns e YIM SpIs 1O1IN}so0g ‘(€ °*31,{) wioyed ssourenbs as19A -suel) AYSI[S & YIM UdAD UISIeUI [eIUDA ‘UOT}eLIOS OU SPY dBPod [esIOG B1IO}JEJLU Je}OS jeutwmopqeisog usulopqeisog A qui] yuniy II quay yunsy, J qui] yunsy, seuuguUyy se[NuUusJuUy peoH 1°4$ ,02SOd “WW ,DJpanovsadns “pw g[ewisy DjoUssOusyIed sinjePy so1seds MoU ‘7AOUMIUS XIUYJOLIDW Jajoereyo ‘(ouLING) Vaso’ "FW pue (AOUIIUIS) DJvajnovA1adNs ‘PF poyefel 94} YIM so1deds MoU ‘MAOUMUS X1IUYJOAIDPF JO uosteduicD— | IQR 125 VOLUME 110, NUMBER 1 JO SMOI OM] O} DUO YIM UISIeU [esIOGg ‘POZIUNOIJO[OS A][IABVOHY ‘pojetnuess AjTasreoo DoRJANS ‘UOTJ[NIOI JO seT[OWIR] Yor) yA ‘UOISOI S[Pprw oy} 1e BJaS JJOS SUC] B pu BIAS AIOSUaS [eESeq 9UC ‘POZIUTJOIO[IS SSo] ‘soTeOS UT pesueue jou so[nurds jo siajsnjo re[nuuajUy “WIN]][ISuas [eseq noyM pur oje[d oyI]-o]eOs [eordeqns INOYWM [BOIUOS UsUTOpQe}sod jo puod [eIsIq "eT [OUI -e] [eotdeqns ouo ATuo YIM Suttiade ‘1apusts ‘(p861) YouLIOy ur eyep poystiqnd woj poureyqo wmniddiyde pur sayeu oimjew uO UONPULIOJU] , ‘IO[OO UMOIG YyIeq ‘oeTIded papuno1 ‘(€Z ‘8I4) IOJOo uMOIq “IO[OD MOT[OX “poz -TUO1a]OS A[qQ99,J ‘S][29 pepunol Jo oimjdjnog uiniddiydq ‘aejos AlOsuas [eseq OM], ‘sg—nutds Suons Jo siaysnjo POZIUTOIO[OS ATIAVOY YJIM POI9AO0O apis JOUU] “"WIN][ISsuds [eseq pue aed aJe[nonuep [eordegns syI]-97eos WIM ‘uorjoefoid repnqn) oyur AT[eIsIp Surpusixq "S$]So19 [esiaAsuey Teordeqns sey) SutAues ‘3uons YSIMOT[EX ‘oep[ided pepunoi jo mor auo ATUO YUM UISIeUI [eSIOG “‘PpezIUToJa[Os ATIAROY ‘uloyjed oyI[-Ysou & YIM poyefnueIs Ajour7 ‘(O€ ‘SIq) UOTSe1 o[pprur ay} Iv BIOS Os Sug] & pur ‘uoNdefoid oyI][-por & Wor Seslie YOIYM JO 9U0 ‘avjos AlOSUaS [eseq OM], ‘(87 ‘S[4) d0RjINs JoUUT oy) uo Ajfewitxoid sojnurds jo sdnoizs sory) pue So[Njes oyI]-siey suo] Jo dnois ve ore as10y) peojsur saynurds Jo siojsnjo Ou ore a1dy], ‘(CE “Steq) oyeUloJ UT Jey) 0} Ie[IUIIs JeYMouIOS odeys ‘(re “Stq) souvieodde oyt]-uoods ev uoniod sty) 0} Sur -A13 dy ye BSplI JUSSI oUO SuUTARY ‘SUONS ejnuuguy uswiopqrisog yooy A1oyeindoa yDASOd “PW yDIDajnovAadns “pw saideds Mou ‘Moudus x14YJoAIDP Jajovleyo soyeul oinjeyy a ee ee ee ae ee ee ae = ‘ponunuog—'] 21qeL 126 only published data available is the M. ele- gans Sars. The description, which is a young synonym of M. triserialis according to Smirnov (1992), is featured by a long cylindrical distal part of postabdomen, no claws and a copulatory hook of trunk limb I with three transversal crests at tip. There are no described males of the related M. superaculeata from the type locality, but Korinek (1984) analyzed a population from Cuba that he assigned to this taxon, where males may be distinguished by the presence of three transversal crests at tip of copula- tory hook and postabdomen, extending dis- tally into a tubular projection, with a sub- apical scale-like plate and without any claws. M. rosea males are also different since there are a conical postabdominal dis- tal part (Werner 1927) with terminal claws (Korinek 1984) and a slender copulatory hook tapering distally with only one sub- apical lamella. Macrothrix rosea resembles the ephip- pial structure and reticulation of M. smir- novi, which is characterized by a mesh-like pattern on the surface and differs because the dorsal margin of the former has only one row of rounded papillae whereas M. ro- sea includes from one to two rows. How- ever, the sculpture is markedly different in M. superaculeata which possesses rounded cells; itis feebly sclerotized and lightly col- ored (Korinek 1984). There are five species of this genus re- ported from Mexico including M. smirnovi as well. The Mexican cladoceran fauna, as was monographed by Frey (1982b), includ- ed only two species: M. laticornis (Jurine) and M. rosea (Jurine). Recently, this num- ber has increased because of the record of M. triserialis s.1. (Ciros & Elias 1995) and by the description of M. mexicanus (Ciros et al. 1995). However, it is worthwhile to point out that current knowledge of the tax- onomy and geographical distribution of Macrothrix in this region is still scarce and some old reports should be analyzed with caution. It is possible, according to the improve- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ment in research on macrothricid cladocer- ans, that some species known in the past as cosmopolitan would be geographically re- stricted as was demonstrated by Frey for diverse chydorids. This statement is vindi- cated because macrothricids are the cladoc- erans most closely related to chydorids (Frey 1988). Future analysis should be based on larger samples containing ephip- pial females, males and females in all in- stars. Acknowledgments We are indebted to Prof. Nikolai N. Smirnov for providing specimens of some Macrothrix species from his collection, and for reading the manuscript and checking our English. We are also thankful to Prof. H. J. Dumont for the permission to check specimens of M. triserialis from his collec- tion; to CONABIO (Mexico) for financial support (grant H-112). Finally we would like to thank to Dr Jgrgen Olesen and to an anonymous reviewer for checking and im- proving our manuscript. Literature Cited Brandorff, G. O., W. Koste, & N. N. Smirnov. 1982. The composition and structure of rotiferan and crustacean communities of the lower Rio Nha- munda, Amazonas, Brazil.—Studies on Neo- tropical Fauna.and Environment 17:69—121. Ciros-Perez, J., & M.-Ehas-Gutierrez. 1995. Nuevos _registros. de cladéceros (Crustacea: Anomopo- da) en México.—Revista de Biologia Tropical 44:297-304. , M. Silva-Briano, & M. Elias-Gutierrez. 1995. A new species of Macrothrix (Anomopoda: Ma- ‘crothricidae) from central Mexico.—Hydrobiol- ogia 319:159—166. Dumont, H. J. 1994. On the diversity of the Cladocera in the tropics.—Hydrobiologia 272:27-38. , & I. Van de Velde. 1977. Cladocéres et Con- chostracés récoltés par le professeur Th. Monod dans la moyenne du Niger en décember 1972 et janvier 1973.—Bulletin de LInstitute Fran- cais D’ Afrique Noire 39:75-93. Frey, D. G. 1982a. Questions concerning cosmopoli- tanism in Cladocera.—Archiv fiir Hydrobiolo- gie 93:484-502. . 1982b. Cladocera. Pp. 177—186 in S. H. Hul- bert & A. Villalobos-Figueroa, eds., Aquatic bi- VOLUME 110, NUMBER 1 ota of Mexico, Central America and the West Indies. Aquatic Biota SDSU Foundation Vol XV, San Diego University Press, 529 pp. . 1986. The non-cosmopolitanism of chydorid Cladocera: implications for biogeography and evolution. Pp. 237—256 in R. H. Gore and K. L. Heck, eds., Crustacean biogeography. Bal- kema, Rotterdam. . 1988a. Are there tropicopolitan macrothricid Cladocera?—Acta Limnologica Brasiliensia II: 513-525. 1995. Changing attitudes toward chydorid anomopods since 1769.—Hydrobiologia 307: 43-55. Fryer, G. 1974. Evolution and adaptive radiation in the Macrothricidae (Crustacea: Cladocera): a study in comparative functional morphology and ecology.—Philosophical Transactions of Royal Society, London Biological Sciences B 269:137-274. Haney, J. EF, & D. J. Hall. 1973. Sugar-coated Daph- nia: a preservation technique for Cladocera.— Limnology and Oceanography 18:331-—333. 127 Korinek, V. 1984. Cladocera. in Hydrobiological sur- vey of the lake Bangwelulu Luapula river basin. Scientific results 13, fasc 2, Cercle hydrobiol- ogique de Bruxelles, Bruxelles, 117 pp. Sars, G. O. 1901. Contributions to the knowledge of the freshwater Entomostraca of South America, as shown by artificial hatching from dried ma- terial. Part I: Cladocera.—Archiv for Mathe- matik og Naturvidenskab Kristiania 23:1—102. Smirnov, N. N. 1971. Chydoridae of the world’s fau- na. Fauna of the USSR. New series No. 101. Crustacea. 1(2), Leningrad, 531 p. (English transl. by A. Mercado. Israel Program for Sci- entific Translations, Jerusalem, 1974). 1976. Macrothricidae and Moinidae fauna mira. Fauna SSSR, novaya seriya, N 112. Rkoobraznye T.I,3 Leningrad, Nauka. 237 pp. . 1992. The Macrothricidae of the world. SPB Academic Publishing, The Hague, 143 pp. Werner, C. EF 1927. Wachstum und Formentwicklung der Cladocere Macrothrix rosea.—Wilhem Roux Archiev Entwicklungsmechanik Organis- men 109:241—252. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):128-142. 1997. Some deep-sea Pycnogonida from the Argentine slope and basin C. Allan Child Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Five stations from the Argentine Slope and Basin in the South Atlantic resulted in a rich collection of thirteen pycnogonid species, the ma- jority new to that area of the Atlantic Ocean. Ten of the thirteen species are from the family Nymphonidae and four of these are new: Heteronymphon caecigenum, Nymphon centrum, N. dentiferum, and N. vacans. Distributions are given for all species along with pertinent remarks. The new species are described, illustrated, and compared with likely congeners. One additional spe- cies, Nymphon scotiae, is illustrated to contribute to knowledge of its mor- phology where the type figures were diagramatic and not entirely accurate. This small collection was omitted from a previous report on deep-sea pycnogonids from the North and South Atlantic Basins (Child 1982) because it was not known to exist at that time and did not come to my attention until many years later. It consists of five additional stations made by the At- lantis IT, of the Woods Hole Oceanographic Institution, in the South Atlantic off Argen- tina. None of these stations is duplicated in the previous report listed above. Seventeen species were listed in the first report from the Argentine Basin with six species and one genus described as new of the seven- teen (35% new). It was noted (Child 1982: 1) that Argentine Basin fauna is poorly known. Knowledge of the Basin’s fauna is only slightly improved with the addition of the thirteen species listed in this report, four of which are described as new (31%) and four additional species recorded for the first time herein from the Argentine Basin. The new species are; Heteronymphon caecigenum, Nymphon centrum, Nymphon dentiferum, and Nymphon vacans. Known species re- corded for the first time in the Argentine Basin and slopes are; Nymphon longicollum Hoek, N. inferum Child, N. scotiae Stock, and a questionable specimen of Colossen- deis scoresbii Gordon. The other five spe- cies are either cosmopolitan deep water spe- cies or have apparently invaded this basin from nearby deeps or Subantarctic deep- seas. Family Ammotheidae Dohrn, 1881 Genus Cilunculus Loman, 1908 Cilunculus acanthus Fry & Hedgpeth, 1969 Cilunculus acanthus Fry & Hedgpeth, 1969:126—127, figs. 207—209.—Stock, 1978:197 [key].—Child, 1982:9; 1994a: 34. Material examined.—Slope E of Cabo San Antonio, Argentina, 36°55.7’S, 53°01.4'’W, 2707 m, sta. 245A, epibenthic sled, 14 Mar 1971, 1 6 with eggs, 1 6,5 2, 4 juv. Distribution.—The species is apparently confined to the Scotia Sea and Argentine basin, based on what few records there are of captures. The type was collected in the Drake Passage. This capture extends the distribution of C. acanthus a little to the north in the Argentine Basin where it has been reported (Child 1982) and adds noth- ing new to a known depth range of 2450— 2818 m. VOLUME 110, NUMBER 1 Remarks.—This is a rare species, but perhaps only scarce because of the few deep-sea collections which have come from the Argentine basin. It is rare in the Scotia Sea where many more trawl samples have been made and from which many pycno- gonids are known. Members of this genus are more commonly found in temperate regions. The distinctive truncate conical cement gland tube and delicate slender dorsomedi- an tubercles easily identify this species among the many rather plain Subantarctic pycnogonids known from the Scotia Sea. Family Austrodecidae Hodgson, 1907 Genus Pantopipetta Stock, 1963 Pantopipetta longituberculata (Turpaeva, 1955) Pipetta longituberculata Turpaeva, 1955: 324-327, fig. 2. Pantopipetta brevicaudata Stock, 1963: 336-338, figs. 9, 10a.—Hedgpeth & McCain, 1971:219, fig. 1E, 220, table 1, 222 [key], 223-225, figs. 3, 4, table 3. Pantopipetta longituberculata-complex.— Stock, 1981:465—466 [text]. Pantopipetta longituberculata.—Child, 1982:49—50 [literature]; 1994b:88-89, fig. 17. Material examined.—Basin E of Mar del Plata, Argentina, 38°16.9'S, 51°56.1'W, 4382—4402 m, sta. 242, epibenthic sled, 13 Mar 1971, 27 specimens. Slope E of Cabo San Antonio, 36°55.7'S, 53°01.4'W, 2707 m, sta. 245A, epibenthic sled, 14 Mar 1971, 30 specimens. Basin E of Valdez Peninsula, 43°33.0'S, 48°58.1’W, 5208-5223 m, sta. 247, epibenthic sled, 17 Mar 1971, 30 spec- imens. Basin E of Mar del Plata, 37°40.9’S, 52°19.3'W, 3906-3917 m, sta. 256, epi- benthic sled, 24 Mar 1971, 16 specimens. Basin E of Cabo San Antonio, 37°13.3’S, 52°45.0'W, 3305-3317 m. sta. 259, epi- benthic sled, 26 Mar 1971, 14 specimens. Distribution.—This species is fairly com- mon in very deep waters of the Pacific and Atlantic basins, and appears to be common 128 in the Argentine basin. It has been found as far south as the South Shetland Islands of the Antarctic. It has one of the broadest depth ranges of any known pycnogonid: 567—6700 m. Family Colossendeidae Hoek, 1881 Genus Colossendeis Jarzynsky, 1870 Colossendeis ?scoresbii Gordon, 1932 Colossendeis scoresbii Gordon, 1932:18— 2ie os. SC. 60. Ob , Oc. OC. 7a. b= Child, 1995b:90, fig. 8. not Colossendeis megalonyx scoresbii Fry & Hedgpeth, 1969:18 [key], 32, 33, figs. 7, 8, 17-20, 23. Material examined.—Basin E of Mar del Plata, Argentina, 38°16.9’S, 51°56.1’'W, 4382—4402 m, sta. 242, epibenthic sled, 13 Mar 1971, 1 specimen. Distribution.—The species is only known from a few stations generally north of the Falkland Islands in 128—303 m. This specimen extends this distribution to the northeast and into very much deeper wa- ters. Remarks.—This small specimen is pos- sibly not C. scoresbii due to the gross depth differences in this capture and those of the type specimens and the records of Child (1995b:90). This specimen also apparently lacks eyes. They are not discernable in the rather tall ocular tubercle. The ninth palp segment is shorter than usual for C. scores- bii, and the tarsus and propodus are longer in relation to the long claw of the Atlantis IT specimen. It is closer to C. scoresbii than to any other known species. It has similarity to C. angusta Sars in the short proboscis, although C. angusta has a shorter proboscis in relation to trunk length. The ninth palp segment of C. angusta is also longer. It is as long as the terminal seg- ment while the ninth segment of this Atlan- tis IJ specimen is shorter than the tenth. This specimen is possibly a new species but in light of the great variation known to most species in this large genus, this single spec- imen must remain as a doubtful record of 130 C. scoresbii until more specimens are col- lected from this position in the South At- lantic. Family Nymphonidae Wilson, 1878 Genus Heteronymphon Gordon, 1932 Heteronymphon caecigenum, new species Fig. 1 Material examined.—Basin E of Valdez Peninsula, Argentina, 43°33.0’S, 48°58.1’W, 5208-5223 m, sta. 247, epibenthic sled, 17 Mar 1971, 1 ¢, holotype, USNM 234719; 13,4 9, 1 juv., paratypes, USNM 234720. Distribution.—Known only from the Ar- gentine basin, in 5208-5223 m. Description.—Size moderately small, leg span about 22 mm. Trunk fully segmented. Trunk and lateral processes smooth, gla- brous, segments fully articulated, well sep- arated by intervals equal to their diameters or slightly greater. Lateral processes slightly longer than their diameters. Ocular tubercle and eyes lacking, but hump containing lat- eral sensory papillae at anterior of cephalic segment suggests a low ocular tubercle. Oviger implants large, placed just anterior to but not touching first lateral processes. Neck short, not as long as wide, moderately expanded at anterior. Proboscis typical, gradually tapering to rounded oral surface, not as long as chelifore scapes. Abdomen short, distally a truncate cone with later- odistal pair of short setae. Chelifores slender, scapes slightly longer than proboscis, armed laterally and distally with row of short setae. Chela palm as long as fingers, armed with many short lateral and ventral setae. Fingers robust, sharply curved distally, armed with 13 short re- curved teeth on movable finger and ten sim- ilar teeth on immovable finger. Palp segments moderately short, only slightly longer than chelifore scapes, typical for genus. Second segment short, only 0.6 as long as third, the longest segment. Fourth slightly shorter than second, fifth 0.25 lon- ger than fourth, distal three segments with few very short distal and ventral setae. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Oviger (female) fourth segment subequal to fifth, sixth about 0.6 as long as fifth, all with few short lateral setae increasing in numbers on distal segments. Strigilis distal segments increasingly shorter than those more proximal, each armed with three-four short ectal setae and short endal denticulate spines in formula 6: 5: 5: 6, with terminal claw lacking teeth, hardly longer than distal denticulate spine. Spines with four lateral lobes per side. Legs moderately slender, with few short dorsal and ventral setae, only three-four dorsal setae longer than segment diameters. Second tibia the longest segment, with first tibia slightly longer than femur. Tarsus about 0.75 length of propodus, both slightly curved, armed with row of very short sole Spines and few short dorsal setae. Claw half propodal length, robust, moderately curved, auxiliaries lacking. Male characters: slightly smaller size, femoral cement glands not evident, oviger fifth segment almost twice length of fourth. Measurements (holotype in mm):— Trunk length, 2.46; trunk width across 2nd lateral processes, 1.1; proboscis length, 0.83; abdomen length, 0.63; third leg, coxa 1, 0.32; coxa 2, 1.0; coxa 3, 0.41; femur, 1.78; tibia 1, 2.03; tibia 2, 2.59; tarsus, 0.83; propodus, 1.08; claw, 0.53. Etymology.—The name (Latin, caecigen- us, meaning born blind) refers to the lack of any evidence of eyes in the new species. Remarks.—This is the third known blind species in the genus Heteronymphon, the first being H. profundum Turpaeva, 1956, and second, H. abyssale (Stock, 1968). The latter species was moved from the genus Nymphon (Turpaeva, 1970:1723) because the sensory papillae commonly associated with an ocular tubercle are situated at the extreme anterior of the cephalic segment, suggesting that an ocular tubercle would be found there had one existed in H. abyssale. Most members of this genus live in deep waters and have inconspicuous. or not fully formed eyes which are unpigmented as be- fits a lack of need for eyes in those habitats. VOLUME 110, NUMBER 1 131 4 eve }. — . -. a a date SSA shen e™ — — Fig. 1. Heteronymphon caecigenum, new species, holotype: A, Trunk, dorsal view; B, Trunk, lateral view; C, Third leg; D, Oviger, with enlargements of terminal segment and denticulate spine; E, Chela. This new species and the two previously genus Nymphon is the placement of the oc- known have progressed to a fully blind sta-__ular tubercle and its associated sensory pa- tus, going beyond the partly formed but un-__ pillae. These are found in the extreme an- pigmented eyes of other species. The prin-_ terior of species in the genus Heteronym- cipal difference between this genus and the phon but are further posterior in species of 132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON B cry ites POU SURLY CERN cays Jar ad eee Sa Ae Sale sees ToS Fig. 2. Nymphon centrum, new species, holotype: A, Trunk, dorsal view; B, Ocular tubercle from right side; C, Third leg; D, Palp; E, Chela; EK Oviger; G, strigilis, enlarged. Nymphon. Where there are no eyes and the Genus Nymphon Fabricius, 1794 ocular tubercle is only a morphological sug- Nymphon centrum, new species gestion, the sensory papillae must exist in Fig. 2 the location where there would be an ocular tubercle if one existed. With these three Material examined.—Slope E of Cabo species, the papillae are anterior as in the San Antonio, Argentina, 36°55.7’S, other members of Heteronymphon, strong 53°01.4'W, 2707 m, sta. 245A, epibenthic evidence that they belong in that genus. sled, 14 Mar. 1971, 1 6 holotype, USNM VOLUME 110, NUMBER 1 234715; 3 3 with eggs, 6 6, 7 2 ovig., 4 2, 10 Juv. paratypes, USNM 234716. Distribution.—Known only from the type-locality in 2707 m. Description.—Size moderately small for genus, leg span 23.5 mm. Trunk fully seg- mented, unadorned. Lateral processes slightly longer than their diameters, sepa- rated by intervals equal to their diameters or less, each armed with single long me- diandorsal spine, spines often missing. Ovi- ger attachment lobes against first lateral processes, filling most of short neck. Ocular tubercle a slight bulge only, directly dorsal to oviger bases, sensory papillae not evi- dent. Proboscis short, slightly tapering dis- tally. Abdomen very long, extending well beyond first coxae of fourth leg pair, slight- ly inflated medially, armed with six short dorsolateral setae. Chelifores large with wide cylindrical Scapes armed with fringe of distal setae and one proximolateral seta. Chelae only mod- erately curved inward, palm shorter than fingers, armed with two-three setae. Fingers strongly curved distally, overlap at tips, armed with 33 short pointed teeth on mov- able finger and 24 slightly larger pointed teeth and six-seven ectal setae on immov- able finger. Palps moderately short. Second segment longest, with few distal setae, third segment about 0.75 length of second, with few distal setae, fourth/fifth subequal, short- er than third, armed with many scattered setae longer than segment diameters. Oviger segment four well curved, longer than first three combined. Fifth almost twice length of fourth, slightly curved, armed with row of scattered ectal setae as long as segment diameter. Sixth segment short, armed with similar row of ectal setae. Strigilis segments increasingly short distal- ly, armed With ectal setae and few dentic- ulate spines in formula 6: 3: 3: 3:, the spines with one or two lateral lobes. Ter- minal claw slightly curved, longer than ter- minal segment, armed with seven endal teeth and one distal ectal tooth. Legs long, slender. First coxae with long 133 middorsal spines matching those of the lat- eral processes. Second coxae longer than usual, about 2.5 times length of first and third. Femora with small dorsodistal tuber- cle bearing long spine. Cement glands prox- imoventral, with five-six tiny internal bulbs each with external pore. First tibiae the lon- gest of major segments, armed with several dorsal and lateral tubercles bearing long spines, with few other shorter spines. Sec- ond tibiae with shorter spines only and row of short ventral setae. Tarsus slightly longer than propodus, both armed with row of short sole spines. Claw well curved, about half length of tarsus. Auxiliaries lacking. Female characters: size slightly larger ex- cept for ovigers in which segment four is subequal to segment five. Strigilis with few more denticulate spines. Measurements (holotype in mm).— Trunk length, 2.64; trunk width across 2nd lateral processes, 1.46; proboscis length, 0.91; abdomen length, 1.3; third leg, coxa 1, 0.46; coxa 2, 1.03; coxa 3, 0.44; femur, 21012) tibtays1)2:52: tibiav2; 2:34 tarsus, 0.95; propodus, 0.82; claw, 0.46. Etymology.—The name (Latin: centrum, a noun in apposition, meaning center or the midpoint of a circle,) refers to the large middorsal spines placed centrally on each lateral process and first coxa. Remarks.—A member of the N. australe- group (Child 1995a), this new species com- pares with those few species of the group that are blind and uniunguiculate. It fits into the group key (Child 1995a:6—7) next to N. compactum Hoek, and has some similarities with N. hampsoni Child, also from the Ar- gentine basin, and N. inornatum Child, from the Antarctic Weddell Sea. It is prob- ably closest to N. compactum, but has a very different oviger morphology from that species. The fifth oviger segment of N. compactum is distally inflated and the sixth segment is much longer than that of this new species. The chelifore scapes of N. compactum are much longer and the chelae have many more teeth than in this species. N. centrum also has middorsal lateral pro- 134 cess spines which, although common in this group, are almost always in groups or only placed dorsodistally. N. hampsoni has sim- ilar lateral process spines, but they are in pairs and placed dorsodistally. It also has an ocular tubercle mound with conspicuous sensory papillae, longer chelifores, an ovi- ger similar to that of N. compactum, and it has vestigial auxiliary claws. This new species has a general habitus similar to that of N. inornatum. However, that species has an ocular tubercle almost twice as long as its diameter, palps with dif- fering segment lengths, legs with more and longer major segment spines, and other small differences. Nymphon dentiferum, new species Fig. 3 Material examined.—Basin E of Mar del Plata, Argentina, 37°40.9'S, 52°19.3'W, 3906-3917 m, sta. 256, epibenthic sled, 24 Mar 1971, 1 2 holotype, USNM 234721; 1 2 ovig., 4 juveniles, paratypes, USNM 234722. Basin E of Valdez Peninsula, Ar- gentina, 43°33.0’S, 48°58.1'W, 5208-5223 m, sta. 247, epibenthic sled, 17 Mar 1971, 15 @, 5 juveniles, paratypes, USNM 234723. Distribution.—Known from its type-lo- cality, off Mar del Plata, Argentina, in about 3900 m, the new species was also collected E of the Valdez Peninsula in about 5200 m. Description.—Moderate sized, leg span about 48 mm. Trunk ovoid, fully segment- ed, lateral processes separated by about their diameters, moderately short, glabrous. Neck long in relation to most other Nym- phon species, oviger implants halfway be- tween first lateral processes and chelifore insertion, anterior to rounded hump repre- senting ocular tubercle which is just ante- rior to first lateral processes. Sensory pa- pillae of ocular tubercle prominent, on el- evated surface, nipple-shaped. Proboscis cylindrical, very slightly inflated at tip, lips flat. Abdomen short, not extending to tip of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON first coxae of fourth leg pair, armed with four short dorsodistal setae. Chelifores large, scapes short cylinders armed with two lateral setae distally. Chelae long, slender, palms slightly longer than scapes, fingers longer than palms, well curved, overlap at tips, armed with 26 slen- der sharp teeth on movable finger, 17 longer curved teeth on immovable finger, and one short seta at movable finger base. Palps rather long in relation to proboscis, slender, very lightly armed with few short setae, third segment slightly longer than second, fifth about 0.25 longer than fourth, distal two segments longer in combination than third. Oviger (female) fourth segment little longer than third which has small lateral bulge proximal to midpoint. Sixth segment about 0.6 length of fifth. Strigilis segment seven slightly longer than eighth which is subequal to ninth and tenth. Denticulate spines in formula 8: 5: 5: 6. Spines very short except for one distal spine of each segment which is twice longer than those more proximal, with more lateral serrations. Terminal claw longer than terminal seg- ment, slender, well curved, armed with ten very long sharp teeth. Legs robust, moderately long, major seg- ments armed with rows of slender dorsal and lateral setae, some longer than segment diameters. Second coxae distally inflated, with large prominent ventrodistal sex pores. Femora of ovigerous female moderately swollen in proximal 0.75 of their lengths. Second tibiae longest, more slender than first tibiae or femora. Tarsus about 0.6 prop- odus length, both of equal diameter, slender, curved, armed with row of very short ven- tral spines and row of slightly longer dorsal setae. Claw long, slender, slightly curved, about 0.25 as long as tarsus. Male characters unknown. Measurements (holotype in mm).— Trunk length, 5.71; trunk width across 2nd lateral processes, 3.32; proboscis length, 2.46; abdomen length, 0.86; third leg, coxa 1, 0.97; coxa 2, 1.84; coxa 3, 0.96; femur, VOLUME 110, NUMBER 1 135 Fig. 3. Nymphon dentiferum, new species, holotype: A, Trunk, dorsal view, with enlargement of sensory papilla; B, Palp; C, Chela, with enlargement of finger tips; D, Oviger, with enlargement of terminal segment and claw; E, Distal leg segments, enlarged. 5.14; tibia 1, 4.5; tibia 2, 6.09; tarsus, 1.01; propodus, 1.74; claw, 0.24. Etymology.—The name (Latin: dentifer- um, diminutive of dens, a tooth or denticle) draws attention to the enlarged terminal denticulate spine on each of the four stri- gilis segments which is an unusual charac- ter in the genus Nymphon. Remarks.—The strigilis of at least one species of Nymphon (N. floridanum, in Child, 1979:33) was seen to be used as an instrument for cleaning appendages and an 136 enlarged distal spine might offer some ad- vantage in this scraping function. This cleaning or scraping function was shown experimentally by Prell (1910:13, fig. 5B) with specimens of Nymphon leptocheles Sars, 1891. The proboscis, trunk, and abdomen of this species are very like those of N. hadale Child, particularly in the neck length, place- ment of the oviger bulges, and the sensory papillae representing the vestigial ocular tu- bercle. The appendages are each quite dif- ferent from N. hadale. The chelifore scapes of the new species are much shorter than those of N. hadale; and the chelae fingers of that species have many more teeth than the fingers of N. dentiferum. The combined length of the distal two palp segments of N. hadale is shorter than the third segment while the two are much longer than the third in the new species. The oviger strigilis of N. dentiferum has a very enlarged distal denticulate spine en each segment while those of N. hadale are more or less equal in size. Nymphon inferum Child, 1995a Nymphon inferum Child, 1995a:40—42, figs. 12ZA—F Material examined.—Slope E of Cabo San Antonio, Argentina, 36°55.7'S, 53°01.4'W, 2707 m, sta. 245A, epibenthic sled, 14 Mar 1971, 1 9. Distribution.—This species was known only from Subantarctic and Antarctic wa- ters among the island groups on both sides of the Scotia Sea. The nearest specimens to the above record are from South Georgia Island. All are deep-sea and range from 2450 to 3873 m. The Cabo San Antonio record marks a lengthy range extension to the north but adds nothing to its known depths. Remarks.—It is not surprising to find some of the Antarctic species with ranges extending into the South Atlantic. There will probably be some which extend into the South Pacific, but deep-sea collections PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from that distant region are as rare as are the species known from there. This relatively giant species (comparable only to N. charcoti of Antarctica in that re- spect) is difficult to confuse with any other from the Scotia Sea and South Atlantic re- gion. It is blind, uniunguiculate, and with a leg span of 160+ millimeters, its size sur- passes any other Nymphon known among species with these characters. Nymphon laterospinum Stock, 1963 Nymphon laterospinum Stock, 1963:322, 323, fig. 1; 1978a:211, 212, fig. 9a—d. Child, 1982:42—43. Material examined.—Argentine Basin E of Cabo San Antonio, 37°13.3'S, 52°45.0’W, 3305-3317 m, sta. 259, epibenthic sled, 26 Mar 1971, 25+ specimens. Distribution.—This distinctive species is known from the North and South Atlantic basins on both sides of the Mid-Atlantic Ridge and has a wide depth range of 1894— 4715 m. The above record is very near an- other Atlantis IT capture in nearly the same depth. Remarks.—The tall slender lateral pro- cess tubercles of this species along with its characters of uniunguiculate claws and blind ocular mound serve to differentiate it from all other deep water Nymphon, at least in the South Atlantic. Nymphon longicollum Hoek, 1881 Nymphon longicollum Hoek, 1881:40—41, pl. 3, figs. 1-3, pl. 15, fig. 11.—Gordon, 1944:18, table 2, 19 [key].—Stock, 1965: 22 [list, key].—Child, 1995a:43. Material examined.—Argentine basin E of Mar del Plata, 38°16.9’S, 51°56.1’W, 4382-4402 m, sta. 242, epibenthic sled, 13 Mar 1971, 3 2, 4 Juv. Same locality, 37°40.9'S, 52°19.3'W, 3906-3917 m, sta. 256, epibenthic sled, 24 Mar 1971, 1 2, 1 Juv. ; Distribution.—This species was known only from the Chilean basin for more than VOLUME 110, NUMBER 1 one hundred years until many more speci- mens were collected in the Subantarctic is- lands of the South Atlantic and from the Antarctic in the Weddell and Ross Seas. It was also found on the southern New Zea- land slope in 2612 m (Child 1995a:43). It is therefore known in many Southern Hemi- sphere localities in a wide depth range of 508—4069 m. These records extend its range to the north of the Subantarctic lo- calities, into the Argentine basin and in- crease its known depth slightly to 4402 m. Remarks.—The tall, erect, conical, blind ocular tubercle serves as a good recognition character for this species. It is conspicuous, broad based, and its shape is not found on any other known Southern Hemisphere Nymphon. Nymphon longicoxa Hoek, 1881 Nymphon longicoxa Hoek, 1881:38-—39, pl. 2, figs. 1-5, pl. 15, figs. 8, 9.—Child, 1995a:43—44 [literature]. Material examined.—Slope E of Cabo San Antonio, Argentina, 36°55.7'S, 53°01.4’W, 2707 m, sta. 245A, epibenthic sled, 14 Mar 1971, 2 2 without legs. Distribution.—This species has been re- corded from south of New Zealand to the SE Pacific, Scotia Sea, Ross Sea, and South Atlantic off Argentina. It has a known wide depth distribution of 318—3000 m. Remarks.—A diagnosis of the species has been provided by Child (1995a:44). These specimens are without legs, but there can be little doubt in their identification. There is a small rounded ocular tubercle which may or may not have eyes. The palps are typical of the species; very slender, long, and delicate. The lateral processes are glabrous, the proboscis has the three bumps arranged radially around its distal circum- ference, and the oviger bases are well an- terior to the first lateral processes and crowd most of the neck lateroventrally. There are very many chelae finger teeth. The tarsus and propodus are variable but usually subequal in length although it is im- 557 possible to tell from these legless speci- mens. Nymphon scotiae Stock, 1981 Fig. 4 Nymphon stocki Turpaeva, 1974:282, fig. 1 [preoccup.:Utinomi, 1955:10]. Nymphon scotiae Stock, 1981:458 (foot- note). Material examined.—Slope E of Cabo San Antonio, Argentina, 36°55.7’S, 53°01.4'W, 2707 m, sta. 245A, epibenthic Sled, 14 Mar 1971, 2 6 with eggs, 9 ¢, 19 2, 24 juveniles. Distribution.—This species was de- scribed from two specimens collected in the Scotia Sea in 2960-2980 m. It has not been captured since until the present record in the Argentine basin. This record places it well north but near the same longitude and in a similar depth. Remarks.—This is a slender graceful species which is only recorded with new material for the second time. Illustrations are provided herein for this rare species be- cause the only figures of its unique char- acters are not readily available, are diagra- matic and slightly inacurate. Nymphon typhlops (Hodgson, 1915) Chaetonymphon typhlops Hodgson, 1915: 144; 1927:327-329, fig. 6. Nymphon_ typhlops.—Gordon, 1944:19 [key].—Stock, 1965:22 [key].—Child, 1995a:20—21. Nymphon spicatum Child, 1982:46—48, fig. 15. Material examined.—Slope E of Cabo San Antonio, Argentina, 36°55.7'’S, 53°01.4'W, 2707 m, sta. 245A, epibenthic sled, 14 Mar 1971, 2 2, 4 Juv. Distribution.—This species was de- scribed from Antarctic specimens and the few other specimens known were collected either in Antarctic waters or in the Argen- tine slope and basin. The specimen de- scribed by Child (1982) as N. spicatum was 138 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Nymphon scotiae Stock. male: A, Trunk, dorsal view; B, Palp; C, Chela; D, Third leg; E, Oviger, with enlargement of terminal segment and claw. Ovigerous female: F Proximal leg segments. VOLUME 110, NUMBER 1 collected just slightly to the SE of the spec- imens in hand but in deeper waters. The known depth range for this species is 2450— 3822 m. Remarks.—This is another of the blind deep-sea species that either has vestigial auxilliary claws or none at all. The Argen- tine specimens all appear to lack auxiliaries and have a few small differences from the type specimens. The ocular tubercle is vari- able in its length and it appears to be di- rected slightly toward the posterior in most but not all specimens. The paired dorso- median trunk spines are often broken off or missing. The distal two palp segments are slightly longer in the types, the strigilis has several more denticulate spines, the chelae fingers a few more teeth, and the propodal claw is flanked by tiny auxiliaries. Nymphon vacans, new species Fig. 5 Material examined.—Slope E of Cabo San Antonio, Argentina, 36°55.7'S, 53°01.4'W, 2707 m, sta. 245A, epibenthic sled, 14 Mar 1971, 1 2, holotype, USNM 234717; 2 6,5 2, 7 juv. paratypes, USNM 234718. Distribution.—Known only from the type locality in 2707 m. Description.—Size moderate, leg span about 36 mm. Trunk fully segmented, gla- brous. Lateral process length about 1.5 times their diameters, separated by slightly less than their diameters, armed with two long dorsodistal spines except for second lateral processes which have three. Neck short, crowded with oviger implants. Very slight bulge of ocular tubercle placed dor- sally and slightly posterior to oviger bulges, sensory papillae evident laterally, eyes en- tirely lacking. Proboscis a cylinder with slight distal constrictions, oral surface flat. Abdomen of moderate length, tapering dis- tally to tiny tip, extending almost to distal rim of fourth leg first coxae, armed with four short dorsodistal setae. Chelifores large, scapes cylindrical, slightly longer than proboscis, with slight 139 inward curve, armed with lateral and distal setae of varying lengths. Chelae long, slen- der, palm about 0.75 length of fingers, armed with short scattered setae. Fingers curved sharply at tips which overlap, im- movable finger armed with short ectal setae and about 30 short teeth of varying sizes. Movable finger without setae, with about 37 teeth similar to those of immovable finger. Palps with many short setae increasing in numbers on distal segments. Second seg- ment slightly longer than third, fourth and fifth subequal in length, combined length longer than either second or third segment. Oviger (female) fourth segment slightly longer than fifth, sixth about 0.75 length of fifth, both with short lateral setae. Strigilis segments each shorter than last, each armed with ectal setae and endal denticulate spines in formula 8: 6: 4: 5:, spines slender, with three lateral lobes. Terminal claw slightly shorter than terminal segment, armed with six endal teeth. Legs moderately spinose, few spines lon- ger than segment diameters. Second tibiae the longest segments, with first tibiae very slightly shorter and femora shortest of ma- jor segments. Propodus about 0.8 as long as tarsus, both slender, armed with dorsal row of short setae and ventral row of very short sole spines. Claw about half propodal length. Auxiliaries lacking. Male characters: size slightly smaller, oviger fourth segment quite curved, fifth about 0.3 longer, almost straight, slightly swollen distally. Fewer denticulate spines, each with only two lateral lobes. Cement glands and pores not evident. Measurements (holotype in mm).— Trunk length, 3.55; trunk width across 2nd lateral processes, 1.8; proboscis length, 1.97; abdomen length, 1.0; third leg, coxa 1.0.6: coma .2. LLB coxa, 3, 0.88: femur, 3.17; tibia: 1, 3.86; tibia 2, 3.89; tarsus, 1.64; propodus, 1.3; claw, 0.66. Etymology.—The name (Latin: vacans, meaning empty, void, or clear) refers to the slight bulge of the ocular tubercle which is devoid of eyes. 140 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ¢ Fig. 5. Nymphon vacans, new species, holotype: A, Trunk, dorsal view; B, Ocular tubercle from right side; C, Third leg; D, Palp; E, Chela, with enlargement of several teeth; KE Oviger, with enlargement of denticulate spine and terminal claw. Male: G, Oviger; H, Strigilis, enlarged. VOLUME 110, NUMBER 1 Remarks.—The habitus of this species is superficially quite close to that of N. cen- trum, discussed elsewhere in this report, and to N. hampsoni Child, N. compactum Hoek, and indeed to most members of the N. australe-group of related species (Child 1995a). The similarities with N. centrum occur in the trunk habitus of the same shape with long spines on the lateral processes, although the spines are placed differently on each species. The many small differ- ences are in the proboscis and chelifore scape lengths; long in N. vacans and much shorter in N. centrum. The abdomen of this new species is shorter than that of N. cen- trum, and its palp segments are longer than those of the latter species. Nymphon vacans agrees with N. hamp- soni in lateral process length and place- ment, the dorsolateral spination, proboscis and chelifore length, and with other simi- larities in legs and palps. The major differ- ences occur in the short male ovigers with club-shaped fifth and sixth segments in N. hampsoni and also in N. compactum. The ovigers of N. vacans have much longer fifth segments in relation to the fourth, and they are only slightly inflated distally and the sixth segment is much shorter in relation to the fifth. The other major difference be- tween all of these species is in abdominal length. This varies with each species and is longest in N. centrum and decreases in length from N. hampsoni, then N. vacans, and is shortest in N. compactum. Nymphon species indeterminate Material examined.—Basin E of Mar del Plata, Argentina, 38°16.9'S, 51°56.1’'W, 4382-4402 m, sta. 242, epibenthic sled, 13 Mar 1971, 1 2, 2 Juv. Same locality, 37°40.9’S, 52°19.3'W, 3906-3917 m, sta. 256, epibenthic sled, 24 Mar 1971, 1 juv. Remarks.—The female from sta. 242 is badly damaged and the other specimens are too immature for identification. 141 Acknowledgments I wish to thank Joel W. Hedgpeth for re- linquishing his right to describe this collec- tion and for contributing it to me for ex- amination and report. All specimens are de- posited in the National Museum of Natural History, Smithsonian Institution. Literature Cited Child, C. A. 1982. Deep-sea Pycnogonida from the North and South Atlantic basins.—Smithsonian Contributions to Zoology 349:1—54. . 1994a. Antarctic and subantarctic Pycnogon- ida I. The family Ammotheidae.—Biology of the Antarctic Seas XXIII. Antarctic Research Series 63:1—48. . 1994b. Antarctic and subantarctic Pycnogon- ida II. The family Austrodecidae.—Biology of the Antarctic Seas XXIII. Antarctic Research Series 63:49-99. . 1995a. Antarctic and subantarctic Pycnogon- ida III. The family Nymphonidae.—Biology of the Antarctic Seas XXIV. Antarctic Research Series 69:1—68. . 1995b. Antarctic and subantarctic Pycnogon- ida IV. The families Colossendeidae and Rhyn- chothoraxidae. Biology of the Antarctic Seas XXIV.—Antarctic Research Series 69:69—111. Dohrn, A. 1881. Die Pantopoden des Golfs von Nea- pel und der Angrenzenden Meeresabschnitte.— Flora und Fauna des Golfes von Neapel 3:1— 22s Fabricius, J. C. 1794. Entomologia Systematica Emendata et aucta 4:416—417. Hafniae. Fry, W. G., & J. W. Hedgpeth. 1969. Pycnogonida, 1. Colossendeidae, Pycnogonidae, Endeidae, Am- motheidae. Fauna of the Ross Sea 7.—Memoirs of the New Zealand Oceanographic Institute 49: 1-139. Gordon, I. 1932. Pycnogonida.—Discovery Reports 6:1-138. . 1944. Pycnogonida.—Reports of the British, Australian and New Zealand Antarctic Research Expedition (ser. B) 5(1):1-72. Hedgpeth, J. W., & J.C. McCain. 1971. A review of the pycnogonid genus Pantopipetta (family Austrodecidae, emend.) with the description of a new species. Pp. 217-229 in G. A. Llano and I. G. Wallen, eds., Biology of the Antarctic Seas, IV.—Antarctic Research Series 17:217- 229. Hodgson, T. V. 1907. Pycnogonida.—National Ant- arctic Expedition 1901-1904. 3(Zoology and Botany): 1-72. . 1915.. The Pycnogonida collected by the 142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Gauss in the Antarctic regions, 1901-3; prelim- inary report——Annals and Magazine of Natural History ser. 8, 15(85):141-149. . 1927. Die Pycnogoniden der Deutschen Siid- polar-Expedition 1901—03.—Deutsche Siidpo- lar-Expedition, Ser. II, Zool. 19:303-—358. Hoek, P P C. 1881. Report on the Pycnogonida dredged by HMS Challenger 1873—76.—Re- ports of the Scientific Results of Exploring Voy- age of HMS Challenger 3(10):1—167. Jarzynsky, T. 1870. Praemissus catalogus Pycnogon- idarum, inventarum in mari Glaciali, ad oras Lapponiae rossicae et in mari Albo, anno 1869 et 1870.—Annales de la Société des Natural- istes de St. Pétersbourg 1:319—320. Loman, J. C. C. 1908. Die Pantopoden der Siboga- Expedition.—Siboga Expeditie Monographie 40:1-88. Prell, H. 1910. Beitrage zur Kenntnis der Lebensweise einiger Pantopoden.—Bergens Museums Aar- bok (N.R.), 1909 (10):1-—30. Stock, J. H. 1963. South African deep-sea Pycnogon- ida with descriptions of five new species.—An- nals of the South African Museum 46(12):321— 340. . 1965. Pycnogonida from the southwestern Indian Ocean.—Beaufortia 13(151):13-33. . 1968. Pycnogonida collected by the Galathea and Anton Bruun in the Indian and Pacific Oceans.—Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjgbenhavn 131:7—65. . 1978. Abyssal Pycnogonida from the north- eastern Atlantic Basin, Part I—Cahiers de Biol- ogie Marine 19:189-219. . 1981. Abyssal Pycnogonida from the Walvis Basin, southern Atlantic.—Cahiers de Biologie Marine 22:453-—471. Turpaeva, E. P. 1955. [New Species of Sea Spiders (Pantopoda) from the Kurile-Kamchatka Trench].—Trudy Instituta Okeanologii P. P. Shirshov, Akademiya Nauk SSSR 12:323-327. [In Russian] . 1956. [Pantopoda of the genus Heteronym- phon from the northwest part of the Pacific Ocean.]|—Byulleten Moskovskovo Obshchestva Ispytatelei Prirody, Otdel Biologicheskii, 61(2): 67-72. [In Russian] . 1970. Belonging of Nymphon abyssale Stock to the genus Heteronymphon.—Zoologicheskii Zhurnal 49(11):1723-1725. [In Russian + En- glish title] 1974. [The pycnogonids of the Scotia Sea and surrounding waters.]—Trudy Instituta Okeanologii P. P. Shirshov, Akademiya Nauk SSSR 98:277-—305. [In Russian] Utinomi, H. 1955. Report on the Pycnogonida. col- lected by the Soyo-Maru Expedition made on the continental shelf bordering Japan during the years 1926—1930.—Publications of the Seto Marine Biological Laboratory V(1):3—42. Wilson, E. B. 1878. Synopsis of the Pycnogonida of New England.—Transactions of the Connecti- cut Academy of Arts and Sciences 5:1—26. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):143-—149. 1997. Syllides eburneus, a new species, with notes on other members of the genus (Polychaeta: Syllidae) from the coast of New England and New Brunswick Nathan W. Riser Marine Science Center, Northeastern University, Nahant, Massachusetts 01908, U.S.A. Abstract.—Syllides eburneus new species, is described from the subtidal zone of the southern Gulf of Maine and the intertidal zone of the northern part. Morphological data obtained from living individuals of Syllides convoluta Web- ster & Benedict, 1884, S. benedicti Banse, 1971, S. longocirrata (Orsted, 1845), and S. setosa Verrill, 1882 occuring in the region are presented to clarify dis- crepancies which have arisen from the use of fixation artifacts encountered in preserved specimens. The role of protraction and retraction in the location of the foregut in specific setigers is discussed. Syllides setosa Verrill, 1882 is accepted as a valid species, whose morphology has been misinterpreted. The polychaete genus Syllides is com- monly encountered in sandy interstitial en- vironments in which there is little indica- tion of anaerobic conditions, although there may be some silt or finely fragmented algal debris. Living individuals can be readily identified, and while many of the identify- ing characters can not be seen or clearly observed in preserved animals setation fre- quently is distinctive. Syllides convoluta Webster & Benedict, 1884, S. benedicti Banse, 1971, and S. longocirrata (Orsted, 1845) are found in the intertidal zone of the Gulf of Maine. S. longocirrata and S. se- tosa Verrill, 1882 (usually as S. japonica Imajima, 1966) have also been recorded from depths of 20—40 m, in environmental surveys in the region. A previously unde- scribed species has been encountered at depths of 7-30 m in Nahant Bay, Massa- chusetts, frequently in large numbers, and in the intertidal interstitial community in the Northern Gulf of Maine. Materials and Methods Specimens were obtained from sand col- lected for meiofaunal studies. Intertidal samples were dug from coarse sand beaches with a garden trowel at low tide level dur- ing periods of extreme low tide. The sub- tidal samples were taken from the piles of sand kicked out from under rocks by lob- sters digging retreats at depths of 7—40 m. Sediment was washed with fresh sea water and decanted onto 153 wm screens from which animals were removed for sorting. The sediment was then extracted with 7.5% MgCl,, and decanted onto the screens from which the animals were washed into fresh sea water. Specimens of the new species from the intertidal were not numerous, 3—8 per 8-ounce bag of sand. Subtidal samples yielded 15-35 individuals per 8 ounces, the largest numbers occuring at depths of 20 m. All measurements were obtained from liv- ing specimens. Syllides eburneus new species Figs. 1-5, 7 Diagnosis.—Small species attaining body length of less than 5 mm, up to 30 setigers. Prostomium hemispherical, slight- ly broader than long, projecting anteriorly beyond bases of lateral antennae. With two 144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. of posterior eyes, dorsal view; 2, Optical section of foregut region of immature specimen, ventral view; 3, Surficial dorsal view. pairs of eyes, one pair at bases of lateral antennae, second pair approximately same size as anterior pair, at posterior ventrolat- eral corners of prostomium. Tentacular seg- ment with many large greenish vacuoles. Antennae, tentacular cirri, and dorsal cirri of first three setigers, pseudoarticulate; dor- sal cirri of following setigers with up to 14 articles, large ivory white vacuoles in each article. Antennae, tentacular cirri and dorsal cirri about same length, caudal cirfi longer. Palps approximately same length as prosto- mium. Ventral papilla of palp digitiform. Pygidial stylus 16 wm long, with a stereo- cillium at either side of the tip. Ventral cir- rus of each parapod digitiform, extending slightly beyond setal lobe. Aciculum ta- pered slightly to rounded apex; a thin, Figs. 4, 5. Syllides eburneus new species. 4, Api- cal end of simple seta; 5, Falciger blades and apices of shafts. | Syllides eburneus new species. Slightly compressed living specimens. 1, Optical section at level curved, pointed aciculum also present in parapods of reproductive setigers. Simple setae denticulated on convex surface toward hemispherical apex (Fig. 4), usually only one per setiger, one or two thin, pointed, simple setae in last three setigers. Falciger blades unidentate, three to six in each par- apod, in three distinct sizes, large blades ~48 wm long, serrated at base, medium blades ~36 wm long, shortest blades ~14 wm long. Apex of shafts of falcigers some- times with a subapical tooth. Shafts of fal- cigers and simple setae of first three setigers thinner than those of following setigers. Capillary (swimming) setae begin on setig- er 10. Lips of pharynx nipple-like with large vacuole in base. Pharynx/proventric- ulus length ratio ~1:1.4, proventriculus width/length ratio ~1:1.9; ventriculus in 5 with no apparent caeca. Muscles of proven- triculus thin, lattice-like, approximately 35 rows posterior to proventricular organ, muscles not in rows anterior to organ. Go- nads begin in setigers 7-9; 1—2 yolky, pink ova, 65 wm diameter, per gonad; sperm 8 wm long, pear-shaped with pointed acro- some constituting about one-tenth of that length. Etymology.—Latin eburneus, like ivory or ivory colored, from Latin ebur, ivory; Cth iA A oe - e VOLUME 110, NUMBER 1 Figs. 6, 7. 145 v 6, Syllides setosa. Apices of shafts and bases of blades of falcigers. SEM 4000; 7, Syllides eburneus new species, Apices of shafts, and bases of blades of falcigers, SEM X3600. referring to the ivory colored vacuoles in the dorsal cirri. Holotype-—USNM 054107. Massachu- setts; Nahant Bay, North side of Egg Rock. Coarse shelly sand; 10 m depth. 12 Sep. 1976. Paratype-—USNM 054108. Same loca- tion and date. Observations.—Antenna and _ tentacular cirri of living sexual individuals were 0.19— 0.22 mm long. The dorsal cirri were up to 0.5 mm long with as many as 14 articles; the terminal article bearing scattered ster- eocilia at its apex, and each of the other articles with a ring of such cilia apically, and a single stereocilium in the middle of the posterior surface as is characteristic of other species of the genus. Caudal cirri tend to coil and attain lengths of 1.9 mm. They consist of a few long articles, rarely more than eight, usually six. Ventral cirrus (Fig. 2) of each parapod 57-76 wm long. The digitiform palpal papilla was 12-17 pm long with a few stereocilia scattered over the margin of the apex and was directed medially from the ventral edge of each palp near the palpal base. This structure is char- acteristic of Syllides species and was re- ferred to by Webster & Benedict (1884) as a “‘short conical cirrus”’ in their description of S. convoluta and as “‘small projecting pa- pillae”’ in the (1887) description of S. lon- gocirrata. It appears to be a sensory struc- ture incapable of use in food manipulation. The palps of preserved members of the ge- nus usually form a funnel in front of the prostomium with the palpal papilla directed inward, but in the living state, they fre- quently flatten so that the papillae project laterally. Greenish vacuoles (gold or amber in transmitted light), concentrated in a col- lar behind the tentacular cirri (Fig. 3), are 146 characteristic of the genus. Frequently such vacuoles are also present behind the pos- terior eyes in the prostomium. The nuchal organs are present in a sulcus anterior to the collar. The thickened backs of the blades of the falcigers hook at the tip, but a subapical tooth is absent; the shafts are rather simple with the apical end extending as a single pointed prong sometimes with a subapical tooth (Figs. 5, 7) and the articular side with a flat termination that may be slightly ele- vated to either side. The ivory-white con- tents of the vacuoles in the antennae and dorsal cirri become transparent after a few days in specimens maintained in sea water, but can be used for preliminary identifica- tion while sorting fresh samples. The pres- ence of two pairs of eyes (Fig. 1) is a pri- mary character for the species (A posterior dorsal pair is absent), but in immature spec- imens, with fewer than 20 setigers, only the anterior pair of eyes (corresponding to ocel- li) are pigmented. Gonads occurred from setiger 7—9 through 23-25 in specimens collected in October and November, with swimming setae beginning in setiger 10. The species seems to be an annual, breed- ing in late autumn and early winter. Inter- tidal specimens, collected in January and February at Liberty Point, Robbinston, Maine and Pagan Point, St. Andrews, New Brunswick, have consistently had fewer than 12 setigers. Remarks,—vThe anterior pair of eyes of S. eburneus are always pigmented and rel- atively large, compared with the ocelli of most other species, and the posterior pair can be located on small immature speci- mens even though they may not be pig- mented, but both pairs are pigmented and obvious in sexual individuals. Syllides ed- entula (Claparéde, 1868), a Mediterranian species in which the dorsal cirri are not ar- ticulate, was described as having two pairs of eyes and the specimens from the Gala- pagos, tentatively identified as that species by Westheide (1974) only one pair. West- heide’s specimens consisted of up to 12 se- tigers and some specimens were sexual. It PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON would appear that the Mediterranean S. ed- entula and that from the Galapagos are not conspecific. Preserved specimens of S. con- voluta, in which the eyes have lost pigmen- tation, and preserved S. eburneus are diffi- cult to distinguish without comparative ma- terial. Both species belong to the group of Syllides species in which the proventriculus is located in setiger 5, simple setae do not taper to a point, and when sexual, measure 3—5 mm. The two species are very similar. Variability of characters used to distinguish different species may “in reality simply represent intraspecies variability”’ as noted by Reish (1977), but the eye pattern and the » pigment in the dorsal and caudal cirri is a constant found in intertidal and subtidal S. eburneus. These characters are not restrict- ed to populations and appear to the author to signify separate species. Protractors and retractors of the foregut occur as muscle stays running from the body wall to attachment points on the phar- ynx and proventriculus of syllid poly- chaetes as illustrated and described by Ma- laquin (1893). The fine stays involved with eversion of the pharyngeal tip are weakly developed in the genus Syllides, while the protractors attached to the base of the phar- ynx and proventriculus are strongly devel- oped and draw back the body wall so that the proventriculus comes to occupy many or even all of the anterior setigers. The number of setigers occupied by the various regions of the foregut is not a fixed char- acter, and in preserved specimens varies ac- cording to degree of contraction of the an- terior body region. The stays (retractors) at the base of the proventriculus tend to fix the posterior location of the organ although it may be pressed into the ventricular setiger. Such compression against the intestine may bulge the ventriculus giving the impression that ventricular caeca are present. The ratio of length of pharynx to proventriculus can be affected by the amount of eversion or expansion (Fig. 1) of the pharynx, but the relative lengths are of use in distinguishing some sympatric species in which setation is VOLUME 110, NUMBER 1 closely similar such as S. benedicti and S. convoluta. The proventricular organ is an eliptical epidermal gland containing a strongly azanophilous secretion which tends to harden and fracture in histological preparations. The intestine is fixed in posi- tion anteriorly, and thus, the ventriculus oc- curs in a specific setiger. The ventriculus occurs in setiger 5 of S. convoluta and S. eburneus and setiger 8 of S. benedicti and S. longocirrata. Syllides eburneus is sympatric with Ophryotrocha gracilis Huth, 1934, Schis- tomeringos caecus (Webster & Benedict, 1884), and an eyeless species of Protodor- villea which are primarily subtidal species in the southern Gulf of Maine but are also commonly encountered in the intertidal of the northern part of the region. Other Syllides species from the Gulf of Maine.—The method of collection of sub- Strate does not allow for distinction be- tween epipsammic and mesopsammic spe- cies. The various species of Syllides main- tain themselves in the interstitial environment by wrapping around grains of sand, thus, few are encountered in the sea water rinses, and extraction with MgCl, is required. Substrate containing S. benedicti or S. convoluta routinely yielded 10-30 specimens per 8 ounce sand sample, thus over 100 individuals of each of these spe- cies have been examined and measured since the start of this study. Syllides convoluta has been the most fre- quently encountered member of the genus in intertidal coarse sand between the south- ern shore of Cape Cod, Massachusetts and Georgetown, Maine. When freshly collect- ed, the gut is dark purple to black in color, but in individuals maintained in clean sea water, it becomes reddish brown to tan. Fig. 3 of Banse (1971) is drawn from the syn- type slides of S. convoluta. The membrane- like hood covering the tips of the acicula and simple setae is not apparent as a result of the clearing and mounting of these spec- imens. The apices of these structures in fresh, unfixed specimens tend to be round 147 as a result of the constriction of the shaft near the apex and the visible presence of the hood over the tip, producing a ball-like shape. Ridges in the wall of the hood (form- ing the apex in Banse’s fig. 3b in which the hood is not apparent) are evident under high magnification. The ridges in the wall of the hood of the aciculum are weakly produced as in Banse’s fig. 3a, making the ball-like appearance of the apex more pronounced. (Oil immersion is required to see these ridg- es in S. eburneus and even then, they are frequently not apparent due to the smaller diameter of the distal ends of setae and acicula in this species.) Syllides benedicti Banse, 1971 is the dominant intertidal interstitial member of the genus North of Georgetown, Maine. Fig. 6h of Banse (1971) shows a ‘“‘basal spur’’ which is actually the thickened mar- gin of the cutting edge of the blade of S. benedicti. Beyond this thickened region, the edge is very thin. Serrations are not evident on any of the blades. There are four blade sizes, usually two small (15-20 pm long), one or two medium (27—29 wm), two with thickened ridge on blade (56—58 p.m), and two or three large (64—70 pm). San Martin et al. (1985) ascribed material from the pla- ya de Toja of Spain to S. benedicti; how- ever, although the serrations on the blades could be interpreted as intraspecific varia- tion, the length of the largest falciger blades, and the pharynx/proventriculus length ratio, indicate a distinct and different species. The simple setae of S. benedicti are similar to those of S. convoluta and S. ebur- neus. Ability to see the ridges at the apex is a factor of the relative diameter of that region, and thus, the ridges are pronounced in S. benedicti, obvious in S. convoluta and almost indiscernible in S. eburneus. The gut of freshly collected specimens of S. bene- dicti has been brown or tan. The muscle stays anchoring the intestine of S. benedicti originate in setiger 9. Gonads sometimes were observed in setigers 10 and 11, but in most sexual individuals first occurred in 12. Ova were pink. 148 In the type description of Syllides setosa Verrill, 1882, the “‘dark colored’’ pharynx is described as “‘apparently unarmed, but sometimes showing a pale, oblong spot, that might be taken for a feeble tooth.” Hartman (1942) reported that the type ma- terial could not be found, however a portion of a specimen from the “‘surface, July 26, 1883” and identified by Verrill as S. setosa (USNM 10080) is available and was ex- amined for the present paper. Emerton’s drawing of S. setosa (Hartman, 1944 PI. 24, fig. 11) shows a structure in the pharynx that has been interpreted as a tooth by var- ious workers in spite of Verrill’s statement. From time to time, Verrill mentioned in his papers that drawings accompanying some of his papers had to be corrected [appar- ently as a result of artistic license on the part of the illustrator as has been noted by Banse (1969) for the figure of Acrocirrus leidyi Verrill, 1882 in Hartman (1944), and is evident in the dorsal cirri, setae and se- tation, absence of palps etc, in Emerton’s figure of S. setosa]. Palps are not present, the articles of the dorsal cirri are few in number, the basal article is almost half the total length of the cirrus, and capillary setae begin on setiger 8 in Emerton’s figure. Ver- rill further stated that “‘simple long setae begin singly on the eighth or ninth setig- erous segment; fascicles of capillary setae appear on the eighteenth segment in our largest example.’’ Verrill described some polychaete species from the drawings as noted by Hartman (1944) and it is possible that he may have used Emerton’s figure and inadvertedly recorded eighteen instead of 8 in his description. However, neither 8 nor 18 seem to be correct. He stated that the description of S. setosa was ‘‘from life.’ At the end of the description, he mentions col- lecting specimens about 3 mm long “‘per- haps distinct from the above” with capil- lary setae beginning ‘“‘on the tenth body segment.’’ Banse (1971) noted that the sim- ple setae of the specimens which he ex- amined began in the first setiger and the capillary setae in the tenth as in the Nahant PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bay individuals. The benthic specimens from Nahant Bay have been taken routinely since 1976 between March and November at which time divers could be safely em- ployed. Specimens with gonads have ranged in length from 3.2—11.5 mm. Mor- phological differences which could not be accounted for by size (age or growth) were not evident in any of the more than forty specimens which were measured or exam- ined. Careful measurements using a cali- brated ocular micrometer have yielded no measurable differences other then total length and breadth. Verrill’s statement that the smaller epitokes might belong to a dif- ferent species may be correct, however, it is possible that the species lives and grows through more than one reproductive epi- sode or season, adding setigers after the ini- tial spawning. The heavily pigmented phar- ynx is very black in specimens over six mm long which might be an indication of aging. The simple setae of S. setosa are geniculate, flexing at the origin of the denticulations. The denticulated side terminates below the apical tooth in such a way that the apex often appears to be bidentate. The denticu- lated side is thin, much like the cutting edge of the falcigers and the thickened back hooks at the tip forming the apical tooth. The cutting edges of the falcigers are very thin, but ridges resulting in fine denticula- tion are modified producing two or three ‘‘basal spines”’ (Fig. 6). Subapical spines on the shafts of the falcigers are not distinctly visible with light nor Nomarski microscopy, but are demonstrable with SEM (Fig. 6). Imajima (1966) figured strong subapical serrations on the falciger shafts of S. japon- icus and comparable serrations are obvious on the Pacific coast specimen (USNM 45264) from Lopez Island, Washington identified by Banse (1971), and are distinct- ly figured by Westheide (1974) for S. ja- ponica edentula and by San Martin et al. (1985) for Syllides sp A. The notoacicula of S. setosa taper gently toward the apex which is expanded as a mushroom-like cap in dorsal and ventral view, but is flexed in VOLUME 110, NUMBER 1 lateral view. A slender accessory aciculum occurs adjacent to the notoaciculum in each parapod from setiger 10 through the intes- tinal region. Single slender, tapered neu- roacicula with rounded apices also occur in these parapodia. Intact dorsal cirri consist of up to twenty articles. The proventriculus is anchored in setiger 8, which is occupied by the ventriculus. Perkins (1981) furnished a table of characters recorded for the “‘S. japonicus” complex of species. He con- cluded that possibly five species were pres- ent, and that S. japonicus Imajima 1966 “perhaps should not be a member of the genus,”’ with which I concur, since Imajima specifically stated that “‘a subterminal mid- dorsal tooth’’ was present in the pharynx. Unfortunately, this has never been con- firmed. The sperm of S. setosa differ from those of other species in the present study in that the head is almost round, total length of head, about 4 ym and diameter slightly over 3 pm with a small pointed acrosome. Syllides longocirrata has only been en- countered in the intertidal of the Gulf of Maine at Pagan Point, along the seaward side of the causeway to Ministers Island, and at Joe’s Point, St. Andrews, New Brunswick, in 25 years of routine met- ofaunal sampling and has been associated with S. benedicti and S. eburneus at both locales. It falls into the same size range as S. benedicti and S. setosa, from which it can be readily distinguished by the distribution and morphology of the simple setae. Acknowledgments The author is indebted to the many stu- dents who throughout all months of the year for many years, have collected sand for me while SCUBA diving for their own research activities. Material for comparative purposes was supplied by the USNM. Literature Cited Banse, K. 1969. Acrocirridae n. fam. (Polychaeta Se- dentaria)—Journal of the Fisheries Research Board of Canada 26:2595—2620. 149 1971. A new species, and additions to the descriptions of six other species of Syllides Orsted(Syllidae: Polychaeta).—Journal of the Fisheries Research Board of Canada 28:1469-— 1481. Claparéde, E. 1868. Les annélides chétopodes du gol- fe de Naples.—Mémoires de la Société de Phy- sique et d’Histoire naturelle de Genéve 19:313- 584. Hartman, O. 1942. A review of the types of poly- chaetous annelids at the Peabody Museum of Natural History, Yale University.—Bulletin of the Bingham Oceanographic Collection 8:1—98. . 1944. New England Annelida. Pt.2. Includ- ing the unpublished plates by Verrill with re- constructed captions.—Bulletin of the Ameri- can Museum of Natural History 82:327-344. Imajima, M. 1966. The Syllidae (polychaetous anne- lids) from Japan (III). Eusyllinae.—Publications of the Seto Marine Biological Laboratory 14: 85-116. Malaquin, A. 1893. Recherches sur les Syllidiens. Morphologie, anatomie, reproduction, dévelop- pement.—Mémoires de Société des Sciences et Arts, Lille, pp. 1-477. Perkins, T. H. 1981. Syllidae (Polychaeta), principally from Florida, with descriptions of a new genus and twenty-one new species.—Proceedings of the Biological Society of Washington 93:1080— 1172. Reish, D. J. 1977. The role of life history studies in polychaete systemtics. Pp. 461—476 in D. J. Reish and K. Fauchald, Essays on polychaetous annelids in memory of Dr. Olga Hartman. Allan Hancock Foundation, University of Southern California, Los Angeles. San Martin, G., G. Gonzalez, & E. L6pez-Jamar. 1985. Aspectos sistematicos y ecoldgicos sobre de las algunas especies de Silidos (Polychaeta:Sylli- dae) de las costas gallegas.—Boletin del Insti- tuto Espafiol de Oceanografia 2:27—36. Verrill, A. E. 1882. Notice of the remarkable marine fauna occupying the outer banks off the south- ern coast of New England, and of some addi- tions to the fauna of Vineyard Sound.—Amer- ican Journal of Science 24:360-—371. Webster, H. E., & J. E. Benedict. 1884. The Annelida Chaetopoda from Provincetown and Wellfleet, Massachusetts.—Report of the United States Commissioner of Fisheries 1881. pp. 699-747. . 1887. The Annelida Chaetopoda from East- port, Maine.—Report of the United States Com- missioner of Fisheries 1885, pp. 707-755. Westheide, W. 1974. Interstitielle Fauna von Galapa- gos. XI. Pisionidae, Hesionidae, Pilargidae, Syl- lidae (Polychaeta).—Mikrofauna des Meeres- bodens 44:195-338. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):150—153. 1997. Luriculus minos (Platyhelminthes: Rhabdocoela: Luridae) from the Canary Islands Wolfgang Sterrer Bermuda Natural History Museum, Flatts FLBX, Bermuda Abstract.—Previously known from Crete (Eastern Mediterranean), the stat- ocyst-bearing turbellarian Luriculus minos (Sterrer, 1992) was found in subtidal fine sand off the island of Gran Canaria (eastern Atlantic). Of six species of Luridae known, this is the first to be reported from outside its type locality. Significant biometric differences in the sclerotized structures of the reproduc- tive system between the Cretan and Canarian populations suggest that contem- porary means of dispersal are sufficient for island colonization, but insufficient to prevent this species from diverging into distinct populations. Marcus (1950) described the marine mi- croturbellarian Lurus evelinae which com- bined the general organization of Dalyel- lioida-Provorticidae with the possession of a Statocyst, a unique feature that prompted Sterrer & Rieger (1990) to erect the family Luridae. Five additional species have been described to date (Sterrer & Rieger 1990, Sterrer 1992, Faubel et al. 1994), and the ultrastructure of spermiogenesis (Rohde & Watson 1993a), of sensory receptors (Roh- de & Watson 1993b), and of the statocyst (Rohde et al. 1993) has been documented. Because L. evelinae was described by Mar- cus as having paired testes as well as sep- arate germaria and vitellaria, Faubel et al. (1994) proposed the genus Luriculus for those species that have germovitellaria and a single testis. The family Luridae thus comprises the species Lurus evelinae Mar- cus, 1950, from the Atlantic coast of Brazil; Luriculus castor (Sterrer & Rieger, 1990) from the Atlantic coast of the United States; Luriculus tyndareus (Sterrer & Rieger, 1990) from Fiji; Luriculus minos (Sterrer, 1992) from Crete; and Luriculus australien- sis Faubel et al., 1994, from New South Wales. Lurus pollux Sterrer & Rieger, 1990, also from the Atlantic coast of the United States, remains a species inquirenda since neither its vitellaria nor its testes are known. No species of Luridae has ever been re- ported outside its type locality. During a brief collecting trip to Gran Ca- naria in February 1996 I found specimens that clearly belonged to Luriculus minos (Sterrer, 1992) but differed consistently in details of the male copulatory organ. Sam- ples were collected by snorkeling over the shallow bottom and scooping sand into a bucket. Specimen extraction, documenta- tion and analysis follow Sterrer (1971). This study is based on observation of live animals in squeeze preparation. Luriculus minos (Sterrer, 1992) - (Figs. 1, 2; Table 1) Localities.—Gran Canaria, Las Palmas, Playa de Las Canteras, fine sand with Sparse seagrass (Cymodocea nodosa), at 2-3 m depth, collected 16 Feb 1996; 5 speci- . mens. Arinaga, harbor basin, fine sand be- tween and in Cymodocea nodosa at 4-5 m ~ depth, collected 21 Feb 1996; 9 specimens. - Description.—Mature specimens (Fig. 1A) are 245—420 wm long (vs. 275-500 pm in Crete). The statocyst (Figs: 1B, 2C) con- tained 3 statoliths in 3.specimens and 4 stat- oliths in one, which gives a mean statolith — VOLUME 110, NUMBER 1 C Fig. 1. Luriculus minos. A, Habitus of free-swimming adult; B, Statocyst; C, Bursa mouthpiece; D, Scler- otized parts of reproductive system. Phase contrast micrographs of living specimens from Gran Canaria. Abbre- viations: bm bursa mouthpiece, pf perforation of vagina mouthpiece, s male copulatory stylet, vm vagina mouth- piece. number of 3.25 for the sample (3.22 in male stylet, a vagina mouthpiece, and a bur- Crete). sa mouthpiece (Fig. 1D), are nearly identi- The complex sclerotized structures of the cal between Cretan and Canarian specimens reproductive system, which consist of a except for two consistent differences (Table 152 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Biometric comparison of vagina mouth- piece of individual specimens from the Canaries and Crete. Specimens from Canaries 1 x (wm)* y (wm)* 2 1133 65 3 13 62 13 60 Mean 13.00 62.33 Crete x’ (wm)** y’ (wm)** 1 Dy, 52 2D, 21 a, 3 19 50 4 20 51 Mean 20.50 51.25 * See Fig. 2A. ~*' See Fig, 2B: 1). The distal, funnel-shaped portion of the vagina mouthpiece (Fig. 2A; y) is 60—65 um (mean 62.33 wm) long in Canarian specimens but only 50—52 wm (mean 51.25 —] 2 Sart Fatt ! AS ‘ANA a wiih ym) in those from Crete (Fig. 2B; y’). The “lateral perforation”? is 13.0 wm long in Canarian specimens (Fig. 2A; x) vs. 19-22 wm (mean 20.5 wm) in Cretan specimens (Fig. 2B; x’). The distal width is the same in both populations. An additional yet less consistent difference is that the distal end of the male stylet, a prominent papilla made up of concentric rings, reemerges from the vagina mouthpiece not straight, as in most Cretan specimens, but always bent at a 90° angle. The proximal end of the vagina mouthpiece flares into a shallow saucer into which the bursa mouthpiece also opens. Shaped like a roll of unequal-sized coins, the bursa mouthpiece is considerably more robust in Canarian than in Cretan speci- ~ mens (Figs. 1C, 2D-—F). Discussion.—Global distribution and speciation of marine interstitial sand fauna continue to pose the paradox of a high de- gree of taxonomic uniformity, including ap- Fig. 2. Luriculus minos. A, distal portion of vagina mouthpiece of a specimen from Gran Canaria; B, the same of a specimen from Crete; C, statocyst; D—K proximal end of vagina mouthpiece (vm) and bursa mouth- piece (bm) of 3 specimens from Gran Canaria. The scales apply to A—C and D-E respectively. Abbreviations: bm bursa mouthpiece, pf perforation of vagina mouthpiece, vm vagina mouthpiece; x, y and x’, y’ indicate distances measured (see Table 1). VOLUME 110, NUMBER 1 parent cosmopolitan species, combined with the absence of a ready dispersal mech- anism. Some authors consider contempo- rary means of dispersal—such as passive water-column transport, rafting on drifting objects, or clinging to “‘marine snow’’—as sufficient to explain all existing patterns (Giere 1993), whereas others invoke plate tectonics, coupled with very slow specia- tion rates, as the primary vector for global distribution (Sterrer 1973). Differences be- tween populations of Luriculus minos at Crete and the Canary Islands suggest that contemporary means of dispersal must have been sufficient for this species to colonize islands such as Crete and the Canaries, but not powerful enough to prevent its frag- mentation into significantly different popu- lations. Acknowledgments I am grateful to Dr. Angel Luque Esca- lona and Lic. Javier Pérez Fernandez of the Universidad de Las Palmas for providing lab facilities and helping with sample col- lection. Literature Cited Faubel, A., K. Rohde, & N. A. Watson. 1994. Sandy beach meiofauna of Eastern Australia (Southern 153 Queensland and New South Wales). I. Luricu- lus australiensis, gen. et sp. nov. (Luridae: Dal- yelliida: Platyhelminthes).—Invertebrate Tax- onomy 8:1009—1015. Giere, O. 1993. Meiobenthology. Springer-Verlag, Berlin, 328 pp. Marcus, E. 1950. Turbellaria Brasileiros (8).—Bole- tins da Faculdade de filosofia, ciéncias e letras, Universidade de Sao Paulo (Zoologia) 15:5— 19OL: Rohde, K., & N. A. Watson. 1993a. Ultrastructure of spermiogenesis and sperm of an undescribed species of Luridae (Platyhelminthes: Rhabdo- coela).—Australian Journal of Zoology 41:13-— £9: . 1993b. Ultrastructure of sensory receptors of an undescribed species of Luridae (Platyhelmin- thes: Rhabdocoela).—Australian Journal of Zo- ology 41:53-65. , & A. Faubel. 1993. Ultrastructure of the stat- ocyst in an undescribed species of Luridae (Pla- tyhelminthes: Rhabdocoela: Luridae).—Austra- lian Journal of Zoology 41:215—224. Sterrer, W. 1971. On the biology of Gnathostomuli- da.—Vie et Milieu, Suppl. 22:493-508. . 1973. Plate tectonics as a mechanism for dis- persal and speciation in interstitial sand fau- na.—Netherlands Journal of Sea Research 7: 200-222. . 1992. Lurus minos, the first species of Luri- dae (Turbellaria: Rhabdocoela) from the Old World.—Proceedings Biological Society Wash- ington 105:636—639. , & R. M. Rieger. 1990. New species of the statocyst-bearing marine dalyellioid genus Lu- rus Marcus (Luridae nov. fam., Turbellaria- Rhabdocoela).—Cahiers de Biologie Marine 31:485—500. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):154. 1997. Selection of a type species for Farfantepenaeus Burukovsky (Crustacea: Decapoda: Penaeidae) Rudolf N. Burukovsky Department of Hydrobiology, Kaliningrad State Technical University, Sovetsky Avenue 1, 236000 Kaliningrad, Russia. Abstract.—Penaeus brasiliensis Latreille, 1817, is selected as the type spe- cies of the subgenus Farfantepenaeus Burukovsky, of the genus Penaeus. The complex penaeid shrimp genus Pe- naeus Fabricius, 1798, has over time, been divided into a number of subgenera, includ- ing the nominate subgenus Penaeus, Meli- certus Rafinesque-Schmaltz, 1814, Litopen- aeus Pérez Farfante, 1969, Fenneropenaeus Pérez Farfante, 1969, and Farfantepenaeus Burukovsky, 1972. In a paper on the systematics and distri- bution of species of Penaeus, Burukovsky (1972) proposed the new subgenus Farfan- tepenaeus, included in it the species P. duorarum, P. brasiliensis, P. aztecus, P. californiensis, P. brevirostris, and P. pau- lensis, but did not designate a type species. According to Article 13b of the Internation- al Code of Zoological Nomenclature (1985), any genus-group name published after 1930 must be accompanied by the fix- ation of a type species, to make the name available. This brief note is published to correct this situation. Family Penaeidae Genus Penaeus Fabricius, 1798 Farfantepenaeus, new subgenus Penaeus (Farfantepenaeus) Burukovsky, 1972:10, 13, 17.—Pérez Farfante, 1988:9. Type species: By present designation, Penaeus brasiliensis Latreille, 1817:156. Type locality: Brazil [‘“‘cétes du Brésil’’ ]. Literature Cited Burukovsky, R. N. 1972. Nekotorye voprosy siste- matiki i rasprostraneniya krevetok roda Penae- us. Rybokhozyaistvennye issledovaniya v At- lanticheskom okeane. [Some problems of the systematics and distribution of shrimps of the genus Penaeus|.—Trudy AtlantNIRO, Kalinin- grad 42:3-21. Fabricius, J. C. 1798. Supplementum Entomologiae Systematicae. Hafniae, 572 pp. Latreille, P A. 1817. Pénée. Jn Nouveau dictionnaire d’histoire naturelle applique aux arts, principa- lement l’agriculture et l’?economie rurale ... Volume 25:152—156, Paris. Deterville. Pérez Farfante, I. 1969. Western Atlantic shrimps of the genus Penaeus.—Fisheries Bulletin, U.S. 67(3):461-591. . 1988. Illustrated key to penaeoid shrimps of commerce in the Americas.—NOAA Technical Report NMFS 64:1-32. Rafinesque-Schmaltz, C. S. 1814. Précis des décou- vertes et travaux somiologiques de M. r. C. S. Rafinesque-Schmaltz entre 1800 et 1814 ou choix raisonné de ses principales découvertes en zoologie et en botanique, pour servir s’introd- uction 4 ses ouvrages futurs. Palermo, Royale Typographie Militaire, 55 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(1):155—156. 1997. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE Applications published in the Bulletin of Zoological Nomenclature The following Applications were published on 30 September 1996 in Vol. 53, Part 3 of the Bulletin of Zoological Nomenclature. Comment or advice on any of these applications is invited for publication in the Bulletin and should be sent to the Executive Secretary (I.C.Z.N.), c/o The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. Case No. 2978 Plumularia Lamarck, 1816 (Cnidaria, Hydrozoa): proposed conservation by the designation of Sertularia setacea Linnaeus, 1758 as the type species. Lirobarleeia Ponder, 1983 (Mollusca, Gastropoda): proposed designation of Alvania nigrescens Bartsch & Rehder, 1939 as the type species. Arca pectunculoides Scacchi, 1834 and A. philippiana Nyst, 1848 (currently Bathyarca pectunculoides and B. philippiana; Mollusca, Bivalvia): proposed conservation of the specific names. Parapronoe crustulum Claus, 1879 (Crustacea, Amphipoda): proposed con- servation of the specific name. Meristella Hall, 1859 (Brachiopoda): proposed designation of Atrypa laevis Vanuxem, 1842 as the type species. Hemidactylus garnotii Duméril & Bibron, 1836 (Reptilia, Squamata): pro- posed conservation of the specific name. Bombycilla cedrorum Vieillot, [1808] and Troglodytes aedon Vieillot, [1809] (Aves, Passeriformes): proposed conservation of the specific names. The 124th Annual Meeting of the Biological Society of Washington will be held on Thursday, 8 May, 1997, at 11:00 in the Waldo Schmitt Room, National Museum of Natural History, Washington, D. C. 156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Opinions published in the Bulletin of Zoological Nomenclature The following Opinions were published on 30 September 1996 in Vol. 53, Part 3 of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be ob- tained free of charge from the Executive Secretary, I.C.Z.N., c/o The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. Opinion No. 1843. Stictostroma Parks, 1936 (Porifera, Stromatoporoidea): conserved, and Stic- tostroma gorriense Stearn, 1995 designated as the type species. 1844. Aplysia juliana Quoy & Gaimard, 1832 (Mollusca, Gastropoda): specific name conserved. 1845. Tropidoptera Ancey, 1889 (Mollusca, Gastropoda): Endodonta wesleyi Sy- kes, 1896 designated as the type species. 1846. Eophacops Delo, 1935 and Acernaspis Campbell, 1967 (Trilobita): con- served. 1847. Diplocentrus mexicanus Peters, 1861 (Arachnida, Scorpiones): rediscovered holotype confirmed as the name-bearing type. 1848. Cubaris murina Brandt, 1833 (Crustacea, Isopoda): generic and specific names conserved. 1849. Livoneca Leach, 1818 (Crustacea, Isopoda): the original spelling confirmed as correct, and the spelling Lironeca rejected. 1850. Nepa rustica Fabricius, 1781 and Zaitha stollii Amyot & Serville, 1843 (currently Diplonychus rusticus and Belostoma stollii; Insecta, Heteroptera): specific names conserved. 1851. XANTHOLININI Erichson, 1839 and QUEDIINI Kraatz, [1857] (Insecta, Coleop- tera): given precedence over some senior synonyms; Quedius Stephens, 1829: Sta- phylinus levicollis Brullé, 1832 designated as the type species. 1852. Melissodes desponsa Smith, 1854 and M. agilis Cresson, 1878 (Insecta, Hy- menoptera): specific names conserved. 1853. Xerammobates Popov, 1951 (Insecta, Hymenoptera): Ammobates oxianus Popov, 1951 designated as the type species. 1854. Rhabdomeson Young & Young, 1874 (Bryozoa): Rhabdomeson progracile Wyse Jackson & Bancroft, 1995 designated as the type species. 1855. Agonus Bloch & Schneider, 1801 (Osteichthyes, Scorpaeniformes): con- served; and AGONIDAE Kirby, 1837 (Insecta, Coleoptera): spelling emended to AGON- UMIDAE, So removing the homonymy with AGONIDAE Swainson, 1839 (Osteichthyes, Scorpaeniformes). 1856. Lycognathophis Boulenger, 1893 (Reptilia, Serpentes): conserved. 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CONTENTS Cretaceous anuran and dinosaur footprints from the Patuxent Formation of Virginia Robert E. Weems and Jon M. Bachman A new species of the genus Platymantis (Amphibia: Ranidae) from Luzon Island, Philippines ; Walter C. Brown, Angel C. Alcala, and Arvin C. Diesmos A new species of chaenopsid fish, Emblemariopsis ruetzleri, from the western Caribbean off Belize (Blennioidei), with notes on its life history Diane M. Tyler and James C. Tyler Pontoniine shrimps (Decapoda: Caridea: Palaemonidae) of the northwest Atlantic. V. Pericli- menes mclellandi, a new species, a gorgonian associate from Pine Cay, Turks and Caicos Islands, British West Indies Richard W. Heard and Stephen Spotte New records and distribution ranges of shrimps (Crustacea: Decapoda: Penaeoidea and Caridea) in Chilean waters Ingo S. Wehrtmann and Alberto Carvacho A new species of the genus Petrolisthes Stimpson, 1858 (Crustacea: Decapoda: Anomura: Porcellanidae) from Yonaguni Island, the Ryukyu Islands Masayuki Osawa A new species of Pachycheles from the Hawaiian Islands (Crustacea: Decapoda: Porcellanidae) Alan W. Harvey and Elizabeth M. De Santo A new genus for the Central American crab Pinnixa costaricana Wicksten, 1982 (Crustacea: Brachyura: Pinnotheridae) Ernesto Campos and Mary K. Wicksten New records of marine Isopoda from Cuba (Crustacea: Peracarida) Brian Kensley, Manuel Ortiz, and Marilyn Schotte Redescription and first record of Cymbasoma tenue (Isaac, 1975) (Copepoda: Monstrilloida) in the Mediterranean Sea E. Suarez-Morales and N. Riccardi Amboleberis cubensis, a new species of myodocopine ostracode from the vicinity of Cuba | (Crustacea: Ostracoda: Cylindroleberididae) Rogelio Lalana and Louis S. Kornicker Macrothrix smirnovi, a new species (Crustacea: Anomopoda: Macrothricidae) from Mexico, a member of the M. triserialis-group J. Ciros-Pérez and M. Elias-Gutiérrez Some deep-sea Pycnogonida from the Argentine Slope and Basin C. Allan Child Syllides eburneus, a new species, with notes on other members of the genus (Polychaeta: Syllidae) from the coast of New England and New Brunswick Nathan W. Riser Luriculus minos (Platyhelminthes: Rhabdocoela: Luridae) from the Canary Islands Wolfgang Sterrer Selection of a type species for Farfantepenaeus Burukovsky (Crustacea: Decapoda: Penaeidae) Rudolf N. Burukovsky International Commission on Zoological Nomenclature Notice: Annual Meeting 18 107 LS 128 143 150 154 | 3) 155 THE BIOLOGICAL SOCIETY OF WASHINGTON 1996-1997 Officers President: Stephen D. Cairns Secretary: Carole C. Baldwin President-elect: Richard P. Vari Treasurer: T. Chad Walter Elected Council John A. Fornshell Rafael Lemaitre Alfred L. Gardner Diana Lipscomb Susan L. Jewett James N. Norris Custodian of Publications: Storrs L. Olson Se PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: Frederick M. Bayer Invertebrates: Jon L. Norenburg Frank D. Ferrari Plants: David B. Lellinger Rafael Lemaitre Insects: Wayne N. Mathis Vertebrates: Gary R. Graves Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $25.00 (for USA and non-USA addresses) include subscription to the Pro- ceedings of the Biological Society of Washington. Annual dues are payable on or before January 1 of each year. Renewals received after January 1 must include a penalty charge of $3.00 for reinstatement. Library subscriptions to the Proceedings are: $40.00 for USA and non-USA addresses. Non-USA members or subscribers may pay an additional $25.00 to receive the Proceedings by Air Mail. The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. 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This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):157—166. 1997. Review of the genus Schizopathes (Cnidaria: Antipatharia: Schizopathidae) with a description of a new species from the Indian Ocean Dennis M. Opresko Oak Ridge National Laboratory, 1060 Commerce Park (107), Oak Ridge, Tennessee 37830, U.S.A. Abstract.—The genus Schizopathes is reviewed and its relationship with oth- er genera in the family is discussed. The genus contains two previously de- scribed species, S. affinis Brook and S. crassa Brook, and one new species described here as S. amplispina. Schizopathes amplispina can be differentiated from S. affinis and S. crassa by its larger spines and more closely spaced pinnules. Schizopathes amplispina is known only from the type locality in the western Indian Ocean. In the course of examining the antipa- tharian collections in the National Museum of Natural History (USNM) at the Smith- sonian Institution, several colonies of a spe- cies of Schizopathes were located which upon closer study were found to represent a new species. This is the first new species added to the genus since its establishment by Brook in 1889. The type specimens, which were collected by the R/V Anton Brunn in the Indian Ocean east of Mada- gascar, are deposited at the Smithsonian In- stitution, Washington, DC. Schizopathidae (Brook, 1889) Diagnosis.—Antipatharians with polyps transversely elongated and subdivided by ‘““mesogloeal septa’ into three sections, each bearing one pair of tentacles (Brook 1889). Middle section of each polyp with Sagittal tentacles, mouth, stomodeum (ac- tinopharynx), and six primary and four sec- ondary mesenteries. Lateral sections of each polyp with pair of lateral tentacles and single transverse mesentery containing re- productive cells. Remarks.—The Schizopathinae was orig- inally established by Brook as one of two subfamilies in the family Antipathidae. In- cluded in the subfamily were the genera Schizopathes, Bathypathes, Cladopathes, and Taxipathes. The basis for subfamily recognition by Brook was the division of each polyp along the transverse axis into three sections each bearing one pair of ten- tacles, and which Brook regarded as di- morphic zooids (i.e., two gonozooids and one gastrozooid for each polyp). In Schi- zopathes these divisions of a polyp are as- sociated externally with “‘peristomal invo- lutions”’ (constrictions of the coenenchyme between the sagittal and lateral tentacles) and internally with ‘“‘mesogloeal septa” (also referred to as peristomal partitions by later workers) placed at right angles to the transverse axis of the polyp (Brook 1989). Most later workers have not considered this a case of true dimorphism but rather one of polyp modification and specialization, with the reproductive tissues being isolated in the lateral chambers of the coelenteron (Thompson 1905, van Pesch 1914). There are other antipatharian genera such as Par- antipathes in which the polyps are elongat- ed along the transverse axis and in which the reproductive organs are located at the distal and proximal ends of the primary 158 transverse mesenteries; however, all the genera originally placed in the subfamily Schizopathinae presumably had ‘‘meso- gloeal septa’’ which further isolated the dis- tal and proximal coelenteral chambers of each polyp. It should be noted, however, that Brook did not provide as detailed a de- scription of the internal anatomy of the pol- yps of Bathypathes, Cladopathes, and Tax- ipathes as he did for Schizopathes. A\l- though the polyps of these genera are elon- gated transversely (in some cases to a greater degree than that occurring in Schi- zopathes), the occurrence of internal “‘me- sogloeal septa’’ was not specifically men- tioned by Brook. Brook did report, how- ever, that the polyps of Bathypathes and Taxipathes, like those of Schizopathes, pos- sessed 10 mesenteries, six primary and four secondary; the number, location and ar- rangement being identical to that occurring in the family Antipathidae. The polyps of Cladopathes were described as having six primary mesenteries but no secondaries. In his revision of the order, Schultze (1896) divided the family Antipathidae into three subfamilies based on the number of mesenteries in the polyps; the Dodekamer- ota with 12 mesenteries, the Dekamerota with ten mesenteries, and the Hexamerota with six mesenteries. The Dekamerota were further divided into two unnamed tribes, one of which contained Schizopathes, Bath- ypathes, and Taxipathes. The genus Cla- dopathes was placed by Schultze (1896) in the new subfamily, the Hexamerota. van Pesch (1914) renamed Schultze’s Dekamer- ota as the Heterotaeniales and established the name Ptuchaephora for the tribe con- taining Schizopathes, Bathypathes and Tax- ipathes. Other workers, however, have taken the opposite approach and elevated the Schizopathinae to the family level (Hickson 1907, Pax 1918). With the recent removal of the Dendrobrachiidae from the Antipatharia (see Opresko & Bayer 1991), family level distinctions within the order need to be reevaluated. Based on the pro- nounced transverse elongation of the pol- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON yps, the recognition of the Schizopathidae as a distinct family seems appropriate; how- ever, further investigation is needed into the importance of the ““mesogloeal septa” as a family level character. Schizopathes Brook, 1889 Schizopathes Brook, 1889:146.—Hickson, 1907:6.—Cooper, 1909:308.—Pax, 1918: 468. Bathypathes.—van Pesch 1914:27 (part).— Pasternak, 1977:157 (part).—Zhou & Zou, 1992:46 (part). Type species.—Schizopathes crassa Brook, 1989. Brook (1889) does not spe- cifically designate a type species for any of his new genera; however, he makes the statement that the type species of all but one of the genera were examined microscopi- cally (Brook 1889:75). Of the three species that Brook referred to Schizopathes, only S. crassa was evaluated histologically (Brook 1889:46); therefore, by inference, it can be concluded that S. crassa is the type species of the genus. Diagnosis.—Colony monopodial, un- branched, but pinnulate. Pinnules simple, arranged alternately and bilaterally; de- creasing in length towards top of corallum. Basal section of stem not pinnulated, but modified to serve as holdfast in soft sedi- ments; upper part laterally compressed in plane at right angles to plane of pinnules; lower part curved away from polyp side of colony, sometimes spatula-like near base. Spines usually simple (rarely bifid), smooth, and triangular in lateral view. Pol- yps crowded, arranged in a single row usu- ally on the upper side of the pinnules, and elongated transversely; transverse diameter (from distal side of distal lateral tentacles to proximal side of proximal lateral tenta- cles) usually greater than 2.5 mm. Remarks.—Brook originally placed three species in Schizopathes, S. crassa, S. affinis, and S. conferta. The type material of the first two species consisted of complete col- onies with the unique basal holdfast intact. VOLUME 110, NUMBER 2 Schizopathes conferta was described from an incomplete specimen in which the bot- tom and top parts of the corallum were missing. Although the polyps of S. conferta were described by Brook as being very sim- ilar to those of the other two species in the genus, the absence of the holdfast, and the very narrow internal angle formed by the lateral rows of pinnules suggest that S. con- ferta may actually be a specimen of the re- lated genus Bathypathes in which the cor- allum is attached permanently to the sub- strate by a basal plate. Brook stated that he could not determine if S. conferta had been attached by a basal plate. Excluding S. conferta from considera- tion, features that are consistent among spe- cies of this genus are: the unique basal holdfast; the regular decrease in the length of the pinnules higher up on the corallum; the simple, triangular, laterally compressed spines; and the transversely elongated pol- yps. Features which have been used to sep- arate species of the genus include the size of the polyps and spines and the density of the pinnules. In all Schizopathes species there is a strong tendency for the pinnule density to increase towards the top the cor- allum and for the spines and polyps to be largest along the distal parts of the pinnules; however, these characteristics vary consid- erably from specimen to specimen and even with the same colony. It should be noted that some earlier workers such as van Pesch (1914) relegated the genus Schizopathes to subgeneric status within Bathypathes, and Pasternak (1977) even synonomized two of the three species described by Brook (S. crassa and S. affin- is) with Bathypathes patula Brook. The lat- ter decision was based on the assumption that the type of holdfast formed by the col- ony was simply a function of the type of substrate on which the planulae settled. Al- though there are similarities in terms of pin- nulation pattern and size of the polyps and spines, the very distinctive basal holdfast of all the species of Schizopathes (that have been described from complete specimens) 159 would argue for their being recognized as a separate genus. Schizopathes crassa Brook, 1889 Fig: 1 Schizopathes crassa Brook, 1889:147-—148, pl. VII, figs. 1-5. Bathypathes (Schizopathes) affinis.—van Pesch, 1914:27—29 (part). ?Schizopathes crassa.—Hickson, 1907:6 (incomplete colony). Material examined.—SW Indian Ocean, S of Tasmania, 58°06’S, 144°55’E, 3089-— 3164 m, USNS Eltanin sta. 126, 1 Oct. 1965 (1 specimen, USNM 78796).—Ant- arctica, NW of Balleny Ids, 64°59’S, 160°36’E, 2836-2864 m, USNM Eltanin sta. 1957, 7 Feb. 1967 (1 specimen, USNM 78820).—SE Indian Ocean, 55°01’S, 39°55'E, 2886-3040 m, USNS Eltanin sta. 1537, 8 Feb. 1966 (2 specimens, USNM 78810). Description.—Colony monopodial, un- branched, but pinnulate (Fig. 1B). Pinnules simple, arranged alternately in two lateral rows along the stem, and decreasing in length from base to apex. Maximum length of pinnules about 20 cm on colonies 30 cm tall (total length of stem) and about 30 cm on colonies 55 cm tall. Pinnules in each row usually 10—12 mm apart on lower parts of stem; becoming more closely spaced to- wards apex of corallum. Interior angle formed by lateral rows of pinnules 180° over most of corallum; <180° for lower- most pair, and greater or less than 180° for several pairs at top of corallum. Distal angle of pinnules 55—60° on lower parts of cor- allum but <45° near apex. Spines small, triangular and compressed; typically 0.06—0.08 mm tall (from midpoint of base to apex; range, 0.04—0.1 mm); ar- ranged in rows with varying degrees of reg- ularity; 3—5 spines per millimeter. Largest spines usually on polyp side of pinnules; abpolypar spines slightly smaller than po- lypar spines. 160 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. height 30 cm. Polyps (Fig. 1A) 4—6 mm in transverse diameter on pinnules; <4 mm at basal end of pinnules and on stem. Polyps arranged uniserially, with two per centimeter. Discussion.—The unpinnulated section of the stem of the type specimen of S. cras- sa was reported by Brook (1889) to be only 3.3 cm long. The unpinnulated stalks of the Eltanin specimens range from 6 to 8 cm. The maximum size of the spines in both the type and in the Eltanin specimens is about 0.1 mm. According to Brook (1889), the rows of spines on the flattened part of the stalk of the type are confined to the narrow anterior edge; however, in the Eltanin spec- Schizopathes crassa Brook, USNM 78820. A. Pinnule with polyps; scale equals 5 mm. B. Corallum; imens they are present on the wider sides as well. Brook (1889) reported that the “‘zooids”’ in the type specimen measured 3 mm across the base of the tentacles, and the illustra- tions given of the type indicate that the total transverse width of the polyps is 6 mm. Pol- yps of a similar size (commonly 5 mm, but up to 6 mm in transverse diameter) are present on the pinnules of the specimen from Eltanin sta. 126, with smaller polyps (as small as 3 mm or less) occurring at the base of the pinnules and on the stem. In the specimens from Eltanin sta. 1957 and 1537, most of the polyps on the pinnules are 4— VOLUME 110, NUMBER 2 4.5 mm and only a few are as large as 5 mm. As in the case with the other Eltanin specimen, smaller polyps (3—3.5 mm) occur at the base of the pinnules and on the stem. The specimen of S. crassa from Eltanin sta. 126 (USNM 78796) is 56 cm tall with pinnules up to 31 cm long. It is unique in that the pinnules in each lateral row are spaced up to 16 mm apart on the lowermost part of the corallum and as much as 12 cm apart near the apex of the corallum. In com- parison, the pinnules in the type and in the other Eltanin specimens are 10—12 cm apart on the lower part of the stem and 7-8 cm apart near the apex of the corallum. Comparisons.—See description of Schi- zopathes affinis Brook. Distribution.—The type locality, 35°39’S, 50°47'W, is off Montevideo, Uru- guay (Challenger sta. 323). All the Eltanin specimens were collected in the southern oceans. Schizopathes affinis Brook, 1889 Fig. 2 Schizopathes affinis Brook, 1889:148—150, pl. IX, figs. 1-6.—Cooper, 1909: 310, pl. 41, fig. 1-2. Bathypathes (Schizopathes) affinis.—van Pesch 1914:27 (part)—Zhou & Zou, 1992:46—47. Material examined.—Western Atlantic, Bahamas, Tongue of the Ocean, 25°16.00’ N, 77°45.3' W, 2780 m, R/V Columbus Is- elin sta. 172, 8 Feb. 1974 (2 specimens, USNM 59011).—Western Atlantic, Baha- mas, Tongue of the Ocean, 25°16’ N, 77°42.16' W, 2911 m, R/V Columbus Iselin sta. 288, 11 Nov 1974 (1 specimen, USNM 59009). Description.—Colony monopodial, un- branched, but pinnulate (Fig. 2B). Pinnules simple, up to 20 cm long in colonies 30 cm tall, and arranged alternately in two lateral rows along stem; decreasing in length to- ward apex of corallum; and inclined up- ward (distal angle formed with stem ~60° for lowermost pairs, decreasing to 30° or 161 less for those near apex). Lowermost pin- nules mostly 8-10 mm apart in each lateral row, decreasing to 5-6 mm apart near top of corallum. Interior angle formed by pin- nules in opposing rows generally 180°. Spines small, triangular and compressed; arranged, with varying degrees of regular- ity, in axial rows (4—5 rows visible in lat- eral view); 0.15—0.20 mm apart in each row (about six per millimeter). Spines usually 0.03—0.04 mm tall (from midpoint of base to apex), but occasionally up to 0.08 mm near the distal end of pinnule. Abpolypar spines equal to or slightly smaller than po- lypar spines. Polyps (Fig. 2A) uniserially arranged, on front or upper side of pinnules; 3.0—4.5 mm in transverse diameter; about three polyps per centimeter. Polyps slightly reduced in size at base of pinnules and on stem. Discussion.—Several features of the type specimen of S. affinis, including the pro- nounced triangular shape of the corallum, the relatively long unpinnulated stalk, and the very distinctly curved hook at the end of the holdfast, are also evident in two of the Columbus Iselin specimens. The unpin- nulated stalk is 12 cm in the type specimen and 10—12 cm in the Columbus Iselin spec- imens from sta 172, but only 7.5 cm in the specimen from Columbus Iselin sta. 288. Other features of the type, such as the size of the spines and polyps, were not specifi- cally described by Brook; however, based on the illustrations given, it can be estimat- ed that the spines in the type are not more than 0.06 mm and the polyps not more than 4.5 mm in transverse diameter. In the Co- lumbus Iselin material, the spines on the pinnules are mostly 0.03—0.04 mm, al- though in places they reach 0.08 mm, and the polyps (only present in the specimen from Columbus Iselin sta. 288) are 2.4—3.2 mm in transverse diameter. Brook (1889) states that the spines on the flattened section of the unpinnulated stalk are confined to the wider lateral sides. In the Columbus Iselin material, the spines oc- cur on the lateral sides on the upper portion PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bip. height 28.5 cm. of this section of the stalk, but along the narrow front edge further down towards the base. Comparisons.—According to Brook (1889), S. affinis can be differentiated from S. crassa by a greater curvature in the hooked base, and by a more pronounced tri- angular shape caused by a more abrupt de- crease in length of the pinnules from the lower to the upper parts of the corallum. The Columbus Iselin and Eltanin specimens tend to follow this pattern. For colonies of comparable size (about 30 cm), the ratio of the length of the lowermost pinnules to those 10 cm higher up on the axis is less than 0.5 in S. affinis and more than 0.5 in S. crassa. An analysis of a much larger suite of specimens would be needed to de- termine if this pattern is consistent for spec- imens of all sizes. From the limited number of specimens examined, it appears that col- Schizopathes affinis Brook, USNM 59009. A. Pinnule with polyps; scale equals 3 mm. B. Corallum; onies of S. crassa reach a larger maximum size then S. affinis (i.e., >50 cm in height). The major features that can be used to distinguish S. affinis from S. crassa are the size of the spines and polyps. The pinnular spines in S. affinis are typically 0.03—0.04 mm, but occasionally reach 0.08 mm. In contrast, the pinnular spines in S. crassa are usually 0.06—0.08 mm and sometimes as large as 0.10 mm. Brook (1889) reported that the polyps of S. affinis were smaller than those in S. cras- sa. Although polyp size was not specifically mentioned in the type description of affinis, the type illustrations do indicate a slight dif- ference in polyp size between the two spe- cies (maximum of 6 mm in crassa and 4.5 mm in affinis). In the Eltanin specimens of S. crassa the polyps on the pinnules are 4— 6 mm in transverse diameter, whereas those in the Columbus Iselin specimens of S. af- VOLUME 110, NUMBER 2 finis are not larger than about 3.2 mm. These data support Brook’s conclusions. Brook (1889) also states that the spines in S. affinis are more numerous than those in S. crassa. Although estimates made from the type illustrations indicate a spine den- sity of about 3/mm in both species, the Co- lumbus Iselin and Eltanin specimens indi- cate a density of about 6/mm for S. affinis and 3—5/mm for S. crassa. Distribution.—Cosmopolitan. Four of the type specimens and the colony de- scribed by van Pesch (1914) were collected in the western Pacific, off the Banda Is- lands. The fifth type specimen was obtained west of the Admiralty Islands. The species has also been reported from the Indian Ocean (Cooper 1909) and the South China Sea (Zhou & Zou 1992). The Columbus Is- elin specimens were collected in the West- ern Atlantic, near the Bahama Islands. Schizopathes amplispina, new species Figs. 3—4 Material examined.—Indian Ocean, east of Madagascar, 21°18’S, 36°18’E, 1510- 1600 m, R/V Anton Brunn sta. 399C, 10 Oct. 1964 (3 specimens; holotype USNM 96966, paratypes USNM 96967). Diagnosis.—Corallum monopodial, un- branched, but pinnulate (Fig. 3). Pinnules simple, arranged alternately and bilaterally; spaced 6—8 mm apart in each row on lower section of stem, decreasing to 2—5 mm apart near apex. Spines on basal half of pinnules small, triangular and compressed; 0.04— 0.10 mm from midpoint of base to apex (Fig. 4D); becoming larger (0.12—0.18 mm) and less symmetrical on upper parts of pin- nules (Figs. 4A—B). Polyps transversely elongated; 3.0—3.5 mm from distal side of distal lateral tentacles to proximal side of proximal lateral tentacles; arranged uniser- ially, with about three polyps per centime- ter. Description.—Holotype about 42 cm tall; unpinnulated stalk 8 cm; tip of stem and basal end of stalk missing. Stem com- 163 pressed laterally in plane at right angles to plane of pinnules; most strongly com- pressed along midsection of unpinnulated stalk (axis about 1 mm wide across anterior edge, but 3.5 mm along sides). Unpinnu- lated stalk sigmoidal; upper portion curved slightly away from polyp side of corallum, midsection curved strongly anteriorly, low- er section curved strongly posteriorly. Basal end of holdfast pointing away from abpo- lypar side of corallum; flattened, spatula- like near tip, with ridge extending down center of one side. Lowermost pinnules 24 cm long; pin- nules 10 cm higher on corallum about 19 cm long; those 10 cm higher about 16 cm; and smallest pinnules near apex 2—3 cm long. Pinnules spaced 6-8 mm apart on lower parts of corallum, decreasing to 2-3 mm apart near apex; inclined upward (distal angle formed with stem 55—60° for lower- most pairs, decreasing to about 30° at apex). Largest pinnules about 1.0 mm in diameter at base and 0.5—0.6 mm in diameter near midpoint. Rows of pinnules nearly opposite (interior angle about 180°) over most of corrallum; lowermost pair forming acute in- terior angle; uppermost pairs anterolateral or posterolateral in position. Polypar spines on basal half of pinnules (axis diameter 0.5—1.0 mm) triangular and symmetrical (Fig. 4D); 0.04—0.10 mm tall (from midpoint of base to apex); abpolypar spines 0.04—0.07 mm. Polypar spines on distal parts of pinnules (axis diameter 0.3-— 0.4 mm) large, less symmetrical, sometimes pointing proximally (Figs. 4A—B); 0.12- 0.18 mm tall (from midpoint of base to apex); abpolypar spines 0.10—0.12 mm. Bi- fid and double spines present on distal parts of pinnules, but usually absent on lower sections. Spines arranged in axial rows of varying regularity; 2—4 rows visible in lat- eral view on basal section of pinnules; 4—5 rows visible along distal section. Spines in each row mostly 0.25—0.35 mm apart (range 0.2—0.4 mm); with 4—5 spines per millimeter in each row. Rows of spines on unpinnulated stalk confined to wider sides 164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON PigSy (2—3 rows on each side) above, but with rows converging near anterior edge further down. Spines absent on lowermost 2 cm of stalk. Polyps transversely elongated; 3—3.5 mm from distal side of distal lateral tentacles to proximal side of proximal lateral tentacles; arranged uniserially, usually on the distal or anterior side of the pinnules, with about three polyps per centimeter. Discussion.—The two paratypes are about the same size as the holotype and are Schizopathes amplispina, holotype USNM 96966; height 42 cm. very similar in several other respects. The unpinnulated section of the stem is 8—9 cm long, the lowermost largest pinnules are 24—26 cm long, those 10 cm higher are 18— 20 cm and those 10 cm higher are 14—16 cm. The pinnules are as crowded together in the paratypes as they are in the holotype, the distance between adjacent pinnules in each lateral row never being more than 8 mm, even for the lowermost pairs. In all three of the type specimens the stem and the lower sections of the larger pinnules are VOLUME 110, NUMBER 2 tS ae * oe — . : ~ mses pi Sil AE OIE Seeders et O Fig. 4. Schizopathes amplispina, holotype USNM 96966. A. Distal section of pinnule; scale equals 0.1 mm. B. Distal section of pinnule; scale as in A. C. Midsection of pinnule; scale equals 0.1 mm. D. Basal section of pinnule; scale equals 0.1 mm. 166 compressed laterally such that their widest (anterior to posterior) axis is at right angles to the plane containing the pinnules. As in the holotype the largest spines in both paratypes occur along the distal half of the pinnules and on the side of the pin- nules corresponding to the polyp side of the corallum. On the basal sections of the larger pinnules, where the axis is compressed lat- erally, the rows of spines are not always evenly distributed around the circumference of the axis, and in some places the appear- ance is given that spines are missing from one side (Fig. 4C). Near the basal end of holdfast, the stem is compressed laterally in the same direction as the upper part of the stem in one specimen, but flattened at right angles to the upper part in the other two specimens. In the latter case, there is a small keel (defined by the central axial ca- nal) extending down the center of one side. The basal end of the holdfast ends in a rounded point. In none of the specimens are polyps pres- ent on all parts of corallum, and only in the holotype can the polyps be measured with any degree of reliability. In the holotype the polyps at the distal ends of the pinnules measure 3—3.5 mm in transverse diameter; they are about the same size or slightly smaller in the middle of the pinnules. Comparisons.—This species can be dif- ferentiated from S. crassa and S. affinis by its much larger spines (0.08—0.18 mm vs. 0.1 mm or less) and its much more closely spaced pinnules (7—8 mm apart in S. am- plispina vs. 8—16 mm apart in the other spe- cies). Furthermore, both the pinnules and the stem are much more strongly com- pressed in S. amplispina than in the other two species. In the size of its polyps (max- imum of 3.5 mm transverse diameter near the tips of the pinnules), S. amplispina re- sembles S. affinis (polyps 3—4.5 mm) more than S. crassa (polyps 4—6 mm). Etymology.—Latin amplius, larger, and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON spina, spine; in reference to the large spines on the distal sections of the pinnules. Distribution.—Known only from the type locality. Acknowledgments The author wishes to thank S. Cairns for his helpful suggestions and for taking the scanning electron micrographs (SEM), and S. Braden of the Smithsonian Institution for preparing the samples for the SEM. This work was supported in part by the Smith- sonian Institution and by Oak Ridge Na- tional Laboratory, Oak Ridge, TN. Literature Cited Brook, G. 1889. Report on the Antipatharia.—Reports of the Scientific Results of the Voyage of the Challenger, Zoology 32:5—222. Cooper, C. F 1909. Antipatharia. Reports of the Percy Sladen Trust Expedition to the Indian Ocean.— Transactions of the Linnean Society of London. (Zool. Ser. 2) 12:301—323. Hickson, S. M. 1907. The Alcyonaria, Antipatharia and Madreporaria, collected by the “Huxley” from the north side of the Bay of Biscay, Au- gust, 1906.—Journal of the Marine Biological Association 8:6—14. Kinoshita, K. 1910. On a new antipatharian Hexapa- thes heterosticha n. gen. and n. sp.—Annota- tiones Zoologicae Japonenses 7:231—234. Opresko, D. M., & E M. Bayer. 1991. Rediscovery of the enigmatic coelenterate Dendrobrachia, (Octocorallia: Gorgonacea) with descriptions of two new species.—Transactions of the Royal Society of South Australia 115:1—19. Pasternak, E A. 1977. Antipatharia——Galathea Re- port 14:157-164. Pax, E 1918. Die Antipatharien.—Zoologische Jahr- bucher 41:419—478. Schultze, L. S. 1896. Beitrag zur Systematik der An- tipatharien.—Abhandlungen der Senckenber- gischen naturforschenden Gesellschaft 23:1—40. Thompson, J. 1905. Report on the Antipatharians, Scotia Collection.—Proceedings of the Royal Physical Society of Edinburgh 16:76—79. van Pesch, A. J. 1914. The Antipatharia of the Siboga Expedition.—Siboga Expeditie Monographie 17:1-258. Zhou, J., & R. Zou. 1992. Studies on the antipathar- ians from Zhongsha Islands waters.—Tropical Oceanology 11:45—52. (in Chinese) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):167—185. 1997. Cnidae of Scleractinia Débora de Oliveira Pires Departamento de Invertebrados, Museu Nacional/Universidade Federal do Rio de Janeiro, Quinta da Boa Vista s/n, 20940-040, Rio de Janeiro, RJ, Brasil Abstract.—Corals and corallimorpharians are considered cnidarians with simple cnidae. This work shows that such a statement is relative, and depends upon the accuracy of the observation. The cnidae of tentacles and mesenterial filaments of 60 species belonging to 17 families of Scleractinia and three fam- ilies of Corallimorpharia were studied using Normarski optics. The diversity of the cnidae (morphological patterns and their varieties) is described and il- lustrated, and their distributions are given: In addition, a new type of cnida from the mesenterial filaments of Agariciidae is recorded. Synonym lists of the names used by other authors for the cnidae of Scleractinia and Corallimorpharia are also included, and aspects of their terminology are discussed. Diversity in the cnidae of Scleractinia confirms the potential of soft tissues as an important source of characters that should be explored in taxonomic studies. When used with other characters, the descriptions of cnidae will contribute to the improve- ment of studies on coral taxonomy and systematics. The knowledge of the cnidae in terms of their diversity, size and distribution, has been considered as a useful taxonomic character for many groups of Cnidaria (see Pires & Pitombo 1992). The nematocysts represent categories that are usually con- stant (Gravier—Bonnet 1987, Pires 1988) and their structural complexity provides many features for comparison (Tilbury & Cameron 1989). The systematics of the Scleractinia, the largest order of the sub- class Hexacorallia (=Zoantharia) (Wells 1956), with approximately 225 recent gen- era and 1500 species (Cairns 1990), is not fully satisfactory (Chevalier & Beauvais 1987), being based primarily on skeletal morphology (Wells 1956). One of the reasons for the neglect of cor- al cnidae studies is the lack preservation of their tissue. It is not routine in most scien- tific collections to preserve scleractinian coral tissues, especially in formalin, be- cause the skeleton is the primary or only taxonomic source of characters. Problems with nematocyst nomenclature, some of which have been recently discussed by En- gland (1991), have also made it difficult to understand and compare studies that have included data on cnidae. The goal of this paper is to describe the diversity of cnidae occurring in tentacles and mesenterial filaments of Scleractinia and to propose a detailed terminology to classify subtypes of coral cnidae. This study is based on a large array of taxa and on features that can be observed through light microscopy, which is easily accessible to most researchers. To help in the understand- ing and interpretating other papers, syn- onym lists of terms used for cnidae by other researchers are included. The cnidae of the Corallimorpharia were also studied here due to their great similarities with the Scler- actinia, as has been previously pointed out (Schmidt 1974, den Hartog 1980). The cnidae are structures unique to the 168 Cnidaria and it seems clear that they must have a distinct place in phylogenetic spec- ulations (Vervoort 1987). This paper con- tributes to explore some characteristics of coral cnidae, with the aim of providing new perspectives for systematics studies. Methods and Materials The cnidae of tentacles and mesenterial filaments of Scleractinia were studied be- cause these structures were found to have the most diversified cnidae (Pires, pers. obs.) Most observations were made on ma- terial fixed in 4% formalin and stored in 70% alcohol or 4% formalin. Some species from Brazil were maintained alive for ex- aminating discharged cnidae. Decalcifica- tion of the skeleton was obtained using a solution of 10% formic acid and 5% for- malin. Measurements were made with an eyepiece micrometer and illustrations were done using camera lucida. Preparations were examined with interference contrast, using a magnification of 1250 X. The syn- onyms of the cnidae are cited in the original publication language. For the evaluation of the diversity of the cnidae, the species listed below were examined. The taxa are listed in the same order used by Wells (1956) for Scleractinia (except Fungiacyathidae), and den Hartog (1980) for Corallimorpharia. The families are indicated in parentheses: Scleractinia.—Suborder Astrocoeniina: Stephanocoenia michelinii (Astrocoeni- idae); Madracis decactis (Pocilloporidae); Suborder Fungiina: Agaricia agaricites, A. fragilis, Pavona sp., P. gigantea, Lep- toseris cucullata (Agariciidae); Siderastrea stellata (Siderastreidae); Cycloseris mar- ginata (?), Lithophyllum mokai, Fungia (Cycloseris) sinensis, Fungia (Verrillofun- gia) repanda, Fungia (Verrillofungia) con- cinna, Fungia (Fungia) fungites, Herpoli- tha limax, Polyphyllia talpina, Halomitra pileus, Zoopilus echinatus (Fungiidae); Leptopenus antarcticus (Micrabaciidae); Goniopora tenuidens, Porites astreoides PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Poritidae); Fungiacyathus sp. (Fungiacy- athidae); Suborder Faviina: Favia gravida, F. lep- tophylla, Favites abdita, Platygyra daeda- lea, Colpophyllia natans, Cladocora debilis, C. arbuscula, Montastrea caverno- sa (Faviidae); Astrangia rathbuni, Phyllan- gia americana (Rhizangiidae); Oculina pa- tagonica, Madrepora sp. (Oculinidae); Meandrina braziliensis (Meandrinidae); Mussismilia hartti, M. braziliensis, M. his- pida, Scolymia wellsi, Lobophyllia hempri- chii, Lobophyllia sp. (Mussidae); Pectinia paeonia (Pectiniidae); Suborder Caryophylliina: Caryophyllia ambrosia caribbeana, C. cornuformis, Del- tocyathus italicus, D. calcar, D. eccentri- cus, Dasmosmilia lymani, D. variegata, Rhizosmilia maculata, Physogyra lichten- steini (Caryophyllidae); Flabellum sp., Ja- vania cailleti (Flabellidae); Suborder Dendrophylliina: Balanophyllia europaea, Balanophyllia sp., Dendrophyllia sp., (Dendrophylliidae). Corallimorpharia.—Corynactis califor- nica (Corallimorphidae), Ricordea florida (Ricordeidae), Discosoma carlgreni (Dis- cosomatidae). The studied material is deposited in the Collection of Cnidaria of the Museu Na- cional/Universidade Federal do Rio de Ja- neiro, Rio de Janeiro (MNRJ) (Appendix), except Pavona gigantea (USNM 81542), Pectinia paeonia (USNM 90471) and Mad- repora sp. from the National Museum of Natural History, Smithsonian Institution, Washington D. C. Results The cnidae of tentacles and mesenterial filaments of the studied material can be grouped into five basic morphological pat- terns: spirocysts, holotrichs, b-rhabdoids, p-rhabdoids D (sensu Schmidt 1974) and agaricysts. Four of these cnidae were al- ready well described in the literature (Weill 1934, Mariscal 1974, 1984; Schmidt 1974) and a new type is described (agaricysts). VOLUME 110, NUMBER 2 Table 1.—Diversity of cnidae and their varieties from tentacles and filaments of Scleractinia and Cor- allimorpharia. Spirocysts Holotrichs holotrich I [with one or two size classes] holotrich I (Discosoma var.) [close to holotrich I] holotrich II B-rhabdoids b-rhabdoid (1) b-rhabdoid (Discosoma var.) [close to b-rhabdoid (1)] b-rhabdoid (Favites var.) [close to b-rhabdoid (1)] b-rhabdoid (2) b-rhabdoid (Dendrophylliina var.) [close to b-rhabdoid (2)] b-rhabdoid (3) b-rhabdoid (Physogyra var.) [close to b-rhabdoid (3)] P-rhabdoids D p-rhabdoid D(1) p-rhabdoid D(2) [with one or two size classes] p-rhabdoid D(3) p-rhabdoid D(4) p-rhabdoid D(5) p-rhabdoid D(6) Agaricysts However, subvarieties of previously de- scribed types (namely holotrichs, b-rhab- doids, and p-rhabdoids D) are herein intro- duced and characterized. Table 1 shows the types of cnidae here observed and their sub- varieties. Table 2 shows the range of mea- surements of the cnidae observed. Tables 3-5 show main differences between sub- varieties of holotrichs, b-rhabdoids, and p-rhabdoids D. Their distribution in differ- ent taxa of Scleractinia and Corallimor- pharia is tabulated elsewhere (Pires & Cas- tro, 1997). Spirocysts (Figs. 1A, 3A) Description.—Capsule elongate and thin- walled, with a long tubule of uniform di- ameter, coiled in numerous spirals. Tubule without spines. Remarks.—The spirocysts are especially 169 abundant in tentacles, although they may be seen in the other structures of the polyps of both Scleractinia and Corallimorpharia. Holotrichs Description Capsule of varying shape and size (Table 2), from ellipsoid to very elongated cylinder. Tubule without differen- tiated parts - without the funnel-shaped ‘““V” notch; long and helicoidal inside the capsule. Discharged tubule bears spines along almost all its length, arranged in distinct coils, and at right angles to the tubule surface. Remarks.—Two morphological varieties of holotrichs were recognized in Coralli- morpharia and Scleractinia. holotrich I (Figs. 1B, 2, 3B, 3C, Table 3) — tangled cnidae (cnidae glomiferae) sen- su Gosse 1860 — nematocyst type III sensu Matthai 1914 —nematocyst type IIIb and IIIc sensu Matthai 1928 —macrocniden sensu Seifert 1928 —haplonéme isorhize holotriche sensu Weill 1934 —nematocyst B sensu Abe 1938 —holotrich sensu Carlgren 1940 —miacrobasic p-mastigophore sensu Cu- tress 1955 —isorhize haploneme holotriche sensu Schmidt 1969 —haploneme holotriche I sensu Schmidt 1972 —haplonem holotrich I sensu Schmidt 1974 —haploneme holotrichous isorhiza sensu Mariscal 1974 — penicilli E sensu den Hartog 1980 — holotrichous isorhiza sensu Song 1988 —holotrich sensu England 1991 — holotrich I sensu Pires & Pitombo 1992 Description.—Capsule of varying shape and size (Table 2), from ellipsoid to very elongate cylinder; of rich contrast, with a very distinct tubule, usually arranged in many regular coils. Tubule isodiametric in dis- 170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ESN M Fig. 1. Cnidae of tentacles of Scleractinia and Corallimorpharia, scale = 10 wm. A, spirocyst of Leptopenus antarcticus; B, holotrich I of Phyllangia americana; C, holotrich I var. of Discosoma carlgreni; D, holotrich I of Discosoma carlgreni; E, b-rhabdoid (1) of Fungia (Verrillofungia) concinna; F, b-rhabdoid (1) of Discosoma carlgreni; G, b-rhabdoid (1) var. of Favites abdita; H, b-rhabdoid (2) of Discosoma carlgreni; I, b-rhabdoid (3) of Stephanocoenia michelinii; J, b-rhabdoid (3) var. of Physogyra lichtensteini; K, p-rhabdoid D(1) of Montastrea cavernosa; L, p-rhabdoid D(2) of Fungia (Verrillofungia) concinna; M, p-rhabdoid. D(3) of Siderastrea stellata; N, p-rhabdoid D(6) of Halomitra pileus. charged state, with spines of equal size pres- (Figs. 1B, 2, 3B, 3C). Tubule of holotrich I ent along whole length, except a small naked of some Corallimorpharia presents -an abrupt- basal portion that often is present; latter por- ly tapered and spineless distal tip. tion easily seen in undischarged capsules Remarks.—Holotrichs I are sparingly pres- VOLUME 110, NUMBER 2 17a Table 2.—Morphometrics of cnidae (except spirocysts) from tentacles and mesenterial filaments of Scleractinia and Corallimorpharia. All measurements are in pm. mcl = minimum capsule length; MCL = maximum capsule length; mcw = minimum capsule width, MCW = maximum capsule width; C = number of capsules measured; S = number of species in which the cnida was observed. a OS IS a Cnidae mcl MCL mcw MCW c S Fig a Tentacles holotrich I (1 size) 26:3 82.4 3.8 18.4 312 15 1B holotrich I (2 sizes—small) 33.8 69.0 113 30.0 a3 4 A) holotrich I (2 sizes—large) 3135 145.0 213 60.0 73 4 2 holotrich I (Discosoma var.) 125 21.0 Ee 6.0 4 | i holotrich II 21.9 42.0 5.0 12.0 20 1D b-rhabdoid (1) 16.3 56.8 1.3 6.3 1074 44 1E b-rhabdoid (Discosoma vat.) 15.0 26.0 5.0 7.0 6 1F b-rhabdoid (Favites var.) 11.9 18.8 ZS 4.4 px 1 1G b-rhabdoid (2) TS the3 1.5 25 10 l 1H b-rhabdoid (3) 13.1 632 2:5 8.1 618 25 1] b-rhabdoid (Physogyra var.) 24.4 35:0 9.4 25 oF 1 lJ p-rhabdoid D(1) 24.0 88.1 4.0 8.1 1144 a2 1K p-rhabdoid D(2) 20.0 OT 3.8 1321 641 pA LL, p-rhabdoid D(3) 10.6 25.0 Sul 70 139 6 1M p-rhabdoid D(6) 18.1 49.4 4.4 8.1 53 4 IN Filaments, holotrich I (1 size) 17.6 101.9 4.8 29.6 709 p 3B holotrich I (2 sizes—small) 73 60.0 4.8 20.0 457 22 3C holotrich I (2 sizes—large) 40.6 186.3 8.0 Ran 596 22 a holotrich II 8.1 29.4 20 12.5 497 14 3D b-rhabdoid (1) 15.0 S10 2.0 5.0 42 a 3E b-rhabdoid (Favites var.) 13.8 29.4 3A 5.0 pe 1 3F b-rhabdoid (2) 5.6 28.8 13 4.8 1038 49 3G b-rhabdoid (Dendrophylliina var.) E35 16.3 aul 4.4 38 1 3H p-rhabdoid D(2) (1 size) 16.0 59.4 3.8 15.6 957 pe) 4A p-rhabdoid D(2) (2 sizes—small) 16.9 TL 4.0 11.9 666 25 4B p-rhabdoid D(2) (2 sizes—large) 34.4 136.0 5.0 18.1 736 25 4B p-rhabdoid D(3) 8.1 32.0 235 £053) 843 41 4C p-rhabdoid D(4) 51.3 2.5 9.4 15.6 155 6 4D p-rhabdoid D(5) fea 96.9 ‘7S 28.1 24 1 4E Agaricyst 25.0 63.8 5.6 jen) 104 4 7 ent in the tentacles of corals. They occur in Micrabaciidae, Poritidae, Fungiacyathidae, Rhizangiidae, Oculinidae, in some Cary- ophylliidae (Caryophylliinae), Flabellidae and Dendrophylliidae, and are common in the tentacles of corallimorpharians. They occur in two size classes in the tentacles of Micraba- ciidae, Flabellidae and Discosomatidae (Fig. 2). These size classes are present in both kinds of tentacles of Discosomatidae (mar- ginal and discal), which uniquely presented also another distinct morphological variety of holotrich I. The latter shows poor contrast and is rather small (=holotrich I Discosoma var., Table 1, 2; Fig. 1C). The occurrence of two distinct size class- es of holotrich I is very common in the mesenterial filaments of some Faviina, Car- yophylliina and Dendrophylliina (Fig. 3C). They also occur in two size classes in the mesenterial filaments of Corallimorpharia of the families Corallimorphidae and Dis- cosomatidae. The anterior end of smaller ones is usually slightly tapered. 172 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig; 2. holotrich II (Figs. 1D, 3D, Table 3) —haplonéme anizorhize homotriche sen- su Weill 1934 —haploneme holotrich II sensu Schmidt 1972 (part) —homotrich sensu den Hartog 1980 —homotrich sensu England 1991 Description.—Capsule cylindrical, little refractive, of varying size (Table 2); content is composed of a long tubule, irregularly 1 Me ie PLM ¢ LZ — a7 4 ’ a. cS SSAA ” Two size classes of holotrichs I from discal tentacles of Discosoma carlgreni, scale = 10 wm. coiled in small turns, filling whole undis- charged capsule. Basal portion of tubule slightly enlarged and tapering toward distal end. Spines smaller than those of holotrichs I and shorter as the width of tubule decreas- es. Sometimes the slightly enlarged proxi- mal part of tubule, which bears longer spines, is visible in undischarged capsule. When discharged, this portion is smaller than length of capsule. Table 3.—Main differences between holotrichs I and holotrichs II. Contrast of Spines along Spines in the capsule Capsule sizes Discharged tubule Undischarged tubule discharged tubule undischarged state I high may occur in two isodiametric regular coils, not equal in size clearly visi- distinct size filling the whole ble classes capsule II low only one size basal portion irregular coils, fill- shorter as the —_undistinct class slightly en- ing the whole tubule width larged capsule decreases VOLUME 110, NUMBER 2 ¢.. ‘ » 7 o C Py SP 5 ' i 5 = ay as re ~_ et D - < = a i . F gids y ra AAS = iy it A‘ e. ety = aa) Aa~ : on od ‘ 74 Zz B Fig. 3. 173 Cnidae of mesenterial filaments of Scleractinia, scale = 10 wm. A, spirocyst of Colpophyllia natans; B, holotrich I (1 size class) of Cycloseris marginata (?); C, holotrich I (2 size classes) of Phyllangia americana; D, holotrich II of Porites astreoides; E, b-rhabdoid (1) of Balanophyllia sp.; E b-rhabdoid (1) var. of Favites abdita; G, b-rhabdoid (2) of Zoopilus echinatus; H, b-rhabdoid Dendrophylliina var. Remarks.—This cnida is very common in the mesenterial filaments of Poritidae, Faviidae, and Mussidae. It also occurs in the marginal tentacles of Discosomatidae. B-rhabdoids Description.—Capsule of varying shape and size (Table 2). Tubule with no differ- entiated parts - without funnel-shaped ““V”’ notch. Spines of proximal part of tubule slightly longer than those of distal portion. In the undischarged capsules, proximal part is visible as a rod. Tubule tapers very grad- ually; distal portion bears tiny spines. Remarks.—Three morphological varie- ties of b-rhabdoids were here recognized (b-rhabdoids (1), (2) and (3)). b-rhabdoid (1) (Figs. 1E, 3E, Table 4) — spirulae sensu Stephenson 1928 — basitriche sensu Weill 1934 — heteroneme b-rhabdoide sensu Schmidt 1969, 1972 —heteronem b-rhabdoid sensu Schmidt 1974 — basitrich sensu England 1991 Description.—Capsule cylindrical and elongated, sometimes slightly curved. Basal portion of tubule slightly enlarged, and can be seen in undischarged state as a thin axial 174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 4.—Main differences of b-rhabdoids subvarieties. Capsule contrast Capsule shape (1) high elongated cylinder, sometimes slightly curved (2) high oval (3) low cylindrical rod, almost of same size of capsule length. Width of tubule diminishes toward distal end. Tubule arranged in few distinct regular coils, somewhat horizontally arranged in- side capsule. When discharged, basal por- tion bearing larger spines is a little longer than entire length of capsule. Remarks.—B-thabdoid (1) is very com- mon in the tentacles of most scleractinian families. They may also be present in the mesenterial filaments of Dendrophylliidae. A similar morphological variety of b-rhabdoid (1) was observed in the tentacles (Fig. G) and mesenterial filaments (Fig. 3F) of Favites ab- dita. That variety is more ellipsoid in shape, the tubule being longer and arranged in many coils. The coils fill all the interior of the cap- sule. The marginal and discal tentacles of Discosoma carlgreni contain another variety of b-rhabdoid, similar of b-rhabdoid (1). It is more translucent, the tubule is arranged in fewer coils, and usually the axial rod is oblique inside the capsule (Fig. 1F). b-rhabdoid (2) (Figs. 1H, 3G, Table 4) — mastigophore microbasique sensu Weill 1934 — microbasic b-mastigophore sensu Carl- gren 1940 — b-rhabdoide sensu Schmidt 1969 —b-rhabdoid sensu Schmidt 1972, 1974 — spirulae 2 sensu den Hartog 1980 —miicrobasic b-mastigophore sensu En- gland 1991 Description.—Capsule oval, very small Size of enlarged basal portion of undischarged Arrangement of distal portion tubule of undischarged tubule almost equal to capsule length few distinct regular coils approximately % of cap- sule length approximately 15 of capsule length few distinct regular coils many small irregular coils, filling the whole interior of the capsule (Table 2), and of strong contrast. Basal part of tubule clearly distinguished in undis- charged state, this rigid part approximately % of capsule length. Thinner part of tubule bears tiny spines; these latter arranged in few conspicuous turns. Thinner part of tu- bule often coiled down to rigid basal part. Remarks.—B-rhabdoids (2) occur in the mesenterial filaments of all examined Scler- actinia, and in some Corallimorpharia (Cor- allimorphidae), and exceptionally in the discal tentacles of Discosomatidae (Fig. 1H). A variety of b-rhabdoid (2) of larger capsule and shorter tubule occurs in the mesenterial filaments of Dendrophylliidae (Fig. 3H, Table 2). b-rhabdoid (3) (Fig. 1I, Table 4) — basitrichs sensu Weill 1934 — microbasic b-mastigophore sensu Carl- gren 1940 — b-rhabdoide sensu Schmidt 1969, 1972 — b-rhabdoid sensu Schmidt 1974 — spirulae 1 sensu den Hartog 1980 —microbasic b-mastigophore sensu En- gland 1991 Description.—Capsule of varying shape, but usually cylindrical, and slightly refrac- tive. Proximal part of tubule bears spines somewhat longer than those from distal part. This region can be distinguished in un- discharged capsule as a rod; approximately Y; of the length of the capsule; thicker than rods of b-rhabdoids (1) and (2). Distal part of tubule long; irregularly arranged; fills VOLUME 110, NUMBER 2 whole interior of undischarged capsule. Ba- sal portion of tubule, approximately % of capsule length, in discharged state, with spines somewhat longer than those from thinner part of tubule. Remarks.—This variety can be confused with holotrich II, especially in the undis- charged state. Sometimes it is difficult to distinguish the slightly larger tubule basal portion, a characteristic of b-rhabdoids. Den Hartog (1980) considered them homolo- gous, since he observed transitional stages between both types. This variety occurs in the tentacles of Scleractinia (Astrocoeniidae, Pocillopori- dae, Agariciidae, Fungiidae, Faviidae Mon- tastreinae, Rhizangiidae, Oculinidae, few Caryophylliina, Dendrophylliidae) and Cor- allimorpharia (Ricordeidae and Corallimor- phidae). A morphological variety was observed in the tentacles of Physogyra lichtensteini. The proximal part of its tubule is excep- tionally thick and very distinct (Fig. 1J) P-rhabdoids D —chambered cnidae (cnidae cameratae) sensu Gosse 1860 — type II sensu Matthai 1914, 1928 —hétéronéme mastigophore microba- sigue sensu Weill 1934 —nematocyst A sensu Abe 1938 — microbasic p-mastigophore sensu Carl- gren 1940 — p-rhabdoid D sensu Schmidt 1974 — penicilli D sensu den Hartog 1980 —microbasic p-mastigophore D sensu England 1991 Description.—Capsule of varying con- trast, size and shape, from oval to very elongate cylinder. Tubule with two clearly differentiated parts: larger basal shaft and tapered distal tubule. These two regions de- limited by sudden change in diameter of tu- bule and in length of spines; seen as funnel- shaped “V’’ notch in undischarged cap- sules. Shaft often isodiametric; usually ori- 175 ented straight inside capsule. In few cases it presents discrete coil in its base or, more rarely, shaft entirely coiled. This disposition of tubule varies inside capsule. Both regions, shaft and tubule, usually coarsely spined. When discharged, spines arranged at perpendicular to shaft surface; distinctly longer than shaft diameter. Tubule long; bearing spines rather smaller than those of shaft. Remarks.—This cnida is exclusive to and very common in Scleractinia and Coralli- morpharia. Six morphological varieties of p-rhabdoids D were here recognized (p-rhabdoids D(1), D(2), D(3), D(4), D(5) and D(6)). p-rhabdoid D(1) (Fig. 1K, Table 5) —nematocyst IIb sensu Matthai 1928 — penicilli D1 sensu den Hartog 1980 —miicrobasic p-mastigophores type III sensu Thomason & Brown 1986 Description.—Capsule elongate; cylin- drical; slightly curved; of rich contrast. Shaft short (approximately 4% of capsule length) in undischarged state. A protruding tip commonly projects at anterior end of capsule. Spines of tubule clearly distin- guished even in undischarged state. Remarks.—This variety has a restricted distribution, occurring in the tentacles of most Scleractinia and commonly in the ten- tacles of Corallimorpharia, but not Disco- somatidae. p-rhabdoid D(2) (Figs. 1L, 4A, 4B, Table 5) —nematocyst IId sensu Matthai 1928 — penicilli D3 sensu den Hartog 1980 — microbasic p-mastigophores type II sensu Thomason & Brown 1986 Description.—Capsule often cylindrical; of rich contrast. Spines of shaft arranged in obvious turns—approximately % of capsule length. Tubule long, irregularly arranged in- side capsule. Remarks.—This variety occurs in the 176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 5.—Main differences of p-rhabdoids D subvarieties. Capsule contrast Capsule shape D(1) high elongated cylinder, slightly curved D(2) high cylindrical D(3)___ low, almost cylindrical to oval transparent D(4) high cylindrical D(5)_ low cylindrical D(6) low cylindrical Length of undischarged shaft approximately % of capsule length approximately % of capsule length approximately % of capsule length approximately equal to capsule length approximately equal to capsule length approximately 15 of capsule length Diameter of undischarged shaft isodiametric isodiametric isodiametric isodiametric anisodiametric with three distinct parts isodiametric Spines in undischarged tubule clearly distinguish- able clearly distinguish- able almost indistinct clearly distinguish- able clearly distinguish- able slightly distinguish- able mesenterial filaments of all Scleractinia and Corallimorpharia. In the mesenterial fila- ments of Astrocoeniidae and Pocilloporidae it is characteristically thick and club- shaped. It occurs in two distinct size classes in the mesenterial filaments of most Faviina (Fig. 4B). Exceptionally, it is also present in the tentacles of some Fungiidae (Fig. 1L). p-rhabdoid D(3) (Figs. 1M, 4C, Table 5) — penicilli D2 [?] sensu den Hartog 1980 Description.—Capsule thin-walled; little refractive; almost transparent; often ovoid in shape. Shaft thin; approximately % of capsule length. Tubule short; occupying small portion of capsule. Spines of shaft and tubule almost indistinct in undischarged State. Remarks.—This variety is very common in the mesenterial filaments of most Scler- actinia and Corallimorpharia. It also occurs in the tentacles of Siderastreidae and Agar- iciidae. p-rhabdoid D(4) (Fig. 4D, Table 5) Description.—Capsule cylindrical; of rich contrast. Shaft long, in undischarged state filling almost whole capsule length; its basal portion slightly to coarsely coiled. Tu- bule long; irregularly coiled. Spines clearly distinguished in both shaft and tubule. -Remarks.—This variety occurs in mes- enterial filaments of some Fungiina and Caryophylliina. When present, it substitutes the larger class of D(2), which is common in the mesenterial filaments of some Favi- ina (Fig. 4B). Only undischarged capsules were observed. The length of the shaft of this variety is similar to those belonging to the ‘“‘macrobasic state’? of Weill (1934). The inclusion of this variety in p-rhabdoid D is tentative, pending the examination of discharged capsules. p-rhabdoid D(5) (Fig. 4E, Table 5) — type IV sensu Abe 1938 Description.—Capsule very refractive in undischarged state. Shaft anisodiametric with three distinct parts: 1) short coiled ba- sal portion, very thin and apparently naked, clearly distinguished in undischarged state; 2) middle part with abrupt enlargement, coiled, coarsely armed with long spines, and diminishes toward distal part, both in width of shaft and length of spines, com- prises the longest and thickest part of shaft; and 3) distal part, gradually thicker, with very deep “‘funnel-shaped ’’V“ notch. Tu- bule relatively short; irregularly coiled; with tiny spines. VOLUME 110, NUMBER 2 A B C wi a | a q e si N Lv? =—— y, (4 O a, = << ——— SS Ral os 38 eB a CU D E Fig. 4. Cnidae of mesenterial filaments of Scleractinia, scale = 10 wm. A, p-rhabdoid D(2) (1 size class) of Madracis decactis; B, p-rhabdoid D(2) (2 size classes) of Favia leptophylla; C, p-rhabdoid D(3) of Cladocora debilis; D, p-rhabdoid D(4) of Polyphyllia talpina; E, p-rhabdoid D(5) of Goniopora tenuidens. Remarks.—This peculiar and very char- acteristic variety was found only in the mesenterial filaments of Goniopora tenui- dens. Only undischarged capsules were ex- amined. It clearly belongs to the “‘macro- basic state’’ of Weill (1934). The inclusion of this variety in p-rhabdoid D is tentative, pending the examination of discharged cap- sules. p-rhabdoid D(6) (Fig. 1N, Table 5) Description.—Capsule cylindrical; little refractive. Shaft thin; approximately Y; the capsule length. Spines of shaft and of tu- bule poorly distinguished in undischarged state. Tubule irregularly coiled; almost fills entire capsule. Remarks.—Only undischarged capsules were examined. This variety can be con- fused with p-rhabdoids D(3); however, be- sides its larger capsule, it presents a pro- portionally shorter shaft, and the tubule is 178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A Pigs. . ~ Kalas pee ot C D Agaricysts from the mesenterial filaments, scale = 10 pm. A, discharged capsule and spineless tubule of Agaricia agaricites; B, discharging capsule and tubule with few attached and detached spines of Agaricia agaricites; C, undischarged capsule of Pavona sp.; D, undischarged capsule of Leptoseris cucullata. somewhat longer than in p-rhabdoids D(3). It was only observed in the tentacles of spe- cies of Fungia and Halomitra. Agaricysts (Figs. 5, 6) Description.—Capsule elongated; thick- walled; usually has a somewhat crushed ap- pearance. Capsular content only slightly re- fractive. Tubule occupies almost whole length of capsule; arranged in two or three parallel bands (Fig. 5); running along lon- ger axis of capsule. Apparently some spines are found around some of these bands (Figs. 5C, 5D). Discharged tubule long; an- isodiametric (Fig. 5A). Proximal part of tu- bule approximately 2.4 mm in width, 4.0 —~ mm in median region, and 1.6 mm close to tapering tip. Discharged tubule may seem to be spineless; however, spines can be ob- served on tubule surface without a distinct arrangement. Few unattached spines close to discharged tubule also present (Fig. 5B). Spines long; seem to loose contact with tu- bule when it discharges. Discharged tubule forms almost right angle with longer axis of capsule (Fig. 6). Remarks.—This is a previously unde- scribed cnida, which was observed only in the mesenterial filaments of members of Agariciidae. Mariscal et al. (1977) classi- fied the coelenterate cnidae in two major groups; helicoptychonemes and heteropty- chonemes. The first group presents the un- VOLUME 110, NUMBER 2 Fig. 6. Agaricyst from the mesenterial filaments of Agaricia agaricites, scale = 10 ym. Discharged tubule forming almost a right angle with the longer axis of the capsule. discharged thread helically folded to form multiple pleats in length and three pleats in circumference and includes the nematocysts and spirocysts. The second one have the un- discharged thread not helically folded, with a variable number of pleats in circumfer- ence but none in length, and includes the ptychocysts. The agaricysts are similar to the heteroptychonemes described by Mar- iscal et al. (1977). Both present the unev- erted tubule not helically arranged inside the capsule. When discharged, both tubules are anisodiametric, with longitudinal ridges along their length. Commonly, both tubules evert at an angle of about 60 to 90 degrees with the plane of the capsule. Discussion A detailed study of the cnidae can pro- vide characters for a better understanding of the different groups of Cnidaria. This holds true even among taxa in which cnidae have 179 been previously considered of less taxo- nomic value. Even spirocysts, which show less structural variation, can be used as a source for comparative studies. Weill’s major work (1934) was a turning point in the study of nematocysts. However, its broad scope did not allow him to detail the cnidae of all cnidarian groups. In Scler- actinia, for example, he superficially ex- amined only eight species. He overlooked many cnidae, and he did not examine their distribution within the polyp. As the knowl- edge of the morphological structure of the cnidae of some groups accumulated, it be- came clear that some of Weill’s categories included a wide variety of morphological types. The “‘rhabdoides”’ of Weill (1934), for example, represent a group of different types of nematocysts with few characteris- tics in common, and a most diverse struc- ture. Schmidt (1974) divided the ‘‘rhabdoi- des” of Weill (1934) into four basic types: p-rhabdoids A, B, C and D. He subdivided these in some subcategories, and created a new terminology. Although his terminology is poorly known, his typology was accepted by some workers (den Hartog 1980, Ost- man 1983, England 1991, Pires & Pitombo 1992, Schlenz & Belém 1992). England (1991), pointed out the difficulties in using and adopting a codified alphanumeric sys- tem, without the descriptive elements of Weill’s classification. Nevertheless, Schmidt’s system is the one that most close- ly represents the diversity of the Anthozoa cnidae, and therefore is partially adopted in this paper. According to Schmidt (1972, 1974) Scleractinia presents only one type of p-rhabdoid belonging to his category D. When discharged, the shaft of this type is rarely longer, frequently equal or shorter, than the length of the capsule (Schmidt 1972), corresponding to the microbasic state of Weill. However, the p-rhabdoids D(4) and D(5) cited in this paper, even when examined only undischarged, belong to the macrobasics of Weill, in contradic- tion to Schmidt’s definition of his type D. 180 Moreover, p-rhabdoids D present the same basic structure as p-rhabdoids A. Both of them have the length of the shaft and cap- sule ratio as described above. I believe the p-rhabdoids A and D belong to the same unit, as suggested by den Hartog (1980). Problems in fitting cnidae into a previous system of classification led different authors to propose their own system (see synony- mies here). However, some of these au- thors, and others who used these systems, did not always clearly define and illustrate the types they were working with. As a re- sult, the usefulness of some papers on cni- dae was partly lost. Misunderstandings re- lated to the use of different terminologies must be avoided by using more precise de- scriptions and illustrations, based on large arrays of material. Corals and corallimorpharians present the four basic morphological patterns of cnidae found in Hexacorallia, with the most diversified cnidae: spirocysts, holotrichs, b- and p-rhabdoids. Agaricysts were observed only in Agariciidae. In agaricysts, the ar- rangement of the undischarged tubule with- in the capsule and the appearance of the tubule following discharge look very simi- lar to that of the heteroptychonemes de- scribed by Mariscal et al. (1977). The study of its ultrastructure in eletronic microscopy would be very helpful for describing it more precisely. The evidence of the diversity occurring in the cnidae of Scleractinia presented in this paper support the important role of soft tissues as a source of useful taxonomic characters. The inclusion of morphological descriptions of cnidae in further studies will contribute to improving studies on coral taxonomy and systematics. Acknowledgments This study would not be possible without the donations of specimens obtained through Dr. J. C. den Hartog (Nationaal Na- tuurhistorisch Museum, Leiden), Dr. A. M. S. Vanin (Instituto Oceanografico, Univer- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sidade de Sao Paulo, Sao Paulo), Dr. S. D. Cairns (National Museum of Natural His- tory, Smithsonian Institution—USNM, Washington, D. C.), Dr. C. Hand (Bodega Marine Laboratory, California) and Dr. A. Herrera (Academia de Ciéncias de Cuba, Habana). I gratefully acknowledge useful sugges- tions to the manuscript given by Dr. E. Schlenz, Dr. E L. da Silveira, Dr. A. E. Migotto, and A. C. Marques (Universidade de Sao Paulo, Sao Paulo), Dr. C. B. Castro and Dr. G. W. Nunan (Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro) and Dr. S. D. Cairns (USNM). I thank Dr. R. N. Mariscal (Flor- ida State University, Florida), who contrib- uted with helpful discussion on the agari- cysts. Dr. E. Schlenz offered invaluable help with translations of papers in German. I also thank Dr. S. D. Cairns for facilities during my visit to the USNM and for the identification of some ahermatypic corals. Finally, I thank Funda¢gao Universitaria José Bonifacio—FUJB and Conselho Nacional de Desenvolvimento Cientifico e Tecnol6- gico—CNPgq for financial support and Fun- dacao Coordenacgao de Aperfeigoamento de Pessoal de Nivel Superior—CAPES, Brazil, for the doctorate fellowship and for travel support to visit the USNM, Washington. Literature Cited Abe, N. 1938. Feeding behaviour and the nematocysts of Fungia and 15 other species of corals.—Pa- lao Tropical Biological Station Studies 3:469-— SQ, Cairns, S. D. 1990. Antarctic Scleractinia. Pp.1—78 in J. W. Wagele & J. Sieg, eds., Synopses of the Antarctic Benthos. Vol. 1. Koeltz Scientific Books, Koenigstein. Carlgren, O. 1940. A contribution to the knowledge of the structure and distribution of the cnidae in the Anthozoa.—Lunds Universitets Arsskrift. N. FE Avd. 2, Bd 36, Nr 3:1—62. Chevalier, J. PR, & L. Beauvais. 1987. Pp. 679-764 in P. P. Grassé, ed., Systématique. Traité de Zool- ogie. Anatomie, systématique, biologie. Tome III. 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Lenhoff, eds., Coelen- terate Biology: Reviews and New Perspectives. Academic Press, New York. . 1984. Cnidaria: cnidae. Pp. 57—68 in J. Ber- eiter-Hahn, A. G. Matoltsy & K. S. Richards, eds., Biology of integument. Vol. 1. Inverte- brates. Springer-Verlag, Berlin. WEewaiConkim dé CoH, Bigger 1977. The ptychocyst, a major new category of cnida used in tube construction by a cerianthid anemone.— Biological Bulletin 152:392—405. Matthai, G. 1914. A revision of the recent colonial astraeidae possessing distinct corallites. Reports of the Percy Sladen Trust Expedition to the In- dian Ocean in 1905 under the leadership of Mr. J. Stanley Gardiner, Vol. VI.—The Transactions of the Linnean Society of London (Zoology) 2nd series, 17:1—140. . 1928. A monograph of the recent meandroid Astraeidae.—Catalogue of the Madreporarian corals in the British Museum (Natural History) 7:1—288 Ostman, C. 1987. Pp. 67-82 in J Bouillon, E Boero, E Cicogna, & P. E S. Cornelius, eds., New tech- niques and old problems in hydrozoan system- atics. Modern Trends in the Systematics, Ecol- ogy and Evolution of Hydroids and Hydrome- dusae, Oxford University Press, New York, 328 Pp. Pires, D. O. 1988. Tricnidactis errans n. gen, Nn. sp. (Cnidaria, Actiniaria, Haliplanellidae), from Guanabara Bay, Rio de Janeiro, Brazil.—Revis- ta Brasileira de Biologia 48(3):507-516. , & C. B. Castro. 1997. Scleractinia and Cor- allimorpharia: an analysis of cnidae affinity.— Proceedings of the 8th International Coral Reef Symposium, In press. , & E B. Pitombo. 1992. Cnidae of the Bra- 181 zilian Mussidae (Cnidaria: Scleractinia) an their value in taxonomy.—Bulletin of Marine Sci- ence 51(2):231—244. Schlenz, E., & M. J. C. Belém. 1992. Phyllactis cor- reae n. sp. (Cnidaria, Actiniaria, Actiniidae) from Atol das Rocas, Brazil, with notes on Phyllactis flosculifera (Lesueur, 1817).—Bole- tim de Zoologia, Universidade de Sao Paulo 12: 91-117. Schmidt, H. 1969. 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Tilbury, R., & A. M. Cameron. 1989. Cnidom of Goniopora species in 5th International Confer- ence on Coelenterate Biology. Programme and abstracts. P. 95. July 10-14, 1989, University of Southampton, Southampton. Vervoort, W. 1987. Pp. 83-103 in J. Bouillon, EF Boe- ro, EK Cicogna, & P. FE S. Cornelius, eds., Eval- uation of taxonomic characters in the Hydroida, particularly in the Thecata (=Leptomedusae). Modern Trends in the Systematics, Ecology and Evolution of Hydroids and Hydromedusae, Ox- ford University Press, New York, 328 pp. Weill, R. 1934. Contribution a l’étude des cnidaires et de leurs nématocystes. I, Il.—Travaus de la Station zoologique de Wimereux 10 & 11:1- AON: Wells, J. W. 1956. Scleractinia. Pp. 328-477 in R. C. Moore, ed., Treatise on Invertebrate Paleontol- ogy, part EK Geological Society of America and Kansas University Press, Lawrence, 498 pp. Appendix Collection data of studied material from the Museu Nacional, Universidade Federal do Rio de Janeiro 182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (MNRJ). Donations of some specimens were obtained from the Instituto Oceanogrdafico, Universidade de Sao Paulo (IO/USP), through Dr. A. M. S. Vanin; from the Nationaal Natuurhistorisch Museum (Rijksmuseum van Natuurlijke Historie) (RMNH), through Dr. J. C. den Hartog; from the National Museum of Natural His- tory, Smithsonian Institution (USNM), through Dr. S. D. Cairns; from the Bodega Marine Laboratory (BML) through Dr. C. Hand and from the Academia de Cién- cias de Cuba (ACC), through Dr. A. Herrera. Agaricia agaricites (Linnaeus, 1758). MNRJ 02686. Brazil, Bahia, Nova Vigosa, Coroa Vermelha Reef, coll. D. O. Pires 26 Jan 1991, det. D. O. Pires 3 Nov 1994. Agaricia agaricites (Linnaeus, 1758). MNRJ 02051. Brazil, Bahia, Prado, Coroa de Cumuruxatiba Reef, coll. C. B. Castro Jan 1992, det. D. O. Pires Jan 1992. Agaricia agaricites (Linnaeus, 1758). MNRJ 02052. Brazil, Bahia, Prado, Coroa de Cumuruxatiba Reef, coll. C. B. Castro Jan 1992, det. D. O. Pires Jan 1992. Agaricia agaricites (Linnaeus, 1758). MNRJ 02684. Brazil, Bahia, Abrolhos, SE Pedra Lixa, 16 m, coll. D. O. Pires 25 Jan 1991, det. D. O. Pires 3 Nov 1994. Agaricia agaricites (Linnaeus, 1758). MNRJ 02685. Brazil, Bahia, Abrolhos, SE Pedra Lixa, 16 m, coll. D. O. Pires 25 Jan 1991, det. D. O. Pires 3 Nov 1994. Agaricia fragilis Dana, 1848. MNRJ 02681. Brazil, Bahia, Prado, Aprofundados de Cumuruxatiba Reefs, coll. P. S. Young 30 Jan 1991, det. D. O. Pires 1 Nov 1994. Astrangia rathbuni Vaughan, 1906. MNRJ 02600. Bra- zil, Rio de Janeiro, Angra dos Reis, Queimada Pe- quena Island, coll. Coelenterology/MNRJ 6 Dec 1992, det. D. O. Pires 31 Aug 1994. Astrangia rathbuni Vaughan, 1906. MNRJ 01093. Bra- zil, Santa Catarina, Porto Belo, Quatro Ilhas Beach, coll. D. O. Pires 25 Jan 1986, det. D. O. Pires. Astrangia rathbuni Vaughan, 1906. MNRJ 02615. Bra- zil, Rio de Janeiro, Guanabara Bay, near “‘lage do Cacao’, coll. P. S. Young 19 Jun 1991, det. D. O. Pires 8 Sep 1994. Balanophyllia europaea (Risso, 1826). MNRJ 00090. Spain, Mediterranean, Cala Reona, Cabo Depalos, 1-2 m, coll. H. Zibrowius, 9 Apr 1981, det. H. Zi- browius. Obs.: donation RMNH. Balanophyllia sp. MNRJ 02377. Brazil, Bahia, Prado, Aprofundados de Cumuruxatiba Reefs. 17°00’S, 039°05'W, 18 m, coll. C. B. Castro 30 Jan 1991, det. S. D. Cairns. Obs.: donation USNM (USNM 90329). Caryophyllia ambrosia caribbeana Cairns, 1979. MNRJ 02376. Brazil, Sao Paulo, off Couves Island. 24°35.05’S, 044°12’W, 600 m, coll. R/V Prof. W. Besnard 7 Dec 1988, det. P. S. Young & D. O. Pires 2 Mar 1994. Obs.: donation IO/USP. Caryophyllia cornuformis Pourtalés 1868. MNRJ 02442. Brazil, Sao Paulo, off Couves Island. 24°35.05'S, 044°12’W, 600 m, coll. R/V Prof. W. Besnard 7 Dec 1988, det. S. D. Cairns May 1991. Obs.: donation IO/USP. Cladocora arbuscula (Lesueur, 1821) MNRJ 02550. USA, off Florida, Gulf of Mexico. 25°45.53'N, 082°31.37’W coll. Continental Shelf Ass. MMS/ BLM 9 May 1983, det. S. D. Cairns. Obs.: donation USNM (USMN 71945). Cladocora debilis Milne Edwards & Maime, 1849. MNRJ 02407. Brazil, Sao Paulo, off Couves Island. 24°23.03'S, 044°18'W, 240 m, coll. R/V Prof. W. Besnard 8 Dec 1988, det. P. S. Young & D. O. Pires 7 Mar 1994. Obs.: donation IO/USP. Corynactis californica Carlgren, 1936. MNRJ 02130. USA, California, Bodega Harbor, coll. C. Hand, det. C. Hand. Obs.: donation BML. Corynactis sp. MNRJ 01453. Brazil, Rio de Janeiro, _Rio de Janeiro, Cagarras Islands, 20.5 m, coll. C. C. Ratto 19 Nov 1988. (?) Cycloseris marginata (Boschma, 1923). MNRJ 02118. Indonesia, SW Sulawesi, coll. B. W. Hoek- sema, 2 May 1985. Obs.: donation RMNH. Dasmosmilia lymani (Pourtalés, 1871). MNRJ 02372. Brazil, Sao Paulo, off Couves Islands. 24°25’S, 044°16.05’W, 320 m, coll. R/V Prof. W. Besnard 7 Dec 1988, det. P. S. Young & D. O. Pires 2 Mar 1994. Obs.: donation IO/USP. Dasmosmilia variegata (Pourtalés, 1871). MNRJ 02373. Brazil, Sao Paulo, off Couves Islands. 24°25'S, 044°16.05'W, 320 m, coll. R/V Prof. W. Besnard 7 Dec 1988, det. P. S. Young & D. O. Pires 2 Mar 1994. Obs.: donation IO/USP. Deltocyathus calcar Pourtalés, 1874. MNRJ 02378. Brazil, Sado Paulo, off Sao Sebastiao Island. 24°42.05’S, 044°30’W, 320 m, coll. R/V Prof. W. Besnard 6 Dec 1988, det. P. S. Young & D. O. Pires 3 Mar 1994. Obs.: donation IO/USP. Deltocyathus calcar Pourtalés, 1874. MNRJ 02380. Brazil, Sao Paulo, off Vitdéria Island. 24°31’S, 044°28'W, 240 m, coll. R/V Prof. W. Besnard 8 Dec 1988, det. P S. Young & D. O. Pires 3 Mar 1994. Obs.: donation IO/USP. Deltocyathus eccentricus Cairns, 1979. MNRJ 02409. Brazil, Sao Paulo, off Vit6ria Island. 24°41.01’S, 044°18.05’W, 510 m, coll. R/V Prof. W. Besnard 7 Dec 1988, det. S. D. Cairns May 1994. Obs.: do- nation IO/USP. Deltocyathus eccentricus Cairns, 1979. MNRJ 02411. Brazil, Sao Paulo, off Couves Island. 24°35.05’S, 044°12’W, 600 m, coll. R/V Prof. W. Besnard 7 Dec 1988, det. S. D. Cairns May 1994. Obs.: donation IO/USP. Deltocyathus italicus (Michellotti, 1838). MNRJ 02412. Brazil, Sado Paulo, off Vitoria Island. VOLUME 110, NUMBER 2 24°41.01’S, 044°18.05’W, 510 m, coll. R/V Prof. W. Besnard 7 Dec 1988, det. S. D. Cairns May 1994. Obs.: donation IO/USP. Deltocyathus italicus (Michellotti, 1838). MNRJ 02413. Brazil, Sado Paulo, off Couves Island. 24°35.05’S, 044°12'W, 600 m, coll. R/V Prof. W. Besnard 7 Dec 1988, det. S. D. Cairns May 1994. Obs.: donation IO/USP. Dendrophyllia sp. MNRJ 02352. Cabo Verde, S coast of Sao Tiago, SE Porto. 14°55’N, 023°30’W, 15 m, coll. 5 Jun 1982. Obs.: CANCAP VI 1982, Sta. 6 DO1 “‘Tydan’’, donation RMNH. Discosoma carlgreni (Watzl, 1922). MNRJ 00331. Brazil, Espirito Santo, Guarapari, Trés Ilhas Archi- pelago, coll. S. Rosso et al. 17 Jul 1981, det. E. Schlenz & M. J. C. Belém 15 Oct 1981. Discosoma carlgreni (Watzl, 1922). MNRJ 02259. Brazil, Bahia, Abrolhos Archipelago, coll. E B. Pi- tombo 23 Nov 1993, det. S. M. Pinto 8 Dec 1993. Favia leptophylla Verrill, 1868. MNRJ 02280. Brazil, Bahia, Abrolhos, Parcel das Paredes Reef, 5-10 m, coll. D. O. Pires et al. 24 Jan 1991, det. D. O. Pires 29 Dec 1993. Favia leptophylla Verrill, 1868. MNRJ 02574. Brazil, Bahia, Abrolhos, SE Pedra Lixa Reef, coll. Coelen- terology/MNRJ 25 Jan 1991, det. D. O. Pires 26 Aug 1994. Favites abdita (Ellis & Solander, 1786). MNRJ 02342. Australia, Queensland, Swain Reefs Complex, Sweetlip Reef, 3—5 m, coll. P. Alderslade 4 Jul 1981, det. E B. Pitombo 25 Jan 1981 Obs.: donation RMNH. Flabellum sp. MNRJ 02720. Off Guiana. 07°56’N, 057°12'W, 618 m, coll. ““Luymes” Guyana Shelf Expedition 1 Sep 1970. Obs.: donation RMNH. Fungia (Cycloseris) sinensis Milne Edwards & Haime, 1851. MNRJ 02116. Indonesia, SW Sulames, coll. B. W. Hoeksena, 2 May 1985, det. B. W. Hoeksema 2 May 1985. Obs.: donation RMNH. Fungia (Fungia) fungites (Linnaeus, 1758). MNRJ 02679. Indonesia, S Sulawesi, N side of Samalona, 2-5 m, coll H. Moll 20 Nov 1980. Obs.: donation RMNH. Fungia (Verrillofungia) repanda Dana, 1846. MNRJ 02120. Indonesia, S Sulawesi, N side of Samalona, coll. H. Moll, 20 Nov 1980, det. H. Moll, 20 Nov 1980. Obs.: donation RMNH. Fungia (Verrillofungia) concinna (Verrill, 1864). MNRJ 02113. Indonesia, reef S. of Pilau Tikus, NW Java, coll. B. W. Hoeksema, 7 May 1983, det. B. W. Hoeksema, 7 May 1983. Obs.: conation RMNH. Fungiacyathus sp. MNRJ 02391. Brazil, Sao Paulo, off Vitoria Island. 24°31'S, 044°28’W, 250 m, coll. R/V Prof. W. Besnard 8 Dec 1988. Obs: donation IO/USP. Fungiacyathus sp. MNRJ 02393. Brazil, Sao Paulo, off Ilha de Sao Sebastiao, 320 m, coll. R/V Prof. W. Besnard 6 Dec 1988. Obs.: donation IO/USP. 183 Goniopora tenuidens (Quelch, 1886). MNRJ 02459. Australia, Queensland, Swain Reefs Complex, Sweetlip Reef, 3—5 m, coll. P. Alderslade 4 Jul 1981, det. D. O. Pires May 1994. Obs.: donation RMNH. Halomitra pileus (Linnaeus, 1758). MNRJ 02119. In- donesia, North of Pulau Tikus, NW Java, coll. B. W. Hoeksema 11 May 1983, det. B. W. Hoeksema 11 May 1983. Obs.: donation RMNH. Herpolitha limax Esper, 1797. MNRJ 02115. Indone- sia, W of Pulau Tikus, NW Java, coll. B. W. Hoek- sema 10 May 1985, det. B. W. Hoeksema, 10 May 1985. Obs.: donation RMNH. Herpolitha limax Esper, 1797. MNRJ 02111. Indone- sia, N side of Pulau Pajung, coll. B. W. Hoeksema 12 May 1983, det. B. W. Hoeksema 12 May 1983. Obs.: donation RMNH. Javania cailleti (Duchassaing & Michelotti, 1864). MNRJ 02383. Brazil, SAo Paulo, off Sao Sebastiao Island. 24°35.04’S, 044°33.03'W, 184 m, coll. R/V Prof. W. Besnard 6 Dec 1988, det. PR S. Young & D. O. Pires 3 Mar 1994. Obs.: donation IO/USP. Javania cailleti (Duchassaing & Michelotti, 1864). MNRJ 02384. Brazil, SAo Paulo, off Vitéria Island. 24°23.02'S, 044°24.08’W, 180 m, coll. R/V Prof. W. Besnard 20 Jul 1987, det. PR. S. Young & D. O. Pires 3 Mar 1994. Obs.: donation IO/USP. Leptopenus antarcticus Cairns, 1989. MNRJ 02554. Antarctica, Ross Sea, approx. 200 Km W of Pennel Bank. 74°55’S, 174°12’W, 2022-2060 m, coll. R/V Eltanin, Usarp 7 Feb 1988, det. S. D. Cairns. Obs.: donation USNM (USNM 47482, 2 paratypes). Leptoseris cucullata (Ellis & Solander, 1786). MNRJ 02561. Venezuela, Curagao, Isle Bay, 20 m, coll. R. P. M. BAK. Obs.: donation P. S. Young (MNRJ). Lithophyllum mokai Hoeksema, 1989 MNRJ 02114. Indonesia, SW Sulawesi, coll. B. W. Hoeksema 2 May 1985. Obs.: donation RMNH. Lobophyllia hemprichii (Ehrenberg 1834). MNRJ 02129. Seychelles, Alphonse Atoll, SE of the la- goon. 07°02'S, 052°44’E, 0-8 m, coll. R/V Tyro Seychelles Expedition 4 Jan 1993, det. J. C. den Hartog. Obs.: Netherlands Indian Ocean Expedition, donation RMNH. Lobophyllia sp. MNRJ 02677. Australia, Queensland, Swain Reefs Complex, Sweetlip Reef, 3—5 m, coll. P. Alderslace 4 Jul 1981. Obs.: donation RMNH. Madracis decactis (Lyman, 1859). MNRJ 02125. Bra- zil, Bahia, Parcel das Paredes Reefs, SE Pedra Lixa, coll. Coelenterology/MNRJ 25 Jan 1991, det. D. O. Pires 25 Jan 1991. Madracis decactis (Lyman, 1859). MNRJ 02070. Bra- zil, Rio de Janeiro, Angra dos Reis, Queimada Pe- quena Island, coll. D. O. Pires et al. 6 Dec 1992, det. D. O. Pires 28 Dec 1992. Madracis decactis (Lyman, 1859). MNRJ 02124. Bra- zil, Bahia, Abrolhos, Parcel das Paredes Reef, 10 m, coll. Coelenterology/MNRJ 24 Jan 1991, det. D. O. Pires 24 Jan 1991. 184 Madracis decactis (Lyman, 1859). MNRJ 02127. Bra- zil, Bahia, Prado, Aprofundados de Cumuruxatiba Reefs, coll. Coelenterology/MNRJ 30 Jan 1991, det. D. O. Pires 30 Jan 1991. Madracis decactis (Lyman, 1859). MNRJ 02680. Bra- zil, Rio de Janeiro, Angra dos Reis, ‘“‘Ponta do Can- tador’’, coll. E B. Pitombo Feb 1989, det. D. O. Pires 1 Nov 1994. Meandrina braziliensis (Milne Edwards & Haime, 1848). MNRJ 02275. Brazil, Bahia, Abrolhos, Par- cel das Paredes Reef, 5—10 m, coll. D. O. Pires et al. 24 Jan 1991, det. D. O. Pires 21 Dec 1993. Meandrina _ braziliensis (Milne Edwards & Haime, 1848). MNRJ 02276. Brazil, Bahia, Abrolhos, Par- cel das Paredes Reef, 5—10 m, coll. Coelenterology/ MNRJ 24 Jan 1991, det. D. O. Pires 21 Dec 1993. Meandrina braziliensis (Milne Edwards & Haime, 1848). MNRJ 02277. Brazil, Bahia, Abrolhos, Par- cel das Paredes Reef, 5—10 m, coll. D. O. Pires et al. 24 Jan 1991, det. D. O. Pires 21 Dec 1993. Meandrina braziliensis (Milne Edwards & Haime, 1848). MNRJ 02278. Brazil, Bahia, Abrolhos, Par- cel das Paredes Reef, coll. D. O. Pires et al. 24 Jan 1991, det. D. O. Pires 21 Dec 1993. Montastrea cavernosa (Linnaeus, 1767). MNRJ 01441. Brazil, Bahia, Abrolhos Archipelago, Santa Barbara Island, 5.2 m, coll. A. C. Marques & E D. Amaral 7 Mar 1989, det. E D. Amaral 5 Apr 1989. Montastrea cavernosa (Linnaeus, 1767). MNRJ 01442, Bahia, Abrolhos Archipelago, Santa Barbara Island, 4 m, coll. A. C. Marques & E D. Amaral 7 Mar 1989, det. E D. Amaral 5 Apr 1989. Montastrea cavernosa (Linnaeus, 1767). MNRJ 01520. Brazil, Bahia, Abrolhos Archipelago, Santa Barbara Island, 3 m, coll. E D. Amaral & S. M. Pinto 5 Aug 1989, det. E D. Amaral 15 Aug 1989. Montastrea cavernosa (Linnaeus, 1767). MNRJ 02350. Brazil, Bahia, Abrolhos, Parcel das Paredes Reef, 5-10 m, coll. Coelenterology/MNRJ 24 Jan 1991, det. D. O. Pires 21 Dec 1993. Montastrea cavernosa (Linnaeus, 1767). MNRJ 02612. Brazil, Bahia, Abrolhos, SE Pedra Lixa, 16 m, coll. Coelenterology/MNRJ 25 Jan 1994, det. D. O. Pires 08 Sep 1994. Montastrea cavernosa (Linnaeus, 1767). MNRJ 02676. Brazil, Bahia, Abrolhos, Parcel das Paredes Reef, 5—10 m, coll. D. O. Pires 24 Jan 1991, det. D. O. Pires 19 Oct 1994. Mussismilia braziliensis (Verrill, 1868). MNRJ 01704. Brazil, Bahia, Abrolhos Archipelago, Santa Barbara Island, coll. EK B. Pitombo 3 Apr 1989, det. E B. Pitombo 18 Apr 1990. Mussismilia braziliensis (Verrill, 1868). MNRJ 01710. Brazil, Bahia, Abrolhos Archipelago, Santa Barbara Island, Portinho, coll. E B. Pitombo 5 Apr 1989, det. EK B. Pitombo 18 Apr 1980. Mussismilia braziliensis (Verrill, 1868). MNRJ 01717. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Brazil, Bahia, Morro de S4o Paulo, coll. E B. Pi- tombo 22 Jan 1989, det. EF B. Pitombo 24 Apr 1980. Mussismilia hartti (Verrill, 1868). MNRJ 01707. Bra- zil, Bahia, Abrolhos Archipelago, Santa Barbara Is- land, coll. EK B. Pitombo 4 Apr 1989, det. E B. Pi- tombo 18 Apr 1990. Mussismilia hartti (Verrill, 1868). MNRJ 01709. Bra- zil, Bahia, Abrolhos Archipelago, Siriba Island, coll. FE B. Pitombo 5 Apr 1989, det. E B. Pitombo 18 Apr 1980. Mussismilia hartti (Verrill, 1868). MNRJ 01746. Bra- zil, Bahia, Morro de Sao Paulo, coll. EF B. Pitombo 14 Mar 1990, det. EF B. Pitombo 26 Jun 1990. Mussismilia hispida (Verrill, 1902). MNRJ 01711. Brazil, Bahia, Abrolhos Archipelago, Santa Barbara Island, coll. EF B. Pitombo 5 Apr 1989, det. FE B. Pitombo 18 Apr 1980. Mussismilia hispida (Verrill, 1902). MNRJ 01718. Brazil, Pernambuco, Porto de Galinhas, coll. FE B. Pitombo 6 Mar 1990, det. E B. Pitombo 26 Apr 1990. Mussismilia hispida (Verrill, 1902). MNRJ 01719. Brazil, Rio de Janeiro, Angra dos Reis, Grande Is- land, coll. E B. Pitombo 31 Apr 1990, det. E B. Pitombo. Oculina patagonica de Angelis, 1908. MNRJ 02351. Spain, Mediterranean, Portman, appr. 20 km E of Cartagena, 1 m, coll. H. Zibrowius 10 Apr 1981, det. H. Zibrowius 31 Jan 1994. Obs.: donation RMNH. Pavona sp. MNRJ 02576. Indonesia, S. Sulawesi, N side of Samalona, 2—5 m, coll. H. Moll 20 Nov 1980. Obs.: donation RMNH. Phyllangia americana Milne Edwards & Haime, 1850. MNRJ 02563. Brazil, Rio de Janeiro, Angra dos Reis, Botinas Islands, 2 m, coll. E B. Pitombo 30 Jul 1991, det. D. O. Pires 26 Aug 1994. Phyllangia americana Milne Edwards & Haime, 1850. MNRJ 01056. Brazil, Bahia, Morro de Sao Paulo. 3 m, coll. E B. Pitombo 4 Mar 1987, det. P. S. Young 24 Apr 1987. Phyllangia americana Milne Edwards & Haime, 1850. MNRJ 02601. Brazil, Rio de Janeiro, Angra dos Reis, Queimada Pequena Island, coll. Coelenterol- ogy/MNRJ 6 Dec 1992, det. D. O. Pires 31 Aug 1994. Phyllangia americana Milne Edwards & Haime, 1850. MNRJ 02611. Brazil, Bahia, Abrolhos, Nova Vicosa Reef. 18°0047'S, 039°16.57'W, 9 m, coll. C. C. Rat- to & C. B. Castro 23 Aug 1994, det. D. O. Pires 8 Sep 1994. Physogyra lichtensteini Milne Edwards & Haime, 1851. MNRJ 02272. Seychelles, Alphonse Atoll, parte SE of the lagoon, STA SEY 788. 07°02’S, 052°44’'E, 8 m, coll. R/V Tyro 4 Jan 1993, det. J. C. den Hartog 16 Dec 1993. Obs.: Netherlands In- dian Ocean Expedition 1992/93, donation RMNH. Physogyra lichtensteini Milne Edwards & Haime, VOLUME 110, NUMBER 2 1851. MNRJ 02273. Seychelles, St. Joseph Atoll, NW Rim lagoon, STA SEY 754. 05°24’S, 053°19’E, coll. R/V Tyro 26 Dec 1992, det. J. C. den Hartog 16 Dec 1993. Obs.: Netherlands Indian Ocean Ex- pedition 1992/93, donation RMNH. Platygyra daedalea (Ellis & Solander, 1786). MNRJ 02346. Australia, Queensland, Swain Reefs Com- plex, Sweetlip Leef, 3-5 m, coll. P. Alderslade 4 Jul 1981, det. E B. Pitombo 25 Jan 1994 Obs.: donation RMNH. Polyphyllia talpina (Lamarck, 1801). MNRJ 02112. Indonesia, Spermonde Archipelago, SW Sulawesi, coll B. W. Hoeksema 1985, det. B. W. Hoeksema 1985. Obs.: donation RMNH. Porites astreoides Lamarck, 1816. MNRJ 02281. Bra- zil, Bahia, Abrolhos, Parcel das Paredes Reef, SE Pedra Lixa, 2—16 m, coll. D. O. Pires et al. 25 Jan 1991, det. D. O. Pires 29 Dec 1993. Porites astreoides Lamarck, 1816. MNRJ 02354. Bra- zil, Bahia, Nova Vicosa Reef, coll. C. B. Castro & P. S. Young Aug 1993, det. C. C. Ratto 31 Jan 1994. Porites astreoides Lamarck, 1816. MNRJ 02713. Bra- zil, Bahia, Abrolhos, Parcel das Paredes Reef, 5—10 m, coll. Coelenterology/MNRJ 24 Jan 91, det. D. O. Pires 20 Mar 1995. Porites astreoides Lamarck, 1816. MNRJ 02714. Bra- zil, Bahia, Abrolhos, Parcel das Paredes Reef, 5—10 m, coll. Coelenterology/MNRJ 24 Jan 91, det. D. O. Pires 20 Mar 1995. Rhisosmilia maculata (Pourtalés, 1874). MNRJ 02390. Brazil, Bahia, Prado, Aprofundados de Cumuruxa- tiba Reefs, coll. C. B. Castro 30 Jan 1991, det. S. D. Cairns 4 Mar 1994. Ricordea florida Duchassaing & Michelotti, 1860. MNRJ 02108. Cuba, La Juventud Island, Punta del 185 Este Reef, 10 m, coll. A. Herrera 26 Aug 1992, det. M. J. C. Belém 21 Aug 1992. Obs.: donation ACC. Scolymia wellsi Laborel, 1967. MNRJ 01198. Brazil, Bahia, Parcel dos Abrolhos Reef, Rosalina Ship wreck., 17 4 m, coll. Coelenterology/MNRJ 8 Oct 1987, det. C. C. Ratto 3 Nov 1987. Scolymia wellsi Laborel, 1967. MNRJ 01713. Brazil, Bahia, Abrolhos Archipelago, Guarita Island, coll. E B. Pitombo 6 Apr 1989, det. E B. Pitombo 23 Apr 1980. Scolymia wellsi Laborel, 1967. MNRJ 01714. Brazil, Bahia, Abrolhos Archipelago, Guarita Island, coll. E B. Pitombo 6 Apr 1989, det. E B. Pitombo 24 Apr 1990. Siderastrea radians (Pallas, 1766). MNRJ 01555. Mexico, Quintana Roo, Punta Brava, coll. FE D. Amaral 11 Nov 1989, det. E D. Amaral 7 Dec 1989. Siderastrea stellata Verrill, 1868. MNRJ 02675. Bra- zil, Bahia, Abrolhos, Parcel das Paredes Reef, 5—10 in,.coll,..D,.O-. Pires..24 Jan 1991, -det..D.,O;. Pires 19 Oct 1994. Siderastrea stellata Verrill, 1868. MNRJ 02271. Bra- zil, Rio de Janeiro, Cabo Frio, Buzios, coll. E Pi- tombo et al. 2 Feb 1993, det. E B. Pitombo 5 Dec 1993. Siderastrea stellata Verrill, 1868. MNRJ 02283. Bra- zil, Bahia, Abrolhos, Parcel das Paredes Reef, SE Pedra Lixa coll. D. O. Pires 25 Jan 1991, det. D. O. Pires 25 Jan 1991. Stephanocoenia michelinii Milne Edwards et Haime, 1848. MNRJ 02121. Brazil, Bahia, Parcel das Pa- redes Reef, SE Pedra Lixa, coll. Coelenterology/ MNRJ 25 Jan 1991, det. D. O. Pires 25 Jan 1991. Zoopilus echinatus Dana, 1846. MNRJ 02117. Indo- nesia, SW Sulawesi, coll. B. W. Hoeksema 2 May 1985. Obs.: donation RMNH. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):186—197. 1997. Gnathostomulida from the Canary Islands Wolfgang Sterrer Bermuda Natural History Museum, Flatts FL BX, Bermuda e-mail wsterrer @ bbsr.edu Abstract.—Five species of Gnathostomulida are reported from Gran Canaria (northeastern Atlantic). One, Paragnathiella trifoliceps new genus, new spe- cies, represents a new genus in the family Mesognathariidae, another a new species, Austrognathia clavigera; one is known from the northwestern North Atlantic (Labidognathia longicollis), and the remaining two are cosmopolitan (Haplognathia rosea and H. ruberrima). As part of an ongoing investigation of the taxonomy and biogeography of Gna- thostomulida (Sterrer et al. 1986, Sterrer POST aS tbe 199 te 299d, 1992-0997), a phylum of microscopic worms inhabiting marine sand, I collected sediment samples in the Canary Islands (northeastern Atlan- tic) in February 1996. Of ten samples from eight localities on two islands, only one lo- cality produced Gnathostomulida (Table 1). Samples were collected by hand, snorkeling or SCUBA, and extracted as described ear- lier (Sterrer 1971, 1991la). The way in which specimens and species are analyzed and described follows Sterrer (1991a). This includes the use of a relative scale of 100 units (U) for the body length, various in- dices for length-width ratios, and the mean (X), standard deviation (SD), maximum (Max), minimum (Min), and number (n) of measurements. Order Filospermoidea Sterrer, 1972 Composition.—Two families, Haplogna- thiidae Sterrer, 1972, and Pterognathiidae Sterrer, 1972 emend. Sterrer, 1991a. Family Haplognathiidae Sterrer, 1972. One genus, Haplognathia Sterrer, 1970. Haplognathia rosea (Sterrer, 1969) (Figs. 1A, 2A) Material.—One juvenile from Gran Ca- naria, Playa de Las Canteras (sample C1). Description.—The faintly reddish speci- men had jaws 20 pm long, with 9 wm long rostral apophyses (apophysis index 0.45). Of the basal plate, only the rostral outline could be recorded, as well as the lack of teeth or thorns. Discussion.—Sterrer (1997) merged H. rosacea Sterrer, 1970 with H. rosea, which is now known from the North Sea (Sterrer 1969), the northwestern Atlantic (Sterrer 1997), and the southern Pacific (Sterrer 1991a, 1991c). This species, which is dif- ficult to distinguish from, and possibly hy- bridizes with H. ruberrima, has been rede- fined as having jaws with rostral apophyses less than half as long as the jaws (index 0.5 or smaller), and a basal plate that lacks thorns. In the northwestern Atlantic, jaw length ranges from 15 to 23 wm, with a mean of 18.89 wm. Haplognathia ruberrima (Sterrer, 1966) (Figs. 1B—G, 2B-—C, Table 2) Material.—Nine specimens, of which one adult, from Gran Canaria, Playa de Las Canteras (sample C1). Description.—The only adult was 2150 wm long and 75 wm wide at U 41.9 (index VOLUME 110, NUMBER 2 187 28.67). The smallest intact specimen mea- sured 310 pm by 35 um, with a rostrum 140 pm by 35 pm; these are the dimensions of a recently hatched juvenile (Sterrer 1997). Most of the specimens were colour- Gnathostomulida spp 5 1 i oe : = less to faintly pink; only one was red. All 9 3 ie 9 ec had a basal plate in the shape of a trans- = Ba83 * 9 . verse, ventrally concave buckle (length 6— E a eee 9 wm, width 8-13 pm; index 0.57), with 5 e 2 O a 5 7 3 Es rows of regularly spaced thorns on the dor- E a 6 aie Z a a E sal surface. The jaws are 22-29 um long Ae ee ee fa (X = 25.57 pm), with well delimited rostral See eee ee oe apophyses whose length usually equals or ea eis =e Se exceeds the jaw length. The jaws have one Se eeew cs & ses Ss & s . eer ae an. ao strong tooth, possibly with numerous small, SESSEEEEEE thorn-like bristles surrounding it. A pair of pharyngeal glands, with fine granula and large vacuoles, is typical for this species Ss a ; a E = = Es (Fig. 1G). ; AIM EEE RMNMH E Discussion.—The most cosmopolitan ao 2 = ae . . . cre species of the entire phylum is now known from the North Sea and Mediterranean eee op lol 0 © (Sterrer 1969), the northwestern Atlantic 2 r 2 2 2 2 2 2 2 2 2 (Sterrer 1997), and the Pacific islands of Pik Fs Ba Gs Ae Peles Be i Be De Hawaii (Sterrer 1991b) and Fiji (Sterrer Be ee ee es eyed Cy 1991a). It has been redefined as having jaws with rostral apophyses at least half as long as the jaws (index 0.5 or greater), and a basal plate that always bears thorns. The Canaries specimens fall in every respect within the limits of this highly variable spe- cies. Sample Cl C C3 C4 C C7 C e C10 Ll Order Bursovaginoidea Sterrer, 1972 Composition.—Two suborders, Sclero- peralia Sterrer, 1972, and Conophoralia Sterrer, 1972. Locality Suborder Scleroperalia Sterrer, 1972 Composition.—Eight families, Agna- thiellidae Sterrer, 1972; Mesognathariidae Sterrer, 1972; Clausognathiidae Sterrer, 1992; Gnathostomariidae Sterrer, 1972; Rastrognathiidae Kristensen & N@grrevang, 1977; Problognathiidae Sterrer & Farris, 1975; Onychognathiidae Sterrer, 1972; and Gnathostomulidae Sterrer, 1972. Las Palmas, Playa de Las Canteras Las Palmas, Playa de Las Canteras Arinaga Las Palmas, Playa de Las Canteras Puerto de San Nicolas Playa del Ingles Playa Ojo de Garza Playa de las Burras Melenara Island La Papagaya Table 1.—Localities and samples. Gran Canaria Lanzarote 188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ‘esi Fig. 1. ruberrima, basal plate and jaws of several specimens; E shows the same specimen as D but in left lateral view. All to the same scale. Family Agnathiellidae Sterrer, 1972 Composition.—One genus, Agnathiella Sterrer, 1971. Paragnathiella, new genus Diagnosis.—Agnathiellidae with jaws but without a basal plate. Type species: P. trifoliceps, new species. Paragnathiella trifoliceps, new species (Figs. 3A—-L, 4A—E, Table 3) Synonymy.—‘“‘Genus IV” and ‘“‘Genus V" in. Stemer 197 2: on | twee we tooul <>) arty: Wb D = 4Ea es Haplognathia rosea and H. ruberrima. A, H. rosea, jaws and rostral edge of basal plate; B—G, H. Etymology.—From the Greek par (be- side), to indicate a close relationship with Agnathiella; and Latin trifoli- (three- leaved) and -ceps (-headed), in reference to the clover-shaped rostrum. Material.—18 specimens, of which 2 adult, from Gran Canaria, Playa de Las Canteras (17 from sample Cl; one from sample C9). Holotype.—USNM 174368, one adult in Squeeze preparation. Type locality.—Gran Canaria, Las Pal- mas, Playa de Las Canteras, fine sand with Sparse sea grass (Cymodocea nodosa) at VOLUME 110, NUMBER 2 189 Fig. 2. Haplognathia rosea and H. ruberrima, phase contrast micrographs. A, H. rosea, jaws and rostral edge of basal plate (cf. Fig. 1A); B, H. ruberrima, jaws; C, H. ruberrima, jaws, and basal plate with thorns (upper right). 2-3 m depth; sample collected 16 Feb 1996. Diagnosis.—About 1500 wm long Par- agnathiella (body index 13.42) with clover- shaped rostrum (rostrum index 2.40). Jaws 14-17 wm long bearing 14-20 delicate teeth. Male stylet to 70 pm long. Description.—Organization and behav- ior: Colourless and very opaque due to many round, greenish epidermal inclusions. Animals move rather swiftly, often carrying the posterior third of the body curled up, and are able to swim backward. Adults are to 1500 pm long and 120 pm wide at U 49.84 (index 13.42); the posterior end ta- pers into a short tail region. The rostrum of the only adult measured was 120 pm long and 50 wm wide at U 8.0 (index 2.40). The rostrum is somewhat clove-shaped, with a 25 wm long frontal lobe separated from lat- eral lobes by conspicuous ciliary pits. It seems that there is one pair of apicalia (to 16 wm long). Sensory cirri are difficult to analyze since they are poorly defined and usually join the ciliary beat. There is one pair each of frontalia (42 wm), ventralia (30 wm), dorsalia (23 wm), lateralia (40 wm), and postlateralia (35 wm). Sensory bristles originate well apart from each other, and do not coincide in dorsal view. Digestive tract: The transversely oval mouth extends from U 7.3 to U 8.0. A basal plate is lacking. The prefrontal epidermis is thickened, and contains many round inclu- Table 2.—Haplognathia ruberrima. Measurements and statistics. x SD Max Min n Body length of adults 2150.00 2150 2150 l Body width of adults 75.00 tf ia l Body index of adults 28.67 28.67 28.67 ] Rostrum index of adults 4.33 4.33 4.33 1 Jaw length 25-57 4 29 fe 7 Apophysis index 0.52 0.03 0.56 0.48 7 Basal plate length 6.33 0.52 Z 6 6 Basal plate width 11.50 2a 13 8 6 Basal plate index 0.57 0.12 0.75 0.46 6 190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figé 3 left lateral view; D-I, jaws of several specimens; F—H, strongly squeezed, I, left lateral view; J, bursa; K, male copulatory organ; L, sperm. Scales apply to A, B—C, D-J, and K-L, respectively. sions. The jaws are 14-17 wm long (X = 15.31 wm), delicate, forceps-shaped and la- mellar. In resting position the jaws point dorsorostrally at a 50° angle from the hor- izontal (Fig. 31). The symphysis is narrow, and there is a pair of rostral apophyses that converge ventro-medio-caudally. Under low squeezing the jaws appear to have only one strongly curved terminal tooth. Stron- ger squeezing reveals that what appears as a terminal tooth is only the dorsal endpoint Paragnathiella trifoliceps. A, habitus of adult; B, rostrum of adult, ventral view; C, rostrum of adult, of a delicate, caudo-ventrally descending lamella set with 14—20 (X = 18.00) teeth, of which all except the 2 or 3 dorsal-most are very short. The pharynx bulb is 12—17 wm (X = 14.40 wm) long behind the sym- physis. The gut cells appear strongly vac- uolized. There is a “‘lateral system”’ of un- known function, 1.e., tissue strands between gut and epidermis which originate on either side of the pharynx and continue into the tail region. VOLUME 110, NUMBER 2 19] Ce Fig. 4. Paragnathiella trifoliceps, phase contrast micrographs. A—B, habitus of free-swimming specimen; C-E, jaws of three specimens, E strongly squeezed to show teeth (cf. Fig. 3H). Male system: The paired, tubular testes are 200 wm long, extending from U 73.3 to U 86.7. They empty into a muscular, ovoid vesicula seminalis 28 zm long and 22 wm wide, which in turn connects with the penis (Fig. 3L). Located between U 87.35 and U 91.48, the male stylet, made up of concen- trically arranged rods, is 69-70 wm long and 5 wm wide. Sperm is irregularly round to oval, 2—3 wm in diameter, with no dis- cernible filaments (Fig. 3J). Female system: The only adult measured had an egg 280 um long which extended from U 40.0 to J 58.7. The bursa system lies immediately posteriorly, extending from U 60.38 to U 64.71. It consists of an anterior bell-shaped bursa, 34—60 wm long (X = 44.00 pm) and 27-28 wm wide (X = 27.75 ym), and an unstructured, globular prebursa about 50 wm in diameter. The bur- sa is made up of stacked, layered cells as is usual for the suborder Scleroperalia, but cristae are lacking, and a bursa mouthpiece, although probably present, is not obvious (Fig. 3K). The bursa is usually tightly packed with sperm. A vagina was not ob- served. Discussion.—This species has been known to me since 1966 when I found one specimen at Banyuls-sur-Mer (southern France). Another specimen encountered in Rovinj (Adriatic) in 1967 suggested at first that I was dealing with two separate species or even genera, to which I referred as “‘Ge- nus IV”? and “Genus V”’ (Sterrer 1972). Further material from Tunis in 1971, Rovinj and Crete in 1991, and now Gran Canaria, have convinced me that we are dealing with a single species. The possession of a cutic- ular bursa and stylet assigns it to Bursova- ginoidea-Scleroperalia, and the lack of a ba- sal plate, lamellar forceps structure of jaws, and loose arrangement of the sensorium place it at the lower end of the suborder. The jaws most resemble those of lower Scleroperalia, i.e., Clausognathia Sterrer, Gnathostomaria Ax, Mesognatharia Ster- rer, Labidognathia Riedl, and Tenuignathia Sterrer. With the two latter genera, Parag- nathiella also shares the possession of an v2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.—Paragnathiella trifoliceps. Measurements and statistics. X Body length of adults 1460.00 Body width of adults 110.00 Body index of adults 13.42 Rostrum index of adults 2.40 Jaw length 15231 Penis stylet length 69.50 Sperm length 3.00 Sperm width 2.38 Sperm index 1.31 unpaired apical ciliary pit; with Clausog- nathia, Tenuignathia, Rastrognathia Kris- tensen & N@rrevang and Agnathiella Ster- rer, it shares the lack of a basal plate. In terms of its habitus, the clover-shape of the rostrum, and the unique possession of an extra pair of sensory cirri (postlateralia), the new genus is nearly identical with the jaw- less Agnathiella. Since the loss of basal plate and/or jaws may have occurred sev- eral times independently in lower Sclero- peralia, I propose that the new genus be united with Agnathiella in the family Ag- nathiellidae. Family Mesognathariidae Sterrer, 1972 Composition.—The 3 genera Mesogna- tharia Sterrer, 1966, Labidognathia Riedl, 1970, and Tenuignathia Sterrer, 1976. Labidognathia longicollis Riedl, 1970 (Figs. sA—E, 6A—B, Table 4) Material.—Nine specimens, of which 3 are adults, from Gran Canaria, Playa de Las Canteras (sample C1). Description.—Colorless. One adult mea- sured 700 wm by 65 pm at U 42.9 (index 10.77). The basal plate is shield-shaped, 14—18 pm long and 10—13 wm wide (index 1.31), with a rostral concavity and an oval knob at the caudal end. In most specimens the rostral concavity clearly showed 8-12 longitudinal striations, whereas each of the lateral wings was set with 9-12 shorter teeth. Jaws are lamellar, 20—23 wm long (X SD Max Min n 56.57 1500 1420 2 14.14 120 100 De 2.24 15.00 11.83 2 2.40 2.40 1 0.85 17 14 he 0.71 70 69 2 0.00 3 3 8 O52 3 2 8 0.26 1.50 1.00 8 = 21.00 wm), and provided with a some- what coarser dorsal row of 13—15 teeth, and a finer ventral row of 10-15 teeth. Both rows are composed in such a way that a robust tooth always alternates with two del- icate teeth (Fig. 5F). The male stylet was 35—48 wm long (X = 43.00 wm). Discussion.—This is the first record of the species outside the northwest Atlantic, where it has been reported from North Car- olina (Riedl 1970), and from Florida, Be- lize, Puerto Rico and Panama (Sterrer in press). The Canaries specimens differ from their western Atlantic counterparts in the longer male stylet (35-48 wm vs. 23-35 yum), but especially in the dentition of the basal plate. Riedl (1970) described ‘a row of short ridges or projections, perhaps 6 to 8 in number,” on the central part of the ros- tral contour, and Sterrer (1997) recorded specimens from Puerto Rico with nine teeth on each of the lateral lobes. Yet nowhere have I seen specimens in which both the central ridges (or striations) and the lateral teeth were so consistently conspicuous. Suborder Conophoralia Sterrer, 1972 Composition —One family, Austrogna- thiidae Sterrer, 1971. Family Austrognathiidae Sterrer, 1971 Composition.—Three genera, Austrog- nathia Sterrer, 1965 emend. Sterrer, 1991a; Austrognatharia Sterrer, 1971 emend. Ster- rer, 1991a; and Triplignathia Sterrer, 1991d. VOLUME 110, NUMBER 2 ys Le RL tty, hin > es Fig. 5. 193 Brae Labidognathia longicollis. A-E, jaws and basal plates of several specimens, strongly squeezed; FE detail of jaw teeth in dorsal row. A-E to the same scale; F not to scale. Austrognathia clavigera, new species (Figs. 7A—O, 8A-—C, Table 5) Etymology.—From the Latin clavus (nail), and gerere (to carry), in reference to the sperm (conuli) which give the appear- ance of nails or thumbtacks. Material.—Twenty-eight more speci- mens, of which 7 adults, from sample Cl. Holotype.—USNM 174367, one adult in squeeze preparation. Type locality.—Gran Canaria Las Pal- A mas, Playa de Las Canteras, fine sand with sparse sea grass (Cymodocea nodosa) at 2— 3m depth; sample collected 16 Feb 1996. Diagnosis.—Stout Austrognathia (body index 7.63) with squarish rostrum (rostrum index 0.81) and many bundles of spindle- shaped epidermal rhabdoids. Basal plate 6.30 wm long, 20.09 wm wide (index 0.32), with inconspicuous median and flat lateral lobes. Jaws 20.24 ym long, with 4.78 teeth in dorsal and 8.14 teeth in ventral row; pos- Fig. 6. Labidognathia longicollis, phase contrast micrographs. A—B, basal plate and jaws of two specimens, strongly squeezed. 194 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Oe G Fig. 7. Austrognathia clavigera. A, habitus of adult; B, rostrum of another specimen; C, rhabdoid bundle; D-G, basal plate and jaws of four specimens, strongly squeezed; H, posterior part of body showing male reproductive system; I-M, mature conuli; N, immature conulus; O, bursa conulus. Conuli in K and L, M and N, and J and O, respectively, are from the same specimen. Scales apply to A-B, C-—G, H, and I—O, respectively. VOLUME 110, NUMBER 2 195 Fig. 8. Austrognathia clavigera, phase contrast micrographs. A, basal plate and jaws, strongly squeezed, so that the anterior edge of the basal plate is facing posteriorly; B, rhabdoid bundles; C, terminal conuli. terior-most dorsal tooth usually rooted. Conuli to 52 4m long and 24 wm wide (in- dex 1.96), without cingulum; hat much wid- er than cone and circumscribing 250°. Description.—Organization and behav- ior: Colorless, and rather opaque from large numbers of epidermal rhabdite bundles. Animals glide slowly, but prefer to attach themselves to detritus particles to which they may cling with tenacity. Adults are 420-900 wm long and 80-90 wm wide at U 47.26 (index 7.63). The rostrum is 45— 50 wm long and 50—70 um wide at U 6.57; it is typically rather square, with nearly par- allel lateral contours. The sensorium—dif- ficult to ascertain since the cirri join the cil- iary beat almost continuously—consists of two pairs of single apicalia (to 20 ym long), and the compound frontalia (48 wm), ven- tralia (SO wm), dorsalia (38 wm), and later- 196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 4.—Labidognathia longicollis. Measurements and statistics. Xx SD Max Min n Body length of adults 700.00 700 700 1 Body width of adults 65.00 65 65 1 Body index of adults OWT 10.77 10.77 1 Rostrum index of adults 1.88 1.88 1.88 1 Jaw length 21.00 1.15 23 20 Fi Basal plate length 15.43 1.40 18 14 7 Basal plate width 11.86 OF 13 10 7 Basal plate index 13 0.11 1.45 Mee / gh Penis stylet length 43.00 7.00 48 35 3 alia (40 wm). Occipitalia and a pair of cil- lary pits are probably present. Throughout the body, but especially in the tail region, the epidermis contains ovoid bundles of spindle-shaped rhabdoids (Fig. 7C) 6 wm long and 2 wm wide which presumably serve an adhesive function. Digestive tract: The basal plate measures 5-8 pm in length and 18—22 wm in width (index 0.32). Its rostro-lateral lobes are fair- ly low, and a median lobe is inconspicuous or lacking. Caudally the basal plate is set with 8—13 (X = 10.18) rather uniform teeth. Jaws are 18-22 wm long (X = 20.24), and have a 7 wm long pear-shaped cauda. In addition to a strong terminal tooth there are two rows of teeth: a longer ventral row of 5-12 (X = 8.14) long teeth, and a shorter dorsal row of 4—7 (X = 4.78) short teeth. In the ventral row, a stronger tooth usually alternates with two weaker teeth. In the dor- sal row, the caudal-most tooth is always rooted. Male system: The single testis is 50-120 ym long, extending dorso-caudally from U 71.73 to U 83.98. The penis, from U 84.62 to 94.58, has three regions characterized by medium, coarse, and fine granula, respec- tively, with the finely-granular part sur- rounding the male pore (Fig. 7H). The testis contains up to 11 conuli which come in at least two size classes. Of a total of 46 con- uli encountered in 7 specimens, 41 were what I call “‘terminal’’ conuli (Fig. 7I—M): to 52 wm long and 24 wm wide (index 1.98), with a towering hat that describes up to 250° of a circle, and whose width sig- nificantly exceeds that of the cone. A cin- gulum is lacking. Only five conuli, seen in three specimens in addition to terminal con- uli, were smaller and more slender (30 wm by 10 um), and endowed with a less prom- inent hat (Fig. 7N). Female system: Located between U 39.75 and U 54.38, a mature egg may be 85-110 wm long. A bursa conulus, 22 wm Table 5.—Austrognathia clavigera. Measurements and statistics. x Body length of adults 645.00 Body width of adults 85.00 Body index of adults 7.63 Rostrum index of adults 0.81 Jaw length 20.24 Basal plate length 6.30 Basal plate width 20.09 Basal plate index 0.32 Sperm length 43.63 Sperm width Dip dee \ Sperm index 1.96 SD Max Min n 155715 900 420 6 5.48 90 80 6 2.06 M:.25 3:25 6 0.08 0.90 0.71 4 1.05 DIBA 18 2s 0.70 8 5 23 1.38 22, 18 23 0.05 0.40 O25 23 3.56 52 38 16 1.01 24 ZA 16 0.14 2.26 1.67 16 VOLUME 110, NUMBER 2 by 17 wm, was seen in one specimen (Fig. 7O). Discussion.—The genus Austrognathia Sterrer, 1965 currently comprises 8 valid species: A. hymanae Kirsteuer, 1970; A. mi- croconulifera Farris, 1977; A. nannulifera Sterrer, 1991la; A. novaezelandiae Sterrer, 1991a; A. riedli Sterrer, 1965; A. christi- anae Farris, 1977; A. singatokae Sterrer, 199la; and A. macroconifera Sterrer, 1991c. A comparison with A. clavigera sets the first four species apart as having conuli with a maximum length of only 9-20 pum. Of the remaining species, A. macroconifera and A. singatokae have conuli quite differ- ent from those of A. clavigera. The conuli of A. christianae and A. riedli are similar to those of A. clavigera except that in the former two species the hat circumscribes no more than 180°, and does not significantly exceed the width of the cone. According to a recently proposed typology of austrog- nathian mouth parts, the new species be- longs in the “‘novaezelandiae”’ group char- acterized by “‘a basal plate with a flattened or absent central lobe and rounded lateral lobes ... and jaws in which the dorsal row has 3 or more teeth of which the caudal- most is usually rooted’’ (Sterrer 1991d). In aspects such as the shape of rostrum and conuli, as well as the possession of many adhesive papillae, A. clavigera most closely resembles the species reported by Riedl (1966) from the Red Sea and provisionally named A. riedli forma maris-rubri of which, however, neither the structure nor the dimensions of jaws and basal plate are known. Acknowledgments I am grateful to Dr. Angel Luque Esca- lona and Lic. Javier Pérez Fernandez of the 197 Universidad de Las Palmas for providing lab facilities and helping with sample col- lection, and Dr. Karl Wittmann (University of Vienna, Austria) for collecting a sample in Lanzarote. Literature Cited Riedl, R. 1966. Faunistische Studien am Roten Meer im Winter 1961/62, III. Die Aufsammlungen in Suez und Al-Ghardaga, nebst einigen Bemer- kungen tiber Gnathostomulida.—Zoologische Jahrbiicher, Systematik 93:139-157. 1970. On Labidognathia longicollis, nov. gen., nov. spec., from the West Atlanic Coas (Gnathostomulida).—Internationale Revue der gesamten Hydrobologie 55:227—244. Sterrer, W. 1969. Beitrage zur Kenntnis der Gnathos- tomulida. I. Anatomie und Morphologie des Ge- nus Pterognathia Sterrer—Arkiv fdr Zoologi 22:1-125. . 1971. On the biology of Gnathostomulida.— Vie et Milieu, Suppl. 22:493-508. . 1972. Systematics and evolution within the Gnathostomulida.—Systematic Zoology 21: 151-173. 1991a. Gnathostomulida from Fiji, Tonga and New Zealand.—Zoologica Scripta 20:107— 128. 1991b. Gnathostomulida from Hawaii.— Zoologica Scripta 20:129-136. . 1991c. Gnathostomulida from Tahiti.—Zool- ogica Scripta 20:137—146. 1991d. Triplignathia adriatica, new genus and species, and a typology of mouth parts in Austrognathiidae (Gnathostomulida). Proceed- ings of the Biological Society of Washington 104:640-—646. 1992. Clausognathiidae, a new family of Gnathostomulida from Belize.—Proceedings of the Biological Society of Washington 105:136— 142. . 1997. Gnathostomulida from the subtropical northwest Atlantic.—Studies on the fauna of Curagao (in press). , M. Mainitz, & R. M. Rieger. 1986. Gnathos- tomulida: enigmatic as ever—Pp. 181-199 in S. Conway Morris, J. D. George, R. Gibson, & H. M. Platt, eds., The origins and relationships of lower invertebrates—The Systematics As- sociation Special Volume No. 28:1—397. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):198—202. 1997. A new species of Stenoninereis (Polychaeta: Nereididae) from the Gulf of Mexico J. A. de Le6n-Gonzalez and V. Solis-Weiss (JAL-G) Laboratorio de Zoologia de Invertebrados, Facultad de Ciencias Biolégicas, U.A.N.L., Ap. Postal 5“‘F’’, San Nicolas de los Garza, N.L. 66451, México; (VS-W) Laboratorio de Poliquetos, Instituto de Ciencias del Mar y Limnologia, U.N.A.M., Ap. Postal 70-305, México, D.F 04510, México Abstract.—The genus Stenoninereis is newly recorded from Mexican waters. The only formerly known species of the genus, Stenoninereis martini Wesen- berg-Lung, 1958, is recorded from Laguna de Terminos, State of Campeche, Mexico, where it was found associated with mangrove roots. Stenoninereis tecolutlensis n. sp. was collected in Tecolutla, State of Veracruz, under oyster Shells attached to mangrove roots in an estuarine zone. Both specimens are described and illustrated, and a Key is provided in order to distinguish between them. The genus Stenoninereis was established in 1958 by Wesenberg-Lund for those spe- cies with dorsal cirri formed by elongate basal cirrophores and subulate distal cirro- styles. Up until now, only the type species (S. martini and synonym WNicon lackeyi Hartman, 1958) was known, from localities in the Great Caribbean region, western Gulf of Mexico, Cuba and North Carolina (Wes- enberg-Lund 1958, Pettibone 1971, Wil- liams et al. 1976, Hartmann-Schroder 1977, Gardiner & Wilson 1979). The material analyzed for this study was collected in Laguna de Terminos, State of Campeche from sediment associated with mangrove roots, and in Estero de Larios, Tecolutla, State of Veracruz under oyster Shells attached to mangrove roots. The Specimens were fixed with formalin, pre- served in 70% alcohol and stained with a methyl blue solution to highlight some im- portant features. The type material is de- posited in the Smithsonian Institution (USNM, Washington, D.C., U.S.A.) as well as in the polychaete collections of the In- stituto de Ciencias del Mar y Limnologia (CPICML-UNAM) and Facultad de Cien- cias Biologicas, Universidad Aut6énoma de Nuevo Leén (UANL), both in Mexico. A diagnosis of S. martini is given, in order to complement the original description and to compare it to the new species herein de- scribed. Stenoninereis Wesenberg-Lund, 1958 Type species.—Stenoninereis martini Wesenberg-Lund, 1958. Diagnosis.—Prostomium small, rounded and distally notched, with paired frontal an- tennae, biarticulate palps and two pairs of eyes, anterior pair crescent-shaped, and posterior pair rounded. Four pairs of tentac- ular cirri. Pharynx with paired jaws without paragnaths or papillae. First two pairs of parapodia subbiramous. Dorsal cirri with long basal cirrophores and short cirrostyles. Biramous parapodia with notopodia bi- lobed, with lower acicular lobes and upper ligules reduced in posterior parapodia; neu- ropodia with bluntly conical acicular lobes in anterior region, becoming more elongate in middle setigers, and shorter, more point- VOLUME 110, NUMBER 2 ed in posterior region. Ventral cirri cirri- form. Notosetae homogomph and sesqui- gomph spinigers. Neurosetae heterogomph spinigers, sesquigomph and heterogomph falcigers with thin blades. Pygidium with a pair of expanded lobes and a pair of anal cirri. Stenoninereis martini Wesenberg-Lund, 1958 Fig. 1 Stenoninereis martini Wesenberg-Lund, 1958: 9, figs. 2—4; Pettibone, 1971: 39, figs. 23, 24; Williams, et al., 1976: 83; Hartmann-Schréder, 1977; Gardiner & Wilson, 1979: 165 fig. 2a—h. Nicon lackeyi Hartman, 1958: 263, figs. 1—5. Material examined.—México, San Juli- an, Laguna de Terminos, Campeche, 1/Mar/ 1984 (18 specimens) (CPICML-UNAM POH-39-39). Description.—Best preserved specimen complete (33 setigers), olive-green, with small pigmented dots, on the most dorsal part of the body, 6 mm long and 1 mm wide including parapodia. Prostomium pentago- nal, slightly notched frontally. Two pairs of eyes: anterior pair crescent shaped, poste- rior pair small and rounded. Frontal anten- nae cirriform and not longer than distal margin of palps. Palps globular, biarticulate with elongate conical palpostyles. Peristo- mium slender, with four pairs of tentacular cirri, anterior dorsal pair reaching up to se- tiger 6. Pharynx with one pair of jaws armed with 9 teeth. First two parapodia subbiramous; noto- podia reduced to small notoacicula. Follow- ing parapodia biramous; anterior ones (Fig. 1A) with long dorsal cirri consisting of elongate basal cirrophore and short piriform cirrostyle, which becomes longer towards the body end (Fig. 1B, C). Notopodia tri- lobate, superior lobe short and digitiform, inferior lobe subulate, with small presetal lobe at base of upper lobe. Superior noto- podial lobes reducing in size in posterior setigers (Fig. 1C). Neuropodia with bluntly 199 conical acicular lobe in anterior region (Fig. 1A), becoming more elongate in middle se- tigers (Fig. 1B) and shorter, more pointed in posterior region (Fig. 1C). Supracicular notosetae are sesquigomph spinigers, infra-acicular notosetae are hom- ogomph spinigers; both with slender appen- dix, serrated on the inner edge. Supracicular neurosetae are heterogomph and _ sesqui- gomph spinigers; infracicular ones are ses- quigomph spinigers: one heterogomph spi- niger with a short blade strongly serrated on at least three quarters of its length on its inner margin (Fig. 1D); heterogomph fal- cigers with long blade, spinulose, distally hooked. Blades of the most ventral hetero- gomph falcigers are one fourth of the length of the superior ones. Pygidium subterminal, with pair of lat- eral flattened, wide lobes, and pair of long anal cirri. Anal opening ventral. Distribution.—Stenoninereis martini has been reported from the Greater Caribbean region (type locality: San Martin Island; and Sarasota, Florida), western Gulf of Mexico (Texas), Cuba, North Carolina, and herein from Laguna de Terminos, Campe- che, Mexico, where it was collected in soft bottoms associated with mangrove roots. Stenoninereis tecolutlensis, new species Fig. 2 Material examined.—Mexico, Estero de Larios, Tecolutla, Veracruz, 19/Nov/1994 (Holotype: USNM-174870, and 2 para- types: UANL-3980 and CPICML-UNAM POH-39-002). Description.—Holotype complete, green- yellowish, 15 mm long and 2 mm wide with 49 setigers. Palps and eyes contour- pigmented; peristomium and following three setigers with transverse dark stripe, less conspicuous on following segments. Prostomium rounded, deeply incised frontally, with a pair of small, cirriform frontal antennae, longer than palps. Two pairs of eyes: anterior pair crescent-shaped, posterior pair rounded. Biarticulate palps — 200 Py ~ rd mS. ‘ : = as . m . > Br > 4 ie ee 2 Nt fe on fr, ea s-) ’ se - AS ta 33) : d ee te boy ie, 7 =8 * oq Se os, te, as ? Rae Q rs Boe hs Se Gg ‘ eb iSe tes, 4“ — "te, eS . oo oo Ay “YY y N 7D +) ae oY * a ve ». 5) . » vi - . “, rQ $° . A 1 2 aed Se ao . Vos FA ry A ae * : we on 3 7 ve ca ~ < eS Re ea r . Yb) - uty ae Bs a creel c* fet XN. 7 oi a Of! oh &] = “ “4 ny AW. Pa G eS? sh! 7 0 Se 2 ie = a oe ald Fig. 1. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON "SAS oe SO ree — * o> I “a. oe Tes av : a =f, aes ~S . as : we e @ BS. ges FES yr we Bea Nee - st -£ oe” ve =) 28 Sg ie a “ye ars She vee OH ngs as wes e “how Ss Cer) Bd 4 < lone ~ bs . . . se oy wn aD +. aa Ff, ban EY s Coma (5 i So -}. ok Set TPS S oe ee Sad : cr Stenoninereis martini. A. 6th parapodium; B. 18th parapodium; C. 28th parapodium; D. heterogomph spiniger from parapodium of middle segments. (A, B, C anterior view, setae omitted). Scale: A-C = 60 pm; D = 8 pm. with globular palpophores and conical pal- postyles. Peristomium narrow, with four pairs of tentacular cirri, posterodorsal pair longest reaching setiger 12; anterodorsal pair laterally expanded (Fig. 2A). Pharynx with a pair of jaws armed with five minute teeth. First two parapodia uniramous, the fol- lowing biramous. In anterior and middle parapodia (Fig. 2 B—C) two long and slen- der lobes form notopodium, superior one being shorter. Posterior parapodia bearing only an acicular lobe (Fig. 2D). Cirrophores of the dorsal cirri in anterior and middle VOLUME 110, NUMBER 2 | 201 Fig. 2. Stenoninereis tecolutlensis new species A. Anterior end, dorsal view; B. 10th parapodium; C. 25th parapodium; D. Posterior parapodium; E. Neuropodial infracicular heterogomph spiniger from parapodium of middle segments; E Neuropodial infracicular heterogomph falciger from parapodia of middle segments. (B, C, D: anterior view, setae omitted). Scale: A = 0.5 mm; B — D = 60 pm; E, F = 8 pm. 202 segments 1.5 times longer than the cirros- tyles; on posterior parapodia cirrostyle 1.5 times longer than the cirrophore, but the cirrophore is basally stouter. Ventral cirri longer on posterior parapodia. Setation is similar throughout the body. Notosetae are homogomph spinigers in su- pracicular position and two sesquigomph spinigers in subacicular position. Supraci- cular neurosetae are heterogomph and ses- quigomph falcigers with smoothly serrated blades. Infracicular ones in dorsal position with one pair of heterogomph spinigers strongly denticulate (Fig. 2E), and hetero- gomph falcigers with elongate terminal hooked blades (Fig. 2F); inferior ones one third of superior ones’ length. Anus terminal, with two flat winglike ex- pansions, and a pair of anal cirri. Geographical distribution.—Known only from Estero de Larios, Tecolutla, Ve- racruz, Mexico, where it was collected un- der oyster shells attached to mangrove roots in very shallow estuarine waters. Etymology.—Stenoninereis tecolutlensis is named after the type locality, Tecolutla, Veracruz. Remarks.—The genus Stenoninereis was erected by Wesenberg-Lund (1958) for only one species: S. martini. S. tecolutlensis dif- fers from the type species in the shape of the prostomium and the size of the eyes: in S. martini, the prostomium is rounded with small eyes while in S. tecolutlensis the pro- stomium is deeply incised frontally, with large eyes. They further differ in the shape and distribution of the supracicular double notopodial lobe (short and present to the end of the body in S. martini, long and missing on posterior parapodia in S. teco- lutlensis); the relative length of the cirrop- hore and the cirrostyle, the number and size of teeth on the jaws, (nine in S. martini and five minute teeth in S. tecolutlensis), the difference in size and area covered by the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON denticulation of the heterogomph spinigers, S. martini have homogomph spinigers in in- fracicular position, while S. tecolutlensis have sesquigomph spinigers; and the rela- tive size of the ventral cirri on posterior par- apodia (short in S. martini and longer in S. tecolutlensis). Key to Stenoninereis 1. Prostomium rounded, supracicular dou- ble notopodial lobes short, present throughout body, notopodium with hom- ogomph spinigers in infracicular position Paes he's) ee ray a a. S. martini — Prostomium deeply incised frontally, su- pracicular double notopodial lobes miss- ing on posterior parapodia, notopodium with sesquigomph spinigers in infraci- cular position . i... 6c sea S. tecolutlensis Literature Cited Gardiner, S. L., & W. H. Wilson, 1979 (1977). New records of polychaete annelids from North Car- olina with the description of a new species of Sphaerosyllis (Syllidae).—The Journal of Eli- sha Mitchell Scientific Society 93(4):159-172. Hartman, O. 1958. A new nereid worm from Warm Mineral Springs, Florida with a review of the genus Nicon Kinberg.—Journal of Washington Academic Science 48:263—266. Hartmann-Schréder, G. 1977. Die Polychaeten del Kubanisch-Rumanischen Biospeologischen Ex- pedition nach Kuba 1973.—Résultats des ex- péditions biospéologiques cubano-roumaines, Cuba 2:51-63. Pettibone, M. H. 1971. Revision of some species re- fered to Leptonereis, Nicon and Laeonereis (Polychaeta: Nereididae).—Smithsonian Contri- butions to Zoology 104:1—-53. Wesenberg-Lund, E. 1958. Lesser antillean poly- chaetes chiefly from brackish water with a sur- vey and a bibliography of fresh and brackish- water polychaetes.—Studies on the Fauna of Curacao and other Caribbean Islands 8:1—41. Williams, G. E., M. J. Poff, & J. T. MzBee. 1976. Western Gulf of Mexico records of Stenoniner- eis martini Wesenberg-Lund 1958 (Polychaeta, Nereidae) with contributions to its habitat ecol- ogy.—Contributions in Marine Sciences 20:83— 85. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):203—209. 1997. Boguea panwaensis, a new species from Thailand: the first member of the Bogueinae (Polychaeta: Maldanidae) to be found outside northeast America C. Meyer and W. Westheide Spezielle Zoologie, Fachbereich Biologie/Chemie, Universitat Osnabriick, D-49069 Osnabriick, Germany Abstract.—A new boguein polychaete species, Boguea panwaensis, is de- scribed from Cape Panwa, Phuket, Thailand. It is closely related to the North American B. enigmatica, but differs in number, arrangement and structural details of its chaetae. The first species of the maldanid subfam- ily Bogueinae was described from Bogue Sound, North Carolina, by Hartman (1945), who first considered it as belonging to the Oweniidae. Later on (Wolf 1984) it was re- corded from different localities around the coasts of North America at depths down to 100 m, usually between 2 and 20 m, in sand and sand-gravel. With the description of the closely related species Boguella ornata Hartman & Fauchald, 1971, the two authors erected the family Bogueidae and men- tioned that it differs from oweniids in pos- sessing avicular or terebelloid uncini (Hart- man & Fauchald 1971). Wolf (1983) rede- scribed the taxon and reduced its rank to Bogueinae, a subfamily of the Maldanidae, emphasising that they share morphological and ontogenetic features especially charac- teristic of this family. Independently of Wolf (1983), Nilsen and Holthe (1985) ar- gued in the same direction by stating that the boguein uncini could well be derived from those of the Rhodininae, a subfamily of the Maldanidae. Holthe (1986) con- firmed this in his phylogenetic discussion of the Bogueidae. The new Boguea species presented is the first one occurring outside the North American area. Boguea panwaensis, new species Figs. 1, 2, 3 A-EK 4 A-E Material examined.—Two complete specimens and one anterior end with 5 chaetigers. Location: Thailand, Phuket, Cape Panwa, near the Phuket Marine Bio- logical Center (PMBC), at a depth of about 10—15 m in the centre of the bay west of the aquarium (7°52’N, 98°22’E); mature Specimen and anterior end found in October 1994, immature specimen in March 1995. Probably living in a layer (1—3 cm) of very fine, mostly oozy organic material on top of fine sand with organic material, few shells and little gravel, partly with clay texture. Methods.—Extraction of the polychaetes was carried out with a solution of 8% MgCl, isotonic to sea water. Specimens were observerd in living condition before they were fixed in Bouin’s fluid, and then transferred to 70% ethanol. For SEM in- vestigations the dehydrated specimens were critical-point dried with CO, as interme- dium, mounted with a carbon film on alu- minium stubs, sputter-coated with gold and examined in a Zeiss 962 SEM. Type material.—Holotype with 24 chae- tigers (length 10 mm), mature; deposited in the Phuket Marine Biological Center, Ref- erence Collection, Thailand (No. PMBC 13577). Paratype with 23 chaetigers (length 4 mm), immature; SEM preparation, depos- ited in the collection of the Senckenberg Museum, Frankfurt (No. SMF 5633). The new species was named after Cape Panwa in the southern region of the island 204 kom Fig.l, uncinus). Scale bar = 1 mm. of Phuket, where it was found by the au- thors. Material for comparison.—Boguea en- igmatica Hartman, 1945 (extracted from: ZMUC-POL-541; Bogue Sound, North Carolina 6 Aug. 1962). Description.—Length about 10 mm, up to 24 chaetigerous segments, width about 0.5 mm. Colour light brownish-translucent. Body divided into three regions differenti- ated by the shape of the segments and the distinctness of their boundaries, type and number of notochaetae, number of uncini and their arrangement in one or two rows. Prostomium, peristomium and pygidium without appendages. Anterior region.—Through chaetiger 7. Prostomium and peristomium fused. Pro- stomium anteriorly rounded. No nuchal or- gans visible. Borderline between peristo- mium and first chaetiger indistinct. Chaetig- ers 2 to 7 contractible, so that their anterior PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Boguea panwaensis, new species. Holotype with oocytes, lateral view (on chaetiger 2 a single juvenile part may be wider than their posterior part (Fig. 1) and the preceding chaetiger is part- ly recessed into the following one. When relaxed, chaetigers longer than wide. Para- podia absent, except groups of notopodial and neuropodial chaetae, emerging in the middle of each chaetiger, and arranged in one or two rows. Types of notopodial chae- tae in this region include simple, smooth capillaries, serrated capillaries and short, stiff spines. In chaetiger 3 four serrated cap- illary chaetae with minute imbrications and one smaller additional capillary. In the fol- lowing chaetigers only some of the capil- laries show this special structure (Table 1). In chaetiger 3 one additional row of four short spines anterior to these capillaries. Neuropodial chaetae may be present from chaetiger 2 backwards. One small ros- trate uncinus in chaetigers 2 to 4 (Fig. 3D, E) in the juvenile specimen (Fig. 2). In the mature specimen (Fig. 1) the latter are miss- VOLUME 110, NUMBER 2 Table 1.—Distribution of types of notopodial chae- tae in the holotype of Boguea panwaensis, new spe- cies. Chaetiger Types of chaetae on the notopodia 1-2 smooth capillary chaetae 5) stiff, short spines; serrated capillary chae- tae 4-7 serrated and smooth capillary chaetae 8-13 imbricated capillary chaetae 14-17 serrated capillary chaetae; capillary chae- tae with triangular imbrications; slightly curved, flattened serrated chaetae 18-22 short spines; smooth capillary chaetae 23-24 short spines ing, with the exception of one in chaetiger 2. The neuropodial chaetation commences in chaetiger 5, consisting of a single row of avicular uncini in chaetigers 5 to 7, their number varying between 7 and 12. Middle region.—Borderlines between chaetigers 8 to 13 indistinct. Chaetigers lon- aye 205 ger than wide. Neuropodial chaetae resem- ble those in the anterior and posterior regions, and notopodial chaetae are capil- leries with minute imbrications. These chaetae are shorter than in the anterior re- gion and only up to four in number. From chaetiger 8 onwards two rows of neuropo- dial uncini on both sides. Anterior row comprising 3 to 6 on chaetiger 8, increasing in number up to 12 in the following chae- tigers. Posterior row with 7 to 15 uncini in the mature specimen. Each uncinus with a rostrum and two rows of distinct teeth (=capitium in the terminology of Holthe 1986), several additional small teeth later- ally and between the larger teeth, visible only by SEM. Additionally some hair-like projections emerging from the lateral side of the uncini cover the tip of the rostrum (Figs. 3E 4D). Posterior region.—From chaetiger 14 backwards. Number of notopodial chaetae Fig. 2. Boguea panwaensis, new species. Paratype. Immature, lateral view. Scale bar = 0.5 mm. 206 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Big. 3. A-F Boguea panwaensis, new species. A, Serrated capillary chaeta from chaetiger 15 with slender, curved imbrication. Scale bar = 5 pm. B, Capillary chaeta from chaetiger 16 with triangular imbrications in the middle region. Scale bar = 5 wm. C, Slightly curved, flattened serrate chaetae from chaetiger 14. Scale bar = 10 pm. D, Juvenile rostrate uncinus from chaetiger 4 (D and E from paratype). E, Juvenile rostrate uncinus from chaetiger 2. Scale bar in D and E = 10 pm. EK Uncinus from chaetiger 15. Scale bar = 2 pm. G, Boguea enigmatica Hartman, 1945. Paratype, uncinus from chaetiger 5. Scale bar = 2 wm. first increasing up to 10, then decreasing in the posteriormost chaetigers, last two seg- ments achaetous. Some of the imbricated capillary chaetae on chaetiger 14 to 17 pos- sessing a specific structure not present in those of the anterior and middle region: in light microscopy these chaetae appear featherlike in lateral view with a series of fine pinnulated imbrications. From SEM preparations, pinnules from one type of chaetae revealed to be cylindrical in the proximal part of the chaetae; distally they are arranged in pairs, gradually forming tri- angular structures (Figs. 3B, 4E). The im- brications project dorsally from the ventral sides of the chaetal axes. The two rows of imbrications are covered on the ventral side with a series of additional forward directed imbrications, which build a characteristic keel (Fig. 4B). On chaetiger 14 to 17 an- other type of capillary chaetae with single rows of slender, slightly curved pinnules on each side exist (Figs. 3A, 4C). These pin- nules are bent forward on the ventral side of the chaetae and directed dorsally. They differ from the chaetae with the triangular imbrications in that the appendages of these chaetae are longer and remain separated up to the distal end of the chaetae. There are also some curved, flattened serrated chaetae on these chaetigers (Fig. 3C; see also Wolf 1983, fig. 11). In the following segments, from chaetiger 18 on, they are replaced by smooth and shorter capillary ones. From chaetiger 18 backwards relatively small spines occur, gradually decreasing in num- ber; they are present even on one or two of the posteriormost segments, which bear no other chaetae. These spines resemble those of chaetiger 3, but are smaller. Number of uncini reduced to approxi- mately 8 in each row in chaetiger 14, de- creasing gradually in number to chaetiger 23, with the 2 rows merging towards the posterior end of the body. Uncini in the last VOLUME 110, NUMBER 2 207 ra Fig. 4. Boguea panwaensis, new species. A, Spines and serrated capillary chaeta from chaetiger 3, arrow points to a broken-off additional capillary chaeta. Scale bar = 20 pm. B, Serrated capillary chaeta with view on the ventral edge formed of imbrications on ventral side. Scale bar = 5 ym. C, Capillary chaeta from chaetiger 15 with slender, curved imbrications. Scale bar = 5 ym. D, Uncinus from chaetiger 15 with several additional teeth on the lateral side. Scale bar = 2 pm. E, Chaeta from chaetiger 16 with triangular imbrications in the distal region. Scale bar = 5 um. segments of the juvenile specimen small and rostrate. Pygidium without appendages, anus ter- minal. In mature animals eggs of light brownish colour can be seen from the mid- dle region to the posterior chaetiger (Fig. Ly. Discussion.—The new boguein species was discovered at Phuket, Thailand. The two other described species (Boguea enig- matica Hartman, 1945; Boguella ornata Hartman & Fauchald, 1971) were recorded only from Northeast America. To our knowledge this is the first species found outside this region, with the exception of another undescribed species at the coast of Mauritania, probably belonging to Boguella (Dr. H. Michaelis, pers. comm.). The new species from Thailand belongs to the genus Boguea. It shares the following characters with B. enigmatica: The body is divided into three regions. Distribution and types of chaetae are almost identical. A dis- tinct borderline between peristomium and first chaetiger is not distinguishable, al- though Wolf (1983) mentioned slight lateral 208 indentations. The chaetae on the anterior chaetigers show the same minute distally rounded imbrications. The first segment that bears a single row of uncini is the fifth chaetiger. Both species of Boguea possess posterior notopodial spines and small ros- trate uncini. The most characteristic difference of Bo- guea panwaensis is the double row of un- cini starting on chaetiger 8, whereas B. en- igmatica has double rows from chaetiger 9 backwards. In addition, on the third chae- tiger of the new species four capillary chae- tae and four spines exist, which were not described for B. enigmatica. Four to six acicular spines are also mentioned by Wolf (1984) in his description of Boguea sp. A from the Gulf of Mexico, however, his drawings differ from the structures ob- served by light microscopy in the speci- mens from Thailand. The length of the acic- ular spines in B. panwaensis is approxi- mately the same, whereas Boguea sp. A (Wolf 1984) show spines of varying length. These characteristic spines occur in the ho- lotype and in the incomplete specimen of the present material, but are missing in the immature specimen. The specific structures of the capillary chaetae in the species from Thailand at first—when compared with the drawings of the corresponding chaetae published by Wolf (1983)—seemed clearly to differenti- ate it from B. enigmatica. However, exam- ination of a paratype of the latter species by SEM made clear that these differences de- rive largely from the lower resolution achievable with light microscopy; the struc- ture of the chaetae is fundamentally the same in the two species. Certain differences in detail nevertheless did appear in this SEM comparison: serrated capilleries with paired and triangular imbrications (Figs. 3B, 4E) could not be found in B. enigma- tica. The uncini show a similar arrangement of teeth in both species, with additional small teeth between the larger teeth on the lateral side. In the area under the rostrum, several smaller teeth are also present, but PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON they are much more numerous in B. pan- waensis (Figs. 3E 4D) than in B. enigma- tica (Fig. 3G). The hair-like projections are only found on B. panwaensis. The small rostrate uncini are quite similar in both spe- cies. Acknowledgments We are especially grateful to Mr. Somsak Chullasorn and Mr. Anuwat Nateewathana, Phuket Marine Biological Center for their hospitality and allowing us to use their fa- cilities. We would like to thank Dr. Mary Petersen and Dr. Danny Eibye-Jacobsen from the Zoologisk Museum, University of Copenhagen, for making available speci- mens of Boguea enigmatica (extracted from: ZMUC-POL-541; Bogue Sound, North Carolina 6 Aug. 1962). We are grate- ful to Monika C. Miller for her help in pre- paring the SEM pictures, Dr. Deborah Dex- ter, San Diego, for her kind guidance of one of us (C. Meyer) to the sampling sites at the beaches of Phuket, two Thai students and Claire Bradshaw and Jason Shephard from Edinburgh for providing diving-sam- ples from the Cape Panwa Bay. We are deeply obliged to Prof. Dr. Nathan W. Riser, Nahant, for going through the English text. Literature Cited Fauchald, K. 1977. The polychaete worms. Defini- tions and keys to the orders, families and gen- era.—Natural History Museum Los Angeles County Science Series 28:1—190. Hartman, O. 1945. The marine annelids of North Car- olina.—Bulletin of Duke University Marine Station 2:1—54. , & K. Fauchald. 1971. Deep water benthic polychaetous annelids off New England to Ber- muda and other North Atlantic areas, Part 2.— Allan Hancock Monographs in Marine Biology 6:1—327. Holthe, T. 1986. Evolution, systematics, and distri- bution of the Polychaeta Terebellomorpha, with a catalogue of the taxa and a bibliography.— Gunneria 55:1—236. Nilsen, R., & T. Holthe. 1985. Arctic and Scandina- vian Oweniidae (Polychaeta) with a description of Myriochele fragilis sp.n., and comments on the phylogeny of the family.—Sarsia 70:17-—32. VOLUME 110, NUMBER 2 Wolf, P. S. 1983. A revision of the Bogueidae Hart- man and Fauchald, 1971, and its reduction to Bogueinae, a subfamily of Maldanidae (Poly- chaeta).—Proceedings of the Biological Society of Washington 96:238—249. 1984. Family Bogueidae (Hartman & Fau- chald, 1971). Chapter 16, Pp. 16-1 to 16-9 in J. 209 M. Uebelacker & P. G. Johnson, eds., Taxonom- ic guide to the polychaetes of the Northern Gulf of Mexico, Vol 2. Final reports to the Minerals Management Service, Contract 14-12-001- 29091. Barry A. Vittor & Associates, Inc. Mo- bile, Alabama. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):210—226. 1997. Triathrix montagni and T. kalki, a new genus and two new species of Cletodidae (Crustacea: Copepoda: Harpacticoida) from California and the Gulf of Mexico J. Michael Gee and Robert Burgess (JMG) Plymouth Marine Laboratory, Prospect Place, West Hoe, Plymouth PL1 3DH, United Kingdom (RB) University of Texas Marine Sciences Institute, 750 Channelview Drive, Port Aransas, Texas 78373, U.S.A. Abstract.—Triathrix new genus (Copepoda, Harpacticoida, Cletodidae) is established to accommodate T. montagni new species from the California con- tinental shelf and T. kalki new species from the Gulf of Mexico. The genus is characterised by a triangular, sharply pointed rostrum; a chitinous extension bearing four socles on the posterior border of the cephalothorax; no socles on the preanal somite; a median tube pore on the operculum; seta I of the caudal ramus implanted posterior to seta II; antennal exopod with three setae; man- dibular palp with five setae; maxillulary coxal endite with one seta and basal complex with eight setae; maxillary proximal syncoxal endite with two ele- ments, allobasal endite with a spine and two setae, endopodal setae not fused at base; maxillipedal syncoxa without seta; no sexual dimorphism on male P3; female P5 with baseoendopodal lobe and exopod long, rectangular and equal in length. The two species can be distinguished easily from each other by the size of the dorsal extension to the cephalothorax, the shape of the dorsal median socles on the free prosomites, the length of setae on the Pl and male P5 baseoendopodal lobe and the length to width ratio of the caudal rami and the female P5. Enhydrosoma nicobarica Sewell, 1940 is also included in the genus as T. nicobarica new combination. Sewell (1940) described Enhydrosoma nicobarica Sewell, 1940 from a single im- mature specimen found in weed washings in Nankauri Harbour in the Nicobar Islands off the north coast of Indonesia. Within the genus Enhydrosoma Boeck this species is unique in that the exopod of the antenna has three setae: a large, plumose seta on the lat- eral margin along with a large, plumose and a small, naked seta on the distal margin. In his consideration of the genus, Gee (1994) dismissed E. nicobarica, in his preliminary assessment of the genus, on the grounds that it was a juvenile (copepodid V), al- though he considered that an antennal ex- opod bearing three setae was probably the plesiomorphic condition in Cletodidae based on this condition being found in Lim- nocletodes, Borutsky and, reportedly, in Acrenhydrosoma perplexa (T. Scott, 1899). More recently, Fiers (1996) has shown that a trisetose antennal exopod also occurs in the copepodid I stage of E. lacunae Jaku- bisiak, 1933 (and other Enhydrosoma spe- cies) but the small distal seta is lost in co- pepodid II and subsequent developmental stages. He concluded, from this, that the an- tennal exopod structure of E. nicobarica is not a juvenile feature and that the species should be removed from Enhydrosoma and placed as species inquirenda within the family. ; During recent monitoring studies of the effects of oil and gas platforms on the sur- VOLUME 110, NUMBER 2 rounding benthic fauna on the Californian continental shelf (Hyland et al. 1990, Mon- tagna 1991) and the northern part of the Gulf of Mexico (Montagna & Harper 1996), two new species of Cletodidae have been discovered which, in the adult, have an antennal structure exactly akin to that described by Sewell (1940) for E. nicobar- ica. In this paper, we describe these species and demonstrate that they, and E. nicobar- ica, should be placed in a new genus within the Cletodidae. Methods Before dissection the habit is was drawn, and body length measurements made, from whole specimens temporarily mounted in lactophenol. Specimens were dissected and the parts mounted in lactophenol under cov- erslips sealed with Bioseal. All drawings were prepared using a camera lucida on a Nikon Optiphot 20 differential interference contrast microscope. The terminology for body and appendage morphology follows that of Huys et al. (1996). Abbreviations used in the text and figures are P1—P6 for pereiopods 1—6 exopod (endopod)-1(-2-3) to denote the proximal (middle, distal) seg- ments of a ramus. Body length was mea- sured from the base of the rostrum to the median posterior border of the anal somite (i.e., excluding the caudal rami). Family Cletodidae T. Scott, 1904 Our concept of the Cletodidae is based on that defined by Por (1986) after he re- moved many of the genera included in this family by Lang (1948). Triathrix, new genus Enhydrosoma Boeck, 1872, p. 53 (part.) Diagnosis.—Cletodidae. Body semi-cy- lindrical with well defined somites, tapering posteriorly without clear distinction be- tween prosome and urosome. Cephalotho- rax with distinct chitinous extension on dor- sal posterior margin carrying 4 sensillum- 211 bearing socles. Free-prosomites and uro- somite-1 with 6 sensillum-bearing socles and 4 sensilla without socles, dorso-lateral socles usually very pronounced; uroso- mites-2 to -4 with 4 sensillum-bearing so- cles and 4 sensilla without socles. Female genital double-somite with a continuous cu- ticular rib marking line of fusion; genital field with vestigial P6s, bearing 1 seta, cov- ering Separate gonopores; minute copula- tory pore adjacent to cuticular rib. Preanal somite without any socles. Urosomites, ex- cept anal somite, with a double row of spi- nules on ventral posterior margin and a pair of lateral tube pores; anal somite with ad- ditional pair of ventro-lateral tube pores, and single tube pore on operculum. Caudal rami much longer than wide, tapering pos- teriorly; minute seta I implanted posterior to seta II; seta III implanted more or less medially on outer margin and seta VII prox- imal to seta III on inner dorso-lateral mar- gin; ventral and lateral tube pore present in median portion of ramus. Rostrum well-de- veloped, distinctly triangular in shape with sharply pointed recurved tip. Female anten- nule 5-segmented; setal formula 1[1], 2[8], 3[8+aesthetasc], 4[1], 5[11+aesthetasc]. Male antennule 6-segmented. subchirocer; segment-4 swollen, with row of lanceolate setules but without modified setae; setal torme@la “I[i]....219+-tobe _porel,, 315], 4[13+aesthetasc], 5[2+tube pore], 6[9+aesthetasc]. Antennal allobasis with 2 pinnate setae on abexopodal margin; exo- pod a well-developed segment with 3 setae (1 pinnate seta on lateral margin and 1 pin- nate and a small naked seta on distal mar- gin). Labrum without central tuft of setules. Mandibular palp with 5 setae. Maxillulary coxal endite with 1 seta; basis with 8 setae. Maxilla with 2 syncoxal endites, proximal endite with 2 elements, distal endite with 3 elements; allobasal endite with fused spine and 2 setae; endopod represented by 2 setae not fused at base. Maxillipedal syncoxa without seta; endopodal claw pinnate with a long accessory seta and a short finger-like projection at base. Male P3 without sexual 2 Wa dimorphism. Setal formula of P1-P4 as fol- lows: Exopod Endopod Pl 0:0:022 0:0/111 P2 0:0:022 0:020 P3 0:0:122 0:021 P4 0:0:122 0:021 P5 exopod articulating with baseoendopod; in female endopodal lobe and exopod long, rectangular, equal in length and each with 3 setae; in male rami with 2 setae but en- dopodal lobe reduced. Male P6 a single asymetrical plate without setae. Sexual di- morphism in antennule, P5, P6, urosome. Females with 1 egg-sac, males with 1 sper- matophore. Type species.—Triathrix montagni new species, by designation. Other species.—Triathrix kalki new spe- cies and 7. nicobarica (Sewell 1940) new combination. Etymology.—The generic name is de- rived from the Greek noun treis (plural tria) meaning three and thrix meaning hair and refers to the condition of the exopod of the antenna. Gender.—Feminine. Triathrix montagni, new species. Figs. 1-6C Material examined.—Holotype, an adult female (dissected) USNM 278221; para- types, 8 females (2 dissected) and 10 males (3 dissected) USNM 278822. All material collected by Dr. P. Montagna at CAMP site R4, 15 km off the central California coast, 34°43’N, 121°13’'W, from a coarse silt sed- iment at approx. 90 m depth. Female.—Body (Fig. 1). Length 0.59-— 0.67 mm (X 0.62 mm, n = 6), almost cy- lindrical but flattened ventrally in urosome; PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON body surface glabrous. Cephalothorax ta- pering anteriorly, relatively deep dorso-ven- trally; ornamented with sensilla and tube pores as in Fig. 1; anterior ventral borders with lateral extensions (Fig. 1A) slightly re- curved dorsally (Fig. 1B); dorsal posterior border with large chitinous extension car- rying 4 sensillum-bearing socles; lateral and ventral border with 14 sensilla. Poste- rior border of free prosomites and urosom- ite-1 with 6 sensillum-bearing socles and 4 sensilla without socles; dorso-lateral socles very pronounced, increasing in size poste- riorly, each with an associated tube-pore. Genital double-somite with continuous me- dian cuticular rib marking line of fusion, with 4 lateral sensillum-bearing socles and 2 dorsal sensilla; posterior border with, dor- sally 2 sensilla, laterally 6 sensillum-bear- ing socles (ventral pair with an associated tube-pore) and, ventrally a double row of spinules and 2 sensilla (Fig. 2E). Urosom- ite-4 posterior border as for genital-double somite except that only 4 lateral sensillum- bearing socles. Pre-anal and anal somite without socles or sensilla except for a pair associated with smooth operculum (Fig. 1A); both somites with a pair of ventro-lat- eral tube pores; anal somite with additional pair of ventral tube-pores (Fig. 2C) and 1 tube pore on operculum (Fig. 1). Rostrum (Figs. 1, 3E) well-developed, fused to cephalothorax, triangular, tapering to sharply pointed, strongly recurved tip; with 2 lateral sensilla but, apparently, no median ventral tube pore. Caudal rami (Figs. 1, 2C) divergent, slightly swollen at base and tapering pos- teriorly; 4.1—4.4 times longer than maxi- mum width; with tube pore mid-ventrally and laterally immediately posterior to seta III (Fig. 2C). Seta I inserted on ventro-lat- eral margin, posterior to seta II; seta III in- serted laterally at 40% of ramus length; seta IV very small, fused at base to large seta V; seta VI small; triarticulate seta VII in- serted at 28% of ramus length. Genital field (Fig. 2D). Vestigial P6s ex- tremely reduced with 1 seta. Gonopores 213 VOLUME 110, NUMBER 2 | ies. Female body A, dorsal view; B, lateral view. Ww spec Fig. 1. Triathrix montagni, ne 214 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 58) FSF. Ss ae Powys: moran) — Ji — KSA ZS , | : I AN IH As IIE AD i ah ah Hl ma A Cc LN oA | in hs IN IN A\ f x j I Nh me nN ‘| y aan) i ae TUE GTS ae, LN AK ps hi Fig. 2. Triathrix montagni, new species. A, female P5; B, male P5; C, female caudal ramus, ventral view; D, female genital field. E, portion of posterior border of female genital double-somite. VOLUME 110, NUMBER 2 215 — es a | \ Fig. 3. Triathrix montagni new species. A, female antennule dislocated; B, female antennule segmentation only; C, male antennule dislocated; D, male antennule segmentation only; E, rostrum, dorsal view. 216 completely separate and under P6s Gudging by attachment of remains of egg-sac arrow- ed in Fig. 2D). A pair of large tube-pores immediately posterior to gonopores. Copu- latory pore well posterior near median cu- ticular rib, minute and maybe protected by integumental fold. Antennule (Fig. 3A-—B) short, stout, 5-segmented. Segment 1 with 3 rows of spi- nules and 1 large, bipinnate seta. Segment 2 with 4 pinnate and 4 naked setae. Seg- ment 3 with 8 naked setae and an aesthetasc fused at base to one of setae. Segment 4 small with 1 naked seta. Segment 5 with 2 strongly pinnate setae and 7 naked setae on lateral margins; distal margin with a trithek of 2 naked setae and an aesthetasc. Antenna (Fig. 4A). Coxa well-developed with row of setules. Allobasis with 2 strongly pinnate setae on abexopodal mar- gin. Exopod well-developed, 1-segmented with row of setules round distal margin and with 3 setae (a large bipinnate seta on me- dian lateral margin, a large bipinnate seta and a small naked seta in common socket on distal margin). Endopod with 2 rows of strong spinules on anterior margin and a dentate hyaline frill on posterior distal mar- gin; armature consists of 2 spines and a seta subdistally and, on distal margin, 3 finely pinnate spines, 2 geniculate setae, a small seta fused to base of large inner spine, and a tube pore. Mandible (Fig. 4B). Syncoxa robust with row of setules near base of palp; gnathobase with uni- and multi-cuspid teeth and 1 pin- nate seta at distal inner corner. Palp well- developed, 1-segmented, with 5 bipinnate setae (3 on distal and 2 on inner margin). Maxillule (Fig. 4C). Praecoxa with row of marginal setules; arthrite with 2 large tube setae on anterior surface, a row of se- tules on inner margin and, on distal margin, 4 cuspid teeth and a pinnate seta. Coxa well-defined with only 1 pinnate seta on distal margin. Basis with a row of spinules on anterior surface and 8 marginal elements (2 pinnate and 1 naked setae on distal en- dite, 2 naked setae on sub-distal endite, a PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pinnate seta representing endopod, and a pinnate and a naked seta representing exo- pod). Maxilla (Fig. 4D). Syncoxa with 2 mar- ginal rows of spinules and 2 endites; prox- imal endite with 1 fused pinnate spine and 1 slender seta, distal endite with a fused pinnate spine and 2 naked setae. Allobasal endite with a fused pinnate spine and 2 na- ked setae. Endopod represented by 2 setae not fused at base. Maxilliped (Fig. 4E). Syncoxa with 2 rows of spinules but no seta. Basis relative- ly short, oval, with a row of spinules on palmar and outer margin. Endopod repre- sented by a minutely pinnate, recurved spine, a long, well developed accessory seta, and a small finger-like projection. _ P1—P4 (Fig. 5) exactly same in both sexes (Fig. 5B—C). Intercoxal sclerites slender, curved. Protopods ornamented on anterior face as in Fig. 5, same in P1—P4 except that P1 basis with a strong bipinnate seta on inner margin and P2—P4 coxa with an extra row of small spinules centrally. Exopods 3-segment- ed, segments not elongate; Pl (Fig. 5A) with plain inter-segmental hyaline frills, P2—P4 with dentate frills; no tube pores on any seg- ments; exopod-1 to 3 with 3 rows of spinules on outer margin, exopod-2 and -3 with a row of setules on inner margin; outer terminal seta of exopod-3 much longer than distal outer spine and 2 terminal pinnate setae with a comb of pinnules at tip. Endopods 2-seg- mented, endopod-2 much longer than endo- pod-1; both segments with spinules on outer margin, endopod-2 with setules on inner mar- gin; P1 endopod-2 with 3 armature elements, middle one a pinnate seta with a comb-like tip (Fig. 5A). P5 (Fig. 2A). Limbs well separated and connected by slender intercoxal sclerite (not illustrated). Small basal part of baseoendo- pod with a tube pore on anterior surface; outer margin with long pedicel bearing out- er seta; endopodal lobe long, slender (4 times longer than wide), rectangular in shape and reaching to distal margin of ex- opod, with 3 rows of spinules on inner and VOLUME 110, NUMBER 2 217 = SSS aa > Fig. 4. Triathrix montagni new species. A, antenna; B, mandible; C, maxillule; D, maxilla; E, maxilliped. 218 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SNK SS SK SSS — —— SS == Se : Fig. 5. Triathrix montagni new species. A, P1; B, female P3 basis and endopod; C, male P3; D, female P4 protopod and endopod. outer margin, 1 row terminally on posterior rectangular; ornamented with 2 rows of se- surface, and tube pore on anterior surface tules on outer margin, 1 row around distal between inner lateral and inner sub-distal margin, and a tube pore proximally on an- seta; bearing 3 setae inserted and orna-_ terior surface; bearing 3 setae inserted and mented as in Fig. 2A. Exopod separate, ar- ornamented as in Fig. 2A. ticulating with margin of baseoendopod; Male.—As female except in urosome, long, slender (4 times longer than wide), antennule, PS and P6. VOLUME 110, NUMBER 2 Body (Fig. 6) length 0.60—-0.71 mm (X 0.64 mm, n = 8), slightly more slender than female. Urosomite-2 and -3 completely sep- arate. P6 (Fig. 6C) on posterior margin of urosomite-2, an asymmetrical oval plate with row of small spinules but no setae. Antennule (Fig. 3C—D) 6-segmented, sub-chirocer. Segment 1 with 3 rows of spi- nules and a large bipinnate seta on anterior margin. Segment 2 with nine setae (5 pin- nate, 4 smooth) and a tube pore. Segment 3 with 8 setae (%? pinnate, ¥? smooth). Segment 4 strongly swollen with 13 setae and an aesthetasc fused to base of 1 seta on palmar margin; a row of strong setules on dorsal surface. Segment 5 with 2 setae and a tube pore. Segment 6 with a distal trithek and 7 other setae. PS (Fig. 2B). Elements well separated and connected by slender sclerite (Fig. 6C). Baseoendopod small, endopodal lobe vir- tually non-existant, with a tube pore on in- ner margin and 2 terminal setae, outer seta about half length of inner. Exopod separate, articulating with margin of baseoendopod, slender (3 times longer than wide), rectan- gular, with 3 rows of spinules on inner and outer margin and a tube-pore proximally on anterior face; with 2 bipectinate setae dis- tally, outer about half length of inner. Etymology.—This species is dedicated to our friend and colleague Dr. Paul Montagna who provided the specimens from his col- lections. Variation.—Amongst the 19 specimens examined there was no variation in the structure and proportion of the appendages or the major features of the body ornamen- tation. Triathrix kalki, new species Figs. 6D-8 Material examined.—Holotype, an adult female (dissected) USNM 278223; para- types, 4 females (1 dissected), 8 males (1 dissected) and 19 copepodids USNM 278224. All material collected by Dr. P. Montagna at Flower Gardens site, SE of 219 Galveston Texas, in the Gulf of Mexico, 27°54'N, 93°34'W, from a silty sand sedi- ment at 110—160 m depth. The following description is confined to those characters either not described for, or which differ from, T. montagni. Female.—Body (Fig. 7A—B) length 0.44-0.54 mm (X 0.51 mm, n = 4). Chitin- ous extension on posterior border of ceph- alothorax less pronounced and socles more equal in size than in 7. montagni. Median dorsal sensillum-bearing socles on free pro- somites and anterior urosomites fused to- gether into Y-shaped structure, extended dorso-lateral socles without tube pore. Anal operculum (Fig. 8A) fringed with fine se- tules. Caudal rami (Fig. 8A—B) slender, taper- ing posteriorly, 6 times longer than maxi- mum width in ventral view. Setae II and VII inserted slightly more posteriorly (14% and 33% respectively) than in T. montagni. Labrum (Fig. 8C). With a pore and small row of setules medially, oral margin with a short row of long spinules laterally and minute denticles medially. P1—P4 (Fig. 8D-E). Outer terminal seta of P1 exopod-3 (arrowed in Fig. 8D) short- er than outer distal spine. Terminal setae of exopod-3 and endopod-2 without setule comb at tip. Median row of small setules on coxa of P2—P4 (Fig. 8E) slightly more proximal in position than in T. montagni. P5 (Fig. 8F). Baseoendopodal lobe and exopod 6 times longer than wide. Distal in- ner seta on endopodal lobe shorter (in ab- solute length) than proximal inner seta. Male.—As female except for urosome, antennule, PS and P6. Body length 0.45—0.52 mm (X 0.48 mm, n = 8) and slightly more slender than fe- male. Urosomites-2 and -3 completely sep- arate. P5 (Fig. 7C). Exopod 4 times longer than wide (more slender than in 7. montagni). Outer seta on endopodal lobe not reaching distal margin of exopod and only slightly shorter than inner seta. Etymology.—tThe species is dedicated to 220 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Twat) (AFI >) TA) ANTM ae r ae ee Fig. 6. Triathrix montagni new species, male urosome A, dorsal view; B, lateral view; C, ventral view. Triathrix kalki new species, female copepodid V cephalothorax and first two free prosomites, D, dorsal view, E, lateral view. 221 VOLUME 110, NUMBER 2 Fig. 7. Triathrix kalki new species. A, female body, dorsal view; B, female body lateral view; C, male P5. 222 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON % FA A. —=— a “ny aN Lay 2 —~ Mp GP Fig. 8. Triathrix kalki new species. A, Female operculum and caudal ramus, dorsal view; B, female caudal ramus, ventral view; C, labrum; D, P1, E, P2 protopod; E female PS. VOLUME 110, NUMBER 2 our good friend and colleague Mr. Rick Kalke, the leader of the team engaged in the analysis of meiofauna samples from the GOMEX project. Variation.—Amongst the 13 adult spec- imens examined there was no variation in the structure and proportion of the append- ages or the major features of the body or- namentation. Discussion The specimens described above can be placed within the Cletodidae on the basis of the following combination of characters: well defined somites with no hyaline frills but with sensilla and socles on the posterior border; an operculum with a pair of asso- ciated sensillum-bearing socles; an anten- nule which is 5-segmented in the female and 6-segmented in the male; a 1-segment- ed antennary exopod with at most three se- tae; rami of the mandible and maxillule fused to the basis; a maxilla with two syn- coxal endites; a chelate maxilliped without setae on the basis; the Pl with a 2-seg- mented endopod in which endopod-2 is much longer than endopod-1; endopod-1 of P1—P4 without an inner seta; Pl and P2 ex- opod-3 with four setae/spines; the P5s in both sexes not fused medially; the male P6 an asymetrical plate without armature ele- ments. Gee (1994) and Gee & Huys (1996) re- gard the structure of the antenna to be sig- nificant in phylogenetic considerations within the Cletodidae. The antenna of the above species (and of E. nicobarica) is the most primitive condition found in the fam- ily with two setae on the abexopodal mar- gin of the allobasis and a 1-segmented ex- opod bearing three setae, two on the distal margin and one on the lateral margin. Lim- nocletodes also retains the primitive struc- ture of the antennary exopod but has lost one seta on the abexopodal margin of the allobasis. In the original description of Acrenhydrosoma perplexa, Scott (1899) il- lustrates (plate XI, fig. 15) an exopod with 223 three setae but this are not homologous with the above because there is only one seta on the distal margin and two setae on the lat- eral margin. Fiers (1996) is of the opinion that the proximal lateral seta may have been confused with a large spinule, especially in the light of the fact that the other two spe- cies of Acrenhydrosoma Lang (see Lang, 1965 and Schizas & Shirley 1994) have only one terminal and one lateral seta. All other genera within the Cletodidae have ei- ther (a) a well developed exopod with two setae (one distal and one lateral) and one or no setae on the abexopodal margin of the allobasis; or (b) a small cylindrical exopod with one seta and with two or one setae on the abexopodal margin of the allobasis. Although the structure of the antenna in the species described in this paper is ple- siomorphic they are placed in a separate ge- nus on the basis of the following autapo- morphies: A chitinous plate with four so- cles on the posterior margin of the cepha- lothorax. Most genera of cletodid usually have more than four socles arranged around the margin of the cephalothorax but in no species has a chitinous extension to the cephalothorax been described; No socles on the preanal somite. The arrangement of so- cles on the urosomites has not always been accurately recorded but for all species and genera that we have examined in detail there are always two non-sensillum bearing socles on the posterior border of this so- mite; A median tube pore on the opercu- lum. The presence of a pore in this position has not been recorded before in the Cleto- didae but again this could be the result of lapses in observation or resolution of mi- croscopes used by previous workers; Seta I of the caudal ramus inserted posterior to seta II. In all other genera in the family, where the condition is known, seta I and seta II arise in very close proximity on the lateral margin of the ramus; The maxillu- lary coxal endite with only | seta. For many genera and species of Cletodidae the mouthparts are unknown. However, amongst those that are known, there are two © 224 setae on the coxal endite, except in species of Enhydrosoma where the coxal endite is fused to the basis and in Cletodes millero- rum Hamond, 1973 and C. pseudodissimilis Coull, 1971; Maxillulary basal complex with eight setae; three on the distal endite, two on the proximal endite, one represent- ing the endopod and two the exopod. It is thought that the primitive condition in the family is 10 or 11 setae on the basal com- plex, (six or seven on the basal endites and two each representing the exopod and en- dopod). This is the condition found in Cle- todes, Intercletodes Fiers, Strongylacron Gee & Huys and Schizacron Gee & Huys. In Enhydrosoma, Enhydrosomella Monard, Kollerua Gee, Stylicletodes Lang and Acrenhydrosoma (referred to below as the Enhydrosoma group) there is a maximum of six setae on the basal complex (four on the basal endites and one each representing the exopod and endopod). In addition to the above autapomorphies, the genus Triathrix is also characterized by the following features: The sharply pointed, triangular rostrum. The phylogenetic signif- icance (and polarity) of this character is dif- ficult to assess but no other cletodids have a rostrum shape identical to these species and there is evidence (Gee & Huys, 1996) that the rostrum has a characteristic shape in each genus; A mandibular palp with five setae. The primitive condition in the family is Six setae on the mandibular palp (two on the inner, three on the distal and one me- dially on the outer margin), found in some species of Cletodes, Intercletodes and Mon- ocletodes. Triathrix has lost the seta on the outer margin, as has Strongylacron and oth- er species of Cletodes with five palp setae, except C. millerorum, which appears to have lost the proximal seta on the inner margin. Enhydrosoma curticauda Boeck, 1872, the remaining three species of Cle- todes, Schizacron and Acrenhydrosoma maccalli Schizas & Shirley, 1994, have four setae on the palp, all having lost the seta on the outer margin and one on the distal mar- gin. All other genera or species where the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON condition is known, have three setae on the palp, the additional lost seta beings the proximal seta on the inner margin; Proxi- mal maxillary syncoxal endite with two ar- mature elements. The primitive condition in the family is three elements on both syn- coxal endites and is found in most species where the condition has been reliably re- ported. In a number of species of Enhydro- soma there are two elements on the distal endite but, except in Cletodes reductus Moore, 1977 there are no reliable reports of two elements on the proximal endite; Max- illary allobasal endite with a spine and two setae. Again the primitive condition within the family is a spine and three setae on this endite although all genera in the Enhydro- soma group appear to have the Triathrix condition; The two setae representing the maxillary endopod are not fused at the base. This is probably the primitive condi- tion in the family and, as far as we know, the basal fusion of these two setae is only found in genera in the Enhydrosoma group; Maxillipedal coxa without a seta. Most gen- era in the family have retained the seta on the coxa but it has been lost in a large num- ber of the species at present placed in En- hydrosoma (including the type species); The shape of the female P5 in which the baseoendopodal lobe and the exopod are rectangular in shape and equally long. Only in Stylicletodes is the P5 similarly shaped but in this genus there are four or five setae on each ramus compared to only three in Triathrix. The rami of the female P5 in Strongylacron and Schizacron are equal in length and bear three setae but are well sep- arated (giving the limb a characteristic U-shape) and the exopod is fused to the baseoendopod. The female P5 exopod in most species of Cletodes is long and rect- angular but always bears five setae and the endopodal lobe is usually much reduced. Within the family, the morphology and setal arrangement of the P5 in both sexes are im- portant in defining generic boundaries and are undoubtably of great phylogenetic sig- nificance (Fiers, 1996) but at the moment VOLUME 110, NUMBER 2 the relationships of these characters are dif- ficult to understand; No sexual dimorphism on swimming legs. Within the Cletodidae, sexual dimorphism, when present, is found on the P3 endopod, although in three spe- cies of Enhydrosoma (E. longifurcatum Sars, 1909, E. latipes (A. Scott, 1909) and E. pericoense Mielke, 1990) it is also found on the P3 exopod. The range of sexual di- morphism on the P3 endopod, its origins and homologies are discussed in detail by Gee (1994), Gee & Huys (1996) and Fiers (1996). Briefly however, the former two au- thors regard the 3-segmented P3 endopod in the male of Cletodes, Strongylacron, Schizacron and Enhydrosoma curvirostre (T. Scott, 1894) as a development (apomor- phy) from an originally 2-segmented ramus (plesiomorphic condition) and interpret the absence of sexual dimorphism in some spe- cies of Cletodes, Intercletodes and Mono- cletodes as a secondary loss of sexual di- morphism. The latter author, on the other hand regards the 3-segmented ramus on the male P3 as the plesiomorphic condition and absence of sexual dimorphism as the most advanced state. We have not examined the single known specimen of E. nicobarica which is a male copepodid V and the description of Sewell (1940) does not detail the structure of the mouthparts. Nevertheless, the shape of the rostrum, the setation of the antennal exopod and the caudal ramus, and the structure of the juvenile P5, all suggest that this speci- men is a juvenile Triathrix. There is no ev- idence of the posterior extension to the cephalothorax in the drawing of the dorsal view of the body of E. nicobarica (Sewell, 1940 fig. 85A). However, it is known (EF Fiers, pers. comm.) that one of two further new species belonging to this genus from Campeche Bank, Yucatan Peninsula, south- ern Gulf of Mexico, has an even smaller dorsal extension than that of 7. kalki. Fur- ther, an examination of the copepodid V specimens in our sample of T. kalki, shows that the posterior extension of the cepha- lothorax is not developed in the last juve- 225 nile stage (Fig. 6D-E). In addition, it should be noted from Fig. 6D-E, firstly, that there are eight socles on the cephalo- thorax and ten on the free prosomites in co- pepodid V rather than the four and six re- spectively found in the adult and, secondly that the median dorsal socles are well sep- arated in copepodid V rather than fused as in the adult of this species. Thus, it is clear that the distinctive body features of the spe- cies described in this paper may only ap- pear in the adult and that the copepodid V Stage is the same as that drawn by Sewell (1940) for E. nicobarica. Further, Sewell il- lustrated the P1 endopod-2 of E. nicobarica with only two elements (Sewell, 1940, fig. 85F) and noted that the inner distal element of P2 exopod-3 was a spine-like element (rather than a normal seta as in JT. montagni and T. kalki). Developmental studies on Cletodidae (Fiers, 1991, 1996) indicate that all setae present in the adult are also present in copepodid V but may differ slightly in appearance. Thus the absence of an inner seta on Pl endopod-2 in E. nicobarica clearly distinguishes it from the two species described above but the form of the distal elements on P2-endopod-3 may be just a juvenile precursor of a normal seta and not a distinguishing feature. Thus, it is clear that E. nicobarica shouid be transferred to the new genus TJriathrix as T. nicobarica new combination. Finally, Triathrix appears to be limited in its distribution to the Indo-Pacific Oceans and the western Atlantic with three species known to inhabit the Gulf of Mexico region east of the central American isthmus, one the Californian Pacific coast and one the In- dian Ocean. Woodring (1966) presents ev- idence from the fossil molluscan record suggesting that the whole of the Caribbean area and tropical eastern Pacific formed a single biogeographic province during the Miocene. Jones & Hasson (1985) suggest that the Central American isthmus gradu- ally emerged from about the time of the late Miocene with final closure ocurring in the southern part of the isthmus some three | 226 m.y.a. in the late Pliocene or early Pleisto- cene. The present known distribution of Triathrix suggests that the genus was in ex- istence well before that date and that during more recent times speciation within the Gulf of Mexico region has been greater than that in the eastern Pacific region. How- ever, it must be remembered that the har- pacticoid copepod fauna of the Caribbean and Central American region is extremely poorly known, especially for sublittoral communities. Therefore the present known distribution of species on either side of the isthmus may be more a function of sam- pling effort than the pace of evolution. Acknowledgments We wish to thank Dr. Paul Montagna for making all the specimens available to us and for his encouragement and financial as- sistance which has made this work possible. We would also thank Dr. Frank Fiers for making available to us information on two undescribed species from Campeche Bank and Dr. Rony Huys for checking the avail- ability of the generic name. Literature Cited Fiers, F 1991. Three new harpacticoid copepods from the Santa Maria Basin off the Californian Pa- cific coast (Copepoda, Harpacticoida).—Beau- fortia 42(2):13—47. 1996. Redescription of Enhydrosoma lacu- nae Jakubisiak, 1933 (Copepoda, Harpacticoi- da); with comments on the Enhydrosoma spe- cies reported from West Atlantic localities, and a discussion of cletodid development.—Sarsia 81:1-27. Gee, J. M. 1994. Towards a revision of Enhydrosoma Boeck, 1872 (Harpacticoida: Cletodidae sensu Por); A re-examination of the type species, E. curticuada Boeck, 1872, and the establishment of Kollerua gen. nov.—Sarsia 79:83-—107. , & R. Huys. 1996. An appraisal of the taxo- nomic position of Enhydrosoma_ buchholzi (Boeck, 1872), E. bifurcarostratum Shen & Tai, 1965, E. barnishi Wells, 1967 and E. vervoorti Fiers, 1987 with definition of two new genera PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Copepoda, Harpacticoida, Cletodidae).—Sarsia In press. Huys, R., J. M. Gee, C. G. Moore, & R. Hamond. 1996. Marine and brackish water harpacticoid copepods Part 1. Synopses of the British Fauna (Ed. R. S. K. Barnes & J. H. Crothers), Field Studies Council Press, UK. 51:1—352. Hyland, J., D. Hardin, I. Crecelius, D. Drake, P. Mon- tagna, & M. Steinhauer. 1990. Monitoring long-term effects of offshore oil and gas devel- opment along the southern California outer shelf and slope: background environmental con- ditions in the Santa Maria Basin.—Oil and Chemical Pollution 6:195—240. Jones, D. S., & P. E Hasson. 1985. History and de- velopment of the marine invertebrate faunas separated by the Central American Isthmus. Pp. 325-356 in E G. Stehli & S. D. Webb, eds., The Great American Biotic Interchange. Ple- num Press. Lang, K. 1948. Monographie der Harpacticiden I: 1— 896, figs 1-361; II: 897-1682, figs. 362-607, maps 1—378. Hakan Ohlsson, Lund. 1965. Copepoda Harpacitcoidea from the California Pacific Coast—Kungliga Svenska VetenskapsAkademiens handlinger 10:1—560. Montagna, P. A. 1991. Meiobenthic communities of the Santa Maria Basin on the California conti- nental shelf—Continental Shelf Research 11: 1355-1378. , & D. E. Harper Jr. 1996. Benthic infaunal long-term response to offshore production plat- forms.—Canadian Journal of Fisheries and Aquatic Sciences. In Press. Por, EF D. 1986. A re-evaluation of the family Cleto- didae Sars, Lang, (Copepoda, Harpacticoida). In G. Schriever, H. K. Schminke, & C.-T. Shih eds., Proceedings of the Second International Conference on Copepoda, Ottawa, Canada, 13-17th August, 1984.—Syllogeus 58:420—425. Schizas, N. V., & T. C. Shirley. 1994. Acrenhydro- soma maccalli, a new species from Auke Bay, Alaska (Copepoda, Harpacticoida).—Crusta- ceana 67:329-—340. Scott, T. 1899. Notes on recent gatherings of micro- Crustacea from the Clyde and the Moray Firth 17th Annual Report of the Fisheries Board of Scotland for 1898, Part III: 248-273. Sewell, R. B. S. 1940. Copepoda, Harpacticoida.— The John Murray Expedition 1933-34, Scientif- ic Reports, British Museum (Natural History) 7: 117-382. Woodring, W. P. 1966. The Panama land bridge as a sea barrier—Proceedings of the American Philosophical Society 110:425—433. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):227—235. 1997. New records of the genus Hansenomysis in Japan with description of a new species (Crustacea: Mysidacea: Petalophthalmidae) Manuel Rafael Bravo and Masaaki Murano (MRB) Tokyo University of Fisheries, Department of Aquatic Biosciences, 4-5-7 Konan, Minato-ku, Tokyo 108, Japan; (MM) Institute of Environmental Ecology, METOCEAN Co. Ltd., Riemon 1334-5, Ooigawa-cho, Shida-gun, Shizuoka 421-02, Japan Abstract.—A new species, Hansenomysis japonica, and a species tentatively identified as ?7Hansenomysis lucifugus (Faxon, 1893), of the mysid family Pe- talophthalmidae, were collected from Japanese waters. Hansenomysis japonica is clearly distinguishable from the nearest species of the genus, H. violacea (Birstein & Tchindonova, 1958), by the long acute horns of the eyeplate, the narrower antennal scale, the segmented carpopropodus of the endopod of the eighth thoracopod, and the longer and narrower telson. The new species is the fifth described species of Hansenomysis in the Pacific Ocean. ?Hansenomysis lucifugus has not previously been recorded from Japan and western Pacific, if the identification is correct. A key to the species of Hansenomysis is also included. The genus Hansenomysis was established by Hansen in 1887, under the name Arc- tomysis, to incorporate A. fyllae collected southwest of Greenland. However, Arcto- mysis was already allocated to a different species (Czerniavsky 1883), and Stebbing (1893) changed the name to Hansenomysis. Since the establishment of the genus, the classification of its species has experienced notable changes. Hansenomysis lucifugus and H. violacea, were initially described as the two only species of the genus Scol- ophthalmus. Birstein & Tchindonova (1970) transferred these two species of Scolophthalmus to Hansenomysis. Later, Murano & Krygier (1985) transferred five Hansenomysis species to Bacescomysis, which was established by them for B. pa- cifica, based mainly on the exopod of uro- pod which is a 2-segmented plate in the for- mer genus, but unjointed in the latter. The most recent species of Hansenomysis, H. carinata, was described by Casanova (1993) for a single male specimen from the New Caledonian area. Presently, the genus Hansenomysis con- sists of 15 species. The new species, Han- senomysis japonica, is the 16th species of the genus. ?Hansenomysis lucifugus, is re- ported for the first time from Japan and the western Pacific. Table 1 shows the latitu- dinal occurrence, the adult body length, and the habitat of each species of Hansenomy- SiS. The type specimens of H. japonica are deposited in the National Science Museum, Tokyo (NSMT). Order Mysidacea Boas, 1883 Suborder Petalophthalmida Tchindonova, 1981 Family Petalophthalmidae Czerniavsky, 1882 Genus Hansenomysis Stebbing, 1893 Arctomysis.—Hansen, 1887:210. Scolophthalmus.—Faxon, 1893:219; 1895: 224-226. Diagnosis.—Carapace very short. Eyes fused in single plate, without visual pig- | 228 Table 1.—Latitude (n°), adult body length (mm), and habitat or depth (m) of the species of Hansenomysis Stebbing, 1893 (Mauchline & Murano 1977, Lagardére 1983, Casanova 1993). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Species Latitude Body length Habitat/depth Occurrence H. angusticauda O. S. Tattersall, 1961 75S >26 mesopelagic Ross Sea, Palmer Archipelago H. antarctica Holt & Tattersall, 1906 53S—76S 20-23 100—400 Antarctic H. armata Birstein & Tchindonova, 1958 =SON—35N 13 2960 Kurile-Kamchatka Trench H. carinata J.-P. Casanova, 1993 23S 950-1000 New Caledonia H. chini Bacescu, 1971 8S >12 2000 Peru Trench H. falklandica O. S. Tattersall, 1955 50S—53S 12-15 200—400 Southern Oceans H. fyllae (Hansen, 1887) 7ON—-40N 16-17 150—1500 North Atlantic H. japonica new species 35N 12 590 Japan H. lucifugus (Faxon, 1893) 2?35N-—0 42 7742-2000 Off Galapagos, ?Japan H. menziesi Bacescu, 1971 8S 22 2000 Peru Trench H. nouveli Lagardére, 1983 S56N—44N 14-18 1913-2498 Bay of Biscay H. pseudofyllae Lagardére, 1983 48N-44N 14.4 1950-4829 Bay of Biscay H. rostrata Birstein & Tchindonova, 44N 32-35 bathypelagic Kurile-Kamchatka 1970 Trench H. spenceri Bacescu, 1971 8S 17 2000 Peru Trench H. tropicalis Bacescu, 1967 8S >8 2000 Peru Trench H. violacea (Birstein & Tchindonova, 43N 19 bathypelagic Kurile-Kamchatka 1958) Trench ments or with small pigmented area. Dorsal Type species.—Hansenomysis fyllae surface of proximal region of antennular peduncle having what is identified as a sen- sorial organ called the “Tattersall organ”’ (Bacescu 1971). Antennal scale lanceolate with spines and setae. Maxilla and maxil- lule normal. First and 2nd thoracopods ro- bust; endopods of 3rd—5th thoracopods slender, with chelate structure terminally; endopods of 6th—8th thoracopods slender with dactylus and nail together forming long slender claw. Pleopods of female uni- ramous; lst—4th pleopods unsegmented; 5th pleopod longest, 2 or 3-segmented. Pleo- pods of male biramous; 1st with endopod unsegmented, exopod segmented; 2nd pleo- pod with exopod segmented and modified, endopod segmented. Endopod of uropod 2-segmented, without spines on its inner margin; exopod of uropod 2-segmented, proximal segment with spines on outer mar- gin. Telson elongate, entire, without plu- mose setae on apex, posterior part of the lateral margins armed with long strong spines separated by groups of short spines. (Hansen, 1887) Hansenomysis japonica, new species Figs. 1, 2, 3A—D Type specimens.—Holotype (NSMT-Cr 11910), adult male 12.0 mm; paratype (NSMT-Cr 11911), juvenile 8.8 mm; 17 Oct 1990, Sagami Bay (35°09.0'N, 139°24.6’E), 590 m, sledge net. Description of male.—Body robust, elon- gate. Carapace without spines, covering lat- erally part of 7th thoracic somite, and dor- sally all but 6th—8th somites; anterior mar- gin broadly rounded without rostral projec- tion, leaving fused eyes uncovered (Fig. 1A); anterolateral corner sharply pointed. Eyes fused in single plate with 2 acute median horns, outer margin undulated. Eye- plate with 2 fused rounded bulks of visual pigments away from eyeplate margin (Fig. 1A). Antennular peduncle robust; first seg- ment longest, basal dorsal surface with well-developed Tattersall organ (Fig. 1A), VOLUME 110, NUMBER 2 2nd segment about same length as 3rd in dorsal view, shorter in ventral view (Fig. 1B), with blunt process armed with setae at distal outer corner. Outer flagellum very ro- bust, clearly separated into subsegments, each subsegment with rounded inner mar- gin armed with 2 rows of tight setae (Fig. 1B). Antennal scale lanceolate, nearly 5 times as long as the maximum width, extending beyond distal end of antennular peduncle for 0.75 of its length, setose all round ex- cept for proximal 40% of outer margin. Outer distal edge of naked margin with 4 spines that gradually increase in length (Fig. 1C). Peduncle slightly shorter than scale but considerably longer than anten- nular peduncle, 3-segmented, Ist segment very short, 3rd segment about half length of 2nd. Sympod with one spine at base of scale (Fig. 1C). Mandible with strong lacinia mobilis; palp large and slender, 3-segmented, Ist segment shortest, 2nd segment about twice as long as 3rd (Fig. 1D). Labrum symmet- rical, pentagonal, wider than long, without frontal spiniform process (Fig. 1E). Maxil- lule with 7 spines and 1 seta on outer lobe. These spines bear small spinules on mar- gins. Inner lobe with 7 setae, apical 3 large and plumose (Fig. 1F). Maxilla with distal segment of endopod longer than wide, densely setose on inner margin and scarcely setose on outer margin; proximal segment with 4 setae on inner margin; exopod large, with 26 setae on margin (Fig. 1G). First thoracopod small and robust, with- out exopod; endopod with short preischium and dactylus, ischium, merus and carpopro- podus similar in length; dactylus with 3 long plumose spines on distal margin, car- popropodus bearing single, long plumose Spine on inner margin, merus with 5 plu- mose spines on inner margin, ischium with 5 shorter plumose spines on inner margin, preischium and basis with plumose setae but not spines on inner margins (Fig. 1H). Second thoracopod robust, endopod with 1 spine on outer margin of ischium, inner 229 margin produced into very large lamellar lobe armed with many simple setae, preis- chium shortest, merus longest with expand- ed inner distal part, dactylus with long and slender nail (Fig. 2A). Third to 5th thoracic endopods long and slender, forming minute chelate structure terminally, but concealed by crown of long setae (Fig. 2B). Endopod of 3rd thoracopod with carpopropodus un- segmented and about equal to merus in length (Fig. 2C). Endopod of 5th thoraco- pod with carpopropodus longer than merus and divided in 2 subsegments by oblique articulation, proximal subsegment very short (Fig. 2D). Sixth to 8th thoracic en- dopods long and slender, dactylus and nail together forming long slender claw. Endo- pod of 8th thoracopod with carpopropodus separated into 3-subsegments by oblique ar- ticulations, proximal subsegment very short but 2nd shortest, merus longer than carpo- propodus; penis cylindrical (Fig. 2E). Tho- racic exopods distal to basal plate 9-seg- mented in 2nd limb, and 10-segmented in 3rd to 8th limbs; 1st segment longest. Sixth pleonite about 1.7 times as long as Sth. Pleopods developed, biramous. First pleopod with exopod 9-segmented, endo- pod unsegmented, expanded distally, not reaching distal end of lst segment of exo- pod (Fig. 2F). Second pleopod (Fig. 2G) with 7-segmented exopod, Ist segment thick, 2nd segment extended, provided with 2 short simple setae and one strong spinous seta that is spinulose in distal part. This spi- nous seta extending beyond distal end of exopod. Endopod 9-segmented, 1st segment thick and long (Fig. 2G, H). Third pleopod with both rami 9-segmented (Fig. 21). Fourth pleopod with 9-segmented exopod; 8-segmented endopod, Ist segment very long, almost reaching distal end of 3rd seg- ment of exopod (Fig. 3A). Fifth pleopod with 9-segmented exopod; endopod unseg- mented, almost reaching 6th segment of ex- opod (Fig. 3B). Uropods slender, long. Endopod without statocyst, slightly extending beyond distal 230 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON : : 1mm es ey 0.6 mm SS Cre 0.3 mm omgramenaprer tame” ee CR 9 0.4 mm : F H Fig. 1. Hansenomysis japonica, new species. Holotype, adult male. A, anterior part in dorsal view; B, antennular peduncle in ventral view; C, antenna; D, mandible; E, labrum; FE maxillule; G, maxilla; H, 1st thoracopod. Abbreviation, TO: Tattersall organ. VOLUME 110, NUMBER 2 2a Fig. 2. Hansenomysis japonica, new species. Holotype, adult male. A, 2nd thoracopod; B, chela of endopod of 3rd thoracopod; C, 3rd thoracopod; D, 5th thoracopod; E, 8th thoracopod and penis; FE Ist pleopod; G, 2nd pleopod; H, distal part of modified seta on exopod of 2nd pleopod; I, 3rd pleopod. Zaz edge of telson, 2-segmented, lst segment 4.5 times longer than 2nd, which is lanceo- lated, setose all round without spines on in- ner margin. Exopod shorter than endopod, 2-segmented, lst segment about 6 times longer than 2nd, armed in distal half of out- er margin with 2 small, regularly spaced spines, and 3—4 closely set spines near dis- tal end. The latter spines lengthen gradually towards extremity, inner margin setose, 2nd segment setose all round (Fig. 3C). Telson entire (Fig. 3C), long and narrow, almost 3 times longer than 6th pleonite and about 4.2 times as long as broad, distal third tapered posteriorly in 3 steps, each step marked by strong spine; between these spines a series of 3—6 smaller spines. Re- mainder of lateral margin armed with 11-— 12 small spines regularly spaced, proximal 0.2 of lateral margin unarmed (Fig. 3C). Apex without plumose setae, truncate with 9 spines, central spine about same length as outermost spines; penultimate pair of ter- minal spines longest; two pairs of smaller spines on each side of central spine. Mar- ginal spines moderately barbed (Fig. 3D). Etymology.—The species name ‘“‘japoni- ca”’ refers to the collecting locality. Remarks.—Hansenomysis japonica closely resembles H. violacea in general body form, but is easily distinguishable from it by the long acute horns of the eye- plate, the narrower antennal scale, the seg- mented carpopropodus of the endopod of the eighth thoracopod, and the longer and narrower telson. With Hansenomysis ar- mata Birstein & Tchindonova, 1958, H. lu- cifugus, H. rostrata, and H. violacea, the new species is the fifth species of the genus recorded from the Pacific Ocean (Table 1). ?Hansenomysis lucifugus (Faxon, 1893) Fig. 3E-—G Scolophthalmus lucifugus.—Faxon, 1893: 219; 1895:226, pl. LV, fig. 1.—TIllig, 1930:556.—W. M. Tattersall, 1951:243. Material.—Immature female 14.3 mm, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 14 May 1995, Sagami Bay (35°05.9’N, 139°32.0’E), 742 m, sledge net. Remarks.—Hansenomysis lucifugus was established by Faxon (1893) without illus- trations, but a later redescription (Faxon 1895) included illustrations. His descrip- tions and illustrations, however, are brief, so that we cannot compare the present im- mature specimen with his type specimen. The following characters of the present Specimen agree well with those of the type specimen: (1) carapace is produced to form an acute rostrum, anterolateral margins armed with two spines, one behind the ex- ternal margin of the antennule, the other at the anterior inferior angle; (2) eyeplate bears two “‘spines’’; (3) antennal peduncle with second and third segments about equal in length (Fig. 3E); (4) endopod of uropod slender with distal end extending beyond the telson and exopod (Fig. 3F). A differ- ence is found in the fifth female pleopod. In the original description it is two-seg- mented whereas in the present our juvenile specimen it is unsegmented (Fig. 3G). Hansenomysis lucifugus closely resem- bles Hansenomysis rostrata; they are the only two Hansenomysis species having the anterior margin of frontal carapace pro- duced in an acute rostrum. Hansenomysis rostrata, however, differs from the former species in the telson which is ovate in shape and which does not bear large spines on the central region of apex. Distribution.—Hitherto known only from the type locality, eastern Pacific off Gala- pagos. This is the first record of Hanseno- mysis lucifugus for Japan and western Pa- cific, if the identification is correct. Key to species of the genus Hansenomysis (Modified from Bacescu 1971) 1. Carapace with spines. ...1. 02.2.7 242 2 Carapace without spines ........... 6 2. Posterolateral angles of pleonites pro- duced in form of spine-like processes a LS ae Ns inci 3 Posterolateral angles of pleonites not VOLUME 110, NUMBER 2 233 WO AAAs SSS 0.3 mm 0.6 mm Cc aw 0.2 mm D G 1mm = 0.6 mm E Fig. 3. Hansenomysis japonica, new species. Holotype, adult male. A, 4th pleopod; B, Sth pleopod; C, uropod and telson; D, apex of telson. 7Hansenomysis lucifugus (Faxon, 1893). Immature female. E, anterior part in dorsal view; FE uropod and telson; G, Sth pleopod. 234 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON produced in form of spine-like process- 12. Outer margins of antennal scale and ex- CSincs leer Paes. Aah. eee as. By: 4 opod of uropod with spines located 3. Eyeplate with 2 lateral processes among setae .... H. fyllae (Hansen, 1887) > Sere. 2 beng H. menziesi Bacescu, 1971 Outer margins of antennal scale and ex- Eyeplate with 1 central process ...... opod of uropod without spines located PC ee st H. nouveli Lagardére, 1983 aMONng SCtAE .... nus valent 4. Outer margin of antennal scale with 13. Distalmost spine of outer margin of an- spines located among setae ......... tennal scale and of exopod of uropod H. armata Birstein & Tchindonova, 1958 extending beyond apices of respective Outer margin of antennal scale without lamina. Outer margin of exopod of uro- spines located among setae ......... 5) pod with 8 spines ........ H. chini 5) Outer smaroin -of antennal -scale*swith, =. |. ee a ee ee Bacescu, 1971 11-15 spines. Telson ovate ......... Distalmost spine of outer margin of an- H. antarctica Holt & Tattersall, 1906 tennal scale and of exopod of uropod Outer margin of antennal scale with not extending beyond apices of respec- 17—23 spines. Telson long and narrow tave, lamina Fino 324 ok -» « pOAg on 14 H. angusticauda O. S. Tattersall, 1961 14. Outer margin of exopod of uropod with 6. Eyeplate with anterolateral or median less than PO spies i... ... 2c sce eee 15 OMS OF MFOCESSES e455 wns sys Bae Possess 7 Outer margin of exopod of uropod with Eyeplate straight frontally or with weak more than 10 spines. Exopod of uropod lateral or median emargination ...... r2 longer than endopod .... H. carinata 7. Anterolateral or median hors or’pro- 0s ee ee ee ee Casanova, 1993 cesses of eyeplate not acute ........ 8 15. Apex of telson rounded and narrow. Anterolateral or median horns or pro- Antennal scale with spines on distal cesses of eyeplate acute ........... 9 half of outer margin .... H. spenceri Bacescu, 1971 Apex of telson broadly rounded. Anten- nal scale with spines on proximal half of outer margin H. rostrata Birstein & Tchindonova, 1970 Ce ee eS 8. Apex of telson rounded and narrow. Outer margin of antennal scale with spines located among setae H. falklandica O. S. Tattersall, 1955 Apex of telson broadly rounded. Outer margin of antennal scale without spines located among setae .... H. violacea hares Se (Birstein & Tchindonova, 1958) 9. Apex of telson somewhat rectangular with truncate apex. Carapace without oe © © © e ee ee © © © © © Acknowledgments One of the authors, MB, wishes to extend his most sincere thanks to the Ministry of rostral projection 2.1. ..... ee. 10 Education, Science, Sports and Culture of Apex of telson rounded. Carapace with Japan for granting to him the opportunity, or without rostral projection ........ 11 by means of a fellowship, of carrying out 10. Antennal scale shorter than peduncle. Outer margin of exopod of uropod un- armed except for 2 spines confined near distal suture JH. tropicalis Bacescu, 1967 Antennal scale longer than peduncle. Outer margin of exopod of uropod armed H. japonica new species 11. Outer margin of antennal scale without spines located among setae. Carapace with rostral projection .. H. lucifugus i Ae A ct ar BR (Faxon, 1893) Outer margin of antennal scale with spines located among setae. Carapace without rostral projection fy ae H. pseudofyllae Lagardére, 1983 the present study. Literature Cited Bacescu, M. 1967. Further mysids from the Pacific Ocean collected during the XIth cruise of R/V ‘“‘Anton Bruun’’, 1965.—Revue Roumaine de Biologie, Série de Zoologie, 12(3):147—159. . 1971. Contributions to the mysid Crustacea from the Peru-Chile Trench (Pacific Ocean).— Anton Bruun Report 7:1—24. Birstein, Y. A., & Y. G. Tchindonova. 1958. The deep sea mysids of the northwest Pacific Ocean.— Trudy Instituta Okeanologii, Akademiya Nauk SSSR 27:258-355. 1970. New mysids (Crustacea, Mysidacea) VOLUME 110, NUMBER 2 from the Kuril-Kamchatka Trench.—Trudy In- stituta Okeanologii 86:277-—291. Casanova, J.-P. 1993. Crustacea Mysidacea; les Mys- idacés Lophogastrida et Mysida (Petalophthal- midae) de la région néo-calédonienne.—Mé- moire d’Muséum national d’Histoire naturelle 156(0):33-53. Czerniavsky, V. 1883. Monographia Mysidarum Im- primis Imperii Rossici. in Transactions. of St. Petersburg Naturalists’ Society, 18(3):1-102 + pls. 1-31. Faxon, W. 1893. Reports on the dredging operations off the west coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commis- sion Steaner ‘Albatross’, during 1891, Lieut.— Commander Z. L. Tanner, U.S.N., Command- ing. Preliminary descriptions of new species. of Crustacea.—Bulletin of the Museum of Com- parative Zoology 24:217—220. . 1895. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands to the Gulf of California in charge of Alexander Agassiz, car- ried on by the U.S. Fish Commission Steamer “Albatross” during 1891. XV. The stalk-eyed Crustacea.—Memoirs of the Museum of Com- parative Zoology 18:1—292. Hansen, H. J. 1887. Oversigt over det vestlige Gron- lands Fauna af Malakostrake Havkrebsdyr.— Videnskabelige Meddelelser fra Dansk Natur- istrisk Forening 1 Kjgbenhevn 9:5—226, tables 2-7, 1 map. 235 Holt, E. W. L., & W. M. Tattersall. 1906. Preliminary notice of the Schizopoda collected by H.M.S. ‘Discovery’ in the Antarctic region.—The An- nals and Magazine of Natural History, Ser. 7, 17(97):1-11. IHlig, G. 1930. Die Schizopoden der Deutschen Tief- see-Expedition.—Deutschen Tiefsee-Expedition 1898-1899, 22(6):1—229. Lagardére, J.-P. 1983. Les Mysidacés de la plaine abyssale du golfe de Gascogne I. Familles des Lophogastridae, Eucopiidae et Petalophthalmi- dae.—Bulletin d’Muséum national d’Histoire naturelle, Paris, 4° sér, 5, section A, n° 3:809-— 843. Mauchline, J., & M. Murano. 1977. World list of the Mysidacea, Crustacea.—Journal of the Tokyo University of Fisheries 64(1):39-88. Murano, M., & E. E. Krygier. 1985. Bathypelagic mysids from the northeastern Pacific.—Journal of Crustacean Biology 5(4):686—706. Stebbing, T. R. R. 1893. A history of Crustacea.— International Science Serie of London 74:1-— 466. Tattersall, O. S. 1955. Mysidacea.—Discovery Re- ports 28:1—190. . 1961. Report on some Mysidacea from the deeper waters of the Ross Sea.—Proceedings of the Zoological Society of London 137(4):553- Dad: Tattersall, W. M. 1951. A review of the Mysidacea of the United States National Museum.—The United States National Museum, Bulletin 201: 1-292. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):236—241. 1997. Nanomysis philippinensis, a new species (Crustacea: Mysidacea) from brackish waters of the Philippines Masaaki Murano Institute of Environmental Ecology, METOCEAN Co. Ltd., Riemon 1334-5, Ooigawa-cho, Shida-gun, Shizuoka 421-02, Japan Abstract.—A new species, Nanomysis philippinensis, is described based on specimens from the Philippines. N. philippinensis is easily distinguished from the two known species of the genus, N. siamensis and N. insularis, by the numbers of setae on the first segment of the exopods of the male third and fourth pleopods, the shape and the marginal spine number of the telson, and the size of body. Specimens of a new species of Nano- mysis were found in plankton samples col- lected with a scoop net at a cove in Panay Island, the Philippines and in those stored at the Southeastern Asian Fisheries Devel- opment Center (SEAFDEC) in Iloilo of the same island. In this paper, the description of the new species is given. The type spec- imens are deposited in the National Science Museum, Tokyo (NSMT). Genus Nanomysis Tattersall, 1921 Nanomysis Tattersall, 1921: 408—409.—li, 1964: 426. Diagnosis.—Carapace fringed with spi- nules on anterior margin. First, second and fifth pleopods of male rudimentary, unjointed and of the same form as those in female. Third pleopod of male biramous with unjointed inner ramous and 3-jointed outer ramous; outer ramous longer than inner, third joint terminating in single strong seta. Fourth pleopod of male biramous; inner ramous unjointed; outer ra- mous very long, 4-jointed, penultimate joint with long seta, ultimate joint short, termi- nating in 2 long, slender setae. Antennal scale narrowly lanceolate, 2-jointed, setose along entire margin, with apical part narrow but not pointed. _ Carpopropodus of endopods of third to eighth thoracic limbs 3- or 4-jointed. Endopod of uropod without spines on in- ner margin. Telson short; posterior margin convex, Straight or concave, not split, armed with a comb of spines between last lateral spines; lateral margins armed with spines. Type species.—Nanomysis siamensis Tat- tersall, 1921. Remarks.—The original diagnosis of the genus was given by Tattersall (1921) when the genus was established for Nanomysis siamensis and amended by Ii (1964) who took into account the description of the sec- ond species, Nanomysis insularis, described by Nouvel (1957). With the addition of the present new species, the generic diagnosis is modified again as mentioned above. Modifications are made in three points, the addition of character in the anterior margin of the carapace, the number of subsegments in the carpopropodus of thoracic endopods, and the shape of the posterior margin of the telson. Nanomysis philippinensis, new species Figs. 1, 2 Type material.—Holotype (NSMT-Cr 11912), adult female with embryos, 3.2 mm; allotype (NSMT-Cr 11913), adult VOLUME 110, NUMBER 2 male, 2.6 mm; paratypes (NSMT-Cr 11914), 5 adult females with embryos (2.8— 3.0 mm) and 8 adult males (2.1—3.0 mm); Batan Bay, Panay Island, Philippines, 1 Dec 1979 collected with scoop net above eel grass bed at depth of about 1 m. Other material.—Two adult females with embryos (3.0, 3.1 mm), 7 adult males (2.2— 2.9 mm) and 1 immature male, Hamtik, Pa- nay Is., 5 Apr 1976, 5 m deep, CM net (75 cm in diameter, 0.49 mm in mesh size); 2 immature females and 1 juvenile, off Ne- gros Occidental School of Fisheries, Negros Is., Philippines, 28 Jun 1976, plankton net tow, 1 juvenile (1.4 mm), Oton Beach, Pa- nay Is., 25 Aug 1976, plankton net tow; 10 immature females (up to 2.5 mm) and 4 im- mature males (up to 1.9 mm), same as type specimens; 3 adult females (2 with embry- os) (2.9—3.2 mm), 11 adult males (2.2—2.8 mm), 11 immature females (up to 2.8 mm) and 3 immature males, mouth of Altavas River, Banga Cove, Batan, Panay Is., 2 Dec £979: Body length.—Adult female 2.8—3.2 mm, adult male 2.1—2.9 mm. Description.—Carapace slightly pro- duced into broadly rounded rostrum, frontal margin fringed with about 60 tiny spinules throughout (Fig. 1B); anterolateral corners rounded; posterior margin emarginate, leav- ing last thoracic somite exposed (Fig. 1A). Eye large; cornea globular, slightly broader than eyestalk; eyestalk without pap- illiform process (Fig. 1B). Antennular peduncle of female: first seg- ment as long as following 2 segments to- gether, armed at outer distal corner with several setae, one of which is plumose, lon- ger and directed backward; second segment connecting obliquely with third segment, with single plumose seta at median distal corner; third segment with plumose seta at median distal corner (Fig. 1A, C). Anten- nular peduncle of male: more robust than that of female, first segment as long as third, third segment with processus mascu- linus small (Fig. 1B). Antennal scale slender, lanceolate, ex- 257 tending beyond distal margin of antennular peduncle for % of its length in female and %4 in male, 5 times as long as greatest width, setose all round, distal suture marked off at distal %, distal segment more than 3 times as long as broad (Fig. 1D). Antennal pe- duncle more than half as long as scale, sec- ond segment longest, occupying about half of peduncle length (Fig. 1D). Antennal sympod with strong spinelike process at outer distal corner (Fig. 1D). Mandibular palp relatively small, sparse- ly setose, second segment armed on exter- nal margin with 7 setae of which distal one differs slightly in shape from others, third segment about half as long as_ second, armed with 8 barbed setae arranged regu- larly on distal % of external margin (Fig. 1E). Coxa of mandibles as shown in Fig. 1E Maxillule: outer lobe armed with 9 strong spines on distal margin and 3 thick setae on inner surface; inner lobe armed with 3 stout and one slender setae on apex, 2 setae on inner margin and 3 setae on outer margin (Fig. 1G). Maxilla: exopod small and slender, 3 times as long as broad, with only 2 setae, one at apex plumose, long and thick, the other on subapex of outer margin short, margin of exopod fringed with fine hairs; terminal segment of endopod oval, 1.2 times longer than broad (Fig. 1H). Endopod of first thoracopod robust; en- dite of basis large, armed with 4 to 5 thick and hairy setae on inner margin and apex; preischium with 3 to 4 similar setae on in- ner margin (Fig. 11). Endopod of second thoracopod robust; ischium fused with preischium, merus equal to combined length of carpopropodus and dactylus, car- popropodus becoming broader distally, twice as long as maximum breadth at distal end; dactylus broader than long, bearing 5 strong barbed setae in addition to slender setae (Fig. 1J). Endopods of third to eighth thoracic limbs becoming more slender to- wards posterior pairs; dactylus small, half length of slender terminal claw; in third to seventh pairs carpopropodus divided into 3 subjoints of which the middle is shortest — 238 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON F Fig. 1. Nanomysis philippinensis, new species. A, C—J, holotype (female); B, allotype (male). A, whole body, dorsal; B, anterior end, dorsal; C, antennular peduncle; D, antenna; E, mandibular palp; FE laciniae and molar parts of mandibles; G, maxillule; H, maxilla; I, first thoracic limb; J, endopod of second thoracic limb. VOLUME 110, NUMBER 2 (Fig. 2A—C); in eighth thoracopod, carpo- propodus divided into 4 subjoints, armed with strong antler-shaped spine at middle and distal end of outer margin of first sub- joint and at outer distal end of second and third subjoints (Fig. 2D). Exopods of tho- racic appendages 9-jointed in first to sev- enth pairs (Fig. 11, Fig. 2A—C) and 8-joint- ed in eighth; basal plate with outer distal corner rounded. Abdominal somites without furrows or spines, last somite slightly longer than pre- ceding one, 1.3 times as long as broad (Fig. 1A). Third and fourth pleopods of male mod- ified. Third pair extending backwards be- yond middle of sixth abdominal somite, bi- ramous; endopod unsegmented and short; exopod 3-jointed: first joint somewhat curved outwardly much longer than second and third joints combined, armed with a small thin seta at outer distal corner, sec- ond joint 4 times as long as broad, armed with a small thin seta at outer distal corner, third joint % as long as second, terminating into long, stout, naked seta which is more than twice as long as joint (Fig. 2E). Fourth pair biramous; endopod unseg- mented and short; exopod elongate, ex- tending posteriorly near distal end of tel- son, 4-jointed: first joint longest, 2.5 times longer than endopod, furnished with 5 short setae regularly spaced on distal ¥, of outer margin, second joint slightly shorter than first, unarmed, third joint slightly shorter than second, armed at outer distal corner with stout seta which is 1.5 times as long as joint and extending beyond tip of terminal setae, terminal joint short, armed on apex with 2 setae being equal in length but different in structure, one 2-jointed indistinctly near base, basal joint swollen (Fig. 2F). Uropod setose along entire margin; en- dopod more than twice as long as telson, tapered, without spines on inner ventral margin; exopod slightly longer than en- dopod, with straight outer margin (Fig. 1A). 239 Telson short, trapezoid, 0.7 as long as last abdominal somite, 1.2 times longer than maximum width, posterior margin about % of maximum width at base, con- cave or straight, armed with long spine at each corner between which 5 to 10 short, slender spines are inserted, average num- ber of distal inserted spines is 6.5 in male (n = 6) and 7.2 in female (n = 5); lateral margin straight or slightly concave, armed with 5—9 short spines along entire length (Fig. 2G). Etymology.—The species is named after the locality in which it was collected. Remarks.—The present specimens be- long clearly to the genus Nanomysis, which was established by Tattersall in 1921, in having the carapace with spinules on the anterior margin, the slender anten- nal scale, 3-segmented exopod of the male third pleopod, 4-segmented exopod of the male fourth pleopod, and the trapezoid tel- son. Nanomysis philippinensis, new spe- cies, is clearly different from two known species of Nanomysis, N. siamensis Tatter- sall, 1921, and N. insularis Nouvel, 1957, as follows. The first segment of the exopod of the male third pleopod is armed with only a single thin seta at the outer distal corner in the new species, while armed on outer margin with 3 and 5 long setae in N. siamensis and N. insularis, respectively. The first segment of the exopod of the male fourth pleopod is armed with 5 setae on the outer margin in N. philippinensis, as against 3 in N. siamensis (Tattersall did not describe this character, but illustrated it in plate XV, fig. 10) and 7 setae in N. insularis. The posterior margin of the tel- son is concave or straight and with 5—10 shorter spines between longer spine at each corner in N. philippinensis, whereas it is convex and with 12 shorter spines in N. siamensis and 15 in N. insularis. The num- ber of spines arming the lateral margin of the telson is 5-9 in N. philippinensis as against 10 in N. siamensis and 11-12 in N. insularis. The present new species is con- siderably smaller (2.8—3.2 mm in the adult 240 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a iw WW WN igs) loca le ae < H ¢ WOK Fig. 2. Nanomysis philippinensis, new species. A—D, G, holotype (female); E, F allotype (male). A, third thoracic limb; B, sixth thoracic limb; C, seventh thoracic limb; D, extremity of endopod of eighth thoracic limb; E, third pleopod (seta of first and second joint closely oppressed to following segment); E fourth pleopod; G, telson. VOLUME 110, NUMBER 2 female, 2.1—2.9 mm in the adult male) than the two known species (5 mm in the adult male of N. siamensis and 4.5 mm in N. insularis). Acknowledgments I express my thanks to the ex-Vice-di- rector of SEAFDEC, Mr. N. Hoshino, for giving me the opportunity of examining the plankton material stored in SEAFDEC. 241 Literature Cited li, N. 1964. Fauna Japonica, Mysidae (Crustacea). Biogeographical Society of Japan, Tokyo, 610pp. Nouvel, H. 1957. Mysidacés provenant de deux échantillons de ((Djembret)) de Java.—Zoolo- gische Mededelingen 35:315-331. Tattersall, W. M. 1921. Zoological results of a tour in the Far East. Mysidacea, Tanaidacea and Is- opoda.—Memoirs of the Asiatic Society of Bengal 6:403—433, pls. 15-17. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):242—248. 1997. A new species of bopyrid isopod, Pseudione chiloensis, a parasite of Nauticaris magellanica (A. Milne-Edwards, 1891) (Crustacea: Decapoda: Hippolytidae) Ramiro Roman-Contreras and Ingo Wehrtmann Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Aut6noma de México (UNAM), P.O. Box 70-305, Mexico, D.E 04510 (IW) Alfred Wegener Institut. Am Handelshafen 12, 27560 Bremerhaven, Germany Abstract.—Pseudione chiloensis sp. nov., is described from the central- southern Chilean region, parasitic on the hippolytid shrimp Nauticaris magel- lanica (A. Milne-Edwards, 1891). Females of Pseudione chiloensis are similar to Pseudione affinis (Sars, 1882), Pseudione indica Chopra, 1930 and Pseu- dione parviramus Adkison, 1988 in having a wide frontal lamina, first antennae 3-segmented, 5 pairs of biramous pleopods, uropods uniramous, and eyes pres- ent; Pseudione chiloensis differ from these species in the absence of ornamen- tations on internal crest of oostegite 1, female’s size, and setose maxillipedal palp. Males of Pseudione chiloensis are similar to those of Pseudione affinis, Pseudione indica, and Pseudione parviramus in having eyes present, first an- tennae 3-segmented, pleon of six pleomeres, midventral tubercules present on pereomeres, 5 pairs of uniramous pleopods, and last pleomere bifurcated; but differ from these species in having the second antennae 6-segmented, (7-seg- mented in the former species). Although males and females of Pseudione chil- oensis share some characters with these species, the morphological differences and host lead us to propose a new species. Resuimen.—Pseudione chiloensis n. sp., es descrita de la regi6n centro-sur de la costa chilena, parasito del hipolitido Nauticaris magellanica (A. Milne- Edwards, 1891). Las hembras de Pseudione chiloensis son similares a Pseu- dione affinis (Sars, 1882), Pseudione indica Chopra, 1930 y Pseudione parvir- amus Adkison, 1988 en que tiene la primera antena 3-segmentada, 5 pares de pleépodos birrameos, urépodos unirrameos y ojos presentes. Pseudione chil- oensis difiere de esas especies en la ausencia de ornamentaciones en la cresta interna del oostegito 1. Pseudione magna Shiino, 1951 difiere de Pseudione chiloensis, Pseudione affinis, Pseudione indica, y Pseudione parviramus en que tiene la segunda antena 4-segmentada, lamina frontal angosta y ausencia de ojos. Los machos de Pseudione chiloensis son similares a los de Pseudione affinis, Pseudione indica y Pseudione parviramus en la presencia de ojos, pri- mera antena 3-segmentada, pleén de seis ple6meros, tubérculos medio-ventra- les en los pereOmeros, 5 pares de pleépodos unirrameos y el ultimo pleémero bifurcado. Los machos de Pseudione chiloensis difieren de Pseudione magna en la primera y segunda antenas que son 2- y 4-segmentadas, respectivamente, en la ultima especie; en Pseudione parviramus las segundas antenas son 7-seg- mentadas, y 6-segmentadas en Pseudione chiloensis. Aunque esta especie com- parte caracteristicas con las especies mencionadas, al mismo tiempo presenta caracteres y hospedero particulares para ser propuesta como una especie nueva. VOLUME 110, NUMBER 2 Three species of the genus Pseudione have been described from the Chilean wa- ters: Pseudione tuberculata Richardson, 1904 parasitizing Neolithodes diomedae (Benedict); Pseudione paucisecta Richard- son, 1904 parasitic on Munida curvipes Benedict, and Pseudione battstroemi Stuar- do, Vega, & Céspedes, 1986 parasitic on Callianassa uncinata H. Milne-Edwards. We report a new species of Pseudione par- asitizing Nauticaris magellanica (A. Milne- Edwards, 1891), an hippolytid shrimp from the South American coasts including the Falkland Islands (Boschi 1979). In Chile Nauticaris magellanica is one of the most abundant shrimps associated with mussel raft cultures (Aracena & Lé6pez 1973, Wehrtmann & Albornoz 1997), and has been collected from holdfasts of the kelp Macrocystis pyrifera from the southern region (Ojeda & Santelices 1984). Howev- er, despite intensive sampling efforts over the past 4 years along the entire coast of Chile, only eleven parasitized individuals of N. magellanica were obtained exclusively from Putemutn, Chiloé, central southern Chile. Nauticaris marionis Bate, has been re- corded in New Zealand (Page 1985) as a host for Pseudione affinis, this being the first record of Pseudionidae parasitizing the genus Nauticaris. Although our specimens share attributes with other species of Pseu- dione, there are differential characters which justify a new species. Two individ- uals male and female, were chosen as type specimens and prepared for SEM photo- graphs. Order Isopoda Suborder Epicaridea Family Bopyridae Rafinesque, 1815 Subfamily Pseudioninae R. Codreanu, 1967 Genus Pseudione Kossmann, 1881 Pseudione chiloensis, new species (Figs. 1-15) Holotype female (dry).—USNM-274301; allotype male (dry): USNM-274302 (both 243 male and female, mounted on a stub for MEB). Material examined _ (Paratypes).— USNM-274256 (2 specs.), USNM-274257 (1 spec.); ICMyL-UNAM-4999 (1 spec.), ICMYL-UNAM-S000 (7 specs.). One adult male and 1 female, 5 Jun 1993; 1 female with cryptoniscus larvae associ- ated, 5 Jun 1993; 2 males and 2 females, 12 May 1994; 1 male and 1 female, 5 Jun 1994; 5 males and 5 females, 13 May 1994; 1 male and 1 female, 26 Jun 1994. Two fully developed individuals male and fe- male, were used for descriptions and se- lected as type specimens. Type _ locality.—Putemtin (42°25'S; 73°43'’W), Chiloé Island, central-southern Chile. Host.—Nauticaris magellanica; most in- fected individuals were males (n = 10; total length varying between 14.93 and 19.15 mm; X = 17.96 mm); the only non-oviger- ous female infected measured 22.61 mm to- tal length. Habitat.—The shrimps parasitized were associated with massive cultures of Mytilus chilensis, and were collected from approx- imately 5 m depth (salinity 29.5—29.7 ppm; temperature 9.0—10.5°C). Description of female (Figs. 1—8).— Length 4.0 mm, width 2.9 mm; head tri- angular, slightly wider than long, deeply set into first pereomere but discernable (Fig. 2); barbula with two lateral projections on each side; anterolateral borders of head almost rounded, frontal lamina wide, rounded (Figs. 1, 2); maxilliped anterolaterally rounded, falcate, 3—4 setae on upper border, unarticulated palp (Fig. 3). First antenna 3 segmented; basal segment globose; second segment cylindrical, almost as high as for- mer, 5 small aesthetascs on upper border; third segment cylindrical, smooth, 0.75 slender than previous, 5 aesthetascs on tip (Fig. 4). Second antenna 5 segmented, basal segment subovoid, other segments cylindri- cal, almost as high as the former but slen- der; fourth and fifth segments smooth, api- cal segment with a tuft of 5 setae (Fig. 5); 244 distal segment projects slightly beyond the border of frontal lamina; squamous struc- tures on surface of both antennae; eyes present. Pereon margins forming smooth curve; thin, conspicuous rectangular coxal plates on pereomeres 1—4 conspicuously exceeding border; tergal projections nar- row, pigmented on short side. Oostegite 1 covering head and partially anterior portion of brood pouch; anterior lobe rounded, higher than posterior, no ornamentation on internal ridge (Fig. 6). Strong pigmentation on oostegites of short side; some chromat- ophores dispersed on oostegites 2—4 of op- posite side. Oostegites 2—4 foliate, fifth oostegites fringed, slender, extending across posterior region of brood pouch, and over- lapping opposite one. First two pereopods smaller than last five, small subcuadrangu- lar carina on basis of first four pairs (Fig. 7); last three pairs of pereopods not cari- nated. All articles distinct, small scales on ventral surface of legs, four small setae on carpus, dactyli deeply set into propodi. Pleomeres completely separated dorsal and laterally, ending in lateral, foliate, and rounded plates. Five pairs of biramous fo- liate pleopods, partially covering ventral surface of pleon (Fig. 1); border of pereo- meres 5—7 dorsally folded and directed for- ward on short side (Fig. 8); endopods short- er and narrower than exopods, lanceolate, both rami progresively larger from first to fifth pleopods; uropods uniramous present. Description of male (Figs. 9—15).— Length 0.96 mm, width 0.26 mm. Head an- teriorly rounded, wider than long, partially fused with first pereomere but clearly dis- cernable; anterolateral borders rounded; eyespots irregular, conspicuous (Fig. 9). First antenna three segmented, globose ba- sal segment with 2 small setae on external surface; cylindrical medium-sized, second segment; third segment 0.6 smaller than second, ending in a tuft of 6 aesthetascs, small scales on surface (Fig. 10). Second antenna 6-segmented, almost 3 times larger than first one. Basal segment truncated, subpyramidal; second one higher than first; PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON third and fourth segments cylindrical, smooth, slender than former, some setae on upper border; fifth segment slender and shorter than previous; distal segment with tuft of 5—6 setae (Fig. 11) reaching first per- eomere; both antennae with scarce scales on surface. Pereomeres all similar, deeply sep- arated laterally and ventrally, tips rounded and reflexed ventrally, with small scales on borders, mid-ventral tubercules on pereo- meres (Fig. 12). All pereopods similar in size, not carinated (Fig. 13); dactyli inserted into propodi (Fig. 14). Five distinct pleo- meres separated dorsal, ventral, and later- ally; five pairs of sessile tuberculiform ple- opods on pleomeres 1—5, prominent and conspicuous in ventral view, anal cone present; sixth pleomere bifurcated, ending in short setae, sparse small scales on the tips (Fig. 15); no uropods. Etymology.—The specific name is in ref- erence to Chiloé, the type location. Gender masculine. Variations.—Females varied in pigmen- tation patterns with oostegites 1—5 totally pigmented, and other individuals with oos- tegites only partially pigmented on the short side; in males the sixth pleomere are bi- lobed, but others with button-shaped or Y-shaped pleomere. Discussion This century many authors have cited the need for a revision for the genus Pseudione. The literature shows an immense diversity of forms and the genus parasitizes many families of Anomura and Brachyura, but a few species are known to infect certain gen- era of Caridea (Chopra 1930). We decided to include our material in the genus Pseu- dione because of the morphological char- acteristics are in agree with Sars (1899), Bourdon (1968), and Markham’s (1985) de- scriptions; and, although Danforth (1971) quoted that ‘‘approximately 60 species of Pseudione so far described, identification of a new form is quite difficult’’, many species of Pseudione have since been described 245 VOLUME 110, NUMBER 2 1ew. ventral v first and second 4999), paratype (ICMYL-UNAM- I, h setose palp (arrow). F 1g Ef 1€S hiloensis, new spec tone C. female. Pseud. Figs. 1-6. Fig. 2, head of same > 4-5 igs iped wit . Fig. 3, maxill 1e¢w dorsal v > in mm) (scales iew lv ite, interna antennae. Fig. 6, first oosteg 246 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 7-15. Pseudione chiloensis. Fig. 7, second pereopod (female) Fig. 8, holotype female (dry; USNM- 274301), dorsal view. Figs. 9-15 male. Fig. 9, allotype, dorsal view (dry; USNM-274302). Fig. 10-11, first and second antennae. Fig. 12, pleon (broken) in ventral view. Figs. 13-14, seventh leg. Fig. 15, sixth pleomere and anal cone; (scales in mm). VOLUME 110, NUMBER 2 from American waters (Stuardo et al. 1986, Adkison 1988, Adkison & Heard 1995). Females of Pseudione chiloensis are sim- ilar to those of Pseudione affinis, Pseudione indica Chopra, 1930 and Pseudione parvir- amus, in having a wide frontal lamina, eyes present, first antennae 3-segmented, five pairs of biramous pleopods, and uropods uniramous (Sars 1899, Chopra 1930, Bour- don 1968, Adkison 1988). Pseudione chil- oensis is similar to Pseudione magna Shi- ino, 1951 in that the first antennae is 3-seg- mented, pleopods biramous, and uropods uniramous; but the second antennae are 4-segmented, the frontal lamina narrow, and the eyes are absent in the later species. Pseudione chiloensis differs from P. par- viramus, P. affinis, P. indica, and P. magna in the absence of ornamentation on internal crest of oostegite 1, and presence of setose maxillipedal palp. The body size of females is different in the former species as follows: 10.0 and 14.5 mm in P. affinis (Sars 1899, Bourdon 1968), 20.2 mm in P. magna (Shiino 1951), and 9.0—11.0 mm in P. parviramus (Adki- son 1988). Pseudione chiloensis however, is smaller (4.0 mm) than the former species and more similar in size to P. indica (2.0 mm; Shiino 1951). Males of P. chiloensis are similar to those of P. affinis, P. magna, P. parvira- mus, and P. indica in having eyes present, pleon six-segmented, and five pairs of tub- erculiform pleopods; although the males of P. magna differs from P. chiloensis, P. af- finis, P. indica, and P. parviramus in hav- ing the first antennae 2-segmented, while these structures are 3-segmented in the re- maining species. The second antennae of the males of P. chiloensis are 6-articulated, 7-segmented in P. affinis, P. indica and P. parviramus, and 4-segmented in P. magna. The males of P. chiloensis have conspic- uous midventral tubercles on pereomeres sixth and seventh, and on the first two pleo- meres, and the final pleomere produced into bifurcated and setose structure, similarly to P. indica (Markham 1994), but not uropods 247 in Adkison & Heard’s sense (Adkison & Heard 1995). In addition, the males of P. indica closely resembles that of Pseudione cognata Markham, 1985 (Markham 1994). Acknowledgments The study was partially financed by In- stituto de Ciencias del Mar y Limnologia, Universidad Nacional Aut6noma de Méxi- co (ICMYL-UNAM) (RRC), the ‘“‘German Academic Exchange Service’’, the ‘‘Deut- sche Gesellschaft fiir Technische Zusam- menarbeit’”’ and the Universidad Austral de Chile (DID-Project No. E-91-1 and S-94- 53) ((W). We are grateful to Y. Homelas (ICMyL-UNAM) for SEM micrographs, from which the figures were inked by R. Mendoza (Facultad de Ciencias, UNAM). J. Clark, Collection Manager, Division of Crustacea, United States Natural Museum (USNM) provided facilities to one of us (RRC) during a short visit to the Division in July 1995. We appreciate the cooperation with the “Instituto de Fomento Pesquero’”’ which allowed one of us (IW) to carry out the sampling program at its station in Pu- temun, Chiloé, Chile. J. Savitz (Chesapeake Bay Foundation) kindly revised our En- glish. FE Ferrari, Associate Editor of PBSW, and three anonymous referees improved greately the final manuscript. Literature Cited Adkison, D. L. 1988. Pseudione parviramus and Apo- robopyrus collardi, two new species of Bopyr- idae (Isopoda: Epicaridea) from the Gulf of Mexico—Proceedings of the Biological Society of Washington 101:576—584. , & R. W. Heard. 1995. Pseudione overstreeti, new species (Isopoda: Epicaridea: Bopyridae), a parasite of Callichirus islagrande (Decapoda: Anomura: Callianassidae) from the Gulf of Mexico.— Gulf Research Report 9(2):105—110. Aracena, P. O., & M. T. Lépez. 1973. Observaciones biol6gicas en organismos encontrados en sub- stratos artificiales. Caleta Leandro, Talcahuano, Chile. I. Crustacea, Decapoda, Macrura.—Tra- bajos V Congreso Latinoamericano de Zoologia 1:40-—48. Boschi, E. E. 1979. Geographic distribution of Ar- gentinian marine decapod crustaceans.—Bulle- 248 tin of the Biological Society of Washington 3: 134-143. Bourdon, R. 1968. Les Bopyridae des Mers Euro- péennes.—Mémories du Muséum National D’Histoire Naturelle, Paris. Serie A, Zoologie, Tome L, Fascicule 2:76—424. Chopra, B. 1930. Further notes on Bopyrid isopods parasitic on Indian Decapoda Macrura.—Re- cords of the Indian Museum 32(2):113-147. Danforth, Ch. G. 1971. Two Bopyrids (Isopoda) from New Guinea.—Bulletin Southern California Academy of Sciences 70(2):99-—102. 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. 1994. Crustacea Isopoda: Bopyridae in the MUSORSTOM collections from the tropical Indo-Pacific I. Subfamilies Pseudioninae (in part), Argeniinae, Orbioninae, Athelginae and Entophilinae. Pp. 225-253 in A. Crosnier, ed., Résultates des Campagnes MUSORSTOM, Volume 10—Mémoirs du Muséum national d’ Histoire Naturelle, Paris 161:225—253. Ojeda, E P, & B. Santelices. 1984. Invertebrate com- munities in holdfasts of the kelp Macrocystis PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pyrifera from southern Chile.—Marine Ecology Progress Series 16:65-73. Page, R. D. M. 1985. Review of the New Zealand Bopyridae (Crustacea: Isopoda: Epicaridea).— New Zealand Journal of Zoology 12:185-—212. Richardson, H. 1904. Contributions to the natural his- tory of the Isopoda.—Proceedings of the United States National Museum 27(1350):1—89. Sars, G. O. 1882. Oversight af Norges Crustaceer med Forelobige Bemaerkninger Over de Nye Eller Mindre Bekjendte Arter. I. (Podophthalmata- Cumacea-Isopoda-Amphipoda).—Christiana Videnskabelige Solk. Forhandling 18:1—124. . 1899. An account of the Crustacea of Nor- way. II. Isopoda.—Publications of the Bergen Museum, Bergen, pp. 195-205. Shiino, M. S. 1951. Some bopyrid parasites found on the Decapod Crustaceans from the waters along Mie Prefecture.—Report of Faculty of Fisher- ies, Prefectural University of Mie 1(1):26—40. Stuardo, J., R. Vega, & I. Céspedes. 1986. New bo- pyrid isopod parasitic on Callianassa uncinata H. Milne-Edwards: with functional and ecolog- ical remarks.—Gayana Zoologia 50(1—4):3-15. Wehrtmann, I. S., & L. Albornoz. 1997. Larval de- velopment of WNauticaris magellanica (A. Milne-Edwards, 1891) (Decapoda, Hippolyti- dae), reared under laboratory conditions.—Bul- letin of Marine Science. (in press). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):249—255. 1997. Observations on Hexapanopeus schmitti Rathbun from Brazil (Crustacea: Decapoda: Xanthidae) C. Sankarankutty and Raymond B. Manning Departamento de Oceanografia e Limnologia, Universidade Federal do Rio Grande do Norte, Praia de Mae Luiza S/N, Via Costeira, Natal RN-59014-100, Brazil; Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Observations are presented on members of a population of Hex- apanopeus schmitti Rathbun from Pitangui, Rio Grande do Norte, Brazil. This species is very small, apparently not exceeding 10 mm in carapace length. It is illustrated in detail as are the gonopods of the four species of Hexapanopeus known from Brazil. A key to Brazilian species of Hexapanopeus is provided. Field studies along the coast of Rio Gran- de do Norte, Brazil by one of us (C:S.) yielded a series of specimens of Hexapan- opeus schmitti Rathbun, 1930, originally described from material from localities in Brazil and Uruguay. The population record- ed here shows wide variation in color pat- tern that has not been recorded previously. Gonopods are illustrated for the four Bra- zilian species of Hexapanopeus, and a key to those species is presented. Abbreviations used in the accounts below include: cb, carapace breadth; cl, carapace length; mm, millimeter; P1-5, first to fifth pereopods (P1 is the cheliped, P2-5 the walk- ing legs). USNM is an acronym for the Na- tional Museum of Natural History, Smithson- ian Institution, Washington, D.C., where some of the newly-collected specimens are deposited. The majority of the specimens are in the collections of the Museu do Mar “On- ofre Lopes” at the Universidade Federal do Rio Grande do Norte, Natal, Brazil. Family Xanthidae MacLeay, 1838 Hexapanopeus Rathbun, 1898 Hexapanopeus schmitti Rathbun, 1930 Figs. 1-4, 5e—h Hexapanopeus schmitti Rathbun, 1930:393, pl. 169, figs. 3-5. Type locality Bay of Rio de Janeiro [22°54’S, 43°14’W], Rio de Janeiro State, Brazil—Coelho & Ra- mos, 1972:191 [listed].—Melo, 1985:105 [Pernambuco, Rio de Janeiro, and Sao Paulo states, Brazil].—Melo et al., 1989: 15 [Parana].—Bakker et al., 1989:137, figs. 1-10 [Parana; larval develop- ment].—Melo, 1996:360 [Brazil]. Material.—Brazil: Rio de Janeiro State, bay of Rio de Janeiro (22°54’S, 43°14’W), leg. W. L. Schmitt, 1925: 1 ¢ (holotype, USNM 59831).—Rio Grande do Norte State, intertidal region of Pitangui and es- tuary of River Potengi (5°47’S, 35°16’W), leg. C. Sankarankutty, 1995: 12 66, 12 2 2 (Museu do Mar “Onofre Lopes”; 1 d, 1 ovigerous 2, USNM 284138). Size.—Males, 3.4 by 4.6 mm (cl by cb) to 5.7 by 7.5 mm; non-ovigerous females, 3.0 by 3.8 mm to 5.7 by 7.8 mm; ovigerous females 4.1 by 5.3 mm to 5.3 by 7.7 mm. The holotype measures 9.4 by 12.8 mm (Rathbun, 1930:394). Color (Fig. 4).—Variable but most live specimens are greenish-grey or pink in col- or; fronto-orbital border and anterolateral margins of carapace often bordered with white; fingers of chelae black except for tips. 250 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fis. 1. showing rim around anterolateral and frontal margins; b, Right chela, outer face; c, Left chela, outer face; d, Abdomen of male; e, Cheliped (P1), dorsal view; f, Third walking leg (P4), posterior face; g, Gonopod. Scale a-f, 1 mm; scale g, 0.5 mm. The majority of the freshly preserved specimens presented a uniform greyish green coloration while it was not uncom- mon to see a large variety of color patterns, some of which are depicted in Fig. 4. The carapace may be totally whitish with a few dark spots as in Fig. 4b; greyish green on most of its surface with a broad whitish band across the frontal and anterolateral regions (Fig. 4c, e). The carapace may also have a greenish background with a range of stripes extending posteriorly (Fig. 4a, d). Hexapanopeus schmitti Rathbun. Composite figure based on several specimens. Pitangui. a, Carapace Chelipeds normally assume the color of the carapace, but in some cases both chelae were whitish or in one rare case only the small chela was whitish. In a majority of cases, the merus, carpus, and the exopod of the third maxilliped had a fine whitish bor- der. Dactyli of all of the walking legs were whitish and in some cases the distal half of the propodus also was whitish. Remarks.—Martin & Abele (1986) sur- veyed gonopods (first male pleopods) of crabs related to Panopeus H. Milne Edwards, VOLUME 110, NUMBER 2 251 Fig. 2. Hexapanopeus schmitti Rathbun. Male, cl 5.0 mm, Pitangui. Dorsal view. Scale = 1 mm. 1834, and they pointed out that the tips of the gonopods of two species of Hexapanopeus Rathbun, 1898, the type species, H. angustif- rons (Benedict & Rathbun, 1891), and H. paulensis Rathbun, 1930, differ from those of Panopeus s.s. The apices of the gonopods of those species of Hexapanopeus do not resem- ble those of Panopeus s.s., as they lack the strongly trilobed apex, with a long, sharp ac- cessory process, a shorter rounded process, and a lateral tooth, often bifid. The apices of the gonopods of the holotype of H. schmitti Rathbun, 1930 (Fig. 5e—h), shown here, are similar in shape to those of H. angustifrons (Fig. 57) but differ in having apical spines. The gonopod of H. paulensis (Fig. 5c, d) dif- fers from that of species of Panopeus s.s. in having a trilobed apex with much smaller lobes. In contrast, the apices of the gonopods of H. beebei Garth, 1961, from the eastern Pa- cific, have a long, tapering lateral tooth, a similarly shaped accessory process, and a rounded median process with three terminal spines (Martin & Abele 1986:fig. 3d). This agrees with the shape of the tip of the gon- opods in the western Atlantic H. caribbaeus (Stimpson 1871) (Fig. 5a, b), the fourth nominal species of Hexapanopeus from Brazilian waters, in which the median pro- cess is oOmamented with more terminal spines. As pointed out by Martin & Abele (1986:185), Hexapanopeus as currently un- derstood contains at least two very distinct 252 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rig. 3: outer face; c, First walking leg (P2); d, Second walking leg (P3); e, Third walking leg (P4); f Fourth walking leg (P5); g, Abdomen. Scales = 1 mm. types of gonopods, that found on H. an- gustifrons and that found on H. caribbaeus. Indeed, as pointed out below, there may be three types of gonopod in different repre- sentatives of the genus. The gonopod of H. paulensis from Brazil resembles that of H. paulensis from North America (see Williams 1984:410, fig. 3310) in shape but differs in ornamentation. In the Brazilian specimens the accessory process Hexapanopeus schmitti Rathbun. Male, cl 5.0 mm, Pitangui. a, Right chela, outer face; b, Left chela, is covered with much larger spines. The gonopod of H. paulensis illustrated in Wil- liams (1965:200, fig. 183E) apparently is based on another, undetermined species; it resembles that of H. schmitti. The gonopod of H. angustifrons (from Williams 1965:200, fig 183D) is shown here (Fig. 57) for comparison with the gon- opods of the other three species of the ge- nus known from Brazil. It appears to rep- VOLUME 110, NUMBER 2 253 Fig. 4. Hexapanopeus schmitti Rathbun. Diagrammatic representation of common color patterns of speci- mens from Rio Grande do Norte. resent a third type of gonopod within the genus, one with poorly developed lobes that also lacks distal spines. We have seen no material of H. angustifrons from Brazil, but it has been recorded from Maceio, Bahia, Rio de Janeiro, and Santa Catarina by Melo (1985), and from Pernambuco by Coelho Filho, Coelho Santos, & Coelho (1994). Melo (1996:357) gave its range in Brazil as from Pernambuco to Santa Catarina. Rathbun (1930:393) commented on the thick, beveled front in H. schmitti. Actually the margin of the front is a continuation of a margin or shelf (Figs. la, 2), 0.1 mm wide, that extends across the front from the base of each fifth anterolateral tooth. The surface of the carapace rises from the inner edge of this shelf. The shelf is evident in H. paulensis but not on A. angustifrons or H. caribbaeus. It is much more prominent in our smaller specimens of H. paulensis than in ones as large as the holotype. 254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. from Brazil. a, b, Hexapanopeus caribbaeus (Stimpson), male, cl 7.0 mm, Santa Catarina, USNM 61803; c, d, Hexapanopeus paulensis Rathbun, male holotype, cl 7.0 mm, Santos, USNM 61136; e, f, Hexapanopeus schmitti Rathbun, male holotype, cl 9.4 mm, Bay of Rio de Janeiro, USNM 59831; g, h, Hexapanopeus schmitti Rathbun, male, cl 5.0 mm, Pitangui; i, Hexapanopeus angustifrons (Benedict & Rathbun, 1891) (from Williams 1965:fig. 183D; not based on specimen from Brazil). Scale = 0.2 mm (a—h only). Hexapanopeus caribbaeus lacks the large basal tooth on the dactylus of the cheliped that is found in H. angustifrons, H. paulen- sis, and H. schmitti. Most species of Hexapanopeus have five well developed anterolateral teeth. The two Species that have the anterior four teeth well developed and the fifth vestigial, H. beebei and H. caribbaeus, also have a long, taper- ing lateral process on the apex of the gon- opod, and in these two characteristics differ from the type species of Hexapanopeus, H. angustifrons. This suggests that these two species should be removed from Hexapan- opeus and placed in a new genus, an action that is beyond the scope of the present pa- per. Rathbun (1930:394) pointed out that “Small specimens are easily mistaken for Panopeus permudensis on account of the similarity of the granulate lines of the car- apace, but they can be identified by the character of the lateral teeth, the thick front, Sternal (a, c, e, g, i) and abdominal views (b, d, f, h) of apex of gonopod of species of Hexapanopeus the texture of the palms, and the extension of the color of the finger in the male.” Key to Brazilian species of Hexapanopeus 1. Anterolateral teeth 5, fifth well devel- oped, lateral. Apex of gonopod blunt, lacking long, tapering lateral process .. — Anterolateral teeth 4, fifth minute, al- most posterolateral. Apex of gonopod with long, tapering lateral process Te Ae AMES heehee" H.. caribbaeus 2. Carpus of cheliped not markedly tuber- culate. Color of fixed finger of chela ex- tending proximally and dorsally onto Palint sao. 5 0: pit). eign. 3 8. eee 3 — Carpus of cheliped with about 15 irreg- ularly placed tubercles. Color of fixed finger of chela barely extending onto palm. [Apex of gonopod trilobate] .-.. LiF center teat is de moter ty agin _ H. paulensis 3. Edge of front thick, beveled. Apex of gonopod a single rounded lobe, with 2— 3rapical spines) 2 At. De H. schmitti VOLUME 110, NUMBER 2 — Edge of front not thick or beveled. Apex of gonopod subtriangular, lacking apical spines H. angustifrons Acknowledgments C. Sankarankutty thanks the Office of Fellowships and Grants, Smithsonian Insti- tution, for supporting his visit to Washing- ton, D.C., to work on this project, and the National Council for the Development of Science and Technology, Brazil (CNPq) for its support. Figures 2-5 were prepared by Lilly King Manning. This is contribution no. 405 from the Smithsonian Marine Sta- tion at Link Port. Support of that program for studies on the systematics of xanthid crabs is gratefully acknowledged. Literature Cited Bakker, C. de, M. Monti, K. Anger, & L. Loureiro Fernandes. 1989. Larval development of Hex- apanopeus schmitti Rathbun, 1930 (Decapoda, Brachyura, Xanthidae) reared in the labora- tory.—Neritica, Pontal do Sul, PR 41/2:137- 164. Benedict, J. E.. & M. J. Rathbun. 1891. The genus Panopeus.—Proceedings of the United States National Museum 14:35-—385, pls. 19-24. Coelho, P. A., & M. de Aratjo Ramos. 1972. A con- stituigao e a distribuicao da fauna de decapodos do litoral leste da América do Sul entre as lat- itudes de 5°N e 39°S.—Trabalhos Oceanogrdafi- cos, Universidade Federal de Pernambuco, Re- cife 13:133-236. Coelho Filho, P. A., Coelho Santos, M. A., & Coelho, P A. 1994. Estudo dos Xanthidae (Crustacea- Decapoda-Brachyura) da Praia de Piedade, Ja- boata—PE.—Revista Nordestina de Zoologia 1(1):125-151, pls. 1-6. 235 Garth, J. S. 1961. Brachygnatha Brachyrhyncha. Non- intertidal brachygnathous crabs from the west coast of tropical America, Part 2. Eastern Pa- cific Expeditions of the New York Zoological Society, XLV.—Zoologica 46(3):133-159, pl. 1. Martin, J. W., & L. G. Abele. 1986. Notes on male pleopod morphology in the brachyuran crab family Panopeidae Ortmann, 1893, sensu Guin- ot (1978) (Decapoda).—Crustaceana 50(2): 182-198. Melo, G. A. S. de. 1985. Taxonomia e padroes distri- bucionais e ecolégicos dos Brachyura (Crusta- cea: Decapoda) do litoral sudeste do Brasil. Un- published Ph.D. Dissertation, Instituto de Bio- ciéncias, Universidade de Sao Paulo, 215 pp., 32 figs., 27 tables. . 1996. Manual de identificagao dos Brachyura (caranguejos e siris) do litoral Brasileiro. Sao Paulo, Pléiade, FAPESP: 604 pp. , V. G. Veloso, & M. C. de Oliveira. 1989. A fauna de Brachyura (Crustacea, Decapoda) do litoral do Estado do Parana. Lista preliminar.— Neritica, Pontal do Sul, PR 41/2:1-31. Milne Edwards, H. 1834. Histoire naturelle des Crus- tacés, comprenant l’anatomie, la physiologie et la classification de ces animaux. Agasse, 1:xxv + 468 pp. Rathbun, M. J. 1898. The Brachyura of the Biological Expedition to the Florida keys and the Bahamas in 1893.—Bulletin from the Laboratories of Natural History of the State University of lowa 4(3):250-294, pls. 1-9. . 1930. The cancroid crabs of America.—Unit- ed States National Museum Bulletin 152:xvi + 609 pp., pls. 1-230. Stimpson, W. 1871. Notes on North American Crus- tacea in the museum of the Smithsonian Insti- tution, No. III.—Annals of the Lyceum of Nat- ural History of New York 10(4-—5):92-136. Williams, A. B. 1965. Marine decapod crustaceans of the Carolinas.—Fishery Bulletin, U.S. 65(1):1- 298. . 1984. Shrimps, lobsters, and crabs of the At- lantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Wash- ington, D.C., xvii + 550 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):256—262. 1997. Eunephrops luckhursti, a new deep-sea lobster from Bermuda (Crustacea: Decapoda: Nephropidae) Raymond B. Manning Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Eunephrops luckhursti, the fourth species of the genus, is de- scribed from material trapped at depths of about 800 m off Bermuda. It resem- bles E. cadenasi Chace, from the Bahamas and Caribbean Sea, in having a median carina on the second to fifth abdominal somites, but differs in having much shorter scaphocerites, more acute pleura on the abdominal somites, and in color in life. The species named below was taken dur- ing exploratory trapping operations in deep water off Bermuda conducted by John P. (Sean) Ingham, Pathfinder Fisheries Ltd., and Brian Luckhurst, of the Division of Fisheries, Bermuda. William McCallan, an- other local fisherman using deep water traps, has been documenting and studying the catches from these traps since 1984. The traps used off Bermuda are described in Luckhurst (1986:209): ““The traps used for this exploratory fishing are Antillian (sic) arrowhead traps of the type presently employed for catching reef fish and spiny lobsters in inshore waters. Several different trap sizes have been tested ranging in size from 3 X 3 X 1.5 feet (91 X 91 X 46 cm) to 8 X 8 X 4 feet (244 X 244 X 122 cm) ... The traps are constructed of 2 inch (5 cm) hexagonal mesh wire, either galvanized or vinyl coated. The frames are made prin- cipally of steel reinforcing rod with some stick supports. Traps are baited with various chopped reef fish ... The traps have been fished over a wide depth range to the west and south of the Bermuda platform.”’ Types are deposited in the National Mu- seum of Natural History, Smithsonian In- stitution (USNM) and the Bermuda Natural History Museum (BNHM). Abbreviations used in the account below include: Al, antennule; A2, antenna; cl, postorbital carapace length; cm, centime- ters; fm, fathoms; m, meters; mm, miulli- meters; MXP3, third maxilliped; P1-5, per- eopods (P1-3 are chelate, Pl is the major cheliped, P4-5 are walking legs); sta, sta- tion; tl, total length, measured from tip of rostrum to posterior margin of telson. All measurements are in millimeters. Family Nephropidae Dana, 1852 Subfamily Nephropinae Dana, 1852 Eunephrops Smith, 1885 Eunephrops luckhursti, new species Figs. 1, 2, 3a, b, 4 Type material.—Holotype: Off Bermuda, 32°14.55'N, 64°47.74'W, depth 450 fm (824 m), leg. S. Ingham, crab trap, 18 Feb 1991: 1 2, cl 51 mm (USNM 284136). Paratypes: Off south shore of Bermuda, depth 820 m, S. Ingham sta 7, 7 Dec 1991: 1 3, cl 57 mm (BNHM 1991-090-012).— Off Bermuda, 1992, no other data: 3 9? 2, cl 39 and 54 mm (USNM 284137) and cl 44 mm (BNHM). Diagnosis.—Carapace lacking submedi- an postcervical spines. A2 peduncle with outer spine at base of scaphocerite. Scapho- cerite minute, extending to or barely over- reaching base of penultimate segment of A2. P2 and P3 with fingers about 4%—% as VOLUME 110, NUMBER 2 long as palm. Abdominal somites 2—6 each with longitudinal median carina dorsally. Description.—Rostrum extending be- yond end of antennular peduncle. Lower margin of rostrum unarmed, appearing smooth, with few, very low, serrations vis- ible only under magnification. Anterior pair of lateral rostral teeth larger than and di- rected more anterolaterally than posterior pair. Postsupraorbital spines smaller than supraorbital, 1 placed behind each supra- orbital, both situated anterior to small post- orbital and small postantennal spines. Su- praorbital ridge absent. Postcervical spines absent. Carapace covered with tubercles, larger anteriorly; surface sparsely pubescent between tubercles. First abdominal somite with shallow, transverse groove. Anterolateral angle of pleuron acute, sharp but not spiniform. Ab- dominal somites 2—6 each with longitudinal median carina dorsally. Longitudinal groove over bases of pleura faint. Pleura of somites 3—6 acute, forming sharp point. Telson longer than wide, lateral margins bulging slightly near midlength, tuberculate there. Posterolateral spines strong. Scaphocerite minute, barely extending to or overreaching base of penultimate seg- ment of A2 peduncle. Apex usually round- ed, rarely acute. Larger spine present on A2 peduncle at base of scaphocerite. MXP3 extending about to end of ros- trum. Merus and carpus each with blunt, lower distal spine, that of merus larger. Major chelipeds (P1) very strong, heavy, subequal, longer than carapace and rostrum combined, extending beyond rostrum with distal part of merus. Chela length about 3 times width; lateral surfaces with broad lon- gitudinal carina formed by tubercles, dis- talmost on inner surface largest, each carina bordered by upper and lower shallow, lon- gitudinal depressions. Upper margin of palm formed by single carina, irregularly tuberculate, lower margin bicarinate to about middle of fixed finger. All carinae or- namented with large and small blunt spines. Dactylus tuberculate dorsally, slightly 257 shorter than palm, shorter than fixed finger. Cutting edge of chela evenly tuberculate or with some larger teeth and smaller, uniform tubercles. Carpus about half as long as che- la (measured dorsally), with 2 rows of spines on outer face, distalmost spines larg- est; dorsal surface with low row of rounded spines; upper, inner margin sparsely spined, distalmost largest of all spines on carpus; lower margin with single row of spines; other spines of various sizes placed on sur- face and distal margin. Merus as long as palm (measured on outer edge), inner and outer edges each with irregular row of spines, each with distal tooth. P2 extending beyond rostrum with chela. Fingers about %4%—% palm length. Carpus twice as long as dactylus. Merus more than twice as long as carpus. P3 extending beyond rostrum with chela and distal part of palm. Fingers about 4—% palm length. Carpus about %4 as long as palm and half as long as merus, latter as long as chela. P4 extending beyond rostrum with dac- tylus. Carpus about twice as long as dac- tylus. Merus twice as long as carpus. PS short but overreaching penultimate segment of A2 peduncle. In both sexes, sternite of P2 ending in narrow, acute, bifid process, sharper in males; sternites of P3 and P4 wider, longer, and rounded. Gonopod as figured. Color (Fig. 4).—Carapace, Al and A2 peduncles, and chelipeds almost uniform red. Al and A2 flagellae pink proximally, beige or clear distally. Ischium and basal part of merus of P2-5 red, carpus and prop- odus clear, extremities pink. Ridges on ab- dominal somites, telson, and uropods red, uropodal setae clear. Measurements.—Male, cl 57 mm (cl plus rostrum 81 mm; tl 170 mm); females, cl 39, 44, 51, and 54 mm (cl plus rostrum 55, 57, 70, and about 63 mm, respectively; tl 11.7, 12.5, 15, ca. 15 cm, respectively). The larg- est specimen, the male, is 170 mm long; other measurements of largest male: P2 258 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Eunephrops luckhursti, new species, female holotype, cl 51 mm. Dorsal view. palm length 24.2 mm, movable finger Distribution.—Known only from Ber- length 6.9 mm; P3 palm length 25.8 mm, muda in depths of 800-824 m. movable finger length 5.9 mm. Etymology.—Named for Brian Luck- VOLUME 110, NUMBER 2 259 Fig. 2. Eunephrops luckhursti, new species, a—f, Female holotype, cl 51 mm; g, Male paratype, cl 57 mm. a, Chela; b, P2; c, P3; d, P4, e, P5; f Tail fan; g, Gonopod. (Setae omitted). hurst, a fishery scientist with the Bermuda Division of Fisheries, whose interest in ex- ploratory deep trapping off Bermuda led to the discovery of this species. Other species taken in these trapping operations off Ber- muda are reported in Luckhurst (1986) and Manning & Holthuis (1986), and Luckhurst & Manning (pers comm). Remarks.—The species of Eunephrops are all known from localities in the north- western Atlantic. Eunephrops luckhursti is the fourth species of Eunephrops to be rec- ognized. It agrees with E. cadenasi Chace, 1939, and differs from both E. bairdii Smith, 1885 and E. manningi Holthuis, 1974, in having a distinct median carina on abdominal somites 2—5. It also differs from E. cadenasi in having much shorter scapho- cerites (Fig. 3b, E. luckhursti; 3d, E. cad- enasi) and in color as well (see below). Eu- nephrops luckhursti is a smoother species than E. cadenasi, with much smaller tuber- cles on the carapace and chelipeds; the spi- nulation of the major chelae is much less pronounced than in E. cadenasi. The pleura of abdominal somites 2—6 are acute and more pointed in E. luckhursti (Fig. 3a) than in E. cadenasi (Fig. 3c). The color of E. luckhursti is quite differ- ent from that of E. cadenasi (see Paulmier 1993:pl. 24; Poupin 1994:pl. 2a,b). In E. cadenasi the chelipeds and abdominal ridg- es are primarily beige or yellow; the cara- pace is marked with yellow anteriorly; and the walking legs are clear or beige proxi- mally, red distally. In E. luckhursti the car- 260 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. dorsal view to show antennal scale (indicated by arrow). a, b, Eunephrops luckhursti, new species, female holotype, cl 51 mm; c, d, E. cadenasi Chace, ovigerous female, cl 66 mm, Caribbean Sea, USNM 170675. apace, chelipeds, and abdominal ridges are red (Fig. 4). The five known specimens of E. luck- hursti are somewhat smaller than those of E. cadenasi, the largest specimen having a total length of 170 mm, a carapace length (including rostrum) of 81 mm. The holo- type of E. cadenasi is 224 mm long (Chace 1939). Holthuis (1974) reported males with carapace lengths (including rostrum) of 46 to 135 mm, with the carapace lengths of females somewhat smaller, 40 and 50 mm. Paulmier (1993) recorded a specimen 490 mm long; that specimen has a carapace length of 110 mm. Poupin (1994) reported Specimens with a carapace length (includ- ing rostrum) of 39 to 116 mm. The smallest female, cl 39 mm, has an extra rostral spine on the right side, both scaphocerites very small and rounded, and both major chelae evenly toothed. The fe- male with cl 44 mm has the rostrum bifid at the apex and the left scaphocerite longer than the right, extending about to the mid- dle of the penultimate segment of the pe- duncle. The largest female, cl 54 mm, has a broken rostrum; the scaphocerite is round- ed on one side, pointed on the other; both a, c, Outline of abdominal pleura 2—5 in lateral view; b, d, Basal segment of antenna in oblique chelae have enlarged teeth as well as evenly spaced smaller ones. The length of the fingers and palm of both P2 of all available specimens of E. luckhursti (n = 5 specimens, 9 legs) and E. cadenasi (n = 8 specimens, 13 legs) were measured. In E. luckhursti the propodus ranged from 3.1 to 3.8 times as long as the dactylus (mean 3.4 times). In EF. cadenasi ranged from 2.9 to 3.9 times as long as the dactylus (mean 3.5 times). These two spe- cies and E. manningi cannot be separated using this feature, but it can be used to dis- tinguish all three of these species from E. bairdii. The depth ranges of E. luckhursti and E. cadenasi are similar, the former having been taken in depths around 800 m, the lat- ter in slightly shallower water. The holotype of E. cadenasi was taken at a depth of 300— 315 fm (549-576 m), the allotype at 250 fm (458 m) (Chace 1939). Holthuis (1974) noted that E. cadenasi had been taken in depths between 373—434 and 591 m. Paul- mier (1993) suggested that its optimal depth was below 450 m, and Poupin (1994) stud- ied one lot taken at 607 m. Specimens of E. cadenasi in the USNM collections were VOLUME 110, NUMBER 2 261 Fig. 4. by Brian Luckhurst. taken in depths of 275 to 340-380 fm (503 to 622-695 m). Material of E. luckhursti was compared directly with the following specimens of E. cadenasi from the Leeward Islands in the USNM collections: USNM 170673, female, cl 65.3 mm, west of St. Martin, depth 360 fm (659 m); USNM 170674, ovigerous fe- male, cl 76.3 mm, northeast of Nevis, depth Eunephrops luckhursti, new species. Female holotype, cl 51 mm. Off Bermuda. Color in life. Photo 317 fm (580 m); USNM 170675, ovigerous female, cl 66.4 mm, northeast of St. Kitts, depth 344 fm (630 m); USNM 170676, male, cl 53.6 mm, north of St. Kitts, depth 365 fm (668 m); USNM 170677, male, cl 78.5 mm, off Dominica, depth 275 fm (503 m); USNM 170678, female, cl 85.9 east of St. Kitts, depth 350-370 fm (641-677 m); USNM 170679, 2 females, cl 57.9 and 89.9 262 mm, north of St. Kitts, depth 340-380 fm (622-695 m). Key to Species of Eunephrops (modified from Holthuis 1974) 1. Carapace with submedian postcervical spines. No spine present on antennal pe- duncle at base of scaphocerite. P2 with fingers slightly less than half length of palm... 95425 2 eee are E. bairdii — Carapace lacking postcervical spines. A spine present on antennal peduncle at base of scaphocerite. P2 with fingers less than % length of palm ............. 2, 2. Abdominal somites 2—5 with single transverse groove, lacking distinct me- dian carina: 22723 ae E. manningi — Abdominal somites 2—5 with distinct median carina 3. Scaphocerite pointed anteriorly, long, extending to base of ultimate segment of antennal peduncle. Pleura of abdominal somites rounded ............ E. cadenasi — Scaphocerite usually rounded anteriorly, short, not extending to middle of penul- timate segment of antennal peduncle. Pleura of abdominal somites pointed E. luckhursti, new species Acknowledgments I thank John P. (Sean) Ingham, William McCallan, Brian Luckhurst, and Lisa Greene, Bermuda Museum of Natural His- tory, for their help in the field and for mak- ing these specimens available for study. Brian Luckhurst provided color photo- graphs of E. luckhursti. The line drawings were prepared by Lilly King Manning. My studies on the systematics of western At- lantic lobsters have been supported by the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Smithsonian Marine Station at Link Port, Florida, a facility of the National Museum of Natural History. This is contribution 418 from that facility. This is contribution No. 8, Bermuda Biodiversity Project (BBP), Bermuda Aquarium, Natural History Mu- seum and Zoo. Literature Cited Chace, FE A., Jr. 1939. Preliminary descriptions of one new genus and seventeen new species of deca- pod and stomatopod Crustacea. Reports on the scientific results of the first Atlantis Expedition to the West Indies, under the joint auspices of the University of Havana and Harvard Univer- sity.—Memorias de la Sociedad Cubana de His- toria Natural 13(1):31-54. Holthuis, L. B. 1974. The lobsters of the superfamily Nephropidea of the Atlantic Ocean.—Biologi- cal Results of the University of Miami Deep- Sea Expeditions, 106.—Bulletin of Marine Sci- ence 24(4):723-884. Luckhurst, B. 1986. Discovery of deep-water crabs (Geryon spp.) at Bermuda—a new potential fishery resource.—Proceedings of the Thirty- Seventh Annual Gulf and Caribbean Fisheries Institute, Cancun, Mexico, November 1994: 209-211. Manning, R. B., & L. B. Holthuis. 1986. Notes on Geryon from Bermuda, with the description of Geryon inghami, new species (Crustacea: De- capoda: Geryonidae).—Proceedings of the Bi- ological Society of Washington 99:366—373. Paulmier, G. 1993. Crustacés profondes capturés aux casiers aux Antilles frangaises—Catalogue de l'Institut Frangais de Recherche pour |’ Exploi- tation de la Mer (FREMER), February 1993: 1-34. Poupin, J. 1994. Faune marine profonde des Antilles Frangaises. Récoltes du Navire Polka faites en 1993. Paris, ORSTOM Editions, Collection Etu- des et Théses, 79 pp., colored pls. 1-5. Smith, S. I. 1885. Description of a new crustacean allied to Homarus and Nephrops.—Proceedings of the United States National Museum 8:167— 170. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):263-—271. 1997. A new crayfish of the genus Orconectes from western Tennessee (Decapoda: Cambaridae) Christopher A. Taylor and Mark H. Sabaj Center for Biodiversity, Illinois Natural History Survey, 607 E. Peabody Drive, Champaign, Illinois 61820, U.S.A. Abstract—A new crayfish, Orconectes pagei, is described from northern and eastern flowing tributaries of the Tennessee River in western Tennessee. The species occurs in small to medium streams with sand substrate. Form I males of O. pagei differ from all other members of the genus Orconectes in being both pigmented and possessing a first pleopod with a short, laterally flattened central projection and a short, dorsoventrally flattened mesial process. Recent re-examination of crayfishes col- lected from western Tennessee in the late 1970’s and early 1980’s housed in the IIli- nois Natural History Survey’s Crustacean Collection, revealed the presence of several unidentifiable form II males and females from two locations in the Big Sandy River drainage. Subsequent field work in that drainage in 1996 resulted in the collection of several form I males of that unidentifia- ble taxon. That species, described herein as Orconectes pageéi, is assigned to the noni- mate subgenus Orconectes. This subgeneric assignment is based on the overall similar- ity in the shape of the form I male pleopod of O. pagei to other members of the sub- genus. O. pagei represents the only epigean member of this subgenus and raises the to- tal number of taxa assigned to it to seven. The remaining members of Orconectes (Or- conectes) are Orconectes (O.) australis aus- tralis (Rhoades, 1941), O. (O.) australis packardi Rhoades, 1944, O. (O.) incomptus Hobbs & Barr, 1972, O. (O.) inermis iner- mis Cope, 1872, O. (O.) inermis testii (Hay, 1891), and O. (O.) pellucidus (Tellkampf, 1844). A subgenus that contains both epi- gean and troglobitic members is not uncom- mon in the family Cambaridae. Other de- scribed subgenera with both ecotypes in- clude Cambarus (Erebicambarus), C. (Jug- icambarus), Procambarus (Austrocamba- rus), and P. (Ortmannicus). Orconectes pagei, new species Figs 1 & 2, Table 1 Diagnosis.—Body and eyes pigmented. Rostrum flat anteriorly, slightly concave posteriorly, terminating in long acumen; median carina absent. Rostral margins thickened, slightly converging distally; ter- minating in spines (see Variation). Areola 25.0-31.5% (X = 28.3, n = 40, SD = 1.4) of total length of carapace. Narrowest part of areola just anterior of midpoint, 5.8—15.0 (X = 9.2, n = 40, SD = 1.9) times as long as wide with 2 to 4 (mode = 3, n = 40, SD = 0.5) punctations across narrowest part. One large cervical spine on each side of carapace. Postorbital ridges well developed, terminating in large spines. Suborbital an- gle weakly developed. Antennal scale broadest slightly proximal to midlength. Is- chia of third pereiopods of form I and form II males with hooks; hooks overreaching basioischial articulation in form I males only. Chela with 3 rows of tubercles along mesial margin of palm; small tufts of setae over mesial margin of palm, fingers, and dorsomesial surface; dorsal surfaces of fin- gers with well defined longitudinal ridges. Ventral surface of chela with tubercle at 264 base of dactyl. First pleopods of form I male symmetrical, extending to bases of third pereiopods when abdomen flexed. First pleopod of form I male without dis- tinct shoulder on cephalic surface at base of central projection; central projection cor- neous, constituting 8.0-11.0% (X = 9.2, n = 4, SD = 1.3) of total length of first pleo- pod, flattened laterally and bladelike, taper- ing rapidly to a sharply pointed tip; mesial process equal in length and corneous, flat- tened dorsoventally and bladelike, tapering to acute tip. Central projection and mesial process of form I first pleopod divergent, forming wide gap between distal tips. An- nulus ventralis immovable, subrhomboidal; cephalic half with median trough, lateral prominences forming anterior margin of fossa; fossa shallow with narrow lateral width; sinuate sinus running from center of fossa to caudal edge. Description of holotypic male, form I.— Body slightly compressed laterally, thorax slightly wider than abdomen (16.3 and 14.8 mm, respectively). Greatest width of cara- pace slightly larger than height at caudo- dorsal margin of cervical groove (16.3 and 14.4 mm respectively). Postorbital carapace length 91.0% of length of carapace. Areola 9.7 times longer (10.7 mm) than wide (1.1 mm) with 2 punctations across narrowest part: length of areola 30.9% of length of carapace. Rostrum with scattered puncta- tions and setae, posterior half slightly ex- cavated; margins slightly converging ante- riorly, fringed with setae and terminating in small rounded tubercles. Acumen terminat- ing in corneous spine reaching nearly to end of antennular peduncle. Postorbital ridges well developed, terminating in corneous spines. Suborbital angle poorly developed. Cervical spine large and corneous; dorsal and branchiostegal areas of carapace punc- tate. Abdomen longer than carapace (40.5 and 34.6 mm, respectively). Cephalic section of telson with 1 movable and 1 immovable Spine in each caudolateral corner. Protopod- ite of uropod with spine extending over en- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dopodite and spine in caudolateral corner extending over exopodite. Caudal margin of cephalic section of exopodite with numer- ous spines (15) and 1 movable spine in cau- dolateral corner. Lateral margin of endo+ podite terminating in spine; endopodite with prominent median ridge terminating in premarginal spine. Dorsal surfaces of telson and uropods setiferous. Antennal scale broadest slightly proximal to midlength; thickened lateral margin ter- minating in large corneous spine. Left an- tennal scale 8.5 mm long, 3.7 mm wide. Mesial surface of palm of left chela with 3 rows of tubercles, 10 tubercles in middle row, 6 in dorsal-most row, and 4 in ventral- most row; dorsal and lateral surfaces of palm covered with many small setiferous punctations. Ventral surface of palm of che- la with corneous spine at base of dactyl. Dorsal and ventral surfaces of finger of propodus with submedian longitudinal ridg- es flanked by setiferous punctations, small tuft of setae at base; basal half of opposable margin with 5 weakly developed tubercles. Dorsal and ventral surfaces of dactyl with submedian longitudinal ridges flanked by setiferous punctations. Fingers with subter- minal corneous tips. Carpus with deep oblique furrow dorsal- ly; mesial surface with 1 tubercle at distal end, large corneous procurved spine just distal to midlength, small corneous spine at midlength; ventral surface with 2 corneous spines at midlength of distal margin. Dor- sodistal surface of merus with 2 large cor- neous spines; ventral surface with 2 large corneous spines just distal to midlength of ventrolateral margin and mesial row of 9 tubercles, some corneous; row terminating in large corneous spine. Ischium with 2 small corneous spines on mesial margin. Hook on ischium of third pereiopod only; hook simple, overreaching basioischial ar- ticulation and not opposed by tubercle on basis. Right fifth pereiopod absent. First pleopod as in Diagnosis, reaching just ce- phalic to bases of third pair of pereiopods when abdomen flexed. VOLUME 110, NUMBER 2 265 Fig. 1. Orconectes pagei, new species: a, Mesial view of first pleopod of form I male; b, Caudal view of first pleopods of form I male; c, Lateral view of first pleopod of form II male; d, Dorsal view of carapace; e, Annulus ventralis; f, Dorsal view of left chela. Figures la, 1b, 1d, and 1f are of holotype (INHS 5785); figure lc is of morphotype (INHS 5777); figure le is of allotype (INHS 5772). 266 Description of allotypic female.—Differ- ing from holotype as follows. Areola con- stituting 28.5% of length of carapace and 8.7 times longer than wide. Margins of ros- trum terminating in corneous spines. Mesial margin of palm of left chela with secondary row of 5 weakly developed tubercles on dorsal surface lateral to primary row of 7 tubercles. Ventral surface of left merus with large spine in distolateral corner, mesial margin with row of 7 spines terminating in large corneous spine. Ventral surface of right merus with 2 spines in distolateral cor- ner, 2 spines at midlength, and a mesial row of 4 weakly developed spines terminating in 2 large corneous spines. Sternum between third and fourth pereio- pods narrowly V-shaped. Postannular scler- ite % as wide as annulus ventralis (de- scribed in Diagnosis). First pleopod unira- mous, barely reaching annulus when abdo- men flexed. Description of morphotypic male, form II.—Differing from holotype as follows. Areola constituting 28.0% of length of car- apace and 14.8 times longer than wide. Margins of rostrum terminating in corneous spines. Mesial margin of palm of left chela with secondary row of 6 weakly developed tubercles on dorsal surface lateral to pri- mary row of 6 tubercles, mesial margin of palm of right chela with secondary row of 6 weakly developed tubercles on dorsal sur- face lateral to primary row of 8 tubercles. Ventral surfaces of meruses with mesial row of 7 tubercles terminating in large cor- neous spines. Hook on ischium of third pereiopod not overreaching basioischial articulation. Left second pereiopod detached. First pleopod of uniform texture; both terminal elements noncorneous and of equal length with rounded distal ends. Size.—The largest specimen examined is the holotype, a 34.6 mm CL form I male. Females (n = 6) range in size from 20.0 to 27.4 mm CL. Form I males (n = 4) range in size from 16.3 to 34.6 mm CL. Form II PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON males (n = 30) range in size from 15.6 to 29.8 mm CL. Color.(Fig. 2).—Dorsal and lateral sur- faces of cephalothorax and abdomen tan to light brown and mottled with dark brown patches of varying size. Dorsal surface of each abdominal segment with 2 dark brown patches that form a pair of parallel bars ex- tending from the posterior edge of the ceph- alothorax to the 5th abdominal segment when abdomen is fully extended. Dorsal surfaces of chela, carpus, and merus tan to light brown in color with dark brown patch- es. Dorsal surfaces of pereiopods with sim- ilar coloration and mottling pattern. Fingers of chelae with red tips. Ventral surfaces of chelae, cephalothorax, and abdomen cream to white. Type locality.—Motrris Creek at Tennes- see Hwy. 424, 0.5 km W jct. W/Tennessee Hwy. 114, 1.6 km NE Yuma, Carroll Coun- ty, Tennessee. Holotype was collected among woody debris in midchannel, 25 m downstream of Hwy. 424 bridge. At the time of collection, Morris Creek ranged in width from 5—8 m with an average depth of 0.4 m. Substrate at the type-locality was sand. Woody debris piles occurred com- monly in the creek both upstream and downstream of the bridge. Disposition of types.—The holotype, al- lotype, and morphotype are in the Illinois Natural History Survey Crustacean Collec- tion (catalogue numbers INHS 5785, INHS 5772, and INHS 5777, respectively), as are the following paratypes: 14 form II males and 1 female (INHS 5764); paratypes con- sisting of 2 form I males, 2 form II males, and 2 females (USNM 130530), and one form I (USNM 284135) are deposited at the National Museum of Natural History, Smithsonian Institution, Washington, D.C. The localities and dates of collection are provided in the following Range and spec- imens examined section. Range and specimens examined.—Or- conectes pagei is confined to streams drain- ing the Cretaceous McNairy Sand and Coon Creek formations which occur as a VOLUME 110, NUMBER 2 Fig. 2. A 25.7 mm carapace length ¢ II Orconectes pagei collected from Hunting Creek, 1.6 km E Bruceton, Carroll Co., Tennessee on 9 May 1996. 268 Table 1.—Measurements (mm) of Orconectes pa- gei, new species. Morpho- Holotype Allotype type Carapace: Total length 28.8 27.4 21.1 Postorbital length 26m 2070 eS Width 1G35e° 13:2 9.6 Height 14.4 13.1 8.9 Areola: Width 1 0.9 0.4 Length 10.7 7.8 Se) Rostrum: Width 523 4.4 323 Length 9.2 TES 6.0 Chela, left: Length, palm mesial margin 10.1 S52 4.4 Palm width 11.0 5.9 4.6 30.5 LGi9e” (F7ED U2 Length, lateral margin Dactyl length Abdomen: Width 14.8 13.5 9.1 Length AQ Sr) -3328- 250 narrow north-south band along the eastern edge of the crest of the Coastal Plain in Henry, Benton, Carroll, and Henderson counties, Tennessee (Fig. 3). The northern- most record for this species is Eagle Creek, a tributary of the Tennessee River in Henry County; the southernmost record being from Middleton Creek, a tributary of White Oak Creek in Henderson County. Orconec- tes pagei is most common in tributaries of the northern flowing Big Sandy River. Its propensity for streams with pure sand sub- strate of the McNairy Sand and Coon Creek formations most likely restricts its distri- bution to headwater streams of the Beech River and White Oak Creek drainages. Downstream portions of both of these drainages, and drainages south of White Oak Creek (e.g. Beason, Snake, Lick, and Chambers creeks), flow through high level alluvial deposits and are characterized by predominantly gravel substrates. In east- ward flowing drainages of the Tennessee River south of White Oak Creek, O. pagei PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON is replaced by Orconectes (Faxonius) wrighti Hobbs, 1948. A total of 81 specimens from 14 locali- ties have been examined from the following Tennessee counties: Benton County: 1) INHS 780, Big Sandy River at TN Hwy. 69, 3.2 km W Big Sandy, 26 Apr 1978 (2 6 II, 1 @); Carroll County: 2) USNM 148879, Hollow Rock Branch at Bruceton, 7 Jun 1978 (1 3 ID; 3) INHS 5764, Morris Creek at TN Hwy. 424, 1.6 km NE Yuma (type locality), 9 May 1996 (14 ¢ II, 1 @, all paratypes); INHS 5785, INHS 5784, 15 Jul 1996 (holotype and 1 6 I, 8 d IL, 3 &,); 4) INHS 5772, INHS 5771, Hunting Creek at Old S.R. 1, 1.6 km E Bruceton, 9 May 1996 (allotype and 4 <6 II,); Henderson County: 5) USNM 130530, Owl Creek at Rt. 20, 1 km E Lexington, 16 Apr 1969 (2 6 I, 2 o Il, 2 &, all paratypes); 6) INHS 5641, Haley Creek at Davis Rd., 8 km E Lexington, 11 Jun 1996 (1 ¢ I, 3 juvenile 3,2 2,3 juvenile 2); 7) INHS 5781, Mid- dleton Creek at TN Hwy. 100, 3.4 km NNE Roby, 11 Jun 1996 (5 6 II, 5 2); Henry County: 8) INHS 5766, McGowen Branch at India Rd., 1.6 km NE Paris, 8 May 1996 (2 3 II); 9) INHS 5777, INHS 5776, Barnes Fork Creek at Reynoldsburg Rd., 7.2 km SSE Paris, 8 May 1996 (morphotype and 2 6 II, 2 2); 10) INHS 5778, USNM 284135, Gin Creek at Copper Springs Rd., 6 km WSW Big Sandy, 16 Apr 1996 (1 d I,1 4 I paratype to UNSM); 11) INHS 778, trib. Barnes Fork Creek 0.5 km downstream TN Hwy. 77, 1.6 km N Van Dyke, 4 May 1981 (1 3 ID; 12) INHS 5634, West Sandy Creek at TN Hwy. 641/69, 4.2 km S Oakwood, 16 Oct 1996 (8 6 I, 4 6 I, 2 juvenile 2); 13) INHS 5626, Holly Fork Creek at TN Hwy. 79, 1.2 km SW Nobles, 17 Oct 1996 (3 3 I, 3 6 ID; 14) INHS 5637, Eagle Creek at TN Hwy. 79, 4 km SW Oak Hill, 17 Oct 1996 (1 6 I). Etymology.—Named in honor of Dr. Lawrence M. Page, Principal Scientist and Curator of Fishes at the Illinois Natural His- tory Survey. Dr. Page has contributed great- ly to our knowledge of midwestern cray- VOLUME 110, NUMBER 2 Figis. fishes and continues to show an intense in- terest in the conservation and systematics of all aquatic organisms. This broad interest has been inherited by many of his co-work- ers and students. Habitat and life-history notes.—During field sampling, all individuals were collect- Known range of Orconectes pagei. Type locality denoted by star. 1 = Mississippi River, 2 = Ten- nessee River, 3 = Big Sandy River. ed from small to medium-sized streams with sand substrate. Within these creeks, the species occurred exclusively in woody debris piles composed primarily of fallen tree limbs and bark in midchannel and along stream banks in areas with slow to moderate current. Orconectes pagei is 270 strongly associated with sand substrate; the species was never collected from streams within its range with even the smallest amount of gravel or cobble substrate. Form I males have been collected in the months of April, July, and October. Field collection efforts from April through July 1996 revealed form I males to be uncom- mon, accounting for only four of the 51 specimens collected. In October, form I males were much more common, account- ing for 12 of the 21 specimens collected. Juveniles were observed in June and Oc- tober. Ovigerous females were collected in the months of April and May. Two oviger- ous females collected on 8 May 1996 mea- sured 18.5 and 14.5 mm CL and carried 101 and 43 eggs, respectively. Eggs were spher- ical and ranged from 1.7 to 1.8 mm in di- ameter. Crayfish associates.—The following spe- cies were collected from habitats containing O. pagei: Cambarus (Depressicambarus) striatus Hay, 1902, Orconectes (Triselles- cens) validus (Faxon, 1914), Procambarus (Ortmannicus) viaeviridis (Faxon, 1914), and P. (O.) acutus (Girard, 1852). Variation.—Size of spines on rostral margins appears to be inversely proportion- al to carapace length. Juveniles and small individuals (ca. <25 mm CL) have large, well developed spines while larger individ- uals, such as the holotype, have margins that terminate in rounded tubercles. Comparisons.—Orconectes pagei differs from all other members of Orconectes in the shape of the form I male gonopod and by being pigmented. The gonopod of O. pa- gei iS unique to pigmented members of the genus in possessing all of the following characteristics: terminal elements short, central projection comprising less than 12% of total length of gonopod; elements diver- gent; central projection laterally flattened, tapering rapidly to acute tip distally; mesial process flattened dorsoventrally and blade- like. Relationships.—In his subgeneric reor- ganization of the genus Orconectes, Fitz- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON patrick (1987) stated that in crayfishes, most external morphological features are difficult to use for inferring intergroup re- lationships because they are readily modi- fied to adapt to environmental conditions. While still variable, Fitzpatrick (1987) sug- gests that structures associated with am- plexus are less susceptible to environmental modification and offer reliable characters for subgeneric classification. Since the form I male gonopod of Orconectes pagei is most similar to those of members of the subgenus Orconectes, we follow Fitz- patrick’s contention and tenatively assign O. pagei to this subgenus. Within the sub- genus Orconectes the form I gonopod of O. pagei is most similar to Orconectes inermis testii, a troglobitic species that occurs in south-central Indiana. Orconectes pagei presents somewhat of a dilemma in that it is pigmented, has developed eyes, possesses hooks on ischia of the third pereiopods only, and occurs in epigean habitats roughly 230 miles south of the known range of O. inermis testii. Future genetic analysis planned for O. pagei and other members of the genus will shed new light on interspe- cific and intersubgeneric relationships with- in Orconectes and possibly determine whether this unique species deserves its own subgeneric status. Acknowledgments We are grateful to J. W. Armbruster, T. J. Near, and J. M. Serb for field assistance. We are also indebted to K. Reed for graciously providing records from and access to the crustacean collections at the National Mu- seum of Natural History, Smithsonian In- stitution. Special thanks to K. S. Cummings for photographic assistance with Fig. 2. Literature Cited Cope, E. D. 1872. On the Wyandotte Cave and its fauna.—American Naturalist 6(7):406—422. Faxon, W. 1914. Notes on the crayfishes in the United States National Museum and the Museum of Comparative Zodlogy with descriptions of new species and subspecies to which is appended a VOLUME 110, NUMBER 2 catalogue of the known species and subspe- cies.—Memoirs of the Museum of Comparative Zodlogy at Harvard College 40(8):351—427. Fitzpatrick, J. FE, Jr. 1987. The subgenera of the craw- fish genus Orconectes (Decapoda: Cambari- dae).—Proceedings of the Biological Society of Washington 100:44—74. Girard, C. 1852. A revision of the North American Astaci, with observations on their habits and geographical distribution.—Proceedings of the Academy of Natural Sciences of Philadelphia 6: 87-91. Hay, W. P. 1891. The crustaceans of Indiana.—Pro- ceedings of the Indiana Academy of Science 1891:147-150. . 1902. Observations on the crustacean fauna of Nickajack Cave, Tennessee, and vicinity.— Proceedings of the United States National Mu- seum 25(1292):417—439. Hobbs, H. H., Jr. 1948. A new crayfish of the genus 241 Orconectes from southern Tennessee (Decapo- da, Astacidae).—Proceedings of the Biological Society of Washington 61(15):85—91. , & T. C. Barr, Jr. 1972. Origins and affinities of the troglobitic crayfishes of North America (Decapoda: Astacidae), II: genus Orconectes.— Smithsonian Contributions to Zoology 105. Rhoades, R. 1941. Notes on some crayfishes from Al- abama caves, with the description of a new spe- cies and a new subspecies.—Proceedings of the United States National Museum 91(3129):141- 148. . 1944. The crayfishes of Kentucky, with notes on variation, distribution and descriptions of new species and subspecies.—American Mid- land Naturalist 31:111—-149. Tellkampf, T. A. 1844. Ausflug nach der Mammuth- hdhle in Kentucky.—Das Ausland. 168:671-— 672; 169:675-676; 170:679-680; 171:683- 684; 172:687—688; 173:691—692; 174:695-696; 175:699-—700. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):272—279. 1997. A new crawfish of the genus Distocambarus, subgenus Fitzcambarus (Crustacea: Decapoda: Cambaridae) from South Carolina J. E Fitzpatrick, Jr. and Arnold G. Eversole (JFF) Department of Biological Sciences, University of South Alabama, Mobile, Alabama 36688, U.S.A., and Tulane Museum of Natural History, Belle Chasse, Louisiana 70037, U.S.A.; (AGE) Department of Aquaculture, Fisheries, and Wildlife, Clemson University, Clemson, South Carolina 29634-0362, U.S.A. Abstract.—A new crawfish, Distocambarus (Fitzcambarus) hunteri, is de- scribed from an apparently limited distribution in the Saluda River basin of South Carolina, U.S.A. The new species is distinguished by a large bladelike central projection and a mesial process that extends about half the length of the central projection; the mesial margin of the palm in males is shorter than the width of that structure and the postannular sternite of females is subequal in length to the annulus ventralis. The closest relative to the new species seems to be D. (F.) youngineri Hobbs & Carlson. In the lengthy paper containing the de- scription of Procambarus (Distocambarus) devexus, later to become the first member of the genus Distocambarus, Hobbs (1981: 308-309) commented on the serendipitous discovery of the initial specimens. Despite 40 years of intense collecting in Georgia, he had never encountered the species and was “reasonably certain’’ that the first ju- veniles collected were members of Cam- barus (Depressicambarus) latimanus (LeConte 1856). Subsequent collecting, particularly by Dr. Paul Carlson in South Carolina, demonstrated sufficient diversity prompting Hobbs & Carlson (1983) to el- evate the subgenus Distocambarus to a ge- neric rank, which Hobbs (1983) then divid- ed into two subgenera. By 1985 when Carl- son had moved from South Carolina and Hobbs’ advancing age had curtailed his field activities, the number of known spe- cies had risen to four (Hobbs 1989; Hobbs & Carlson 1985). All members of the genus are primary burrowers of somewhat limited distributions, which probably explains why so few records existed for a genus that not only displays unexpected diversity but also occurs in two different watersheds (Saluda and Savannah) in Georgia and South Car- olina. In 1994 the second author collected sev- eral burrowing crawfishes as part of a study of the rare crawfish species of South Car- olina. He sent them to the first author for identification, and among them was a fifth member of Distocambarus. In the text that follows, the following ab- breviations are used: TCL, total carapace length; PCL, postorbital carapace length; TU, Tulane University Museum of Natural History, Belle Chasse, LA; USNM, US. National Museum of Natural History, Smithsonian Institution, Washington, D.C.; UADC, University of Alabama Decapod Collections, Tuscaloosa, AL. Distocambarus (Fitzcambarus) hunteri, new species Fig. 1 Diagnosis.—Pigmented; eyes small, fac- eted. Rostrum subspatulate with tiny, bead- like acumen and lacking marginal spines. Carapace rarely with single cervical spine or tubercle on one or both sides, spine usu- VOLUME 110, NUMBER 2 ally absent. Branchiostegal spines small but acute. Suborbital angle small to obtuse. Areola 37.1% to 47.9% (avg. 40.9%) of TCL, 43.5% to 54.1% (avg. 47.8%) of PCL; 10.5 to 14.8 (avg. 13.1) times longer than wide, with 1 or 2 punctations across narrowest part. Chela width less than length of mesial margin of palm, with row of 6 to 9 tubercles, flanked dorsolaterally by sec- ond row of 3 to 7 spiniform to squamiform tubercles; dactyl of Form I male about 1.3 times length of mesial of palm, and chela 68.8% to 74.5% (avg. 69.4%) of TCL; length of carpus greater than inner margin of palm. First pleopod of Form I male with small but distinct shoulder somewhat prox- imal to cephalic base of corneous, platelike, subquadrangular central projection, which projection oriented about 45° distolaterally to main axis of appendage; mesial process tapering from base to acute tip, noncor- neous, extending distally about half dis- tance of central projection; cephalic process only suggested by swelling near cephalo- mesial base of central projection; distal third of appendage inclined caudad; proxi- momesial base of pleopod usually with tiny spur. Annulus ventralis of female with cephal- omedian margin membraneous, allowing hingelike motion; annulus much elevated (ventrally) except for cephalomedian exca- vation; sinus originating near midline in ce- phalic third, then transcribing gentle arc to be lost just before reaching caudal margin. Postannular sclerite subtriangular, and sub- equal in length to annulus; first pleopods represented by tubercles at best. (Measure- ments of type specimens in Table 1.) Description of holotypic male, form I.— Cephalothorax (Fig. 1b, f) subovate, com- pressed laterally; maximum width of cara- pace 1.1 times height at caudodorsal margin of cervical groove; abdomen slightly nar- rower than carapace. Areola 13.1 times lon- ger than wide with only one punctation in narrowest part. Cephalic section of carapace 1.6 times length of areola, latter constitut- ing 38.3% of TCL (46.7% of PCL). Cara- 273 Table 1.—Measurements (mm) of type specimens of Distocambarus (Fitzcambarus) hunteri, new spe- cies. The asterisk (*) indicates an estimation; left bran- chiostegite broken. Holotype Allotype Morphotype Carapace Height 5 es 11.4 abs Width 12.4 12:1 13.07 Entire length 27.4 25:9 vA Postorbital length 22.5 22.0 23.9 Areola Width 0.8 0.8 0.9 Length 10.5 10.4 10.4 Rostrum Width 12 6.1 6.3 Length =e 2.5 =a! Chela Length of mesial margin of palm 12 S.J 7.0 Width of palm 6.9 6.0 6.5 Length of lateral margin 18.3 14.0 16.3 Length of dactyl 9.9 8.6 O11 Carpus of cheliped Width 4.7 5.8 6 Length 8.9 Fe V3 Abdomen Width 10.4 i 10.2 Length 212 228 22.9 pace mostly punctate, except sparse granu- lose tuberculations in extreme cephalolater- al region. Rostrum broadly spatulate, with apex produced into tiny, tuberculate acu- men reaching distal margin of ultimate po- domere of antennular peduncle; rostrum moderately depressed. Subrostral ridge moderately developed and evident in dorsal aspect to near midlength of rostrum. Sub- orbital angle obtuse but evident; branchios- tegal spine small but acute. Fifth abdominal pleuron gently rounded distally with no ev- idence of acute caudo- or cephaloventral angles. Telson with single fixed acute spine in each caudolateral corner of cephalic por- tion. Epistome (Fig. 1k) with cephalic lobe broadly subtriangular, about as wide as long, with small cephalomedian projection; margins of lobe slightly elevated (ventrally) 274 and obtuse submedian longitudinal fovea at caudal union of lobe with main body; ep- istomal zygoma broadly arched. Antennular peduncle lacking spine on ventral surface of proximal podomere; antennal peduncle with spine on ventral surface of basal po- domere reduced to tubercle; tip of flagellum reaching nearly to caudal margin of cara- pace. Antennal scale (Fig. lg) about 1.5 times longer than wide, widest slightly dis- tal to midlength, lateral portion thickened and ending distally in small but stout spine; greatest width of lamellar area 1.8 times width of thickened lateral area. Third maxilliped with palpus, when ex- tended, reaching nearly to distal margin of antennal peduncle; ventromesial surface of ischium with dense mat of long stiff setae occurring in tufts, which mat completely obscuring dentate opposable margin. Right chela (Fig. 17) subelliptical in sec- tion, strongly depressed, palm somewhat narrower than length of mesial margin of palm; length of latter 44.5% of total length of chela; most of palm studded with squa- mous and subsquamous tubercles; mesial margin of palm with row of 8 tubercles, flanked dorsally by less developed row of 5 tubercles. Both fingers with well defined dorsomedian ridges flanked by setiferous punctations. Opposable margin of fixed fin- ger with row of 4 subequal tubercles along proximal third of finger and single row of minute denticles lying between tubercles and reaching to base of corneous tip of fin- ger; additional prominent tubercle ventral to denticle row just distal to midlength of fin- ger; lateral margin weakly costate. Oppos- able margin of dactyl with 2 large tubercles in proximal third, distalmost distinctly larg- er, with single row of minute denticles ex- tending from tubercles to base of corneous tip of finger; mesial margin with 2 weak Squamous tubercles in basal portion, fol- lowed by indistinct row of setiferous punc- tations; setae of ventrolateral punctations of margin not conspicuously long or stiff. Carpus of cheliped 1.1 times length of inner margin of palm; dorsal surface sparse- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ly punctate, with slightly sinuous sulcus; mesial surface with 4 subacute spiniform tubercles in proximal half, flanked dorsally by 3, more rounded, tubercles; ventral mar- gin with stout subspiniform distolateral tu- bercle, and weak squamous tubercles in me- sial corner, lateral one articulating with con- dyle on propodus. Merus with small dorsodistal subspini- form tubercle and tuberculate ventral mar- gins, lateral row of 9 irregular tubercles and mesial row of 12, tubercles of both rows increasing in size distally. Ischium with row of 5 tubercles along mesial margin; suffla- men absent. Chela of second pereiopod with conspic- uous row of long stiff setae on lateral and mesial margins; carpus with similar row on mesial margin, lateral margin with few long setae; merus with only single tuft of about 10 long setae situated on dorsomedian mar- gin. Hook on ischium of third pereiopod (Fig. 127) stout but simple, slightly over- reaching basioischial articulation and not opposed by strong tubercle on basis. Coxae of all pereiopods except first with conspic- uous tufts of plumose setae masking sternal and coxal features; small rounded cau- domesial eminence on fourth periopodal coxa. First pleopods (Figs. la, c, b) symmet- rical, bases not contiguous, tips reaching coxae of third pereiopods when abdomen flexed; prominent caudoproximal and prox- imomesial lobes present; pleopods flexed slightly distal to midlength and lacking sub- apical setae. Terminal elements as in “‘Di- agnosis.”’ Mesial lobe of proximal podomere of uropod bearing acute spine, spine lacking on lateral lobe; mesial ramus with disto- median spine, spine small and not reaching distal margin. | Description of allotypic female.—Except for secondary sexual characters, differing from holotype in following respects: Bran- chiostegal spine more prominent; areola with 2 punctations in narrowest part; tuber- culations of upper surface of chela more VOLUME 110, NUMBER 2 ite h pasate vi ‘ , - oR 23 iA f Fig. 1. Distocambarus (F.) hunteri, new species (all from holotypic male, Form I, except d, e from mor- photypic male, Form II, and j from allotypic female): a, d, Mesial aspect of first pleopod; b, Lateral aspect of carapace; c, e, Lateral aspect of first pleopod; f Dorsal aspect of carapace; g, Antennal scale; h, Caudal aspect of first pleopods; i, Dorsal aspect of distal podomeres of cheliped; j, Annulus ventralis and adjacent sternites; k, Epistome; J, Ventral aspect of proximal podomeres of left third through fifth pereiopods. 275 276 sparse. Sternal sclerites of third and fourth pereiopodal segments broadly excavate and densely setose laterally; setae of coxa and sternite of second segment shorter and less dense. Annulus ventralis (Fig. 1j) broadly ex- cavate cephalically and movable through arc of about 15 to 20°; 1.8 times wider than long; sinus arising in fossa. Postannular sclerite about 40% width of annulus; ce- phalic portion somewhat overreaching ster- nite of antecedent segment. First pleopods absent. Description of morphotypic male, form IT.—Except in secondary sexual characters, differing from holotype in following re- spects: Areola with 2 punctations in nar- rowest part; mesial margin of palm with row of 5 tubercles, flanked dorsally by sec- ond row of 5; both lobes of proximal po- domere of uropod with acute spine, that of lateral lobe tiny; tuberculation of upper sur- face of chela intermediate between holotype and allotype. Hook on ischium of third pereiopod rep- resented by large but low tubercle and not overreaching more proximal segment. First pleopods (Figs. ld, e) with incomplete ju- venile suture delineating basal part; termi- nal elements much more blunt than those of form I male and noncorneous. Color notes.—Ground color varies from lighter brown in recently molted individuals to darker brown in older intermolt ones. Staining from burrow soils and groundwa- ter apparently masks exoskeleton colors. Cephalic sections of carapace more uni- formly brown and less mottled than thorac- ic sections; mottling fading to reddish brown posteriorly in cephalic section and ventrally in thoracic section. Ground color of abdomen lighter than that of carapace; terga with paired, almost black, elongate spots, which become smaller and less ob- vious caudally; pleura with similarly col- ored markings giving appearance of stripe above lighter brown lateral margins; telson and uropods lacking distinct markings, and of lighter color than other abdominal seg- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ments. Distal portion of merus and carpus of cheliped ash brown and bearing dark brown tubercles; dorsal surface of palmar area of propodus paler brown; fingers of chela with reddish brown to red finger tips; movable finger darker of the two. Remain- ing pereiopods bluish to ash brown with more flesh-colored bases. Ventral surface of abdomen and pleopods brown to flesh-col- ored. Specimens held in laboratory and fed ar- tificial formulated diet retain the mottled appearance and abdominal markings. How- ever, ground color of these animals has a distinctly blue hue, blue replaces brown in the chelipeds, and other pereiopods are pale blue. Ventral surfaces of the pereiopods are tan to bluish cream and the ventral abdo- men is a pale blue. These observations are compatible with the suggestions put forth by Fitzpatrick (1987) concerning the effect of diet, and perhaps light, on coloration. Disposition of types.—The holotypic male, Form I, the allotypic female, and the morphotypic male, Form II are in the col- lections of the National Museum of Natural History, USNM (282786, 282787, and 282788, respectively); paratypic series are in the Tulane Museum of Natural History (TU 6746, 1 oI, 1 2, 1 dimm; TU 6747, 2 2, 2 2imm) and the University of Ala- bama (UADC 450.01, 1 dII, 1 2; UADC 451.01, 1 oI, 1 @). Type locality.—Burrows in runoff drain- age just west of State Route 391 at junction of State Route 194; 34°05'26.8’N, 81°34’88.1"W; Saluda River drainage, Sa- luda Co., South Carolina. Here the sandy substrate was wet (water table <15 cm) from recent rains, but numerous chimneys were evident among the thick mat of roots from associated vegetation. Except for one female that was associated with recently re- leased young, each burrow was occupied by a single animal, and the creatures were ac- tively moving in the burrow. The overstory was composed of loblolly pine (Pinus tae- da), red maple (Acer rubrum), willow oak (Quercus phellos), and American elm (UI- VOLUME 110, NUMBER 2 mus americana); the midstory was red ma- ple and dogwood (Cornus florida); and the understory was red oak seedlings (Q. fal- cata), sumac (Rhus glabra), honeysuckle (Lonicera sp.), smilax (Smilax sp.), Amer- ican holly Ulex americana), and grapevine (Vitis sp.). At the type locality, one speci- men of Cambarus (Depressicambarus) la- timanus was collected from a burrow. Range and specimens examined.—This species seems to be confined to a very lim- ited portion of the Saluda drainage in Sa- luda County, South Carolina. This is not wholly surprising. It closest relative, Dis- tocambarus (Fitzcambarus) youngineri Hobbs & Carlson 1985, likewise has a very limited distribution, being confined to New- berry County, just north of the Saluda River (Hobbs & Carlson 1985, Eversole 1995). The other members of the genus all have limited distributions. During the course of this study, the second author conducted ex- tensive searches throughout the upper por- tions of the several drainages associated with the southern part of the state and no other collections of this species were found. Several other burrowing species, including other Distocambarus spp. were recorded, however. Fourteen specimens from the type local- ity have been studied (counts below), plus four juveniles kept alive in the laboratory for collection of maturation data. The sec- ond site produced two adults. Saluda County, South Carolina: (1) type locality, 1 SII, 1 2, 24 Sep 1994, A. G. Eversole, C. D. Baumann, colls.; 1 CII, 1 2, 1 dimm, 2 2imm, 18 Oct 1995, A. G. Eversole et al., colls.; 2 ¢I, 3 2, 1 dimm, 1 Simm, 20 Nov 1995, A. G. Eversole et al., colls.; (2) burrows on W side of Wyses Ferry Rd, about 0.6 mi N of St Rte 194, 1 31, 1 2, 4 Oct 1994, C. J. Kempton and C. D. Baumann, colls. Variations.—The small sample, almost all from the same locality, gave little infor- mation on variation beyond that encom- passed by the descriptions of the primary types. Most conspicuous, as is usual, was 277 the varying numbers and types of tubercu- lation associated with the podomeres of the pereiopods; additional collections from oth- er sites may indicate a wider range of ex- pression of other characters. Size and life history notes.—The largest specimen was a form I male of 29.4 mm TCL (25.6 mm PCL); the smallest mature animal, a female, was 18.0 TCL (14.0 mm PCL). No ovigerous females or females bearing young were collected, although a female collected on 18 October 1995 and another on 20 November 1995 each had re- cently released young in its burrow, the only recorded instances, save one noted be- low, of an adult animal sharing its burrow. Form I males were collected in October and November. Ecological notes.—Distocambarus (Fitz- cambarus) hunteri is a primary burrower and as characteristic of other members of the genus, the burrows extend to the water table. Under some circumstances, vertical shafts of the burrows may reach more than a meter in depth and the main burrow may have an array of side shafts connecting with the surface at capped openings. Many of the burrow shafts were adjacent to tree and shrub roots, making excavation difficult. Freshly capped burrows appeared as if the crawfish individually stacked uniformly made balls (approximately 5 to 6 mm diam) of excavation material into a chimney shaped mound. The chimneys reached 10 mm in height in some cases, but both the architecture and chimney height were sub- ject to weathering. Although weathered chimneys appeared as only a clod of exca- vated soil, their different color was notice- able against the forest floor. The species seems to be a solitary animal because on only one occasion (Oct) was more than one adult (1 oI, 1 2) excavated from a burrow. On two occasions, however, a female was collected from a burrow that contained free- ranging juveniles, eight in one burrow and six in the other. Unfortunately, all of our collections come from September through November, and it is not possible to say that _ 278 recruitment occurs solely in the fall of the year. The exoskeleton on the specimen col- lected on 10 October 1995 appeared soft, a sign that crawfish were rapidly growing and molting at that time of year. The preferred habitat seems to be a mixed pine-hardwood woodland, with a comparatively high water table. Relationships.—This crawfish is most closely related to D. (F.) youngineri, the nearest populations of which occur about 9.5 airmi (15.3 air km) northwest of the sites where D. hunteri is found. The pleo- pods of the Form I males are quite similar, both having a large subquadrangular central projection that is bladelike and oriented somewhat oblique to the cephalocaudal axis of the appendage. In D. hunteri, the central projection is disposed mesially almost 45° to this axis, whereas in D. youngineri it is disposed about 30°. The cephalic process of D. hunteri is scarcely recognizable, but is usually visible, albeit as a vestigial struc- ture, in D. youngineri. The mesial process of the new species extends only half as far distally as does the central projection, while in D. youngineri the process extends at least two-thirds as far. The areola of D. hunteri is wider and, accordingly, frequently has an extra punctation in its narrowest part. In D. hunteri the areola is 10.5 to 14.8 (avg. 13.1) times longer than wide and it constitutes 37.1% to 47.9% (avg. 40.9%) of TCL. The comparable numbers in D. youngineri are 13.0 to 24.0 (avg. 17.6) and 37.6% to 41.9% (avg. 38.9%) of TCL. The bran- chiostegal spine, usually absent or extreme- ly obtuse in D. youngineri, is small but ev- ident in D. hunteri. In Form I males of the latter, the second tubercle of the opposable margin of the dactyl is distinctly larger than the more proximal one, and the mesial mar- gin of the palm is longer than the palm is wide. In D. youngineri, the primary tuber- cles of the opposable margin of the dactyl are subequal in size, and the mesial margin of the palm is shorter than the palm width. The ventral surface of the ischium of the third maxilliped of D. hunteri is provided PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with so dense a mat of long stiff setae that the denticulate opposable margin is com- pletely obscured. In the female, the cephal- omedian margin of the annulus ventralis is less deeply and widely excavate, and a less extensive membranous area permits a movement through an arc of 15° to 20°, as opposed to 30° to 45° in D. youngineri. In the new species, too, the sternite between the fourth pereiopods is broadly excavate, rather than having a fissure-like configura- tion. Females of D. hunteri can also be dis- tinguished by the much longer postannular sclerite that is subequal in length to the an- nulus and is subtriangular in outline. The large, bladelike central projection of the first pleopod of the first form male, coupled with the comparative length of the mesial process, will serve to separate D. hunteri from the other members of the genus. Sev- eral features of the new species (e.g., the ratio of palm width to mesial margin length, areola width, and postannular sclerite of fe- males) may require a redefinition of the subgenus, but we deem it prudent to defer such until more specimens of the several species are available to permit a greater ap- preciation of the limits and extent of vari- ation. Etymology.—This crawfish is named in honor of Dr. W. D. Russell-Hunter, formerly of the Department of Biology, Syracuse University, Syracuse, New York, and the Marine Biology Laboratory, Woods Hole, Massachusetts, for his friendship, support and guidance during the formative years of the second author’s education. We are pleased to propose this taxon honoring Dr. Russell-Hunter for all his contributions to invertebrate zoology. Acknowledgments The authors extend their appreciation to Chris J. Kempton, Christopher D. Baumann and Larry Woodward for their assistance in making field collections. We are also in- debted to John E. Cooper whose critical reading of the manuscript greatly enhanced VOLUME 110, NUMBER 2 its quality. Collection efforts were support- ed by the U.S. Fish and Wildlife Service and the South Carolina Agricultural Exper- iment Station. Literature Cited Eversole, A. G. 1995. Distribution of three rare cray- fish species in South Carolina, USA.—Fresh- water Crayfish 8:113-—120. Fitzpatrick, J. F, Jr. 1987. Notes on the so-called “blue color phase”’ in North American cambar- id crawfishes (Decapoda, Astacoidea).—Crus- taceana 52:316—319. Hobbs, H. H., Jr. 1981. The crayfishes of Georgia.— Smithsonian Contributions to Zoology 318:1- 549. 1983. Distocambarus (Fitzcambarus) carl- soni, a new subgenus and species of crayfish 279 (Decapoda: Cambaridae) from South Caroli- na.—Proceedings of the Biological Society of Washington 96:429-—439. . 1989. An illustrated checklist of the Ameri- can crayfishes (Decapoda: Astacidae, Cambari- dae, and Parastacidae).—Smithsonian Contri- butions to Zoology 480:1—236. , & PH. Carlson. 1983. Distocambarus (De- capoda: Cambaridae) elevated to generic rank, with an account of D. crockeri, new species from South Carolina.—Proceedings of the Bi- ological Society of Washington 96:420—428. , & . 1985. A new member of the ge- nus Distocambarus (Decapoda: Cambaridae) from the Saluda Basin, South Carolina.—Pro- ceedings of the Biological Society of Washing- ton 98:81-89. LeConte, J. 1856. Description of new species of As- tacus from Georgia.—Proceedings of the Acad- emy of Natural Sciences of Philadelphia 7:400-— 402. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):280—284. 1997. Neogonodactylus campi, a new species of stomatopod crustacean from the Caribbean Sea, with additional records for N. caribbaeus (Schotte & Manning) Raymond B. Manning Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Neogonodactylus campi, a sublittoral species, is described from two localities in the Caribbean Sea. It is the second western Atlantic species to be recognized that has both dorsal spinules on the telson and a spined posterolateral angle on the fifth abdominal somite. It differs from N. caribbaeus (Schotte & Manning) in having much shorter anterior submedian carinae on the telson and a longer basal portion on the rostral plate. The first records for N. caribbaeus from Florida and the Bahamas are provided. Recognition of a new species of Gono- dactylus from Tobago (Schotte & Manning 1993) that resembled G. spinulosus Schmitt in having some of the carinae of the telson armed with dorsal spinules prompted me to reexamine all of the material in the crusta- cean collections of the National Museum of Natural History identified as G. spinulosus. The new species described below was found among that material, as were the ad- ditional specimens of Neogonodactylus car- ibbaeus listed below, most of which had been identified as G. spinulosus in Manning (1969). All of the American species formerly re- ferred to Gonodactylus were transferred to the genus Neogonodactylus, type species Gonodactylus oerstedii Hansen, 1895, by Manning (1995). Abbreviations used in the account below include: AWCLI, abdominal width-cara- pace length index (abdominal width divided by carapace length < 100), fm (fathoms), ft (feet), leg. (collector), m (meters), mm (millimeters), n (number), sta (station). All of the specimens are in the crustacean collection of the National Museum of Nat- ural History, Smithsonian Institution, Washington, D.C. (USNM). The number following the number of specimens is total length, measured on the midline. Neogonodactylus campi, new species Figs. 1, 2 Material.—Dominican Republic: Navi- dad Bank, 20°11’'N, 68°52’W, depth 14-15 fm (26-27 m), 8 ft tumbler dredge, M/V Oregon sta 5474, 12 Jun 1965: 1 2, 36 mm (holotype, USNM 126013).—Off Navidad Bank, 20°04’'N, 68°53'W, depth 20-24 fm (37—44 m), 8 ft tumbler dredge, M/V Ore- gon sta 5471, 12 Jun 1965: 1 2, 28 mm (paratype, USNM 126014). St. Lucia: West of Pointe du Cap, 14°06'N, 61°05’W, depth 16 fm (29 m), tumbler dredge, M/V Oregon sta 5946, 8 Mar 1966: 1 3, 26 mm (paratype, USNM 126034). Diagnosis.—Size relatively small, total length of adults less than 40 mm. Rostral plate slightly longer than broad, basal part obtusely rounded anterolaterally, anterior margins sloping to slender median spine, latter more than 1.5 times as long as basal part of plate. Ocular scales relatively broad, flattened, separate. Thoracic so- mites lacking dark pigment dorsally. Ab- VOLUME 110, NUMBER 2 281 Fig. 1. Neogonodactylus campi, new species, female holotype, 36 mm. a, Rostral plate and ocular scales; b, Sixth abdominal somite, telson and right uropod, dorsal view (distalmost uropod spine hidden by distal segment of exopod); c, Fifth abdominal somite, lateral view. Scale = 1 mm (a), 2 mm (3b, c). domen lacking distinctive dark pigment dorsally; anterior 4 somites unarmed pos- terolaterally, fifth somite with sharp pos- terolateral spines; sixth somite with 6 carinae, each with sharp posterior spine; AWCLI of male 852, of females 857. Tel- son of oerstedii-type, with dorsal tuber- cles on carinae, latter sharp, intermediates cristate in female; anterior tubercles of telson each produced into erect spinule; median carina ending in sharp spine; ac- cessory median carinae forming anchor posteriorly, extending anteriorly about % length of median carina, with 1—3 dorsal and no posterior tubercles; knob evenly rounded, with 4—5 tubercles; anterior sub- median carinae short, not extending pos- teriorly beyond bases of accessory medi- ans, with terminal spine followed poste- riorly by single sharp, erect tubercle; sub- median marginal teeth slender, movable apices present, with 3—7 sharp dorsal tu- bercles, inner margin lined with spiniform denticles; carina of intermediate tooth cristate, usually unarmed (with 5—6 spi- nules in 1 specimen only); accessory in- termediate carinae cristate, with 1-5 spines dorsally, 1 usually terminal; inner intermediate denticle with short dorsal ca- rina, spined posteriorly, both intermediate denticles with sharp apical spinule; lateral teeth sharp, unarmed dorsally. Uropod with 9-11 graded, movable spines later- ally. Size.—Total lengths of male (n = 1), 26 mm; of females (n = 2), 28 and 36 mm. Other measurements of female holotype, to- tal length 36 mm: carapace length 7.7 mm; rostral plate length 2.8 mm, width 2.5 mm; fifth abdominal somite width 6.6 mm; tel- son length 5.5 mm, width 5.3 mm. Remarks.—Neogonodactylus campi is the second species to be recognized from localities in the western Atlantic in which: (a) the fifth abdominal somite is armed pos- terolaterally; (b) some dorsal carinae of the telson are armed with sharp spinules or tu- bercles; and (c) there are no patches of dark pigment on the sixth thoracic somite and abdomen. In these features it resembles WN. 282 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Neogonodactylus campi, new species, female paratype, 28 mm. Telson, dorsal view. Scale = 2 mm. caribbaeus (Schotte & Manning, 1993) and differs from N. spinulosus (Schmitt, 1924). It differs from N. caribbaeus in having shorter anterior submedian carinae on the telson armed with a posterior spine, fol- lowed posteriorly by a sharp tubercle, as well as a longer basal part of the rostral plate. Neogonodactylus campi differs from N. minutus (Manning, 1969), the only other Species in the western Atlantic with spi- nules or tubercles on the dorsal carinae of the telson, in having many more dorsal tu- bercles on those carinae. Variation in size and distribution of dor- sal tubercles on the telson of two different females is shown in Figs. 1b and 2. Etymology.—Named for my colleague and friend David K. Camp, Florida Depart- VOLUME 110, NUMBER 2 283 Picts: ea See Neogonodactylus caribbaeus (Schotte & Manning), Florida, female, 36 mm. a, Rostral plate; b, ocular scales; c, sixth abdominal somite, telson, and uropod; d, posterolateral angle of fifth abdominal somite, sixth abdominal somite, and base of uropod, lateral view. Scale = 1 mm (a, b), 2 mm (c, d). ment of Environmental Protection, Florida Marine Research Institute, St. Petersburg, Florida. David has added much to our knowledge of the diverse and important Marine invertebrate fauna of Florida through his own research on stomatopods and other groups, and through his efforts to have the extensive Hourglass collections of invertebrates studied by others and pub- lished by the Institute under his guidance. Recognition of his contributions to our knowledge of systematics of marine crus- taceans is long overdue. Neogonodactylus caribbaeus (Schotte & Manning, 1993) Fig. 3 Gonodactylus spinulosus.—Manning, 1969: 299 [part, all specimens listed below ex- cept for those from the Bahamas; not G. spinulosus Schmitt, 1924]. Gonodactylus caribbaeus Schotte & Man- ning, 1993:568, fig. 1. Material.—Florida: Monroe County, % mile (ca 400 m) south-southwest of Alli- gator Reef Light [24°51'N, 80°38'W], depth 284 15 ft (about 5 m), leg. W. A. Starck, I, 26 Aug 1961: 1 2, 36 mm (USNM 124639).— 200 yards (ca 183 m) southwest of Alligator Reef Light, depth 15—20 ft (4.6—6 m), leg W. A. Starck, II, 7 Jan 1962: 1 6, 25 mm (USNM 119320). Bahama Islands: Providence Channel, off Abaco Island, 25°50'N, 77°10'W, depth 12 fm (22 m), R/V Silver Bay sta 5133, 1 Oct 1963: 1 3d, 30 mm, 1 2, 21 mm (USNM 126033). U.S. Virgin Islands: St. John, Coral Har- bor, base of Coral Bay (18°21'N, 64°43’W), leg. L. P- Thomas, 20 Dec 1958: 1 6, 25 mm (USNM 124636). Barbuda: Martello Tower (13°53’N, 60°53’W), reefs off south coast, Smithson- ian-Bredin Expedition sta 92-56, leg. D. V. Nicholson, 7 Apr 1956: 2 22, 17 and 20 mm (USNM 124633). Mexico: Quintana Roo, Ascension Bay (19°40'N, 87°30'W), behind central part of Nicchehabin Reef, Smithsonian-Bredin Ex- pedition sta 67—60, 13 Apr 1960: 1 juvenile 2, 14 mm (USNM 124634).—Same local- ity, depth 4—6 ft (1-2 m), Smithsonian-Bre- din Expedition sta 72-60, 14 Apr 1960: 1 juvenile 2, 12 mm (USNM 124635). Size.—Total lengths of males (n = 3), 25—30 mm; of females (n = 6), 12—36 mm. The specimens studied by Schotte & Man- ning (1993) were up to 33 mm long. Remarks.—The material reported here extends the range of this species from To- bago in the southern Caribbean to other lo- calities in the Caribbean, and provide the first records for the species from Florida, the Bahamas, the U.S. Virgin Islands, Bar- buda, and Mexico. In addition to differing from N. campi in having longer anterior submedian carinae PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON on the telson, extending posteriorly beyond the bases of the accessory median carinae and less produced posterior spines on the anterior submedian carinae, N. caribbaeus has a rostral plate with a markedly different Shape. In N. caribbaeus (Fig. 3a) the basal part of the plate is quite short and broadly rounded laterally, whereas in N. campi (Fig. la) the basal part of the plate is longer and has a distinct angle anterolaterally. Acknowledgments Studies on the systematics of stomato- pods are supported by the Smithsonian Ma- rine Station at Link Port, Florida, a facility of the National Museum of Natural History, Smithsonian Institution, Washington, D.C.. This is contribution no. 410 from that sta- tion. The figures were prepared by my wife Lilly. The manuscript was improved mate- rially by the careful reading of two anony- mous reviewers. Literature Cited Hansen, H. J. 1895. Isopoden, Cumaceen und Sto- matopoden der Plankton-Expedition.—Ergeb- nisse der Plankton-Expedition der Humboldt- Stiftung 2(Gc):1—105, pls. 1-8. Manning, R. B. 1969. Stomatopod Crustacea of the western Atlantic.—Studies in Tropical Ocean- ography, Miami 8:380 pp. . 1995. Stomatopod Crustacea of Vietnam: The legacy of Raoul Seréne.—Crustacean Re- search, Carcinological Society of Japan, Special Number 4:339 pp., 38 colored plates. 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. Schotte, M., & R. B. Manning. 1993. Stomatopod Crustacea from Tobago, West Indies.—Proceed- ings of the Biological Society of Washington 106:566—581. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):285-—300. 1997. A new subgenus and four new species of Gliricola (Phthiraptera: Gyropidae) from Caribbean hutias (Rodentia: Capromyidae) Roger D. Price and Robert M. Timm (RDP) Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108, U.S.A. (Current address) 4622 Kinkead Ave., Fort Smith, Arkansas 72903-1947, U.S.A.; (RMT) Natural History Museum and Department of Systematics & Ecology, University of Kansas, Lawrence, Kansas 66045-2454, U.S.A. (direct reprint requests to RMT). Abstract.—A new subgenus, Hutiaphilus (Phthiraptera: Gyropidae), is described for five previously named species of Gliricola (G. armatus, G. capromydis, G. cubanus, G. ewingi, and G. omahonyi) and four new species (G. rabbi, with the type host Geocapromys ingrahami; and G. pinei, G. schwartzi, and G. wernecKi, all with the type host Mysateles melanurus melanurus). We redescribe and illus- trate the previously described species, and provide a key for the identification of these nine species. The nine species of Hutiaphilus are restricted to the caviomorph rodent family Capromyidae, the West Indian hutias. This chewing louse-host as- sociation is parallel to other louse-host associations we have documented for ca- viomorph rodents in that there are two (and in one case, three) species of lice on each host species and typically two even on single host individuals. Hutiaphilus is a derived clade well supported by several synapomorphic features. Its position within the genus Gliricola suggests that the family Capromyidae may be nested within what is now recognized as the Neotropical family Echimyidae. Resumen.—Se describe Hutiaphilus, un nuevo subgénero de Gliricola (Phthir- aptera: Gyropidae). Se incluyen en Hutiaphilus cinco especies de Gliricola ya descritas (G. armatus, G. capromydis, G. cubanis, G. ewingi y G. omahonyi) y quatro especies nuevas (G. rabbi, con el hospedero tipico Geocapromys ingra- hami; y G. pinei, G. schwartzi y G. wernecki, las tres con el hospedador tipico Mysateles melanurus melanurus). Se redescriben las especies anteriormente des- critas, incluyendo ilustraciones. Ademas se presenta una clave de identificacién de las nueve especies, que se restringen a la familia de roedores caviomorfos Cap- romyidae, las jutias antillanas. Las relaciones piojo/hospededor de estas especies se aproximan a las que hemos documentado de otros roedores caviomorfos, por tener dos (y en un caso hasta tes) especies de piojos en cada especie hospededor, incluso en un solo individuo hospededor. Hutiaphilus es un clado derivido bien apoyado por varios caracteres sinapomorficos. Su posicién dentro del género Glir- icola sugiere que la familia Capromyidae posiblemente se encuentre filogenéti- camente dentro de lo que actualmente se reconoce como la familia neotropical Echimyidae. The chewing louse genus Gliricola rodent families Capromyidae, Caviidae, and Mjéberg (Phthiraptera: Gyropidae) is re- Echimyidae—caviomorph rodents of the stricted in its distribution to the New World Caribbean islands and Central and South | 286 America. Thirty-five species of Gliricola are now recognized (Price & Timm 1993). Emerson & Price (1975) and Price & Timm (1993) provide species descriptions, illus- trations, brief reviews of species, host dis- tributions, and literature citations for a number of these taxa. Of the 35 species of Gliricola, 5 are from 3 species of West In- dian hutias (Rodentia: Capromyidae). Wer- neck (1944) described Gliricola capromy- dis and G. cubanus from the Cuban hutia, Capromys pilorides (Say). In the same pa- per, G. ewingi and a second subspecies of G. capromydis, G. c. armatus [herein con- sidered a full species], were described from a second species of Cuban hutia, Capromys prehensilis [now Mysateles prehensilis (Poeppig)]. Werneck (1951) subsequently described the fifth and final species, G. omahonyi from the Bahamian hutia, Geo- capromys ingrahami (J. A. Allen). The hutias are a Caribbean radiation of mid-sized, terrestrial caviomorph rodents that occupied many of the larger and small- er islands of the Greater and Lesser Antilles and the Bahamas in the West Indies. The capromyids are a complex and poorly un- derstood radiation of rodents that are in need of revision. Modern authors recognize from 20 to 40 Recent species in the family, grouped in 6 to 8 genera in 4 subfamilies (Hall 1981, Nowak 1991, Woods 1989, 1993). The majority of these species are now extinct, with many of the extinctions having taken place since the arrival of Eur- opeans on these islands in the 16th century (Woods 1989). We recently obtained a number of Gliri- cola lice from specimens of hutias collected in the 1950’s by the late Albert Schwartz and his colleagues. These hutia specimens have only recently become available for study. This collection is extremely signifi- cant in that it contains a host, Mysateles me- lanurus (Poey), from which lice were un- known previously, and it provides addition- al collections of Gliricola from poorly rep- resented hosts. This prompted us to re-examine as many specimens of Gliricola PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from capromyids that we could locate in collections and to collect additional mate- rial from specimens of hutias in the collec- tions of the Field Museum and University of Kansas Natural History Museum. The purposes of this paper are to: de- scribe a new subgenus for the nine species of Gliricola we now recognize from West Indian capromyids; redescribe the five pre- viously named species; describe four new species of Gliricola; provide a key for the identification of the nine species in the new subgenus; and discuss the host relationships and geographic distribution of the louse ge- nus Gliricola on capromyids. In the following descriptions, all mea- surements are in millimeters. The scientific names of the hosts follow Hall (1981), with updates by Woods (1993). The deposition of the holotypes is given in each new spe- cies description; as numbers allow, para- types will be placed in the collections of the National Museum of Natural History (Washington, DC), the Field Museum (Chi- cago, IL), University of Minnesota (St. Paul, MN), and Oklahoma State University (Stillwater, OK). Acronyms designating museum collections where the hosts are de- posited are: FMNH = Field Museum of Natural History; KU = University of Kan- sas Natural History Museum (Lawrence, KS); USNM = National Museum of Nat- ural History. Genus Gliricola Mjdberg Micropus Denny, 1842:247 (nec Meyer & Wolf 1810:280). Type species: Gyropus gracilis Nitzsch. Gliricola Mjéberg, 1910:292. Type species: Gyropus gracilis Nitzsch. Paragliricola Ewing, 1924:29. Type spe- cies: Paragliricola guadrisetosa Ewing. Clay (1970), in discussing supraspecific relationships within the suborder Ambly- cera, placed Gliricola as the principal com- ponent of the subfamily Gliricolinae (fam- ily Gyropidae). The only other members of this subfamily are two species of Mono- VOLUME 110, NUMBER 2 thoracius Werneck described, respectively, from Myoprocta acouchy (Erxleben) and Kerodon rupestris (Wied) from Brazil and one species of Pitrufguenia Marelli from Myocastor coypus (Molina) from Chile (Werneck 1948). Werneck (1948) and Clay (1970) discuss these louse taxa and the characters distinguishing them. Morphologically, members of Gliricola may be characterized by the following com- bination of features: grossly much as in Fig. 29, except for terminalia; antennae clubbed, mostly concealed beneath head, with third segment pedunculate; with two-segmented maxillary palps; without ventral spinous head processes; with only five pairs of ab- dominal spiracles, those on segment VIII absent; with single thin tarsal claw on each leg; prothorax distinctly separated from fused meso/metathorax; and little sexual di- morphism except that associated with ter- minalia and dimensions. Hutiaphilus, new subgenus Type species.—Gliricola capromydis Werneck Description.—The features that unique- ly characterize species of Hutiaphilus are the male dorsal terminalia with a conspic- uous bilobed plate (Fig. 4), and the male genitalia with variably complicated me- sosomal features and large parameres each bearing two conspicuous subapical setae (Fig. 5). This separation is rein- forced by the geographical distribution of the hosts, all of which are hutia species on the West Indies islands. The remaining 30 Gliricola species, all herein considered to be in the nominate subgenus Gliricola, are found on the main- land of Central and South America, on a considerably different array of hosts. No males of these louse taxa have any sugges- tion of the bilobed plate on the dorsal ter- minalia. Their male genitalia are quite dif- ferent, with the single exception of those of G. mirandai described by Werneck (1935) from Isothrix bistriatus Wagner in Bolivia. 287 This genitalic similarity led Werneck (1951) to include his five taxa from hutias with G. mirandai as forming a well-defined group within Gliricola. We have studied specimens of G. mirandai and have con- cluded, in spite of this genitalic similarity, that this species is a member of the nomi- nate subgenus. Both sexes of G. mirandai have a markedly different sternum II and sternal and pleural chaetotaxy; the male lacks the bilobed dorsal plate on the last segment; and the female terminalia are of a grossly different configuration. Etymology.—The new subgenus pro- posed herein, Hutiaphilus, derives its name from the common name of the hosts, hutias, in combination with the substantive suffix phila, derivable from philos (dtdos), “‘lov- ing.’’ The gender is masculine. In the following species descriptions, setal counts are provided only for selected abdominal terga and sterna, as the com- promised quality of many of the available specimens and the large number of setae involved made quantification difficult; the counts are given only to reflect the overall pattern of abundance. The principal fea- tures for separation of males involve de- tails of the genitalia and the dorsal and ventral terminalia; for females, the chae- totaxy associated with the terminalia, es- pecially that of the subgenital plate and anus; and for both sexes, the shape and chaetotaxy of sternum II, the size of spir- acles, and dimensions. Abbreviations for dimensions are: POW, preocular width; TW, temple width; HL, head length; PW, prothorax width; MW, metathorax width; AWIV, abdomen width at segment IV; TL, total length; GW, male genitalia width at paramere base; GPL, male genitalia paramere length; GL, male genitalia length measured from anterior end of outer demarcated portion of basal apo- deme to tip of paramere (see Fig. 5). Brack- eted information under material examined are additions that we have made to the orig- inal data. For split drawings with a median 288 vertical line, dorsal is to the left and ventral to the right. Gliricola (Hutiaphilus) capromydis Werneck Figs. 1—5 Gliricola capromydis Werneck, 1944:394. Type host: Capromys pilorides (Say). Male.—Sternum II shaped as in Fig. 2, constricted and weakly pigmented medially. Setae on tergum IV, 45-73. Setae on ster- num II, 4—7; IV, 10-15; V, 14-20; VII, 22— 35. Spiracle diameter, 0.010—0.015. Dorsal terminalia as in Fig. 4; dorsal sclerite with prominent widely spaced lobes; short spi- niform setae on each lobe, other setae fine. Ventral terminalia as in Fig. 3, with termi- nal portion bearing scattered small projec- tions. Genitalia as in Fig. 5; prominent api- cally pointed triangular mesosome. Dimen- sions: POW, 0.18—0.20; TW, 0.23-0.26; HL, 0.23—0.27; PW, 0.17—0.19; MW, 0.22— 0.24; AWIV, 0.35-0.40; TL, 0.96—1.12; GW, 0.13—0.16; GPL, 0.15—0.17; GL, 0.31- 8 sj Female.—Sternum II as for male. Setae on tergum V, 55. Setae on sternum II, 7; IV, 19; V, 25. Spiracle diameter, 0.020—0.025. Terminalia as in Fig. 1; dorsal anterior setae distributed across segments; subgenital plate with total of 32 setae, none of these very long; setae around anus as shown. Di- mensions: POW, 0.21; TW, 0.28; HL, 0.28; PW, 0.20; MW, 0.28; AWIV, 0.52; TL, 1.42. Type material.—Holotype male, allotype female, ex Capromys pilorides: Cuba: [Pi- nar del Rio], San Diego de los Bafios, coll. W. M. Mann #2202; in collection of the Na- tional Museum of Natural History (Wash- ington, DC). Other material examined.—3 males, ex Capromys pilorides [pilorides]: Cuba: Las Tunas, 29 km W of Victoria de las Tunas, Samalloa Farm, 15 Jun 1952, coll. E. T. Willis, Albert Schwartz #2327 2 (KU 147702); 1 male, same data, except Albert Schwartz #2328 2° (KU 147703); 1 male, same data, except Albert Schwartz #2329 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2 (KU 147704). 1 male, ex Capromys pi- lorides [pilorides|: Cuba: [Camagiiey], Ca- magtiey, 10 Mar 1913, coll. C. Sheldon #10154X% 36 (USNM 181232). Remarks.—The male of this species is readily distinguished from all others of the subgenus by the details of the genitalia, es- pecially the shape of the mesosome, in con- junction with the type of dorsal terminal plate, the sculpturing of the ventral terminal segment, and the structure of sternum II. The female is recognizable by the chaeto- taxy of the terminalia and the chaetotaxy and shape of sternum II. Gliricola (Hutiaphilus) cubanus Werneck Figs. 6—9 Gliricola cubanus Werneck, 1944:397. Type host: Capromys pilorides (Say). Male.—Sternum II shaped as in Fig. 6, of fairly equal width throughout. Setae on tergum IV, 57—75. Setae on sternum II, 7— 8, much shorter medially; IV, 16—24; V, 25— 31; VII, 32—40. Spiracle diameter, 0.010— 0.015. Dorsal terminalia near to Fig. 18; dorsal sclerite with prominent close-set lobes; short spiniform setae on each lobe, other setae fine. Ventral terminal segment as in Fig. 7, with terminal portion bearing weak transverse lines. Genitalia as in Fig. 9; with complex mesosomal structures, broad and rounded apically, with prominent elongate often convoluted sclerite anterior to these. Dimensions: POW, 0.17—0.19; TW, 0.23—0.25; HL, 0.22—0.24; PW, 0.17—0.19; MW, 0.20-0.22; AWIV, 0.36—0.40; TL, 0.96—1.10; GW, 0.15-—0.18; GPL, 0.15- 0.16; GL, 0.29-0.31. Female.—Sternum II as for male. Setae on tergum V, 44—48. Setae on sternum II, 7-9; IV, 19-21; V, 25-26. Spiracle diame- ter, 0.013. Terminalia as in Fig. 8; dorsal anterior setae distributed across segments; subgenital plate with total of 27—32 short, 5—6 very long setae; broad setae around anus, as shown. Dimensions: POW, 0.19— 0.20; TW, 0.24—0.26; HL, 0.23—0.25; PW, VOLUME 110, NUMBER 2 289 Figs. 1-12. 1-5, Gliricola capromydis: (1) female terminalia; (2) sternum II; (3) male ventral terminalia; (4) male dorsal terminalia; (5) male genitalia (GL = genitalia length). 6-9, Gliricola cubanus: (6) sternum II; (7) male ventral terminalia; (8) female terminalia; (9) male genitalia. 10-11, Gliricola ewingi: (10) female terminalia; (11) male genitalia. 12, Gliricola armatus: male genitalic mesosome. 0.19-0.20; MW, 0.23-0.25; AWIV, 0.44— Sheldon #10154X ¢ (USNM 181232); in 0.49; TL, 1.18—1.22. collection of University of California Type material.—Paratype male, ex Cap- (Berkeley). romys pilorides [pilorides]: Cuba: [Cama- Other material examined.—1 male, 1 fe- gtiey], Camagiiey, 10 Mar 1913, coll. C. male, ex Capromys pilorides [pilorides): 290 Cuba: Las Tunas, 29 km W of Victoria de las Tunas, Samalloa Farm, 15 Jun 1952, coll. E. T. Willis, Albert Schwartz #2327 2 (KU 147702); 4 males, 1 female, same data, except Albert Schwartz #2328 2 (KU 147703); 4 males, 2 females, same data, ex- cept Albert Schwartz #2329 2 (KU 147704). Remarks.—There are considerable differ- ences between this taxon and Gliricola cap- romydis, which co-occurs on individuals of Capromys pilorides. The male mesosomal sclerites are profoundly different, as well as the shape of sternum II, the close-set lobes of the dorsal terminal plate, and the sculp- turing of the ventral male terminalia; the female, with 2—3 very long setae on each side of the subgenital plate, is easily sepa- rated from G. capromydis. Gliricola (Hutiaphilus) ewingi Werneck Figs. 10, 11 Gliricola ewingi Werneck, 1944:399. Type host: Capromys [=Mysateles] prehensilis [prehensilis| (Poeppig). Male.—Sternum II as in Fig. 2, constrict- ed and weakly pigmented medially. Setae on tergum IV, 83-86. Setae on sternum II, 7; IV, 22-24; V, 27; VII, 35-36. Spiracle diameter, 0.015. Dorsal terminalia near to Fig. 18; dorsal sclerite with prominent close-set lobes; short spiniform setae on each lobe, other setae fine. Ventral termin- alia as in Fig. 7, with terminal portion bear- ing weak transverse lines. Genitalia as in Fig. 11; complex of mesosomal structures as shown, broadly rounded, with H-shaped anterior sclerite. Dimensions: POW, 0.17— 0.18; TW, 0.23—0.24; HL, 0.22; PW, 0.19-— 0.20; MW, 0.21—0.23; AWIV, 0.43-0.44; TL, 1.14; GW, 0.20; GPL, 0.17—0.19; GL, 0.38—0.41. Female.—Sternum II as for male. Setae on tergum V, 42—45. Setae on sternum II, 8; IV, 17-18; V, 20—23. Spiracle diameter, 0.018—0.025. Terminalia as in Fig. 10; dor- sal anterior setae distributed across seg- ments; subgenital plate with total of 28-32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON setae, none of these very long; several un- interrupted transverse rows of projections across base of subgenital plate; setae around anus as shown. Dimensions: POW, 0.17— 0.19; TW, 0.23-0.25; HL, 0.22—0.24; PW, 0.20; MW, 0.23; AWIV, 0.44; TL, 1.21. Type material.—Paratypes 2 males, 1 fe- male, ex Mysateles prehensilis [prehensi- lis]: Cuba: [Pinar del Rio], San Diego de los Banios, 8 Apr 1900, Palmer and Riley #137 6 (USNM 103887); in collection of University of California (Berkeley). Other material examined.—1 male, ex Mysateles prehensilis [prehensilis]: Cuba: Pinar del Rio, 4 December 1940, coll. I. Perez Viqueras. 1 female, ex M. p. prehen- silis: Cuba: Havana, Feb 1886, coll. C. B. Cory (FMNH 15021). Remarks.—The male genitalia of Gliri- cola ewingi are unique in possessing the H-shaped anterior mesosomal sclerite and in being larger in all dimensions than for any of the other species of the subgenus. The female is separable on the basis of the structure and chaetotaxy of its terminalia; the rows of projections at the base of the subgenital plate are much as those of the first new species described below, but with much smaller spiracles and a different shape of sternum II. Gliricola (Hutiaphilus) armatus Werneck Figs. 12—15 Gliricola capromydis armatus Werneck, 1944:397. Type host: Capromys [=My- sateles| prehensilis [prehensilis] (Poep- pig). Male.—Sternum II apparently near Fig. 2, constricted and weakly pigmented me- dially, but difficult to discern. Setae on ter- gum IV, 54. Setae on sternum II, 8; IV, 19; V, 24; VII, 27. Spiracle diameter, 0.015. Dorsal terminalia as in Fig. 14; dorsal scler- ite with prominent widely spaced lobes; short spiniform setae on each lobe and along medioposterior border. Ventral ter- minalia as in Fig. 15, with terminal portion bearing weak groups of projections basally, VOLUME 110, NUMBER 2 transverse lines of fine projections apically. Genitalia close to Fig. 5; mesosomal struc- ture as in Fig. 12, more slender and apically blunt. Dimensions: POW, 0.17; TW, 0.23; HL, 0.23; PW, distorted; MW, 0.23; AWIV, 0.38; TL, distorted; GW, 0.15; GPL, 0.14; GL, 0.28. Female.—Sternum II as for male. Setae on tergum V, 43-51. Setae on sternum II, 8; IV, 22-24; V, 25-27. Spiracle diameter, 0.015. Terminalia as in Fig. 13; only lateral dorsal anterior setae on terminal segments; subgenital plate with total of 36—41 setae, none of these very long; setae around anus as shown. Dimensions: POW, 0.18—0.20; TW, 0.24—0.26; HL, 0.23—0.25; PW, 0.21; MW, 0.23-—0.24; AWIV, 0.48—-0.50; TL, 1.28—1.29. Type material.—Holotype male, allotype female, ex Mysateles prehensilis [prehen- silis|: Cuba: [Pinar del Rio], San Diego de los Bafios, 8 Apr 1900, coll. Palmer and Riley #137 56 (USNM 103887); in collec- tion of University of California (Berkeley). Other material examined.—1 female, ex Mysateles prehensilis [prehensilis|: Cuba: Pinar del Rio, 4 December 1940, coll. I. Perez Viqueras. 1 female, ex M. p. prehen- silis: Cuba: Havana, Feb 1886, coll. C. B. Cory (FMNH 15021). Remarks.—This species co-occurs with Gliricola ewingi on individuals of Mysate- les prehensilis. While the male genitalia are close to those of G. capromydis, the unique chaetotaxy of the dorsal terminal plate and the sculpturing of the ventral terminal seg- ment will separate males of these species. The female, with the virtual absence of any median anterior setae on the last 2 abdom- inal terga, is unique; this is further sup- ported by differences in the ventral termin- alia chaetotaxy. Gliricola (Hutiaphilus) omahonyi Werneck Figs. 16—20 Gliricola o’mahonyi Werneck, 1951:309. Type host: Geocapromys ingrahami (J. A. Allen). 291 Male.—Sternum II as in Fig. 17, of equal width throughout, curve relatively flattened. Setae on tergum IV, 55—62. Setae on ster- num II, 0O—2; IV, 10-11; V, 15—16; VII, 21. Spiracle diameter, 0.010. Dorsal terminalia as in Fig. 18; dorsal sclerite with prominent closely-set lobes; short spiniform setae on each lobe, sparse fine setae elsewhere. Ven- tral terminalia as in Fig. 19, with terminal portion bearing weak transverse lines ba- sally, weak transverse lines of fine projec- tions apically. Genitalia as in Fig. 20; with ovoid mesosomal structure as shown, only small slender sclerite anterior to this; para- meres with sharp barb on outer margin. Di- mensions: POW, 0.16—0.17; TW, 0.22; HL, 0.19—0.20; PW, 0.15—0.16; MW, 0.18-—0.19; AWIV, 0.32; TL, 0.84; GW, 0.11; GPL, 0.12; GL, 0.25—0.26. Female.—Sternum II as for male. Setae on tergum V, 31—38. Setae on sternum II, 2; IV, 10-11; V, 15-17. Spiracle diameter, 0.010—0.013. Terminalia as in Fig. 16; dor- sal anterior setae distributed across seg- ments; subgenital plate with total of 24—30 setae, none of these very long; setae around anus as shown. Dimensions: POW, 0.15— 0.18; TW, 0.21—0.24; HL, 0.19-0.21; PW, 0.16—0.18; MW, 0.19-—0.22; AWIV, 0.37— 0.40; TL, 0.91-0.99. Material examined.—1 male, 3 females, ex Geocapromys ingrahami: Bahamas: Lit- tle Way Cay, 12 Dec 1985, coll. K. C. Jor- dan. 1 male, 1 female ex Geocapromys in- grahami ingrahami: Bahamas: [Plana Keys], East Plana Cay, 22°27'N, 73°32’W, 4 Mar 1953, coll. George B. Rabb and E. B. Hayden, Jr. #136 ¢ (KU 60655). 3 fe- males, G. i. ingrahami: Bahamas: Plana Keys, East Plana Cay, Feb 1891, coll. D. P. Ingraham (FMNH 5624); 1 female, same data, except (FMNH 15022). Remarks.—The reduced chaetotaxy of sternum II and the unique male genitalia, in conjunction with the close-set lobes of the male terminal dorsal plate and the chaeto- taxy of the female terminalia, easily distin- guish Gliricola omahonyi from the other species of Hutiaphilus. Even though we . 252 were unable to study any type material of G. omahonyi, the description and illustra- tions furnished by Werneck (1951) left no doubt as to the correct identity of our ma- terial. In his initial description, Werneck (1951) used the spelling o’mahonyi for this spe- cies. Under articles 27, 32(c)(vi), and 32(d) of the International Code of Zoological No- menclature, no apostrophes are to be used in scientific names. Names containing apos- trophes are now considered to be spelled incorrectly, and as such are to be changed to omit the apostrophe (ICZN 1985). We herein correct the spelling of the specific epithet to omahonyi. Gliricola (Hutiaphilus) rabbi, new species Figs. 21—23 Type host.—Geocapromys ingrahami (J. A. Allen) Male.—Sternum II much as in Fig. 6. Se- tae on tergum IV, 63—64. Setae on sternum II, 6—8; IV, 13-14; V, 17-19; VII, 21-22. Spiracle diameter, 0.010. Dorsal terminalia as in Fig. 21; dorsal sclerite with prominent widely spaced lobes; short spiniform setae on each lobe, fine setae elsewhere. Ventral terminalia as in Fig. 22, with terminal por- tion bearing conspicuous uninterrupted transverse lines of projections basally, weak scattered heavier projections apically. Gen- italia as in Fig. 23; with tapered blunt me- sosomal structure as shown only small slen- der sclerites anterior to this. Dimensions: POW, 0.18—0.19; TW, 0.24—0.25; HL, 0.23; PW, 0.19; MW, 0.22; AWIV, 0.38; TL, 0.99-1.01; GW, 0.12; GPL, 0.12—0.13; GL, 0.27-0.28. Female.—Sternum II as for male. Setae on tergum V, 51. Setae on sternum II, 7; IV, 12; V, 13. Spiracle diameter, 0.010. Termin- alia distorted preventing adequate illustra- tion, but grossly near Fig. 16; dorsal ante- rior setae extending across segments; sub- genital plate with total of 25 setae, none of these very long, and basally with 4 unin- terrupted transverse rows of projections, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON much as for male. Dimensions: POW, 0.18; TW, 0.25; HL, 0.23; PW & MW, distorted; AWIV, 0.47; TL, 1.20. Type material.—Holotype male, ex Geo- capromys ingrahami: Bahamas: Little Way Cay, 12 Dec 1985, coll. K. C. Jordan; in collection of Oklahoma State University (Stillwater, OK). Paratypes: 1 male, same data as holotype; 1 female, ex Geocapro- mys ingrahami ingrahami: Bahamas: [Plana Keys], East Plana Cay, 22°27'N, 73°32’'W, 4 Mar 1953, coll. George B. Rabb and E. B. Hayden, Jr. #136 5 (KU 60655). Etymology.—This species is named in honor of George B. Rabb, Director of the Chicago Zoological Society (Brookfield Zoo), and collector of the original host from which we recovered this species of louse over four decades later and in recognition of his long commitment to conservation and public education. Remarks.—The presence in both sexes of Gliricola rabbi of uninterrupted transverse rows of small projections near the base of the ventral terminal segment sets this spe- cies apart from all other Hutiaphilus except the female of G. ewingi. Separation is fur- ther supported by the unique male genitalia. It is interesting that our entire series of lice from Geocapromys ingrahami bore the identification of Gliricola omahonyi. The fact that there were two easily separated species involved had escaped the identi- fier’s attention. This represents the third in- stance of a pairing of species of Hutiaphilus on the same host individual. Gliricola (Hutiaphilus) schwartzi, new species Figs. 24—29 Type host.—Mysateles melanurus melan- urus (Poey) Male.—As in Fig. 29. Sternum II as in Fig. 25. Setae on tergum IV, 53-62. Setae on sternum II, 7—8, shorter median setae re- cessed from posterior margin; IV, 15—17; V, 16-18; VII, 20-25. Spiracle diameter, 0.013—0.015. Dorsal terminalia as in Fig. VOLUME 110, NUMBER 2 293 Figs. 13-25. 13-15, Gliricola armatus: (13) female terminalia; (14) male dorsal terminalia (15) male ventral terminalia. 16—20, Gliricola omahonyi: (16) female terminalia; (17) sternum II; (18) male dorsal terminalia; (19) male ventral terminalia; (20) male genitalia. 21-23, Gliricola rabbi: (21) male dorsal terminalia; (22) male ventral terminalia; (23) male genitalia. 24—25, Gliricola schwartzi: (24) male genitalia; (25) sternum II. 28; dorsal sclerite with prominent widely spaced lobes; short spiniform setae on each lobe, sparse fine setae elsewhere. Ventral terminalia as in Fig. 27, with terminal por- tion bearing weak short transverse lines ba- sally, weak short transverse lines of fine projections apically. Genitalia as in Fig. 24; with slender, apically truncate mesosomal structure as shown. Dimensions: POW, 0.17—0.18; TW, 0.24—0.25; HL, 0.19-—0.20; 294 PW, 0.17—0.18; MW, 0.19—0.21; AWIV, 0.36—0.37; TL, 0.92—0.96; GW, 0.13-—0.14; GPL, 0.14; GL, 0.28—0.30. Female.—Sternum II as for male. Setae on tergum V, 44-57. Setae on sternum II, 7-9; IV, 12-16; V, 16—20. Spiracle diame- ter, 0.015—0.025. Terminalia as in Fig. 26; dorsal anterior setae extending across seg- ments; subgenital plate with total of 28—35 setae, none of these very long, but with short row of 5—8 longer submarginal setae associated with U-shaped pigmentation; se- tae around anus as shown. Dimensions: POW, 0.18—0.21; TW, 0.26—0.28; HL, 0.21- 0.24; PW, 0.19-0.22; MW, 0.23-0.26; AWIV, 0.44-—0.52; TL, 1.13-—1.34. Type material.—Holotype male, ex My- sateles melanurus melanurus: Cuba: Hol- guin; Santa Maria, Gibara, 24 August 1951, coll. M. Diaz-Piferrer, Albert Schwartz #3500 3 (KU 147709); in collection of the National Museum of Natural History (Washington, DC). Paratypes: 4 males, 3 fe- males, same data as holotype; 3 females, Cuba: [Santiago de Cuba], 22 km S of Bueycito, 26 Dec 1954, coll. Ventura, Al- bert Schwartz #3026 6 (KU 147707); 4 fe- males, Cuba: Holguin, Cueto, Guira River, 11 Jul 1949, coll. M. Diaz-Piferrer, Albert Schwartz #3499 6 (KU 147708). Etymology.—This species is named after the late Albert Schwartz in honor of his ca- reer in documenting the fauna of the Carib- bean islands and in recognition of his as- sembling the exceedingly valuable collec- tion of hutias, which made this study pos- sible. Schwartz’s efforts in training and supporting students, his foresight in assem- bling zoological collections, and his written works are an irreplaceable contribution to our knowledge of the evolution of this fau- na and its conservation. Remarks.—The unique male genitalic mesosome, with its slender shape and trun- cate apical margin, the female ventral ter- minalia with the longer submarginal setae, and the anal setae as shown, along with de- tails of the shape and chaetotaxy of sternum II and the male terminalia, afford separation PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from the other taxa. This species represents the first of three new species reported here from the host taxon Mysateles melanurus. Gliricola (AHutiaphilus) pinei, new species Fig. 35 Type host.—Mysateles melanurus melan- urus (Poey) Male.—Unknown. Female.—Sternum II close to Fig. 2. Se- tae on tergum V, 38—44. Setae on sternum II, 7-8; IV, 20—27; V, 26—29. Spiracle di- ameter, 0.010. Terminalia as in Fig. 35; dor- sal anterior setae extending across seg- ments; subgenital plate with total of 25—28 short, 6 very long setae; setae around anus as shown. Dimensions: POW, 0.16; TW, 0.21-—0.22; HL, 0.19—0.20; PW, 0.18—0.19; MW, 0.20—0.21; AWIV, 0.41-0.42; TL, 0.97—1.06. Type material.—Holotype female, ex Mysateles melanurus melanurus: Cuba: Holguin; Sarta Maria, Gibara, 24 Aug 1951, coll. M. Diaz-Piferrer, Albert Schwartz #3500 3 (KU 147709); in collection of the National Museum of Natural History (Washington, DC). Paratypes: 2 females, same data as holotype. Etymology.—This species is named for Ronald H. Pine in honor of his long com- mitment to the study of biodiversity through both field collections and published works. His efforts have greatly increased our understanding of Central and South American mammals. Remarks.—Although the male is un- known for Gliricola pinei, the female is sufficiently different with its three very long setae on each side of the subgenital plate and the unusual anal chaetotaxy to justify its recognition as a distinct species. Its dimensions also are consistently smaller than either of the other two species of My- sateles melanurus. This species co-occurred with G. schwartzi on the same host individ- ual. VOLUME 110, NUMBER 2 Gliricola (Hutiaphilus) werneckKi, new species Figs. 30-34 Type host.—Mysateles melanurus melan- urus (Poey) Male.—Sternum II as in Fig. 30, of equal width throughout, but with fainter median area. Setae on tergum IV, 65. Setae on ster- num II, 8, all prominent; IV, 19; V, 23; VII, 29. Spiracle diameter, 0.023. Dorsal termin- alia as in Fig. 32; dorsal sclerite with prom- inent slender widely spaced lobes; short spiniform setae on each lobe, sparse fine setae elsewhere. Ventral terminalia as in Fig. 31, with terminal portion bearing weak interrupted transverse lines basally, weak short transverse lines of fine projections apically. Genitalia as in Fig. 34; with broad, parallel-sided mesosomal structure as shown, apically rounded with median in- dentation. Dimensions: POW, 0.18; TW, 0.24; HL, 0.23; PW, 0.18; MW, 0.19; AWIV, 0.37; TL, 1.09; GW, 0.14; GPL, 0.15; GL, 0.31. Female.—Sternum II as for male. Setae on tergum V, 43—47. Setae on sternum II, 8, lengths as for male; IV, 15-16; V, 22—24. Spiracle diameter, 0.025—0.028. Terminalia as in Fig. 33; dorsal anterior setae extend- ing across segments; subgenital plate with total of 29-32 short, 9-11 very long setae; setae around anus as shown. Dimensions: POW, 0.20—0.21; TW, 0.27-—0.28; HL, 0.26—0.28; PW, 0.22—0.23; MW, 0.24-0.26; AWIV, 0.48—0.50; TL, 1.35—1.36. Type material.—Holotype male, ex My- sateles melanurus melanurus: Cuba: [San- tiago de Cuba], 22 km S of Bueycito, 26 December 1954, coll. Ventura, Albert Schwartz #3026 3 (KU 147707); in collec- tion of the National Museum of Natural History (Washington, DC). Paratypes: 3 fe- males, same data as holotype. Etymology.—This species is named in honor of Fabio Leoni Werneck in recogni- tion of his pioneering work on the taxono- my of chewing lice of mammals, including extensive studies on the taxa of Gliricola. 295 Remarks.—The female of Gliricola wer- necki, with its 4—6 very long setae on each side of the subgenital plate, its anal chae- totaxy, its large dimensions, and other mi- nor features, is separable from that of G. pinei and others of the subgenus. As has proven to be the case with all species for which male genitalia are known, the geni- talic details of G. wernecki are unique. This louse species also co-occurred with G. schwartzi on the same individual of Mysa- teles melanurus, but from a different local- ity than that with the pairing of G. schwart- zi and G. pinei. We cannot comment further on this now, but it may be an indication of a potential problem with contamination or host identifications. Discussion The amblyceran family Gyropidae is re- stricted to the Neotropics and contains eight genera, which are found on an array of ro- dents (primarily caviomorphs), as well as on night monkeys (Primates: Cebidae) (Price & Timm 1995). Clay (1970) recog- nized three subfamilies within the Gyropi- dae: Gyropinae, Protogyropinae, and Gliri- colinae. The Gliricolinae contains the cur- rently known 39 species of Gliricola, in- cluding 30 in the nominate subgenus and 9 in the new subgenus Hutiaphilus. The Hu- tiaphilus are found only on the Caribbean hutias of the family Capromyidae; they are absent from all other families of New World caviomorph rodents. Where accurate records are available, gy- ropids appear to be extremely host specific (Price & Timm 1993, 1995). The older lit- erature, especially for South American ro- dents, provides numerous examples where several species of lice have been described from a single host species. Most of these have been difficult for us to evaluate when host individuals were collected in different parts of a species range, hosts have not been preserved for confirmation of identifica- tions, and workers in different time periods and in different countries had different no- — 296 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 33 an fas) dd *}s eee | AY ? a (e7 eo \n . nat a te / 1\ ra WY if 44 ~ SCT ae Figs. 26-35. 26-29, Gliricola schwartzi: (26) female terminalia; (27) male ventral terminalia; (28) male dorsal terminalia; (29) male. 30—34, Gliricola wernecki: (30) sternum II; (31) male ventral terminalia; (32) male dorsal terminalia; (33) female terminalia; (34) male genitalia. 35, Gliricola pinei: female terminalia. tions as to species limits. An extreme ex- ample of this is the complex array of names for Gliricola found on the spiny rats of the genus Proechimys (Emerson & Price 1981). There is little doubt that some of the named species of Gliricola are associated. with hosts whose names have been applied er- roneously. Recently, however, we have shown that two, and even three, species of chewing lice VOLUME 110, NUMBER 2 co-occur on two other groups of cavio- morph rodents. Individuals of the spiny tree rat, Mesomys hispidus (Desmarest), in Peru have two species of Gliricola, a situation parallel to our findings on hutias presented herein (Price & Timm 1993). In the New World porcupines (Erethizontidae), we doc- umented that two, and on occasion three, species of chewing lice of the genus Eutri- chophilus Mjoberg occur on several of the South American porcupine species (Timm & Price 1994). These parallel findings are remarkable in that Gliricola and Eutricho- philus are not closely related, with the for- mer belonging to the suborder Amblycera and the latter to the suborder Ischnocera. Historically, it was considered rare to find multiple species of chewing lice on a single mammal host species (Hopkins 1957), whereas on birds it is typical to find chew- ing lice of several species belonging to two, three, or more genera on a single host spe- cies. We herein confirm that each species of Gliricola (subgenus Hutiaphilus) is restrict- ed to a single host species (high host spec- ificity). Additionally, each host species has two (and in one case, three) species of lice, and two species of Hutiaphilus are typically found on single host individuals, confirm- ing that these chewing lice can be truly sympatric. Four subfamilies are recognized in the West Indian rodent family Capromyidae. We now have lice from only one of the four subfamilies, the Capromyinae, which in- cludes three extant genera: Capromys, Geo- capromys, and Mysateles. Woods (1989) hypothesized that the Capromyidae almost certainly evolved from South American echimyid rodents in the central Antilles (Hispaniola) and that the subfamily Capro- myinae is a derived clade that originated on Cuba and secondarily dispersed to other is- lands in the Greater Antilles as well as to islands in the Bahamas. Although the fossil record for the capromyids goes back only to the Pleistocene, Woods dates the origin of the Capromyinae as “‘after the early Mio- 207 cene”’ or “‘as late as the Pliocene’? (Woods 1989:760). Our discovery of lice of the subgenus Hutiaphilus on all species of capromyines that we have studied to date (four species, representing the three extant genera of the Capromyinae) suggests that these lice are broadly distributed on these rodents and differentiated as their hosts dispersed and speciated on the various islands of the west- ern Antilles. We herein recognize the sub- genera Gliricola and Hutiaphilus as sister clades within the genus Gliricola. Lice of the subgenus Gliricola are broadly distrib- uted on the Central and South American ro- dents of the family Echimyidae, and as we now recognize the subgenus, it is restricted to the echimyids. Thus, the relationships of the lice fully support Woods’s hypothesis that capromyids evolved from echimyids. Although at present we are recognizing the subgenera Gliricola and Hutiaphilus as sis- ter clades, further resolution of the relation- ships of these species within the genus Glir- icola may suggest that the family Capro- myidae is nested within what is now rec- ognized as the Neotropical family Echimyidae. The South American nutria or coypu, Myocastor coypus, has been included as a member of the family Capromyidae (see Hall 1981, Nowak 1991). To date only a single species of chewing louse, Pitruf- guenia coypus Marelli, is known to parasit- ize M. coypus. Based upon the morphology and relationships of this louse in the dis- tinctive genus Pitrufguenia, we concur with recent classifications that treat nutria as a separate lineage (family Myocastoridae) of caviomorph rodents. The host family Capromyidae is a Carib- bean endemic, being found throughout many of the islands of the Lesser and Greater Antilles, as well as the Bahamas. The family contains 20 to 40 Recent spe- cies, although at least 70% of these are ex- tinct. The causes of extinctions are complex and certainly involve overhunting by the Amerindian civilizations, who utilized hu- 298 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tias aS a major portion of their diet (Wing 1989). The introduction of feral cats to the islands by Europeans also contributed sig- nificantly to the extinction of hutias (Mor- gan 1985). Although we have much to learn about the chewing louse genus Gliricola, our rec- ognition of one clade, Hutiaphilus, restrict- ed to the rodent family Capromyidae (sub- family Capromyinae), is an important step towards clarifying host-distributional rela- tionships. Given that chewing lice have now been found on only 4 of the 20 to 40 capromyid species, we strongly suspect that numerous new species of Gliricola await discovery. Unfortunately, a number of the species of capromyids are now extinct. Al- though teeth and/or skeletal remains for all of the species of extinct hutias are known from the fossil record, we will never be able to reconstruct the true diversity of the par- asitic chewing lice from these interesting Caribbean endemic rodents. Key to the Species of Gliricola (Hutiaphiluxs) Males (excluding G. pinei) 1. Dorsal plate of terminalia with close-set HODES CRIS: 1) eats el bee ce eae z — Dorsal plate of terminalia with widely spaced lobes (Figs. 4, 14, 21, 28, 32) 4 2. Genitalia (Fig. 20) without conspicuous anterior mesomeral sclerites, with barb on outer margin of each paramere. Ex Geocapromys ingrahami ............ MARE AA ok G. (A1.) omahonyi Werneck — Genitalia otherwise (Figs. 9,11)..... 3 3. Genitalia with long slender often con- voluted anterior mesosomal sclerite (Fig. 9). Ex Capromys pilorides .......... ot Pont: yaar ARAN, G. (H.) cubanus Werneck — Genitalia with H-shaped anterior meso- somal sclerite (Fig. 11). Ex Mysateles prehensilis. ...... G. (H.) ewingi Werneck 4. Dorsal plate of terminalia with short spi- niform setae along medioposterior mar- gin (Fig. 14). Ex Mysateles prehensilis 2k A A G. (H.) armatus Werneck — Dorsal plate of terminalia with fine setae along medioposterior margin (Figs. 4, Ph @Siv32)) js. elated iieol.eketeee 3D . Genitalia with mesosome triangular, ei- ther pointed (Fig. 5) or blunt (Fig. 23) CE aa Uy He ROY Ea AN. 6 Genitalia with mesosome either slender and apically truncate (Fig. 24) or broad and apically rounded (Figs. 34) ..... 7 . Genitalia as in Fig. 23; ventral terminal segment basally with uninterrupted rows of close-set projections (Fig. 22). Ex Geocapromys ingrahami ............ BRT. Ade ve. Joe G. (H1.) rabbi n. sp. Genitalia as in Fig. 5; ventral terminal seg- ment without such rows of projections (Fig. 3). Ex Capromys pilorides ...... Rk ca chee G. (H.) capromydis Werneck . Genitalia with mesosome slender, api- cally truncate (Fig. 24). Ex Mysateles melanurus ...... G. (H.) schwartzi n. sp. Genitalia with mesosome broad, apically rounded, with indentation (Fig. 34). Ex Mysateles melanurus’.. 20.0. 22 eee Females . Subgenital plate laterally with 2—6 very long prominent setae (Figs. 8, 33, 35) .. 2 Subgenital plate without such setae .. = . Subgenital plate laterally with 4—6 very long setae (Fig. 33); spiracle diameter greater than 0.020. Ex Mysateles melan- HUES. SP AL AL ce G. (H..) wernecki n. sp. Subgenital plate laterally with only 2—3 very long setae (Figs. 8, 35); spiracle di- ameter less than’0.015 .. 2... eee 3 . Anterior anal setae (Fig. 8) long, prom- inent; temple width more than 0.23; total length greater than 1.15. Ex Capromys PHOrides: 38s 4 G. (H.) cubanus Werneck Anterior anal setae (Fig. 35) short, less conspicuous; temple width less than 0.23; total length less than 1.10. Ex My- sateles melanurus .. G. (A) pinei n. sp. . Without median anterior setae on last 2 terga (Fig. 13). Ex Mysateles prehensilis...... G. (H.) armatus Werneck With median anterior setae on last 2 ter- Od Wher tiasne se «a ee a. eee >) . Basal portion of subgenital plate with several uninterrupted rows of small pro- jections (Rig. 10)0.. . 28 OCS 6 VOLUME 110, NUMBER 2 — Without such rows of small projections 6. Sternum II as in Fig. 2; spiracle diameter greater than 0.015. Ex Mysateles prehen- Si EE is G. (H1.) ewingi Werneck — Sternum II as in Fig. 6; spiracle diameter less than 0.015. Ex Geocapromys ingra- GET Ch ER Oh Se ede. GAMES G. (H.) rabbi n. sp. 7. Sternum II (Fig. 17) with total of 2 or less setae. Ex Geocapromys ingrahami eM a Bets G. (H1.) omahonyi Werneck — Sternum II (Figs. 2, 25) with at least 6 UIE Sw 2 yee eae Tg ee eneeee 8 8. Subgenital plate with medioposterior row of 5—8 longer submarginal setae as- sociated with U-shaped pigmentation (Fig. 26); sternum II as in Fig. 25. Ex WIVSGICICS THELGHUTUS 0. 2.x. ccen bine 9 th. RS eee oe G. (—1.) schwartzi n. sp. — Subgenital plate without such row of longer setae (Fig. 1); sternum II much as in Fig 2. Ex Capromys pilorides ...... G. (H1.) capromydis Werneck “ee © © © © ee ew Acknowledgments This project was supported in part by a grant from the National Science Foundation (DEB-9301021). Lawrence R. Heaney, AlI- fred Newton, Jr., and Bruce D. Patterson graciously assisted in obtaining material de- posited at the Field Museum. Robert P. An- derson and Linda K. Gordon provided in- formation on specimens housed at the Na- tional Museum of Natural History (Wash- ington, DC). We thank Barbara L. Clauson and Lance A. Durden for their constructive comments on this manuscript and Robert Anderson for translating our abstract into Spanish for the resumen. The skill of Errol D. Hooper and Thor Holmes in preparing the host specimens, thereby allowing us to undertake this study, is greatly appreciated. We also thank Nancy Adams, National Mu- seum of Natural History, and Cheryl B. Barr, Essig Museum of Entomology, Uni- versity of California (Berkeley, CA), for lending us critical material of the Werneck species. 299 Literature Cited Clay, T. 1970. The Amblycera (Phthiraptera: Insec- ta).—Bulletin of the British Museum (Natural History), Entomology 25:73-98. Denny, H. 1842. Monographia anoplurorum Britan- niae. Henry G. Bohn, London, xxvi + 262 pp. [+xxvi plates]. Emerson, K. C., & R. D. Price. 1975. Mallophaga of Venezuelan mammals.—Brigham Young Uni- versity Science Bulletin, Biological Series 20(3):1-77. . 1981. A host-parasite list of the Mallophaga on mammals.—Miscellaneous Publications of the Entomological Society of America 12:1-72. Ewing, H. E. 1924. On the taxonomy, biology, and distribution of the biting lice of the family Gy- ropidae.—Proceedings of the United States Na- tional Museum 63(20):1—42. Hall, E. R. 1981. The mammals of North America. Vol. 2. 2nd Edition. John Wiley & Sons, New York, xv + 601-1181 + 90 pp. Hopkins, G. H. E. 1957. The distribution of Phthir- aptera on mammals. Pp. 88-119, in First sym- posium on host specificity among parasites of vertebrates. Institut de Zoologie, Université de Neuchatel, Neuchatel, 324 pp. 1985. International Code of Zoological No- menclature, 3rd ed. International Trust for Zoo- logical Nomenclature in association with the British Museum (Natural History), London, 338 Pp. Meyer, B., & J. Wolf. 1810. Taschenbuch der deutsch- en Voégelkunde, oder Kurze beschreibung aller vogel Deutschlands. Frankfurt am Main, Fried- rich Wilmans. Mjoberg, E. 1910. Studien iiber Pediculiden und Mal- lophagen.—Zoologischer Anzeiger 35:287- 293. Morgan, G. S. 1985. Taxonomic status and relation- ships of the Swan Island hutia, Geocapromys thoracatus (Mammalia: Rodentia: Capromyi- dae), and the zoogeography of the Swan Islands vertebrate fauna.—Proceedings of the Biologi- cal Society of Washington 98:29—46. Nowak, R. M. 1991. Walker’s mammals of the world. Vol. II. 5th ed. Johns Hopkins University Press, Baltimore, pp. 643-1629. Price, R. D., & R. M. Timm. 1993. Two new species of Gliricola (Phthiraptera: Gyropidae) from the spiny tree rat, Mesomys hispidus, in Peru.—Pro- ceedings of the Biological Society of Washing- ton 106:353-358. 1995. The chewing louse genus Aotiella (Phthiraptera: Gyropidae) from South American night monkeys, Aotus (Primates: Cebidae).— ICZN. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Proceedings of the Entomological Society of Washington 97:659-—665. Timm, R. M., & R. D. Price. 1994. Revision of the chewing louse genus Eutrichophilus (Phthirap- tera: Trichodectidae) from the New World por- cupines (Rodentia: Erethizontidae).—Fieldiana: Zoology (New Series) 76:1—35. Werneck, FE L. 1935. Notas para o estudo da ordem Mallophaga.—Memorias do Instituto Oswaldo Cruz 30:417—435. . 1944. Cinco espécies novas do género “*Glir- icola’”’? (Mallophaga, Gyropidae).—Revista Brasileira de Biologia 4:391—399. . 1948. 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Morphology, systematics, and distribution of Meoma ventricosa grandis and M. ventricosa ventricosa (Echinodermata: Echinoidea: Brissidae) along Mexican coasts Francisco A. Solis-Marin, Alfredo Laguarda-Figueras, and Antonio Leija-Tristan (FAS-M, AL-F) Laboratorio de Sistematica y Ecologia de Equinodermos, Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Aut6énoma de México, Apdo. Post. 70-305, México, D. F 04510; (AL-T) Departamento de Ecologia, Facultad de Ciencias Bioldégicas, U.A.N.L., Apdo. Post. 365, San Nicolas de los Garza, N. D. 66451, México Abstract.—Two subspecies of the genus Meoma are known from the coasts of México, Meoma ventricosa grandis and M. ventricosa s.s. These conspic- uous spatangoids, easily recognized by their incomplete subanal fasciole, are confined to the tropical region of the American seas and are thus characteristic of this region. The systematics of M. ventricosa grandis and M. ventricosa s.s. are reviewed. A new northern distribution limit is now recognized for M. ven- tricosa grandis, extending its range to Magdalena Bay, Baja California Sur, México. Resumen.—En las costas de México, existen dos subespecies del género Meoma: Meoma ventricosa grandis y M. ventricosa ventricosa. Estos conspi- cuos espatangoides que se reconocen facilmente por su fasciola subanal incom- pleta, estan confinados a la regién tropical de los mares americanos siendo eminentemente caracteristicos de la misma. En este trabajo se revisa la siste- matica de las dos subespecies. Por otra parte, se amplia el rango de distribucién de Meoma ventricosa grandis hasta Bahia Magdalena, Baja California Sur, México, que es el limite norte de su distribucién. Species of the order Spatangoida live on muddy and sandy bottoms in all oceans, ex- hibiting wide bathymetric distributions, from the intertidal zone down to 5000 m (Chesher 1970). The family Brissidae Gray, 1855, is represented along the Mexican coasts by 15 species, most of which have wide bathymetric distributions and are abundant in deep waters, they are Brissop- sis alta, B. columbaris, B. pacifica, B. at- lantica; Plethotaenia spatangoides; Brissus latecarinatus, B. obesus, B. unicolor, B. elongata; Plagiobrissus grandis, P. pacifi- cus; Meoma ventricosa grandis, M. ventri- cosa ventricosa; Metalia maculosa, M. no- bilis and M. pectoralis. Meoma ventricosa is the most widely dis- tributed species of brissid in Mexican wa- ters: M. ventricosa grandis Gray, 1851 is distributed along the Mexican Pacific coast and M. ventricosa s.s. (Lamarck, 1816) in the south east of the Gulf of Mexico and the Caribbean Sea. M. ventricosa s.s. is abundant in the Mexican Caribbean Sea, while M. ventricosa grandis is scarce with- in Mexican waters (Caso 1949, 1961, 1983). Their thin test and digging habits make them difficult to collect and preserve with conventional sampling methods. There are few published works referring to the taxonomy of the genus Meoma Gray. Mortensen (1951) mentioned the existence of two recent species of the genus for the American coasts: Meoma ventricosa (La- — 302 marck) and M. grandis Gray. Chesher (1970) expanded the genus Meoma by in- cluding a new species. He also considered the previously known recent species as two subspecies of M. ventricosa: Meoma ven- tricosa s.s. and M. ventricosa grandis. A\- though the taxonomic status of these sub- species has been reviewed by Chesher (1970), some aspects of the classification at this level have not yet been completely re- solved. ; The purpose of this work is to clarify as much as possible the taxonomic status of the subspecies of Meoma ventricosa and to doc- ument a new distribution range for M. ven- tricosa grandis along the Pacific Coast. Material and Methods Measurements of the test of Meoma ven- tricosa were taken using some of the stan- dard measurements suggested by Chesher (1969, 1970). All measurements were made with vernier calipers to the nearest 0.1 mm. A total of 39 specimens was used in this work and data from 18 specimens were as- sembled for the statistical analysis (nine of each subspecies). To ameliorate the effects of growth and allometry, specimens of sim- ilar sizes of both subspecies were chosen. The range in the test length of M. ventri- cosa s.s. is from 94.84 to 145.41 mm (X 122.06 mm) and M. ventricosa grandis is from 110.25 to 130.81 mm (X 119.92 mm). Variations between subspecies were anali- zed by simple and multiple regression anal- yses. Throughout the study, statistical sig- nificance is set at the 95 per cent level using Student’s ¢ test values. A total of 33 measurements was chosen to assess variation in test shape and the characteristics of some important structures. The variables and their abbreviations are defined as follows: test length (TL); test width (TW); test height (H); height of the periproct (AH); width of the periproct (AW); distance from the adoral median sec- tion of the subanal fasciole to the periproct (SAFT); width of the base of 5 tubercles PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON - for each ambulacra zone (I, I, HI, IV, V) located in a perimeter of 20 mm around the apical system (T), and width of the peri- stome (PW). Material used in this study is housed in the Laboratorio de Sistematica y Ecologia de Equinodermos of the Instituto de Cien- cias del Mar y Limnologia at the Univer- sidad Nacional Aut6noma de México (ICMyL-UNAM). Results and Discussion Taxonomy Family Brassidae Gray, 1855 Genus Meoma Gray, 1851 Diagnosis.—Large echinoids, with peri- petalous fasciole markedly indented be- tween the paired petaloids, forming an acute angle in interambulacra 2 and 3. Test oval, broad, with a moderate frontal de- pression, high, thick-walled, regularly arched, with highest point almost central. Oral surface flat. Posterior end obliquely di- rected downwards, truncated. Frontal am- bulacrum slightly depressed, with podial pores arranged in 2 regular rows distally. Paired ambulacra sunken, narrow, straight; only the anterior ones are slightly curved backwards at their distal ends. Interporifer- ous zones narrow, densely covered by small tubercles. Apical system slightly anterior, of the ethmolytic type with 4 genital pores; madreporite prolonged backwards so as to separate the posterior genital and ocular plates. Peristome situated anteriorly, wide, semilunar, slightly sunken. Labrum anteri- orly extended, densely covered with tuber- cles. Large periproct, located at the trun- cated posterior end of the test and over- hung. Sternum short and narrow, densely covered with large tubercles. Long epister- num (nearly half the length of sternum). Anal fasciole absent. Subanal fasciole bi- lobed in Meoma frangibilis and M. caden- ati, aborally degenerate in adults of M. ven- tricosa s.s. and M. ventricosa grandis (only the adoral transverse branch remains differ- entiated). Five types of pedicellariae: glob- VOLUME 110, NUMBER 2 iferous, tridentate, ophicephalous, triphyl- lous and rostrate. Type species:—Meoma grandis = ventricosa grandis Gray, 1851). In Mexico, this genus is represented by two subspecies: Meoma ventricosa grandis (West Coast) and M. ventricosa ventricosa (East Coast). (M. Meoma ventricosa ventricosa (Lamarck, 1816) Figs. 1-3, Table 1 Spatangus ventricosus.—Lamarck 1816: 29; Blainville 1827: 89. Brissus ventricosus.—Gray 1825: 431. Meoma ventricosa.—Mortensen 1951: 529; Fontaine 1953: 8; Mayr 1954: 6; Hy- man 1955: 548, 556; Madsen 1957: 476; Caso 1961: 309; Kier & Grant 1965: 38; Fischer 1966: U592; Chesher 1969: 72-110. Meoma ventricosa ventricosa.—Chesher 1970: 737-745; Serafy 1979: 94-98; Paw- son 1986: 537; Hendler et al. 1995: 243— 245. Material examined.—2 specimens, Puer- to Morelos, Quintana Roo, México (20°54'07"N, 86°52'05”W), ICMyL-UNAM 4.95.0; 1 specimen, Puerto Morelos, Quintana Roo, México (20°51'03”N, 86°51'07”W), ICMyL-UNAM 4.95.1; 1 specimen, Puer- to Morelos, Quintana Roo, México (20°50’06"N, 86°52'03”W), ICMyL-UNAM 4.95.2; 3 specimens, north of Cabo Catoche, Quintana Roo, México (22°32'08"N, 87°06'09"W), ICMyL-UNAM 4.95.3; 3 spec- imens, north of Cabo Catoche, Quintana Roo, México (22°56'11”N, 87°16'01”W), ICMyL- UNAM 4.95.4; 3 specimens, north of Cabo Catoche, Quintana Roo, México (22°33'01"N, 87°05'9"W), ICMyL-UNAM 4.95.5; 4 spec- imens, north of Cabo Catoche, Quintana Roo, México (22°32'08’N, 87°06'09”"W), ICMyL- UNAM 4.95.6; 1 specimen, northwest of Cabo Catoche, Quintana Roo, México (22°48'06"N, 87°13'07’W), ICMyL-UNAM 4.95.7; 1 specimen, north of Cabo Catoche, Quintana Roo, México (22°56’01’N, 87°16'01"W), ICMyL-UNAM 4.95.8; 4 spec- 303 imens, northwest of Cabo Catoche, Quintana Roo, México (23°14'09"N, 87°28'02’W), ICMyL-UNAM 4.95.9; 3 specimens, north- west of Cabo Catoche, Quintana Roo, Méx- ico (23°14’09"N, 87°28'02”W), ICMyL- UNAM 4.95.10; 2 specimens, northwest Isla Contoy, Quintana Roo, México (22°07’08’N, 86°52'03”W), ICMyL-UNAM 4.95.11; 1 specimen, Puerto Morelos, Quintana Roo, México (20°51’08"N, 86°51'06”W), ICMyL- UNAM 4.95.12. Diagnosis.—Test broad, with elongate outline. Peristome broad (14 to 18% of TL). Periproct vertically elongated; distance from lower portion of periproct to adoral portion of subanal fasciole usually equal to, or greater than, the vertical diameter of the periproct. Description.—Test broad anteriorly emarginated; the posterior contour is almost truncated and oblique in its end. Reddish brown test, covered with short and striated spines. Naked test brownish, brown or earth brown color. Apical system anterior, eth- molytic, each of the four genital plates bears a gonopore; two posterior plates are larger and more widely separated. Large tu- bercles on aboral surface scant, widely dis- persed, distributed within the peripetalous fasciole. Small tubercles numerous, distrib- uted over entire surface. Tubercles of the oral surface relatively enlarged, irregularly distributed. Anterior and posterior ambula- cral grooves deep and narrow. Length of ambulacral grooves variable: some speci- mens with ones on right side longer, others with those of left side longer. Anterior am- bulacral grooves curved backward. Poste- rior ambulacral grooves slightly longer than anterior ones. Scarce miliar granules occur on external edge of poriferous zone, be- tween pairs of pores. Peristome anterior, sunken, narrow (14.4 to 17.6% of TL); an- terior edge semicircular, keeled labrum. Periproct vertically elongated (17.75 mm mean height, 11.73 mm mean width). Per- ipetalous fasciole narrow, angular in course around petals. Subanal fasciole incomplete © 304 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Comparison of denuded tests of both subspecies. 1. Meoma ventricosa ventricosa, dorsal view; 2, peristome; 3, periproct. (Test length 134.56 mm), Puerto Morelos, Quintana Roo, México (20°51'03’N, 86°51'07"W), ICMyL-UNAM 4.95.1). 4. Meoma ventricosa grandis, dorsal view; 5, peristome; 6, periproct., (test length 122.19 mm), Playa Las Gatas, Zihuatanejo, Guerrero, México, (ICMyL-UNAM 4.57.0). in adults, enclosing a small area compared Bathymetric distribution.—From the in- with M. ventriocosa grandis. tertidal to 200 m. Distribution.—Fort Lauderdale, Florida, Holotype.—Unknown. . and the Bahamas Islands, southward Type locality.—‘‘Antilles’’ (Lamarck through the Greater and Lesser Antilles; in 1816). the Gulf of México from southern Florida westward to Central America; the Orinoco Meoma ventricosa grandis Gray, 1851 River is the southern limit of this subspe- Figs. 4-6, Table 1 cies (Hendler et al. 1995). In Mexican wa- ters it has been collected in Mérida, Yuca- Meoma grandis.—Gray 185: 132; Verrill tan (Serafy 1979); Cabo Catoche, Isla Con- 1867: 302; Agassiz 1872-74: 142; H. L. toy and Puerto Morelos, Quintana Roo. Clark 1917: 220; L. Boone 1928: 12; Zie- VOLUME 110, NUMBER 2 Table 1.—Statistical summary. Coefficients of sim- ple and multiple regression analysis of M. ventricosa s.s. (n = 9) and Meoma ventricosa grandis (n = 9). (Abbreviations in section of methods). Statistical sig- nificance level 95%. Meoma ventricosa Meoma ventricosa ventricosa grandis Correlation Correlation Variables coeff. ig coeff. r TL-TW 0.991 0.983 0.605 0.366 TL-H 0.957 0.917 OS5it 0.261 TW-H 0.937 0.878 0.596 0.355 TL-PW 0.987 0.974 —0.078 0.006 TL-TW-H t-values r t-values r TL-TW 4.501 0.999 2.856 0.998 TL-H 1.188 0.647 senhenne 1937: 236; U. S. Grant & L. G. Hertlein 1938: 130; H. L. Clark 1940: 344; J. Steinbeck & E. E Ricketts 1941: 401; H. L. Clark 1948: 344; Caso 1949: 354; Mor- tensen 1951: 526; Madsen 1957: 476; Caso 1961: 300—303; Fischer 1966: U592; Caso 1983: 66. Kleinia nigra.—Agassiz 1863-1869: 27. Meoma nigra.—Verrill 1867: 251; 1870: 93: Macropneustes grandis.—Cooke C. W. 1959: 83. Meoma ventricosa grandis.—Chesher 1970: 745. Material examined.—2 specimens, Aca- pulco, Guerrero, México, ICMyL-UNAM 4.57.1; 7 specimens, Playa Las Gatas, Zih- uatanejo, Guerrero, México, ICMyL- UNAM 4.57.0; 1 specimen, outside of the Laguna de Yavaros, Sonora, México, ICMyL-UNAM 4.57.2, and 2 specimens from Magdalena Bay, Baja California Sur, México, not in catalogue. Diagnosis.—Test with broad outline; peristome broad (19 to 22% of TL). Peri- proct almost circular. Distance from lower portion of periproct to adoral portior of su- banal fasciole usually less than the vertical diameter of the periproct. Description.—Dark brown test, covered with short, striated spines. Test wide and oval, anteriorly emarginated; posterior test 305 contour almost truncated and oblique in its end. Some specimens somewhat flattened, but majority arched at apex. Naked test light brown, dark brown or earth gray color. Apical system anterior, ethmolytic, each of the four genital plates possesses a gono- pore; two posterior plates larger and more widely separated. Large tubercles on aboral surface scant and widely dispersed, distrib- uted within the peripetalous fasciole. Small tubercles numerous, distributed over entire surface. Tubercles of the oral surface rela- tively enlarged, irregularly distributed, more tightly packed towards the anterior. Anterior and posterior ambulacral grooves wide (compared to those of Meoma ventri- cosa ventricosa). Anterior ambulacral grooves curved backward; in some speci- mens, length varies less than those of the posterior region (minimal variation 30 mm). Posterior ambulacral grooves slightly longer than anterior ones. Scarce, small miliary granules occur on the external edge of the poriferous zone, between pairs of pores. Peristome broad (19.1 to 21.1% of TL), anterior, sunken. Anterior edge formed by two lines converging on its anterior end, forming an obtuse angle, with lateral parts formed by small vertical lines. Labrum prominent, semicircular, not keeled; the posterior prolongation short. Periproct al- most circular (16.8 mm mean high, 14.2 mm mean width). Remarks.—Contour of peripetalous fas- ciole is highly variable in the observed specimens. The portion across the anterior interambulacral region varies less than oth- er parts; nevertheless, in some specimens, the middle transverse portion is almost straight and in others it is slightly curved. Also, in some specimens it is folded at the level of the ambulacral groove; it generally folds over a small distance, forming a right angle on each side. In other specimens there are two bends, the second one larger than the first, near the anterior lateral grooves. As in M. ventricosa ventricosa, some of spines in the interambulacral zones are slightly curved. Large, primary spines are | 306 dark brown. In adult specimens, the average between the anterior genital pores is 1 mm, and between the posterior pores 2 mm. Distribution.—Northern part of the Gulf of California (Angeles Channel, 29°01'05”N and 113°29’05”’W) to Punta Choco, Puerto Utria, Colombia; Galapagos and Revillagi- gedo Islands. In the Gulf of Panama, out- side Tortolita and Taboguilla Islands, at depths of 0.5-10 m (Mortensen 1951). In Mexican waters it has been collected in Punta Arena, Punta Gorda, Cabo San Lu- cas, Baja California; on the seaward side of Laguna de Yavaros, Sonora; Bahia Tena- catita, Jalisco; Playa Las Gatas, Zihuatane- jo, Acapulco, Guerrero; Bahia Tangola Tan- gola, Bahia Santa Cruz, Oaxaca (Caso 1949, 1961, 1983). Chesher (1970) reports it from Bahia Santa Inés, Bahia Concep- cion, Gulf of California; Manzanillo, Coli- ma; Acapulco, Guerrero; Puerto Huatulco, Oaxaca. Geographic range extension.—Bahia Magdalena, Baja California Sur (24°35’00"N and 112°03'15”W); two adult specimens col- lected in the spring of 1991, on sandy bot- toms, with an Ekman—Birge dredge at 26 m. Bathymetric distribution.—From the in- tertidal to 100 m. Holotype.—Meoma grandis (=M. ventri- cosa grandis Gray, 1851), The Natural His- tory Museum, London, Catalogue number 1949.10.24.6. Type locality.— Australia (locality which was taken from original label, the label no longer exists). The species is western Mex- ican and the locality Australia is therefore wrong (S. Halsey pers. comm. 1996). The original material was not labeled “‘Acapul- co, México’’ as pointed out by Chesher (1970). Statistical Analysis Simple regression and correlation analy- sis showed that between the variables LT— TW, TW-—H, TW-H (Table 1) in the case of Meoma ventricosa s.s. the correlation co- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 94 164 114 124 134 144 154 TW and H - Fig. 7. Multiple regression analysis of the test length (TL), test width (TW) and test high (H) of Meo- ma ventricosa ventricosa. (r? = 0.999, n = 9). efficients and the 7* were highly significant. Otherwise, in the case of M. ventricosa grandis this correlation was relatively weak (Table 1). The above results were confirmed by multiple regression analysis of TL with respect to TW and H, which indicated that in the case of Meoma ventricosa s.s. fit of the regression line in the plot was more sig- nificant (Fig. 7) than in the case of M. ven- tricosa grandis (Fig. 8). The shape of the test in M. ventricosa s.s. tended to be more broad and anteriorly emarginated than in M. ventricosa grandis, which bears a wide and oval shape. The simple regression analysis between TL and PW for M. ventricosa s:s. showed a highly significant correlation, while that of M. ventricosa grandis was not significant (Table 1). The simple regression analysis of AH with AW for M. ventricosa s.s. showed a correlation coefficient (0.713) and r* (0.508) that indicated AH was sig- nificantly higher than AW. This is easy to recognize by simple observation of the specimens (Fig. 3). The simple regression analysis of AH with AW for M. ventricosa grandis showed a correlation coefficient (0.311) and r? (0.097) that indicated lack of VOLUME 110, NUMBER 2 11@ 114 118 1lé@e 126 138 i134 TW and H Fig. 8. Multiple regression analysis of the test length (TL), test width (TW) and test high (H) of Meo- ma ventricosa grandis. (r* = 998, n = 9). significance: the periproct is almost circular (Fig. 6). For the correlation analysis be- tween AH and SAFT, the 7? (M. ventricosa s.s. = 0.227 and M. ventricosa grandis = 0.181) didn’t show a significant difference. Descriptive statistics showed no signifi- cant difference between the two subspecies in analysis of TL versus width of the base of the tubercles for each ambulacral zone. r’ was 0.99 in all cases. This character over- laps in both subspecies: M. ventricosa s.s., Me 14034 tnax, = 4.4. min,=,0.7 .and-SD = 0.152; M. ventricosa grandis, X = 1.02, max = 1.4 min = 0.7 and SD = 0.114. Discussion Mortensen (1951) state that “‘only the characters of the primary tubercles of the aboral side, the frontal ambulacrum, and the peripetalous fasciole seem to afford reliable distinction between the two species’’. Chesher (1970) state that the differences between M. ventricosa s.s. and M. ventri- cosa grandis are the distance from the per- iproct to the adoral portion of the subanal fasciole compared with the vertical diame- ter of the periproct, and the distance from 307 the apical system to the lateral portion of the peripetalous fasciole. In the present work, the statistical anal- yses showed following characters to be im- portant in separating the two subspecies: 1) the shape of the test—in M. ventricosa s.s. it is broad and anteriorly emarginated, whereas in M. ventricosa grandis it is wide and oval; 2) the shape and size of the peri- stome—in M. ventricosa s.s. it is narrow, with the anterior edge semicircular and a keeled labrum, whereas in M. ventricosa grandis it is broad, with anterior edge formed by two lines converging on its an- terior end and forming an obtuse angle, with lateral parts formed by small vertical lines, prominent labrum, that is semicircular and not keeled; 3) the shape of the peri- proct—vertically elongated in M. ventrico- sa s.s. and almost circular in M. ventricosa grandis. We agree with Chesher (1970) that the distance from the periproct to the adoral portion of the subanal fasciole compared with the height of the periproct is important to separate the two subspecies. Statistical analysis showed that the size of the aboral tubercles is not useful in dis- tinguishing the two subspecies, as was also pointed out by Chesher (1970). It was shown by the same author that “‘this char- acter is subject to ecological variation’’. A new distribution range is now recog- nized for M. ventricosa grandis, extending to Magdalena Bay, Baja California Sur, México. This may be the northern limit of its distribution. This locality shows a spe- cial “‘faunistic conglomerate”’ that is tran- sitional sub-tropical, with mingling of trop- ical and temperate faunas (Solis-Marin 1991). This transitional fauna does not pre- vail along the entire Baja California coast; in Magdalena Bay, a qualitative faunistic change is evident (Williams 1974). This change is explained by the influence of the California current (Loeb et al. 1983). Acknowledgments We are grateful to Dr. R. Mooi (Califor- nia Academy of Sciences) and Dr. P. K. 308 Donovan (University of the West Indies) for the revision and helpful comments on this manuscript and to Sheila Halsey (The Nat- ural History Museum, London) for infor- mation on type material of Meoma ventri- cosa grandis. We thank Cynthia Ahearn (National Museum of Natural History) for bibliographic assistance, and Dr. José Luis Soto, Dr. Elva Escobar-Briones and Bi0l. Carlos Illescas, of the ICMyL, UNAM, México for making available specimens of Meoma ventricosa ventricosa collected on the OGMEX project. Marco A. Martinez help us with statistical program. A. Bieler Antolin took the photographs. Literature Cited Agassiz, A. 1863-1869. Preliminary report on the echini and starfishes dredged in deep water be- tween Cuba and the Florida Reef.—Bulletin of the Museum of Comparative Zoology 1(1—13): 253-308. . 1872-74. Revision of the Echini.—Illustrated Catalogue of the Museum of Comparative Zo- ology at Harvard University 7(1):1—242; (3): 381-629. Blainville, H. M. 1827. Oursins.—Dictionnaire des Sciences Naturelles 37:59—245. Boone, L. 1928. Echinoderms from the Gulf of Cal- ifornia and the Perlas Islands.—Bulletin of the Bingham Oceanographic Collection, Yale Uni- versity 2(6):1—14. Caso, M. E. 1949. Contribucion al conocimiento de los equinodermos litorales de México.—Anales del Instituto de Biologia. México (20):351-354. . 1961. Los equinodermos de México. Unpub- lished Doctoral Thesis, Facultad de Ciencias, Universidad Nacional Aut6énoma de México, 388 pp. . 1983. Los equinoideos del Pacifico de Méx- ico. Parte cuarta, ordenes Cassiduloida y Spa- tangoida.—Anales del Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Au- t6noma de México, Publicacién especial 6:66— pe Chesher, R. H. 1969. Contribution to the biology of Meoma ventricosa (Echinoidea: Spatango- ida).—Bulletin of Marine Science 19(1):72— 110. . 1970. Evolution in the genus Meoma (Echi- noidea: Spatangoida) and a description of a new species from Panama.—Biological Results of the University of Miami Deep-Sea Expedition 68:731-761. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Clark, H. L. 1917. Hawaiian and other Pacific Echini. The Echinoneidae, Nucleolitidae, Urechinidae, Echinocorythidae, Calymnidae, Pourtalesiidae, Palaeostomatidae, Aeropsidae, Palaeopneusti- dae, Hemiasteridae and Spatangidae.—Memoirs of the Museum of Comparative Zoology at Har- vard College 46(2):85—283. . 1940. Notes on echinoderms from the west coast of Central America.—Eastern Pacific Ex- peditions of the New York Zoological Society, No. 21, Zoologica (New York) 25:331-352. . 1948. A report of the echini of the warmer eastern Pacific, based on the collections of the ““Velero”’? IJ.—Allan Hancock Pacific Expedi- tion 8(5):225-351. Cooke, C. W. 1959. Cenozoic echinoids of eastern United States —United States Geology Survey, Professional Papers 321:1—106. Fischer, A. G. 1966. Spatangoids. Pp. U543-U628 in R. C. Moore. ed. Treatise on invertebrate pale- ontology. Part U, Echinodermata 3, 2:695 pp. Fontaine, A. 1953. The shallow-water echinoderms of Jamaica, part. 3. The sea urchins (class Echi- noidea).—Natural History Notes Jamaica 61:39. Grant, U. S. IV, & L. G. Hertlein. 1938. The west American Cenozoic Echinoidea.—University of California Publications in Mathematical and Physical Sciences 2:1—225. Gray, J. E. 1825. Attempt to divide the Echinida or sea-eggs into natural families.—Annals of Phi- losophy 26:423—431. . 1851. Description of some new genera and species of Spatangidae in the British Muse- um.—Annals and Magazine of Natural History Ser. 2, 7:130—134. 1855. An arrangement of the families of Echinida, with descriptions of some new genera and species.—Proceedings of the Zoological Society of London 1855:35-39. Hendler, G., J. E. Miller, D. L. Pawson, & P. M. Kier. 1995. Sea stars, sea urchins & allies: Echino- derms of Florida & the Caribbean. Smithsonian Institution Press, Washington, D.C., 390 pp. Hyman, L. H. 1955. The Invertebrates, vol. 4: Echi- nodermata, the Coelomate Bilateria. New York, Mc. Graw Hill, 763 pp. Kier, P. M. & R. E. Grant. 1965. Echinoid distribution and habits, Key Largo Coral Reef Preserve, Florida.—Smithsonian Miscellaneous Collec- tions 149(6):68 pp. Lamarck, J.B. PR A. de M. de. 1816. Histoire naturelle des animaux sans vertébres. Volumen 3. Bail- liére, Paris, 1-59 pp. Loeb, V. J., PR E. Smith, & H. G. Moser. 1983. Icth- yoplankton and zooplankton abundance patterns in the California Current areas. CalCOFI, Re- port 24:109-131. Madsen, FE J. 1957. On a new species of Meoma, and VOLUME 110, NUMBER 2 on a few other echinoids from tropical West Af- rica.—Bulletin de Institute fra Africa Noire, Serie A 19(2):474—481. Mayr, E. 1954. Geographic speciation in tropical echi- noids.—Evolution 8(1):1—18. Mortensen, Th. 1951. A monograph of the Echino- idea. V. 2. Spatangoida 2, Reitzel, Copenhagen, 593 pp. Solis-Marin, E A. 1991. Composicién y distribuci6én es- pacio-temporal de los macroinvertebrados bentén- icos del complejo lagunar Magdalena-Almejas, de la costa occidental de Baja California Sur. Unpub- lished Professional Thesis, Universidad Michoa- cana de San Nicolas de Hidalgo, 95 pp. Serafy, D. K. 1979. Memoirs of the Hourglass Cruis- es. V (III): Echinoids (Echinodermata: Echino- idea). St. Petesburg, Florida: Florida Departa- ment of Natural Resources, 120 pp. 309 Steinbeck, J., & E. F Ricketts. 1941. Sea of Cortez. A leisurely journal of travel and research. New York: The Viking Press, 598 pp. Verrill, A. E. 1867. The echinoderms of Panama and West coast of America.—Transactions of the Connecticut Academy of Arts and Sciences 1(@2):251--322. . 1870. Descriptions of echinoderms and cor- als from the Gulf of California—American Journal of Science 49(2):93—100. Williams, A. B. 1974. The swimming crabs of the genus Callinectes (Decapoda, Portunidae).— United States Natural Marine Fisheries Ser- vices. Bulletin (3):685—798. Ziesenhenne, F C. 1937. Echinoderms from the west coast of Lower California. Zoologica, New York, Zoological Society (22):236 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):310-—313. 1997. New species of Paratriacanthodes spikefish (Triacanthodidae: Tetraodontiformes) from the South China Sea James C. Tyler National Museum of Natural History, Smithsonian Institution (MRC-106), Washington D.C. 20560, U.S.A. Abstract.—A new species of deep-water triacanthodid spikefish from the MacClesfield Bank in the South China Sea between the Philippines and Viet- nam is described in the subfamily Triacanthodinae as Paratriacanthodes abéi. It differs from the other two species of that western Pacific and Indian Ocean genus in several meristic and proportional characteristics and in coloration. Among fishes received from the Fisheries Research Station of Hong Kong in 1972 for deposit in the Smithsonian Institution’s Na- tional Museum of Natural History (USNM) is the single specimen of a new species of the genus Paratriacanthodes described be- low. It had been identified for shelving pur- poses when received as the more common of the two species then known. Methods.—Descriptive data are in con- formity with those of the revision of the family by Tyler (1968). Paratriacanthodes abei, new species Fig. 1 Holotype.—USNM 342571, 49.9 mm SL, mature male, MacClesfield Bank, South China Sea, 16°19.3’—16°22.2'’N, 114°29’— 114°24.5’E, 200-210 fathoms (366-384 m), 20 Jun 1964, RV Cape St. Mary, cruise 3/64, station 55, Agassiz trawl. Diagnosis.—A member of the Triacan- thodinae (posterior process of pelvis flat- tened and tapering posteriorly; supraoccip- ital flattened, as determined from a radio- graph) referable to Paratriacanthodes by the unique combination in that genus of: short snout; conical teeth in a single series; short pseudobranch with few lamellae; dor- sal-fin spines decreasing gradually in length posteriorly; and gill opening moderate. Dif- fers from the two previously known species of the genus, P. retrospinis Fowler and P. herrei Myers, by having: 16 dorsal-fin rays (versus usually 15 in the other two species); 14 anal-fin rays (versus usually 13); 13 pec- toral-fin rays (versus usually 14); 7-8 ol- factory lamellae (versus 10-14); greater body depth (54% SL versus about 40—47% SL); longer first dorsal-fin spine (38% SL versus about 28—32% SL); and color pattern with wavy reticulate stripes (versus alter- nating dark and light horizontal lines). Description.—See Table 1 for meristics of the fins, teeth, gill rakers, and olfactory and pseudobranch lamellae of the new spe- cies in comparison to those of related spe- cies. The intact right side (left side has a large cut) is shown in Fig. 1. Body depth 54.3% SL. Head length 38.9% SL, with a slightly concave profile. Snout short, 12.4% SL. Distance between tip of snout and base of first dorsal-fin spine 47.7% SL. Orbit 15.6% SL; width of rela- tively flat interorbital 7.0% SL. Postorbital length of head 9.4% SL. Gill opening rel- atively short, reaching ventrally only to a level about one-fifth down pectoral-fin base, 4.8% SL. Lamellae of pseudobranch reaching ventrally to a level just above top of pectoral-fin base. Width of slightly su- praterminal mouth 7.0% SL. A single series of strong, bluntly conical teeth in each jaw. Vertebrae 8 + 12 = 20. Length of pelvic-fin spine 34.3% SL; VOLUME 110, NUMBER 2 311 Table 1.—Meristics of the species of Paratriacanthodes and Mephisto. P. retrospinis P. herrei P. abei M. fraserbrunneri No. Spec. Range Average No. Spec. Range Average Holotype No. Spec. Range Dorsal rays 65 14-16 15.0 3 15 — 16 5 16 Anal rays 65 12-14 13.0 3 13 — 14 5 14 Pectoral rays* 128 13-15 139 6 14-15 14.2 13 9 14 Pelvic rays* 130 1 — 6 1-2 Ly | 10 ] Olfactory lamellae 38 10-13 11.8 2, 13-14 = 7-8 2 9-10 Pseudobranch lamellae 36 12-16 13.5 5 21-24 22.3 12 2 18-19 Gill rakers 37. 17-23 20.3 3 19-21 20.0 18 2 19 Teeth, upper jaw 45 10-18 14.7 3 13-17 15.0 15 2 17 Teeth, lower jaw 45 15-24 19.2 3 16-19 Lig Pes, Zz pe 19-20 * Usually two fins from the same specimen. length of unbranched rudimentary ray 1.0% (fully erect, at a right angle to pelvis). Ex- SL. Basal flange of pelvic-fin spine only ternal surface of posterior process of pelvis slightly grooved, apparently with a single flattened, with upturned lateral edges and position of spine erection against pelvis tapering to a blunt point posteriorly. Length Fig. 1. Paratriacanthodes abei, new species, holotype, USNM 342571, 49.9 mm SL, South China Sea: the anal fin is folded up against the side of the body; the pelvic-fin spines from both sides of the body are visible, that of the left side less fully erected than the other. 2 of posterior pelvic process 36.3% SL; its width between pelvic-fin spines 8.8% SL. Pectoral-fin length 17.2% SL; length of up- permost ray 5.1% SL. Lengths of dorsal-fin spines, from first to last, 38.0, 33.3, 24.4, 16.8, 9.4, and 4.0% SL. Soft dorsal-fin base 20.0% SL; longest ray 21.0% SL. Anal-fin base 17.6% SL; longest ray 17.4% SL. Anal-fin origin at level of about third dor- sal-fin ray. Caudal-peduncle depth 10.0% SL; its length 17.0% SL. Caudal-fin length 30.5% SL. Caudal-fin rays 12, with 10 un- branched. Most of the scales of the body have a row of three upright spinules, the center one the longest, but some scales have only a single central spinule, and the lateral-line scales have a pair of spinules surrounding the pore; the spinules are all simple. There is a broad patch of spinulose scales on the upper lip but none on the lower lip. Spinulose scales extend out most of the length of the soft dorsal-fin rays and along all but the dis- tal one-tenth of the length of the dorsal-fin spines. Some of the spinulose scales are slightly enlarged laterally on the first dor- sal-fin spine, but none of them is retrorse. The color pattern is faded, but melano- phores indicate a pattern of wavy reticular stripes, one below the spiny and soft dorsal fins and another with very irregular outlines from the eye, over the pectoral-fin base, and onto the top of the abdomen, with an irreg- ular pattern in the region between these two wavy stripes. There are no well-defined horizontal lines or spots in the pattern. The peritoneum is tan, lightly speckled with darker color. The color in life is unknown, but if the new species is like other triacanthodins for which living color is known, the wavy re- ticulate stripes are probably pale blue and the background color pinkish red. Etymology.—The patronym is given in honor of the late Dr. Tokiharu Abe (3 Apr 1911-9 Aug 1996), of the Museum of To- kyo University and of the Fish Museum at Tsukiji Fish Market, an authority on the fishes of Japan, and especially on the te- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON traodontiform fugus; he always shared his enthusiasm, knowledge, and specimens with other researchers having similar inter- ests. Remarks.—One of the two previously described species of Paratriacanthodes, P. herrei Myers, is known from only the three type specimens from the Philippines, and until recently the other species, P. retros- pinis Fowler, was known from only twelve specimens, the four type specimens from the South China Sea and eight others from Japan and east Africa (Tyler 1968, Amaoka 1982). However, a large series of 65 spec- imens of P. retrospinis has recently been collected by ORSTOM expeditions around New Caledonia in the South Pacific, and the species has been redescribed by Matsuura & Tyler (1997) on the basis of these 65 specimens. Therefore, the degree of vari- ability in meristics of P. retrospinis can be compared with the meristics of the new spe- cies. The dorsal-, anal-, and pectoral-fin rays of P. abei are, respectively, 16, 14, and 13, whereas in P. retrospinis only 3 of 65 spec- imens (7 from Tyler 1968, and 58 from Matsuura & Tyler 1997) have 16 dorsal-fin rays, only 2 of 65 have 14 anal-fin rays, and only 6 of 128 pectoral fins counted have 13 rays; all three specimens of P. herrei have 15 dorsal-, 13 anal-, and 14 pectoral-fin rays, like the large majority of specimens of P. retrospinis. There is no overlap in the number of olfactory lamellae between P. abei (7-8) and the other two species (10— 14) of the genus. There also is no overlap in the propor- tional measurements of body depth and dor- sal-spine length between P. abei and the other two species of the genus. Relative body depth decreases somewhat with in- creasing body length from juveniles to adults in triacanthodids. The average body depth in P. retrospinis is 46% SL at sizes between 22-40 mm SL and 40% SL at sizes between 51—110 mm SL. Thus, P. re- trospinis at all sizes has a more slender body than P. abei (54% SL), as do the three VOLUME 110, NUMBER 2 specimens of P. herrei, with an average depth of 47% SL for the 66-71 mm SL specimens. The length of the first dorsal-fin spine in P. retrospinis ranges from 28-33% SL (X¥ 31% SL) and in P. herrei from 31— 33% SL (X 32% SL), whereas it is 38% SL in P. abei. There are some similarities between P. abei and one or the other previously de- scribed species of the genus. The number of pseudobranch lamellae is relatively low in both P. abei (12) and P. retrospinis (12— 16), versus P. herrei (21-24). The gill opening is relatively shorter in both P. abei (4.8% SL) and P. retrospinis (3.7—6.6% SL, X 5.3), versus P. herrei (7.0—7.6% SL, X 7.3). The postorbital portion of the head is relatively shorter in P. abei (9.4% SL) and P. retrospinis (8.0—-10.5% SL, X 9.2), ver- sus P. herrei (10.8—11.7% SL, X 11.2). The pelvic-fin ray is relatively shorter in P. abei (1.0% SL) and P. retrospinis (1.5—3.8% SL, X 2.4), versus P. herrei (5.1-7.2% SL, X 6.3). Conversely, both P. abei and P. herrei lack retrorse barbs on the first dorsal- and pelvic-fin spines, whereas these are well de- veloped in P. retrospinis. Paratriacantho- des abei and P. herrei have only slightly grooved flanges at the base of the pelvic- fin spine and only a single position of erec- tion, whereas this flange is well grooved in P. retrospinis, allowing for multiple posi- tions of erection. Paratriacanthodes abei shares several features with the single species of Mephis- to, M. fraserbrunneri Tyler, including num- bers of dorsal- and anal-fin rays (respec- tively 16 and 14 in both species) and low numbers of olfactory lamellae (7-8 in P. abei and 9-10 in M. fraserbrunneri). Both P. abei and M. fraserbrunneri have rela- tively irregular wavy stripes or blotches on the body versus more regular horizontal lines in P. retrospinis and P. herrei. These similarities between P. abei and M. fraser- brunneri presumably are either primitive or independently derived. The two species can S13 most easily be distinguished by the much greater depth of the gill opening in M. fras- erbrunneri (12-14% SL, reaching to a level slightly below the pectoral-fin base, versus 5% SL and reaching only to the upper re- gion of the pectoral-fin base in P. abei). All three species of Paratriacanthodes are known to occur in either the South Chi- na Sea (P. abei and P. retrospinis) or the adjacent Philippines (P. herrei). Only P. re- trospinis is known to have a wider distri- bution outside of this area, with records from Japan and China to South Africa and into the western Pacific as far as New Cal- edonia (Matsuura & Tyler 1997). A speci- men of P. retrospinis from Fiji examined for this paper extends the range even further to the east (University of the South Pacific cat. no. 4393, 33.7 mm SL, from 485 m depth on the Suva Barrier Reef, 30 Septem- ber 1981, R/V Nautilus). Acknowledgments The manuscript was improved by sug- gestions received from Jeffrey T. Williams, Smithsonian Institution, and an anonymous reviewer. The specimen of Paratriacantho- des retrospinis from Fiji was examined on loan through the courtesy of Johnson Seeto and Uday Raj, University of the South Pa- cific. Literature Cited Amaoka, K. 1982. Tetraodontiformes. Pp. 302-306, 409-410 in O. Okamura, K. Amaoka, & E Mi- tani, eds., Fishes of the Kyushu—Palau Ridge and Tosa Bay. Japan Fisheries Resource Con- servation Association, Tokyo. Matsuura, K., & J. C. Tyler. 1997. Tetraodontiform fishes of New Caledonia (south—western Pacif- ic) collected by ORSTOM, mostly in deep wa- ter (100-650 m). Pp. 1-35 in B. Séret, ed., Ré- sultats des campagnes MUSORSTOM.—Me- moires du Muséum National d’Histoire Natu- relle (in press). Tyler, J. C. 1968. A monograph on plectognath fishes of the Superfamily Triacanthoidea.—Academy of Natural Sciences of Philadelphia, Monograph 16:1-364. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):314-319. 1997. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 3. Parentage of Lesbia ortoni Lawrence Gary R. Graves Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Lesbia ortoni Lawrence, 1869, collected in the Quito Valley, Ec- uador, is shown to be a hybrid between Lesbia victoriae and Ramphomicron microrhynchum, sympatric inhabitants of Andean forest edge and shrublands from Colombia to Peru. The hybrid exhibits a blended mosaic of plumage characters of the parental species. Although the parental species differ signif- icantly in size, the external measurements of the hybrid approximate those predicted by least squares regression. Ornithological literature of the 19th cen- tury is littered with the descriptions of doz- ens of enigmatic trochiline taxa from South America (Gould 1861, Salvin 1892, Bou- card 1893) that now are treated as plumage mutations or hybrids (e.g., Berlioz & Jouan- in 1944, Greenway 1978, Graves 1990). Unfortunately, these taxonomic disposals were often too brief and insufficiently doc- umented to permit considered rejection of alternate hypotheses. Consequently, the sta- tus of a significant number of nominal taxa, now all but forgotten, in fact is unresolved. The spectacular holotype of Lesbia or- toni Lawrence, 1869, was sent to Professor Orton of Vassar College from the Quito Valley of Ecuador (see Greenway 1978). It was deposited in the American Museum of Natural History (AMNH) on loan, 21 Oc- tober 1921; ownership was finally trans- ferred to the AMNH in March 1965 (fide M. LeCroy, pers. comm.). Lesbia ortoni was considered a valid species for more than 50 years (Elliot 1879, Salvin 1892, Boucard 1893, Oberholser 1902, Cory 1918, Chapman 1926). Mulsant & Verreaux (1876) erected the genus Zodalia, with or- toni as the type species. Simon (1921) treat- ed ortoni as a junior synonym of Zodalia glyceria, believing it to represent the adult male plumage of that taxon (Peters 1945). Finally, in a terse appraisal of several puz- zling taxa, Meyer de Schauensee (1947) pronounced as hybrids both ortoni and Z. glyceria (Lesbia victoriae X Ramphomi- cron microrhynchum). This opinion was followed implicitly by subsequent authors (e.g., Morony et al. 1975, Greenway 1978, Fjeldsa & Krabbe 1990, Sibley & Monroe 1990, Collar et al. 1992). Meyer de Schauensee’s proposal could be correct, but rigorous documentation of Lesbia ortoni is a critical and necessary first step in unrav- eling the parentage of other enigmatic An- dean taxa believed to represent hybrids (Graves, unpubl.). Here I provide a detailed hybrid diagnosis of Lesbia ortoni employ- ing the methods and assumptions outlined in Graves (1990). Materials and Methods The unsexed holotype of L. ortoni (AMNH 156651), a relaxed taxidermy mount with glass eyes, lacks the left wing (at the time of my first examination of the specimen in March 1986). The greatly elon- gated rectrices, large brilliant gorget, purple dorsal plumage, and unstriated maxillary ramphothecum indicate that the specimen is an adult male in definitive plumage (Figs. 1, 2). The unique appearance of Lesbia or- VOLUME 110, NUMBER 2 315 Fig. 1. toni cannot be attributed to mutation or de- velopmental variation of any known taxon. It must then represent a hybrid or a valid taxon. As hybrids have no standing in zoo- logical nomenclature, the burden of proof lies with the taxonomist to reject conclu- sively the hybrid origin of L. ortoni before bestowing species status on it. As the re- sults will show, I was unable to reject the hypothesis of hybridity. Assuming a hybrid origin of L. ortoni, the pool of potential parental species (=geographic pool) includes the species of hummingbirds (n = 48; see Appendix 1 in Graves 1996b) known to occur regularly above 2000 m elevation in the Ecuadorian Andes (Chapman 1926, Fjeldsa & Krabbe 1990). I compared L. ortoni directly with males of the potential parental species and the holotype of Chalcostigma purpureicau- da at the American Museum of Natural His- tory (AMNH 483931). Notes, photographs, and videotape of L. ortoni were compared with the holotypes of Zodalia glyceria (The Natural History Museum, BM[NH] 1888.7.25.184), Zodalia thaumasta (Na- tional Museum of Natural History, Smith- sonian Institution, USNM 173911), and He- liangelus zusii (Academy of Natural Sci- ences of Philadelphia, ANSP 159261; see Graves 1993a). The taxonomic status of C. purpureicauda, Z. glyceria, and Z. thau- masta will be addressed in future papers. Color descriptions were made under nat- ural light. Measurements of wing chord, bill length (from anterior extension of feathers), Lateral view of the type of Lesbia ortoni Lawrence (AMNH 156651). and rectrix length (from point of insertion of the central rectrices to the tip of each rectrix) were taken with digital calipers and rounded to the nearest 0.1 mm (Table 1). Measurements and least squares regression lines were projected on bivariate plots to illustrate size differences (Wilkinson 1989). The hybrid diagnosis was approached in a hierarchical manner. The presumed paren- tal species of L. ortoni were hypothesized through the comparative analysis of plum- age pattern and color and feather shape. As a second step, the restrictive hypothesis was tested with the quantitative analysis of size and external proportions. Concordance of results is regarded as strong support for the hypothesis (Graves 1990, 1993b, 1996a; Graves & Zusi 1990). Results and Discussion Characters of Lesbia ortoni (hereafter hy- brid) that permit its parental species to be identified include: large brilliant gorget; forked tail (depth = 59.7 mm); tips of out- ermost rectrices ““‘bowed’’ in cross section; purple feather tips on dorsal body plumage; lack of brilliant frontlet or crown; and short tibial plumes. None of the potential parental species alone exhibits this character com- bination. Four species (Ocreatus underwoodii, Lesbia victoriae, L. nuna, Aglaiocercus kin- gi) in the geographic source pool have deeply forked tails. Ocreatus can be elimi- nated from consideration because the hy- 316 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Ventral view of the type of Lesbia ortoni Lawrence (AMNH 156651). brid shows no evidence of racket-tipped rectrices or elongated tibial plumes. The hy- brid also lacks evidence of the awl-shaped bill (dorsal profile) and brilliant crown of Aglaiocercus kingi. Thus, by the process of elimination, one of the “‘trainbearers’”’ (Les- bia victoriae or L. nuna) is implicated in the parentage of L. ortoni. Plumage char- acters of the hybrid are consistent with this hypothesis, but it is doubtful that the spe- cific identity of the Lesbia parent can be determined from plumage color or pattern alone. Determination of the other parent seems straightforward. Only one species in the geographic pool, Ramphomicron micror- hynchum, has the rich purple dorsal plum- age (including crown) exhibited by the hy- brid. In fact, the hybrid appears to exhibit a blend of definitive male plumage char- acters of R. microrhynchum and the train- bearers (Lesbia sp.). No other combination (8 = 1128 possible pairs of species) of spe- cies in the geographic pool could have pro- duced the phenotype of the hybrid (Appen- dix). External measurements.—Examination of external measurements enables identifi- Table 1.—Ranges and means (+ one standard deviation) of measurements (mm) of adult male Lesbia victo- riae, Ramphomicron microrhynchum, and the hybrid, Lesbia victoriae * Ramphomicron microrhynchum (= Lesbia ortoni Lawrence, 1869; AMNH 156651). Measurements of L. nuna are included for comparison. victoriae* nuna* microrhynchum? ( 12 (n = 12) (n = 12) n= Hybrid Wing chord 58.7-62.2 50.3-52.8 47.3—-53.5 55.8 O03. Tal 18: = 106 54.67 2216 Bill length 13.5-15.3 7.5—9.0 5.9-7.0 10.1 PAE SG 8.2 + 0.4 6:5727053 Rectrix 1 22.1—24.9 19.8—22.4 24.3—27.0 2S 23.5 O19 210 2107 26.1 + 0.8 Rectrix 2 26.2—31.1 25.9-28.1 28.4—34.1 31.3 28. Galen 2120 2 O.7 St 2 les Rectrix 3 39.2—44.4 36.0-40.4 35.7-41.0 40.5 ADE hte M7 38.0 = 1.4 Sr sealed Rectrix 4 62.1—68.5 531.3—57.6 41.5-47.3 56.1 65:2. 12:3 54.3 2 0b:9 43.4 + 1.5 Rectrix 5 149.0—189.0 94.1—109.0 46.0-51.0 85.0 Lisa" = 104 YL is Meer | 4321 55 4: 4 Collected in Ecuador. VOLUME 110, NUMBER 2 65 Wing Length 5 10 15 20 Bill Length 28 27 26 25 Length Rectrix 1 24 23 22 * 10 15 20 Bill Length o)) Fig. 3. Sy 46 we Bill Length Length Rectrix 3 h ro) 200 % 150 100 Length Rectrix 5 50 45 50 55 60 6 Wing Length an Bivariate plots of mensural characters of males in definitive plumage: Lesbia victoriae (circles); Ramphomicron microrhynchum (diamonds); and their hybrid (triangle) (=Lesbia ortoni, AMNH 156651). Least squares regression lines and 95% confidence bands are illustrated for comparison. cation of the Lesbia parent. Lengths of the hybrid’s bill and wing chord exceed the cu- mulative range of those measurements for males of L. nuna and R. microrhynchum (Table 1). Morphological luxuriance has never been observed in trochiline hybrids; thus, L. nuna is not one of L. ortoni’s pa- rental species. On the other hand, measure- ments of the hybrid fell between the char- acter means of R. microrhynchum and L. victoriae. Of particular interest, bill and rec- trix 1 measurements exhibit negative allom- etry in the pooled sample, whereas those of the bill and rectrix 3 show positive allom- etry (Fig. 3). In both cases, hybrid values approximate those predicted by the least squares regression line. Geographic overlap.—The geographic and elevational ranges of the parental spe- cies, Lesbia victoriae and Ramphomicron microrhynchum, overlap broadly in the An- des (Fjeldsa & Krabbe 1990). Lesbia vic- toriae inhabits forest edge and dry brushy slopes at 2600—4000 m elevation, whereas R. microrhynchum prefers cloud forest edge and paramo in more humid regions at 1700-3400 m (Graves 1985, Fjeldsa & Krabbe 1990). Both species, particularly the former, are fairly common residents in the Quito region and were well represented in 318 19th century collections from Ecuador (Ob- erholser 1902, L6nnberg & Rendahl 1922, Chapman 1926). Acknowledgments I thank R. C. Banks, A. T. Peterson, T. S. Schulenberg, and R. L. Zusi for review- ing the manuscript. I am grateful to the cu- rators and staff of the American Museum of Natural History, New York, and The Natural History Museum, Tring, for per- mitting me to examine collections in their care. Mary LeCroy kindly researched the accession record for L. ortoni. Photographic prints were prepared by the Smithsonian photographic services. Travel was support- ed by the Research Opportunities Fund and the Department of Vertebrate Zoology, Smithsonian Institution. Literature Cited Berlioz, J., & C. Jouanin. 1944. Liste de Trochilidés trouvés dans les collections commerciales de Bogota.—Oiseau 14:126—155. Boucard, A. 1893. Genera of humming birds. Vol. 3, Part 2. Bournemouth, London. Chapman, E M. 1926. The distribution of bird-life in Ecuador.—Bulletin of the American Museum of Natural History 55:1—784. Collar, N. J., L. P Gonzaga, N. Krabbe, A. Madrojfio Nieto, L. G. Naranjo, T. A. Parker, II, & D. C. Wege. 1992. Threatened birds of the Americas: The ICBP/IUCN Red Data Book, 3rd edition, part 2. International Council for Bird Preser- vation, Cambridge, U.K., 1150 pp. Cory, C. B. 1918. Catalogue of birds of the Americas. Part 2, No. 1.—Field Museum of Natural His- tory Zoological Series 13:1—315. Elliot, D. G. 1879. A classification and synopsis of the Trochilidae.—Smithsonian Contributions to Knowledge, No. 317. Fjeldsa, J., & N. Krabbe. 1990. Birds of the high An- des. Zoological Museum, University of Copen- hagen, Denmark, 876 pp. Gould, J. 1861. An introduction to the Trochilidae or family of humming-birds. Published by the au- thor, London. Graves, G. R. 1985. Elevational correlates of specia- tion and intraspecific geographic variation in Andean forest birds.—Auk 102:556—579. 1990. Systematics of the “‘green-throated sunangels”’ (Aves: Trochilidae): valid taxa or PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON hybrids?—Proceedings of the Biological Soci- ety of Washington 103:6—25. . 1993a. Relic of a lost world: a new species of sunangel (Trochilidae: Heliangelus) from ““Bogota.’’—Auk 110:1-8. . 1993b. A new hybrid manakin (Dixiphia pi- pra X Pipra filicauda) (Aves: Pipridae) from the Andean foothills of eastern Ecuador.—Pro- ceedings of the Biological Society of Washing- ton 106:436—441. 1996a. Hybrid wood warblers, Dendroica striata X Dendroica castanea (Aves: Fringilli- dae: Tribe Parulini) and the diagnostic predict- ability of avian hybrid phenotypes.—Proceed- ings of the Biological Society of Washington 109:373-—390. 1996b. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 2. Hybrid origin of Erioc- nemis soderstromi Butler—Proceedings of the Biological Society of Washington 109:764-769. , & R. L. Zusi. 1990. An intergeneric hybrid hummingbird (Heliodoxa leadbeateri X Helian- gelus amethysticollis) from northern Colom- bia.—Condor 92:754—760. Greenway, J. C., Jr. 1978. Type specimens of birds in the American Museum of Natural History. Part 2.—Bulletin of the American Museum of Nat- ural History 161:1—305. Lawrence, G. N. 1869. Characters of some new South American birds, with notes on other rare or little known species.—Annals of the Lyceum of Nat- ural History 9:265—275. Lonnberg, E., & H. Rendahl. 1922. A contribution to the ornithology of Ecuador.—Arkiv for Zoologi 14:1-87. Meyer de Schauensee, R. 1947. New or little-known Colombian birds.—Proceedings of the Acade- my of Natural Sciences of Philadelphia 99:107— 126. 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., & E. Verreaux. 1876. Histoire Naturelle de Oiseaux-mouches ou Colibris, constituant la famille des Trochilidés. Part 3. Bureau de la So- ciété Linnéenne, Lyon. Oberholser, H. C. 1902. Catalogue of a collection of hummingbirds from Ecuador and Colombia.— Proceedings of the United States National Mu- seum 24:309-342. Peters, J. 1945. Check-list of birds of the world. Vol. 5. Museum of Comparative Zoology, Cam- bridge, Massachusetts, 306 pp. Salvin, O. 1892. Catalogue of the birds in the British Museum, Vol. 16. London, 703 pp. Sibley, C. G., & B. L. Monroe, Jr. 1990. Distribution and taxonomy of birds of the world. Yale Uni- VOLUME 110, NUMBER 2 versity Press, New Haven, Connecticut, 1111 Pp. Simon, E. 1921. Histoire naturelle des Trochilidae (synopsis et catalogue). Encyclopedia Roret, L. Mulo, Paris. Wilkinson, L. 1989. SYSTAT: the system for statis- tics. SYSTAT, Inc., Evanston, Illinois, 822 pp. Appendix General comparative description of definitive plum- ages of male Lesbia victoriae, Ramphomicron micro- rhynchum, and the hybrid, L. victoriae X R. micro- rhynchum (=Lesbia ortoni Lawrence, 1869; AMNH 156651). Descriptions of structural colors are unusu- ally subjective, as color seen by the observer varies according to the angle of inspection and direction of light. For this reason I use general color descriptions. Dorsal feathering (capital and spinal tracts) of vic- toriae to the upper tail coverts is medium dull green; feather bases are gray and some lateral barbs are nar- rowly fringed with buff. These plumage areas of mi- crorhynchum are deep purple; feather bases are gray separated from the purple terminal discs by a narrow greenish band. The dorsum of the hybrid appears an amalgam of victoriae and microrhynchum, more closely resembling the latter species. The greenish subterminal band of crown and back feathers of the hybrid is wider and the purple terminal disc narrower than in microrhynchum, imparting a mottled purple and green appearance to the crown, hindneck and back. Scapulars are mottled green and purple; the rump and uppertail coverts of the hybrid are more uniformly purple, whereas a few feathers on the sides of the lower back are tipped with green. Purple feather tips occur ventro-laterally to the auriculars, sides of the neck, and sides of the rump. In victoriae, a brilliant medium-green gorget ex- tends from the chin to the upper breast the posterior border of the gorget is broadly lanceolate in shape. Feathers of the lores, auriculars, sides of neck, breast, and flanks are green, finely margined (10 X magnifi- cation) with buff in fresh plumage; feathers along the midline below the gorget and on the abdomen are ex- tensively fringed with buff. Vent plumes are white; undertail coverts are buff with a muted and elongated grayish spot along the rachis; tibial plumes are buff. The ventral color and pattern of microrhynchum are similar to victoriae, but the lores and auriculars are rich purple, the gorget more rounded and proportion- ally smaller. Gorget color is light green, subtly paler than in victoriae. Feathers of the breast and sides of microrhynchum are narrowly margined with grayish- buff, especially on the abdomen and along the midline 319 below the gorget. Vent plumes in microrhynchum are white; undertail coverts are dark purplish brown, some tinted with green, and all broadly margined with buff or buffy—white; tibial plumes are brownish-black, nar- rowly tipped with buff. The hybrid’s gorget is light green, intermediate in color, size, and shape between those of the parental species. Barbs of auricular feathers are green tipped with purple. The lores are dull green with some purple reflections. The remainder of the underparts are inter- mediate in appearance between those of the parental species. Vent plumes are white, whereas the short tibial plumes are buff or light brown. Undertail coverts are buff with gray bases. In victoriae, the rectrices (dorsally) are black with brownish-purple reflections in bright light, conspicu- ously (rectrices 1—4) or inconspicuously (rectrix 5) tipped with dark green. The proximal % of the lateral vane of rectrix 5 is gray (dorsally) and grayish-white (ventrally). The proximal % of the rachis (rectrix 5) is grayish—white on the ventral surface. Tips of the out- ermost rectrices (5) of victoriae are slightly subspatu- late and “‘bowed”’ in cross-section. The rectrices of microrhynchum are black with bronzy—purple reflec- tions, especially on rectrix 1; the rachises are blackish— brown. In cross-section, the outermost rectrices (5) are nearly flat. The rectrices of the hybrid are intermediate between those of the parental species: (rectrix 1)—black with purplish tint proximally, shading to bronze, then cop- pery to coppery-purple at the tip; (rectrices 2—4)— black with bronzy reflections turning to coppery-pur- ple at the tip; (rectrix 5)—black with faint bronzy re- flections but lacking the coppery-purple tip present on the other rectrices. The lateral vane of rectrix 5 is mar- gined dorsally with buffy—white to within 25 mm of the feather tip. Ventrally, the rachis is white or very pale buff proximally, becoming dark brownish—black near the middle of the long axis of the feather. In cross- section, rectrix 5 of the hybrid is intermediate in cur- vature between that shown by victoriae and micror- hynchum. Primaries and secondaries are dull dark brown in victoriae, and blackish-brown with purplish reflections in microrhynchum. The flight feathers of the hybrid are intermediate in color and iridescence. In victoriae, the greater wing coverts are green (same as back), the pri- mary coverts are dark brown tipped with green, and the tiny coverts along the leading edge of the wing are green broadly edged with buff. Greater and primary coverts are blackish-brown in microrhynchum,; the leading edge coverts are brownish-black, some nar- rowly margined with light brown. Wing coverts of the hybrid are intermediate in color and pattern. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):320-—325. 1997. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 4. Hybrid origin of Calothorax decoratus Gould Gary R. Graves Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Calothorax decoratus Gould 1860 is shown to be a hybrid be- tween Acestrura heliodor and Acestrura mulsant. Plumage characters of the hybrid are a blended combination of those of the parental species. External measurements of the hybrid approximate the values expected from least squares regression of parental measurements. This is the first conclusively documented case of hybridization among the diminutive woodstars (Acestrura). The systematics and taxonomy of the Andean woodstars (Trochilidae: Acestrura) have yet to be resolved. In particular the validity of Calothorax decoratus Gould 1860 (hereafter Acestrura decorata), de- scribed from a unique specimen (No. 1888.7.25.203 in the Natural History Mu- seum (formerly British Museum [Natural History]), has been questioned since Salvin (1892: 408) pronounced it ‘“‘a species of doubtful value, intermediate between A. mulsanti and A. heliodor.’’ Since then, A. decorata either has been placed in the syn- onymy of A. heliodor (Cory 1918) or con- sidered as a doubtful species or hybrid (Berlioz & Jouanin 1944, Peters 1945, Meyer de Schauensee 1949, Morony et al. 1975). In this paper I confirm the hybrid origin of A. decorata using the methods outlined in Graves (1990) and Graves & Zusi (1990). Materials and Methods Species of Acestrura are sexually dimor- phic. The unsexed type of A. decoratus ap- pears to be in definitive male plumage and lacks striations on the ramphothecum at 10 xX magnification, indicating adulthood. Gould (1860) believed that the specimen was collected in Antioquia, Colombia. However, as pointed out by Berlioz & Jouanin (1944), specimens exported from ‘“‘Bogota”’ and other Colombian localities during the 19th century could have been collected nearly anywhere in northwestern South America. Assuming a hybrid origin of A. decorata, the pool of potential paren- tal species would include all the humming- birds known to occur in Colombia (Hilty & Brown 1986), and possibly Ecuador (Chap- man 1926, Fjeldsa & Krabbe 1990) and Venezuela (Meyer de Schauensee & Phelps 1978). The diminutive type of Acestrura decor- ata as noted by Gould (1860), Salvin (1892), and Hartert (1922) is, in fact, so similar in appearance to A. heliodor and A. mulsant that I focused my analysis on adult males of the five unquestioned species of Acestrura (heliodor, astreans, mulsant, ber- lepschi, bombus; see Graves 1986), and on other species of “‘woodstars’’ known from northwestern South America (MVyrmia mi- crura, Chaetocercus jourdanii, Philodice mitchellii, Calliphlox amethystina) (taxon- omy of Sibley & Monroe 1990). I com- pared the type of A. decorata directly with series of these species in the Natural His- tory Museum (BM[NH]), Tring. Photo- graphs, videotape, and notes later were compared with specimens in the American VOLUME 110, NUMBER 2 321 yams a T Tr SCART yy eee | * BL pumed . ‘ ee VOW asa 2 ri os a 7. ce qa Ventral and lateral views of male Acestrura heliodor (top), A. mulsant (bottom), and their putative hybrid, A. decorata (BM[NH] 1888.7.25.203; =Calothorax decoratus Gould). 322 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Ranges and means (+ standard deviation) of measurements (mm) of adult males of Acestrura heliodor, A. mulsant, and the hybrid, A. heliodor X A. mulsant [= A. decorata (Gould)]. Hybrid heliodor mulsant BM [NH] (n = 12) (n = 9) 1888.7.25.203 Wing chord 26.8—29.0 37.6—39.8 324 21S = 0.6 238) S278.8 Bill length 11.8—13.4 15.8-17.3 14.3 t25-2 05 16:3°20.5 Rectrix 1 8.3-9.2 14.8-16.9 11.8 8.8 + 0.4 15.6 + 0.6 Rectrix 2 10.8-13.4 18.0—20.6 13.8 jel ame 8 9 13.0 = 20.1 Rectrix 3 19.1-21.1 22.8—26.1 Za 200° "0:6 "2455212 Rectrix 4 17.8-19.8 22.8-25.4 DAe2 1I8.8= 0606, 243 = 08 Rectrix 5 12.7-14.7 19.3—21.6 oye) 13.6 = O16 ~~ 20:45 O:8 Museum of Natural History (AMNH), New York, and the National Museum of Natural History (USNM), Smithsonian Institution, Washington, D.C. Color descriptions were made under nat- ural light (Appendix). Measurements of wing chord, bill length (from anterior ex- tension of feathers), and rectrix length (from point of insertion of the central rec- trices to the tip of each rectrix) were taken with digital calipers and rounded to the nearest 0.1 mm (Table 1). Least squares re- gression lines and confidence intervals were projected on bivariate plots to illustrate size differences (Wilkinson 1989). The hybrid diagnosis was approached in a hierarchical manner. I hypothesized the presumed parental species of A. decorata through the comparative analysis of plum- age pattern and color, feather shape, and bill curvature. As a second step, the restrictive hypothesis was tested with the quantitative analysis of size and external proportions. Agreement of results is regarded as strong support for the hypothesis (Graves 1990, 1993, 1996; Graves & Zusi 1990). Results and Discussion Gould (1860:309) noted ‘“‘this species [Acestrura decorata] might easily be mis- taken for Calothorax [Acestrura] heliodori --- it differs from it in several particulars,— in being larger, in having the frill in front of the throat not so prolonged on the sides (in which respect it more nearly resembles C. mulsanti), the two centre tail-feathers finer and more spiny, and the bill much lon- ger.’’ Could a rare developmental or genetic aberrancy affecting size of A. mulsant or A. heliodor be responsible for the unique spec- imen of A. decorata? Neither dwarfism nor gigantism has ever been documented within the Trochilidae, and bona fide cases of those phenomena are very rare within birds (Buckley 1982). Details of the plumage col- or of A. decorata differ from that of both A. heliodor and A. mulsant (Fig. 1, Appen- dix), indicating that it is not a scale version of either species. Thus, we may conclude that A. decorata represents a hybrid or a valid taxon. My analysis presented below shows that the hybrid hypothesis cannot be refuted. Characters of Acestrura decorata rele- vant for hybrid diagnosis include: brilliant violet-purple gorget with elongated lateral feathers; white feathers on chin; grayish- white pectoral band below gorget; white feathers along midline of lower breast; lack of rufous pigment in rectrices; and widths of rectrices 4 and 5 <% those of rectrices 1-3. This combination of characters can only be derived from Acestrura heliodor and A. mulsant among the nine species (36 possible pairs of species) in the restricted pool (see Appendix). The plumage of A. de- corata is almost perfectly intermediate in appearance between A. heliodor and A. mulsant, whose geographic ranges and ele- vational distributions overlap in the Cordil- lera Central and Cordillera Oriental of the Colombian Andes. Hybridization between other species could not have produced the plumage char- acters present in the type of Acestrura de- corata. Chaetocercus jourdanii, Acestrura bombus, Myrmia micrura, Philodice mitch- ellii, and Calliphlox amethystina can be eliminated as possibilities because all pos- VOLUME 110, NUMBER 2 Bill Length 20 30 40 50 Wing Chord 30 Length Rectrix 1 & Rectrix 3 (top) = =) ibe) NO re) oO ‘o) Oo 12 14 16 18 20 22 Length Rectrix 5 325 30 25 20 15 Length Rectrix 1 & Rectrix 3 (top) 5 20 30 40 50 Wing Chord 30 25 20 15 10 Length Rectrix 1 & Rectrix 3 (top) Bill Length Fig. 2. Bivariate plots of selected measurements (see Table 1) of male Acestrura heliodor (diamonds), A. mulsant (filled circles), and their putative hybrid (triangle), A. decorata (BM[NH] 1888.7.25.203; =Calothorax decoratus Gould). sess rufous, buff, or cinnamon-colored pig- mentation on the rectrices and ventral plumage; A. decorata lacks any trace of such coloration. The geographic range of Acestrura astreans, which is restricted to the Sierra Nevada de Santa Marta, does not overlap the ranges of the other eight species in the restricted pool. Acestrura astreans also possesses a dark red gorget, which is not expressed in the type of A. decorata. The poorly-known woodstar A. berlepschi has a limited range in the lowlands of west- ern Ecuador (Collar et al. 1992) and co-oc- curs only with woodstar species possessing buff or rufous plumage characters (e.g., Myrmia micrura). These facts constitute ample reason for removing A. astreans and A. berlepschi from the list of potential pa- rental species. External measurements.—Measurements of Acestrura decorata fall between the character means of the hypothesized parents (Table 1, Fig. 2). Despite the significant dif- ference in size between the parental species (character means for A. mulsant are 22-— 77% larger than those for heliodor), both 324 species and A. decorata share unusual tail proportions where rectrices 1, 2, and 5, are substantially shorter than rectrices 3 and 4. This offers additional support for the con- clusion drawn from plumage pattern and color, that A. decorata is a hybrid between A. mulsant and A. heliodor, the first docu- mented case of hybridization involving spe- cies of Acestrura. As such, A. decorata (=Calothorax decoratus Gould) should be removed from the synonymy of A. heliodor (contra Cory 1918) and relegated to the growing list of documented trochiline hy- brids. Acknowledgments I thank Robert Pr¥s-Jones and Michael Walters for permitting me to examine the type of A. decorata in The Natural History Museum, and the curators and staff of the American Museum of Natural History for access to their collections. I appreciate Richard Banks, Robert Prys-Jones, Tom Schulenberg, Michael Walters, and Richard Zusi for commenting on the manuscript. I thank Harry Taylor of The Natural History Museum photographic services for taking photographs and making prints. Travel was supported by the Department of Vertebrate Zoology and the Research Opportunities Fund, Smithsonian Institution. Literature Cited Berlioz, J., & C. Jouanin. 1944. Liste des Trochilidés trouvés dans les collections commerciales de Bogota.—L Oiseau (n.s.) 14:126—155. 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, E M. 1926. The distribution of bird-life in Ecuador.—Bulletin of the American Museum of Natural History 55:1—784. Collar, N. J., L. PB. Gonzaga, N. Krabbe, A. Madrojio Nieto, L. G. Naranjo, T:. A. Parker III, & D. C. Wege. 1992. Threatened birds of the Americas: The ICBP/IUCN Red Data Book, 3rd edition, part 2. International Council for Bird Preser- vation, Cambridge, U.K., 1150 pp. Cory, C. B. 1918. Catalogue of birds of the Americas. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Part 2, No. 1.—Field Museum of Natural His- tory Zoological Series 13:1-315. Fjeldsa, J., & N. Krabbe. 1990. Birds of the high An- des. Zoological Museum, University of Copen- hagen, Denmark, 876 pp. Gould, J. 1860. Descriptions of twenty-two new spe- cies of humming-birds.—Proceedings of the Zoological Society of London 28:304—312. Graves, G. R. 1986. Systematics of the gorgeted woodstars (Aves: Trochilidae: Acestrura).— Proceedings of the Biological Society of Wash- ington 99:218-—224. 1990. Systematics of the “‘green-throated sunangels’’ (Aves: Trochilidae): valid taxa or hybrids?—Proceedings of the Biological Soci- ety of Washington 103:6—25. . 1993. A new hybrid manakin (Dixiphia pipra x Pipra filicauda) (Aves: Pipridae) from the Andean foothills of eastern Ecuador.—Proceed- ings of the Biological Society of Washington 106:436—441. 1996. Hybrid wood warblers, Dendroica striata X Dendroica castanea (Aves: Fringilli- dae: Tribe Parulini) and the diagnostic predict- ability of avian hybrid phenotypes.—Proceed- ings of the Biological Society of Washington 109:373-—390. , & R. L. Zusi. 1990. An intergeneric hybrid hummingbird (Heliodoxa leadbeateri < Helian- gelus amethysticollis) from northern Colom- bia.—Condor 92:754—760. Hartert, E. 1922. Types of birds in the Tring Museum. B. Types in the general collection. Trochili.— 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. Meyer de Schauensee, R. M. 1949. Birds of the Re- public of Colombia. Part 2.—Caldasia 5:381-— 644. , & W. H. Phelps, Jr. 1978. A guide to the birds of Venezuela. Princeton University Press, Princeton, New Jersey, 424 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. Peters, J. 1945. Check-list of birds of the world. Vol. 5. Museum of Comparative Zoology, Cam- bridge, Massachusetts, 306 pp. Salvin, O. 1892. Catalogue of the birds in the British Museum, Vol. 16. London, 703 pp. Sibley, C. G., & B. L. Monroe, Jr. 1990. Distribution and taxonomy of birds of the world. Yale Uni- versity Press, New Haven, Connecticut, 1111 Pp. Wilkinson, L. 1989. SYSTAT: the system for statis- tics. SYSTAT, Inc., Evanston, Illinois, 822 pp. VOLUME 110, NUMBER 2 Appendix Comparative descriptions of diagnostic plumage characters of adult male Acestrura heliodor, A. mul- sant, and their hybrid, BM[NH] 1888.7.25.203 (=Cal- othorax decoratus Gould 1860). Descriptions of struc- tural colors are unusually subjective, as color seen by the observer varies according to the angle of inspection and direction of light. For this reason I use general color descriptions. In heliodor, plumage of the capital and spinal tract is dark green, toned on the back, rump and uppertail coverts with bluish-green. Bluish reflections are slight- ly less evident on homologous plumage tracts in mul- sant and the hybrid. Gorget color in heliodor exhibits postmortem color change in specimens (Graves 1986), becoming pinker, less purple over time. The gorget color of the hybrid and 19th century specimens of heliodor and mulsant is pinkish-purple. Most specimens of heliodor possess a few buffy—white barbs on feathers at the anterior apex of the gorget. The pale chin spot of mulsant is significantly larger, whereas that of the hybrid is inter- mediate in size. The bases of gorget feathers in helio- i a) dor are grayish—buff separated from the pinkish-purple tip by a narrow transitional zone tinted green. Gorget feathers are patterned similarly in mulsant and in the hybrid, but the transitional bands are dark green and greenish—black, respectively. Lateral gorget feathers are much more elongated in heliodor (pinkish—purple tip, width = 1.6 mm, length = 4.7 mm) than in mul- sant (width = 2.8 mm, length = 2.1 mm); gorget di- mensions in the hybrid are intermediate (2.7 X 2.7). The underparts of heliodor (posterior to the gorget) are similar in pattern but significantly darker than those of mulsant (Fig. 1). The hybrid is intermediate in ap- pearance—in the darkness of the pectoral band, the width and intensity of the midline stripe, and in the distribution of white on the lower belly. Rectrix color of heliodor: rectrix 1 is dark green; rectrix 2 is black medially and blackish—green later- ally; rectrices 3—5 are black. Rectrix 1 of mulsant is dark green, the others (2—5) are black. Rectrices of the hybrid are intermediate in color: rectrix 1 is dark green with a blackish tip; rectrix 2 is black medially and blackish—green laterally; the outermost rectrices (3—5) are black. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1 10(2):326., 19873 INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE Applications published in the Bulletin of Zoological Nomenclature The following Applications were published on 20 December 1996 in Vol. 53, Part 4 of the Bulletin of Zoological Nomenclature. Comment or advice on any of 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, U.K. Case No. 2950 Pseudofoenus Kieffer, 1902 (Insecta, Hymenoptera): proposed designation of Foenus unguiculatus Westwood, 1841 as the type species. 2987 Geopeltis Regteren Altena, 1949, Geoteuthis Miinster, 1843, Jeletzkyteuthis Doyle, 1990, Loligosepia Quenstedt, 1839, Parabelopeltis Naef, 1921, Paraplesioteuthis Naef, 1921 and Belemnoteuthis montefiorei Buckman, 1880 (Mollusca, Coleoidea): proposed conservation. Gladiolites geinitzianus Barrande, 1850 (currently Retiolites geinitzianus; Graptolithina) proposed designation of a neotype. Trematospira Hall, 1859 (Brachiopoda): proposed designation of Spirifer multistriatus Hall, 1857 as the type species. Nothosaurus Minster, 1834 (Reptilia, Sauropterygia): proposed precedence over Conchiosaurus Meyer, [1833]. Opinions published in the Bulletin of Zoological Nomenclature The following Opinions were published on 20 December 1996 in Vol. 53, Part 4 of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be ob- tained free of charge from the Executive Secretary, I.C.Z.N., % The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. Opinion No. 1857. Metabla tothrix Sugonjaev, 1964 (Insecta, Hymenoptera): Blastothrix iso- morpha Sugonjaev, 1964 designated as the type species. 1858. Nectria Gray, 1840 (Echinodermata, Asteroidea): Nectria ocellata Perrier, 1875 designated as the type species. 1859. Nine specific names of southern Afrotropical birds conserved. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(2):327-—328. 1997. BIOLOGICAL SOCIETY OF WASHINGTON 124th Annual Meeting, 9 May 1997 President Stephen Cairns called the meet- ing to order at 11:00 a.m. in the Waldo Schmitt Room, National Museum of Natu- ral History. In summarizing the year’s ac- tivities, Steve noted that in addition to the Council meeting held annually just prior to the general meeting, the Council met on 11 February 1997 and voted to donate a com- plete set of the Proceedings to the Labato- rio de Invertebrados Bentonicos, Estacion Mazatlan, providing that institution would cover shipping and handling costs. The Council also authorized use of Society funds for compilation of an additional five sets of the Proceedings, which the Society hopes to sell for about $3,000 per set, plus postage. Wheldon & Wesley will advertise the availability of these sets, and receive 10% of profits from sales. In the future, the Society may not be able to compile addi- tional complete sets because many issues are missing, and the large amount of pho- tocopying necessary to produce a set is pro- hibitive. 7 Treasurer Chad Walter summarized Soci- ety finances for the period between 1 January and 31 December 1996. Total income was $72,372.30 ($40,130.10 from publication charges, $23,678.00 from dues and subscrip- tions, and $8,564.20 from sales of back issues and interest on Society accounts). Expendi- tures totalled $71,616.90 ($64,167.03 for publication costs, $7,118.70 for management costs, and $331.17 for bank charges). The net gain was $755.40 for this period (Table 1). The treasurer’s report also indicated that the number of Proceedings’ pages for which the Society received no remuneration was down in Volume 109 to 27.6%, vs. 45.6% in Vol- ume 108 and 42.1% in Volume 107. The President then summarized a report that Finance Committee Members Dick Banks and Austin Williams had presented to the Council. The report suggested that the Society (1) should look into better in- vestment of endowment funds to increase the rate of return (currently less than 5%); (2) is subsidizing too many pages in the Proceedings which is depleting the General Fund residual; (3) should change the by- laws to indicate that interest from the en- dowment fund automatically be re-invested into the endowment fund, and that proceeds from sales of back-issues of the Procedings be used for general operating costs (these are currently the practices, but by-laws mandate the opposite, i.e., that proceeds from sales of back-issues by deposted into the endowment fund, and earnings from the endowment be used in the general opera- tions of the Society); and (4) should adhere to the by-laws in the future by operating under an approved budget. Prior to the gen- eral meeting, the Council (1) voted to in- crease the amount authors are charged for a page in the Proceedings from $60.00 to $65.00, starting in 1988, (2) approved a motion allowing the Treasurer to continue current practices involving use of proceeds from sales of back-issues of the Proceed- ings until by-laws can be amended, and (3) approved a 1997 budget for the Society drafted by the Finance Committee. President Cairns thanked Dick Banks, Chad Walter, and Austin Williams for their extraordinary efforts in maintaining and evaluating Society finances. Council member Rafael Lemaitre then read the Editor’s report, which was pre- pared by Proceedings’ editor Brian Rob- bins. The report indicated that four issues of Volume 109 were published, comprising 70 papers and 770 pages. There were 80 submissions in 1996, down from the four 328 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Summary Financial Statement for 1996. General Fund Endowment Fund Total Assets ASSETS: JANUARY 1, 1996 30,356.86 T2477 106,078.63 TOTAL RECEIPTS FOR 1996 68,744.73* 31627 S74 12,372.38 TOTAL DISBURSEMENTS FOR 1996 71,616.90 00.00 71,616.90 ASSETS: DECEMBER 31, 1996 27,484.69 79,349.34 106,834.03 NET CHANGES IN FUNDS = 2 ONes |, 3:62 7.57 755.40 * A clause in the By-laws concerning the Endowment Fund states ““The annual earnings from this Fund (as well as the proceeds from the sale of back issues) shall (may) be used in the general operations of the Society.” The income from sales of back issues and bulletins inventory ($4,216) was not deposited into the Endowment Fund (Calvert Account) but into the General Fund (Riggs and Douglas Accounts). ** The interest generated by this Fund was not used. previous years (86—93). However, as of 1 The meeting was adjourned at 11:14. May 1997, there were 41 submissions, up from 34 in 1996 and 36 in 1995. There is Respectfully submittted, no current backlog for accepted papers. | Carole C. Baldwin Secretary 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 an Abstract in an alternate language when appropriate. Submission of manuscripts.—Submit three copies of each manuscript in the style of the Proceedings to the Editor, complete with tables, figure captions, and figures (plus originals of the illustrations). Mail directly to: Editor, Proceedings of the Biological Society of Wash- ington, National Museum of Natural History NHB-108, Smithsonian Institution, Washington, D.C. 20560. 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CONTENTS Review of the genus Schizopathes (Cnidaria: Antipatharia: Schizopathidae) with a description of a new species from the Indian Ocean Dennis M. Opresko Cnidae of Scleractinia Débora de Oliveira Pires Gnathostomulida from the Canary Islands Wolfgang Sterrer A new species of Stenoninereis (Polychaeta: Nereididae) from the Gulf of Mexico J. A. de Leén-Gonzdlez and V. Solis-Weiss Boguea panwaensis, a new species from Thailand: the first member of the Bogueinae (Poly- chaeta: Maldanidae) to be found outside northeast America C. Meyer and W. Westheide Triathrix montagni and T. kalki, a new genus and two new species of Cletodidae (Crustacea: Copepoda: Harpacticoida) from California and the Gulf of Mexico J. Michael Gee and Robert Burgess New records of the genus Hansenomysis in Japan with description of a new species (Crustacea: Mysidacea: Petalophthalmidae) Manuel Rafael Bravo and Masaaki Murano Nanomysis philippinensis, a new species (Crustacea: Mysidacea) from brackish waters of the Philippines Masaaki Murano A new species of bopyrid isopod, Pseudione chiloensis, a parasite of Nauticaris magellanica (A. Milne-Edwards, 1891) (Crustacea: Decapoda: Hippolytidae) Ramiro Roman-Contreras and Ingo Wehrtmann Observations on Hexapanopeus schmitti Rathbun from Brazil (Crustacea: Decapoda: Xanthi- dae) C. Sankarankutty and Raymond B. Manning Eunephrops luckhursti, a new deep-sea lobster from Bermuda (Crustacea: Decapoda: Nephro- pidae) Raymond B. Manning A new crayfish of the genus Orconectes from western Tennessee (Decapoda: Cambaridae) Christopher A. Taylor and Mark H. Sabaj A new crawfish of the genus Distocambarus, subgenus Fitzcambarus (Crustacea: Decapoda: Cambaridae) from South Carolina J. F. Fitzpatrick, Jr. and Arnold G. Eversole Neogonodactylus campi, a new species of stomatopod crustacean from the Caribbean Sea, with additional records for N. caribbaeus.(Schotte & Manning) Raymond B. Manning A new subgenus and four new species of Gliricola (Phthiraptera: Gyropidae) from Caribbean hutias (Rodentia: Capromyidae) Roger D. Price and Robert M. Timm Morphology, systematics, and distribution of Meoma ventricosa grandis and M. ventricosa ventricosa (Echinodermata: Echinoidea: Brissidae) along Mexican coasts Francisco A. Solis-Marin, Alfredo Laguarda-Figueras, and Antonio Leija-Tristan New species of Paratriacanthodes spikefish (Triacanthodidae: Tetraodontiformes) from the South China Sea James C. Tyler Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 3. Parentage of Lesbia ortoni Law- rence Gary R. Graves Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 4. Hybrid origin of Calothorax de- coratus Gould Gary R. Graves International Commission on Zoological Nomenclature Biological Society of Washington: 124th Annual Meeting 135% 167 186 198 203 210 227 236 242 249 256 263 212 280 285 301 SiO 314 320 326 327 B ‘tx -PROCEEDINGS _OF THE BIOLOGICAL SOCIETY : OF WASHINGTON | EWiTH OF lay 1; OCT 29 {997 THE BIOLOGICAL SOCIETY OF WASHINGTON 1996-1997 Officers President: Stephen D. Cairns Secretary: Carole C. Baldwin President-elect: Richard P. Vari Treasurer: T. Chad Walter Elected Council John A. Fornshell Rafael Lemaitre Alfred L. Gardner Diana Lipscomb Susan L. Jewett James N. Norris Custodian of Publications: Storrs L. Olson PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: Frederick M. Bayer. Invertebrates: Jon L. Norenburg Frank D. Ferrari Plants: David B. Lellinger Rafael Lemaitre Insects: Wayne N. Mathis Vertebrates: Gary R. 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Payment for membership is accepted in US dollars (cash or postal money order), checks on US banks, or MASTERCARD or VISA credit cards. Manuscripts, corrected proofs, editorial questions should be sent to: EDITOR BIOLOGICAL SOCIETY OF WASHINGTON NATIONAL MUSEUM OF NATURAL HISTORY WASHINGTON, D.C. 20560, U.S.A. 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 Periodicals postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, P.O. Box 1897, Lawrence, Kansas 66044. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):329-—331. 1997. Identification and restriction of the type locality of the Manzano Mountains cottontail, Sylvilagus cognatus Nelson, 1907 (Mammalia: Lagomorpha: Leporidae) Jennifer K. Frey, Robert D. Fisher, and Luis A. Ruedas (JKE LAR) Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, New Mexico 87131, U.S.A.; (RDF) Biological Resources Division, U.S. Geological Survey, National Museum of Natural History, Washington, D.C. 20560, U.S.A. Abstract.—The locality from which the holotype of Sylvilagus cognatus Nel- son, 1907 was collected is identified and restricted. Nelson (1907:82) reported the type lo- cality of Sylvilagus cognatus (currently re- garded as S. floridanus cognatus) as “10,000 feet altitude, near summit of Man- zano Mountains, New Mexico’’. Poole & Schantz (1942) subsequently reported the type locality as ““Tajique, near summit of Manzano Mountains, altitude 10,000 feet, Valencia Co., N. Mex.”’ The reference to Tajique as the specific locality where the holotype was collected has caused some question (Frey 1996). Tajique is a town at 2043 m on the eastern base of the Manzano Mountains located in the Tajique land grant, Torrance Co., New Mexico. In the south- western U.S., cottontail taxa often are as- sociated with distinctive vegetative com- munities. Accordingly, precise and accurate locations for type localities are important both to systematic and ecological studies of southwestern cottontails. Nelson (1907) reported the holotype of Sylvilagus cognatus as collected by A. Rea in February 1905 and deposited in the Bi- ological Survey Collection, USNM, as No. 136569; no additional specimens were mentioned. The USNM has three additional specimens of this taxon labeled as topo- types, also collected by A. Rea. Rea was not an employee of the Biological Survey of the museum and there are no field notes associated with these specimens. At that time, specimens received from non-employ- ees were catalogued first into an interme- diate catalogue (the “‘X catalogue’’) before formal cataloguing in the USNM catalogue. The four specimens of cognatus were X-ca- talogued on three separate occasions; there- fore, they either were received at different times or processed at different times: first a single entry, later another, and finally two together, including the holotype (see Table 1). All four were catalogued as being from *“Tajique, Manzano Mountains’’. The first entry, but not the later ones, has had added to it, in a different hand, “‘10,000 ft.’’ These four specimens were then entered into the USNM catalogue at two different times, a single specimen first, and then later a series of three. Again all were entered in this cat- alogue as being from ‘““Tajique, Manzano Mountains’’. The first entry has no eleva- tion; the series of three, which includes the holotype, all have ‘10,000 ft.”’ as part of the locality. Finally, all four specimen la- bels, which probably were produced at USNM rather than in the field, bear ‘**10,000 ft.’’ as part of the locality (vari- ances with respect to locality among cata- logues and specimen tags are summarized in Table 1). It thus would seem that Poole & Schantz (1942) included Tajique as part of the locality because it appears on all the labels and in both of the catalogues. Nelson (1907) may not have included Tajique in the original description because he had rea- 330 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Data recorded on the specimen tags, in the intermediate ‘‘X catalogue,’ ° and in the museum catalogue of the Division of Mammals, U.S. National Museum for the holotype and three topotypes of Sylvilagus cognatus. Source Date Locality Remarks X No. 4999 17 Jan 1905 New Mexico: Tajique 10,000 ft. (Manzano Mts.) topotype USNM 135755 17 Jan 1905 New Mexico: Tajique skin tag 17 Jan 1905 New Mexico: Tajique Manzano Mts. 10,000 ft. X No. 5190 7 Feb 1905 New Mexico: Manzano Mts. (Tajique) topotype USNM 136567 7 Beb 1905 New Mexico: Manzano Mts. Tajique (10,000 ft.) skin tag 7. Beb,1905 New Mexico: Manzano Mts. Tajique (10,000 ft.) X No. 5330 Feb 1905 New Mexico: Tajique, Manzano Mts. Topotype? USNM 136568 Feb 1905 New Mexico: Manzano Mts. Tajique (10,000 ft.) skin tag Feb 1905 New Mexico: Manzano Mts. Tajique (10,000 ft.) Xo No: 5331 Feb 1905 New Mexico: Tajique, Manzano Mts. Type? USNM 136569 Feb 1905 New Mexico: Manzano Mts. Tajique (10,000 ft.) skin tag Feb 1905 New Mexico: Manzano Mts. Tajique (10,000 ft.) 4 Remarked for both entries ‘‘Skulls may be mismatched; received without tags.”’ son to believe that these specimens were not actually from Tajique. In his revision of the North American rabbits, Nelson (1909) reported five specimens from the Manzano Mountains and repeated the type locality in the synonymy as it was reported in the orig- inal 1907 description. However, he later re- marked that “‘The topotypes of the present form from Tajique ranch [italics ours], at 10,000 feet altitude, near the highest part of the Manzano Mountains, vary [... ], buta specimen from a short distance away and lower down on the east slope of the extreme south end of the range is larger ... and agrees with those from Tajique’’ (Nelson 1909:192). We could not find ‘“Tajique Ranch”’ on any map or gazetteer. We were unable to find any correspon- dence concerning the actual place of cap- ture of the rabbits in the Smithsonian Insti- tution archives. However, Rea did send some miscellaneous bones to the Smithson- ian Institution that he had uncovered in a cave on (or near) his ranch and they were cataloged as being from “‘Manzano Mts.” or “‘Tajique, Manzano Mts.”’ Further, in 1904 Rea sent a snake (Pituophus) to the Smithsonian Institution that subsequently was cataloged as being from ‘“‘Tajique, ab. 10,000 alt’’. In a letter from A. K. Fisher to L. Stejneger in regards to this snake from Rea “‘captured near Tajique at about 10,000 ft. altitude,’’ Fisher, discussing the unusual capture, quotes from Rea “speaking gen- erally of rattlesnakes they are not found up here ... suppose the climate is not warm enough for them. Rea’s mailing address was given as ““Tajique, Valencia Co., New Mex- ico.”’ Torrance Co. was created from por- tions of Valencia and other counties on 16 March 1903 with additional portions of Va- lencia Co. (including most of the Manzano Mountains) added to Torrance Co. on 2 February 1905 (Coan 1965). Thus, it may be assumed that everything sent to the Smithsonian Institution by Rea was given the locality of Tajique because that was his mailing address rather than where the spec- imens were collected. James H. Gaut, an employee of the Bi- ological Survey, worked in the Manzano Mountains for three months during the fall and early winter of 1903 (Bailey 1928). Gaut’s field notes (Smithsonian Institution Archives Record Unit 7176) describe camps made “‘near Tajique, altitude 7800 feet.’’ Bailey (1928), in describing localities visited by Gaut describes Tajique, as “A VOLUME 110, NUMBER 3 town [italics ours] on the eastern slope of the Manzano Mountains, 30 miles southeast of Albuquerque; 7500 feet.’ This corre- sponds with the present town of Tajique and suggests that during the early 1900’s Ta- jique commonly referred to the town. Inter- estingly, Gaut’s field notes also describe a camp “‘at an elevation of 10,500 feet at Mr. A. Rea’s ranch on the summit of the moun- tain due west of Tajique.’’ The collector of the holotype and owner of this ranch prob- ably are one and the same. Further, Gaut certainly knew the difference between Ta- jique and Rea Ranch. He cataloged a Mus- tela and several Peromyscus as being from “Summit of Ridge, Rea’s Ranch, 10,000 ft.”’ as opposed to others from “‘East slope near Tajique, 8400 ft.’’. A map of the Cibola National Forest in the Manzano Mountains (USDAFS, 1938) labels a “house, cabin, or other building”’ symbol as “‘Rea.’’ We believe this refers to the ranch owned by A. Rea from which the type specimen of S. cognatus was collected and herein restrict the type locality of S. cognatus to the vicinity of Rea Ranch. It is located 1.9 km N and 13.4 km W of Tajique (TON, RS5E, NE1/4 of NW 1/4 Sec. 9, N34°45'05.39”, W106°25'18.04”) on the northeast side of Bosque Peak, at 2880 m (= 9450 ft.) elevation. Bosque Peak is the highest peak of the north end of the Man- zano Mountains; the relatively flat top of the peak covers approximately 2.5 km”. Vegetation in the vicinity of the restricted type locality (vegetative communities fol- low Dick-Peddie, 1993) includes Upper Montane Coniferous Forest (i.e., mixed co- niferous forest) characterized by Douglas- fir (Pseudotsuga menziesii) and white fir (Abies concolor), small patches of Subal- pine Coniferous Forest characterized by En- gelmann spruce (Picea engelmannii) and corkbark fir (Abies lasiocarpa var. arizoni- 331 ca), Subalpine-Montane Grassland, and Montane Scrub on dry, steep western slopes. The vegetative community in the vi- cinity of Rea Ranch is consistent with the observation that Sylvilagus floridanus is typically associated with montane forests in New Mexico. In contrast, Tajique, at a low- er elevation, is a transition between Colo- rado Pifion (Pinus edulis)-One-seed Juniper (Juniperus monosperma) Woodland, One- seed Juniper Savanna, and Plains-Mesa Grassland. Acknowledgments Partial financial support for this project was provided by a grant to T. L. Yates from the Biological Resources Division, U.S. Geological Survey. We thank M. A. Bogan, M. S. Burt, R. W. Dickerman, A. L. Gard- ner, L. R. Heaney, and R. P. Reynolds for helpful comments on earlier versions of this paper. Literature Cited Bailey, E M. 1928. Birds of New Mexico. New Mex- ico Department of Game and Fish, 807 pp. Coan, C. E 1965. The county boundaries of New Mexico. New Mexico Legislative Council Ser- vices, Santa Fe, 27 pp. Dick-Peddie, W. A. 1993. New Mexico vegetation; past, present, and future. University of New Mexico Press, Albuquerque, 244 pp. Frey, J. K. 1996. Mammalian type localities in New Mexico.—Occasional Papers, Museum of Southwestern Biology 7:1—21. Nelson, E. W. 1907. Descriptions of new North Amer- ican rabbits.—Proceedings of the Biological So- ciety of Washington 20:81-84. . 1909. The rabbits of North America.—North American Fauna, 29:1—314. Poole, A. J., & V. S. Schantz. 1942. Catalog of the type specimens of mammals in the United States National Museum, including Biological Surveys Collection.—Bulletin of the United States National Museum 178:1—705. U.S. Department of Agriculture Forest Service (US- DAFS). 1938. Cibola National Forest (Man- zano Division) New Mexico. Map. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):332—337. 1997. A new species of Tantilla (Serpentes: Colubridae) from northeastern Guatemala Jonathan A. Campbell and Eric N. Smith Department of Biology, The University of Texas at Arlington, Arlington, Texas 76019, U.S.A. Abstract.—Tantilla tecta is described from near Laguna Yaxha in north- eastern Guatemala. This species belongs to the taeniata group and is the only member of this group known from the Petén region. It is characterized by: a narrow pale middorsal stripe restricted to the vertebral scale row; a narrow pale lateral stripe on adjacent portions of the third and fourth scale rows; a broad pale collar that is uninterrupted medially; and 54 subcaudals in the single known specimen. It is most similar to T. jani of the Pacific versant of northern Middle America and to T. slavensi of the Tuxtla region of southern Veracruz, Mexico, but may be differentiated from these species by the condition of the pale lateral stripes on the body, the extent of the pale nuchal collar, the col- oration of the paraventral scale row, the number of ventrals and subcaudals, and relative tail length. | Resumen.—Se describe el colubrido Tantilla tecta de las inmediaciones de la Laguna Yaxha, Petén, Guatemala. Esta especie pertenece al grupo taeniata y es el Unico miembro del grupo conocido de la regi6n petenera. Se caracteriza por poseer: una linea dorsal angosta restringida a la hilera vertebral de escamas; una linea lateral palida en las areas adjacentes de las hileras de escamas tres y cuatro; un collar palido y ancho no interrumpido dorsalmente; y 54 subcaudales en el unico especimen conocido. Se parece mas a T. jani de la vertiente pacifica de Guatemala y Chiapas y a T. slavensi de la regi6n de Los Tuxtlas en el sur de Veracruz, México, pero se diferencia de estos por varias caracteristicas, incluyendo la condicion de la linea lateral palida del cuerpo, la extencidn del collar palido, la coloracién de la hilera de escamas paraventrales, el numero de escamas ventrales y subcaudales, y el largo relativo de la cola. The extensive lowlands of the Petén re- gion of Guatemala have been the focus of a number of herpetological investigations summarized by Campbell (1998), Duellman (1963), Lee (1980, 1996), and Stuart (1934, 1935, 1937, 1958). Relatively few species of reptiles probably remain to be discovered in this region, although our knowledge of the distributions and life histories for most Species remains poor. Nevertheless, certain small, secretive snakes are notorious for their ability to escape detection. Among these are members of the genus Tantilla, of which several species are widespread in the Petén region. The taeniata group of Tantil- la has a Middle American distribution and is composed of a dozen species, including the one herein described. The fact that half of the species in this group have been de- scribed since 1971, and that over half the species remain known from fewer than a dozen specimens, provides some indication of the difficulty of finding these snakes. It is likely that the combination of secretive habits, disjunct distributions, and perhaps rarity in nature contribute to the infrequen- cy with which these snakes are encoun- tered. A single specimen of the taeniata group was recently discovered near Laguna Yaxha VOLUME 110, NUMBER 3 in Petén, a locality from which no member of this group was known previously. We have compared this specimen with material housed in the University of Texas at Ar- lington and the University of Kansas col- lections, as well as carefully consulted the thorough descriptions provided by Pérez- Higareda et al. (1985), Savitsky & Smith (1971), Wilson (1982, 1983), and Wilson & Meyer (1971). This specimen cannot be al- located to any known species and we there- fore propose that this species be known as Tantilla tecta, new species Figs. 1-2 Holotype.—The University of Texas at Arlington (UTA) R-41160 (previously UVG 1742), an adult female from the slope flanking the NE side of Laguna Yaxha, Pe- tén, Guatemala (17°03'43”"N, 89°23'12”W). Collected by Cristian Granizo on 29 Jun 1992. This locality lies in Tropical Dry For- est at about 220 m. Diagnosis.—A small species of Tantilla of the taeniata group (sensu Wilson, 1983) that may be distinguished from all other members of the genus by having: a narrow pale middorsal line restricted to the verte- bral scale row; a narrow pale lateral line on adjacent portions of the third and fourth scale rows that extends onto the tail; a broad pale collar that is not interrupted me- dially; and 54 subcaudals in the single known specimen. In T. briggsi, T. cuesta, T. cuniculator, and T. tayrae the pale mid- dorsal line is absent or restricted to a few scales on the anterior portion of the body, and in 7. taeniata the middorsal stripe usu- ally is expanded laterally to include all of the vertebral scale row and adjacent por- tions of the paravertebral rows. Tantilla taeniata can also be distinguished from T. tecta by the pale coloration on the top of the head which is distinctly paler than the dark borders of the nuchal collar. The dor- sum of the head in T. fecta, in contrast, is about the same as the borders of the collar. Tantilla flavilineata, T. oaxacae, and T. re- 333 ticulata differ from T. tecta in having a broader pale lateral stripe located on the fourth dorsal scale row and adjacent por- tions of the third and fifth rows. Tantilla tecta differs from all species in the taeniata group except TJ. jani and T. slavensi in having a narrow pale middorsal line confined to the vertebral scale row. Tantilla jani differs from T. tecta in having less distinct pale lateral stripes that usually terminate on the posterior part of the body; a pale collar that includes the posterior por- tions of the parietals, posterior temporals, and ultimate supralabial; a pale postocular spot that includes the lingual margin of the fifth supralabial; the first pair of infralabials usually in contact; 37—47 subcaudals in fe- males; and a relative tail length from 15 to 18% of the total length (versus 23% in fe- male holotype of T. tecta). Tantilla slavensi may be distinguished from T. tecta in hav- ing part of the paraventral scale row pale (versus ground color from dorsum extend- ing onto ventrals); a pale nuchal collar that is interrupted medially (versus not inter- rupted) and no more than one scale in length (versus two); a pale lateral line that becomes obscure on the base of the tail (versus evident even on distal portion of tail); the pale lateral line with a narrow dark brown border above, but not distinctly bor- dered below (versus a distinct dark border below pale lateral line which is darker than the border above); and 158—159 ventrals in two known females (versus 148 in single female). A higher number of ventrals is present in females of 7. flavilineata (152- 164) and T. reticulata (162—173) than in T. tecta (148), whereas slightly lower numbers are present in females of 7. oaxaca (145), T. tayrae (140-146), and T. cuesta (147). A higher number of subcaudals is present in females of 7. taeniata (58-65) and T. reticulata (58-70), and fewer subcaudals are present in females of T. flavilineata (43—49), T. jani (37-47), T. striata (31-34), T. oaxacae (48), T. cuniculator (48-53), T. tayrae (44-46), and T. cuesta (45) than in T. tecta (54). Undoubtedly, as many of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rigi: Head length 6.7 mm from front face of rostral to posterior end of mandible. these species become known from more ad- equate samples, the ranges for some seg- mental counts will overlap. Description.—An adult female, 222 mm in total length; tail length 51 mm (23% of total); head length 6.7 mm from front face of rostral to posterior end of mandible; head width 4.3 mm at broadest point (at level of angle of mouth); head scarcely distinct from neck; snout truncate in dorsal view; eye small, snout about 2.4 times as long as horizontal distance across eye; pupil sub- circular; rostral about 1.6 times broader than high; internasals 1.7 times wider than long, laterally contacting anterior and pos- Dorsal (upper) and lateral (lower) aspects of the head of Tantilla tecta, holotype (UTA R-41160). terior nasals; prefrontals large, slightly wid- er than long, laterally contacting posterior nasal, prefrontal, and narrowly contacting second supralabial; median prefrontal su- ture 0.4 times as long as frontal; frontal about 1.3 times longer than wide; parietals about 1.8 times longer than wide, median suture about 0.9 of frontal length; nasals completely divided, nostril located in pos- terior portion of anterior nasal; no loreal; 1/1 preoculars; 2/2 postoculars; temporals 1 + 1, separating fifth, sixth, and seventh su- pralabials from parietal; supralabials 7/7, the first contacting nasals, the second con- tacting postnasal, prefrontal, and preocular, VOLUME 110, NUMBER 3 335 Fig. 2. the third contacting preocular, the third and fourth contacting orbit, the fifth and sixth contacting anterior temporal, and the sev- enth the largest and contacting anterior and posterior temporals; mental about twice as broad as long, contacting anterior pair of chinshields; anterior chinshields well de- veloped, about twice as long as wide; pos- terior chinshields well differentiated from gulars, about half of size of anterior chin- shields; posterior chinshields separated from first ventral by two gulars and two preventrals; infralabials 6/6, first four pairs contacting anterior chinshields, fourth pair largest; dorsal scales smooth, in 15 longi- tudinal rows throughout length of body, no apical pits apparent; dorsal scales in 4 rows Tantilla tecta, holotype (UTA R-41160), 222 mm TL (reproduced from UTA Slide No. 17660). at level of tenth subcaudal; ventrals 148; anal divided; subcaudals 54, paired; anal glands extending posteriorly the length of four subcaudals. In preservative (alcohol after formalin), ground color brown; a beige middorsal stri- pe arising about two scale lengths behind collar, extending most of the snout-vent length but becoming obscure on the poste- rior fourth of body; middorsal stripe occu- pying about median third of vertebral scale row, edged with dark brown laterally; beige lateral stripes on upper portion of scale row 3 and lower portion of scale row 4, edged with dark brown above and below, begin- ning about four scale lengths behind collar and extending to the tip of the tail; ground 336 color below lateral stripe a bit darker than that on either side of the middorsal stripe; dorsal ground color extending to lateral portions of ventrals and subcaudals, parav- entral scale row not pale; venter of body and tail cream-colored; top of head dark brown with pale spot on upper portion of rostral, internasals, and anterior two-thirds of prefrontals; a distinctive white nuchal collar immediately behind parietals and sec- ondary temporals, collar 1—2 scales long, bordered posteriorly with black, merging with pale coloration of venter, not inter- rupted middorsally, a small intrusion of nu- chal collar on lower posterior part of ulti- mate supralabial; a small, indistinct spot on adjacent portions of first and second su- pralabials; a prominent white postocular spot including most of fifth supralabial (but not lingual margin) and adjacent portions of lower postocular and anterior temporal; in- fralabials mostly pale except along lip mar- gin and posterior portion of individual scales, fourth supralabial also with a dark margin medially. The right maxillary bears 13 small teeth, which increase in size posteriorly; a small diastema separates the last two teeth which are enlarged and have lateral grooves. All maxillary teeth are compressed anteropos- teriorly and bladelike. Etymology.—The trivial name is derived from the Latin tectus, meaning concealed or secret, in allusion to the ability of this spe- cies to escape detection despite many years of field work in the region. Remarks.—The taeniata group of Tantil- la, as presently envisioned (Wilson 1983, Pérez-Higareda et al. 1985), consists of 12 species, including the species described here: briggsi, cuesta, cuniculator, flavili- neata, jani, oaxacae, reticulata, slavensi, striata, taeniata, tayrae, and tecta. Based on the presumably derived condition of the pale longitudinal stripes on the body, T. tec- ta appears to be most closely related to T. Jani and T. slavensi. Both T. jani and T. slavensi are geographically remote from T. tecta and occur in more mesic, less seasonal PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON habitats (Tropical and Premontane Wet For- ests). Tantilla jani is found at elevations of 305-960 m (Wilson 1985) along the Pacific versant of Middle America from eastern Oaxaca, Mexico, into Guatemala. Tantilla slavensi occurs in the Los Tuxtlas region of southern Veracruz, Mexico, over 600 km to the WNW of the type-locality for 7. tecta, and is known from 50—800 m (Pérez-Hi- gareda et al. 1985). Although the holotype of Tantilla tecta comes from an area usually considered to be covered with Tropical Dry Forest, Laguna Yaxha is often thought to be near the boundary between Tropical Dry and Tropical Moist Forest, with the latter extending to the south shore of the lake. Certainly, the demarcation between these two types of forest is not clear and many of the hillsides and depressions to the north of the lake are covered with patches of for- est that might be associated with Tropical Moist Forest. Nevertheless, the entire re- gion around Laguna Yaxha experiences a highly seasonal climate with most of the precipitation falling from May to December and a dry season extending from January through April. The only species of Tantilla with which T. tecta is possibly sympatric in the Petén faunal area of Campbell & Vannini (1989) are 7. moesta, T. cuniculator, and T. schis- tosa. Tantilla moesta has a long pale nuchal collar that extends posteriorly from the pa- rietals for a length of at least three dorsal scales and usually includes the posterior- most three supralabials, the dorsum is uni- formly dark brown or black without any longitudinal striping, and the venter is uni- formly dark. Tantilla cuniculator has only the slightest indication of (or completely lacks) a pale middorsal stripe, the pale lat- eral stripe is less distinct than in T. tecta, the pale nuchal collar is usually broader, and the pale postocular spot usually in- cludes the lingual margin of the fifth supra- labial. Tantilla schistosa has a uniformly colored dorsum without longitudinal stripes and 24-40 subcaudals. Tantillita canula and T. lintoni also occur in Petén, but they VOLUME 110, NUMBER 3 are easily distinguished from T. tecta in lacking pale nuchal collars and pale lateral stripes, and having the middorsal stripe (if present) ill-defined and confined to the pos- terior part of the body and the tail. Acknowledgments We thank Michael Dix of the Universi- dad del Valle de Guatemala (UVG) for al- lowing us to report on the material in his care. Permits for conducting research in Guatemala were granted by officials of the Consejo Nacional de Areas Protegidas (CONAP); we are especially grateful to Lic. Oscar E Lara and Licda. Mygdalia Garcia de Soldorzano. This material is based in part upon work supported by the Texas Ad- vanced Research Program under Grant No. 003656-001 to JAC. Literature Cited Campbell, J. A. 1998. The Amphibians and Reptiles of northern Guatemala, Yucatan, and Belize.— The University of Oklahoma Press, Norman, Oklahoma, in press. , & J. P Vannini. 1989. Distribution of am- phibians and reptiles in Guatemala and Be- lize.—Proceedings of the Western Foundation of Vertebrate Zoology 4:1—21. Duellman, W. E. 1963. Amphibians and reptiles of the rainforests of southern El] Petén, Guatemala.— University of Kansas Publications, Museum of Natural History 15:205—249. Lee, J. C. 1980. An ecogeographic analysis of the herpetofauna of the Yucatan Peninsula.—Mis- 337 cellaneous Publications, Museum of Natural History, Univiversity of Kansas 67:1—75. 1996. The Amphibians and Reptiles of the Yucatan Peninsula. Cornell Univiversity Press, Ithaca, New York. 500 pp. Pérez-Higareda, G., H. M. Smith, & R. B. Smith. 1985. A new species of Tantilla from Veracruz, Mexico.—Journal of Herpetology 19:290—292. Savitsky, A. H., & H. M. Smith. 1971. A new snake from Mexico of the taeniata group of Tantil- la.—Journal of Herpetology 5:167-171. Stuart, L. C. 1934. A contribution to a knowledge of the herpetological fauna of El] Peten, Guatema- la.—Occasional Papers of the Museum of Zo- ology, University of Michigan 292:1-18. . 1935. A contribution to a knowledge of the herpetology of a portion of the savanna region of central Petén, Guatemala.—Miscellaneous Publications, Museum of Zoology, University of Michigan 29:1-—56. . 1937. Some further notes on the amphibians and reptiles of the Peten forest of northern Gua- temala.—Copeia 1937:67-—70. 1958. A study of the herpetofauna of the Uaxactun-Tikal area of northern E] Peten, Gua- temala.—Contributions from the Laboratory of Vertebrate Biology, University of Michigan 75: 1-30. Wilson, L. D. 1982. A review of the colubrid snakes of the genus Tantilla of Central America.—Mil- waukee Public Museum Contributions in Biol- ogy and Geology 52:1-77. . 1983. A new species of Tantilla of the taen- iata group from Chiapas, Mexico.—Journal of Herpetology 17:54—59. . 1985. Tantilla jani (Giinther).—Catalogue of American Amphibians and Reptiles, Society for the Study of Amphibians and Reptiles, 369.1. ., & J. R. Meyer. 1971. A revision of the taen- iata group of the colubrid snake genus Tantil- la.—Herpetologica 27:11—40. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):338-—365. 1997. Geographic variation in the frog genus Vanzolinius (Anura: Leptodactylidae) W. Ronald Heyer Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Vanzolinius discodactylus, a forest-dwelling frog species of west- ern Amazonia in South America, varies in characters of color pattern, mor- phology, and advertisement call. Analysis of this variation indicates that very local (site) differentiation results in mosaic patterns of differentiation, largely obfuscating larger geographic patterns. Comparison of available genetic esti- mates of differentiation for V. discodactylus are consistent with the morpho- logically and advertisement call-based conclusions. A previously studied forest- dwelling lizard and another forest-dwelling frog also demonstrate local differ- entiation patterns suggesting that the variation in V. discodactylus may repre- sent a general pattern for forest-dwelling amphibians and reptiles in Amazonia. During examination of specimens for a study of Leptodactylus species (Heyer 1994), several Vanzolinius specimens were encountered. Dr. Claude Gascon found Vanzolinius to be relatively common along the Rio Jurua in Brazil and used the species to test the riverine barrier hypothesis (Gas- con et al. 1996). A cursory examination of these additional materials suggested that there was considerable variation, which might profitably be studied. The purpose of this paper is to analyze geographic variation in Vanzolinius. Materials and Methods As many adults, larvae, and recordings of advertisement calls as possible were as- sembled from major museum collections (Appendix 1). The sex of individuals was determined either by examination of vocal slits, or dis- section to examine gonads. The following categories are used: adult male—vocal slits present; juvenile male—testes present, but vocal slits not broken through; adult fe- male—oviduct folded at least in part; ju- venile female—ovaries present, but oviduct straight; juvenile—condition of gonads in- determinate (in some cases, gonads had been removed by previous workers). Analyses differ depending on the type of data gathered for the characters examined. The following descriptions of characters are arranged by analytical groups. Color patterns and external morpholog- ical features of adult form individuals.— These qualitative traits are categories re- corded as either binary or multistate char- acters. For the latter, states were added to the series as they were encountered during the data-taking phase. The states within each series have no intended or implied re- lationships or transformation series. Inter- mediate conditions between states were re- corded with the first letter of the state that most nearly approached the condition ob- served in the specimen examined. Dorsal snout pattern: Three basic states were encountered: a relatively uniform dark pattern (Fig. 1A); a variegated pattern (Fig. 1B); and a uniform light pattern (Fig. 1C). Light postorbital eye stripe: A series of symbols define the distinctiveness of the postorbital eye stripes: — [absent]; (+) [in- distinct]; + [distinct]; +! [sharply defined]. VOLUME 110, NUMBER 3 339 Fig. 1. Dorsal snout pattern standards. In cases where the two sides of the head differed, both conditions were recorded. Light subocular bar: The distinctiveness of the bar was noted by the same symbols as for the previous character, except the +! category was not encountered. Dorsal pattern: Dorsal pattern variation forms a continuum among the more dis- tinctive states recorded. The states recog- nized are: State A—either an uniform dor- sum (brown or tan) or indeterminately blotched (Fig. 2A); State B—the dorsum with very distinct dark markings in the in- terorbital and interscapular areas (Fig. 2B); State C—distinct interorbital blotch, well defined chevron markings anteriorly and blotches posteriorly on the body (Fig. 2C); State C-1—as previous state except chev- rons continuous; State D—a distinct darker straight edged band extending from behind eyes on full extent of dorsum (Fig. 2D); State D-1—as previous state, except sides irregular. Dark mid-dorsal pin stripe: An interrupt- ed dark mid-dorsal pin stripe was recorded as either present or absent. Throat and chest pattern.—Variation in this character is continuous among the states encountered: State A—-variegated pattern (Fig. 3A); State A-l—as previous state, but light; State B—uniform light pat- tern (Fig. 3B); State B-1—as previous state, but lateral portions darker; State C—dark speckled pattern (Fig. 3C); State C-1—as previous state, but dark spotting more ex- tensive; State D—dark pattern (Fig. 3D); State E—dark pattern with light spots (Fig. SE). Belly pattern: Variation in this character is continuous among the states encountered: Fig. 2. Dorsal pattern standards. 340 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Se ee Fig. 3. Throat and chest pattern standards. State A—speckled pattern (Fig. 4A); State A-1—almost uniform cream pattern; State B—indistinctly mottled, more intense an- teriorly (Fig. 4B); State B-1, as previous State, but lighter; State C—distinctly mot- tled, rather uniform over belly (Fig. 4C); State C-l—as previous state, but lighter; State C-2—as State C but dark anteriorly and no melanophores posteriorly; State D— distinctly variegated dark and light pattern (Fig. 4D). Posterior thigh pattern: Variation in this character is continuous among the states en- countered: State A—indistinctly mottled (Fig. 5A); State B—indistinctly mottled with indistinct dark longitudinal band (Fig. 5B); State C—distinctly mottled (Fig. 5C); State D—speckled with indistinct dark lon- gitudinal band (Fig. 5D); State E—speckled with distinct dark longitudinal band (Fig. 5E); State F—speckled with dark longitu- dinal band bordered above by light longi- VOLUME 110, NUMBER 3 341 Fig. 4. Belly pattern standards. tudinal stripe (Fig. 5F); State F!—as for State F except light stripe very distinct. Outer tarsal pattern: Data were taken on the distinctiveness of the outer tarsal pattern relative to the dorsal tarsal pattern. How- ever, variation turned out to be minimal and scoring could not be done consistently. These data are not analyzed further. Dorsolateral fold condition: There is rel- atively little variation in this character and the variation that exists is difficult to eval- uate in terms of the impact preservation has on recognition of fold condition. Most in- dividuals have no dorsolateral folds. In a few individuals, a short ridge or elongated warts lie in the dorsolateral fold region pos- terior to the eye. The variation in this char- acter is not analyzed further. Male secondary sexual characters: All males lack secondary sexual characters of thumb or chest pads or spines or male arm hypertrophy as found in Leptodactylus. Male vocal sac: Variation in this char- acter is minimal and difficult to evaluate in terms of preservation artifact. In most males, the vocal sac is single and internal; in a few males, the single vocal sac has ex- ternal indications of weak lateral folds. Variation of this character is not analyzed further. Textures: Data were taken on textures of the dorsum, the upper shank, the outer tar- sus, and sole of foot. In all cases the degree of development of shagreen and tubercles was difficult to categorize consistently and differentiate from preservation artifact. The Bis) 5: Posterior thigh pattern standards. 342 A B C Fig. 6. Digit tip dorsal outline standards. dorsum is tuberculate, with either small black-tipped or white-tipped tubercles, usu- ally more densely packed posteriorly. The dorsum may also be somewhat granular, have a shagreen, or be smooth. The upper shank and outer tarsus consistently have tu- bercles, black and/or white tipped, and the surfaces may also be shagreened. The foot is either smooth or the outer margin has a few black/white tipped tubercles and/or a shagreen. Variation for these characters is not analyzed further. Finger tip dorsal outline: For both finger and toe tip shapes, the outline shapes stan- dardized by Savage (1987) for Eleuthero- dactylus were used. For both finger and toe tip dorsal outlines, only three of Savage’s (1987) shapes were encountered corre- sponding to his unexpanded even (Fig. 6A), expanded even (Fig. 6B), and expanded pointed or lanceolate (Fig. 6C) states. The third finger tip is the most expanded and is the digit from which data were taken. Very little variation was encountered, suggesting that observer variation in interpreting shape was probably as large as actual variation. The conditions recorded ranged from not expanded, just A, A, A-B, or A-C. Variation is not analyzed further for this character. Toe tip dorsal outline: The same stan- dards were used for toe tips as for finger tips (Fig. 6). Data were recorded for both the third and fourth toe tips. The conditions for the third and fourth toe dorsal outlines are very similar. For 165 individuals, the conditions are identical, for 9 individuals the fourth toe dorsal outline is perceptibly more expanded than for the third, and for PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 75 individuals, the third toe dorsal outline is perceptibly more expanded than the fourth. As the third toe tip is a bit more expanded in the total sample, data are an- alyzed only for the third toe tip. Dorsal toe grooves: The dorsal surfaces of the toes typically have from 2—5 grooves involving the epidermis and dermis. The grooves are almost parallel to each other along the long axis of the toe, but radiate slightly outward from the proximal toe tip to the distal tip. Usually the grooves extend almost the entire length of the expanded portion of the toe tip, but the grooves do not reach the tip of the toe. Counting the exact number of grooves is not always pre- cise as the grooves are sometimes incom- plete and preservation artifact can obscure the definition of the grooves. Data were taken for both the third and fourth toes. As for the dorsal outline, the conditions within individuals are similar, but typically the third toe has one more groove than the fourth. For 96 individuals, the third and fourth toes have the same number of grooves, for 18 individuals, the fourth toe has more grooves than the third, and for 132 individuals the third toe has more grooves than the fourth. Because the raw data indicate that the variation in the fourth toe mirrors that seen in the third, only the data for the third toe are analyzed further. Analysis of preceding characters: The preceding characters are recorded as dis- crete entities even though variation is main- ly continuous. Because the data are discrete, chi-square analyses are used to determine whether occurrence frequencies of states differ significantly. The 0.05 convention is used to determine significance. Data were adjusted, when necessary, to reach a mini- mum cell size of an average expected fre- quency of 5 (Hayek 1994:239). Data were first examined to determine whether states for adult males and females differed signif- icantly. If they did not, then data recorded for juveniles were added to both the male and female data to provide more robust data sets for statistical analysis among geograph- VOLUME 110, NUMBER 3 ic regions (see definitions of regions be- low). Measurement data and analyses.—Mea- surements were taken on the following vari- ables, as defined by Gascon et al. (1996): snout-vent length (SVL), nostril separation, eye width anterior, eye width posterior, head width, head length, eye to nostril dis- tance, thigh length (=femur length of Gas- con et al. 1996), shank length (=tibia length of Gascon et al. 1996), foot length, tym- panum diameter (=tympanum height of Gascon et al. 1996), eye length, maximum width of third finger, and maximum width of fourth toe. Measurements were taken with a Helios dial calipers and recorded to the nearest 0.1 mm. Only adult specimens are used for the measurement data analyses. As males and females are sexually dimorphic in size, they are analyzed separately (L. C. Hayek, C. Gascon, and W. R. Heyer (unpublished data) discuss multivariate analyses on mor- phometric data on Vanzolinius.) The data are analyzed using the software program SYSTAT 5 (Wilkinson et al. 1992). Larval data.—To my knowledge, the only larvae available are those reported on by Duellman (1978) from a single locality in Ecuador (Mera, Pastaza). Specimens KU 121362—121363 are larvae ranging from Gosner stages 30-38. Specimens KU 121360—121361 are just metamorphosed individuals. There is no internal evidence from study of these specimens to either es- tablish that they are Vanzolinius or they are not. Dr. John Lynch collected the specimens and informs me (pers. comm.) after he con- sulted his field notes, “‘... it appears that I guessed on the identification ... on the basis of habitat selection. Hence, don’t trust the identification.’’ As nothing can be de- termined about geographic variation based on these specimens (even assuming they are Vanzolinius discodactylus), larvae are not treated further in this paper. Advertisement call data and analyses.— Recordings of single individuals from five localities are available for analysis. 343 Brazil: Acre; Nova Vida, Rio Jurua, USNM Tape 256 Cut 12. Recorded at 1900 h on 17 March 1992 by Claude Gascon at a temperature of 25°C, no voucher speci- men. Brazil: Amazonas; Altamira, Rio Jurud, USNM Tape 255 Cut 2. Recorded at 1915 h on 17 November 1991 by Claude Gascon at a temperature of 25°C, voucher INPA 5021. Brazil: Amazonas; Barro Vermelho, Rio Jurua, USNM Tape 254 Cut 5. Recorded at 1900 h on 27 October 1991 by Claude Gas- con at a temperature of 24.4°C, voucher INPA 3352. Brazil: Amazonas; Jainu, Rio Jurud, USNM Tape 254 Cut 13. Recorded at 1740 h on 2 November 1991 by Claude Gascon at a temperature of 26.1°C, no voucher specimen. Ecuador: Napo; Limoncocha, USNM Tape 18 Cut 1. Recorded at 2000-2034 h on 9 July 1971 by Ronald Heyer at an air temperature of 23.4°C, water temperature 23.6°C, voucher LACM 92001. Calls were analyzed using Canary 1.2 software (Charif et al. 1995) on a Power Macintosh 8500 computer. Calls were dig- itized for analysis at a sample rate of 22050 Hz and a sample size of 16 bits. Call rate was determined directly from recordings for periods ranging from 45 to 180 s per re- cording. Other call parameters were taken from a combination of waveform, audios- pectrogram (=spectrogram as used in Ca- nary manual), and spectrum analyses based on ten calls for each individual. Most of the recordings had considerable noise. Many parameters were taken from filtered calls. The filter around option was used for de- termining some parameters for USNM Tape 256 Cut 12 (filtered around 520—5000 Hz), USNM Tape 255 Cut 2 (filtered around 500-5000 Hz), USNM Tape 254 Cut 5 (fil- tered around 330—4000 Hz), and USNM Tape 18 Cut 1 (filtered around 500—4500 Hz). Definition of geographic areas for anal- ysis.—The primary purpose of this study is 344 to determine the nature of geographic vari- ation found within Vanzolinius discodacty- lus. Sample sizes are insufficient to analyze the data from each locality independently. Localities were plotted on a map and lo- calities were grouped on the basis of geo- graphic proximity (Fig. 7). The rationale used for grouping localities involved trying to maximize three criteria simultaneously: to have as many groups as possible in order to characterize geographic variation; to have as many individuals as possible in each group to permit robust statistical anal- yses; and to maintain geographic integrity. With respect to geographic integrity, a rule of thumb of keeping localities within the Same major river drainage basins was gen- erally applied (Areas A, B, C, D, E G, H), but not exclusively so (Area E). Initially 10 geographic area groupings were made. When the data were examined for these groupings to see if they were sufficient, three of the groups lacked sufficient data for analysis. Two Colombian localities were sufficiently isolated from each other as well as other samples to be placed in their own groups; unfortunately, the specimens from both localities are faded such that data are incomplete for them. Thus, data for the Co- lombian localities of Caldas; Villa Maria and Caqueta; Florencia, are not included in the geographic analyses (Fig. 7, upper two Squares). A single Peruvian locality (Uca- yali, Yarinacocha) also formed a distinct geographic group by itself and contains one faded specimen, unsuitable for further geo- graphic analysis (Fig. 7, lower square). Eight geographic groupings remain and are identified by letter in further discussion: (A) northern Amazonian Ecuador; (B) southern Amazonian Ecuador; (C) Amazonian Peru; (D) the Brazil-Colombia border region; (E) easternmost known localities for Vanzoli- nius in Amazonian Brazil; (F) the mid-re- gion of the Rio Jurua of Brazil; (G) the up- per region of the Rio Jurua in the Brazilian State of Amazonas; and (H) the upper re- gion of the Rio Jurua in the Brazilian State of Acre. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON As there are but five individuals available from Region D, (1 female, 1 male, 3 juve- niles), Region D data are omitted from the analyses by areas, unless otherwise noted. Results Dorsal snout pattern.—This was the only character for which some individuals clear- ly demonstrated two states (this exception involved only conditions B and C both oc- curring in the same individual), suggesting partial independent genetic control of this character. The states (Appendix 2) were collapsed for analysis to three: (1) pure A, (2) any B, and (3) any C. Because several individuals had both states (2) and (3), the total number of state conditions analyzed exceeds the number of individuals exam- ined for this character only. The chi-square analysis by sex was not significant (x7 = 1.35; df = 2; P = 0.50 > 0.30). Thus, male, female, and juvenile data were combined to analyze by geographic area. The chi-square analysis by geographic area is significant (y7 = 69.92; df = 12; P < 0.001). In partitioning the results, regions A+B are distinct from regions E+ F+G+H. Region C is not distinct from either of the other two area groupings. Light postorbital eye stripe.—For indi- viduals having different states on either side of the head, the more distinctive state was scored for statistical analysis (e.g., an in- dividual recorded as having state (+) on one side of the head and + on the other was treated as having the + state for statis- tical analysis). The chi-square analysis by sex was significant (x7 = 6.43; df = 2; P = 0.05 > 0.02); therefore the variation among geographic areas has to be analyzed sepa- rately by sex. For females, the chi-square analysis by geographic area is significant (Vy? = 42.43; df = 12; P < 0.001). In par- titioning the significance, regions B+E are distinct from regions A+C+F+G+t+H. In order to meet the minimum expected cell size criterion for statistical robustness for males, the data had to be collapsed by rec- VOLUME 110, NUMBER 3 345 Fig. 7. Map of northwestern South America showing known distribution of Vanzolinius discodactylus. Guy- ana and Brazil are truncated by the 60°W meridian; Bolivia and Brazil by the 15°S parallel. Squares are single localities excluded from analysis of geographic variation. Circles and ellipses labelled A—H indicate groupings of localities (dots) used for analysis of geographic variation. A dot may represent more than one locality. 346 ognizing but two character states: absent (— state) or present (combined (+), +, +! states). The chi-square results are signifi- cant (x7 = 27.80; df = 6; P < 0.001). In partitioning the results, the significance is due entirely to the distinctiveness of the Area B specimens. Light subocular bar.—For individuals having different states on either side of the head, the more distinctive state was scored for statistical analyses. The chi-square anal- ysis by sex was significant (x? = 6.85; df = 2; P = 0.05 > 0.02). In order to meet the assumption of minimum expected cell size for females, the data for Area H were deleted. The resultant chi-square analysis is not significant (x7 = 15.49; df = 10; P = 0.20 > 0.10). To meet the minimum ex- pected cell size for the male data, the states were collapsed to two: absent (— state) and present ((+) and + states). The resultant chi-square analysis is not significant (7? = 9.59; df = 6; P = 0.20 > 0.10). Dorsal pattern.—For statistical analyses, the first state recorded for intermediate con- ditions is used (e.g., a specimen recorded as having condition A—B is scored as having condition A for the statistical tests). The chi-square analysis by sex is not significant (7 = 2.93; df = 3; P = 0.50 > 0.30); fe- male, male, and juvenile data were com- bined to analyze by geographic area. To meet the assumption for minimum expected cell size, States D and D-1 are combined; however, Area D is included. The chi- Square result among areas is significant (y7 = 203.09; df = 28; P < 0.001). In parti- tioning the results the following area group- ings are distinct from each other: A; B; C+D+E; F+G+tH. Dark mid-dorsal pin stripe.—The chi- Square analysis by sex is significant (;* = 5.86; df = 1; P = 0.02 > 0.01). To meet the minimum expected cell size criterion for females by area, Area H was deleted. The chi-square result is significant (x? = 11.47; df = 5; P = 0.05 > 0.02). Partitioning the results indicates the following area group- ings are distinct from each other: A+B; PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON C+E+F+G. For males, the chi-square analysis of character states by area is not significant (x* = 9.43; df = 6; P = 0.20 > 0.10). Throat and chest pattern.—As for dorsal pattern, the first state recorded for inter- mediate conditions is scored for the statis- tical analyses. State E was scored for only one individual and is not considered in the analyses. As only two known-sex individ- uals have State B-1, and only one known- sex individual has State C-1, State B-1 is combined with State B and State C-1 is combined with State C in the analyses. The chi-square analysis by sex is significant (j = 23.20; df = 4; P < 0.001). To satisfy the minimum expected cell frequency assump- tion for females, drastic manipulations are required. Areas B and H are excluded. States A-1 and A are combined. The resul- tant chi-square analysis is significant (77 = 23.39; df = 8; P = 0.01 > 0.001). In par- titioning the results, only areas E+F are dis- tinct. Drastic manipulations are also re- quired to analyze the male variation. States C and C-1 are deleted. Areas C, E, G, and H are deleted. States A-1 and A are com- bined. The resultant chi-square test is not significant (x? = 1.90; df = 4; P = 0.90 > 0.50). Belly pattern.—As for dorsal pattern, the first state recorded for intermediate condi- tions is scored for statistical analyses. To meet the minimum expected cell frequency assumption for the analysis by sex, the fol- lowing states are combined: A and A-1; B and B-1; C, C-1, and C-2; State D is delet- ed. The resultant chi-square analysis is sig- nificant (x7 = 8.60; df = 2; P = 0.02 > 0.01). To analyze females, the same state combinations described above are used and Areas B and H are deleted to meet the min- imum expected cell size assumption. The chi-square result is statistically significant (7 = 56.20; df = 8; P < 0.001). Partition- ing the results indicates that the following area groupings are distinct from each other: A; C+E; F+G. To meet the minimum ex- pected cell size assumption for males, only VOLUME 110, NUMBER 3 the combined states (as above) from Areas B, E, and F could be analyzed. The chi- square results are significant (x7 = 24.43; df = 4; P = 0.001); however, there are too few areas involved for meaningful parti- tioning. Posterior thigh pattern.—As for dorsal pattern, the first state recorded for interme- diate conditions is scored for statistical anal- yses. As only three individuals have State F!, it is combined with State E The chi- Square analysis by sex is not significant ({ = 9.95; df = 5; P = 0.10 > 0.05); females, males, and juveniles are combined to ana- lyze character states by geographic areas. The chi-square analysis by area is significant (y? = 101.30; df = 30; P < 0.001). Parti- tioning the results indicates the following geographic area groupings are distinct from each other_A +B; C; E; F+G; H. Third toe tip dorsal outline.—Only one individual was scored with any indication of State C, and it was an intermediate State B-—C; for statistical analyses, that individ- ual was scored as having State B. As only two individuals had the unexpanded state, that state is combined with the “Just A” State for analyses. The chi-square analysis by sex is not significant (v7 = 6.90; df = 4; P = 0.20 > 0.10); females, males, and juveniles are combined for analysis of state frequencies among geographic areas. The chi-square analysis by area is significant (jy = 158.78; df = 24; P < 0.001). In parti- tioning the results, the following area groupings are distinct from each other: A; BC) Ex E+G+H: Third toe tip dorsal grooves.—Because so few individuals had only one groove (Appendix 2), the individuals are combined with those having two grooves. The result- ing chi-square analysis by sex is not signif- icant (x = 0.58; df = 3; P = 0.80 > 0.70); females, males, and juveniles are combined for analysis of state frequencies among geo- graphic areas. The resultant chi-square anal- ysis is significant (y7 = 34.90; df = 18; P = 0.01 > 0.001). Partitioning the results in- dicates that the following geographic 347 groupings are distinct from each other: C; Ey A-EB+F+G+H: Measurements.—Because only two adult males represent Area G, they are deleted from the analyses. MANOVA analyses were run on the two data sets (male and female). All univariate F tests were statistically significant (P < 0.001). The multivariate tests are also sig- nificant for both data sets (e.g., Wilks’ Lambda for male data = 0.022, Fi) 337 = 4.136, P < 0.001; for female data Wilks’ Lambda = 0.025, Fig4) 31. = 3.501, P < 0.001). Thus, there is significant variation of the measurement data among the geo- graphic areas. In order to understand the nature of the geographic variation, discriminant function analyses were performed on untransformed data and are discussed separately for the male and female data. The male data post-classification results (Table 1) indicate that sizes and shapes are distinctive for each area, with lesser dis- tinctiveness in morphologies between Areas A and B. A plot of the first two canonical scores (Fig. 8) indicates that the Area A and B samples overlap each other to a greater extent than any of the other samples. The male measurement data support the follow- ing area groupings as distinct from each GinerwA+Bo3AC: EB: F: H. The female data post-classification re- sults (Table 2) also indicate that the mor- phologies are distinctive within each re- gion, with less differentiation between Ar- eas C and E and among Areas E G, and H. The plot of the first two canonical scores (Fig. 9) demonstrates extensive overlap of specimen data for Areas C and E and Areas E G, and H. The female measurement data indicate the following area groupings to be distinct from. edch- -ofher:..A; Bs: -C-+E; F+G+H. Comparison of the male and female data | underscore that the males and females from Area C differ markedly in their association with the other samples. The Area C males are quite similar to those of Area H but dif- 348 FACTOR 2 2 Fig. 8. fer strikingly from those of Area E. Con- versely, the Area C females are quite sim- ilar to those of Area E but distinct from those of Area H. These patterns suggest that size and shape have responded to separate selection pressures for males and females in Vanzolinius. Advertisement calls.—The call from Li- moncocha, Ecuador has been analyzed and described previously (Straughan & Heyer 1976). Calls consist of individual notes which are partially pulsed. The basic call param- eters are similar among all five recordings O PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2 4 FACTOR | Graph of first and second canonical factors for Area A-C, E-E H males. (Table 3). The exact durations of the calls are difficult to determine, as it appears that there is microphone ringing. The most variation among calls involves: packaging of pulses; relative sharpness of attack; one or two distinct broadcast fre- quency bands; and maximum broadcast fre- quency energy. (The recording qualities are insufficient to adequately evaluate harmon- ic structure.) There is a continuum of vari- ation from a note that has relatively weak partial pulses to a note that has a few strong pulses, almost or entirely complete, each of which may or may not be partially pulsed VOLUME 110, NUMBER 3 Table 1.—Discriminant function analyses for male Vanzolinius measurement data by geographic areas. Number of observations classified into areas A B Cc E F H Area A 5 D4 1 6) 0) Area B 1 18 0) ] 0) 0) Area C 6) 0) 9 0 0 1 Area E 0 0 10) 12 0) 6) Area F 0) 0) 0 0) 12 6) Area H 0) 0 6) 0) 0 6 FACTOR 2 -5 349 Table 2.—Discriminant function analyses for female Vanzolinius measurement data by geographic areas. Number of observations classified into areas A B e E F G H Area A if 0) 0 0 0 0) 0 Area B 6) 4 @) 0 0 0 0) Area C 6) 6) 5 1 1) 0 0) Area E 1 6) 5 22 0 0 0 Area F 0) 0 1 1) 14 ] 4 Area G 0) 0) 0) 0 6) 3 z Area H 6) 0) 0 6) 0 ] 2 O S FACTOR | Fig. 9. Graph of first and second canonical factors for Area A-C, E-H females. The polygon for Area H is not labeled and occurs entirely within polygon FE 350 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.—Advertisement call data for Vanzolinius specimens from five localities. Geographic area A F F F H USNM Tape & Cut 18,1 254,5 254,13 DD 32. 256,12 Call rate per s 0.8 1.1 aS) 1.4 0.9 Mean call duration (s) 0.14 0.14 O12 0.13 0.14 Modal # of strong pulses 1 > 4+ 1 1 Mean # of total partial pulses 16.8 Dei 18.3 139 18.5 Very sharp attack sta — — nF ate Portion of call with most energy First half First half | Middle to First half First half to middle _—_ to middle first half Two distinct broadcast bands — f ar ar weak Modal value of most intense frequency — 1630 1560 1730 1980 for lower broadcast band Modal value of most intense frequency 2850 2570 2410 2590 2590 for higher broadcast band Modal values of total frequency range 2260-3190 1330-2930 1160-2830 1320-3020 1490-3010 1250- —3090 Harmonics — — weak weak? weak? (Fig. 10). All calls are frequency modulat- ed, increasing over a short time span. The relative sharpness of attack varies from moderate to sharp (Fig. 11). During the fre- quency rise at the beginning of the call, there is variation in how much energy is broadcast during the beginning of the call (1 or 2 broadcast bands, Fig. 12). The dom- inant frequencies of frog calls are known to vary with temperature, typically in an in- creasing manner (Duellman & Trueb 1986: 104). Although there is relatively little vari- ation of environmental temperatures at times of recordings, the few data points suggest an inverse relationship with tem- perature with maximum broadcast frequen- cy energy (Fig. 13). This, in turn, suggests that the variation in dominant frequencies is due to something other than temperature. Given such small sample sizes, it is not clear how much of the observed variation among calls is due to individual variation versus population or regional differentia- tion. The recordings from the paired sites of Barro Vermelho and Jaint (see Gascon et al. 1996) share distinctive call features of pulse packaging and sharpness of attack; these features are not found in the other call from the same geographic area (F), Alta- mira. Thus, this call variation appears to be very local. Two other features vary among the geographic areas, the presence of one or two broadcast frequency bands and domi- nant broadcast frequencies (Table 3). Differentiation Patterns Color patterns, morphology, measure- ments, calls.—Two trends are apparent from the preceding results (summarized in Table 4). 1) There is a strong component of dif- ferentiation at the local geographic area as defined by Areas A-—H in this study. There is a suggestion that local differentiation is more pronounced throughout the region covered by Areas A-E than for the region covered by Areas F—H. 2) There is also a component of geo- graphically related differentiation. Areas A and B share a set of states; Areas C, (D), and E share a set of states (evidence for including D based on only one available character, however); and Areas EK G, and H also share a set of states. The above characterizations are conser- vative. The actual levels of differentiation in Vanzolinius doubtless are greater than those demonstrated in this study. In many cases, insufficient sample sizes forced col- VOLUME 110, NUMBER 3 A 351 FP Sie Se | 100 ms Fig. 10. Vanzolinius advertisement call, wave forms. A. Weakly pulsed call, USNM Tape 256, Cut 12. B. Strongly pulsed call, USNM Tape 254, Cut 5. (Both calls filtered.) lapsing of either the extent of variation ob- served, deletion of geographic areas, or both during the statistical analyses. It is likely that larger sample sizes would rein- force the patterns described above and per- haps better delineate the patterns of differ- entiation among areas C-D-E and F-G-H. Enzymes.—Gascon et al. (1996) pub- lished protein starch gel electrophoretic results for 20 presumptive loci for sam- ples corresponding to Areas E, E G, and H as defined in this study. Using their published data, the samples sizes for each geographic area are E = 41, F = 24,G = 13, and H = 2. Results using Nei’s (1972) genetic distance values in a multidimen- sional scaling analysis (Wilkinson et al. 1992) indicate a general similarity to the morphologically and advertisement call based results. Each area shows some ge- netic-estimate differentiation (Fig. 14). There is not complete concordance of ge- netic-estimate differentiation with geog- raphy in that adjacent areas E and F are the most distinctive area pair in the data set (Fig. 14). As indicated with advertisement call data, the actual area/population structure unit where differentiation occurs is at a finer scale than the size of the geographic areas A—H used to group samples for mor- phological and call analyses. In order to examine the effect of genetic-estimate dif- ferentiation among sites where sample 352 Frequency in Kilohertz PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON B 0.4 O 0.4 0.2 Time in Seconds Fig. 11. Vanzolinius advertisement call, audiospectrograms. A. Sharp attack, USNM Tape 254, Cut 5. B. Very sharp attack, USNM Tape 256, Cut 12. (Neither call filtered.) Sizes were appropriate, all sites for which at least 5 individuals were available were analyzed using Nei’s distances (1972) in a multidimensional scaling analysis (Wil- kinson et al. 1992). The results indicate that there is differentiation among locali- A p -40 Ne D -60 cD) = 3) cD) ips -80 -100 O cD, 4 Fig. 12. ties (Fig. 15, note however, that the sep- aration on the y-axis is 1/10 that of the x-axis). As might be expected, one of the nearby pairs of localities from the same side of the river (Gascon et al. 1996, lo- cality numbers 6 and 7, Fig. 1, Vira-Volta B p eae ok “ea 6 O Zz 4 6 Kilohertz Vanzolinius advertisement call, spectrum analyses. A. Call with virtually all of the broadcast energy in a single peak (p, maximum energy at 2750 Hz), USNM Tape 18, Cut 1. B. Call with significant energy in a second broadcast peak (primary peak, p, with maximum energy at 2590 and 2920 Hz, secondary peak, s, with maximum energy at 1700 Hz), USNM Tape 255, Cut 2. (Both calls filtered.) VOLUME 110, NUMBER 3 26:5 Temperature in C 353 23 2400 2450 2500 2550 2600 2650 2700 2750 2800 2850 Peak Frequency in Hz Fig: (13: varzea and terra firme sites) shows essen- tially no differentiation. However, the amount of differentiation between the oth- er nearby pair of localities from the same side of the river (Gascon et al.’s locality numbers 10, Altamira and 11, Jainu (=Barro Vermelho), both varzea) are about as different from each other as any Plot of temperature versus maximum broadcast energy in Vanzolinius calls. other pair of localities analyzed (Fig. 15). This distinctiveness is due to the Altamira sample. The pronounced genetic-estimate differentiation at this scale is unexpected. Such small-scale differentiation (in a geo- graphic sense) results in an overall mosaic pattern of differentiation. Such a mosaic pattern of differentiation obfuscates any Table 4.—Unique or shared character states among geographic area samples. Upper matrix with number of shared distinctive states. Diagonal with number of unique character states/total number of states analyzed. Lower matrix with total number of characters compared between areas. A B G D E F G H A SITS 2 2 2 2 2 B 10 3/10 2 1 I C 12 10 5/12 1 5 5) 3 2 D l 1 1 0/1 1 34 Fz 10 12 1 4/12 3 Zz l 13 10 12 1 12 1/13 9 7 G 11 S| 11 l 1] 11 0/11 6 H 10 9 9 1 9 10 8 2/10 354 DIMENSION 2 -2 -| PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON O 2 DIMENSION | Fig. 14. Multidimensional scaling results for electrophoretic data given in Gascon et al. (1996), grouped by geographic areas as used in present study. larger pattern of geographic differentia- tion. The very local genetically based differ- entiation pattern provides corroboration and explanation of the results based on the mor- phological and advertisement call analyses presented previously. Discussion Distribution.—As indicated previously, specimens USNM _ 146971-146973 from Colombia: Caldas; Villa Maria are faded. I have no doubt that they are Vanzolinius, however. As seen from the distribution based on other known localities (Fig. 7), the interandean locality of Villa Maria is curious. The specimens were part of the collection donated to the Smithsonian by Hermano Niceforo Maria. The specimens were originally in the Museo La Salle col- lection as numbers 45, 45a, and 45b. Ad- jacent to these specimen records in the USNM catalogue ledger, Museo La Salle numbers 44, 44a, 44b, 44c, 44d, and 231 are also listed from the same locality. Dr. John Lynch identified the 44 series (USNM 146974-146978 respectively) as Eleutherodactylus fitzingeri and 231 (USNM 146979) as Eleutherodactylus sp. Lynch and Myers (1983) reported E. fit- zingeri from Nicaragua through Panama, the Choc6é of Colombia, and the interan- dean valleys of Colombia, but not from the Amazonian versant of the Andes in Colombia (see their Map 6, p. 535). Al- though Lynch & Myers (1983:560—561) do not include USNM 146974-146978 in their list of specimens examined, the lo- cality of Villa Maria falls within the dis- tribution described by them. Thus, some specimens from the La Salle collection from Villa Maria are geographically ap- propriate. In order to determine whether a transcription error had occurred in rela- tion to the locality data for the Vanzoli- nius specimens (USNM 146971-—146973), I asked Dr. Lynch who was in charge of the Museo La Salle herpetological collec- tion in Bogota. My intention was to de- termine the original catalogue entries for these specimens. Dr. Lynch (pers. comm.) informed me that the Director of the Mu- VOLUME 110, NUMBER 3 0.3 0.2 0.1 DIMENSION 2 ~2 -3 325 -| O DIMENSION | Fig. 15. localities represented by 5 or more specimen samples. seo La Salle, in an effort to reduce dupli- cate information, had a student write new bottle labels for specimens in the herpe- tological collection and the Director dis- carded the original catalogues. Given the suspect nature of a species of amphibian with a western Amazonian distribution having a single disjunct population in an interandean Colombian valley, the locality of Villa Maria, Caldas, Colombia must be treated with extreme suspicion and not in- cluded in any distributional analyses until new collections verify the presence of Vanzolinius in Colombian interandean valleys. Thirty six percent of the localities listed in Appendix 1 represent new locality re- cords since 1990. The present patchy dis- tribution (Fig. 7) certainly is due in part Multidimensional scaling results for electrophoretic data given in Gascon et al. (1996), for individual to collecting artifact. The current data in- dicate that the species has a distribution limited to western Amazonia. Exact knowledge of distributional limits of the species can only be approximated at this point, however. Sufficient collecting ef- forts have been undertaken in two critical areas for which absence data have some validity. Vanzolinius discodactylus has not been found in the Manaus, Brazil re- gion, nor in the State of Madre de Dios, Peru. Thus the eastern and southern dis- tributional limits likely fall between the currently known localities and Manaus and the Mant National Park. Differentiation patterns.—Al|most all studies of variation of the Amazonian her- petofauna have been directed at the spe- cies level. This is certainly appropriate, as 356 much work remains before the species limits of the Amazonian herpetofauna are well defined, particularly for amphibians. A classic exception is the detailed study by Vanzolini and Williams (1970) on Anolis chrysolepis, where they studied geographic variation at the population level over the entire range of forms that had been asso- ciated with A. chrysolepis. A portion of their methods is pertinent to the present dis- cussion (1970:25): The investigation of each character was begun by the joint consideration of 13 “‘major samples” and carried forward by the analysis of “‘transects’’—series of localities more or less linearly arranged between major samples. In this manner a two dimensional differentiation pattern of each character was obtained. The organization of the major sam- ples and transects was decided initially only on the basis of the materials avail- able and of the geometry of a map of the area of study. The results of a first analysis were then used to adapt the methods to permit better clarification of the patterns perceived. For example, the evidence (in fact known beforehand but not taken into consideration in the first study) of the existence of a well differ- entiated form, previously believed to be a well set-off species, in the area from Surinam to eastern Para, led to a pre- liminary arrangement of the Guianan and Guiano-Brasilian transects, and to their subsequent modification in order to show to better advantage the phe- nomena of transition between these populations and adjacent ones. This sort of feedback and even bias in the analysis is not only unavoidable but highly desirable in studies of geo- graphical differentiation in South America. If we had a perfect network of localities, each one represented by good samples—with statistically suffi- cient numbers of males and females, and with all age classes represented—a PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON System of isophenes would emerge from the analysis, which might even be computerized. In fact, the available collections are, as is usual with museum collections, not made for a specific purpose, an irregular and patchy representation of the group range, and the study must begin by the setting of preliminary hypotheses, to be tested. In our case the hypothesis is, in all instances, that differences between the major samples correspond to geographic patterns and not to mosaics, and this hy- pothesis must be tested by adaptations of the analysis to the materials available, to the numbers of specimens and their po- sition on the map, in other words, by the consideration of transects. Although the number of localities and samples for Amazonian amphibians and reptiles has certainly increased since 1970, the overall assessment as described by Van- zolini and Williams still holds true. The or- ganization of current studies on differenti- ation patterns of Amazonian amphibians must still be decided on the basis of mate- rials available and the geometry of a map of the study area. As a consequence, direct comparison of results among studies is not possible at present. However, the compari- sons that can be drawn suggest some state- ments that can be tested for generality with additional studies. Vanzolini & Williams (1970) found dif- ferences between samples from the geo- graphic areas represented by the areas de- fined in this paper as A+B and C+D+F+G+H (see their map 8, p. 180). They also found that samples from the nearby localities of Limoncocha and Santa Cecilia, Ecuador differed in some characters (e.g., fourth toe lamellae in males, p. 38). The only other study I am aware of that treats intraspecific geographic variation for an Amazonian amphibian that occurs in the same region with Vanzolinius dis- codactylus is that of Duellman and Mo- VOLUME 110, NUMBER 3 rales (1990) for Edalorhina perezi. Duell- man & Morales (1990) analyzed variation of dorsal texture and belly pattern of E. perezi by river drainage systems. Com- parison of their map (fig. 1:21) with the geographic areas used in this study indi- cate coincidence of the following: Region A (this study) = Napo drainage samples (their study); Region B = Pastaza drain- age samples; Regions C+D = Amazonas drainage samples. Their data (table 2:23) indicate the following: The Amazonas drainage and Napo drainage samples are similar for dorsal texture conditions, but both differ from the Pastaza drainage sample; the Napo and Pastaza drainage samples are similar for ventral pattern condition and differ from the Amazonas drainage samples. Thus the two characters they analyzed in detail show independent patterns of variation and both demonstrate patterns of differentiation at the regional level. Anolis chrysolepis, Edalorhina_ perezi, and Vanzolinius discodactylus are all forest denizens within Amazonia. All three taxa demonstrate some level of geographic dif- ferentiation at rather restricted regional lev- els. Much of the variation is difficult to put in a broad geographic context in all three species studies, however, suggesting a strong differentiation at the very local level which results in a mosaic pattern of differ- entiation that tends to obfuscate any larger scale geographic patterns. This is under- scored by the genetic estimate data avail- able for Vanzolinius. Rather than eschew the null hypothesis of differentiation being a mosaic, rather than having a geographic basis as implied by Vanzolini & Williams (1970:25), perhaps we need to embrace the mosaic concept. One avenue that needs exploration is to determine the lower spatial limits of dif- ferentiation. At present, all we can say is that differentiation occurs at the geo- graphic level that collectors have tradi- tionally used to define distinct, but near- by, collecting localities. Does differenti- 357 ation occur at even a finer scale than that for forest associated Amazonian amphibi- ans? Prognosis for further studies.—The corroboration of results from studies of differentiation of Vanzolinius discodacty- lus based on morphological features and on genetic estimate data is most encour- aging. We should be able to undertake strictly morphological analyses of other taxa of forest associated Amazonian am- phibians with confidence that the resultant patterns of differentiation reflect evolu- tionary processes and are not strictly phe- notypic responses to localized environ- mental conditions. Acknowledgments The patience of the following curators and collection managers, who allowed me to retain materials initially borrowed for an- other study in most cases, is very much ap- preciated: AMNH (standard codes per Lev- iton et al. 1985), Charles W. Myers; CAS- SU, Robert C. Drewes and Jens V. Vindum; FMNH, Harold K. Voris and Alan Resetar; ICNMNH, Maria Christina Ardila-R. and Pedro M. Ruiz-Carranza; INPA, Gloria Moreira; KU, William E. Duellman and John E. Simmons; MCZ, John E. Cadle, José P. Rosado, and Ernest E. Williams; MZUSP, Ana Maria M. Ramos-Costa and P. E. Vanzolini; RMNH, Marinus S. Hoog- moed; TCWC, James R. Dixon and Kath- ryn Vaughan. Lily R. Rodriguez allowed me to examine a specimen prior to its being catalogued. Claude Gascon, Biological Dynamics of Forest Fragments Project, Manaus, Brazil, most generously shared his collection, data, and manuscript with me. This paper would not have been undertaken without Gascon’s specimens and help. Lee-Ann C. Hayek (Smithsonian Insti- tution) provided statistical advice several times during the study. Ralph Chapman (Smithsonian Institution) performed the 358 multidimensional scaling analyses from data provided him. George R. Zug (Smithsonian Institution) critically reviewed the manuscript. The research for this paper was sup- ported by the Smithsonian Institution’s IESP - Neotropical Lowlands Research Program, Richard P. Vari, Principal Inves- tigator. Literature Cited Charif, R. A., S. Mitchell, & C. W. Clark. 1995. Ca- nary 1.2 User’s Manual. Cornell Laboratory of Ornithology. Ithaca, New York, 229 pp. Duellman, W. E. 1978. The biology of an equatorial herpetofauna in Amazonian Ecuador.—Univer- sity of Kansas Museum of Natural History Mis- cellaneous Publication 65:1—352. , & V. R. Morales. 1990. Variation, distri- bution, and life history of Edalorhina perezi (Amphibia, Anura, Leptodactylidae).—Stud- ies on Neotropical Fauna and Environment 25:19—30. Gascon, C., S. C. Lougheed, & J. P. Bogart. 1996. Genetic and morphological variation in Vanzo- linius discodactylus: A test of the river hypoth- esis of speciation.—Biotropica 28:376—387. Hayek, L. C. 1994. Analysis of amphibian biodiver- sity data. Pp. 207—269 in W. R. Heyer, M. A. Donnelly, R. W. McDiarmid, L. C. Hayek, and M. S. Foster, eds., Measuring and Monitoring PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution Press: Washington, D.C., 364 pp. Heyer, W. R. 1994. Variation within the Leptodactylus podicipinus-wagneri complex of frogs (Am- phibia: Leptodactylidae).—Smithsonian Contri- butions to Zoology 546:1—124. Leyiton;, A..E.,,R:; H: Gibbs,, Jn, E.-Healy ae Gane: Dawson. 1985. Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in her- petology and ichthyology.—Copeia 1985: 802-832. Nei, M. 1972. Genetic distance between popula- tions.—American Naturalist 106:283—292. Savage, J. M. 1987. Systematics and distribution of the Mexican and Central American rainfrogs of the Eleutherodactylus gollmeri group (Amphib- ia: Leptodactylidae).—Fieldiana, Zoology, New Series 33(1375):1—57. Straughan, I. R., & W. R. Heyer. 1976. A functional analysis of the mating calls of the Neotropical frog genera of the Leptodactylus complex (Am- phibia, Leptodactylidae).—Papéis Avulsos de Zoologia 29(23):221—245. Vanzolini, P. E., & E. E. Williams. 1970. South Amer- ican anoles: The geographic differentiation and evolution of the Anolis chrysolepis species group (Sauria, Iguanidae).—Arquivos de Zool- ogia 19:1—298. Wilkinson, L., M. Hill, J. PR Welna, & G. K. Birken- bevel. 1992. SYSTAT for Windows: Statistics, Version 5 Edition. SYSTAT, Inc.: Evanston, Il- linois, 750 pp. VOLUME 110, NUMBER 3 Appendix 1. Specimens examined. 339 Geographic area assignments in parentheses. BRAZIL. Acre: Nova Vida, (H), INPA 4490; Poron- gaba, (H), INPA 4271, 4287, 4291, 4294, 4296, 4350-— 4351; Porto Walter, (H), MZUSP 51574-51575; So- bral, (H), INPA 4348. Amazonas: Altamira, (F), INPA 3571-3573, 3584-3585, 3649, 3651, 5010, 5021, ao51.9- 5060, 5062-5063, 5103,. 5132, .5166,..5173, 5177-5178, 5214, 5217-5218; Barro Vermelho, (F), INPA 3076, 3092, 3109, 3154, 3161, 3163, 3177- Sra ol oG. 3229. 3352, 33585, 3387, 3397, 3399, 3405, 3411, 3454, 3507; Benjamin Constant, (D) CAS- -SU 11835; Condor, (G), INPA 2573, 2587, 2608, 2640, 2642, 2644, 2883, 2904-2905, 2912, 2919-— 2921; Igarapé Tucuxi, Auati-Parana, (E), MZUSP 28124; Nova Empreza, (G), INPA 2371, 2373, 2388, 2433, 2503; Nova Olinda, (F), INPA 3041; Parana (near Penedo), (G), INPA 2291, 2399; Penedo, (G), INPA 2410-2412, 2514; Rio Itacoai, (D), MZUSP 9810; Vira-Volta, (E), INPA 5694, 5696-5703, 5720, 2) SAaey 31 SO) DLa2y497 30-0199, -9191;-5761, 5763, 5766-5767, 5790, 5797-5803, USNM 348954-— 348983. COLOMBIA. Amazonas: Leticia, left margin of Rio Loreto Yacu (D), ICNMNH 11274; headwaters of Rio Caiwima, ca 70 km NNE Puerto Narifio, (D), MCZ 97025, 97033. Caldas: Villa Maria (not assigned to area), USNM 14697 1-146973. Caquetda: Florencia (not assigned to area), USNM_ 147036— 147037. ECUADOR. Napo: Coca, 290-320 m, (A), KU 158609, 175463—175464; near Laguna Taracoa, 30 km downriver from Coca, ca 250 m, (A), MCZ 94884-— 94885, MZUSP 56380; Payamino, (A), USNM 196882; Rio Yasuni (150 km upstream from Rio Napo) (A), KU 175132-175133; Santa Cecilia, 340 m, (A), KU 104666, 109163, 111403, 111424, 111429, 119347, 119350, 126241-126242, 143518, 149295- 149308, 152396, 175460. Pastaza: Mera, 1140 m, (B), KU 119303-119318, 178274-—178282; Montalvo, 250 m, (B), RMNH 23990; Puyo and environs, (B), KU 119319-119322, 202647-—202648, MCZ 90385, USNM 196878-196881, 343426—343427;: Shell Mera (B), KU 99069, 99081-99082, 99085, 99090. PERU. Loreto: Aldeia dos Indios Bora, 2 km N mouth of Zumun, (C), MZUSP 54189-54191; Estir6n, Rio Ampiyaco (C), AMNH 115691, MZUSP 23083, 24006, 24782-24786, 24793, 24797, 24803-24804, 24810, 24814, 24816, 24825; Moropon, (C), TCWC 41484; Requena, Jenaro Herrera, 140 m, (C), LR 4382 (to be deposited in Museo de Historia Natural, Univ- ersidad Nacional Mayor de San Marcos); Yanamona, (C), TCWC 41739. Ucayali: Yarinacocha, Rio Uca- yali, (not assigned to area), FMNH 56285. 360 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Appendix 2. Color patterns and morphological data state distribution by sex among geographic areas. cece ae ae ros Dorsal Snout Pattern pepe a VOLUME 110, NUMBER 3 361 Light Postorbital Eye Stripe Light Subocular Bar Dorsal Pattern 362 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Dark Mid-Dorsal Pin Stripe pa Pee eo ee Throat & A 7 5 10 3 Chest Pattern VOLUME 110, NUMBER 3 363 Belly Pattern i re 364 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON eae aimee Posterior A 2 3 4 1 2 1 Thigh Pattern 2 rae ae ee ee Third Toe N.E.+ || Tip Dorsal Outline VOLUME 110, NUMBER 3 365 Third Toe Tip Dorsal Grooves PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):366—372. 1997. Two new species of salamanders (Caudata: Plethodontidae) of the genera Bolitoglossa and Nototriton from Parque Nacional La Muralla, Honduras James R. McCranie and Larry David Wilson (JRM) 10770 SW 164th Street, Miami, Florida 33157-2933, U.S.A.; (LDW) Department of Biology, Miami-Dade Community College, Kendall Campus, Miami, Florida 33176, U.S.A. Abstract.—Two new species of salamanders of the genera Bolitoglossa and Nototriton are described from the isolated cloud forests of Parque Nacional La Muralla, Honduras. The Bolitoglossa is a member of the B. dunni group and differs from all other group members by having well-defined yellow spots on the lateral surfaces of the body. The Nototriton appears to be most closely related to N. barbouri and can be distinguished from that species by its shorter tail and smaller size. Parque Nacional La Muralla, in the northwestern portion of Departamento de Olancho, Honduras, is herpetologically one of the best known national parks in Hon- duras (Espinal et al. 1997). Espinal et al. (1997) included some recently collected salamanders from the park under the name Nototriton ‘“‘barbouri.’’ Subsequent study of these salamanders demonstrated that they were likely an undescribed species related to N. barbouri. Thus, in July 1996, we re- turned to Parque Nacional La Muralla hop- ing to collect additional specimens of these salamanders. About mid-day on 29 July, we Set up a camp at 1430 m elev. at the nearest known source of drinking water to the No- totriton locality. As the Nototriton locality was still a four hour walk from camp, we decided to collect that afternoon and night in the environs of our campsite and go to the Nototriton locality the following day. Unexpectedly, we discovered a brillantly colored undescribed species of Bolitoglossa around the campsite. The following day we made a successful visit to the previously known Nototriton locality. Study of this new material confirmed the distinctness of the Muralla Nototriton. We herein provide a formal description of each of these two salamanders. Methods and Material All measurements were made to the near- est 0.1 mm with dial calipers under a dis- secting microscope. Abbreviations used are SVL (snout to posterior end of vent), HL (head length; snout to gular fold), HW (head width), TL (tail length), HLL (hind limb length), FLL (forelimb length), CLL (combined forelimb and hind limb lengths), HFW (hind foot width), and NL (nostril length). For ease of comparison, the format for the ‘Description’ and ‘Measurements of the holotype’ sections for the Bolitoglossa follow that of McCranie & Cruz (1996), whereas those sections for the Nototriton follow the format of McCranie & Wilson (1997). The numbers in parentheses in the color in life descriptions refer to the color codes in Smithe (1975). McCranie & Cruz (1996) listed the adult material of the Bo- litoglossa dunni group recently examined and McCranie & Wilson (1997) listed all material of Nototriton recently examined. The following specimens of Nototriton bar- bouri were re-examined for this study: AMNH 54949; USNM 339700-12. Addi- tionally, a recently collected specimen of WN. barbouri (USNM 497552: Depto. Atlanti- da, Honduras) was compared to the new VOLUME 110, NUMBER 3 Fig. 1. species of Nototriton (thus a total of 27 specimens of N. barbouri have been ex- amined). Systematics Bolitoglossa decora, new species Fig. 1 Holotype.—National Museum of Natural History, USNM 500000, an adult female from along the trail to Cerro de Enmedio near the Monte Escondido campground (15°05'N, 86°44'W), Parque Nacional La Muralla, 1440 m elev., Departamento de Olancho, Honduras, collected 29 Jul 1996 by D. Almendarez, J. R. McCranie, and L. D. Wilson. Original number LDW 11032. Paratypes.—USNM 497534, an adult fe- male, USNM 497533, 497535, both appar- ently immature males, all from the same hillside as the holotype, 1440-1550 m elev. Referred specimens.—USNM 497536— 38, all subadult females, all from the same hillside as the holotype, 1430-1500 m elev. Diagnosis.—Bolitoglossa decora is a member of the B. dunni group as defined by Elias (1984). Bolitoglossa decora can be distinguished from all other members of the 367 $ , / ee oe tee ae Sisal at aga ent BAg Adult female holotype of Bolitoglossa decora (USNM 500000), SVL 62.1 mm. group—B. carri McCranie & Wilson, B. ce- laque McCranie & Wilson, B. conanti McCranie & Wilson, B. cuchumatana (Stu- art), B. dunni (Schmidt), B. engelhardti (Schmidt), B. helmrichi (Schmidt), B. lon- gissima McCranie & Cruz, B. porrasorum McCranie & Wilson, and B. rostrata (Broc- chi)—by having well-defined Buff-Yellow (in life) or pale yellow (after a short time in preservative) spots on the lateral surface of the body. Individuals of some of the above mentioned species may have pale flecks on the lateral surfaces or pale dor- solateral stripes, but not well-defined pale spots laterally. Additionally, B. decora has less webbing (two to slightly over two pha- langes on both sides of digit III on both forelimbs and hind limbs free of webbing) than all of the above named species except for B. longissima, B. rostrata, and some specimens of B. celaque (see table 1 in McCranie & Cruz 1996, for a comparison to the remaining Honduran members of the B. dunni group). Bolitoglossa decora also has shorter limbs than B. longissima (male HLL/SVL 24.9-26.4%, X = 25.7 versus 31.9% in one male Jlongissima; female 368 HLL/SVL 24.1—25.3%, X = 24.7 versus 31.0-31.9%, X = 31.6 in longissima). Description.—Relatively large (SVL 36.5—40.2, X = 38.4 in two apparently im- mature males; 61.0—62.1, X = 61.6 in two adult females) member of B. dunni group; snout nearly truncate to broadly rounded in dorsal aspect, broadly rounded in profile; females more robust than relatively slender males; labial protuberances well developed in males, weakly developed in females; mental gland very weakly developed in largest male, mental gland not evident in smaller male; head relatively narrow (HW/ SVL 15.7—16.4%, X = 16.1 in two males; 16.1-16.3%, X = 16.2 in two females); eyes slightly protuberant, not or only barely visible beyond margin of jaw when viewed from below in both sexes; postorbital groove shallow, extending posteriorly from eye before turning sharply ventrally to con- nect with gular fold, another groove pro- ceeding sharply ventrally just posterior to mandible and extending irregularly across throat anterior to gular fold; sublingual fold absent; maxillary teeth moderately abun- dant (47-54, X = 50.5 in two males; 68— 70, X = 69.0 in two females), extending beyond level of center of eye; vomerine teeth abundant (23-31, X = 27.0 in two males; 29—31, X = 30.0 in two females), in long, single or slightly irregular, arched se- ries extending slightly beyond level of me- dial border of choanae; premaxillary teeth (2 in both males; 5—6, X = 5.5 in two fe- males) enlarged, piercing lip or located just posterior to lip in males, not enlarged, lo- cated posterior to lip in females; tail later- ally compressed, constricted basally; tail relatively short (TL/SVL 74.9% in one male: 75.7-77.0%, X°= 764 im two’ fe- males); limbs slender, long, adpressed fore- limb and hind limb slightly overlapping to limb interval of 1/2 costal fold in two males, limb interval 1/2—1 costal fold in two females (HLL/SVL 24.9-26.4%, X = 25.7 in two males; 24:1-25.3%,X = 247 in two females); webbing reduced, with two to slightly over two phalanges on both sides PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of digit III on both forelimbs and hind limbs free of webbing; digit tips bluntly rounded, bearing well-developed subdigital pads; relative length of digits on forelimbs I200 mm SL) are known for Anomalops (McCosker & Rosenblatt 1987), and Phthanophaneron harveyi is known from a 20-mm SL juvenile, the 67.7-mm SL ho- 374 lotype, and a 204-mm SL specimen (Ro- senblatt & Montgomery 1976, McCosker & Rosenblatt 1987, Allen & Robertson 1994). Large flashlight fishes typically are taken in deeper water than small ones (e.g., Mc- Cosker & Rosenblatt 1987), and may be more common than their poor representa- tion in fish collections suggests. Further collecting efforts are needed to find small specimens of the new species, the holotype of which is from an area of the Cook Is- lands where small specimens of both An- omalops and Photoblepharon occur. Our purposes here are to describe the new spe- cies and discuss its relationships in the con- text of an existing phylogenetic hypothesis of anomalopid genera. Methods Measurements were made with needle- point dial calipers or an ocular micrometer to the nearest 0.1 mm. Terminology of structures associated with the light organ follows Johnson & Rosenblatt (1988). Gill- raker counts include all rudiments. Scale bars in illustrations represent 1 mm. We fol- low Johnson & Rosenblatt (1988) in using the name “‘Trachichthyoidei’”’ for the clade comprising the Anomalopidae, Monocentri- dae, and Trachichthyidae, the monophyly of which was proposed by Zehren (1979) and Moore (1993). The Monocentridae and Trachichthyidae were considered the first and second outgroups, respectively, for the phylogenetic analysis of anomalopid genera based on the hypothesized monophyly of the Trachichthyoidei and a proposed sister- group relationship between the Monocen- tridae and Anomalopidae (Konishi & Oki- yama 1997). Following Zehren (1979), Moore (1993) suggested a sister-group re- lationship between the Monocentridae and Trachichthyidae, but the single character cited as evidence involves the infraorbital series, which is modified in all anomalopids to accommodate the light organ. Further study is needed to test Konishi & Okiya- ma’s (1997) Anomalopidae + Monocentri- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dae hypothesis, which also is based on a single character: presence of a separate spi- nous dorsal fin in monocentrids and all an- omalopids except Photoblepharon. Recog- nizing that neither hypothesis of trachich- thyoid interrelationships is well supported, we initially included the sister group of trachichthyoids, the Anoplogastridae + Di- retmidae (Zehren 1979, Moore 1993, Bald- win & Johnson 1995), as an additional out- group in our analysis, but this inclusion had no effect on the topology of the tree and will not be discussed further. All characters included in the phylogenetic analysis were weighted equally, and multistate characters were treated as unordered. Protoblepharon, new genus Diagnosis.—An anomalopid with 21 gill rakers on the first arch, about 145 body scale rows, a small (14.5% HL) rotatable light organ, no postorbital papillae, and a very small gap between the lacrimal and nasal for passage of the stalk, which is twisted and not broadly exposed posterior- ly. The following combination of characters also is useful in distinguishing Protobleph- aron: a separate spinous dorsal fin, a pel- vic-fin spine, two anal-fin spines, 59—60 en- larged lateral-line scales, an elastic shutter, and a broad strap-like ethmomaxillary lig- ament with no groove or swelling. Type species.—Protoblepharon rosen- blatti, new species. Etymology.—From the Greek protos, first, and blepharon, eyelid, in reference to the cladistic position of the genus as first in the lineage of flashlight fishes that occlude the light organ with an erectable shutter. Protoblepharon rosenblatti, new species Fig. 1 Holotype.—AMS — 1.24275-001. A 229-mm SL female specimen caught by hook and line northwest of Matavera, Rar- otonga, Cook Islands (21°12'S, 159°45’W), at 274 m on 30 Nov. 1983, and donated by VOLUME 110, NUMBER 3 N. Sims of the Cook Islands Fisheries De- partment. Description.—Counts and measure- ments, in mm, of the holotype: Dorsal-fin rays VI-I, 14; anal-fin rays II, 11; pectoral- fin rays 11151; pelvic-fin rays 1,5; caudal-fin rays 10, 10+9, 9 (all procurrent rays spi- nous except the posteriormost in the upper and lower caudal-fin lobes); branchiostegals 8; gill rakers on first arch 21 (5+ 12 rakers, plus two flat plates at dorsal end of epi- branchial and two at anterior end of cera- tobranchial); pored lateral-line scales 59 (60 on right side); scale rows above lateral line ca. 18; abdominal scutes 9; vertebrae 14+ 16. Head length 83.4; predorsal length 96.9; prepelvic length 112; body depth at origin of dorsal fin 81.9; caudal-peduncle depth 23.3; caudal-peduncle length 50.5; snout length 23.7; eye diameter 19.4; orbit diameter 20.5; light-organ length 12.1; pec- toral-fin length 51.9; pelvic-fin length 41.2; first dorsal-spine length 12.2; third dorsal- spine length 17.0; sixth dorsal-spine length 6.8; seventh dorsal-spine length 20.0; first anal-spine length 5.8; second anal-spine length 12.3. Body compressed (width 1.9 in depth) and deep (depth at origin of dorsal fin 1.8 in length without head). With mouth open, profile sloping gradually from occiput to snout, somewhat convex in region of mes- ethmoid, then dropping slightly to symphy- sis of upper jaw; upper-jaw symphysis at level of horizontal through middle of eye. Nostrils anterior and completely dorsal to eye with mouth open, the anterior with thickened posterior rim. With jaws forced closed, mouth oblique, lower jaw originat- ing anteriorly near horizontal through mid- dle of eye, and maxilla extending posteri- orly to vertical through middle of eye. Pos- terior supramaxilla ovoid, anterodorsal sur- face with small pointed process extending anteriorly along posterodorsal edge of small anterior supramaxilla. Posterior supramax- illa covering most of posterior portion of maxilla, the posteroventral corner of max- illa exposed and covered with tiny black pa- 37D pillae. Distinct notch at symphysis of pre- maxillae, presumably accommodating small dentigerous knobs at symphysis of dentaries when mouth closed. Premaxillae, including most of lateral and medial surfaces, covered with bands of villiform teeth; no teeth at symphyseal notch. Each dentary with nar- row band of villiform teeth posteriorly, patch of slightly larger teeth near symphy- sis extending onto lateral and medial sur- faces. Vomer edentulous, palatines with well-developed bands of villiform teeth. Bones of head and pectoral girdle cov- ered with numerous, rugose to minutely ser- rate ridges. Cleithrum with large exposed surface posteriorly, margin smooth. Supra- cleithrum almost completely beneath oper- cle, only posterodorsal corner exposed, margin smooth. Anterior infraorbitals en- larged, covering anteroventral corner of or- bit, and slightly flared laterally forming a medially sloping plate. Laterosensory ca- nals of head appearing as channels of dark skin surrounded by bone, skin covered with small black papillae and perforated fre- quently by pores. Eye small, diameter 4.1 in head. No fleshy papillae on posterior rim of orbit. Lu- minous organ below eye small, length 6.9 in head. Light organ free posteriorly, sup- ported by fibrocartilaginous cup anteriorly, which is ligamentously bound to a fibro- cartilaginous stalk. Organ capable of being rotated downward into pocket below eye and medial to infraorbitals. When occluded, dorsal margin of light organ well below in- fraorbital rim. Black elastic shutter mem- brane attached along outer margin of sub- orbital pocket. Numerous small, spinoid scales (cf. Rob- erts 1993, = Ct’ of Johnson 1984) covering body, about 145 lateral body rows but dif- ficult to count because of irregular distri- bution of scales. Lateral line covered by en- larged scales, and abdomen with series of about 9 enlarged, keeled scutes. Head most- ly scaleless, a few thick, heavily sculptured scales at anterodorsal corner of opercle, scales coalescing on cheek to form strong 376 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pig. 1. AMS. bony covering. Gular region naked, with low, pigmented, transverse ridges. Most rakers on first gill arch long and lath-like, length of first raker below angle nearly two-thirds diameter of eye. Four an- teriormost and two dorsalmost rakers on first arch and rakers on remaining arches less than half that long. Pseudobranch well developed, about 27 filaments. Spinous dorsal fin lower than soft dorsal fin, the longest spine (seventh) 1.7 in length of longest unbroken soft ray (the fifth). Length of first spine 1.2 in second, second 1.1 in third and fourth, fifth 1.3 in first, sixth 1.7 in first, and seventh 0.6 in first. First anal spine shorter than any dorsal Spine, second anal spine about equal in length to first dorsal spine. Longest anal soft ray (second) about equal to longest dor- sal soft ray. Caudal fin deeply forked. Pectoral-fin base nearly horizontal, fin di- rected posterodorsally. Pectoral-fin length about 1.6 in head, third through sixth rays the longest. Pelvic fin shorter than pectoral, 2.0 in head, and falling well short of anus. Color in alcohol dark brown to black; fins, posterior trunk, and cheek darkest. Lateral- line scales pale. Occlusion mechanism (Fig. 2).—The cup Holotype of Protoblepharon rosenblatti, AMS 1.24275-001, 229 mm SL. Photograph by C. Bento, supporting the light organ is connected to a fibrocartilaginous stalk that is continuous with its contralateral member across the snout, with no attenuation at the commis- sure. Anteriorly, the stalk lies flat against the snout, with the broad surface facing out- ward; posteriorly, where the stalk passes through a small gap between the lacrimal and nasal, it twists such that its broad sur- face lies nearly in the horizontal plane (Fig. 2B). The posteroventral portion of the stalk and anteroventral corner of the cup each terminate in a short ventral process, and these are loosely joined dorsally by a short ligament. There is no ventral stalk hook. The shutter is slightly thickened near its an- terodorsal corner, but there is no discrete shutter knob. The cup supports the anterior end of the light organ and extends posteri- orly along about two-thirds of its ventral surface (Fig. 2B). A large section of it is exposed anterolaterally between the light organ and stalk; anteromedially, the cup forms a medially projecting shelf. The stalk muscle is not differentiated into dorsal and ventral bundles and inserts on the ligament connecting the stalk and cup. The Ligament of Diogenes originates on the rostral carti- lage, curves around the ethmomaxillary lig- VOLUME 110, NUMBER 3 ament as a broad strap, and then narrows posteriorly before inserting broadly on the ventral process and anteromedial shelf of the cup. The ethmomaxillary ligament de- scends anteroventrally from its origin on the mesethmoid to insert on the maxilla with a short branch to the palatine. This lig- ament is broad and flat, lacks a groove where it is crossed by the Ligament of Di- ogenes, has no pronounced forward flexure, and bears no swellings. Etymology.—lIt is our pleasure to name this species in honor of Dr. Richard H. Ro- senblatt, a mentor to one of us (GDJ), friend and valuable colleague to all of us. His contributions to the systematics and functional morphology of flashlight fishes have shed much light on the evolution and biology of the Anomalopidae. Remarks.—States for P. rosenblatti of characters not associated with the light or- gan used by Johnson & Rosenblatt (1988) and Rosenblatt & Johnson (1991) to recon- struct the evolutionary history of anomal- opid genera are given in Table 1, those as- sociated with the light organ in Table 2. In Table 3, we list states of characters among anomalopids and outgroups not considered in previous publications that are useful in diagnosing P. rosenblatti: number and rel- ative size of lateral-line scales, number of gill rakers, relative size of light organ, least distance between nasal and lacrimal, and orientation of fibrocartilaginous stalk. The relatively small size of the light organ of P. rosenblatti could be a function of the large size of the holotype, as comparisons with Phthanophaneron harveyi indicate an in- verse relationship between relative size of light organ and body (31.2% HL in the 67.7-mm SL holotype and 22.7% HL in the 204-mm SL specimen). McCosker (1982) noted a similar trend in Kryptophanaron al- fredi (44.7% HL in a 25-mm SL specimen, 34.9% HL in an 89-mm SL fish). Relationships.—Protoblepharon is most similar to Parmops in the primitive nature of the occlusion mechanism, es- pecially the absence of a shutter knob and S77 stalk hook. These features are important in the erection of the shutter in Phthano- phaneron, Kryptophanaron, and Photo- blepharon (Johnson & Rosenblatt 1988), and it is thus unclear how the shutter mechanism operates in either Parmops or Protoblepharon. Protoblepharon \acks the primary diagnostic feature of Par- mops, expansion of the first four infraor- bital bones to form a medially sloping Shelf that protrudes laterally well beyond the margin of the orbit (Rosenblatt & Johnson 1991). It shares with Parmops, Phthanophaneron, Kryptophanaron, and Photoblepharon many features recognized as synapomorphies of that lineage by Johnson & Rosenblatt (1988): two supra- maxillae, transverse ridges on the gular isthmus, large v-shaped lateral dentary tooth patches, over 100 lateral body scale rows, reflective lateral-line scales, an erectable (or at least elastic) shutter, and a stalk that is continuous across the snout. It lacks another derived feature of that group, a groove in the ethmomaxillary ligament (secondarily absent in Phthano- phaneron). It is thus most parsimonious to hypothesize that Protoblepharon is the sister group of Parmops + Phthanophan- eron + Kryptophanaron + Photoblephar- on. A previously undescribed character corroborating the placement of Proto- blepharon below Parmops involves the configuration of the lacrimal, nasal, and stalk. In most anomalopids, the stalk is broadly exposed at the commissure (Figs. 2B, C; 3B, C), and there is no attenuation where it joins its contralateral member. In Protoblepharon, the lacrimal and nasal are separated by only a small gap (least distance between them ca. 1.6% HL; Fig. 3B), and the stalk twists before passing through this gap such that the broad sur- face is nearly in the horizontal plane, and the narrow margin formerly in a ventral position is exposed laterally (Fig. 2B). In the Parmops clade, the lacrimal is sepa- rated from the nasal by a large gap (4.0% HL or more, Fig. 3C), and the broad sur- 378 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Big. 2, blepharon rosenblatti, AMS 1.24275-001, (C) Kryptophanaron alfredi, USNM 343635. Right side. face of the stalk is exposed along its entire length (Figs. 2C, 3C). Anomalops also has a very small gap between the lacrimal and nasal (ca. 1.0% HL, Fig. 3A), and the two bones are in contact with one another in the outgroups. A large gap between the lacrimal and nasal is thus an additional synapomorphy of Parmops, Phthan- ophaneron, Kryptophanaron, and Photo- Light organ and associated structures in (A) Anomalops katoptron, USNM 293340, (B) Proto- blepharon. A small gap is autapomorphic for the Anomalopidae. Although both have small gaps, the conditions in An- omalops (figs. 2A, 3A) and Protoblephar- on (figs. 2B, 3B) are different in that the stalk is never broadly exposed in Anom- alops (it is attenuated at the commissure) and does not twist posteriorly but nearly disappears from view in the region of the VOLUME 110, NUMBER 3 Table 1.—States in Protoblepharon rosenblatti of Johnson & Rosenblatt’s (1988) characters not associ- ated with the light-organ complex. 1. Vertebrae bearing epineurals* 1, 2, 10-14 2. Branchiostegals spiny 3. Openings in pars jugularis ? 4. Parasphenoid flanges ? 5. Swimbladder stay — 6. Postorbital papillae 0) 7. Cephalic sensory canal covering __Papillose 8. Lateral-line tubes Closed 9. Midventral scutes Continuous 10. Dorsal Fin VI-I,14 Supraneurals 0/0/1+ 1/ 11. Supramaxillae 2 12. Transverse ridges on gular isthmus + 13. Lateral dentary tooth patch arse V"’ 14. Body scale rows ca. 145 15. Reflective or transparent lateral- line scales + 16. Pelvic spine Ss 17. Anal spines II 18. Vertebrae 14 + 16 19. Corner of maxilla Papillae * Epipleurals of Johnson & Rosenblatt (1988) are epineurals (Patterson & Johnson 1995). small lacrimal/nasal gap as it turns abrupt- ly ventrad (Johnson & Rosenblatt 1988). The different associations between the lacrimal and nasal possibly are also re- flected in the morphology of the C-shaped process of the lacrimal (Zehren 1979), 379 which is probably smaller in Proto- blepharon than in the Parmops lineage because of the closer association of the lacrimal and nasal (and hence the lateral ethmoid). Additional specimens of Proto- blepharon that can be cleared and stained are needed to examine osteology. A cladogram depicting relationships among the six anomalopid genera is shown in Figure 4. We constructed the tree using the Branch and Bound option of Swofford’s (1991) PAUP 3.0 with the matrix in Table 4. All of the characters (Tables 1, 2) used by Johnson & Rosenblatt (1988) and Ro- senblatt & Johnson (1991) initially were in- cluded in the analysis, but many are not in- formative and were eliminated from the matrix. Johnson & Rosenblatt (1988) and Rosenblatt & Johnson (1991) included all characters in Tables 1 and 2 on their clado- grams, but noted that many features asso- ciated with the light organ could not be po- larized by outgroup comparison because of the absence of comparable conditions in the outgroups. They surmised that the fully ro- tatable light organ of Anomalops and the complex shutter mechanism of the non-ro- tatable light organ of Photoblepharon rep- resent highly specialized conditions within the family, and interpreted the less special- Table 2.—States in Protoblepharon rosenblatti of Johnson & Rosenblatt’s (1988) characters associated with the light organ. I Attachment of Ligament of Diogenes on cup Medial Il Attachment of Ligament of Diogenes anteriorly Rostral Cartilage Il Cup with medial shelf Moderate IV Insertion of stalk muscle dorsally Ligament Vv Stalk with inward flexure at cup articulation — VI Rotation pad a VII Postocular skin flap — Villa Erectile shutter VIIIb Shutter knob — IX Stalk hook a xX Stalk continuous across snout a Xla Ethmomaxillary ligament with groove — XIb Ethmomaxillary ligament with medial swelling XII Hook and shutter knob intimately associated NA XIII Cup process attached to stalk hook by ligament NA XIV Organ rotatable BE 380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.—Comparison of some diagnostic features of Protoblepharon rosenblatti among anomalopids and other trachichthyoids. Data are from the literature or specimens examined in this study (single values represent counts or measurements of a single individual). For light organ length and least distance between nasal and lacrimal, range of values given only for taxa where data from a wide range of adult sizes was available. HL = head length, ant. = anteriorly, NA = not applicable. Least distance between Light organ nasal & Gill rakers length lacrimal Taxon Lateral-line scales on Ist arch (% HL) (% HL) Orientation of stalk Anomalops 57-59 28-323 35.4 1.0 Not broadly exposed (Not enlarged) Protoblepharon 59-60 Dae 14.5 1.6 Broadly exposed ant., (Enlarged) twisted posteriorly Parmops 30 30 35.6 4.8 Broadly exposed (Enlarged) Phthanophaneron 38 22-24» 22.7—31.2 4.0-4.6 Broadly exposed (Enlarged) Kryptophanaron 32—34¢ 24—28° 36.3—44.74 Si) Broadly exposed (Enlarged) Photoblepharon 3) 25—30° 48.6 5.6 Broadly exposed (Enlarged) . Monocentridae 12—15' 19-21° NA 0) NA (Not enlarged) Trachichthyidae 25-638 15-44» NA 0) NA (Enlarged in some) 4 Shimizu (1984), this study. > Counts include flat plates at dorsal end of epibranchial, anterior end of ceratobranchial, or both. © Colin et al. (1979), this study. 4 McCosker (1982). © Abe & Haneda (1973). f Smith (1986). ® Gomon (1994). h Kotlyar (1980). A NASAL LACRIMAL eae = Fig. 3. Diagrammatic illustrations of the anterior snout region in three anomalopids, showing different con- figurations of the nasal, lacrimal, and fibrocartilaginous stalk. (A) Anomalops katoptron, USNM 293340, (B) Protoblepharon rosenblatti, AMS 1.24275-001, (C) Kryptophanaron alfredi, USNM 343635. Dotted line in Anomalops shows anterior extent of skin covering stalk, that in Protoblepharon represents region where stalk is twisted. VOLUME 110, NUMBER 3 5 6 1 8, 9, 381 16,: Ethmomaxillary ligament with swelling 13,: Shutter knob present 14,: Stalk hook present 4,: One postorbital papilla 17,: Large gap between lacrimal & nasal 15,*:Ethmomaxillary ligament with groove 18,: 15 or more dorsal rays ; 2 Supramaxillae : Gular ridges present : Large V-shaped dentary tooth patch : >100 body scale rows : Pale lateral-line scales 1,: Light organ present 2,: Neural & haemal spines support procurrent rays 3,: Interarcual cartilage present 17,: Lacrimal & nasal separate but close Fig. 4. Cladogram showing hypothesized relationships among anomalopid genera. * = ambiguous character resolved using ACCTRAN. Treelength = 29 CI = 0.83 RI = 0.83. ized states in Parmops and Phthanophan- eron as ancestral for the family. They po- larized features of the light organ based on these assumptions. The PAUP analysis did not make such assumptions, and recognized that the occlu- sion mechanisms in either Anomalops or cladistically primitive members of the Pro- toblepharon lineage could be ancestral for the family. Nevertheless, the phylogeny of Johnson & Rosenblatt (1988) and Rosen- blatt & Johnson (1991) emerged as the sin- gle most parsimonious tree based on 18 in- formative characters (Table 4). We concur with Johnson & Rosenblatt (1988) that the occlusion mechanisms of Anomalops and Photoblepharon probably represent the most derived conditions within the family and that the occlusion mechanism of Pro- toblepharon, which lacks a well defined shutter knob, a stalk hook to engage the shutter knob, and a groove and swelling in the ethmomaxillary ligament, is ancestral. Relative to outgroups Monocentridae and Trachichthyidae, other features of Proto- blepharon that might corroborate a Proto- blepharon-like anomalopid ancestor include the small size of the light organ (14.5% HL vs. 22.7—48.6% HL in all other anomalopid genera), absence of postorbital papillae (eight in Anomalops, one in other anomal- opids, none in the outgroups), and low number of gill rakers (21 or fewer in Pro- toblepharon, monocentrids, and some trachichthyids, 24 or more in other flash- light fishes). Acknowledgments We thank N. Sims of the Cook Island Fisheries Department for donating the spec- imen of the new flashlight fish to the Aus- tralian Museum. J. E. McCosker, J. A. Moore, R. H. Rosenblatt, V. G. Springer, 382 Table 4.—Character matrix used in constructing cladogram in Figure 4, followed by brief description of character states. See text and Johnson & Rosenblatt (1988) for further descriptions of characters. ? = miss- ing data, 9 = not applicable, ¥, = polymorphism. 125 6-10 i=15_ e=1s Trachichthyidae HVOOO™ -COL0O 17,999) 910% Monocentridae 00000 00000 00999 900 Anomalops LEV2O 0 O10 ALOOO 010 Protoblepharon 20d. 41210 00000 O10 Parmops rece wi 2AM O26 Phthanophaneron 1 2A ot 1 2a OA OO Di: Kryptophanaron deel tee OO OLN. £27 Photoblepharon Mia LL 12. 1. Light organ absent (0), present (1). 2. Neural and haemal spines of fourth preural ver- tebra do not support procurrent caudal rays (0), support procurrent caudal rays (1). 3. Interarcual cartilage absent (0), present (1). 4. Postorbital papillae zero (0), one (1), eight or nine (2). 5. Supramaxillae one (0), two (1). 6. Transverse gular ridges absent (0), present (1). 7. Lateral dentary tooth patch small (QO), large v- shaped (1). 8. Lateral body scale rows <50 (QO), 50-100 (1), >100 (2). 9. Pale (reflective?) lateral-line scales absent (0), present (1). . Pelvic-fin spine present (0), absent (1). *11. Corner of maxilla papillose (0), with bony or- namentation (1), smooth (2). . Gill rakers 21 or fewer (0), 24 or more (1). 13. Shutter knob absent (0), present (1). 14. Stalk hook absent (0), present (1). . Ethmomaxillary ligament without groove (0), with groove (1). 16. Ethmomaxillary ligament without swelling (0), with discrete swelling (1). 17. Nasal connected to lacrimal (0), not connected but close (least distance between bones 1.0— 1.6% headlength)—(1), separated by large space (least distance between bones >4.0% headlength)—_(2). 18. Dorsal-fin rays 14 or fewer (0), 15 or more (1). *Informative only as autapomorphy of one or more genera; ** ambiguous character. and H. J. Walker made helpful comments on a draft of this manuscript. Literature Cited Abe, T., & Y. Haneda. 1973. Description of a new fish of the genus Photoblepharon (Family Anomal- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON opidae) from the Red Sea.—Sea Fisheries Re- search Station, Haifa 60:57—62. Allen, G. R., & D. R. Robertson. 1994. Fishes of the tropical eastern Pacific. Crawford House Press, Bathurst, Australia, 332 pp. Baldwin, C. C., & G. D. Johnson. 1995. A larva of the Atlantic flashlight fish. Kryptophanaron al- fredi (Beryciformes: Anomalopidae), with a comparison of beryciform and stephanoberyci- form larvae.—Bulletin of Marine Science 56:1— 24. Colin, P L., D. W. Arneson, & W. E Smith-Vaniz. 1979. Rediscovery and redescription of the Ca- ribbean anomalopid fish Kryptophanaron alfre- di Silvester and Fowler (Pisces: Anomalopi- dae).—Bulletin of Marine Science 29:312-319. Gomon, M. FE 1994. Family Trachichthyidae. Pp. 399-410 in M. E Gomon, J. C. Glover, & R. H. Kuiter, eds., The fishes of Australia’s south coast. State Print, Adelaide, 992 pp. Harvey, E. N. 1922. The production of light by the fishes Photoblepharon and Anomalops.—Car- negie Institute of Washington Publication 312: 45-60. Haygood, M. G., & D. H. Cohn. 1986. Luciferase genes cloned from the unculturable luminous bacteriod symbiont of the Caribbean flashlight fish, Kryptophanaron alfredi.—Gene 45:203-— 209. Johnson, G. D. 1984. Percoidei: development and re- lationships. Pp. 464—498 in H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall, Jr., & S. L. Richardson, eds., Ontogeny and systematics of fishes——American Society of Ichthyologists and Herpetologists Special Publication 1:1—760. , & R. H. Rosenblatt. 1988. Mechanisms of light organ occlusion in flashlight fishes, family Anomalopidae (Teleostei: Beryciformes), and the evolution of the group.—Zoological Journal of the Linnean Society 94:65—96. Konishi, Y., & M. Okiyama. 1997. Morphological de- velopment of four trachichthyoid larvae (Pisces: Beryciformes), with comments on_trachich- thyoid relationships.—Bulletin of Marine Sci- ence 60:66—88. Kotlyar, A. N. 1980. Systematics and distribution of trachichthyid fishes (Trachichthyidae, Beryci- formes) of the Indian Ocean.—Transactions of the P. P. Shirshova Institute of Oceanology 110: 177-224. McCosker, J. E. 1982. Discovery of Kryptophanaron alfredi (Pisces: Anomalopidae) at San Salvador, Bahamas, with notes on anomalopid light or- gans.—Revista De Biologia Tropical 30:97-99. , & R. H. Rosenblatt. 1987. Notes on the bi- ology, taxonomy and distribution of anomalopid VOLUME 110, NUMBER 3 fishes (Anomalopidae: Beryciformes).—Japa- nese Journal of Ichthyology 34:157—164. Moore, J. A. 1993. The phylogeny of the Trachi- chthyiformes (Teleostei: Percomorpha). Pp. 114-135 in G. D. Johnson & W. D. Anderson, Jr., eds., Phylogeny of the Percomorpha.—Bul- letin of Marine Science 52:1—629. Patterson, C., & G. D. Johnson. 1995. The intermus- cular bones and ligaments of teleostean fish- es.—Smithsonian Contributions to Zoology 559:1-83. Roberts, C. D. 1993. Comparative morphology of spined scales and their phylogenetic signifi- cance in the teleostei.—Bulletin of Marine Sci- ence 52:60-113. Rosenblatt, R. H., & G. D. Johnson. 1991. Parmops coruscans, a new genus and species of flashlight fish (Beryciformes: Anomalopidae) from the South Pacific.—Proceedings of the Biological Society of Washington 104:328-—334. 383 , & W. L. Montgomery. 1976. Kryptophan- eron harveyi, a new anomalopid fish from the eastern tropical Pacific, and the evolution of the Anomalopidae.—Copeia 1976:510—515. Shimizu, T. 1984. Order Beryciformes. Pp. 108-110 in H. K. Masuda, K. Amaoka, C. Araga, T. Uyeno, & T. Yoshino, eds., The fishes of the Japanese Archipelago. Tokai University Press: 1-437. Smith, M. M. 1986. Family No. 128: Monocentridae. Pg. 413 in M. M. Smith and P. C. Heemstra, eds., Smiths’ Sea Fishes. Macmillan South Af- rica (Publishers) (Pty) Ltd., Johannesburg, 1047 Pp. Swofford, D. L. 1991. PAUP: Phylogenetic analysis using parsimony, version 3.0s. Computer pro- gram distributed by the Illinois Natural History Survey, Champaign, Illinois. Zehren, S. J. 1979. Pisces: Teleostei. The comparative osteology and phylogeny of the Berycifor- mes.—Evolutionary Monographs I, 389 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):384-387. 1997. Status of Platycephalus cantori Bleeker, 1879 (Teleostei: Platycephalidae) Leslie W. Knapp and Hisashi Imamura (LWK) Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (HI) Hachinohe Branch, Tohoku National Fisheries Research Institute, 25—259 Same, Hachinohe, Aomori 031, Japan Abstract.—Platycephalus cantori Bleeker, 1879, P. maculosus Peters, 1869 and P. bobbosok Bleeker, 1853 are here regarded as junior synonyms of P. carbunculus Valenciennes in Cuvier & Valenciennes, 1833. Lectotypes are se- lected for P. cantori and P. maculosus. With a brief description based on a spec- imen from Bombay, Valenciennes in Cuvier & Valenciennes (1833) described Platyce- Phalus carbunculus (currently Eurycephal- us carbunculus according to Imamura, 1996). Cantor (1849) identified specimens from Pinang as Platycephalus carbunculus and gave a more extensive description. Giinther (1860) erroneously listed dried ‘skins from Dr. Cantor’s collection as types of P. carbunculus and synonymized P. car- bunculus under P. malabaricus Cuvier, 1829. Bleeker (1879) insisted that Cantor’s P. carbunculus was not P. carbunculus Va- Table 1.—Counts and proportional measurements of holotype of Eurycephalus carbunculus and syntypes of Platycephalus cantori. E. carbunculus Platycephalus cantori (Holotype) (Syntypes) Standard length (SL) mm 128.4 133.6 135.4 Head length (HL) 43.1 44.3 43.2 Counts Dorsal-fin rays I-VII-11 I-VIII-11 I-VII-? Anal-fin rays 12 12 12 Pectoral-fin rays 19 19 19 Pelvic-fin rays ik 5 1) 1 re) Lateral line scales 54 55 55 Proportions as % SL Head length 33.6 3) o4 513 Snout length 8.8 9.0 8.1 Orbital diameter 8.6 9.4 8.3 Interorbital width 1:9 je es Length of caudal peduncle 8.8 O72 8.9 Depth of caudal peduncle 4.4 4.1 4.5 Pectoral-fin length yell 15.4 2 Proportions as % HL Snout length 26.2 PLAS 29.5 Orbital diameter 2535 28.2 Dh) Interorbital width 5.8 ae 4 ? Upper jaw length 34.1 359 34.5 Lower jaw length 30:3 aye) 53.9 VOLUME 110, NUMBER 3 > SS ae ——__ —s >> ~~ 2>>>>55>~ 385 > ae oo > —_— a7 = > N > pe aad ae > els — > = —_—_, => = oF Fig. 1. mm. lenciennes and renamed it P. cantori. After examining the primary type specimens of P. carbunculus and P. cantori, de Beaufort & Briggs (1962) followed Bleeker in rec- ognizing the two species. Murty (1982) list- ed Thysanophrys cantori but noted the lack of authentic records from India. We believe de Beaufort and Briggs were incorrect and we regard P. cantori Bleeker, 1879, P. bo- bossok Bleeker, 1860 and P. maculosus Pe- ters, 1869 as junior synonyms of P. car- bunculus Valenciennes in Cuvier & Valen- ciennes, 1833. Counts and measurements follow Hubbs & Lagler (1958). Institutional abbreviations are taken from Leviton et al. (1985). A characteristic series of pimple-like pro- tuberances on the upper margin of the eye (Fig. 1), with one elongated as a simple ten- tacle, is present in the holotype of Euryce- phalus carbunculus (MNHN 6875). This series is also readily visible in the putative Dorsal view of head of holotype of Eurycephalus carbunculus, MNHN 6875. Scale bar indicates 10 types P. bobossok (RMNH 5919) and the syntypes of P. maculosus (ZMB 5145) but, unfortunately, cannot be seen on the dried skins (syntypes) of P. cantori (BMNH 1860.3.19.268-9). However, the mottled coloration on the back and sides of the body (Fig. 2), short snout, lack of any interoper- cular flaps, presence of a preorbital spine, supraorbital ridge well-serrated above the eye, and three or more suborbital spines confirm that the putative syntypes of P. cantori are identifiable as E. carbunculus. The same combination of characters is ev- ident in the type specimens of P. bobossok and P. maculosus. A comparison of counts and proportional measurements of the ho- lotype of E. carbunculus and the syntypes of P. cantori appears in Table. 1. The close agreement in proportional measurements and the combination of 11 dorsal soft rays, 12 anal-fin rays, 19 pectoral-fin rays and a 386 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Lateral view of lectotype of Platycephalus cantori (upper, 133.6 mm SL) BMNH 1860.3.19.268 and paralectotype (lower, 135.4 mm SL) BMNH 1860.3.19.269. pelvic-fin ray count of 1,5 are further evi- dence of their strong similarity. It seems appropriate here to designate lectotypes for Platycephalus cantori Bleek- er and P. maculosus Peters. We select BMNH 1860.3.19.268 (134 mm SL, caudal fin largely broken, interorbital space entire) as the lectotype and BMNH 1860.3.19.269 (135 mm SL, caudal fin mostly complete, interorbital space truncated) as the paralec- totype of P. cantori. ZMB 5145 (93 mm SL) is selected as the lectotype of P. ma- culosus Peters and ZMB 32760 (129 mm) as a paralectotype. As Peters (1869), in a footnote, compared several features of the above two specimens from Singapore with ZMB 724 (1, 103 mm) from the Celebes, we regard the latter to be part of the type series and, is here designated as a second paralectotype. Bleeker (1853:461) de- scribed Platycephalus bobossok from a sin- gle specimen (148 mm in length) from Ba- tavia. In his later revision of Platycephalus, Bleeker (1879:24) listed 4 specimens of P. bobossok (118—180 mm in length) from Ba- tavia and other localities. Three specimens (143, 146, 156 mm total length) are cur- rently found in the type collection (RMNH 5919). Of these, the specimen 146 mm in total length most closely approximates Bleeker’s holotype. Acknowledgments The authors gratefully acknowledge as- sistance in recataloging lectotypes received from Hans-J. Paepke (ZMB) and Oliver Crimmen and Anne-Marie Woolger (BMNH). Literature Cited Bleeker, P. 1853. Diagnostische beschrijvingen van nieuwe of weinig bekende vischsoorten van Ba- tavia. Tiental I-VI.—Natuurkundig Tijdschrift voor Nederlandsche Indie IV:451—516. . 1879. Revision des especes insulindiennes du genre Platycephalus.—Natuurkundige Verhan- delingen Koninklijke Akademie, v. XIX (1878): 1-31. Cantor, T. 1849. Catalogue of Malayan Fishes.—Jour- nal of the Asiatic Society of Bengal 18(2):981— 1443. Cuvier, G., & A. Valenciennes. 1829. Histoire natu- VOLUME 110, NUMBER 3 relle des poissons. E G. Levrault, Paris, vol. 4, 518 pp. , & . 1833. Histoire Naturelle des Pois- sons. E G. Levrault, Paris, v. 9:429-—512. de Beaufort, L. F, & J. C. Briggs. 1962. The fishes of the Indo-Australian Archipelago. XI. Scler- oparei, Hypostomides, Pediculati, Plectognathi, Opisthomi, Discocephali, Xenoterygii. E. J. Brill, Leiden, 1-481. Giinther, A. 1860. Catalogue of the Acanthopterygian fishes in the collection of the British Museum. 2. Squamipinnes, Cirrhitidae, Triglidae, Tra- chinidae, Polynemidae, Sphraenidae, Trichiuri- dae, Scombridae, Carangidae, Xiphiidae. Lon- don, xxi + 548 pp. Hubbs, C. L., & K. E Lagler. 1958. Fishes of the Great Lakes Region.—Cranbrook Institute of Science Bulletin 26:213 pp. 387 Imamura, Hisashi. 1966. Phylogeny of the Family Platycephalidae and Related Taxa (Pisces: Scor- paeniformes).—Species Diversity 1(2):123- 23Se Leviton, A. E. et al. 1985. Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology.—Copeia 1985(3): 802-832. Murty, V. S. 1982. On the fishes of the family Pla- tycephalidae of the seas around India.—Journal of the Marine Biological Association of India (1975) 17(3):679-694. Peters, W. (C.H.). 1869. Uber die von Hrn. Dr. EF Ja- gor in dem ostindischen Archipel gesammelten und dem Konigl. Zoologischen Museum uber- gebenen Fische.—Monatsberichte der Konig- lich-Preussische Akademie der Wissenschaften zu Berlin for 1868:254-281. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):388—392. 1997. Pseudothelphusa ayutlaensis, a new species of freshwater crab (Crustacea: Brachyura: Pseudothelphusidae) from Mexico Fernando Alvarez and José Luis Villalobos Colecci6n Nacional de Crustaceos, Instituto de Biologia, Universidad Nacional Aut6noma de México, Apartado Postal 70-153, México 04510 D.F, México Abstract.—Pseudothelphusa ayutlaensis, new species, is described from the State of Guerrero, Mexico. The new species is placed in the genus Pseudo- thelphusa based on the presence of a first gonopod with the characteristic broadly rounded mesial process and a well developed subtriangular lateral pro- cess. The unique orientation of the mesial and lateral processes of the first gonopod distinguishes P. ayutlaensis from other species in the genus. The genus Pseudothelphusa de Saussure, 1857, is one of the most diverse within the family Pseudothelphusidae Rathbun, 1893, with 22 species (Alvarez & Villalobos 1996, Alvarez et al. 1996), distributed ex- clusively in Mexico, and one, P. puntarenas Hobbs, 1991, from Costa Rica. The genus is distributed along the Pacific slope from Sonora to Guerrero, throughout central Mexico, and along the Gulf of Mexico Slope in Veracruz (Rodriguez 1982, Alvarez 1989). Although species of Pseudothelphu- sa exhibit a great variety of gonopod mor- phologies, all exhibit a broadly rounded mesial process and subtriangular lateral lobe (Rodriguez 1982). In the majority of species of Pseudothelphusa the mesial pro- cess of the first gonopod is oriented proxi- mally, descending from the apex towards the base of the gonopod and is reniform in shape, while the lateral process projects lat- erally. In P. ayutlaensis, new species, the mesial process is oriented distally, extend- ing upwards from the apex; and the lateral process is projected cephalically. All the specimens are deposited in the Coleccion Nacional de Crustaceos, Instituto de Biol- ogia, Universidad Nacional Aut6noma de México (CNCR). Carapace width and car- apace length are abbreviated as cw and cl, and expressed in millimeters. Pseudothelphusa de Saussure, 1857 _Pseudothelphusa ayutlaensis, new species Figs. 1, 2 Holotype.—¢, cw 24.3 mm, cl 16.1 mm; junction of Pinela and Tonala rivers, Mun- icipio de Ayutla de los Libres, Guerrero (16°52'N, 99°12’W), 18 Dec 1987, coll. J. P. Gallo; CNCR 8715. Paratypes.—2 ¢, cw 23.0, 22.0 mm, cl 15.0, 14.4 mm; same locality, date, and col- lector as holotype; CNCR 8715. 2 2, cw 36.3, 28.2 mm, cl 23.5, 18.6 mm; same lo- cality, date, and collector as holotype; CNCR 8715. Description.—Dorsal surface of carapace covered with small papillae (Fig. 1a). In- ferior frontal border continuous, thick, slightly sinuous in dorsal and frontal views, extending laterally to form superior margin of orbits (Fig. 1b). Superior frontal border formed by the folding of the carapace with an irregular row of blunt tubercles, divided by median groove, inclined towards the center (Fig. 1b). Postfrontal lobes present as two distinct elevations. Median groove deep, extending posteriorly beyond post- frontal lobes. Cervical groove wide and deep, curved slightly posteriorly, reaching anterolateral margin (Fig. la). Cardiac re- gion of carapace weakly marked. Antero- lateral margin between orbit and cervical VOLUME 110, NUMBER 3 389 Fig. 1. Pseudothelphusa ayutlaensis, new species, male holotype: a, dorsal view; b, frontal view; c, ventral view. Carapace width 24.3 mm, length 16.1 mm. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 390 Poe ASE unig iew; C, mesial v b Scale bars . > iew lateral v a, ht chela . . , a-e left first gonopod ies f, left th new spec Pseudothelphusa ayutlaensis, view Fig. 2. 1 mm, f a-e = &, mg . > d maxilliped 1r . > ] apica > € . ° , caudal view d = 10 mm. . ? ic view 5 mm, cephal g VOLUME 110, NUMBER 3 groove with irregularly placed denticles, posterior to cervical groove with uniform denticles. Ratio exopod/ischium of third maxilliped 0.73. Ischium of third maxilli- ped trapezoidal in shape; merus narrower than ischium (Fig. Ic). Right chela the larg- est; fingers not gaping, teeth worn out in holotype. Both chelae with blunt denticle at base of movable finger. First gonopod slender, with twisted sperm channel (Fig. 2a). In caudal view (Fig. 2d) with constriction at two thirds its length below cavity of apex. Marginal su- ture sinuous (Fig. 2b). Marginal process ab- sent from apex of gonopod, except for one sharp tooth at base of mesial process (Fig. 2c). Mesial process rounded, oriented dis- tally (Fig. 2b). Lateral process extending in laterocephalic direction, wide proximally, tapering distally, ending in two sharp tips (Fig. 2d). Cavity of apex oval shaped (Fig. 2e). Crest of cavity of apex low caudally, higher cephalically. Etymology.—The specific name ‘“‘ayu- tlaensis’”” makes reference to the county, Ayutla de Los Libres, in the State of Guer- rero where the species is distributed. Remarks.—The new species is placed in the genus Pseudothelphusa based on the torsion of the sperm channel of the first gonopod, and the presence of a typically rounded mesial process. Pseudothelphusa ayutlaensis is closest to those species of Pseudothelphusa distributed in western Mexico that lack a marginal process in the apical region of the gonopod (e.g., P. re- chingeri Pretzmann, 1965; P. sonorae Rod- riguez & Smalley, 1969; and P. galloi A\l- varez & Villalobos, 1990) and to those spe- cies in which the marginal process is re- duced to a series of small teeth (e.g., P. jouyi Rathbun, 1893; P. lophophallus Rod- riguez & Smalley, 1969; and P. nayaritae Alvarez & Villalobos, 1994). The new spe- cies also resembles P. galloi in the shape of the lateral process of the first gonopod. Although this process is oriented differently in both species (cephalically in P. ayutlaen- sis and laterally in P. galloi), in both spe- 391 cies it is roughly triangular and ends in two sharp tips. In southern Guerrero Pseudo- thelphusa ayutlaensis and P. galloi share the same drainage system, 10 km away from each other. Acknowledgments We thank Rolando Mendoza for the drawings and Juan Pablo Gallo for donating the specimens described in this study. Literature Cited Alvarez, FE 1989. Smalleyus tricristatus, new genus, new species, and Pseudothelphusa parabelli- ana, new species (Brachyura: Pseudothelphusi- dae) from Los Tuxtlas, Veracruz, Mexico.— Proceedings of the Biological Society of Wash- ington 102:45—49. , & J. L. Villalobos. 1990. Pseudothelphusa galloi, a new species of freshwater crab (Crustacea: Brachyura: Pseudothelphusidae) from southwestern Mexico.—Proceedings of the Biological Society of Washington 103: 103-105. oars . 1994. Two new species and one new combination of freshwater crabs from Mexico (Crustacea: Brachyura: Pseudothelphu- sidae).—Proceedings of the Biological Society of Washington 107:729—737. , & . 1996. Especie nueva de cangrejo de agua dulce del Género Pseudothelphusa (Brachyura: Pseudothelphusidae) de Guerrero, México.—Anales del Instituto de Biologia, Universidad Nacional Aut6noma de México, Serie Zoologia 67(2):297—302. : , & E, Lira. 1996. Decapoda. Pp. 103-129 in J. Llorente, A. N. Garcia-Aldrete, & E. Gonzalez, eds., Biodiversidad, Taxonomia y Biogeografia de Artré6podos de México: Hacia una Sintesis de su Conocimiento. Instituto de Biologia, Universidad Nacional Aut6noma de México-CONABIO, 660 pp. Hobbs, H. H., III. 1991. A new pseudothelphusid crab from a cave in southern Costa Rica (Decapoda: Brachyura).—Proceedings of the Biological So- ciety of Washington 104:295-298. Pretzmann, G. 1965. Vorlaufiger Bericht tiber die Familie Pseudothelphusidae.—Anzeiger der Mathematisch-Naturwissen-Schaftlichen Klasse der Osterreichische Akademie der Wissenschaf- ten 179:14-—24. Rathbun, M. J. 1893. Descriptions of new species of American freshwater crabs.—Proceedings of the United States National Museum 16:649-— 661. 392 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rodriguez, G. 1982. Les crabes d’eau douce del Instituto de Biologia, Universidad Nacional d’Amérique. Famille des Pseudothelphusi- Auténoma de México 40:69-112. dae.—Faune Tropicale 22:1—223. Saussure de, H. 1857. Diagnoses de quelques Crus- , & A. E. Smalley. 1969. Los cangrejos de tacés nouveaux des Antilles et du Mexique.— agua dulce de México de la familia Pseudo- Revue et Magazin de Zoologie Pure et Appli- thelphusidae (Crustacea: Brachyura).—Anales quée 9:304—306. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):393-398. 1997. Austinixa, a new genus of pinnotherid crab (Crustacea: Decapoda: Brachyura), with the description of A. hardyi, a new species from Tobago, West Indies Richard W. Heard and Raymond B. Manning (RWH) University of Southern Mississippi, Institute of Marine Sciences, Ocean Springs, Mississippi 39566-7000, U.S.A.; (RBM) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—The new genus Austinixa is recognized for Pinnixa cristata Rath- bun and six other species formerly assigned to Pinnixa White, 1846. An eighth species, A. hardyi, new species, is described from Tobago, West Indies. Mem- bers of Austinixa can be distinguished from species of Pinnixa sensu stricto by the transverse ridge that completely crosses the posterior surface of the cara- pace, the strongly deflected fingers of the chelae, and the much longer, slen- derer, and smoother third walking leg. Members of Austinixa often are asso- ciated with callianassid shrimps. Austinixa hardyi, new species, found associ- ated with a member of the callianassid genus Callichirus Stimpson, 1866, is the only described species of Austinixa having dense patches of setae on the carapace of the male. Collections of infaunal decapods from Tobago were made by one of us (R.H.) in 1992 and 1993. One of the species collected proved to be an undescribed member of the pinnotherid genus Pinnixa White, 1846. It is named and it and related species are placed in a new genus below. Abbreviations used in the accounts in- clude: Al, antennule; cl, carapace length; Mxp3, third maxilliped; Pl, cheliped; P2- 5, first to fourth walking legs. Carapace size is expressed as length 2 > 3; third article markedly lon- ger on medial than lateral side. Basal article stout, bearing longitudinal row of setae and with curved setose ridge extending from base of stylocerite to groove between sty- locerite and basal article. Stylocerite (Fig. 2c) strong, reaching to distal edge of second peduncular article on medial side, gently curved on lateral side but nearly straight to only slightly curved along medial border. Flagella inserted side by side but usually with lateral flagellum crossing over medial flagellum, which curves downward and backward. Peduncular and flagellar articles with setae as illustrated. Antennae (Fig. la, 2b, d, e) with thick, stout peduncular articles. Flagellum usually sweeping backward, ranging in length from slightly shorter to slightly longer than car- apace length. Antennal scale (Fig. 2d, e) large, broadly oval, setose only along me- dial and distal border, with supportive lon- gitudinal dorsal ridge and lacking notch PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON along distolateral border. Base of antennal scale and peduncle (Fig. 2e) not forming groove to receive leading edge of carapace border. Mandibles (Fig. 3a, b) with 2-segmented palp; first segment with 1 long plumose seta on distodorsal border; second segment heavily setose. Cutting edge (incisor pro- cess) gently tapering, slightly produced, with 5—6 small sharp teeth giving way to row of smaller teeth along descending (ven- tral) border. Posterior tooth (molar process) blunt, simple, divergent from incisor pro- cess. Maxillule (Fig. 3c) with palp well devel- oped, incipiently bilobed, bearing 2 small distal short setae and 1 longer, plumose sub- terminal seta. Basal endite stout, curved strongly inward, with innermost setae in well defined row sweeping backward and inward, and with setae around distal and medial borders. Proximal endite large, nar- row basally (at point where palp is at- tached) but expanded distally and bearing some 20 to 30 stout spines along medial border. Maxilla (Fig. 3d) with endites reduced, nearly obsolescent. Basal endite composed of 2 roughly similar setose lobes; distal en- dite narrow basally and expanded distally, fringed with setae. Palp narrow, following curve of dorsal edge of distal endite. Sca- phognathite large, flattened, bearing dense- ly plumose setae on all borders, expanded distally, giving rise at posterior terminus to many long, stout, microscopically serrate setae that we presume sweep over and clean gill surfaces in life. Blade with scattered short setae. First maxilliped (Fig. 3e) reduced, with components nearly completely fused into flattened, triangular, phylliform limb; basal and distal endites setose; palp narrow, ex- tending beyond endites. Epipod short, stout, incipiently bilobed, with anterior dorsal bulge and posteroventral triangular termi- nus. Second maxilliped (Fig. 3f) flattened but becoming pediform, composed of 5 heavily VOLUME 110, NUMBER 3 403 SS = Fig. 2. Rimicaris aurantiaca, new species, female holotype, LACM 93-46.3. a, frontal region, dorsal view; b, left frontal region, lateral view; c, base of right antennule and its stylocerite, dorsal view; d, right antennal scale, dorsal view; e, same, lateral view; f, telson and right uropods, dorsal view. Scale bar = 2.0 mm. 404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SSS = S ; WS =<, i) Wi WS | 1 Ni \\ Mi Ik R WN\\ A 4 \ \\ Why XN \ Ret | HN iN SAN | wW \\ ly Rimicaris aurantiaca, new species, female holotype, LACM 93-46.3. a, b, right and left mandibles; Big..3: c, maxillule; d, maxilla; e, first maxilliped; f, second maxilliped. Not drawn to scale. VOLUME 110, NUMBER 3 setose articles. Dactylus with distinct row of short, evenly-sized setae forming brush along medial edge. Epipod strongly arched dorsally, with small protruberance at apex of arch. Third maxilliped (Fig. 4a, b, 5a, b) ped- iform, composed of 2 short proximal arti- cles and 3 longer more distal articles; bas- almost of 2 proximal articles bearing ar- throbranch and small bilobed epipod. Distal 3 articles (articles 3, 4, and 5, numbered proximal to distal) differning in length such that 3 > 5 > 4 (with 5 being distalmost article, presumably dactylus or fused dac- tylus + propodus). Distal 2 articles with well developed rows of stout setae (Fig. 4b); dactylus with stout spines distally. Dis- tal spines and setae including variety of ser- rate, stout, plumose, and other types of spines and setae (Fig. 5a, b). Pereiopod 1 (Fig. 4c, d) short, stout, ap- pearing slightly twisted. Chela stout, curved downward and inward; movable finger (dactylus) approximately % length of prop- odus, and bearing comb row of minute spines along cutting edge. Carpus expanded distally, lacking well developed cluster of cleaning spines or setae at distoventral bor- der. Pereiopod 2 (Fig. 4e, f, 5c) slender, ap- proximately equal in length to pereiopod 1. Chela with row of spines on cutting edges of dactylus and propodus; these spine rows on each finger terminating in long spine di- rected toward opposing finger; spines over- reaching opposing finger when closed (Fig. 4f, 5c). Length of dactylus approximately half that of propodus. Pereiopods 3—5 (Fig. 4g) stout, similar to one another, slightly increasing in length from P3 to P5. Propodus with transverse rows of setae along ventral border. Dactylus short, stout, recurved, bearing numerous distal and ventral spines (Fig. 4h, 5d), and with distinct basal keel nearly obscured by protruding spines of propodus (Fig. 5e). Coxae of PS with small, ventrally- and slightly anteriorly-directed spine between them (Fig. 4i, 5f); spine more or less 405 straight on ventral border but bulging up- ward (toward body) along anterodorsal bor- der (Fig. 41). Pereiopods lacking exopods. Gill formula: pleurobranchs on pereio- pods 1-5, arthrobranchs on maxilliped 3 and on pereiopods 1-4. Abdomen (Fig. la) gently curving to nearly straight behind carapace. Abdominal pleura of somite 2 expanded and covering those of somites 1 and 3; posteroventral borders of pleura of somites 2-5 becoming increasingly acute from somite 2 to somite 5, but always smooth edged, lacking den- ticles or serrations. Telson (Fig. 2f) with 8 or 9 spines on each side in row beginning at proximal fourth of telson and extending posteriorly and laterally; progressively more posterior spines directed more laterally than dorsally. Extremity of telson with pair of heavy spines flanking row of shorter and thinner spines and setae. Uropods elongate, oval, lower branch with well developed diaresis; both rami heavily setose. Measurements in mm.—Total length of the 20 specimens examined (including the holotype) ranged from 21.5 to 26.8 mm. Color.—In life the species is bright or- ange to reddish orange, with the ocular re- gion reflecting light and appearing either gray or bright white (Nuckely et al. 1996: 101, figs. 2B, C), depending upon the angle of reflected light. Storage in ethanol causes these colors to fade, although a light orange color of the carapace and abdomen was still detectable some 28 months after preserva- tion. Most obviously pigmented are the ar- eas where the cuticle is more transparent; these include an oval region on the dorsal surface of the carapace that also contains the ocular apparatus (Fig. 1b), a slightly ventrally protruding area of the sternal cu- ticle just posterior to the fifth pereiopods and anterior to the first abdominal somite, and to a lesser extent the dorsal and lateral surfaces of each abdominal somite. Through each of these areas small orange- 406 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ Nh | x \\M ay | | (\" Fig. 4. Rimicaris aurantiaca, new species, female holotype, LACM 93-46.3. a, right third maxilliped, lateral view; b, magnified view of same, medial view; c, right pereiopod 1 (cheliped), lateral view; d, higher magnifi- cation of same, medial view; e, pereiopod 2; f, same, higher magnification of chela and carpus; g, right pereiopod 3, lateral (posterior) view; h, same, higher magnification of dactylus. Scale bar = 2.0 mm (a, c, e, g), 1.0 mm (bd. iis VOLUME 110, NUMBER 3 : . a iJ Adee. Ne He > 4, Fig. 5. 407 Rimicaris aurantiaca, new species, selected SEM images. a, b, distalmost article of third maxilliped, showing diversity of setal and spine types; c, chela of right pereiopod 2 (note long spine at tip of each finger); d, dactylus and part of propodus of left pereiopod 4, lateral view; e, same, higher magnification of ventral keel (arrow in d). f, spine between coxae of pereiopod 5, anterior is toward left of photograph. Scale bars: = 200 um (a, c), 100 um (b, e), and 500 (d, f). Sizes in mm of the two specimens used in these photographs were as follows: carapace length = 10.1, total length = 26.7, carapace width = 5.4 for the larger; carapace length = 9.5, total length = 26.1, carapace width = 5.2 for the smaller. colored oil droplets are visible even in pre- served specimens. Etymology.—The specific name is from the adjectival and feminine form of auran- tium, a Neolatin neuter noun meaning orange (Brown 1955: 207). This choice of epithet is in reference to the distinctive coloration of this species in life. The name also honors Syracuse University, students and alumni of which are nicknamed the ‘“‘Orangemen,”’ where the original research on the fascinating visual components of the new species was completed at the Department of Bioengineer- ing and Neuroscience and at the Institute of Sensory Research (e.g., O’Neill et al. 1995, Nuckley et al. 1996). 408 Remarks.—The details of the unique ‘enlarged dorsal eye specialized for detect- ing light in a very dim environment instead of the expected compound eye”’ possessed by this species have been presented by Nuckley et al. (1996). This organ is visible in life (see color photographs in Nuckley et al. 1996) as a branched lobe extending backward from the front of the carapace, and just below the surface of the carapace, with each lobe being paddle-shaped and posteriorly rounded. This organ in R. au- rantiaca differs from that in R. exoculata in that in the latter species it is considerably more elongate (e.g., Van Dover et al. 1989). Discussion It is reassuring to find that previously noted differences in neuroanatomy, physi- ology, and ecology are in agreement with taxonomic separations based on morpho- logical characters. Nuckley et al. (1996), based primarily on the details of the unusu- al visual apparatus, referred to this new spe- cies as Rimicaris sp., feeling that it was more similar to R. exoculata than to any described species of the closely related ge- nus Chorocaris. They also commented on ecological differences between Chorocaris and Rimicaris, noting that Rimicaris sp. (now R. aurantiaca, new species) occurs in dense swarms at the vent site, as does R. exoculata, whereas no species of Choro- caris exhibits this behavior, or at least not to this degree. The new species shares with R. exoculata the highly unusual dorsal eye, with very similar retinal anatomy (O’ Neill 1995, Nuckley et al. 1996, S. Chamberlain, pers. comm.). Although species of Choro- caris share some of the same visual com- ponents, there are important neuroanatom- ical differences, the most salient of which is that the visual array is always oriented anteriorly (rather than dorsally) in all spe- cies of Chorocaris examined to date (Kuen- zler et al. 1997), whereas both R. exoculata and R. aurantiaca have a dorsally directed visual array that receives input through the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cuticle of the shrimp’s carapace (Kuenzler et al. 1997, and S. Chamberlain, pers. comm.). This difference may be tied to the observed differences in behavior. Shrimp capable of receiving optical input only from the anterior end, which could be blocked by swarming behavior, might be less likely to exhibit swarming than would a species with a dorsal eye, where optical input might be obstructed to a somewhat lesser degree by swarming. The new species also shares with R. ex- oculata some small but significant external morphological details, such as an antennal scale that has a smooth (unnotched) border on its anterolateral margin, a cheliped car- pus lacking a well developed cleaning brush (i.e., with at most two or three setae in the location where such a brush is found in other carideans, including all known spe- cies of Chorocaris; Martin et al. unpub- lished data), and a distinctive brush-like pad of setae on the distal segment of the second maxilliped (see also Van Dover et al. 1988). This last character, the scraping setal brush of the second maxilliped, is potentially of high interest. According to Van Dover et al. (1988) this brush in R. exoculata is used to scrape bacteria from the more anterior ap- pendages (the first two pereiopods and the third maxilliped), and although the evi- dence to date remains somewhat equivocal, R. exoculata may depend mostly or even exclusively on vent bacteria for nutrition (see Van Dover et al. 1988, Casanova et al. 1993, Gebruk 1993, Segonzac et al. 1993, Van Dover 1995). Thus, this character may be an important generic character separat- ing the two genera on morphological as well as ecological and functional grounds. The new species differs from R. exocu- lata in ecology, coloration, and morpholo- gy. Rimicaris aurantiaca prefers an ambient temperature of about 10°C compared to a higher ambient temperature of 28°C pre- ferred by R. exoculata (Nuckley et al. 1996); they are also less active within a swarm than are individuals of R. exoculata and are only rarely seen swimming singly VOLUME 110, NUMBER 3 in the water column. According to Nuckley et al. (1996), “‘the small orange shrimp ag- gregates in swarms of hundreds, probably thousands of individuals on the sides of black smoker chimneys at the Beehive Mound of the Snake Pit site at a depth of 3500 meters.”’ Coloration is markedly different, both from R. exoculata and from previously de- scribed species of Chorocaris, with the new species appearing bright orange compared to a drab whitish or gray color exhibited by R. exoculata and Chorocaris. The color ap- pears to come from the numerous oil drop- lets visible through the cuticle of the shrimp, and indeed upon dissection the shrimp exudes some of these droplets, which remain orange-colored. Although Creasey et al. (1996) attribute the orange coloration to the presence of “‘an oily, lipid- rich hepatopancreas,”’ the oil droplets ob- viously occur in areas where no hepatopan- creas is found, as well as in the region of the hepatopancreas. Photographs in Nuck- ley et al. (1996: 101, fig. 2A—C) show the orange coloration and the difference in col- or from the sympatric and light gray col- ored R. exoculata very clearly. It is inter- esting to note that in the description of Opaepele loihi from hydrothermal vents on the Loihi Seamount of Hawaii, Williams & Dobbs (1995) refer to the color of that spe- cies as ““intensely orange (astaxanthin pig- ment),’’ although in the case of O. loihi the color comes apparently from an accumula- tion of particles of iron oxyhydroxide (Wil- liams & Dobbs 1995). Morphological differences in the new species include a “‘normal’”’ (not greatly in- flated) carapace; eyestalks that, although re- duced and fused medially, are closer to the original caridean eyestalk condition and are at least recognizable as such; a frontal re- gion that does not form a protective oper- culum with the carapace; a third maxilliped composed of three longer and two shorter articles; and a small, blunt rostrum. All of these characters are more consistent with previous descriptions of species in the ge- 409 nus Chorocaris Martin & Hessler, 1990. In- deed, if we employ the most recent key to the genera of the Bresiliidae (Williams & Dobbs 1995) the new species keys out as a member of the genus Chorocaris. The char- acters that Martin & Hessler (1990) used to distinguish Rimicaris from their newly de- scribed genus Chorocaris seem to be, for the most part, unique to R. exoculata. Rimicaris aurantiaca had been observed previously by Segonzac et al. (1993), who referred to the Beehive Mound at this site as being “densely covered with adult Rim- icaris and juveniles identified by their red color’ (English translation). At least some of these “juveniles”? we now know to rep- resent this new species, and this was in fact suspected by Segonzac et al., who noted, in an addendum to that paper, that a “‘new spe- cies with features intermediate between Rimicaris exoculata and Chorocaris chac- ei’ was present at this site, and that the juvenile stages were very similar in the three species. However, there are sufficient differences between adult R. exoculata and R. aurantiaca that there can be no doubt as to their separate identity. Special mention should be made of the recent study by Creasey et al. (1996) on genetic composition of populations of R. exoculata. In that paper, the following men- tion is made of the new species: “ . . . With- in the Snake Pit vent field, small shrimp with an orange colouration due the presence of an oily, lipid-rich hepatopancreas have been observed within swarms of R. exocu- lata’ (Creasey et al. 1996: 474). In that same paragraph these authors refer to the small orange Snake Pit species (undoubtedy R. aurantiaca) as Chorocaris sp., citing Van Dover (1995). However, it should be pointed out that Creasey et al. (1996) did not use any Snake Pit specimens in their comparison, and that the specimens they re- fer to in the remainder of their paper as Chorocaris sp. were from the TAG and Broken spur sites and therefore are attrib- utable to Chorocaris fortunata Martin & Christiansen, 1995. Thus their conclusion 410 ‘“‘that all morphotypes of R. exoculata ex- amined, including those previously inter- preted as representing separate species, are conspecific’”’ does not apply to R. aurantia- ca, but rather only to the two different pop- ulations of R. exoculata at the TAG and Broken Spur sites, the latter of which had been suggested by Murton et al. (1995) to contain a different species of Rimicaris based on slight morphological differences. The somewhat intermediate assemblage of characters (i.e., some shared with R. ex- oculata and others with species of Choro- caris) might justify creation of yet another bresiliid shrimp genus from the hydrother- mal vents. We refrain from doing so in this paper, believing that discovery of additional species is almost certainly forthcoming and will shed light on the entire assemblage, and that a conservative approach is war- ranted until such time that more is known. In the meantime we recognize that the ge- neric diagnosis for Rimicaris as emended in this paper leaves it a rather poorly delimited genus on morphological grounds, as it must be to accommodate two shrimp species that exhibit so many morphological differences. Unfortunately, description of the new species does not add appreciably to the body of information that would eventually lead to recognition or rejection of the fam- ily Alvinocarididae, as proposed by Chris- tofferson (1989) and employed by Chris- tofferson (1991) and Saint Laurent (1993, in Segonzac et al. 1993) to accommodate the genera and species of bresiliids known from hydrothermal vents and cold seeps. Thus, as have Martin & Hessler (1990), Chace (1992), Holthuis (1993), Williams & Dobbs (1995), Martin & Christiansen (1995), and Van Dover (1995), we retain the older, albeit recognized to be somewhat artificial, limits of the caridean family Bre- siliidae. Acknowledgments This work was funded by the National Science Foundation via grant number ESI PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 9552906 to K. Wise and S. Lafferty in sup- port of the Museum Research Apprentice Program of the Natural History Museum of Los Angeles County, and by a Research Experiences for Undergraduates award (DEB 9642010, supplemental to DEB 9320397) to J. W. Martin. We thank the staff of the Education Branch of the Natural History Museum, especially S. Lafferty and J. Trochez, and the other participants in the MRAP program, as well as Todd Zimmer- man, George Davis, and Karen Wise in the museum’s Research and Collections Banch, for their assistance, support and encourage- ment. We also thank Alicia Thompson for her help with the SEM component of the study, Cindy Lee Van Dover for providing the specimens upon which this report is based, and especially S. C. Chamberlain, R. N. Jinks, P. J. O’Neill, D. J. Nuckley, and other members of the Department of Bioen- gineering and Neuroscience and the Insti- tute for Sensory Research at Syracuse Uni- versity for keeping us informed as to de- velopments in their neurological and phys- iological studies of vent shrimp. The manuscript was significantly improved by comments from Rafael Lemaitre, Austin Williams, Ted Bayer (etymology of the spe- cies name), and Cindy Van Dover, to all of whom we are grateful. Literature Cited Brown, R. W. 1956. Composition of scientific words. Smithsonian Institution Press, Washington, D.C., 882 pp. Casanova, B., M. Brunet, & M. Segonzac. 1993. Limpact d’une épibiose bactérienne sur la mor- phologie fonotionelle de crevettes associées a Vhydrothermalisme médio-atlantique.—Cahiers de Biologie Marine 34:573-588. Chace, FE A., Jr. 1992. On the classification of the Caridea (Decapoda).—Crustaceana 63:70-—80. Christoffersen, M. L. 1986. Phylogenetic relationships between Oplophoridae, Atyidae, Pasiphaeidae, Alvinocarididae fam. n., Bresiliidae, Psalido- podidae and Disciadidae (Crustacea Caridea Atyoidea).—Boletim de Zoologia, Universidade de Sao Paulo 10:273-—281. . 1991. A new superfamily classification of the Caridea (Crustacea: Pleocyemata) based on VOLUME 110, NUMBER 3 phylogenetic pattern.—Zeitschrift fur Zoologis- che Systematik und Evolutionsforschung 28(1990):94—106. Creasey, S., A. D. Rogers, & P. A. Taylor. Genetic comparison of two populations of the deep-sea vent shrimp Rimicaris exoculata (Decapoda: Bresiliidae) from the Mid-Atlantic Ridge.—Ma- rine Biology 125:473—482. Galkin, S. V., & L. I. Moskalev. 1990. Hydrothermal fauna of the Mid-Atlantic Ridge.—Okeanologia 30:842-847. Gebruk, A. V., N. V. Pimenov, & A. S. Savvichev. 1993. Feeding specializations of bresiliid shrimps in the TAG site hydrothermal commu- nity.—Maarine Ecology Progress Series 98:247— 253. Holthuis, L. B. 1993. The Recent genera of the cari- dean and stenopodidean shrimps (Crustacea, Decapoda), with an appendix on the order Am- phionidacea.—Nationaal Natuurhistorisch Mu- seum, Leiden, Netherlands, 328 pp. Kuenzler, R. O., J. T. Kwasniewski, R. N. Jinks, R. C. Lakin, B.-A. Battelle, E. D. Herzog, L. Kass, G. H. Renninger, & S. C. Chamberlain. 1997. Retinal anatomy of new bresiliid shrimp from the Lucky Strike and Broken Spur hydrothermal vent fields on the Mid-Atlantic Ridge. Journal of the Marine Biological Association (United Kingdom) (In press). Martin, J. W., & J. C. Christiansen. 1995. A new spe- cies of the shrimp genus Chorocaris Martin and Hessler, 1990 (Crustacea: Decapoda: Bresili- idae) from hydrothermal vent fields along the Mid-Atlantic Ridge.—Proceedings of the Bio- logical Society of Washington 108:220—227. , & R. R. Hessler. 1990. Chorocaris vandov- erae, a new genus and species of hydrothermal vent shrimp (Crustacea, Decapoda, Bresiliidae) from the western Pacific.—Contributions in Sci- ence, Natural History Museum of Los Angeles County 417:1-11. Murton, B. J.. C. L. Van Dover, & E. Southward. 1995. Geological setting and ecology of the Broken Spur hydrothermal vent field 29°10'N on the Mid-Atlantic Ridge. Pp. 33-41 in L. M. Parson, C. L. Walker, & D. R. Dixon, eds., Hy- drothermal vents and processes. Special Publi- cations of the Geological Society of London, Geological Society, London, 000 pp. Nation, J. L. 1983. A new method using hexamethyld- 411 isilazane for preparation of soft tissues for scan- ning electron microscopy.—Stain Technology 38:347-351. Nuckley, D. J., R. N. Jinks, B.-A. Battelle, E. D. Her- zog, L. Kass, G. H. Renninger, & S. C. Cham- berlain. 1996. Retinal anatomy of a new spe- cies of bresiliid shrimp from a hydrothermal vent field on the mid-Atlantic Ridge.—Biolog- ical Bulletin 190:98—110. O’Neill, P. J., R. N. Jinks, E. D. Herzog, B-A. Battelle, L. Kass, G. H. Renninger, & S. C. Chamberlain. 1995. The morphology of the dorsal eye of the hydrothermal vent shrimp, Rimicaris exocula- ta.—Visual Neuroscience 12:861—875. Segonzac, M. 1992. The hydrothermal vent commu- nities of the Snake Pit area (Mid Atlantaic Ridge; 23°N, 3480 m).—Comptes Rendus Heb- domadaires des Séances de 1’ Académie des Sci- ences, Paris 314:593—600. , M. de Saint Laurent, & B. Casanova. 1993. L’énigme du comportement trophique des crev- ettes Alvinocarididae des sites hydrothermaux de la dorsale médio-atlantique.—Cahiers de Biologie Marine 34:535—571. Van Dover, C. L. 1995. Ecology of Mid-Atlantic Ridge hydrothermal vents. Pp. 257-294 in L. M. Parson, C. L. Walker, & D. R. Dixon, eds., Hydrothermal Vents and Processes.—Geologi- cal Society Special Publication 87. , B. Fry, J. E Grassle, S. Humphris, & P. A. Rona. 1988. Feeding biology of the shrimp Rimicaris exoculata at hydrothermal vents on the Mid-Atlantic Ridge.—Marine Biology 98: 209-216. , E. Z. Szuts, S. C. Chamberlain, & J. R. Cann. 1989. A novel eye in “‘eyeless’’ shrimp from hydrothermal vents of the Mid-Atlantic Ridge.—Nature 337:458—460. Williams, A. B. 1988. New marine decapod crusta- ceans from waters influenced by hydrothermal discharge, brine and hydrocarbon seepage.— Fisheries Bulletin 86:263—287. , & EC. Dobbs. 1995. A new genus and spe- cies of caridean shrimp (Crustacea: Decapoda: Bresiliidae) from hydrothermal vents on Loihi Seamount, Hawaii.—Proceedings of the Biolog- ical Society of Washington 108:228—237. , & P. Rona. 1986. Two new caridean shrimps (Bresiliidae) from a hydrothermal vent on the Mid-Atlantic Ridge.—Journal of Crustacean Bi- ology 6:446—462. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):412—416. 1997. Occurrence of three species of mud shrimps in aquiculture ponds on Caribbean coasts of Venezuela and Colombia, with a redescription of Upogebia omissago Williams, 1993 (Decapoda: Upogebiidae) Austin B. Williams National Marine Fisheries Service Systematics Laboratory, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, U.S.A. Abstract.—Three species of mud shrimps, Upogebia brasiliensis, U. omissa, and U. omissago, are reported from ponds maintained for the commercial cul- ture of penaeid shrimps on the Caribbean coasts of Venezuela and Colombia, South America. Upogebia omissago is redescribed and illustrated based on sexually mature individuals collected from the ponds. Catalogued lots of each species collected are listed. Three species of mud shrimps, Upogebia brasiliensis Holthuis, 1956, U. omissa Go- mes Corréa, 1968, and U. omissago Wil- liams, 1993 have been found in discrete or intermingled populations in ponds main- tained for the commercial culture of penae- id shrimps on the Caribbean coasts of Ven- ezuela and Colombia, South America. The ponds are stocked by pumping or flooding with seawater at appropriate tidal stages, and numerous organisms, including larvae and perhaps juveniles of the mud shrimps, are introduced with the incoming water. Upon attainment of the juvenile stage, the mud shrimps burrow into substrate of the ponds. Supplemental aquicultural feeding promotes growth of the cultured penaeid shrimps as well as the mud shrimps, pro- ducing sexually mature individuals of the latter in at least some cases. The description of Upogebia omissago Williams, 1993 was based on only five sub- adult specimens from Luis Correia, Praia do Coqueiro, Piaui, northeastern Brazil. Now that larger fully mature specimens are avail- able which exhibit features not completely developed in the originally described type series, the description can be emended. Re- marks on other upogebiid species that occur in these ponds are included. Specimens examined are deposited in the crustacean research collections of the Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C., (USNM), and the University of Southwest- ern Louisiana Department of Zoology, La- fayette (USLZ). Upogebia omissago Williams, 1993 Fig. 1 Material studied.—Venezuela: USNM 251468, 6 6, 2 2 ovig., shrimp farm *““Siembra Mar,’’ approximately 3 km from airport of Barcelona [Estado Anzoategui], from R. Lemaitre, 27 Oct 1992. USLZ 3581, 1 45,3 2 ovig., ‘““Siembra Mar,”’ pond 09, coll. Sergio Nates, 16 Dec 1992; USLZ 3583, 3 2, same, 14 Jun 1994; USLZ 3586, 5 3, 5 @, same, coll. Eduardo Viso, Apr 1993. Blanco Rambla (1995) also listed U. omissago from this locality. Diagnosis.—Projections to either side of rostrum ending in acute spine. Postocular spine present. Abdominal sternites un- armed. Telson subrectangular. Carpus of cheliped with | long strong inferior spine and 1 short strong spine above it on me- siodistal margin. Merus of pereopod 2 bear- ing 1 proximal mesioventral spine and 1 subdistal dorsal spine; merus of pereopod 3 VOLUME 110, NUMBER 3 with 1 subdistal dorsal spine; merus of pe- reopod 4 spineless. Description.—Rostrum triangular, short, basal width greater than median length; straight to slightly downcurved in lateral view; tip exceeding eyestalks; dorsal pair of strong subapical spines followed on each side by 2—3 remote spines and spine mesial to them at base; rostral armature merging with field of similar spines on flattened an- terior cephalothoracic shield, all spines hid- den in dense cover of setae anteriorly, but spines less strongly developed and setae less dense posteriorly; median area of ros- trum spineless but obscured by setae; spine field abruptly ending posteriorly and fol- lowed by smooth gastric region with nar- rowing median extension reaching anteri- orly between armed area to either side; pos- teriorly divergent lateral ridge bearing crest of 13—15 spines and setae, strongest on pro- cess lateral to rostrum and decreasing pos- teriorly. Shoulder lateral to cervical groove bearing 1—2 blunt or acute tubercles below intersection with thalassinidean line, latter continuing to posterior margin of carapace. Postocular spine present. Abdominal sternites unarmed. Telson subrectangular, low transverse proximal ridge confluent with inconspicu- ous lateral ridge at either side. Eyestalk stout, deepest at about mid- length in lateral view, slightly concave dor- sally, convex ventrally, horizontal to obliquely erect in repose; prominent termi- nal cornea narrower than diameter of stalk; few tiny spiniform tubercles or spinules scattered on mesiodorsal aspect of stalk posterior to cornea, and ventral margin vari- ably smooth or bearing 1—2 obsolescent spines, occasionally with single well-devel- oped spine near cornea. Antennular peduncle reaching to about midlength of terminal article of antennal peduncle; proximal 2 articles together lon- ger than terminal article. Antennal peduncle with less than % its length extending beyond tip of rostrum; ar- 413 ticle 2 bearing slender subdistal ventral spine; scale moderate, oval. Maxilliped 3 bearing epipod. Epistomial projection rather broad in lat- eral view, usually bearing 2 small unequal apical spines, sometimes | spine. Chelipeds with ventral margin of ischium bearing 1—2 spines. Merus with row of 5— 6 spines on ventral margin; single subdistal dorsal spine reaching level of postocular spine. Carpus trigonal, shallow longitudinal groove laterally, strong spine at anterior ventrolateral corner preceded by 1-2 spines; mesiodorsal crest of 6—9 crowded small spines, partly obscured by setae, in irregular row behind prominent dorsal spine on anterior margin, row flanked by 1 or 2 spines laterally and cluster of spines at proximal end mesially, and 4-6 short spines obscured by setae on anterodorsal margin mesial to articulation with propo- dus; 1 strong spine near middle of antero- mesial margin, shorter stout spine dorsal to it, and strong slender spine at distoventral corner. Chela length about 2.6—3.2 times chela height; spineless dorsal ridge termi- nating anteriorly near stout subdistal spine mesial to it, ridge not always uniformly straight; mesiodorsal row of about 8-15 small spines beginning with more or less erect spines proximally that tend to cluster at either side of row and become obsoles- cent at about %—%4 length of row; similar but more defined row of tubercles or spines below this row on upper mesial surface; distomarginal spine on mesial dactylar con- dyle and row of smaller spines ventral to it on distal margin; spine below lateral dac- tylar condyle; lower mesial surface of palm with obsolescent spines scattered or tending to form a row, and a few setae; sinuous row of crowded tubercles along proximomesial margin. Fixed finger shorter than dactyl and more slender, though stout in basal %, slightly downcurved at juncture with palm and tapering abruptly to slender tip; pre- hensile edge with 4—5 teeth; not as well de- veloped in female as in male. Dacty! lon- gitudinally ridged and setose; that of male 414 Fig. 1). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON MAY) ula r Y OAK AK WY Rl AW is XY) AY ; wi AKA ANAK) SUN Y la aut Dt 7 TANI Bin y Upogebia omissago, USNM 251468, 2, a, cephalic region, lateral; b, anterior carapace, dorsal; c, parts of abdominal segment 6, telson, and uropods, dorsal; d, cheliped, right lateral; e, chela and carpus, right mesial; f—-i, pereopods 2—5. Scale = 3 mm. with corneous tip preceded on prehensile edge by strong tooth, followed by multi- dentate crest bracketed on proximal end by larger tooth, and toothless section basally; concave mesial aspect in both sexes bearing 2 unequal rows of subcircular flattened and crowded tubercles, most numerous in upper row. Pereopod 2 reaching about to midlength of palm; carpus with acute distodorsal spine and smaller subdistal ventral spine; merus bearing slender subdistal dorsal spine and strong proximal mesioventral spine. Merus of pereopod 3 with O—1 slender distodorsal spine and ventral spines tending to cluster near ischio-meral articulation; ischium un- armed; coxa of female with low spine lat- eral to gonopore, that of male with smaller gonopore, its functional status uncertain. Pereopod 4 with merus unarmed. Pereopod 5 unarmed. Uropod with acute spine on protopod above base of mesial ramus; mesial rib of lateral ramus bearing smaller acute proxi- mal spine, distal margin of both rami bear- ing close-set row of tiny spines and spini- VOLUME 110, NUMBER 3 form granules except for short mesial sector on each. Measurements in mm.—Selected speci- mens larger than in type series; 6, anterior carapace length 10.2, carapace length 14.7, chela length 8.3, chela height 3.1; d, same, PA 66.6, 11552*5: 22 -Y. same, 9:2,. 13.7, 7.4, 2.3. Remarks.—Upogebia omissago from the Venezuelan shrimp ponds attains a larger size and is more variable in morphology than specimens in the type series from a single locality in northeastern Brazil. To comparative remarks by Williams (1993), the following should be added. The species is similar to U. vasquezi Ngoc-Ho, 1989 (distributed from SE Florida to Brazil) in having two spines on the anteromesial mar- gin of the carpus of the cheliped and in hav- ing spines on the lower mesial face of the chela palm, but in U. omissago the latter are relatively small and either scattered or in a weakly developed row compared to stronger, well aligned spines on U. vasque- zi. Upogebia omissago resembles U. car- eospina Williams, 1993 (known from Ceara, Brazil) in having the dorsal ridge of the chela spineless, but spines or tubercles in the mesiodorsal rows are far less numer- ous than in U. careospina. Upogebia om- issago differs from both of the above con- geners in having the carpus of the cheliped armed with a crest of numerous dorsal spines flanked by 1 or 2 supernumerary lat- eral spines and a cluster of crowded mesial spines at the proximal end of the row. Each of these species has a spineless merus on pereopod 4. Rostral and adjacent dorsal spines are relatively stronger on U. omis- sago than in the others, and the spineless gastric area is far more abruptly defined, with the anterior median spineless exten- sion being characteristic of U. omissago. Finally, only in U. omissago are there tiny mesiodorsal-ventral spinules and sharp tu- bercles on the eyestalks, and sharp tubercles on the shoulder of the cervical groove be- low its juncture with the thalassinidean line. Notes.—Occurrence of other species of 415 Upogebia taken from burrows in penaeid shrimp culture ponds on the Caribbean coasts of Venezuela and Colombia are as follows. Upogebia brasiliensis Holthuis, 1956: USLZ 3580, 23, 1 @ ovig., shrimp farm ““Siembra Mar,” approximately 3 km from airport of Barcelona [Estado Anzoategui], Venezuela, coll. Eduardo Viso, Aug. 1993; USLZ 3582, 3 2, same, coll. Eduardo Viso, Aug 1993; USLZ 3586, 1 2, same, coll. Eduardo Viso, Apr 1993 (occurring with U. OmMIiSSago). Upogebia omissa Gomes Corréa, 1968: USNM 251469, 4 3, 1 & ovig., shrimp farm: Colombiana de Acuacultura, S.A., pond P15, Bahia de Barbacoas, Cartagena, Colombia, coll. Sergio E Nates, 4 Feb 1992, yabby pump; USLZ 3584, 1 6,2 2 Ovig., shrimp culture pond, Universidad de Oriente, Isla de Margarita, Venezuela, coll. E. Viso & O. Pichardo, 23 Jul 1994; USLZ 3588, 2 2 ovig., same, coll. E. Viso, O. Pichardo, S. Nates, 23 Jul 1994; USLZ 3585, 1 &, ‘“‘Colombiana’”’ shrimp farm, Cartagena, Colombia, coll. D. L. Felder, 26 Get 1992: USEZ,.3587;,2..6...2, 2 vie. 1 juv., Parque Nacional Mochima, Chimana Grande, La Ensenada, Venezuela, coll. S. Nates, 26 Dec 1993. Acknowledgments I am indebted to shrimp farm owners, collectors, and institutions listed in Mate- rials Studied and Notes for providing the specimens that made this report possible. Keiko Hiratsuka Moore enhanced the text with her excellent illustrations. Critical comments on the manuscript were provided by Juan P. Blanco, B. B. Collette, D. L. Felder, and Sergio E Nates. Literature Cited Blanco Rambla, J. P. 1995. Additional records of ghost shrimps (Decapoda: Thalassinidea) from Venezuela.—Caribbean Marine Studies, 4:59— 75. Gomes Corréa, M. M. 1968. Sobre as espécies de ‘“‘Upogebia’’ Leach do litoral brasileiro, com 416 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON descrigao de uma espécie nova (Decapoda, Cal- velles de la famille des Upogebiidae (Crustacea, lianassidae).—Revista Brasiliera de Biologia, Thalassinidea).—Bulletin du Muséum National 28(2):97-109. d’Histoire Naturelle, Paris, section A, series 4, Holthuis, L. B. 1956. Three species of Crustacea De- 11(4):865—-878. capoda Macrura from southern Brazil, including Williams, A. B. 1993. Mud shrimps, Upogebiidae, a new species of Upogebia.—Zoologische Me- from the western Atlantic (Crustacea: Decapo- dedelingen, Leiden, 34(11):173-181. da: Thalassinidea).—Smithsonian Contributions Ngoc-Ho, N. 1989. Description de trois especies nou- to Zoology, No. 544:77 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):417—421. 1997. Esanpotamon namsom, a new genus and species of potamid crab (Crustacea: Decapoda: Brachyura) from a waterfall in northeastern Thailand Phaibul Naiyanetr and Peter K. L. Ng (PN) Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand; (PKLN) School of Biological Sciences, National University of Singapore, Kent Ridge, Singapore 119260, Republic of Singapore Abstract.—A new genus and species of potamid crab, Esanpotamon nam- som, is described from a waterfall in northeastern Thailand. The new genus is closest to Demanietta Bott, 1966, but can be distinguished by several important carapace, male abdominal and gonopodal characters. Waterfall crabs of the family Potamidae are extremely speciose in Thailand, and about 30 species are now known (Naiyanetr 1992, Ng & Naiyanetr 1993). Several years ago, the senior author collected a number of specimens from a waterfall in northeast- ern Thailand which were initially referred to the genus Demanietta Bott, 1966. As the authors were involved in a study of all the Thai species of Demanietta, the specimens were kept aside for this revision. A recent re-examination of the north- eastern Thai specimens showed that they represent a new species. While the external features of this new species superficially re- semble species of Demanietta, the male ab- domen and gonopods of these northern Thai specimens differ very markedly. This ob- servation, together with due consideration of their zoogeography (the genus Deman- ietta s. Ss. is known only from western and southern Thailand) has compeled the au- thors to establish a new genus for this new species. The present paper describes this new ge- nus and species, here named Esanpotamon namsom. The abbreviations G1 and G2 are utilized for the male first and second pleo- pods (gonopods) respectively. The terms used in the text follow those recommended by Ng (1988). Specimens examined are de- posited in the Zoology Collection of the Department of Biology, Chulalongkorn University, Bangkok (CUMZ); and Zoolog- ical Reference Collection, School of Bio- logical Sciences, National University of Singapore (ZRC). Taxonomy Family Potamidae Ortmann, 1896 Esanpotamon, new genus Diagnosis.—Carapace much broader than long, dorsal surface flat; postorbital and epigastric cristae confluent, rugose, very low, indistinct; anterolateral margin confluent with external orbital angle. Exo- pod of third maxilliped with well developed flagellum which is longer than maximum width of merus. Ambulatory legs short. Male abdomen broadly triangular; tip of tel- son reaching to imaginary line joining mid- points of bases of chelipeds. G1 sinuous; terminal segment subcylindrical, curving upwards (towards anterior of carapace when lying in situ), distal part sharply ta- pering, tip appearing spine-like, subdistal dorsal surfaces with numerous long, very stiff setae, proximal dorsal part with small, swollen flap. G2 longer than G1, elongated distal segment longer than half-length of basal segment. 418 Type species.—Esanpotamon namsom, new species, by present designation. Remarks.—Of all Indo-Chinese potam- ids, the external morphology of Esanpota- mon most closely approaches that of De- manietta Bott, 1966 (type species Potamon manii Rathbun, 1904) (sensu Ng & Naiya- netr 1993), especially with regards to the broad and relatively smooth, flattened car- apace, and the smooth surfaces of the che- lipeds. Thus far, four species of Demanietta are known, 1.e., D. manii (Rathbun 1904), D. smalleyi (Bott 1966), D. merguensis (Bott, 1966) and D. tritrungensis (Naiyanetr 1986); however, several additional Thai and Burmese waterfall species remain unde- scribed (unpublished data). All known De- manietta species occur along the Tenasser- im Range which borders southern Thailand and southeastern Burma, and the Phuket Range in southeastern Thailand. Esanpotamon, however, differs from De- manietta (sensu Ng & Naiyanetr 1993) in several key characters. The postorbital cris- ta of Esanpotamon is very weak and poorly defined, whereas it is distinct and sharp in Demanietta. The external orbital angle and anterolateral margins are confluent without any epibranchial tooth, whereas there is a distinct epibranchial tooth and a broad ex- ternal orbital angle present in Demanietta. Compared to known Demanietta species, the male abdomen of Esanpotamon is broadly triangular with segments 5 and 6 proportionately much broader than 9. Most significantly, the G1 is very different, with the subterminal segment gradually tapering distally, the terminal segment subcylindri- cal with a small proximal dorsal fold and has numerous long, stiff subdistal setae. In known Demanietta species, the subterminal segment has a distinct “‘neck’’, the terminal segment is conical, gently tapering, and has a broad, low dorsal fold with scattered short, soft setae (Bott 1966, 1970; Ng & Naiyanetr 1993). These differences warrant the establishment of a new genus for this new species. In addition, the distributions of Esanpotamon and Demanietta are quite PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON distinct. Demanietta is known only from western and southern Thailand, whereas Es- anpotamon occurs in northeastern Thailand. Etymology.—The generic name is de- rived from an arbitrary combination of Esan, the name of the northeastern part of Thailand where the type species occurs, and Potamon, the name of the type genus of the family Potamidae. Gender neuter. Esanpotamon namsom, new species Figs. 1, 2 Material examined.—Holotype: male (carapace width 31.3 mm, carapace length 22.7 mm), Sam Teb Waterfall, Nam Som District, Udon Thani Province, northeastern Thailand, coll. PB Naiyanetr, 26 Oct 1991 (ZRC 1997.772). Paratypes: 1 male, 3 fe- males (largest carapace width 23.4 mm, car- apace length 18.3 mm), same data as ho- lotype (ZRC 1997.773—776). 6 males, 4 fe- males same data as holotype (CUMZ). Diagnosis.—As for genus. Description.—Carapace much broader than long; dorsal surfaces almost flat, smooth, glabrous; epigastric cristae rugose, very weak (not sharp); epigastric cristae slightly anterior of postorbital cristae; epi- gastric cristae separated by broad Y-shaped groove; anterolateral margin cristate, lined with small, rounded granules, strongly con- vex; external orbital angle small, confluent with anterolateral margin, not separated by discernible cleft (sometimes with very small notch visible in smaller specimens); anterolateral and posterolateral regions lined with oblique striae, those on antero- lateral regions stronger; posterolateral mar- gins almost straight, strongly converging towards gently sinuous posterior carapace margin; frontal margin gently sinuous, with shallow, broad cleft visible when viewed frontally; supraorbital margin lined with low, rounded granules; infraorbital margin beaded with low, rounded granules; subor- bital, pterygostomial and sub-branchial regions smooth; H-shaped median carapace depression shallow but distinct, confluent VOLUME 110, NUMBER 3 419 Co PROD ann. Wed ae « oa as / a fo ag a i mire ee —_ Fig. 1. Esanpotamon namsom, new genus and species. Holotype male (carapace width 31.3 mm, carapace length 22.7 mm) (ZRC 1997.772). A, left side of carapace; B, left third maxilliped (setae denuded); C, left third ambulatory leg (setae denuded); D, left fourth ambulatory leg (setae denuded); E, carpus of right cheliped (dorsal view); E anterior thoracic sternites; G, abdomen (setae denuded). Scales = 5.0 mm. 420 eZ, via > PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Esanpotamon namsom, new genus and species. Holotype male (carapace width 31.3 mm, carapace length 22.7 mm) (ZRC 1997.772). A, left G1 (ventral view); B, left G1 (dorsal view); C, terminal segment of left G1 (ventral view); D, terminal segment of left G1 (dorsal view); E, left G2. Scales = 1.0 mm. with shallow cervical grooves which reach to area between postorbital cristae and junc- tion of antero- and posterolateral regions; shallow lateral depression present between cardiac and intestinal regions. Orbits trans- verse; eyes and cornea well developed. Is- chium of third maxilliped rectangular, with shallow median sulcus; margins of merus cristate with median depression; exopod slender, with long flagellum. Male chelipeds with one chela much larger than other; female chelipeds sub- equal; outer surfaces of palm smooth to weakly punctate; fingers gently curving, subequal in length to palm, cutting edges with numerous broad teeth and denticles; VOLUME 110, NUMBER 3 carpus with broad, strong, sharp tooth on inner distal angle, with smaller tooth at its base positioned at approximately right an- gles. Ambulatory legs of normal length; fourth leg shortest; second leg longest; me- rus with low (usually blunt) subterminal tooth but without subterminal spine, surface lined with weak transverse striae and some- times weakly punctate. Male anterior thoracic sternites smooth or weakly punctate; sternites 3 and 4 com- pletely fused without trace of sutures; su- ture between sternites 2 and 3 distinct, al- most straight. Male abdomen with segment 3 broadest; segments 3—6 progressively nar- rower and less trapezoidal; lateral margins of segment 6 gently convex; telson trian- gular, lateral margins almost straight, tip rounded. Female abdomen broadly oval. G1 sinuous; subterminal segment rela- tively slender, gradually tapering towards distal part, gently curving outwards; ter- minal segment, subcylindrical, approxi- mately 0.4 times length of subterminal seg- ment, median part broadest, gently curving upwards (towards anterior of carapace when lying in situ), distalmost part sharply tapering, tip appearing spine-like, subdistal surfaces with numerous long, very stiff se- tae, proximal half with small but well de- veloped, swollen dorsal flap. G2 approxi- mately 1.2 times length of Gl, distal seg- ment very long, approximately 0.7 times length of basal segment. Remarks.—Esanpotamon namsom is the smallest known potamid crab known from Thailand (see Ng & Naiyanetr 1993), reach- ing carapace widths of only about 31 mm. The largest female specimen available (23.4 by 18.3 mm, ZRC 1997.774) is not fully mature, and from the shape of its abdomen, is probably only a few moults from reach- ing adulthood. This suggests that the holo- type male has reached the adult size. In any case, the well developed G1 of the male holotype certainly indicates it is mature. Like other waterfall crabs, E. namsom 421 lives under rocks in clear, fast flowing streams (Naiyanetr 1978, 1988). Etymology.—The name is derived from the name of the district where the species occurs, Nam Som. The name is used as a noun in apposition. Acknowledgments The study was supported in part by a grant (RP 950360) to the second author from the National University of Singapore. Literature Cited Bott, R. 1966. Potamiden aus Asien (Potamon Savig- ny und Potamiscus Alcock) (Crustacea, Decap- oda).—Senckenbergiana Biologica 47:469—509. 1970. Die Siisswasserkrabben von Europa, Asien, Australien und ihre Stammesgeschichte. Eine Revision der Potamoidea und Parathelphu- soidea (Crustacea, Decapoda).—Abhandlungen Senckenbergischen Naturforschenden Gesells- chaft 526:1—338. Naiyanetr, P, 1978. The use of waterfall crabs as the evidence to indicate the relationship between the mountains of Thailand.—Abstracts of the National Conference on Agriculture and Bio- logical Science, Animal Science Section, Ka- setsart University, Bangkok, 1 pg. . 1986. Crab of Tenasserim Range Part IIl.— Abstracts of a Seminar on Wildlife of Thailand, Faculty of Forestry, Kasetsart University, Bang- kok 7:11.1-11.3. 1988. Freshwater crabs in Thailand.—In Book published in memory of the Royal Cre- mation of Associate Professor Dr. Praphun Chi- tachumnong, Chulalongkorn University, Phais- alsilpa Press, Bangkok, 15 pp. . 1992. Validation of five new species of Thai potamid crabs (Crustacea: Decapoda: Brachy- ura).—Raffles Bulletin of Zoology 40(1):1-7. Ng, P. K. L. 1988. The freshwater crabs of Peninsular Malaysia and Singapore. Department of Zool- ogy, National University of Singapore, Shinglee Press, Singapore, 156 pp. , & P. Naiyanetr. 1993. New and recently de- scribed freshwater crabs (Crustacea: Decapoda: Brachyura: Potamidae, Gecarcinucidae and Par- athelphusidae) from Thailand.—Zoologische Verhandelingen 284:1—117. Ortmann, A. 1896. Das system der Decapoden- Krebse.—Zoologische Jahrbuchner (Systemat- ics) 9:409—453. Rathbun, M. J. 1904. Les crabes d’eau douce.—Nou- velles Archives du Muséum d’ Histoire Naturel- le, Paris (4)6:225-312, pls. 9-18. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):422—425. 1997. Tridentella ornata (Richardson 1911), new combination: records of hosts and localities (Crustacea: Isopoda: Tridentellidae) Brian Kensley and Richard W. Heard (BK) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (RWH) Invertebrate Zoology Section, Gulf Coast Research Laboratory, PO. Box 7000, Ocean Springs, Mississippi 39564 Abstract.—Tridentella williamsi Delaney, 1990, is shown to be a junior syn- onym of Aega ornata Richardson, 1911. As Tridentella ornata, the species is recorded from five different host fish species, from the Gulf of Mexico and the Caribbean. The presence of pleotelsonic pits is documented and briefly discussed. The distribution and host information of the 14 known species of Tridentella are provided. Over a period of several years, isopods taken from the nasal cavities of groupers from the Gulf of Mexico were accumulated, and eventually made available to us for study. A review of literature on the family Aegidae from the Atlantic showed that our material was conspecific with a Richardson (1911) species, and also with a recently de- scribed Caribbean species of Tridentella. This note clarifies the taxonomy of the spe- cies, and records new localities and fish hosts. Family Tridentellidae Bruce, 1984 Tridentella Richardson, 1905 Tridentella ornata (Richardson, 1911) Fig. 1 Aega ornata Richardson, 1911:624, figs. 1. Tridentella williamsi Delaney, 1990:643, figs. 1-3. Type material examined.—Holotype of Aega ornata, USNM 42377, ¢ tl 8.0 mm, from Pagrus pagrus, southern United States, coll. R/V Albatross, 1885.—Holo- type of Tridentella williamsi, USNM 239198, 3 tl 8.5 mm, from Epinephelus mystacinus, British Virgin Islands.—Para- type of Tridentella williamsi, USNM 239199, ¢ tl 8.5 mm, from Epinephelus fla- volimbatus, British Virgin Islands. Non-type material examined.—USNM 253284, 1 ¢ tl 7.3 mm, 1 ovigerous 2 tl 9.2 mm, 2 non-ovigerous ¢ tl 9.1 mm, 10 mm, from nasal cavity of Mycteroperca mi- crolepis, 85-100 miles WNW of Clearwa- ter, Florida, 10 Mar 1984.—-USNM 253285, 6 tl 11.2 mm, from Mycteroperca phenax, Big Elbow, Florida, 132 m, 30 May 1978.—USNM 253286, 1 ovigerous ¢ tl 12.7 mm, from Epinephelus flavolimbatus, Gulf of Mexico, 26°00'N, 84°20'W, 157— 168 m, Feb 1984.—-USNM 253287, 1 @ tl 8.5 mm, from nasal cavity of Mycteroperca microlepis, SW of Panama City, Florida, 25 m, 10 July 1977.—USNM 253288, 1 ¢ tl 7.2 mm, 1 juvenile 5.0 mm, from nasal cav- ity of Mycteroperca phenax, 36 miles SW of Panama City, Florida, 53 m, 17 May 1971.—USNM 253289, 1 @ tl 9.3 mm, from nasal cavity of Mycteroperca phenax, north—east Gulf of Mexico, 13 Feb 1977. Diagnosis.—Eyes large, well pigmented, not contiguous. Frontal lamina elongate, five-sided, apically acute, apex meeting acute rostrum, 2 long sides slightly con- cave. Pereonites 4—7, and pleonites 1-—5 with row of small tubercles along posterior margin; pleonites 3 and 4 with additional short median row of tubercles; pleonite 5 VOLUME 110, NUMBER 3 423 rig. 1. indicate pits, scale = 100 yu; C, pleotelsonic pit opening enlarged, scale = 50 pw; D, cross-section through pleotelsonic pit, arrow indicates unsclerotised section of wall, scale = 50 i. with 2 additional rows of tubercles. Pleo- telson triangular, basal width about 1.3 times middorsal length, with irregular row of large tubercles across base; shallow me- dian longitudinal groove running from base to apex, flanked by row of large tubercles on each side, with invaginated pit on each side near base; lateral parts of pleotelson incised into 4 broad flattened sections; pos- terior margin rounded, crenulate. Remarks.—The locality for the single specimen of Aega ornata described by Richardson (1911) is given as ‘“‘southern United States’’. Comparison of this speci- men with the type material and figures of Tridentella williamsi from the British Vir- gin Islands (Delaney 1990), and our mate- rial from the Gulf of Mexico reveal no dif- ferences in the very characteristic pereonal and pleonal ornamentation. Equally, no dif- ferences in appendage structure could be detected. (In Richardson’s description, the Tridentella ornata: A, pleonite 6 and pleotelson, scale = 500 yp; B, pleotelson enlarged, arrows captions for figures 3 and 4 of the first and second maxillae are reversed). Females of the genus Aega Leach, 1815, lack the nar- row, twisted, and strongly spined maxilli- ped that characterizes the males of the ge- nus. The pair of basal pits on the pleotelson are each marked on the surface by a puck- ered slit-like opening, which leads to a small flask-shaped chamber, whose walls are formed by the invaginated exoskeleton. At the base of the chamber is a non-scler- otized region of the exoskeleton. No other structure is visible, and there was no sign of a statolithic inclusion in the specimen sectioned. The pits are sited in the same po- sition as the paired pleotelsonic statocysts of anthurid isopods (see Barnard 1925), and although this feature might tempt one to speculate on a link between the Flabellifera and Anthuridea, these pits should be seen as one possible stage in the development of 424 the type of innervated statocysts seen in the anthurids. Of the 14 species of Tridentella de- scribed, only five have been recorded as as- sociated with fish. These involve five dif- ferent fish families. In the case of 7. ornata under discussion here, the species has been taken from five different host fishes belong- ing to two separate fish families. These facts support the thesis proposed by Dela- ney & Brusca (1985:730), that these iso- pods should be regarded as micropredators, rather than parasites. Tridentellids are thought to lurk in the benthos, and pounce on almost any passing fish, to take a meal of blood and drop off again. The following is a list of the described species of Tridentella, with localities, depth and host records: Tridentella acheronae Bruce, 1988. New Zealand, Kermadec Is., 424-1006 m. Host not recorded. Tridentella cornuta Kussakin, 1979. North- west Pacific, 20-50 m. Host: Hemitrip- terus villosus (Cottidae). Tridentella glutacantha Delaney & Brusca, 1985. North Farallon Islands, Santa Cat- alina Islands, off Los Angeles, 128—360 m. No host recorded. Tridentella japonica Thielemann, 1910. Off Tokyo, Japan. No host recorded. Tridentella laevicephalax Menzies, 1962. Southern Chile, 24 m. No host recorded. Tridentella ornamenta (Menzies & George, 1972). Peru-Chile Trench, 907—935 m. No host recorded. Tridentella ornata (Richardson, 1911). South—eastern United States, Gulf of Mexico, Caribbean Sea, 25-168 m. Hosts: red porgy, Pagrus pagrus (Spari- dae); yellowedge grouper, Epinephelus flavolimbatus, misty grouper, E. mysta- cinus, gag grouper, Mycteroperca micro- lepis, scamp, M. phenax (Serranidae). Tridentella quinicornis Delaney & Brusca, 1985. Off Santa Barbara Islands and Farnsworth Bank, California, 53 m. No host recorded. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Tridentella recava Bowman, 1986. New York Bight, 100—300 m. In burrows of tilefish, Lopholatilus chamaeleonticeps (Malacanthidae). Tridentella saxicola (Hale, 1925). New South Wales, Queensland, Australia, 11— 146 m. No host recorded. Tridentella sculpturata Kussakin, 1955. Northwest Pacific, 70—96 m. Hosts: En- ophris diceraus and Alchichthys elonga- tus (Cottidae). Tridentella tangaroae Bruce, 1988. New Zealand, 90—94 m. No host recorded. Tridentella virginiana (Richardson, 1900). Nova Scotia to Florida; Gulf Stream off Florida, 220-550 m. No host recorded. Tridentella vitae Bruce, 1984. Fiji, 360 m. Host: Pristipomoides flavipinnis (Lutjan- idae). Acknowledgments The collecting efforts of David Camp, Steven Candelari, and Gene Nakamara have made this contribution possible; we are grateful to them. Ms. Cheryl Bright and Dr. Jon Norenburg of the Department of Inver- tebrate Zoology, National Museum of Nat- ural History, carried out the sectioning and slide preparation, and photography, respec- tively, of the pleotelsonic pits; we thank them sincerely for this assistance. Literature Cited Barnard, K. H. 1925. A revision of the family An- thuridae (Crustacea Isopoda), with remarks on certain morphological peculiarities.—Journal of the Linnean Society of London, Zoology 36: 109-160. Bowman, T. E. 1986. Tridentella recava, a new iso- pod from tilefish burrows in the New York Bight (Flabellifera: Tridentellidae).—Proceed- ings of the Biological Society of Washington 99:269-273. Bruce, N. L. 1984. A new family for the isopod crus- tacean genus Tridentella Richardson, 1905, with description of a new species from Fiji.— Zoological Journal of the Linnean Society 80: 447-455. . 1988. Two new species of Tridentella (Crus- tacea, Isopoda, Tridentellidae) from New Zea- VOLUME 110, NUMBER 3 land.—Records of the National Museum of New Zealand 3(7):71-79. Delaney, P.M. 1990. Tridentella williamsi, a new spe- cies of isopod crustacean from the British Vir- gin Islands, Western Atlantic (Flabellifera: Tri- dentellidae).—Proceedings of the Biological Society of Washington 103:643-648. , & R. C. Brusca. 1985. Two new species of Tridentella Richardson, 1905 (Isopoda: Flabel- lifera: Tridentellidae) from California, with a re- diagnosis and comments on the family, and a key to the genera of Tridentellidae and Coral- lanidae.—Journal of Crustacean Biology 5:728-— 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 of the U.S.S.R.—the warm water families of Isopo- da.—Travaux Trudy Zoologicheskogo Instituta, Akademiya NAUK USSR 18:228-234. . 1979. On the isopod crustaceans (Isopoda) of the Sea of Okhotsk.—Transactions, Academiya NAUK CCCP [Investigations of Pelagic and Bottom Organisms from the Far-Eastern Seas] 15:106—122. Leach, W. E. 1815. A tabular view of the external characters of four classes of animals, which 425 Linné arranged under Insecta; with the descrip- tion of the genera comprising three of these classes into order, etc., and descriptions of sev- eral new genera and species.—Transactions of the Linnean Society of London 2:306—400. Menzies, R. J. 1962. The zoogeography, ecology, and systematics of the Chilean marine isopods.— Lunds Universitets Arsskrift, N. F (2)57(11):1- 162. , & R. Y. George. 1972. Isopod Crustacea of the Peru—Chile Trench.—Anton Bruun Report 9:1-124. Richardson, H. 1900. Synopses of North American invertebrates. 7. The Isopoda.—American Nat- uralist 34:207—230. . 1905. A monograph on the isopods of North America.—Bulletin of the United States Nation- al Museum 54:1-—727. . 1911. Description of a new species of Aega from the Atlantic coast of the United States.— Proceedings of the United States National Mu- seum 40:623-624. Schioedte, J. C., & E Meinert. 1879. Symbolae ad Monographiam Cymothoarum Crustaceorum Is- opodum Familiae 1. Aegidae.—Natur-historisk Tidsskrift (3)13:281—378. Thielemann, M. 1910. Beitrage zur Kenntnis der Is- opodenfauna Ostasiens.—Mtinchens Abhan- dlungen der Akademie Wissenschaft math.- phys. KI. suppl.-bd 2, 3:1—109. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):426—438. 1997. Doxomysis acanthina, a new leptomysinid (Crustacea: Mysidacea) from the northern Great Barrier Reef, Australia, with extensions to the known distributions of D. australiensis W. M. Tattersall, 1940 and D. spinata Murano, 1990, and a key to the genus Doxomysis M. S. Talbot Department of Biological Sciences Macquarie University, N.S.W. 2109, Australia Abstract.—Doxomysis acanthina is described from the lagoon at Lizard Is- land, northern Great Barrier Reef, Queensland, Australia. The spines on the palp of its maxillary endopod are distinctive in having 2 types of secondary spinule, long and slender and small and thorn-like. D. spinata, previously found only in the Northern Territory, is provisionally identified from Lizard Island, and the known range of D. australiensis 1s extended into northern Queensland waters. The telson of all three species exhibit sexual dimorphism. A key to the genus Doxomysis is given. Nine species belonging to the genus Dox- omysis, tribe Leptomysini, have been taken in the Australian region. W. M. Tattersall (1936) recorded D. littoralis Tattersall, 1922 from the vicinity of Low Isles, Great Barrier Reef. Some of the specimens were faintly spinulose and Pillai (1973) tentative- ly referred these to D. longiura Pillai, 1963. D. australiensis was described under the name of Afromysis australiensis from Bro- ken Bay, New South Wales by Tattersall (1940). It was recorded from Morton Bay, southern Queensland by Bacescu & Udres- cu (1982), who also described D. proxima from the same area. Panampunnayil (1986) described D. johnsoni from Western Aus- tralia in the coastal waters of the extreme southwest and Murano (1990) described D. brucei and D. spinata from Port Essington, Northern Territory. Pillai (1973) recorded the occurrence of 9 specimens of D. quad- rispinosa in the North Australian Basin. They were taken individually on 4 different cruises and were found mostly to the east of Christmas Island or off the North West Cape of Western Australia. Three species of Doxomysis, D. spinata (identified provisionally), D. australiensis and the new species D. acanthina described here, were found in a survey of the mysid fauna of the Lizard Island region of the Great Barrier Reef during the years 1975— 1980. These records extend the known dis- tributions of D. spinata and D. australiensis eastwards and northwards respectively into the waters of northeastern Queensland. Murano’s (1990) description of D. spinata was made from a single adult male. The females of this species from Lizard Island reveal that it exhibits the same type of sex- ual dimorphism in the structure of the tel- son as that found in D. longiura by Pillai (1973) and as is also seen in D. australien- sis (Bacescu & Udrescu 1982, fig. 5) and in D. acanthina. The genus Doxomysis is one of a group of 9 allied genera in the tribe Leptomysini. Although full descriptions are not available for many of the 48 species that belong to these genera, an affinity among them is sug- gested by resemblances in the following characters: the enlargement and elaboration of the palp on the endopod of the maxilla; the similarity in the structure of the modi- fied terminal and subterminal setae of the exopod of the fourth male pleopod; the VOLUME 110, NUMBER 3 well-defined cleft in the telson and the over- all resemblance in the structure and pattern of distribution of telson spines. The genera are listed below: Afromysis Zimmer, 1916 (6 species) Australomysis W. M. Tattersall, 1927 (5 species) Bathymysis W. M. Tattersall, 1907 (2 spe- cies) Doxomysis Hansen, 1912 (15 species) Hyperiimysis Nouvel, 1966 (1 species) Timysis Nouvel, 1966 (2 species) Nouvelia Bacescu & Vasilescu, 1973 (3 species) Pseudoxomysis Nouvel, 1973 (1 species) Tenagomysis Thomson, 1900 (13 species) Fenton (1991) pointed out the need to reassess the status of this group of genera and discussed problems arising from the lack of information on diagnostic features in many of the species. Members of the ge- nus Afromysis are characterised by the markedly crescentic form of the maxillary palp, a feature that distinguishes them from members of the other genera in the group. In the genera Doxomysis, Hyperiimysis and Pseudoxomysis (the doxomysid sub-group), the palp is broader than long and typically fan-like in appearance, while in the genera Tenagomysis, Iimysis, Australomysis, Bath- ymysis and Nouvelia (the tenagomysid sub- group), it is longer than broad and usually obovate in shape. Nouvelia also contains a single species in which the palp is broader than long and resembles that of the doxo- mysid sub-group. The doxomysid sub-group may be fur- ther divided on the basis of the structure of the marsupium of the female, which is formed of only two pairs of oostegites in the genus Doxomysis, while in both Hyper- limysis and Pseudoxomysis it is made up of three pairs and in both, there is also a small projection at the base of the fifth pair of thoracic limbs, which, in the case of Pseu- doxomysis, has been interpreted as a rudi- mentary forth pair of oostegites. Although smaller, the basal projection in Hyperiimy- 427 sis could be given the same interpretation. This feature is probably indicative of a close affinity between the two genera, as are others they have in common, such as a spine on the anterior border of the labrum and secondary spinules on the distal spines of the maxillary palp. Such similarities sug- gest that the species concerned should be placed in the same genus. However, differ- ences in the structure of their thoracic limbs may mitigate against doing so, as the car- popropodus is made up of 3 articles in Pseudoxomysis and two articles in Hyperi- imysis. Additionally, in the the latter genus the carpo-propodus of the eighth thoracic limb is enlarged and modified to form a prehensile structure. Among the members of the tenagomysid sub-group, Australomysis and Bathymysis both lack the pair of plumose setae present in the telson cleft of all the other genera. They both contain species with dorso-ven- trally flattened eyes and they are not clearly distinguished from each other anatomically, apart from the fact that the distal spines of the maxillary palp are barbed in Bathymysis and simple in Australomysis. They also have widely separate distributions and con- trasting habitats. Australomysis has been found in shallow inshore waters along the south-west, south and east coasts of Austra- lia and in the surf zone on the Pacific coast of central Japan (Fukuoka & Murano 1994), while both species of Bathymysis were taken at depth in the Atlantic (W. M. Tattersall 1907, 1951). The genus /imysis was erected to accom- modate those Tenagomysis species that had an anterior spine on the labrum and a tarsus made up of four articles (Nouvel 1966). As indicated by Fenton, 1991, 7. tanzaniana (Bacescu 1975) should probably be trans- ferred to this genus. Its tarsus is made up of only three articles, but it has a spine on the anterior border of the labrum. The genus Nouvelia is distinguished by a tarsus consisting of only three articles and a gap in the row of lateral spines on the telson. It includes N. nigeriensis (O. Tatter- 428 sall, 1957), a species in which the palp on the maxilla is broader than long and which might, therefore, be better placed in the ge- nus Doxomysis. At least 10 species are known to have secondary spinules on the spines that bor- der the distal margin of the maxillary palp. In the descriptions of 3 of these, the spi- nules are shown in figures of the maxillae, but not mentioned in the text. They are present in the members of the genus Bath- ymysis and in 3 species of Tenagomysis, T. australis, T. tasmaniae and T. bruniensis (Fenton 1991), in Hyperiimysis madagas- cariensis (Nouvel 1966, fig. 10), in Pseu- doxomysis caudaensis (Nouvel 1973, fig. 10) and in 3 species of Doxomysis, D. spi- nata, where they are long and slender (Mur- ano 1990, fig), D. rinkaiensis, which has | small spinules on the expanded tips of some of the spines (Valbonesi & Murano 1980, fig 3D) and D. acanthina, described below, in which there are 2 distinct sets of spi- nules, one, long and slender, towards the bases of the spines and the second, short and close- set nearer the tips. Among the species under discussion, D. acanthina is the only one so far found with more than 1 type of secondary spinule. As D. murariui was described from 1 damaged female and D. sanuriensis from two damaged specimens, the status of these two species is difficult to ascertain. The il- lustrations of both show secondary spinules on the spines of the maxillary palps (Ba- cescu 1993, figs. 1E, maxillule (sic) and 2B, maxillule (sic)). This does not correspond with the account of the spines in the de- scription of D. sanuriensis, however, while in the description of D. murariui, the spines are not discussed. The elongate shape of the maxillary palp of the latter species suggests that it may have a greater affinity with the tenagomysid group of species than with the doxomysid group. Doxomysis acanthina, new species Figs. 1-3 Material examined.—179 specimens were taken from the Lizard Island lagoon, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON during 1975-1980. Of these 113 were caught in nets and traps deployed close to the sandy floor, 60 in plankton hauls made in the lagoon center at night and 3 in traps placed above living coral. 3 additional spec- imens were taken in a night haul in the open channel between Lizard Island and Eagle Cay. Size range.—Length, measured from an- terior border of eyes to end of telson; 55 adult males, 3.7-6.5 mm.; 27 immature males, 2.5—3.8 mm.; 4 brooding females (up to 8 larvae in marsupium), 5.0—5.5 mm.; 12 adult females, marsupium empty, 4.5—5.5 mm.; 32 immature females, 2.8—5.0 mm.; 49 juveniles, 1.5—3.5 mm. Type series.—Type specimens deposited in the Australian Museum, Sydney, para- types deposited in the National Museum of Natural History, Smithsonian Institution, Washington D.C. Types and Paratypes all collected in the Lizard Island lagoon, 14°40'S, 145°27'E. Holotype: adult male and slide prepara- tion of right maxilla, AM P42693, light trap, sandy lagoon floor, depth 11 m, 28 May 1975, 1922-1927 hrs (Sta. #STL-75- L10). Allotype: adult female, AM P42694, plankton net pushed above lagoon floor, depth 9 m, 31 May 1975, 1600 hrs (Sta #STL-75-P8). Paratypes: 2 adult males, AM P43138, 3 adult males, USNM 259765, light trap light, sandy lagoon floor, depth 11 m, 28 May 1975, 1922-1927 hrs (Sta. #STL-75-L10), 1 adult female, USNM 259766, light trap on living Porites coral, E of Palfrey Island, depth 1 m, 4 Jan 1975, 2228—2233 hrs (Sta. #STL-75-T11). Description.—Body slender. Integument spiny, small scale-like spinules conspicu- ously dense on carapace, less dense on ab- domen, eyestalks, sparse on bases of anten- nae, pleopods. Carapace small, rounded, ex- posing last 2 thoracic segments. Rostrum short, bluntly pointed, reaching base of eye- stalks (Figs. 1A, B). Eyes prominent, fairly dark, hemispheri- VOLUME 110, NUMBER 3 F Big. 1. Labrum. E, Right mandible and palp. E Left mandible. cal, greater in diameter than width of eye- Stalks. Antennular peduncle of male stout, first article less than 0.5 total length, third ar- ticle expanded, appendix masculina coni- cal with prominent sensory bristles (Fig. 1A). Antennular peduncle of female slen- der, first segment 0.5 total length (Fig. 1B). 429 0.5mm AB 0.1mm C O.Lmm DEF Doxomysis acanthina. A, Anterior end of male. B, Anterior end of female. C, Left antenna. D, Antennal scale narrow, overreaching an- tennular peduncle by about 0.25 of its length, distal segment present, suture incon- spicuous (Fig. 1C). Labrum rounded, without anterior spine (Fig. 1D). Mandible with well-developed incisor process, lacinia mobilis, spine row and spi- nulose molar process, palp typical of genus, 430 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sag e 7 ee Fig. 2. Doxomysis acanthina. A, Maxillule. B, Segment 2 of endopod of maxilla. C, Thoracic limb 1. D, Thoracic limb 2. E, Endopod of thoracic limb 4. apart from relatively slender second article ment 3 armed with 10 short straight spines (Figs. 1E, F). (Fig. 2A). Maxillule with inner lobe of segment 1 Palp of maxillary endopod expanded to ending in 2 prominent notched spines, seg- form characteristic plate, bordered by 7—8 VOLUME 110, NUMBER 3 stout spines, each carrying a few long slen- der secondary spinules and 3-6 small, thorn-like, closely spaced spinules, in 2 lat- eral rows on distal part of each spine (Fig. 2B). Thoracic limb 1 with robust endopod, row of 10 strong curved setae on mastica- tory endite of merus, epipod with a long seta near base (Fig. 2C). Dactylus of tho- racic limb 2 armed with 7 prominent serrate spines (Fig. 2D). Thoracic endopods 3-8 with oblique articulation between carpus and propodus, propodus made up of 2 sub- equal subsegments, dactylus slender, ending in long sharp nail (Fig. 2E). Marsupium of female formed of 2 pairs of oostegites borne on thoracic limbs 7 and 8. Segment 6 of abdomen elongate, almost twice as long as segment 5. Pleopods of male biramous, sympod slightly spinulose, exopods with 7 seg- ments. Endopod of pleopod 1 short, un- segmented with single distal seta, 5 lateral setae, pseudobranchial lobe ending in 4 setae with bulbous bases (Fig. 3A). Pleo- pods 2-5 with 6-segmented endopods. Exopod of pleopod 4 modified: basal seg- ment with shallow keel on inner edge; 4th segment with small, naked seta on inner distal margin; 5th segment with large, ro- bust, curved spine, extending from outer distal margin to reach almost 0.6 length of terminal spines, distal 0.3 with outer row of close-set, secondary spinules; 6th segment with slender, naked distal seta on outer border, adjacent to strong, slightly curved spine, extending beyond terminal setae, row of sharp, slender secondary spinules on distal 0.6 of outer margin; 7th segment small, ending in 2 long subequal setae (Figs. 3B, C). Pleopod 5 with prom- inent conical distolateral process on basal segment of endopod (Fig. 3D). Uropod slender, all margins setose, exo- pod almost twice as long as telson, endopod slightly more than 0.75 length of exopod, spine row of endopod with 22-23 spines, closely-set, alternately small and large in proximal part near statocyst, grading distal- 431 ly to larger, more widely-spaced spines near apex (Fig. 3E). Telson about equal in length to pleonite 6, cleft slightly more than 0.25 total telson length. Telson of male strongly tapering, width at apex 0.5 width at base, apical lobes narrow, margins slightly concave, 11-12 lateral spines, terminating in 4 prominent apical spines, the second being the longest, cleft with 8—9 small spines on each side, 2 plumose setae projecting from base (Fig. 3F). Telson of female with less pronounced taper than that of male, width at apex 0.6 that of base, apical lobes broad, margins markedly concave, 12 lat- eral spines, grading into 5 large, some- what spatulate apical spines, central apical spines subequal, 9-10 cleft spines on each side (Fig. 3G). Chromatophore pattern.—Observed in 2 females hand-netted above the pale sandy lagoon floor. Eyestalk; 1 white chromatophore and 1 red chromatophore. Cephalothorax; 1 large, highly branched white chromatophore and 1 red chromat- ophore mid-dorsally above proventricu- lus, 1 white mid-lateral chromatophore on each side, 1 red mid-ventral chromato- phore. Marsupium; 1 white dorso-lateral chromatophore on each oostegite, 1 red ventro-lateral chromatophore on anterior part of each posterior oostegite. Abdo- men; 1 white mid-dorsal chromatophore on each segment, that of last segment large and highly branched, situated above sixth abdominal ganglia, 1 white and | red mid-ventral chromatophore on each segment. Telson; | red dorsal chromato- phore. Note on habitat.—The major portion of the D. acanthina catch was made up of 113 specimens found living just above the sandy floor of the lagoon, while 60 were caught in plankton tows taken in the la- goon at night, an indication that this spe- cies migrates into the water column dur- ing the hours of darkness. Only 3 speci- mens were caught above living coral and 3 in offshore plankton tows made at night 432 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ISK a h ZK) {| 4 eh A N x LO” Pa ees | Pimm ABD y \ 0.1 DIME EG Fig. 3. Doxomysis acanthina. A, Pleopod 1. B, Pleopod 4. C, Terminal 3 segments of exopod of pleopod 4. D, Pleopod 5. E, Left uropod of male. K Right uropod and telson of male. G, Telson of female. in the channel between Lizard Island and Doxomysis spinata Murano, 1990 Eagle Cay. Fig. 4. Etymology.—From the Greek ‘“‘akanthi- nos’’, thorny, referring to the lateral rows Material examined.—326- specimens, of small, thorn-like secondary spinules on provisionally assigned to this species, were the spines of the maxillary palp. caught at Lizard Island, 308 in a light trap VOLUME 110, NUMBER 3 433 Fig. 4. Doxomysis spinata. A, Segment 2 of endopod of maxilla. B, Anterior end of female. C, Telson of female. D, Telson of male. on the reef flat, 17 in a light trap on living Acropora formosa and a single female in a hand-net just above the sandy lagoon floor. Size range.—Length, measured from an- terior border of eyes to end of telson, 93 adult males, 3.5—5.7 mm; 36 immature males, 2.8-4.0 mm; 38 brooding females (up to 9 larvae in marsupium), 4.5—6.0 mm; 17 adult females, marsupium empty, 4.5— 5.5 mm; 141 immature females, 3.0—5.2 mm; | juvenile, 1.7 mm. Description of Lizard Island speci- mens.—These correspond in most features with Murano’s (1990) description of D. spi- nata, which was made from a single male, 5.8 mm in length, taken in Port Essington, Northern Territory. Their integument is smooth and they have the long slender sec- 434 ondary spinules on the spines of the max- illary palp, noted in D. spinata (Fig. 4A). The secondary sexual characters of the males resemble those of the type specimen. They have a similarly robust antennular pe- duncle with a prominent appendix mascu- lina and the exopod of pleopod 4 is modi- fied in the same way as in the type speci- men, although mature individuals have a slightly longer inner seta on segment 7 than is shown in Murano’s Fig. 4C. It is possible, however, that in spite of their similarity to D. spinata, the specimens from Lizard Island will prove to belong to a separate species, because they differ from the type specimen in the number of spines on the endite on the merus of the endopod of thoracic limb 1, in the number in the spine row on the endopod of the uropod and — in the number of spines on the telson. The counts were made on 10 of the Lizard Is- land males, varying in length from 4.2—6.8 mm, with the smaller specimens tending to have a smaller number of spines than the larger. The spine counts were as follows: LI. specimens Holotype Endite on thoracic limb 1— 8—10 7 Endopod of uropod— 21-33 37. Each side of telson, lateral spines— 14—20 12 apical spines— 3 3 cleft spines— 10-16 9 These differences suggest that the Lizard Is- land specimens may be distinct from D. spi- nata, but since the range of variation in specimens from the type locality cannot as yet be assessed, they have been relegated to the same species provisionally, pending the collection of further material in the North- ern Territory. An examination of the type specimen of D. spinata has confirmed that it corresponds with the Lizard Island spec- imens, except in the features detailed above. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON In the females from Lizard Island the an- tennule has a slender basal segment equal in length to the combined lengths of the second and third segments (Fig. 4B). The marsupium is formed of 2 pairs of ooste- gites. The telson is moderately tapered and has 16 lateral spines, 5 subequal apical spines and 12 cleft spines on each side (Fig. 4C). It differs from the male telson (Fig. 4D) in being slightly less tapered, in lack- ing a large posterior apical spine and in having a shorter cleft with fewer cleft spines. Note on habitat.—325 of the Lizard Is- land specimens of D. spinata were trapped in the vicinity of coral, while only 1 was netted above the sandy lagoon floor. Doxomysis australiensis (W. Tattersall, 1940) Material examined.—During May 1976, 4 specimens of D. australiensis were taken at Lizard Island. They included a single fe- male, caught in a net pushed above the la- goon floor and an immature male and 2 ju- veniles caught in 2 plankton tows made at night in the channel between Lizard Island and Eagle Cay. Its smooth integument dis- tinguishes it from D. acanthina and its lack of secondary spinules on the maxillary palp separates it from D. spinata (Table 1). Size range.—Length, measured from the anterior border of the eyes to the end of the telson, 1 immature male, 4.0 mm; 1 brood- ing female (12 larvae in marsupium), 6.5 mm; 2 juveniles, 1.5 and 2.4 mm. Note on distribution.—This species was described from Broken Bay, New South Wales, and has been recorded from More- ton Bay, southern Queensland (Bacescu & Udrescu 1982). The Lizard Island material extends its known range into the waters of northern Queensland. The 4 specimens of D. australiensis in the Lizard Island samples were taken on 2 days in mid-May 1976, one in association with D. acanthina near the lagoon floor and the other three at night in offshore plankton VOLUME 110, NUMBER 3 Table 1.—Comparison between D. acanthina, D. spinata and D. australiensis. 435 D. acanthina D. spinata D. australiensis Integument Carapace spiny smooth smooth Abdomen spiny smooth smooth Max. palp Spine no. 1-8 8-10 10 Secondary spinules 2 types 1 type absent lst type long, slender long, slender 2nd type close-set absent Male pleop. 4 Exop. seg. 7 Distal setae both long, stout 1 long, 1 small both curved, moderate Exop. seg. 6 Length greater than 2 X 2 MSese7 6 X seg. 7 sex. 7 Exop. seg. 5 Distal seta length Exop. seg. 1 Keel Uropod Endop. spines Telson Lat. spines Apic. spines Cleft spines Sexual dimorphism Male telson: Apic. lobes Apic. spines Fem. telson: Apic. lobes Apic. spines greater than distal seta present on inner mar- tapered pointed rounded spatulate less than distal seta absent greater than 2 X distal seta keel not recorded type, 37 L. Is., 21-33 42-45 type, 12 L. Is., 14—20 19-21 type, 3°L/Is., 5 4 type, 9 L. Is., 10-16 by tapered tapered pointed pointed less rounded tapered more pointed pointed tows. Doxomysis australiensis occurs in the coastal waters of central New South Wales and southern Queensland, where it was the second most abundant mysid in samples from Moreton Bay examined by Bacescu & Udrescu (1982). It may be reaching the northern limit of its range in the region of Lizard Island. The small number of speci- mens taken and the fact that strong south- easterly trade winds were blowing at the time, raise the possibility that it is carried into the area intermittently, when trade winds dominate the weather pattern and en- hance the northward flow of the coastal cur- rent. The species of Doxomysis.—The genus Doxomysis is made up of the following 15 species: D. acanthina n. sp.—Lizard Island, Queensland, Australia. D. anomala W. M. Tattersall, 1922—An- daman Islands, India. D. australiensis (W. M. Tattersall, 1940)— E. coast, Australia; Mossel Bay, South Africa. D. brucei Murano, 1990—Port Essington, N.T., Australia. D. hanseni Colosi, 1920—Malay Archipel- ago. 436 D. johnsoni Panampunnayil, 1986—W. coast, Australia. D. littoralis W. M. Tattersall, 1922—An- daman Islands, India; Great Barrier Reef, Australia. D. longiura Pillai, 1963—Kerala, India. D. microps Colosi, 1920—Galapagos Is., Ecuador. D. murariui Bacescu, 1993—Bali, Indone- sia. D. proxima Bacescu & Udrescu, 1982— Moreton Bay, Australia. D. quadrispinosa (illig, 1906)—Tropical Indo-Pacific. D. rinkaiensis Valbonesi 1980—Tanabe Bay, Japan. D. sanuriensis Bacescu, 1993—Bali, Indo- nesia. D. spinata Murano 1990—Port Essington, N.T., Australia. c& Murano, Doxomysis zimmeri Colosi, 1920 has been omitted from the above list, as it was considered to be a synonym of D. quadris- pinosa (illig 1906) when reviewed by W. Tattersall (1922, 1951) and Pillai (1973). They argued that its description lacked characters distinguishing it from the latter species. Sri Lanka, the type locality of D. zimmeri, is also within the known distri- bution range of D. quadrispinosa. Key to the Species of Doxomysis la. Integument of carapace and abdomen SUIROOUNG tens eet se neat ae ae eaten ate chee 2 b. Integument of carapace and abdomen SHODDY! 5 5 Se ee one ee 12 2a. Median dorsal tubercle on posterior surface of carapace D. johnsoni b. Dorsal surface of carapace lacking projections 3a. Telson with 1 stout and 2 small ter- minal spines on each apical lobe, cleft wide and shallow D. anomala b. Telson with 3—8 stout terminal spines on each apical lobe, cleft narrow and (NOS) Oye ata i cate ba a ee hohe a a wow ee 4 4a. Row of spines on lateral margin of tel- son with large gap, at least half as long AS Me lSOM 2B) 8 es eal ME Be Sen ae ee 5 b. Sa. 6a. 7a. 8a. 9a. 10a. lla. 12a. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Spine row on lateral margin of telson without gap, or with small gap, 1 quar- ter telson length, or less Telson with 6 spines on each lateral margin, a large gap between the 3 proximal spines and the 3 distal spines D. murariui Telson with set of 8—9 small, evenly spaced spines on distal extremity of each lateral margin D. hanseni Telson with 9 spines on each lateral margin, a gap between the 4 proximal and the 5 distal spines, endopod of uropod with row of 29 spines D. sanuriensis At least 12 spines on each lateral mar- gin of telson, 32—50 spines on endo- ee © © © © © © © © © © © © © © © © ew ew ee eee © © we we eo © © © © © © © © © © © © © ee ee ee ll pod_of uropod ... .. 05.55 7 Telson with rounded apical lobes, about 8 broad, spatulate terminal spines on each lobe ......... D. brucei Telson with tapered apical lobes, 3—4 long, pointed terminal spines on each lobe! :. 260). wed oo Ae 8 Spines on distal margin of maxillary palp with secondary spinules....... 2 Spines on distal margin of maxillary palp without secondary spinules ... 10 Spines on maxillary palp expanded to- wards tip, small secondary spinules located distally on expanded part, 48— 50 spines on endopod of uropod .... D. rinkaiensis Spines on maxillary palp taper to a point, long slender secondary spinules more proximally located, fewer than 40 spines on endopod of uropod .... Lh. TEAS OL A a Ce D. spinata Lateral and terminal spines total about 17 on each side of telson, maxillary palp with 9 blunt spines . D. littoralis . Lateral and terminal spines total 25— 29 on each side of telson, maxillary palp with 10 or 12 sharp spines ... 11 Maxillary palp with 10 spines, about 47 spines on endopod of uropod .... D. australiensis eo 2 © © © © © © © © © © © we we ee el . Maxillary palp with 12 spines, about 32 spines on endopod of uropod .... ee ae ee ee neem ee rer eee D. proxima Gap in lateral spine row of telson, or spines confined to distal part of mar- HE? LOC ee EE a. ee lise VOLUME 110, NUMBER 3 b. Lateral spine row of telson without gap Eyes large, rounded, diameter approx- imately twice that of eyestalk Fl SOIR Se iP em D. quadrispinosa b. Eyes small, flattened, diameter similar to, tliat of Eyestalke oc. D. microps Antennal scale reaches end of anten- nular peduncle, maxillary palp with 9 simple spines D. longiura b. Antennal scale extends beyond anten- nular peduncle by 1 third of its length, maxillary palp with 7—8 spines bear- ing both long slender and short thorn- like secondary spinules ... D. acanthina 13a. 14a. Acknowledgments I wish to thank the Division of Crustacea in the Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution and Professor H. Choat and Associate Professor C. Alexan- der of the Department of Marine Biology, James Cook University, for accommodation and support. I particularly wish to acknowl- edge my great debt to the late Dr. T. E. Bowman for all the guidance and encour- agement he gave me. I am also grateful to Mrs. M. K. Ryan, for her patient instruction and generous help with the illustrations and to Dr. B. Kensley, Dr. J. W. Reid, Dr. FE A. Chace, Jr., Dr. & Mrs. J. Greenwood and Dr N. Tait for assistance and advice and to Ms. K. Coombes of the Northern Territory Museum for arranging for access to the type specimen of D. spinata. Dr. E H. Talbot’s support was invaluable. Literature Cited Bacescu, M. 1975. Contributions to the knowledge of the mysids (Crustacea) from the Tanzanian wa- ters.—Science Journal, University of Dar es Sa- laam 1(2):39-61. 1993. New contributions to the Mysidacea (Crustacea, Peracarida) of Indonesia: Doxomy- sis genus.—Travaux du Muséum d’ Histoire na- turelle Grigore Antipa 33:285—290. , & A. Udrescu. 1982. New contribution to the knowledge of the Mysidacea from Australia.— 437 Travaux du Muséum d’Histoire naturelle Gri- gore Antipa 24:79—96. , & Vasilescu. 1973. New benthic mysids from the littoral waters of Kenya: Mysidopsis keny- ana n.sp. and Nouvelia natalensis mombasae n.g., n.sp.—Revue Roumaine de Biologie, Serie de Zoology 18:249-256. Colosi, G. 1920. Raccolte planctoniche fatte dalla R. Nave “Liguria” del 1903-1905, Crostacei— 4—-Misidacei 2(9):229-—260. Fenton, G. E. 1991. Three new species of Tenago- mysis from the coastal waters of south-eastern Tasmania (Crustacea: Mysidacea: Mysinae: Leptomysini)—Memoirs of the Museum of Victoria 52:325-335. Fukuoka, K., & M. Murano. 1994. A new species of the Genus Australomysis (Crustacea, Mysida- cea) from Japan.—Proceedings of the Japanese Society of Systematic Zoology 51:18—24. Hansen, H. S. 1912. Report on the scientific results of the Expedition to the tropical Pacific— Steamer “‘Albatross’’ The Schizopoda.—Mem- oirs of the Museum of Comparative Zoology Harvard 35(4):173—296. Illig, G. 1906. Bericht tiber die neuen Schizopoden- gattungen und Arten der Deutchen Tiefsee-Ex- pedition 1898-1899, 1, Mysiden.—Zoologisch- er Anzeiger 30:194-—211. Murano, M. 1990. Three new leptomysids (Mysida- cea) from northern Australia.—Crustaceana 59(3):23 1-244. Nouvel, H. 1966. Mysidaces recoltes par S. Frontier a Nosy-be 3. Hyperiimysis madagascariensis n. gen., n.sp., Leptomysini.—Bulletin de la Société d’Histoire Naturelle de Toulouse 102(2—3):493— 505: . 1973. Pseudoxomysis caudaensis n. gen., Nn. sp., crustace, mysidace (Leptomysini) de la Mer de Chine meridionale.—Bulletin de la Société d’Histoire Naturelle de Toulouse 109:131—141. Panampunnayil, S. U. 1986. New mysids from the South Australian coastal waters: Paranchialina secunda sp. nov.; Leptomysis longisquama sp.nov. and Doxomysis johnsoni sp.nov.—Jour- nal of Plankton Research 8(6):1183-—1195. Pillai, N. K. 1963. On a new mysid from the inshore waters of the Kerala coast.—Journal of the Ma- rine Biological Association of India 5:258—262. . 1973. Mysidacea of the Indian Ocean. Papers on the Zooplankton Collections of the TOE.— Handbook to the International Zooplankton Collections 4:1—125. Tattersall, O. S. 1957. Report on a small collection of mysidacea from the Sierra Leone estuary together with a survey of the genus Rhopalopthalmus IUllig and a description of a new species of Tenagomysis from Lagos, Nigeria.—Proceedings of the Zoolog- ical Society of London 129:181—128. 438 Tattersall, W. M. 1907. Preliminary diagnosis of six new Mysidae from the west coast of Ireland.— Annals and Magazine of Natural History 7(19): 106-118. . 1922. Indian Mysidacea.—Records of the In- dian Museum 24:445—504. . 1927. Australian opossum shrimps (Mysida- cea).—Records of the South Australian Muse- um 3(3)235-—257. 1936. Mysidacea and Euphausiacea.—Sci- entific Reports of the Great Barrier Reef Ex- pedition 5:143-176. . 1940. Report on a small collection of Mysi- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dacea from the coastal waters of New South Wales.—Records of the Australian Museum 20: 327-340. . 1951. A review of the Mysidacea of the Unit- ed States National Musem.—Bulletin of the United States National Museum 201:1—292. Valbonesi, A., & M. Murano. 1980. Mysidae of shal- low water in Tanabe Bay.—Publications of the Seto Marine Biological Laboratory 25(1/4): 211-226. Zimmer, C. 1916. Beitrage zur Kenntnis der Meeres- fauna Westafricas. Crustacea, 4: Cumacea und Schizopoden.—Hambourgh 2(1):55—66. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):439—446. 1997. Acanthomysis bowmani, a new species, and A. aspera Ii, Mysidacea newly reported from the Sacramento-San Joaquin Estuary, California (Crustacea: Mysidae) Richard F Modlin and James J. Orsi (RFM) Department of Biological Sciences, The University of Alabama in Huntsville, Huntsville, Alabama 35899, U.S.A.; (JJO) Department of Fish and Game, Bay-Delta and Special Waters Projects Division, 4001 North Wilson Way, Stockton, California 95205-2486, U.S.A. Abstract.—Acanthomysis bowmani, a new species is described and named. Geographical range is extended for A. aspera li (1964) to include the Sacra- mento-San Joaquin Estuary. Evidence indicates that both species were recently introduced into this estuarine system. W. M. Tattersall (1932) reported five spe- cies of Mysidacea in San Francisco and San Pablo Bays. Distribution of one of these species, Neomysis mercedis Holmes, 1897 extends upstream into the Sacramento-San Joaquin Delta. Neomysis mercedis, an epi- benthic species, shares this Delta with a re- cently described species, Deltamysis ho- Imquistae Bowman & Orsi (1992). This pa- per discusses two species of Acanthomysis that have begun to appear in plankton sam- ples collected since 1992, A. bowmani, a new species, first caught 8 July 1993, and A. aspera Ii (1964), first caught 5 August #992. Methods.—All samples were collected with a tow-net mounted in a frame con- structed of steel pipe and towed for 10 min- utes from the bottom to the surface in sev- eral steps depending on water depth. The mouth of the net was 0.3 m in diameter, bag length equaled 1.5 m, and mesh size was 505 wm. Acanthomysis bowmani, new species Figs. 2—4 Material examined.—All specimens from Suisun Bay, California (Fig. 1), J. J. Orsi, 11 Apr 1996. Holotype male, 11.6 mm total length (USNM 282745); Allotype, female, 12.5 mm (USNM 282746); Para- types (USNM 282747) 7 males and 1 dis- sected male (7.7-11.8 mm), 4 females (10.5—11.8 mm), and 12 juveniles (3.5—8.1 mm). Description.—Body slender, elongate. Carapace (Fig. 2A, B) with anterior margin produced into sharp triangular rostrum ven- trally supported with distinct keel and 2 struts at base, posterior margin emarginate exposing somites 7 and 8, anterio- and pos- teriolateral lobes rounded. Narrow, nearly inconspicuous furrow encircles carapace about % the distance posterior from ros- trum. Eyes prominent, stalked; short pig- mented stripe near base of stalk dorsome- dially; cornea large, kidney-shaped, dorsal margin scalloped, conspicuous ocular tooth on anterosuperior edge (Fig. 2A, B). Antennular peduncle (Fig. 2C) stout, 3-segmented; combined length of segments 1 and 2 equal length of segment 3; segment 1 with group of 4 pinnate setae distolater- ally; segments 2 and 3 without conspicuous setae; ventrolateral male lobe on segment 3 conspicuous, heavily setose with fine sim- ple setae. Antennal peduncle (Fig. 3A) 3-segment- ed; segments 1 and 3 subequal in length; segment 2 about 1.5 times longer than seg- ment 1 or 3, with 3 stout naked and | pin- nate setae distolaterally; segment 3 with 4 440 Y bed Carquinez Straits Bay San Francisco Bay s ew Map San Francisc A\ = 24 2 St San Pablo on Sea Se cin PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Location |: Pig. 1: Game. Stations 20 and 24 in Suisun Bay are where specimens of Acanthomysis aspera were first collected and stations 64 and 78 are where A. bowmani was first obtained. stout naked and 2 pinnate setae distolater- ally, 2 small pinnate setae ventrally. Anten- nal scale blade-like, setose all around, about 1.8 times longer than peduncle, lateral mar- gin nearly straight, medial margin slighty convex near base, 8.4 times as long as wide near base, rounded articulated tip 0.05 times scale length. Right and left mandibles (Fig. 3B) with bicuspid incisors, left lacinia mobilis with 3 robust cusps, right monocuspid; each mandible with 3 robust setose accessory blades; left molar with medial surface slightly papillose, base heavily papillose and armed with clump of short, stout setae; right molar basal surface armed with row of strong, tooth-like projections and clump of short, stout setae. Mandibular palp (Fig. 3C) 3-segmented, segment 1, inconspicu- ous; second proximal segment robust, tri- angular in cross section, 1.5 times longer than distal segment, lateral margin with Sacramento-San Joaquin Esturary with sampling stations of the California Department of Fish and stout, naked setae, medial margin with 4 widely separated, stout setae; distal segment lateral margin armed with spines furnished with lateral rows of spinules, medial margin with 6 widely spaced naked setae, termi- nating in single robust, naked spine. Maxillule (Fig. 3D) typical of genus, out- er lobe with hump-like process on anterior margin, posterior margin naked. Maxilla (Fig. 3E) exopod blade-like, 4 times as long as broad at greatest width; endopod 2-seg- mented, proximal segment with rectangular patch of minute papillae near lateral mar- gin, distal segment margins setose. Labrum (Fig. 3F) and paragnaths (Fig. 3G) typical of genus. Endopod of first (Fig. 3H) and second (Fig. 31) thoracic limbs typical of genus. Endopod of third thoracic limb (Fig. 4A) with ischium 1.8 times longer than merus, ischium as long as carpo-propodus, carpo- propodus 9-segmented, dactylus small VOLUME 110, NUMBER 3 44] Fig. 2. Acanthomysis bowmani, new species: A. Anterior profile; B. Dorsal view; C. Antennule peduncle. Male, 11.6 mm. 442 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ My lye ip ——_—— a AC-1 07mm NN B 0.05 7, \\ Pm RWS rh ON il \ Fig. 3. Acanthomysis bowmani, new species: A. Antenna and antennal scale; B. Right and left mandibles; C. Mandibular palp; D. Maxillule; E. Maxilla; F Labrum; G. Paragnaths; H. Endopod of 1st thoracic limb; I. Endopod of 2nd thoracic limb. Male, 11.6 mm. VOLUME 110, NUMBER 3 about 0.2 times length of strong terminal claw; exopod (Fig. 4B) about 2/3 length of endopod, 9-segmented. Structure of endo- pods and exopods of thoracic limbs 4 to 8 similar to 3. Penis (Fig. 4C) robust, length about 1.7 © times width, each with pair of terminal lobes containing an aggregate of naked se- tae, posterior margin with 7 naked setae around edge of an articulation and small plumose setae near base, anterior margin with 1 naked and 3 pinnate setae. Male pleopods 1 to 3 and 5 rudimentary, unjointed; pleopod 1 smallest (Fig. 4D) and 5 longest (Fig. 4F); pleopod 4 biramous (Fig. 4E), endopod unjointed, exopod long, 2-segmented, segment 1 about 1.5 times longer than endopod and 1.2 times longer than segment 2 which terminates in 2 long, robust, spinulose nails, a long, robust seta and | minute spine. Uropod (Fig. 4G) exopod blade-like, lat- eral margin slightly concave, medial margin slightly convex, about 1.4 times longer than endopod; endopod margins tapering distal- ly, 2 distally directed spines along medial margin near statocyst ventrally. Telson (Fig. 4H) linguiform, 2.4 times longer than width at base, each lateral mar- gin with 3—4 spines near base, a space and 18-19 spines of equal length along distal 2/3, ultimate marginal spines 3.3 times lon- ger than other marginal spines and as long as pair of terminal spines. Remarks.—Close resemblance exists be- tween A. bowmani, n. sp., and two species reported from the western Pacific Ocean by li (1964), A. sinensis and A. longirostris (Table 1). In addition to two subtle char- acter variations, e.g., arrangement of the long spines on or near the apex of the telson and the difference in the number of distal telsonal marginal spines, the near equal length of the two segments that comprise the exopod of male fourth pleopod and a longer antennal scale separates A. bowmani from the other two species (Table 1). Pres- ently A. sinensis is known only from the East China Sea off the mouth of the Yang- 443 tze River, while A. longirostris has a more cosmopolitan distribution in oriental waters. It has been reported from Ariake Bay, Ja- pan, Port Kusan and Haejun Bay, Korea (Ii 1964, Jo & Ma 1996), and waters off the North China Coast (Shen et al. 1989). To provide an additional taxonomic char- acter and to ease total length measurements, the relationship between carapace length of A. bowmani (measured along the dorsal midline from rostral tip to posterior margin) and its total length (measured from rostral tip to posterior margin of telson) was de- termined using a least square linear regres- sion. This relationship is described by the equation, total length = 3.69 (carapace length) — 0.07; with a sample size of 28 specimens, the correlation coefficient equaled 0.98. Ecological notes.—Acanthomysis bow- mani is most abundant in June and July in Suisun Bay (Fig. 1) at about 2.0%coS. Water temperature at which type specimens were collected was 16° C, specific conductivity equaled 340 wS/cm, <0.5%0S. Etymology.—Named in honor of the late Dr. Thomas E. Bowman, curator of Crus- tacea U.S. National Museum of Natural History and our close colleague, who con- tributed greatly to the systematic knowl- edge of the Mysidacea. Acanthomysis aspera Ili, 1964 Acanthomysis aspera, reported from coastal waters around Japan (Mauchline & Murano 1977), China (Shen et al. 1989) and Korea (Jo & Ma 1996) has begun to appear in collections from the western end of Suisun Bay, downstream of A. bowmani sampling locations (Fig. 1). This is the first record of the occurrence of this species in the eastern Pacific. Dr. Tom Bowman veri- fied the identification of the specimens. Synanthropic introductions.—Historical and biogeographic evidences strongly sug- gest that A. bowmani and A. aspera were introduced to the Sacramento-San Joaquin Estuary with the flushing of ship ballast wa- 444 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Acanthomysis bowmani, new species, A. Endopod of 3rd thoracic limb; B. Exopod of 3rd thoracic limb; C. Penis; D. Pleopod 1; E. Pleopod 4; E Pleopod 5; G. Uropod endopod (upper), exopod (lower); H. Telson. Male, 11.6 mm. VOLUME 110, NUMBER 3 445 Table 1.—Comparison of diagnostic characteristics of Acanthomysis bowmani, n. sp., taken from the waters of the Sacramento-San Joaquin Estuary, California, with those noted by Ii (1964) for A. sinensis, and A. lon- girostris from waters off the western Pacific Ocean. Characters A. bowmani A. sinensis A. longirostris Rostrum Eye stalk Antennal scale Carpopropodus 4th pleopod of male Uropod Telson Apex of telson Reaches middle of anten- nular peduncle first seg- ment Proximal half without spi- nules 8.4X as long as broad 9 segments 2-segmented, Ist 1.2 longer than 2nd Endopod with 2 spines near statocyst 3—4 basal marginal spines, 18—19 distal marginal spines Narrowly truncated, with 2 terminal spines and 2 Reaches middle of anten- nular peduncle first seg- ment Proximal half without spi- nules 5.0X as long as broad 10 segments 2-segmented, Ist 30 lon- ger than 2nd Endopod with 2 spines near statocyst 4 basal marginal spines, 13-14 distal marginal spines Broadly truncated, with 4 terminal spines Long, reaches second joint of antennular peduncle Proximal half with spi- nules 7.0X as long as broad 9-11 segments 2-segmented, Ist 22 lon- ger than 2nd Endopod with 2-3 spines near statocyst 3 basal marginal spines, 20 distal marginal spines Narrowly truncated with 2 terminal spines long ultimate marginal spines Distribution Coastal Offshore Coastal ter (Carlton 1979). Acanthomysis aspera has been reported from coastal regions of Japan (li 1964), China (Shen et al. 1989) and Korea (Jo & Ma 1996), while morpho- logical characteristics of A. bowmani show very close affinity with congeneric species that also have only been reported from the western Pacific. Although the Sacramento- San Joaquin Estuary has been continuously and systematically surveyed for the past 25 years, neither species occurred in any sam- ples taken at the same collecting sites and at similar times prior to 5 August 1992. Other crustaceans foreign to the Sacra- mento-San Joaquin Estuary have been re- ported. Ferrari & Orsi (1984) found a co- pepod, Limnoithona sinensis Burckhardt, previously reported from the Yangtze River delta and described a new species, Oithona davisae, which is also common in the West- ern Pacific. Likewise, Bowman and Orsi (1992) suggest that Deltamysis holmquistae may also have been introduced. A possible site of introduction of the Acanthomysis spp. is the Port of Oakland, since it receives a considerable amount of shipping from the Far East. Research on these exotic species is currently underway because they may have a detrimental affect on the endemic Mysidacea. Acknowledgments We would like to thank all those from the California Department of Fish and Game that assisted in the collection and process- ing of the field material. Sally Skelton first detected A. bowmani in the samples being processed from Suisun Bay. The Interagen- cy Ecological Program provided support for the sampling work. Literature Cited Bowman, T. E., & J. J. Orsi. 1992. Deltamysis ho- Imquistae, a new genus and species of Mysi- dacea from the Sacramento-San Joaquin Estu- ary of California (Mysidae: Mysinae: Hetero- mysini).—Proceedings of the Biological Soci- ety of Washington 105:733-742. Carlton, J. 1979. Introduced invertebrates of San Francisco Bay. Pp. 427—444 in T. Conomos, ed., San Francisco Bay: the urbanized estuary. Pa- cific Division, American Association for the 446 Advancement of Science, San Francisco, Cali- fornia, 493 pp. Ferrari, EF D., & J. Orsi. 1984. Oithona davisae, new species, and Limnoithona sinensis (Burckhardt, 1912) (Copepoda: Oithonidae) from the Sacra- mento-San Joaquin Estuary, California.—Jour- nal of Crustacean Biology, 4:106—126. 1964. Fauna Japonica, Mysidacea (Crustacea). Biogeographical Society of Japan, Tokyo, 610 Pp. Jo, S.-G., & C.-W. Ma. 1996. Mysidacea (Crustacea from the West Coast of Korea.—Journal of the Korean Fishery Society, 29:805—827. hi, Né PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Mauchline, J., & M. Murano. 1977. World list of the Mysidacea, Crustacea.—Journal of the Tokyo University of Fisheries 64:39-88. Shen, C. J., J. Y. Liu, & W. Shaowu. 1989. Mysidacea in waters off the North China Coast.—Studia Marina Sinica 30:189-—227. Tattersall, W. M. 1932. Contribution to a knowledge of the Mysidacea of California. II: the Mysi- dacea collected during the survey of San Fran- cisco Bay by the U.S.S. Albatross in 1914.— University of California Publications in Zoolo- gy 37:315-347. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):447—456. 1997. Lamellibrachia satsuma, a new species of vestimentiferan worms (Annelida: Pogonophora) from a shallow hydrothermal vent in Kagoshima Bay, Japan Tomoyuki Miura, Junzo Tsukahara, and Jun Hashimoto (TM) Faculty of Fisheries, Kagoshima University, 4-0-20, Shimoarata, Kagoshima 890, (JT) Faculty of Sciences, Kagoshima University, 1-21-35, Korimoto, Kagoshima 890, and (JH) Japan Marine Science and Technology Center, 2-15, Natsushima, Yokosuka 237, Japan Abstract.—Vestimentiferan tube worms were found forming clumps in 82— 110 m in Kagoshima Bay, southern Japan, during a series of surveys exploring the biological community associated with shallow hydrothermal vents in 1993. The newly found tube worm is described here as Lamellibrachia satsuma, a new species. It differs from other congeneric species in having a short vesti- mentum, a short obturaculum, up to 4 pairs of lamellar sheaths and up to 19 pairs of branchial lamellar sheaths. About 20 living worms were maintained in a laboratory for more than 400 days. Release of eggs from the opening of the tube was observed at the beginning of the maintenance experiment. Trocho- phore-like larvae were also examined and photographed. Kagoshima Bay is characterized by two gigantic calderas, the Aira and the Ata, which form the northern and the southern areas of the bay, respectively. The northern bay-head area is about 18 km long and 20 km wide and is separated from the southern area by an active volcano, Mt. Sakurajima, but connected by a shallow and narrow strait (40 m deep and 2 km wide). The erup- tion forming the Aira Caldera that is the present bay-head area is thought to have oc- curred about 22,000 years ago (Aramaki & Ui 1966). In the east part of the head area, two hydrothermal vent sites at depths of about 80 m and 200 m are recognized by the appearance of gas bubbles that reach the surface and are called “‘Tagiri’’ by local fishermen (Oki & Hayasaka 1978). The word “‘Tagiri’’ originates from a Japanese word meaning ‘boil and bubble’. Although ‘““Tagiri”’ sites are quite shallow when com- pared with other hydrothermal vents, the chemical characteristics of these sites are believed to be the same or close to those of hydrothermal vents where associated bio- logical communities are found. In 1993, during a series of surveys exploring the community associated with submarine fu- maroles in Kagoshima Bay, we discovered a world of vestimentiferan worms forming clumps at depths of 82—110 m and collected a batch of living worms by means of a small dredge attached to a deep towed cam- era system (Hashimoto et al. 1993). The first vestimentiferan species, Lamel- librachia barhami, was described as a unique pogonophoran worm from the Northeast Pacific at a depth of 1125 m (Webb 1969). In a study of L. luymesi col- lected at 500 m depth off Guyana, the class Vestimentifera was placed in the phylum Annelida (van der Land & N@rrevang 1975). Jones (1985) proposed, however, a new phylum, Vestimentifera, for the vesti- mentiferan tube worms in working on above species and various other species from deep-sea vents and seeps. Mafie-Gar- zon & Montero (1985) also proposed a sep- arate phylum under the name Mesoneuro- phora with the description of a new species of Lamellibrachia. Describing two new vestimentiferan species, Southward (1991) 448 reclassified the vestimentiferan worms as a subclass of the class Pogonophora within the phylum Annelida. The affiliation of ves- timentiferans to Annelida is also suggested by the hemoglobin structure (Suzuki et al. 1989) and by the amino acid sequence of Elongation Factor-la (Kojima et al. 1993), the analysis of 28S ribosomal DNA dem- onstrated that the Vestimentifera form a monophyletic group to the exclusion of the polychaete Melinna and the periviate po- gonophore Siboglinum used for comparison (Williams et al. 1993). Rouse & Fauchald (1995) recently proposed that the name Ar- ticulata be used to include the Vestimenti- fera and Pogonophora as well as the Clite- lata, the Polychaeta, and the Euarthropoda and Onychophora. In this study, we follow the classification proposed by Southward (1991) and place the vestimentiferan worms in the phylum Annelida. To date, eight genera and 13 species of vestimentiferans have been described from hydrothermal vents, cold seeps, and other deep-sea bottoms (Jones 1985, Southward 1991, Southward & Galkin 1997). Two of them were amalgamated into a single spe- cies on the basis of careful examination of morphology and allozymes (Southward et al. 1995). Vestimentiferans are known from depths of 300—3270 m, in the eastern Pa- cific and the Gulf of Mexico (Jones 1985), the Lau Basin (Southward 1991), the Ma- nus Basin (Southward & Galkin 1997), off Guyana (van der Land & N@grrevang 1975), off Uruguay (Mafie-Garzon & Montero 1986), and from the eastern Atlantic (Dan- do et al. 1992). Around Japan, several types of vestimentiferans have been also ob- served by manned or unmanned deep-sea submersibles and captured from deep-sea chemosynthetic communities in Sagami Bay (Hashimoto et al. 1989) and the mid- Okinawa Trough (Hashimoto et al. 1995) between depths of 690—1370 m in tectoni- cally active zones. The vestimentiferans in Kagoshima Bay represent the shallowest occurrence of the group in the world. Re- cently, partial nucleotide sequences of mi- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tochondrial DNA were analyzed from sev- eral local populations of Japanese vestimen- tiferans (Kojima et al. 1995). This analysis suggested the presence of more than one genetically distinguishable populations of Lamellibrachia species-complex around Ja- pan. The authors also suggested that the populations differed in their vertical distri- bution. In this study, a shallow water spe- cies from Kagoshima Bay is described. Material and Methods More than a hundred vestimentiferan specimens were collected by a small dredge 40 cm wide X 10 cm high X 50 cm deep, attached about 1 m beneath the bottom of a deep towed color television camera sys- _ tem operated by R/V Kaiyo, in the northern semi-closed area of Kagoshima Bay in Feb- ruary, 1993. Other additional specimens were collected by an unmanned deep-sea research vehicle Dolphin 3K seven months later, at the same site. Specimens for taxonomic study were fixed in 10% formalin and transferred to 70% ethanol. Some dozens of worms were dissected from their tubes on the vessel and the remainder were kept intact. Specimens for chemical analysis were frozen at —80°C, a part of which were used in this study to calculate the dry/wet weight ratio of the soft body. Living worms were washed very carefully by using a brush to eliminate sed- iments and to examine newly settled larvae that might attach on the adult tubes. Every ten to twenty cleaned specimens were trans- ferred into a 10-liter bottle and kept in the refrigerator of the research vessel at about 5°C. The living animals were transported to the laboratory of the Faculty of Fisheries at Kagoshima University within a week after the sampling. The worms were maintained in a glass tank 30 cm wide X 18 cm long x 24 cm deep filled with 10 1 of filtered seawater (Millipore 1.2 wm filter). The tank was maintained at 16°C in an-incubator be- cause the temperature of the bottom sea- water at the collecting site is constant VOLUME 110, NUMBER 3 throughout the year at about this value. The water was changed for every 7—14 days to maintain a salinity of about 34 parts per thousand and a pH value of about 8.2, even though this last value altered by the addi- tion of sodium sulfide. The worms were provided on average with 1.6 g of sodium sulfide (Na,S-9H,O) as a source of hydro- gen sulfide twice a day. The maximum con- centration of hydrogen sulfide was calcu- lated to be about 0.7 mM per 1 of sea water (not analyzed). The concentration in culture was, therefore, about 35 times higher than the concentration of hydrogen sulfide in the field, where only 0.02 mM per 1 was re- corded. The types are deposited in the National Science Museum, Tokyo (NSMT), Japan Marine Science and Technology Center (JAMSTEC), the National Museum of Nat- ural History, Smithsonian Institution (USNM), the Los Angeles County Museum of Natural History (LACM-AHP), the Mu- seum National d’ Histoire Naturelle de Paris (MNHN), and the Australian Museum, Syd- ney (AM). Family Lamellibrachiidae Webb, 1969 Genus Lamellibrachia Webb, 1969 Lamellibrachia satsuma, new species Figs. 1-5 Material examined.—Holotype (NSMT- Pc-H3), 15 paratypes (5: NSMT-Pc-P4, P5, P6, P7, P8; 10: USNM_ 175102-175111), Kagoshima Bay, Deep Tow Camera Obser- vation DT-13, 6 Feb 1993, 31°39.55’N, 130°48.07'E, 98 m; 13 paratypes (5: MNHN UE798—UE802, 8: JAMSTEC Ves- 0270-77-93), DT-15, same site, 6 Feb 1993, 102 m; 6 paratypes (3: MNHN UE803-— UE805, 3: JAMSTEC Ves-0278-80-93), Dolphin 3K Dive 154, 12 Sep 1993, 31°39.83'N, 130°48.97’E, 101 m; 40 para- types (10: USNM_ 175092-175101; 10: LACM-AHF POLY 1873—POLY 1882; 10: AM W23599-—W235608; 10: JAMSTEC Ves-0281-90-93), Dolphin 3K Dive 157, 12 Sep 1993, 31°39.80’N, 130°48.05’E, 122 m; 449 7 paratypes (JAMSTEC Ves-0291-97-93), Dolphin 3K Dive 164, 16 Sep 1993, 31°39.70'N, 130°48.02’E, 110 m. Measurements.—Tube length 60—1000 mm (X 317 mm, n = 71); opening width of top collar 2.5—-8.7 mm (X 5.6 mm, n = 78), bottom width of top collar 1.7—7.2 mm (X 3.9 mm, n = 81), width of basal end 0.5— 2.4 mm (X 1.1 mm, n = 53); tube wet weight 0.18-4.16 g (X 1.35 g, n = 53). Body length 45—443 mm (X 219 mm, n = 53); body wet weight 0.08-1.57 g (X 0.68 g, n = 53) (dry/wet weight ratio 0.06—0.28, X 0.23, n = 37, measured differently using materials for chemical analysis). Obturacu- lar length 1.8—9.8 mm (X 5.2 mm, n = 64); obturacular width 1.0—5.6 mm (X 4.1 mm,. n = 53). Vestimental length 7.2—24.0 mm (X 17.2 mm, n = 64). Vestimental length/ Obturacular length ratio 2.1-8.3 (X 3.5, n = 64); Vestimental length/obturacular width ratio 2.3—9.7 (X 4.5, n = 53). Obtur- acular lamellar sheaths 0—4 pairs (X 2.77, n = 57). Sex ratio 1.39:1 (32 males: 23 fe- males). Description.—Anterior face of obtura- culum of adult bare, lacking secreted struc- tures (Figs. la, b, 2a); with up to 19 pairs of branchial lamellae (Figs. la, b, 2a, b); each lamella formed by a single series of fused branchial filaments with pinnules; branchial lamellae hidden by up to four pairs of peripheral lamellar sheaths (Figs. la, b, 2a, b); sheaths composed of fused fine filaments, with only extreme distal tips of filaments free (Figs. la, b, 2a, b); obtur- aculum lenticular in transverse section (Fig. 2b), lacking dorsal groove, with ventral ridge, distally; with single, medial excreto- ry pore opening in groove at base of ob- turaculum. Anterior margin of vestimentum forming short sheath or collar extending around base of obturaculum; central dorsal surface of vestimentum of male (holotype) with paired ciliated grooves, diverging at anterior ends (Fig. 1a); postero-ventral mar- gin of vestimentum broadly incised (Fig. la, b); ventral surface with numerous small papillae topped by cuticular plaques 35—63 450 Fert PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lamellibrachia satsuma, new species.—Holotype. a, Anterior end, in dorsal view; b, Same, in ventral view. Body fixed after removed from the tube. Scale = 3 mm. 4m in diameter. Trophosome very long, with numerous small papillae topped by cu- ticular plaques 51—82 zm in diameter. Op- isthosome of selected specimen with 33 segments (Fig. 3a—c), anterior 27 with a sin- gle row of setae (Fig. 3b, d—f); most setae with two groups of denticles, anterior group bearing 6—9 denticles, posterior group with. 10-15 denticles in 3—4 rows (Fig. 3e-f). Tube with obvious growth collars and ir- regularly placed light and dark bands (Figs. 2a, 4). Anterior parts of tubes more straight than posterior in large specimens; posterior parts coiled in general. Tubes tangled to- gether in tight clusters, making large hemi- spherical clumps on thick sediments; clumps sometimes more than 10 m in di- ameter. Etymology.—The specific epithet satsu- ma, a noun in apposition, refers to the old province on Kyushu island. Japanese name of the species, ‘“‘satsuma-haorimushi’’ is also composed of the provincial name and the group name of vestimentiferan worms. Remarks.—Among the five described species of the genus Lamellibrachia, L. sat- suma and L. barhami Webb, 1969 differ from the other three by the number of pe- ripheral lamellar sheaths, i.e. up to four pairs in the first two species, six pairs in L. VOLUME 110, NUMBER 3 451 Fig. 2. Lab-reared specimens and embryos of Lamellibrachia satsuma.—a, Anterior end with the obtura- culum, branchiae and the lamellar sheaths extending from the tube, scale = 3 mm; b, Transverse section near base of obturaculum, scale = 1 mm; c, Regenerated posterior ends of tubes attached to the glass bottle surface, scale = 10 mm; d, Released egg, scale = 0.1 mm; e, Trochophore-like larva about two-weeks old, in anterior view (upper hemisphere), scale = 0.1 mm; f, Same, in latero-posterior view (lower hemisphere with telotroch- like ciliated area), scale = 0.1 mm. luymesi van der Land & N@rrevang, 1975, three to four pairs of lamellar sheaths, but seven pairs in L. victori Mafe-Garz6n & only one or two pairs are large enough to Montero, 1985, and 8-16 pairs in L. col- cover the branchial region in most case. umna Southward, 1991. In L. satsuma, One very small specimens considered as a more than half of examined specimens has juvenile of L. satsuma has no lamellar PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Seok TN | Tis ua ’ hee Gee Y's we ,, = See = Veh ae Fig. 3. Lamellibrachia satsuma, new species.—a, Regenerated opisthosome from a lab-reared specimen; b, Anterior portion of the same enlarged; c, Posterior portion of the same enlarged; d, Portion of opisthosome with a single row of setae; e, and f, Setae. Scale bars = 0.1 mm for a—c and 0.002 mm for d-f. VOLUME 110, NUMBER 3 & \ 157-20 ai" ’ 453 Fig. 4. Lamellibrachia satsuma, new species.—Anterior portion of tubes of selected paratypes. sheaths. Lamellibrachia satsuma has up to 19 pairs of branchial lamellae; this number is smaller than in L. barhami which has as many as 25 pairs of branchial lamellae (Webb 1969; Jones 1985). The ratio of ob- turacular length to width ovewrlaps in these two species, but more variable in L. bar- hami (Fig. 5). Lamellibrachia satsuma is slightly smaller in size and has a shorter vestimentum and obturaculum (Fig. 5). The diameter of cuticular plaques on the vestimentum and the trophosome are mea- sured for L. columna (vestimental plaques 65-90 wm, trunk plaques 70—120 wm) and for L. barhami (60-115 wm, 115-160 wm, respectively) by Southward (1991, also her personal communication). These plaques are smaller in L. satsuma (35—63 wm, 51- 82 wm, respectively) than the above two species. The tube form is slightly different in L. barhami and L. satsuma. The tubes are twisted throughout the length in L. bar- hami, whereas the anterior parts of tubes are relatively straight in L. satsuma. Lamelli- brachia satsuma forms very crowded clumps on the sediment and the tubes ap- pear to be apart from one another in grow- ing straightly (Fig. 2 in Hashimoto et al. 1993). These two species differ from each other also in their vertical and ecological distribution, i.e. L. satsuma lives in volca- nic vents at depths of 98-110 m and per- haps also in cold seep sites of about 300 m depth, whereas L. barhami is presently known only in cold seep sites at depths of 1100—2000 m. Observation of living worms in labora- tory.—The worms extended the anterior part of their body from the tube, exposing the obturaculum and the lamellar sheaths (Fig. 2a). The anterior end was extended whether light was on or off, even when the strobe light was flashed. However, the 454 w@ L. barhami @ L. satsuma Obturacular Length (mm) Obturacular Length / Width 0 10 20 30 40 Vestimental Length (mm) Fig. 5. Relationship between obturacular length and vestimental length (lower) and between obtura- cular length/width ratio and vestimental length (upper) in Lamellibrachia satsuma (closed circle, present study) and L. barhami (closed square, data from Jones 1985). worms were very sensitive to vibration and withdraw into the tube when the tank was shaken. Retraction was very quick. Among 43 living specimens maintained in the laboratory tank, six died in the first 75 days. Some of the remaining specimens were occasionally removed and dissected to check their condition. About 20 specimens were maintained alive for more than 400 days. During the maintenance experiment in the laboratory, the worms regenerated their damaged posterior ends. Regenerated opis- thosomes were observed at least two months after capture. The posterior ends of tubes were repaired and elongated to attach PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON to a glass bottle which was used as a sinker for a bundle of worm tubes (Fig. 2c). Elon- gation of the anterior ends of tubes was not observed. Spawning.—Some bottles of specimens were kept in the refrigerator of the research vessel at about 5°C for less than 6 days after their capture before transfer to the rearing tank. In a bottle containing about ten spec- imens, one released eggs from its tube opening when it was transferred to a tank filled with seawater at about 15°C (room temperature). The newly released eggs were photographed (Fig. 2d) and recorded on videotape. The eggs ranged in diameter from 0.10—0.11 mm (X 0.102, n = 11). The eggs were neutrally buoyant in seawater and formed a cloud-like mass drifting in the ‘water, but most eggs transferred to a shal- low petri dish floated at the surface. Cleav- age of these eggs did not occur in the next five days. Observation was then abandoned. Larvae.—In another bottle kept in the re- frigerator, developing embryos were found five days after collection. Most of them were in a swimming, blastula-like stage. These embryos were kept in a small dish at room temperature ranging from 14.0 to 15.5 °C. Most of the embryos developed into trochophore-like larvae (Fig. 2e, f) 0.10 mm in diameter and 0.12—0.13 mm in length (n = 7). The larva had a well-ciliated anterior hemisphere with a prototroch-like ciliated band (Fig. 2e). A telotroch-like cil- iated area was also well developed (Fig. 2f). A dark mass observed inside the larva (Fig. 2f) was thought to be comparable with the primitive gut of the early polychaete troch- ophore, but no blastopore or proctodaeum were observed. The gut-like structure of the larva was completely closed. No larvae had settled after two weeks of observation, when most larvae became inactive. Discussion Kojima et al. (1995) recognized three ge- netic types of Lamellibrachia in the waters around Japan by differences in the amino VOLUME 110, NUMBER 3 acid sequences of the mitochondrial cyto- chrome c oxidase I. Some Lamellibrachia specimens collected from Kagoshima Bay (82—110 m) and those from the Kanasunose Bank of the Nankai Trough (300 m) cluster into one of their types, living shallower than 300 m, which is the species described here as L. satsuma. This is thought to be distributed widely on the Southwest coast of Japan in less than 300 m deep. Judging by our preliminary examination of a single specimen from Sagami Bay, another types of Lamellibrachia from deeper sites (Koji- ma et al. 1995) is probably distinguishable from L. satsuma in morphological charac- ters such as the body size, tube form, and the number of lamellar sheaths. As shown in our maintenance experi- ment, it is not difficult to keep the mouth- less worms alive if a suitable chemical en- ergy source such as hydrogen sulfide is sup- plied. The expanded, blood-red branchiae forming flower-like fans at the top of the tubes may also serve to take up oxygen and carbon dioxide from the water. On the other hand, the source of nitrogen required for their growth or maturation is not yet iden- tified. Since no growth except for some re- generation of posterior parts was observed in reared specimens, a suitable source of ni- trogen such as nitrate or dissolved amino acid might have been scarce in the labora- tory conditions. Young et al. (1996) reported observations on the development of two vestimentiferans Lamellibrachia sp. and Escarpia sp. in the Gulf of Mexico. Lamellibrachia satsuma has eggs as large as those of the former but some differences can be seen. The larva of L. satsuma has a teminal telotroch-like cil- iated area and an internal gut-like dark mass of cells, which were not found in the two species reported by Young et al. (1996). It is not yet clear if these differences are spe- cies-specific or not. Our study on the early development of L. satsuma is not yet com- pleted and more details will be published elsewhere. The length of the planktonic lar- val life of L. satsuma suggested by the 455 slightly buoyant eggs and a trochophore- like larva able to swim for more than two weeks is enough to explain their distribu- tion in several sites along the Southwest coast of Japan washed by the strong Ku- roshio Current. The Kagoshima Bay popu- lation of L. satsuma might have been formed originally by pioneer larvae re- leased from a neighboring population out- side the bay. The larvae might have settled at the active volcanic vents which are thought to have been more numerous and active in the bay during its formative period than at present. The northern part of the bay has been almost closed by the formation of Mt. Sakurajima during 13,000 years (Ara- maki & Ui 1966), since the postulated col- onization by the pioneer worms. Acknowledgments We thank the captain and crew of R/V Kaiyo and R/V Natsushima, and the oper- ation team of Dolphin 3K for their coop- eration and enthusiasm throughout the sur- veys. Our special thanks are also due to M. Nedachi, K. Oki, and H. Sakamoto of Ka- goshima University, J. Ossaka of Tamaga- wa University, and K. Fujikura of JAM- STEC for their enthusiasm and assistance of the surveys. The manuscript was im- proved by the careful review and correc- tions by E. C. Southward of Marine Bio- logical Association, J. L. Norenburg of Smithsonian Institution, and an anonymous reviewer. This work was supported in part by grant-in-aid for the first author from the Ministry of Education, Science, Sports, and Culture, Japan (No. 06839021). Literature Cited Aramaki, S., & T. Ui. 1966. The Aira and Ata pyro- clastic flows and related calderas and depres- sions in southern Kyushu, Japan.—Bulletin of Volcanology 29:29-—47. Danndo, P. R., A. J. Southward, E. C. Southward, D. R. Dixon, A. Crawford, & M. Crawford. 1992. Shipwrecked tube worms.—Nature 356:667. Hashimoto, J., S. Ohta, K. Fujikura, & T. Miura. 1995. Microdistribution pattern and biogeography of 456 the hydrothermal vent communities of the Min- ami-Ensei Knoll in the Mid-Okinawa Trough, Western Pacific.—Deep-Sea Research, Part I, Oceanographic Research Papers 42:577—598. , I. Tanaka, H. Hotta, S. Matsuzawa, & H. Sakai. 1989. Deep-sea communities dom- inated by the giant clam, Calyptogena soyoae, along the slope foot of Hatsushima Island, Sa- gami Bay, Central Japan.—Palaeogeography, Palaeoclimatology, and Palaeoecology 71:179-— 192. , TI. Miura, K. Fujikura, & J. Ossaka. 1993. Discovery of vestimentiferan tube-worms in the euphotic zone.—Zoological Science 10:1063-— 1067. Jones, M. L. 1985. On the Vestimentifera, new phy- lum: Six new species, and other taxa, from hy- drothermal vents and elsewhere.—Bulletin of the Biological Society of Washington 6:117— 158. Kojima, S., J. Hashimoto, & S. Ohta. 1995. The dis- tribution and the phylogenies of the species of genus Calyptogena and those of vestimentifer- ans around Japan.—JAMSTEC Journal of Deep Sea Research 11:243-—248. (In Japanese with English abstract). ‘ , M. Hasegawa, S. Murata, S. Ohta, H. Seki, & N. Okada. 1993. Close phylogenetic relationship between Vestimentifera (tube worms) and Annelida revealed by the amino acid sequence of elongation factor-1a.—Journal of Molecular Evolution 37:66—70. Mané-Garzon, E, & R. Montero. 1985. Sobre una nueva forma de verme tubicola—Lamellibra- chia victori n. sp. (Vestimentifera)—Proposi- ci6n de un nuevo phylum: Mesoneurophora.— Revista de Biologia del Uruguay 8:1—28. Oki, S., & S. Hayasaka. 1978. Geological study on Kagoshima Bay, South Kyushu, Japan. Part IV—A note on the peculiar mode of occurrence of foraminifers in the bottom sediments of the bay-head area.—Report of Faculty of Science, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Kagoshima University, in Earth Science and Bi- ology 11:1-11. Rouse, G. W., & K. Fauchald, 1995. The articulation of annelids.—Zoologica Scripta 24:269-301. Southward, E. C. 1991. Three new species of Pogon- ophora, including two vestimentiferans, from hydrothermal sites in the Lau Back-arec Basin (Southwest Pacific Ocean).—Journal of Natural History 25:859-881. , & S. V. Galkin, 1997. A new vestimentiferan (Pogonophora: Obturata) from hydrothermal vent fields in the Manus Back-are (Bismarck Sea), Papua New Guinea, Southwest Pacific Ocean.—Journal of Natural History 31:43—55. , V. Tunnicliffe, & M. Black. 1995. Revision of the species of Ridgeia from northeast Pacific hydrothermal vents, with a redescription of Ridgeia piscesae Jones (Pogonophora: Obturata = Vestimentifera).—Canadian Journal of Zool- ogy 73:282—295. Suzuki, T., T. Takagi, K. Okuda, T. Furukohri, & S. Ohta. 1989. The deep-sea tube worm hemoglo- bin: Subunit structure and phylogenetic rela- tionship with annelid hemoglobin.—Zoological Science 6:915—926. van der Land, J., & A. Ngrrevang. 1975. The system- atic position of Lamellibrachia (Annelida, Ves- timentifera).—Zeitschrift fiir zoologische Sys- tematik und Evolutionsforschung, Sonderheft 1975:86—101. Webb, M. 1969. Lamellibrachia barhami, gen. nov., sp. nov. (Pogonophora), from the northeast Pa- cific.—Bulletin of Marine Science 19:18—47. Williams, N. A., D. R. Dixon, E. C. Southward, & P. W. H. Holland. 1993. Molecular evolution and diversification of the vestimentiferan tube worms.—Journal of the Marine Biological As- sociation of the United Kingdom 73:437—452. Young, C. M., E. Vazquez, A. Metaxas, & P. A. Tyler. 1996. Embryology of vestimentiferan tube worms from deep-sea methane/sulphide seeps.—Nature 381:514—516. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):457—470. 1997. The polychaetous annelids from oil platforms areas in the southeastern Gulf of Mexico: Phyllodocidae, Glyceridae, Goniadidae, Hesionidae, and Pilargidae, with description of Ophioglycera lyra, a new species, and comments on Goniada distorta Moore and Scoloplos texana Maciolek & Holland Alejandro Granados-Barba and Vivianne Solis-Weiss Laboratorio de Ecologia Costera (poliquetos), Instituto de Ciencias del Mar y Limnologia, UNAM Apdo. Postal 70-305. México, D.F, 04510, México Abstract.—The distribution and composition of the polychaete families Phyl- lodocidae, Glyceridae, Goniadidae, Hesionidae and Pilargidae of the oil plat- forms area of Campeche Sound, southeastern Gulf of Mexico are analyzed. They involve the identification of 14 species and the revision of some of the material reported by authors in Uebelacker & Johnson (1984) for these families. Ophioglycera lyra, a new goniadid species is described, diagnosis of genus Ophioglycera is emended. Goniada distorta Moore, 1903 is removed from Ophioglycera and maintained in the genus Goniada. Ancistrosyllis sp. B of Wolf, 1984 is reported as A. commensalis Gardiner, 1976, and Naineris sp. A of Taylor, 1984 is reported as Scoloplos texana Macioleck & Holland, 1978. Despite the fact that the Campeche Sound, in the southeastern Gulf of Mexico, is the most important area of oil extraction activity coupled with the most important shrimp fishery zone in Mexico, its fauna is poorly known. A few years ago, regional surveys were undertaken under our direc- tion to study the benthic fauna of the area surrounding the offshore oil platforms. Part of the results of the extensive polychaete collections made during these studies are presented here. Previous taxonomic reports on other polychaetes collected during these studies include Granados-Barba (1994), Granados-Barba & Solis-Weiss (1994), So- lis-Weiss et al. (1994, 1995) and Granados- Barba & Solis-Weiss (1997). When doubts arose over the taxonomic position of several species, comparisons were made with material collected in the northern and eastern continental shelves of the Gulf of Mexico by different authors who collaborated in the Taxonomic Guide of the Polychaetes of the Northern Gulf of Mexico by Uebelacker & Johnson (1984) and deposited in the Smithsonian Institution (USNM) collection, since affinities with the study area environment were obvious. The appropriate remarks are herein reported where relevant. Study area.—The study area is located in Campeche Sound, between 18°46’—20°03'N and 91°33'—92°34'W, covering the offshore oil platforms area and part of the continen- tal shelf down to about 200 m depth (Fig. 1). The sediment in the oil platforms area is mainly mud, although there are some iso- lated patches of sandy mud (Granados-Bar- ba 1994). Materials and methods.—Sampling was done on board the R/V Justo Sierra, as part of the interdisciplinary projects IMCA- DINAMO during expeditions IMCA—1 (I-1, March 1988), IMCA—2 (I-2, Septem- ber 1988), IMCA—3 (1-3, March 1989), DINAMO—1 (D-1, March 1990) and DIN- AMO—2 (D-2 November 1990). Sixteen stations are herein considered (Fig. 1). The 458 20 30) ® w I I I & I I Fa N N Ne \\ wr \ \\ Grijalva- Usumacinta’ = River Fig. 1. Study area. 1 Sampling stations. coordinates and depths of each station are presented in Table 1. The polychaetes were collected with a 0.1 m?* Smith-MclIntyre grab. At each station, about 40 liters of sed- iment were screened through a 0.5-mm Table 1.—Positions (Latitude and Longitude) and depths (m) of the sampling stations. Latitude Longitude Station (N) (W) Depth 1 19°15’ 92°08’ 312 De 19°23’ 92°21’ 75.6 3 19°18’ 92°28’ 102.6 4 TOPS; 92°28’ 71.8 5 19°05’ S223) 32.6 6 18°49’ 92°16’ 16 di 18°50’ 92°06’ 16 8 19°04’ 91°42’ 16.2 9 19°04’ O33! 16.6 10 19°20’ 91°39’ Ble? 11 19°20’ 91°49’ 32.4 12 19°33" 91°54’ 56 8 19°42’ 92°00’ 70.2 14 19°43’ O27 10" 98.2 15 19°52’ 92°16’ 148.4 16 20°03’ 92°09’ PATA PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON CAMPECHE SOUND oe Champoton River ~ TERMINOS £5. LAGOON _~~ Jj: 0 5 10 Nautic males sieve, and specimens fixed in 10% forma- lin. In the laboratory, specimens were washed, sorted, and transferred to 70% eth- anol. Depth, salinity, and temperature were recorded at each station with a Niels-Brown Cab: For each species, selected synonyms, habitat and reported worldwide distribution are included. Environmental factors mea- sured in this and previous studies that are included when available are cited with the following abbreviations: D = depth (m); T = temperature (°C); S = salinity (%); OM = organic matter in the sediment (% of or- ganic carbon) and OD = dissolved oxygen (ml/1). The specimens are deposited in the Polychaetological Collection of the Institu- to de Ciencias del Mar y Limnologia (CPICML), UNAM, Mexico, City. The type material for Ophioglycera lyra is de- posited in the following museums: National Museum of Natural History, Smithsonian Institution (USNM), Washington, D.C.; Los Angeles County Museum of Natural His- tory (LACMNH-AHB), California, USA. VOLUME 110, NUMBER 3 Family Phyllodocidae Orsted, 1843 The family Phyllodocidae is represented by about 31 genera and 306 species. In this study, 11 specimens belonging to two gen- era and four species were collected. Pleijel (1991:232) clarified the authorship of the family. See also Blake (1994a). Genus Paranaitis Southern, 1914 Paranaitis gardineri Perkins, 1984 Paranaitis gardineri Perkins, 1984:563, figs. 4a—1.—Granados-Barba, 1994:112. Paranaitis polynoides.—Gardiner, 1976: 110.—Gathof, 1984:19.21, fig. 19.18 [not Anaitis polynoides Moore, 1909]. Material examined.—5 specimens: Mar 1989, sta. 5(1), 9(1), 11(1); Mar 1990, sta. 16(1); Oct—Nov 1990, sta. 16(1). Remarks.—Perkins (1984) erected the species P. gardineri to include specimens reported by Gardiner (1976) as P. polyno- ides separating Atlantic Ocean specimens from P. polynoides Moore, 1909, based on the fact that P. gardineri has dorsal cirri narrower and shorter than the acicula that are not covering the dorsum in the posterior region, in addition to the shape of the anal cirri, which are long and filiform rather than short, thick and cylindrical. The specimens examined in this study agree with the de- scription of P. gardineri. Perkins further re- marked that the North Carolina specimens were twice as large as the ones from Flor- ida. In this study, the specimens are about half the size of the Florida specimens. Previously reported habitat.—Intertidal to 125 m, in mud, muddy sand, coarse to fine sand, sand with gravel and shells, T = 20-26; S = 36.48—37.43; OM = 0.19-1.17. Occurrence.—In mud, D = 16—127; T = 20; S = 36.48; OM = 1.17. Distribution in Mexico.—Southern Gulf of Mexico. Distribution.—North Carolina; Florida; northern Gulf of Mexico. 459 Genus Phyllodoce Lamarck, 1818 Phyllodoce (Phyllodoce) arenae Webster, 1879 Phyllodoce (Anaitides) arenae.—Pettibone, 1963:82, fig. 18a—Day, 1973:23.—Gar- diner, 1976:117, figs 8d. Phyllodoce arenae.—Gathof, 1984:19.21, figs. 19.18a—e.—Granados-Barba, 1994: 112. pill 5p. Phyllodoce (Anaitides) panamensis.—Day, 1973:24, fig. 3n—p.—Gardiner, 1976:117. Phyllodoce panamensis.—Granados-Bar- ba, 1994:114. (not Phyllodoce panamen- sis Treadwell, 1917.) Material examined.—4 specimens: Sep 1988, sta. 13(1); Mar 1989, sta. 9(3). Material of other species examined.— Phyllodoce panamensis USNM 16831, Ho- lotype from Port Chame, Panama, Sta. 362, 30°11'48"N, 88°43’40"W. USNM 51028, 2 specimens from Beaufort, North Carolina, USA 34°24'N, 75°57'W, 80 m. Remarks.—Day (1973) separated North Carolina specimens he reported as P. pan- amensis from P. arenae based on the dif- ference in color patterns of both species. Whereas in the former there is a middorsal continuous band, the latter is characterized by the presence of fusiform dark dorsal spots in the intersegmental furrows. In one of the specimens examined for this study, a middorsal continuous band is present, so that it was formerly identified as P. pana- mensis by one of us (Granados-Barba 1994). However, a recent comparison be- tween Day’s specimens, the holotype of P. panamensis and our specimens showed that P. panamensis Treadwell is considerably larger than the specimens from the Gulf of Mexico and North Carolina and that it does not have the middorsal band. We could con- clude that both our specimens and Day’s North Carolina specimens are P. arenae. Previously reported habitat.—Intertidal to 200 m, in mud, sandy mud, muddy sand and coarse sand with shells, T = 27.5; S = 36.21—36.76; OM = 0.64-1.01; DO = 3.45. 460 Occurrence.—In mud, D = 16-70. Distribution in Mexico.—Tamiahua La- goon, Veracruz; southern Gulf of Mexico. Distribution.—New England; North Car- olina; northern Gulf of Mexico. Phyllodoce (Anaitides) madeirensis Langerhans, 1880 Phyllodoce (Anaitides) madeirensis.—Day, 1973:23.—Gardiner, 1976:115, figs. 7q, 8a—c. Anaitides madeirensis.—Gathof, 1984: 19.39, figs. 34a-e. Phyllodoce madeirensis.—Mountford, 1991:161, figs. 2—3a—c.—Pleiel, 1991: 257; 1993a:33.—Granados-Barba, 1994: 113. Material examined.—1 specimen: Mar 1990, sta. 13(1). Remarks.—The specimen examined agrees with the redescription of Mountford (1991) of this species; the revision by this author of some phyllodocids from Puerto Rico led to the synonymy of P. oculata and P. madeirensis, and the separation of P. er- ythrophylla and P. madeirensis, based on color patterns. See also the list in Pleijel (i991): Previously reported habitat.—Intertidal to 200 m, on rocks and corals, in silt, clay, mud, sandy mud, muddy sand, sand and fine sand, T = 14-31; S = 34.44-37.4; OM = 0.18-5.5; DO = 1.03-—5.4. Occurrence.—In mud, D = 70; OM = 1.47. Distribution in Mexico.—Southern Baja California; Gulf of California; Jalisco; Co- lima; Guerrero; Gulf of Tehuantepec; Isla de Enmedio Veracruz; central and southern regions of the Gulf of Mexico. Distribution.—Cosmopolitan in intertidal areas and continental shelves of tropical seas; South Africa; pacific coast of Panama; North Carolina; northern Gulf of Mexico; Puerto Rico. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Phyllodoce (Anaitides) mucosa Orsted, 1843 Phyllodoce (Anaitides) mucosa.—Pettibo- ne, 1963:81, fig. 18f.—Gardiner, 1976: It. fis, 7: Anaitides mucosa.—Gathof, 1984:19.33, fig. 19.28. Phyllodoce mucosa.—Pleijel, 1991:259; 1993:51, figs. 33—34.—Granados-Barba, 1994:114. Material examined.—1 specimen: Sep 1988, sta. 15(1). Remarks.—The examined specimen agrees with description of Gathof (1984). Previously reported habitat.—Intertidal to 425 m, in mud, sand, muddy sand, sand and shells, and gravel, T = 13-15; S = 35- 35.46; OM = 3-6.9; DO = 0.8-3.09. Occurrence.—In mud, D = 148; T = 21; S = 36.42; OM = 1.15. Distribution in Mexico.—Baja Califor- nia; west of Baja California Sur; Gulf of California; Guerrero; southern Gulf of Mexico. Distribution.—Northern Europe; Azores Islands; western North America from Alas- ka to Mexico; western Africa; New En- - gland; North Carolina; northern Gulf of Mexico; Cuba. Family Glyceridae Grube, 1850 The family Glyceridae is represented by three genera and about 80 species. For this study, we collected 15 specimens belonging to one genus and three species. Genus Glycera Savigny in Lamarck, 1818 Glycera americana Leidy, 1855 Glycera americana.—Pettibone, 1963:213, figs. 54a—e.—Gardiner, 1976:161, figs. 171-n.—Gilbert, 1984a:32.15, fig. 32.12.—Granados-Barba, 1994:152.— Hilbig, 1994a:200, fig. 6.1. Material examined.—7 specimens: Mar 1988, sta. 7(1); Sep 1988, sta. 9(1), 141); Mar 1989, sta. 6(2), 7(1), 9(1). VOLUME 110, NUMBER 3 Remarks.—Everted branchiae were ob- served from setiger 14, but we consider that they could be retracted in anterior setigers. In very small specimens, the branchiae could not be seen; however, all the other characters agree with description of G. americana. Previously reported habitat.—Intertidal, continental shelf and slope, in mud, sandy mud, muddy sand and fine to medium sand, D = 22-106; T = 13-28; S = 35.06—37.19; OM = 0.47-3.9; OD = 1.04-5.4. Occurrence.—In mud and muddy sand, D = 16—98; T = 22-28; S = 35.51-37.19; OM = 0.39-1.05. Distribution in Mexico.—Baja Califor- nia; west of Baja California Sur; Gulf of California; Veracruz; Tamiahua Lagoon, Veracruz; central and southern regions of the Gulf of Mexico. Distribution.—Magellan Strait, New Zealand; Australia; Eastern Pacific Ocean, from southern Canada to Peru; Western At- lantic Ocean from New England to Argen- tina; northern Gulf of Mexico; Bermuda. Glycera robusta Ehlers, 1868 Glycera robusta.—Hartman, 1950:69, pl. 10, figs. 7-8.—Pettibone, 1963:218, figs. 54f—-g.—Gardiner, 1976:162, fig. 170.— Gilbert, 1984a:32.22, figs. 32.20a—i.— Granados-Barba, 1994:153. Material examined.—2 specimens: Mar 1990, sta. 2(1), 9(1). Remarks.—In one of the specimens, six to seven long structures resembling bran- chiae or cirri were observed on the first 10 setigers (very similar in shape to those found among the cirratulids); they do not follow a determined pattern, since some can be seen emerging above or below the dorsal cirrus, while others emerge from below the ventral cirrus. Hilbig (pers. comm.) sug- gests that these structures could be algae or fungi, or some other parasite/commensal. Previously reported habitat.—Intertidal to 380 m, in mud, sandy mud, sand, sand 461 with gravel and shells, T = 27.5-28; S = 36.5—36.7; OM = 0.59-1.6; DO = 3.9. Occurrence.—In mud, D = 16-75; OM = 0.59-1.66. Distribution in Mexico.—Gulf of Cali- fornia; southern Gulf of Mexico. Distribution.—Central California to Mexico; New England; North Carolina; northern Gulf of Mexico. Glycera tesselata Grube, 1863 Glycera tesselata.—Hartman, 1950:77, pl. 10, fig. 11.—Granados-Barba, 1994:154, pl. 23a.—Hilbig, 1994a:208, fig. 6.5. Material examined.—6 specimens: Sep 1988, sta. 2(2), 3(1), 15(1); Mar 1989, sta. 14(1); Oct-Nov 1990, sta. 16(1). Remarks.—Glycera tesselata resembles closely G. capitata, G. abranchiata, and Glycera sp. F of Gilbert (1984a), but differs from the first by the presence of bilobed, rather than entire postsetal lobes, from the second by the presence of smooth instead of striated proboscidial papillae, and from the last by the presence of only one, rather than two types of proboscideal papillae. These species were compared to material of both named species collected elsewhere. Previously reported habitat.—Continen- tal shelf down to 97 m, in mud with gravel, coarse to fine sand, T = 13-25.8; S = 34.92—37.4; OM = 0.52-6.9; DO = 0.63- 5.4. Occurrence.—In mud, D = 75-148; T = 19-24.5; S = 36.42-36.48; OM = 1.12- 133: Distribution in Mexico.—Baja California Peninsula; Revillagigedo Islands; Sinaloa; Gulf of Tehuantepec, Oaxaca; Tamiahua Lagoon, Veracruz; southern Gulf of Mexi- co; Quintana Roo. Distribution.—Mediterranean Sea; Red Sea; from England to Morocco; India; Ja- pan; Canada; California; Atlantic coast of Panama; North Carolina. 462 Family Goniadidae Kinberg, 1866 The family Goniadidae is represented by nine genera (Hilbig, 1994) and about 75 species. In this study 14 specimens, in two genera and two species, were collected. Genus Goniada Audouin & Milne Edwards, 1833 Goniada cf. maculata Orsted, 1843 Goniada maculata.—Hartman, 1950:20, pl. 1, figs. 7-8.—Pettibone, 1963:225, fig. 58.—Day 1973:51.—Gardiner, 1976:167, figs. 19c—f.—Gilbert, 1984b:33.11, fig. 33.8.—Granados-Barba, 1994:157.—Hil- big, 1994b:226, fig. 7.5. Material examined.—2 specimens: Mar — 1989 sta. 15(1); Mar 1990 sta. 15(1). Remarks.—The specimens examined agree with description of Gilbert (1984b) for this species. However Hartman (1950: 20) recorded 39-41 setigers with unira- mous parapodia (rather then 25). For this reason the identity of the species must be considered doubtful. We think that speci- mens from the Gulf of Mexico could be a new species, but we need to examine type specimens in order to confirm it. In some of the specimens, most of the body was dark brown, but we think this could be the result of fixation. One specimen had pairs of ventrolateral black spots located in the intersegmental furrows from setigers 24—25 to the end of the fragment. Previously reported habitat.—Intertidal to 3020 m, in silt, silty sand, clay, mud, sandy mud, muddy sand, sand with shells, medium to fine sands and sands, T = 16— 28; S = 35.3-37.6; OM = 0.18—0.96; DO = 1.14-3. Occurrence.—In mud, D = 148; OM = 118: Distribution.—Northern Japan; western Europe; Alaska; South Africa; California, New England; North Carolina; northern Gulf of Mexico. Distribution in Mexico.—West of Baja California Sur; Gulf of Tehuantepec; La- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON guna de Tampamachoco, Veracruz; central and southern regions of the Gulf of Mexico. Genus Ophioglycera Verrill, 1885, emended Type species: Ophioglycera gigantea Verrill, 1885. Diagnosis.—Body long, cylindrical; pro- stomium with 9 or 10 annulations and four antennae of equal length; no eyes. No chev- rons on pharynx. Notosetae acicular; neu- rosetae compound spinigers and lyrate setae can be present. Remarks.—This diagnosis includes the presence of lyrate setae first observed in Ophioglycera lyra; such setae had not pre- viously been reported in this genus. Ophioglycera lyra, new species Figs. 2—3 Ophioglycera sp. A.—Gilbert, 1984b: 33.19, figs. 33.16a—j; Granados-Barba, 1994:158. Material examined.—12 specimens: Mar 1988 sta. 1(1), 7(1), 12(3); Sep 1988 sta. 1(1); Mar 1990 sta. 1(1), 7(1), 1101); Mar 1990 sta. 1(1), 7(1), 10(1). Type locality.—Sta. 10, holotype, USNM 175433, Gulf of Mexico, 10, 19°20.9'N, 91°39.6'W, 13 Mar 1990. Paratypes: Sta. 1, 19°15.2'N, 92°08.3’W, 9 Mar 1988, (1) USNM | 175434... Sta. . 7, 18730nie 92°06.2'W, 8 Mar 1988, (1) USNM 175435. Sta. 1, 19°15.2’N, 92°08.3'W, 25 Sep 1988, (1) USNM 175436. Sta. 1, 19°15.2'N, 92°08.3'W, 12 Mar 1989, (1) LACM-AHF POLY 1886. Sta. 7, 18°50.1’N, 92°06.2’W, 11 Mar 1989, (1) LACM-AHF POLY 1887. Sta. 11, 19°20'N, 91°49.8'W, 3 Mar 1989, (1) CPICMLPOP-42-002. Sta. 6, 18°49.2'N, 92°16.2'’W, 9 Mar 1990, (1) CPICMLPOP- 42-001. Sta. 12, 19°33.3’N, 91°54.7'W, 11 Mar 1988, (3) CPICMLPOP-42-003. Gulf of Mexico specimens identified as Ophioglycera sp. A by Gilbert 1984b: USNM 089828, southern Louisiana, Gulf of Mexico, Sta. 0O3P 28°40'02’N, VOLUME 110, NUMBER 3 90°14'43”W. USNM 089829, Texas, Gulf of Mexico, Sta. S-52, 26°10'N, 97°01'W. Material of other species examined.— Goniada distorta Moore 1903, USNM 15720 Honshu Island, Japan, sta. 3739 (moved to Ophioglycera by Hartman, 1950: 36). Ophioglycera gigantea Verrill 1873, USNM_ 13417 syntype, Newport Harbor, sta. 901, surface, 1980. Description.—The description is based on the holotype unless specified otherwise. Holotype a large specimen, 69 mm long and 1.8 mm wide without parapodia, almost complete, with 175 setigers. Paratypes and additional material incomplete with 45—117 setigers, 9-29 mm long X 0.2—0.8 mm wide. Body long, cylindrical, tapering to- wards posterior end, last 10 to 12 setigers very small, crowded. Parapodia and part of dorsum pigmented giving appearance of ophiuroid arm (Fig. 2h). Prostomium sem- iconical with ten annulations (Fig. 2a) (8— 10 in paratypes and additional material) four semiglobular antennae, distal pair dis- tinctly shorter. Basal ring of prostomium laterally expanded as small lappets, no eyes. Pharynx without chevrons (Fig. 2a), very long (6.5 mm long, almost 2 mm wide on holotype and somewhat damaged) covered with papillae. Proboscidial papillae of two forms: triangular with large base and beak slightly recurved (Fig. 2b-—d); and very small and rounded. Macrognaths (not visi- ble in holotype) with 3-5 teeth, microg- naths 20—27 in dorsal arc, 7—8 in ventral arc (Fig. 2e). First 65 setigers uniramous (49-52 in paratypes, this character some- what size related) with three transitional se- tigers (O—3 in paratypes), then biramous. From setiger 65 two midventrally located black dots present close to intersegmental furrows (Fig. 2f), in longitudinal midventral groove; dots and groove absent from last 27—30 segments. Middle anterior region with two dorso- lateral pale bands (darker than the body col- or) subdivided maximally into 10 thin lon- gitudinal bands (Fig. 2g); bands decreasing in number but increasing in width down to 463 three in middle region; thereafter two and finally one much broader, diffuse band pres- ent towards the end of body. Dorsal and ventral cirri with dark pigmentation and en- tire notopodium pigmented on biramous se- tigers. Pigmentation in paratypes similar but less conspicuous, with smaller pig- mented areas and dorsolateral pigmentation not in form of longitudinal bands. First parapodia with pair of dorsolateral cirri and single presetal lobe. Single prese- tal lobe present either in first or in first and second parapodia in paratypes, apparently related to size; first two setigers of speci- mens from northern Gulf of Mexico having that single presetal lobe (Gilbert 1984b). Thereafter, two digitiform slender presetal lobes present, dorsal one longer, at first dis- tinctly so, then subequal. Postsetal lobes shorter than presetal lobes, slender, digiti- form, broader at base, similar on all para- podia. Dorsal cirrus broad basally, pointed distally, more rounded (bulbous) in unira- mous parapodia (Fig. 3a—b), more flattened in biramous parapodia (Fig. 3c—d). Ventral cirri elongate, pointed distally, longer than neuropodial lobes (Figs. 3a—b), becoming broader in transitional setigers, thereafter tapering and pointed distally (Fig. 3c—d). Two acicular notosetae and one acicula of same width in notopodia (Fig. 3c). Neuro- podia with two acicula and compound spi- nigers (Fig. 3e—h) throughout the body; bi- ramous parapodia with additional one to four lyrate setae in upper position (Fig. 31). Remarks.—Ophioglycera lyra, n. sp. dif- fers from related species by having bira- mous parapodia starting on setigers 49-65, two long acicular notosetae, and one to four superior lyrate neurosetae. This species is similar to O. gigantea from which it differs in being smaller, having a much larger num- ber of uniramous setigers, and having a dif- ferent setal composition, particularly the ly- rate setae not previously reported for the genus. Gilbert (1984b) stated that Ophioglycera sp. A and G. distorta Moore, 1903 were ‘‘most similar’’ because they had a similar 464 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.05 mm Ea 1.0 mm f 0.05 mm Fig. 2. Ophioglycera lyra n. sp. a) Anterior end with everted proboscis; b—d) Proboscidial papillae; e) Distal portion of the proboscis, dissected; f) Middle body (biramous parapodia) showing coloration patterns, ventral view; g) Anterior setigers showing ventrolateral bands; h) Anterior setigers showing dorsal coloration patterns. number of uniramous parapodia. However, upon examination of the holotype of G. dis- torta (which is in good condition and an- teriorly dissected) we found that this spe- cies has chevrons, and thus cannot be even maintained in the genus Ophioglycera, which lacks them by definition. Those two Species are therefore not comparable or VOLUME 110, NUMBER 3 465 0.05 mm 0.01 mm 0.05 mm e f g h i Fig. 3. Ophioglycera lyra n. sp. a) Uniramous parapodium from setiger 17, posterior view; b) Uniramous parapodium from setiger 18, anterior view; c) Biramous parapodium from setiger 74, posterior view; d) Biramous parapodium from setiger 75, anterior view; e—h) Compound spinigers from setiger 18; i) Lyrate neuroseta from setiger 74. 466 close. Hartman (1950:36) did not examine type material of Goniada distorta when placing it in the genus Ophioglycera, prob- ably believing that if Moore 1903 did not report chevrons for the species, they did not exist. After examination of the holotype, where the chevrons are clearly seen, we maintain Goniada distorta in the genus Goniada as originally described by Moore. Etymology.—tThe specific name refers to the lyrate setae of the new species, the first to be reported in the genus. Previously reported habitat.—15—98 m, in silt, mud, sand, and silty sand. Occurrence.—In mud, D = 16—56; T = 24-27; S = 35.94—37.19; OM = 0.68—1.65. Distribution in Mexico.—Southern Gulf of Mexico. Distribution.—Northern Gulf of Mexico. Family Hesionidae Grube, 1850 The family Hesionidae is represented by 30 genera and about 150 species (Hilbig 1994c). In this study six specimens in one genus and one species were collected. Genus Podarkeopsis Laubier, 1961 Podarkeopsis levifuscina Perkins, 1984 Podarkeopsis levifuscina Perkins, 1984: \/5, ne. 10: Gyptis vittata.—Taylor, 1971:155.—Day, 1973:25 [not Webster & Benedict, 1887]. Gyptis brevipalpa.—Gardiner, 1976:119, figs. 8q—t, 9a.Uebelacker, 1984:28.27, figs. 28.26a—e.—Granados-Barba, 1994: 134, pl. 19a—h [not Oxydromus brevipal- pa Hartmann-Schréder, 1959]. Material examined.—6 specimens: Mar 1989, sta. 11(1); Mar 1990 sta. 9(2); Oct— Mov 19907sta. lh); 92): Remarks.—The genera of hesionids are currently being reviewed by Pleijel (Fau- chald, pers. comm.) and the position of most species thus remains doubtful. Previously reported habitat.—10—189 m, clay, mud, sandy mud and sand, T = 14.5— 28; S = 35.45-37.22; OM = 0.44-6.9; DO = 3.09-3.17. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Occurrence.—In mud and sandy mud, D = 16-32; T = 28; S = 36.94; OM = 0.59. Distribution in Mexico.—Gulf of Cali- fornia; ZTamiahua Lagoon, Veracruz and Terminos Lagoon, Campeche; southern Gulf of Mexico. Distribution.—North Carolina; Florida; northern Gulf of Mexico. Family Pilargidae Saint-Joseph, 1899 The family Pilargidae is represented by 10 to 13 genera (Blake 1994b) and about 56 species. In this study 25 specimens in three genera and four species were collect- ed. Genus Ancistrosyllis McIntosh, 1879 -Ancistrosyllis commensalis Gardiner, 1976 Ancistrosyllis commensalis Gardiner, 1976: 123, figs. g—k. Ancistrosyllis sp. B.—Wolf, 1984:29.17, fig. 29.12.—Granados-Barba, 1994:137, pl. 20b. Material examined.—1 specimen: Mar 1989 sta. 8(1). Additional material examined.—Ancis- trosyllis commensalis USNM 052902, ho- lotype, Banks Channel, Wrightsville Beach, North Carolina, USA, sta. 11. USNM 052903, paratypes, 3 specimens from Banks Channel Wrightsville Beach, North Caroli- na, USA, sta. 10, in burrows of Notomastus lobatus. USNM 86930, 1 specimen from Texas, USA, sta. S49-6, as Ancistrosyllis sp. B Wolf, 1984. Remarks.—The examination of the ho- lotype of A. commensalis and Wolf’s voucher material of Ancistrosyllis sp. A from Texas and its comparison to our ma- terial shows that all are indeed the same species, the only difference being their size. Specimens from North Carolina are larger than those from the southern Gulf of Mex- ico, which in turn are larger than those from the northern Gulf of Mexico. - Previously reported habitat.—Intertidal to 30 m, commensal with Notomastus lob- VOLUME 110, NUMBER 3 atus, in muddy clay, muddy silt and mud, T = 27; S = 37.04; OM = 0.9. Occurrence.—In mud, D = 16. Distribution in Mexico.—Southern Gulf of Mexico. Distribution.—North Carolina; northern Gulf of Mexico. Genus Cabira Webster, 1879 Cabira incerta Webster, 1879 Cabira incerta.—Pettibone, 1966:178, figs. lla—c, 12a—e.—Wolf, 1984:29.5, figs. 29.2a—f.—Salazar-Vallejo & Orensanz, 1991:272, figs. 2e—f—Granados-Barba, 1994:138. Material examined.—2 specimens: Mar 1989 sta. 7(1); Mar 1990 sta. 2(1). Remarks.—A large number of broken se- tae were present in the specimens collected during this study, especially on anterior segments, making proper identification dif- ficult. Previously reported habitat.—1-—75 m, in mud, muddy sand, silty sand and very fine to medium sand. Occurrence.—In mud, D = 16—70; OM = 1.66. Distribution in Mexico.—Gulf of Te- huantepec, Oaxaca; southern Gulf of Mex- ico. Distribution.—Chesapeake Bay; north- ern Gulf of Mexico. Genus Sigambra O. E Miiller, 1858 Sigambra tentaculata (Treadwell, 1941) Ancistrosyllis tentaculata Treadwell, 1941: 1 figs,.1—3. Sigambra tentaculata.—Pettibone, 1966: 182, figs. 14a—f, 15a—e.—Gardiner, 1976: 121, fig. 9c.—Wolf, 1984:29.8, figs. 29.6a—h.—Granados-Barba, 1994:138, pl. 20a.—Blake, 1994b:285, fig. 10.6. Material examined.—19 specimens: Mar 1988, sta. 6(1), 14(1); Sep 1988, sta. 4(1), 9(1);;Mar 1989; sta. 6(1), 7(1), 9(1), 10(1), 11(1); March 1990, sta. 2(2), 5(3), 9(1), 12(3), 16(1). 467 Remarks.—On all specimens examined in this study, the notopodial hooks start on setiger 4, except for two specimens in which they start on setiger 5. Previously reported habitat.—Intertidal to 5121 m, in sandy silt, mud, sandy mud, muddy sand, sand, and sand with gravel and shells, T = 13-31; S = 34.44-37.72: OM = 0.18-3.72; DO = 0.54—4.32. Occurrence.—In mud and sandy mud, D = 16-127; T = 21-28; S = 35.51-36.99; OM = 0.39-1.58. Distribution in Mexico.—Baja California Peninsula, Gulf of California, Sinaloa, Co- lima, Jalisco, Gulf of Tehuantepec, Tamia- hua Lagoon, Veracruz, and central and southern regions of the Gulf of Mexico. Distribution.—Black Sea, Red Sea, Cal- ifornia, northwestern USA, New England to North Carolina, northern Gulf of Mexico, northeastern South America, and South Af- rica. Sigambra wassi Pettibone, 1966 Sigambra wassi Pettibone, 1966:186, figs. 17a—-f, 18a—e.—Wolf, 1984:29.8, figs. 29.4a—j.—Granados-Barba, 1994:139. Material examined.—1 specimen: Sep 1988, sta. 6(1). Previously reported habitat.—11—37 m, in sandy mud and silty sand. Occurrence.—In mud, D = 16; T = 21-— 28; S = 35.59-36.52; OM = 1.1-1.32. Distribution in Mexico.—Southern Gulf of Mexico. Distribution.—Chesapeake Bay, northern Gulf of Mexico, and Cuba. Family Orbiniidae Hartman, 1942 Some time after the galley proofs for the paper “The polychaetous annelids of the oil platforms area from the southeastern Gulf of Mexico: Orbiniidae and Cossuridae”’ by Granados-Barba & Solis-Weiss (1997) were sent to the press, we had the opportunity to examine material reported as Naineris sp. A by Taylor (1984) deposited in the Smith- 468 sonian Institution and the paratypes of Scol- oplos texana. The results of our examina- tion of this material are presented below. Scoloplos (Scoloplos) texana Maciolek & Holland, 1978 Scoloplos texana Maciolek & Holland, 1978:161, figs. 1-4.—Taylor, 1984:1.31, feo. 1.32. Naineris sp. A.—Taylor, 1984:1.5, figs. 1.2a—f.—Granados-Barba, 1994:29, pl. 3d.—Granados-Barba & Solis-Weiss, Sie Material examined.—8 specimens: Sep 1988, sta. 6(1); Mar 1989, sta. 7(1), 9(1), 11(1); Oct-Nov 1990, sta. 6(1), 7(1), 9(1), 10(1). Additional material examined.—Scolo- plos texana USNM 52733, paratype, Cor- pus Christi Bay, Texas, Gulf of Mexico, USA, sta. 147-1, 22°49'N, 92°08'22’W. USNM 52732, paratypes, 2 specimens from Corpus Christi Bay, Texas, Gulf of Mexico, USA, sta. 122-6, 27°48'38"N, 97°20'17’"W. BLM 2423C, Jul 1976, B-23-1, 2 speci- mens as Naineris sp. A., USNM 090117, 1 specimen from southern Louisiana, Gulf of Mexico, USA, sta. O04P 28°34'09’N, 90°24'32"W, as Naineris sp. A. Material of other species examined.— Scoloplos treadwelli Eisig, 1914 USNM 16067, as Aricia cirrata Treadwell, 1901, Mayaguez Harbor, Blackbuoy, Puerto Rico, sta. (133)6061. USNM 16066, as A. cirrata Treadwell, 1901, Mayaguez Harbor, E of P. Algarrobo, Puerto Rico, sta. (136)6066. Remarks.—Taylor’s (1984) Naineris sp. A, are in fact Scoloplos texana also cited in Taylor (1984). The confusion probably stems from the definition of the prostomium shape for the genus Naineris (rounded or Square). The prostomium of S. texana is not clearly pointed nor triangular as happens with most other species of Scoloplos neither it is rounded or square as is typical of Na- ineris; however, the presence of branchiae in abdominal setigers and the characteristic Shape of the abdominal neuropodia (es- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pecially the setal insertion), so reminiscent of S. texana, left no doubt as to the proper identity of the former Naineris sp. A. Specimens of S. treadwelli and S. texana are similar; they basically differ in that in the first, the neuropodial acicular spines present in anterior setigers are arranged in two rows, whereas in the second there is only a single row. Maciolek & Holland (1978), stated that, characteristically, in S. texana, the branchi- ae are never present in the thorax whereas in S. treadwelli they can start anywhere from last two thoracic to the first two ab- dominal segments. The latter arrangement was not observed in any of the specimens of this study. Previously reported habitat.—15—45 m, in clay, mud, sandy mud and fine sand, T = 28; S = 35.59-36.94; OM = 0.59-1.65. Occurrence.—In mud and sandy mud, D = 15-33; T = 28; S = 35.59-36.94; OM = 0.59-1.65. Distribution in Mexico.—Gulf of Cali- fornia, southern Gulf of Mexico. Distribution.—Northern Gulf of Mexico. Acknowledgments We would like to thank all the partici- pants of the projects IMCA-DINAMO as well as the R/V Justo Sierra’s crew for their help in the field. DGAPA-UNAM (project IN209789) and CONABIO (project PO/52) financed part of this study for which we are grateful. Kristian Fauchald and Linda Ward allowed us to examine the voucher material reported in the volumes edited by Uebe- lacker and Johnson and other material de- posited in the USNM for which they are thanked. In addition Kristian Fauchald kindly revised the last version of the MS for which he is deeply thanked. We also thank Ma. Eugenia Zamudio Resendis and Ignacio Palomar Morales who prepared the illustrations for this paper.° Finally, we would like to thank B. Hilbig and the other anonymous reviewer whose comments def- VOLUME 110, NUMBER 3 initely improved the quality of the manu- script. Literature Cited Blake, J. A. 1994a. Phyllodocidae. Pp 115-186 in J. Blake & B. Hilbig, eds. Taxonomic Atlas of the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 4. The Annelida Part 1. Oligochaeta and Polychaeta: Phyllodocida (Phyllodocidae to Paralacydoni- idae). Santa Barbara Museum of Natural His- tory Publications, 377 pp. . 1994b. Pilargiidae. Pp. 271-294 in J. Blake & B. Hilbig, eds. Taxonomic Atlas of the ben- thic fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 4. The Annelida Part 1. Oligochaeta and Polychaeta: Phyllodo- cida (Phyllodocidae to Paralacydoniidae). Santa Barbara Museum of Natural History Publica- tions, 377 pp. Day, J. H. 1973. New Polychaeta from Beaufort, with a key to all species recorded from North Caro- lina. NOAA Technical Reports NMFS Circ. 375, 139.-pp. Gardiner, S. L. 1976. Errant Polychaete Annelids from North Carolina.—Journal of Elisha Mitchell Science Society 91:77—270. Gathof, J. M. 1984. Phyllodocidae. Pp. 19.1—19.42 in J. M. Uebelacker & P. G. Johnson, eds., Taxo- nomic guide to the Polychaetes of the Northern Gulf of Mexico. Final report to the Minerals Management Service, contract 14-12-001- 29091, Barry A. Vittor & Assoc., Inc., Mobile, Alabama. Gilbert, K. M. 1984a. Glyceridae. Pp. 32.1—32.26 in J. M. Uebelacker & P. G. Johnson, eds., Taxo- nomic guide to the Polychaetes of the Northern Gulf of Mexico. Final report to the Minerals Management Service, contract 14-12-001- 29091, Barry A. Vittor & Assoc., Inc., Mobile, Alabama. . 1984b. Goniadidae. Pp. 33.1—33.19 in J. M. Uebelacker & P. G. Johnson, eds., Taxonomic guide to the Polychaetes of the Northern Gulf of Mexico. Final report to the Minerals Man- agement Service, contract 14-12-001-29091, Barry A. Vittor & Assoc., Inc., Mobile, Ala- bama. Granados-Barba, A. 1994. Estudio sistematico de los anélidos poliquetos de la regién de plataformas petroleras del Sur del Golfo de México.—Tesis Maestria, Facultad de Ciencias, Universidad Nacional Aut6noma de México, 287 pp. , & V. Solis-Weiss. 1994. New records of pol- ychaetous annelids (Order: Eunicida) from the southeastern Gulf of Mexico.—Bulletin of Ma- rine Science 54(2):420—427. 469 , & . 1997. The polychaetous annelids of the oil platforms area from the southeastern Gulf of Mexico: Orbiniidae and Cossuridae.— Bulletin of Marine Science. 60(2), March 1997. Hartman, O. 1950. Goniadidae, Glyceridae and Neph- tyidae.—Allan Hancock Foundation Pacific Ex- peditions 15(1):1—181. Hilbig, B. 1994a. Glyceridae. Pp. 197-214 in J. Blake & B. Hilbig, eds., Taxonomic atlas of the Ben- thic Fauna of the Santa Maria Basin and west- ern Santa Barbara Channel. Vol. 4. The Anne- lida Part 1. Oligochaeta and Polychaeta: Phyl- lodocida (Phyllodocidae to Paralacydoniidae). Santa Barbara Museum of Natural History Pub- lications, 377 pp. . 1994b. Goniadidae. Pp. 215—230 in J. Blake & B. Hilbig, eds., Taxonomic atlas of the Ben- thic Fauna of the Santa Maria Basin and west- ern Santa Barbara Channel. Vol. 4. The Anne- lida Part 1. Oligochaeta and Polychaeta: Phyl- lodocida (Phyllodocidae to Paralacydoniidae). Santa Barbara Museum of Natural History Pub- lications, 377 pp. . 1994c. Hesionidae. Pp. 243-269 in J. Blake & B. Hisbig, eds., Taxonomic atlas of the Ben- thic fauna of the Santa Mariabasin and western Santa Barbara Channel. Vol. 4. The Annelida Part 1. Oligochaeta and Polychaeta: Phyllodo- cida (Phyllodocidae to Paralacydoniidae). Santa Barbara Museum of Natural History Publica- tions, 377 pp. Maciolek, N. J., & J. S. Holland. 1978. Scoloplos tex- ana: a new orbiniid polychaete from South Tex- as, with notes on the related species Scoloplos treadwelli Eisig.—Contribution to Marine Sci- ence 21:161—169. Moore, J. P. 1903. Polychaeta from the coastal slope of Japan and from Kamchatka and Bering Sea.—Proceedings of the Academy of Natural Sciences of Philadelphia 55:401—490, pls. 23- 2s . 1909. The polychaetous annelids dredged by the USS “Albatross” off the coast of southern California in 1904. I. Syllidae, Sphaerodoridae Hesionidae and Phyllodocidae.—Proceedings of the Academy of Natural Sciences of Phila- delphia 61:321-351. Mountford, K. N. 1991. Redescription of Phyllodoce erythrophylla (Schmarda, 1861) and P. madei- rensis Langerhans, 1880 (Polychaeta: Phyllo- docidae), with comments on some related taxa.—Ophelia Supplement 5:157—168. Perkins, T. H. 1984. New species of Phyllodocidae and Hesionidae (Polychaeta), principally from Florida.—Proceedings of the Biological Society of Washington 97:555-582. Pettibone, M. H. 1963. Marine polychaete worms of 470 the New England region.—United States Na- tional Museum Bulletin 227(1):1—356. . 1966. Revision of the Pilargidae (Annelida: Polychaeta) including descriptions of new spe- cies, and redescription of the pelagic Podarmus ploa Chamberlin (Polynoidae).—United States National Museum Bulletin 118(3525):155—207. Pleijel, E 1991. Phylogeny and classification of the Phyllodocidae (Polychaeta).—Zoologica Scrip- ta 20:225-261. . 1993. Polychaeta Phyllodocidae Marine In- vertebrates of Scandinavia number 8. Scandi- navian University Press, Norway, 159 pp. Salazar-Vallejo, S. I., & J. M. Orensanz. 1991. Pilar- gidos (Annelida: Polychaeta) de Uruguay y Ar- gentina.—Cahiers de Biologie Marine 32:267— 219. Solis-Weiss, V., V. Rodriguez-Villanueva, A. Grana- dos-Barba, V. Ochoa-Rivera, L. Miranda-V4z- quez, & P. Hernandez-Alcantara. 1994. Anne- lid polychaete populations of the Order Euni- cida from the southern Gulf of Mexico.—Mé- moires du Muséum. National d_ Histoire Naturelle 162:559—566. , A. Granados-Barba, V. Rodriguez- Villanueva, L. Miranda-Vazquez, V. Ochoa-Rivera, & P. Hernandez-Alcantara. 1995. The Lumbrineri- dae of the continental shelf in the Mexican por- tion of the Gulf of Mexico.—Mitteilungen des Hamburgischen Zoologischen Museum Institut Band 92, Ergbd S, pp. 61-75. Taylor, J. L. 1984. Orbiniidae. Pp. 1.1—1.38 in J. M. Uebelacker & P. G. Johnson, eds., Taxonomic guide to the Polychaetes of the Northern Gulf of Mexico. Final report to the Minerals Man- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON agement Service, contract 14-12-001-29091, Barry A. Vittor & Associates. Inc., Mobile, Al- abama. Treadwell, A. L. 1917. Polychaetous annelids from Florida, Puerto Rico, Bermuda, and the Baha- mas.—Carnegie Institute of Washington, De- partment of Marine Biology Papers 11:255— 268. . 1941. Polychaetous annelids from the New England Region, Porto Rico and Brazil.— American Museum Novitates New York (1138): 4 pp. Uebelacker, J. M. 1984. Hesionidae. Pp. 28.1—28.39 in J. M. Uebelacker & P. G. Johnson, eds., Tax- onomic guide to the Polychaetes of the North- ern Gulf of Mexico. Final report to the Minerals Management Service, contract 14-12-001- 29091, Barry A. Vittor & Assoc., Inc., Mobile, Alabama. , & P. G. Johnson (eds.). 1984. Taxonomic guide to the Polychaetes of the Northern Gulf of Mexico. Final report to the Minerals Man- agement Service, contract 14-12-001-29091, Barry A. Vittor & Assoc., Inc., Mobile, Ala- bama. Webster, H. E., & J. E. Benedict. 1887. The Annelida Chaetopoda from Eastport, Maine.—U.S. Com- mision Fishery of Washington Reports 1885: 707-755. Wolf, P. S. 1984. Pilargidae. Pp. 29.1—29.41 in J. M. Uebelacker & P. G. Johnson, eds., Taxonomic guide to the Polychaetes of the Northern Gulf of Mexico. Final report to the Minerals Man- agement Service, contract 14-12-001-29091, Barry A. Vittor & Assoc., Mobile, Alabama. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):471—475. 1997. Stratiodrilus (Annelida: Polychaeta: Histriobdellidae) associated with a freshwater decapod, with the description of a new species A. Cecilia Z. Amaral and Eloisa Helena Morgado Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, C. Postal 6109, 13083-970 Campinas, SP, Brasil Abstract—A new species of Histriobdellidae (Annelida; Polychaeta) from Brazil, belonging to the genus Stratiodrilus (commensal polychaete) is de- scribed. Stratiodrilus arreliai is distinguished by the characteristics of the body appendages, and mandible apparatus as well as by its biogeographical distri- bution. A brief discussion is made on the association between the genus Stra- tiodrilus and the anomuran genus Aegla. Freshwater decapod crustaceans are known to show a wide range of ectosym- biont invertebrates. These include protozo- ans, platyhelminths, nematodes, rotifers, annelids and small crustaceans. The occur- rence and distribution of these associations as well as nutritional aspects of the physi- ology of some of these symbionts have been investigated (Cannon & Jennings 1987). There are records of species of Stratiod- rilus as symbionts of the following decapod genera: Parastacus, Cherax, Trichodacty- lus, Astacoides and, more frequently, Asta- copsis and Aegla (Haswell 1900, Cordero 1927, Harrison 1928, Mouchet 1932, Vila & Bahamonde 1985, Nonato 1985, Cannon & Jennings 1987). In Brazil, the genus Stra- tiodrilus has been noted as commensal on the brachiuran Trichodactylus petropolitan- us by Nonato (1985) and, in the present work, on the anomuran genus Aegla. Among the several epibionts that are commonly associated with the Aeglidae, are the polychaetes that coexist with tem- nocephalid platyhelminths (Moyano et al. 1993). Several families of the Class Poly- chaeta have commensal species that live in the tubes or cavities of other polychaetes and crustaceans, in corals, and in echino- derms. Among these, the family Histriob- dellidae includes some species with odd characteristics in morphology and life his- tory. These are divided in only two genera, Histriobdella and Stratiodrilus, epizootic polychaetes respectively of marine and freshwater invertebrates. The small poly- chaete Stratiodrilus occurs frequently among the branchial lamellae of crabs and feeds on the microflora that grows in the branchial cavity of the host (Cannon & Jen- nings 1987). In this paper we record the association between Aegla and Stratiodrilus and de- scribe the new species S. arreliai from Southeast Brazil. We also discuss briefly the main interspecific characters of the ge- nus. The Genus Stratiodrilus Stratiodrilus presents no distinction into prostomium and peristomium; head with seven appendages, five of these being ten- tacles (one medium and two lateral pairs). The second pair is bisegmented having sen- sorial (tactile) function (Haswell 1900). The other two appendages (anterior limbs) may aid in locomotion, according to Haswell (1900). The mandible apparatus is black and ex- tremely complex. It is composed of differ- ent pieces arranged in two sets: the upper and lower jaws. The upper jaws consist of 472 a median piece, “‘the fulcrum’’, and two sets of lateral pieces composing the rami. The lower jaws are paired throughout. The body is constricted at regular inter- vals, and may be described as imperfectly divided into six segments (Haswell 1900). The first segment (“‘neck’’) is short and has no cirri. The second, third and fifth seg- ments have each a mammiform elevation that bears the lateral cirri of the body (Cl, C2, C3). The caudal region bears the pos- terior limbs which are not retractile and have one or two pairs of posterior cirri (C4, C5), which may be single or double. These characters are of great taxonomic signifi- cance to the species level. The tentacles and the cirri alike are tipped with fine sensory cilia. The sexes are separate and dimorphic. The male has a pair of retractile ventral claspers in the fourth segment of the body very conspicuous, which are totally absent in the female. In the female the eggs are also easily distinguished as rounded vol- umes of different sizes. Stratiodrilus arreliai, new species Fig. 1 The record of individuals of Stratiodrilus completely confined to the branchial cham- bers of anomurans comes from specimens of Aegla perobae collected at Gruta da Per- oba in the municipality of Sao Pedro, Sao Paulo state (22°55'S; 47°87'W). This ano- muran species was described in 1977 and, to date, has not been recorded for other lo- calities (Bond-Buckup & Buckup 1994). The study of the above mentioned Stra- tiodrilus specimens led us to the conclusion that they belong to a new species, which we called Stratiodrilus arreliai. The new Species show some similarities with the ones already described, but is distinguished mainly by the characteristics of the body appendages, the disposition of the mandible apparatus and by its geographical distribu- tion. Material examined.—A total of 18 spec- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . oak («| Fig. 1. Stratiodrilus arreliai new species (ventral view); anterior limbs (A1); 1st to 5th pair body cirri (C1—5); clasper (CL); mandible (Md); maxilla (Mx); penis (P); posterior limbs (P1); tentacles (T). imens and 5 eggs of Stratiodrilus were found in only one individual of Aegla per- obae (ten and eight individuals in either side of the branchial chamber). Five of these (three males and two females) were stained and slide mounted for microscopical study. The holotype is deposited in the ‘“Museu de Histéria Natural da Universi- dade Estadual de Campinas’” (MHN-BPO- 54). Individuals of the new species are very small, like individuals of the other species of the genus. The total length of males and females, was in average 0.681 mm (7 = 3) and 0.745 mm (n = 2), respectively. Mea- sures for mandible apparatus averaged 0.145 (n = 3) and 0.144 mm (n = 2), re- spectively for males and females. One em- VOLUME 110, NUMBER 3 473 Table 1.—Species of Aegla recorded in the state of Sao Paulo and their sites of occurrence. Species Site A. marginata Bond-Buckup & Buckup 1994 A. castro Schmitt 1942 A. franca Schmitt 1942 A. strinatii Tiirkay 1972 . paulensis (Schmitt 1942) . leptochela Bond-Buckup & Buckup 1994 . perobae Hebling & Rodrigues 1977 . cavernicola Tiirkay 1972 > > Sb DP Db . microphthalma Bond-Buckup & Buckup 1994 Iporanga (cave) Itatinga and Ourinhos (river) Perus and Franca Registro (river), Iporanga (cave) and Eldorado Paulista (river and cave) Perus and Paranapiacaba Sao Pedro (cave) Iporanga (cave) Iporanga (cave) Iporanga (cave) bryo and one immature were also measured (total length 0.137 and 0.370 mm, and man- dible apparatus 0.081 and 0.099 mm, re- spectively). The tentacles of the head are elongated, those of the first anterior pair (T2) being as long as the median one. The second pair (T3) is three times as long as the first, with one distinct division (Fig. 1). The three pairs of lateral cirri (C1, C2, C3) are long and unsegmented. On the posterior border of each limb there is a long cylindrical ten- tacle (cirrus C4). The posterior cirri (C4) emerge from a support that is an extension of the body. The mandible apparatus is very long, reaching the first segment. In the em- bryo, the mandible apparatus is fully de- veloped, being almost the size of the whole egg. Etymology.—The name of the new spe- cies was given in honor of Arrelia (Wal- demar Seyssel), a famous Brazilian clown. This is in accordance with the meaning of the family name Histriobdellidae, which stands for animal similar to a stage-player (clown-like, in a free translation). The Host Anomuran Aegla The genus Aegla has its distribution re- stricted to the temperate and subtropical regions of South America. It is character- ized as being the only anomuran genus in- habiting exclusively freshwater. It occurs in lagoons, streams, swift flowing rivers and cave rivers (Bond-Buckup & Buckup 1994). In the neotropical region its limits of distribution are, to the North, the munici- pality of Franca (Sao Paulo state, Brazil), and, to the South, the province of Ultima Esperanza, in Chile. Of the 34 species at present recorded from Brazilian freshwaters, 29 may be con- sidered endemic. As noted by Bond-Buck- up & Buckup (1994) this fact supports the hypothesis that most species of Aegla have small areas of occurrence. There are records of nine species for the state of Sao Paulo (Table 1). Detailed information on the genus Aegla may be found in Bond-Buckup & Buckup (1994) which present a review of the family Aeglidae as well as the description of twen- ty new species. Discussion To this moment, seven species of Stra- tiodrilus have been described (including one described here), plus one in Chile that was mentioned by Moyano et al. (1993) as new, but remains undescribed (Table 2). The main differences between these spe- cies, regarding the structure and number of the cirri (C1—C5) are presented in Fig. 2. Stratiodrilus arreliai seems to be most closely related to S. aeglaphilus, mainly for being a small sized species and for having long appendages. About the terminology used to characterize the cirri C4 and C5, the term ‘forked’? does not seem adequate, since all the references about the described 474 S. tasmanicus Haswell 1) novaehollandiae Haswell Bisegmente Bisegmented PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pp C4 double non-segmented C5 absent C4 single bisegmented C5 absent (dorsal lobe) jens arreliai n.sp. In haswelli Harrison S. Aeglaphilus Vila & Bahamondi S. platensis Cordeiro S. pugnaxi Vila & Bahamondi Stratiodrilus sp. (in Moyano et al. 1993) Fig. 2. species (including Moyano et al. 1993) in- dicate that they are double. Therefore, the term “‘forked”’ was replaced by ‘“‘double’”’ in this work, as it was mentioned in the discussion by Moyano et al. (1993). After consulting Haswell (1900) and Lang (1949), we concluded that the correct num- ber of segments is six instead of five as mentioned by Vila & Bahamonde (1985). Non-segmented Non-segmented Distal, forked Non-segmented Non-segmented Non-segmented Non-segmented C4 single non-segmented basal support CS absent C4 single bisegmented C5 absent small tubercle C4 single non-segmented C5 absent small tubercle C4 double non-segmented C5 absent C4 double C5 double non-segmented C4 double CS single non-segmented fee As bods oe Main differences among species, concerning the cirri (C1—C5). Besides the structural differences as com- pared to fig. 3 of Moyano et al. (1993), these species may also be distinguished by their host species and geographical distri- bution. The genus Stratiodrilus (Haswell 1900) includes epizootic species exclusive to freshwater, with a high degree of spe- cialization and peculiar geographical distri- butions. This supports the alleged geologi- VOLUME 110, NUMBER 3 475 Table 2.—Species of Stratiodrilus and their hosts, with their geographical distribution. Species Host Geographic distribution S. tasmanicus Haswell 1900 Astacopsis franklinii (Gray) Tasmania S. tasmanicus Haswell 1900 A. franklinii tasmanicus Er- Tasmania ickson S. novaehollandiae Haswell 1913 Astacopsis serratus Shaw Australia S. platensis Cordero 1927 Aegla laevis (Latreille) Uruguay, Argentina S. haswelli Harrison 1928 Astacoides madagascarien- Madagascar sis Milne Edwards S. aeglaphilus Vila & Bahamonde 1985 Aegla laevis (Latreille) Chile S. pugnaxi Vila & Bahamonde 1985 Parastacus pugnax (Poep- Chile pig) S. arreliai new species Aegla perobae Hebling Brazil & Rodrigues cal relationship between Australia, Mada- gascar and South America, as mentioned by Harrison (1928) and Vila & Bahamonde (1985). The geographical distribution of Stra- tiodrilus, with such similar representatives in such distant continents and the coexis- tence on the same host with Temnocephala both in South America and in Australia, is no coincidence, as was mentioned by Cor- dero as early as 1927. The study of these distribution patterns may bring valuable in- sights to the comprehension of the vicari- ance model of biogeography of these groups, as well as to the understanding of the origin of the environments in which they are found. Acknowledgments We are deeply grateful to Dr. E. K Non- ato and Jorge Rodolfo Lima for their incen- tive, valuable suggestions and for providing relevant literature. Literature Cited Bond-Buckup, G., & L. Buckup. 1994. A familia Ae- glidae (Crustacea, Decapoda, Anomura).—Ar- quivos de Zoologia, Museu de Zoologia da Universidade de SAo Paulo, Sao Paulo 32(4): 159-346. Cannon, L. R. G., & J. B. Jennings. 1987. Occurrence and nutritional relationships of four ectosym- biotes of the freshwater crayfishes Cherax dis- par Riek and Cherax punctatus Clark (Crusta- cea: Decapoda) in Queensland.—Australian Journal of Marine and Freshwater Research 38: 419-427. Cordero, E. H. 1927. Un nuevo Arquianélido, Stra- tiodrilus platensis n.sp., que habita sobre Aegla laevis (Latr.). Nota preliminar.—Physis, 7:574— 578. Harrison, L. 1928. On the genus Stratiodrilus (Ar- chiannelida: Histriobdellidae), with a descrip- tion of a new species from Madagascar.—Re- cords Australian Museum 16:116—121. Haswell, W. A. 1900. On a new Histriobdellidae.— Quarterly Journal of Microscopical Science N.S. 43:299—335. . 1913. Notes on Histriobdellidae.—Quarterly Journal of Microscopical Science N.S., 59:91-— 99. Lang, K. 1949. Morphology of Stratiodrilus platensis Cordero (Histriobdellidae).—Arkiv for Zoologi 42A(23):1-30. Mouchet, S. 1932. Notes sur la biologie du Galathéide Aeglea laevis (Latr.). Bulletin de la Societé Zoologique de France 57:316—340. Moyano, H. I., C. Franklin, & S. Gautéia. 1993. Sobre las espécies chilenas de Stratiodrilus Haswell 1900 (Polychaeta, Histriobdellidae).—Boletin de la Sociedad de Bioliologia de Concepcion Chile, 64:147—157. Nonato, E. E 1985. Biologia de Stratiodrilus platensis Cordero 1927 (Annelida, Polychaeta). Pp. 42 in Resumos XII Congresso Brasileiro de Zoologia. Campinas, Brasil, 12. Vila, P. I., & N. Bahamonde. 1985. Two new species of Stratiodrilus, S. aeglaphilus and S. pugnai (Annelida, Histriobdellidae) from Chile.—Pro- ceedings of the Biological Society of Washing- ton, 98:347-350. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):476—482. 1997. A new polyclad flatworm, Tytthosoceros inca (Plathyhelminthes: Polycladida: Cotylea: Pseudocerotidae), from Chilean coastal waters. Juan A. Baeza, David Véliz, Luis M. Pardo, Karin Lohrmann, and Chita Guisado Universidad Catolica del Norte, Sede Coquimbo, Facultad de Ciencias del Mar, Departamento de Biologia Marina, Casilla 117, Coquimbo, Chile Abstract.—A new polyclad species, Tytthosoceros inca is described from Chilean coastal waters. It resembles Pseudoceros luteus (Plehn, 1898) Hyman 1953 incerta sedis but differs in having ear-like pseudotentacles, a brown mar- ginal band, and dark brown spots on the dorsal surface of the body. Tyttho- soceros inca is associated with the colonial ascidians Ciona intestinalis and Pyura chilensis, living on or near them. This is the first record of the genus from Chilean coastal waters and the second report of the family from cold temperate waters, although this time from the southern hemisphere. Pseudocerotidae is one of the most pop- ulous families of polyclad flatworms (Hy- man 1954, Newman & Cannon 1994). Mar- cus (1950) cited an extensive list of over 70 species, mainly Pseudoceros sensu lato, de- scribed from around the world. Hyman (1959a) extended the list to approximately 120, adding many species overlooked by Marcus (1950) and others described by her. Another 60 new species described by Hy- man (1959b), Faubel (1984), Prudhoe (1981, 1989), and Newman & Cannon (1994, 1995, 1996a, 1996b) must now be added to the list of species described world- wide, increasing the total number of known species to at least 200. Most species of the family occur in trop- ical and subtropical waters of the Indo-Pa- cific region (Hyman 1954). They have been reported less frecuently from warmer tem- perate waters around the world, and only one species, Pseudoceros canadensis Hy- man, 1953 from cold temperate waters around British Columbia, Canada (Hyman 1953). However, this species is considered incerta sedis due to its incomplete descrip- tion (Faubel, 1984; Newman & Cannon, 1984). At present, the following 14 pseudocer- otid genera are recognized; Thyzanozoon Grube, 1840, Acanthozoon Collingwood, 1876, Pseudoceros Lang, 1884, Yungia Lang, 1884, Nymphozoon Hyman, 1959, Cryptoceros Faubel, 1984, Cryptobiceros Faubel, 1984, Monobiceros Faubel, 1984, Pseudobiceros Faubel, 1984, Parapseudo- ceros Prudhoe, 1989, Bulaceros Newman & Cannon 1996, Tytthosoceros Newman & Cannon 1996, Maiazoon Newman & Can- non 1996, and Phrikoceros Newman & Cannon 1996. They can be easily disti- guished by details of the male and female reproductive systems (presence or absence of prostate and seminal vesicle, and the number of gonopores and complexes), the alimentary system (presence or absence of anal pores), and by external characters, such as the shape of the body, pseudoten- tacles, pharynx and pharyngeal lobes, and the arrangement of the cerebral and pseu- dotentacular eyes (Newman & Cannon 1994, 1995, 1996a, 1996b). Nevertheless, species recognition and differentiation within each genus presents problems in this group, like in many others that lack solid structures. Marcus (1950), VOLUME 110, NUMBER 3 Hyman (1954, 1959a), Faubel (1984), Prud- hoe (1989), and Newman & Cannon (1994) have commented extensively on the subject. Authors based species identification pri- marily on body colour pattern (Hyman 1954, 1959a, Prudhoe 1989, Newman & Cannon 1994). Aspects of the male copu- latory apparatus, such as the relationship between size of the seminal and prostatic vesicles, cannot be used, due to the extreme similarity of the complex within each genus and considering that some features can change during development or can be de- formed by histological procedures (see Prudhoe 1989, Newman & Cannon 1994). In Chile, the only study on polyclad flat- worms is that of Marcus (1954), based on specimens collected by the Lund’s Univer- sity Chile Expedition. No other research has been conducted and Pseudocerotidae spe- cies have not been reported from the coun- try to date. During exploratory diving in northern Chile we observed a pseudocerotid poly- clad which belongs to the recently de- scribed genus, Tytthosoceros Newman & Cannon, 1996. This is the first record of the genus from Chilean coastal waters (Marcus 1951) and the second report of the family that we have noticed from cold temperate waters, in this case in the southern hemi- sphere. Materials and Methods Tytthosoceros inca specimens were col- lected from lantern nets with Argopecten purpuratus (Lamarck, 1819) (Pectinidae) located along a dock at the Universidad Ca- tdlica del Norte, in Bahia La Herradura, Co- quimbo (29°58’S, 71°22'’W), Chile. Flat- worms were observed on the fouling of the lanterns, typically in association with the ascidian Ciona intestinalis (Linnaeus, 1767). Four specimens were mounted in toto af- ter fixation in Bouin-Hollande, stained with Delafield’s hematoxylin, and cleared with xylol. 477 For histology, specimens were fixed in Bouin-Hollande, embedded in paraffin, se- rial sectioned at seven micrometers, and stained with Harris hematoxylin and eosin. Photographs were taken using a Nikon Bio- phot microscope and drawings of the male and female copulatory apparatus were made. The specimens selected as type material were fixed in Bouin-Hollande, preserved in 70% ethanol, and deposited in the Museo Nacional de Historia Natural, Santiago, Chile (MNHNC) and in the Sala de Siste- matica, Departamento de Ecologia, Ponti- ficia Universidad Catdlica de Chile, Santi- ago (SSUC). Results and Discussion Tytthosoceros inca, new species Figs. 1-3 Material examined.—North Chile: Bahia La Herradura, Coquimbo, 2 m, D. Véliz & L. M. Pardo coll., from scallop lantern nets associated with the ascidian Ciona intestin- alis, 26 Feb 1996, holotype (MNHNC-PL. 11143), 2 paratypes (MNHNC-PL. 11144 and 11145), and 1 specimen (SSUC 6765); Bahia La Herradura, from lantern nets, 1 m, J. A. Baeza coll., Oct 1994, 7 specimens (4 cleared and 3 sectioned); Bahia La Herrad- ura, associated with Pyura chilensis Moli- na, 1782 (Tunicata), 0.5 m, R. Acufia, P. Romero and J. C. Villarroel coll., Apr 1993, 7 specimens. Description.—External morphology: Mature living specimens are extremely soft and thin, reaching a maximum length and width of 11 cm and 6 cm, respectively. The body is ovoid in small animals, becoming elongated-ovoid in worms over 5 cm long. The body margins are broadly folded, cau- dal and anterior margins are rounded. The dorsal surface is smooth. A pronounced middorsal ridge extends along the animal, ending blindly some distance before the posterior margin (Figs. 1, 2A). Marginal pseudotentacles are weakly developed and ear-like, formed as is usual by the upfolding 478 Fig. 1. Tytthosoceros inca, new species. Photo- graph of living organism, dorsal view. Scale bar in centimeters. of the anterior margin (Figs. 2B, C). Each pseudotentacle has a pronounced auricular groove ventrally (Fig. 2C). On living animals, the mouth opens in the mid-ventral line, in middle of the phar- ynx. The pharynx begins on the second ninth of the animal, occupying about one fourth or one fifth of body length. The male gonopore opens beneath the posterior end of the pharynx over a promontory. The fe- male pore is located close behind it, about one half the distance between the mouth and the male apperture. The ventral sucker is conspicuous and circular, lying anterior to the middle of the body. The distance be- tween the sucker and female apperture is about three or four times the distance be- tween the male and female pores (Fig. 2A). Color: The ground color of the dorsal PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON surface on living animals is brownish-yel- low with a narrow marginal white stripe and a broader border-band of pale brown around the body, except on the pseudoten- tacles (Fig. 1). The dorsal surface also shows many dark brown dots with no spe- cial arrangement, decreasing in size to the margins but increasing in number. A narrow black stripe extends along the center of middorsal ridge. This stripe begins at the proximal end of the pharynx, immediately behind cerebral eyespot, ending somewhat before the ridge (Figs. 1, 2B). Only the tip presents the white marginal stripe on the pseudotentacles. Along the inner border of each pseudotentacle, a dark brown stripe exists, narrowing towards the tips and wid- ening at the pseudotentacular bases. Both ‘stripes are not connected with the middorsal one. Another pair of dark brown stripes ex- tends marginally between pseudotentacular bases (Fig. 2B). Ventrally, the color is brownish-yellow with a marginal white stri- pe and pale brown border-band around the body. Eyes: Ocelli of chalice type. Cerebral eyespot present as a single oval cluster in small animals or horseshoe shaped in worms over 6 cm length (Fig. 2B). In all animals over 6 cm length, a pair of pre- cerebral eyes lies deep in the parenchyma some distance before the cerebral cluster. Few pseudotentacular eyes extend along the dorsal surface of pseudotentacles. There are also eyes scattered dorsally, between pseu- dotentacular bases (Fig. 2B). Ventrally, pseudotentacular eyes are more numerous, extending as two irregular clusters along the margins of pseudotentacles, ending as a pair of inverted triangles between the pseu- dotentacular bases (Fig. 2C). Digestive system: Ruffled pharynx deep- ly folded and contained inside the pharyn- geal chamber formed by the anterior por- tion of the central ridge. Behind the phar- ynx chamber, the ridge contains the main intestinal branch that extends to the poste- rior end of the body, giving rise along its course to numerous branches that divide to VOLUME 110, NUMBER 3 a | 1 479 Fig. 2. Tytthosoceros inca, new species. A, ventral view of body; B, anterior portion of body in dorsal view, showing pseudotentacle morphology, eye arrangement and color pattern; C, pseudotentacles, ventral view (MNHNC-PL. 11143). Scale bars in centimeters (A) and micrometers (B and C). Abbreviations: ag = auricular groove, fg = female gonopore, i = main intestine, mg = male gonopore, ms = marginal inter-pseudotentacular stripe, mo = mouth, ph = pharynx, ps = dorsal pseudotentacular stripe, s = sucker, v = vas deferens. form a dense anastomosed network. It seems that the main intestine does not ex- tend further over the pharynx, but some diverticules can be found. Copulatory apparatus: The vas deferens form a prominent network extending for- ward and backward, running along each side of the main intestine. A single male copulatory complex exists (Fig. 3). Two sperm ducts connect spermiductal vesicles with the posterolateral extremes of an oval- elongated seminal vesicle. At its proximal end it narrows to form a sinuous ejaculatory duct that joins the base of a conical penis papilla. At the same location on papilla the ovoid prostate or prostatic vesicle is con- nected by a short prostatic duct. The semi- nal vesicle is about three times the size of the prostate. The penis papilla is armed with a short and wide cuticular stylet both surrounded by a penis sheath. The length: width ratio of the stylet is 1:3.5. The male antrum is narrow and deep. The female gonopore leads into a deep female antrum that opens into a laterally expanded cement pounch. On all sec- tioned specimens, the vagina was never observed connected with any cement gland. From the cement pounch the vagi- na ascends, surrounded by the cement gland. The vagina then curves and re- ceives the uteri (Fig. 3). Geographical’ distribution.—Known from Bahia La Herradura (29°58’S, 480 pv - aS =} i) Gey) S A Hiss s) idan ena Ratti tdi “iu Uy, S TITIAN mg # ote oS PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SV i ] 3%, a ii, A UANGVOVONAV MNT ACGLTDTLCZCIGHIANNNS RTT fg -—————_; 10 Fig. 3. Tytthosoceros inca, new species. Male and female copulatory apparatus, sagital view. Scale bar in micrometers. Abbreviations: cg = cement gland, cp = cement pounch, ed = ejaculatory duct, fa = female antrum, fg = female gonopore, ma = male antrum, mg = male gonopore, mw = muscular wall, pd = prostatic duct, pp = penis papilla, ps = penis sheath, pv = 71°22'’W), Guanaqueros Bay (30°04’S, 71°23'W), and Tongoy Bay (30°14’S, 71°28'’W), north Chile. Taxonomical remarks.—The presence of small ear-like pseudotentacles, the arrange- ment of dorsal eyes, and the short and wide penial stylet allow us to classify the studied type specimens as Tytthosoceros, a genus recently erected by Newman & Cannon (1996a). This genus has a single reproduc- tive system, but on its overall morphology it resembles Pseudobiceros Faubel, 1984 wich possesses paired male systems (New- man & Cannon 1994). Studied specimens can be distinguished from Bulaceros New- man & Cannon 1996, which has two cere- bral eye clusters (not a horseshoe shaped cerebral eyespot), a flat body shape (with- out a pronounced ridge) and a weakly scler- otised, small, and narrow penial stylet. Moreover, the pronounced central ridge of the body, the ear-like tentacles, and the dor- sal eye arrangement of type specimens dis- tinguish 7. inca from Pseudoceros, which prostatic vesicle, s = spermiductal vesicles, sv = seminal vesicle, v = vagina. stylet, sd = sperm ducts, spv = has a flat body shape, simple pseudotenta- cles, and dorsal eyes disposed in two to three scattered lines across the anterior pseudotentacular rim (Newman & Cannon 1994). With regard to the overall body shape and the colour pattern (ground colour of the body and medial black stripe), Tytthosocer- os inca resembles only one species, Pseu- doceros luteus (Plehn, 1898) previously re- ported from Monterrey Bay and Corona del Mar, California coast (Hyman 1953). The latter is considered as incerta sedis, since Newman & Cannon (1994) did not consid- ered it as a member of Pseudoceros sensu stricto when they reviewed the taxonomy of the group. Both species can be distinguished by de- tails of pseudotentacular morphology and colour pattern. While in P. luteus the ten- tacles are broad and flap-like (Marcus 1953), in 7. inca they are weakly developed and ear-like, with respect to body marks, P. luteus lacks the pale brown border-band and VOLUME 110, NUMBER 3 the dark brown spots on the dorsal surface observed on the body of 7. inca. Also, the middorsal stripe in P. luteus forks before the cerebral eyes and continues along the anterior margin of the body to connect with a pair of large oval spots (Plehn 1898, Hy- man 1953), while in 7. inca this stripe ends immediately behind the cerebral eyes and is not associated with any other markings. All features mentioned above have been considered as important diagnostic charac- ters by Marcus (1950), Hyman (1954), Prudhoe (1984), and Newman & Cannon (1994, 1995, 1996a, b), allowing us to dif- ferentiate Tytthosoceros inca from Pseu- doceros luteus and to describe the former as a new species. Hyman (1953) also reports a mottled gray color variant of Pseudoceros luteus. In our opinion, it is probably another species due to colour pattern differences with P. lu- teus. It is also appropriate to compare the new species with this color variant. P. lu- teus and the color variant of P. luteus pres- ent the same morphological differences with respect to T. inca. Concerning body marks, the color variant can be distin- guished from 7. inca because it lacks the middorsal black stripe and the pale brown border-band found in the new species. The main diagnostic characters that differ between this new species and other Tyttho- soceros are; the pronounced male antrum, the extremely short prostatic duct and the long ejaculatory duct, the long and narrow stylet (largest in T. inca, length:width ratio 1:1.35, compared to 1:2 for T. lizardensis Newman & Cannon 1996 and T. nocturnus Newman & Cannon 1996), and the pres- ence of a pair of pre-cerebral eyes lying deep in the parenchyma in all animals over 6 cm length (not present in others conge- ners). Due to these differences, it is possible that the present specimens warrant a new genus (Newman, pers. comm.). However, due to the lack of complete morphological and anatomical information on previously described pseudocerotids, we considered our specimens as members of Tytthosocer- 481 os. We believe that more information is needed on several species in order to erect a new genus. Etymology.—The name inca refers to the Incan civilization that occupied the northern and north-central area of Chile during the Pre-Hispanic age. Ecological remarks.—This species has also been observed in Bahia La Herradura and Tongoy in shallow waters, living close to or on the colonial ascidian Pyura chilen- sis. It probably feeds directly on the ascid- ians or may prey on the bivalves when in- habiting lantern nets. The former feeding habit has been previously described for oth- er species of the family (Hyman 1951, Newman & Cannon 1994). Tytthosoceros inca also can be observed in dense aggre- gations on the bottom or swimming towards the surface with undulating movements. This is the first record of the genus Tyr- thosoceros from coastal Chilean waters. Since until now only one species of Pseu- docerotidae has been cited for cold temper- ate waters, this appears to be the second record for the family, although this time from the southern hemisphere. Acknowledgment The authors thank Dr. Wolfgang Stotz for providing and translating papers in German, and Edgardo Zamorano and Dr. Raymond Bienert for correcting the English version of the manuscript. We also acknowledge Carlos Gallardo and especially Dr. Leslie Newman (Queensland Museum, Australia), whose comments made substantial contri- butions to the present manuscript. Literature Cited Faubel, A. 1984. The Polycladida, Turbellaria. Pro- posal, and establishment of a new system. Part II. The Cotylea.—Mitteilungen aus dem ham- burgischen Zoologischen Museum und Institut, Hamburg 81:189-—259. Hyman, L. H. 1951. The Invertebrates II. Platyhel- minthes and Rhynchocoela. McGraw-Hill Book Company, New York, 550 pp. . 1953. The polyclad flatworms of the Pacific 482 coast of North America.—Bulletin of the Amer- ican Museum of Natural History, New York 100:265-—392. . 1954. The Polyclad genus Pseudoceros, with special reference to the Indo-Pacific region.— Pacific Science 8:219—225. 1959a. A further study of micronesian po- lyclad flatworms.—Proceedings of the United States National Museum 108:543-—597. 1959b. Some Australian polyclads (Turbel- laria)—Records of The Australian Museum, Sydney 25:1-17. Marcus, E. 1950. Turbellaria Brasileiros (8).—Boletin da Faculdade de Filosofia Ciencias e Letras Universidada de Sao Paulo, Zoologia 15:5—192. . 1954. Reports of the Lund University Chile Expedition 1948-1949. 11.—Acta Universita- tits Lundensis 49(13):1—115. Newman, L. J., & L. R. G. Cannon. 1994. Pseudo- ceros and Pseudobiceros (Platyhelminthes, Po- lycladida, Pseudocerotydae) from eastern Aus- tralia and Papua New Guinea.—Memortrs of the Queensland Museum 37(1):205—266. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1995. Colour pattern variation in the tropical flatworm, Pseudoceros (Platyhelminthes, Poly- cladida), with descriptions of three new spe- cies.—The Raffles Bulletin of Zoology 43(2): 435-446. 1996a. Bulaceros, new genus, and Tyttho- soceros, new genus, (Platyhelminthes: Poly- cladida) from the great barrier reef, Australia and Papua New Guinea.—The Raffles Bulletin of Zoology 44(2):479—4972. 1996b. New genera of pseudocerotid flat- worms (Platyhelminthes, Polycladida) from Australian And Papua New Guinean coral reefs.—Journal of Natural History 30:1425— 1441. Plehn, M. 1898. Drei neuen Polycladiden.—Jenaische Zeitschrift fiir Naturwissenschaft, Jena 31:90— oo Prudhoe, S. 1981. Polyclad turbellarians from the southern coasts of Australia—Records of the South Australian Museum 18:361—384. 1989. Polyclad turbellarians recorded from African waters.—Bulletin of the British Muse- um (Natural History), Zoology 55:47—96. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):483—486. 1997. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE Applications published in the Bulletin of Zoological Nomenclature The following Applications were published on 26 March 1997 in Vol. 54, Part 1 of the Bulletin of Zoological Nomenclature. Comment or advice on any of 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, U.K. Case No. 2906 Anomalina d’Orbigny, 1826 (Foraminiferida): proposed designation of A. ariminensis d’Orbigny in Fornasini, 1902 as the type species. 2940 Riisea and riisei Duchassaing & Michelotti, 1860 (Cnidaria, Anthozoa): proposed conservation as the correct original spellings of generic and specific names based on the surname Riise. Umbellula Cuvier, [1797] (Cnidaria, Anthozoa): proposed conservation as the correct original spelling, and corrections to the entries relating to Umbellularia Lamarck, 1801 on the Official Lists and Indexes of Names in Zoology. Galba Schrank, 1803 (Mollusca, Gastropoda): proposed designation of Buc- cinum truncatulum Miller, 1774 as the type species. Roeslerstammia Zeller, 1839 and Acrolepiopsis Gaedike, 1970 (Insecta, Lepidoptera): proposed conservation by the designation of Alucita erxlebella Fabricius, 1787 as the type species of Roeslerstammia; and A. erxlebella and Tinea imella Hiibner, [1813] (currently Roeslerstamnia erxlebella and Monopis imella): proposed con- servation of the specific names by the designation of a neotype for A. erxlebella. Euchroeus Latreille, 1809 and Chrysis purpurata Fabricius, 1787 (currently E. purpuratus) (Insecta, Hymenoptera): proposed conservation of usage; and Chrysis gloriosa Fabricius, 1793: proposed suppression of the specific name. Diemenia atra Macleay, 1884 (currently Demansia atra; Reptilia, Serpentes): proposed conservation of the specific name. Trigonocephalus pulcher Peters, 1863 (currently Bothrops pulcher) and Bo- throps albocarinatus Shreve, 1934 (currently Bothriechis oligolepis albocarinatus) (Reptilia, Serpentes): proposed conservation of the specific and subspecific names by the designation of a neotype for T. pulcher. 484 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Applications published in the Bulletin of Zoological Nomenclature The following Applications were published on 30 June 1997 in Vol. 54, Part 2 of the Bulletin of Zoological Nomenclature. Comment or advice on any of these ap- plications is invited for publication in the Bulletin and should be sent to the Exec- utive Secretary (I.C.Z.N.), % The Natural History Museum, Cromwell Road, Lon- don SW7 5BD, U.K. Case No. 2943 Aporcelaimus Thorne & Swanger, 1936 (Nematoda): proposed designation of Dorylaimus superbus de Man, 1880 as the type species. 2996 Pila Réding, 1798 and Pomacea Perry, 1810 (Mollusca, Gastropoda): pro- posed placement on the Official List, and AMPULLARIIDAE Gray, 1824: proposed confirmation as the nomenclaturally valid synonym of PILIDAE Preston, 1915. Disparalona Fryer, 1968 (Crustacea, Branchiopoda): proposed conservation. Dasineura Rondani, 1840 (Insecta, Diptera): proposed designation of Tipula sisymbrii Schrank, 1803 as the type species. Varanus teriae Sprackland, 1991 (Reptilia, Squamata): proposed conserva- tion of the specific name. Hydrosaurus gouldii Gray, 1838 (currently Varanus gouldii) and Varanus panoptes Storr, 1980 (Reptilia, Squamata): proposed conservation of the specific names by the designation of a neotype for H. gouldii. VOLUME 110, NUMBER 3 485 Opinions published in the Bulletin of Zoological Nomenclature The following Opinions were published on 26 March 1997 in Vol. 54, Part 1 of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be obtained free of charge from the Executive Secretary, I.C.Z.N., % The Natural History Mu- seum, Cromwell Road, London SW7 5BD, U.K. Opinion No. 1860. Acanthoteuthis Wagner in Miinster, 1839 and Muensterella Schevill, 1950 (Mollusca, Cephalopoda): placed on the Official List. 1861. Octopus vulgaris Cuvier, [1797] and Loligo vulgaris Lamarck, 1798 (Mol- lusca, Cephalopoda): specific names conserved. 1862. Aspidiphorus Ziegler in Dejean, 1821 (Insecta, Coleoptera): conserved as the correct original spelling, and SPHINDIDAE Jacquelin du Val, [1861]: given precedence over ASPIDIPHORIDAE Kiesenwetter, 1877 (1859). Sphaerocera Latreille, 1804 and Borophaga Enderlein, 1924 (Insecta, Dip- tera): conserved: Musca subsultans Linnaeus, 1767: specific name placed on the Official List. Chaetodacus latifrons Hendel, 1915 (currently Bactrocera latifrons; Insecta, Diptera): given precedence over Dacus parvulus Hendel, 1912. Eudistoma Caullery, 1909 (Tunicata): given precedence over Paessleria Mi- chaelsen, 1907. Hydromantes Gistel, 1848 (Amphibia, Caudata): Spelerpes platycephalus Camp, 1916 designated as the type species. Phyllophis carinata Ginther, 1864 (currently Elaphe carinata; Reptilia, Ser- pentes): specific name conserved. 486 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Opinions published in the Bulletin of Zoological Nomenclature The following Opinions were published on 30 June 1997 in Vol. 54, Part 2 of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be obtained free of charge from the Executive Secretary, I.C.Z.N., % The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. Opinion No. 1868. Patella longicosta Lamarck, 1819 (Mollusca, Gastropoda): specific name conserved. 1869. Monstrilla Dana, 1849 and Thaumaleus Kr@yer, 1849 (Crustacea, Copepo- da): conserved. 1870. Sicus Scopoli, 1763 and Myopa Fabricius, 1775 (Insecta, Diptera): conserved by the designation of Conops ferrugineus Linnaeus, 1761 and C. buccatus Linnaeus, 1758 as the respective type species; and Coen- omyia Latreille, 1796 placed on the Official List. Iodotropheus sprengerae Oliver & Loiselle, 1972 (Osteichthyes, Percifor- mes): holotype replaced by a neotype. Siboma atraria Girard, 1856 (currently Gila atraria; Osteichthyes, Cyprin- iformes): specific name conserved. HEMIDACTYLIINI Hallowell, 1856 (Amphibia, Caudata): conserved. Aptornis Owen, 1848 (Aves): conserved as the correct original spelling. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(3):487. 1997. REVIEWERS The following people reviewed manuscripts for the Proceedings in 1996. P. Alderslade, M. V. Angel, R. Bamber, K. Banse, E M. Bayer, D. Berner, J. A. Blake, J. P. Blanco, G.-O. Brandorff, N. L. Bruce, S. D. Cairns, E. Campos, L. Cannon, V. Cappola, M. D. Carleton, A. Carvacho, F A. Chace, Jr., T. Y. Chan, R. Chapman, J. E. Cooper, R. Cowie, K. Crandall, M. M. Criales, R. E. Crombie, N. Cumberlidge, P. J. E Davie, R. Davies, M. Dojiri, S. Donovan, L. Durdan, P. C. Dworschak, A. Farjon, R. Farris, K. Fauchald, D. L. Felder, R. M. Feldmann, G. E. Fenton, F. D. Ferrari, F Fiers, O. S. Flint, Jr, C. H. J. M. Fransen, S. Gardiner, A. L. Gardner, S. R. Gelder, C. J. Glasby, C. Goulden, M. J. Grygier, D. Guinot, G. Hartman, P. Hastings, J. Hedgpeth, M. E. Hendrickx, W. R. Heyer, B. Hilbig, S. Hiruta, H. H. Hobbs, III, D. Holdich, P. Holroyd, N. Hotton, A. G. Humes, D. A. Jones, B. Kensley, I-H. Kim, W. Kobusch, F Krapp, R. K. Kropp, J. D. Kudenov, D. B. Lellinger, R. Lemaitre, A. Mackie, C. Magalhaes, R. B. Manning, R. Mariscal, J. C. Markham, W. N. Mathis, R. W. Mc- Diarmid, P. A. McLaughlin, J. McLelland, R. Mooi, D. Moore, R. Moser, G. Moura, T. A. Munroe, E. Murdy, P. Naiyanetr, K. Nakamura, R. Nelson, L. Newman, P. K. L. Ng, J. Norenburg, S. Ohtsuka, J. Olesen, J. Pakaluk, © Park, J. £. Patton, T. Perkins, M. H. Pettibone, W. W. Price, C. E. Ray, N. A. Rayner, Y. R. Reddy, R. P. Reynolds, C. B. Robbins, R. Rosenblatt, D. Russell, N. V. Schizas, G. Shinn, J. Smith, E. Southward, P. J. Spangler, D. H. Staples, J. Stock, K. C. Stuck, M. Tavares, C. Taylor, E Thompson, A. Tsukagoshi, S. Tyler, J. C. von Vaupel Klein, B. Werding, M. K. Wicksten, A. B. Williams, J. T. Williams, D. E. Wilson, R. Winter- bottom, T. Woolridge, K. Wouters, A. H. Wynn, D. A. Yun. “@ = ? - =) ia . a4 S| . & 2 < Pe ; Ce rT \y M6 : Nitta ‘ oe ee ee ee eee oe me “ole Pang gia . ret ee i et mee a nd bal Casal up eee . i ‘ j —r is ° a , in ee fi Oleg ee ah a ) 4 a eere 5D 4 » 7 ye ‘ ie J \ } oc Vag ' ® by 4 ‘-¢ ict € » Loe #45 } g ct he ; _ Ce May tae “ita > «eae ~ ' 1 - — F . es 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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Ronald Heyer Two new species of salamanders (Caudata: Plethodontidae) of the genera Bolitoglossa and Nototriton from Parque Nacional La Muralla, Honduras James R. McCranie and Larry David Wilson Protoblepharon rosenblatti, a new genus and species of flashlight fish (Beryciformes: Anom- alopidae) from the tropical South Pacific, with comments on anomalopid phylogeny Carole C. Baldwin, G. David Johnson, and John R. Paxton Status of Platycephalus cantori Bleeker, 1879 (Teleostei: Platycephalidae) Leslie W. Knapp and Hisashi Imamura Pseudothelphusa ayutlaensis, a new species of freshwater crab (Crustacea: Brachyura: Pseu- dothelphusidae) from Mexico Fernando Alvarez and José Luis Villalobos Austinixa, a new genus of pinnotherid crab (Crustacea: Decapoda: Brachyura), with the de- scription of A. hardyi, a new species from Tobago, West Indies Richard W. Heard and Raymond B. Manning A new species of Rimicaris (Crustacea: Decapoda: Bresiliidae) from the Snake Pit hydrother- mal vent field on the Mid-Atlantic Ridge Joel W. Martin, James Signorovitch, and Hema Patel Occurrence of three species of mud shrimps in aquiculture ponds on Caribbean coasts of Venezuela and Colombia, with a redescription of Upogebia omissago Williams, 1993 (De- capoda: Upogebiidae) Austin B. Williams Esanpotamon namsom, a new genus and species of potamid crab (Crustacea: Decapoda: Brachyura) from a waterfall in northeastern Thailand Phaibul Naiyanetr and Peter K. L. Ng Tridentella ornata (Richardson 1911), new combination: records of hosts and localities (Crus- tacea: Isopoda: Tridentellidae) Brian Kensley and Richard W. Heard Doxomysis acanthina, a new leptomysinid (Crustacea: Mysidacea) from the northern Great Barrier Reef, Australia, with extensions to the known distributions of D. australiensis W. M. Tattersall, 1940 and D. spinata Murano, 1990, and a key to the genus Doxomysis M. S. Talbot Acanthomysis bowmani, a new species, and A. aspera li, Mysidacea newly reported from the Sacramento-San Joaquin Estuary, California (Crustacea: Mysidae) Richard F. Modlin and James J. Orsi Lamellibrachia satsuma, a new species of vestimentiferan worms (Annelida: Pogonophora) from a shallow hydrothermal vent in Kagoshima Bay, Japan Tomoyuki Miura, Junzo Tsukahara, and Jun Hashimoto The polychaetous annelids from oil platforms areas in the southeastern Gulf of Mexico: Phyl- lodocidae, Glyceridae, Goniadidae, Hesionidae, and Pilargidae, with description of Ophio- glycera lyra, a new species, and comments on Goniada distorta Moore and Scoloplos texana Maciolek & Holland Alejandro Granados-Barba and Vivianne Solis-Weiss Stratiodrilus (Annelida: Polychaeta: Histriobdellidae) associated with a freshwater decapod, with the description of a new species A. Cecilia Z. Amaral and Eloisa Helena Morgado A new polyclad flatworm, Tytthosoceros inca (Platyhelminthes: Polycladida: Cotylea: Pseu- docerotidae) from Chilean coastal waters Juan A. Baeza, David Véliz, Luis M. Pardo, Karin Lohrmann, and Chita Guisado International Commission on Zoological Nomenclature Reviewers, 1996 329 332 338 366 393 384 388 393 399 412 417 422 426 439 447 457 47] 476 483 487 THE BIOLOGICAL SOCIETY OF WASHINGTON 1996-1997 Officers President: Stephen D. Cairns Secretary: Carole C. Baldwin President-elect; Richard P. Vari Treasurer: T. Chad Walter Elected Council John A. Fornshell Rafael Lemaitre Alfred L. Gardner Diana Lipscomb Susan L. Jewett James N. Norris Custodian of Publications: Storrs L. Olson PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: Frederick M. Bayer — Invertebrates: Jon L. Norenburg Frank D. Ferrari Plants: David B. Lellinger Rafael Lemaitre Insects: Wayne N. Mathis Vertebrates: Gary R. 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Payment for membership is accepted in US dollars (cash or postal money order), checks on US banks, or MASTERCARD or VISA credit cards. | Manuscripts, corrected proofs, editorial questions should be sent to: EDITOR BIOLOGICAL SOCIETY OF WASHINGTON NATIONAL MUSEUM OF NATURAL HISTORY WASHINGTON, D.C. 20560, U.S.A. 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 Periodicals postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, P.O. Box 1897, Lawrence, Kansas 66044. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):489-501. 1997. Pachyrotula, a new genus of freshwater sponges from New Caledonia (Porifera: Spongillidae) Cecilia Volkmer-Ribeiro and Klaus Riitzler (CV-R) Museu de Ciencias Naturais, Fundacéo Zoobotanica do Rio Grande do Sul, Rua Dr. Salvador Franga, 1427, 90.690-000, Porto Alegre, RS, Brasil; (KR) Department of Invertebrate Zoology, National Museum of Natural History Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Characteristics found during a review of the type material of Spongilla (Stratospongilla) raceki Riitzler from New Caledonia point to a new, monotypic genus, Pachyrotula. This new genus is closely related to Hetero- rotula Penney & Racek and to Houssayella Bonetto & Ezcurra De Drago. Heterorotula also occurs in New Caledonia and is represented by the species H. caledonensis new species and H. multidentata (Weltner). These species, along with Oncosclera diahoti (Riitzler), new combination, are indicators of a rare and particular freshwater sponge fauna of this island. For the past three decades, the freshwater sponges of New Caledonia have been known only from the specimens described by Riitzler (1968). It recently became clear that this material needed to be reexamined, especially in the light of Penney & Racek’s (1968) comprehensive study of all gem- mule-producing genera of freshwater sponges and Racek’s (1969) extensive work on Australia’s freshwater sponges, which provided new insight into the generic and specific relationships of these faunas and revealed their heretofore unsuspected rich- ness. That work and several subsequent studies on the freshwater sponges of South America indicated the need for a taxonomic and systematical updating of the New Cal- edonia description. As a result of this up- dating, a new genus is being proposed, with Spongilla (Stratospongilla) raceki Riitzler as the type species; also, Ephydatia multi- dentata (Weltner) forma caledonensis Riutz- ler is elevated to a full species in the genus Heterorotula Penney & Racek, and Spon- gilla (Stratospongilla) diahoti Riitzler is transferred to the genus Oncosclera Volk- mer-Ribeiro. Material and Methods The material examined consisted of types and paratypes of Spongilla (Stratospongil- la) diahoti, S. (Stratospongilla) raceki, and Ephydatia multidentata f. caledonensis (Riitzler 1968), all deposited at the National Museum of Natural History, Washington D.C. (USNM); slides of Houssayella igua- zuensis Bonetto & Ezcurra de Drago, 1966, provided by I. Ezcurra de Drago; and slides of species of the genus Heterorotula pro- vided by A. A. Racek; the latter deposited at Museu de Ciencias Naturais (MCN) of Fundacgao Zoobotanica do Rio Grande do Sul, Porto Alegre, Brazil. A minute fragment of Spongilla (S.) ra- ceki with gemmules was dissociated after boiling in nitric acid and washed several times in distilled water. When completely dry, the suspended clean spicules were dropped on a stub and coated with gold in preparation for scanning electron micro- scopic (SEM) observations. Some dry gem- mules were hand-sectioned under a stereo- microscope and their halves glued to a stub and also gold-coated for SEM examination. Photomicrographs were obtained with an 490 MCN JEOL-5200 microscope equipped with a Pentax SF7 35 mm camera. Systematics Pachyrotula, new genus Type species.—Spongilla (Stratospongil- la) raceki Riitzler, 1968. Genus monotypic. Diagnosis.—Spongillidae with stout, heavily spined birotulate gemmoscleres with knobby rotules progressively reduced from the inside to the outside of the gem- mular coat. Description.—Sponges forming slender, greyish, small, irregular, soft crusts with gemmules loosely distributed near the basal spongin plate. Oscula and pores inconspic- uous. The skeleton consists of a dense po- lygonal network of spicules bound together by scanty spongin. No main fibers are dis- cernible. Three classes of megascleres. The most abundant are the alpha megascleres, which are finely spined oxea, abruptly and acutely pointed, and with a denser spine arrange- ment at the extremities. The beta mega- scleres measure half to two-thirds the length of the alpha megascleres, are curved or even bent oxea or styles, are strongly spined with abruptly and sharply pointed extremities that are enhanced by a distal swelling and a concentration of minute spines turned toward the point. These mega- scleres are tangentially packed around the gemmules under a loose and haphazard ar- rangement of the alpha megascleres. The third class consists of very rare, smooth, thin, long oxea with differing mucronated extremities. Microscleres absent. Gemmoscleres are dumb-bell-shaped bir- otules, with knobby rotules ranging from ir- regularly sculptured, tuberculate, or spiny bulbs to merely knobbed expansions of the shaft (Figs. 1, 2). As the rotules become smaller, the shafts become longer. Sharp- ened, lanceolated extremities of the shaft may protrude from one or both of the ro- tules or terminal knobs, thus morphologi- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cally grading into beta megascleres. Shafts are spined, some are displaying a few tub- ercules capped by rosettes of spines. Gemmules abundant, located at the basal part of the sponge. Inner layer thick; pneu- matic layer thin, with small, irregular air spaces and only a few short gemmoscleres radially embedded in it. The longer gem- moscleres form a loose radial arrangement and, together with the derived acanthoxea (beta megascleres), turn more tangential to- ward the gemmule periphery. The outer ro- tules and the largest part of the shafts of most gemmoscleres protrude at the gem- mular surface. One single foramen in each gemmule. Pachyrotula raceki (Ritzler), new combination Figs. 1-5, 9 Spongilla (Stratospongilla) raceki.—Ritz- ler, 1968:60, figs. 5—9. Holotype.—USNM 23882, New Caledo- nia; Ferdinand Starmiihler leg. 18 Sep 1965. Paratype.—USNM 23883, same location as for holotype. Type locality.—River Le Diahot, New Caledonia (Fig. 9). Description.—Soft, grayish, tiny patches 1 mm thick on the lower surface or in con- cavities of stones. Sponge surface smooth, porous. Oscula inconspicuous. Skeleton a confused arrangement of megascleres with scanty spongin at the crossing points. A tangential packing of the megascleres around the gemmules produces a thickening of the skeleton at the basal portion of the sponge. Alpha megascleres are microspined, straight to more often slightly curved oxea with abruptly pointed, spined extremities. Oxea vary noticeably in length. Spines at tips turned outward. These megascleres form the skeleton mesh and also occur in a loose and confused manner around each gemmule. Beta megascleres are short, slightly to VOLUME 110, NUMBER 4 49] Fig. 1. Gemmoscleres of Pachyrotula raceki (Riitzler) (SEM). a, Immature or poorly developed spicules; b, Average development; c, Largest class of gemmoscleres exhibiting differentiated rotular reduction; d, Further differentiation; e, Same as d, reduced scale. Scales = 10 wm. 492 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig: (Riitzler) gemmosclere in Fig. 1b (lower right); note different size of the rotules (SEM). Scale = 5 pm. Enlarged rotules of Pachyrotula raceki strongly curved or even bent, strongly spined oxea or styles with very abruptly pointed extremities. Usually marked by a subterminal swelling or bulb with a heavier covering of spines turned to the spicule ex- tremities. Some beta megascleres are em- bedded in the gemmular wall together with the gemmoscleres, but for the most part are tangentially packed around the gemmules. Long, smooth thin megascleres are very rare but certainly belong to the sponge, as became evident once the series grading from very thin and short ones was per- ceived in the original preparations. These spicules may be up to four times longer and thinner than the alpha megascleres. How- ever, they are too scarce to determine their locale in the skeleton. Gemmoscleres stout, strongly spined bir- otulates of conspicuously large range of lengths. With thick shafts and, terminally, with spool- to knob-like rotules irregularly ornamented by spined lobes, rosettes, or tub- ercules. The shorter gemmoscleres display larger, more conspicuous rotules, the outer one usually smaller than the inner one. The bulbous endings of the longer gemmoscleres gradually taper, thus changing the spicules into straight or gradually curved oxeas with piercing points enhanced by a slight subter- minal swelling of the shaft. Such spicules grade into what are here classed as beta me- gascleres because they are not embedded in the gemmular wall but are packed around the gemmules (Figs. 1, 2, 3b; and Riitzler 1968, figs. 6-8). Gemmules abundant. They occur from the base to the middle portion of the thin crusts. Because the sponge was in the pro- cess of producing gemmules, it is possible to describe the manner in which the gem- mular wall is formed. Wherever gemmules are about to be formed, and even prior to the congregation of the thesocytes, a gath- ering of cells secreting the alpha mega- scleres takes place, joined shortly thereafter by spicule-forming beta megasclerocytes and gemmosclerocytes. By the time the in- ner gemmular wall is formed, a large num- ber of the short and longer gemmoscleres are already present and beginning to be- come inserted at the base of the inner gem- mular wall. During the formation of the thin pneumatic coat around the gemmosclere layer, more and more gemmoscleres con- tribute to this coat. At the same time, the beta and alpha megascleres congregate tightly around the forming gemmule, until only a round mass of alpha megascleres can be perceived. The presence of some gem- moscleres (particularly shorter ones) radi- ally embedded in the pneumatic coat (Fig. 3) indicates that a radial orientation was abandoned for a tangential one. The outer gemmular wall is thin; the foraminal tube is short and does not reach beyond the lon- ger birotulates. For spicule and gemmule measurements, refer to the original descriptions (Riitzler 1968). | Habitat.—The sponge encrusts the lower surface of stones in running and standing waters with extreme low conductivity (28— 56) and slightly acidic to neutral pH (6.6— i). Remarks.—Penney & Racek (1968) ele- vated Annandale’s subgenus Stratospongil- la to a genus, defining its gemmoscleres (p. 40) as ‘“‘more or less strongly bent amphi- strongyles .. . or slightly curved spined am- phioxea, or a combination of both’’ and its VOLUME 110, NUMBER 4 ¥ ’ id ~ Fig. 3. Gemmular wall in Pachyrotula raceki (Rtitzler) (SEM). a, A cross-section exposes the irreg- ularly smooth surface of the inner coat and the irreg- ular air spaces of the pneumatic coat, with some short birotulates radially embedded and some longer ones tangentially embedded, both kinds projecting beyond the thin outer gemmular coat; b, Gemmular surface, with rotule of one short gemmosclere projecting and several beta megascleres tangentially embedded in the growing pneumatic coat. Scales = 10 wm. microscleres as “shorter and slender am- phioxea.”’ They suggested, however, that Stratospongilla species lacking microscleres might be grouped under a new genus. Volk- mer-Ribeiro (1970) followed up on this suggestion by defining the genus Oncos- clera, with Oncosclera jewelli (Volkmer 1963) as the type species. Oncosclera spe- cies not only lack microscleres but differ markedly from Stratospongilla in their gemmular structure. At the time the genus Oncosclera was defined, it was assigned several South American species previously described as Stratospongilla by Bonetto & Ezcurra de Drago (1966). Also included in 493 Fig. 4. (Riitzler) (SEM). a, Detail of the loose arrangement of the alpha megascleres cemented by scanty spongin; b, Cross section of the sponge crust showing the loose skeletal arrangement of the megascleres. Scales = 50 wm (a); 100 pm (b). Skeletal arrangement in Pachyrotula raceki the genus were the Oriental and Ethiopian Stratospongilla species reported to be lack- ing microscleres (Penney & Racek 1968). A reexamination of the spicular set and gemmular coat of Spongilla (Stratospongil- la) raceki shows its gemmoscleres to be bir- otulates, modified by a graded reduction in the size of their rotules that depends on their position relative to the inside or out- side of the gemmular coat; this morpholog- ical series ends in highly variable acanthox- eote or acanthostylote forms. Just as such gemmoscleres cannot be considered stron- gyles, the gemmular structure of Pachyro- tula has no parallel in the large mammilate gemmules that are cemented to the substrate in both Stratospongilla and Oncosclera. Neither of these genera show any trace of radially arranged gemmoscleres that are ev- ident in Pachyrotula new genus. Nor are the size, the shape, or the localization of the A494 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON -,9 >? Q Oe be sme voa dd 9 e eo e990 “o ° Xe e%PF ves vo . 2 BS SO De, ce OF b Fig. 5. Camera lucida drawings of spicules of Pachyrotula raceki (Riitzler). a, Alpha megascleres grading to beta megascleres; b, Gemmoscleres. Scale = 50 wm. acanthoxea or acanthostyles around the gemmules characteristic of microscleres. Rather, they point to a second category of megascleres. The selective process respon- sible for the production of such megascleres also caused the birotulate spicules in the gemmular wall to depart from a radial pat- tern. The new genus is closer to genus Het- VOLUME 110, NUMBER 4 erorotula Penney & Racek, 1968, which at the time of its discovery was known only from Australia and New Zealand. Only a recent study of some spongillite (ore) de- posits from Brazil (Volkmer-Ribeiro & Motta 1995) has brought to light the first neotropical species of the genus. The new genus clearly stands apart from Heteroro- tula, whose diagnostic characteristic con- sists of gemmoscleres with “slender, usu- ally granulated shafts and terminally with comparatively wide and flat rotules of mod- erately to greatly varying diameter, the in- ner invariably larger than the outer’’ (Pen- ney & Racek 1968:96). Furthermore, the very distribution of Pachyrotula raceki, which appears to be the most widely oc- curring species on New Caledonia (Fig. 9) and is also known for its broad range of habitats, indicates that it cannot be a mere ecomorph of a Heterorotula species. Heterorotula caledonensis, new species Figs. 6, 9 Ephydatia multidentata (Weltner) f. cale- donensis Riitzler, 1968:63, figs. 13-19. Holotype.—USNM 23884 (the gemmule slide preparation figured in Riitzler, 1968, fig. 15), sta. FNK 36 (in Riitzler 1968:58), New Caledonia; Ferdinand Starmiihlner leg. 25 Jul 1965. Paratypes.—USNM 39463 (5 slides), sta. FNK 36 (in Riitzler 1968:58) New Caledo- nia, Ferdinand Starmiihiner leg. 25 Jul 1965. Remarks on the syntypic series.—The de- Scription of the new species is based on several fragments from three specimens. The fragments in alcohol and five slides (two of them with entire gemmules) repre- senting the three specimens are deposited in the USNM. Type locality.—Sta FNK 36 (in Riitzler 1968:58), near La Foa, southwest New Cal- edonia (Fig. 9). Diagnosis.—Heterorotula species with stout megascleres and stout, highly variable spined to granulated gemmoscleres that make up a series, running from quite long 495 acanthoxea with bulbous, strongly spined extremities to more typical birotules, to quite short, irregularly shaped “‘aster-like”’ spicules. The series of regular, birotulate Heterorotula gemmoscleres tends to form freak rotules and displays granulated to al- most smooth shafts. Description.—Ritzler’s (1968:65) de- tailed description of Ephydatia multidentata f. caledonensis holds for Heterorotula ca- ledonensis new species. In addition, a series of very short “‘aster-like’’ gemmoscleres not referred to originally may be seen in Riitzler’s, 1968, fig. 16 alongside the gem- moscleres in the pneumatic coat. Such bir- otulate-derived gemmoscleres are abundant in the original slides examined. Remarks.—Ephydatia multidentata Welt- ner 1895 was one of the species that Penney & Racek (1968) transferred to their new ge- nus Heterorotula. Racek (1969) found it to be one of the most common freshwater sponge species in eastern Australia and re- ported considerable variation in its spicular components. On New Caledonia, H. multi- dentata (Weltner 1895) occurs almost sym- patrically with H. caledonensis new species (sta. FNK 36 and 44, both described in Riit- Zler, 1968) and in quite similar water-qual- ity and substrate conditions. The great vari- ations in the gemmoscleres of H. caledo- nensis new species could not be found in specimens of H. multidentata from either eastern Australia or New Caledonia and are now considered to be of such magnitude that they denote a new species. Oncosclera diahoti (Riitzler), new combination Fig. 9 Spongilla (Stratospongilla) diahoti Riitzler, 1968:59, figs. 2—4. Holotype.-—USNM 23881 (5 fragments), New Caledonia, Ferdinand StarmiihIner leg. 16 Sept 1965. Type locality.—River Le Diahot near Quénia, sta. FNK 105, North New Cale- donia (Fig. 9). 496 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Camera lucida drawings of spicules of Heterorotula caledonensis, new species. a, Megascleres; b, acanthoxeote gemmoscleres; c, birotulate gemmoscleres (c’, head-on view of rotules); d, “‘aster-like’’ gemmos- cleres. Scale = 50 pm. VOLUME 110, NUMBER 4 497 Fig. 7. Camera lucida drawings of the spicules from several specimens of Heterorotula multidentata (Welt- ner) from Australia. a, Megascleres, b, Gemmoscleres. Scale = 50 pm. 498 Fig. 8. Tornote megascleres; b, Strongylote megascleres. c, Beta megascleres; d, gemmoscleres; e, ““Aster-like”’ mi- croscleres. Scale = 50 pm. Remarks.—From Riitzler’s (1968) de- tailed description and illustrations of Spon- gilla (S.) diahoti, particularly of the gem- mular structures, it appears the species be- longs to the genus Oncosclera Volkmer-Ri- beiro, 1970, which by definition contains Stratospongilla-like species that lack mi- croscleres and have their gemmoscleres PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Oat o ° o e Kg Reg a Ca “SS Fo 92 0 C09 2» co°o o ae) 99 Camera lucida drawings of the spicules in Houssayella iguazuensis Bonetto & Ezcurra de Drago. a, loosely and tangentially arranged in the out- er gemmular coat. Oncosclera diahoti stands very close to O. navicella from South America, which also has small oxea with a middle thickening as gemmoscleres. However, the gemmoscleres of O. navicella all have a strong middle curvature that makes them look like small boomerangs. VOLUME 110, NUMBER 4 } FNK 112 8 jo FNK 105 Ox Nouvelle FNK 76 Q Fig. 9. Map of New Caledonia showing the col- lecting stations and the distribution of the freshwater sponge fauna. Square = Pachyrotula raceki (Riitzler); asterisk = Oncosclera diahoti (Riitzler); open circle = Heterorotula caledonensis n. sp.; triangle = Hetero- rotula multidentata (Weltner 1895). Adapted from Riitzler (1968). Discussion In updating the only known survey of New Caledonian freshwater sponges, we found three genera: Pachyrotula new ge- nus, Heterorotula Penney & Racek, 1968, and Oncosclera Volkmer-Ribeiro, 1970. We also found four species: P. raceki (Riitzler 1968), H. multidentata (Weltner 1895), H. caledonensis new species, and O. diahoti (Riitzler 1968) new combination. The new genus occurs in the northern and southern extremities of New Caledonia (Fig. 9), in both lotic and lentic environments, whereas the genus Heterorotula is found near the southern third of the island and the genus Oncosclera occurs only on the northern part of the island. Most of the freshwater sponge fauna of New Caledonia seem to have evolved around the genus Heterorotula (with two Species) and the new monotypic genus Pa- chyrotula, which stands closer to Hetero- rotula than to any other known genus of freshwater sponge on account of its thick, dissimilar-sized rotules of gemmoscleres with scalloped profiles, its lack of micro- scleres, and the fact that the outer part of 499 the pneumatic layer is reinforced with a second category of megascleres of evident birotulate origin. Five species of Heterorotula have also been recorded in Australia (Penney & Ra- cek 1968, Racek 1969), one species in New Zealand (Penney & Racek 1968), and one species in South America (Volkmer-Ribeiro & Motta 1995). The highest species diver- sity occurs in Australia, where one species is restricted to the arid central part and thus to standing saline waters, another occurs only in the swampy area east of the Divid- ing Range, and the other three have a marked east-west distribution (Racek 1969). Australia’s species show a number of variations, such as the occurrence in some H. multidentata specimens of a spic- ule that does not seem to fit the category of a microsclere and the reinforcement in some species of the outer pneumatic layer with normal megascleres or with a second category of megascleres (Racek 1969, Fig. gp): Heterorotula’s presence in South Amer- ica was first reported by Volkmer-Ribeiro & Motta (1995), in their description of H. fistula from southwestern Brazil. Hetero- rotula spicules have recently been found in pond sediments in the northeastern and southern coastal areas of the country (Volk- mer-Ribeiro, unpublished). This suggests that the genus may also be present in the eastern part of South America. The distribution pattern of the genus Het- erorotula thus resembles an arch extending from eastern South America to New Cale- donia. The area richest in species is Austra- lia. Zinmeister (1979, 1982) has suggested that such a faunistic distribution may have existed in the past, in the Low Tertiary Weddelian Province, as indicated by a fossil molluscan fauna. At that time, the Antarc- tic-Australian bloc occupied a central po- sition in a splitting land arch that had iso- lated the South Circum-Pacific for a con- siderable period of geological time. It may be that the genus Pachyrotula stands for the eastern branch of an old pa- 500 rental stock with Heterorotula relationship. This suggestion is supported by the simi- larity in characteristics, the central position of the Antarctic and Australian in the Wed- delian Province, and the fact that the high- est Heterorotula species diversity occurs in Australia. Could a western branch of that group also have evolved in South America? This question brings to mind the monotypic genus Houssayella with H. iguazuensis Bo- netto & Ezcurra de Drago, 1966, described from the lower Parana River rocky bottom in Argentina and later from the lower Uru- guay River in southern Brazil (Volkmer-R1- beiro 1971, Fig. 8). In H. iguazuensis, the short birotulates with thick shafts are ar- ranged radially in the gemmules. This spe- cies is also known for its irregularly sculp- tured granulated rotules of dissimilar size, and for a series of beta megascleres that form a palissade packing around the gem- mules; this series of beta megascleres ends in short, almost birotulate spicules quite similar to the ones found in P. raceki. The megascleres in H. iguazuensis vary from ty- lote to strongylote and are spined, with a concentration of spines near both ends. A third category of highly variable strongy- lote spicules with bulbed ends approaches the shape of beta megascleres of Hetero- rotula caledonensis new species. At the same time, H. iguazuensis has aster-like mi- croscleres that closely resemble the aster- like gemmoscleres found in H. caledonen- sis New species. No fossil or extant freshwater sponges have yet been reported from Antarctica. However, the freshwater sponge genera considered in this discussion may well have evolved from an ancient Weddelian Heter- orotula-like stock. Conclusions The geographical distribution and the characteristics shared by the freshwater sponge genera Heterorotula (Penney & Ra- cek 1968), Pachyrotula new genus, and Houssayella Bonetto & Ezcurra de Drago, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1966, indicate that these genera may be the recent branches of an ancient fauna that thrived during the early Tertiary in the southern Circum-Pacific continents, which by then had already begun to drift. This re- gion, also known as the Weddelian Prov- ince (Zinmeister 1979, 1982), included southern South America, Antarctica, Aus- tralia, New Zealand, and New Caledonia. Heterorotula caledonensis new species may have the largest number of character- istics indicative of the ancient group, such as long megascleres; long birotulate gem- moscleres that include transitions to beta megascleres; short gemmoscleres, some of which grade into an “‘aster-like”’ spicule sit- uated in the gemmule or outside it; and the regular, spiny to granulated birotulates with thick, unequal and ragged rotules. Acknowledgments Grant 453930-95.6 of the National Coun- cil for the Scientific and Technological De- velopment of Brazil (CNPq) enabled C. Volkmer-Ribeiro to examine material at the National Museum of Natural History from 18 to 31 March 1996. We are indebted to A. A. Racek for donating a slide collection of Australian freshwater sponges, to I. Ezcurra de Drago for providing samples of Houssayella iguazuensis, and to Frederick W. Harrison for valuable comments on the manuscript. Thanks also to Milene M. da Silva and Rejane Rosa (MCN) and Molly K. Ryan (NMNH) for assisting in the prep- aration of the figures, and to Cleodir J. Mansan (MCN) for operating the SEM. This paper is dedicated to the memory of A. A. Racek, eminent freshwater-sponge systematist who died in early 1997. Literature Cited Bonetto, A. A., & Ezcurra de Drago. 1966. Nuevos aportes al conocimiento de las Esponjas Argen- tinas.—Physis 26(71):129—140. Penney, J. T., & A. A. Racek. 1968. Comprehensive revision of a worldwide collection of freshwater sponges (Porifera: Spongillidae).—United States National Museum Bulletin 272:1—184. VOLUME 110, NUMBER 4 Racek, A. A. 1969. The freshwater Sponges of Aus- tralia (Porifera: Spongillidae).—Australian Journal of Marine and Freshwater Research 20: 267-310. Riitzler, K. 1968. III. Freshwater sponges from New Caledonia.—Cahiers ORSTOM, Sér. Hydro- biologie 2:57—66. Volkmer-Ribeiro, C. 1970. Oncosclera, a new genus of freshwater sponges with redescription of two species.—Amazoniana 2:435-—442. 1971. Houssayella iguazuensis Bonetto & Ezcurra de Drago, 1966 (Porifera—Spongilli- dae) in Iti River, Rio Grande do Sul, Brazil.— Iheringia, Série Zooldgica 40:53-—60. , & J. EM. Motta. 1995. Esponjas formadoras 501 de espongilitos em lagoas no Triangulo Mineiro e adjacéncias, com indicacgao de preservacao de habitat.—Biociencias 3(2):183—205. Zinsmeister, W. J. 1979. Biogeographic significance of the Late Mesozoic and Early Tertiary mol- luscan faunas of Seymour Island (Antarctic Peninsula) to the final breakup of Gondwana- land. Pp. 349-355 in J. Gray & A. Boucot, eds., Biogeography, Plate Tectonics and the Chang- ing Environment. Proceedings, 37th Annual Bi- ology Colloquium and Selected Papers. Oregon State University Press, Corvallis. . 1982. Late Cretaceous—early Tertiary mol- luscan biogeography of the southern Circum Pacific.—Journal of Paleontology 56:84—102. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):502—510. 1997. The genus Julavis de Laubenfels (Porifera: Halichondrida) Rob W. M. van Soest and Helmut Lehnert (RVS) Institute for Systematics and Population Biology (Zoological Museum), University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands (HL) Institut & Museum fiir Geologie und Palaontologie, Goldschmidtstr. 3, 37077 Gottingen, Germany Abstract.—Julavis jamaicensis new species is reported from Northern Ja- maica. This represents the second record of the genus and the first from the Atlantic basin. Julavis de Laubenfels 1936 was erected to accomodate Tedania levis Kirkpatrick 1900 from the Funafuti Atoll, Central Pacific. Its status was never discussed prior to the present paper and order and family allocation (Poecilosclerida: Acarniidae) have remained tentative. It is proposed to allocate it to the order Halichondrida, family Desmoxyidae, on the basis of confused skeletal architecture, possession of acanthose diactines and wispy trichodrag- mas. The genus is distinguished from other genera of Desmoxyidae by its smooth strongylostyles and by the strongylote nature of the acanthose diactines. The contents of the family Desmoxyidae is discussed and compared with re- lated families. Kirkpatrick (1900) described from deep waters (90—125 m) off the atoll of Funafuti, Central Pacific, a thinly encrusting sponge with the unusual skeletal structure of a felt- work of spined strongyles with scattered smooth strongyles, combined with frayed trichodragmas. He named the species Te- dania levis but admitted that it was “ only placed in Tedania provisionally, .. . should probably come under a new genus near Tedania... ’”’. De Laubenfels (1936) in his monograph of the orders, families and genera of the Porifera erected a new genus Julavis (origin of the name not ex- plained) without having seen the original material. His definition for the new genus was: “... two distinct categories of spiny strongyles (sic) for megascleres and raphi- des for microscleres.”’ This is a misrepre- sentation of Kirkpatrick’s description, as there are smooth and spiny megascleres rather than two spiny types. Since then, as with so many other “paper” genera of de Laubenfels, no further reference was made in the literature on either Tedania levis or the genus Julavis. Recently, one of us (HL) collected an or- ange encrusting sponge off the north coast of Jamaica, with characters closely similar to those of Tedania levis. This material is described below as a species new to science and compared to the type specimen of T. levis borrowed from the collections of the Natural History Museum, London (BMNH). In view of the fact that there are now two species answering to the definition of Ju- lavis, it is proposed to consider the genus as valid. Its family and order allocation has to be changed from Poecilosclerida: Acar- niidae, as proposed by de Laubenfels, to Halichondrida: Desmoxyidae; this will be discussed below. Julavis jamaicensis, new species Figs. 1-3 Material examined.—Holotype: Zoolog- ical Museum Amsterdam, Porifera collec- tion, reg.no. ZMA POR.11520. Jamaica, off Chalet Caribe, Montego Bay in 20 m depth, dried specimen. Description.—Thinly encrusting on the VOLUME 110, NUMBER 4 surface of a large dead sclerosponge, Cer- atoporella nicholsoni (Porifera: Ceratopo- rellidae). The type specimen was sawed out of the basal skeleton of a large (25 cm di- ameter) specimen of the sclerosponge (Fig. 1). It is about 1 mm thick, lateral expansion at least 10 X 15 cm. Surface in dry con- dition smooth, hard, crumbly, difficult to section tangentially. No visible oscules or openings. Colour light orange to beige. Skeleton (Figs. 2A—B) in dry condition a packed feltwork of confusedly arranged acanthostrongyles with occasional single smooth long spicules in a position perpen- dicular to the surface. The outermost acan- thostrongyles tend to form an irregular pal- isade with apices pointing outward at dif- ferent angles. Rarely, the smooth long spic- ules form bundles. Trichodragmas are scattered throughout the spicule mass. The organic parts are greatly reduced (no doubt due to shrinking). There is no recognizable spongin, but the narrow space between the spicules is definitely fibrous. Spicules (Figs. 2C—E, 3, Table 1) include acanthostrongyles, strongylostyles and tri- chodragmas. Acanthostrongyles (Figs. 2C, 3.1—3) are in majority entirely covered by coarse blunt spines; they are somewhat ir- regularly shaped, often curved, with apices slightly narrower than the shaft. Occasion- ally they are almost entirely smooth (Fig. 3.2) or smooth asymmetrically at one of the ends (Fig. 3.3); juvenile spicules are finely acanthose, almost smooth, oxea-like (Fig. 2C, middle spicule). Size 127—258 X< 8—20 wm. Strongylostyles (Figs. 2D, 3.4) are smooth, style-like but conically rounded at one end and often rather bluntly pointed or Stair-stepped at the other. Occasionally the conical end is rhabdose, i.e. abruptly curved. They are rare both in sections and in dissociated spicule mounts and invari- ably broken. Size up to at least 850 X 4-8 wm. Trichodragmas (Figs. 2E, 3.5—6) form wispy, straight or S-curved bundles of 12- 20 raphides. Bundles may be entirely sheathed and then superficially resemble 503 oxea-like spicules (Fig. 3.5). Size 52-152 x 2-8 pm. Ecology: the sclerosponge encrusted by the new species was collected in a cave. Comparison with Tedania levis.— The type specimen of TJ. levis, BMNH 1900.10.19.16, is a thin crust of 1.5 mm on a small piece of coralline alga. It has the Same appearance and consistency as Julavis jamaicensis. Differences between the two species are mostly confined to details of the spicules (Table 1). The acanthostrongyles form the same felted mass as in J. jamai- censis, but the uppermost are tangentially arranged (Fig. 4A—B), rather than in a pal- isade. The acanthostrongyles (Figs. 4C, 5.1—2) are much more strongly curved and distinctly longer and thinner than those of J. jamaicensis: 211-340 X 7-14. The long smooth spicules (Figs. 4E, 5.3) closer to typical strongyles, although they also show somewhat unequal ends. Their sizes appear to be somewhat longer: up to 1385 xX 4-12 wm. A shorter category of styles reported by Kirkpatrick (245 ym) appears to be for- eign. The trichodragmas (Figs. 4D, 5.4—6) are similar but longer than those of J. ja- maicensis: 144—203 X 3-12 wm. These skeletal differences in themselves do not form an impressive load of evidence for specific distinctness between the two specimens because specific variation is un- known. However, the wide geographic sep- aration of the recorded specimens supports these small morphological differences and we erect the new species with confidence. Generic allocation.—De Laubenfels (1936) erected Julavis on the basis of the incompatibility of the described characters of Tedania levis with those of the genus Tedania. It is clear from recent discussions on the contents of Tedania and the family Tedaniidae (e.g., in Desqueyoux-Fatindez & van Soest 1996) that de Laubenfels was correct in removing T. levis from Tedania. The synapomorphy for Tedaniidae, i.e., on- ychaetes, is lacking in T. levis. With two related species known, use of the genus Ju- lavis is certainly justified. It remains a rare 504 Tr mi ay Fae - Ae AY Big: i. genus with only two records at opposite sides of the world, perhaps indicating a wider Tethyan distribution. Family and order allocation.—The ma- jor problem with Julavis is the family and order allocation. De Laubenfels (1936) as- signed Julavis to the family Acarniidae, erected for an odd assortment of genera, in- cluding among others Sceptrintus (now Hadromerida: Latrunculiidae), Janulum (Poecilosclerida: Raspailiidae), Jelissima (Poecilosclerida: Myxillidae) and Jones (Poecilosclerida: Coelosphaeridae). The type genus of the family Acarniidae is Acarnia Gray, 1867, erected for type spe- cies Hymeniacidon cliftoni Bowerbank, 1864, which is a junior synonym of Clath- ria frondifera (Lamarck 1814) (see Hooper & Wiedenmayer 1994: 256). Thus, Acar- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Julavis jamaicensis new species, photo of holotype encrusting Ceratoporella nicholsoni. niidae de Laubenfels, 1936 is a junior syn- onym of Microcionidae Carter, 1875. Julav- is needs to be removed from Acarniidae/ Microcionidae because it lacks the ectoso- mal subtylostyles, toxas and palmate iso- chelae characteristic for this family. The skeletal architecture and spicule complement would allow allocation to both Poecilosclerida (e.g., family Crellidae) and Halichondrida (e.g., families Desmoxyidae and Halichondriidae). Crellidae have a surface crust of acan- thoxeas or acanthostyles and a choanosomal skeleton consisting of bundles of smooth tornotes, which may be oxea-like, strongly- like or style-like. Normally there are chelate microscleres and short echinating acantho- styles, but these may be absent. Allocation of Julavis to Crellidae is not warranted for 505 VOLUME 110, NUMBER 4 Ah UNV SS YRWe et fe i Eo Of — — 2. oo 4%! . i ; (Yet aN . < A ae BCS 4 SY J *| Ses << ‘XN Ws surface view of ectosomal skeleton; C, various growth stages of acanthostrongyles; D, trichodragmas; E, smooth style and stron- > Fig. 2. Julavis jamaicensis n. sp., drawings of skeleton and spicules. A cross section; B, gylote modification. 506 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. spined acanthostrongyle (650); 3.2, virtually smooth ‘‘acanthostrongyle”’ (<650); 3.3, partly smooth, partly spined acanthostrongyle (X650), 3.4, style (<650); 3.5, sheathed trichodragma 2200); 3.6, irregularly fanned- out trichodragma (X2200). the following reasons: Crellidae tend to be soft sponges with surface areolae or veinal canals visible; the tangential surface crust is generally thin (single spicule layer) and strictly tangential; the acanthose spicules are never strongyles; the tornotes are sel- dom longer than 300 wm and never over 600 wm; and, no trichodragmas have been recorded from Crellidae. Halichondriidae and Desmoxyidae are considered closely related families in a re- defined order Halichondrida by van Soest et al. (1990). The generic content of these Julavis jamaicensis n. sp., Scanning electronic microscope (SEM) photos of the spicules. 3.1, a fully two families is still in debate: for example, Ptilocaulis, assigned to Desmoxyidae by van Soest et al. (1990) on account of its gross morphological similarity to certain Higginsia species, was shown to be very close to Axinella of the family Axinellidae by Alvarez et al. (1997); the allocation of Axinyssa and Myrmekioderma to Halichon- driidae is disputed by Hooper & Bergquist (1992) and Hooper & Lévi (1993). Myrmekioderma shares several features with Julavis: spined diactinal surface spic- ules (oxeas in Myrmekioderma), smooth VOLUME 110, NUMBER 4 507 Table 1.—Comparison of spicule types and sizes (wm) of Julavis levis (Kirkpatrick 1900) and J. jamaicensis n.sp. (ranges, means in italics). Spicule type Julavis levis Julavis jamaicensis Acanthostrongyles 211—280.5—340 x 7-11.1-14 127—185.2—258 X 8-14.1—20 Strongylostyles* <1385 x 4-12 <850 X 4-8 Trichodragmas 144—178.3—200 X 3-—3.8-12 52—108.3-152 X 2-3.7-8 * Most often broken in the spicule slides. diactinal choanosomal spicules (oxeas in Myrmekioderma), and wispy _ trichodrag- mas. The two genera are nevertheless con- sidered to be distinct because spination of the surface spicules in Myrmekioderma is very fine (and may be occasionally absent) and the choanosomal spicules are arranged in definite tracts. The surface of the known species of Myrmekioderma shows a char- acteristic groove pattern (see van Soest et al. 1990). A further genus sharing features with Ju- lavis is Heteroxya Topsent, 1904. Van Soest et al. (1990) tentatively considered it a ju- nior synonym of Myrmekioderma, but the evidence for that assumption is weak. The genus is defined as having spined and smooth oxeas in a confused mass, with smaller spined oxeas forming a dense pal- isade at the surface. No trichodragmas have been reported from the type and only as- signed species, H. corticata Topsent, 1904. The strongylote and stylote spicules as well as the possession of trichodragmas easily differentiate Julavis from Heteroxya. Another genus that needs to be consid- ered is Higginsia (Desmoxyidae), which likewise shares with Julavis diactinal spined surface spicules (oxeas) over much longer smooth choanosomal spicules. The two genera are considered distinct because the spined oxeas of Higginsia have a dis- tinct angular curve in the middle. These spicules often do not form a surface crust but are scattered among the choanosomal megascleres. The choanosomal skeleton consists of well-defined tracts of smooth megascleres. Julavis links Higginsia and Myrmekio- derma by its possession of coarsely spined diactinal surface spicules (shared with Hig- ginsia) and wispy trichodragmas (shared with Myrmekioderma). It is sufficiently dis- tinct from both in possessing acanthostron- gyles in a thick surface feltwork. Accord- ingly we propose to include Julavis in a re- arranged and redefined family Desmoxyi- dae, which in disagreement with van Soest et al. (1990) receives Myrmekioderma and the closely similar Didiscus from the family Halichondriidae, and loses Ptilocaulis to the family Axinellidae. Family Desmoxyidae.—Halichondrida with a surface skeleton consisting of spined diactinal spicules (oxeas or strongyles); the choanosomal skeleton is formed either by a confused or perpendicular arrangement of single spicules or interconnected bundles perpendicular to the surface. Higginsia Higgin, 1877 Gun. syn. Des- moxya Hallmann 1917): spined oxeas with abrupt angular curve in the middle; choan- osomal skeleton an elaborate system of bundles of megascleres. Halicnemia Bowerbank, 1866: spined oxeas with abrupt angular curve in the mid- dle; two categories of choanosomal mega- scleres one of which is erect on the sub- strate while the other surrounds the first. Heteroxya Topsent, 1904: Surface skele- ton a palisade of smaller spined oxeas, though which perpendicular subectosomal longspined oxeas protrude; choanosomal skeleton a confused mass of smooth and spined oxeas. Myrmekioderma Ehlers, 1870: surface oxeas rugose or finely spined; choanosomal megascleres in several length categories ar- ranged in a system of interconnected tracts; microscleres wispy or straight trichodrag- 508 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Julavis levis (Kirkpatrick), holotype BMNH 1900.10.19.16, drawing of skeleton and spicules. A, surface view of ectosomal skeleton; B, cross section; C, various growth stages of acanthostrongyles; D, tri- chodragmas; E, smooth strongyle. VOLUME 110, NUMBER 4 509 Fig. 5. thostrongyle (300); 5.2, detail of part of acanthostrongyle (800); 5.3, terminal part of smooth strongyle (X800); 5.4, sheathed trichodragma (300); 5.5, partly split-open trichodragma (300); 5.6, detail of trichod- ragma (X2200). mas; surface has characteristic sinuous grooves. Didiscus Dendy, 1922: surface oxeas ru- gose or finely spined, possessing two un- equally sized discs assymmetrically along the shaft; choanosomal megascleres in sev- eral length categories arranged in a system of interconnected tracts; surface has char- acteristic sinuous grooves. Julavis de Laubenfels, 1936: surface spicules are coarsely spined strongyles forming a thick felted mass at the surface; choanosomal skeleton reduced, consisting Julavis levis (Kirkpatrick), holotype BMNH 1900.10.19.16, SEM photos of the spicules. 5.1, acan- of long strongylostyles arranged singly per- pendicular to the surface; wispy trichodrag- mas. Acknowledgments HL was funded by the Deutsche For- schungsgemeinschaft (Re 665/10-1) and thanks Prof. Dr. Joachim Reitner, the pro- ject leader. Thanks also to the dive-buddies Greg Lee and Jean Luc Solandt and to the ‘““Reefkeepers’’, Montego Bay who sup- plied boat and tanks. This is contribution 510 Nr. 591 of the Discovery Bay Marine Lab- oratory, Jamaica. Ms Clare Valentine graciously sent us the specimen of Tedania levis on loan. Louis Van der Laan (ZMA) made the habit pho- tograph; Jan Vermeulen (ZMA) made the SEM photos. John Hooper (Brisbane) gave valuable advice. Literature Cited Alvarez, B., R. W. M. van Soest, & K. Rititzler. 1997. Revision of the Central West Atlantic Axinel- lidae.—Smithsonian Contributions to Zoology (in press). Desqueyroux-Fatindez, R., & R. W. M. van. Soest. 1996. A review of Iophonidae, Myxillidae and Tedaniidae occurring in the South East Pacific (Porifera: Poecilosclerida)—Revue Suisse de Zoologie 103(1):3-79. Hooper, J. N. A., & PR. Bergquist. 1992. Cymbas- tela, a new genus of lamellate coral reef spong- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON es.—Memoirs of the Queensland Museum 32(1):99-137. , & C. Lévi. 1993. Axinellidae from the New Caledonian lagoon (Porifera: Demospon- giae).—Invertebrate Zoology 7(6):1395—1472. , & E Wiedenmayer. 1994. Porifera. in: A. Wells, ed., Zoological Catalogue of Australia. Volume 12. Melbourne, CSIRO, Australia, 612 Pp. Kirkpatrick, R. 1900. Descriptions of sponges from Funafuti—Annals and Magazine of Natural History (7)6:345-362. Laubenfels, M. W. de. 1936. A discussion of the sponge fauna of the Dry Tortugas in particular and the West Indies in general, with material for a revision of the families and orders of the Porifera.—(Publications of the Carnegie Insti- tute Washington 467) Publications of the Dry Tortugas Laboratory 30:1—225. Soest, R. W. M. van, M. C. Diaz, & S. A. Pomponi. 1990. Phylogenetic classification of the Hali- chondrids (Porifera, Demospongiae).—Beaufor- tia 40(2):15—62. Topsent, E. 1904. Spongiaires des Acores.—Résultats des Campagnes Scientifiques du Prince Albert ler de Monaco 25:1—280. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):511-519. 1997. Narella nuttingi, a new gorgonacean octocoral of the family Primnoidae (Anthozoa) from the eastern Pacific Frederick M. Bayer Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Specimens originally reported by Nutting (1908) as Stachyodes dichotoma Versluys from Albatross explorations in Hawaiian waters are shown to be misidentified and are now described as a new species, Narella nuttingi. In the eastern Pacific, four species of Na- rella have been reported (as Stachyodes) from the Hawaiian Islands: Stachyodes an- gularis Nutting, 1908; S. regularis Wright & Studer, 1889; S. bowersi Nutting, 1908, and Stachyodes dichotoma Versluys, 1906 (Nutting 1908). The species described by Nutting (1908: 576) as Stachyodes angularis is referable to the genus Calyptrophora, not to Narella. The specimen reported by Nutting (1908: 577) as S. regularis Wright & Studer con- sists of an incomplete terminal branch with remnants of only nine whorls of polyps, but it is adequate to demonstrate that it repre- sents S. dichotoma Versluys. The specimens reported as S. dichotoma by Nutting (1908:577) were incorrectly identified. Nutting’s (1908) plate 43, fig. 5, captioned “‘Stachyodes bowersi Nutting,” does not depict bowersi, but the specimen reported as “‘Stachyodes dichotoma’”’ (Nut- ting 1908:577) from Albatross sta. 4013 and retained by Nutting at the State Uni- versity of Iowa under the name Calypteri- nus allmani Wright & Studer, a species that he did not report from Hawaii. The type specimen of S$. bowersi (USNM 25377) was not illustrated and does not harbor speci- mens of the ophiuroid Ophiocreas, a spec- imen of which is clearly shown in Nutting’s photograph (1908: pl. 43, fig. 5) and spe- cifically mentioned in the figure explanation (1908:600). In the text (1908:577) Nutting mentions the occurrence of Ophiocreas on S. dichotoma, not on S. bowersi. Narella Gray, 1870 Narella Gray, 1870:49.—Deichmann, 1936:168.—Bayer, 1956:222; 1961:295 (key only); 1981:937 (key only). Stachyodes Studer, 1887:49; 1901:40.— Wright & Studer, 1889: xlvii, 53.—Ver- sluys, 1906:86.—Thomson & Henderson, 1906:35.—Kinoshita, 1907:233; 1908: 45.—Thomson & Russell, 1910:142.— Kukenthal, °1912:59; 1915:052:; 1919: 452; 1924:308. Calypterinus Wright & Studer, 1889:xlviii, 54. Type species.—Primnoa regularis Du- chassaing & Michelotti, 1860. Diagnosis.—Primnoidae forming colo- nies branched pinnately, dichotomously, or trichotomously, rarely unbranched; polyps in pairs or whorls, usually directed basad; sclerites of polyps comprising three or four pairs of large scales surrounding the polyp body but not solidly fused to form rings, and an operculum of 8 roughly triangular scales; coenenchyme with a layer of large scales that may be polygonal, elongate, sometimes tapered and almost fusiform. Narella nuttingi, new species Figs. 1-5 Stachyodes dichotoma.—Nutting 1908:577. Stachyodes bowersi Nutting 1908: pl. 43, fig. 5. 512 Not Stachyodes dichotoma Versluys 1906: 88. Material examined.—Kauai Island, Ha- waii, Hanamaulu warehouse bearing N. 82°45', W 3.7’, 419-399 fath. (= 767-730 m), bottom temperature 41.0°K USFC str. Albatross sta. 4013, 20 Jun 1902. One near- ly complete colony with attached ophiuroid, holotype, USNM 91864 (SEM 2336; Figs. 1, 2). This specimen erroneously illustrated as Stachyodes bowersi by Nutting, 1908:pl. 43, fig. 5. Kauai Island, Hawaii: Ukula Point bear- ing S. 82°30’, E. 13.1’, 423-438 fath. (= 774-802 m), bottom temperature 41.0°E USFC str. Albatross sta. D-4030, 24 Jun 1902. One colony somewhat broken, para- type, USNM 25376 (SEM 2335; Figs. 4, 5). Kauai Island, Hawaii: Hanamaulu Ware- house bearing S. 33°, W. 9.5’, 500-385 fath. (915-693 m), bottom temperature 40°F USFC str. Albatross sta. D-3989, 11 Jun 1902. One unbranched terminal branchlet, USNM 22561. Kauai Island, Hawaii: Hanamaulu Ware- house bearing N. 74°30’, W. 6.6’, 671-957 fath. (1228-1751 m), bottom temperature 38.4°F USFC str. Albatross sta. D-3989, 11 Jun 1902. Detached branches, USNM 25375. Kauai Island, Hawaii: Ukula Point bear- ing N. 65°30’, W. 7.4’, 508-557 fath. (= 930-1019 m), bottom temperature 40°F USFC str. Albatross sta. D-4007, 17 Jun 1902. One colony somewhat broken, para- type, USNM 77293 (SEM 2338; Figs. 3, 4). North of Maui Island, Hawaii: 21°07.0'N, 156°12.7'W, no other particulars as to collection recorded. Three large, somewhat damaged colonies, with ophiu- roids attached, USNM 56790. Cross Seamount: 18°38.7'N, 158°16.8’W, 1420 m, Pisces V dive PV238, station CR201 (sample 1), 29 Aug 1993, coll. Scott France and J. Ewann Agenbroad. One in- complete colony with attached ophiuroid, USNM 94451. Cross Seamount: 18°38.7’'N, 158°16.8’W, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1350 m, Pisces V dive PV238, station CR203 (sample 1), 29 Aug 1993, coll. Scott France and J. Ewann Agenbroad. One in- complete colony with attached ophiuroid, paratype, USNM 94452 (Fig. 1). Cross Seamount: 18°38.7'N, 158°16.8'W, 1205 m, Pisces V dive PV238, station CR205 (sample 1), 29 Aug 1993, coll. Scott France and J. Ewann Agenbroad. One in- complete colony with attached ophiuroid, USNM 94453. Diagnosis.—Bushy Narella colonies with branching initially verticillate, subse- quently becoming dichotomous and approx- imately uniplanar in the distal branches; 3- 5 polyps per whorl; commonly 8-1/2 to 11 (but sometimes as few as 5-6) whorls in 3 cm of branch length; none of the three pairs of body plates adaxially closed; all body plates regularly curved, without keels or crests, their free margins without serrations or sharp projections; one pair of small ad- axial buccal scales; apical keel on inner face of opercular scales low, absent from the larger abaxial scales; adaxials narrow, small; cortical sclerites elongate, usually narrow but a few may be wider, without external crests; tentacles with very few minute, flat rods; coenenchyme with a sin- gle layer of elongate oval plates, becoming thick and irregular on the large branches, locally with several strong, hemispherical projections longitudinally arranged along the midline. Description.—The colonies (Fig. 1) are bushy, the branching of the lower parts ver- ticillate with 3 or 4 lateral branches in wide- ly spaced whorls, in the distal branches be- coming dichotomous and tending to lie in one plane so the colonies are compressed- flabellate rather than flat fans. The axis is strongly calcified, round, longitudinally striated, with little or no metallic luster or iridescence. Specimens having the proximal part of the trunk preserved show thick sec- ondary deposits of white calcareous mate- rial around and above the holdfast. Polyps are directed basally, up to 3 mm in height (measured parallel with the branch SCIENTIFIC APPARATUS MERRICK AVE. NEARS’ il SCIENTIFIC APPARATUS MERRICK AVE. — ee 3 ry 5 warm 1 2 | itu Paratype colony, USNM > 2 — : — SCIENTIFIC CONSO - L APPARATUS (5 AVE NEARS 3 4 a Pesan eee = lS) = ° pO + Ne co — O Z Y =) > eS = ° Oo o a. > Sel = 0 am 2 i Narella nuttingi, new species. VOLUME 110, NUMBER 4 Fig: ft PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2336 06 10Ky i: Femme 40233606 1e@KyY 1. 76mm Fig. 2. Narella nuttingi, new species. Part of terminal branch of holotype colony, USNM 91684. SEM 2336, stereo pair. VOLUME 110, NUMBER 4 i Fémm 2338 8610KY | 1. 76mm Fig. 3. Narella nuttingi, new species. Part of terminal branch of paratype colony, USNM 77293. SEM 2338, stereo pair. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 5 So SAP > e335 16K 3M th e333 1QK¥ » 43mm Fig. 4. Narella nuttingi, new species. Top, Whorl of polyps of paratype colony, USNM 77293, showing prominent operculum and absence of dorsolateral crest on body scales. SEM 2338. Bottom, Detail of coenen- chyme showing elongate, plate-like sclerites; SEM 2335, stereo pairs. VOLUME 110, NUMBER 4 =~ 2335 10KY Fig. 5. Narella nuttingi, new species. Part of terminal branch of paratype colony, USNM 25376. SEM 2335, stereo pair. 518 axis), in whorls of 5 on the larger branches, sometimes only 3 near the tips of the ter- minal branchlets (Figs. 2-5); 8% to 11 whorls occur in 3 cm of branch length. Pol- yps with 3 pairs of non-annular body plates, all with uniformly and moderately expand- ed margins regularly curved and without marginal spines, teeth, or conspicuous ser- rations. The two basal plates are the largest, regularly curved around the base of the pol- yp, without a dorsolateral angle or crest, with moderately flared, smooth margins; one of the plates overlaps the other where they meet along the abaxial midline, its thin edge somewhat reflexed. The medial pair of body plates are the smallest, smoothly curved around the body and without dor- solateral crests. The buccal pair have smooth, moderately flared free margins; one pair of small, thin, adaxial buccal scales lie below the adaxial operculars. The operculum is high, the abaxial scales tall, broad, rounded apically, the outer sur- face longitudinally concave but without a corresponding well-defined keel on the in- ner surface; the lateral scales are narrower, more pointed, and have a low keel on the inner surface; the adaxials are narrow, with a low, ridge-like keel. The tentacles have a few minute, irregular flat rods. The sclerites of the coenenchyme (Fig. 4) are more or less elongate, flat plates with no trace of longitudinal crests or keels even near the polyps, but some of those on the large branches may have several hemi- spherical projections along the midline. Etymology.—This species is named for the late Prof. C. C. Nutting, who first un- knowingly reported specimens in his report on Hawaiian octocorals collected by USFC steamer Albatross. Variation.—The polyps vary somewhat in size among different colonies, and the spacing of whorls varies locally both within a single colony and among different colo- nies (compare Figs. 2, 3 and 5). Distribution.—Eastern Pacific, vicinity of Hawaiian Islands and nearby seamounts, 730-1420 m. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Commensals.—Most colonies have a large ophiuroid, Asteroschema caudatum H. L. Clark, entwined among the branches. Nutting (1908:577) reported that colonies examined by him “had coiled around its branches a simple-armed basket fish, prob- ably belonging to the genus Ophiocreas.”’ Comparisons.—Although the polyps of N. nuttingi resemble those of N. dichotoma (Versluys) as originally illustrated (Versluys 1906:89, 90), the coenenchymal scales lack the characteristic ridges of that species but instead may have several strong, hemi- spherical projections along the midline, the adaxial opercular scales are very reduced in size, and branching is not in one plane. Na- rella orientalis (Versluys), which has simi- lar polyps, differs in having broad cortical scales and more strongly developed adaxial opercular scales. The polyps of Narella megalepis (Ki- noshita 1908) are generally similar to those of N. nuttingi, but the opercular scales have a strong apical keel on the inner surface, the adaxial operculars are broad and well- developed, and the cortical sclerites are thick, irregular polygonal plates; moreover, the overall colonial form is not known be- cause all existing specimens are fragmen- tary. As originally illustrated (Kinoshita 1908, pl. 3, fig. 21), the branching is di- chotomous although it was described as ““beinahe federartig.’’ It seems unlikely that the complete colonies would have the open- ly bushy aspect of N. nuttingi. The identity of specimens collected in Japanese waters by USFC str. Albatross and reported as Stachodes megalepis by Nutting (1912:59) cannot be determined because the speci- mens were not returned to the U.S. National Museum for permanent storage. It is un- likely that they were N. megalepis as Nut- ting (1912:60) mentioned the presence of worm galleries formed by parasitic anne- lids, a feature not reported by Kinoshita (1908:47—49) or Utinomi (1979:1015). Acknowledgments I am grateful to Dr. Scott C. France and Ms. J. Ewann Agenbroad Berntson, who VOLUME 110, NUMBER 4 collected some of the material reported herein. The commensal ophiuroid was iden- tified by Dr. David L. Pawson. The stereo- graphic scanning micrographs illustrating this paper were made by Mr. Walter R. Brown, head of the SEM Laboratory, Na- tional Museum of Natural History. Literature Cited Bayer, E M. 1956. Octocorallia. Pp. 163-231, figs. 134-162 in R. C. Moore, ed., Treatise on In- vertebrate Paleontology Part E Coelenterata. Lawrence, Kansas: Geological Society of America and University of Kansas Press. xx + 498 pp., 358 figs. . 1961. The shallow-water Octocorallia of the West Indian region. A manual for marine biol- ogists.—Studies on the fauna of Curacao and other Caribbean islands 12:1—373. . 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: 901-947. 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. Gray, J. E. 1870. Catalogue of the lithophytes or stony corals in the collection of the British Museum. London: British Museum. [i—iv] + 1—51 pp. Kinoshita, K. 1907. Vorlaufige Mitteilung tiber einige neue japanische Primnoidkorallen.—Annota- tiones Zoologicae Japonenses 6(3):229—237. . 1908. Primnoidae von Japan.—Journal of the College of Science, University of Tokyo 23(12): 1-74, pls. 1-6. Kiikenthal, W. 1912. Die Alcyonaria der deutschen Sudpolar-Expedition 1901-1903.—Deutsche Sudpolar-Expedition 1901-1903, 13. Band. Zoologie 5(3):289—349, pls. 20-23. 1915. System und Stammesgeschichte der Primnoidae.—Zoologischer Anzeiger 46(5): 142-158. 1919. Gorgonaria.—Wissenschaftliche Er- ay Ie) gebnisse der deutschen Tiefsee- Expedition auf dem Dampfer “‘Valdivia’’ 13(2):1—946, pls. 30— 89. 1924. Gorgonaria.—Das Tierreich 47:i— Xxvill + 1—478, 209 figs. Berlin and Leipzig, Walter de Gruyter & Co. Nutting, C. C. 1908. Descriptions of the Alcyonaria 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— Sh: Studer, Th. 1887. Versuch eines Systemes der Al- cyonaria.—Archiv fiir Naturgeschichte 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. Calcutta: The Indian Museum. i—xvi + 1-132 pp., pls. 1-10. , & E. S. Russell. 1910. Alcyonarians collect- ed on the Percy Sladen Trust Expedition by Mr. J. Stanley Gardiner. Part 1, the Axifera.—Trans- actions of the Linnean Society of London (2) 13(2):139-164, pls. 6-14. Utinomi, H. 1979. Redescriptions and illustrations of some primnoid octocorals from Japan.—Pro- ceedings of the Biological Society of Washing- ton 91(4):1008—1025. Versluys, J. 1906. Die Gorgoniden der Siboga Expe- dition II. Die Primnoidae. Siboga-Expeditie, Monographie 13a. 1-187 pp., figs. 1-178, pls. 1—10, chart. Wright, E. P., & Th. Studer. 1889. Report on the Al- cyonaria.—Report on the scientific results of the voyage of H. M. S. Challenger during the years 1973-76 under the command of Captain George S. Nares, R. N., E R. S. and the late Captain Frank Tourle Thomson, R. N. Prepared under the superintendence of the late Sir C. Wy- ville Thomson, Knt., EF R. S. & c. Regius Pro- fessor of Natural History in the University of Edinburgh, director of the civilian scientific staff aboard and now of John Murray, LL. D., Ph. D. & c. one of the naturalists of the expe- dition. Zoology 31 part 64. i-lxxvii + 1-314 pp., 49 pls. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):520-—536. 1997. Annotated list of Veronicellidae from the collections of the Academy of Natural Sciences of Philadelphia and the National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A. (Mollusca: Gastropoda: Soleolifera) José W. Thomé, Patricia H. dos Santos, and Luciana Pedott Laborat6ério de Malacologia, Instituto de Biociéncias, PUCRS, Av. Ipiranga, 6681, prédio 12; 90619-900 Porto Alegre, RS-Brazil. Abstract.—The list of veronicellid slugs presented in this paper is restricted to species identified by criteria previously proposed by the first author. We were able to distinguish 30 species or subspecies, classified in 12 genera. In- cluded are the following: Belocaulus angustipes; Colosius propinquus; C. pulcher; Diplosolenodes occidentalis; D. olivaceus; Heterovaginina peruviana; Laevicaulis alte (with new illustrations); L. natalensis brauni; L. stuhlmanni (with new illustrations); Latipes cnidicaulus; Leidyula dissimilis; L. floridana (with new data and illustration); L. goodfriendi; L. kraussi; L. moreleti; L. portoricensis; L. trichroma; Phyllocaulis gayi; P. soleiformis; Sarasinula du- bia; S. linguaeformis; S. plebeia; Simrothula columbiana; S. prismatica; Va- ginulus taunaisii; Veronicella bahamensis; V. cubensis; V. davisi; V. sloanei; V. tenax. The family Veronicellidae Gray, 1840 comprises terrestrial mollusks without shell, commonly called slugs. The distribution is pantropical and over 300 specific names have been registered, most of them syn- onyms (Hoffmann 1925, Forcart 1953, Thomé 1975a, 1993b). The phylogenetic position of the Veron- icellidae is uncertain. Usually they are clas- sified within the order Soleolifera Simroth, 1890. Some North American authors clas- sify them in the order Systelommatophora Pilsbry, 1948. Others propose placing them in another order of Pulmonates: Archaeo- pulmonata (G6tting 1974). Zilch (1959/60) put the Soleolifera as an Order of the sub- class Euthyneura. Colosi (1922) and Hy- man (1967) placed the same as an Order of the class Opistobranchia. Thomé (1993b) discusses the classifica- tion status of American Veronicellidae, points out the necessity of embryological and ontogenetic studies, and suggests new groups of characteristics to be analyzed. He comments on the characteristics tradition- ally used and is of the opinion that there are not enough to determine conclusively the species, and that they do not permit a con- sistent phylogenetic classification. The objective of the present work is to identify species of Veronicellidae from two collections through the classical anatomical characteristics proposed by Thomé (1993b). This list is relevant due to the importance of these North American institutions’ col- lections for comparative study of Veroni- cellidae. Materials and Methods The analyzed material belongs to the fol- lowing Museums: The Academy of Natural Sciences of Philadelphia (ANSP) with 110 lots and 464 specimens; and The National Museum of Natural History, Smithsonian Institution, Washington (USNM) with 97 lots and 272 specimens. Some of these lat- VOLUME 110, NUMBER 4 ter lots were intercepted in the United States by US Department of Agriculture (USDA) and bear the occurrence number. The specimens were dissected according Thomé & Lopes (1973) and the anatomical characteristics used for identification were based on Thomé (1988a, 1993b). We have included illustrations only for those species where new data made it nec- essary for their complete description. The radula and the jaw of some speci- mens were analyzed by Scanning Electron Microscope (SEM), using the method of Gotting (1985). The synonymy is restricted to main ref- erence works. The occurrences are those cited in Thomé (1993b) with new data from these collections. Results Belocaulus angustipes (Heynemann, 1885) Vaginula angustipes Heynemann, 1885: 275-77. Belocaulus angustipes; Lopes-Pitoni & Thomé, 1981:586—93; Thomé, 1993b:70. Occurrence.—Argentina; Brazil: Rio Grande do Sul; Paraguay; Colombia; (intro- duced: Honduras; U.S.A: Florida, Alabama, Louisiana, Texas). Examined lots—ANSP A1438. U.S.A: Florida, Leon County, Tallahassee; L. T. Rose leg; 12 Oct 1967; (Elberta Carte Co. property, East side of the international air- port, Dr. Lake Bradford Rd); (3 specimens). USNM 574919. Colombia; 5 Nov 1943; (on Anthurium sp., USDA 200372); (1 specimen). USNM 575531. U.S.A: Ala- bama, Mobile; L. Hubricht leg; Jun 1961; (Mobile Co., Old Church Street, USDA 237188); (3 specimens). USNM 576363. U.S.A: Louisiana, New Orleans; 12 Sep 1962; (on lawn, USDA 244560); (1 speci- men). USNM 758251. U.S.A: Louisiana, New Orleans; D. Dundee leg; 1963; (4 specimens). USNM 769131. U.S.A: Texas, Brownsville; D. L. Dunagan leg; 27 Mar 1968; (La Verne’s Nursery, USDA 278478); (8 specimens). USNM 769164. S21 U.S.A: Texas, Raymondville; R. Kloepping leg; 11 Apr 1968; (Caldwell Nursery, USDA 278478); (1 specimen). USNM 769169. U.S.A: Texas, Brownsville; 29 Apr 1969; (USDA 288982); (1 specimen). USNM 769170. U.S.A: Texas, Harington; R. Kloepping leg; 12 Apr 1968; (Rainbow Nursery, USDA 278478); (1 specimen). USNM 769175. U.S.A: Texas, Edinburg; W. J. Bone leg; 08 Feb 1968; (Mac’s Nurs- ery; Hidalgo Co., USDA 273170); (9 spec- imens). USNM 769176. U.S.A: Texas, Ray- mondville; R. Kloepping leg; 30 Jan 1968; (La Verne’s Nursery; USDA 273170); (5 specimens). USNM 769178. U.S.A: Texas; W. J. Bone leg; 8 Feb 1968; (McAllen; Waugh Nursery, USDA 273170); (1 speci- men). USNM 769180. U.S.A: Texas, San Benito; O. V. Mullins leg; 30 Jan 1968; (Bougainville Nursery, USDA 273170); (6 specimens). USNM 769187. U.S.A: Texas, Brownsville; B. C. Stephenson leg; 30 Jan 1968; (Tropical Nursery; Cameron Co., USDA 273170); (2 specimens). USNM 769196. U.S.A: Texas, Corpus Christi; E. FE Sublett leg; 23 Apr 1968; (Backyard Nurs- ery, Nueces Co., USDA 278478); (1 spec- imen). USNM 769198. U.S.A: Texas, San Benito; O. V. Mullins leg; 12 Apr 1968; (Brown Lee Farm; Cameron Co., USDA 278478); (6 specimens). USNM 769199. U.S.A: Texas, San Benito; D. L. Dunagan leg; 9 Apr 1968; (Bougainville Nursery, Cameron Co., USDA 278478); (3 speci- mens). USNM 791487. U.S.A: Florida, Pensacola; J. P. E. Morrison leg; Dec 1961; (1 specimen). Colosius propinquus (Colosi, 1921) Vaginula propinqua Colosi, 1921:157; Co- losi, 1922:499—500. Angustipes (Angustipes) pulcher; Kraus, £95555: (0-D.). Occurrence.—Ecuador; U.S.A: Puerto Rico. Examined lot.—ANSP A1440. Porto Rico: Yauco river, Yaves Lares; H. B. Ba- ker; (8 specimens). Colosius pulcher (Colosi 1921) Vaginula pulchra Colosi, 1921:157; Colosi, 1922:496—98; Thomé, 1970b:23—5, 27-9, BT: Angustipes (Angustipes) pulcher; Kraus, 1953:63 (p.p.). Colosius pulcher; Thomé, 1993b:72. Occurrence.—Ecuador; Antilles: Domin- ican Republic. Examined lots—ANSP A1092. St. Do- mingo; W. M. Gabb leg; (7 specimens). USNM 769142. Ecuador; 18 Mar 1969; (in- tercepted at Miami on bromeliads); (1 spec- imen). Diplosolenodes occidentalis (Guilding 1825) Onchidium occidentale Guilding, 1825:322. Diplosolenodes occidentalis; Thomé, 1985: 411-17; 1993b:71. Occurrence.—Guyana; Venezuela; Co- lombia; Panama; Costa Rica; El Salvador; Nicaragua; Honduras; Antilles: Dominica, W. I.; Bahamas; U. S. A: Hawaii. Examined lots ——ANSP A1076. Nicara- gua; W. G. Binney leg; (2 specimens). ANSP A1079. Panama: Punta de Pifia, R. E. B. Mc Kenney leg; 11 Jan 1907; (1 spec- imen). ANSP A1080 (=71911). Dominica, W. I.; T. D. A. Cockerell leg; (3 specimens). ANSP A1083. Costa Rica: Rio Surubrer, Bonnefil farms; P. Calvert leg; 16 Oct 1909; (2 specimens). ANSP A1086 (=46583). Colombia: Aracataca; (1 specimen). ANSP A1095 (=57954). Dominican Republic; G. E. & A. H. Verrill leg; 1892; (1 specimen). ANSP A1098 (=140994). Venezuela: Pal- ma Sola; H. B. Baker; 1920; (1 specimen). ANSP A1107a (=45180). A. A. Hinkley leg; 1914; (4 specimens). ANSP A1123. Panama: Flamenco Island, Canal Zone; N. L. H. Krauss leg; 29 Oct 1948; (1 speci- men). ANSP A7018. British Guiana, (4°0’ N, 58°0’ W); H. Lang leg; (1 specimen). ANSP A7019. British Guiana, (4°0'’ N, 58°0’ W); (2 specimens). ANSP A7022. British Guiana: Georgetown, (6°46’ N, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 58°10’'W); Lang leg; (1 specimen). USNM 574369. U.S.A: Hawaii, Oahu Island, Eka- hanui Gulch; G. S. Starkey leg; 08 May 1945; (from Army Medical Museum, USDA 171519); (1 specimen). USNM 575267. Panama: Cathedral Plaza; J. Zetek leg; (USDA 215629); (1 specimen). USNM 575897b. Honduras; 24 Jan 1961; (inter- cepted at Charleston on bananas, USDA 234701); (1 specimen). USNM 575973. Ba- hamas; 20 Oct 1959; (intercepted at Miami on orchids, USDA 227708); (7 specimens). USNM 575975. El Salvador; 15 Jun 1960; (intercepted at Miami on orchids, USDA 231346); (1 specimen). USNM 576190. Nicaragua: Ticuantepe, El Mamon; 20 May 1957; (on plants, USDA 216243); (1 spec- imen). USNM 805216. Honduras: Bantua; G. W. Van Hyning leg; 03 Oct 1929; (1 specimen). Diplosolenodes olivaceus (Stearns 1871) Veronicella olivacea Stearns, 1871:1. Diplosolenodes olivaceus; Thomé & Lopes- Pitoni, 1976:710—12. Occurrence.—Nicaragua. Examined lot—USNM 39160. Nicara- gua: Hacienda Polvén, Departamento de Leon. J. A. McNiel leg; 1885. (Lectotype). Heterovaginina peruviana (Kraus 1953) Vaginina (Heterovaginina) peruviana Kraus, 1953:63—5. Heterovaginina peruviana; Thomé, 1969: 357; Occurrence.—Peru. Examined lot—USNM 574941. Peru; 01 Feb 1954; (intercepted at New Orleans, on cabbage from Peru, USDA 201313); ( specimen). Laevicaulis alte (Férussac 1821) (Figs 2, 5, 8, 11) Vaginulus alte Férussac, 1821:14. Laevicaulis alte; Forcart, 1953:63-8. Occurrence.—Asia (type locality: Pon- VOLUME 110, NUMBER 4 dichery, India); Oceania; Africa; (intro- duced?: Bermudas; U.S.A: Texas, Hawaii). Examined lots.—ANSP A1130/1 (=98631). Bermudas: near Harrington House; S. Brown leg; Jun 1909; (45 spec- imens). ANSP A1130/2. Bermuda, near Harrington House, Parish; 18 Feb. 1908; (12 specimens). ANSP A10173 (=28314). U.S.A: Texas, Brownsville; 25 Sep 1983; (1515 Flamingo, Cameron Co.); (7 speci- mens). USNM_ 187499. U.S.A: Hawaii, Hilo; D. B. Kuhms leg; 17 Aug 1901; (2 specimens). USNM 574368. U.S.A: Ha- waii, Oahu Island, Ekahanui Gulch; W. B. Lewellen and W. W. White leg; 31 Jan 1945; (on sewerage system, from Army Medical Museum, USDA 171519); (1 spec- imen). USNM 574371. U.S.A: Hawaii, Oahu Island, Honolulu; (USDA 171297); (2 specimens). USNM 574943. U.S.A: Ha- wali, Midway Island; Lulhilson leg; 13 Nov 1953; (intercepted at Honolulu, on Poinset- tia in soil from Midway, USDA 200540); (3 specimens). USNM 576340. U.S.A: Ha- wali, Oahu Island, Kailua; 1 Aug 1960; (USDA 232396); (1 specimen). USNM 769190. U.S.A: Hawaii; 21 Oct 1968; (in- tercepted at San Antonio, Texas with ship- ment of Achatina fulica to S. W. State Teachers College, San Marcos, Texas, USDA 278478); (1 specimen). Laevicaulis natalensis brauni (Simroth 1913) Vaginula (Annulicaulis) brauni Simroth, 1913:187. Laevicaulis natalensis brauni; Forcart, 1953:71—4. Occurrence.—Africa: South Africa. Examined lot—ANSP A1106 (=47326). Africa, South Africa: Port Shepstone; H. C. Burnup; (2 specimens). Laevicaulis stuhlmanni (Simroth 1895) (Figs 1, la, 4, 7, 10) Vaginula stuhlmanni Simroth, 1895: 61. Laevicaulis stuhlmanni stuhlmanni; Forcart, 1953:74-6. 523 Occurrence.—Africa: Zaire. Examined lots.—ANSP A1137. Rotachu- ru (?); Dr. J. Beguait; Sep 1914; (2 speci- mens). ANSP A1138. Belgian Congo (=Zaire); 1919; (3 specimens). Latipes cnidicaulis (Baker 1926) Vaginulus (Latipes) cnidicaulis Baker, 1926:29-31; Thomé, 1988c:25-7. Latipes cnidicaulis; Thomé, 1993b:72. Occurrence.—Guyana. Examined lot——ANSP A7020. (=A5927); Guiana: Kamakusa; H. Lang leg; Jan 1923. (Holotype). Leidyula dissimilis (Cockerell 1892) Veronicella dissimilis Cockerell, 1892:134. Leidyula dissimilis; Thomé, 1988b:808—09; 1993b:71. Occurrence.—Antilles: Jamaica. Examined lots—ANSP A1451. H. B. Baker; 20 Jul 1933; (7 specimens). ANSP A1453. H. B. Baker; 8 Sep 1933; (1 spec- imen). ANSP A1456. 27 Jun 1933; (3 spec- imens). ANSP A12219 (=291339a); Ja- maica: Somerset; H. B. Baker leg. (Holo- type of Veronicella leptothali Baker, 1935. ANSP A12220 (=291339b); same data. (6 specimens). (Paratypes of Veronicella lep- tothali Baker, 1935). Leidyula floridana (Leidy & Binney in Binney 1851) (Figs: 36,9, 12) Vaginulus floridanus Leidy & Binney in Binney, 1851:198, 251. Leidyula floridana; Baker 1925:167-—71; Thomé, 1993b:74. Occurrence.—Antilles: Dominica W. L.; Cuba. U.S.A: Puerto Rico, Florida, Texas. New data.—The jaw presents about 26 flattened laths, sometimes imbricate and of different sizes. The laths have small trans- verse furrows on the upper third. On some laths the furrows are lengthwise. The mid- region of the jaw is larger than the ends (Figs. 524 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-9. Penis: (1, la) Laevicaulis stuhlmanni; (2) L. alte; (3) Leidyula floridana; penial gland; (4) Laev- icaulis stuhlmanni; (5) L. alte; (6) Leidyula floridana; pedal gland: (7) Laevicaulis stuhlmanni; (8) L. alte; (9) Leidyula floridana. AC tip of the penis, PE penis, MR retractor muscle, DA anterior deferential duct, PA papilla, TU tubules, OP opening. 13, 14). On 3 radulas examined, there are ca. 94 rows and ca. 95 teeth in each, with an average of about 8.900 teeth (Figs. 15,16). Length, width, and distance from the base to the greatest width for each of 3 rachidian and 3 lateral teeth of the first file on the right was measured (Table 1) and ratios between the means also were calculated (Table 2). Examined lots —ANSP A1059 (=154994). U.S.A: Florida, Lignum Vitae Key; H. A. Pilsbry; 28 Mar 1931; (2 specimens). ANSP A1060. U.S.A: Florida, St. Petersburg; (2 specimens). ANSP A1061. Cuba: Matanzas Jumari; H. A. Pilsbry; (6 specimens). ANSP A1067. Cuba: Cienfuegos; H. A. Pilsbry; (2 specimens). ANSP A1068 (=197812). Cuba: VOLUME 110, NUMBER 4 325 Figs. 10-12. Hermaphroditic region: (10) Laevicaulis stuhlmanni; (11) L. alte; (12) Leidyula floridana. AP accessory purse, CP copulation purse, DCP duct of the copulation purse, OV oviduct, RE rectum, DL joining duct, DPD distal posterior deferential duct, DPP proximal posterior duct deferential duct, PR prostate. 526 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 639394609 14 Figs. 13-14. Leidyula floridana. (13) jaw 50X; (14) median region of the jaw 150. Paratype USNM 120394. Cienfuegos; H. A. Pilsbry; (6 miles from specimens). ANSP A1074 (=137807). Cuba: Cienfuegos) (9 specimens). ANSP A1069a. San José, Spiritus; H. A. Pilsbry; (1 speci- Cuba: Zaza del Medio, Prov. Santa Clara; H. men). ANSP A1081 (=137803)..Cuba: Sanc- A. Pilsbry (2 specimens). ANSP A1071. ti Spiritus; H. A. Pilsbry; (52 specimens). Cuba: San Juan de Letran; H. A. Pilsbry; (4 ANSP A1090 (=63528). U.S.A: Florida, Key VOLUME 110, NUMBER 4 = 907 Figs. 15-16. Leidyula floridana. Median region of the radula 500X, (15) paratype USNM 805188; median region of the radula 500X, (16) paratype 805201. L lateral tooth, R rachidian tooth. Largo; H. C. Manchette leg; (7 specimens). ANSP A1091. U.S.A: Florida, Miami; H. A. Pilsbry; 1899; (5 specimens). ANSP A1093. Cuba: Marianas; M. C. & S. N. Rhoads leg; 1899; (12 specimens). ANSP A1094 (=85610). U.S.A: Florida, Miami; (10 spec- imens). ANSP A1096 (=91399). U.S.A: Florida, Mond Key; C. B. More. leg; (1 spec- imen). ANSP A1101 (=93280). U.S.A: Flor- ida, Sugar Loaf Key; H. A. Pilsbry; 1907; (1 specimen). ANSP Al1111. U.S.A: Florida, Dry Tortugas; T: Van Hyning leg; (2 speci- 528 Table 1.—Length, width and distance of the base to the greatest width for each of 3 first lateral teeth and 3 rachidians of Leidyula floridana, in .m. Laterals Rachidians Lots Length Width Distance Length Width Distance 1094B 49.28 39.93 34.02 33.32 16.93 19.46 49.10 41.28 35.24 29.91 17.36 20.67 54:04] 39'S S5598 S069 1712 22.73 Mean 50.80 40.34 34.95 31.30 17.13 20.95 805188 69.93 44.65 45.42 40.77 17.85 26.75 66:30) 46102) 427.22 -939:°71)) 1ia3 27-24 68.24 44.38 42.85 39.77 18.08 25.26 Mean 68.15 45.01 43.49 40.08 17.88 26.41 10944 “SIs eater Sow ly 428.87 422.24 2075 A932, ASNAy 31:53.) 29.85) 18.59" 18 71 SS OS— 4erso8 54-12 293067 19:00 19795 Mean 52.04 42.55 33.78 29.26 19.94 19.80 Overall means 56.99 42.63 37.40 33.54 18.31 22.38 mens). ANSP A1113 (=185244). U.S.A: Florida, St. Petersburg; (1 specimen). ANSP A1119. U.S.A: Florida, Paradise Key, (18 mi W. of Paradise Key - Dade Co. Florida); O. C. Van Hyning leg; 8 Jul 1930; (1 specimen). ANSP A1129 (=156474). U.S.A: Florida, Collier Co.; O. C. Van Hyning leg; 5 Aug 1930; (1 specimen). ANSP A1159. 12 Jul 1933; (1 specimen). ANSP A1163. U.S.A: Florida; 2 Jul 1936; (1 specimen). ANSP A1439. Puerto Rico: Humanaco, Baker; (5 specimens). ANSP A1448. Sierra Movalos; H. B. Baker; (1 specimen). ANSP A8020. U.S.A: Texas, Corpus Christi; G. M. Davis leg; Aug 1979; (Nueces Co., 27°43’N, 97°25'W) (1 specimen). ANSP A9345. U.S.A: Florida, Lignum Vitae Key; Mar 1931; (24°53'N, 80°42’W); (12 specimens). USNM 120394. U.S.A: Florida, H. H. Bun- ning leg; (3 specimens). USNM 574579. Puerto Rico: Mayaguez; M. Aviles leg; 23 Jun 1941; (8 specimens). USNM 576488. Dominica, at Clarke Hall; P Spangler leg; (USDA 249592); (2 specimens). USNM 769159. U.S.A: Texas, Brownsville; D. L. Dunagon leg; 27 Mar 1968; (USDA 278478); (2 specimens). USNM 769168. U.S.A: Texas, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Ratios between the averages of the radula teeth of Leidyula floridana. Ql= Q2= Q3= Q4= Q5= Q6= Lots LI:RI Lw:Rw LI:Ld RI:Rd Ll:Lw RI:Rw 1094B 1.62." 2335 1.45 1.49°- 1.25 1.82 $0518S7 1.70) 325i P65 esd 1 Sloe. 24 10944 LWT ORNS i:54. (47. 122546 Mean E69. 72233 1.51 LA9. 1.32 84 Ld = lateral distance; Ll = lateral length; Lw = lateral width; Rd = rachidian distance; Rl = rachidian length; Rw = rachidian width. Raymondville; R. Kloepping leg; 11 Apr 1968; (USDA 278478); (2 specimens). USNM 769195. U.S.A: Texas, Brownsville; O. V. Mullens leg; 2 Feb 1968; (USDA 273170); (4 specimens). USNM 769197. U.S.A: Texas, Brownsville; 2 May 1967; (USDA 266325) (2 specimens). USNM 805188. Cuba: El Guama; 10 Mar 1900; (1 specimen). USNM 805201. U.S.A: Florida, Monroe County, Pinecrest; J. P E. Morrison leg; 27 Feb 1966; (1 specimen). USNM 805202. U.S.A: Florida, Monroe County; Pi- necrest; ; hl mi W; J. P. E. Morrison leg; 27 Feb 1966; (2 specimens). Leidyula goodfriendi Thomé, 1994 Leidyula goodfriendi Thomé, 1994:169—72. Occurrence.—Antilles: Jamaica. Examined lots —ANSP A1154. 22 Jun 1933; (2 specimens). ANSP A1161. 1 Aug 1933; (1 specimen). Leidyula kraussi (Férussac 1823 in Férussac & Deshayes, 1820—1851) Vaginulus kraussii Férussac, 1823:96x in Férussac & Deshayes, 1820-1851. Leidyula kraussi; Thomé, 1990:513-18; 1993b:70. Occurrence.—Antilles: Haiti; Jamaica; U.S.A Examined lots —ANSP A?? (=62126). Jamaica: Bog Walk; J. Fox leg; 1891; (5 specimens). ANSP A1115. Jamaica: Port Antonio; Wm. J. Fox; 1891; (2 specimens). ANSP A1133 (=85633). Haiti; C. J. Simp- VOLUME 110, NUMBER 4 son leg; (1 specimen). ANSP A1153. 24 Aug 1933; (1 specimen). ANSP A1148 (=62130). Jamaica: Bog Walk; Wm. J. Fox.; 1891; (1 specimen). ANSP A1150. 25 Aug 1933; (3 specimens). ANSP A1152. 26 Jun 1933; (1 specimen). ANSP A1160. 7 Jul 1933; (1 specimen). Leidyula moreleti (Fischer 1871) Vaginula moreleti Fischer, 1871:168—69, #75: Leidyula moreleti; Thomé, 1971:32-—4, 1993b:71. Occurrence.—Colombia; Nicaragua; Honduras; Guatemala; Mexico; U.S.A: Florida; Antilles: Cuba. Examined lots ANSP A1054 (=63941). U.S.A: Florida, Cape Romano; (1 speci- men). ANSP A1066. Mexico: Vera Cruz; H. B. Baker; (Custotolapam farm); (43 speci- mens). ANSP A1107b (=45180). Guate- mala; A. A. Hinkley leg; 1914; (4 speci- mens). ANSP A1110. Cuba: Nueva Gerona, Isle of Pines; G. W. H. Soelnei leg; 18 Jun 1917; (3 specimens). ANSP A1112. Mexico: Mecos; A. A. Hinkley; (1 specimen). ANSP Al1117 (=45670). Guatemala: Livingston, A. A. Hinkley; (2 specimens). ANSP A1135 (=45188). Guatemala: Jacola; A. A. Hinkley leg; 12 Feb 1917; (3 specimens). ANSP A1146 (=61309). Mexico: Tabasco; (1 spec- imen). ANSP A9344. Mexico: Campeche, Peninsula Yucatan; 28 Jul 1932; (18°50’N, 91°20’W); (2 specimens). USNM 120395. Nicaragua: Polvon; W. G. Binney leg; 1890; (1 specimen). USNM 574589. Mexico; 12 Dec 1949; (intercepted at Brownsville, Tex- as, USDA 185542); (1 specimen). USNM 574940. Colombia; 1945; (on plants, USDA 202348); (1 specimen). USNM 575079. Co- lombia: Bello; C. Carmona leg; 10 Feb 1956; (on vegetables, USDA 209666); (3 specimens). USNM 575194. Mexico: San Luis Potosi, Tamazunchale; 30 Jul 1957; (in- tercepted in Laredo, Texas, on orchid plants, USDA 215961); (1 specimen). USNM 576004. Honduras; 27 Sep 1961; (in seed corn, USDA 241044); (7 specimens). 329 USNM 620158. Mexico: San Luis Potosi, Huichihuayan; (on orchid plants, USDA 212287); (1 specimen). USNM 769137. Mexico; 8 Apr 1967; (intercepted at Brownsville, Texas on orchids, USDA 266325); (1 specimen). Leidyula portoricensis (Semper 1885) Vaginula portoricensis Semper 1885:302 Leidyula portoricensis; Thomé 1975b:18; Thomé & Mansur 1990:534-35. Occurrence.—Antilles: Us A. Examined lots.—ANSP A1439. Porto Rico: Rio Humanaco; H. B. Baker; (2 spec- imens). ANSP A1442. Porto Rico: Monto- ro; H. B. Baker; (1 specimen). ANSP A1447. Sa Valuera; H. B. Baker; (2 speci- mens). Puerto Rico, Leidyula trichroma Baker, 1935 Veronicella (Leidyula) kraussii trichroma Backer, 1935:84, 88. Leidyula trichoma; Thomé, 1988c:32-33. Occurrence.—Antilles: Jamaica. Examined lots —ANSP A1058. Jamaica: Catadupa station, Montego Bay, St. James; H. B. Baker leg; (holotype); ANSP A12218. Same data (paratype). Phyllocaulis gayi (Fischer 1871) Vaginula gayi Fischer, 1871:172. Phyllocaulis gayi; Thomé, 1976:71-5; 1993b:70. Occurrence.—Chile; (introduced?: Mex- ico). Examined lot—USNM 805195. Chile: Valdivia; E. P. Reed leg; (2 specimens). Phyllocaulis soleiformis (Orbigny 1835) Vaginulus soleiformis Orbigny, 1835:2. Phyllocaulis soleiformis; Thomé, 1976:75— 6; 1993b:70. Occurrence.—Argentina; Uruguay; Bra- zil: Rio Grande do Sul. 530 Examined lots —ANSP A1085 (=69656). Uruguay: Maldonado; W. H. Rush leg; (1 specimen). USNM 576353. Argentina: 30 Km E of Azul; Lord and Jellison leg; May 1962; (USDA 245400); (1 specimen). Sarasinula dubia (Semper 1885) Vaginula dubia Semper, 1885:296; Thomé, 1972:252—53. Sarasinula dubia; Thomé, 1993b:71. Occurrence.—Brazil: Rio Grande do Sul, Sao Paulo; Venezuela; Colombia; Hondu- ras; Mexico; U.S.A: Florida, Texas; Antil- les: Trinidad and Tobago; Dominica, W.L.; Dominican Republic; Haiti; French Poly- nesia: Society Islands, Tahiti. Examined lots —USNM 769177. U.S.A: Texas, Raymondville; O. V. Mullens leg; 2 Feb 1968; (Willacy Co., Caldwell Nursery, USDA 273170); (3 specimens). USNM 574646. Dominican Republic: Santiago de los Caballeros; J. Jimines leg; 1951; (USDA 190012); (2 specimens). USNM 574676. Trinidad and Tobago: Trinidad, (British W. Indies); 1951; (USDA 191870); (4 specimens). USNM 575465. Haiti; 2 Dec 1958; (intercepted at New York, on Cala- dium plants, USDA 224155); (1 specimen). USNM 575897a. Honduras; 24 Jan 1961; (intercepted at Charleston on bananas, USDA 234701); (2 specimens). USNM 576001. U.S.A: Florida, Miami; 13 Sep 1961; (on the tree Persea americana, USDA 239104); (1 specimen). USNM 576337. Honduras: La Lima; 28 Oct 1958; (on a banana plantation, USDA 234081); (8 specimens). USNM 769138. U.S.A: Texas, Brownsville, (on with general nursery plants, USDA 273170); (10 specimens). USNM 769157. U.S.A: Texas, Brownsville; D. L. Dunagan leg; 16 Apr 1968; (Neal Robinson’s Nursery, USDA 278478); (7 specimens). USNM 769163. U.S.A: Texas, Raymondville; R. D. Kloepping leg; 11 Apr 1968; (Caldwell Nursery, USDA 278478); (3 specimens). USNM 769177. U.S.A: Tex- as, Raymondville; O. V. Mullens leg; 2 Feb 1968; (Willacy Co., Caldwell Nursery, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON USDA 273170); (3 specimens). USNM 769179. U.S.A: Texas, Brownsville; D. L. Dunagan leg; 26 Mar 1968; (Pan American Nursery, USDA 278478); (2 specimens). USNM 805184. Society Islands: Tahiti; 28 Sep 1899; (2 specimens). USNM 805204. Dominica; Portsmouth; 4 Aug 1929; (E side of watershed); (14 specimens). Sarasinula linguaeformis (Semper 1885) Vaginula linguaeformis Semper, 1885:307- 08; Thomé, 1973:35-6. Sarasinula linguaeformis; Thomé, 1993b: Tal, Occurrence.—Brazil: from Sao Paulo, Minas Gerais to the Northeast, Amazonas; Paraguay; Peru; Guyana. Examined lots—ANSP A7021. British Guiana: Georgetown, (6°46’N, 58°10’W); H. Lang; (specimens had been dried out and treated with tri-sodium phosphate, upgraded to 70% ETOH); (2 specimens). USNM 381391. Paraguay; Felippone leg; (4 spec- imens). USNM 769152. Peru; 17 Jan 1969; (intercepted at Miami on bromeliads, USDA 278478); (1 specimen). USNM 805252. Brazil: Lassance, MG; D. M. Cochran leg; 22 Mar 1935; (Marthyin Corn, USDA 132173); (6 specimens). Sarasinula plebeia (Fischer 1868) Vaginulus plebeius Fischer, 1868:145; Thomé, 1971:34—6. Sarasinula plebeia; Thomé, 1993b:71. Occurrence.—Chile; Brazil: from Rio Grande do Sul to Paraiba; Venezuela; Co- lombia; Costa Rica; El Salvador; Honduras; Guatemala; Mexico, U. S. A: Puerto Rico, Florida, Texas; Antilles: St. Lucia; Domin- ica, W. I.; Virgin Islands; Dominican Re- public; Jamaica; Cuba. Also cited to Aus- tralia; Asia; Africa. | Examined lots. —ANSP A9346. Vene- zuela: Carabobo State, Campo Alegre, (4 miles east of Caracas); (00021548, 10°35’N, 66°52’W, EDP, 21548); (26 spec- imens). USNM 574451. Venezuela: Sucre, VOLUME 110, NUMBER 4 Cariaco; G. Marcuzzi leg; Sep 1948; (USDA 180965); (3 specimens). USNM 574834. Virgin Islands: Saint Thomas; 1952; (USDA 195560); (2 specimens). USNM 574838. Virgin Islands: Saint John; 4 Aug 1952; (USDA 195744); (8 speci- mens). USNM 574942. Dominican Repub- lic; 7 Jan 1954; (USDA 201313); (1 spec- imen). USNM 575265. Costa Rica; 2 Oct 1957; (USDA 216763); (1 specimen). USNM 575366. Venezuela: Anzoatequi, San Tome; T. Briceno leg; (USDA 216763); (1 specimen). USNM 576341. Costa Rica; 1960; (USDA 231087); (1 specimen). USNM 578214. Costa Rica: Golfito; M. Kosztarali leg; 21 Feb 1970; (USDA 295099); (1 specimen). USNM 768944. Nicaragua; 10 Nov 1969; (intercepted at Miami on bromeliads, USDA 283854); (1 specimen). USNM 768945. El Salvador: Santa Tecla; 3 Jun 1969; (intercepted at Mi- ami on weeds, USDA 283854); (4 speci- mens). USNM 768946. Mexico; V. C. Fort leg; 23 Oct 1969; (intercepted at Fort La- redo, Texas, on orchids); (2 specimens). USNM 769135. El Salvador; 8 Jan 1968; (intercepted at New Orleans on orchids, USDA 273170); (1 specimen). USNM 769143. Puerto Rico: Bayamon; 16 Sep 1966; (intercepted at San Juan on Saint Au- gustine grass, USDA 266325); (1 speci- men). USNM 769156. Puerto Rico: Cupey Alto; 23 Oct 1968; (intercepted at San Juan on Dossaena leaf, USDA 278478); (1 spec- imen). USNM 769186. U.S.A: Texas; W. J. Bone leg; 8 Feb 1968; (McAllen, Hidalgo Co., Waugh Nursery, USDA 273170); (1 specimen). USNM 805203. Honduras; H. E. Ostmark and C. Evers leg; 13 Aug 1962; (on bananas, USDA 243607); (5 speci- mens). USNM 805204. Dominica: Ports- mouth; 4 Aug 1929; (E side of watershed); (14 specimens). USNM 805235. Unknown (St. Lucia?); 10 Aug 1929; (1 specimen). Simrothula columbiana (Simroth 1914) Vaginula columbiana Simroth, 1914:300-— 03; Thomé, 1970a:76—8; 1979:144—45. 531 Simrothula columbiana; Thomé, 1993b:72. Occurrence.—Colombia. Examined lot.-—ANSP A1441. (?) Cabo Rojo, (R33, Cabo Rojo); H. B. Baker; (1 specimen). Simrothula prismatica (Simroth 1914) Vaginula prismatica Simroth, 1914:314— 16; Thomé, 1972:261-63. Simrothula prismatica; Thomé, 1993b:72. Occurrence.—Colombia. Examined lot-—USNM 575078. Colom- bia: Bello. C. Carmona; 10 Feb 1956; (USDA 209666); (2 specimens). Vaginulus taunaisii Férussac, 1821 Vaginulus taunaisii Férussac, 1821:13. Vaginulus taunayi; Thomé, 1971:36—9; £993b:72: Occurrence.—Brazil: Rio de Janeiro. Examined lot—USNM 805194. Brazil: Rio de Janeiro, Niemeyer Avenue; 7 Feb 1935; (1 Specimen). Veronicella bahamensis Dall, 1905 Veronicella schivelyae bahamensis Dall, 1905:446. Veronicella bahamensis; Thomé & Lopes- Pitoni, 1976:712—13; Thomé, 1993b:72. Occurrence.—Bahamas. Examined lot—USNM 180415. Baha- mas: Little Abaco; O. Bryant leg; (lectoty- pe); USNM_ 180738. Bahamas: Nassau, Johnson Place; O. Bryant leg; 6 Jul 1904; (3 specimens). Veronicella cubensis (Pfeiffer 1840) Onchidium cubense Pfeiffer, 1840:250. Veronicella cubensis; Thomé, 1993a:113- 21; 1993b:70 Occurrence.—Antilles: Dominica, W.L.; Dominican Republic; Haiti; Cuba; Baha- mas; U.S.A. Puerto Rico, Louisiana, Ha- wail. 5 by PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Examined lots—ANSP A1083. Cuba: Havana, La Salud; H. A. Pilsbry; 1904; (5 specimens). ANSP A1443. Araciba river; H. B. Baker; (4 specimens). ANSP A1446. Catano; H. B. Baker; (1 specimen). ANSP A1741. Cuba: Punto Angre; (2 specimens). USNM 575898. Bahamas: Andros Island; 11 Feb 1961; QGntercepted at Port Ever- glades, on bananas plantation); (1 speci- men). USNM 576360. Puerto Rico; 17 Oct 1962; (on plants); (1 specimen). USNM 709075. Dominica; T. M. Clarke and J. E G. leg; 13 Jan 1965; (at Clarke Hall); (1 specimen). USNM 769160. U.S.A: Louisi- ana, New Orleans; 29 Jun 1970; (on side- walk near Tuoro Hospital); (1 specimen). USNM 805193. Haiti: Furcy; A. Curtiss; 20 Jun 1944; (with eggs); (1 specimen). Veronicella davisi Thomé, Santos & Pedott, 1996 Veronicella davisi Thomé, Santos & Pedott, 1996:265-—74. Occurrence.—Antilles: Jamaica; Cuba. Examined lots.—ANSP A1065a. Cuba: Majagua; H. A. Pilsbry (Holotype); ANSP A1065. Same data; (6 specimens) (Para- types); ANSP A1069. Cuba: Prov. Santa Clara, Zaza del Medio; H. A. Pilsbry; (6 specimens). ANSP A1072. Jamaica; J. B. Henderson; (1 specimen). ANSP A1073 (=137804). Cuba: Trinidad; H. A. Pilsbry; (5S specimens). ANSP A1075. Cuba, 15 Km South of Sancti Spiritus; H. A. Pilsbry; (2 specimens). ANSP A1077. (=137806). Cuba: Trinidad; H. A. Pilsbry; (1 speci- men). ANSP A1155. J. 5. Som.; 22 Jun 1933; (1 specimen). ANSP A1156. 20 Jul 1933; (1 specimen). ANSP A1157. 7 Jul 1933; (1 specimen). ANSP A1158. 22 Jun 1933; (1 specimen). ANSP A1748. J. 5. Som.; 22 Jun 1933; (2 specimens). Veronicella sloanei (Cuvier 1817) Onchidium sloanei Cuvier, 1817:411. Veronicella sloanei; Thomé, 1988a:13-—35; 1993b:70. Occurrence.—Colombia: Isla de Provi- dencia; Nicaragua; Bermudas; Bahamas; (U.S.A: Massachusetts—introduced?); An- tilles: Barbados; Jamaica; Cayman Islands. Examined lots—ANSP A1070. Grand Cayman; 17 Apr 1929; (Sta 4; Pinchat Ex- ped); (2 specimens). ANSP A1078. Isabel, Old Providence Id; 1929; (Pine Hot Exped); (1 specimen). ANSP A1104 (=57956). Nic- aragua: Machuca; Dr. J. E Bransford; (1 specimen). ANSP A1108. Bermuda: Fras- cati Hotel; A. Gulick; (3 specimens). ANSP A1109. Jamaica: Kingston Bath; T. D. A. Cockerell; (7 specimens). ANSP A1114. Ja- maica: Kingston; T. D. A. Cockerell; May 1892; (1 specimen). ANSP A1118. Jamai- ca: Kingston; T. D. A. Cockerell; (1 speci- men). ANSP A1120. Grand Cayman; 1929; (Pinchat Exped); (1 specimen). ANSP A1124. Jamaica: Kingston; T. D. A. Cock- erell; (2 specimens). ANSP A1125. Jamai- ca: Kingston; T. D. A. Cockerell; (1 speci- men). ANSP A1126. Jamaica: Kingston; T- D. A. Cockerell; (1 specimen). ANSP A1130 (=98631). Bermuda: near Harring- ton House; S. Brown; Jun 1909; (1 speci- men). ANSP A1144 (=65396). Jamaica; J. B. Henderson Jr.; (2 specimen). ANSP A1145. Jamaica: Port Antonio; W. J. Fox; 1891; (4 specimens). ANSP A1149 (=69360). Nantucket (introduced ?); Dr. B. Sharp; Aug 1896; (on bananas); (1 speci- men). ANSP A1437 (=60964). Bermudas: St. George; Heilprind Expedition leg; 1888. (Holotype of Vaginulus schivelyae Pilsbry, 1890). ANSP A1452. H. B. Baker; 5 Sep 1933; (3 specimens). ANSP A1455. H. B. Baker; 5 Jul 1933; (1 specimen). ANSP A1747. J. 33 .H.; 20 Aug 1933; (15 speci- mens). ANSP A1748. Bahamas; Mich. Goodrich; 21 Aug 1933; (1 specimen). USNM 574916. Jamaica; (British West In- dies); 24 Jul 1953; (intercepted at Miami, Florida, on orchids, USDA 199469); (1 specimen). USNM 791486. Jamaica; (1 specimen). USNM_ 805186. Jamaica: Kingston; C. R. Orcutt leg; 29 Aug 1927; (2 specimens). USNM 805190. Jamaica: Saint Thomas, Antully Parish; C. R. Orcutt VOLUME 110, NUMBER 4 leg; Aug 1927; (3 specimens). USNM 805192. Jamaica: Antully; C. R. Orcutt leg; (10 specimens). Veronicella tenax Baker, 1931 Veronicella (Tenacipes) tenax Baker, 1931: 131-34. Veronicella tenax; Thomé, 1988c:30-2; 1993b:72. Occurrence.—Antilles: Antigua and Bar- buda; Cuba. Examined lot.—ANSP A12221 (=291340a). Cuba: Ensefiada de San Vi- cente, Cueva de Tiburon, Pinar del Rio; H. P. Pilsbry leg; 1928; (Holotype). ANSP A12222 (=291340b). same data; (2 speci- mens); (Paratypes). ANSP A1164. Antigua; A. P. Brown; (5 specimens). Acknowledgments We thank Dr. George M. Davis, Chair- man of the Department of Malacology, The Academy of Natural Sciences of Philadel- phia and Dr. Robert Hershler, Associate Cu- rator of the Division of Mollusks, Depart- ment of Invertebrate Zoology, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, for kindly loaning us their collections for an extended period. We are grateful to: Falavena, Mansur, Martins, Fonseca, Montenegro and others, who helped us with the identification and illus- tration of some lots, while working in the Laboratory, as grantees; Cleodir José Man- san, of the Funda¢cao Zoobotanica do Rio Grande do Sul, for help with the scanning microscopy; and William Belton, for kindly revising the English. Literature Cited Baker, H. B. 1925. North American Veronicellidae.— Proceedings of the Academy of Natural Sci- ences of Philadelphia 77:157—184. . 1926. 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Os géneros da familia Veronicellidae nas Américas (Mollusca; Gastropoda).—lher- ingia, Porto Alegre, Série Zoologia (48):3—56. 1976. Reviséo do género Phyllocaulis Co- losi, 1922 (Mollusca; Veronicellidae).—Iherin- gia, Porto Alegre, Série Zoologia (49):67—90. . 1979. A redescription of Types of six Species of neotropical Veronicellidae (Mollusca; Gas- tropoda) in the British Museum (Natural His- tory).—Bulletin of the British Museum (Natural History), Zoology 35(2):135-147. . 1985. Redescrigaéo dos tipos de Veronicelli- dae (Mollusca; Gastropoda) neotropicais. X. Os tipos de Diplosolenodes occidentalis (Guilding, 1825) no British Museum (Natural History), Londres.—Revista Brasileira de Zoologia, Sao Paulo 2(6):411—417. . 1988a. Recaracterizagao de Veronicella slo- anei (Cuvier, 1817) (Mollusca; Gastropoda; Ve- ronicellidae).—lIheringia, Porto Alegre, Série Zoologia (67):13-35. 1988b. Lista das Veronicellidae (Mollusca; Gastropoda) da Jamaica.—Revista Brasileira de Biologia, Rio de Janeiro 48(4):807—810. . 1988c. Redescrigao dos tipos de Veronicel- lidae (Mollusca; Gastropoda) neotropicais: XI. Espécies na ““Academy of Natural Sciences’’, Philadelphia.—lIheringia, Porto Alegre, Série Zoologia (69):25—40. . 1990. Recaracterizagao de Leidyula kraussi (Férussac 1823) (Mollusca; Gastropoda; Veron- icellidae).—Revista Brasileira de Biologia, Rio de Janeiro 50(2):513-—518. . 1993a. Erneute Beschreibung von Veronicel- la cubensis (Pfeiffer 1840) (Gastropoda; Veron- icellidae).—Archiv fiir Molluskenkunde 122: 113-121. . 1993b. Estado atual da sistematica dos Ve- ronicellidae (Mollusca; Gastropoda) america- nos, com comentarios sobre sua importancia econdmica, ambiental e na saide.—Biociéncias, Porto Alegre 1(1):61-—75. . (1993) 1994. Nova Leidyula Baker da Ja- maica (Mollusca; Soleolifera; Veronicelli- dae).—Revista Brasileira de Zoologia, SAo Pau- lo 10(1):169-172. , & V.L.R. Lopes. 1973. Aulas Praticas de Zoologia, I. Dissecagao de um molusco gastr6- pode desprovido de concha.—lheringia, Porto Alegre, série Divulgagao (3):34—45. , & V.L. Lopes-Pitoni. 1976. Redescri¢gao dos tipos de Veronicellidae (Mollusca; Gastropoda): espécies no “‘National Museum of Natural His- tory, Smithsonian Institution’’, Washington, E.U.A.—Revista Brasileira de Biologia, Rio de Janeiro 36(3):709-714. 535 , & G. G. Mansur. 1990. Recaracterizacao de Leidyula portoricensis (Semper, 1885) (Veron- icellidae; Gastropoda). In: REUNIAO ANUAL DA SBPC, 42, 08/13-07-90, Porto Alegre. An- ais (Communicagées)... SBPC, Sao Paulo 567pp. pp. 534-35. (Suplemento de Ciéncia e Cultura). , P. H. dos Santos, & L. Pedott. (1995) 1996. Nova espécie de Veronicella Blainville, 1817 (Mollusca; Gastropoda; Soleolifera) da América Central—Biociéncias, Porto Alegre 3(2):265-— 274. Zilch, A. 1959/1960. Gastropoda, Teil 2: Euthyneura. In: Schindewolf, O. H. (org.). Handbuch der Pa- laozoologie. Gebriider Borntrager,. Berlin, 6: 834pp. Appendix On the spelling and date of Vaginulus taunaisii Fér- ussac, 1821 (Veronicellidae; Soleolifera; Gastropoda; Mollusca).—A number of differences continue to exist among malacologists regarding the spelling and date of the name proposed more than a century ago by the Frenchmen, d’Audebard de Férussac (father and son). Even now, through correspondence, an argument on the subject has been put forth by a well-known col- league. I do not pretend to have found the full solution, but I hope to contribute to clarification of the contro- versy with a view to achieving a consensus. The generic name Vaginulus was proposed by Fér- ussac in 1821 (p. 6,9,13). In my opinion it is a valid name, with correct, latinized, original spelling, al- though use of the masculine suffix (from the Latin: vagina + ulus, ula, ulum = diminutive) (Jaeger 1978: 275) may be considered unsuitable, but not wrong. The spelling is in conformity with the International Code of Zoological Nomenclature (ICZN) 1985, art. 11(b)G), 32 (c)(i1), table 2 on P. 207. To replace Vaginulus, the generic name Vaginula was proposed by Berthold, 1827, according to Neave (1940:625). Fischer (1871:148) also suggested chang- ing it due to error (fautif) and among others, Deshayes (1930) himself, in Férussac & Deshayes 1820-1851: 96° and even Hoffman (1925:172) insisted on the sub- stitution, without justifying their position. Neverthe- less, it should be considered an unjustified emendation (ICZN 1985, art. 33 (b) (iii) and, therefore, Vaginula Berthold, 1827 (non Vaginula Sowerby, 1820 (Hoff- mann, 1925:172); Vaginula Risso, 1826 (Neave 1940: 625) must be considered a junior objective synonym. The specific name taunaisii (Férussac 1821:13, and Férussac 1821 and 1823, in Férussac & Deshayes 1820-1851, pl. 8A, fig. 7, pl. 8B, fig. 1-2, and p. 96%") is said to be an homage to the Frenchman, Taunay (son), whose modern patronym appears to have been erroneously latinized. Fischer (1871:170) and Hoffman (1925:173), amongs others, have already proposed the correct spelling as taunayi (ICZN 1985, art. 11, (h) (i) 536 (3), 31 (a) Gi) and (ii)). The original species name, taunaisii, has been used by very few authors, the most recent of these being Martens (1867). In spite of this, taking into consideration the opinions of editors and consultants, and based on art. 32 (c) (ai) of the ICZN (1985), I propose a return to the original spelling of the specific name, with the specific name taunayi be- coming an “incorrect subsequent spelling”’ (art. 33 (c) ICZN 1985). Regarding the date of the confusing publication of the Férussacs, the most solid work of dating by exter- nal evidence must be that of Kennard (1942a, 1942b), who accepts the date of 17 July 1822 for the pages containing the original description of Vaginulus and V. taunaisii. Nevertheless, the date on the cover of the Férussac work is January 1821 (janvier 1821) and in the second edition of the great work of Férussac & Deshayes (1820-1851), page 96! (edited by Deshayes PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON in 1830) is it expressly stated that the disputed work of Férussac was published in June of 1821 (juin 1821), and therefore distributed with six months delay, but not more than a year and a half, as Kennard figured. Also Herrmannsen (1849), Fischer (1871), and Hoff- mann (1925), among others, mention the date of 1821 as valid for the taxa in question, which also appears to me to be valid external evidence. The species Vaginulus taunaisii (=taunayi) Férus- sac, 1821 has been well redescribed (Thomé 1971:36— 39, 41, fig, 15-17, 22), and the genus recharacterized (Thomé 1975b:7-9, 28-32), which should now be con- sidered monotypic. Thus the argument has little sig- nificance for the stability and universality of the taxa, for there are no taxonomic implications for their de- termination and classification. This is also true with reference to the maintenance of the family name, which should be Veronicellidae, as has been demon- strated by Thomé (1979:138—-141). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):537-551. 1997. Revision of the scaleworm genus Eulagisca McIntosh (Polychaeta: Polynoidae) with the erection of the subfamily Eulagiscinae and the new genus Pareulagisca Marian H. Pettibone Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—A new subfamily of Polynoidae, Eulagiscinae, is established for Eulagisca McIntosh, to include the type species E. corrientis, E. gigantea Mon- ro, E. puschkini Averincev, and two new species, E. uschakovi and E. macnabi, all from the Antarctic and Subantarctic regions, and the new genus Pareula- gisca for P. panamensis (Hartman), new combination, from the central Pacific Panama region. McIntosh (1885) described Eulagisca corrientis from deep water in the South At- lantic and South Indian Oceans. Monro (1939) described some large polynoids from the deep Antarctic as E. gigantea. Hartman (1939) added E. panamensis from shallow water of Pacific Panama, and Av- erincev (1972) added E. puschkini from Shallow water in the Antarctic Davis Sea. The prostomia resemble those of the mem- bers of the subfamilies Lepidonotinae Wil- ley, 1902 and Lepidastheniinae Pettibone, 1989, by having the lateral antennae insert- ed on anterior extensions, on the same level as the ceratophore of the median antenna, or subterminally, without distinct cerato- phores. The parapodia differ from those of both subfamilies by having the notopodia and neuropodia well developed, with pro- jecting digitiform acicular lobes, rather than projecting acicular lobes lacking. The elytra of some specimens, identified as E. gigan- tea, and some new material, showed some differences and are referred to two new spe- cies: E. uschakovi and E. macnabi. Some distinctive characters of species of Eulagisca, and not shown by E. panamen- sis, referred herein to the new genus Pareu- lagisca, include the following: long palps of the prostomium with six longitudinal rows of papillae; prominent conical facial tubercle on the upper lip, ventral to the cera- tophore of the median antenna; tentaculo- phores of segment I, lateral to the prosto- mium, with projecting digitiform acicular lobe and group of long setae; and 15 pairs of elytra (16 pairs in P. panamensis). In addition to the collections in the Na- tional Museum of Natural History, Smith- sonian Institution (USNM), the specimens covered herein are deposited in the follow- ing Museums: Natural History Museum, London (BMNH); Natural History Museum of Los Angeles County (LACM-AHF): Zoological Institute Academy of Sciences, Leningrad (ZIASL); Zoological Museum, Hamburg (ZMH): Zoologisk Museum, Universitet i Oslo (ZMUO). Family Polynoidae Kinberg, 1856 Eulagiscinae, new subfamily Diagnosis.—Body elongate, with seg- ments moderate in number (up to 41). El- ytra and bulbous elytrophores 15 (Eulagis- ca) or 16 (Pareulagisca) pairs, on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, 32 (33). Elytra large, oval. with or without papillae, microtubercles, macrotubercles and globular vesicles or papillae. Dorsal cirri, on non-elytrigerous segments, with 538 cylindrical cirrophores posterodorsal to no- topodia, with distal styles; dorsal tubercles, in line with elytrophores, bulbous or indis- tinct. Prostomium bilobed, with paired palps and three antennae; median antenna with large ceratophore in anterior notch of prostomium, with distal style; lateral anten- nae inserted terminally or subterminally on anterior extensions of prostomium, without distinct ceratophores; two pairs of eyes. First or tentacular segment not visible dor- sally; tentaculophores lateral to prostomi- um, each with dorsal and ventral tentacular cirri, similar to median antenna, with or without digitiform acicular lobe and nu- merous or single setae; with or without dis- tinct facial tubercle on upper lip. Second or buccal segment with nuchal fold, first pair of elytra and elytrophores, biramous para- podia, and ventral buccal cirri, longer than following ventral cirri. Biramous parapodia with both noto- and neuropodia with prom- inent projecting digitiform acicular lobes and numerous noto- and neurosetae. Noto- setae of one kind, stout (Eulagisca) or two kinds: stout and slender, capillary (Pareu- lagisca). Ventral cirri short, tapering. Phar- ynx with two pairs of jaws and eleven pairs of dorsal and ventral papillae (not examined in Pareulagisca). Pygidium with pair of anal cirri. Remarks.—The prostomia of the Eulag- iscinae agree with the Lepidastheniinae and the Lepidonotinae in having the lateral an- tennae inserted terminally or subterminally, without distinct ceratophores, and disagree- ing with the Harmothoinae having distinct ceratophores and inserted ventrally. The nu- chal folds on the second or buccal segment in Eulagiscinae are distinctive. The bira- mous parapodia, with both rami having prominent digitiform acicular lobes and nu- merous noto- and neurosetae in Eulagisci- nae, differ from the other subfamilies. Key to the Genera and Species of the Subfamily Eulagiscinae 1. Elytra 16 pairs, on segments 2, 4, 5, 7, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON alternate segments to 23, 26, 29, 32, 33. Elytra without tubercles and border or surface papillae (Fig. 5G). Prostomium with long palps, without 6 longitudinal rows of papillae (Fig. 5A). Tentaculo- phore of segment I without projecting acicular lobe, with single seta (Fig. 5A); facial tubercle weakly developed. Noto- setae of two kinds: 1) stout, acicular, smooth or with faint spinous rows (Fig. 5D); and 2) slender, capillary, smooth or with few spines (Fig. SE)... 2.2 sae Pareulagisca, n.gen.: P. panamensis (Hartman 1939), n. comb. (Fig. SA—G) — Elytra 15 pairs, on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, 32. El- ytra variable. Prostomium with long palps with 6 longitudinal rows of papil- lae (Fig. 2A) Tentaculophores of seg- ment I with projecting digitiform acicu- lar lobe and group of long setae; with prominent conical facial tubercle (Fig. 2A). Notosetae of one kind, stout, with long spinous regions, tapering to blunt tips" (Figs. iC; '2C,;'3C) 4D) t. 7 ADS Ee Eulagisca McIntosh 1885. ...... 2 2. Neurosetae tapering to sharp tips (Fig. 3D; Averincev 1972,..pl: 13:9—11)5 saa — Neurosetae tapering to slightly hooked tips (Figs. 1D, 2D, 4E) 3. Elytra with sharply pointed tubercles and globular vesicles with few to numerous terminal spines (Figo3E)) 0. 2 ee ... E. uschakovi, new species (Fig. 3A—E). — Elytra without border or surface papillae; with numerous microtubercles on anterior part (Averincev 19772, pl. 13:3): Soe .. E. puschkini Averincev, 1972 (pl. 13:8—12). 4. Elytra with border papillae (Figs. 1E, 4F) — Elytra without border papillae; surfaces with conical microtubercles, larger conical tubercles, and globular vesicles (Fig. 2F) .... E. gigantea Monro, 1939 (Fig. 2A—P) 5. Surface of elytra with conical papillae on anterior part (Fig. 1E) . E. corrientis McIntosh, 1885 (Fig. 1A—E) — Surface of elytra with conical microtuber- cles and larger spiny tubercles (Fig. 4F) .... E. macnabi, new species (Fig. 4A—F). Genus Eulagisca McIntosh, 1885 Type species.—Eulagisca corrientis Mc- Intosh, 1885, by monotypy. Gender, femi- nine. ee © © © © © © 8 ee ee VOLUME 110, NUMBER 4 Diagnosis.—Body rather large, wide, flattened, slightly tapering anteriorly and more so posteriorly, with segments up to 41. Elytra and bulbous elytrophores 15 pairs, on segments 2, 4, 5, 7, alternate seg- ments to 23, 26, 29, 32. Elytra large, cov- ering dorsum, easily detached, with or with- out microtubercles, macrotubercles, bul- bous vescicles, and fringes of papillae. Dor- sal cirri with cylindrical cirrophores and long styles on non-elytrigerous segments; dorsal tubercles bulbous. Prostomium bi- lobed, with paired long palps each with six longitudinal rows of papillae, and three an- tennae; median antenna with ceratophore in anterior notch of prostomium, with long style; lateral antennae inserted on anterior extensions of prostomium, on same level as median antenna, without distinct cerato- phores; eyes 2 pairs, large, laterally on mid- dle of prostomium, nearly contiguous. First or tentacular segment indistinct dorsally; tentaculophores lateral to prostomium, each with projecting digitiform acicular lobe, group of setae, and long dorsal and ventral tentacular cirri. Prominent conical facial tu- bercle ventral to ceratophore of median an- tenna. Second segment with semilunar nu- chal pad and nuchal fold covering posterior part of prostomium, first pair of elytro- phores, biramous parapodia, and long ven- tral buccal cirri lateral to ventral mouth. Parapodia biramous, with numerous golden setae. Notopodia well developed, with prominent digitiform acicular lobes; neu- ropodia with long digitiform presetal acic- ular lobes and shorter rounded postsetal lobes. Notosetae numerous, stouter than neurosetae, with long spinous regions, ta- pering to blunt tips; neurosetae more nu- merous and finer than notosetae, with long spinous regions, tapering to rather long bare tips. Ventral cirri short, subulate. Pharynx with two pairs of chitinous jaws and 11 pairs of dorsal and ventral papillae. Pygid- ium with pair of anal cirri. Nephridial pa- pillae short, cylindrical, beginning on seg- ment VI. Remarks.—As indicated by Benham 539 (1921:45), McIntosh’s choice of Eulagisca for E. corrientis was a poor one, since it differs in so many ways from Lagisca, in- cluding the type of the prostomium, para- podia, and elytra. Eulagisca corrientis McIntosh, 1885 Fig. 1 Eulagisca corrientis McIntosh, 1885:91, Fig. 1, pl. 13:fig. 4, pl. 7A:figs. 3, 4, Not Eulagisca corrientis—Benham, 1921: 43, pl. 6:figs. 36-38, pl. 7:figs. 39-42. [?] Eulagisca corrientis.—Monro, 1930:48, fig. 1la—e (no elytra); 1936:90 (no ely- tra). Not Harmothoe corrientis (?).—Augener, 1932:19. (= Harmothoe sp.). Not Eulagisca corrientis.—Stiller, 1996:30, pl. 4:figs. 1-5. Material examined.—South Atlantic: off east coast of South America, south of Buen- as Aires, 37°17'S, 53°52'W, 1097 m, green sand, Challenger Sta. 320, 14 Feb 1876, syntype (BMNH 1885.12.1.71). South In- dian Ocean: Between Kerguelen and Heard Islands, 52°04’S, 71°22'E, 274 m, coarse gravel, Challenger Sta. 150, 2 Feb 1874, syntype (BMNH 1885.12.1.83). Remarks.—The species is somewhat doubtful, since it was based on two syn- types, a smaller complete specimen but without elytra, from Challenger Sta. 320, and an anterior fragment of a larger speci- men with two elytra, from Sta. 150. There is a possibility that the two syntypes belong to different species but the type material does not allow one to clarify this point. Monro (1930:48) examined MclIntosh’s type specimens and referred some speci- mens from off the South Shetlands to E. corrientis, but without elytra and thus doubtful. Monro (1936:90) added a record from off the Falkland Islands, but the spec- imen also lacked elytra. Stiller (1996:30) added E. corrientis from the eastern Wed- dell Sea. It does not agree with the two syn- types in the BMNH. The specimens described as E. corrientis 540 by Benham (1921:43) appear to be a mix- ture of Eulagisca species and need to be reexamined. The specimen that Augener (1932:19) questionably referred to Har- mothoe corrientis was examined from the Olso Museum (ZMUO); it is in poor con- dition, the prostomium is harmothoid and different from that of Eulagisca; it is re- ferred to Harmothoe sp. Description.—Incomplete syntype with 19 segments, 29+ mm long and 24 mm wide with setae. Smaller complete syntype with 35 segments, 34 mm long and 16 mm wide with setae. Dorsum with brown pig- mentation. Elytra and bulbous elytrophores 15 pairs, in usual arrangement (McIntosh 1885: pl. 13, fig. 4). Elytra subreniform, surface with soft conical papillae and long delicate clavate papillae near posterior and lateral borders (Fig. 1E). Dorsal cirri with cylindrical cirrophores and papillate styles extending beyond setae; dorsal tubercles nodular. Bilobed prostomium wider than long, with long papillate palps and three anten- nae; median antenna with large ceratophore in anterior notch, with style missing; lateral antennae inserted on anterior extensions of prostomium, with rather long styles with subterminal enlargements and filamentous tips; two pairs of large closely approximat- ed eyes on lateral sides; tentaculophores lat- eral to palps and prostomium, each with projecting digitiform acicular lobe and four setae directed anteriorly, and dorsal and ventral tentacular cirri, similar to lateral an- tennae; prominent conical facial tubercle ventral to ceratophore of median antenna (Fig. 1A; McIntosh 1885: pl. 13, fig. 4). Segment II with large semilunar nuchal fold, first pair of large elytrophores, bira- mous parapodia and long ventral buccal cir- ri (Fig. 1A; McIntosh: pl. 13 fig. 4). Biramous parapodia with notopodia and neuropodia about equal in size, both with projecting acicular lobes; notopodia round- ed basally, with projecting acicular lobes on lower sides; neuropodia subconical with projecting presetal acicular lobes and short- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON er rounded postsetal lobes (Fig. 1B; McIn- tosh 1885: fig. 1). Notosetae numerous, forming radiating bundle of four lengths, slightly stouter than neurosetae, short, slightly curved and longer, straight, with long spinous regions, tapering to short bare tips (Fig. 1C; McIntosh 1885: pl. 7A fig. 3). Neurosetae numerous, forming fan- shaped bundle, with long spinous regions, tapering to slightly hooked bare tips; upper ones more slender and upper few with del- icate secondary tooth (Fig. 1D; McIntosh 1885: pl. 7A fig. 4). Ventral cirri slender, with few papillae, tapering, extending slightly beyond basal part of neuropodia (McIntosh 1885: fig. 1). Distribution.—South Atlantic and South Indian Oceans, in 274—1097 meters. Eulagisca gigantea Monro, 1939 Fig. 2 Eulagisca gigantea Monro, 1939:103 (part), Fig. 4a—d, f-—h (not e), Sta. 30 only.—Uschakov, 1962:174 (part), Sta. 283, pl. 9E, G, H (mot F); 1966:174 (part), Sta. 282, pl. 9E, G, H (not F) (En- glish translation).—Hartman, 1967:21 (part, not Sta. 1003).—Not Averincey, 1972:131 (=E. uschakovi, new spe- cies).—Not Desbruyéres, 1976:85, pl. 1: figs. a-c (=Eulagisca, new species?).— Hartmann-Schroder & Rosenfeldt, 1990: 91 (part), pl. 1:3—5 (not 1, 2).—Not Still- er, 1996:30, pl. 10:figs. 1-5. (=E. us- chakovi, new species). Material examined.—Antarctic: Off Princess Elizabeth Land, 66°48'S, 71°24’E, 540 m, British, Australian, New Zealand, Antarctic Research Expedition Sta. 30, 27 Dec 1929, lectotype (BMNH 1941.3.3.16, largest of three syntypes of E. gigantea). Drake Passage, between South America and Antarctica, 56°06’S, 66°19'W, 384—494 m, Eltanin Sta. 740, 18 Sep 1963, 1 spec- imen (USNM 57466, ident. by Hartman 1967). South Shetland Islands, 61°25’S, 56°30'W, 300 m, Eltanin Sta. 993, 13 Mar 1964, 1 specimen (USNM 57467, ident. by VOLUME 110, NUMBER 4 Fig. *. 541 Eulagisca corrientis, A-D, syntype from Sta. 320 (BMNH 1885.12.1.71): E, syntype from Sta. 150 (BMNH 1885.12.1.83): A, Dorsal view of anterior end, right tentaculophore damaged; missing: styles of median and lateral antennae, palps, and tentacular cirri; B, Right elytrigerous parapodium, anterior view, aciculae dotted, style of ventral cirrus missing; C, Tips of long and short notosetae; D, Middle and upper neurosetae, with detail of tip, E, Right elytron, with detail of papillae. Scales = 1.0 mm for A; 0.5 mm for B; 0.1 mm for C,D; E (not to scale). Hartman 1967). South Orkney Islands, 60°50'S, 42°55’ W, 298-302 m, Eltanin Sta. 1082, 14 Apr 1964, 1 specimen (USNM 57469, ident. by Hartman 1967). 59°53’S, 32°19'W, 523-671 m, Islas Orcada Cruise 1876, USARP Sta. 130, 25 Feb. 1976, 1 specimen (USNM 74575). Remarks.—The type material of Eula- gisca gigantea Monro (1939) consists of three syntypes. They proved to be a mixture of at least two species. The largest syntype from Sta. 30, with the elytral tubercles and vesicles as on Fig. 4c, d and not Fig. 4e by Monro (1939), was selected as the lectoty- pe. The next largest syntype from Sta. 107, with elytral tubercles as shown on Fig. 4e, was selected as the holotype of E uschak- ovi, new species. The smallest syntype from Sta. 39, which lacked elytra, was selected as a paratype of E. uschakovi, based on the similarity of the neurosetae. The two specimens from the Continental plateau off Kerguelen, in 172—262 m, de- scribed and figured by Desbruyéres (1976) 542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON as E. gigantea, differ: the elytra have a well-developed lateral fringe of long papil- lae and the surface is covered with conical spiny vesicles (pl. 1A—C), thus differing from E. gigantea, and probably indicating a new species. The specimens were not available for examination to complete the description. One of the specimens from Elephant Is- land, Antarctica, in 280—432 m, referred to E gigantea by Hartmann-Schréder and Ro- senfeldt (1990:91, pl. 1; 3-5, not 1, 2), agree with E. gigantea, based on the photos of the elytra; the other specimen is referred to E. uschakovi, new species. Stiller (1996: 30, pl. 10:figs. 1-5) reported E. gigantea from the eastern Weddell Sea. Based on the description and figures, it is added to E. us- chakovi, new species. Additional records of E. gigantea, with elytra lacking, are omitted. They must be referred to Eulagisca sp. Description.—Lectotype with 38 seg- ments, 148 mm long, 44 mm wide with se- tae. Specimen from South Shetland Island (USNM 57467) 68 mm long, 46 mm wide with setae, 38 segments. Specimen from South Orkney Island (USNM 57469) 155 mm long, 94 mm wide with setae, 38 seg- ments. Body large, flattened, with long par- apodia (parapodia and setae longer than body width). Body colorless except for trac- es of brown transverse bands, with golden setae. Fifteen pairs of elytra, on bulbous ely- trophores, large, thick, subreniform, cover- ing dorsum, darker along posterior border, without border papillae (Fig. 2F; Monro 1939: fig. 4b). Surface of anterior part of elytra with numerous conical microtuber- cles, middle part with larger sharply pointed conical tubercles and posterior part with curved conical tubercles (rarely 2-pronged), some smaller spines, and soft globular ves- icles, dotted yellowish distally (Fig. 2F; Monro 1939: fig. 4c, d, not e; Uschakov, 1966: pl. 9E, not F). Dorsal cirri with cy- lindrical cirrophores, bulbous on posterior basal parts, with styles extending to end of neurosetae, with short clavate papillae. Dor- sal tubercles bulbous, posterior ones on seg- ments 33—38 projecting posteriorly. Bilobed prostomium wider than long, median antenna with large ceratophore in anterior notch, with style missing (probably long, similar to but longer than lateral an- tennae); lateral antennae inserted on ante- rior extensions of prostomium, without dis- tinct ceratophores, with styles rather long, finely papillated, with subterminal enlarge- ments and filamentous tips; palps long, Stout, tapering, with six longitudinal rows of papillae; two pairs of rather large eyes with lenses on posterior half of prostomi- um; tentaculophores (segment I) lateral and anterior to prostomium, with prominent digitiform acicular lobes, group of long se- tae (up to 10) directed anteriorly, and long dorsal and ventral tentacular cirri, similar to but longer than lateral antennae; with prom- inent conical facial tubercle below cerato- phore of median antenna (=subtentacular cirrus by Monro) (Fig. 2A; Monro 1939: fig. 4a). Segment II with prominent semi- lunar nuchal pad and conical fleshy nuchal fold covering posterior part of prostomium (latter may be pulled back), first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri, inserted basally, similar to tentacular cirri (Fig. 2A; Monro 1939: fig. 4a). Biramous parapodia with small subconi- cal notopodia with projecting acicular lobes; larger neuropodia with subtriangular presetal acicular lobes and shorter subcon- ical postsetal lobes (Fig. 2B; Monro 1939: fig. 4f). Notosetae numerous, forming ra- diating bundle, about as stout as neurosetae, of three lengths, some shorter, curved, some about as long as neurosetae, stout, acicular, with long spinous regions, tapering to bare entire tips (Fig. 2B, C; Monro 1939: fig. 4g; Uschakov 1962: pl. 9G). Neurosetae nu- merous, forming fan-shaped bundle, with long spinous regions and rather long, slight- ly hooked entire bare tips; upper ones more slender, with longer spinous regions (Fig. 2D, E; Monro 1939: fig. 4h; Uschakov VOLUME 110, NUMBER 4 543 Fig. 2. Eulagisca gigantea, A. Specimen from South Shetland Island (USNM 57467); B-E lectotype (BMNH 1941.3.3.16): A, Dorsal view of anterior end, pharynx partially extended; missing; right palp, median and right lateral antennae, right dorsal and ventral tentacular cirri, left parapodium of segment 2 not shown; B, Right elytrigerous parapodium, anterior view; C, Notoseta from same; D, Middle neuroseta from same; E, Upper neuroseta from same; FE Elytron, with detail of microtubercles, larger tubercles, and soft globular vesicles. Scales = 2.0 mm for A; 2.0 mm for B; 0.1 mm for C—E; 5.0 mm for FE 1966: pl. 9H). Ventral cirri short, subulate, 1962:174 (part: Sta. 232, pl. 9F only; extending slightly beyond basal part of neu- 1966:174 (part: Sta. 232, pl. 9F only (En- ropodia (Fig. 2B). glish translation).—Hartman, 1967:21 Distribution.—Antarctic, Off Princess (part: Sta. 1003 only).—Averincev, 1972: Elizabeth Land, Davis Sea, Drake Passage, 131, pl. 13:1—-7.—Hartmann-Schréder & Scotia Sea, South Shetlands, South Ork- KRosenteiat, 1990:91) (part, pl. 171, 2, not neys, in 200—900 meters. 3-5). Not Monro, 1939 (lectotype).— Stiller, 1996:30, pl. 10: figs. 1-5. Fic. 3 Material examined.—Antarctic: Off a? MacRobertson Land, 66°45’S, 62°63’E, 219 Eulagisca gigantea Monro, 1939:103 (part: m, British, Australian, New Zealand Ant- Sta. 39 & 107, Fig. 4e only).—Uschakov, arctic Research Expedition Sta. 107, 16 Feb Eulagisca uschakovi, new species 544 1931, holotype (BMNH 1941.3.3.18, as next longest syntype of E. gigantea). Off Enderby Land, 66°10’S, 49°41’E, 300 m, B.A.N.A.R.E. Sta. 39, 17 Feb 1930, para- type (BMNH 1941.3.3.17, as smallest syn- type of E. gigantea). Bransfield Strait, 62°41'S, 54°43’W, 210- 220 m, Eltanin Sta. 1003, 15 Mar 1964, paratype (USNM 57468, as E. gigantea by Hartman, 1967). Off Palmer Archipelago, 62°09'S, 57°49’'W, 220-229 m, Eastwind Sta. 66, 19 Feb 1966, 2 paratypes (USNM 35289). Scotia Sea, 61°19’S, 44°26’W, 274— 280 m, Islas Orcada Sta. 123, 22 Feb 1976, 1 specimen (USNM 74576). Ross Sea, 75°50'S, 173°08’W, 476 m, EI- tanin Sta. 38, 31 Jan 1968, 2 paratypes (USNM 58350). Types.—Holotype (BMNH 1941.3.318, next largest syntype of E. gigantea): 110 mm long, 70 mm wide with setae, 39 seg- ments, with elytra. Paratype (BMNH 1941.3.3.17, smallest syntype of E. gigan- tea): 56 mm long, 36 mm wide with setae, 37 segments, without elytra but neurosetae agree with holotype. Two paratypes from Ross Sea (USNM 58350): 140-190 mm long, 85-100 mm wide with setae, 39 seg- ments, with elytra. Paratype from Palmer Archipelago (USNM 58351): 81 mm long, 48 mm wide with setae, 38 segments, with elytra (figured). Description.—Body brownish on mid- dorsum, on bases of elytrophores and dorsal tubercles. Dorsum with ciliated bands (ca 14 per segment), continuing on elytro- phores and dorsal tubercles (Averincev 1972: pl. 13:1). Elytra large, thin (compared with E. gi- gantea), subreniform, splashed with brown- ish pigmentation; surfaces with numerous microtubercles on anterior part, sharply pointed tubercles on middle part, and pos- terior part with additional globular vesicles with few to numerous terminal spines, and some papillae but no thick fringe (Fig. 3E; Monro 1939; fig. 4e; Uschakov 1962; pl. OF; Averincev 1972: pl. 13.5; Hartmann- Schréder & Rosenfeldt 1990: pl. 1:1, 2; PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Stiller 1996, pl. 10 fig. 3). Dorsal cirri with large cylindrical cirrophores, inflated basal- ly, with long papillate styles extending be- yond setae; dorsal tubercles nodular (Fig. 3B; Averincev 1972: pl. 13:2). Bilobed prostomium wider than long; median antenna with large ceratophore in anterior notch, with long, finely papillated style with subterminal enlargement and fil- amentous tip; lateral antennae inserted on anterior extensions of prostomium, without distinct ceratophores, styles similar to but shorter than median antenna; palps long, stout, tapering, with six longitudinal rows of papillae; two pairs of rather large eyes on posterior half of prostomium; tentacu- lophores (segment I) lateral and anterior to prostomium, with digitiform acicular lobes, group of long setae directed anteriorly, and long dorsal and ventral tentacular cirri, sim- ilar to but longer than antennae; with prom- inent conical facial tubercle below ceratop- hore of median antenna (Fig. 3A; Averin- cev 1972:pl. 13:1; Stiller 1996:pl. 10, fig. 1). Segment II with prominent semilunar nuchal pad and conical nuchal fold cover- ing posterior part of prostomium, first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri, inserted ba- sally, similar to tentacular cirri (Fig. 3A; Averincev 1972:pl. 13:1; Stiller 1996: pl. 10, fig. 1). Biramous parapodia with smaller and shorter notopodia and larger neuropodia, both with projecting digitiform acicular lobes (Fig. 3B). Notosetae numerous, short to longer, slightly stouter than neurosetae, with long spinous regions and short bare tapered tips (Fig. 3C; Averincev 1972: pl. 13:4; Stiller 1996: pl. 10, fig. 4). Neurosetae numerous, with long spinous regions, all ta- pering to long, slender, bare, pointed tips (Fig. 3D; Averincev 1972: pl. 13:3; Stiller 1996: pl. 10, fig. 5). Ventral cirri short, ta- pered, papillate (Fig. 3B). Etymology.—The species is named for the late P. V. Uschakov, in recognition of his vast contributions to the study of the Polychaeta. VOLUME 110, NUMBER 4 545 | { \ Fig. 3. Eulagisca uschakovi, new species, A—D, paratype (BMNH 1941.3.3.17); E, paratype (USNM 58351): A, Dorsal view of anterior end, pharynx partially extended; missing: palps, styles of median and lateral antennae, dorsal and ventral tentacular cirri, except left one; segment 2 with nuchal fold partially folded back, right parapodium not shown: B, Right cirrigerous parapodium, posterior view; C, Tip of notoseta; D, Tips of two neurosetae; E, Right elytron, with detail of microtubercles, sharply pointed tubercles, spinous globular vesicles, and papillae. Scales = 1.0 mm for A; 1.0 mm for B; 0.1 mm for C, D; 2.0 mm for E. 546 Remarks.—The polynoids from the Da- vis Sea in 11—920 m, referred by Averincev (1972:131, pl. 13:1—7) to Eulagisca gigan- tea, were not available for examination. The description and figures agree with E. uschakovi. One of the specimens from El- ephant Island (Antarctica) in 280-437 m, re- ferred to E. gigantea by Hartmann-Schréd- er and Rosenfeldt (1990:91, pl. 1:1, 2, not 3-5), also agree with E. uschakovi. The elytra of E. uschakovi differ from the other species of Eulagisca in having sharp- ly pointed tubercles and globular vesicles with few to numerous terminal spines. The neurosetae of E. uschakovi agree with E. puschkini Averincev by having neurosetae tapering to sharp tips, rather than ending in slightly hooked blunt tips, as in the other species of Eulagisca. This also applies to the record of E. gigantea by Stiller (1996, pl. 10:figs. 3, 5) from the eastern Weddell Sea. Distribution.—Antarctic, off MacRobert- son Land, off Enderby Land, Davis Sea, Bransfield Strait, off Palmer Archipelago, Scotia Sea, Elephant Island, South Orkneys, Ross Sea, Weddell Sea, in 10—920 meters. Eulagisca macnabi, new species Fig. 4 Material examined.—Antarctic: Off South Orkneys, 61°00’S, 44°58'W, 283-329 m, Eastwind, Sta. 29, 11 Feb 1966, D. L. Pawson and D. E Squires, coll., holotype (USNM 58352). Description.—Holotype 62+ mm long, 38 mm wide with setae, segments 24, plus small regenerating posterior end. Body flat- tened, with long parapodia (about as long as body width); dorsum with transverse cil- iated bands (ca 10 per segment) extending on bases of elytrophores and dorsal tuber- cles (Fig. 4A—C). Elytra large, subreniform, covering dor- sum, with fringes of long border papillae laterally, posteriorly and medially; surfaces splotched with brownish pigmentation, con- ical microtubercles on anterior part, small PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON to larger spiny tubercles posteriorly and medially, some rounded tubercles with me- dial filamentous extensions (Fig. 4F). Dor- sal cirri with cylindrical cirrophores with glandular bulbous lobes on anterior and posterior sides and distal brown bands; styles missing; dorsal tubercles nodular (Fig. 4C). Bilobed prostomium with large cerato- phore of median antenna in anterior notch, style missing; lateral antennae inserted on anterior extensions of prostomium, with long papillate styles and filamentous tips; two pairs of lateral eyes moderate in size; palps long, tapering, with six longitudinal rows of papillae; tentaculophores lateral to prostomium and palps, each with projecting digitiform acicular lobe, missing dorsal and ‘ventral tentacular cirri, and bundle of long setae (ca. 14); conical facial tubercle ventral to ceratophore of median antenna; dark brown bands on ceratophore of median an- tenna, lateral extensions of prostomium for lateral antennae, and tentaculophores (Fig. 4A). Segment I with semilunar nuchal pad and subconical nuchal fold, partially pulled back, first pair of large elytrophores, bira- mous parapodia, and long ventral buccal cirri, similar to lateral antennae (Fig. 4A). Biramous parapodia with prominent pro- jecting digitiform acicular lobes on both rami. Notopodium smaller than neuropo- dium, rounded, with projecting acicular lobe on lower side; larger neuropodium with subconical presetal acicular lobe and shorter subconical postsetal lobe (Fig. 4B, C). Notosetae numerous, forming radiating bundle of three lengths, stouter than neu- rosetae, acicular, with spinous rows and rather long bare tips (Fig. 4B, D). Neuro- setae numerous, forming fan-shaped bun- dle, with long spinous regions, upper ones more slender, all with rather long, slightly hooked bare tips (Fig. 4B, E). Ventral cirri short, tapered, papillate (Fig. 4B, C). Ne- phridial papillae beginning on segment VI. Etymology.—The species is named for the late Dr. James A. Macnab, a dedicated and inspirational teacher. VOLUME 110, NUMBER 4 547 Fig. 4. Eulagisca macnabi, new species, holotype (USNM 58352): A, Dorsal view of anterior end, missing: styles of median and right lateral antennae, right and left dorsal and ventral tentacular cirri, right ventral buccal cirrus, and elytra; nuchal fold partially pulled back; B, Right elytrigerous parapodium, anterior view, aciculae dotted; C, Right cirrigerous parapodium, posterior view, style of dorsal cirrus missing; D, Notoseta from same; E, Middle and upper neurosetae from same; FE Right elytron, with detail of microtubercles, spiny vesicles, and papillae. Scales = 2.0 mm for A-C; 0.1 mm for D, E; 2.0 mm for EF 548 Distribution.—Antarctic, South Orkneys, 300 meters. Remarks.—The elytra of E. macnabi dif- fer from the other species of Eulagisca by the well developed border fringes of papil- lae and the surface spiny tubercles. Eulagisca puschkini Averincev, 1972 Eulagisca puschkini Averincev, 1972:131, pl. 13:8—12. Remarks.—A single specimen was col- lected by aqualung in the Davis Sea, in 32 meters, on rocky bottom, among sponges and hydroids (Type deposited in ZIASL). The type was not available for examination. The following was reported by Averincev. The large specimen is 175 mm long, 55 mm wide without setae, and 85 mm wide with setae, number of segments ?. The an- terior end, including the prostomium, palps, tentaculophores, facial tubercle, nuchal fold, and pharynx, agree with Eulagisca gi- gantea. The elytra differ from the other spe- cies of Eulagisca by lacking border and sur- face papillae, by having numerous micro- tubercles on the anterior part, with a red- brown iridescent spot on the medial half (pl. 13:8). The biramous parapodia have large digitiform processes with inner acic- ulae. The notosetae are thicker than the neurosetae, with transverse spinous rows and short bare tips (pl. 13:12). The neuro- setae have subdistal spinous rows and end in tapered bare tips, and few with small sec- ondary tooth (pl. 13:9—11). The elytra of E. puschkini differ from other members of Eulagisca by lacking bor- der papillae, surface tubercles and vesicles, and with only numerous microtubercles on the anterior half. The neurosetae of E. pus- chkini, with tapered bare tips, agree with those of E. uschakovi new species. Distribution.—Antarctic, Davis Sea, in 12 meters. Pareulagisca, new genus Type species.—Eulagisca panamensis Hartman, 1939. Gender: feminine. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Diagnosis.—Body subrectangular, taper- ing slightly anteriorly and posteriorly, with segments up to 37. Elytra and bulbous ely- trophores 16 pairs, on segments 2, 4, 5, al- ternate segments to 23, 26, 29, 32, 33. El- ytra large, covering dorsum, smooth, with- out tubercles or papillae. Dorsal cirri on non-elytrigerous segments, with cylindrical cirrophores, bulbous basally, with papillate styles extending to tips of setae; dorsal tu- bercles slightly bulbous. Prostomium bi- lobed, with two long finely papillate palps and three papillate antennae; ceratophore of median antenna large, in anterior notch of prostomium, lateral antennae inserted on anterior extensions of prostomium subter- minal to ceratophore of median antenna; two pairs of lateral eyes, moderate in size. . Tentaculophores of segment I lateral to pro- stomium, each with single seta, without projecting acicular lobe (as in Eulagisca), and papillate dorsal and ventral tentacular cirri; facial tubercle weakly developed. Sec- ond or buccal segment with thick, fleshy nuchal fold, first pair of elytrophores, bi- ramous parapodia, and long ventral buccal cirri, lateral to ventral mouth. Parapodia bi- ramous, noto- and neuropodia about equal in length, both subconical with projecting acicular lobes. Numerous notosetae forming radiating bundles, nearly as long as neuro- setae, of two kinds: stout, acicular (about as stout as neurosetae), smooth or with faint spinous rows; slender, capillary, smooth or with few spines. Numerous neurosetae stout, with long spinous regions and long, bare, slightly hooked entire tips, with or without minute secondary tooth. Ventral cirri short, tapered, papillate. Pygidium with pair of papillate anal cirri. Nephridial papillae beginning on segment IV. Etymology.—Pareulagisca from Greek para = near + Eulagisca in allusion to the similarity of the two genera. Pareulagisca panamensis (Hartman 1939), new combination Fig. 5 Eulagisca panamensis Hartman, 1939:31, pl. 3:figs. 38—42. VOLUME 110, NUMBER 4 549 | ti iz Zz hy. : SEEN Ze Eg ge Zc BSG Zi SENG Fig. 5. Pareulagisca panamensis, holotype of Eulagisca panamensis (LACM-AHF 10): A, Dorsal view of anterior end; B, Right elytrigerous parapodium, anterior view, aciculae dotted; C, Right cirrigerous parapodium, posterior view; D, Three stout notosetae from same; E, Two slender, capillary notosetae from same; K Lower and upper neurosetae from same; G, Right elytron. Scales = 2.0 mm for A; 1.0 mm for B, C; 0.1 mm for D—F; 2.0 mm for G. Material examined.—Pacific: Panama, _ subrectangular, flattened, tapering slightly Pifias Bay, 07°34'N, 78°12'W, shore, rock, anteriorly and posteriorly. Elytra and bul- Velero Sta. 436, 28 Jan 1935, holotype bous elytrophores 16 pairs (not 15, as in- (LACM-AHF 10). dicated by Hartman) on segments 2, 4, 5, Description.—Holotype 42 mm long, 15 7, alternate segments to 23, 26, 29, 32, 33, mm wide with setae, 37 segments. Body dorsal cirri present on posterior four seg- 550 ments. Elytra large, covering dorsum, fleshy, smooth, without tubercles or papil- lae, with rust-colored semilunar pigmenta- tion on posterior half (Fig. 5G; Hartman 1939, pl. 3:fig. 42). Dorsal cirri with cylin- drical cirrophores, bulbous basally, and long styles extending to about tips of setae, with short papillae on distal half and brown pigmentation on basal three-fourths; dorsal tubercles slightly bulbous (Fig. 5C; Hart- man 1939: pl. 3, fig. 39). Bilobed prostomium suboval, without ce- phalic peaks; ceratophore of median anten- na in anterior notch of prostomium, very large, with long papillated style enlarged subdistally, with filamentous tip; lateral an- tennae inserted on anterior extensions of prostomium, subterminal to large cerato- phore of median antenna, with shorter styles; palps long, thick, tapered, finely pap- illated; two pairs of eyes moderate in size; tentaculophores lateral to prostomium, each with single seta, dorsal and ventral tentac- ular cirri, similar to median antenna; facial tubercle indistinct (Fig. 5A). Second or buccal segment with triangular nuchal fold, first pair of bulbous elytrophores, biramous parapodia, and long ventral buccal cirri, similar to tentacular cirri (Fig. 5A). Notopodia of biramous parapodia almost as long as neuropodia, subconical, with pro- jecting acicular lobes on lower sides; neu- ropodia subconical, with longer presetal acicular lobes and shorter, rounded postsetal lobes (Fig. 5B, C; Hartman 1939: pl. 3, fig. 39). Notosetae numerous, forming radiating bundle of two kinds: acicular, about as stout as neurosetae, mostly smooth and shorter, curved, with 6-12 widely-spaced spinous rows (Fig. 5D: Hartman 1939: pl. 3, fig. 40); and slender, tapering to pointed and capillary tips, smooth or with few widely- spaced spines (Fig. 5E). Neurosetae numer- ous, forming fan-shaped bundle, lower shorter to upper longer, with long spinous regions and rather long bare, slightly hooked entire tips, upper few with minute secondary tooth (Fig. 5F; Hartman 1939: PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Comparison of characters for Eulagisca and Pareulagisca. Eulagisca Pareulagisca Elytra 15 pairs 16 pairs Palp 6 longitudinal rows __ without longitudi- of papillae nal rows of pa- pillae Tentacu- with projecting acic- without projecting lophore ular lobe and acicular lobe group of long se- and single seta tae Notosetae of one kind, long, of 2 kinds: 1) stout, acicular 2) slender, cap- illary spinous rows, ta- pering to blunt tips pl. 3, figs. 38, 41). Ventral cirri short, ta- pered, with short papillae (Fig. 5B, C). Distribution.—Central Pacific Ocean, Panama, intertidal. Remarks.—Pareulogisca panamensis is unusual in having 16 pairs of elytra, instead of the more common 15 pairs, and in hav- ing two distinct types of notosetae: stout and slender, capillary. The large size of the ceratophore of the median antenna is also unusual. Pareulagisca differs from Eulagisca as shown in Table 1. Literature Cited Augener, H. 1932. Antarktische und Antiboreale Po- lychaeten nebst einer Hirundinee.—Scientific results of the Norwegian Antarctic Expedition 1927-28. Det Norske Videnskaps-Akademii Oslo, Pp. 1-88. Averincev, V. G. 1972. [Benthic polychaete worms of the Errantia from the Antarctic and Subantarctic in the material of the Soviet Antarctic Expedi- tions].—Akademiia Nauk SSSR Zoologisches- kii Institut 5:88—293. [In Russian] Benham, W. B. 1921. Polychaeta. Australian Antarc- tic Expedition 1911—14, Under the leadership of Sir Douglas Mawson, D.Sc.B.E.—Scientific Reports, Series C. Zoology & Botany 6(3):1-— 128. Desbruyéres, D. 1976. Bionomie benthique du plateau continental de l’archipel des Iles Kerguelen- Macrofaune 3. Contribution a 1’étude des Pol- ynoidae (Annelides Polychétes) des Terres aus- trales et Antarctiques Frangaises.—CMFRA Prospections en Océnographie Biologique et VOLUME 110, NUMBER 4 Bionomie Benthique aux abords des Iles Ker- guelen et Crozet No. 39:83—96. Hartman, O. 1939. Polychaetous annelids. Part I. Aphroditidae to Pisionidae.—Allan Hancock Pacific Expeditions 7(1):1—156. . 1967. Polychaetous annelids collected by the USNS Eltanin and Staten Island cruises, chiefly from Antarctic Seas.—Allan Hancock Mono- graphs in Marine Biology 2:1—387. Hartmann-Schréder G. 1986. Die Polychaeten 56 Re- ise der ‘“‘Meteor”’ zu den South Shetland-Inseln (Antarktis).—Mitteilungen aus dem hambur- gishchen zoologischen Museum und Institut 83: 71-100. , & P. Rosenfeldt. 1990. Dis Polychaeten der “Walther Herwig”’ Reise 68/1 nach Elephant Is- land (Antarktis) 1985 Teil 1: Aphroditidae bis Cirratulidae.—Mitteilungen aus dem Hambur- gishchen zoologischen Museum und Institut 87: 89-122. Kinberg, J. G. H. 1856. Nye slagter och arter af An- nelider.—Ofversigt af Konglia Vetenkaps-Aka- demiens Forhandlingar, Stockholm 12:381—388. McIntosh, W. C. 1885. Annelida Polychaeta.—Report on the Scientific Results of the Voyage of H. M. S. Challenger ... 1873-76 ... Zoology 12(34):1-554. Monro, C. C. A. 1930. Polychaete worms.—Discov- ery Reports 2:1—222. 351 . 1936. Polychaete worms. II.—Discovery Re- ports 12:59-198. 1939. Polychaeta.—Reports British Austra- lian and New Zealand Antarctic Research Ex- pedition, 1929-1931, series B 4(4):87—156. Pettibone, M. H. 1989. A new species of Benhami- polynoe, (Polychaeta:Polynoidae:Lepidastheni- inae) from Australia, associated with the unat- tached stylasterid coral Conopora adeta.—Pro- ceedings of the Biological Society of Washing- ton 102:300-—304. Stiller, M. 1996. [Distribution and biology of the Aphroditides and Polynoids (Polychaeta) in the eastern Weddell Sea and the Lazarev Sea (Ant- arctica).].—Reports on Polar Research, No. 185 Heft Nr. 185/1996:1—200. [in German] Uschakov, P. V. 1962. [Polychaetous Annelids of the Families Phyllodocidae and Aphroditidae from the Antarctic and Subantarctic Waters.—Bio- logical Results of Soviet Antarctic Expedition (1955—1958).] Issledovaniia fauny morei Zoo- logicheskii Institut Akameniia Nauk SSSR, I((X):129—189. [in Russian] . 1966. English translation of 1962 by Israel Program Scientific Translations, 131-188. Willey, A. 1902. Polychaeta.—Report on the Collec- tions of Natural History made in the Antarctic Regions during the Voyage of the “Southern Cross”’ 12:262-283. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):552-—557. 1997. Protodrilus gelderi, a new species of infralittoral, interstitial polychaete from Massachusetts Bay Nathan W. Riser Marine Science Center, Northeastern University, Nahant, Massachusetts 01908, U.S.A. Abstract.—A new species of the interstitial meiofaunal polychaete genus Protodrilus (P. gelderi) is described from relatively stable, subtidal, shelly sub- strate that occurs in patches along the north shore of Massachusetts Bay, U.S.A. Males and females differ in total body length as well as in the distribution of salivary glands and gamete producing regions of the body. With the exception of the absence of segmentally arranged ciliated bands, males of the new species are morphologically quite similar to P. ciliatus. However, the females differ markedly from all previously described species in that the ovaries are present from the first segment to the posterior end of the body. Interstitial meiofaunal polychaetes have commonly been encountered in coarse shel- ly substrate collected by divers at shallow depths in Massachusetts Bay. Polygordius appendiculatus Fraipont, 1887, P. lacteus (Schneider, 1868), and P. triestinus Hem- pelmann, 1906 are routinely present in large numbers in such substrate at depths of 3-— 30 m. In some of these samples from Na- hant Bay and Broad Sound at depths of 6— 20 m, a previously undescribed species be- longing to the genus Protodrilus, usually numbering in excess of 50 per 8 ounce (vol- ume) plastic bag of substrate, has also been present. Other species of Protodrilus have been encountered during intertidal meio- faunal studies along the shores of the Gulf of Maine, but this species appears to be strictly subtidal. Ruebush (1939) reported Protodrilus leuckarti Hatschek, 1880 from specimens collected intertidally by Dr. Don- ald Zinn at Branford, Connecticut, USA. No further details were presented in the pa- per; however, Jouin (1970) stated that it was more likely that the specimens were P. cil- iatus Jagersten, 1952 and this unconfirmed report remains the only record of a species of the genus in New England. The follow- ing description of the new species from the subtidal is derived primarily from living specimens. Materials and Methods Substrate was collected by Scuba divers using garden trowels or large Modiolus Shells for scooping into plastic bags. The bagged samples were maintained in the lab- oratory on a running seawater table to pre- vent temperature shock prior to sorting. The sediment was extracted with 7.5% MgCl, and decanted onto 153 wm screens from which the animals were washed into shal- low dishes containing fresh sea water. More than 200 living individuals have been ex- amined since the species was first encoun- tered in 1976 in an effort to determine dis- tribution of nephridia and location of sperm ducts. Weather conditions have prevented collecting between February and April. Sta- bilization of substrate following winter storms has necessitated an annual search to relocate areas occupied by animals of inter- est. The new species has only been obtained between June and January during which time, small and sexually immature speci- mens have been uncommon. Anesthetized specimens were fixed in Hollande’s cupri-picri-formal-acetic and VOLUME 110, NUMBER 4 Figs. 1, 2. Protodrilus gelderi, living specimens. 1. lobes, scale =0.08 mm. whole mounts were stained with Mayer’s alcoholic HCl carmine, in some instances counterstained with indigo-carmine. Suit- able fixed individuals were selected for sec- tioning following embedding in Steedman’s polyester wax. Mounted sections were treated and subsequently stained with Hei- denhain’s Azan or Alcian blue buffered to pH 2.5 (Humason 1967). Family Protodrilidae Czerniavsky, 1881 Protodrilus gelderi, new species Figs. 1-8 Type specimens.—Zoologisches Museum der Universitat Hamburg: holotype female P-23532 and allotype male P-23533, whole mounts on one slide. Muséum National d’ Histoire Naturelle de Paris: paratype male UE 622, female UE 623. American Muse- um of Natural History: paratype male AMNH 4194, female AMNH 4193. U. S. National Museum of Natural History: para- type male USNM 175429, Female USNM 175428. Type locality——Coarse shelly sand, off north side of Egg Rock, 20 m depth. Nahant Bay, MA., 42°26’N; 70°54'W. Description.—Body slightly transparent; 353 Dorsal view of anterior prostomial region. 2. Caudal red coloration associated with buccal mass; gut green to brown. Up to 44 segments in mature individuals, females to 9 mm in length and males to 7 mm. Head broader than following region of body; midbody slightly broader than head, tapering to pos- terior end. Caudal lobes broad, fan-shaped, with four or more large stereocilia (Fig. 2) projecting from posterior margin. Prosto- mium bluntly rounded with four stereocilia around apex. Tentacles up to 1.2 mm long, extending back to segment 6. Ocelli absent. Pair of anterior sensory organs (‘“‘stato- cysts’’) ~16 wm diameter to either side of prostomial apex. Nuchal organs large, slit- like, and lateral, at bases of tentacles. A sin- gle very long stereocilium about half head diameter, on dorsal surface immediately be- hind top of each nuchal organ. Small, cili- ary patches present on head and body, cil- iary annulations absent. A stereocilium is present laterally near the anterior and pos- terior end of each segment, and one is sometimes present in the middle of the lat- eral margin. Curved (gourd shaped) bacil- lary gland cells ~30 pm long and 7 wm maximum diameter, in thick double row across head behind nuchal organs, scattered 554 in epidermis of anterior body segments be- coming more abundant posteriorly, concen- trated along sides of ventral ciliated furrow. Pygidial stylus not evident. Salivary glands extend into segment 12 in most males, to 19 maximum; at least into segment 15 and up to 30 in females. Ovaries occur from segment | or 2 into penultimate or antepen- ultimate segment. Oviducts and dorsal or- gans absent. Testes from segment 11 to an- tepenultimate segment. Male lateral organ in segment 7, large, round, surrounded by gland cells, followed by a continuous cili- ated furrow from segment 8 to 17-18; spermducts in 12 through 15. Etymology.—The species is dedicated to Dr. Stuart R. Gelder who as a predoctoral student carried out part of his dissertation investigations at our laboratory and intro- duced me to histochemical procedures which could be applied to the study of in- terstitial polychaetes. Observations.—Total length and number of segments of mature animals are variable features frequently attributable to posterior regeneration. The anterior sensory organs contain an amorphous body and are not readily visible. In sections, these bodies ap- pear as a gel, but in whole mounts are di- vided into two or three parts. The large, lat- eral, slit-like nuchal organs are the only ob- vious area on the head bearing motile cilia, as the cilia tufts on the head are insignifi- cant. SEM indicates that the stereocilia on each body segment are arranged in three in- terrupted rings. A row of pigment cells ex- tends across the prostomium (Fig. 1) im- mediately behind the nuchal organs. The bacilliary gland cells stain strongly with Al- cian blue pH 2.5, and are cyanophilous with Azan. In sections, it can be seen that the cell bodies displace the other epidermal cells and bulge beneath them, so that the epidermis could be described as pseudostrat- ified. They can be distinguished by their rods from other mucous cells which also expand beneath the surficial epidermal cells. Mucous cells are abundant in and along the sulcus between segments, and are PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dominant (Figs. 6, 8) outside the layer of serous cells which discharge into the lateral organ in segment 7 of the males. These se- rous cells are also subepidermal with long necks which pass between the ciliated sup- port cells of the organ (Fig. 6). Parenchyme cells fill the area, laterally, between the epi- dermis and longitudinal muscles. However, the cells are absent near the middorsal and midventral lines of the body. There are usu- ally nine diagonal muscle bundles on either side in each segment of mature individuals. Salensky (1907) stated “ungefahr 6—7”’ in the Protodrilus species which he investi- gated, noting that the number was much smaller than in the genus Polygordius. The arrangement of these muscles in sectioned material is similar to the diagram by Pier- antoni (1908: pl. 2, fig. 2). Relaxation of these muscles allows a flattening of the body, and expansion of the ventral ciliated creeping groove. The strong retractor muscles of the buc- cal mass usually contract following fixa- tion, so that the esophagus is arched as it passes over the mass (Fig. 5) to open at the anterior margin of the mouth. As a result, the initial group of salivary glands are drawn forward along the sides of the arch. In living individuals, the base of the esoph- agus is a straight tube extending back from the arched portion. The epithelium of the esophagus is initially composed of tall, thin, simple columnar cells with long cilia (Fig. 5). The longest region is formed by cuboi- dal cells with relatively large roundish nu- clei, and the cilia are spread apart. This is possibly an artifact resulting from the stretching of this region, although there is no intergradation (Fig. 5) with the cells of the anterior or posterior regions. The pos- terior straight part of the esophagus is lined with very tall ciliated columnar cells which are closely packed together. Variation in the number of segments con- taining salivary glands may be the result of posterior regeneration and is most notice- able in females. The number of salivary gland cells is reduced, often to one or two, VOLUME 110, NUMBER 4 355 Figs. 3-8. Protodrilus gelderi, segments 9-13 of male; dorsal view, whole mount, Mayer’s carmine with indigo carmine counterstain. 4. Anterior segments behind buccal mass of holotype female; ovaries (0) of right side in focus, sg = salivary glands, Mayer’s carmine. 5. Longitudinal section through esophageal region; Azan stain. 6. Slightly oblique section through aperture of male organ in segment 7; Azan stain. 7. Male organ in segment 7 with longitudinal trough extending posteriorly from it; whole mount, Mayer’s carmine with indigo carmine counterstain. 8. Oblique section through aperture of male organ. Alcian blue pH 2.5 stain. along each side of the midgut of the last few segments in which they occur. The pair of testes in segment 11 (Fig. 3) have rarely contained spermiogenetic stages. The male lateral organ in segment 7 is large and obvious as is the ciliated furrow which begins immediately behind it (Fig. 7). The sulcus between segments 7 and 8 is densely ciliated with closely packed mu- cous cells. Only euspermatozoa have been observed. The first pair of ovaries in preserved 556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON specimens, frequently project forward against the buccal cavity as the result of the retraction of the buccal mass drawing the esophagus and initial salivary glands up over it. The first two pairs of ovaries are smaller than subsequent ones and contain fewer oocytes. They are often difficult to observe in cleared, preserved specimens and whole mounts (Fig. 4). Oocytes are small, attaining a diameter of ~25 wm with large germinal vesicles. There are slightly over 50 oocytes in each mature pair of ova- ries. Maturation of ovaries is sequential with the smaller, less mature ovaries ante- rior and posterior to the mid-body region. The only ciliary action observed in the body cavity has come from a large pair of metanephridia in segment 8 of both sexes. The metanephridia are located on each side in this segment, and they appear to be the only excretory organs in the body. Protodrilus gelderi either attaches to the substrate with its caudal lobes or swims up- ward with a violent wriggling motion when disturbed, an escape strategy that was re- ported in the description of Protodrilus al- bicans Jouin, 1970. Specimens maintained in the laboratory are most frequently en- countered on the sides of the bowls or glid- ing on the surface film. Individuals live for several weeks in finger bowls of clean sea water at approximately ambient sea tem- perature, but a satisfactory food source has not been discovered to maintain them. Loss of the red pigment around the buccal mass occurs in a few days after isolation. Discussion The histology and organology of the ge- nus Protodrilus were extensively described by Pierantoni (1906) and Salensky (1907). Differences between species are not appar- ent at those levels except for the distribu- tion of cell types in the epidermis. How- ever, such variations may be attributable to different environmental conditions or a technic artifact. Gross anatomy of sexually mature individuals shows variation which allows for the identification of living worms, and often from stained whole mounts of well anesthetized individuals. The secondary sexual characters of males have been the most frequently used features to distinguish species. Females tend to be conservative in the degree of variation ex- cept for oocyte number and size, and the occurrence of cocoon glands. The four stereocilia around the apex of the prostomium appear to be a basic char- acter of the genus, and thus, plesiomorphic. Purschke (1990a, 1990b) examined the so called “‘statocysts’’ located near the apex of the prostomium of a number of species of Protodrilus. He concluded that they were not statocysts and based upon TEM obser- vations were more likely involved in pho- toreception. Purschke (1990b) referred to them as anterior sensory organs to distin- guish them from the phaosome-like poste- rior receptors of Protodrilus adherens Ja- gersten, 1952. They differ from pigmented ocelli, and he postulated that they consti- tuted an autapomorphy for the genus. The opacity of the preoral region of the prosto- mium of living P. gelderi makes it very dif- ficult to observe them with transmitted light, in spite of their size. The contents of the organelle, which appear as a partitioned gel in histological preparations, apparently are the paracrystalline bodies described from TEM observations by Purschke (1990b). Protodrilus gelderi differs from other members of the genus in the presence of ovaries along almost the entire length of the female mid-gut. No species has been de- scribed previously in which ovaries are present anterior to the segment 7, or with salivary glands in females posterior to seg- ment 20. Pierantoni (1906, 1908) noted that Uljanin (1877) and Hatschek (1880) had identified the salivary glands as ovaries, and they had concluded that the species of Protodrilus which they had described were hermaphroditic based upon the subsequent posterior occurrence of testes in the same specimens. VOLUME 110, NUMBER 4 Males are morphologically similar to those of P. ciliatus, differing primarily in the absence of ciliary annulations. Among species lacking such annulations, they are similar to Protodrilus submersus von Nord- heim, 1989 and P. albicans, in both of which, however, salivary glands are absent from the fertile region. Bacillary glands are densely packed in the dorsal epidermis, and nephridia are readily apparent in all seg- ments of living P. submersus (pers. obs.) in contrast to the situation in P. gelderi. Pro- todrilus albicans is described as much larg- er than P. gelderi and is figured with oval, dorsal nuchal organs. Jouin (1970) did not Mention pigmentation of the buccal mass for P. albicans, although she recorded it’s presence for other species described in the same paper. The implication was that the mass was not pigmented. Acknowledgments Intertidal interstitial faunal investigations over the past thirty-five years have been ex- tended into the subtidal by the cooperation of many students who, while diving to do their thesis research, carried plastic bags to obtain “‘any coarse sand”’ they might en- counter. Their willingness to help is sin- cerely appreciated. This paper represents Contribution No. 224 from the Marine Sci- ence Center of Northeastern University. Literature Cited Hatschek, B. 1880. Protodrilus leuckarti, eine neue Gattung der Archianneliden.—Arbeiten aus den So7 Zoologischen Instituten der Universitat Wien 3: 79-92. Humason, G. L. 1967. Animal tissue techniques. 2nd ed. Freeman, San Francisco, 569 pp. Jouin, C. 1970. Recherches sur les Protodrilidae (Ar- chiannélides). 1—Etude morphologique et sys- tématique du genre Protodrilus.—Cahiers de Biologie Marine 11:367—434. Nordheim, H. von 1989. Six new species of Proto- drilus (Annelida, Polychaeta) from Europe and New Zealand, with a concise presentation of the genus.—Zoologica Scripta 18:245—268. Pierantoni, U. 1906. Organi genitali e glandole sali- vari nei Protodrili.—Bollettino della Societa dei Naturalisti in Napoli 20:154—157. 1908. Protodrilus—Fauna und Flora des Golfes von Neapel 31:1—226. Purschke, G. 1990a. Fine structure of the so-called statocysts in Protodrilus adhaerens (Protodrili- dae, Polychaeta).—Zoologischer Anzeiger 224: 286-296. . 1990b. Ultrastructure of the “‘statocysts”’ in Protodrilus species (Polychaeta): Reconstruc- tion of the cellular arganization with morpho- metric data from receptor cells.—Zoomorphol- ogy 110:91-104. Ruebush, T. K. 1939. The occurrence of two rare gen- era Protohydra and Protodrilus, on the east coast of North America.—Science 90:617—618. Salensky, W. 1907. Morphogenetische Studien an Wiirmern. II. Uber die Anatomie der Archian- neliden nebst Bemerkungen itiber den Bau ei- niger Organe des Saccocirrus papillocercus.— Mémoires de 1l’Académie Impériale des Sci- ences de Saint-Pétersbourg. s.8. Classe Physico- Mathématique 19:103—451. Uljanin, W. N. 1877. Observations on the genus Po- lygordius of the Bay of Sebastopol.—Bulletin de la Société des Naturalistes de Moscou 52: 53—96. (in Russian) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):558—-559. 1997. Synonymy of Platicrista cheleusis (Tardigrada: Eutardigrada: Hypsibiidae) R. Deedee Kathman and Clark W. Beasley (RDK) Aquatic Resources Center, P. O. Box 680818, Franklin, Tennessee 37068 U.S.A.; (CWB) Department of Biology, McMurry College, Abilene, Texas 79697 U.S.A. Abstract.—The eutardigrades Platicrista cheleusis Kathman 1990 and Di- phascon craigi Beasley 1990 are declared to be synonymous based on exam- ination of the specimens from both authors. Pilato (1987) revised the genus Diphas- con Plate, 1889 and subdivided it into four genera: Hebesuncus, Diphascon, Mesocris- ta, and Platicrista. Characteristics of the bucco-pharyngeal apparatus were used to establish these new genera, as there tends to be little variability in these apparati with- in genera. Diphascon is separated from Pla- ticrista by the following characters: in Di- phascon the apophyses for the insertion of the stylet muscles are shaped like semilunar hooks, the furcae have thickened processes at the apices, the pharyngeal tube is longer than the buccal tube, and the stylet supports are inserted between half and three-quarters of the buccal tube length; in Platicrista the apophyses are in the shape of wide and flat ridges, the furcae are arch-shaped and taper at their apices, the pharyngeal tube is as long as or slightly shorter than the buccal tube, and the stylet supports are inserted at the end of the buccal tube. Although the four genera established by Pilato (1987) are accepted today, confusion surrounding the descriptions and placement of species within these genera led some au- thors to continue to use only the name Di- phascon of Plate, 1889, for several years. In 1990, Beasley described a new species of tardigrade from Colorado, USA, as Di- phascon craigi. In that same year, Kathman described Platicrista cheleusis from British Columbia Canada. The two descriptions of the holotypes, both deposited in the U.S. National Muse- um (USNM 234018 and USNM 234019), are very close, although there are some dif- ferences. The specimens from British Co- lumbia did not have eyes, whereas those from Colorado did, although they are no longer apparent in the preserved material. The presence of eyes appears to be a vari- able trait, with eyes apparent during certain times of the year in the same species from the same location (pers. obs., senior author; pers. comm., D. R. Nelson, East Tennessee State University, Johnson City, Tennessee). The reason for such variability has not been investigated. Kathman (1990) states that there are “‘two cuticular bars on first three pairs of legs, one beside the external claw and the other at the base between the inter- nal and external claws”. Her drawing, how- ever, indicates that the first of these bars is beside the internal claw, which is the cor- rect position. In describing the two macro- placoids, Beasley (1990) states “‘first (4.8 2) Shorter than the second (22.3 w)’’. His drawing suggests that the first length is probably erroneous; rather, the first is slightly more than twice as long as the sec- ond, based on the description of the para- types, as well as new measurements of specimens from both collections. The other differences in measurements of various characters of the holotypes are most likely a result of the differences in size of the two animals (688 wm versus 530 pm length). Although Beasley found all six specimens VOLUME 110, NUMBER 4 from Colorado above 2900 m, Kathman found her 14 specimens from 150—1540 m. Examination of specimens from both col- lections reveals the same cuticular pattern- ing, although the decrease in size of the polygons in the cuticle from the anterior to posterior end is more noticeable in the spec- imens from British Columbia. In some specimens it is difficult to detect any change in size posteriorad. The sclerotized bar under the claws of the fourth pair of legs is variable among specimens, some- times small and occurring between the two claws (as described by Kathman) and some- times extending under the internal claw and reaching the middle of the bottom of the external claw. Perhaps this variability is a result of the stage of development of the tardigrade. Beasley and Kathman have determined that the two species are synonymous. Ac- cording to the rules of priority of the Inter- national Code of Zoological Nomenclature, the correct name is that of Kathman (1990), which appeared in publication several weeks prior to that of Beasley (1990). It is interesting to note differences that appear to be due to the preservation tech- niques. The two sets of specimens were 559 prepared and mounted in two different me- dia—the ones from British Columbia were removed directly from water and placed in Hoyer’s medium, while those from Colo- rado were fixed in Cuénot’s fixative, dehy- drated in glycerin and mounted in Faure’s liquid. Several years later, the reticulated cuticle is barely visible in Hoyer’s, while it is fairly distinct in Faure’s. However, the eyes are no longer visible in the Colorado specimens. The sclerotized parts, such as the furcae, stylets and other mouthparts have been better preserved in Hoyer’s. Literature Cited Beasley, C. W. 1990. Tardigrada from Gunnison Co., Colorado, with the description of a new species of Diphascon.—The Southwestern Naturalist 35:302—304. Kathman, R. D. 1990. Eutardigrada from Vancouver Island, British Columbia, Canada, including a description of Platicrista cheleusis n. sp.—Ca- nadian Journal of Zoology 68:1880-—1895. Pilato, G. 1987. Revision of the genus Diphascon Plate, 1889, with remarks on the subfamily Ita- quasconinae (Eutardigrada, Hypsibiidae). Pp. 337-357 in R. Bertolani, ed., Biology of tardi- grades.—Selected Symposia and Monographs 1:1-380. Unione Zoologica Italiana, Mucchi Editore, Modena. Plate, L. 1889. Beitrage zur Naturgeschichte der Tar- digraden.—Zoologische Jahrbiicher Abteilung fiir Anatomie der Tiere 3:487—550. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):560—568. 1997. The cladoceran collection of the National Museum of Natural History, Smithsonian Institution, Washington, D.C. Dorothy B. Berner Department of Biology, TU 015-00 Temple University, Philadelphia, Pennslyvania 19122, U.S.A. Abstract.—The cladoceran collection at the Smithsonian Institution National Museum of Natural History is both extensive and diverse. Wet materials (al- cohol- and formalin-preserved) comprise over 18,000 littoral and limnetic zoo- plankton samples and approximately 300 lots of sorted, identified specimens. There are over 20,000 slides, the majority with specimens identified to species and in excellent, useable condition; about three quarters are grouped by taxon, the remainder according to country or specific location. Anomopod taxa (Chy- doridae, Daphnia, and Bosmina) predominate in materials donated by D. G. Frey, R. W. Kiser, and E. S. and G. B. Deevey, respectively, but the collection includes representatives of the Onychopoda, Haplopoda, and Ctenopoda as well. The collection is world-wide in scope. Geographically, coastal regions of North America are particularly well represented; there is considerable material from Central and South America, Denmark, South Africa, and Australia. There are presently types of 52 species comprising about one hundred specimens, half on slides and half in alcohol. The comprehensive library given by D. G. Frey facilitates study of cladocerans at the museum. Following receipt of gifts of sizeable col- lections of cladoceran specimens and sam- ples to the National Museum of Natural History (NMNH) by R. W. Kiser and D. G. Frey, the cladoceran collection at the Nat- ural History Building in Washington, Dis- trict of Columbia, was moved to the Mu- seum Support Center (MSC) in Suitland, Maryland. At the MSC, fluid-preserved (“‘wet’’) specimens and samples are housed in a controlled-environment storage pod, and slides, books, and papers are housed in a nearby room, the David G. Frey Cladoc- eran Library. The NMNH cladoceran collection com- prises one of the finest assemblages of cla- doceran materials in the world. The British Museum (Natural History) holds more types and perhaps greater diversity in spe- cies and sorted specimens. The Canadian Museum of Nature (Ottawa) has a large col- lection of littoral samples, primarily from Canadian waters. However, the USNM cla- doceran-containing, unsorted zooplankton collection is unparalleled, both in number and geographic range of samples. The cla- doceran holdings of other North American museums are meager in comparison. Categories of Material Comprising the Cladoceran Collection The cladoceran materials held by the NMNH are curated by the Department of Invertebrate Zoology. They can be accessed in several ways. A six digit NMNH acces- sion number is assigned all items donated by an individual or otherwise acquired at one time; items given at different times by the same individual will bear different NMNH accession numbers. NMNH cata- logue numbers are given to individual lots of materials such as slides, bottles, etc. when they are catalogued by the Depart- ment of Invertebrate Zoology. Lastly, some items may be accessed by a number des- VOLUME 110, NUMBER 4 ignated by a previous owner of the mate- rials. A few older items in the collection lack accession and catalogue numbers (Ber- ner 1995) and are not detailed below. All NMNH numbers reported here are acces- sion numbers. To facilitate description of the cladoceran collection, I group its diverse materials into two categories: sorted specimens identifia- ble by NMNH accession and catalogue numbers, and larger gifts of multiple items commonly recognized by the donor’s name. Specimens with NMNH Accession and Catalogue Numbers Type material.—Approximately 100 type specimens, about half on slides and half in alcohol, representing 52 taxa. Many are species described by D. G. Frey from the Americas and elsewhere. Some of his para- types, retained in his personal collection and included in his major gift (see below), have not yet been catalogued. Wet material—(n alcohol.) Approxi- mately 90 vials of identified specimens, representing 40 genera, shelved by genus and species. Most are identified to species, some to genus only. There are catalogue cards for about 70 of them. One hundred and five catalogued bottles of unidentified cladocerans. There are also at least 12 vials of unidentified cladocerans in one of E. B. Reed’s gifts (NMNH 383116), located in the ““Copepod holding area’’ at the NMNH. Reed’s material, pri- marily from Colorado, may complement that in the Shantz and Dodds gifts (see be- low). Slide material.—Approximately 365 cat- alogued slides, representing 50 genera. Many of these slides are of specimens col- lected in Brazil by H. Kleerekoper, whose samples are among those donated by R. Kiser (see below). These specimens were mounted, identified, and donated individu- ally by Kiser and have accession and cata- logue numbers different from the Kiser gift. Data.—One set of catalogue cards for 561 (mostly) identified specimens, either hand- written or computer-printed. Cards are ar- ranged by genus and species; those for slides and wet specimens are interspersed. Types are identified by orange cards. The set does not include cards for catalogued bottles of unidentified cladoceran speci- mens, but does include cards for the vials of unidentified E. B. Reed cladocerans. David G. Frey Gift, NMNH 403774 Because of the size and complexity of this gift, its components are evaluated sep- arately below. The gift also included three discrete collections that Frey had acquired: the J. Richard collection and the E. A. Birge collection, which Frey obtained from the University of Wisconsin, Madison, and the E. S. and G. B. Deevey freshwater collec- tion. Frey incorporated the Richard and Birge collections into his own, and they are con- sidered as a subset of his gift. The Deevey freshwater materials are considered as a separate acquisition (see below). The materials donated by Frey reflect his early training (under C. Juday) and life-long interest in limnology, his research on the paleoecology of lakes as revealed by cla- doceran remains, and his world-wide stud- ies on the speciation and distribution of chydorid cladocerans in relation to glacia- tion and continental drift. Frey and his col- leagues sampled zooplankton extensively in the areas he studied: eastern North Ameri- ca, northern Europe, southern South Amer- ica, South Africa, and Australia. Frey also collected indefatigably on his international travels, so most parts of the world are rep- resented to some extent in his collection. Because they are primarily from the littoral zone, his zooplankton samples have a much higher diversity of microcrustaceans and other invertebrates than do limnetic sam- ples. In addition to containing cladocerans, they potentially are a rich source of material for studies of copepods, ostracods, and oth- er littoral organisms, and of the littoral bio- coenosis. Enhancing the sample collection 562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON is Frey’s meticulous documentation, by notes and mounted specimens, of the ma- terial he studied. Wet material.—(3—4% formaldehyde so- lution with a few ml of glycerol added to prevent total drying if evaporation occurs. Some sorted specimens may be in alcohol.) Approximately 9260 bottles of samples and specimens housed in 49 boxes. Per Frey’s accession books, 9193 samples have Frey’s own F—-—-— numbers on the bottle tops; in- side labels bear the same number, sample collection date and number, locality, and name of county, province, or country. Bot- tles comprising Richard and Birge samples are detailed below. Two boxes marked “‘Types”’ contain bot- tles with identified specimens; they are not numbered or listed in Frey’s accession ledg- ers. Twenty of the bottles are marked ‘‘Paratypes”’ and 47 have sorted, identified specimens. All have inner labels giving spe- cies name and collection data. They have not yet been catalogued by the NMNH and therefore have no identifying numbers. Field data for the samples are scattered in 92 small notebooks, many labeled with year and place visited. In these, Frey de- tailed aspects of his trips as well as collec- tion data; often, he later recorded his ac- cession number for each sample in red ink next to its collection data and number. Frey’s samples are recorded (usually by batches in correspondence with dates of collection) in two ‘“‘Collection Accession’’ ledgers that list Frey’s sample number, lo- cality, date, and collector’s name. Samples, Specimens, and some slides received from other collectors are similarly recorded. There is no index for finding samples from a particular locality. Several thousand inventory cards, filed by Frey number, detail contents of samples Frey studied. Chydorid species are listed, other cladocerans are noted by genus, and there are notes on other Branchiopoda, al- gae, etc. Frey requested that anyone remov- ing specimens from a sample note the spe- cies and number of specimens taken on the back of the corresponding card. If a visitor examined a sample Frey had not yet stud- ied, he requested that the researcher make up an appropriate inventory card. These practices are being continued at the NMNH. Slide material.—Several hundred slides of identified specimens (mostly chydorids), the large majority mounted in glycerine-jel- ly or polyvinyl-lactophenol with ringed coverslips, are arranged alphabetically by genus and species in a metal tray cabinet. They are in excellent condition, well-la- beled, usually bearing the Frey number of the original sample. Fourteen large and two small boxes con- tain Frey’s working slides, also in excellent condition, of identified specimens and para- types of particular taxa on which he pub- lished. Twenty-four boxes house specimens, mostly identified, from diverse locales: Southhampton Islands; the Soviet Union (F 1421, a gift from N. N. Smirnov that in- cludes three paratypes); Denmark; Holland; Africa; South America; U.S.A.; Canada; Maine-New Hampshire; Florida & south- eastern U.S.A.; and Monroe County, Indi- ana, ponds. In addition, there are miscella- neous sets of slides in wooden boxes: two boxes of cladocerans from the S. Wright collection (35 slides of poor quality), five boxes from Nova Scotia by G. Doran, and six boxes of various origin and content. Eleven large and 19 small boxes have slides of lake sediments and cores, repre- senting paleolimnological and taxonomical studies by Frey and his students. There is no inventory of Frey’s slides. Library.—The library reflects Frey’s em- phasis on chydorid cladocerans. While strong in all the early cladoceran literature, it lacks many recent papers on non-chydor- id cladoceran taxonomy as well as some important contemporary guides with keys. It is usefully organized: Eleven shelves of bound books, offprints, and photocopies (about 2500 items) of pub- lications relevant to cladoceran taxonomy, evolution, and ecology. These are num- VOLUME 110, NUMBER 4 bered and entered into Frey’s “Accession Book for Literature’? and are arranged on the shelves alphabetically by author. Two shelves of theses and dissertations relevant to limnology and cladoceran tax- onomy. Three shelves of books on limnology, pa- leontology, and taxonomy, most of which have relevance to cladoceran systematics. These do not have Frey accession numbers and are loosely arranged by topic and au- thor. Numerous unbound offprints of papers not directly related to chydorid systematics; these are housed by author in a small, mul- tidrawer file cabinet. There are both a literature accession ledger and an index card file of literature. Papers.—Papers filling several file cabi- nets were received with the Frey gift, prod- ucts of his lifetime of research, teaching, and professional activities. Materials per- taining to cladoceran systematics presently kept in the Frey Cladoceran Library are: Research data for specific ecological, pa- leolimnological, paleotaxonomic, and tax- onomic studies. Generally, a single folder contains all the material related to a single publication: data (counts, measurements, species lists, etc.) from samples that Frey studied, manuscript drafts, and sometimes related correspondence. Research and correspondence on historic, systematic publications, e.g., publication of the fascimile of Lilljeborg’s Cladocera Sue- ciae and of Sars’ unpublished 1861 manu- script. Negatives and prints of scanning electron micrographs associated with the above re- search and publications. Plates, maps, and miscellaneous draw- ings associated with research publications. A set of computer punch cards with data on Frey’s collections. The key to these cards (which are on a shelf of the library) appears to be a large posterboard diagram lying in a map drawer. Frey’s correspondence with other profes- sionals. Many of these are cladocerologists 563 and some letters contain taxonomic infor- mation. The J. Richard and E. A. Birge Materials (Gift of D. G. Frey) The impact of E. A. Birge’s early taxo- nomic studies of North American cladoc- erans, and the roles he and his associate, C. Juday, played in the development of Amer- ican limnology are well-documented (Frey 1963, Beckel 1987). Birge was in commu- nication with the French taxonomist, J. Richard, who had studied material collected from foreign countries in the late 1800’s by numerous persons. When Richard retired, Birge purchased his collection of samples and slides. D. G. Frey acquired that collec- tion as well as samples, slides, and miscel- laneous papers and notebooks belonging to Birge and Juday from the Museum of Zo- ology (formerly, the Wisconsin Geological and Natural History Survey) at Madison, Wisconsin. Some of the samples in the Birge collection apparently were collected in South America by Harriet B. Merrill, an as- sistant to Birge (Hartridge 1995), and had been thought lost (Reid 1991). Frey incor- porated the Richard and Birge samples into his wet collection, but kept the slides sep- arate. During this survey, papers and note- books relevant to these materials were found scattered among Frey’s papers and were assembled together in marked file cab- inet drawers. Richard and Birge described numerous new cladoceran species. The former did not designate or set aside types and although the latter indicated that types would be de- posited for some of the species he described (Birge 1910), they are not to be found in the museum at Madison, Wisconsin (EF Iwen, pers. comm.). Therefore, these ma- terials are of great importance. For instance, during this review the slides from which Birge made the drawings for his description of Wlassicsia kinistiensis were found, but none was designated as a type. Also located was a Slide of Ceriodaphnia acanthina 564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ross, 1896 from the type locality and date; as no type exists for this species, Birge’s slide might be significant. Furthermore, it is possible that specimens from type localities are in certain of the Richard and Birge sam- ples. Because of the significance of the Rich- ard and Birge materials to cladoceran tax- onomy, they warrant further careful study and documentation. Wet material.—(n formalin.) According to Frey’s accession book, samples from the Richard collection are: F 701—1041 (from Africa, Sumatra, South America & Haiti, the Soviet Union, China, and France). Samples belonging to the Birge collec- tion are: F 607—676 (from Wisconsin, South and Central America); 1042—1232 (1902-— 1903, from South America); 1095-1232, 1277-1282, and 1284 (1900-1929, from Wisconsin). Slides.—Six metal, tabletop file cabinets, with 25 trays per cabinet, not all full of slides. Four cabinets are labeled A—D (per- haps by Frey) and have labeled trays; the remaining two were labeled E and F for this review. No inventory of the slides was found. Twenty trays, mostly in cabinets A and FE appear to be Richard’s slides (as deter- mined by the handwriting and French la- bels). The remaining slides apparently are Birge’s. Their condition ranges from excel- lent (specimens remounted in polyvinyl lac- tophenol and relabeled in 1966-1967 by *““J.V.B.”’) to useless: some slides lack cov- erslips and specimens or have uninforma- tive labels. Data.—A drawer of index cards, appar- ently made by Birge, with various data. A subset of the cards relates to the Richard samples. Field notebooks from South American collections, apparently made by H. B. Mer- rill. A bound volume of Birge’s correspon- dence, assembled by the Museum at Mad- ison. His purchase of the Richard collection is documented here. Frey’s analysis of H. B. Merrill’s work, the cladocerans found in each of her sam- ples, and correspondence related to a paper he wrote about it. That paper was not found; Frey apparently sent it to M. L. Har- tridge, Merrill’s niece (cf. Hartridge 1995). A folder of photocopies, made by Frey, from Birge’s (H. Merrill’s?) notebooks and Merrill’s letters to Birge. The originals ap- parently were deposited by Frey at the State Historical Society of Wisconsin, in accord with a request by M. L. Hartridge, Merrill’s niece (M. L. H. pers. comm.). These materials have been put together in drawers marked “‘Birge Collection’’. Edward S. and Georgiana B. Deevey Gift . The Deeveys gave their marine collec- tions directly to the NMNH but their fresh- water materials to D. G. Frey, who kept them apart from his collection. Although re- ceived by the NMNH as part of the Frey gift, the Deevey freshwater materials are being treated as a separate acquisition. A large amount of this material derives from limnetic plankton tows, and is related to the Deevey studies on Bosmina and Eu- bosmina. Florida, Central America, Austra- lia, and New Zealand are particularly well represented. Deevey & Deevey (1971), available in the Frey library, is a valuable reference for locating many of the habitats represented in this material. Wet material.—(In formalin.) Sixteen boxes, all marked ‘“‘Deevey”’, containing glass and plastic vials and bottles of un- sorted zooplankton that are mostly labeled only with a date and lake name. A few bot- tles contain sorted, identified specimens. Locales marked on the boxes are: Wiscon- sin & Michigan; Costa Rica, etc.; Spain; Denmark, Hungary, Poland, Ireland; Viet- nam; Australia & Tasmania; Australia & New Caledonia; New Zealand; Venezuela & Brazil; Nova Scotia, Newfoundland, U.S.A./Canada; Florida (two boxes); Texas, Virginia, Kansas, Vermont, Maine, U.S.A; Unknown Countries. VOLUME 110, NUMBER 4 Reference to samples containing marine and brackish water cladocerans is in the Deevey notebooks labeled Long Island Sound, Long Island 1950, and Tisbury Great Pond located in the Department of Invertebrate Zoology at the Natural History Building in Washington, District of Colum- bia. Slides.—Eight boxes of slides, marked: Rodgers Lake & Queechy Lake, Connecti- cut (two boxes); Mexico-Guatemala; Mex- ico, Guatemala, Texas (contains some slides from New York, Ireland, Denmark); Gua- temala; Linsley Pond (Connecticut), Mexi- co, Guatemala, Texas, (includes some slides from the Killarney Lakes, Ireland); New Zealand; Australia (also contains slides from Tasmania, Poland, and Brazil). Many of the slides are of Bosmina; some have mixed zooplankton. Their general condition was not assessed. Data.—Five large, looseleaf notebooks, marked ‘““‘Deevey’’ and: Florida Lakes; Rodgers & Queechy Lakes; New Zealand Lakes; New Zealand, Tasmania, Australia; Guatemala Lakes. They contain research data on measurements of Bosmina from di- verse habitats, drawings, some information on copepods, and other miscellaneous in- formation. No field notebooks with collec- tion data were found. Alfred A. Doolittle Gift, NMNH 157575 A. A. Doolittle, a teacher at Central High School in Washington, District of Colum- bia, studied cladocerans as an avocation, collecting them in the eastern United States and especially in and near Sebago Lake, Maine, where he summered. Chydorus bi- cornutus, a North American endemic that he described, later triggered D. G. Frey’s studies on the “‘honeycombed”’ species of Chydorus. Doolittle’s material is primarily from Sebago Lake, Maine, and Lake Coo- per, Iowa. There are samples from other lakes in those areas, from various north- eastern U.S.A. states, and a few from Mex- ico. 565 Wet material.—(In alcohol.) Twenty-four jars, labeled by state, containing vials of un- sorted zooplankton samples. (One District of Columbia bottle has two vials of sorted specimens). Vials contain labels with sam- ple dates only; jars contain labels with Doo- little’s name and NMNH accession number. Slides.—One box, mostly of identified cladocerans from lakes in Maine, mainly Sebago Lake. About two thirds appear to be in useable condition. The box is tem- porarily labeled ‘‘Doolittle Collection’’. There is no inventory of the slides, and they have not been catalogued. Data.—Two boxes, one with notebooks and one with index cards. The notebooks contain data on collection sites and meteo- rological data on Lake Sebago. They are labeled: 1904 Memoranda, 1904 Connecti- cut Lakes, 1905 Umbagog Lake, 1906 Se- bago Lake, 1907 Sebago Lake, 1908 Se- bago Lake, 1911 Sebago Lake, 1914 Lake Cooper, 1916 Miscellaneous (this contains collection data through 1921), and Miscel- laneous Notes on Freshwater Cladocera. There are four categories of index cards, only the first two of which can be correlated to data in the collection books: Locations of collections (presently grouped by state), some having lists of spe- cies found and a few with quantitative counts. Species names, listing locations where they were found. A small set of references on fish diet. A miscellaneous set of cards with data concerning material from other collectors and localities. Rufus W. Kiser Gift, NMNH 356548 R. E Kiser was a professor at Centralia Junior College, Washington, whose partic- ular interest was the taxonomy of Daphnia. He collected extensively throughout the Pa- cific Northwest, especially Washington and Oregon. His collection includes samples from Alaska to southern California made by Trevor Kincaid, a copepodologist at the 566 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON University of Washington. He also received South American material from Herman Kleerekoper, a Dutch limnologist who worked in Brazil for a number of years be- fore moving to Canada (J. Reid, pers. comm.). D. G. Frey became interested in Kiser’s collection and persuaded him to do- nate it to the NMNH (cf. the ‘“‘Kiser’’ file among Frey’s papers at the NMNH.) Wet material.—(In alcohol.) Over 8000 vials of unsorted zooplankton samples, stored in about 135 1-2 liter bottles filled with alcohol. All vials contain Kiser’s sam- ple number. Most bottles have an interior list of the vials they contain; outside they are identified by Kiser’s original bottle number. Most of the samples arrived at the NMNH stained by Kiser with Fast Green or Acid Fuchsin, dehydrated and in xylene in preparation for mounting specimens on slides. Others, that had been loaned to D. G. Frey, were in formalin. The formalin and xylene were replaced with alcohol at the NMNH. The bulk of the samples is from west coast states. There are a few samples from New Zealand, Mexico, Brazil (Kleereko- per’s samples), British Columbia, and the north-central U.S.A. states. The majority of the samples above 2500 are from synoptic sampling of Oregon lakes, especially Fern Lake. Slides.—One hundred, seventy-nine box- es of stained, balsam-mounted specimens. Most boxes are organized by genera; Daph- nia predominate. Six boxes have specimens from specific areas (China, Brazil, South America, New Zealand, and Scotland), one box has slides with University of Michigan Zoological Laboratory labels, one has mixed zooplankton from Oregon, and one has copepods. Most of the slides are well-labeled and nearly all bear the sample number from which the specimens came. They appear to be in excellent condition. Within the boxes, slides are not ordered by sample number or by species. Data.—Kiser’s typed index labeled “Plankton Collection of Rufus W. Kiser” giving sample number, location (including county and state or country), other pertinent data, and date. Kiser’s typed inventory of the vials con- tained in each bottle, stating if they were in xylene or formalin. A typed NMNH list of the location of each sample by sample number and bottle number. This list was made by reference to Kiser’s bottle inventory, and may be par- tially incorrect since some vials were mis- placed. Kiser’s sample numbering system started with 100 (cf. his letter of reply to a query from Frey, in the Frey “‘Kiser’’ correspon- dence file). Some slides have numbers less than 100, or are marked ***A (with data not corresponding to Kiser’s sample data). These specimens are from collectors other than Kiser, and there appear to be no sup- porting data for them. Kiser apparently had 46 bound note- books recording information on his sam- ples, and an accession file of 3 X 5 in. cards. (See letter of D. G. Frey to R. B. Manning, 28 May, 1983, in Frey’s corre- spondence.) These are not among the ma- terials at the MSC and Kiser may have kept them. There is no inventory of the slides in the Kiser collection. S. E Light Gift, NMNH 177850 S. E Light was a copepodologist at the University of California, Berkeley, where Mildred S. Wilson was his assistant for a time (Damkaer 1988). This cladoceran ma- terial has been separated from his larger zooplankton collection. It has specimens primarily from the western states as well as some from other localities and collectors (e.g., Wisconsin, from C. Juday). Wet material.—(n alcohol.) Five mason jars and one small bottle, filled with vials of cladoceran specimens sorted by genus and species. In 1983, these sorted cladoc- VOLUME 110, NUMBER 4 eran specimens were found on the shelves of the cladoceran collection but there was no record of them in the files. At that time some of the vials had dried out; they have since been rehydrated. They are not cata- logued. Data.—A list of collection stations and dates and one small box of file cards with SFL on the top; the cards bear sample num- bers and a list of cladoceran species in each. H. L. Shantz Gift In 1903, H. B. Ward, of the University of Nebraska, investigated lakes in the Pikes Peak, Colorado, and Lake Tahoe, Califor- nia, regions; E. A. Birge and C. D. Marsh contributed analyses of the cladocerans and copepods, respectively, in his collections (Ward 1904). H. L. Shantz, also of the Uni- versity of Nebraska, did a limnological study of some of the same lakes in the Pikes Peak area under Ward’s direction (Shantz 1907), but he did not analyze the cladoc- erans in his collections. G. S. Dodds (1924) later published a list of the cladocerans in Shantz’s samples. Shantz’s specimens were found in 1995 among Dodds’ material at the NMNH. These materials are of historic interest because in 1906 several of the Pikes Peak lakes were converted into reservoirs (Shantz 1907) and in 1995 I found that an- other, the type locale for the rare cladoceran Macrothrix montana Birge, has recently been drained. Wet material.—(Un alcohol.) One large, unidentified jar with vials of unsorted zoo- plankton samples; labels inside vials give lake names and collection dates. Jar is now temporarily labeled. Data.—None found. Shantz (1907) gives the lake locales and Dodds (1924) lists the cladocerans in the samples. Gideon S. Dodds Gift, NMNH 061153 G. S. Dodds worked at the University of Colorado. This collection represents an al- titudinal study of Branchiopoda in Colora- do lakes (Dodds 1917). Comparable mate- 567 rial may be in the Shantz Gift (see above) and in E. B. Reed’s material (see below). Wet material.—(In alcohol.) Three hun- dred and one small vials of sorted zoo- plankton specimens (cladocerans and co- pepods), each containing a sample number. From two to four vials, all from one local- ity, are stored together in small bottles, each bearing a catalogue number. Interior labels in the bottles list catalogue and sample numbers, approximate number of speci- mens, locality and county, date, and other pertinent data. Species names are listed on the reverse side of the interior labels. This material is housed in one large jar full of vials and in six boxes holding about 30 bottles each, not in strict order according to catalogue number. Data.—Dodds’ list of sample number, lo- cation (lake name), date, and county and terrain (mountain or plains). Other Sources of Cladoceran Materials The following, while not included in the cladoceran collection, might contain mate- rial useful to cladoceran researchers: The Albatross Expedition collection.— No NMNH accession number. This collec- tion includes freshwater samples made across the United States. The computerized database can be consulted to locate appro- priate material. The M. S. Wilson gifts —Several NNUNH numbers. The catalogue list and materials on the ““Copepod holding area’”’ shelves at the Natural History Building in Washing- ton, District of Columbia, should be con- sulted. Lots under NMNH 319629 might be most useful. The E. B. Reed gifts —Several NMNH numbers. The catalogue list and materials on the “Copepod holding area’’ shelves should be consulted. About 16 lots of un- identified cladocerans under NMNH 383116 have been catalogued. Reed’s col- lections are primarily from Alaska, the Rocky Mountains, and plains lakes near Fort Collins, Colorado. 568 The C. D. Marsh gift—NMNH 120079. There are containers marked “‘Entomostra- ca’’ on the “‘Copepod holding area”’ shelves that may contain cladocerans. Acknowledgments This work was made possible, in part, by Contract Number 95-22-UU-03728 from the NMNH. I thank E D. Ferrari, Research Zoologist, for making this study possible and for constructive comments on the manuscript, J. C. Walker and L. Ong for helping me locate materials, and J. W. Reid for background information on S. Wright, C. Juday, E. A. Birge, H. B. Merrill, and H. Kleerekoper. I appreciate the helpful suggestions of two anonymous reviewers. Literature Cited Berner, D. B. 1995. The cladoceran collection of the United States National Museum of Natural His- tory (The Smithsonian Institution)—Unpub- lished report, 19 pp. Beckel, A. L. 1987. Breaking new waters. A century of limnology at the University of Wisconsin.— Transactions of the Wisconsin Academy of Sci- ences, Arts, & Letters. Special issue, 122 pp. Birge, E. A. 1910. Notes on Cladocera IV.—Trans- actions of the Wisconsin Academy of Sciences, Arts, & Letters 16:1017—1066. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Damkaer, D. M. 1988. Mildred Stratton Wilson, co- pepodologist (1909—1973).—Journal of Crus- tacean Biology 8:131—-146. Deevey, E. S., Jr., & G. B. Deevey. 1971. The Amer- ican species of Eubosmina Seligo (Crustacea, Cladocera).—Limnology and Oceanography 16:201—218. Dodds, G. S. 1917. Altitudinal distribution of Ento- mostraca in Colorado.—Proceedings of the United States National Museum 54:59-87. . 1924. Notes on Entomostraca from Colorado. The Shantz collections from the Pikes Peak re- gion.—Proceedings of the United States Nation- al Museum 65:1-—7. Frey, D. G. 1963. Wisconsin: The Birge-Juday era. Pp. 3—54 in D. G. Frey, ed., Limnology in North America. The University of Wisconsin Press, Madison, 734 pp. Hartridge, M. L. 1995. H. B. Merrill: early Wisconsin scientist and adventurer.—Wisconsin Academy Review 41:16—22. Reid, J. W. 1991. The Stillman Wright collection of Copepoda (Crustacea) from South America in the National Museum of Natural History, Smithsonian Institution.—Proceedings of the Biological Society of Washington 104:736—741. Shantz, H. L. 1907. A biological study of the lakes of the Pikes Peak region-preliminary report.— Transactions of the American Microscopical So- ciety 27:75-98. Ward, H. B. 1904. A biological reconnaissance of some elevated lakes in the Sierras and the Rock- ies.—Transactions of the American Microscop- ical Society 25:127—154. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):569—580. 1997. Aglaodiaptomus atomicus, a new species (Crustacea: Copepoda: Calanoida: Diaptomidae) from freshwater wetland ponds in South Carolina, U.S.A., and a redescription of A. saskatchewanensis (Wilson 1958) Adrienne E. DeBiase and Barbara E. Taylor The University of Georgia, Savannah River Ecology Laboratory, P.O. Drawer E, Aiken, South Carolina 29802, U.S.A. Abstract.—A new calanoid copepod, Aglaodiaptomus atomicus, is de- scribed from freshwater wetland ponds in Aiken County, South Carolina, U.S.A. It is common in Aiken and Barnwell Counties. It has been collected from shallow, acidic wetland ponds, most of which dry periodically. Agla- odiaptomus atomicus resembles A. saskatchewanensis, which is redescribed from Saskatchewan, Canada, and Louisiana, U. S. A. The male of A. atom- icus differs from all of its congeners by possessing a distinctively large distolateral process on the right leg 5, exopod 1. The female differs from its closest congener, A. saskatchewanensis, in possessing nearly symmetrical thoracic wings. In A. saskatchewanensis, the wings are distinctly asymmet- rical. A survey of freshwater wetland ponds on the upper coastal plain of western South Carolina revealed an apparently unde- scribed calanoid copepod species. This spe- cies, which was incorrectly reported as Diaptomus conipedatus Marsh, 1907 by Mahoney et al. (1990), is one of the most common calanoid copepod species occur- ring in Carolina bays and other isolated, shallow, wetland ponds in Aiken and Barn- well Counties, South Carolina. It most closely resembles Aglaodiaptomus sas- katchewanensis Wilson, 1958. However, detailed comparisons with specimens of A. saskatchewanensis proved it to be a sepa- rate, undescribed species. In her original description of Agla- odiaptomus saskatchewanensis, Wilson (1958) provided an illustration of only the male fifth leg and a partial description of the species. Few additional details were added to her 1959 key (Wilson 1959). A more detailed description of A. saskatch- ewanensis is presented here, as well as a description of the new species, A. atomi- cus. Diaptomid copepods vary in the follow- ing morphological characteristics: body length, features of the thoracic somites (especially the 6" somite), length, number of and armature of the antennules in both sexes, number of somites of the female urosome, and segmentation, shape and or- namentation of the fifth leg of both sexes (Wilson 1959). The mouth parts and legs 1—4 of the species presented in this paper correspond to those of Leptodiaptomus si- ciloides (Comita & Tommerdahl 1960). Descriptions and measurements of whole copepods were made in glycerine. Dissect- ed specimens were mounted either in lac- tophenol or in CMC-9. Most were stained with Chlorazol Black E before being mounted. 570 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Family Diaptomidae Baird 1850 Genus Aglaodiaptomus Light, 1939 Aglaodiaptomus saskatchewanensis (Wilson 1958) Figs. 1-3 Diaptomus saskatchewanensis Wilson, 1958:490—491, 493-495, fig. 3.—Diap- tomus (Aglaodiaptomus) saskatchewan- ensis Robertson, 1972:202.—A glaodiap- tomus saskatchewanensis, Chengalath & Shih, 1994:2422. Material.—Saskatchewan: 8 66, 6 29; farm reservoir near Lucky Lake, Saskatch- ewan, Canada, 107°20'N, 50°10’W, 16 Jun 1948; USNM 210784; coll. J. R. Nursall. Louisiana: 33 66, 28 2°; shallow road- side pond 19.4 km south of Natchitoches, Natchitoches Parish, Louisiana, U.S.A., 31°60’N, 93°05’W, 10 Apr 1953; USNM 278233; coll. W. G. Moore. Female.—Length, excluding caudal se- tae, of specimens from Saskatchewan, range: 1.24—-1.64 mm (n = 6), from Loui- Siana, range: 1.34—1.64 mm (” = 28). Body broadest at pedigers 1 and 2 in dorsal view (Fig. 1A). Pedigers 4 and 5 incompletely separated with faint sutures visible laterally. Pediger 5 small with posteriorly-directed wings (Fig. 1B). Left wing with large dor- somedial lobe (Fig. 1C) reaching beyond posterior margin of wing in dorsal view (Fig. 1B). Right wing has 2 poorly-devel- oped lobes (Fig. 1D). Each lobe on both wings tipped with short sensillum. Urosome of 3 segments (Fig. 1B). Genital segment with slight asymmetrical lateral protrusions; left protrusion more rounded than right. Both protrusions tipped with sensillum. Caudal rami (Fig. 1B) nearly twice as long as broad; inner margins hairy. Antennules 25-segmented, extending to middle of caudal setae (Fig. 1A). Append- ages per segment as follows (Roman nu- meral = segment, Arabic numeral = num- ber of setae, sp = spine, a = aesthetasc): I(i;a), 1G +a)) MG ha), LV), Vicia), ViIGCL), -VITCt-+a), VIC + sp), IX(2+a), X(1), XI(2), X1(i+sp+a), XII(1), XIV (1+a), XV(1), XVI(1+a), XVII(1), XVII (1), XIX(1+a), XX(1), XXI(1), XXII(2), XXIII(2), XXIV(2), XXV(5+a). Setae on segments 17, 19, 20, 22 with hooked ends (Fig. 1E); length of seta approximately %4 that of respective segment. Leg 2 endopod 2 with Schmeil’s organ (not figured). Leg 5 (Fig. 1F): Coxa with posterior lat- eral protrusion ending in sensillum. Basis with short lateral seta. Exopod segments 1 and 2 approximately equal in length. Claw of exopod 2 slightly curved with denticles on inner and outer margins. Exopod 3 not articulated. Lateral spine of exopod 2 about % the length of outer seta of exopod 3. Ex- opod 3 inner seta plumose, twice the length of outer seta. Endopod single-segmented, equal in length to exopod 1, two plumose setae and pointed protrusion at tip (Fig. 1G). Male.—Length, excluding caudal setae, of specimens from Saskatchewan, range: 1.24-1.44 mm (n = 8), of specimens from Louisiana, range: 1.28-1.64 mm (n = 33). Body (Fig. 2A) as in female, with 5-seg- mented urosome. Thoracic wings (Fig. 2B) asymmetrical, right wing developed into elongated lobe, left wing small. Each wing with 2 sensilla; sensilla of right wing twice as large as left. Urosome (Fig. 2B): segment 1 asymmetrical in dorsal view, left margin shorter than right; left posterior corner bifid with obliquely directed lobe tipped with sensillum, right posterior corner simple, also tipped with sensillum. Segments 2—4 approximately equal in size; ventral sensilla on segments 2 and 3 visible under high magnification (not figured). Segment 4 asymmetrical, left margin shorter than right. Caudal rami with hairs on inner margins. Antennules (Fig. 2A, C—E) extend to proximal margin of caudal rami. Right an- tennule geniculate between segments 18-19 and 20—21. Segments 18-19 (Fig. 2D), 20- 21 (not figured) and 22—23 (Fig. 2E) fused. Large parallel spiniform processes on seg- ments 10, 11; additional large spine on seg- ment 13 (Fig. 2C). Spines on segments 10 VOLUME 110, NUMBER 4 Fig: 1: dorsal, distal segments of antennule indicated by arrow; B, Urosome, dorsal; C, Left thoracic wing and urosome, lateral; D, Right thoracic wing and urosome, lateral; E, Antennule, segments 19, 20; EK Leg 5, anterior, setae of exopod 3 (thin arrows), lateral seta of exopod 2 (thick arrow); G, Leg 5 endopod, distal end. and 11 equal in length, shorter than spine on segment 13. Spines on segments 15—17 with bifurcated tip (Figs. 2C, D). Segment 17 (Fig. 2D) with flat, digitiform seta ex- tending to center of segment 18—19. Addi- tional modified seta arising from proximal % of segment 18—19, running length of seg- ment. Segment 22—23 (Fig. 2E) with curved distal process; process about as long as seg- ment 24. Aglaodiaptomus saskatchewanensis Wilson, 1958. Paratype female (USNM 278233): A, Habitus, Left leg 5 (Figs. 3A—E): Leg (excluding spines) extending slightly beyond right ba- sis. Coxa, small process tipped with sensil- lum near outer posterodistal margin. Basis with short lateral seta. Exopod segment 1 longer than segment 2; inner margin with pad; distal corner of pad hairy (Fig. 3B). Exopod 2 articulating at distolateral margin of exopod 1. Inner margin of exopod 2 with hairs extending the length of the segment PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Aglaodiaptomus saskatchewanensis Wilson, 1958. Paratype male (USNM 278233): A, Habitus, dor- sal, distal segments of antennule indicated by arrow; B, Urosome, dorsal; C, Right antennule, segments 8—16; D, Right antennule, segments 16-19; E, Right antennule, segments 23-25. (Fig. 3B); hairs extend to cover % of ante- rior surface (Fig. 3C). Outer distal quadrant of exopod 2 covered with minute protuber- ances in both anterior and posterior views. Terminal seta about as long as segment and plumose, terminally placed on inner mar- gin; short, rounded digitiform process placed on outer distal margin, covered with short hairs. Endopod as long as exopod; outer margin of posterior face (Fig. 3D) with crenate longitudinal groove; remaining surface of endopod with small rows or clus- ters of short spinules. Outer *% of anterior face of endopod (Fig. 3E) crenulate; small pad on inner distal edge covered with tiny spinules. Right leg 5 (Fig. 3A): Coxa with disto- medial lobe on posterior surface tipped with sensillum. Basis with narrow, inwardly rounded longitudinal hyaline membrane on VOLUME 110, NUMBER 4 Fig. 3. 573 K < e fen a are eter, we ee Figs. 1-8. Argyrodiaptomus nhumirim, new species, holotype ¢ (MZUSP 12286): 1, Pedigers 4 and 5 and urosome, dorsal (in flattened mount); 2, Right antennule segments 8—15 (most setae omitted); 3, Right antennule terminal segments (most setae omitted); 4, Leg 5, caudal; 5, Pattern of tiny spinules on left leg 5 basipodite, frontal (drawn reversed); 6, Pattern of tiny spinules on right leg 5 basipodite, frontal (reversed); 7, Proximal section of terminal claw of right leg 5, frontal (reversed); 8, Left leg 5 exopodite and endopodite, frontal (reversed). Scales = 100 pm. 584 medial and frontal surfaces (Fig. 5). Left exopodite 1 with proximal haired pad little developed. Left exopodite 2 (Fig. 8) ap- pearing divided on frontal surface, with haired pad on proximal part of medial mar- gin, and ending in narrow, coarsely serrate digitiform process; frontal side with field of tiny denticles proximal to digitiform pro- cess; proximal spine minutely serrate, with hairlike tip, extending only slightly past tip of digitiform process. Left endopodite in- distinctly 2-segmented, with terminal row of 6 stout spinules. Female.—Length of allotype 1.8 mm, of paratype 2.0 mm. Body (Figs. 9-11) stout. Prosome without ornamentation except few pairs of pores, some with hairs. Rostral points (not illustrated) acute. Pedigers 4 and 5 fused, faint fusion line visible laterally only. Posterodorsal margins of pedigers 4 and 5 not elevated. Lateral wings of pediger 5 small, posteriorly directed, right wing slightly larger than left wing, each wing with small dorsal sensillum tipped with fine hair and larger ventral spiniform sensillum, wing widest posteriorly at level of ventral spiniform sensillum. Urosome of 3 seg- ments plus caudal rami. Genital compound segment nearly symmetrical, except right margin slightly produced posteriorly, and right spiniform sensillum placed slightly more anteriorly than left spiniform sensil- lum; left sensillum (Fig. 10) with acute bi- fid tip, right sensillum (Fig. 11) with blunt tip. Urosomite 2 slightly broader than long, proximal half telescoped into genital com- pound segment. Genital operculum (Fig. 12) with narrow crescentic proximal plate and broad distal plate with prominent lat- eral arms set wide apart (terminology after Cicchino 1994). Urosomites and caudal rami with several pairs of pores. Caudal rami about 1.7 times longer than broad, haired on medial margins only. Lateral and lateralmost terminal caudal setae stouter than more medial setae; dorsal seta slender; all setae finely and uniformly plumed. Antennules reaching just past anterior margin of genital compound segment. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Number of segments and armature as in left antennule of male. Antenna, mouthparts, and legs 1—4 as in male. Schmeil’s organ (Fig. 13) mammi- form, borne on caudal surface of leg 2 en- dopodite 2 (compare Fig. 55). Leg 5 (Figs. 14—16): coxopodite stout, with blunt prominence on caudal surface. Basipodite with long convex medial mar- gin, short lateral margin, and short seta at distolateral corner. Exopodite 1 about twice as long as broad, lateral margin slightly convex, medial margin uneven; with pore on caudal surface near midlength of medial margin. Exopodite 2 with small spine at base of exopodite 3, spine as long as exo- podite 3; claw coarsely serrate along most of both margins; claw of each foot curved anteriorly and slightly distally, left claw more strongly curved than right claw (Fig. 14). Exopodite 3 distinct from exopodite 2, with 2 terminal spines of which lateral spine is slightly more than half length of medial spine. Endopodite slightly longer than exopodite 1, unsegmented, bearing ter- minal row of hairs and 2 long, curved sub- terminal spines. Color of living specimens.—Light blue. Etymology.—Named for the Fazenda Nhumirim where the species was collected; proposed as a noun in apposition. Discussion and comparisons.—Argyro- diaptomus nhumirim falls morphologically and geographically among three congeners: A. denticulatus (Pesta, 1927), known from Argentina and Bolivia; A. azevedoi (Wright, 1935), from the Brazilian Northeast and Amazon Basin; and A. robertsonae Dussart, 1985a, from the Brazilian Amazon Basin. The male of A. nhumirim is easily distin- guished from that of A. denticulatus by sev- eral characters of the 5th leg, the latter spe- cies having prominent lobes on the right basipodite medial margin, a long lateral spine on the right exopodite 2, and a long spiniform proximal process on the left ex- opodite 2. The spiniform process on the an- tennular antepenultimate segment is long in A. denticulatus, reaching the end of the pen- VOLUME 110, NUMBER 4 585 9 Figs. 9-16. Argyrodiaptomus nhumirim, new species, allotype 2 (MZUSP 12287): 9, Habitus, dorsal; 10, Pedigers 4 and 5 and urosome, left lateral, with enlarged illustration of bifid spiniform sensillum; 11, Pedigers 4 and 5 and urosome, right lateral, with enlarged illustration of acute spiniform sensillum; 12, Genital operculum; 13, Leg 2 endopodite 2, caudal, showing Schmeil’s organ; 14, Left leg 5, lateral; 15, Right leg 5, caudal, with detail of endopodite; 16, Left leg 5 exopodites 2 and 3, caudal. Scales = 100 pm. 586 ultimate segment. Females of A. denticula- tus usually have a row of spines along the lateral part of the line of fusion between pedigers 4 and 5, although these spines are lacking in some populations (J. C. Paggi, in litt. 1997). Females of A. denticulatus con- sistently have the dorsal sensillum of each wing set on an expansion; a longer, more slender genital compound segment with the left and right spiniform sensilla set at the same level; and the leg 5 with a short, two- segmented endopodite, and no spine lateral to the base of exopodite 3 (after the rede- scription by Dussart 1985a). The male of A. robertsonae differs from that of A. nhumirim in having pedigers 4 and 5 fused, urosomite 1 (the genital so- mite) not inflated, the spiniform process on the right antennule segment 11 reaching the distal end of segment 12, and especially in having the right leg 5 basipodite with a large lobe on its caudal surface, and the dis- tal process of the right exopodite 1 directed laterally. The female of A. robertsonae dif- fers in having the lateral wings of pediger 5 most produced at the level of each dorsal spiniform sensillum, urosomite 2 much broader than long, and leg 5 exopodite 3 with a tiny lateroterminal spine, less than ¥, the length of the medioterminal spine (from Dussart 1985a:Pl. 2 Fig. 10). Argyrodiaptomus azevedoi is morpholog- ically closest to A. nhumirim, but the male of the former differs in a few characters of the right leg 5: there is a large lobe on the caudal surface of the basipodite, the caudal surface of exopodites 1 and 2 lacks pro- cesses, and the terminal claw is doubly an- gled with the middle part straight. These features are consistent in the representations of Wright (1935, 1938), Kiefer (1936), and Brandorff (1972), and were confirmed by inspection of a male from Betume (near Ne6épolis), State of Sergipe, Brazil, 13 March 1983, leg. E. R. dos Santos, identi- fied by C. E. EK da Rocha (USNM 227122). Wright (1935) stated that the left leg 5 en- dopodite of the male is one-segmented, but it is indistinctly divided in the specimen PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from Sergipe. In the female of A. azevedoi (confirmed by inspection of a female from USNM 227122), there is a lateral groove but not a suture line between the fused ped- igers 4 and 5, both spiniform sensilla of the genital compound segment are acute, the antennules extend to the posterior end of the genital compound segment, and the la- teroterminal spine of leg 5 exopodite 3 is less than % the length of the medioterminal spine. The armature of the antennules of the female and the left antennule of the male is identical in both species. Argyrodiaptomus azevedoi has been reported from the north- eastern Brazilian States of Bahia, Ceara, and Paraiba, and the Brazilian Amazon Ba- sin (States of Amazonas and Para). This is the first published record from Sergipe. Males of the remaining congeners, Ar- gyrodiaptomus bergi (Richard 1897), A. falcifer (Daday 1905) (=Diaptomus argen- tinus Wright, 1938), A. furcatus (G. O. Sars 1901) (=D. aculeatus Van Douwe 1911, 1912), A. furcatus f. exilis Dussart, 1985b, A. granulosus Brehm, 1933, A. macrochae- tus Brehm, 1937, and A. neglectus (Wright 1938) all lack denticles on the medial sur- face of the right leg 5 basipodite (although some have areas of minute granules on the caudal surface of this segment). In the fe- males of all these, the leg 5 endopodite is distinctly two-segmented, and except in A. macrochaetus and some populations of A. furcatus, it is much shorter than exopodite 1. All known species of Argyrodiaptomus are South American (Brandorff 1976). The Chinese Argyrodiaptomus ferus Shen & Tai, 1964 and A. cavernicolax Shen & Tai, 1965 were improperly assigned to this ge- nus (Dussart & Defaye 1983, Dussart 1985a). Both are in need of redescription and re-evaluation as to their generic affili- ation, which is beyond the scope of the present article. The Fazenda Nhumirim is maintained for agricultural and ecological research by the Empresa Brasileira de Pesquisa Agropecu- aria (EMBRAPA). It contains numerous permanent and ephemeral freshwater and VOLUME 110, NUMBER 4 saline ponds, locally termed “‘baias’’ and ‘“‘salinas”’ respectively. Of 19 localities in the southern Pantanal investigated by Reid & Moreno (1990), A. nhumirim appeared in only two small, shallow, fishless baias. Argyrodiaptomus macrochaetus Brehm, 1937, new rank Figs. 17-31 Argyrodiaptomus furcatus var. macrochae- tus Brehm, 1937:122-125, figs. 3, 4.— Dussart & Defaye 1983:131. Argyrodiaptomus furcatus macrochaetus.— Dussart 1984:63. Material examined.—é (fully dissected on slide), 2 (fully dissected on slide), and 2 2 and 2 copepodids, preserved whole in ethanol, from R. W. Kiser Collection, Sample 460SM/164/520 (USNM 283139). Accompanying copepod species: Attheyella fuhrmanni (Thiébaud; USNM 283146), Ec- tocyclops rubescens Brady (USNM 283140), Eucyclops ensifer Kiefer (USNM 283141), Mesocyclops longisetus s.s. (USNM 283142), Microcyclops alius (Kiefer; USNM 283143), Neutrocyclops brevifurca (USNM 283145), Paracyclops chiltoni (Thomson; USNM 283114), and Tropocy- clops prasinus (Fischer) s.s. (USNM 283144). d and 2 2 2, ethanol-preserved, Kiser Collection Sample 465B/198 (USNM 283123). Accompanying copepods: A. fuhr- manni (USNM 283126), Austrinodiaptomus kleerekoperi, new species (see following section), E. rubescens (USNM 283125), E. ensifer (USNM 283119), Macrocyclops al- bidus (Jurine) s.s. (USNM 283130), M. al- ius (USNM 283121), Microcyclops anceps (USNM 283122), Microcyclops cf. cei- baensis (USNM 283127), Microcyclops varicans (G. O. Sars) s.s. (USNM 283116), Microcyclops sp. (USNM 283118), P. chil- toni (USNM 283113), and Tropocyclops prasinus meridionalis (Kiefer; USNM 283117). Both samples from temporary pools near Porto Alegre, Rio Grande do Sul, Brazil, Sep 1941, leg. H. Kleerekoper. (Note: the Kiser Collection material arrived 587 at the National Museum of Natural History in xylene, and was transferred to ethanol about 1986.) The following description refers mainly to the specimens in hand, except where in- formation is added from the incomplete de- scription of Brehm (1937). Because of the age and dark color of the specimens, the prosomite pores could not be reliably ob- served in most individuals. Male.—Lengths in mm: 1.09 (USNM 283139), 1.2 (USNM 283123). Body (Fig. 17) slender in dorsal view. Pedigers 4 and 5 distinct. Lateral wings of pediger 5 nearly symmetrical, each with hairlike dorsal sen- sillum and long spiniform ventral sensillum on margin, right wing slightly longer and its ventral sensillum directed slightly more laterally. Right side of urosomite 1 slightly inflated and produced posteriorly; short hair-sensillum present on each posterior corner of urosomite 1. Metasomites, uro- somites, and caudal rami with paired pores (pore pattern may be incomplete as illus- trated). Caudal rami haired on medial mar- gins only. Antennules short, reaching posterior end of prosome. Right antennule (Figs. 18, 19) with 22 segments, geniculate between seg- ments 18 and 19. Segment 8 and (simple) segment 12 each with small socketed spine; spiniform processes of segments 10 and 11 nearly parallel to long axis of antennule, each reaching only to proximal end of suc- ceeding segment; spiniform process of seg- ment 13 large; segments 14-16 without processes; antepenultimate segment with stout curved process reaching nearly mid- length of segment 22 (Fig. 19). Left anten- nule (Fig. 20) with 25 segments; number of setae (Ss), spines (sp), and aesthetascs (a) of each segment as follows: (1) s + a, (2) 3s ian f3)is +a) (4) s::45).8 -7ea,.(6): s\°(7) s antes spr yas + a,.(10) ss, (2 1)-s, (12) s + sp + a, (13) s, (14) s + a, (15) s, (16)-s, G7)osy (18), s;°G19).s a, (20).:s, (21) s, (22) 2s, (23) 2s, (24) 2s, (25) 5s + a. Seta of segment 16 reaching past mid- length of segment 20, no aesthetasc at base 588 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 17-22. Argyrodiaptomus macrochaetus Brehm, 1937, 5 (17, USNM 283123; 18-22, USNM 283139): 17, Habitus, dorsal, with detail of left and right wings of pediger 5; 18, Right antennule segments 8—15 (setae omitted); 19, Right antennule terminal segments; 20, Left antennule segments 16—20 (most setae omitted); 21, Leg 5, caudal, showing Schmeil’s organ; 22, Right leg 5, lateral. Scales = 100 pm. VOLUME 110, NUMBER 4 of seta. Lengths of remaining setae similar to corresponding setae of female (Figs. 27, 28). Antenna, mouthparts, and legs 1—4 like corresponding structures of Austrinodiap- tomus kleerekoperi, new genus, new species (see following section). Schmeil’s organ as in female (compare Fig. 29). Leg 5 (Figs. 21, 22): each coxopodite with large spiniform sensillum on small, caudally directed lobe. Right basipodite with distal part of medial margin slightly lobate, and ornamented only with long lat- eral seta. Right exopodite 1 slightly longer than broad, with small blunt conical lobe on distolateral corner and long acute curved process on distomedial corner. Right exo- podite 2 with small rounded process at dis- tal *% of medial margin; lateral spine sub- terminal, about % length of segment, with serrate medial margin; terminal claw about as long as right basipodite and exopodite combined, gently curved with major cur- vature at distal %, and serrate along medial margin (tip of claw worn, thus claw may actually be somewhat longer and possibly recurved). Right endopodite short, cylindri- cal, indistinctly 2-segmented, with apical row of fine hairs. Left basipodite with prox- imomedial corner slightly dilated, and 3 small groups of denticles on medial surface (denticles not continuing onto caudal or frontal surface). Left exopodite 1 with small haired pad on medial surface; exopodite 2 with larger haired pad on medial surface, and ending in coarsely serrate digitiform process; proximal spine longer than digiti- form process, straight, with blunt tip and fine hairs along medial margin. Left endo- podite indistinctly 2-segmented, with ter- minal row of many fine hairs. Female.—Lengths in mm: 1.6, 1.8, 2.0 (USNM 283139); 1.5, 1.7 (USNM 283123). Body (Figs. 23—25) moderately stout. Ros- tral points (not illustrated) acute. Pedigers 4 and 5 completely fused, line of fusion vis- ible as lateral sulcus. Dorsal margins of pedigers 4 and 5 not elevated. Lateral wings of pediger 5 small, double, each with large 589 dorsal and smaller ventral spiniform sensil- lum on margin; right wing slightly longer than left wing, with larger sensilla. Uro- some of 3 segments plus caudal rami. Gen- ital compound segment long, nearly sym- metrical, except right anterior side slightly more produced than left side, and right sen- sillum located very slightly more posteri- orly than left sensillum. Both sensilla acute (Figs. 24, 25). Genital operculum (Fig. 26) with lateral arms slightly longer than in A. nhumirim, otherwise similar. Urosomite 2 broader than long, telescoped into and com- pletely covered ventrally by genital com- pound segment. Urosomites and caudal rami with several pairs of pores. Caudal rami haired on medial margins only. Antennule (Figs. 27, 28) reaching only posterior end of pediger 4. Number of seg- ments and armature as in left antennule of male, including segment 16 without aesth- etasc. Seta of segment 16 extending past end of antennule. Antenna, mouthparts, and legs 1—4 like those of male. Schmeil’s organ (Fig. 29) bluntly rounded, borne on caudal surface of leg 2 endopodite 2 (compare Fig. 55). Leg 5 (Figs. 30, 31): coxopodite stout, with acutely bifid prominence on caudal surface. Basipodite with long convex me- dial margin, short lateral margin, and long lateral seta. Exopodite 1 slightly more than twice longer than broad, with pore on cau- dal surface near midlength of medial mar- gin. Claw of exopodite 2 (of both 5th legs) broad and coarsely serrate along most of both straight margins; exopodite 2 also with slender spine at base of exopodite 3, this spine as long as and closely appressed to exopodite 3. Exopodite 3 distinct, with 2 stout terminal spines of which lateral spine is about half length of medial spine. En- dopodite reaching end of medial margin of exopodite 1, distinctly 2-segmented, bear- ing subterminal medial row of hairs and 2 long, straight subterminal spines, distal spine especially stout. Color of living specimens.—Notes by R. Thomsen, as reported by Brehm (1937:123, 590 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 23-29. Argyrodiaptomus macrochaetus Brehm, 1937, 2 (USNM 283139): 23, Habitus, dorsal; 24, Pedigers 4 and 5 and urosome, left lateral; 25, Pedigers 4 and 5 and urosome, right lateral; 26, Genital operculum; 27, Antennule segments 1-15; 28, Antennule segments 16-25; 29, Leg 2 endopodite 2, caudal, showing Schmeil’s organ. Scales = 100 pm. VOLUME 110, NUMBER 4 591 30 Figs. 30—40. Argyrodiaptomus macrochaetus Brehm, 1937, 2 (USNM 283139): 30, Leg 5, caudal; 31, Leg 5 endopodite, frontal. Figs. 32-40. Austrinodiaptomus kleerekoperi, new genus, new species, holotype d (MZUSP 12288): 32, Habitus, dorsal, with enlarged detail of lateral wings of pediger 5 (cephalosome broken); 33, Right antennule segments 8—16; 34, Right antennule terminal segments; 35, Leg 2 endopodite 2, caudal, showing Schmeil’s organ; 36, Right Leg 5, oblique caudal; 37, Leg 5, left exopodite 2, oblique caudal; 38, Leg 5, left exopodite, caudal; 39, Leg 5, left endopodite, frontal; 40, Right leg 5, endopodite and exopodite 1, frontal. Scales = 100 pm. 392 translated): “‘The antennas of all animals are marked with 2 or 3 colored bands, each band includes 2—3 segments. Thus in some males, which I have directly under the mi- croscope, on the antenna counting from the end, segments 6, 7, 8, 13 and 14 have a deep red coloration, which can be nearly black, while the other segments remain clear. Also the body shows these remark- able bands: segment 6 and 5 dark, 4 and 3 light, 2 again dark. A female is, in contrast, grass-green.”” Brehm (1937) confirmed this banding in his preserved specimens, but their color had changed to brownish red. The present, long-preserved specimens are light to dark brown with no trace of color banding. Discussion and comparisons.—Brehm (1937) gave a cursory description of A. fur- catus var. macrochaetus, with only two par- tial figures of the antennule and fifth leg of the female. Nevertheless, most characters mentioned by Brehm, including (in the male) the lack of spiniform processes on the right antennule segments 14 and 15, and the lateral spine of the right fifth leg exopodite 2 being much shorter than the segment, and (in the female) the long, strong subterminal spine of the leg 5 endopodite, and especial- ly the strikingly long seta of antennule seg- ment 16 agree exactly with the present specimens. Brehm (1937) stated that leg 5 endopodite of the female had three spines, but he probably mistook the rather stout proximalmost hairs of the subterminal hair row for a spine. In other respects, as observed by Brehm, A. macrochaetus does indeed resemble A. furcatus and A. furcatus f. exilis. Important differentiating characters of A. macrochae- tus include the long setae (especially on segment 16) of the antennule of both sexes; in males, the antennule lacking spiniform processes on segments 14 and 15 and with a long process on the antepenultimate seg- ment, and, in the fifth leg, the large spini- form sensilla on the coxopodites, the small proximomedial lobe of the left basipodite, the long acute distomedial process of the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON right exopodite 1, and the relatively short lateral spine of the right exopodite 2; in fe- males, the double lateral wings of pediger 5, and in the fifth leg, the bifid prominence of the coxopodite and the relatively long, two-segmented endopodite with two strong spines. These distinctive attributes fully jus- tify raising the taxon to species rank. Argyrodiaptomus macrochaetus has been reported only from the original collection, near the mouth of the La Plata River, Uru- guay. The find near Porto Alegre extends its range northeastwards. Brehm (1937) did not specify the habitat, except that it was in fresh water. Apparently this is a species of temporary pools. Cicchino (1994) pointed out the potential utility of the genital operculum in the tax- onomy of diaptomids. She described the genital operculum structure in 10 South American species belonging to 6 genera, not including Argyrodiaptomus. The oper- cula of A. nhumirim and A. macrochaetus differ from all these in having a narrow proximal plate and broad distal plate with prominent lateral arms. The shape of the distal plate and its lateral arms resembles most that of species of Notodiaptomus Kie- fer, 1936, but in Notodiaptomus the proxi- mal plate is about as broad as the distal plate, and subrectangular (Cicchino 1994). Austrinodiaptomus, new genus Diagnosis.—Diaptomidae, Diaptominae. Species of temporary ponds. Left antennule of male and both antennules of female with 2 setae on segment 11 and 1 seta on each of segments 13-19; no setae with hooked ends, but several setae of terminal segments stout, with blunt tips. Legs 1—4 with lateral spine on each exopodite segment, except leg 1 exopodite 2 lacking spine. Schmeil’s organ present on leg 2 endopodite 2. Male: urosomite 1 (genital somite) with small hair-sensillum on right distal margin. Right antennule with socketed spine on each of segments 8 and 12, and spiniform process on each of segments 10, 11, 13, 15, and 16; VOLUME 110, NUMBER 4 antepenultimate segment with hyaline membrane along nearly entire length, but without process. Right leg 5, coxopodite not markedly expanded medially, caudal surface with large, distally directed protru- sion bearing spiniform sensillum; basipo- dite produced posteriorly, ornamented only with tiny granules; endopodite of 1 short segment; exopodite 1 short and broad, ex- opodite 2 slightly expanded distally, bear- ing subterminal lateral spine and long slen- der terminal claw. Left leg 5, coxopodite with small protrusion bearing spiniform sensillum; basipodite slender, lacking sur- face ornament except lateral seta; endopod- ite of 2 short segments; exopodite 1 more than twice as long as exopodite 2, bearing medial haired pad; exopodite 2 narrow, with medial haired pad and ending in short stubby digitiform process and slightly lon- ger proximal spine. Female: Pedigers 4 and 5 distinct, without dorsal process; lateral wings double, nearly symmetrical, each with 2 large spiniform sensilla. Urosome of 3 segments plus caudal rami, urosomite 2 short. Leg 5, coxopodite with large spini- form sensillum borne on conical protrusion on caudal surface, and with or without ac- cessory spiniform process on anterolateral surface; basipodite with lateral seta inserted directly on segment, not on protrusion; ex- opodite 1 usually without ornament except for medial pore (1 specimen of A. kleere- koperi with lateral spiniform process on 1 leg), exopodite 2 with lateral spine, and ex- opodite 3 distinct, with 2 terminal spines; endopodite shorter than or equal to exopo- dite 1, with 2 or 3 short subterminal spines and subterminal oblique row of hairlike spi- nules. Etymology.—From the Latin austrinus, south, prefixed to Diaptomus; gender mas- culine. Type species.—Austrinodiaptomus _ kle- erekoperi, new species. Additional species.—Austrinodiaptomus inexspectatus (Brehm 1958), new combi- nation. Discussion and comparisons.—A com- 593 bination of several characters makes it im- possible to assign the new species to any existing genus of the family, particularly the several genera that include medium to large diaptomids of temporary ponds. These characters are, in both sexes, the two setae on antennule segment 11 and one seta on segments 13-19, the lack of hooked se- tae and the presence of stout blunt setae on some segments of the antennule, and the presence of Schmeil’s organ on leg 2 en- dopodite 2. Characters of the male include the right antennule without a spiniform pro- cess on segment 14 or a process on the an- tepenultimate segment, and in leg 5, the short endopodites, the narrow left exopod- ite, and the right coxopodite without a large medial expansion. Characters of the female include the lack of a dorsal projection on pedigers 4 or 5, and in leg 5, the short stout spines on the endopodite, the presence of a lateral spine on exopodite 2, and the distinct exopodite 3. Of the South American diaptomid gen- era, several (Argyrodiaptomus, Colombod- taptomus Gaviria 1989, Dasydiaptomus De- faye & Dussart, 1993, Notodiaptomus Kie- fer 1936, Prionodiaptomus Light 1939, Rhacodiaptomus, and Scolodiaptomus Reid 1987) possess only 1 seta on antennule seg- ment 11, among other differences. Other genera for which the antennular setation is undescribed (Calodiaptomus Kiefer 1936, Dactylodiaptomus Kiefer 1936) differ in having the male with spiniform processes on segment 14 and the antepenultimate seg- ment of the night antennule, and the female with leg 5 exopodite 3 fused to exopodite 2, among other features. In Aspinus Bran- dorff, 1973b, exopodite 1 of legs 1—4 lacks a lateral spine. Females of /diodiaptomus Kiefer, 1936 have one subterminal spine on the leg 5 endopodite; males possess a spi- niform process on right antennule segment 14. Females of Odontodiaptomus Kiefer, 1936 have single pediger 5 wings and the leg 5 endopodite with only one subterminal spine; males have the right leg 5 endopodite long, and complex lobing on the basipodite 594 and exopodite. In females of Tumeodiap- tomus Dussart, 1979 the urosomite 2 is lon- ger than broad and expanded, and in males there is a pectinate process on the antepen- ultimate segment of the antennule and the lateral spine of the right leg 5 exopodite 2 is inserted in the proximal half of the seg- ment. Therefore, the new genus Austrino- diaptomus is proposed to accommodate the new species and the similar D. inexspecta- tus. Austrinodiaptomus kleerekoperi, new species Figs. 32—59 Diaptomus s.]. inexspectatus Brehm, 1958.—Brandorff 1972:50 (partim). Brandorff 1973a:342 (partim). Diaptomus s.l. inexpectatus.—Brandorff 1976:618 (partim).—Dussart & Defaye 1983:64 (partim).—Dussart 1984:64 (partim).—Battistoni 1995:958 (partim). Rhacodiaptomus inexspectatus.—Brehm 1965:3, 11-14, fig. 1 (partim). Material examined.—¢é holotype (MZUSP 12288) and 2 allotype (MZUSP 12289), each fully dissected on slide. Paratypes: 2 2¢, ethanol-preserved (MZUSP 12290); 2, dissected on slide, and 2 2 (1 broken) and 1 copepodid, ethanol-preserved (USNM 283124). All from R. W. Kiser Collection, Sample 465B/198, temporary pool near Porto Alegre, Rio Grande do Sul, Brazil, Sep 1941, leg. H. Kleerekoper. Ac- companying fauna: see description of A. macrochaetus. Male.—Length 1.2 mm. Body (Fig. 32) slender. Rostral points (not illustrated) acute. Pedigers 4 and 5 distinct. Lateral wings of pediger 5 small, nearly symmet- rical, each with dorsal hair-sensillum and ventral spiniform sensillum, sensilla of right wing slightly larger. Right side of uro- somite 1 slightly inflated and posteriorly produced, with short sensillum on distal corner. Right posterior corner of urosomite 4 produced in triangular process. Cephalo- some broken; metasomites, most uroso- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mites, and caudal rami with paired pores. Caudal rami haired on medial margins only. Antennules reaching posterior end of anal somite. Right antennule (Figs. 33, 34) with 22 segments, geniculate between seg- ments 18 and 19. Segments 8 and 12 each with small socketed spine; slender spini- form processes of segments 10 and 11 ex- tending slightly outward from long axis of antennule and the latter well past base of next process; process of segment 13 large, with tiny subterminal hook; segment 14 without process, but segments 15 and 16 each with small spiniform process, process of segment 16 about twice as long as that of segment 15; antepenultimate segment lacking distal process, but with broad hya- line membrane extending along nearly en- tire length of segment (Fig. 34). Left anten- nule as in female. Seta on segment 16 bro- ken, but reaching at least end of segment 19. Schmeil’s organ (Fig. 35) mammiform, located on caudal surface of leg 2 endopo- dite 2 (compare Fig. 55). Leg 5 (Figs. 36-41): each coxopodite with large sensillum on lobe on caudal sur- face, left lobe small, right lobe large and produced distally (Fig. 41). Right coxopo- dite with distomedial margin not much ex- panded. Right basipodite with proximal part of medial margin slightly expanded, and distal part of caudal surface expanded, with field of tiny granules; lateral seta inserted at distal %. Right exopodite 1 slightly broader than long, lateral margin about twice as long as medial margin, and disto- lateral corner produced in small, distally di- rected mammiform swelling. Right exopod- ite 2 lacking notable surface ornamentation, with distal half of medial margin expanded, broken lateral spine inserted subterminally, and partly serrate terminal claw, gently curved with recurved tip. Right endopodite represented by short conical protrusion with row of hairs along medial surface. Left bas- ipodite long, proximomedial corner slightly expanded, surface smooth except for lateral seta inserted at distal %4. Left exopodite VOLUME 110, NUMBER 4 395 Al 42-44 Figs. 41-45. Austrinodiaptomus kleerekoperi, new genus, new species; 41, holotype 6 (MZUSP 12288): 41, Right leg 5, lateral; 42—45, allotype 2 (MZUSP 12289): 42, Habitus, dorsal, showing tip of antennule and enlarged details of wings of pediger 5; 43, Posterior pedigers and urosome, right lateral; 44, Posterior pedigers and urosome, left lateral; 45, Genital operculum. Scales = 100 wm. 596 (Fig. 38) of 2 segments, exopodite 1 about twice as long as segment 2 and bearing me- dial haired pad; exopodite 2 bluntly quad- rate, with field of tiny hairs on medial sur- face, medial pad with long hairs, slender, curved, naked proximal spine, and short, stubby, finely haired digitiform process. Left endopodite (Fig. 39) of 2 segments, en- dopodite 2 with row of hairs along medial surface. Female.—Lengths of allotype 1.75 mm, of paratypes 1.72, 1.72, and 1.80 mm. Body (Figs. 42—44) stout; metasomites and uro- some with several pairs of dorsal pores. Rostral points (not illustrated) acute. Pedi- gers 4 and 5 distinct. Pedigers 4 with pair of thick transverse bars near anterodorsal margin (Fig. 42) and slightly elevated pos- terodorsal margin (Figs. 43, 44). Lateral wings of pediger 5 large, approximately symmetrical, double, each wing with 2 large spiniform sensilla, dorsal sensillum ventrally curved (Figs. 43, 44). Urosome of 3 segments plus caudal rami. Anterior half of genital compound segment much ex- panded, conical right expansion directed dorsally and posteriorly, left expansion spherical, each with large acute spiniform sensillum; right posterior margin of seg- ment extended in shallow trapezoidal pro- cess (Fig. 43). Genital operculum (Fig. 43). Genital operculum (Fig. 45) with narrow crescentic proximal plate, broad distal plate, and broad, prominent lateral arms set wide apart. Urosomite 2 broader than long, but narrower than other urosomites. Anal so- mite broadened posteriorly. Caudal rami haired on medial margins only. Antennule (Figs. 46—48) reaching caudal rami, with 25 segments. Number of setae (s), spines (sp), and aesthetascs (a) of each segment as follows: (1) s + a, (2) 3s + a, (3)ysese a, G4): SO) so+: al) sG)vs = a; (8) s + sp, (9) 2s + a, (10) s, (11) 2s, (12) s + sp + a, (13) s, (14) s + a, (15) s, (16) s + a, (17) s, (18) s, (19) s + a, (20) s, (21) Sy(22): 28, O3)i 2s, 424) 2se4(25 2Ssu aia: Seta of segment 16 reaching end of anten- nule. Larger setae of segments 18, 21, and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 23-25 stout, with blunt tips; smaller setae of segments 22, 23, and 25 short and slen- der. Antenna, mandible, maxillule, maxilla, and maxilliped (Figs. 49-53) of usual struc- ture in family. Leg 1 (Fig. 54) with 3-segmented exo- podite and 2-segmented endopodite, exo- podite 2 lacking lateral spine. Leg 2 (Fig. 55) with both rami 3-segmented and bluntly rounded Schmeil’s organ (Fig. 56) borne on caudal surface of endopodite 2. Legs 3 and 4 (not illustrated) similar to leg 2 except slightly larger and lacking Schmeil’s organ. Leg 5 (Figs. 57-59): coxopodite stout, with large spiniform sensillum borne on coniform expansion on caudal surface, and spiniform process on anterolateral surface (indicated by arrow in Fig. 59). Basipodite with lateral seta inserted directly on seg- ment, not on expansion. Exopodite 1 in most specimens without ornament except medial pore; allotype female with lateral spiniform process on 1 leg, as in detail of Fig. 57. Exopodite 2 with narrow lateral spine reaching midlength of exopodite 3, and stout claw serrate along most of each margin. Exopodite 3 distinct, with 2 stout terminal spines, lateral spine half length of medial spine. Endopodite as long as exo- podite 1, indistinctly divided at proximal %, with 2 short subterminal spines and oblique subterminal row of fine hairs. Etymology.—This species is dedicated to Herman Kleerekoper, whose early investi- gations of southern Brazilian continental waters formed much of the basis for the subsequent development of the science of limnology in that country. Discussion and comparisons.—Diapto- mus S.1. inexspectatus was incompletely de- scribed from Argentinian material by Brehm (1958, repeated 1960). Later, Brehm (1965) made additional comments on sup- posed D. inexspectatus from old manuscript notes on female specimens collected from Las Garcias, Corrientes, Argentina, and (by H. Kleerekoper) near Porto Alegre, Brazil, and gave a sketch of a female from Porto VOLUME 110, NUMBER 4 597 Figs. 46-50. Austrinodiaptomus kleerekoperi, new genus, new species; 46—48, allotype 2 (MZUSP 12289); 49, 50, paratype 2 (USNM 283124): 46, Antennule segments 1-17; 47, Antennule segments 16—23; 48, Anten- nule segments 24—25; 49, Antenna; 50, Mandible. Scales = 100 pm. 598 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 51-59. Austrinodiaptomus kleerekoperi, new genus, new species; 51-55, paratype 2 (USNM 283124); 56-59, holotype 2 (MZUSP 12289): 51, Maxillule; 52, Maxilla; 53, Maxilliped; 54, Leg 1, frontal; 55, Leg 2, frontal; 56, Leg 2 endopodite 2, caudal, showing Schmeil’s organ; 57, Leg 5, caudal, with detail of aberrant spiniform process on exopodite 2, found on one specimen; 58, Leg 5 endopodite, caudal; 59, Leg 5 coxa- basipodite, left lateral. Scales = 100 pm. VOLUME 110, NUMBER 4 Alegre. Brehm (1965) claimed that his specimens from Brazil corresponded exact- ly with the Argentinian species. Certainly, Brehm’s descriptions of the female of D. inexspectatus agree with the Brazilian spec- imens in hand regarding the distinctive dou- ble wings of pediger 5, the lobate genital compound segment, and certainly the leg 5 exopodite structure and ornament. Howev- er, there are several discrepancies. Accord- ing to Brehm’s (1958, 1960) descriptions, in females of the Argentinian population, the antennules reach the end of the caudal setae, the leg 5 endopodite is about *% the length of exopodite 1 and bears three spines, the lateroterminal spine of exopo- dite 3 is about % as long as the medioter- minal spine, and Brehm saw no spiniform sensillum on the left expansion of the fe- male genital compound segment. Brehm’s (1958, 1960) representations of the right leg 5 of the Argentinian male correspond with some features of the Brazilian specimen, such as the general proportions of the seg- ments and terminal claw, the large coxo- podite lobe, and the placement of the lateral exopodite spine. However, other features such as the right basipodite and exopodite 1 processes, and especially the left fifth leg exopodite do not correspond at all. Similar- ly, Brehm described the male right anten- nule as having a long segment 15 spiniform process (% the length of the process of seg- ment 13). Even though Brehm had difficul- ty in describing the single, poorly preserved male, these discrepancies are difficult to ex- plain as lapses of observation, and appear significant at the species level. Therefore it seems justifiable to propose a new taxon for the Brazilian population, pending redescrip- tion of the Argentine species and improve- ments in knowledge of variation. Brehm (1965) transferred D. inexspecta- tus to the genus Rhacodiaptomus Kiefer, 1936, on the basis of the lobate genital compound segment of the female, while ad- mitting that the structure of the male leg 5 did not agree with the diagnosis of Rhaco- diaptomus. Subsequent authors (Brandorff 599 1972, 1973a, 1976; Dussart & Defaye 1983; Dussart 1984; Battistoni 1995) pre- ferred to retain the species in Diaptomus Westwood, 1836 s.]. Acknowledgments I thank I. H. Moreno for his invitation to visit the southern Pantanal in 1987, and his assistance in collecting, together with G. Morao and [. H. Ishii, then at the Departa- mento de Ciéncias, Universidade Federal do Mato Grosso do Sul (UFMS), Corumba4, and Z. M. Silva Campos and members of the staff of the Fazenda Nhumirim. Travel and logistical support was provided by the Universidade Federal do Mato Grosso do Sul, the Empresa Brasileira de Pesquisa Agropecuaria, and the Organization of American States. The manuscript was much improved by comments from E D. Ferrari, J. C. Paggi, and three anonymous review- ers. As always I am grateful to the Depart- ment of Invertebrate Zoology, National Mu- seum of Natural History, for providing re- search facilities. Literature Cited Baird, W. 1850. The natural history of the British En- tomostraca. The Ray Society, London, 364 pp. + pls. 1-36. Battistoni, P A. 1995. Crustacea Copepoda. Pp. 953-— 971 in E. C. Lopretto & G. Tell, eds., Ecosis- temas de Aguas Continentales: Metodologias para su Estudio. Vol. 3, Ediciones Sur, La Plata. Brandorff, G.-O. 1972. Ein Beitrag zur Calanidenfau- na (Crustacea Copepoda) des Amazonasgebie- tes, mit einem Uberblick iiber die Diaptomidae (Crustacea Copepoda) Sulamerikas. Unpub- lished Diplomarbeit, Max-Planck-Institut fiir Limnologie, Plon, 57 pp. 1973a. Die neotropischer Gattung Rhaco- diaptomus Kiefer (Crustacea, Copepoda), mit der Beschreibung von zwei neuen Arten.— Amazoniana 4:341-—365. 1973b. Neue freilebenden calanoide Cope- poden (Crustacea) aus dem Amazonasgebiet.— Amazoniana 4:205-—218. 1976. The geographic distribution of the Diaptomidae in South America (Crustacea, Co- pepoda).—Revista Brasileira de Biologia 36: 613-627. Brehm, V. 1933. Argyrodiaptomus granulosus nov. 600 spec., ein neuer Diaptomus aus Uruguay.— Zoologischer Anzeiger 104:283—287. 1937. Weitere Mitteilungen tiber die Siis- swasserfauna Uruguays. I. Teil.—Zoologischer Anzeiger 120:120—125. 1958. Bemerkungen zu einigen Kopepoden Siidamerikas.—Sitzungsberichten der Osterrei- chischen Akademie der Wissenschaften, Math- ematische-naturwissenschaftliche Klasse, I, 167:139-171. 1960. Sobre los copepodos hallados por el Profesor Biraben en la Argentina. 67. Comuni- cacion.—Neotr6dpica 6(20):49—54. . 1965. Bericht tiber eine unvollendet geblie- bene Untersuchung der Argentinischen Kope- podenfauna.—Sitzungsberichten der Osterrei- chischen Akademie der Wissenschaften, Math- ematische-naturwissenschaftliche Klasse, I, 174:1-15. Cicchino, G. 1994. Importance of the genital oper- culum in the taxonomy of South American Diaptomidae (Copepoda, Calanoida).—Hydro- biologia 292/293:143—-147. Daday, E. von. 1905. Untersuchungen tiber die Siis- swasser-Mikrofauna Paraguays.—Zoologica 18(44):1-374 + pls. 1-23. Defaye, D., & B. H. Dussart. 1993. Dasydiaptomus nom. nov. for Trichodiaptomus (Copepoda, Cal- anoida, Diaptomidae).—Hydrobiologia 257: 2 Ts Dussart, B. H. 1979. Algunos copépodos de América del Sur.—Publicacion Ocasional del Museu Na- cional de Historia Natural, Santiago de Chile 30:3-13. . 1984. Some Crustacea Copepoda from Ven- ezuela.—Hydrobiologia 113:25—67. 1985a. Another new diaptomid (Crustacea, Copepoda) from the Brazilian Amazon.—Ama- zoniana 9:275—280. . 1985b. Sur quelques copépodes d’ Amérique du Sud. V. Diaptomidae.—Archiv fiir Hydro- biologie 103:201—215. , & D. Defaye. 1983. Répertoire mondial des crustacés copépodes des eaux intérieures. I.— Calanoides. Editions du Centre National de la Recherche Scientifique, Paris, 224 pp. Gaviria, S. 1989. The calanoid fauna (Crustacea, Co- pepoda) of the Cordillera Oriental of the Co- lombian Andes.—Hydrobiologia 178:113—134. Kiefer, E 1932. Versuch eines Systems der Diaptom- iden (Copepoda, Calanoida).—Zoologisches Jahrbuch fiir Systematik 63:451—520. . 1936. Uber die Systematik der siidamerikan- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ischen Diaptomiden (Crustacea, Copepoda).— Zoologischer Anzeiger 116:194—200. Light, S. FE 1939. New American subgenera of Diap- tomus Westwood.—Transactions of the Ameri- can Microscopical Society 58:473—484. Pesta, O. 1927. Ein Beitrag zur Kenntnis der Cope- podenfauna von Argentinien.—Zoologischer Anzeiger 73:67—80. Reid, J. W. 1987. Scolodiaptomus, a new genus pro- posed for Diaptomus (sensu lato) corderoi Wright, and description of Notodiaptomus bran- dorffi, new species (Copepoda, Calanoida), from Brazil.—Journal of Crustacean Biology 7: 364-379. , & I. H. Moreno. 1990. The Copepoda (Crus- tacea) of the southern Pantanal, Brazil—Acta Limnologica Brasiliensia 3:721—739. Richard, J. 1897. Sur quelques entomostracés d’eau douce des environs de Buenos Aires.—Anales del Museo Nacional de Buenos Aires 5:321— 332. Sars, G. O. 1901. Contributions to the knowledge of the fresh-water Entomostraca of South America as shown by artificial hatching from dried ma- terial. II. Copepoda—Ostracoda.—Archiv for Mathematik og Naturvidenskab 24:3—52 + pls. 1-8. . 1903. An account of the Crustacea of Nor- way. 4. Copepoda Calanoida. Bergen Museum, Bergen, 171 pp. + pls. 1-102, suppl. pls. 1-6. Shen, C. J., & A. Y. Tai. 1964. Descriptions of eight new species of freshwater Copepoda (Calanoi- da) from the Delta of the Pearl River, South China.—Acta Zoologica Sinica 16:225—246. (in Chinese; English summary.) 1965. Descriptions of six new species of freshwater copepods chiefly from the Pearl Riv- er Delta, South China.—Acta Zootaxonomica Sinica 2:126—-140. (in Chinese; English sum- mary.) Van Douwe, C. 1911. Neue Stisswasser-Copepoden aus Brasilien.—Zoologischer Anzeiger 37:161— 163. 1912. Zur Kenntnis der Stisswassercopepo- den von Brasilien.—Archiv fiir Hydrobiologie 7:309-321. Westwood, J. O. 1836. Cyclops (Muller).—British Cyclopaedia of Natural History 2:227—228. Wright, S. 1935. Three new species of Diaptomus from Northeast Brazil—Annaes da Academia Brasileira de Sciencias 7:213—233 + pls. 1—4. 1938. A review of the Diaptomus bergi group, with descriptions of two new species.— Transactions of the American Microscopical So- ciety 57:297-315. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):601—607. 1997. Lophogaster muranoi, a new species of mysid from the coastal waters of Argentina (Crustacea: Mysidacea: Lophogastridae) Kouki Fukuoka, Monica S. Hoffmeyer, and Maria D. Vifias (KF) Department of Aquatic Biosciences, Tokyo University of Fisheries, 4-5-7 Konan, Minato-ku, Tokyo 108, Japan; (MH) Instituto Argentino de Oceanografia, Avenida Alem 53, 8000 Bahia Blanca, Argentina; (MV) Instituto Nacional de Investigacion y Desarrollo Pesquero, C.C. 175, 7600 Mar del Plata, Argentina Abstract.—Lophogaster muranoi, a new species, is described from the coast- al waters of Argentina. This new species is characterized by the medium-sized median rostral spine extending slightly beyond the apex of the antennular lobe, the presence of a pair of the postorbital spines on the carapace, and the telson armed with 4 pairs of lateral spines and a narrow apical median plate. This is the third species of the genus Lophogaster from the South Atlantic Ocean. The genus Lophogaster comprises 18 species and 1 subspecies to date, all from tropical to temperate waters of the world. Among them, only two species, L. spinosus Ortmann, 1906 and L. challengeri Fage, 1940, have been represented in the South Atlantic Ocean, the former was recorded from mid-Atlantic waters (Fage 1942) and off Rio de Janeiro (O. Tattersall 1955) and the latter from coastal waters of South Af- rica (G. Sars 1885, Fage 1942, O. Tattersall 1955) and Cape of Lopez, West Africa (O. Tattersall 1955). An undescribed species of Lophogaster collected from coastal waters of San Matias Gulf and Rio de la Plata estuary, Argentina, is the first species of the genus from Ar- gentine waters. In this paper a description of the new species is given. The type specimens are deposited in the National Science Museum, Tokyo (NSMT). Lophogaster muranoi, new species Figs. 1-4 Type _ series.—Holotype (NSMT-Cr 11994), adult male (19.2 mm, tip of the me- dian spine of the rostrum to the posterior end of the telson except apical spines); al- lotype (NSMT-Cr 11995), adult female (18.8 mm); paratypes (NSMT-Cr 11996), 1 adult male (18.0 mm) and 1 adult female (16.0 mm); San Matias Gulf (42°09’S, 64°32'W); depth 170 m; 11 to 22 May £99}: Other material.—1 adult male (16.0 mm), | adult female (16.0 mm), 1 immature male (13.4 mm) and 2 immature females (13.0 and 14.4 mm); same as type speci- mens. 1 adult male (15.8 mm), 2 adult fe- males (13.4 and 16.0 mm), 1 immature male (13.0 mm) and 3 juveniles (7.8-9.6 mm); Rio de la Plata estuary (35°32’S, 53°18’W); depth 52 m; 12 Oct 1995; col- lected with a Bongo net, 45-0 m oblique tow by the R/V Oca Balda of the INIDEP, Mar del Plata, Argentina. Description.—Carapace covered with minute tubercles on anterior half of dorsal surface; rostrum covering eyestalks and proximal 2 segments of antennular pedun- cles, tridentate anteriorly, median process medium-sized extending slightly beyond apex of antennular lobe, lateral spines short; lateral margin of rostrum concave, minutely serrate (Fig. 1A, B); posterior margin of carapace deeply emarginate, leaving eighth or seventh and eighth thoracic somites ex- posed dorsally, furnished dorsally with fine 602 setae; postero-lateral angles of carapace ter- minating in comparatively long, slightly up- ward spine (Fig. 4C); pair of prominent postorbital spines present (Fig. 1A, B). Antennular peduncle robust; first segment as long as broad; second segment short, with outer distal corner produced and tipped with 2 plumose setae; third segment slightly shorter than first and second segments com- bined, distal margin produced into 2 blunt processes with spinules frontally (Fig. 1A, B); antennular lobe from third segment with finely serrated and rounded anterior margin with slight median depression armed with 2 unequal setae (Fig. 1C). Antennal scale cordiform, 1.8 times as long as widest part; outer margin slightly convex, armed with 5 denticles in holotype and 6 in allotype on distal 0.7, terminating in long, straight, acute spine; inner margin setose (Fig. 1B, D). Antennal peduncle com- posed of 3 segments; first segment with acutely pointed small process at inner distal corner, third segment longer than proximal 2 segments combined, inner distal corner end- ing in long, outwardly curved spine (Fig. 1D). Labrum triangular, produced anteriorly into acute spinose process. Mandibular palp long and slender, second segment longest, 6 times as long as broad, third segment 0.7 of second segment in length, armed densely with short setae on distal 0.8 (Fig. 1E). Maxillule with outer lobe armed with 13 strong spines on terminal margin and 1 long seta on basal part, inner lobe rounded with 7 long and 8 short setae (Fig. 1F). Maxilla with exopod oval, armed with many plu- mose setae on whole margin, second seg- ment of endopod twice as long as broad at its base (Fig. 1G). Each sternum of second to seventh tho- racic somites with 2 acutely pointed spine- like processes on midline; anterior one short, posterior one long and forward-di- rected. First thoracic limb short and robust; exo- pod short, leaf-like, 2-segmented, without setae, endopod with expanded merus and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON carpopropodus armed with 3 stout setae each on inner margin in addition to slender setae, dactylus with stout claw; epipod large (Fig. 2A). Second thoracic limb slender; exopod short, about half length of endopod; endopod setose on inner margin, dactylus with small curved claw (Fig. 2B). Third and fourth tho- racic limbs long and slender, exopod similar to that of second limb (Fig. 2C, E); endopod of third limb with dactylus armed with 3 hooked spines on distal half and 3 ordinary spines on proximal half of inner margin (Fig. 2D); endopod of fourth limb with me- rus armed densely with short and slender spines on distal half of inner margin (Fig. 2F), dactylus without spines as seen in third limb. Fifth thoracic limb robust, exopod 14- segmented; endopod with merus longest, 1.2 ‘times longer than carpopropodus and dac- tylus combined (Fig. 2G). Sixth to eight tho- racic limbs stout, exopod 9-16-segmented; endopod with merus short, 0.6 as long as carpopropodus (Fig. 3A—C). Marsupium of female composed of 7 pairs of brood lamellae. Each abdominal somite with well-devel- oped pleural plate; first to fifth somites with short, acutely pointed, median process; sixth somite armed posteriorly with pair of sharp denticles about half length of apical spines of telson (Fig. 4A). Pleopods in both sexes well-developed and biramous; exopods 23—26 segments, longer than 13-19 segments of the endo- pods (Fig. 3D—-H). Uropods shorter than telson; exopod un- divided, outer margin naked, terminating in tooth, inner margin setose; endopod slightly longer than exopod, both inner and outer margins setose (Fig. 4A). Telson 2.2 times longer than last abdom- inal somite, about 3 times as long as broad at its base; lateral margin armed with 4 spines; subapical one long, half the length of apical spines and located away from proximal 3 arranged at regular intervals; apical margin narrow, with pair of strong spines at corners, median plate, 4 spinules, narrow, and pair of plumose setae present VOLUME 110, NUMBER 4 603 Fig. 1. Lophogaster muranoi, new species. A, C—G: holotype (male); B: allotype (female). A, Anterior end, dorsal view; B, anterior end, dorsal view; C, antennular lobe, ventral view; D, antenna; E, mandible; E maxillule; G, maxilla. 604 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON xy J WY Q SS RQ ‘Va wenn ACE G OO inm DE Fig. 2. Lophogaster muranoi, new species. Holotype (male). A, First thoracic limb; B, second thoracic limb; C, third thoracic limb; D, dactylus of third thoracic limb; E, fourth thoracic limb; EK merus of fourth thoracic limb; G, fifth thoracic limb. 605 VOLUME 110, NUMBER 4 Fig. 3. Lophogaster muranoi, new species. Holotype (male). A—C, Sixth to eigth thoracic limbs; D-H, first to fifth pleopods. 606 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON H 1mm : | oun eee ae 1mm C 1mm t————1DG Fig. 4. Lophogaster muranoi, new species. A-C: holotype (male); D—F: young specimen (7.8 mm); G-I: young specimen (8.0 mm). A, Telson and uropod; B, apical part of telson; C, right postero-lateral part of carapace; D, anterior end, dorsal view; E, postero-lateral part of carapace; FE telson; G, anterior end, dorsal view; H, right postero-lateral part of carapace; I, telson. VOLUME 110, NUMBER 4 between outer 2 indentations of median plate; 2 spinose dorsal keels present on pos- terior 0.8 of telson (Fig. 4A, B). Morphological change with growth.—In many species of the genus Lophogaster, geographical variation, sexual dimorphism and morphological change with growth have been observed in several characters (Hansen 1910, Fage 1942, O. Tattersall 1960, Casanova 1993, 1996). In this new species morphological change with growth was observed in the lengths of the median rostral spine and the postero-lateral spine of the carapace, and in the number of lateral spines of the telson (Fig. 4D-I). The me- dian rostral spine of young specimens is much shorter as compared to adults and ex- tends only to the middle of the third seg- ment of the antennular peduncle. The pos- tero-lateral spine of the carapace is also shorter in young specimens than in adults. Three lateral telson spines are present in the smallest specimen (7.8 mm body length) compared to 4 in larger specimens. The mi- croscopic tubercles on the carapace become noticeable in older specimens. The anten- nular lobe, the postorbital spines on the car- apace, and the apical plate of the telson do not change with growth or gender. Etymology.—Named in honor of Dr. Ma- saaki Murano. Remarks.—Lophogaster muranoi, new species, is related to Lophogaster typicus in respect to the rounded antennular lobe, the presence of a pair of the postorbital spines, and the armature of the telson, but differs from the latter species in the following ways. The median rostral spine extends beyond the apex of the antennular lobe in L. muranoi, while it ends in the middle of the distal seg- ment of the antennular peduncle in L. typicus, the spine at postero-lateral angle of the car- apace is much longer in L. muranoi than in L. typicus, and in L. muranoi the apical mar- gin of the telson between the pair of the long apical spines forms a narrow projecting plate while the apical margin does not protrude to a plate in L. typicus. Tattersall & Tattersall (1951) observed 607 that for L. typicus the coarse serrations on the antero-lateral margin of the rostrum, which is seen in younger specimens, is gradually lost as growth proceeds and com- pletely disappears at a length of 13 mm. In L. muranoi the minute serrations on the lat- eral margin of the rostrum are retained con- tinuously from the younger stage (7.8 mm) to the adult (19.2 mm). Lophogaster muranoi is easily distin- guishable from the two known species from the South Atlantic Ocean, L. challengeri and L. spinosus, by the presence of the postorbital spines on the carapace. Acknowledgments We are grateful to Dr. Masaaki Murano, Institute of Environmental Ecology, METOCEAN Co. Ltd., for his critical re- viewing the manuscript. Literature Cited Casanova, J.-P. 1993. Crustacea Mysidacea: Les Mys- idacés Lophogastrida et Mysida (Pethalo- phthamidae) de la région néo-calédonienne. in A. Crosnier, ed., Résultats des Campagnes MU- SORSTOM, Volume 10.—Mémoires du Muse- um national d’Histoire naturelle 156:33-—53. . 1996. Crustacea Mysidacea: Les Lophogas- tridés d’Indonésie, de Nouvelle-Calédonie et des Iles Wallis et Futuna. in A. Crosnier, ed., Résultats des Campagnes MUSORSTOM, Vol- ume 15.—Mémoires du Museum national d’ Histoire naturelle 168:125—146. Fage, L. 1942. Mysidacea Lophogastrida II. The Carls- berg Foundation’s oceanographical expedition round the world 1928-1930 and previous ‘“‘Dana’’ expeditions under the leadership of Prof. Johannes Schmidt.—Dana Report 23:1—67. Hansen, H. J. 1910. The Schizopoda of the Siboga expedition.—Siboga-Expeditie 37:1—123. Sars, G. O. 1885. Crustacea I. in Den Norske Nor- dhavs-Expedition, 1876-78, Christiana, 14, Zool.: 1—280. (not seen) Tattersall, O. S. 1955. Mysidacea.—Discovery Report 28:1-190. 1960. Notes on mysidacean crustaceans of the genus Lophogaster in the U. S. National Museum.—Proceedings of the United States National Museum 112:527—547. Tattersall, W. M., & O. S. Tattersall. 1951. The British Mysidacea.—The Ray Society, London, Vol- ume 136:1—460. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):608—616. 1997. A new species of troglobitic crayfish of the genus Cambarus, subgenus Aviticambarus (Decapoda: Cambaridae), endemic to White Spring Cave, Alabama John E. Cooper and Martha Riser Cooper (JEC) North Carolina State Museum of Natural Sciences, P. O. Box 29555, Raleigh, North Carolina 27626, U.S.A.; (MRC) 209 Lynwood Lane, Raleigh, North Carolina 27609, U.S.A. Abstract.—Cambarus (Aviticambarus) veitchorum is a rare, diminutive spe- cies of troglobitic crayfish endemic to White Spring Cave in the Tennessee River Valley, Limestone County, Alabama. It is most closely related to C. (A.) hamulatus, which occurs in subterranean waters of the Sequatchie Uplift in Alabama and Tennessee, and is less closely related to C. (A.) jonesi, with which it is syntopic and whose range encompasses both sides of the Tennessee River in Alabama. The new species differs from the other Aviticambarus in the re- lationships of the terminal elements of the gonopod (male first pleopod); the shape of the abdominal pleura and presence of spines at their caudoventral angles; the lack of spines or prominent tubercles on the mesial surface of the carpus; the attenuated podomeres of the pereiopods, especially of the cheliped; the morphology of the epistome; and the structure of the annulus ventralis of the female. Six species of troglobitic crayfishes, four described and two undescribed, are known to live in subterranean waters of northern Alabama. The described species are Cam- barus (Aviticambarus) hamulatus (Cope 1881), which occurs in caves of the Se- quatchie Uplift in Blount, Jackson, and Marshall counties; Cambarus (Aviticamba- rus) jonesi Hobbs & Barr, 1960, whose range encompasses both sides of the Ten- nessee River Valley, from Madison County west into Colbert and Lauderdale counties; Orconectes (Orconectes) australis australis (Rhoades 1941), known only from Jackson and Madison counties; and Procambarus (Remoticambarus) pecki Hobbs, 1967, re- ported from three caves along the Tennes- see River, one each in Colbert, Lauderdale, and Morgan counties (see Hobbs et al. 1977). Cambarus jonesi, P. pecki, and the two unnamed species are endemic to Ala- bama, but the ranges of C. hamulatus and O. a. australis extend north into Tennessee. Each of the two undescribed forms is known from but a single cave, one in Mad- ison County and the other in Limestone County. The description of the former spe- cies has been submitted for publication, and the latter is described herein. Only seven specimens of this rare crayfish, six adults and a juvenile, have been collected. Cambarus (Aviticambarus) veitchorum, new species Fig. 1 ‘Three new species.’’—Cooper & Cooper, 1966:39 (p. p.); Cooper, 1967:14 (p. p.). Cambarus (Aviticambarus) sp. A.—Bou- chard, 1976:14, 15. ‘“‘Two crayfishes of subgenus Aviticamba- rus.”’—Hobbs et al., 1977:5 (p. p.). ‘““Two undescribed troglobites.”’—Hobbs et al., 1977:75 (p. p.). C. (A.) sp. A.—Fitzpatrick, 1990:78. Diagnosis.—Albinistic; eyes degenerate, VOLUME 110, NUMBER 4 recessed, lacking pigment. Rostrum acari- nate, excavate, margins narrow, slightly el- evated, and parallel or subparallel to base of long acumen, which delimited by strong marginal spines; margins of acumen con- stricted at base, then broadly concave and strongly converging to acute, cephalodor- sally directed apex; acumen comprising 35.0 to 43.2% (X = 39.4) of rostrum length, latter constituting 23.3 to 25.9% (X = 24.6) of total carapace length (TCL). Areola 5.1 to 7.4 (X = 6.1) times as long as broad, constituting 40.0 to 43.1% (X = 41.3) of TCL and 52.7 to 55.8% (X = 54.6) of post- orbital carapace length (PCL), with 4 to 5 punctations across narrowest part. Cepha- lothorax subovate; carapace 1.4 to 2.3 (X = 1.8) times wider than deep, dorsally punc- tate, laterally granulate. Cervical spines strong, usually 1 or 2 on each side. Bran- chiostegal spine strong; hepatic region with scattered weak tubercles. Suborbital angle obsolete, margin of orbit diagonal. Postor- bital ridge moderately strong, groove ob- solete and represented by row of minute punctations, cephalic margin with strong spine. Antennal scale 2.5 to 2.8 (X = 2.7) times as long as wide, greatest width just distal to midlength; lateral margin thick- ened and terminating distally in strong spine. Chela of cheliped attenuate, width of palm 15.1 to 21.9% (X = 19.1) of total length of chela, latter 58.3 to 76.0% (X = 64.6) of TCL and 64.4 to 98.4% (X = 82.4) of PCL; palm subovate in cross section, 1.3 to 1.9 (X = 1.4) times as wide as deep, 2.3 to 3.1 (X = 2.6) times as long as wide; me- sial margin of palm with 2 or 3 poorly de- fined rows of minute tubercles, 13 to 18 (usually 15-16) in mesial row, length of margin 45.4 to 51.0% (X = 48.4) of total chela length. Fingers without gape, without dense setae at opposable bases; both fingers with longitudinal ridge dorsally and ven- trally; opposable margin of fixed finger usu- ally bearing 3 to 4 tubercles in addition to usual subconical one; mesial margin of dac- tyl punctate, with 1 to 3 small tubercles 609 near base; opposable margin of dactyl usu- ally with 3 to 4 tubercles, one of which larger than others, subconical, and situated ventral to denticles; dactyl length 1.0 to 1.2 (X = 1.1) times length of mesial margin of palm, 2.7 to 3.4 (X = 2.8) times width of palm, and 50.1 to 59.4% (X = 53.8) of total chela length. Carpus of cheliped subrectan- gular in dorsal outline, 1.5 to 2.5 (X = 2.0) times as long as wide, mesial surface with- out spines or prominent tubercles. Merus of cheliped attenuate, 4.0 to 5.7 (X = 4.6) times longer than deep, length 41.4 to 52.1% (X = 45.8) of TCL and 54.5 to 67.4% (X = 60.5) of PCL. Pleura of second through fifth abdominal segments with an- gular caudoventral corners, and spine at caudoventral apex of angle. Mesial ramus of uropod with foreshortened median keel, which lacking distal spine; cephalic section of telson with 2 cephalolateral spines each side. Hook on ischium of third pereiopod, that of form I male slightly overreaching bas- ioischial articulation, not opposed by tuber- cle on basis; coxa of fourth pereiopod of males with prominent caudomesial boss, which somewhat laterally compressed. Gonopods (first pleopods) of form I male symmetrical in caudal aspect, total length 21.0% of TCL, proximomesial apophyses widely separated; proximolateral lobe set off from rest of shaft by transverse groove; left gonopod in lateral aspect with moderate protuberance on cephalic border; central projection corneous, curved at greater than 90° angle to shaft, strongly tapering to sub- acute, proximomesially directed tip; sub- apical notch present, directed proximally; caudal portion of central projection with proximal margin twisted mesially; mesial process curved, somewhat expanded at base but tapering to subacute tip, which directed caudolaterally and extending caudally to level of tip of central projection or beyond; caudal knob absent; in mesial aspect, ce- phalic protuberance obvious, mesial surface lacking setae. Annulus ventralis symmetrical, subovate, 610 1.8 times wider than long; caudal margin free, broadly convex, elevated, and dissect- ed by sinus just dextral to midline, at which point caudosinistral terminus of wall plung- ing into expanded caudodextral terminus; lateral apices of caudal wall moderately ex- panded; cephalic margin of annulus arched, clearly delimited but fused to preannular sternite; cephalolateral margins relatively thick, slightly concave on each side of me- dian arch; cephalic half of annulus with broad central depression, which extending into caudal half as oblique depression on each side of median C-shaped ridge sur- rounding caudosinistral terminus of caudal wall. Postannular sclerite about 3 times as wide as long, with sublinear caudal margin, broadly convex cephalic margin, and domed ventral surface bearing some small tubercles. Female first pleopod absent, slight protuberance at site on each side. Measurements (mm) of type specimens provided in Table 1. Holotypic male, form I.—Cephalothorax (Fig. 1A, D) subovate; carapace 1.7 times wider than deep. Areola 5.1 times as long as broad, constituting 43.1% of TCL (55.8% of PCL), with dense, small punc- tations, 5 or 6 across narrowest part; bran- chiocardiac grooves subparallel throughout most of length, caudally flared, with trans- verse eminence between caudal termini. Rostrum with narrow, somewhat elevated margins, parallel nearly to level of strong marginal spines, where moderately con- verging to bases of spines; margins of acu- men slightly concave, converging strongly to corneous, cephalodorsally directed apical spine reaching distal margin of second ar- ticle of antennular flagellum; acumen com- prising 35.0% of rostrum length, latter con- Stituting 24.0% of TCL; floor (dorsal sur- face) of rostrum with deep longitudinal ex- cavation, apunctate except for small punctations proximal to marginal spines, and row of punctations bearing long, re- cumbent setae just mesial to each marginal ridge; subrostral ridge narrowly visible to base of marginal spines in dorsal aspect. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Measurements (mm) of types of Cam- barus (Aviticambarus) veitchorum, new species. Holo- © Allo- Morpho- type type type Carapace Total length 16.7 14.5 15.9 Postorbital length 12:9 11:0 See Length cephalic section 9:5 S8le oe Width 6.1 [3457 62 Depth 3.5... ppatohin eels Length rostrum 4:0" - 3.6 ) 3a Length acumen 14 14 1.6 Areola Length Tad, ideo =e Width 4 TO -09 Antennal scale Length 3) 3 ee Width 1:3 Ee ies Abdomen Length 18.2 “153 AT3 Width 513 4a Cheliped (right) Length lateral margin chela 12.7 °8.6 10.4 Length mesial margin palm Gf 42 233 Width palm 23 “ae eee Depth palm 1:8. 0 1.6 Length dactyl 64 47 £5.46 Length carpus 3.4. 2a ee Width carpus 2M. V4” 16 Length dorsal margin merus 8.7 GIO) rae Depth merus i9S°*) Sears Gonopod length 34 NA 3.4 Postorbital ridge fairly strong, groove ob- solete and represented by row of small punctations; dorsal margin ridgelike, ce- phalic margin with strong spine. Branchio- stegal spine strong; suborbital angle obso- lete, orbital rim with slight concavity around base of antennal peduncle, dorsal to which rim oblique. Carapace dorsally punc- tate, laterally and dorsolaterally granulate; cephalic section 1.3 times longer than are- ola and constituting 56.9% of TCL; gastric region mostly glabrous, with some scattered punctations; carapace cephalolateral to cer- vical groove with small scattered tubercles, -and row of same along ventral margin of cephalic portion of cervical groove. Cervi- cal spines strong, 1 on each side. Abdomen VOLUME 110, NUMBER 4 611 Fig. 1. Cambarus (Aviticambarus) veitchorum, new species (all from holotypic male, form I, except C, E, from morphotypic male, form II, and G, from allotypic female; setae not illustrated): A, lateral aspect of ceph- _alothorax; B, C, mesial aspect of left gonopod (first pleopod); D, dorsal aspect of cephalothorax; E, F lateral aspect of left gonopod; G, annulus ventralis and associated structures; H, caudal aspect of in situ gonopods; I, lateral aspect of abdomen; J, dorsal aspect of antennal scale; K, epistome; L, basal podomeres of third, fourth, and fifth pereiopods; M, dorsal aspect of distal podomeres of right cheliped. 612 narrower and slightly longer than cephalo- thorax. Pleura of third through fifth abdom- inal segments (Fig. 11) with curved cephal- oventral margins, linear caudal margins, and acute caudoventral corners; second through fifth pleura with spine at apex of caudoventral angle. Proximal podomere of uropod with strong distomedian spine on both lobes, that on mesial lobe longer; me- sial ramus of uropod with foreshortened median keel, which lacking distal spine, and long distal spine on lateral margin; lat- eral ramus with submedian ridge of ce- phalic section terminating in spine at trans- verse flexure, latter bearing row of 12 fixed spines (14 on left) across margin, and 1 large movable spine at lateral corner. Telson with 1 long stationary and 1 smaller mov- able spine at each caudolateral corner of ce-_ phalic section; lateral margins of caudal section somewhat converging caudally, caudal margin rounded. Epistome (Fig. 1K) with cephalic lobe subrectangular in outline, wider than long, margins slightly thickened; cephalic border entire, rounded, without projection; lateral margins subparallel for about % of length of lobe, lateral apices not thickened; lobe without constriction at base; floor (ventral surface) of lobe with 8 minute tubercles, and central concavity extending into central depression of body; depression narrow, with shallow median fovea; lamellae with cephalolateral margins broadly concave, lateral apices narrow, subangular; zygoma weakly arched, pits short, deep. Antennal peduncle with strong, procurved cephalo- lateral spine on basis, and strong, erect ven- tral spine on ischium; antennular peduncle with strong ventral spine at base of distal % of proximal podomere. Antennal scale (Fig. 1J) 2.8 times as long as broad, widest just distal to midlength; lateral margin thick- ened, terminating in long distal spine (tip missing from spine on left side), which di- rected distolaterally, tip reaching base of distal %4 of ultimate podomere of antennular peduncle; lamella about 2.1 times as wide as thickened lateral margin; distal margin of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lamella sloping proximomesially to mesial margin, latter gently curving proximally to widest point, then curving proximolaterally to base. Third maxilliped with distolateral margin of ischium slightly produced, not spinelike; ventrolateral margin of ischium with row of punctations bearing short setae at base of longitudinal ridge; ventral surface of lateral half with scattered punctations bearing short setae, some punctations containing small squamous tubercles; ventral surface of mesial half with long, dense setae ob- scuring most of proximal area; mesial mar- gin with about 18 denticles; tip of exopodite reaching base of distal %4 of merus of en- dopodite. Right mandible with incisor ridge bearing 7 denticles; molar process in two parts, caudalmost with 2 round, adjacent tu- bercles, cephalicmost with single tuberculi- form mound. Chela of cheliped (Fig. 1M) with palm subovate in cross section, 1.3 times wider than deep; mesial margin of palm 2.7 times longer than palm width, and length com- prising about 50% of chela length; latter 76.0% of TCL (98.4% of PCL); dorsal sur- face of palm with small punctations bearing short setae, and several longer recumbent setae present; dorsal articular ridge fairly well defined, with subdistal clump of setae near midwidth; ventral surface of palm with small punctations bearing short setae, most punctations containing small squamous tu- bercles; ventral articular ridge well defined, with broad subdistal tubercle bearing sev- eral long setae at midwidth; lateral margin of palm rounded, punctate; mesial margin with 2 to 3 staggered rows of small tuber- cles, 18 to 19 in mesialmost row. Fingers without gape, without plumose setae at op- posable bases; both fingers in dorsal aspect fairly straight, in lateral aspect gently curv- ing distoventrally. Dactyl with longitudinal ridge dorsally and ventrally, with several small punctations on dorsal ridge near base, and 1 punctation bearing long setae at mid- length; surfaces flanking ridges punctate, without grooves; mesial margin punctate, VOLUME 110, NUMBER 4 but with several minute tubercles near base; dactyl comprising 50.1% of length of chela. Fixed finger with surfaces flanking dorsal and ventral ridges punctate; lateral surface of fixed finger punctate, some median punc- tations large and bearing prominent setae. Opposable margin of dactyl with single row of prominent denticles on entire length of finger, 5 small tubercles on proximal % dor- sal to row of denticles, and 1 larger sub- triangular tubercle ventral to denticles at base of distal 7, of finger; opposable margin of fixed finger with single row of prominent denticles on entire length, 3 small tubercles on proximal Y; of finger dorsal to denticles (second from base largest), and larger sub- conical tubercle ventral to denticles at base of distal *% of finger. Carpus of cheliped (Fig. 1M) subrectan- gular in dorsal outline, 1.9 times longer than wide; dorsal surface with shallow, nearly obsolete sulcus, mesial and lateral to which suface punctate and with short setae; ventral surface with strong distomedian spine, proximomesial to which is row of small tubercles; mesial margin without spines, with small scattered tubercles. Me- rus of cheliped without pronounced disto- dorsal spines or tubercles; dorsal ridge with 2 staggered rows of small tubercles, ex- panding distally into group of tubercles on dorsal, lateral, and mesial surfaces; rest of lateral and mesial surfaces with scattered small tubercles; ventrolateral ridge of merus with 16 or 17 scarcely discernible tubercles, and long distolateral spine near lateral ar- ticular prominence; ventromesial ridge with 16 or 17 small tubercles, slightly larger than those on ventrolateral ridge, and 1 long dis- tal spine; area between ventral ridges punc- tate, with short setae; area between distal ends of ridges with some small tubercles. Ischium with row of 5 or 6 minuscule tu- bercles along ventral ridge, with several deep punctations and 2 minuscule tubercles lateral to them; dorsal surface with minus- cule tubercles. Hook on ischium of third pereiopod (Fig. 1L) simple, distally flattened and moderate- 613 ly acute; tip overreaching basioischial artic- ulation, not opposed by tubercle on basis. Coxa of fourth pereiopod with prominent caudomesial boss. See the “Diagnosis”? for description of gonopod (Fig. 1B, E H). Allotypic female.—Except for secondary sexual characters, differing from holotype in following respects: Carapace 1.8 times wider than deep; areola 5.9 times longer than wide, constituting 40.7% of TCL (53.6% of PCL), with 3 to 4 punctations across natrowest part. Cephalic section of cephalothorax 1.5 times longer than areola and constituting 59.3% of TCL. Acumen comprising 38.9% of rostrum length, latter constituting 24.8% of TCL. Cervical spines 2 on left, 1 on right. Antennal scale 2.7 times longer than wide, tip of spine reach- ing distal margin of second article of anten- nular flagellum. Palm of chela of cheliped 1.5 times wider than deep; mesial margin of palm 2.5 times longer than palm width, and length comprising 48.8% of chela length; latter 59.3% of TCL (78.2% of PCL); mesial margin of palm with 14 to 15 minute tubercles in mesialmost row. Dactyl comprising 54.7% of chela length; mesial surface punctate, without tubercles; oppos- able margin with 2 small tubercles near base, and 1 small subtriangular tubercle ventral to denticles at proximal % of finger. Opposable margin of fixed finger with 2 small tubercles near base, and subconical tubercle at base of distal % of finger. Carpus of cheliped 2.1 times longer than wide; ven- tromesial ridge of merus with 12 or 13 small tubercles, plus large distal spine. Coxa of fourth pereiopod without boss. See the “‘Diagnosis”’ for description of annulus (Fig. 1G). Morphotypic male, form II.—Differing from holotype in following respects: Areola 7.4 times longer than wide, constituting 42.1% of TCL (55.5% of PCL), with 4 punctations across narrowest part. Cephalic section of carapace 1.4 times longer than areola and constituting 57.9% of TCL. Acu- men comprising 43.2% of TCL. Rostrum 614 with right margin and marginal spine ex- tending farther cephalically than left margin and spine. Single small cervical spine on left side of carapace, none on right. Anten- nal scale 2.7 times longer than wide, tip of spine reaching distal margin of fourth arti- cle of antennular flagellum. Palm of chela of cheliped with mesial margin 2.5 times longer than wide, chela length 65.4% of TCL (86.0% of PCL); mesial margin of palm with 14 or 15 minuscule tubercles in mesialmost row. Dactyl comprising 53.8% of chela length; opposable margin of dactyl with 3 small tubercles near base (middle one slightly larger), 1 large subtriangular tubercle ventral to denticles at base of distal ¥; of finger, followed distally by 2 minus- cule tubercles. Opposable margin of fixed finger with subconical tubercle situated just proximal to midlength. Carpus of cheliped 2.0 times longer than wide; ventrolateral ridge of merus with 14 or 15 scarcely dis- cernible tubercles and small distolateral spine. Hook on ischium of third pereiopod moderately well developed, not overreach- ing basioischial articulation. Gonopods (Fig. 1C, E) reaching just be- yond midlength of coxa of third pereiopod when abdomen flexed; left gonopod in lat- eral aspect without juvenile suture; promi- nence on cephalic margin reduced; central projection noncorneous, fairly broad but ta- pering to rounded tip, which directed slight- ly proximocaudally; proximal margin of central projection slightly overlapping dis- tal margin of mesial process; mesial process long, narrow, with constriction near tip; lat- ter curved at slightly greater than 90° angle, tip extending well beyond apex of central projection. Disposition of types.—The holotype, al- lotype, and morphotype are in the crusta- cean collections of the North Carolina State Museum of Natural Sciences (NCSM), Ra- leigh (catalogue numbers NCSM C-2544, C-2545, and C-2546, respectively), as are the following paratopotypes: 1 6 I, 1 2 (C- 2547), 1 6 I, 1j 6 (C-2548). Type locality.—Alabama, Limestone PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON County, subterranean stream in White Spring Cave, NW of Holland Gin (Tanner 7.5' USGS Quadrangle, Sec. 11, TSS, R.4W). The cave is designated AL 242 in the cave cataloguing system of the Alabama Cave Survey, an official project of the Na- tional Speleological Society. White Spring Cave is located in the Ten- nessee Valley north of the Tennessee River, within the southern boundaries of the High- land Rim of the Interior Low Plateaus. The cave is developed in rock of Mississippian age, probably the Fort Payne Chert. The ex- tent of the cave is unknown because the ac- cessible passage is only traversible for about 12 to 18 m beyond either of the two entrances. The northern entrance is at the western edge of a shallow surface depres- ‘sion, in which water from the cave stands during periods of elevated water level. We have observed troglobitic crayfish in the de- pression, where they were exposed to day- light. The stream within the cave flows gen- erally southeast and south, and the water was moving slowly during both of our vis- its. Range and specimens examined.— Known only from the type locality, where the following specimens have been collect- ed: 2 6 I, 2 2, 24 Sep 1967, coll. J. Veitch, MRC, JEC; 1 6 I, 1 6 Il, 1j d, 4 Jul 1968, coll. A. Dobson, R. C. Graham, JEC. Variations.—The range of variation in most characters is provided in the “Diag- nosis”’ and in the discussions of the allotype and morphotype. The following additional variations have been noted. The number of tubercles on the opposable margin of the fixed finger of the chela (exclusive of the larger subconical tubercle) ranges from one in the two smallest animals, to two or three in the others; the number of tubercles on the opposable margin of the dactyl is usu- ally greater and shows more variation, with two in the juvenile male and from three to six in the others. The number of cervical spines varies from one animal to another, and from one side of the body to the other. The juvenile male and the holotype have a VOLUME 110, NUMBER 4 single spine on each side, the other form I males have two spines on each side, the morphotype and the smallest female have one spine on the left side and none on the right, and the allotype has two spines on the left and one on the right. One of the form I males has the usual two spines in the cau- dolateral corner of the cephalic section of the left side of the telson, but a single spine on the right side. The margins of the ros- trum are more strongly convergent in the allotype and the juvenile male, and in both these specimens the merus is slightly longer than it is in the others. Sexual dimorphism is evident in the length of the chela, with form I males having chelae that exceed 75% of TCL, while the chelae of other males and females constitute 60 to 65% of TCL. Size.—Adult TCL ranges from 13.5 mm (a female) to 16.7 mm (a form I male). The mean TCL of the four males is 15.9 mm, and of all six adults is 15.2 mm. Life history notes.—Form I males were found in July and September. The largest of the two females collected in September ex- hibited oocytes (visible through the cara- pace) in a late stage of development, as well as highly developed cement glands. The small female collected at the same time contained oocytes in an early stage of de- velopment. As is the case with nearly all troglobitic crayfishes, no females carrying attached ova or young have been found. Relationships.—Cambarus veitchorum is closely related to C. hamulatus and C. jo- nesi, and is syntopic with the latter. Al- though the gonopods of the form I and form II males of C. veitchorum are more like those of C. jonesi than those of C. hamu- latus, as are the relative proportions of the areola, the overall facies of the new species argues for a somewhat closer relationship to C. hamulatus. The rostrum of both is very similar (and unlike that of C. jonesi), as are the shape of the antennal scale, the general shape and proportions of the chela and the cephalothorax, and the overall 615 structure of the annulus. In all three species the female lacks first pleopods. Cambarus veitchorum is markedly dif- ferent from both its closest relatives in the following characters: the configuration and spatial relationships of the terminal ele- ments of the form I male gonopod, with the central projection directed caudomesially and the proximal margin of its caudal por- tion twisted mesially; the shape of the ven- tral margins of the third through fifth ab- dominal pleura, and the presence of a spine on the caudoventral angle of the second through fifth pleura; the absence of spines or prominent tubercles on the mesial sur- face of the carpus; the attenuated chela, car- pus, and merus; the configuration of the ce- phaiic lobe of the epistome; the reduced number of tubercles on the mesial surface of the dactyl; the greater number and very weak nature of the tubercles on the mesial margin of the palm and on both ventral ridges of the merus; the obsolete groove of the postorbital ridge; the very broad de- pression in the cephalic portion of the an- nulus; the lower number of tubercles on the opposable margins of both fingers; the com- parative dearth of long, erect setae on the palm and fingers of the cheliped (also gen- erally absent in C. hamulatus); and the di- minutive adult size, with the maximum TCL of 16.7 mm, as opposed to 28.9 mm for C. jonesi (3 I, NCSM C-198), and 35.2 mm for C. hamulatus (Hobbs et al. 1977: 76). Crayfish associates.—The only other crayfish known to inhabit White Spring Cave is C. jonesi, which is far more abun- dant there than C. veitchorum. Both species were observed in very close proximity to each other, but no interactions between them were seen. Etymology.—We take great pleasure in naming this species for John D. Veitch and Joyce Veitch of Decatur, Alabama, friends who introduced us to White Spring Cave and its owners, and who served as im- mensely cheerful hosts to a couple of mud- 616 dy cave biologists on more occasions than they would probably care to remember. Suggested vernacular name: White Spring Cave Crayfish. Acknowledgments We are very grateful to John and Joyce Veitch for their hospitality and many kind- nesses; to Mr. and Mrs. Rowe Sanderson, who gave us permission to collect in White Spring Cave; and to Arthur Dobson and Richard C. Graham, who accompanied JEC on the trip that yielded the morphotype. JEC expresses his sincerest gratitude to his co-author, to R. E. Ashton, Jr., J. E. Cooper, Jr., J. Perry, and especially to D. Howard and A. L. Braswell, without whose gener- osity this paper could never have been com- pleted. Joseph F Fitzpatrick, Jr., Steve Bus- ack, and an anonymous reviewer, provided cogent comments on the manuscript. As al- ways, we are immensely grateful to the late Horton H. Hobbs, Jr., for the splendid les- sons he taught (not all of them about dec- apods). Literature Cited Bouchard, R. W. 1976. Crayfishes and shrimps. Pp. 13—20 in H. Boschung, ed., Endangered and threatened plants and animals of Alabama. Bulletin of the Alabama Museum of Natural History No. 2, 93 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cooper, J. E. 1967. Animals in Alabama caves. Pp. 14-16 in J. Veitch. The caves of Alabama. Huntsville Grotto, National Speleological So- ciety, Huntsville, Alabama, 51 pp. , & M. R. Cooper. 1966. Comments on Ala- bama cave biology. Huntsville Grotto Newslet- ter 7(5—6):37—41. Reprinted 1969, pp. 25-30 in W. T. Plummer, ed., Speleo Digest, Section 2. National Speleological Society, Huntsville, Al- abama. Cope, E. D. 1881. Orconectes hamulatus. in E. D. Cope & A. S. Packard, Jr. The fauna of the Nickajack Cave.—American Naturalist 15:881— 882. Fitzpatrick, J. E, Jr. 1990. Decapoda. Pp. 77-80 in S. C. Harris, Preliminary considerations on rare and endangered invertebrates in Alabama.— Journal of the Alabama Academy of Sciences 61(2):64—92. Hobbs, H. H., Jr. 1967. A new crayfish from Alabama caves with notes on the origin of the genera Orconectes and Cambarus (Decapoda: Astaci- dae).—Proceedings of the United States Nation- al Museum 123(3621):1—17. , & T. C. Barr, Jr. 1960. The origins and affin- ities of the troglobitic crayfishes of North Amer- ica (Decapoda, Astacidae), I: the genus Cam- barus.—American Midland Naturalist 64(1): 12-33. , H. H. Hobbs, III, & M. R. Daniel. 1977. A review of the troglobitic decapod crustaceans of the Americas.—Smithsonian Contributions to Zoology 244:1—183. Rhoades, R. 1941. Notes on some crayfishes from Al- abama caves, with the description of a new spe- cies and a new subspecies.—Proceedings of the United States National Museum 91(3129):141— 148. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):617—623. 1997. Two new species and a range extension of mud shrimps, Upogebia, from Pacific Costa Rica and Mexico (Decapoda: Thalassinidea: Upogebiidae) Austin B. Williams National Marine Fisheries Service Systematics Laboratory, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Two new species of mud shrimps are described from the Pacific side of Middle America, Upogebia baldwini from the state of Nayarit, Mexico, and U. vargasae from Puntarenas Province, Costa Rica. The geographic range of the poorly known U. burkenroadi Williams, 1986 is extended from its type locality in the state of Sonora, Mexico to a documented Costa Rican occur- rence. The new species are illustrated, and a key for identification of eastern Pacific species of Upogebia (Williams 1986) is amended to accommodate the new species. Cataloged lots of each species collected are listed. Three species of mud shrimps, genus Upogebia, from the Pacific side of Middle America that have become known since the publication of Williams (1986) require more than routine identification. The first is a unique specimen of a new species collected in mangrove habitat from the state of Nay- arit, Mexico by Aaron Baldwin in 1988. Ad- ditional material has not become available, so it seems inappropriate to delay descrip- tion. A second new species, collected by Rita Vargas in 1995 from a mangrove hab- itat in Costa Rica, is described. Finally, spec- imens collected by J. Cortés from Costa Rica in 1991 are provisionally referred to Upo- gebia burkenroadi Williams, 1986 described from Sonora, Mexico, thus extending the geographic range of that species. Specimens are deposited in the U.S. Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. (USNM), or returned to the University of Costa Rica Museum of Zoology (UCRMZ), San José, as indicated in the species ac- counts. Upogebia baldwini, new species Fig. 1 Material.—USNM 251486. Holotype 2°, Estero Playa Novillero, Nayarit, Mexico, dug from mangrove estero, coll. A. Bald- win, 5 Feb 1988. Diagnosis.—Projections to either side of rostrum ending in acute spine. Postocular spine present. Abdominal sternites un- armed. Telson subrectangular, with trans- verse proximal ridge. Carpus of cheliped with 2 spines on mesiodistal margin, palm with row of spines on mesioventral surface. Merus of pereopod 2 with proximal me- sioventral spine and subdistal dorsal spine; merus of pereopod 4 spineless. Description.—Rostrum triangular, later- al margin shorter than basal width; lower margin nearly horizontal in lateral view but dorsal margin downturned, tip exceed- ing eyestalks; dorsal pair of strong subapi- cal spines followed on each side by 4 erect marginal spines separated by subequal in- tervals; 3 spines mesial to marginal row at base on right side, none on left; central dorsal area hidden under setae. Rostral spines hidden by dense tufts of setae sprouting anterior to their bases, confluent with field of similar postrostral spines be- coming smaller and less hidden posterior- ly. Ridge lateral to field bearing crest of about 15-16 moderate to small spines, 618 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON U FAK: de =~ : Fig.) 1. dorsal; c, telson, left uropod, part of abdominal segment 6, dorsal; d, cheliped, right lateral; e, crest of right cheliped carpus showing spines; f, chela and carpus, right mesial; g—j, right pereopod 2, left 3—4, right, 5. strongest on process lateral to rostrum but decreasing posteriorly, with tendency to cluster at end of row. Shoulder lateral to cervical groove armed with tubercle on left side above intersection with thalassinidean line and with 4—9 obsolescent spines or tu- bercles to either side below intersection, line continuing strongly to posterior mar- gin. Postocular spine present. Abdomen with sternites on segments 1 and 2 wrinkled but not spined. Upogebia baldwini, USNM 251486, 2 Holotype; a, cephalic region, lateral; b, anterior carapace, Telson with transverse proximal ridge confluent with low longitudinal ridge to ei- ther side. Eyestalk stout, deepest at about mid- length, convex ventrally, horizontal in re- pose; cornea prominent, obliquely terminal, narrower than diameter of stalk, row of tiny forward trending spines above margin of cornea on mesial aspect. Antennular peduncle reaching slightly beyond midlength of terminal article of an- VOLUME 110, NUMBER 4 tennal peduncle, proximal 2 articles togeth- er slightly longer than terminal article. Antennal peduncle with almost % of length extending beyond tip of rostrum; scale moderate, oval. Maxilliped 3 bearing epipod. Epistomial projection rather broad in lat- eral view, bearing 2 small apical projec- tions. Chelipeds with ventral margin of ischium bearing 1 spine. Merus with row of 5 spines on ventral margin, proximal 2—3 modest in size, distal spines smaller; single subdistal dorsal spine reaching level of postocular spine. Carpus trigonal, shallow longitudinal groove laterally, anterior ventrolateral cor- ner bearing modest spine; mesiodorsal crest of 4 moderate spines partly obscured by se- tae behind prominent spine on anterior mar- gin; 2 spines on anterodorsal margin mesial to articulation with propodus obscured by dense setae; strong spine near middle of an- teromesial margin, smaller spine dorsal to it, and strong slender spine behind distov- entral corner. Chela length about 5 times chela height; dorsal ridge spineless, stout spine mesial to it subdistally; mesiodorsal row of 13-14 erect to forward-hooked spines not reaching anterior part of palmar length; moderate spine below mesial dac- tylar condyle, single spine on margin below lateral condyle; mesial surface bearing se- tose longitudinal row of obsolescent spines on upper half, lower half bearing horizontal row of about 7 strong spines; transverse low ridge near proximomesial margin intersect- ing ventral ciliated keel. Fixed finger short- er than dactyl, continuing slightly sinuous trend of lower palmar margin and tapering to slender tip, 3 irregular teeth on proximal prehensile edge, strong double tooth in middle of row. Dactyl with corneous tip preceded on prehensile edge by crowded row of coalescent teeth ending in molari- form tooth near toothless basal section; slightly enlarged tooth opposing tip of fixed finger; mesial more or less concave surface bearing few crowded pearly tubercles prox- imally, curved extensor surface bearing 2 619 rows of dense setae, with clustered granules proximally. Pereopod 2 reaching to near midlength of cheliped palm; carpus with small acute dis- todorsal and subdistal ventral spines; merus with strong subdistal dorsal spine and prox- imal mesioventral spine. Pereopod 3 with merus bearing 4 ventral spines and clus- tered tubercles; coxa of female with slender spine lateral to gonopore. Pereopod 4 with spineless merus. Uropod with acute spine on protopod above base of mesial ramus; mesial rib of lateral ramus bearing small acute proximal spine, distal margin of both rami bearing close-set row of tiny spines and spiniform granules except for short mesial sector on each. Measurements in mm.—Holotype @, an- terior carapace length 10.2, carapace length 15.4, chela length 7.3, chela height 2.3. Remarks.—The specimen, dry when re- ceived, was softened in surfactant (Aero- sol® OT Solution) and then preserved in 70% ethanol before study. The eyestalks of Upogebia baldwini somewhat resemble those of U. acanthops Williams, 1986 in having an oblique ante- rior corneal surface, but there is no subter- minal spine mesial to the cornea. Instead, there is a row of small forwardly trending spines dorsal to the cornea along its mesial margin. Other observed conspicuous differ- ences from U. acanthops include: U. bald- wini has no corneous spinules on pleura of abdominal segments 1 and 2, the rostrum has more spines on the lateral margin, the carpus of the cheliped has no spine preced- ing the spine at the anterolateral corner, rows of spines on the dorsal and mesial sur- faces of the palm are more prominent, and the merus of leg 4 is spineless. In comparison with similar eastern Pacif- ic species in the genus, both U. baldwini and U. longipollex (Streets 1871) have the merus of leg 4 spineless, but U. baldwini is not thickly strewn with deciduous corneous spinules on its tail fan, abdominal sternites and pleura, and thoracic parts of the cara- 620 pace to either side of the thalassinidean line posterior to intersection with the cervical groove as is U. longipollex. Etymology.—The species is named for Aaron Baldwin, then of Sea and Shore Mu- seum, Port Gamble, Washington, who col- lected this and other decapod crustaceans from Pacific Middle America. Upogebia vargasae, new species Fig. 2 Material—USNM 251484. Holotype 6, Boca Guarumal [08°52'N, 83°36’W, Puntar- enas Prov.], Costa Rica, Mangroves of Ter- raba-Sierpe, from burrows with thick walls, coll. with Upogebia spinigera (Smith), Rita Vargas, 23 Nov 1995. USNM 251485. Paratypes, 1 6, 3 @ (1 ovig.), same. Both holotype and paratypes from UCRMZ, Cat. No. 2108-02; 1 6 and 4 @ returned to UCRMZ. Diagnosis.—Projections to either side of rostrum ending in laterally divergent acute spine. Postocular spine present. Abdominal sternites armed. Telson subrectangular with strongly spined transverse proximal ridge. Carpus of cheliped with 2 spines on me- siodistal margin, palm without row of spines on mesioventral surface but with cluster of scattered spines there. Merus of pereopod 2 with proximal mesioventral Spine and subdistal dorsal spine; merus of pereopod 4 spineless. Description.—Rostrum triangular, lateral margin shorter than basal width; nearly hor- izontal in lateral view, tip exceeding eye- stalks; dorsal pair of strong subapical spines followed on each side by 2—3 erect margin- al spines separated by subequal intervals, spine mesial to marginal row at base, cen- tral dorsal area hidden under setae spineless and glabrous. Rostral spines, confluent with field of postrostral spines, hidden by dense tufts of setae sprouting anterior to their bas- es. Ridge lateral to field bearing crest of about 10-12 moderate to small spines, strongest on divergent process lateral to rostrum and decreasing almost to obsoles- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cence posteriorly. Shoulder lateral to cer- vical groove armed with O-—2 obsolescent spines or tubercles above and 2-5 spines below its intersection with thalassinidean line, latter continuing strongly to posterior margin, additional 0—2 spines or tubercles above it on lateral aspect of head. Postoc- ular spine present. Abdomen armed with many spinules on sternites and few spinules on margin of pleura, mainly on segments 1 and 2. Telson with proximal transverse proximal ridge bearing row of acute reclining spines directed posteriorly, confluent with low lon- gitudinal ridge to either side occasionally bearing 1 or 2 spines proximally. Eyestalk stout, deepest at about mid- length, convex ventrally, angled slightly up- -ward in repose; cornea prominent, oval, obliquely terminal, narrower than diameter of stalk, few tubercles on mesial aspect near base. Antennular peduncle not quite reaching midlength of terminal article of antennal peduncle, proximal 2 articles together slightly longer than terminal article. Antennal peduncle with almost 7% of length extending beyond tip of rostrum; an- tepenultimate article bearing ventral subter- minal spine; scale moderate, oval. Maxilliped 3 bearing epipod. Epistomial projection rather broad in lat- eral view, bearing 2 small apical projec- tions. Chelipeds with ventral margin of ischium bearing 1 spine. Merus with row of 4—5 rel- atively strong spines on ventral margin; sin- gle subdistal dorsal spine reaching level of postocular spine. Carpus trigonal, shallow longitudinal groove laterally, anterior ven- trolateral corner with strong spine preceded by 2 much smaller spines; mesiodorsal crest of 5—10 moderate spines behind prominent spine on anterior margin partly obscured by setae in proximal part of row, series often asymmetrical; 4 or fewer spines on ante- rodorsal margin mesial to articulation with propodus often obscured by dense setae; strong spine near middle of anteromesial VOLUME 110, NUMBER 4 621 a A 47-7 “ir-2y we a re - . aii a, ay pee Raa 7 ae = Fs mG » (OH AC x = Un ji Jt SS > Fig. 2. Upogebia vargasae, USNM 251484, ¢ Holotype; a, cephalic region, lateral; b, anterior carapace, dorsal; c, parts of abdominal segments 1—2 showing spinules on sternites and edge of pleura, oblique view; d, telson, right uropod, part of abdominal segment 6, dorsal; e, cheliped, right lateral; f, crest of right cheliped carpus showing spines; g, chela and carpus, right mesial; h—-k, right pereopods 2-5. 622 margin, smaller similar spine dorsal to it, and very strong spine behind distoventral corner. Chela length about 2.3—3.2 chela height; low dorsal ridge bearing 9—17 vari- able spines partly obscured by setae, spines on distal part of ridge tending to obsoles- cence, stout spine mesial to ridge subdis- tally; mesiodorsal row of about 10-12 ir- regularly distributed small spines or spini- form tubercles becoming reduced in size or obsolescent near distal % of palmar length and with tendency for development of scat- tered spines around proximal end of row; moderate spine and row of about 4-5 smaller rounded teeth below mesial dactylar condyle, single spine on margin below lat- eral condyle; mesial surface bearing longi- tudinal row of 15 spines on upper half, prominent proximally but obsolescent dis- tally, lower half bearing about 6—8 spines variable in position but tending to arrange- ment in oblique line or tract, holotype male with additional 4—5 mesioventral spines near or on ventral keel; transversely sinuous granulo-tuberculate ridge near proximome- sial margin intersecting ventral ciliated keel bearing 2—4 obsolescent tubercles or spines. Fixed finger shorter than dactyl and more slender, continuing slightly sinuous trend of lower margin of palm and tapering to slen- der tip, 2—4 irregular strong teeth on prox- imal prehensile edge. Dactyl with corneous tip preceded on prehensile edge by strong subdistal tooth and crowded row of 7-8 small coalescent teeth ending in much larg- er tooth near base, basal section toothless; mesial more or less concave surface bearing 2 rows of closely crowded tubercles, pearly on proximal half, upper row running nearly length of finger, lower row only half as long; curved extensor surface bearing 2 rows of setae, clustered rows of tubercles between them proximally. Pereopod 2 reaching to near midlength of cheliped palm; carpus with distodorsal and subdistal ventral spines obsolescent; merus with strong subdistal dorsal spine and prox- imal mesioventral spine. Pereopod 3 with merus bearing 4 ventral spines, a few tu- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON bercles or spiniform tubercles and scattered setae; coxa of female with broad spine lat- eral to gonopore. Pereopod 4 occasionally with obsolescent, proximal, ventrolateral spine on merus. Uropod with acute strong spine on pro- topod above base of mesial ramus; mesial rib of lateral ramus bearing proximal spine, distal margin of both rami bearing row of small spines and spiniform granules except for short mesial sector on each. Measurements in mm.—Holotype 6, an- terior carapace length 11.0, carapace length 15.9, chela length 11.9, chela height 4.9; paratype ovigerous ?, same, 10.5, 15.2, 8:2, 2:6. Remarks.—Upogebia vargasae is most closely allied with U. longipollex and U. - spinigera (Smith 1871), but it differs from each in having a row of strong reclining, posteriorly directed spines on the transverse proximal ridge of the telson, and all spines on the uropods are more strongly developed than in those species. Upogebia vargasae lacks the tiny deciduous spinules that are characteristically distributed over dorsal surfaces of the tail fan in U. longipollex as well as on external surfaces of its abdomi- nal pleura and on thoracic parts of its car- apace to either side of the thalassinidean line posterior to intersection with the cer- vical groove. Upogebia vargasae and U. longipollex have conspicuously developed spinules on abdominal sternites and pleura, but U. spinigera has these only on the pleu- ra. Much like U. longipollex, U. vargasae usually has a spineless merus on pereopod 4, but occasionally there is an obsolescent, proximal, lateroventral spine; that of U. spi- nigera is armed with ventral and ventrolat- eral spines and tubercles. Etymology.—The species is named for Rita Vargas, University of Costa Rica Bi- ology School, who collected this and other thalassinideans from Pacific Costa Rica. Discussion.—In a key to known Eastern Pacific species of Upogebia (Williams 1986:7—10), the following insertion in p. 8 VOLUME 110, NUMBER 4 will aid in identification of new species de- scribed above: 12. Eyestalks with cornea normally rounded AAP eee ees 2 SESS Gants, mile Bw s,m 13 = Eyestalks obliquely truncate distally, cor- nea triangular in lateral view Eyestalk bearing short subterminal spine mesial to reduced, triangular cornea; merus of leg 4 bearing ventral spines and tuber- cles. Panama ....U. acanthops Williams = Eyestalk bearing row of small forward trending spines on dorsal surface mesial to margin of cornea; merus of leg 4 usu- ally spineless. Nayarit, Mexico ..... 2 oe ee U. baldwini n. sp. 13. Telson with proximal transverse ridge spineless - Telson with proximal transverse ridge bearing row of acute reclining spines di- rected posteriorly; merus of leg 4 usually spineless. Costa Rica U. vargasae n. sp. Abdominal segments 1 and/or 2 spined ventrally on sternites or edge of pleura 12a. 13a. = Abdominal segments 1 and/or 2 lacking ventral spines Upogebia burkenroadi Williams, 1986 Material.—University of Costa Rica, UCR 1708. 1d, 1 2, Punta Pitahaya, Guan- acaste, 8—10 m, coll. J. Cortés, 15 Jun 1991. Note on back of label translated from Span- ish, ““Living in holes covered with mud, 5 x 5 mm, chambers approx. 5 mm diameter, connection to exterior covered with sand and some algae.” Measurements in mm.—®, anterior car- apace length 3.9, carapace length 5.3, chela length 3.3, chela height 1.0; d, same, 4.5, 6.5, 4.4, 1.2. Remarks.—Upogebia burkendroadi Wil- liams, 1986, described on the basis of a 623 male (holotype) and female (paratype), with no accompanying ecological information, were the only specimens of the species known, until the material listed above came to my attention. I provisionally refer the Costa Rican material to this species after comparison with the type series. Except for smaller size, the Costa Rican specimens seem almost identical to the types, but a noticeable difference is presence of a cor- neous tip on the prehensile edge of the che- liped dactyl in the type specimens, whereas this feature is reduced or absent in the smaller Costa Rican specimens. Develop- ment of this feature may be an age depen- dent character. Acknowledgments I am indebted to collectors, institutions listed in Materials, and to Ana Dittel, Uni- versities of Delaware and Costa Rica, for providing specimens that made this report possible. Keiko Hiratsuka Moore and Mol- lie S. Oremland enhanced the text with the excellent illustrations. Critical comments on the manuscript were provided by B. B. Col- lette and Ana I. Dittel. Literature Cited Smith, S. I. 1871. List of the Crustacea collected by J. A. McNeil in Central America. Second and Third Annual Report of the Trustees of the Pea- body, Academy of Science, for the years 1869 and 1870, pp. 87-98. Streets, T. H. 1871. Catalogue of Crustacea from the Isthmus of Panama, collected by J. A. Mc- Neil.—Proceedings of the Academy of Natural Sciences of Philadelphia 23:238—243. Williams, A. B. 1986. Mud shrimps, Upogebia, from the eastern Pacific (Thalassinoidea:Upogebi- idae).—Memoirs of the San Diego Society of Natural History, 14:1—60. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):624—628. 1997. A trans-Atlantic record of the fossil tropicbird Heliadornis ashbyi (Aves: Phaethontidae) from the Miocene of Belgium Storrs L. Olson and Cyril A. Walker (SLO) Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560, U.S.A. (CAW) Bird Group, Department of Zoology, The Natural History Museum, Akeman Street, Tring, Hertfordshire, HP 23 6AP, U.K. Abstract.—The fossil tropicbird Heliadornis ashbyi Olson, previously known from the unique holotype from the middle Miocene Calvert Formation of Maryland, is here recorded from a contemporaneous site in Belgium based on a nearly complete humerus. Its trans-Atlantic occurrence at a much higher latitude suggests that Heliadornis may have been the higher-latitude sister group of the extant tropical genus Phaethon, much as within the Sulidae gan- nets of the genus Morus are the higher latitude counterparts of the tropical boobies of the genus Sula. The three living species of tropicbirds (Phaethon, Phaethontidae) are remnants of an ancient and aberrant group that have al- most always been allied with the Pelecani- formes. The best-preserved fossil associat- ed with this group is Prophaethon shrub- solei Andrews (1899), from the early Eo- cene London Clay. This was shown to be so different from modern tropicbirds as to merit its own family, Prophaethontidae (Harrison & Walker 1976). Other very frag- mentary unnamed remains of early Tertiary birds resembling tropicbirds have been re- ported (e.g., Olson 1994), but it is uncertain to which of the two families these may be- long. Apart from Quaternary records of extant species, the first certain fossil record of the Phaethontidae proper was based on three associated bones from the middle Miocene Calvert Formation of Maryland that were described as a new genus and species, He- liadornis ashbyi (Olson 1985), which was considered to have been already too spe- cialized to be on a lineage leading to Phae- thon. This record was somewhat anomalous geographically, as modern tropicbirds are usually found only within the tropics, an exception being Bermuda, where warm wa- ters of the Gulf Stream have permitted Phaethon lepturus to colonize. That the original specimen of Heliadornis ashbyi was not some aberrant wanderer from the south is now confirmed by the discovery of a second specimen from a much higher lat- itude on the opposite side of the Atlantic. Phaethontidae Heliadornis Olson 1985 Heliadornis ashbyi Olson 1985 Referred material.—Left humerus lack- ing the area of the internal tuberosity and bicipital crest and a few fragments of mid- shaft, BMNH A9005 (Figs. 1, 2). Collected in 1979 and presented in 1985 by Jacques Herman of the Service Géolgique de Bel- gique. Locality. Belgium, Antwerp (Anvers) Province, Brussels-Antwerp motorway at the vicinity of the boundary between the communities of Berchem and Wilrijke. Ser- vice Géolgique de Belgique site designation 28W 126’. Horizon.—At the base of the “Sables d’Anvers”’ or Sands of Antwerp, Middle VOLUME 110, NUMBER 4 Fig. 1. Left humeri of tropicbirds in anconal view: A, Phaethon aethereus USNM 558044; B, Heliadornis ashbyi BMNH A9005, referred specimen from Belgium; C, Heliadornis ashbyi USNM 237226, holotype. Scale = 2 cm. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHING Fig. 2. Left humeri of tropicbirds in palmar view: A, Phaethon aethereus USNM 558044; B, Heliadornis ashbyi BMNH A9005, referred specimen from Belgium; C, Heliadornis ashbyi USNM 237226, holotype. Scale = 2 cm. VOLUME 110, NUMBER 4 Miocene. The Anversian Sands fall some- where within European Neogene land mammal stages (MN) 6 through 8 (Chene- val 1996). The sediments at the site are glauconitic sands with phosphatic concre- tions. The associated vertebrate fauna con- sists of rare cetaceans, frequent bones of teleosts, and abundant elasmobranchs. Measurements (mm).—Measurements in parentheses are from the holotype of H. ashbyi. Total length, 97.3; distance from head to distal extent of pectoralis scar 31.2 (30.2); depth through head 5.1 (5.3); prox- imo-distal extent of bicipital intumescence 16.1 (16.0); width and depth of shaft at midpoint, 6.5 X 5.3 (64 X 5.3); distal width, 14.1; greatest diameter of brachial depression, 9.7; depth through external con- dyle, 8.2; greatest diameter of external con- dyle, 7.1. Comparisons.—As far as they preserve portions in common, both the new speci- men and the holotype of Heliadornis ashbyi are identical to one another and distinct from Phaethon in the manner outlined in the original diagnosis, especially in the great difference in the shape of the pectoral crest. Compared with Phaethon aethereus, which is the most similar in size (P. rub- ricauda is larger, P. lepturus smaller), the Belgian specimen of Heliadornis is some- what larger with a straighter, more robust shaft. The distal end is wider because of the greater extension of the entepicondylar area, the brachial depression is wider and deeper, and the external condyle is shorter and more ovoid. Discussion.—The age of the Belgian specimen appears to be almost exactly equivalent to that of the holotype of Helia- dornis ashbyi, which came from Bed 11 of the Plum Point Member of the Calvert For- mation and is thus earliest Middle Miocene (Langhian) in age (Ward 1992: figs. 2, 3). The Belgian specimen, coming from the base of the Anversian Sands, is possibly equivalent to MN 6, which is also at least partially Langhian in age (Steininger et al. 1989, fig. 1). In absolute terms, this was 627 approximately 14 million years ago. These specimens provide such a nice trans-Atlan- tic correlation that it is tempting to cite the ornithostratigraphy as evidence for the con- temporaneity of the Calvert Formation and the Anversian Sands. The holotype of Heliadornis ashbyi comes from just below 39° north latitude, Whereas the Belgian specimen is from much higher, just above 51°N. Mlikovsky (1997) has described a second species in this genus, Heliadornis paratethydicus, based on an abraded proximal end of an ulna from the late Miocene at Brunn-V6s- endorf, Austria, in a deposit that formed in the Paratethys Sea at about 48°N. This was referred to Heliadornis because it was ge- nerically distinct from Phaethon and con- sidered a new species because it was larger than H. ashbyi. Thus Heliadornis was ap- parently a truly high latitude tropicbird, the scarcity of which is probably a reflection of strictly pelagic habits similar to those of modern tropicbirds of the genus Phaethon. In the Tertiary, the situation in tropicbirds may have paralleled that seen today in the Sulidae, in which gannets of the genus Mo- rus are found in temperate waters at higher latitudes whereas boobies of the genus Sula occur in the tropics. Heliadornis may there- fore have been the high latitude subtropical counterpart of the tropical Phaethon. The same factors, including the rigors of Pleis- tocene climates and the closing of the Pan- amanian seaway, that may have been re- sponsible for the reduction in diversity of Morus from three species in the Calvert Formation to a single species in the North Atlantic today may also have brought about extinction in the higher latitude lineage of tropicbirds. Acknowledgments We are greatly indebted to Jacques Her- man for collecting the specimen, donating it to The Natural History Museum, formerly the British Museum (Natural History) (BMNH) and for providing information 628 concerning it, and to him, Larry D. Martin, and Jiri Mlikovsky for commenting on the manuscript. The photographs are by John Steiner and Carl Hansen of Smithsonian Photographic Services. Literature Cited Andrews, C. W. 1899. On the remains of a new bird from the London Clay of Sheppey.—Proceed- ings of the Zoological Society of London 1899: 776-775. Cheneval, J. 1996. Tertiary avian localities of Bel- gium. Pp. 535-540 in J. Mlikovsky, ed., Terti- ary avian localities of Europe.—Acta Univer- sitatis Carolinae Geologica 39 (for 1995):519— 846. Harrison, C. J. O., & C. A. Walker. 1976. A reap- praisal of Prophaethon shrubsolei Andrews (Aves).—Bulletin of the British Museum (Nat- ural History) Geology 27:1-—30. Mlikovsky, J. 1997. A new tropicbird (Aves: Phae- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON thontidae) from the late Miocene of Austria.— Annalen des Naturhistorischen Museums in Wien, 98A:151—154. Olson, S. L. 1985. A new genus of tropicbird (Pele- caniformes: Phaethontidae) from the Middle Miocene Calvert Formation of Maryland.—Pro- ceedings of the Biological Society of Washing- ton 98:851—855. . 1994. A giant Presbyornis (Aves: Anserifor- mes) and other birds from the Paleocene Aquia Formation of Maryland and Virginia.—Pro- ceedings of the Biological Society of Washing- ton 107:429—435. Steininger, EK E, R. L. Bernor, & V. Fahlbusch. 1989. European Neogene marine/continental chrono- logic correlations. Pp. 15—46 in E. H. Lindsay, V. Fahlbusch, and P. Mein. European Neogene mammal chronology. New York: Plenum Press. 658 pp. Ward, L. 1992. Molluscan biostratigraphy of the Mio- cene, Middle Atlantic coastal plain of North America.—Virginia Museum of Natural History Memoir 2, 159 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):629—639. 1997. A new species of Cyclemys (Testudines: Bataguridae) from Southeast Asia John B. Iverson and William P. McCord (JBI) Department of Biology, Earlham College, Richmond, Indiana 47374, U.S.A.; (WPM) East Fishkill Animal Hospital, Hopewell Junction, New York 12533, U.S.A. Abstract.—A new species of batagurid turtle, Cyclemys atripons, is described from the mountainous areas of southeastern Thailand and adjacent Cambodia. It differs from other Cyclemys (herein all referred to as C. dentata) by the unigue combination of a head mottled with black dorsally and striped laterally, a nearly immaculate chin, a distinct carapace pattern, a plastron with no or only a few coarse black rays, a coarsely and densely pigmented bridge, a narrow carapace, a long plastral hindlobe, a wide plastral forelobe, a small gular scute, and a long interhumeral seam length. Discriminant function anal- ysis of 17 morphometric characters standardized for body size supported the distinctiveness of the new species from Cyclemys dentata, and indicated that other populations of this complex (on Borneo and in China) may also be mor- phologically distinct. The genus Cyclemys is currently envi- sioned by most authors to comprise two species (C. dentata Gray 1831 and C. tche- ponensis Bourret 1939) with very uncertain distributions (Ernst & Barbour 1989, Iver- son 1992). Cyclemys dentata was originally described by Gray (1831:20) as Emys Dhor (type locality: ‘““Bengal ... Java’’), but he corrected the name to Emys dentata in the Errata of that publication. However, for the next 100 years the species name dhor was used by most authors (including Gray on occasion, e.g., 1870 and 1872). During that period several additional names (now rec- ognized as synonyms) were applied to this taxon: Cyclemys orbiculata Bell 1834; Cis- tudo diardi Duméril & Bibron 1835; Cycle- mys oldhami Gray 1863; Cyclemys ovata Gray 1863; and Cyclemys bellii Gray 1863. Stejneger (in Barbour 1912:143) and Smith (1930:8) both argued that the correct spe- cies name is dentata and nearly all authors subsequent to Smith have used the latter name. As defined by Smith, C. dentata ranges from northeastern India to south- western China, and through southeast Asia to Sumatra, Java, Borneo and the Philip- pines (Taylor 1920, Smith 1930, Zhao & Adler 1993). In 1939, Bourret recorded C. dentata from Hanoi, Vietnam, but described Geoe- myda tcheponensis from central Vietnam, based on a juvenile in the Hanoi Museum. Wermuth & Mertens (1961) first recognized the similarity of G. tcheponensis and C. dentata (even though the two species were originally described in separate genera) and synonymized the former under the latter. In a comprehensive study of the generic rela- tionships among all the pond turtles (emy- dines and batagurines), McDowell (1964) supported that synonymy. However, in 1976, after examining 16 pet trade or mar- ket specimens of Cyclemys, McMorris ar- gued that tcheponensis and dentata were distinct species characterized by color pat- tern differences (the former having a head mottled dorsally and striped laterally, and the latter lacking head and neck stripes and having the dorsum of its head uniform brown). During the following 20 years, most authors have cautiously recognized 630 both, pending further study (e.g., Pritchard 1979, Ernst & Barbour 1989, Iverson 1992). A single anomalous specimen of Cy- clemys with only 11 marginal scutes per side was described from southern Yunnan, China by Kou (1989) as C. tiannanensis, but that name was quickly synonymized with dentata by Das (1991:80) and Zhao & Adler (1993:167). In 1994, distinctive turtles labelled as Cy- clemys dentata began appearing in the pet trade from southeast Asia. According to the exporters, these turtles originated from Ton- le Sap, near Phnom Penh, Cambodia (An- son Wong, pers. comm.). These unique tur- tles clearly belonged to the genus Cyclemys (sensu McDowell 1964), but in order to compare them with other Cyclemys we be- gan examining variation across all known populations of the genus. Our full analysis of variation is not yet complete (Iverson, McCord, van Dijk, Das, and Moll, pers. comm.), but our preliminary morphometric comparisons of over 180 Cyclemys repre- senting all known areas of the range, of most of the type specimens, and of all phe- notypes from typical tcheponensis to typical dentata (sensu McMorris 1976) revealed the existence of an unnamed population ap- parently associated with the Cardamom up- lift in southeastern Thailand and adjacent Cambodia; indicated that tcheponensis is apparently not morphometrically distinct from dentata; and suggested that popula- tions on Borneo and from China may also each be distinctive. Although the new taxon seems to have a very limited distribution, it is now exported from Cambodia in great numbers. There- fore, it seems appropriate to name it now, before our full analysis of variation in the genus is complete, so that it might soon be given appropriate legal protection. Materials and Methods Shell and scute measurements (to the nearest 0.1 mm) were recorded from all available museum and pet trade specimens PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of Cyclemys (sensu McDowell 1964). Be- cause we observed specimens with both the tcheponensis and dentata phenotypes from nearly everywhere across the composite range and because specimens without heads could not be allocated confidently to either phenotype, we lumped specimens by ge- ography, establishing ten populations for analysis: India and Burma (QO), northern Cambodia/Laos/Vietnam (1), southeastern Thailand and adjacent Cambodia (2), Thai- land (3, excluding southeast Thailand), Chi- na (4), Malay Peninsula (5), Borneo (6), Sumatra (7), Java (8), and Philippines (9). Preserved material was borrowed from the American Museum of Natural History (AMNH), the British Museum of Natural History (BMNH), the California Academy -of Sciences (CAS), the Field Museum of Natural History (FMNH), the Museum of Comparative Zoology at Harvard (MCZ), the National Museum of Natural History in Paris (MNHN), the Oxford University Mu- seum (OUM), the National Museum of Nat- ural History in Leiden (RMNH), the Florida Museum of Natural History at the Univer- sity of Florida (UF), the University of Kan- sas Museum of Natural History (KU), and the United States National Museum (USNM), and living material was available in McCord’s private collection (WPM) (Fig. 1). Recorded measurements included max- imum (not midline) carapace length (CL), maximum carapace width (CW), maximum carapace height (CH), maximum (not mid- line) plastron length (PL), maximum (not midline) length of the plastral forelobe from the interabdomino-interpectoral junction to a line across the anterior ends of the gular scutes (FL), maximum (not midline) length of the plastral hindlobe from the interab- dominal-interfemoral junction to a line across the posterior ends of the anal scutes (HL), minimum (median) length of the plastral hindlobe from the interabdominal- interfemoral junction to the anal notch (NHL), plastral forelobe width at the level of the junction of the humeropectoral seam and the plastral margin (PWA), anterior VOLUME 110, NUMBER 4 631 Fig, iL, represent literature or museum locality records. Hatched area in Thailand and Cambodia encircles hypothesized range of Cyclemys atripons. plastral hindlobe width at the level of the junction of the abdominofemoral seam and the plastral margin (PWC), posterior plas- tral hindlobe width at the level of the junc- tion of the femoroanal seam and the plastral margin (PWD), left bridge length from ax- illa to inguinal pocket (BL), maximum ven- tral width across gular scutes (GW), maxi- mum (not midline) length of right gular scute (GL), and lengths of right interhu- meral (IH), interpectoral (IP), interabdom- inal (IAB), interfemoral (IF), and interanal (IAN) seams. Although there is some sex- ual dimorphism in some of these characters, females and males were analyzed together in order to include dried museum speci- mens lacking soft parts and subadults that Composite range map of Cyclemys in southeast Asia (from Iverson, 1992, and unpublished). Dots could not be sexed confidently. Only adult and subadults (>110 mm CL) were includ- ed in this preliminary analysis. The data were standardized for body size by division by carapace length. Although concerns have been expressed about the sta- tistical validity of using ratios rather than residuals in quantitative analyses (Atchley et al. 1975, 1976; among others), multivar- iate analyses of ratios of turtle morphomet- ric data have not yielded results that dif- fered from those employing residuals (e.g., Berry 1978, McCord & Iverson 1991). In addition, the use of ratios offers the advan- tage of working with parameters that can be directly measured and/or compared; it is not possible to compare raw measurements DF1 Fig. 2. Plot of first two canonical axes (discrimi- — nant functions DF1 and DF2) for specimens of Cycle- mys based on discriminant function analysis of the ra- tios of 17 characters (listed in Table 1). First and sec- ond axes account for 44.9% and 27.2% of the varia- tion, respectively. Country codes are India and Burma (0), northern Cambodia/Laos/Vietnam (1), southeast- ern Thailand and adjacent Cambodia (2), Thailand (3, excluding southeast Thailand), China (4), Malay Pen- insula (5), Borneo (6), Sumatra (7), Java (8), and Phil- ippines (9). The arrow marks the adult syntype of Cy- clemys ovata Gray, which is grouped among the other Borneo turtles with a probability of 93%. directly to mean values of a residual. The 17 character ratios produced by this stan- dardization were then submitted to discrim- inant function analysis (DFA) with SPSS software (SPSS, Inc. 1983). ANOVA and pairwise population comparisons for the ra- tios of each variable to CL were made with Fisher’s (protected) least significant differ- ence (PLSD) test with STATVIEW soft- ware (Abacus Concepts 1992). Results and Discussion Both the discriminant function analysis (DFA; Fig. 2) and the analyses of variance of the ratios for the various populations of Cyclemys (Table 1) demonstrated that the population in southeast Thailand and adja- cent Cambodia is morphometrically the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON most distinct of all populations. They also suggested that turtles from Borneo and those from China are divergent. The anal- yses revealed that, compared to other Cy- clemys populations, those from SE Thai- land/Cambodia tend to have a narrower car- apace, a longer plastral hindlobe, a wider plastral forelobe, a narrower and shorter gu- lar scute, and a longer interhumeral seam length; those from Borneo tend to have a deeper shell, a longer hindlobe, a longer gu- lar scute, a shorter interhumeral seam length, a longer interfemoral seam length, and a shorter interanal seam length; and those from China tend to have a very nar- row Shell, a shorter and narrower plastral forelobe, a narrower plastral hindlobe, and a wider gular scute. Untransformed data for the characters that varied most significantly in the ANO- VA’s and DFA (Table 1) were recombined into character ratios and examined to sum- marize variation within the genus Cyclemys (Table 2). That analysis reinforced the dis- tinctiveness of the Thailand/Cambodia pop- ulation and demonstrated significant over- lap among most of the other populations; however, confirmation of the uniqueness of the Borneo and/or Chinese populations must await our additional analyses. Because the Thailand/Cambodia population is both allopatric and morphometrically distinct from other populations of Cyclemys, and because that population also differs in color patterns of the head and shell, we here de- scribe it as: Cyclemys atripons, new species Black-bridged leaf turtle Fig. 3. Holotype-—USNM 81865, a dried shell of a female with limbs but no head, from Thailand, Krat [=Trat], Kao [=Mt.] Kuap (=Khao Kuap), collected 24 December 1929 by Hugh M. Smith. Paratypes—_USNM 94745, an unsexed juvenile from Thailand, Chanthaburi, Kao [Mt.] Sabab (=Khao Sabap) collected 20 VOLUME 110, NUMBER 4 633 Table 1.—Results of univariate analysis of variance of ratios of 17 characters to carapace length across 10 populations of Cyclemys and for Cyclemys atripons (n = 29) versus all other populations (n = 106). Across all populations C. atripons vs. others Character if je if P Maximum carapace width (CW) WAY <0.0001 8.1 0.005 Maximum carapace height (CH) 8.1 <0.0001 11.0 0.0011 Maximum plastron length (PL) 1.9 0.06 6.5 0.012 Maximum forelobe length (FL) 0.6 0.78 4 0.24 Maximum hindlobe length (HL) 312 0.0017 5.8 0.018 Medial hindlobe length (NHL) 3.4 0.001 OF 0.0022 Plastral width (PW1) 4.1 <0.0001 1253 0.0006 Plastral width (PW3) 2 <0.0001 0.3 0.61 Plastral width (PW4) 2.6 0.008 0.4 O55 Bridge length (BL) 2 0.029 4.7 0.032 Gular width (GW) 2.8 0.0046 9.5 0.0025 Gular length (GL) Zell 0.032 8.3 0.0046 Interhumeral seam length (1H) 3.0 0.0025 14.1 0.0003 Interpectoral seam length (IP) 1.2 0.28 25 Ostz Interabdominal seam length (IAB) Syd 0.0022 ibe 0.20 Interfemoral seam length (IF) 4.2 <0.0001 0.03 0.87 Interanal seam length (IAN) 4.3 <0.0001 4.0 0.047 November 1931 by Hugh M. Smith. USNM 53423, a dried shell of a female with sep- arate head, limbs, and viscera in alcohol, and USNM 53424, an unsexed juvenile pre- served in alcohol, from Thailand, Trat, Koh [= Ko = island] Chang, both collected in December 1914 by C. B. Kloss (see Smith & Kloss 1915). MCZ 29571, an unsexed juvenile preserved in alcohol, and MCZ 29572, a subadult (possibly male) in alco- Table 2.—Morphometric characters useful in distin- guishing Cyclemys atripons (n = 29) from other Cy- clemys (n = 106). Character abbreviations are maxi- mum carapace width (CW), maximum shell height (SH), width of anterior plastral lobe (PWA), median length of plastral hindlobe (NHL), maximum gular width (GW), maximum gular length (GL), and inter- humeral seam length (IH). Values are means followed by range in parentheses. All character ratios are highly significantly different (P < 0.0001) by Fisher’s (pro- tected) least significant difference test. Character Cyclemys atripons Other Cyclemys IH/SH 0.293 (0.176—0.425) 0.231 (0.095—0.529) IH/CW 0.132 (0.088-0.199) 0.106 (0.045—0.186) PWA/CW 0.598 (0.503-0.671) 0.554 (0.451-0.651) GL/NHL_ 0.278 (0.201-0.308) 0.305 (0.228—0.377) GW/ PWA 0.414 (0.335-0.503) 0.452 (0.371—-0.562) hol from Thailand, Gulf of Siam, Koh Kong (=Ko Chang), both donated in 1930 by Malcolm Smith. USNM 79515, a com- plete, mummified adult female from Thai- land, Trat, Koh Kut (=Ko Kut), Gulf of Thailand collected 21 May 1929 by Hugh M. Smith. KU 47171 (formerly MCZ 29558), a dried shell of an adult, from ‘‘Bangkok, Siam’ (certainly the shipping point rather than the actual collection lo- cality), donated in 1930 by Malcolm Smith. UF 105992, an adult male preserved in al- cohol, and UF 105993, a dried skeleton of an unsexed subadult; reported to have been collected from Tonle Sap, Phnom Penh, Cambodia [=Kampuchea], but purchased from local people in Phnom Penh by Mr. Anson Wong in the spring of 1994. Diagnosis.—A medium-sized species of Cyclemys (carapace to 224 mm, but usually less than 200; to at least 250 mm in other Cyclemys) with an elongate, basically tri- carinate carapace having an obvious mid- dorsal keel and weak lateral keels and being nearly flat dorsally in adults; a plastron hinged between the pectoral and abdominal scutes (hyoplastral and hypoplastral bones); a very weakly hooked upper tomium with 634 a medial shallow notch or cusp; a relatively wide carapace (maximum width averages 78% of carapace length [CL] in C. atripons; 82% in other Cyclemys); a relatively low carapace (maximum shell height averages 35% of CL in C. atripons; 37% in other Cyclemys); a relatively wide plastral fore- lobe (posterior width averages 46% of CL in C. atripons; 45% in other Cyclemys); a relatively long plastral hindlobe (median length averages 50% of CL in C-. atripons; 49% in other Cyclemys); a small gular scute (maximum length and width average 19% and 14% of CL, respectively, in C. atri- pons; 20% and 15%, respectively, in other Cyclemys); and a relatively long interhu- meral seam (length averages 10% of CL in C. atripons; 8.5% in other Cyclemys) [see also Table 2 and Fig. 3]; a light cream to brown temporal and postorbital stripe on each side of head extending posteriorly onto the neck (flushed with salmon in ju- veniles), and a third narrower light stripe extending from the angle of the jaw to the ventral margin of the tympanum (some- times connected to the postorbital stripe by a short vertical branch along the anterior margin of the tympanum) (basic pattern is similar to that in other Cyclemys); a nearly immaculate cream or yellow chin, lightly flushed with salmon in juveniles and some- times with a few vague black flecks (darkly mottled to almost completely black in other Cyclemys); carapace light olive to brown to nearly black, with coarse black rays radi- ating from the areolae, but the lateral rays on Cl—C3 disappearing with age, leaving the anterior rays in a bold, dense triangular concentration on the anterodorsal half of C1-—C3 (carapacial rays, if visible, evenly distributed across costal scutes in other Cy- clemys); the plastron with no or only a few coarse black rays on a cream to yellow- _ brown to horn-colored background, but with the bridge coarsely and densely flecked or streaked with black (entire plas- tron almost always heavily streaked with black [sometimes completely brown or black] in other Cyclemys). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Description (based on the type series as well as 12 adult females, 8 adult males, 6 unsexed subadults or juveniles alive in McCord’s collection).—Carapace length to at least 191 mm in males and at least 224 mm in females, elongate, moderately tricar- inate with a prominent medial keel and weak lateral keels, not domed (maximum Shell height/CL = 0.308—0.395; mean = 0.352), nearly flat dorsally in full adults, widest at marginal M7 (maximum carapace width/CL = 0.703—0.898; mean = 0.781), with a slightly serrated posterior margin, and with moderately obvious growth annuli (least obvious in old individuals). Margin- als all fairly uniform in length (along car- apace margin); M5, 6.7, or 12 tallest; M9-— 11 usually flared. Cervical scute medium in size, usually longer than wide, indented me- dially along the posterior margin, and oc- casionally wider posteriorly than anteriorly. Vertebrals V2—5 wider than long; V1 usu- ally wider than long, but not contacting seam between M1 and M2; V5 not even close to contacting M10. Prominent medial keel most pronounced on V4 and V5, but also obvious on V1 and V3; lateral keels weak, but most pronounced on costal C3. Carapace olive to brown to nearly black, with seams more darkly marked; coarse black rays radiate from the areolae; how- ever, rays extending to the marginals on costals C1l—C3 fade with age such that adults appear to have a dense triangular concentration of black streaks on the ante- rodorsal half of C1—C3: (Fig. 3); carapacial keels not distinctly colored unless due to abrasion, but medial keel in subadults and most adults tending to lack black pigment, giving the impression of a weak, light mid- vertebral stripe (particularly evident on the nuchal scute). Maximum plastron length shorter than carapace length (PL/CL = 0.899-1.039; mean = 0.975). Plastron very slightly up- turned anteriorly, with hinge present be- tween the hyoplastral and hypoplastral bones (approximately aligned with the seam between the pectoral and abdominal VOLUME 110, NUMBER 4 635 ie ae s+) rs ~eFS Rate ¥ a) - S Fig. 3. Cyclemys atripons. Left (plastral views; top to bottom): carapace of first year turtle, USNM 94745, 78 mm carapace length (CL); subadult, WPM 001, 132 mm CL; adult, WPM 003, 191 mm CL. Right (top to bottom); juvenile, WPM 002, 135 mm CL; subadult, WPM 001, 132 mm CL; adult female, WPM 004, 196 mm CL. 636 scutes). Plastral forelobe width (PWA) at level of junction of humeropectoral seam and lateral plastral margin relatively wide (PWA/CL = 0.420—0.507; mean = 0.464). Median length of plastral hindlobe lobe (NHL) relatively long (NHL/CL = 0.441- 0.536; mean = 0.501). Plastral hindlobe with relatively shallow anal notch. Bridge moderately long (BL/CL = 0.304-0.360; mean = 0.336); axillary and inguinal scutes very reduced or absent. Gular scute rela- tively small (GW/CL = 0.162—0.216; mean = 0.192; GL/CL = 0.108—0.163; mean = 0.139), but interhumeral seam relatively long (IH/CL = 0.065-—0.155; mean = 0.103). Average plastral formula (see also Table 2 and Fig. 1 for diagnostic ratios): interpectoral seam (IP) > interabdominal seam (IAB) = interanal seam (IAN) > gu- lar length (GL) = interfemoral seam (IF) = interhumeral seam (IH). Plastron cream, yellow-brown, or horn colored, with seams more darkly marked with brown; no or only a few short, coarse, black flecks or rays present (Fig. 3); bridge area with obvious coarse black rays or flecks radiating ante- riorly and laterally (not medially) from the scute areolae. Ventral surfaces of M4—M7 more boldly marked with black rays or flecks than on other marginals. Head of medium width; upper jaw weak- ly hooked, but with a shallow median notch; triturating surfaces narrow. A few small tubercles evident between angle of jaw and tympanum. Dorsum of head coarsely flecked with black on a greenish- brown background. A light cream to brown temporal and postorbital stripe on each side of head, extending posteriorly onto the neck (flushed with salmon in juveniles), and a third narrower light stripe extending from the angle of the jaw at least to the ventral margin of the tympanum (sometimes con- nected to the postorbital stripe by a short vertical branch along the anterior margin of the tympanum); these stripes darken with age but are still evident even in old indi- viduals (see Fig. 3). Chin cream or yellow (ightly flushed with salmon in juveniles), PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with a few or no vague black markings; most frequent black mark on chin a short, thin median line subequal to orbit diameter. Neck with six vague light stripes ventrally (parallel to and between the stripes extend- ing posteriorly from the angle of the jaws). Tomia cream to yellow brown to dark gray with several coarse black streaks. Vague black horizontal line across eye (through pupil); iris light green to brown. Anterior surface of antebrachium covy- ered with large, imbricate scales, the largest of which are crescent to spade-shaped; larg- est scales on hindlimb at heel, but generally smaller than largest forelimb scales. Upper parts of limbs and tail covered with fine scales. Exposed dorsal parts of forelimbs dark brown (rarely almost black) with black - flecks; ventral surfaces cream to yellow and unmarked; dorsal surface of digits generally lighter in color than lateral surfaces. Dorsal surface of hindlimbs dark gray-brown to black; ventral surfaces cream colored. Nar- row cream colored stripe on posterior mar- gin of each hind limb, extending at least to heel; dark stripe immediately ventral to it, extending to anal region in all but the oldest Specimens. Cream colored soft parts be- tween hindlimbs and tail washed with salm- on in some juveniles. Recessed areas of in- guinal and axillary regions and between neck and forelimbs uniform cream color. Tail (even in males) relatively short, dark gray to brown, with a black middorsal stripe and a ventrolateral dark brown stripe on each side. Males and females both with a flat or slightly convex plastron. Males with a slightly longer tail than females; vent gen- erally at level of posterior carapace margin in males and anterior to it in females. The single available, incompletely pre- pared skull associated with UF 105993 has the maxillae separated anteriorly by the pre- maxillae; pterygoid barely in contact with the jugal; foramen posterius palatinum small and similar in size to the foramen or- bitonasale; fissura ethmoidalis extremely narrow ventrally, distinctly key-hole VOLUME 110, NUMBER 4 shaped; cranial cavity not ventrally nar- rowed by the processi inferiores parietales; and quadratojugal present as part of post- orbital bar. Etymology.—From the Latin, atri, mean- ing black, and pons, meaning bridge, in ref- erence to the darkly pigmented bridge in this species that contasts so strongly with the usually unmarked plastron. Other material.—Twenty-six live speci- mens in the collection of William P Mc- Cord (WPM 1-26), to be deposited on death in the UF collection. Distribution.—Apparently restricted to the isolated hill country of extreme south- eastern Thailand and southwestern Cam- bodia, in association with the Cardamon mountain range, including the Gulf of Thai- land islands of Koh Chang and Koh Kut (the two largest and most mountainous is- lands in the area). Recent pet trade speci- mens, said to be coming from Tonle Sap, a lowland wetland near Phnom Penh, Cam- bodia, more likely came from the hill coun- try to the southwest. The nearest confirmed record for Cyclemys dentata is from Saka- eret, Thailand (FMNH 183723-39; Inger & Colwell 1977), ca. 275 km to the north- northwest. Remarks.—Our preliminary analyses support the suggestion by Das (1995a, 1995b) that the population on Borneo is distinctive. Should our further analysis con- firm its distinctiveness, the name Cyclemys ovata (Gray 1863:178 from Sarawak) is available (see also Fig. 2). If our prelimi- nary results regarding the distinctiveness of the Chinese population are confirmed, the name Cyclemys tiannanensis Kou 1989:193 (from Yunnan) may be applicable to the Chinese population we sampled, even though the name was based on a single ab- errant specimen. In our preliminary analysis we could find no morphometric character to distinguish tcheponensis from dentata. It has been suggested that color pattern dif- ferences between these two may reflect al- titudinal variation more than regional vari- ation (Frank Yuwono, pers. comm.). How- 637 ever, our final conclusion on the validity of tcheponensis must await completion of our broader analysis (including juveniles). Zoogeography.—Cyclemys atripons is apparently not the only species endemic to the hill country of southeastern Thailand and adjacent Cambodia. According to Peter Paul van Dijk (pers. comm.), at least the following two frogs and two lizards are en- demic to the same area: Rana fasciculispi- na, Rana kohchangae, Isopachys roulei, and Pseudocalotes floweri. Comparative material examined: Cycle- mys dentata (sl): India (BMNH 1930.6.8.4, OUM 8502 [syntype of Cyclemys orbicu- lata], OUM 8513 [syntype of Cyclemys or- biculata and holotype of Cyclemys bellii], RMNH 6073, and USNM 293726); Burma (AMNH 58423, BMNH 1947.3.5.63 [syn- type of Cyclemys oldhami]); China (WPM 6 specimens); Thailand (BMNH 1947.3.4.26 [syntype of Cyclemys oldhami], BMNH 62.8.18.20, FMNH 183727-728, 183730, WS5732, “lS3755, 85737-7738, MCZ 29561-566, 29568—-570, 29573, 43066— 067, 43084, USNM 26249, 94602, 269918- 919, WPM 3 specimens); Laos (USNM 103016); Vietnam (MNHN _ 1948.38, RMNH 4751, USNM 95100, WPM 12 spec- imens); Malay Peninsula (AMNH 49933, FMNH 142501, 166553, 224081-082, 224085—-092, 251501, 251508, USNM 30961, 30964, WPM 8 specimens); Suma- tra (BMNH 1979.221, CAS-SU 8615, RMNH 3838, USNM 37792, WPM 11 specimens); Java (BMNH 1946.1.22.62 and 1946.1.22.63 [syntypes of Emys dentata], MNHN 9107 [syntype of Cistudo diardii], USNM 62576, WPM 2 specimens); Phil- ippines (AMNH 90102, CAS 62166, 134331-—332, 157280, FMNH 51598, KU 47172, 1917/6, MC, 25369, -USNM 229500—501, 496884); and Borneo (BMNH 63.6.21.1 [syntype of Cyclemys ovata}, FMNH 14974, 63276-278, 128257, 128259, 151015, 166554, MCZ 11244, USNM 38534, WPM 16 specimens). 638 Acknowledgments The distinctiveness of this new species would not have come to our attention with- out the collecting efforts of Anson Wong of Malaysia and Oscar Shiu of Hong Kong. The generous loan of specimens by the American Museum of Natural History (AMNH; D. Frost), the British Museum of Natural History (BMNH; especially C. Mc- Carthy), the California Academy of Sci- ences (CAS; J. V. Vindum), the Field Mu- seum of Natural History (FMNH; H. Voris and A. Resetar), the University of Kansas (KU; W. E. Duellman), the Paris Museum (MNHN; R. Bour), the Oxford University Museum (OUM; J. Pickering), the Leiden Museum (RMNH; M. S. Hoogmoed), the University of Florida (UF; D. Auth), and | the United States National Museum of Nat- ural History (USNM; R. Crombie, G. R. Zug) also made this study possible. E. O. Moll allowed JBI access to FMNH speci- mens in his care. P. P. van Dijk and M. Dioli provided photographs and information on Cyclemys from Thailand and northern Cam- bodia, respectively. Comments by I. Das, E. O. Moll, and P. P. van Dijk on an early draft of the manuscript are greatly appreciated. We thank L. Thorpe for her insights into Latin grammar. Financial support was pro- vided by Earlham College, the Joseph Moore Museum of Natural History, and Iverson’s family. Literature Cited Abacus Concepts. 1987. Statview. Abacus Concepts, Inc., Berkeley, California, 466 pp. Atchley, W. R., C. T. Gaskins, & D. Anderson. 1975. Statistical properties of ratios of biological data——American Zoologist 15:829. . 1976. Statistical properties of ratios I. Em- pirical results.—Systematic Zoology 25:137-— 148. Barbour, T. 1912. A contribution to the zoogeography of the East Indian islands——Memoirs of the Museum of Comparative Zoology 44:1—203. Bell, T. 1834. Characters of a new genus of freshwater tortoises (Cyclemys).—Proceedings of the Zoo- logical Society of London 1834:17. Berry, J. E 1978. Variation and systematics in the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Kinosternon scorpioides and K. leucostomum complexes (Reptilia: Testudines: Kinosternidae) of Mexico and Central America. Unpublished Ph. D. dissertation, University of Utah, Salt Lake City, 325 pp. Bourret, R. 1939. Notes herpétologiques sur l’Indochine frangaise. XVI. Tortues de la col- lection du Laboratoire des Sciences Naturelles de l'Université. Description d’une espéce nou- velle-—Annexe au _ Bulletin Général de l’Institution Publique (Hanoi) 1939(6):1—34. Das, I. 1991. Colour guide to the turtles and tortoises of the Indian subcontinent. R & A Publ., Avon, England, 133 pp. 1995a. An illustrated key to the turtles of insular south-east Asia—Hamadryad 20:27— 32. . 1995b. Turtles and Tortoises of India. Oxford University Press, Bombay, India, 176. pp. Dumeéril, A.-M.-C., & G Bibron. 1835. Erpétologie Générale ou Histoire Naturelle des Reptiles. Tome Second, Roret, Paris, 680 pp. Ernst, C. H., & R. W. Barbour. 1989. Turtles of the World. Smithsonian Institution Press, Washing- ton, D.C., 313 pp. Gray, J. E. 1831. Synopsis Reptilium, or short de- scription of the species of reptiles. Treuttel, Wurtz & Co., London, 85 pp. . 1863. Observations on the box tortoises, with descriptions of three new Asiatic species.—Pro- ceedings of the Zoological Society of London 1863:173-179. . 1870. Supplement to the catalogue of shield reptiles in the collection of the British Museum. Part I. Testudinata (Tortoises). Taylor and Fran- cis, London, 120 pp. 1872. Appendix to the catalogue of shield reptiles in the collection of the British Museum. Part I. Testudinata (Tortoises). British Museum, London, 28 pp. Inger, R. F, & R. K. Colwell. 1977. Organization of contiguous communities of amphibians and rep- tiles in Thailand.—Ecological Monographs 47: 229-253. Iverson, J. B. 1992. A revised checklist with distri- bution maps of the turtles of the world. Iverson Publishing, Richmond, Indiana, 363 pp. Kou, Z. 1989. Cyclemys from Yunnan, a description of a new species and a new record to China (Testudinata:Emydidae).—Current Herpetology in East Asia: 193-197. McDowell, S. B. 1964. Partition of the genus Clem- mys and related problems in the taxonomy of the aquatic Testudinidae.—Proceedings of the Zoological Society of London 143:239-279. McCord, W. P., & J. B. Iverson. 1991. A new box turtle of the genus Cuora (Testudines: Emydi- VOLUME 110, NUMBER 4 dae) with taxonomic notes and a key to the spe- cies.—Herpetologica 47:407—420. McMorris, J. R. 1976. The generic reassignment of Geoemyda tcheponensis Bourret.—Chelonia 3(2):10-11. Pritchard, P. C. H. 1979. Encyclopedia of Turtles. T. E H. Publ., Inc., Neptune, New Jersey, 895 pp. Smith, M. A. 1930. The Reptilia and Amphibia of the Malay Peninsula from the Isthmus of Kra to Singapore, including the adjacent Islands (A Supplement to Dr. G. A. Boulenger’s Reptilia and Batrachia, 1912).—Bulletin of the Raffles Museum, Singapore, Straits Settlements 3:1-— 138. 639 , & B. Kloss. 1915. On reptiles and batrachi- ans from the coast and islands of south-east Siam.—Journal of the Natural History Society of Siam 1:237—249. SPSS, Inc. 1983. SPSS* User’s Guide. McGraw-Hill, New York, 949 pp. Taylor, E. H. 1920. Philippine Turtles.—The Philip- pine Journal of Science 16(2):111—144. Wermuth, H., & R. Mertens. 1961. Schildkréten, Kro- kodile, Briickenechsen. G. Fischer, Jena, 422 Pp. Zhao, E.-M., & K. Adler. 1993. Herpetology of Chi- na.—SSAR Contributions to Herpetology 10:1— 522: PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):640—680. 1997. Morphological differentiation among Subsaharan and North African populations of the Lemniscomys barbarus complex (Rodentia: Muridae) Michael D. Carleton and Erik Van der Straeten (MDC) Division of Mammals, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (EVDS) Departement Biologie, Rijksuniversitair Centrum, B-2020 Antwerpen, Belgié. Abstract.—Morphometric evaluation discloses patterns of craniodental vari- ation which demonstrate that populations currently assigned to Lemniscomys barbarus consist of two species: L. barbarus (Linnaeus 1766) and L. zebra (Heuglin 1864). One species-group epithet is herein allocated to the junior synonymy of L. barbarus (ifniensis Morales Agacino 1935), ten to L. zebra (spekei De Winton 1897, dunni Thomas 1903, convictus Osgood 1910, albol- ineatus Osgood 1910, manteufeli Matschie 1911, oweni Thomas 1911b, niger- iae Thomas 1912, olga Thomas & Hinton 1921, nubalis Thomas & Hinton 1923, and orientalis Hatt 1935), and a lectotype is designated for Mus zebra Heuglin (1864). No subspecies are recognized for either species, but the sub- stantial heterogeneity documented within L. zebra advises continued study of those populations. The geographic occurrence of L. barbarus is restricted to scrub vegetation along a narrow coastal strip in Morocco, Algeria, and Tunisia; whereas, L. zebra is widely distributed over dry grassland and savanna biomes south of the Sahara Desert, from Senegal in West Africa to central Sudan in East Africa, and southwards to northcentral Tanzania. Lemniscomys hoogstraali Dieterlen (1991), known only by the holotype from eastcentral Sudan, is re- tained as a third member of the barbarus group. The species L. barbarus represents another example of small mammal endemic to the Barbarian prov- ince (Maghreb) of northwest Africa. African grass mice of the genus Lemnis- comys have been conventionally arranged into three species groups or complexes— barbarus, griselda, and striatus—defined principally by the striping pattern of the dorsal pelage (Hollister 1919, Ellerman 1941, Van der Straeten & Verheyen 1980). While the griselda and striatus species groups have attracted much revisionary at- tention over the past 20 years (Van der Straeten 1975, 1976, 1980a, 1980b, 1981; Van der Straeten & Verheyen 1978, 1979, 1980), the barbarus complex, or striped grass mice, has yet to receive the same careful review, an oversight which this study begins to redress. As with other taxa named in the late 1800s through the early 1900s, forms even- tually associated under Lemniscomys bar- barus had been first described as full spe- cies (for example, zebra Heuglin 1864; spekei De Winton 1897, and dunni Thomas 1903). In 1910, however, Osgood diag- nosed two new races of striped grass mice from British East Africa (albolineatus and convictus) that he assigned to Arvicanthis barbarus and at the same time reassociated zebra Heuglin and spekei De Winton as subspecies of A. barbarus. Whether or not he was continuing the polytypic approach advanced in his 1909 revision of Peromys- cus, Osgood’s nomenclatural assessments VOLUME 110, NUMBER 4 641 barbarus.:/ (1766) & ifniensis . (1935) .. 3 Algeria Niger Sudan olga Zz (1921). f * Dia dunni * (1903) / =) nigeriae £ . / a nubalis hoogstraali : ‘ (1923) ke ey * (i912) § j \ a Nigeria ~ { Ghana *|3| 1 Type Localities of Lemniscomys barbarus Group Pigs 1. j 3 Cameroon ( — = \.zebra (1864) 1 aS ( nek A eg a hs Nr a E ._ Ky ( premen= ) Uganda» F) Kenya albolineatus (" (4910) a ae a. * 7 rp Vag ime manteufeli Lk ey convictus~ spekei io Por * : a Tanzania Map of North and Central Africa depicting the type localities (stars) of the fourteen species-group taxa allocated to striped mice of the Lemniscomys barbarus species group. The date in parenthesis below each name indicates the year of the description. established the precedent for treating striped grass mice south of the Sahara Des- ert as geographic representatives of Linnae- us’ (1766) Mus barbarus, a form described from the Barbary Coast of northwest Afri- ca. The new trinomials that appeared in the early 1900s—b. manteufeli Matschie (1911) and b. nigeriae Thomas (1912)—observed Osgood’s delineation of specific bound- aries, and Thomas’ (1916) subsequent des- ignation of a type species and emended di- agnosis stabilized the contents and usage of Lemniscomys for African grass mice. Still, an inclusive species definition was not universally or immediately adopted. By the time of Allen’s (1939) systematic checklist of African mammals, three nom- inal species of the barbarus complex were thought to inhabit Subsaharan savannas: L. barbarus proper (including albolineatus, convictus, ifniensis, manteufeli, nigeriae, spekei, and zebra), L. dunni (including nu- balis Thomas & Hinton 1923, and olga Thomas & Hinton 1921), and L. oweni (in- cluding orientalis Hatt 1935). Ellerman (1941), as he did for so many sweeping for- mulations of species limits of African small mammals, adopted a broad definition of L. barbarus and so consolidated most of these regional epithets as geographic races (Fig. 1). As an exception, he retained, without explanation, L. olga as a species with a re- stricted distribution in Niger and left the fi- nal nomenclatural step to Rosevear, who in 642 1969 ranked olga as a subspecies of L. bar- barus. Ellerman’s (1941) influential classifica- tion of L. barbarus, amplified by Rosev- ear’s (1969) later reassignment of olga, is generally observed in recent African cata- logues and mammalian checklists (Misonne 1974, Honacki et al. 1982, Corbet & Hill 1991, Musser & Carleton 1993). The only other currently recognized species allocated to the barbarus group is L. hoogstraali, a member from southern Sudan recently de- scribed by Dieterlen (1991) and known only by the holotype (Fig. 1). Dieterlen, however, did not address the substantial variation apparent within L. barbarus, a widely distributed species with isolated populations in northwest Africa and with others occurring Over savannas south of the Sahara Desert, from western (Senegal) to eastern Africa (Kenya and Tanzania). The appreciable variation across this region was remarked upon by Lamine Cheniti & Sami (1989), who noted that their Tunisian spec- imens of L. barbarus more closely resem- bled Central African L. striatus in size than West African L. barbarus (b. nigeriae, b. olga, and b. owent). In this study, we ex- amine the level of differentiation between the northwestern and Subsaharan moieties, the former the geographic source (“‘Barbar- ia’) of Mus barbarus Linnaeus (1766), the type species of the genus, and the latter known by many regional epithets, the oldest of which is Mus zebra Heuglin (1864). Materials and Methods Approximately 650 specimens, repre- senting about 170 principal collecting lo- calities (Appendix 1), form the basis of this report. Specimens examined consist prin- cipally of skins with associated skulls con- tained in the museums, universities, or pri- vate collections listed below, each preceded by their institutional abbreviation. Except for Mus barbarus Linnaeus, which proba- bly does not exist, type specimens of all Species-group taxa we associate with the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lemniscomys barbarus complex (Table 4) have been examined and measured by Van der Straeten. AMNH American Museum of Natural History, New York City BEL Bellier Collection, presently housed in the Universitair Cen- trum, Antwerpen BMNH_ The Natural History Museum, London BZM Museum fiir Naturkunde der Humboldt-Universitat, Berlin CM Carnegie Museum of Natural History, Pittsburgh FMNH_ The Field Museum of Natural History, Chicago HAP D. C. D. Happold Collection of . Mammals, Canberra KBIN Koninklijk Belgisch Instituut voor Natuurwetenschappen, Brussel KMMA Koninklijk Museum voor Midden Afrika, Tervuren MAKB- Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn MCZ Museum of Comparative Zoolo- gy, Harvard University MNCN Museo Nacional de Ciencias Na- turales, Madrid MNHN Museum National d’ Histoire Na- turelle, Paris NHRS Naturhistoriska Riksmuseet, Stockholm RUCA — Universitair Centrum, Antwerpen SMF Forschungsinstitut und Natur- Museum Senckenberg, Frankfurt SMNS _ Staatliches Museum fiir Natur- kunde, Stuttgart USNM_ National Museum of Natural His- tory, Smithsonian Institution, Washington, D. C. ZMA Zoologisch Museum, Amsterdam ZMK Zoologisk Museum, Kgbenhavn Fourteen cranial and two dental dimen- sions, distilled from a larger suite of vari- ables previously defined by Van der Strae- ten & Van der Straeten-Harrie (1977), were recorded in millimeters (mm) to analyse VOLUME 110, NUMBER 4 643 Fig. 2. Six age-classes based on toothwear stages, as defined by Van der Straeten (1980a:Fig. 1) for speci- mens of Lemniscomys. univariate and multivariate variation within and between the populations sampled. To assess variation due to sexual dimorphism or post-weaning growth, gender was re- corded from skin tags and specimens were assigned to one of six age classes. The age classes (Fig. 2) correspond to progressive wear patterns of the upper first molar as recognized by Van der Straeten (1980a). Crania were viewed under a stereomicro- scope when measuring the craniodental variables to 0.01 mm using hand-held dig- ital calipers accurate to 0.03 mm. These 16 measurements, and their abbreviations as used herein, include (see Fig. 3 for land- marks): occipitonasal length (ONL); great- est zygomatic breadth (ZB); breadth of the braincase (BBC); breadth across the occip- ital condyles (BOC); least interorbital breadth (IOB); length of nasals (LN); breadth of the rostrum (BR); postpalatal length (PPL); length of the bony palate (LBP); length of the upper diastema (LD); length of the incisive foramen (LIF); breadth across the upper first molars (BMI1s); breadth of the zygomatic plate (BZP); length of the auditory bulla (LAB); alveolar length of the maxillary toothrow (ALM); and coronal width of the first upper molar (WM1). In addition, standard exter- nal dimensions and weight (in grams) were transcribed from skin tags as given by the 644 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Landmark points for 15 cranial measurements recorded in this study, as portrayed on dorsal, ventral, and lateral views of a schematic Lemniscomys skull (adapted from Van der Straeten & Van der Straeten-Harrie LO Tiere yal): VOLUME 110, NUMBER 4 collector: total length (TOTL), tail length (TL), hindfoot length (HFL), and ear length (EL); subtraction of TL from TOTL yielded head-and-body length (HBL). Several authors have commented on the relative scarcity of individuals of Lemnis- comys barbarus in the field ‘and the diffi- culty of obtaining them in adequate series (for example, Saint Girons & Petter 1965, Rosevear 1969, Happold 1987). We formed a similar impression based on the character of the museum samples we encountered, in which striped grass mice are usually rep- resented by only one to a few specimens from a collecting locality. For certain mul- tivariate analyses, specimens from localities Over a general region were necessarily grouped as eleven operational taxonomic units (OTUs). These OTUs, arranged from northwestern toward eastern Africa, and their sample sizes are: OTU (1).—n = 44, from Morocco, Agadir Province (USNM 475125-—475148, 475150—475169); (2).—n = 15, from Morocco, Khour- ibga and Rabat provinces (USNM 475121, 475122, 475170—475182); OTU OTU (3).—n = 9, from Senegal, all local- ities (USNM 376529-376536, 380463); OTU (4).—n = 24, from Ivory Coast and western Burkina Faso (FMNH 105180; USNM 465398—465400, 465413, 466674—466676, 466678-— 466693); (5).—n = 32, from Ghana and Togo, all localities (USNM 412746, 412747, 420570—420585, 435400- 435405, 438311—438314, 466694-— 466697); . (6).—n = 49, from eastern Burkina Faso, Benin, and westernmost Nige- ria (USNM 397647, 404085—404087, 422058—422064, 422066—422083, 439580—439586, 450852—450855, 465401, 465404—465412); (7).—n = 51, from Nigeria, Jos Pla- teau, Panyam and vicinity (USNM OTU OTU OTU 645 404020, 404039, 404050, 404064, 404073, 404084); (8).—n = 11, from Zaire, Faradje and Niangara (AMNH 49610-— 49612, 49614, 49615, 49623, 49624, 49627, 49630, 49632, 49633); (9).—n = 13, from Uganda, Rhino Camp (AMNH 180121; CM 2850; USNM 165191, 165193, 165194, 165198, 165200—165205, 1654272); (10).—n = 23, from Sudan, Equato- rial Province, various localities CPhMINE” 359313: “35314; 66853, 66854, 66857, 66859, 66860, 67171, 67172, 67175, 67293, 79500-79502; USNM 165192, 299764-299766, 299768—299771, 318003); (11).—n = 15, from southcentral Ke- nya, various localities (AMNH 114446-114448, 187678, 187679; CM 57977, 98258, 102462, 102463; USNM 162884, 181737-181740, 437394). Standard descriptive statistics (mean, range, standard deviation, coefficient of variation) were derived for the OTUs. One- and two-way analyses of variance, discrim- inant functions, and principal component 404022—404028, 404030-— 404044, 404046—404048, 404052—404056, 404058-— 404065-—404069, 404071- 404075—404078, 404080-— OTU OTU OTU OTU analyses were computed using the 16 cra- niodental variables, all of which were first transformed to natural logarithms. Principal components were extracted from the vari- ance-covariance matrix, and loadings are expressed as Pearson product-moment cor- relation coefficients of the components with the original cranial variables. Means and ranges of external variables are provided as a guidance in identification (see Appendix 2) but were not considered in multivariate analyses. Except for the a posteriori input of type specimens (all measured by Van der Straeten) to a discriminant function analysis (OTU series measured by Carleton), nu- merical results presented in the various ta- 646 bles and figures are not based on intermix- ture of craniodental data collected individ- ually by the authors. All analytic procedu- res were conducted using Systat (Version 6.0, 1996), a series of statistical routines programmed for microcomputers. Results Morphometric analyses.—Patterns of variation contingent upon age and gender cohorts within local samples of Lemnisco- mys generally conform to those presented for other species of African muroids (Van der Straeten & Verheyen 1978, Carleton & Robbins 1985, Carleton & Martinez 1991). That is, significant age-related size variation is apparent for certain variables, notably those measured at the cranial extremes (ONL, ZB) or on the rostrum (LN, BR, LIE LD); whereas, f values of age effects de- rived from ANOVAs are smaller and typi- cally insignificant for dimensions taken across the braincase (BBC, IOB, BOC) and on the molars (ALM, WM1). Sexual di- morphism in size or age-sex interactions contribute little to mensural variation within OTUs (Table 1). Critical estimation of such influences upon sample variation is sensi- tive to sample size and particularly to the balance of age and sex representation among the specimens available for mea- surement (Voss & Marcus 1992). Our in- ability to achieve these sampling ideals with the real museum series at hand pre- sumably accounts for certain inconsisten- cies and differences in probability levels at- tained for our two largest OTUs (Table 1). Notably, the lack of significant mean size differences among age cohorts for external skin variables (TOTL, TAIL, HFL) and the occurrence of same for the alveolar molar length of the Moroccan sample (OTU 1) seem counterintuitive and at odds with pre- vious studies. For the two largest, geographically ho- mogeneous samples—Morocco (OTU 1) and Nigeria (OTU 7)—nearly all cranial vari- ables (except WM1) contribute substantial- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—f-values from two-way ANOVAs (sex and age cohort) for 3 external and 16 cranial dimen- sions in two large OTUs of the Lemniscomys barbarus group. OTU 1 (25 M, 19 F) OTU 6 (26 M, 23 F) Inter- Inter- Age Sex action Age Sex action TOME (2:6 0.0 0.6 0.1 0.0 ed TAIL Deo 0.1 0.2 il 0.8 1.3 HFL 0.6 3 02 0.1 0.1 0.4 ONL Sr On! eZ 9 Ae = OM |i ZB S252F oO. cull 2.4 0.4 px BBC 4.6* S207) (030 ks ce? 0:2 IOB ey OL. 07 el 0.5 0.1 LN 1.2 0.1 0.8 6.4** 0.6 4.1* BR Oe ee ae OF, 4.5* OZ EROS PPL ht) 0.5703 Oa ee es ZA LBP 4.5* Oxliyes te2 0.2 0.2 Gs LIF 0.9 0.4 1.8 3.3 O22, Hes LD 4.3% OF 9 = 1O.24*2) 20M L.3 BMis” 7 35:0* OZ. 351 8.98 A oe BZP Dy) 0.1 0.4 9:3**5 O10 1.0 LAB ez Ox 10:2 2D 0.2 1.4 BOC 0.5 O13" 1:3 Lo 0.2 1.0 ALM 0.7 0.7 0.8 8.7 tFs Sew 0.4 WM1 Jes) 2s tal 0.7 itat 0.1 * = P < 0.05; ** = P < 0.01; *** = P S 0.001. ly and positively to variation (74.1%) sum- marized by the first principal component (Table 2). Their covariation accounts for the uniformly higher scores and right-oriented dispersion of the Moroccan specimens rel- ative to those from Nigeria (Fig. 4A). Pear- son correlation coefficients of the original variables and the second principal compo- nent are consistently smaller and occasion- ally negative; no biologically meaningful segregation of specimen scores is apparent among the minor variation (6.0%) repre- sented by this axis. Age-related size in- crease may account for some dispersion along the first component within each geo- graphic sample (Fig. 4B); however, for the comparison of these two samples, age-class influence is statistically negligible (f = 1.99, P = 0.09) relative to the pronounced locality effect (f = 121.1, P = 0.001) upon PC I scores. Little overlap of individual scores be- tween the Moroccan and Subsaharan sam- VOLUME 110, NUMBER 4 Table 2.—Results of principal component analysis performed on OTU 1 (Morocco, n = 43) and OTU 7 (Nigeria, n = 51), the two largest samples available for the Lemniscomys barbarus complex (see Fig. 4). Variable PC I PC II ONL 0.99 =0:02 ZB 0.94 0.09 BBC 0.81 0.05 IOB 0.75 0.29 LR OT 0.13 BR 0.83 0.10 PPL 0.94 —0.08 LBP 0.80 0.47 LD 0:97 =O: LIF 0.86 —0.47 BMIs 0.85 0.06 BZP 0.87 Ong LAB 0.74 0.15 BOC 0.74 =0102 ALM 0.67 0.05 WMI US) 0.02 Eigenvalue 0.049 0.004 % Variance 74.1 6.0 ples is observed in plots of the first two canonical variates derived from discrimi- nant function analysis of the 11 defined OTUs (Fig. 5A, B). Again, the uniformly larger skull dimensions of the Moroccan se- ries are emphasized by the high, positive loadings of nearly all cranial variables on the first canonical variate (Table 3); the width of the first molar as in the principal component analysis, is least influential in affecting the segregation of group means along the first canonical axis. Unlike the clearcut separation of the two Moroccan samples, those drawn from West (OTUs 3- 7) and East (OTUs 8-11) Africa display ap- preciable intermixture of individual speci- men scores projected upon the first two ca- nonical variates (Fig. 5A). Cluster diagrams generated either from Mahalanobis distances among group cen- troids (Fig. 6A) or from log-transformed OTU means (Fig. 6B) similarly underscore the strong phenetic separation of the Mo- roccan samples from those found south of the Sahara Desert. Among the latter, the clustering patterns suggest some association 647 according to geographic propinquity, but certain OTUs (Senegal and Kenya) discor- dantly merge with far distant samples. Posthoc numerical classification of ten type specimens with suitably intact crania was effected using the discriminant func- tion statistics computed from analysis of the 11 OTUs. Of the nine types representing taxa described from Subsaharan Africa, none was “correctly” classified with the OTU closest to their geographic origin; for example, albolineatus and convictus, both named from Kenya, were predicted to have membership in Ivory Coast-West Burkina Faso (OTU 4) and East Burkina Faso-Benin (OTU 6), respectively, and manteufeli, named from Tanzania, was allocated to Su- dan (OTU 10). Nevertheless, all type spec- imens were sensibly associated within the range of multivariate dispersion exhibited by specimens from the general region of their collection: namely, Morocco (ifnien- sis) or the Subsahara (the remaining nine) (Fig. 7). In summary, ordination analyses, wheth- er extracting principal components from original measurement data or canonical var- iates from the centroids of 11 predefined geographic samples, reveal the more robust crania typical of the Moroccan samples as compared to striped grass mice from East and West Africa (Fig. 8). In contrast to our experience with other morphologically sim- ilar, congeneric species, features of shape or proportion are not evidenced by the disper- sion of scores along the secondary axis of ordination; that is, the cranium of Moroccan barbarus seems to be an isometrically over- sized version of that characterizing popu- lations broadly distributed across Subsahar- an savannas. Although not used in multi- variate computations, parameters of bodily dimension and mass also convey the excep- tional size of the Moroccan populations (Fig. 9, Appendix 2). Pelage comparisons.—The upperparts of members of the L. barbarus complex are marked by longitudinally continuous, alter- nating dark and light stripes, a distinctive 648 PC II (6.0 %) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON @ Morocco (OTU 1 0.0 0.1 0.2 0.3 0.4 0.5 PC | (74.1 %) 0.2 ox 0.1 oS Sc c Fe -0.0 O ou -0.1 -0.4 -0.3 -0.2 -0.1 @ Morocco (OTU 1 0.0 0.1 0.2 0.3 0.4 0.5 PC | (74.1 %) Fig. 4. Results of principal component analysis performed on 16 log-transformed craniodental variables measured on 94 specimens representing Moroccan (OTU 1, n = 43) and Nigerian (OTU 7, n = 51) samples of the Lemniscomys barbarus complex: A, projection of individual specimen scores on the first two principal components extracted (see Table 2); B, projection of centroids, as calculated a posteriorly, representing six age classes within each OTU on same principal components (age classes V and VI not represented in the Nigerian sample). pattern which readily contrasts to the single mid-dorsal line of the L. griselda group (single-striped grass mice) or to the variably interrupted or punctulated lines of the L. striatus group (spotted grass mice). Atten- tion is here given to qualitative description of chromatic and pattern variation of the upperparts because such traits have histor- ically received prominence in the diagnoses of taxa, whether named as species or sub- species. General characterization of the pelage in the L. barbarus group follows (Fig. 9). The single vertebral line, which extends from the middle crown to the rump, typically possesses a deeper hue than that of the lat- VOLUME 110, NUMBER 4 649 A @ East Africa A West Africa @ Morocco CV 2 (13.0 %) oO 2 3 -4 6 5 -4 -3 9 4 0 1 2 3 4 5 6 CV 1 (60.7 %) B , @ East Africa A West Africa @ Morocco 3 2 CV 2 (13.0 %) oO -6 -5 -4 -3 -2 -1 0 1 2 3 4 5 6 CV 1 (60.7 %) Fig. 5. Results of discriminant function analysis performed on 16 log-transformed craniodental variables measured on 235 intact specimens representing 11 OTUs of the Lemniscomys barbarus complex (see Table 3): A, Projection of individual specimen scores on first two canonical variates extracted; B, Projection of group centroids on first two canonical variates extracted. 650 Table 3.—Results of discriminant function analysis performed on intact specimens (n = 235) representing 11 OTUs of Lemniscomys (see Fig. 5). Variable CV I CV Il f(OTU) ONL 0.94 0.14 49.0*** ZB 0.94 =O108)) 255.2572 BBC O75 O04 Disees IOB 0.70 0.04 19 Ds LR 0.87 =(0MORY See BR 0.76 —0,06, 21. a5*5 PPE 0.84 O28 33/655 LBP 0.75 0.16 iss Oe s3 LD 0.91 O23 4507 525 |O108 0.79 OF 9ST 5290 Fe BMIs 0.82 —(l DORE BZP 0.70 0.14 14 924s LAB 0.70 —0.13 lowae* BOC 0.65 0.23 13) alasee ss ALM 0.70 —0.09 | SyAlessess WMI1 0.45 —0.49 Or Canonical correlation 0.87 0.64 Eigenvalue 3.25 0.69 % Nariance 60.7 13.0 * = P < 0.05; *** = P < 0.001. eral dark stripes, appearing black as com- pared to dark gray or brown. To either side of the mid-dorsal line occur four to six pri- mary pairs of alternating light and dark stripes (as observed over the lumbar-rump region), whose definition more or less abruptly degrades toward the sides; the dark lines are usually wider than the intervening light ones. The venter is much lighter and unmarked, ranging from dull to bright white, and a clear buff lateral line may de- marcate the dorsal-ventral pelage transition. Fine tawny to rufescent hairs densely clothe the pinnae and accent their contrast to the crown and dorsal pelage. Tops of the fore and hindfeet are covered with white to pale buff hairs. The tail is indistinctly bicolored, the dorsal hairs black and those underneath ochraceous to rufous, resembling the color of the pinnae. Within and among locality samples of the L. barbarus group, one can discern several thematic variations around the chromatic groundplan described above. The distinc- tiveness of the mid-dorsal line varies with PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON respect to the dark lateral stripes, a conse- quence of its color (brown to dusky to black) and width. Thus, the mid-dorsal stri- pe may appear densely black and relatively wide (Figs. 9A, C) or brown and notably thin (Fig. 9B). The boldness of the striping pattern is modulated by the color of the dark bands (medium brown to black) in re- lation to the purity of the pale bands (clear white to warm buff or dull ochraceous) and by the degree of ochraceous suffusion over the middle dorsum. These chromatic subtle- ties interact to heighten or lessen the sharp- ness of the linear contrast (compare Figs. 9A to 9C, & D). The fine aural hairs range from nearly cinnamon, setting the pinnae conspicuously apart from the general dor- sum (Fig. 9D), to dull ochraceous, lessening the visual distinction of the pinnae (Fig. 9A). An important pelage variation involves the number of dorsal stripes, counts of which were mentioned regularly by the ear- ly describers. For example, Thomas (1903: 297) noted that dunni possesses “‘on each side, five uninterrupted buffy stripes, sepa- rated from each other by broad brown bands, each of which is divided down the centre into two by an uninterrupted band of light.”” Osgood (1910:11) characterized convictus as having: “Light stripes pale ochraceous buff and numbering 5—6 of the heavier or primary ones and 4—5 of the sec- ondary ones on each side;’’ and ‘‘Four def- inite primary dark stripes on each side, each divided by a secondary light stripe.’ And Hatt (1935:2) cited the “‘six pairs of lateral dark stripes . . . distinctly split by secondary light stripes’”’ in describing orientalis. The number of stripes is an inexact impression largely mediated by the development of these secondary light lines within the first, and sometimes the second, pair of primary dark lateral stripes (as numbered away from the unpaired vertebral stripe). Individuals with strongly penetrant secondary light stripes appear to be finely streaked with more and narrower light and dark stripes, especially over the middle dorsum (Figs. VOLUME 110, NUMBER 4 A Northwest Africa East & West Africa 50 40 30 20 10 Mahalanobis D’ B Northwest Africa East & West Africa 0.12 0.10 0.08 0.06 0.04 0.02 Euclidean D Morocco (1) Morocco (2) Senegal (3) Zaire (8) Uganda (9) Sudan (10) Ivory Coast-W Burkina (4) Ghana-Togo (5) E Burkina-Benin (6) Nigeria (7) Kenya (11) Morocco (1) Morocco (2) Senegal (3) Zaire (7) Uganda (9) Sudan (10) Ivory Coast-W Burkina (4) Ghana-Togo (5) E Burkina-Benin (6) Kenya (11) Nigeria (7) 651 Fig. 6. Clustering (UPGMA) of 11 geographic samples representing the Lemniscomys barbarus complex, as derived from: A, Mahalanobis’ distances (D?) among the OTU centroids (see Fig. 4); B, Euclidean distances between OTUs based on means of the 16 log-transformed craniodental variables. 9B, D); whereas, those that have obscurely posefully selected for its extreme expres- defined secondary lines seem to have fewer sion of this consolidated pattern._). and broader dark stripes on their upperparts The pelage characteristics reviewed (Figs. 9A, C—the latter specimen was pur- above undoubtedly seemed easily definable 652 @ East Africa 4 3 2 1 orientalis WA oS ° zebra , albolineatus -1 + -2 -3 -4 6 5 -4 -3 -2 “I Figs 7 A West Africa PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON @ Morocco ,/anteufeli con victus, spekei , CV 1 Results of discriminant function analysis presented in Fig. 5, with a posteriori classification of type specimens representing ten species-group taxa of the Lemniscomys barbarus complex. Polygons enclose maximal dispersion of individual scores around group centroids for each geographic region. and unique as exemplified by the holotypes and small hypodigms for the various epi- thets created in the late 1800s and early 1900s. However, as more specimens, larger locality series, and broader geographic rep- resentation have become available, the in- dividuality of these color morphs is ob- served to break down and merge impercep- tibly. By 1969, Rosevear concluded that even subspecific separation of West African nigeriae from oweni, originally assigned to Separate species on the basis of having white versus buffy light stripes as purported by Thomas (1911b, 1912), was indefensi- ble. We agree with Rosevear’s assessment and further note that the occurrence of bold- ly marked striping versus more muted dor- sal patterns—such as advanced to discrimi- nate albolineatus versus convictus (Osgood 1910) or nubalis versus dunni (Thomas 1903, Thomas & Hinton 1923)—can be du- plicated by extreme individual patterns just within USNM Nigerian series. Hatt (1935) identified the absence of facial stripes as a useful feature for sorting specimens of nu- balis from those of orientalis and zebra. Nevertheless, we have found the continua- tion of trunk markings (the fourth lateral pair of light and dark stripes) onto the cheeks as highly variable within Subsahar- an locality samples and doubt the trait’s utility as a diagnostic tool at the level in- tended by Hatt (and see Discussion below). The hispid or spiny quality of the dorsal fur has been also mentioned in some descrip- tions of striped mice, but such textural dif- ferences are difficult to objectively convey and practically apply. And so forth for many of the pelage traits given as diagnos- tic criteria. Certain geographic trends in pelage color and pattern do emerge with the better series VOLUME 110, NUMBER 4 653 Big. 8: Dorsal, ventral, and lateral views of crania and mandibles of young adult Lemniscomys (both Age Class 2): left, L. zebra (USNM 465402), a female from Burkina Faso, 6 mi SE Seguenega; and right, L. barbarus (USNM 475145), a male from from Morocco, 15 km E Agadir. now at hand, but these are presented only as average differences or general impres- sions. Higher numbers of dorsal stripes ap- pear more common within Subsaharan pop- ulations, which usually exhibit five, occa- sionally six, pairs of primary dark and light bands on either side of the mediodorsal line; in contrast, specimens from Northwest Africa possess four or five countable pairs of primary dark and light stripes. The im- pression of more dorsal stripes among Sub- saharan series is visually accentuated by the development of secondary light lines, which usually subdivide the first as well as the second primary dark stripes (Figs. 9B, D); within samples from northwestern Af- rica, secondary light lines are typically ev- ident only within the first primary dark stripes, obscurely defined in the second and those more laterad (Fig. 9A). A pattern con- sisting of a broad, black vertebral line, bor- dered by more consolidated dark stripes and irregularly defined secondary light stripes (Fig. 9C), occurs more often among popu- lation samples from East Africa (the source of taxa such as zebra, spekei, and nubalis); 654 it 4 | Be | Ay . j i 4 { i t PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 9. Variation in dorsal pelage pattern of four specimens of striped grass mice, the Lemniscomys barbarus species group. A. L. barbarus (USNM 475121, LHB = 108 mm), a male from Morocco, 17 km SW Rabat. B. L. zebra (BMNH 23.5.12.81, LHB = 86 mm), a female from Niger, Farak (region of olga Thomas & Hinton 1921). C. L. zebra (USNM 165191, LHB = 89 mm), a female from Uganda, Rhino Camp (region of nominate zebra Heuglin 1864). D. L. zebra (USNM 376535, LHB = 88 mm), a male from. Senegal, Kotiare Naoudé (region of oweni Thomas 1911). a finely streaked dorsum (vertebral stripe narrower and secondary light stripes well delineated within the first and second pri- mary dark bands) is typical of West African populations (taxa such as nigeriae and ow- eni—Fig. 9D). With regard to the degree of pelage sat- uration, we observe a subtle trend toward paler, diluted upperparts in examples from more xeric vegetation zones. This associa- tion is best evidenced by the many USNM examples from West Africa, wherein a darker pelage tone characterizes those spec- imens obtained from localities within Guin- ea savanna compared to the slightly paler aspect of series collected within Sudan sa- vanna. At the darker extreme, somewhat muted colors and subdued contrast mark the dorsum of Moroccan populations (Fig. 9A) that occupy the relatively mesic Mediter- ranean region of Northwest Africa; at the pallid end of this spectrum are the few VOLUME 110, NUMBER 4 specimens (Fig. 9B), including the holotype of olga, that originate from the Sahel zone in West Africa. As a generality, specimens of the L. bar- barus group possess moderately hispid dor- sal fur, especially over the rump, compared to those of the L. striatus complex. Within the L. barbarus group, most taxa, as rep- resented by their type specimens, generally conform in this trait with the exception of the more hispid condition characteristic of olga. Discussion Taxonomic conclusions.—The major pat- tern of craniodental variation uncovered here provides evidence for the specific di- vergence of the Moroccan samples from those representing populations that occur to the south of the Sahara Desert. The former correspond to Lemniscomys barbarus s.s. (Linnaeus 1766), and the oldest name ap- plicable to the latter is L. zebra (Heuglin 1864), whose type locality lies in southern Sudan. Seven of our 11 OTUs were regret- tably coarsely defined in geographic cov- erage, an analytical approach dictated by the generally small sample sizes available from the majority of collecting localities. Notwithstanding the composite nature of most OTUs, we believe that the regions thus broadly sampled are sufficient to sus- tain the principal taxonomic conclusion rec- ommended here—the recognition of north- west African populations (L. barbarus) as a species distinct from those occupying sa- vanna biomes in West and East Africa (L. zebra). Infraspecific taxonomic uncertainties per- sist, however, within the Subsaharan moeity we identify as L. zebra, as intimated by the appreciable size and chromatic variation observed among those samples. The earlier conclusions of Robert T. Hatt of the Amer- ican Museum illustrate the biological prob- lems inherent in understanding that varia- tion. In his reports (Hatt 1935, 1940) on the rodents collected by the AMNH Congo Ex- 655 pedition (1909-1915), Hatt provided diag- nostic criteria for three “‘natural groups” of the L. barbarus group that originated from a single locality, Faradje. Hatt identified the three as separate species—L. zebra, L. dun- ni nubalis, and L. oweni orientalis, the last newly described as a “‘dwarf’’ subspecies of West African oweni. For us, unambigu- ous, repeatable separation of East African specimens into discrete species using Hatt’s identification criteria remains elusive. The mensural diagnostic values (means?) listed by Hatt for each “‘species’’—for instance, foot length, 25 versus 25 versus 23 mm; greatest skull length, 27.5 versus 27.2 ver- sus 26.7 mm; length upper molar row, 4.8 versus 5.1 versus 4.6 mm—span the normal variational spectrum that we find within population and regional samples (Appendix 2). Unfortunately, Hatt provided no mea- sures of sample dispersion nor gave any at- tention to potential age effects for the nu- merical contrasts offered, statistical precau- tions now standardly addressed when eval- uating intersample mean differences. The pelage attributes given (general color, spin- iness, and facial stripes) are subjective to apply precisely and suggest no variation be- tween individuals from large series, which contradicts our experience. Within the Far- adje material, it is notable that Hatt allo- cated the two “pallid, very hispid”’ individ- uals to L. dunni nubalis (named from cen- tral Sudan) and the remainder of the “‘dark, hispid’”’ specimens, totaling 17, to either L. zebra or L. oweni orientalis, the latter two separable by average size. Darker versus paler pelage hues and hispid versus very hispid fur are traits whose expression ap- pears subject to slight variation within lo- cality series and to clinal trends over broad regions, for example, Guinea through Sahel savannas in West Africa. In view of the geographic location of Faradje near the southern limit of Guinea savanna in eastcentral Africa, one might anticipate that the majority of specimens would appear rel- atively dark and moderately hispid. In summary, we interpret the differences 656 tabulated by Hatt (1935, 1940) as examples of intraspecific variation of the one species, L. zebra, not sympatry of three members of the L. barbarus group. In the same papers, Hatt (1935:1—2) previewed this possibility: ““That a more conservative person or one with a greater series of specimens than have been available to me might lump all three as fortuitous variations of a single subspecies, I readily admit, but such a procedure would, . . . with the present evidence, but mask the simple assortment of characters as they have presented themselves and tend to smother rath- er than encourage further investigation of the prob- lem of speciation or, it may be ecologic differenti- ation, in this region.” His was a reasoned and reasonable taxo- nomic viewpoint, given the state of knowl- edge about African striped mice in the 1930s. From our vantage point of the late 20th century, and hoping to “encourage further investigation,’ we draw attention to three alpha-systematic problems not yet ad- equately resolved by the Subsaharan mate- rial at out disposal. For one, the nature of morphological dif- ferentiation between East (zebra) and West African (oweni) populations should be fur- ther appraised drawing upon other kinds of data and improved geographic sampling (Chad and the Central African Republic are apparently unrepresented in museum col- lections). For example, another sibling spe- cies pair of Lemniscomys, L. macculus and L. bellieri of the striatus group, is distrib- uted as East and West African counterparts (Van der Straeten 1975, Van der Straeten & Verheyen 1979, 1980). Some geographic integrity of East versus West African ex- amples of striped grass mice is suggested by the phenetic clusters generated from cra- niodental similarity, but the segregation is not complete: one West African sample (Senegal, OTU 3) groups among East Af- _ rican OTUs, and another from East Africa (Kenya, OTU 11) aligns with those from West Africa (Fig. 6). In light of such am- biguous results, set against the inadequacy of geographic coverage and the apparent want of other diagnostic traits, we prefer at PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON this investigatory stage to retain all Subsa- haran populations, except L. hoogstraali (see below), under the one species L. zebra (Heuglin 1864) without formal delineation of geographic races. For another, the taxonomic status of Tho- mas & Hinton’s (1921) olga merits reex- amination. The form, described from a few Specimens and fewer localities in Sahel Sa- vanna in southcentral Niger, is character- ized by small size, pale color, finely delin- eated striping pattern (Fig. 9B), and slightly more hispid pelage texture. The three spec- imens from Timbuktu, Mali, another local- ity in the Sahel, also recall this morphology. In view of the type specimen’s morpho- metric association with other West African populations (Fig. 7), and while acknowl- - edging the distinctiveness of the form’s pel- age and habitat, retention of olga as a junior synonym of L. zebra seems prudent as a provisional arrangement. Careful geograph- ic transects across Sudan and Sahel wood- lands, both in West and East Africa, are re- quired to illuminate the nature of morpho- logical variation in characters attributed to olga and critically judge its taxonomic sta- tus. The third species of the Lemniscomys barbarus complex herein recognized, L. hoogstraali, was described by Dieterlen (1991) from the Upper Nile Province of southern Sudan (Figs. 1, 12). As noted by Dieterlen, the form contrasts with neigh- boring Sudanese examples of L. barbarus (=L. zebra as here understood) by its larger external and cranial size and by the dorsal striping pattern (see additional commentary under Taxonomic Accounts). The dark stripes of the upperparts are exceptionally black and broad, which, together with the purity of the primary white stripes and in- distinct secondary white stripes, heightens the alternating, zebra-like effect of the ho- lotype (and only known specimen). Such an appearance is suggested, as an extreme of pelage variation, by some individuals from East Africa (see Fig. 9C), but their cranial measurements otherwise fit those recorded VOLUME 110, NUMBER 4 657 Table 4.—Selected cranial dimensions (as measured by Van der Straeten) of thirteen type specimens of species-group taxa assigned to the Lemniscomys barbarus group. All taxa listed are viewed as junior synonyms of L. zebra, except ifniensis, allocated to L. barbarus (Linnaeus 1766), and hoogstraali, retained as a third species of the group. Variables Taxon & authorship Age type) class ONL ZB BBC IOB LN LIF LD BMis BZP ALM zebra Heuglin, 1864 (SMNS 1100a/5422) fe ca 128 PET aS EOL SO" 6.0" SS 36) S06 spekei De Winton, 1897 (BMNH 63.7.7.23) 3 Pee - 1253, tA se ah lode, SiS 69) i624 53105 weds dunni Thomas, 1903 (BMNH 3.2.8.15) 5 27.8 — (EA 8 er. SS ee | |: GR 7 MM TO Sr I i SR tag 5 albolineatus Osgood, 1910 (FMNH 17194) 3 Pees A25 AS Oa Sc7) 6S VS Biggs convictus Osgood, 1910 (FMNH 17206) Pa deo oes: 12, ) 45.) PbS 5.9% 67 | 5.9). S38 25.35 oweni Thomas, 1911 (BMNH 11.6.10.61) Peete bey eA 4 IS OS GS Se OS manteufeli Matschie, 1911 (BZM 16025) 1 260 TSO: 91 -OeseArS 9.1 32 SS 32 PIB SNS nigeriae Thomas, 1912 (BMNH 12.1.16.45) 5 29.1 13.4 bas? a9 lls Se aeeeO . GOeh ST tess olga Thomas & Hinton, 1921 (BMNH 21.2.11.128) 2, ZN ro. Wide aes AS Sb i SL SN Se nubalis Thomas & Hinton, 1923 (BMNH 18.7.2.14) 4 29.2 — 12 PAS. PRS 6.0" GS 6. OO S20 orientalis Hatt, 1935 (AMNH 49626) 5 268 20125 1d:4.. 4:1 1Osthay iS: |. Bsiw va «325 eD ifniensis Morales A., 1935 (MNCN 9638) 6 See eS) Oe ae eS Oke PA BS eee SSS hoogstraali Dieterlen, 1991 (USNM 342078) 5-6 32.0 — CAS PSO 2S Ser as Ga Aes. for eastern series of L. zebra and are clearly smaller than the holotype of L. hoogstraali (Table 4). In dimensions of the skull and molars alone, the type of L. hoogstraali in- stead agrees with those of L. barbarus proper from Morocco (Table 4, Appendix 2). Although based on a single specimen, the combination of large size and dramati- cally striped dorsum, considered with the form’s geographic occurrence approximate to typical examples of L. zebra, leads us to accept Dieterlen’s (1991) judgement of hoogstraali as a species. Additional sam- ples should be sought in order to enhance appreciation of its variation, phyletic rela- tionships, and level of differentiation from L. barbarus and L. zebra. With revision of the striped grass mice, there now exists a basic framework of spe- cies definitions and their distributional lim- its within Lemniscomys, a moderately di- verse genus embracing at least 11 species associated in three groups: namely, barba- rus (barbarus, hoogstraali, and zebra), gri- selda (griselda, linulus, rosalia, and rosev- eari), and striatus (bellieri, macculus, mid- dendorfi, and striatus). Future systematic attention should focus on phylogenetic re- lationships among the species and the evi- dence for accepting these nominal group- ings as monophyletic. While visually ob- vious and thus advantageous for employ- ment in taxonomic keys, the dorsal pelage patterns that have traditionally afforded def- inition of the species groups of Lemnisco- mys may not necessarily convey phyletic affinity. Although readily distinctive as rep- resented by the typical species of each 658 group, these basic striping configurations are not necessarily so well defined among _ all of a group’s members. Some individuals of West African L. linulus, for instance, ex- hibit indistinct lateral spotting in addition to the pronounced middorsal stripe, a pattern remniscent of the upperparts of some L. bellieri. For another example, the Nigerian form fasciatus, now synonymized under L. striatus (see Van der Straeten & Verheyen 1980), was long confused with L. barbarus (now = L. zebra) as a result of the nearly continuous union of its lateral spots. In contrast to their external dissimilarity, one is impressed by the fundamental cra- niodental resemblance among the species of Lemniscomys. Certain morphological traits may further characterize these groups, at least as exemplified by their typical species. Examples of L. barbarus possess well in- flated ectotympanic bullae relative to those of L. griselda (intermediate) and L. striatus (smallest); the degree of inflation may cor- respond to the openness of the habitats oc- cupied by each species. The molars of L. griselda are broad and robust compared to those of L. barbarus and especially L. stria- tus. Along with its heavier, more rounded cusps, the t3 of M1 in L. griselda is weakly defined and the t9 appears as a short spur off t8; the t3 of M2 is minute and irregu- larly formed. The cheekteeth of L. striatus seem more cuspidate, with the t3 of Ml separated from t2 by a distinct cleft and the t9 formed as a small albeit discrete cusp apart from t8; the t3 of M2 is small but consistently formed. In general, the molars of L. barbarus more closely resemble those of L. griselda. Such observations should be extended to the other species of Lemnisco- mys and additional morphological charac- ters sought in order to test the monophyly of the currently defined species groups. Investigation of chromosomal homolo- gies promises to shed light on kinship groups within Lemniscomys, as suggested by the karyotypic differences so far report- ed. Based on a single specimen from Al- geria, Filippucci et al. (1987) reported the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON diploid number (2N) of L. barbarus as 54 and the number of autosomal arms (AN) as 58, dominated by a graded series of 23 small to large acrocentric pairs, along with two pairs of small metacentrics and one large submetacentric pair. This diploid count agrees with those reported for L. ze- bra from Burkina Faso (Gautun et al. 1986, as L. barbarus), Ivory Coast (Matthey 1954, also as L. barbarus), and from Came- roun (Van der Straeten, pers. com.); the Cameroun animals even display the same fundamental number (AN = 58), although having three pairs of small metacentrics and apparently lacking the large submetacentric pair. These chromosomal traits appreciably differ from karyotypes that Van der Strae- ten & WVerheyen (1978) documented for Ivory Coast samples of L. bellieri QN = 56) and of L. striatus (2N = 44). Such pre- liminary results should be reconfirmed and extended to include other species, and their differences and mechanism of karyotypic change explored with the banding tech- niques now available. Distributions and zoogeographic impli- cations.—Museum specimens of L. barba- rus and L. zebra have originated from hab- itats that can be loosely categorized as sa- vanna, woodland, or scrubland. All such places, in general, have ample grass and-or herbaceous ground cover intermingled with bushes to small trees, but never close-can- opied, high forest. Nonetheless, the vege- tational character where each species occurs can be broadly contrasted. On skin tags and in field catalogs, col- lectors in West Africa have recorded the vegetation of almost all trapping sites of L. zebra as either Guinea or Sudan savanna (or woodland), following Rosevear’s (1953, 1965) terminology. To date, only a handful of specimens come from the Sahel, but few collections have been made in this very arid zone. These vegetational associations differ in their dominant plant species, physiog- nomy of the low-canopied woods, and den- sity of ground cover, features which in turn correspond to amount of annual rainfall, VOLUME 110, NUMBER 4 length of dry season, and prevailing lowest relative humidity (Guinea savanna, more mesic, to Sahel savanna, more arid). To the east, areas where striped grass mice live have been characterized as grass and scrub (Uganda—Delany 1975), drier savannas and steppes (East Africa—Kingdom 1974), or simply as savanna (annotations on USNM skin tags of Sudanese specimens collected by H. W. Setzer). Our localities from Sudan, Zaire, and Uganda also lie within vegetation zones mapped by Rosev- ear (1953, 1965) as Guinea, Sudan, or Sahel savanna. At Zizonkame, about 8° north latitude in Benin (Fig. 10, Top), degraded Guinea sa- vanna contained prolific grasses three to four feet high, periodically burned, as well as some cultivated plots (cassava) and oil palms (field notes of J. W. LeDuc and C. B. Robbins, USNM Mammal Division ar- chives). Here, individuals of L. zebra were obtained with a variety of small mammals, including: Atelerix albiventris, Galago se- negalensis, Xerus erythropus, Thryonomys swinderianus, Tatera sp., Taterillus graci- lis, Cricetomys gambianus, Arvicanthis nil- oticus, Dasymys rufulus, Lemniscomys striatus, Mastomys natalensis, Myomys dal- toni, Mus musculoides, and Uranomys rud- di. In the vicinity of Cella, about 11°30’ north latitude in Burkina Faso (Fig. 10, Bottom), examples of L. zebra were trapped amongst dry grasses that formed a uniform ground cover, interspersed with shrubs and low-canopied trees, which together com- posed a plant community typical of Sudan savanna. The less diverse set of small mam- mals collected at this site included: Atelerix albiventris, Galago senegalensis, Tatera sp., Taterillus gracilis, Steatomys pratensis, Arvicanthis niloticus, and Mastomys natal- ensis. Populations of L. zebra and L. stria- tus were found to co-occur at the locality within Guinea savanna, but only examples of L. zebra were collected in the drier Su- dan savanna. Populations of L. barbarus also inhabit various nonforested, relatively dry habitats, 659 but along the coastal region and contiguous low plateaus of Northwest Africa, where the severity of drought is moderated by the Atlantic Ocean and Mediterranean Sea. In Morocco, notations on specimen tags iden- tify collecting places as “‘rocky outcrop in vegetated coastal dunes,’’ “‘coastal scrub,” ‘sandy hummocks with dense vegetation,” “juniper scrub,” ‘“‘argon-sage-grassland,”’ and “argon savanna.’ Such Moroccan lo- cales denote a spectrum of climatic domains classified as arid, semiarid, or even humid (see fig. 6 in Saint Girons & Petter 1965: 18). At two places along the valley of the Souss River in Agadir Province, Morocco (about 30°30’ north latitude), specimens of L. barbarus were obtained with those of Atelerix algirus, Gerbillus sp., Gerbillus hoogstraali, Meriones shawi, Mastomys er- ythroleucus, and Mus spretus. The collector, R. E. Vaden, described the site at 16 km W Aoulouz (Fig. 11, Top) as “‘more coastal type of vegetation .. . predominantly argon trees and grass ... clumps of assorted shrubs, from sage through thorn-bushes (acacia?) up to small argons, maximum of about 6 feet in height ... in between [shrubby clumps], it’s grassy.” The locality at 5 km S Taroudannt (Fig. 11, Bottom), he characterized as “‘heavily-vegetated sandy dune area .. . dominant shrubs are the same thorny bushes seen all along . . . some sage, but not like the last camp [16 km W Aoulouz]—telatively smaller ... plenty of grass both under the shrubs and in the open areas” (R. E. Vaden field notes, USNM Mammal Division). At the latter site, sam- ples of L. barbarus were captured in the grassy areas. The regional distributions of L. zebra and L. barbarus do concord with previously recognized biogeographic subregions of Af- rica, as distilled from both plant and animal studies (Chapin 1932; Moreau 1952, 1966; Rosevear 1953, 1965; Davis 1962; Brenan 1978; Quézal 1978; Rautenbach 1978; Crowe & Crowe 1982; see Skinner & Smithers 1990:xxii, for their helpful map 660 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 10. Two examples of West African vegetation inhabited by populations of Lemniscomys zebra. Top, degraded Guinea Savanna in the vicinity of Zizonkame (07°55'N, 02°01’E), Benin, in the early rainy season (middle April); examples of both L. striatus and L. zebra were captured here. Photograph by James W. LeDuc. Bottom, Sudan Savanna in the vicinity of Cella (11°38’N, 00°22’W), Burkina Faso, in the late dry season (middle March); only specimens of L. zebra were trapped here. Photograph by Robert E. Vaden. VOLUME 110, NUMBER 4 661 Fig. 11. Two examples of Moroccan coastal vegetation inhabited by populations of Lemniscomys barbarus. Top, mixed argon-sage-grassland at 16 km W Aoulouz (30°42’N, 08°18’W), Agadir Province; snow covered peaks of the High Atlas Mountains are visible in the left background. Bottom, heavily vegetated dunes with thorny shrubs, dense brush, and grasses at 5 km S Taroudannt (30°26'N, 08°54’'W), Agadir Province; samples of L. barbarus were mainly collected in the grassy areas. Photographs by Robert E. Vaden. 662 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON J7PP 00 8 t ° esis orocco » ea Algeria Niger @ /Burkina re e Ivory ~ Coast Penane) Lemniscomys: barbarus re) zebra e hoogstraali 4 Aunise . 2 Vuni: gla cA 7 ¢ ~ e A Fig. 12. Distribution map of African striped mice of the Lemniscomys barbarus species group, based on 649 specimens examined by the authors (see Taxonomic Accounts) from 177 principal collecting localities (see Appendix 1). and tabular comparison of biotic classifi- cations). Although the formal names adopt- ed for ecophysical subdivisions may differ, particularly between plant and animal in- vestigations, the principal areas (whether called subregions, zones, districts, domains, etc.) are remarkably complementary across these several works on African biogeogra- phy. For the present discussion, we employ the terminology formulated by Davis (1962) for Ethiopian Africa, as modeled af- ter the earlier avifaunal studies of Chapin (1932) and Moreau (1952), with the addi- tion of a Barbarian province as delineated by Heim de Balsac (1936), Moreau (1952), and Quézal (1978, called Mediterranean- Northern African Domain). Museum specimens of L. zebra vouch its occurrence across a swath of savanna bio- topes to the south of the Sahara Desert, from Senegal to central Sudan and south- wards to southern Kenya and northcentral Tanzania (Fig. 12). These localities are vir- tually coterminous with three biotic zones within tropical Africa that have been des- ignated as Sudanese Arid (Sahel and Sudan Savannas of Rosevear 1953), Northern Sa- vanna (Guinea Savanna of Rosevear 1953), and Somali Arid. At the western terminus of its distribution in Senegal, L. zebra does not enter the Sahara Desert proper (not present in USNM series from Mauritania). The abrupt southward projection of its range in Benin and western Nigeria (The Dahomey Gap), almost to the Gulf of Guin- ea, conforms to the anthropogenic prolif- VOLUME 110, NUMBER 4 eration of invasive Guinea savanna (Booth 1958, Robbins 1978). At its eastern limits, the extension of L. zebra into Kenya and Tanzania follows an arid tract of acacia steppe called the Somali Arid Zone. Al- though this eastern African region of rela- tively high endemism shares species also native to either northern or southern savan- na biomes (Moreau 1966, Brenan 1978, Coe & Skinner 1993), it displays a slightly stronger biogeographic affinity with the Southern Savanna Zone based on nonforest bird distributions (Crowe & Crowe 1982). However, as so far known, populations of L. zebra do not inhabit any part of the Southern Savanna Zone (Davis 1962, Rau- tenbach 1978—including both his Southern Savanna Woodland and Southern Savanna Grassland), whose northernmost extent reaches western Tanzania (see map in Skin- ner & Smithers 1990:xxii). Locality records of L. barbarus outline a comparatively restricted distribution that adheres to the coastal sectors of Morocco, Algeria, and Tunisia and is bounded by the upper elevations of the Atlas Mountains (Fig. 12). In Morocco, its distribution cov- ers the coastal lowlands and the contiguous Central Plateau, extending inland only to the northern and northwestern foothills of the Anti Atlas, High Atlas, and Middle At- las Mountains (also see Aulagnier & Thev- enot 1986:80). USNM locality records for Morocco document an altitudinal range from sea level to about 750 m, as interpo- lated from topographic maps. In Algeria and Tunisia, the species occurs mainly along the narrow Mediterranean coastal strip but also among the folded ridges of the Tell Atlas, low mountains which closely approach the seacoast (and see Bernard 1969:109, and Kowalski & Rzebik-Kowal- ska 1991:257). Biogeographers have previously recog- nized the area inhabited by Lemniscomys barbarus as the Barbarian Zone (Heim de Balsac 1936, Moreau 1952), a biological is- land isolated on the north and west by the Atlantic Ocean and Mediterranean Sea and 663 on the south and east by the Sahara Desert, which reaches the coastline in Mauritania and in Libya. The northwest rim of the Af- rican continent, by virtue of its maritime geological setting coupled with climatic os- cillations of the Plio-Pleistocene and ac- companying vegetational shifts, has formed a mixing bowl for species spreading from Afrotropical, European, and Asian land- scapes. Indeed, the high diversity and strik- ing endemism recorded for plants and ani- mals of the area bear testimony to its re- fugial character and recurring opportunity for taxonomic differentiation (Heim de Bal- sac 1936, Moreau 1966, Quézal 1978, Cheylan 1990). Although the Barbarian Zone today har- bors a predominance of temperately adapt- ed species of Palearctic affinities, Heim de Balsac (1936) had regarded the majority of mammals living there as having Ethiopian origins. Such an interpretation contrasted to the large Palearctic contingent observed among the avifauna, an apparent zoogeo- graphic anomaly expanded by Moreau (1966). However, Heim de Balsac had con- flated under his ‘‘Indo-Ethiopian”’ category not only broadly distributed paleotropical mammals (many carnivores) but also xeric- adapted species occurring across the deserts of the Sahara, Arabian Peninsula, and Mid- dle East. The many gerbilline rodents num- bering among the latter prejudice the inter- pretation of Moroccan mammals as pre- dominantly ‘Ethiopian’? sensu Heim de Balsac. Subsequent biogeographic studies of Barbarian mammals have supported their substantial Palearctic pedigree (Saint Gi- rons & Petter 1965, Agusti 1989, Aulagnier 1990, Cheylan 1990, Kowalski & Rzebik- Kowalska 1991), a finding concordant with the broad picture of Northwest African bio- diversity as disclosed by other organisms (Moreau 1966, Quézal 1978). Still, the mo- saic of tropical and temperate habitats long offered within the Barbarian region has fos- tered some isolation and differentiation of Subsaharan groups, and Linnaeus’ Barbary Mouse, Lemniscomys barbarus, represents 664 Table 5.—Rodent species? indigenous to the Bar- barian Zone> of Northwest Africa, with indication of their biogeographic affinity or probable origin. Palearctic Saharo- Taxon (* = endemic) Ethiopian European Sindean Sciuridae Atlantoxerus getulus* > Xerus erythropus xX Muridae: Gerbillinae Gerbillus campestris Gerbillus hesperinus* Gerbillus hoogstraali* Gerbillus jamesi* Gerbillus maghrebi* Gerbillus occiduus* Meriones shawi i a ana Muridae: Murinae Apodemus sylvaticus xX Lemniscomys barba- rus* xX Mastomys erythroleu- cus x Mus spretus »4 Myoxidae Eliomys melanurus xX Dipodidae Jaculus orientalis xX Hystricidae HAystrix cristata XxX? Totals 5) 3 8 a Species compiled from systematic reviews of mam- mals for Morocco (Aulagnier & Thevenot 1986; Saint Girons & Petter 1965), Algeria (Kowalski & Rzebik- Kowalska 1991), and Tunisia (Bernard 1969), with the taxonomy updated to accord with Wilson & Reeder (1993). > The limits of the Barbarian Zone have been vari- ously defined, but are here understood to include the coastal lowlands of Morocco, Algeria, and Tunisia, the inland high plateaus, and the Atlas Mountains (for ex- ample, per Quézal 1978). Such a regional definition thus excludes typical desertic formations of the Sahara and, correspondingly, many rodents, especially gerbil- lines. another of these endemics, one of the few rodent species plausibly derived from Af- rotropical (Ethiopian) relatives (Table 5). Other rodent species with satellite popula- tions in Northwest Africa similarly merit re- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON newed study of their differentiation and tax- onomic status (for example, Apodemus syl- vaticus, Mastomys erythroleucus, many species of Gerbillus, Meriones shawi, and Eliomys melanurus). The historical biogeography of Africa’s savanna landscapes has received less atten- tion in contrast to that devoted to forest- dwelling vertebrates and their patterns of distribution, species diversity, and endem- ism (for example: Moreau 1966, Crowe & Crowe 1982, Carleton & Robbins 1985, Grubb 1990, and references cited by each). Nevertheless, biotic affinities among savan- na biomes, as currently delimited and un- derstood, provide sufficient insight to frame a series of hypotheses to explore the evo- lutionary diversification of species of Lem- niscomys. 1.—Are single-striped mice (griselda- rosalia-roseveari), largely confined to the Southern Savanna Zone, the sister group to the striatus and barbarus complexes of northern savannas and woodlands? That is, does possession of multiple stripes, ar- ranged either in continuous (barbarus group) or in punctulated lateral lines (stria- tus group), form a synapomorphy for those more northern mice? Is L. linulus, a single- striped form isolated in far West Africa (Van der Straeten 1980a), the vicariant sis- ter species to L. griselda and its kin in the Southern Savanna Zone, or does the former represent an instance of evolutionary con- vergence, having been derived from a stria- tus-like ancestor through suppression of lat- eral striping? 2.—If the striatus group (distribution centered in Guinea savanna) and barbarus group (more common in Sudan vegetation) are so closely related, how can one explain their divergence? Other studies have di- vulged little support for the latitudinally graded, contiguous savanna bands found south of the Sahara Desert as refugia or centers of endemism (Moreau 1966, Brenan 1978). Evidence from ornithological diver- sity and centers of endemism (Crowe & Crowe 1982), however, provides some sup- VOLUME 110, NUMBER 4 port for a nonforest refugium in eastern equatorial Africa (within the contemporary Somali Arid Zone), where the progenitor of one complex (striatus?) may have differ- entiated in isolation from the other (bar- barus?) in arid habitats fringing the Sahara, before secondary contact and distributional overlap. 3.—The autochthonous origin of L. bar- barus in the Barbarian Zone is parsimoni- ously explained as allopatric speciation af- ter final severance (late Pliocene—early Pleistocene?) from Subsaharan populations (ancestral to L. zebra) along the West Af- rican coast. The ranges of other Afrotropi- cal species with apparently conspecific seg- ments in southern Morocco similarly inti- mate former connection (Xerus erythropus) or document present continuity (Acomys ca- hirhinus, Mastomys erythroleucus) of pop- ulations across this westernmost region. Al- though reasonable as a first explanation, caution is advised, for yet other corridors of contact with Subsaharan Africa may have existed during pluvial intervals of the Pleis- tocene: for example, via the highlands that occur through the middle of the Saharan Desert or along the eastern Mediterranean coast (Quézal 1978). In either case, deter- mining the nearest kinship of L. hoogstraali (eastcentral Sudan) and the status of East versus West African moieties of L. zebra will assume importance in understanding the evolutionary derivation of L. barbarus. Such phylogenetic questions and their geographic scale elegantly lend themselves to investigation using genetic sequencing and phylographic reconstruction (for ex- ample, see Matthee & Robinson 1997, on the springhare, Pedetes). Taxonomic Accounts Lemniscomys barbarus (Linnaeus 1766) Mus barbarus Linnaeus, 1766: unpaginated addenda at end of volume (type locality, ‘‘Barbaria’’ = Morocco).—Trouessart, 1881:124. 665 Arvicanthis barbarus, Trouessart, 1897: 498.—Trouessart, 1904:388. Lemniscomys barbarus, Thomas, 1916:68 (subsequent designation as type species).— Cabrera, 1921:58.—Misonne, 1974: 20 (part)—Honacki et al., 1982:520 (part).—Aulagnier & Thevenot, 1986:80 (part).—Le Berre, 1990:276 (part).— Corbet & Hill, 1991:180 (part).—Musser & Carleton, 1993:601 (part). Lemniscomys barbarus barbarus, Cabrera: 1932:272.—Allen, 1939:391.—Ellerman, 1941:131.—Saint Girons & Petter, 1965: 37.—Kowalski & Rzebik-Kowalska, 1991:259. Lemniscomys barbarus ifniensis Morales Agacino, 1935:390 (type locality, Span- ish Morocco [Morocco], Sidi Ifni, Ifni; holotype-MNCN 9638).—Allen, 1939: 391.—Ellerman, 1941:131.—Saint Gi- rons & Petter, 1965:37.—Misonne, 1974: 20 (listed in synonymy without indication of rank).—Aulagnier & Thevenot, 1986: 80 (listed in synonymy without indication of rank).—Le Berre, 1990:276 (listed in synonymy without indication of rank).— Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Emended diagnosis.—A species of Lem- niscomys characterized by longitudinally continuous dorsal stripes; upperparts gen- erally more muted in tone, alternating dark and light stripes less sharply contrasting due to suffusion of ochraceous over middle dorsum; primary dark stripes on either side of mediodorsal line typically four in num- ber, fifth stripe absent or indistinct; second- ary light line usually present and continu- ous within first lateral dark stripe, incom- plete or poorly defined in second lateral dark stripe; size relatively large and skull robustly developed, as observed in adult di- mensions of both the body (total length > 235 mm) and cranium (occipitonasal length > 29.0 mm; zygomatic breadth > 13.5 mm). Distribution.—Coastal region of Moroc- co, Algeria, and Tunisia, northwest and 666 north of the Atlas Mountains (Fig. 12). Oth- er locality records based on owl-pellet re- coveries or reliable sightings of L. barbarus have been reported for Morocco (Aulagnier & Thevenot 1986), Algeria (Kowalski & Rzebik-Kowalska 1991), and Tunisia (Ber- nard 1969), but these sites do not alter the general picture of distribution as docu- mented by the specimens we examined and enumerate below. Remarks.—Although some specimens used by Linnaeus in his Systema Naturae still exist and may be regarded as types (see Addendum on L. striatus), Van der Straeten has searched unsuccessfully for a likely type of Mus barbarus in the Naturhistoriska Riksmuseet, Stockholm, and the museum of the University of Uppsala, Sweden. Since L. barbarus is the only species of Lemnis- . comys isolated in northwest Africa, wholly allopatric to other congenors, identification poses no problems and restriction of the type locality to some place other than “‘Bar- baria,”” regarded as equivalent to Morocco (Allen 1939, Ellerman 1941), is unwarrant- ed at this time. Specimens examined.—124, as follows. Algeria: Alger (BMNH 12.3.12.144-— 12.3.12.151, 19.7.7.1952; BZM 1664; MNHN 1975.117); Hammam Meskoutine (BMNH 20.7.19.51); Hammam Rirha (BMNH 12.6.12.152—12.6.12.155); Oran (ZMA 8653, 8654; MNHN_ 1953.830— 1953.832, 1955.618, 1955.619, 1961.894, 1961.1063); Reghaia (MAKB 54.111, 54.112). Algeria, locality unknown (BMNH 44.50, 44.51, 19.7.7.1952, MNHN 1956.616). Morocco: Enzel (BMNH 98.7.4.29); For- ét de Néfifik (MNHN 1957.499, 1958.248); 31.5 km on road from Marrakesh to Casa- blanca (MNCN 9657-9659); 15 km WNW Marrakesh (SMF 47.781); Meknés (MNHN 1964.377); Oued Sebo (BMNH 76.1077); Oued Cherrat (ZMA 4834). Agadir Prov- ince, 8 km S Tiznit (USNM 475125); 15 km E Agadir (USNM 475126—475148); 16 km W Aoulouz (USNM 475150—475164); 5 km S Taroudannt (USNM 475165— PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 475168); 8 km N Tamri (USNM 475169). Khouribga Province, 5 km S Oued Zem (CM 45286; USNM 475170—475182). Ra- bat Province, 17 km SW Rabat (USNM 475121, 475122). Safi Province, 13 km E Essaouira (USNM 475123, 475124); 15 km SW Rabat (USNM 475149); 5 km NE Es- saouira (USNM 482104, 483092—483094). Sidi Ifni Province, Ifni (MNCN 9638). Tan- ger Province, 3 km S Cap Spartel (USNM 475183); Tanger (MNCN 9654). Morocco, locality unknown (MNCN 9655, 9660, 9661; MNHN 1950.405, 1950.407, 1950.872). Tunisia: Algeriet (NHRS M810); Cédria (Potinville) (MAKB 71.210—71.214); La Marsa (MAKB 71.215). Lemniscomys hoogstraali Dieterlen, 1991 Lemniscomys hoogstraali Dieterlen, 1991: 11 (type locality, Sudan, Upper Nile Province, 12 miles N Niayok, Paloich, about 10°22’N—32°33’E; holotype, USNM 342078).—Musser & Carleton, 1993: 601. Emended diagnosis.—Linear markings of upperparts sharply contrasted, the pri- mary dark stripes wide and black, offset by narrower, almost white primary light stripes; secondary light stripes weakly ex- pressed and incomplete, hence primary dark stripes undivided; size of skull closely similar to L. barbarus, notably larger rela- tive to L. zebra. Distribution.—Known only from the type locality in eastern Sudan (Figs. 1, 12). Remarks.—Dieterlen (1991:11) diag- nosed this new form as a “Large species of the Lemniscomys barbarus group. Tail length small (101% of head and body length); dorsal coloration rich in contrast and with broad blackish stripes; skull rela- tively large; incisive foramina ending an- terior to M!'; zygomatic plate indented be- low.’’ However, the discriminatory value of several of these traits is unhelpful when compared to large series of L. barbarus and L. zebra. Many specimens of the latter two species have the forward edge of their zy- VOLUME 110, NUMBER 4 gomatic plate undercut ventrally. In fact, the protuberance of the anterodorsal edge of the plate typically becomes accentuated in older specimens, and, as noted by Dieterlen, the holotype of L. hoogstraali is an old in- dividual (age class 5—6). Similarly, the ab- solute and relative length of the incisive fo- ramina is foreshortened in older individuals of L. barbarus, just meeting the anterior roots of the first molar instead of extending conspicuously between them. Contrary to Dieterlen’s assertion that L. hoogstraali ex- ceeds even L. barbarus sensu stricto in cra- nial size, the measurements of the holotype are typical of old adult specimens of L. bar- barus and fall within the variational limits recorded for our Moroccan samples (Table 4, Appendix 2). Finally, the terminal caudal segment of the holotype’s skin appears missing, which may partially account for the exceptional shortness (101%) of its rel- ative tail length (proportion of TL/HBL ranges from 115 to 123% in samples of L. barbarus and L. zebra—Appendix 2). Notwithstanding the caveats attached to certain diagnostic features advanced by Dieterlen, we concur with his naming of the specimen as a new species. Although sim- ilar to L. hoogstraali in cranial size, no specimen of true L. barbarus exhibits such strongly delineated upperparts and broad, richly black stripes. Furthermore, the robust size of L. hoogstraali, as judged by both hindfoot length (28 mm) and most cranial dimensions (Table 4), is certainly greater than any surrounding Sudanese examples of L. zebra. And while some skins of East Af- rican L. zebra may approach the consoli- dated striping evident in L. hoogstraali, none that we have seen quite matches it. More and larger series must be assembled to rigorously evaluate the characters and status of Dieterlen’s new form, so far known only by the holotype, its condition imperfect (tail tip missing; skull with both zygoma incomplete, right bulla crushed, and pterygoid processes broken off). Specimens examined.—1, the holotype. 667 Sudan: Upper Nile Province, 12 miles N Niayok, Paloich (USNM 342078). Lemniscomys zebra (Heuglin 1864) Mus zebra Heuglin, 1864:10 (type locality, Bahr el Ghazal [Sudan], “‘Lande der Req- Neger, Djur und Bongo;”’’ lectotype, SMNS 1100a).—Trouessart, 1881:124. Arvicanthis zebra, Trouessart, 1897:499.— Trouessart, 1904:388. Arvicanthis barbarus zebra, Osgood, 1910: 10. Lemniscomys barbarus zebra, Hollister, 1919:144.—Allen, 1939:392.—Hatt, 1940: 511.—Ellerman, 1941:131.—Setzer, 1956:511.—Kingdon, 1974:620.—Mi- sonne, 1974:20 (listed in synonymy with- out indication of rank).—Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Lemniscomys zebra, Kershaw, 1924b:25. Arvicanthis spekei De Winton, 1897:318 (type locality, Tanganyika Territory [Tan- zania], Unyamuezi; holotype, BMNH 63.7.7.23).—Trouessart, 1904:388. Arvicanthis barbarus spekei, Osgood, 1910: 10. Lemniscomys barbarus spekei, Matschie, 1911:338.—Allen, 1939:391.—Ellerman, 1941:131.—Kingdon, 1974:620.—Mi- sonne, 1974:20 (listed in synonymy with- out indication of rank).—Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Arvicanthis dunni Thomas, 1903:297 (type locality, Western Kordofan [Sudan], Kaga Hills, about 120 mi [193 km] W El Obeid; holotype, BMNH 3.2.8.15).— Trouessart, 1904:388. Lemniscomys dunni dunni, Allen, 1939: 392.—Setzer, 1956:512. Lemniscomys barbarus dunni, Ellerman, 1941:131.—Dekeyser, 1955:204.—Mi- sonne, 1974:20 (listed in synonymy with- out indication of rank).—Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Arvicanthis barbarus convictus Osgood, 668 1910:10 (type locality, British East Afri- ca [Kenya], Voi; holotype, FMNH 17206). Lemniscomys barbarus convictus, Hollister, 1919:146.—Allen, 1939:391.—Ellerman, 1941:131.—Kingdon, 1974:620.—Mi- sonne, 1974:20 (listed in synonymy with- out indication of rank).—Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Arvicanthis barbarus albolineatus Osgood, 1910:11 (type locality, British East Afri- ca [Kenya], Ulukenia Hills, Lukenya Mountain; holotype, FMNH 17194). Lemniscomys barbarus albolineatus, Hol- lister, 1919:144.—Allen: 1939:391.—Ell- erman, 1941:131.—Kingdon, 1974:620.— Misonne, 1974:20 (listed in synonymy without indication of rank).—Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Lemniscomys albolineatus, Kershaw, 1924a: 536. Arvicanthis oweni Thomas, 1911b:120 (type locality—French Gambia [Senegal], upper Gambia River, Gemenjulla, 50 ft [15 mJ]; holotype, BMNH 11.6.10.61). Lemniscomys oweni oweni, Allen, 1939: 394. Lemniscomys barbarus oweni, Ellerman, 1941:131.—Dekeyser, 1955:204.—Ro- sevear, 1969:336.—Misonne, 1974:20 (listed in synonymy without indication of rank).—Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Lemniscomys barbarus manteufeli Mat- schie, 1911:338 (type locality, Tangan- yika Territory [Tanzania], south coast of Lake Victoria, Mwanza; holotype, BZM 16025).—Allen, 1939:391.—Ellerman, 1941:131.—Kingdon, 1974:620.—Mi- sonne, 1974:20 (listed in synonymy with- out indication of rank).—Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Arvicanthis barbarus nigeriae Thomas, 1912:272 (Nigeria, Panyam, 4000 ft [1220 m]; holotype, BMNH 12.1.16.45). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lemniscomys barbarus nigeriae, Allen, 1939:391.—Ellerman, 1941:131.—Ro- sevear, 1953:102.—Dekeyser, 1955: 204.—Rosevear, 1969:336.—Misonne, 1974:20 (listed in synonymy without in- dication of rank).—Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Lemniscomys olga Thomas & Hinton, 1921:9 (type locality, Damergou [Ni- ger], Takoukout, 1550 ft [472 m], about 15°N and 09°E; holotype, BMNH 21.2.11.128).—Ellerman, 1941:131.— Dekeyser, 1955:204. Lemniscomys dunni olga, Allen, 1939:392. Lemniscomys barbarus olga, Rosevear, 1969:335.—Misonne, 1974:20 (listed in synonymy without indication of rank).— Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Lemniscomys dunni nubalis Thomas & Hinton, 1923:267 (type _ locality, Southern Kordofan [Sudan], Nuba Country, Talodi, 1300 ft [396 m]; holo- type, BMNH 18.7.2.14).—Allen, 1939: 392.—Hatt, 1935:1; 1940:513.—Setzer, 1956:512. Lemniscomys barbarus nubalis, Ellerman, 1941:131.—Misonne, 1974:20 (listed in synonymy without indication of rank).— Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Lemniscomys oweni orientalis Hatt, 1935:2 (type locality, Belgian Congo [Zaire], Faradje; holotype, AMNH 49626).—AI- len, 1939:391.—Hatt, 1940:513. Lemniscomys barbarus orientalis, Eller- man, 1941:131.—DMisonne, 1974:20 (list- ed in synonymy without indication of rank).—Musser & Carleton, 1993:601 (listed in synonymy without indication of rank). Emended diagnosis.—A species of Lem- niscomys characterized by longitudinally continuous dorsal stripes; upperparts gen- erally brighter, alternation of dark and light stripes better defined compared to L. bar- barus; primary dark stripes on either side VOLUME 110, NUMBER 4 of mediodorsal line typically five in num- ber, with secondary light lines usually pres- ent and continuous within first and second dark laterals; external and cranial size me- dium and skull moderate in build (total length < 230 mm; occipitonasal length < 29.0 mm; zygomatic breadth < 13.5 mm). Distribution.—Grassy woodlands and sa- vannas south of the Sahara Desert, from Senegal in the west to southern Sudan in the east, southwards through northeastern- most Zaire, northern Uganda and western Kenya, to northcentral Tanzania (Fig. 12); altitudinal range from near sea level to 1220 m in West Africa (Panyam, Nigeria) and to 1065 m in East Africa (north of Dodoma, Tanzania). Remarks.—Heuglin (1864) did not indi- cate a type specimen for Mus zebra, a com- mon omission for the period, nor is his text clear about the number of specimens avail- able when he described the new species. During his visits (in 1979, 1983, and 1989) to the Staatliches Museum fiir Naturkunde, Stuttgart (SMNS), Van der Straeten located two specimens of Lemniscomys that bear evidence of association with Heuglin’s orig- inal description, a mounted skin and skull (SMNS 1100a) and a complete specimen in alcohol (SMNS 1100b). Labels attached to the mounted skin and fluid specimen each contain the information ‘“‘Bongo; Von Heuglin, 1865,’ and the notation as type, the last datum obviously appended more re- cently. Heuglin’s original characterization of the new form provides little help, for it only mentions coloration, pattern of stripes, and external measurements, all of which could have been plausibly derived from e1- ther a whole mount or a fluid-preserved car- cass, or from both. He did note that he was not yet able to examine the molars, which admission suggests that the skull had not been cleaned (Heuglin provided cranial measurements for other species described in the same paper). Both Stuttgart specimens are currently listed as a “Holotypus”’ of Lemniscomys barbarus zebra in the SMNS card catalog (Fritz Dieterlen, personal com- 669 munication), but they should be properly regarded as syntypes of Heuglin’s (1864) Mus zebra. The fully mounted skin with cleaned skull, however, is more useful for ascertain- ing diagnostic traits of the form, and we designate this specimen (SMNS 1100a) as lectotype of Mus zebra Heuglin (1864). The fluid specimen (SMNS 1100b) becomes the de facto paralectotype of the binomen (see Recommendation 73E International Code of Zoological Nomenclature, Third Edition, 1985). The skin is mounted in a more or less realistic posture, and its label indicates male, but the sex cannot be confirmed by inspection of the skin. Although faded, pre- sumably from years of museum exhibit, the mounted skin clearly displays the alternat- ing dark and light (six to seven) dorsal Stripes typical of the dorsal pelage of the barbarus group. Associated with the skull is an older but now invalid museum num- ber, ““Skelett-Katalog 5422,” an extraneous number dating from the outmoded practice of assigning separate registrations to oste- ological and skin preparations, even for the same individual. The light molar wear (age class 1-2) suggests a young, perhaps im- mature, animal. Except for slight breakage of the pterygoid processes and mandibular rami, the skull of the lectotype is otherwise intact and in good condition. According to a posteriori probabilities of membership de- rived from discriminant function analysis, specimen number SMNS 1100a predictably intermingles with other Subsaharan samples that we identify as L. zebra (Fig. 7); in par- ticular, the lectotype was marginally clas- sified (P = 0.45) with specimens from Zaire (OTU 8), a group assignment likely due to its young age and correspondingly smaller size as compared to most Sudanese exam- ples used in our analyses. The form Arvicanthis fasciatus Wrough- ton (1906), named from southern Nigeria and occasionally listed as a subspecies of L. barbarus (Allen 1939, Rosevear 1969), was reidentified as a synonym of L. striatus by Van der Straeten & Verheyen (1980). 670 Rosevear (1969:333) erred in listing the type locality of L. olga Thomas & Hinton (1921) as Farniso, Nigeria, and citing the type specimen as BMNH 21.2.11.87. Van der Straeten has not been able to locate this specimen in the BMNH; the type locality (Niger, Takoukout) and registration number (BMNH 21.2.11.128) given in our synon- ymy agree both with the authors’ original publication and with the provenience as written on the skin tag of the holotype. Although L. zebra probably occurs in westernmost Ethiopia, Yalden’s et al. (1996) report of its presence (as L. barba- rus) at Arba Minch is questionable. Rupp (1980) had allocated these specimens (SMNS 16751, 16754), both young individ- uals, to L. striatus but also confided his dif- ficulty in discerning the striping configura- tion as typical of that of striatus or of bar- barus. The damaged condition of their skins and skulls precludes exact determination, but Van der Straeten tentatively refers the two specimens to L. macculus based on their smaller auditory bullae as determined from his subsequent examination in 1983. Specimens examined.—524, as follows. Benin: Atacora Region, Kouande (USNM 439584, 439585); Porga (USNM 439586); Soubroukou (USNM 439582, 439583). Bor- gou Region, Bimbereke (USNM 422058— 422060); Guene (USNM 422061, 422062); Nikki (USNM 422081—422084); Segbana (USNM 422063—422080). Central Region, Diho (USNM 422057); Zizonkame (USNM 439580, 439581). Benin, locality unknown (MNHN 1913.13). Burkina Faso: Arly (USNM 450853, 450854); 5 mi N Boussouma (USNM 465401); 6 mi S Cella (USNM 465404, 465405); 1 km N Cella (USNM 465406— 465410); Djipologo (USNM 466682-— 466684); Fo (USNM 466674—466679); 5 km SW Koutoura (USNM 466681); Nasso (RUCA 1278, 1281-1283, 1289, 1345- 1347, 1349-1356, 1370-1380, 1393- 1398, 1414-1416, 1459, 1470, A.1—-A.9, A.12—A.15); Natiaboani (USNM 450855); 3 km SE Nayoure (USNM 465411); Nian- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON goloko (BEL 13243); 9 mi S Nobere (USNM 450852); 27 km ENE Orodara (USNM 466680); Ougarou (USNM 465412); Oulo (USNM 465413, 465414); 6 mi SE Seguenega (USNM 465402, 465403). Cameroun: Boukma (RUCA 2.134); Fort Foureau (MNHN 1956.678); 35 km S Gar- oua (AMNH 241233); 35 km S and 10 km E Garoua (CM 58903, 58905, 58909); Isiri (RUCA 1087); Kali (RUCA 1098, 1099, 1114); Koum (RUCA 1018, 1066, 1095, 1122); Maroua, 20 km NW Dogba (CM 14975, 14978); Mora (MAKB 73.393- 73.395, 73.397, 73.398); Sir (RUCA 1.479, 1.482, 1.492, 1.549, 1.591, 1.592, 1.595). Ghana: Damango (USNM 466694-— 466697). Eastern Region, Legon, Accra Plains (USNM 412745—412747). Northern Region, Gambaga (USNM 435405); Na- bogo (USNM 435400—435402); Sakpa (USNM 420577—420585). Upper Region, Bangwon (USNM 420573—420576); Pirisi (USNM 420570); Pulima (USNM 420571, 420572); Shishe (USNM 435403, 435404). Ivory Coast: Bambela (BEL 13177, 13178); Bandama, near Tiebila (FMNH 105180); Bouna (BEL 3798; MNHN 1971.757, 1971.758); Kafiné (BEL 13695); Kong (USNM 465398—465400); Ouango Fitini (BEL 3802, 3932, 13902, 13903, 13931, 13945, 13949, 13984, 23354, 23368, 23373, 23375, 23377;, 23a 23382); Tyenko (USNM 466685—466687); Yama (USNM 466688—466693). Kenya: Garissa District, Galma Galla, 270 ft (AMNH 187678, 187679); Macha- kos District, Kathekani, 760 m (CM 102462, 102463); Kilabasi (AMNH 114445); Voi District, Kenya Karanzi (AMNH 114446); Machakos District, Ki- boko, 3100 ft (USNM 437394); Machakos District, 11 km N and 17 km E Kibwezi (CM 98258); Machakos (BMNH 34.10.26.11, 34.10.26.12); Maungu (CM 57977); Masi, Sand River (AMNH 114449); 50 mi S Moctow (AMNH 114447, 114448); Mtoto Andei (USNM 181737—181740); near Tana River (MCZ VOLUME 110, NUMBER 4 16222); Taveta (BMNH 10.7.2.115); Ulu- kenia Hills, Athi Plains (USNM 162884); Ulukenia Hills, Lukenya Mountain (FMNH 17194); Voi (BMNH 10.6.2.125, 29.3.17.167; FMNH 17206). Mali: Timbuktu (FMNH 44713; ZMK 3885, 3886). Niger: Farak, 1475 ft (BMNH 25.5.12.80, 25.5.12.81, 39.2083, 39.2084); Gerari (Tes- sawa) (BMNH 25.5.12.83); Takoukout (BMNH 21.2.11.128). Nigeria: Bauchi (BMNH 68.491); Borgu (HAP 1318, 1322); Dada (USNM 404085-— 404087); Igbo Ora (USNM 404041); Kab- wir (BMNH 12.9.10.3, 13.5.2.3, 13.5.2.4, 14.11.8.3); 1 mi S Kabwir (USNM 375975); Kishi (HAP 1026); Mada River, 3 mi E Gudi (USNM 404043); 22 mi S Maiduguri, Bornu Ranch (USNM 379011-6); New Bussa (HAP 1068, 1069, 1073); Panyam, 4000 ft (BMNH_ 12.1.16.42—12.1.16.47, 12.4.3.34, 12.4.3.36); Panyam Fish Farm, 2 mi N Pan- yam, Jos Plateau (USNM 404020—404040, 404044-—404084); Shangunu (USNM 379647); Upper Ogun Ranch, 12 mi N Ise- yin (USNM 404042); Wawa (HAP 901, 938, 943, 1778, 1779); Yankari (HAP 1106); Za- ria (HAP 585). Senegal: Bandia (MAKB 76.269); Ge- menjulla (BMNH 11.6.10.61, 11.6.10.63-— 11.6.10.67); Kotiare Naoude, 27 km NE Tambacounda (USNM_ 376533-—376536); Koussanar (USNM 376529-—376532). Sudan: Agur (BMNH 28.3.11.51, 28.3.11.52); Badigeru Swamp, 20 mi E Mongalla (BMNH 20.4.26.22); Bahr el Zeraf (MAKB 27-30); Bahr al Ghazal (BMNH_ 17.10.4.18); Bongo (SMNS 1100a, 1100b); Delami (BMNH 29.5.19.26); Duk Majok (BMNH 8.4.2.41, 8.4.2.42); El Fasher (BMNH 23.1.1.302, 23.1.1.303, 23.1.1.305); Fashoda (BMNH 1.8.8.26); Gallabat (BMNH 28.1.11.151); Gondokoro (USNM 165192); Ikoto, 2500 ft (USNM 299764; ZMK 11942, 11944, 11947); Imurok (ZMK 11934-11937); Juga Juga (BMNH 20.7.30.18); Kadugli (SMF 33.218); Kaga Hills, about 120 mi W El Obeid (BMNH 3.2.8.15); Nuba 671 Mountains, G. Koalib, Kudring (BMNH 29.5.19.27; FMNH 35313, 35314); Kuna (BMNH 23.1.1.308); Li Rangu (USNM 318003); Loa (ZMK 14084); Maridi (SMNS 27396); Molongori (ZMK 14096); Nimule, 1800 ft (FMNH 67293; USNM 299771); Nile-Congo watershed (BMNH 21.1.8.7); Nuba Mountains (BMNH 18.7.2.16); Tadoro (SMF 33.217); Talodi, 1300 ft (BMNH 18.7.2.13, 18.7.2.14); Ter- rakekka (BMNH 28.3.11.53); Torit, 2000 ft (FMNH 66853, 66854, 66857, 66859, 66860; 67171, 67172, 67175,. 79500— 79502; USNM 299765-—299770; ZMK 14081-14083, 14088, 14092, 14100, 14104); 30 km W Torit (SMNS 27398, 27399, 27401, 27402); 80 km E Juba-Torit (SMNS 27397); Umm Keddada (BMNH 233 BARB O95 23ST 1 se 12314. Acad 2). Southern Sudan, locality unknown (SMNS 27405, 27406). Tanzania: Banagi (KBIN 15575); 50 mi N Dodoma, 3500 ft (CM 57782); Jumbe Kadala, Ussnoro (AMNH 55533; BMNH 24.1.1.180); Manyara Lake (BMNH 71.1246, 71.1247, 71.1249); Mawere (AMNH = 83916-83920); Mdjengo’s (BMNH 24.1.1.178, 24.1.1.179); Mtali’s (MCZ 22958); Mwanza, south coast of Lake Victoria (BZM 16025); Ndogowe (BMNH 24.1.1.181); Pooma (MCZ 22957); Unyamuezi (BMNH 63.7.7.23). Togo: Dapango (USNM 438311- 438313); Paio (RUCA 551); Pewa (USNM 438314). Uganda: Apeluk (BMNH 46.757- 46.760); Apoka, Kidepo Valley National Park, Karamoja (BMNH 71.274); near Kaiso Valley, Bunyono (BMNH 71.277); west of road to Kananarock, Karamoja (BMNH 71.272); Karamoja (BMNH 71.273); Kidepo Valley National Park (BMNH 70.1065, 70.1066); near Loi Jome (AMNH 180121); Lorengikipi (AMNH 119166, 119169, 119170); south of Loru- pei River crossing and post 18, Karamoja (BMNH 71.275, 71.276); Malera (BMNH 46.761); Nabilatuk (BMNH 63.928, 63.929); Nabumali (BMNH 67.900, 672 67.901); Rhino Camp, Lado Enclave (CM 850; USNM 165191, 165193, 165194, 165197, 165198, 165200—165205, 165422). Zaire: Faradje (AMNH 49610-49615, 49617-49619, 49621-49633; KMMA 9040, 9191, 13015, 13016); Niangara (AMNH 49608, 49609); Tingasi (AMNH 2150). Addendum on the Type of Mus striatus Linnaeus Among the types housed by the Natur- historiska Riksmuseet, Stockholm (NHRS), is a specimen of Lemniscomys that probably served as the basis of Linnaeus’ (1758) de- scription of Mus striatus, its habitat of ori- gin then given as “‘India.”’ In correcting the type locality of striatus to Sierra Leone, Thomas (1911a) did not remark on the ex- istence of any original specimen, and as of 1969, Rosevear wrote that no type is known to exist. The preparation located by Van der Straeten in 1982 is a whole carcass (with skull still inside) preserved in alcohol. No formal registration number was associated with the fluid specimen, nor did Linnaeus mention any catalog number, per the non- standardized descriptive conventions of the earliest taxonomy. In a paper bag affixed to the bottle is an old label, upon which occurs the handwritten scientific name, ““Mus Striatus,’’ both in cursive and in printed let- ters; the number 48 appears on the outside of this bag. A newer label (one copy on the bottle and one inside with the specimen) contains the information: “Mus striatus Linne Typ. f.beshr.i.Mus. Ad Sp 1.10 Syst Nat 10.62 Gamla saml. 48.’’ This line pro- vides the bibliographic essentials for the name: the reference to Linnaeus’ original description that appeared in a publication (1754) of the Museum Regis Adophi Fred- erici, Volume I, page 10; and the Systema Naturae, 10th edition (1758), page 62, which references the former. Dr. Bo Fern- holm, Department of Vertebrate Zoology, Naturhistoriska Riksmuseet, informed us PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON that ‘“‘Gamla saml. is short for gamla sam- lingen and means Old Collection and refers to the fact that this specimen was # 48 of the old collection (i.e., before 1839) of the Royal Academy of Science, which was our mother institution until 1965 when the mu- seum was separated as an independent gov- ernment institution.” The small size and proportions of the specimen clearly indicate a juvenile animal. The sex of the immature individual appears to be male, although gender is not indicated on any tag. The pelage color seems diluted, perhaps from decades of leaching in old al- cohol, and the stripes of the specimen im- mersed in fluid appear continuous. Never- theless, seven pairs of dark and light punc- tulated stripes, distributed in the manner of - striatus, are discernable on either side of the now brown (once black?) mediodorsal stri- pe when the specimen was removed from the jar and partially dried. The first through third light stripes exhibit distinctly separat- ed spots, but the light markings of the fourth through seventh lines are set close together, suggesting an almost continuous effect. The belly is wholly white, like most Lemniscomys. Measurements obtained from the fluid specimen, as taken by Van der Straeten in 1982, include: length of head and body, 49.1 mm; tail length, 34.5 mm; hindfoot length, without claw, 13.4 mm. Its immaturity is underscored by comparison to these same dimensions in adult L. s. striatus from Ivory Coast, West Africa: length of head and body, 118.1 mm; tail length, 114.1 mm; hindfoot length, 26.0 mm (Van der Straeten & Verheyen 1978). Troussart’s (1881, 1898) early listings of Mus striatus supply circumstantial evidence that the Stockholm specimen is the one ac- tually viewed by Linnaeus. That author in- dicated “‘Mus striatus L. (uv.)” as a syn- onym of M. barbarus, an interpretation consistent with the age and condition of the fluid specimen examined a century later by Van der Straeten. In the absence of any con- trary evidence, this individual (NHRS A53.2048) in the Naturhistoriska Riksmu- VOLUME 110, NUMBER 4 seet, Stockholm, should be considered the holotype of Mus striatus Linnaeus (1758) by reason of monotypy (see Article 73, In- ternational Code of Zoological Nomencla- ture, Third Edition, 1985). Acknowledgments The many museum staff members who have allowed access to or made available for loan specimens under their care were essential to the conduct of this study: G. G. Musser (AMNH); J. Ingles and P. Jenkins (BMNH); B. D. Patterson (FMNH); R. An- germann (BZM); D. Happold (HAP); X. Misonne (KBIN); W. Van Neer (KMMA); R. Hutterer (MAKB); M. Rutzmoser (MCZ); J. Barreiro (MNCN); E Petter (MNHN); B. O. Stolt (NHRS); D. Kock (SMF); E Dieterlen (SMNS); P. Van Bree (ZMA); and H. Baagoe (ZMK). We are in- debted to Dr. Bo Fernholm and Mr. Jan Englund, NHRS, Stockholm, and to Fritz Dieterlen, SMNS, Stuttgart, all of whom made special effort to accomodate our questions about type material contained in their respective museums. Craig A. Lud- wig, Data Coordinator for Vertebrate Zo- ology, USNM, kindly generated species listings for various African localities, and S. Jones helped in numerous ways with clerical assistance. We especially thank Carl Hansen, NMNH Office of Photoservices, for photographing and composing the color plate, and D. EK Schmidt, Mammal Division, USNM, who undertook the exacting cranial photography and distribution maps; their talents significantly enhanced our presen- tation and documentation of the original data. Finally, we acknowledge the manu- Script reviews undertaken by M. E. Holden (AMNH) and R. Hutterer (RMNH); their efforts and thoughtful comments improved our interpretation and conveyance of that data. Any errors of fact or omission remain our own. Literature Cited Aggundey, I. R., & D. A. Schlitter. 1984. Annotated checklist of the mammals of Kenya. I. Chirop- 673 tera.—Annals of Carnegie Museum 53:119— 161. Agusti, J. 1989. On the peculiar distribution of some muroid taxa in the western Mediterranean.— Bollettino della Societa Paleontologica Italiana 28:147-154. Allen, G. M. 1939. A checklist of African Mam- mals.—Bulletin of the Museum of Comparative Zoology, Harvard 83:1—763. Aulagnier, S. 1990. Zoogéographie et statut des car- nivores sauvages du Maroc.—Vie et Milieu 40: 150-155. , & M. Thevenot. 1986. Catalogue des mam- miferes sauvages du Maroc.—Travaux de l'Institut Scientifique Cherifien, Série Zoolo- gique 41:163 pp. Bernard, J. 1969. 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Revised checklist, zooge- ography and conservation.—Tropical Zoology 9:73-164. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Appendix 1.—Geographic coordinates of principal collecting localities mapped in Figure 12. Coordinates are presented either as extracted from the country gaz- etteers published by the United States Board on Geo- graphic Names (= G) or as they appear on original specimen tags and in field catalogs, as designated by the collector (= C). To verify the equivalence of older placenames and currently recognized coordinates, we generally cross-referenced the cartographic informa- tion provided by Aggundey & Schlitter (1984), Davis & Misonne (1964), Delany (1975), Happold (1987), Rosevear (1965), Setzer (1956), and Swynnerton & Hayman (1951) with that contained in the USBGN gazetteers. Where directional and distance modifiers are given following a comma, the collector’s coordi- nates apply specifically to that point from the named village or town. Algeria Alger 36°47'N, 03°03’E G Hammam Meskou- tine 36°27'N, 07°16'E G Hammam Rirha 36°23'N, 02°24’E G Oran 35°42'N, 00°38’E G Reghaia 36°44'N, 03°21’E G Benin Bimbereke 10°14’N, 02°40’E C Diho 08°05'N, 02°31’E C Guene 11°44’N, 03°13’E C Kouande 10°20’N, 01°41’E C Nikki 09°56'N, 03°13’E C Porga 11°02’N, 00°58’E C Segbana 10°56'N, 03°42’E C Soubroukou 09°41'N, 01°38’E C Zizonkame 07°55'N, 02°01’E C Burkina Faso Arly 11°34’N, 01°26’E C Boussouma, 5 mi N_ 12°57’N, 01°05’W C Cella, 1 km N 11°38’N, 00°22’W C Cella, 6 mi S 11°32’N, 00°22’W C Djipologo 10°56'N, 03°07’'W C Fo 11°53'N, 04°31'W C Koutoura, 5 km SW 10°19’N, 04°53’W C Nasso 11°13’N, 04°26’'W G Natiaboani 11°42’N, 00°30’E C Nayoure, 3 km SE 12°15'N, 00°16’E C Niangoloko 10°17’N, 04°55'W G Nobere, 9 mi S 11°26’N, 01°10’W C Orodara, 27 km ENE 11°04'N, 04°41’W C Ougarou 12°10’N, 00°56’E C Oulo 11°54'N, 02°58’W C Seguenega, 6 mi SE 13°24’N, 01°55'W C Cameroun Boukma 08°32/N, 13°55'E G Fort Foureau 12°05'’N, 14°56’E G VOLUME 110, NUMBER 4 677 Appendix 1.—Continued. Appendix 1.—Continued. Garoua, 35 km S & Essaouira, 13 km E 31°30'N, 09°40’W C Kenya Karanzi Kiboko Kibwezi, 11 km N Not located ODS. 37 427 EC Bs. Pa) Yankari 09°45'N, 10°30’E G & 17kmE 02°19’'S, 38°07’E C wae 11°04'N, 07°42’E G Kilabasi 03°58'S, 38°57’E G Lukenya Mountain 01°28’S, 37°03'E G Senegal Machakos OFSUS, 37° 16 EG Bandia 14°37'N, 17°02'W G Maktau (50 mi S) 03°24’S, 38°08’E G Gemenjulla (Dién- Masi, Sand River Not located oundialla) 13713'N: 13°07 W iG Maungu 03°33'S, 38°45’E G Kotiari Naoudé 13°54'N, 13° 27°W G Mtito Andei 02°41’S, 38°10’E G Koussanar 13°52'N, 14°05'W G Taveta Uoe ee eee G Sadan we es ee oie Agur 11°35'N, 30°28’E G Mali Badigeru Swamp 05°20’N, 32°02 EG Timbuktu (Timbouc- Bahr el Derof Not located tou) 16°49'N, 02°59’'W G Bahr al Ghazal 07°46'N, 27°40’E G Delami (Dalami) £1°52’N, 30°28’E G Morocco Duk Majak 09°05’N, 27°51'E G Agadir, 15 km E 30°24'N, 09°28'’W C El Fasher (Al Fa- Aoulouz, 16 km W 30°42’N, 08°18’W C shir) 13°38'N, 25°21’E G Cap Spartel, 3 km S 35°46’N, 05°55'W C Fashoda 09°53'N, 32°07’E G Enzel (Anzel) 33°13'N, 05°58’W G Gallabat (Qallabat) 12°58’N, 36°09’E G Essaouira, 5 km NE 31°31'N, 09°46’W C Gondokoro 04°54'N, 31°40’E G Shagunu Upper Ogun Ranch Wawa 10 km E 09°06'N, 13°29’E C Ifni 29°23'N, 10°10’W G Isiri 08°24'N, 14°36’E G Marrakesh, 15 km Kali 08°23'N, 14°21'E G WNW 31°53’N, 08°07'W ? Koum 08°23'N, 14°31’E G Meknés 33°54'N, 05°33’W G Maroua 10°35'N, 14°20'E G Néfifik 33°43'N, 07°21'W G Mora 11°03’N, 14°09’E G Oued Cherrat 33°50'N, 07°07'W G Sir 10°36'N, 13°41'E G Oued Sebo 34°04’N, 04°56’W C eats Oued Zem, 5 km S 32°49’N, 06°35’W C rig ies Rabat, 15 km SW = 33°57’N, 06°57’W C Bangwon Daa, ora S Rabat, 17 km SW __33°55’N, 06°59’W C Damango splice eed Cc Tau. Suan Mi 30°20'N, 09°50’W C eas Se Taroudannt, 5 km S 30°26’N, 08°54’W C Legon eS Tiznit, 8 km S 29°38'N, 09°43’W C Nabogo 09°45’N, 00°49’W C Pirisi 10°07'N, 02°27'W C Niger Pulima 10°51'N, 02°03’W C Farak 15°18'N, 08°55'E G Sakpa 08°52'N, 02°21'W C Gerari 13°46'N, 07°55'E G Shishe 10°42'N, 00°13’W C Takoukout 15°07'N, 08°30’E G Ivory Coast Nigeria Bambela 09°37'N, 03°54'W G Bauchi 10°16’N, 09°50'E G Bouna 09°19’N, 02°53’W G Borgu 09°52'N, 04°04’E G Kafiné 08°31'N, 05°19’W G Dada 11°34’N, 04°29’E C Kong 09°09’N, 04°37’W C Igbo Ora 07°26'N, 03°17’E G Ouangofetini 09°34'N, 04°03’W G Kabwir 09°24'N, 09°34’E G Tiebila (Tyébila) 09°45’N, 05°50’W G Kishi 09°05'N, 03°51'E G Tyenko 08°14'N, 07°24'W C Mada River 08°54'N, 08°17’E C Yama 09°36'N, 06°18’W C Maiduguri, 22 mi S 11°33’N, 13°16’E C K New Bussa 09°53'N, 04°31'E G enya sa bay anes Panyam 09°27'N, 09°09'E G Galma Galla 01°11'S, 40°47’E G Panyam Fish Farm 09°27'N, 09°12’E C Kathekani 02°37'S, 38°09’E C 10°21'N, 04°28’E C 08°09'N, 03°30’E C 09°55'N, 04°27'E G SNS... | eee 678 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Appendix 1.—Continued. Appendix 1.—Continued. Ikoto 04°06’N, 33°06’E G Lorengikipi 02°24’N, 33°54’E G Imurok 04°19'N, 32°24’E G Lorupei 03°48'N, 33°40’E C Juga Juga 13°48’'N, 25°25’E G Malera 01°27'N, 34°03’E G Kadugli 11°00'N, 29°44’E G Nabilatuk 02°03'N, 34°35’E G Kaga Hills 13°50'N, 27°45’E G Nabumali 00°59’N, 34°13’E G Kudring 11°30’N, 30°06’E ? Rhino Camp 02°58'N, 31°24’E G Kuna (2354 N, 32351 2'G Tana River, near 00°08'N, 38°50’E C Li Rangu 04°43'N, 28°22’E G Zai Loa 03°48’N, 31°57’E G era ito ee Maridi 04°55’N, 29°28’E G Faradje pee 2s G Molongori 04°10'N, 32°52'E G Nugneara 0342.02) ae Ninule 03°36’N, 32/033 EG Tingasi 03°24'N, 27°55’E G Nuba Mountains 10°40’N, 32°11’E G Paloich 10°28’N, 32°32’E ? Tadoro 10°41’N, 30°01’E G Talodi (Talawd1) 10°38'N, 30°23’E G Terrakekka (Terake- ka) 05°26'N, 31°45’E G Torit 04°24'N, 32°34’E G Torit, 30 km W Not plotted Juba-Torit, 80 km E Not plotted Umm Kaddadah 13°36'N, 26°42'E G Tanzania Banagi 02°16’S, 34°S51'E G Dodoma, 50 mi N_ Not located Jumbe Kadala (Nda- la) 04°46’S, 33°16’E G Manyara Lake 03°30’'S, 35°50’E G Mawere Not located Mdjengo’s (Misinko) 04°45'S, 34°40’E G Mtali’s 04°22’S, 34°36’E G Mwanza 02°31'S, 32°54’E G Ndogowe 05°28'S, 34°22’E G Pooma 04°59'S, 34°44'E G Unyamwezi 04°—06°20’S, 32°-33°40’E G Togo Dapango 10°52’N, 00°13’E C Paio (Payo) 10°14'N, 00°41’E C Pewa 09°17’N, 01°14°E C Tunisia Algeriet Not located Cédria (Potinville) 36°42’N, 10°23’E G La Marsa (Al Mar- sa) 36°53’'N, 10°20’E M Uganda Apeluk (Ajeluk) 01°30’N, 33°50’E G Apoka 03°44’N, 33°43’E C Kaiso Valley 01°31'N, 30°58’E C Kananarock 03°58'N, 33°45'E C Karamoja 03°43’N, 33°36’E C Kidepo Valley Natl. Park 03°56'N, 33°42'E G Loi Jome (Lotome) 02°24'N, 34°31'E G VOLUME 110, NUMBER 4 Appendix 2.—Descriptive statistics for selected ex- ternal and cranial variables and OTUs of the Lemnis- comys barbarus group. Species and OTU n Mean Range Total length Lemniscomys barbarus OTU 1 BT ver 23852 211-266 OTU 2 i> 25158 193-273 Lemniscomys zebra OTU 4 2a? 2182 196-236 OTU 5 Ate & 2G 192-232 OTU 7 47 204.8 172-230 OTU 10 Ly) 203.3 170-231 OTU 11 ES LARS 196-230 Tail length Lemniscomys barbarus OTU 1 37. 127.0 110-145 OTU 2 14 = =139.7 130-148 Lemniscomys zebra OTU 4 23 = 120.4 105-134 OTU 5 27... 147 105-130 OTU 7 ager Vie 91-130 OTU 10 | aes al Na | 98-162 OTU 11 Py 7Hs9 109-120 Hindfoot length Lemniscomys barbarus OTU 1 44 27.1 24—29 OTU 2 15 28.7 27-30 Lemniscomys zebra OTU 4 ZS Dae 24-27 OTU 5 a2 25.3 23-27 OTU 7 oH) 2015 23-27 OTU 10 21 24.4 23-25 OTU 11 12 24.3 22-27 Weight Lemniscomys barbarus OTU 1 44 45.8 28-56 OTU 2 15 35.5 38-69 Lemniscomys zebra OTU 4 £1 a A 5 te 18-31 OTU 5 32 25:9 18—34 OTU 7 oi 25.6 20-34 Occipitonasal length Lemniscomys barbarus OTU 1 44 50.1 28.3-32.3 OTU 2 14 31.2 30.0—31.2 Lemniscomys zebra OTU 4 22 27.8 26.1—29.1 OTU 5 25 By fee | 26.1—29.3 10.5 14.5 0.9 0.6 0.8 0.9 679 Appendix 2.—Continued. Species and OTU n Mean Range SD OTU 7 51 27.2 25.9-29.1 0.8 OTU 10 12 2a 25.7—28.6 1.1 OTW 11 42 pe ("| 26.4—29.1 0.8 Zygomatic breadth Lemniscomys barbarus OTU 1 43 14.2 13.6—-15.1 0.4 OTrer2 14 14.7 13.7-15.7 0.5 Lemniscomys zebra OTU 4 EAR 12.7 11.9-13.4 0.4 OTU 5 Di, 12.7 11.8-13.9 0.4 OTU 7 a1 13.0 12.2-—13.8 0.4 OTU 10 12 12.6 11.7-13.3 0.5 OTU 11 12 13.3 12.6—-14.1 0.5 Breadth of braincase Lemniscomys barbarus OTU 1 44 12.4 11.8-13.1 0.3 OTU 2 14 12:5 12.1-12.9 0.2 Lemniscomys zebra OTU 4 pies 11.8 11.1-12.5 0.3 OTU 5 25 11.8 11.2—12.3 ) i OTU 7 51 11.8 11.2-12.4 0.3 OTU 10 by 11.8 11.5-12.1 0.2 OTU 11 12 12.0 11.5-12.5 0.3 Interorbital breadth Lemniscomys barbarus OTU 1 44 4.7 4.4-5.1 0.2 OTU 2 14 4.9 4.5-5.3 0.2 Lemniscomys zebra OTU 4 pip) 4.5 4.0-4.9 0.2 OTU 5 2S 4.5 4.14.8 0.2 OTU 7 x1 4.3 4.0-4.8 0.2 OTU 10 12 4.3 4.1-4.6 0.2 OTU 11 12 4.3 3.9-4.5 0.2 Length of nasals Lemniscomys barbarus OTU 1 44 iS 10.1—12.6 0.5 OTU 2 14 12 11.6—-13.2 0.4 Lemniscomys zebra OTU 4 pH. 10.6 9.7-11.3 0.4 OTU 5 2D 10.6 9.6—-11.7 0.5 OTU 7 51 10.3 9.3-11.4 0.5 OTU 10 12 10.0 9.1-10.9 0.6 OTU 11 12 10.5 9.9-11.0 0.4 Postpalatal length Lemniscomys barbarus OTU 1 44 10.5 9.6—11.5 0.5 Org'z 13 10.7 10.4—10.9 0.2 680 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Appendix 2.—Continued. Appendix 2.—Continued. Species and OTU n Mean Range SD Species and OTU n Mean Range SD Lemniscomys zebra Alveolar length of maxillary toothrow OTU 4 2D 9.4 8.5-10.3 0.4 Lemniscomys barbarus OTU 5 23 9.4 8.8-10.3 0.4 OTU 1 44 5.59 5.22-5.98 0.18 OTU 7 51 9.1 7.6-10.0 0.4 OTU 2 15 5:60). °5.37-5:83 "Ons OTU 10 12 9.3 8.5—10.1 0.6 : Omi 12 9.8 92104 . 05 EEWSS LI OREN WT 5 OTU 4 24 531 4.95-5.81 0.23 Length of incisive foramen OTU 5 32 5.25 5.02--5.59 0.14 Lemniscomys barbarus OTU 7 51 5.31 4.95-5.74 #045 OTU 1 44 5.9 5.4-6.6 0:2 OTU 10 De 5.09 4.89-5.26 0.11 OTU 2 14 6.2 By /=OS) 0.2 OTU 11 15 5.33 5.07—5.62 0.17 Lemniscomys zebra Width of first upper molar OTU 4 DD 5.4 4.7-6.0 0.3 Lemniscomys barbarus OTU 5 25 5.5 4.9-5.9 0.3 OTU 1 44 1.73 1.611. 86 eis OTU 7 51 5.2 4.5-5.9 0.3 OTU 2 15 1.74 ‘“L6s=ss7 as OTU 10 12 37 A 5—5.6 0.3 } . OTU 11 12 5.4 5.0-5.8 0.3 BECO ee j OTU 4 25 1.66 1.54—1.82 0.07 LETE DG. GAscsand * Vonus 32 1.68 157218 ong Lemniscomys barbarus OTU 7 51 1.71° 1.54—1.84 0.07 OTU 1 44 7.4 6.7—-8.2 0.3 OTU 10 22 1.61 1.48-1.73 0.06 OTU 2 14 7.8 73-85 0.3 OTU 11 15 1.68 1.56—1.80 0.05 Lemniscomys zebra OTU codes: 1, Morocco, Agadir Province; 2, Mo- OTU 4 22 6.5 6:0-7 2 0.3 rocco, Khouribga and Rabat provinces; 4, northern OTU 5 25 6.6 6.0—7.4 0.3 Ivory Coast and western Burkina Faso; 5, Ghana and OTU 7 51 6.3 5.8—7.0 0.3 Togo; 7, Nigeria, Northern Region, Jos Plateau; 10, OTU 10 We 6.4 5.86.9 0.3 Sudan, Equatorial Province; 11, Kenya. OTU 11 12 6.6 5.9-7.1 0.4 Palatal breadth across MIs Lemniscomys barbarus OTU 1 44 6.2 5.9-6.6 0.2 OTU 2 15 6.2 5.9-6.5 0.2 Lemniscomys zebra OTU 4 Dp, 5.8 5.5—-6.3 0.2 OTU 5 25 5.8 5.5-6.3 0.2 OTU 7 5). 5.8 5.3-6.2 0.2 OTU 10 |) 5.6 5.2—5.9 0 Om 1 2 oe 5.3—5.9 0.2 Breadth of zygomatic plate Lemniscomys barbarus OTU 1 44 3.8 3.44.3 0.2 OTU 2 14 3.9 3.44.3 0.2 Lemniscomys zebra OTU 4 2D 3.4 3.0—3.9 0.2 OTU 5 Des) 3.4 3.1-3.8 O72 OTU 7 5)) 3.4 3.0—3.8 0.2 OTU 10 12 33 3.1—3.7 O72 OBUyit 12 35 3.3—3.8 0.1 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110(4):681—692. 1997. A phylogenetic analysis of the southern pines (Pinus subsect. Australes Loudon): biogeographical and ecological implications Dean C. Adams and James F Jackson (DCA) Department of Ecology and Evolution, State University of New York at Stony Brook, Stony Brook, New York 11794-5245, U.S.A.; (JFJ) Department of Biology, University of Southwestern Louisiana, Lafayette, Louisiana 70504, U.S.A. Abstract.—A parsimony analysis on morphological characters was per- formed to estimate the phylogenetic relationships of the taxa of Pinus subsect. Australes. The Adams consensus tree placed the Caribbean species as a mono- phyletic clade with P. rigida—P. serotina as its sister taxon. Based on this phylogeny, area cladograms were constructed and compared to geologic clado- grams constructed from plate-tectonic evidence. This comparison and an an- cestral area analysis indicate that colonization of the tropics most likely oc- curred from Florida to Hispaniola, rather than by the circumferential-Gulf route. Subsequent dispersal events to Central America, Cuba, and the Bahamas are proposed to explain the geographic distribution of P. caribaea. Ecological comparisons within subsect. Australes found that sister species are not syntopic and that syntopic species are not sister species. Although some North American sister species are ecologically quite different, there is low ecological diversity among the Caribbean species. Pinus is one of the most widespread gen- era of plants in the northern hemisphere (Mirov 1967:307—308; Strauss & Doerksen 1990). Its species are found from Central America and Sumatra to the Arctic circle (Little & Critchfield 1969, Strauss & Doerksen 1990, Farjon 1996), occupy xeric to mesic habitats, and comprise one of the dominant vegetation types on the earth. They are important economically, being used for fuel and lumber, and are frequently planted for commercial purposes in parts of the world where they do not naturally occur (Mirov 1967:451). While often considered early successional species well-adapted to poor soils (Govindaraju 1984), pines are ac- tually ecologically diverse and are found from sea level to the timberline, from sea- sonally wet savannas to deserts, and from monotypic stands to multispecies climax forests where there is co-occurrence with hardwoods. McCune (1988) characterized the ecological diversity for North American pines and defined five ecological groups: (1) fire-resistant species, (2) mesophytic shade-tolerant species, (3) stress-tolerant species, (4) fire-resilient species, and (5) southern mesic species. Other studies provide a more quantita- tive assessment of the interaction between pines and their environment. In particular, studies of bark ontogeny and fire have shown most pines to be particularly well adapted to surviving surface fires (Har- mon 1984, D. C. Adams 1994). Recent work by Jackson and Adams demonstrates the evolution of negative bark allometry (D. C. Adams & Jackson 1995) in species whose habitats are characterized by fre- quent surface fires. This finding is consis- tent with models of defensive structure evolution, where resources are allocated 682 earlier to defense as the likelihood of a mortality factor increases. Although the ecological roles of pines are relatively well defined, aspects of the classification and systematics of the genus Pinus have been uncertain (Mirov 1967: 540, Strauss & Doerksen 1990). Shaw (1914) published the first major taxonomic work on pines, where he recognized two subgen- era, Strobus and Pinus. Pilger (1926) pre- sented a second classification of the pines, classifying many of Shaw’s (1914) varieties as species and further dividing the subgen- era into eleven sections. Because of his heavy reliance on needle number, however, Pilger’s classification is considered to be a step backwards (Mirov 1967:526). Duffield (1952) revised the classification of the pines, using information from hybridization . studies. He divided Shaw’s “‘Group Austra- les’’ into two “‘Groups,”’ the eastern species (XI) and the western species (XII). In Shaw’s (1914) classification, most of the southern pines (P. echinata, P. elliottii, P. glabra, P. palustris, and P. taeda) as well as the Caribbean and many western species, were included in the “Group Australes’’. Duffield also added three species from Shaw’s “Group Insignes”’ to his “‘Group XI’: P. rigida, P. serotina, and P. pungens. Little & Critchfield (1969) reviewed the many classification schemes for Pinus. In particular, they restored the name Australes, at the subsectional rank, to Duffield’s ““Group XI,’ which now included the eight southern pines and three Caribbean species. With the advent of phylogenetic meth- odology, many subsections of the genus Pi- nus have been reexamined in an evolution- ary context, so that the relationships among many of the North American taxa are now better known (Wheeler et al. 1983, Strauss & Doerksen 1990, Malusa 1992, Govinda- raju et al. 1992, Krupkin et al. 1996). The Species comprising subsect. Australes, how- ever, have not been examined in a phylo- genetic context. In particular, it is of interest to determine whether the Caribbean taxa comprise a monophyletic clade and, if so, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON what its relationship is to the North Amer- ican species. Mirov (1967:555) thought that insular varieties of P. caribaea originated from a Central American progenitor, but without a phylogenetic analysis he could not specify whether this was the original colonization of the Caribbean islands by subsect. Australes or was part of a subse- quent radiation within the Caribbean re- gion. Farjon (1996) on the other hand, sug- gested that the Caribbean taxa were of a North American origin, and that the Central American P. caribaea originated from Ca- ribbean immigrants. Because of these out- standing questions, we felt that an exami- nation of the relationships of the taxa in subsect. Australes was needed, and there- fore performed a phylogenetic analysis of the southern pines using morphological characters. In addition, we generated an area cladogram for the geographic regions occupied by the taxa and compared this to geologic evidence in order to better under- stand the vicariance and dispersal of the taxa in the Caribbean. Methods Phylogenetic analysis.—The eight spe- cies of subsect. Australes in the eastern United States and certain geographically defined OTUs of the three Caribbean spe- cies were used in this study (Appendix I). Pinus caribaea from the Bahamas, Cuba, Belize, and Honduras-Nicaragua were treat- ed as separate OTUs. Because P. caribaea in the uplands of Belize and Honduras may be subject to hybridization with P. oocarpa (Williams 1955) we used only specimens from lowland localities. Pinus elliottii var. densa was also treated as a separate OTU. The subsection Australes is included within the hard pines (subg. Pinus) as classified by Little & Critchfield (1969). While the re- lationships among the hard pines are still somewhat controversial (see Strauss & Doerksen 1990, Govindaraju et al. 1992), we chose to use two taxa (P. virginiana and P. clausa) from subsect. Contortae as out- VOLUME 110, NUMBER 4 Table 1.—Data matrix for the taxa of Pinus used in this study. Descriptions of characters and character codes are found in Appendix II. ~ DD WWD Species . caribaea—Bahamas . caribaea—Belize . caribaea—Cuba caribaea—Hondu- ras—Nicaragua clausa cubensis echinata elliottii var. elliottii elliottii var. densa glabra occidentalis palustris pungens rigida . serotina taeda . virginiana Character states 0000101011 1101311112 7 0001111011 1101411111 8 0000111001 1100511111 9 0001111011 1101411112 6 0010101110 3100100000 2 0000111011 1111211014 7 0000011111 1100200000 2 1000110011 1100211012 2 1000111011 1100211013 3 0000011111 1100100000 2 0000101011 1101611113 3 1001100000 2000401113 0 1101100000 3100201005 O 0000111011 3100411102 1 0000111011 2101511102 0 1101100000 3100301100 4 0010101110 3100101000 5 groups, based on the phylogenies of Gov- indaraju et al. (1992) and Farjon (1996). Twenty one morphological characters were scored from herbarium specimens (LAE MO, NY), with supplementary infor- mation from North American specimens collected by us and from literature sources (Farjon & Styles 1997, Radford et al. 1964); species were assigned values based on an average of several specimens. Four- teen characters were based on cone mor- phology and seven on needle morphology (Appendix II). Four of these characters were coded as ordered, multi-state charac- ters (Table 1; Appendix II). Binary scoring was employed for the remainder. While rec- ognizing that some of the characters are quantitative (Stevens 1996), we believe bi- nary scoring adequately approximates char- acter states among taxa where we have used it. We generated a phylogenetic hypothesis through Wagner parsimony using the branch and bound algorithm in PAUP ver- sion 3.1 (Swofford 1991). Wagner parsi- mony attempts to reconstruct an evolution- ary tree by minimizing the number of changes of the character states along the 683 tree (Kluge & Farris 1969, Strauss & Doerksen 1990). If one assumes that a spe- cies from a closely related group reflects previous character states, it can be used as an outgroup to root the tree and to polarize the character states as well (Wiley 1981). Biogeographic analysis.—Comparing the current distributions of taxa with their phylogenetic history to elucidate patterns, as well as to evaluate the relative plausibil- ities of vicariance and dispersal, is the pur- pose of cladistic biogeography (Nelson & Platnick 1981, Wiley 1988, Morrone & Cri- si 1995). To examine the biogeography of subsect. Australes in a phylogenetic con- text, we used the method outlined by Brooks & McLennan (1991). After the phy- logenetic relationships were estimated, the geographic areas containing the taxa were defined: eastern North America (NA), the Bahamas (B), Cuba (Cb), Hispaniola (H), the Yucatan peninsula (Y), and Honduras- Nicaragua (Ch). The relationships of these geographic areas were then determined by representing the relationships of the taxa by a matrix containing additive binary codes, replacing the species with their respective geographic areas, and performing a parsi- mony analysis. Brooks Parsimony Analysis (BPA) has been criticized for its treatment of widespread taxa as synapomorphous for areas (Kluge 1988) and as a method for generating general area cladograms (Nelson & Ladiges 1991). Our goal was to evaluate alternative historical biogeographies rather than to produce a general area cladogram. Having treated the species population of each major Caribbean locale as an OTU, our data set did not, by definition, have widespread taxa, but there was a redundant distribution with two OTUs in Cuba. We chose to derive area cladograms by BPA with (inclusive ORing: Cressey et al. 1983) and without redundant distributions in order to evaluate the redundancy (Brooks 1990). The ancestral area of the Caribbean taxa was estimated by the technique of Bremer (1992). This involves comparing areas in regard to the numbers of necessary gains 684 and losses in the area cladogram by Camin- Sokal parsimony and under the assumption that each area is ancestral. The putative an- cestral area best supported by the biogeo- graphic evidence is that having the highest ratio of gains to losses. Phylogenetically-based cladograms are intended for comparison with geological area cladograms. Much work has been done on the biogeography of the Caribbean flora and fauna (e.g., Rosen 1976, 1978, 1985; Guyer & Savage 1986; Page & Lydeard 1994; Hedges et al. 1994). However, due to the complex geologic history of the region (see Pindell & Barrett 1990), many of the interpretations of biogeography are highly controversial and out of date. In light of the increased knowledge of Caribbean geology (Burke 1988, Pindell & Barrett 1990), we felt that using geologic cladograms from previous biogeographic work might be un- wise. We therefore compared the area cladograms for subsect. Australes to geo- logic cladograms generated from recent tec- tonic evidence on the relationships of the geologic regions of the Caribbean. Results Phylogenetic analysis.—We found three most parsimonious trees from our cladistic analysis, each containing 81 steps and a consistency index (CI) of 0.469 (Fig. 1). When uninformative characters were ex- cluded, the consistency index was reduced by only 0.010 to CI = 0.459, implying that most characters were phylogenetically in- formative. From these three most parsimo- nious trees we generated a consensus tree, using the procedure described by E. N. Ad- ams (1972, 1986). This tree contains only that information present in all rival tree to- pologies, and is thus a conservative esti- mate of the true topology. Based on this phylogeny, subsect. Aus- trales is divided into several distinct sub- clades. The smaller subclade is a polytomy containing P. taeda, P. pungens, and P. pa- lustris. A second polytomy is located deep- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON er in the phylogeny and contains the smaller subclade, the larger subclade, and P. echin- ata and P. glabra as single taxa. The re- maining taxa are all found within the larger subclade. That the phylogeny locates P. rig- ida and P. serotina as sister taxa accords with the proposal by Smouse and Saylor (1973), who consider them conspecific. The most significant aspect of this phylogeny is that the Caribbean species are monophyletic within the larger subclade. Within the larger subclade, it is notewor- thy that P. elliottii var. densa is the sister taxon to P. elliottii var. elliottii. Based on the chemical composition of its turpentine, Mirov et al. (1965) and Mirov (1967:555) proposed that P. elliottii var. densa was more closely related to the Caribbean spe- ‘cies, and was a recent arrival to Florida. Our findings suggest that it is in fact more closely related to the mainland slash pine than to any Caribbean taxon. In addition, our phylogeny suggests that P. cubensis is more closely related to P. caribaea than to P. occidentalis, in contradiction to the pro- posal by Mirov (1967:232). Our findings place P. occidentalis basal to the Caribbean subclade. Biogeographic analysis.—Much of the history of the Caribbean region is still un- known, as is evident from the numerous tectonic models of the region (Perfit & Wil- liams 1989). However, the present study fo- cuses on taxa in the Greater Antilles, the Bahamas, the Yucatan peninsula, and Hon- duras-Nicaragua, where there seems to be a general consensus of opinion concerning the geologic history. We therefore present a brief account of the geologic history of the region (summarized in Fig. 2) and use this to generate geologic cladograms for the taxa. Mexico collided with the North Ameri- can plate during the Jurassic (Burke 1988, Pindell & Barrett 1990). The Greater An- tilles were part of a larger body called the Great Arc, which originated in the Pacific during the late Cretaceous and migrated northeast into the Atlantic in the Paleocene VOLUME 110, NUMBER 4 Fig. I. characters of Pinus spp. (Burke 1988, Pindell & Barrett 1990). The Great Arc broke into three segments some- time in the late Cretaceous to early Paleo- cene (Burke 1988). The northern segment became the Greater Antilles, the central be- came the Lesser Antilles, and the southern segment is presumed to have collided with the South American continent. During the late Paleocene, the Greater Antilles began to collide with the Bahaman plate (Pindell & Dewey 1982, Burke 1988, Pindell & Barrett 1990). However, Burke (1988) claims that prior to this, the Greater Antilles collided with the Yucatan. Others 685 P. taeda P. pungens P. palustris P. elliotti P. elliottii-densa P. rigida P. serotina P. occidentalis-Hispaniol P. cubensis P. caribaea-Bahamas P. caribaea-Cuba P. caribaea-Belize P. caribaea-Honduras P. echinata P. glabra P. virginiana P. clausa Adams consensus tree found from three most parsimonious trees based on twenty-one morphological (Pindell & Barrett 1990) claim that al- though there is geologic evidence for such a collision, it cannot be determined whether it was the Greater Antilles or some other geologic body that collided with the Yuca- tan. Strike-slip faults began to separate Cuba and Hispaniola in the mid-Eocene (Pindell & Barrett 1990), and the Honduras- Nicaraguan block (called Chortis) began to collide with the Yucatan shortly after in the Miocene (Perfit & Williams 1989). Though Pindell and Barrett place this collision slightly earlier in the Oligocene, this does not affect our geologic cladograms. Based 686 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Tertiary paleogeography of the Caribbean (redrawn from Perfit and Williams 1989). A. Late Creta- ceous. B. Paleocene. C. Mid-Eocene. D. Late Miocene-Pliocene. AR = Aves Ridge, B = Bahaman Platform, CA = Cuban Arc, CH = Chortis, H = Hispaniola, J = Jamaica, L = Lesser Antilles Arc, NH = North Island of Hispaniola, SH = South Island of Hispaniola. on this geologic information, we have con- structed two area cladograms (Figs. 3a, b). The first (Fig. 3a) represents the relation- ships of the areas if the Yucatan had not collided with the Greater Antilles (sensu Pindell & Barrett 1990), and the second (Fig. 3b) if it had (sensu Burke 1988). The taxon-based area cladogram derived with inclusive ORing has Hispaniola as the sister area to the other Caribbean areas and situates Cuba as the sister area to the Yu- catan, in a derived position (Fig. 4a). The taxon-based area cladogram that treated Cuba as two separate areas historically (ex- clusive ORing) has a pattern of branching identical to that of the taxon cladogram, but with Cuba as a sister area to both the Yu- catan and the Bahamas (Fig. 4b). The an- cestral area analysis (AAA) of Bremer (1992) was carried out in two ways: on an area cladogram with terminal sister taxa not grouped, which corresponds to Fig. 4b; and on an area cladogram that considered there to be no node separating the two OTUs of P. caribaea of Cuba and Belize. The logic of combining the areas of the terminal sister VOLUME 110, NUMBER 4 A NA Y Ch B Cb H ee NA Y Ch B Cb H Fig. 3. Geologic cladograms based on tectonic evolution in the Caribbean. (A) corresponds to hy- pothesis of Pindell and Barrett (1990), where the Yu- catan did not collide with the Greater Antilles; (B) corresponds to Burke’s (1988) hypothesis, which in- cludes this collision. taxa from this node is that the taxa are con- specific. For both approaches to AAA, His- paniola is the most probable ancestral area, and Cuba is the next most probable (Table ZY: The positions of Yucatan-Chortis and Hispaniola differ between the geological and taxon-based area cladograms (Figs. 3 and 4). In the geological area cladograms, Yucatan-Chortis is either the sister area to North America or the basal area in the Ca- ribbean, whereas the taxon-based area cladograms have Yucatan-Chortis separated and both in more derived positions. Hispan- iola is placed near the origin of the Carib- bean subclade by the taxon-based area cladogram, but in a derived position by the geological data. We conclude that the cladogram based on geological contact and separation corresponds poorly to the se- quence of areas colonized by the Caribbean taxa of subsect. Australes. The derived placement of Cuba in the 687 A NA Ch B Y Cb H ING Fig. 4. Area cladograms of six geographic locali- ties based on the phylogeny for Pinus subsect. Austra- les using: (A) inclusive ORing, and (B) exclusive OR- ing. Letters correspond to the different geographic regions: NA = southern United States, Ch = Chortis, B = Bahamas, Y = Yucatan, Cb = Cuba, H = His- paniola. taxon-based area cladogram calculated by inclusive ORing is due to there being two derived taxa (P. cubensis and P. caribaea) on Cuba that determine the location of Cuba on the area cladogram (Brooks & Table 2.—Estimation of ancestral area for the Ca- ribbean subclade of the subsection Australes. Values not in parentheses are for cladogram that reduces con- specific sister taxa. Values in parentheses are for clado- gram without reduction of conspecific sister taxa. G = number of necessary gains under forward Camin-Sokal parsimony. L = number of necessary losses under re- verse Camin-Sokal parsimony. AA = G/L rescaled by division by the largest G/L. AA = mean of AA’s cal- culated by the two reduction alternatives. Area G L G/L AA AA Hispaniola 1 (1) 1 (1) 1.00 (1.00) 1.00 (1.00) 1.00 Cuba 2 (2) 3 (4) 0.67 (0.50) 0.67 (0.50) 0.59 Chortis 1 (1) 2 (2) 0.50 (0.50) 0.50 (0.50) 0.50 Yucatan 1 (1) 3 (4) 0.33 (0.25) 0.33 (0.25) 0.29 Bahamas 1 (1) 4 (4) 0.25 (0.25) 0.25 (0.25) 0.25 688 McLennan 1991:212). The presence of de- rived taxa and the lack of ancestral taxa on Cuba can be explained through intra-Cuban differentiation combined with colonization of other Caribbean areas from Cuba. The AAA clearly supports this explanation. Discussion Based on our phylogenetic hypothesis of the southern pines, several ecoevolutionary trends can be recognized. First, sister spe- cies are not syntopic. For example, the sis- ter clades of P. rigida and P. serotina as well as P. cubensis and P. caribaea repre- sent taxa that are allopatric. Pinus pungens and P. palustris are allopatric as well and occupy different habitats. Second, species that are syntopic are not sister species. Ex- amples of this are P. palustris and P. el- liottii in the eastern Gulf coastal plain flat- woods, P. palustris and P. echinata in the uplands of the western Gulf coastal plain, and P. taeda and P. echinata in the south- ern Piedmont. Major ecological divergence is found be- tween some sister species. For example, P. pungens grows in dense monospecific stands in xeric montane locations and main- tains dominance by mass recruitment after stand-replacing crown fires; its bark grows with positive allometry (Adams 1994). In comparison, P. palustris grows as widely spaced individuals in parkland vegetation subject to frequent surface fires, and early life history stages of this species are fire- resistant (McCune 1988), as typified by its ““grass’’ stage and negative allometry of bark. Curiously, there is a general lack of ecological divergence in the Caribbean sub- clade of subsect. Australes (Smith 1954). This subclade is less diverse in that there are no mountain ridge specialists like P. pungens, there are no shade-tolerant species like P. glabra, and there are no species with a fire-resistant seedling stage like P. palus- tris or P. elliottii var. densa. The purpose of cladistic biogeography is to attempt to- discern whether current dis- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tribution patterns are the result of vicariance or dispersal. In subsect. Australes this issue is related to whether the initial colonization of Caribbean islands was from eastern North America or from Central America. Both the ancestral area analysis and the fact that neither Yucatan nor Chortis branch low on the area cladogram suggest that the Greater Antilles were the first sites of col- onization in the Caribbean. This could have occurred by dispersal from North America in the late-Tertiary or Quaternary, or by vi- cariance from Central America if a com- ponent of Hispaniola collided with, and then separated from, Yucatan during the Pa- leocene (Burke 1988). The issue is one of timing and therefore cannot be decided con- clusively without molecular data on the di- vergence time between North American and Caribbean taxa. However, several facts ar- gue in favor of dispersal to Hispaniola. For example, there is no evidence in Central America of a Caribbean ancestor; P. cari- baea cannot play this role given its derived status. The Caribbean subclade was derived from an ancestor that shared many traits, particularly of cones, with P. serotina. Cur- rently P. serotina has an Atlantic-East Gulf Coastal Plain distribution, not being found west of Mobile Bay. This puts P. serotina geographically closer to the late-Tertiary Greater Antilles than to the putative contact between the Yucatan and the proto-Greater Antilles. In addition, the lack of ecological diversification in the Caribbean clade could suggest an occupation of the region too re- cent to have occurred via Paleocene vicar- iance. Farjon (1996) found that P. occiden- talis and P. caribaea var. hondurensis formed a sister group to all the other neo- tropical pines he considered, a result con- sistent with derivation from a southeastern North American ancestor. Dispersal of Australes to the Greater An- tilles may have been part of a pattern of colonization by xeric-adapted biota. R. P. Adams (1989) presented evidence from leaf morphology and volatile leaf oils that strongly- indicates. an origin of West Indian VOLUME 110, NUMBER 4 Juniperus species from eastern North America, rather than from Mexico. Buck (1990) suggested that several xeric bryo- phyte species found in North America and upland Hispaniola colonized the island via dispersal during glacial episodes of the Pleistocene when over-water distances were less due to lower sea levels (Gascoyne et al. 1979) and when savanna habitats were more widespread than at present because of a more xeric climate (Pregill & Olson 1981). Such a scenario could apply to col- onization of the Greater Antilles by subsect. Australes as well as to inter-island move- ment and colonization of Central America. Mirov (1967:555) hypothesized that P. caribaea colonized the Caribbean islands from Central America. The area cladogram, with Chortis as the sister area to all other areas occupied by P. caribaea, supports this direction of colonization in the species. Dis- persal is implicated because no connection is known to have existed between Chortis and the Greater Antilles. The only possibil- ity of vicariance within the P. caribaea clade would be between the taxonomic sub- units in Cuba and Yucatan. However, any such vicariance would date from early Ter- tiary (Burke 1988), and this appears too longstanding for the small level of differ- entiation between the taxa. It is difficult to ascribe any of the divergences within the Caribbean subclade to vicariance. It was believed by Mirov et al. (1965) that P. elliottii var. densa was a recent im- migrant to Florida from the Caribbean and that it was therefore more closely related to the Caribbean species. Our results place P. elliottii var. densa as the sister taxon to P. elliottii var. elliottii, suggesting that it is more closely related to the mainland spe- cies. Squillace (1966) demonstrated multi- character clinal variation between P. elliot- tii var. elliottii and P. elliottii var. densa in central Florida. We also found that P. cub- ensis is more closely related to P. caribaea than to P. occidentalis. Mirov (1967:232) had proposed that P. cubensis and P. occi- dentalis were closely related taxa. Clearly 689 more work is needed to determine their re- lationship. Klaus (1980) suggested that, for compar- isons through nodes deep within the phy- logeny of Pinus, cones display primitive character states at the apex and derived states at the base. Consideration of intra- cone character variation in the context of the subsect. Australes cladogram provides evidence of this phenomenon at a finer phy- logenetic scale. Pinus glabra, P. elliottii, and P. rigida often have flattened, distally- pointed umbos at the cone base and more erect umbos at the apex. Within the clado- gram, erect umbos characterize the out- group and the P. taeda-P. pungens-P. pa- lustris subclade, whereas flattened umbos are typical in P. serotina and the Caribbean subclade. In P. caribaea var. hondurensis, the umbo spine tends to be unconnected to the keel on basal scales, a character state more widespread in P. cubensis cones, but connected to the keel on apical scales, a state found throughout the cone in taxa de- rived from nodes below P. caribaea vat. hondurensis. A counterexample, however, exists in the relatively erect umbos apically in some P. caribaea var. hondurensis. This character state is derived if the Caribbean clade arose through an ancestor like P. oc- cidentalis, yet it appears at the apex. Our phylogenetic hypothesis of subsect. Australes is a first step in understanding the history of the pines of the southern United States and the Caribbean. Based on this phylogeny, we have identified possible dis- persal and vicariance events and have pro- vided a framework on which future studies may be based. While it is appealing to in- terpret phylogenetic hypotheses as recon- structions of evolutionary history, it must be stressed that they are only estimations of the true topology based on the available data. Studies have shown that most phylo- genetic methods can perform rather poorly in their estimation of true tree topology (Fi- ala & Sokal 1985; Rohlf et al. 1990). We therefore present this phylogenetic hypoth- 690 esis as a tentative estimation of the rela- tionships of the southern pines. Acknowledgments We thank the Herbaria of the New York and Missouri Botanical Gardens and the University of Southwestern Louisiana for providing specimens. We thank A. Farjon for comments on an earlier version of the manuscript, Ursula Jackson for drawing Fig. 2, and W. D. Reese for taxonomic counsel. This work is contribution number 998 from the program in Ecology and Evo- lution at the State University of New York at Stony Brook. Literature Cited Adams, D. C. 1994. 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Pinus caribaea.—Ceiba 4:299— Appendix I Herbarium specimens from which characters were scored. Pinus caribaea Morelet var. bahamensis Barrett and Golfari. BAHAMAS. Grand Caicos: Correll 49463 (NY). New Providence: Degener 18753 (NY). P. car- ibaea var. caribaea Barrett & Golfari. CUBA. Pinar del Rio: Jack 8673 (NY), Leon & Charles 4935 (NY). P. caribaea var. hondurensis B. & G. BELIZE. Belize: Croat 24005 (MO), Kluge s.n. (MO). Corozal: Crane 315 (MO). Orange Walk: Lundell 677 (MQ). Stann Creek: Stevenson 1128 (MO). Toledo: Gentle 3690 (MO). HONDURAS. Gracias a Dios: Clewell 4511 (MO), Nelson & Hernandez 1004 (MO). NICARA- GUA. Zelaya: Marshall & Neill 6559 (MO), Seymour 3650 (MO), Stevens 7638 (MO), Stevens 7753 (MO), Stevens 21679 (MO), Vincelli 625 (MO). P. clausa (Chapm.) Vasey. USA. Florida: Franklin Co., Godfrey 69282 (LAF); Polk Co., Shuey 2317 (LAF); Walton 692 Co., Smith 2071 (LAF). P. cubensis Griseb. CUBA. Oriente: Ekman 3829 (NY), Shafer 4174 (NY). P. echinata Mill. USA. Louisiana: Bienville Par., Thieret 16836 (LAF), Westling 141 (LAF); Evangeline Par., Reese & Reese 1537 (LAF), Thieret 22256 (LAF); St. Helena Par., Allen 1499 (LAF). P. elliottii Engelm. var. densa Little & Dorman. USA. Florida: Dade Co., Small & Carter 1249 (NY); Monroe Co., Brizicky & Stern 378 (NY). P. elliottii var. elliottii L. & D. USA. Louisiana: Lafayette Par., Thieret 17423 (LAF); St. Tammany Par., Thieret 16770 (LAF), Thieret 21879 (LAF); Tangipahoa Par., Thieret 16758 (LAF). P. gla- bra Walt. USA. Louisiana: Livingstone Par., Thieret 16743 (LAF); St. Helena Par., Allen 1500 (LAP), St. Tammany Par., Lynch 940 (LAF); Washington Par., Thieret 16782 (LAF). P. occidentalis Swartz. DOMIN- ICAN REPUBLIC. Zanoni, Mejia, Pimentel & Garcia 32443 (NY). HAITI. Nash 825 (NY). P. palustris Mill. USA. Louisiana: Allen Par., Thieret 10389 (LAF);- Beauregard Par., Thieret 16968 (LAF); Nachitoches Par., Thieret 16811 (LAF), Thieret 17035 (LAF); St. Helena Par., Thieret 17298 (LAF); Tangipahoa Par., Thieret 16782 (LAF); Winn Par., Thieret 16817 (LAF). P. pungens Lamb. USA. Pennsylvania: Franklin Co., Adams s.n. (LAF), Adams 25 (LAF). P. rigida Mill. CANADA. Quebec: Lemieux 1281 (LAF). USA. Maryland: Frederick Co., Windler & Stastny 3900 (LAF). North Carolina: Buncombe Co., Dunton s.n. (LAF); Jackson Co., Duncan 22793 (LAF). Pennsyl- vania: Franklin Co., Adams s.n. (LAF). West Virginia: Hardy Co., Wratchford s.n. (LAF). P. serotina Michx. USA. Alabama: Geneva Co., Kral 33947 (LAF). Geor- gia: McIntosh Co., Duncan 20688 (LAF). South Car- olina: Charleston Co., Ahles 53156 (LAF), P. taeda L. USA. Louisiana: Livingston Par., Thieret 16742 (LAF) Thieret 16753 (LAF), Thieret 16754 (LAF); St. Helena Par., Thieret 17300 (LAF); Tangipahoa Par., Thieret 16756 (LAF). P. virginiana Mill. USA. Alabama: DeKalb Co., Vincent 1281 (LAF). Georgia: Rabun Co., Reade s.n. (LAF). Maryland: Allegany Co., Dun- can 22944 (LAP). North Carolina: Orange Co., Ahles 53117 (LAF). Tennessee: Cheatham Co., Demaree 49183 (LAF). Appendix II Twenty-one morphological characters used in this study. The first fourteen characters describe mature PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON megasporangiate cone morphology and the last seven describe needle morphology. (1) Sum of maximum cone length and maximum cone width: less than 20 cm = 0; greater than 20 cm = 1. (2) Cones subsessile or stalked = 0; sessile = 1. (3) Transverse keel or dorsal surface of cone scale between apophysis and umbo: depressed at junc- tion = 0; continuous at junction = 1. (4) Lateral portion of most umbos not strongly ele- vated above apophysis = 0; strongly elevated above apophysis = 1. (5) Umbo: weakly keeled or unkeeled = 0; strongly keeled = 1. (6) Umbo keel: straight or monotypically curved = O; undulating = 1. (7) Proximal portion of umbo slopes at: high angle or is perpendicular to apophysis surface = 0; low angle from spine to proximal edge of umbo = 1. (8) Proximal portion of umbo: not concave = 0; con- cave = l. (9) Portion of umbo proximal to keel: not larger than distal portion = 0; much larger than distal portion = 1. (10) Junction of distal margin of umbo and apophysis: grooved = O; not grooved = 1. (11) Spine on umbo: unkeeled on apical half = 1; keeled on part of apical half but unkeeled at apex = 2; keeled on all of apical half = 3. (12) Spine on umbo: reflexed = 0; straight or curved outward = 1. (13) Spine: connected to keel = 0; substantially distal to keel = 1. (14) Most umbos erect such that apex of spine is not pointed distally near the level of the apophyseal keel = 0; most umbos flattened and turned dis- tally such that apex of spine is pointed distally near the level of the apophyseal keel = 1. (15) Needle. number: 2:= 1;-2(—3) = 273@2)2— 3 3 = 4; 3(-4) = 5; 3-5 = 6. (16) Hypodermal cells in leaf angles: absent = 0; present = 1. (17) Hypodermis structure: uniform = 0; biform = 1. (18) Endodermal cells: walls normal = 0; walls thick- ened = 1. (19) Resin canals medial = 0; internal = 1. (20) Number of resin canals: coded character. (21) Number of stomata: coded character. 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 an Abstract in an alternate language when appropriate. 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The Society, on request, will subsidize a limited number of contributions per volume. Payment of full costs will facilitate speedy publication. Costs.—Printed pages @ $60.00, figures @ $10.00, tabular material @ $3.00 per printed inch per column. One ms. page = approximately 0.4 printed page. CONTENTS Pachyrotula, a new genus of freshwater sponges from New Caledonia (Porifera: Spongillidae) Cecilia Volkmer-Ribeiro and Klaus Riitzler The genus Julavis de Laubenfels (Porifera: Halichondrida) Rob W.M. van Soest and Helmut Lehnert Narella nuttingi, a new gorgonacean octocoral of the family Primnoidae (Anthozoa) from the eastern Pacific Frederick M. Bayer Annotated list of Veronicellidae from the collections of the Academy of Natural Sciences of Philadelphia and the National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C., U.S.A. (Mollusca: Gastropoda: Soleolifera) José W. Thomé, Patricia H. dos Santos, and Luciana Pedott Revision of the scaleworm genus Eulagisca McIntosh (Polychaeta: Polynoidae) with the erec- tion of the subfamily Eulagiscinae and the new genus Pareulagisca Marian H. Pettibone Protodrilus gelderi, a new species of infralittoral, interstitial polychaete from Massachusetts Bay Nathan W. Riser Synonymy of Platicrista cheleusis (Tardigrada: Eutardigrada: Hypsibiidae) R. Deedee Kathman and Clark W. Beasley The cladoceran collection of the National Museum of Natural History, Smithsonian Institution, Washington, D.C. Dorothy B. Berner Aglaodiaptomus atomicus, a new species (Crustacea: Copepoda: Calanoida: Diaptomidae) from freshwater wetland ponds in South Carolina, U.S.A., and a redescription of A. saskatche- wanensis (Wilson 1958) Adrienne E. DeBiase and Barbara E. Taylor Argyrodiaptomus nhumirim, a new species, and Austrinodiaptomus kleerekoperi, a new genus and species, with redescription of Argyrodiaptomus macrochaetus Brehm, new rank, from Brazil (Crustacea: Copepoda: Diaptomidae) Janet W. Reid Lophogaster muranoi, a new species of mysid from the coastal waters of Argentina (Crustacea: Mysidacea: Lophogastridae) Kouki Fukuoka, Monica S. Hoffmeyer, and Maria D. Vinas A new species of troglobitic crayfish of the genus Cambarus, subgenus Aviticambarus (De- capoda: Cambaridae), endemic to White Spring Cave, Alabama John E. Cooper and Martha Riser Cooper Two new species and a range extension: of mud shrimps, Upogebia, from Pacific Costa Rica and Mexico (Decapoda: Thalassinidea: Upogebiidae) Austin B. Williams A trans-Atlantic record of the fossil tropicbird Heliadornis ashbyi (Aves: Phaethontidae) from the Miocene of Belgium Storrs L. Olson and Cyril A. Walker A new species of Cyclemys (Testudines: Bataguridae) from Southeast Asia John B. Iverson and William P. McCord Morphological differentiation among Subsaharan and North African populations of the Lem- niscomys barbarus complex (Rodentia: Muridae) Michael D. Carleton and Erik Van der Straeten A phylogenetic analysis of the southern pines (Pinus subsect. australes Loudon): biogeograph- ical and ecological implications Dean C. Adams and James F. Jackson Table of Contents, Volume 110 Index to New Taxa, Volume 110 489 502 STI 520 2/6 f/ 552 558 560 569 581 601 608 | CLUS bh A co a Mee tbe a Se) eae di. 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