ties BS Puasa va Meri At soa) Raereyee z texto» Madgseeane EGE NTN ¥ oy etal ah Wah NE » mee dinate yegeatsa.® ices certs Greets ire Ae VR NPS MEY ath DNA ney esPrath stl ST ike ee ceo Perr ora bed ao eka, 4 Nog huge art a es Soy Rady aie 4 eae te OT Brena rede ss ary eh Mop ehh feigh SEY SS rh NE NES, whee rews siedtese peciye ele eee nes yk pees Vee Yipee nner ee acer a ons ee aeeerecen aie cane ae ss! Sede ase? "pe lhe Sligte ee Dae yeteet eee etre treet acta, evar Bye oh sae retest ore Sipe Sage coms ep ape ee Ree Ear Jha eet Fea at de SDE DM ee no) TE re reige seen Pek ae PANN oneal! perenne rss fast? Tha ee sees Fete are ie PROCEEDINGS of the Biological Society of Washington VOLUME 109 1996 Vol. 109(1) published 16 April 1996 Vol. 109(3) published 9 October 1996 Vol. 109(2) published 25 June 1996 Vol. 109(4) published 23 December 1996 WASHINGTON PRINTED FOR THE SOCIETY 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 Ib JEWETT JAMES N. NORRIS TABLE OF CONTENTS Volume 109 Adrianov, Andrey V. and Robert P. Higgins. Pycnophyes parasanjuanensis, a new ki- norhynch (Kinorhyncha: Homalorhagida: Pycnophyidae) from San Juan Island, Wash- HNN SOOT GUND AN ca See eR ee ee ee ee Almany, Glenn R. and Carole C. Baldwin. A new Atlantic species of Acanthemblemaria (Teleostei: Blennioidei: Chaenopsidae): Morphology and relationships —_.___-__________- Alvarez, Fernando, Maria Elena Camacho, and José Luis Villalobos. The first species of Prionalpheus from the eastern Pacific, and new records of caridean shrimp (Crus- tacea: Decapoda: Caridea) from the western coast of Mexico —------- Ashe, James S., Robert M. Timm, and Milton H. Gallardo. Systematics, distribution, and host specificity of Edrabius Fauvel (Insecta: Coleoptera: Staphylinidae) Bayer, Frederick M. The Antarctic genus Callozostron and its relationship to Primnoella (Octocorallia: Gorgonacea: Primnoidae) —__-__-------- === aan Bayer, Frederick M. Three new species of precious coral (Anthozoa: Gorgonacea, genus Gorallivem trom aciticwattes ss he Bayer, Frederick M. The gorgonacean genus Arthrogorgia (Octocorallia: Primnoidae) Bravo, Manuel Rafael and Masaaki Murano. Description of Neobathymysis japonica, a new genus and species, and revision of the genus Bathymysis with a new species from Japany (Crustacea Miysidace asiN fy SiG ac) eee Bueno-Soria, Joaquin and Silvia Santiago-Fragoso. Studies in aquatic insects X: De- scriptions of five new species of the genus Culoptila Mosely (Trichoptera: Glosso- SOmatidae)) fire rms We xc Oa ee ee Chevaldonné, Pierre and Karine Olu. Occurrence of anomuran crabs (Crustacea: Decap- oda) in hydrothermal vent and cold-seep communities: a review __-----_-_-_-_--___-_- Child, C. Allan. Additions to the Pycnogonida fauna of Carrie Bow Cay, Belize, middle PNTIV CNT C Qiks a ae el eee eh A cag ln hae ae oh Se ee Child, C. Allan. Pycnogonida of the western Pacific islands, XII. A recent diving survey of Okinawa; Ryukyullslands\ 22 ee ee eee Child, C. Allan. Pycnogonida of the western Pacific islands, XIII. Collections from Indonesia, Melanesia, and Micronesia Child, C. Allan. The Pycnogonida types of William A. Hilton. II. The remaining un- GES CHI DS AUS PS CLES (cass ere aie I ae en ee ee Child, C. Allan and Michel Segonzac. Sericosura heteroscela and S. cyrtoma, new species, and other Pycnogonida from Atlantic and Pacific hydrothermal vents, with FXO LHSAS OV Lenveloviet cereal) SNA OL SNE Dahlgren, Thomas G. Two new species of Dysponetus (Polychaeta: Chrysopetalidae) ALOMIBL Call yeearrcl zap Ua INS was Gouri Damkaer, David M. Copepod taxonomy: Discovery vs. recognition _________-__---_---------- Desbruyéres, Daniel and Lucien Laubier. A new genus and species of ampharetid poly- chaete from deep-sea hydrothermal vent community in the Azores triple-junction Ferrari, Frank D. and E. L. Markhaseva. Parkius karenwishnerae, a new genus and species of calanoid copepod (Parkiidae, new family) from benthopelagic waters of the Eastern ach eH OCC ea ea a ee Fukuoka, Kouki and Masaaki Murano. Siriella tuberculum, a new species (Crustacea: Mysidacea: Mysidae) from Akajima Island, Ryukyu Islands, Japan __.____________ Garcia-Garza, Maria Elena, Gabino A. Rodriguez-Almaraz, and Thomas E. Bowman. Spelaeomysis villalobosi, a new species of mysidacean from northeastern Mexico (CrustaceayMy sid acca) (ie 1 ae etre ai eee eee ee ee Gelder, Stuart R. Description of a new branchiobdellidan species, with observations on three other species, and a key to the genus Prerodrilus (Annelida: Clitellata) _.______- Gelder, Stuart R. A review of the taxonomic nomenclature and a checklist of the species of the Branchiobdellae (Annelida: Clitellata) Giray, Cem and Gary M. King. Protoglossus graveolens, a new hemichordate (Hemi- chordata: Enteropneusta: Harrimanidae) from the northwest Atlantic 236-247 419-429 715-724 731-743 150 205 605-628 501-511 446-452 286-298 526-532 533-539 540-559 677-686 664-676 575-585 687-694 248-255 264-285 512-516 97-102 256-263 653-663 430-445 Graves, Gary R. Hybrid wood warblers, Dendroica striata X Dendroica castanea (Aves: Fringillidae: Tribe Parulini) and the diagnostic predictability of avian hybrid ENS 11 © iy JS Se a EE SE TE a Sn Graves, Gary R. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 2. Hybrid origin of Eriocnemis soderstromi Butler __...----.-------------------------- a2 Graves, Gary R. and Nancy L. Newfield. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 1. Characterization of Calypte anna X Stellula calliope and the possible effects of egg volume on hybridization potential ___---_---- Handley, Charles O., Jr. New species of mammals from northern South America: Bats of the genera Histiotus Gervais and Lasiurus Gray (Chiroptera: Vespertilionidae) —__ Harvey, Alan W. and Elizabeth M. De Santo. On the status of Pachycheles laevidactylus Ortmann, 1892 (Crustacea: Decapoda: Porcellanidae) —__________-_--_------ Healy, Brenda. Records of Enchytraeidae (Annelida: Oligochaeta) from west Florida. 1. Mesenchytraeus, Cogenettia, Bryodrilus, Hemienchytraeus, Henlea, and DEYOV EMO GIO) Ses Ne ek Se ee ee ee eee Humes, Arthur G. Orecturus amplus, a new species (Copepoda: Siphonostomatoida: Asterocheridae) from an alcyonacean in New Caledonia ____-_---------------------- Karasawa, Hiroaki and Hisayoshi Kato. Dalldorfia Rathbun, 1904 (Crustacea: Decapoda) ROTH HS. INGORE Or Ue DE) =e a ee ee ee Kensley, Brian. Systematics and distribution of the genus Calocarides (Crustacea: De- CRYBYOYGIE AO IIG EY) ) a ee eee Kensley, Brian. A new species of the axiid shrimp genus Acanthaxius from the Carib- bean (Crustacea: Decapoda: Thalassinidea) _ Kensley, Brian and Mary Bursey. Rediscovery of Cymodocella algonense from South Africa (Crustacea: Isopoda: Sphaeromatidae) — = Knapp, Leslie W. Review of the genus Cociella Whitley (Teleostei: Platycephalidae) with the description of three new species _.--- Kornicker, Louis S. and Kenneth G. McKenzie. The adult male of the myodocopid ostracode Philomedes cubitum Kornicker, 1975, from the Strait of Magellan (Crusta- ceaOstracodas My odocopina): 22. ee ee Lechapt, Jean-Paul and David W. Kirtley. Bathysabellaria spinifera (Polychaeta: Sa- bellariidae), a new species from deep water off New Caledonia, southwest Pacific (IBY a ee eee Lemaitre, Rafael and Darryl L. Felder. A new species of ghost shrimp of the genus Sergio Manning & Lemaitre, 1994 (Crustacea: Decapoda: Callianassidae) from the Ganibbeanicoastof (Colombia 2 oa eee See Lewis, Julian J. and Thomas E. Bowman. The subterranean asellids of Texas (Crustacea: WSOPOG aA Elli ae) ee AU eke Be AEE FS SED SO ED oe PN) Tee SANE eS Lips, Karen R. and Jay M. Savage. A new species of rainfrog, Eleutherodactylus phasma (Anura: Leptodactylidae), from montane Costa Rica ____---------------- Lucas, Spencer G. and Robert J. Emry. Early record of indricothere (Mammalia: Per- issodactyla: Hyracodontidae) from the Aral Sea region of western Kazakhstan _____. Lucas, Spencer G. and Robert J. Emry. Late Eocene entelodonts (Mammalia: Artio- dactyla) from Inner Mongolia, Chima ____--__--------------- onan Lucas, Spencer G., Robert J. Emry, and Robert W. Purdy. Marine fossil shark (Chon- drichthyes) from nonmarine Eocene sediments, northeastern Kazakhstan ____-_-______- Manning, Raymond B. and Darryl L. Felder. Nannotheres moorei, a new genus and species of minute pinnotherid crab from Belize, Caribbean Sea (Crustacea: Decapoda: | FUE M SOC DYER UG FAYED) i see ee Re eee Mathis, Wayne N. Australian beach flies (Diptera: Canacidae) McLaughlin, Patsy A. and Janet Haig. A new genus for Anapagrides sensu De Saint Laurent-Dechancé, 1966 (Decapoda: Anomura: Paguridae) and descriptions of four TTS UME [OC CLS fy oe ae oo os 8 a a a a UD Re ce RN McLaughlin, Patsy A. and John P. Hoover. A new species of Aniculus Dana (Decapoda: Anomura: Diogenidae) from Hawait _____-------_-------- anna Ng, Peter K. L. and Boris Sket. The freshwater crab fauna (Crustacea: Decapoda: Brachyura) of the Philippines. IV. On a collection of Parathelphusidae from Bohol - 373-390 764-769 755-163 1-9 707-714 118-137 112-117 44-52 53-69 70-74 91-96 17-33 517-525 560-574 453-463 482-500 744-748 391-396 397-405 349-352 311-317 326-348 75-90 299-305 695-706 Vi Ng, Peter K. L. and Peter Trontelj. Daipotamon minos, a new genus and species of potamid crab (Crustacea: Decapoda: Brachyura) from a cave in China _.-_ Okuno, Junji. Cinetorhynchus manningi, a new shrimp (Crustacea: Decapoda: Caridea: Rhynchocinetidae) from the western Atlantic Pettibone, Marian H. Review of Hermilepidonotus Uschakov, 1974, and two species of polynoidspoly.chaetess(éepidono tin ac) pees Pettibone, Marian H. Revision of the scaleworm genera Acholoe Claparéde, Arctonoella Buzhinskaja, and Jntoshella Darboux (Polychaeta: Polynoidae) with the erection of the new subfamily Acholomacs. 2. 28a ns 8 a ee Rodriguez-Alamaraz, Gabino A. and Ernesto Campos. New locality records of fresh- water decapods from México (Crustacea: Atyidae, Cambaridae, and Palaemonidae) _ Rossman, Douglas A. Identity and taxonomic status of the Mexican garter snake Tham- nophis vicinus Smith, 1942 (Reptilia: Serpentes: Natricidae) Rozbaczylo, Nicolas and Marco A. Méndez. Artacama valparaisiensis, a new species of Terebellidae (Annelida: Polychaeta) from subtidal soft bottoms of Valparaiso Bay, Ci ee ee ee Rozbaczylo, Nicolas, Elba Canahuire, and F. Patricio Ojeda. Presence of Micronereis in Antarctic waters and description of a new species, M. antarctica (Polychaeta: Ne- UCI CH AS PIN LO PP EMV CUT AC) es wre a ee Sandberg, Lennart. Hermit crabs of the genus Paguristes (Crustacea: Decapoda: Di- ogenidae) from the western Atlantic. Part III. Paguristes markhami, a new species fromthe yBahannrayandi@€arcosy ls laricl sy eee ese eee ea eee Savage, Jay M., James R. McCranie, and Mario Espinal. A new species of Eleutherodactylus from Honduras related to Eleutherodactylus bransfordii (Anura: Leptodactylidae) _______ Scanlin, Megan and Janet W. Reid. A new copepod species from California, U.S.A.: Hesperodiaptomus californiensis (Crustacea: Copepoda: Calanoida: Diaptomidae) __ Sluys, Ronald. Reconsiderations of the species status of some South American Planarians (Blatyhelmimthessyiricladida:|Paludicola) Smith, Brian E. and Jonathan A. Campbell. The systematic status of Guatemalan pop- ulations of snakes allied with Ninia maculata (Reptilia: Colubridae) __________ Springer, Victor G. and Helen K. Larson. Pholidichthys anguis, a new species of pho- lodichthyid fish from Northern Territory and western Australia Stark, Bill P. New species of Macrogynoplax (Insecta: Plecoptera: Perlidae) from Peru TN UR ER Uy A 1 a geet ae ee ee Ie Sse eee Tavares, Marcos and Rafael Lemaitre. Lonchodactylus messingi, a new genus and spe- cies of Cyclodorippidae (Crustacea: Decapoda: Brachyura) from the Bahamas _______ Werding, Bernd. Description of a new porcellanid, Petrolisthes gertrudae from the southeastern Caribbean Sea (Crustacea: Decapoda: Procellanidae) —___--_-_---_-_---- Westheide, Wilfried and Giinter Purschke. Leptonerilla diplocirrata, a new genus and species of interstitial polychaetes from the island of Hainan, south China (Nerillidae) __________- Wicksten, Mary K. Neocrangon zacae (Chace, 1937) synonymized with N. resima (Rathbun, 1902), and compared with N. communis (Rathbun, 1899) (Decapoda: Car- Le aes Crary ori Cl Ae) a ree eg eee Woodman, Neal. Taxonomic status of the enigmatic Cryptotis avia (Mammalia: Insec- tivora: Soricidae), with comments on the distribution of the Colombian small-eared Shrewai Gry prolisiCOlombiana ese se ee pec es NO Zecchini, Fulvio, Michael Vecchione, and Clyde F. E. Roper. A quantitative comparison of hectocotylus morphology between Mediterranean and western Atlantic populations of the squid //lex coindetii (Mollusca: Cephalopoda: Oegopsida: Ommastrephidae) _- 476-481 725-730 143-149 629-644 34-38 10-16 138-142 645-652 470-475 366-372 103-111 229-235 749-754 353-365 318-325 464469 306-310 586-590 39-43 409-418 591-599 INDEX TO NEW TAXA Volume 109 (New taxa are indicated in italics; new combinations designated n.c.) CNIDARIA Anthozoa PNT CIENT © OG DA ALELEITO 111 UL go ea Ne tk a a a So eh the a Vogt GAMLOZOSHONTACAIT CAE Smee a a EG od Deo a aN be lease CLA TINGYO NCTA YC ace ee eee (CDT PIC ANE rc 9 oe a a ee ATA] $C LEV AE MA cle N 2 Cap ete ae ent nan Dkk ak eh bl ol oll E ko ein A eae Coralie Kis 2112 OLsiy Came ee ee Rd Fe nai A eee Nt eae SO ae HUITE cact y r e e ANNELIDA Polychaeta EN OET © TE OTN ANE se tad wre Ne ad ed es en Se ae PANPTICET LDN) S Rammer ea 2 fn Rn AU ee sn I ee EE ae hh LTE zener Re Rol et a Eee ev Pe eee Le ane ieee ae ER Ae PNT UAC ATIN ASV AU PLL CLUS TC FUSES 1 0 a eee ee A a BeathhySabelll arias py ir1ife rly gaa ae a ee eo ee ee ee PY SPOTS UUS PE pVCa py ELL CALLUS pte a a I ee OUCLC TOC ULL CULL S las oe te aR ae Pere ta er Sd JS aS eS Mo Se EIN Ds Eenmmilepidonotushelotypus MC.) se ee SDC TOE OTL T LLL Cl mgt was seas A a es I Se eI Se RL nat LUD O CUTICLE Clee eee eg PLN ASD oN Oe es eh SE OR AO LS INAV CTOMELCES ROL LAT CLIC Chap tesa cae ER a ee NEEL LOTTOE 11 Ch gy eet re ee Smee se ees a eso tla 7 Re GEO) OVA OVE a0 EI 01 Ory pes a ese ee me Cee Eke ee oa HTN CEL AST Ne mee eae eI 0 ae NIN Rs es ee (CONST SATIS. ee PE EE ee ee ee ee eee eee INTESEm CHV LACUIS HIATT LE OTe Umm ere ce a ERS a ha Ee a en rs ew eae PAR SIO Teal ONS Co Aa HPAL OH ATR Pe aL Ue ea UR mn Ue rape geen LL PN HNS Ni eam Ce LL O'S Ch aa a a a de Fal et a eh ete a SN A PASTN@ POL OCLAG ENTS WP OCIS a 9 a Mh he ae ee mee MANS COT D TIC EAU S 9 CEL UH Se as a a Roa gl MEZA eA eA LTV SV 079 OS Cae ae a ee ek ere IS CTH COS UTA TLIC tae rel Le al le ll se AA VRCLON OS COU Ch Beata ae Bc Ne age en Se) Sai das SW SN SN Ne Vill PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ARTHROPODA Crustacea Acanthaxius-kipkillent. =e ee ee eee 71 Amniculus hopperae™ ee Ee a ee NE oan oe RCD AY En Re ee 299 Bathiyiny Sis cistaree ta ee a a 504 Gaecidotealbilincatay ee ee ee ee ee 490 Galocanidesicapensisy === a ee ee ee 55 TILL CTI OTS O11 Yea ere RES a Pec so hate IN i Re ed Dao 58 Cinetorhynchusimaninin gin ye eee ee ee ee Ee eee 725 Daldoriiavnagashimae 229 es a ee ee ee eee 46 (Diapotamon! 22S ok See ee Be NO Se Se ee eee 476 TILLILO Sita, wena nce scree ea eRe arene SE SEE oe 2 IR ee 477 Hesperodiaptomusrcalijormnicnsis) a —————————————————————————————Ee 104 BUTE 11 pee ae ree ee er ee ee ee ee ee ee 76 GLBatrOSSGC) a= oe ee 8 es io eee eee ee ee 76 Bale eee ee ee ee ee ee a ee 84 IS CTLEL COS Cha at eee NE BOS ee ee 87 SE ORG Com pene sae ses ted Nes DP A eS ak eed eee 81 TEIn CE OMUSB AC 1 eae EF 483 Honchodaetyliiss ae a ee ee ee eee 464 INES SUG en an ee Ses ee ee 466 Miacrosynoplaxcflinitiy ea: see ee ee ee 321 CLT LUA KAM ee ro a ORI Nem NSSTEAE Es BCE LAD ARO Soe Sega CONES ee ei 321 ET LAT UC CLL Ce Ie aa lara Pe eno Be ee 318 BY LLP CLA G UAL = at a Nes ee Near = SE SIN a eee) aoe 318 IN GQTUTL OU CT C'S poe Sie Ne rT) Oe da 311 TTL OO TC Upendra oe Sle AE hh Ml AE oe eee ae 312 INC OD GLY ITY SUS tae sere wwe ese ee a ae ER cS ek a ON et es eee a 501 if CLD OMUUC ese ca eo ee ee re 502 Orectumusiamp lus. oe ee Fe Ee ee ee eee 112 Bas uiriStese7marK Wari) 2. 5 Sek ENN ad ee 2 i ee 471 ASR SIRI ANE gee oe ee eR ee 266 AE CTRCULES fame oe ew ase a See oN oe ae eal Pe 266 Karenwishnerag 2258 ea ee eee eee 266 Retrolisthessgentrudae, S228 ee ee eee 306 Prionalpheuswiayaritae: a ee ee ee ee 719 SELSTOWSUUUEUS a ee ee eee 453 Sime WawuDenrculinne 2a ee ee ee ee eee 512 Spelacomysissvillalobosi, == ee ee eee eee 97 Sundathelphusavboex. (meee ee Se ee eee 696 SOMOCC ie nara en ee See ee eee a ee ee a 701 PGI A i, ee a a A a eg 702 VCC ITE Ki gee ere ae are ee ee een ee 703 Insecta (Chactocan ace slaty e Siac a i OE eo a 332 COTA CII 1 Co pig ca NN i new RLS) Vs ah ed PIN eh, ele hn 334 LOT SUC GUL Fee sa Be Sn sas re SER RR De ee 334 Culoptilaxae@eria. ox Oa ee ee ee ee ee 448 TLMAY Oe a a re ci cre ee Ea ne ee 451 DGTT CAE a an ee Ed BE ee 448 LETTE, ea I ee 451 JOUO PO creak a Be I cole Sato eI eo 446 Dynomucllaaustralica) ct ee ee eee 336 GAD UULS tes BAL US a I I ee ac 735 CTL CHS USO TITUS, js ai Sol pe A 737 VILLE sh a Sc na 733 VOLUME 109, NUMBER 4 INC ET CAI ACCRLLESTRCLLUNT Cleese aan oe alice Wena nen eh cee rie ee ee A ee 329 | PY ROYGHUMEEY AS) AUCH A NCEE he ee Nr 331 CATITMOCATIACE HCO LI C'S ST amma ean esta anew WN recie era ola sid eas ae eh ee se 345 HEMICHORDATA EOCO SILOS SUS SAV COLCIIS, girs tet ecta oe ae ne Es Sec a eet ee 438 CHORDATA Pisces PN ciratlvennny| UAL 1Ay/ Ver ES ONa ieee tatea rat te ev eee EEE ee et oh aa a 420 Cociellathveeris tice yes eared NB ered eno Relient edie cael eae e se MNO ee ey col alll 27 FRCAL TREN US Viagra Ae Di A Bales ee ee eel ae Sell ee th ta 31 SONTCLLLCTIS US muerte es OOO ope ae: Secs ULE vet tl I NR ga eee oe 29 TP UG Vice HO OS ASS OSV: eT ee ee ees 354 Amphibia Eleutherodactylussl@unaster: exec ce ca ese lee ee 366 FED LLUS 171 Caer mice Bc eS) Se reat et eat A ae hs ae ee eee al 744 Mammalia ASH OCS W/Eter7t DOL Ct earner ek PE a eos, MU ee ee ee ee a a, ote 2 MAS TUUTUN SS CLET CLT LES tape ear ares SA ghee SN tert a cr a de TN Nh oe ere seek I 5 4 a . ’ aa 4 - 1 ee Fi el) ne ALL bac th tid, kta ye = . -< _ rr nal qi = ty : i f Y i * aivdhs mh , i i ’ u at ‘i ir i =o e peers oo i ma a i. | 1 Sj Titles ‘ a) eye ease Sy THE BIOLOGICAL SOCIETY OF WASHINGTON 1996-1997 Officers President: Stephen D. Cairns Secretary: Carole C. Baldwin President-elect: Richard P. Vari Treasurer: TY. Chad Walter Elected Council John 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: George C. Steyskal 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. Library subscriptions to the Proceedings are: $40.00 for USA and non-USA addresses. Non-USA members or subscribers may pay an ad- ditional $25.00 to receive the Proceedings by Air Mail. The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued sporadically) are available. Correspondence dealing with membership and subscriptions should be sent to the Biological Society of Washington, P.O. Box 1897, Lawrence, Kansas 66044, U.S.A. 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 the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, 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 109(1):1—9. 1996 New species of mammals from northern South America: Bats of the genera Histiotus Gervais and Lasiurus Gray (Chiroptera: Vespertilionidae) EQNTHSON A NV . MAY 01. 1996 Division of Mammals, National Museum of NatWral History, Smithsonian Institution, Washington, D.C. 20560NU.S AB R ARIES Charles O. Handley, Jr. Abstract.—Two strikingly differentiated species of northern South American Histiotus and Lasiurus have been long known, but until now have remained undescribed and unnamed. The new Histiotus (H. humboldti) is a relict species, most like H. montanus Philippi & Landbeck, but differing from it and all other Histiotus in small size (forearm <47 mm, maxillary toothrow length <5.7 mm), fragile skull, and weak dentition. Isolated populations occur at medium ele- vations (1500—2200 m) in the Coast Range, in the Mérida Andes, and on Cerro Neblina in Venezuela, and in the Central and Western Andes and upper Cauca Valley in Col6mbia. The new Lasiurus (L. atratus) is a Guayanan endemic known from Venezuela, Suriname, and French Guiana. Medium size (forearm 44-47 mm, max. toothrow |. 4.4—5.0 mm) and black wings relate it to L. varius Poeppig of Chile and L. castaneus Handley of Panama and Costa Rica; but very bright black and white underparts; small antorbital fossa; obsolete lach- rymal process, supraorbital ridge, and basial pits; and well-developed mastoid process distinguish it from both. Mammals and their ectoparasites were collected in Venezuela between 1965 and 1968 by the Smithsonian Venezuelan Pro- ject, supported in part by a contract (DA- 49-MD-2788) of the Medical Research and Development Command, Office of the Sur- geon General, U.S. Army. Numerous pa- pers have described the ectoparasites and mammals of the Project. Throughout these papers undescribed species of mammals have been referred to by alphabetical des- ignations. Some of these have been named subsequently by Handley & Ferris (1972), Handley & Gordon (1980), Handley (1984, 1987). This paper provides formal descrip- tions for insect-eating bats of the genera Histiotus Gervais and Lasiurus Gray. Material and Methods Measurements.—All measurements used in this paper are in millimeters and follow the directions of Kalko & Handley (1994). Cranial measurements were taken with the assistance of a binocular microscope, with dial calipers reading to 0.1 mm. Specimens are deposited in the following institutions: American Museum of Natural History, New York (AMNH); Estacion Biol6gica de Rancho Grande, Maracay (EBRG); Field Museum of Natural History, Chicago (FMNH); National Museum of Natural History, Washington (USNM); Universidad Central de Venezuela, Caracas (UCV). Systematics A new species of Leaf-eared bat, genus Histiotus Gervais One of the first bats netted by the Smith- sonian Venezuelan Project at its inception in July 1965, at Los Venados in the Coast Range overlooking Caracas, was a species 2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of Histiotus. This genus had not been found previously in Venezuela. In a few weeks we caught three more Histiotus at Los Venados and at the Hotel Humboldt higher up on the mountain. These represented a new species that I describe and name here. This species has been mentioned in the literature as Histiotus montanus colombiae Thomas by Tamsitt & Valdivieso (1966) from Cauca, Col6mbia, and as Histiotus sp. A by me (Handley 1976) from Distrito Federal, Venezuela, and by Gardner (1990) from Cerro Neblina, Venezuela. Histiotus humboldti, new species (Fig. 1, Table 1) Synonymy.— Histiotus montanus colombiae Tamsitt & Valdivieso, 1966:102 (not Histiotus co- lombiae Thomas, 1916). Holotype.—USNM 370968, adult female (lactating); skin, skull, and skeleton; col- lected 24 July 1965 by Charles O. Handley, Jr. Original number, SVP 00092. Type locality.—Los Venados, 4 km NNW Caracas, 10°32'N, 66°54’W, 1498 m, Distrito Federal, Venezuela. The holotype was caught 150 m west of park headquar- ters buildings in a mist net set across a Jeep trail in second-growth forest with thick un- derbrush. This area is classified as LOWER MONTANE humid forest (bh-MB) in the Holdridge system (Ewel and Madriz 1968). Etymology.—This impressive bat is named in honor of the great naturalist Al- exander von Humboldt who traveled widely in Venezuela in 1799 and 1800 and de- scribed many Venezuelan mammals (Hersh- kovitz 1987). Distribution.—Histiotus humboldti has a fragmented range, with apparently isolated populations in southwestern Col6mbia (on the lower eastern flanks of the Western An- des at El Tambo and Quisquio and near the head of the Cauca Valley at Popayan), in north-central Col6mbia in the northern part of the Central Andes (La Ceja and Pobla- do), in the Mérida Andes in western Ven- ezuela (near San Juan de Lagunillas), in the Coast Range in northern Venezuela (Los Venados and Pico Avila), and on Cerro Neblina in southern Venezuela. Elevational range, from 1498 m at Los Venados to 2217 m at La Ceja. Ecology.—Histiotus humboldti is a mon- tane species that occurs at medium eleva- tions, lower than H. montanus usually is found at this latitude. Specimens from the Coast Range in northern Venezuela were taken in moist, second-growth, evergreen forest; fairly tall at Los Venados, low and very dense at Hotel Humboldt on Pico Avi- la. Three were taken in forest trails and one was netted in a livestock pen where trees remained, but underbrush had been cleared (Handley 1976). On Cerro Neblina H. hum- boldti was taken in scrubby tepuyan vege- tation in open areas close to rocky sand hills (A. L. Gardner, pers. comm.). Diagnosis.—Histiotus humboldti can be recognized as a Histiotus by its enormous ears, plain (unornamented) snout, Eptesi- cus-like skull, and vespertilionid dentition. It can be distinguished from other Histiotus by its small size, delicate rostrum, fragile zygomata, inflated braincase, and small, weak dentition. Description.—Dorsal coloration bright tan to brown, darker where blackish hair bases show through; underparts buff with fuscous hair bases showing through. Ears very large (28—32); anterior lobe wide (4.3— 5.5) and forming a point where it folds; tra- gus relatively short and broad (9—11 X 3.5— 4.0); tibia and fingers relatively short (tib. 17.5—18.8, F2 39.1—40.6, F3 78.3—80.5, F4 62.6—65.6, F5 56.2—60.0). Skull (Fig. 1) fragile throughout; rostrum short (max. toothrow |. 5.3—5.6), narrow (max. br. 5.9— 6.1), and shallow; facial profile sharply dished; lachrymal ridge strongly developed; supraorbital region bulges, but is not ledged; braincase and area of postorbital constric- tion notably inflated; zygoma fragile, but with a large postorbital process; pterygoid processes thin and delicate. Teeth small and VOLUME 109, NUMBER 1 Fig. 1. Dorsal, ventral, and lateral views of the skull and lateral view of the mandible of Histiotus humboldu., USNM 560627, male, from Cerro Neblina, Amazonas, Venezuela. A. L. Gardner photograph. Scale 7:1. 4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1—Measurements (in millimeters) and mass (in grams) of adult Histiotus humboldti. USNM UCV FMNH 370968* J-03208 72340 Los Venados Cerro Neblina La Ceja Venezuela Venezuela Colombia Total length 109 107 — Tail vertebrae 50 50 — Hind foot (dry) 9 11 10 Ear from notch 30 28 — Forearm 45.7 46.8 46.8 Tibia 17.5 18.0 18.0 Calcar 26.3 23.0 — Mass — 11.5 — Greatest length 16.4 16.5 — Zygomatic breadth 9.6 9.8 — Postorbital breadth 4.7 4.5 — Braincase breadth 8.1 8.0 — Braincase depth 5.8 5.8 — Maxillary toothrow length 5.4 5.5 5.4 Postpalatal length 6.5 6.1 — Maxillary breadth 6.0 6.0 5.9 Canine breadth 4.2 4.2 4.1 * Holotype. weak, with low cusps; dental formula 2/3- 1/1-1/2-3/3 X 2 = 32. Comparisons.—Histiotus humboldti dif- fers in almost every detail from H. macrotis Poeppig and H. velatus I. Geoffroy. It most resembles H. montanus colombiae, a taxon with which it is sympatric in some areas, but these species differ in many details. Coloration, both dorsally and ventrally, is similar in both species. Ears of H. hum- boldti are proportionally about the same size as those of H. m. colombiae, but the anterior lobe is wider and is more pointed than in any H. montanus (not as extreme as in H. velatus, however). The tragus is short- er and broader than in any other Histiotus. The tibia is shorter than in any H. montanus and finger proportions resemble H. velatus more than they do H. m. colombiae. In fact, in all finger measurements, H. humboldti is smaller or averages smaller than H. m. co- lombiae. Skulls of Histiotus montanus and H. ma- crotus are similar to one another and quite different from H. velatus, but the skull of H. humboldti is so distinctive, that com- EBRG USNM USNM USNM FMNH [US370967] 370969 370970 560627 86719 Los Venados_ Pico Avila Pico Avila Cerro Neblina Popoyan Neneausle Venezuela Venezuela Venezuela Colombia 106 110 110 110 — S72) 48 47 51 — 9 10 9 11 11 32 30 31 29 —_— 45.5 45.7 45.5 46.9 46.0 IW/ES) 18.3 17.9 18.8 18.6 26.1 22.8 — 23.8 — — — — 9.5 — 16.0 16.3 16.4 16.9 — 9.2 9.5 9.5 9.4 — 4.5 4.4 4.5 4.5 — 7.8 8.3 7.9 8.0 — 5.6 5.6 5.8 6.2 — 5.5 5.3 Do) 5.6 5.4 6.3 6.4 6.6 6.6 — 6.0 5.9 6.1 6.0 6.1 4.2 4.1 4.1 4.3 4.2 pared with it, the skulls of the other three species are relatively similar to one another. In that comparison the skull of H. hum- boldti looks as though it might even rep- resent a different genus. In fact it bears a strong superficial resemblance to skulls of North American Plecotus E. Geoffroy, Idionycteris Anthony, and Euderma H. A\l- len. Compared with the other species of Histiotus, the skull of H. humboldti is more delicate throughout; braincase and postor- bital are much more inflated; rostrum is much shallower and narrower; dishing of facial profile is much more pronounced; lachrymal ridge is more developed, but the supraorbital ledge is undeveloped; zygo- mata are less flaring and much more fragile; pterygoid processes are more fragile; teeth are relatively tiny, very weak, and have lower cusps. Its relatively delicate skull and weak dentition indicate that Histiotus hum- boldti must have a softer diet than the other species of Histiotus. Remarks.—Linares (1973) reported a specimen of Histiotus in the Muséum Na- tional d’ Histoire Naturelle, Paris (MNHN), VOLUME 109, NUMBER 1 collected in 1894 by Bricefio Gabaldon, near Mérida, Venezuela. Linares regarded the specimen as quite different from Co- l6mbian and Ecuadorean H. montanus be- cause it had smaller canines and upper pre- molars, an enlarged postorbital process on the zygoma, and a dished facial profile. He thought some features of the specimen were reminiscent of H. velatus. However, be- cause of scant material and inadequate knowledge of variation in Histiotus, he ten- tatively identified the specimen as H. m. colombiae (not typical). Some characters that Linares ascribed to the Mérida specimen are characteristic of Histiotus humboldti. However, the suite of measurements of the Mérida specimen place it within the size range of H. m. col- ombiae, and show that it is apparently too large to be H. humboldti. | conclude that Linares was correct in identifying the Ven- ezuelan specimen in MNHN as H. montan- US. Recently Jestis Molinari (pers. comm.) has taken Histiotus humboldti (confirmed by measurements and photographs of the skull) adjacent to an extensive island of xer- ic vegetation at Tierra Negra, 1550 m, ca. 12 km S San Juan de Lagunillas, Estado Mérida, Venezuela. Thus, in Venezuela, Histiotus montanus is known in the Mérida Andes, while H. humboldti appears to have a fragmented distribution at medium ele- vations in the Coast Range, in the Mérida Andes, and on a Guayanan tepui, peripheral to the range of H. montanus. A similar re- lationship is observed in Colombia. This suggests to me that H. humboldti once had a more extensive, continuous range in the mountains of northern South America that has been overrun and fragmented by H. montanus. Specimens examined.—Colombia: Antio- Guias Iba Cela, |ice, 227 wil, 2 ale, (FMNH); Poblado, [ca. 1600 mJ], 1 alc. (AMNH). Cauca: Popayan, 1750 m, 1 alc. (FMNH). Venezuela: Amazonas: Cerro Neblina, Camp II, 2.8 km NE Pico Phelps, 1820 m, 1 skin & skull (USNM), 1 skin & 5 skull (UCV). Distrito Federal: Los Vena- dos, 4 km NNW Caracas, 1498 m, 2 skin, skull & skeleton (USNM); Pico Avila, 5 km NNE Caracas, 2092—2101 m, 1 skin, skull & skeleton (USMN), 1 skin & skull (EBRG). References to other specimens.—Colom- bia: Cauca: El Tambo, 1800 m, 1 skin & skull (Swedish Mus. Nat. Hist., Tamsitt & Valdivieso 1966); Quisquio, 1700 m, | skin & skull (Swedish Mus. Nat. Hist., Tamsitt & Valdivieso 1966). Published measure- ments (fa. 46.1, 47.2; max. toothrow lI. 5.4, 5.6) (Tamsitt & Valdivieso 1966) agree with Histiotus humboldti. A new species of Red Bat, genus Lasiurus Gray In collections of the Field Museum of Natural History, I found a specimen (FMNH 93235) of a strikingly beautiful red bat representing an undescribed species. It was collected in Suriname in 1961 by Harry Beatty. A short time later, in the collections of the Universidad Central de Venezuela in Caracas. I discovered two more specimens of the same species that had been collected in Bolivar state in Venezuela in 1962 by Juhani Ojasti. None of these specimens has been mentioned in the literature, but re- cently a specimen from French Guiana has been reported by Brosset & Charles-Dom- inique (1990) and by Masson & Cosson (1992). Lasiurus atratus, new species (Fig. 2, Table 2) Synonymy.— Lasiurus spec.? Brosset & Charles-Domi- nique, 1990:543. Lasiurus castaneus Masson & Cosson, 1992:476 (not Lasiurus castaneus Hand- ley, 1960). Holotype.-—FMNH 93235, adult male, skin and skull, collected 10 Feb 1961 by Harry A. Beatty. Type locality.—Kaiserberg Airstrip, Zuid 6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. River, Suriname. According to Stephens & Traylor (1985), this is a hilly region that had undisturbed lowland rainforest at the time of Beatty’s visit. It is located about 03°07'N, 56°27’'W, at an elevation of about 278 m. Etymology.—The Latin atratus, dressed Dorsal, ventral, and lateral views of the skull of the holotype of Lasiurus atratus, FMNH 93235, male, from Kaiserberg Airport, Zuid River, Suriname. Scott Steppan photograph. Scale 5.6:1. in black, refers to the black wing mem- branes, which lack the ornate finger outlines that characterize the common red bat (Lasi- urus blossevillii) of South and Central America. Distribution.—Known only from Saiil in southern French Guiana, from the type lo- VOLUME 109, NUMBER 1 ~ Table 2.—Measurements (in millimeters) of adult Lasiurus atratus. Venezuela Suriname Fr. Guiana El Dorado Imataca ~ Kaiserberg Saiil Masson & UCV UCV Ochoa Ochoa Ochoa Ochoa FMNH Cosson 5409 5410 2587 2588 3183 506 93235* (1992) 2 @ 2 3 ic} (c) Total length 112 116 — — — — — — Tail vertebrae 53 57 — — — — = — Hind foot (dry) 10 10 11 11 11 11 10 Ear from notch 13 13 — — — — 13 — Forearm 46.9 46.8 47.6 46.0 45.9 45.1 46.1 45.3 Tibia 21.8 19.5 20.8 21.1 20.1 20.2 19.2 — Calcar 14.1 13.6 16.5 16.5 13.9 14.9 12.7 — Greatest length 13.2 13.0 12.9 27 12.5 12.6 13.0 12.9 Zygomatic breadth 9.9 10.2 9.9 9.6 9.3 10.0 9.4 9.6 Postorbital breadth 4.3 4.3 4.1 4.3 4.1 4.1 4.8 4.2 Braincase breadth 7.9 7.8 7.6 7.9 7.7 Vall 7.7 7.6 Braincase depth 6.4 6.2 6.1 6.1 6.0 6.2 6.5 6.5 Maxillary toothrow length 4.9 5.0 4.9 4.7 4.4 4.8 4.9 4.8 Postpalatal length 5.9 6.1 Delf 5.8 5.5 5.7 5.8 5.7 Maxillary breadth 6.5 6.8 6.8 6.5 6.1 6.8 6.4 6.6 Canine breadth 5.4 5.7 5.5 53) 5.0 5.5 5.4 5.4 cality in southern Suriname, and from east- ern Venezuela (Km. 55 on the highway southeast of El Dorado, and in the Imataca Forest, ca. 28 km E Tumeremo, Bolivar). Elevational range, 100 m in Bolivar to 278 m at Kaiserberg Airstrip. Probably Lasiurus atratus is a Guayanan endemic. Ecology.—Localities in Suriname and Venezuela where this bat was collected in 1961 and 1962 were in undisturbed lowland rainforest, Tropical humid forest (bh-T) in the Holdridge classification (Ewel & Mad- riz 1968). The specimen from French Gui- ana was netted above a small stream on the border of heavy tropical humid forest and a cultivated clearing (Masson & Cosson 1992). Specimens taken in the Imataca For- est in Venezuela by José Ochoa, 1990— 1992, and by Ochoa and Elisabeth Kalko in 1993, were netted over water-filled roadside ditches in secondary forest. Diagnosis.—Lasiurus atratus is charac- terized by red dorsal coloration, black face, contrasting black and white chest, long black wings, medium size, small anteorbital pit, obsolete lachrymal process, slightly at- tenuated exoccipital process, well-devel- oped mastoid process, small median ante- rior mesoterygoid process, obsolete basial pits, and relatively large molars. Description.—A medium-sized Lasiurus (greatest length of skull 12.9—13.2) with long wings (forearm 45.3—46.9); dorsal col- oration bright rufous red, without white or black hair tips; median buffy band of hairs much wider than black basal and red distal bands; face black; chin reddish; throat, chest, and belly contrasting black and white or brown and white (hairs white tipped, with successive black, pale or dark brown, and black bands); prominent white humeral spot; flanks buffy; wings black or blackish, lacking ornamental outlines around fingers and forearm; ears tan; interfemoral mem- brane furred to or near distal edge. Skull (Fig. 2) with conventional shape of red bat group; rostrum broad (max. br. 6.1— 6.8), but very short (max. toothrow 1. 4.4— 5.0) and shallow, sloping sharply down- ward anteriorly; facial profile straight; braincase large and globose, tilted up from palatal plane; sagittal crest low; lambdoidal crest weak and incomplete; mastoid process well-developed; exoccipital process trian- 8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gular, with a somewhat attenuated tip; zy- goma weak, straight on dorsal edge; an- teorbital pit small; lachrymal process and supraorbital ridge obsolete; palate short (max. toothrow |. 4.4—5.0) and wide (max. br. 6.1—6.8); anterior median process of mesopterygoid fossa short and wide; basial pits poorly developed. Dental formula 1/3-1/1-2/2-3/3 xX 2 = 32; anterior upper premolar tiny, wedged between lingual borders of canine and P4; molars moderately large; M3 with fully de- veloped second commissure. Comparisons.—Three species of red bats occur in the Guayana Region. They are graded in size. Lasiurus egregius Peters is large (fa. 50.0, gr. 1. 15.7), L. atratus is me- dium (fa. 45.3—46.9, gr. 1. 12.9-13.2), and L. blossevillii Lesson & Gernot is small (fa. 36-41, gr. 1. 11.5—11.9). In the red phase (the only phase known in L. egregius and L. atratus) dorsal coloration is similar in all three—bright rufous red, with a wide buffy median band on hairs. The face is black in L. atratus; reddish or dusky in the other species. Coloration of the underparts varies widely in the three species—all red like the dorsum in L. egregius; speckled brown or gray and buff, moderately differentiated from the dorsum in L. blossevillii; sharply differentiated black and white in L. atratus. Wings are ornate, with fingers and forearm outlined with reddish in L. blossevillii; plain black in L. atratus and L. egregius. With respect to cranial features, L. atratus shares characters with L. egregius and L. blosse- villii. but it differs from both of these spe- cies in having the anteorbital pit small, the lachrymal process obsolete, and the supra- orbital process poorly developed. Like Lasiurus atratus, its geographically remote relatives, L. castaneus of Central America and L. varius of Chile, have black unornamented wings, but otherwise they are quite different from it. Both have short- er wings and strongly developed lachrymal and supraorbital processes and anteorbital pit. L. castaneus has totally different col- oration—blackish underparts, dark dorsum, and narrow median band on dorsal hairs, as well as more elevated braincase, well-de- veloped basial pits, and much reduced M3. Underparts of L. varius are uniform orange- buff, and it has much more robust molars; cranial characters relate it more closely to L. blossevillii. Specimens examined.—Suriname: Nick- arie: Kaiserberg Airstrip, Zuid River. [ca. 278 mJ], 1 skin and skull (FMNH). Vene- zuela: Bolivar: Km 55 on highway south of El Dorado, 100 m, 2 skins & skulls (UCV); Reserva Forestal Imataca (Unit 5), ca. 28 km E Tumeremo, 140-180 m, 1 skin & skull, 1 skin, skull & skeleton, 2 alcoholics with skulls, 7 alcoholics (collection of J. Ochoa), 2 alcoholics with skulls (USNM). Total 16. References to other specimens.—French Guiana: 4 km N Saiil, 03°40’N, 53°13’W (Masson & Cosson 1992). As described and measured by Masson and Cosson (1992), this specimen can be only Lasiurus atratus (see Table 2). Acknowledgments I am grateful to the curators who allowed me to borrow specimens and/or study col- lections in their care at the American Mu- seum of Natural History. British Museum of Natural History, Field Museum of Nat- ural History, and the Universidad Central de Venezuela. I especially thankful to Jests Molinari, Universidad de los Andes, Méri- da, who sent me photographs and measure- ments of a specimen of Histiotus humboldti he collected near Mérida; and to José Ochoa, Asociacion Venezolana para la Con- servacion de Areas Naturales (ACOANA), Caracas, who made a special effort to cap- ture Lasiurus atratus in Venezuela, loaned me what he caught, and donated specimens to the National Museum of Natural History. My thanks go to A. L. Gardner who has kindly let me reproduce his photographs of the skull of Histiotus humboldti, to Bruce Patterson who arranged for Scott Steppan to photograph the holotype of Lasiurus VOLUME 109, NUMBER 1 atratus, to George Venable who digitized and computer-enhanced the photographs, and to Darelyn Handley, who made the ta- bles and edited and word-processed the manuscript. I appreciate the patience of Al Gardner, Karl Koopman, Tom Munroe, and Don Wilson who read and commented on the manuscript. Literature Cited Brosset, A., & P. Charles-Dominique. 1990. The bats from French Guiana: a taxonomic, faunistic and ecological approach.—Mammalia 54(4):509— 560. Ewel, J. J., & A. Madriz. 1968. Zonas de vida de Venezuela. Ministerio de Agricultura y Cria. Caracas, 265 pp. Gardner, A. L. 1990. Two new mammals from south- ern Venezuela and comments on the affinities of the highland fauna of Cerro de la Neblina. Pp. 411-424 in K. H. Redford & J. E Eisenberg, eds., Advances in Neotropical mammalogy. Sandhill Crane Press, Gainesville, Florida, 614 Pp. Handley, C. O., Jr. 1976. Mammals of the Smithson- ian Venezuelan Project.—Brigham Young Uni- versity Science Bulletin, Biological Series 22(5):1-89. . 1984. New species of mammals from north- ern South America: A long-tongued bat, genus Anoura Gray.—Proceedings of the Biological Society of Washington 97:513—521. . 1987. New species of mammals from north- ern South America: Fruit-eating bats, genus Ar- tibeus Leach. Pp. 163-172 in B. D. Patterson & R. M. Timm, eds., Studies in Neotropical mam- malogy: essays in honor of Philip Hershkov- itz.—Fieldiana: Zoology, n.s. no. 39, Field Mu- seum of Natural History, Chicago, 506 pp. , & K. C. Ferris. 1972. Descriptions of new bats of the genus Vampyrops.—Proceedings of the Biological Society of Washington 84:519— 523. , & L. K. Gordon. 1980. New species of mam- mals from northern South America. Mouse pos- sums, genus Marmosa Gray. Pp. 65-72 in J. FE Eisenberg, ed., Vertebrate ecology in the north- ern Neotropics. Smithsonian Institution Press, Washington, 271 pp. Hershkovitz, P. 1987. A history of the Recent mam- malogy of the Neotropical region from 1492 to 1850. Pp. 11-98 in B. D. Patterson & R. M. Timm, eds., Studies in Neotropical mammalo- gy: essays in honor of Philip Hershkovitz.— Fieldiana: Zoology, n.s. no. 39, Field Museum of Natural History, Chicago, 506 pp. Kalko, E. K. V., & C. O. Handley, Jr. 1994. Evolution, biogeography, and description of a new species of fruit-eating bat, genus Artibeus Leach (1821), from Panama.—dZeitschrift ftir Saugetierkunde 59(5):257-273. Linares, O. J. 1973. Presence de 1’Oreillard d’ Ame- rique du sud dans les Andes Venezueliennes (Chiroptéres, Vespertilionidae).—Mammalia 37(3):433-438. Masson, D., & J. E Cosson. 1992. Cyttarops alecto (Emballonuridae) et Lasiurus castaneus (Ves- pertilionidae), Deux chiroptéres nouveaux pour la Guyane frangaise—Mammalia 56(3):475— 478. Stephens, L., & M. A. Traylor, Jr. 1985. Ornitholog- ical gazetteer of the Guianas.—Bird Depart- ment, Museum of Comparative Zoology, Har- vard University, Cambridge, Massachusetts, 121 pp. Tamsitt, J. R., & D. Valdivieso. 1966. Bats from Co- lombia in the Swedish Museum of Natural His- tory, Stockholm.—Mammalia 30(1):97—104. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):10—16. 1996 Identity and taxonomic status of the Mexican garter snake Thamnophis vicinus Smith, 1942 (Reptilia: Serpentes: Natricidae) Douglas A. Rossman Museum of Natural Science, Louisiana State University, Baton Rouge, Louisiana 70803, U.S.A. Abstract.—The nominal garter snake species Thamnophis vicinus is shown to be a localized color pattern morph of the wide-ranging T. cyrtopsis collaris; this morph is known to occur only in three populations in the Mexican state of Michoacan. Thamnophis vicinus was originally de- scribed (Smith 1942) from the vicinity of Morelia, Michoacan, on the basis of ten specimens differing from 7. cyrtopsis only in lacking a vertebral stripe and in having large dorsal spots arranged in three rows rather than four. Bogert & Oliver (1945), Milstead (1953), Duellman (1961), and Webb (1966) concluded that 7. vicinus is merely a color pattern morph of T. cyrtopsis (specifically of 7. cyrtopsis collaris in the current taxonomy) and not a distinct spe- cies. However, Webb (1978) reexamined most of the type series of 7. vicinus, and six additional specimens from other areas in Michoacan that combined pattern fea- tures of 7. vicinus and T. cyrtopsis collaris. He concluded that the taxonomic status of T. vicinus is uncertain; for that reason, he tentatively resurrected 7. vicinus as a sep- arate species. Materials and Methods Preserved specimens were borrowed from the: American Museum of Natural History (AMNH); California Academy of Sciences (CAS); private collection of E. A. Liner (EAL); Field Museum of Natural His- tory (FMNH); University of Kansas Mu- seum of Natural History (KU); Los Angeles County Museum of Natural History (LACM); Louisiana State University Mu- seum of Natural Science (LSUMZ); Muse- um of Comparative Zoology, Harvard Uni- versity (MCZ); Michigan State University Museum (MSU); Texas Cooperative Wild- life Collection, Texas A&M University (TCWC); Florida Museum of Natural His- tory, University of Florida (UF); University of Illinois Museum of Natural History (UIMNH); University of Michigan Muse- um of Zoology (UMMZ); and Collection of Vertebrates, University of Texas at Arling- ton (UTA). Four standard characters (num- bers of ventrals, subcaudals, and maxillary teeth; relative tail length) were recorded; the results are summarized in Table 1. Sev- eral aspects of color pattern also were noted and are detailed under Results and Discus- sion. Specimens examined include: Colima, LSUMZ 7846; Durango, MSU 4434-36; Guanajuato, CAS 5848; Guerrero, LACM 130112; Hidalgo, MCZ 11432, UMMZ 99085; Jalisco, MSU 9801, 9803 (2 spec.); Michoacan, FMNH 37116—22, 39058-61, 100098 (holotype of 7. vicinus), 126499— 504, LACM 65252, MCZ 56019, 131014, UIMNH 23414, 23435, UMMZ 102510, 104699, 112537, 112541, 119409-12, 121546, UTA R-6050-52; Oaxaca, AMNH 97889, 103091, 103100, 107001, EAL 1797, LACM. 130111-12, LSUMZ 7560, UF 11326—-27; Querétaro, TCWC 53068; Sinaloa, CAS 24077, 24082, KU 40349, 78923, 83413, LACM 130113, MSU 567; Zacatecas, UMMZ 118433. VOLUME 109, NUMBER 1 Table 1.—Variation in four meristic or mensural characters in Thamophis cyrtopsis collaris from Mexico. The Northern Sample includes specimens from northwestern Michoacan and from populations north of Michoacan; the “‘vicinus’’ Morph Sample includes specimens from populations in which any individuals exhibit a T. vicinus color pattern; and the Southern Sample includes specimens from Guerrero and Oaxaca. Ventral and subcaudal data are from Webb (pers. comm.) as well as specimens examined by the author. Values are given as mean + standard deviation (range of variation) and sample size. Southern sample “vicinus” morph sample Northern sample Character 152.8 + 4.50 (149-158) 4 150.2 + 3.17 (144-155) 19 159.9 + 2.95 (155-165) 11 153.1 + 3.94 (147-159) 16 158.1 + 3.65 (152-166) 34 153.8 + 3.59 (146-162) 50 97.8 = 6.50 (86-109) 27 91.9 + 5.93 (80-103) 28 3d 29 Ventrals 89.7 + 5.03 (85-95) 3 87.7 + 2.06 (84—90) 7 oo} 22 Subcaudals 81.4 + 2.56 (77-85) 14 79.9 + 3.73 (74-88) 10 25.5 + 0.68 (24.7—26.7) 8 24.7 + 1.15 (22.7—27.0) 10 27.5 + 1.73 (25-29) 4 26.5 + 1.29 (25-28) 4 (28.7) 1 25.8 + 0.83 (23.9-26.6) 9 28.7 28.3 + 1.12 (26.4—29.6) 8 27.7 + 1.60 (25.1-30.4) 7 26.7 + 0.76 (26-28) 7 Tail/Total Length (%) 29 3d Maxillary Teeth 26.7 + 0.76 (26-28) 7 25.7 + 0.82 (25-27) 10 29 11 Results and Discussion I examined 30 specimens from through- out Michoacan that could be identified as either 7. vicinus or T. cyrtopsis collaris. Partial or complete supression of the ver- tebral light stripe, accompanied by enlarge- ment and frequent fusion of two or more sets of dark dorsal spots or blotches on at least the anterior portion of the body, occurs in specimens from three separate areas in Michoacan: near Morelia, in the northeast; at Tancitaro, in the west-central region; and in the Sierra de Coalcoman, in the south- west (Fig. 1). The 7. vicinus dorsal pattern is by no means uniform in these areas. Most specimens from the vicinity of Morelia are preservative-darkened, hence details of pat- tern are sometimes difficult to discern. No indication of a vertebral stripe is evident on Six specimens (FMNH_ 126499-500, 126504; MCZ 56019; UIMNH 23435; UMMZ 102510), but there appears to be one present on UIMNH 23414, as well as traces of a fragmented, indistinct one on FMNH 100098 (which, ironically, is the holotype of 7. vicinus). In all examples, the anteriormost postnuchal spots are enlarged and fused to form transverse blotches that extend from the venter to the vertebral row and interrupt the lateral stripes. For a vary- ing distance thereafter, the lateral blotches alternate with the dorsolateral spots, the lat- ter being fused transversely across the back to form a single, vertebral row of blotches. This pattern (Fig. 2) contrasts with that usu- ally attributed to 7. cyrtopsis collaris, in which the vertebral light stripe is distinct, both it and the lateral stripes are uninter- rupted, there are no transverse bands on the neck, and the dorsolateral spots are not fused across the back (Fig. 2). The vertebral stripe is present on all eight specimens from the Sierra de Coalcoman, although it is relatively faint in all but one (UMMZ 104699). Six specimens from the vicinity of Dos Aguas (UMMZ 119411-12, 121546; UTA R-6050—-52) have the char- acteristic 7. vicinus blotch pattern anteriorly 12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 102 Fig. 1. 96 Map of southwestern Mexico showing the range of Thamnophis cyrtopsis (diagonal lines) and the areas where animals having a dorsal pattern characteristic of the “vicinus’’ morph have been collected (black blotches). (Fig. 3), whereas the two specimens (UMMZ 104699, 112537) from farther west (toward Coalcomén) have a T. cyrtop- sis collaris blotch pattern on the neck (Fig. i) Finally, a series of 11 specimens (FMNH 37116—-22, 39058—61) from Tancitaro, which lies about halfway between Morelia and the Sierra de Coalcoman, exhibit an ar- ray of dorsal patterns. All but one of the series (FMNH 37117) have a vertebral stri- pe, but it is faint in six of the specimens. In terms of the dorsal blotching, four spec- imens have a pattern more-or-less typical of T. vicinus, four are more characteristic of T. cyrtopsis collaris, and three appear to be intermediate (Fig. 4). None of the authors who have discussed T. vicinus has suggested that it differs from T. cyrtopsis collaris in any respect other than color pattern, although Webb (1978, Table 1) showed that specimens referred to the former have a lower mean number of subcaudals than the latter. The specimens I examined from within the “range” of JT. vi- cinus do, indeed, have fewer subcaudals (and a proportionally shorter tail) than spec- imens of 7. cyrtopsis collaris from north- western Michoacan (LACM 65252; MCZ 131014; UMMZ 119409-10) and farther VOLUME 109, NUMBER 1 13 Fig. 2. Upper: Adult male Thamnophis cyrtopsis collaris (UMMZ 102510) from Pino Gordo (= 37 km by road W Ciudad Hidalgo), Michoacan, Mexico, showing a dorsal pattern characteristic of the “‘vicinus’” morph. Lower: Subadult female Thamnophis cyrtopsis collaris (OMMZ 112541) from Uruapan Parque Nacional, Mi- choacan, Mexico, showing a dorsal pattern characteristic of the subspecies. 14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Re a - ~ a fries T - - ke i Se ee = sal Pa re :, Fig. 3. Upper: Adult female Thamnophis cyrtopsis collaris (JMMZ 121546) from Dos Aguas, Michoacan, Mexico, showing a dorsal blotch pattern characteristic of the “‘vicinus’’ morph. Lower: Subadult male Tham- nophis cyrtopsis collaris (OMMZ 104699) from the Cerro de los Havillos, near Coalcoman, Michoacan, Mexico, showing a dorsal pattern characteristic of the subspecies. VOLUME 109, NUMBER 1 15 Fig. 4. Upper: Adult female Thamnophis cyrtopsis collaris (FMNH 39059) from Tancitaro, Michoacan, Mexico, showing a dorsal blotch pattern characteristic of the “vicinus” morph. Lower: Adult female Thamnophis cyrtopsis collaris (FMNH 37122) from Tancitaro, Michoacan, Mexico, showing a dorsal pattern characteristic of the subspecies. 16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON north in the range of that subspecies, but the mean number of subcaudals is not sig- nificantly less than that of 7. cyrtopsis col- laris from Guerrero and Oaxaca to the south (Table 1). Interestingly, still farther south—in Chiapas and Guatemala—T. cyr- topsis collaris has subcaudal counts that ap- proach those of the population north of Mi- choacan (Webb 1982). Because TJ. vicinus does not differ men- surally or meristically from both geograph- ically adjacent populations of 7. cyrtopsis collaris, and differs only inconsistently in a few aspects of color pattern, it is apparent that 7. vicinus represents nothing more than a variant pattern morph—and a junior syn- onym—of T. cyrtopsis collaris. Analogous situations are not uncommon in other spe- cies of Thamnophis (see Rossman et al. 1996). Acknowledgments I thank the curatorial staffs at the insti- tutions mentioned under Materials and Methods for the loan of specimens in their care. I also am grateful to R. G. Webb for providing a set of his raw data on T. cyr- topsis collaris, M. Kleiner for photographic assistance, J. Boundy for making the map, and D. A. Good for constructive criticism of the manuscript. Literature Cited Bogert, C. M., & J. A. Oliver. 1945. A preliminary analysis of the herpetofauna of Sonora.—Bul- letin of the American Museum of Natural His- tory 83(6):297-426. Duellman, W. E. 1961. The amphibians and reptiles of Michoacan, Mexico.—University of Kansas Publications, Museum of Natural History 15(1): 1-148. Milstead, W. W. 1953. Geographic variation in the garter snake, Thamnophis cyrtopsis.—Texas Journal of Science 5(3):348-379. Rossman, D. A., N. B. Ford, & R. A. Seigel. 1996. The garter snakes: evolution and ecology. Uni- versity of Oklahoma Press, Norman (in press). Smith, H. M. 1942. The synonymy of the garter snakes (Thamnophis), with notes on Mexican and Central American species.—Zoologica 27: 97-123. Webb, R. G. 1966. Resurrected names for Mexican populations of black-necked garter snakes, Thamnophis cyrtopsis (Kennicott).—Tulane Studies in Zoology 13(2):55—70. . 1978. A review of the Mexican garter snake Thamnophis cyrtopsis postremus Smith with comments on Thamnophis vicinus Smith.— Contributions in Biology and Geology, Milwau- kee Public Museum (19):1—13. . 1982. Taxonomic status of some Neotropical garter snakes (genus Thamnophis).—Bulletin of the Southern California Academy of Sciences 81(1):26—-40. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):17—33. 1996 Review of the genus Cociella Whitley (Teleostei: Tear mele ere) with the description of three new species Leslie W. Knapp Department of Vertebrate Zoology, National Museum of Natural History, Smithsonain Institution, Washington, D.C. 20560, U.S.A. Abstract.—The genus Cociella Whitley is diagnosed as having vomerine teeth in two discrete patches, teeth in jaws not depressible, diagonal scale rows slanting downward above lateral line more numerous than lateral-line scales, lateral-line scale pores with a single canal opening to the exterior, iris lappet a simple lobe, side of head bicarinate, suborbital ridge bearing one spine under middle eye and one spine under rear margin of eye (additional spines may or may not be present posterior to eye), and upper preopercular spine distinctly longer than lower spines, bearing a small accessory spine on base. Cociella punctata (Cuvier) is removed from the synonymy C. crocodilus (Tilesius) and the occurence of possible intergrades between C. punctata and C. crocodilus is discussed. Young C. punctata appear to utilize the mangroves as nursury grounds. Three new species tentatively assigned to Cociella are described, C. heemstrai from off the west coast of southern Africa, C. somaliensis from off Oman and Somalia and C. hutchinsi from the Arafura Sea. A key to the species of Cociella is provided that primarily uses differences in arrangement of spines on the suborbital ridge, fin ray counts, number of gill rakers, and scale counts to separate the species. The status of many nominal genera of the Platycephalidae remains unclear. Relatively little has been published in this regard since extensive revision of Japanese flatheads by Matsubara & Ochiai (1955). Matsubara & Ochai (1955) synonymized Platycephalus punctatus (Cuvier in Cuvier & Valenci- ennes, 1829) under Platycephalus crocodi- lus (Tilesius, 1812), restricting the genus Cociella to C. crocodilus. The genus Co- ciella is here regarded as containing C. crocodilus, C. punctata, and three new spe- cies described below. This paper attempts to clarify the limits Cociella and compares features between Cociella and Ratabulus. The taxonomic significance of the pore morphology of the lateral-line scales in the Platycephalidae is well documented. Mat- subara and Ochiai (1955) found that differ- ences in pore structure of the scales were useful as generic characters in Cociella, Onigocia, Platycephalus and Rogadius. This was further corroborated by Hughes (1981, 1985) who, in addition to pore struc- ture, also included other features of lateral- line scale morphology in a comprehensive study of the flatheads that utilized scanning electron microscopy. The configuration of lateral-line scale pores is also considered important here. For example, the scale pores of Thysanophrys have two canals to the exterior and are Y-shaped; those of On- igocia also have two canals to the exterior and resemble short, stubby Ts. Other flat- head nominal genera with two canals in- clude Inegocia, Papillolabium, Rogadius, Sorsogona and Suggrundus. On the basis of having two pore canals, and with the di- agonal scale rows above the lateral line be- ing equal or nearly so to the number of lat- eral-line scales, these genera are removed from consideration here. 18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A second group of genera has lateral line scale pores with one canal to the exterior: Cociella; Elates; Grammoplites; Kumoco- cius; Leviprora; Platycephalus; and Rata- bulus. To separate these from Cociella, it is necessary to use additional characters. The condition found in Cociella is given in pa- rentheses. Vomerine teeth in a single patch is diagnostic for Platycephalus (two patch- es). A single elongate preopercular spine and six dorsal spines is unique to Elates (2— 3 preopercular spines, 9 dorsal spines). Scale rows slanting downward above the lateral line closely approximate the number of lateral line scales in Kumococius and Grammoplites (scale rows are more numer- ous than lateral line scales). In Leviprora, the type of iris lappet (or umbraculum) on the upper surface of the eye is finger-like or cirrose (Matsubara & Ochiai 1955:5, Fig. 2F) and the two upper preopercular spines are subequal (iris lappet a simple lobe, up- per propercular spine distinctly longest). Ratabulus shares many of the diagnostic features given here for Cociella but differs in having depressible teeth in the jaws, a greater number of spines on the suborbital ridge beneath the eye, more elongate later- al-line scales (Fig. 1A) and a smaller and more pointed iris lappet. Although there is some justification for placing Cociella in the synonymy of Ratabulus, such action would require additional evidence that is not available here. Drawings of lateral line scales from the species of Cociella, Ratabulus megace- phalus and Kumococius rodericiensis ap- pear in Fig. 1A—G. These ctenoid scales are small, rather uniformly rectangular, have well-developed radii, and pores that open to the exterior through a single slender canal. The canal of C. heemstrai is somewhat shorter than the canals in other species of Cociella. In Fig. 1A, Ratabulus megace- phalus, the pore and canal is quite similar to those of the other species shown but the scale is more slender and elongate. The scale from Kumococius rodericiensis (Fig. 1G) appears quite similar to those found in Cociella but the spine at the anterior margin of the pore is much more robust. Methods.—Counts follow procedures de- scribed by Hubbs & Lagler (1958:19—24) with the following exceptions: number of diagonal scale rows slanting downward (and backward) starting with the row near- est the anteriormost lateral line scale and ending with row nearest the posterior lateral line scale; number of interpelvic scales are counted in a straight line between the pelvic bases. Measurements (in mm) were taken as follows: interorbital width is the least bony width between the eyes; orbit diame- ter is taken from the lower rear margin of the orbit to the base of the preocular spine; snout length is the distance from the tip of snout to rear base of the preocular spine; head length is taken from the tip of the snout to the rear margin of the head; and standard length is the stright line distance from the tip of snout to the rear margin of the hypural plate. Counts and measure- ments were routinely taken from the left side (unless damaged) while gill rakers were counted on the right side. The material examined was from the following institu- tions (abbreviations in parentheses): Acad- emy of Natural Sciences of Philadelphia (ANSP); American Museum of Natural History, New York (AMNH); Australian Museum, Sydney (AMS); Bernice P. Bish- op Museum, Honolulu (BPBM); California Academy of Sciences, San Francisco (CAS, SU); Commonwealth Science and Industrial Research Organization, Hobart (CSIRO); Field Museum of Natural History, Chicago (FMNH); Hebrew University, Jerusalem (HUJ); Hokkaido University, Hakodate (HUMZ); J. L. B. Smith Institute of Ichthy- ology, Grahamstown (RUSD); Kanudi Fish- eries Research Station, Konedobu, Papua New Guinea (KFRS); Museum of Compar- ative Zoology, Harvard University (MCZ); Muséum National d’ Histoire Naturelle, Par- is (MNHN); Museum fiir Naturkunde der Universitat-Humboldt, Berlin (ZMB); Mu- seum of Zoology, University of Michigan, Ann Arbor (UMMZ); National Museum of VOLUME 109, NUMBER 1 19 / PRELRE KA re SUIT T Yd Fig. 1. Drawings of pored lateral line scales taken from right side (12th scale from front, scale length in parentheses) of species of Cociella and related flatheads: A, Ratabulus megacephalus, USNM 329510, 236 mm SL, G@ mm); B, Cociella somaliensis, USNM 326300, 216 mm SL, (3 mm); C, Cociella heemstrai, USNM 326629, 178 mm SL, (2.9 mm); D, Cociella crocodilus, USNM 329509, 261 mm SL, (3.6 mm); E, Cociella hutchinsi, USNM 327279, 240 mm SL, (4.3 mm); E Cociella punctata, USNM 327189, 216 mm SL, (4.0 mm); G, Kumococius rodericiensis, WAM P26206, 180 mm SL, (4.0 mm). Natural History, Washington D.C. (USNM); National Natuurhistorische Museum, Lei- den (RMNH); Natural History Museum, London (BMNH); Natural History Muse- um, Los Angeles County (LACM); Natur- historisches Museum, Vienna (NMW),; Northern Territory Museum of Arts & Sci- ences, Darwin (NTM); Royal Ontario Mu- seum, Toronto (ROM); South African Mu- seum, Cape Town (SAM); Western Austra- lian Museum, Perth (WAM); Zodlogische Museum, Universtiteit van Amsterdam 20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Interopercular flap (IOP), right side of Co- ciella punctata, USNM 329292, 111 mm SL. (ZMA); and Zoologisk Museum, Koben- havns Universitet, Copenhagen (ZMUC). Cociella Whitley, 1940 Cocius Jordan & Hubbs, 1925:286 (type species Platycephalus crocodilus Tile- sius, 1812, by original designation). Cociella Whitley, 1940:243 (substitute for Cocius Jordan & Hubbs, preoccupied). Diagnosis.—A genus of platycephalid fishes characterized by the following: ocular papillae absent; iris lappet a simple lobe (Figs. 3, 7) or slightly bilobed (latter may be an artifact of preservation); anterior nostril with an elongate dermal flap posteriorly; in- teropercular flap present or absent; pelvic-fin rays I, 4+1; anteriormost 1 to 19 lateral-line scales bearing small spines; scale rows slant- ing downward above lateral line more nu- merous than lateral-line scales; lateral-line scale pores with a single opening to the ex- terior; suborbital ridge bearing one spine un- der middle of eye and one spine under rear margin of eye, additional spines may or may not be present posterior to the eye; a single preocular spine; preopercular spines 2 or 3, uppermost bearing a small accessory spine on base; side of head bicarinate; teeth villi- form, caniniform or granular, in broad bands on tooth-bearing bones; and vomerine teeth in 2 discrete patches. Key to the species of Cociella 1A. Dorsal-fin rays and anal-fin rays usu- ally 11; total number of gill rakers on first archy5—8\ 0s. See eee y 1B. Dorsal-fin rays and anal-fin rays usu- ally 12; total number of gill rakers on first arch: 9-18). 4.4.4. 2.2 eee 4 Fig. 3. of C. crocodilus. Specimen from Kanaiwa, Japan, 285 mm SL; B, Pattern usually found in C. punctata. Specimen from the East China Sea, 340 mm SL. Color pattern variation in Cociella crocodilus (After Matsubara & Ochiai, 1955): A, Pattern typical VOLUME 109, NUMBER 1 2A. Suborbital ridge with 2 spines below eye, several spines behind eye; upper preopercular spine long, nearly reach- ing to opercular margin; preorbital spine slight or lacking a ee ee es C. hutchinsi, new species 2B. Suborbital ridge with 2 spines below eye, no spines behind eye; upper preo- percular spine shorter, reaching about half-way to opercular margin; preor- bital spine usually well developed ... 3 Diagonal scale rows slanting down- ward above lateral line 74-91 (X = 80.6); total gill rakers usually 7 (6-8); interopercular flap absent ..C. crocodilus. 3B. Diagonal scale rows slanting down- ward above lateral line 60-76 (X = 67.1); total gill rakers 5—6, usually 6; interopercular flap present ...C. punctata Total gill rakers 9-11 (X = 10.0); in- terpelvic scale count 16—24 (X = 20.6); caudal fin dusky, with small dark spots on upper half ..C. heemstrai, new species 4B. Total gill rakers 12-18 (X = 15.1); in- terpelvic scale count 25—35 (X = 29.9); caudal fin light, with large dark spots and horizontal streaks throughout 2 Ga C. somaliensis, new species SVAN 4A. Cociella crocodilus (Tilesius, 1812) Fig. 3A Platycephalus crocodilus Tilesius, 1812, pl. 59, fig. 2 (original description, type lo- cality, Nagasaki).—Cuvier in Cuv. & Val., 1829:256 (description taken from Tilesius).—Beaufort & Briggs, 1962: 159-161.—Burgess & Axelrod, 1971: 541, fig. 510.—Burgess & Axelrod, 1974:1006, fig. 302. Platycephalus punctatus.—Ginther, 1880: 66. Platycephalus inermis (not Silurus inermis Houttuyn, 1878) Jordan & Evermann, 1903:361. Thysanophrys crocodilus.—Jordan & Rich- ardson, 1908:638—640, fig. 4.—Jordan & Metz, 1913:54, fig. 49.—Jordan et al., 1913:286, fig. 235. Inegocia crocodilus.—Jordan & Thomp- son, 1914:279. 21 Cocius crocodilus.—Jordan & MWHubbs, ZS) 22%8) [Ic Cociella crocodila.—Matsubara & Ochiai, 1955:87—89, figs. 32, 33 (in part). Anonymous, 1975:214, pl. 208.—Masu- da et al., 1975:342, pl. 146, figs. EK G— Kyushin et al. (eds.), 1982:276, fig. 255.—Masuda et al. (eds.), 1984:322, pl. Ayes), was, (Cy ID). Inegocia crocodila.—Yomiyama & Abe, 1963:69, fig. 201. Material examined (63 specimens).—Ja- pan: UMMZ 183219 (5, 140-174) Niigita. UMMZ 183220 (1, 181) Nigita. SU 23650 (1, 285) Naoetsu. UMMZ 183218 (2, 240— 256) Namerikawa. UMMZ 183222 (1, 250) Namerikawa. FMNH 104717 (3, 142—204) Tokyo. FMNH 71864 (1, 193) Yokohama. MCZ 48821 (1, 140) Yenosima. MCZ 31160 (1, 126) Yenosima. UMMZ 198916 (1, 178) Toba. FMNH 58790 (1, 142) Toba. FMNH 57413 (5, 230-293) Osaka. USNM 151815 (, 239) Kobe. NMW 11179 (2, 332—403) Kobe. ZMB 18778 (2, 251-258) Kobe. USNM 10734 (4, 192-216) Wakan- oura. USNM 62317-18 (2, 112-175) Hi- roshima & Onomichi. UMMZ 183215 (1, 218) northern Kyushu. CAS 120708 (2, 241) Nagasaki. SU 13362 (1, 350) Naga- saki. ZMUC P80233 (1, 146) Nagasaki. Korea, Pusan: UMMZ 183216 (5, 133-— 206). UMMZ 198900 (2, 159-172). USNM 143416 (2, 126-174). FMNH 55763 (1, 251). China: SU 31261 (1, 242). ZMA 112.715 (1, 339) near Chin-huang-tao. MCZ 13789 (1, 253) Shanghai. SU 32766 (1, 218) Ting-hai. USNM 130414 (1, 227) Ningpo. USNM 327194 (1, 245) northern East China Sea. HUMZ 108647 (1, 308) East China Sea. USNM 329509 (1, 261) Dongxiang. UMMZ 198909 (1, 88) Fukien. Taiwan: CAS 30012 (1, 231) Formosa Strait. CAS 15234 (1, 218) Formosa Strait. CAS 107973 (1, 171) Chilung. Hong Kong: ZMUC P80193-194 (2, 80-85). Possible intergrades, C. crocodilus X C. punctata (5 specimens)—MNHN 05-222 (1, 170) Bay d’Along, Gulf of Tonkin. SU 22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Number of diagonal scale rows slanting downward above lateral line in the species of Cociella. 53-55 56-58 59-61 62-64 65-67 68-70 71-73 74-76 77-79 80-82 83-85 86-88 89-91 n x C. crocodilus 7 18 17 9 5 5 61 81.0 C. heemstrai 1 16 24 14 6 1 62 60.7 C. hutchinsi 6 14 5 1 26 62.9 C. punctata 2 1 I@ 29) 34 8923 6 111 67.9 C. somaliensis D; 12 11 7 2 34 59.9 14151 (1, 337) S.W. Kwangtung. HUMZ 109566 (1, 269) South China Sea, off Sa- rawak, Borneo. USNM 32924 (1, 394), Penghu Is., Taiwan. FMNH 47490 (1, 54) Singapore. Description.—(Mean values appear in parentheses). Dorsal-fin rays I, VIII, 10-11 (10.9), usually 11; anal-fin rays 10-12 (11.0), usually 11; pectoral-fin rays 19-22 (20.3); pored lateral line scales 51—60 (54.3), the anteriormost 1—19 (8.2) scales bearing weak spines; number of diagonal scale rows above lateral line slanting down- ward 74-91 (81.0); diagonal scale count 14-20 (16.7); interpelvic scales 28—46 (34.4); total gill rakers on first arch 6—8 (7.1) and branched caudal rays 10-12 (11.3). Least interorbital width going into greatest diameter of orbit 2.3—4.3 (2.9). Nape, opercle and cheek behind eye cov- ered with ctenoid scales; top of head and cheek below eye mostly naked, with a few embedded scales. Preorbital spine present; a pair of small nasal spines. Preopercular spines often two, sometimes three; upper- most longest, reaching about half-way to opercular margin. Infraorbital ridge usually smooth over anterior 4 of eye, bearing 5— 8 small spines posteriorly; suborbital ridge with one spine below middle of eye, a sec- ond spine below rear margin of eye, no spines behind eye. Lateral-line scale shown in Fig. 1D. Interopercular flap absent. Color in alcohol.—Five or six dark bands usually crossing back. Dark spots on dorsum numerous anteriorly (on head, including up- per surface of eye and on anterior trunk reaching below lateral line), more scattered and primarily above lateral line on posterior body. Spinous dorsal fin with broad dusky margin, clear area at base; second dorsal fin with dark spots on rays; anal-fin membranes dusky, rays pale; pectoral fin dusky below, with vertical rows of dark spots on upper half; pelvic fin dusky; caudal fin dusky, with a series of dark blotches or streaks that usu- ally form a broad marginal band. Remarks.—No type specimens are known for Cociella crocodilus. The species is based on an inconotype (Tilesius 1812: pl. 59, fig. 2) and a secondary description based on Ti- lesius given by Cuvier (in Cuvier & Valen- ciennes, 1829:256). A more complete de- scription appears in Matsubara & Ochiai (1955:87—89). Several characters effectively separating Cociella crocodilus and C. punc- tata include differences in the number of di- agonal scale rows slanting downward above the lateral line (Table 1), total numbers of gill rakers (Table 2) and presence or absence of a preopercular flap (Fig. 2). Most C. croco- dilus have from 77 to 91 scale rows (X = 81) while most C. punctata have 56 to 73 scale rows (X = 67.9). Nearly all of C. croc- odilus have 7 or 8 gill rakers (X = 7.1) while most C. punctata have 6 gill rakers or less (X = 6.0). The flap is absent in C. crocodilus but is usually well-developed in C. punctata (Fig. 2). This flap may be partially developed in young specimens and may be difficult to see in larger specimens that were poorly fixed or that have been preserved for many years. Matsubara & Ochiai (1955) illustrated variation in color pattern between a speci- men taken from off Kanaiwa, Ishikawa Pre- fecture, Japan (Fig. 3A) and one from the East China Sea (Fig. 3B). In general, large dark spots on the dorsum are typical for C. VOLUME 109, NUMBER 1 Table 2.—Total number of gill rakers on the first arch in the species of Cociella. 5 6 7 8 9 10 C. crocodilus 1 45 6 C. heemstrai 14 36 C. hutchinsi 15 3 C. punctata 1 84 5 C. somaliensis crocodilus (Fig. 3A), while C. punctata is characterized by having smaller and less prominent dark spots (Fig. 3B). Intergrades.—Possible intergrades be- tween C. crocodilus and C. punctata (Table 3) include three specimens from the north- ern South China Sea, one from off Sarawak, Borneo and one from Singapore (Fig. 4). Interopercular flaps present in some speci- mens from this area appear to be less strongly developed than is typical for C. punctata. A zone of intergradation may ex- tend into the East China Sea and other areas surrounding the South China Sea and it is possible that a few specimens identified by me as C. crocodilus or C. punctata are ac- tually intergrades. This could account for some of the overlap in characters for the two species as shown in Tables 1 and 2. Addi- tional specimens from the South China Sea are needed to clarify relationships between the two species. If integration is occurring between C. crocodilus and C. punctata, it must be very limited as both species appear to be maintaining separate character states. Distribution.—This species is found along the coast of China, Taiwan, Korea and southern Japan (Fig. 4). Limited data indicate that it is taken by trawling at depths from 50 to 90 m and attains a max- imum size of about 400 mm SL. Cociella punctata (Cuvier in Cuvier & Valenciennes, 1829) Figs. 2, 3B Platycephalus punctatus Cuvier in Cuvier & Valenciennes, 1829: 243 (original de- scription, type locality, Ceylon, Vanikoro Is.).—Gtnther, 1860:180.—Sauvage, 23 i2 19 i 15 16 17 16 n x a 71 63 ~=:10.0 18 6.2 90 6.0 1875:307, pl. 36, figs. 5, 5a.—Day, 1876: ZA Ys lls LX, 135 3), Platycephalus malabaricus Cuvier in Cu- vier & Valenciennes, 1829:245 (original description, type locality, Mahé).— Ginther, 1860 (in part): 181. Platycephalus quoyi Bleeker, 1856—1857: 206 (original description, type locality, Ternate and Amboina). Platycephalus fasciatus Ginther, 1872:397 (original description, type locality, Ma- nila Bay). Thysanophrys quoyi.—Fowler, 1927:289.— Herre, 1953:582. Platycephalus crocodilus.—Barnard, 1927: 933 (in part).—Smith, 1950:178 (Cn part)—Beaufort & Briggs, 1962:16i-— 162 (in part).—Fourmanoir, 1957:277 (in part)—Jones & Kumaran, 1980:644, fig. 550. Suggrundus hunti Fowler, 1937:244, figs. 249, 250 (original description, type lo- cality, Rayong, Siam). Grammoplites jacksoni Fowler, 1944:175, figs. 25, 26 (original description, type lo- cality, New Hebrides). Cocius crocodilus.—Herre, 1953:578. Thysanophrys punctatus.—Munro, 1955: 253, fig. 736 (fig. is C. crocodilus, after Jordan et al., 1913). Cociella quoyi.—Munro, 1967:528—529, ing, GPS. Cociella crocodila.—Matsubara & Ochiai, 1955:87—-89, figs. 32, 33 (mn part). Gloerfelt-Tarp & Kailola, 1984:121, figs. A, B.—Dor, 1984:89—90.—Knapp, 1984: no pagination (fig. is C. crocodilus, after Jordan et al., 1913).—Bianchi, 1985a:30 (fig. is C. crocodilus, after Jordan et al., 24 Table 3.—Possible intergrades between Cociella crocodilus and C. punctata. Singapore FMNH 47490 Borneo HUMZ 109566 Northern South China Sea SU 14515 USNM 329294 MNHN 05-222 Character 71 7 (left), 6 (right) Absent 74 76 71 73 Number of scale rows above lateral line Total gill rakers on first arch Absent on left, Present, Absent on left, Present, Development of interopercular flap trace on right Like Fig. 3B trace on right both sides Like Fig. 3A both sides Color pattern PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1913).—Bianchi, 1985b:28 (fig. is C. crocodilus, after Jordan et al., 1913).— Knapp, 1986:483, fig. 155.1.—Baranes & Golani, 1993:305, pl. 7, fig. 23. Material examined (123 specimens).— Syntypes: MNHN 6836 (1, 187) Voyage of Per6dn; MNHN 6851 (1, 209) Trincomalee, Ceylon; MNHN 5852 (1, 193) Vanikoro I. Taiwan: RUSI 38443 (1, 223) Tachi. Oki- nawa: USNM 75448 (1, 127) Naha. Phil- ippines: BMNH 1872.10.18.117 C1, 190) Manila. SU 39846 (1, 55) Manila. SU 9598 (1, 107) Cavite. SU 39021 (1, 122) Nasug- bu. USNM 99762 (1, 210) Leyte I. SU39020 (1, 203) Iloilo. SU 27219 (1, 170) Culion. SU 27218 (1, 190) Dumaguete. SU 29758 (1, 103) Dumaguete. CAS 81318 (©, 37-124) Dumaguete. USNM 329287 (1, 45) Dumaguete. USNM 329511 (1, 42) near Dumaguete. USNM 329512 (1, 79) near Dumaguete. SU 29756 (1, 141) Jolo I. Palau Islands: CAS 81319 (1, 97) Koror I. CAS 83122 (1, 130) Koror I. CAS 83121 (1, 182) Nardueis I. USNM 329288 (1, 243) Nardueis I. CAS 81320 (1, 208) Babelthuap I. USNM 329292 (2, 102-111) Babelthuap I. South Pacific: AMS I.17482-006 (1, 78) Guadacanal I. (1, 26) New Hebrides Is. Pa- pua New Guinea: KFRS F5629.01 (2, 190— 210) Port Moresby. CSIRO 1517 (2, 156— 161) Port Moresby. Indonesia: SU 13756 (1, 330) Manokwari. USNM 327189 (3, 62-216) Semei I. USNM 327289 ((4, 70- 146) Misoél I. USNM 325917 (5, 45-99) Kepuluan Aru, Borear I. WAM 27697.001 (1, 145) Tanimbar I. BPBM 19433 (1, 100) Ambon I. RMNH 5915 (3, 128-182) Am- bon & Ternate Is. ZMA 112.697 (2, 135—- 136) Obi Is. USNM 264806 (1, 72) Bali. USNM 264805 (1, 100) Bali. USNM 264794 (1, 194) Bali. BMNH 1984.1.1.65 (1, 270) Bali. NTM S.11127—042 (1, 116) Bali. NTM S.10733.014 (1, 195) Bali. Gulf of Thailand: CAS 81316 (1, 73) Ko Chang I. CAS 81317 (1, 227) Bangkok fish mar- ket. USNM 32928 (1, 157) Prachuap Kiri Khan. ANSP 62861 (1, 150) Siracha. ANSP. 68247 (1, 75) Rayong. Singapore: VOLUME 109, NUMBER 1 ANDAMAN , 1s. fy HR MALDIVE ; * 1s. 45 ai = cP SEYCHELLES 1S. ¢ Cociella somaliensis C. heemstrai C. hutchinsi C. crocodilus C. punctata Intergrades 2 Ld wf NICOBAR 4; Se ‘* 7 1s, =) ny “ed CEYLON " V | = ro a : Be MARSHALL ee. Migs, 6 Oh @ BidbRoes a bo hoe FiJ} is. ry = 18.130 Fig. 4. Distribution of Cociella crocodilus, C. heemstrai, C. hutchinsi, C. punctata, and C. somaliensis. SU 30809 (2, 61-76). NMW 11171 (2, 146). Malaysia: SU 27794 (1, 174) Sanda- kan. ZMUC P80196 (1, 210) Malacca. ZMUC P80195 (1, 109) Penang. ZMUC P80191 (2, 104-153) Penang. AMS B.5024 (1, 167) Penang. Ceylon: ROM 1878 (1, 130). USNM 327285 (1, 355) Colombo. In- dia: SU 37202 (1, ?) Andaman Is., Port Blair. USNM 327288 (1, 227) Cochin. USNM 329285 (1, 209) Cochin. SU 41735 (2, 136-142) Calicut. AMS B.8128 (1, 205) Malabar. MCZ 4287 (1, 151) Carnara. NMW 11720 (2, 155-232) Bombay. ANSP 101419 (1, 180) Bombay. USNM 327284 (1, 288) near Bombay. Pakistan: LACM 38126 (1, 222) Karachi. USNM 327286 (1, 267) near Karachi. AMS B.8130 (1, 241) Sind. Iran: USNM 327283 (1, 358) Gulf of Oman. Western Indian Ocean: USNM 326291 (1, 200) Zanzibar. MNHN 6848 (1, 220) Mahé. AMNH 88086 (1, 170) Mada- gascar. USNM 327287 (1, 346) Madagas- car. MNHN 1994.505 (1, 275) Madagascar. SU 31387 (3, 118-175) Durban. ANSP 55103 (2, 87-160) Durban. _BMNH 1921.3.1.51 C1, 151) Durban. Gulf of Aden: Uncataloged, J. M. Rose 296-004 (1, 180) Djibouti. Red Sea: NMW 11718 (1, 170) Ghalefca. RMNH 15955 (2, 197-219) Ka- maran. NMW 11719 (1, 162) Kamaran. BPBM 20382 (1, 195) Suakin. NMW 11167 (1, 164) Quseir. USNM 326280 (8, 129-185) Hurghada. HUJ 14019 (2, 303- 320) Gulf of Aqaba. Description. (Mean values appear in pa- rentheses).—Dorsal-fin rays IX or I, VIII, 10-12 (11.0); anal-fin rays 11-12 (11.0); pectoral-fin rays 19—22 (20.6), usually 20 or 21; pored lateral line scales 50—56 (53.7) usually 53 or 54, anteriormost 1—16 (5.5) bearing small spines; number of diagonal scale rows above the lateral line slanting downward 56-76 (67.9); diagonal scale count 10—17 (13.7); interpelvic scales 21— 41 (29.1); total gill rakers on first arch 5—7 (6.0); branched caudal rays 10—26 (11.3). Least interorbital width going into greatest diameter of orbit 1.8—6.0 (3.5). Nape cov- ered with ctenoid scales; top of head, oper- cle and cheek bear embedded scales. Pre- orbital spine present; a pair of small nasal spines. Preopercular spines usually three, sometimes two; uppermost longest, reach- ing about half-way to opercular margin. In- fraorbital ridge usually smooth over ante- rior % of eye, bearing 5—8 (small spines 26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 4.—Young stages of Cociella punctata associated with mangrove habitat. Locality SL (mm) Thailand, CAS 81316 3 Philippines, USNM 329511 42 Philippines, USNM 329512 719 Guadalcanal, AMS I.17482-006 78 Palau, USNM 329292 102-111 Indonesia, USNM 329517 45-98 posteriorly); suborbital ridge with one spine below middle of eye, a second spine below rear margin of eye, no spines behind eye. Lateral-line scale shown in Fig. 1E Intero- percular flap present. Color in alcohol.—Five or six dark bands usually crossing back. Numerous small dark spots on dorsum reaching below lateral line, more widely scattered posteri- orly. Spinous dorsal fin with broad marginal dark band, clear area at base; second dorsal fin with dark spots on rays; anal fin inter- radial membranes dusky, rays pale; pectoral fin dusky on lower half, spotted above (fin entirely spotted in some specimens from western Indian Ocean); pelvic fin dusky; caudal fin variable (entirely dusky in a few), usually with a broad dark marginal band or series of dark spots and horizontal streaks, basal area more or less pale. Remarks.—Young specimens of Cociella punctata appear to be closely associated with mangrove habitat (Table 4). The small- est typically have a broad dark saddle across the back in the area of the spinous dorsal-fin and a narrow saddle near the rear Depth of capture (m) Bottom type 0-0.9 rocky, sand, mud O-1.0 sand O-0.5 sand, silt ? ? 0-0.9 mud, sand, gravel, cobbles O-1.0 ? of the soft dorsal-fin (Fig. 5). In juvenile to adult C. punctata, five or six dark bands may cross the back (Fig. 3B) or in some, the bands may become obscure. A similar pattern is also found in juvenile to adult C. crocodilus. . One specimen of C. punctata (USNM 326291) from Zanzibar was found by East African Marine Fisheries Organization bi- ologists in the stomach contents of a Chwa- ka sole (species unknown, recorded as 370 mm in total length). Distribution.—This species is known from the Red Sea to South Africa, to Tai- wan, Indonesia and to the New Hebrides (Fig. 4). As it is common off Port Moresby, Papua New Guinea, I would expect C. punctata to be found in northern Australian waters but, to my knowledge, none have been taken to date. Perhaps competition from some of Australia’s endemic platyce- phalids has prevented C. punctata from be- coming established. It is frequently taken at shallow depths by seines or with ichthyo- cide and it has been taken by trawl at depths from 23—250 m. Two large specimens (303, Fig. 5. Young Cociella punctata from mangroves, USNM 329511, 42 mm SL; Negros, Philippines. VOLUME 109, NUMBER 1 320 mm SL) were taken in a trap at 300 m in the Gulf of Aqaba (Baranes & Golani 1993:305). Cociella heemstrai, new species Figs. 6, 7 Platycephalus malabaricus (non Cuvier).— Gilchrist & Thompson, 1909:253.—Fow- ler, 1925:255. Platycephalus tentaculatus (non Rup- pell).—Fowler, 1925:255. Platycephalus crocodilus (non Tilesius).— Barnard, 1927:933 (in part).—Fowler, 1934:488 (part)—Smith, 1950:378 (in part)—Fourmanoir, 1957:274 (Gn part). Cociella sp.—Knapp, 1986:483. Material examined.—Holotype, USNM 326281 (formerly RUSI 13761) (472 mm SL) Kenya, 02°38’S, 40°28’E, R/V Fridtjof Nansen, otter trawl, 280 m, 17 Dec 1980, Phillip C. Heemstra. Paratypes (68): Natal: SAM 10514 (Q, 232-245) Durban Bay. SAM 11877 (1, 203) Durban Bay. RUSI 1510 (2, 118-157) Durban. BMNH 1919.4.1.33 (1, 109) Dur- ban. ANSP 54935 (2, 63-136) Durban; 1931. SU 69736 (1, 118) Durban. SAM 10035 (1, 216) Natal. RUSI 16492 (2, 135—- 138) Natal; 1914-20. ANSP 77600 (1, 235) Tugela R. N of Durban. SAM 11876 @, 167—231) South Head, Tugela R., N of Dur- ban; 21 Jan 1901. RUSI 36880 (3, 86-198) Tugela Bank, N of Durban. RUSI 1510 (, 118-157) Durban. Mozambique: RUSI 10516 (1, 122) Inhaca Island near Louren¢go Marques; Aug 1948. USNM 326296 (1, 178) Polana near Lourengo Marques; Feb 1969. RMNH 25144 (1, 172) Lourencgo Marques; 5 Jul 1965. SAM 10822 (1, 215) Lourengo Marques; Jun 1920. SAM 26023 (1, 200) Lourengo Marques; Jun 1920. ANSP 77598 (1, 183) Delagoa Bay. USNM 326297 (1, 175) Delagoa Bay; 6 Feb 1969. USNM 326295 (2, 192—210) Delagoa Bay; 10 Feb 1969. SAM 16725 (1, 202) Delagoa Bay. BMNH 1922.2.9.29 (1, 175) Delagoa Bay. RUSI 39859 (5, 152-206) Maputo Bay (Delagoa Bay); 28 May 1992. USNM 27 Fig. 6. Holotype of Cociella heemstrai, USNM 326281, 172 mm SL; Kenya. 28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. 326294 (11, 83-144) Delagoa Bay, west side; 14 Feb 1969. USNM 326293 (8, 111-— 177) near Beira; 9 Oct 1964. Madagascar: MNHN B.2887 (4, 100-140) Nosy Be. USNM 326292 (3, 79-166) Northwest coast, Baie D’Amboro; 12 Feb 1964. USNM 303749 (4, 140-180) 12°42'12’S, 48°43'06"E; 11 Nov 1988. MNHN 1994.504 (1, 227) 25°03'S, 47°07'E; 12 Mar 1969. Diagnosis.—This species is distin- guished from others in the genus Cociella in having 9-11 gill rakers on the first arch (Table 2) and interpelvic scale count of 16— 24 (X = 20.6). It is further distinguished by the following combination of characters: second dorsal fin and anal fin with 12 rays; ratio of snout length divided by interorbital width ranging from 2.5—4.4 (X = 3.4); only two spines on suborbital ridge below and behind eye; and interopercular flap present. Description.—(Values for holotype given in parentheses). Dorsal-fin rays I, VIII or IX CU, VII), 11-12 (12), usually 12; anal- fin rays 11—13 (12), usually 12; pectoral-fin rays 19-22 (20), usually 20 or 21; pored Paratype of Cociella heemstrai, USNM 303749, 180 mm SL; Mozambique Channel. lateral line scales 52—55 (53), the anterior 3-19 (10) scales bearing weak spines; scale rows above lateral line slanting downward and backward 55—69 (64); diagonal scales 9-15 (12); interpelvic scales 16—24 (20); total gill rakers on first arch 9-11 (9), branched caudal rays 11—14 (11), usually 11 or 12. Measurements for the holotype and paratypes appear in Table 5. Least in- terorbital width going into greatest diameter of orbit 1.8—3.4 times (2.2), usually less than 2 times in specimens over 215 mm SL. Nape, opercle and cheek behind eye cov- ered with ctenoid scales; top of head and cheek below eye mostly naked, with a few embedded scales. Preorbital spine single; a pair of small, reclining nasal spines usually present; infraorbital ridge smooth anterior- ly, bearing 5—7 spines over posterior half of eye; suborbital ridge with one spine below middle of eye and a second spine near rear margin of eye; upper preopercular spine reaching nearly to opercular margin. Lat- eral-line scale shown in Fig. 1C. A narrow, elongate interopercular flap present. Table 5.—Proportional measurements of Cociella heemstrai expressed in thousandths of standard length. Number of specimens given in parentheses after range. Character Holotype Standard length (mm) 172.0 Head length 319.8 Snout length 95.9 Orbit diameter (greatest) 66.9 Interorbital width (least) 30.8 First dorsal spine length 22.1 Second dorsal spine length 130.2 Paratypes Range XG 63.0—245.0 (62) 149.1 301.5—355.0 (62) 333.1 79.8-111.8 (62) 98.6 60.2—82.9 (62) 73.4 24.0—35.9 (62) 29.6 15.0—36.7 (45) 24.5 94.7—166.5 (42) 124.3 VOLUME 109, NUMBER 1 29 Fig. 8. Color in alcohol.—Dorsum grayish or brownish, sometimes with a few small dark spots. Spinous dorsal-fin dusky, with a few large dark spots. Soft dorsal-fin bearing nu- merous large dark spots. Pelvic fin dusky. Pectoral and caudal fins dusky below, with small dark spots near upper margin. Color in life.—Specimens taken from Richards Bay and the Tugela Shelf off Na- tal were observed to have a bright yellow horizontal bar in the middle of the caudal fin. Live specimens seen at the Durban Aquarium had dark bands over the back and 4 or 5 pairs of white spots along the dorsum. The coloration pattern was some- what similar to that exhibited by live spec- imens of Platycephalus indicus of similar S1Ze. Distribution.—Cociella heemstrai is known from Durban, South Africa to Mom- basa, Kenya and Madagascar (Fig. 4). Com- mon in trawl catches at depths to 280 m., it is also taken by seining in shallow estu- aries such as Durban Bay. Etymology.—Named in honor of Phillip C. Heemstra, J. L. B. Smith Institute of Ich- thyology, who, over the years, has contrib- uted many specimens of flatheads to my studies. Cociella somaliensis, new species Fig. 8 Material examined.—Holotype, USNM 326300 (216 mm SL) Western Indian Ocean, Somalia, S of Ras Hafun, M/V Beinta Cruise 19, Sta. 13A, 10°13’'N, Holotype of Cociella somaliensis, USNM 326300, 216 mm SL; Somalia. 51°00'18"E, trawl, 30 m, 8 Feb 1987, Greg Small. Paratypes (38): Somalia: USNM 326299 (8, 199-280) same data as holotype. USNM 302847 (3, 228-290) 10°20'54’N, 51°15'06’E; 8 Feb 1987. USNM 302846 (1, 271 same data as USNM 302847. USNM 326298 (20, 198-272) 10°07'24’N, 51°31'12"E; 6 Feb 1987. BMNH 1993.11.5.1-—3 (3, 252-236) same data as USNM 326298. MNHN 1993- 0265 (1, 213) same data as USNM 326298. MNHN 1993-0266 (1, 236) same data as USNM 326298. Oman: USNM 326301 (1, D3) 2 le SYNE DO 3 Dec nl9s) Diagnosis.—A species tentatively as- signed to Cociella Whitley, it is distin- guished from other members of the genus in having 12—18 gill rakers on the first arch (Table 2) and the ratio of snout length di- vided by interorbital width ranging from 2.0—2.5 (X = 2.2). It is further distinguished by the following combination of characters: an interpelvic scale count of 25-35 (X = 29.9); second dorsal fin and anal fin each with 12 rays; only two spines on suborbital ridge below and behind eye; and interoper- cular flap present. Description. (Values for holotype given in parentheses).—Dorsal-fin rays I, VIII, 12; anal-fin rays 11-13 (12), usually 12; pectoral-fin rays 19—22 (20), usually 20 or 21; pored lateral line scales 52—55 (53), an- terior 6-15 (9) scales bearing weak spines; diagonal scale rows above lateral line slant- ing downward 54—66 (54); diagonal scale count 10—14 (13); interpelvic scales 25—35 30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 6.—Proportional measurements of Cociella somaliensis expressed in thousandths of standard length. Number of specimens given in parentheses after range. Character Holotype Standard length (mm) 216.0 Head length 318.1 Snout length 85.6 Orbit diameter (greatest) 65.7 Interorbital width (least) 38.0 First dorsal spine length 24.1 Second dorsal spine length 145.8 (30); total gill rakers on first arch 12-18 (17); branched caudal-fin rays 11-12 (11), usually 11. Measurements for holotype and paratypes appear in Table 6. Least interor- bital width going into greatest diameter of orbit 1.3—1.8 (1.7) times. Nape, opercle and cheek behind eye covered with ctenoid scales; top of head and cheek below eye mostly naked, with a few embedded scales. Single preocular and preorbital spine; nasal spines usually absent; infraorbital ridge smooth anteriorly, bearing 5—7 small spines over rear half of eye; suborbital ridge with one spine below middle of eye and a second spine near rear margin of eye; upper preo- percular spine curved slightly upward, reaching nearly to or slightly beyond oper- cular margin. Lateral-line scale shown in Fig. 1B. A narrow, elongate interopercular flap present. Color in alcohol.—Dorsum light tan, usually with small scattered dark spots. Spi- nous dorsal-fin dusky, with large dark spots. Soft dorsal-fin pale, with large dark spots. Paratypes Range Xx 199.0—290.0 (38) 241.4 296.9-328.0 (38) 310.6 84.4-93.8 (38) 88.2 55.4-67.4 (38) 62.5 36.4—44.7 (38) 39.6 13.7-28.3 (31) 20.6 119.6—162.0 (33) 146.5 Pectoral and pelvic fins dusky. Anal fin with pale rays, interradial membrane slight- ly dusky. Caudal fin pale, with bold dark spots and elongate bars throughout. Distribution.—This species is found off Somalia and Oman (Fig. 4). It has been taken in the trawl catch at depths from 30— 49 m. Etymology.—Named for the country of Somalia, off whose shores all but one of the known specimens were captured. Cociella hutchinsi, new species Fig. 9 ?Platycephalus malabaricus (non Cu- vier).—Ginther; 1880:41. Suggrundus sp. 1.—Gloerfelt-Tarp & Kail- ola; 1984:123, color fig.—Sainsbury, Kailola & Leyland; 1985:120, color fig. Material examined.—Holotype, CSIRO 1865 (206 mm SL) Australia, Arafura Sea, 09°16'S, 135°00'E; FRV Soela, trawl, 113 m, 27 Jun 1981. Paratypes (26): Arafura Fig. 9. Holotype of Cociella hutchinsi, CSIRO 1865, 206 mm SL; Arafura Sea (after Sainsbury et al. 1984). VOLUME 109, NUMBER 1 31 Table 7.—Proportional measurements of Cociella hutchinsi expressed in thousandths of standard length. Num- ber of specimens given in parentheses after range. Character Holotype Standard length (mm) 206.0 Head length 363.1 Snout length 109.7 Orbit diameter (greatest) 77.7 Interorbital width (least) 20.4 First dorsal spine length 14.6 Second dorsal spine length 112.6* * Indicates broken. Sea: BMNH 1879.5.14.222 (1, 192) 9°59'S, 139°42’E; 10 Sep 1874. AMS I.27847-020 (4, 148-259) 10°02’S, 133°58’E; 17 Nov 1980. AMS 1.21846-002 (2, 254-261) 09°38'S, 134°02’E; 17 Nov 1980. NTM S.11898-008 (1, 235) 10°18’S, 136°30'E; 24 Apr 1986. NTM S.12266-010 (1, 226) 10°20'S, 134°23’FE; 13 Jun 1987. NTM S.11957-005 (2, 175-230) 10°18'S, 134°08’E; 16 Oct 1986. NTM S.11613-017 (2, 173- 218) 10°15’S, 136°20’E; 10 Mar 1985. USNM 327179 (formerly NTM S.11621- 002) (1, 240) 10°08’S, 136°48’E; 17 Mar 1985. Timor Sea: CSIRO 2739 (1, 177) 13°46—48'S, 128°13-14’E; 28 Jun 1980. CSIRO 2740 (1, 175) same data as CSIRO 2739. CSIRO 2741 (1, 165) same data as CSIRO 2739. USNM 327190 (3, 159-165) same data as CSIRO 2739. WAM P30716- O01 (4, 142-184) 13°43.5'S, 128°38.6'E; 26 Dec 1969. USNM 326279 (2, 198-216) 12°04'S, 127°14-16’E; 29 Jun 1979. Diagnosis.—This species is distin- guished from other members of the genus Cociella in having 3—4 spines on the sub- orbital ridge under eye. It is further distin- guished by the following combination of characters: second dorsal fin and anal fin with 11 rays; 6—7 gill rakers on the first arch (Table 2); interpelvic scales 23-34 (X = 28.1); interopercular flap absent; and ra- tio of snout length divided by interorbital width ranging from 4.6 to 6.7 (X = 5.6). Description.—(Values for holotype given in parentheses). Dorsal-fin rays I, VII, I-I, Paratypes Range xX 142.0—261.0 (23) 203.4 351.4—393.8 (23) 373.5 107.1—126.3 (23) 114.1 76.2—90.3 (23) 82.4 17.6—25.1 (23) 20.8 10.6—20.4 (21) 15.4 110.0-142.9 (22) 128.7 Vill d, VIIl), 11; anal-fin rays 11; pectoral- fin rays 20—23 (22), usually 21 or 22; pored lateral line scales 51—55 (54), the anterior- most 2—8 (6) scales bearing weak spines; di- agonal scale rows above lateral line slanting downward 59-68 (63); diagonal scale count 11-14 (12); interpelvic scales 23-34 (24); total gill rakers on first arch 6—7 (6); branched caudal-fin rays 11-12 (12). Mea- surements for the holotype and paratypes ap- pear in Table 7. Least interorbital width go- ing into greatest diameter of orbit 3.2-5.0 (3.8). Nape, opercle and cheek behind eye covered with ctenoid scales, top of head and cheek below eye mostly naked, with few embedded scales. Preorbital spine slight or lacking, a pair of small nasal spines present. Infraorbital ridge usually smooth over ante- rior 4 of eye, bearing 7-10 small spines pos- teriorly; suborbital ridge with one spine be- low middle of eye, a second spine below rear margin of eye and 3—4 spines behind eye; upper preopercular spine reaching near- ly to opercular margin. Lateral-line scale shown in Fig. 1E). Interopercular flap ab- sent. Color in alcohol.—Dorsum brownish; venter white with brownish stippling, sparse on breast, more evident posteriorly. Spinous dorsal fin dusky, with a broad sub- marginal black band. Soft dorsal fin pale, with small brownish spots on rays. Anal fin pale, rays white. Pectoral fin dusky brown, with traces of vertical dark bands, lower margin white. Pelvic fin with whitish base, 32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON stippled with brown, with well-developed submarginal dark band. Caudal fin dusky, sometimes with a series of streaks forming- submarginal dark band. Distribution.—This is species is known from the Timor and Arafura Seas of Aus- tralia. It has been taken by trawling at depths from 39-108 m. Etymology.—Named in honor of J. Barry Hutchins, Western Australian Museum, who has provided substantial assistance to the author’s studies of Australian flatheads. Acknowledgments I am greatly indebted to the following in- dividuals for providing access to specimens and/or other assistance: Gerald R. Allen, Kunio Amaoka, M. Eric Anderson, Marie Louise Bauchot, Adam Ben-Tuvia, Marinus Boeseman, Eugenia B. Bohlke, M. Bou- gaardt; David C. Catania, Simon Chater, Barry Chernoff, Bruce B. Collette, Leonard J. V. Compagno, Martine Desoutter, Wil- liam N. Eschmeyer, Sean Fennessy, Jerome E Finan, Thomas Gloerfelt-Tarp, Daniel Golani, Alastair Graham, Cedric Goliath, Karsten E. Hartel, Phillip C. Heemstra, Bar- bara Herzig, Jean-Claude Hureau, J. Barry Hutchins, Tomio Iwamoto, Robert K. John- son, Patricia J. Kailola, Peter Last, Robert J. Lavenberg, Anthony D. Lewis, Nigel R. Merrett, Douglas W. Nelson, Gareth J. Nel- son, Jorgen Nielsen, Han Nissen, John R. Paxton, John E. Randall, Billy Ranchod, Margaret Rouse, Barry C. Russell, Mark Salotti, William G. Saul, Jeffrey A. Seigal, William E Smith-Vaniz, Pearl M. Sonoda, Arnold Sussumoto, Rex Williams, and Richard Winterbottom. The drawings were skillfully prepared by Penelope Hollen- sworth and Francis W. Zweifel. Harold E. Dougherty and Rafael Lemaitre took the fine photographs. Literature Cited Anonymous. 1975. Illustrations of Chinese marine fish- es. Institute of Oceanography, Academica Sinica & Nature Museum of Shanghai. Peoples Press of Shanghai, 240 pp., 230 pls. Baranes, A., & D. Golani. 1993. An annotated list of deep-sea fishes collected in the northern Red Sea.—Israel Journal Zoology 39(4):299-336. Barnard, K. H. 1927. A monograph of the marine fishes of South Africa. Part 2—Annals South African Museum 21(2):417—1065, pls. 18-28, figs. 19— 82% Bianchi, G. 1985a. Field guide. Commercial marine and brackish water species of Pakistan. FAO, Rome, 200 pp., 24 pls. . 1985b. Field guide. Commercial marine and brackish water species of Tanzania. FAO, Rome, 199 pp., 32 pls. Beaufort, L. FE de, & J. C. Briggs. 1962. The fishes of the Indo-Australian archipelago. E. J. Brill, Lei- den, vol. 11, 481 pp., 100 figs. Bleeker, P. 1856. Achtste bijdrage tot de kennis der ichthyologische fauna van Ternate. (1).—Natu- urkundig Tijdschrift voor Nederlandsch-Indie 12: 191-210. Burgess, W., & H. R. Axelrod. 1971. Pacific marine fishes. Book 2. T. E H. Publications, Inc. Ltd., Hong Kong: 282-560, 529 figs. ,& . 1974. Pacific marine fishes. Book 4. T. E H. Publications, Inc. Ltd., Hong Kong: 848-1110, 466 figs. Cuvier, G., & A. Valenciennes. 1829. Histoire naturelle des poissons. F G. Levrault, Paris, vol. 4, 518 pp. Day, E 1876. The fishes of India, being a natural his- tory of India, Burma and Ceylon. London. Part 2:169—368, pls. 41-78 (+51 A-C). Dor, M. 1984. Checklist of the fishes of the Red Sea. CLOFRES. The Israel Academy of Sciences and Humanities, Jerusalem, 437 pp. Fourmanoir, P 1957. Poissons Teleosteens des eux mal- gaches du Canal de Mozambique.—Mémoires de LInstitute Scientifique de Madagascar, Serie FE vol. 1:1—-316, 17 pls., 195 figs. Fowler, H. W. 1925. Fishes from Natal, Zululand and Portuguese East Africa.—Proceedings Academy of Natural Sciences Philadelphia 77:187—268, figs. 1—4. . 1927. Notes on the Philippine fishes in the col- lection of the Academy.—Proceedings Academy of Natural Sciences Philadelphia 79:255—297. . 1934. Fishes obtained by Mr. H. W. Bell-Mar- ley chiefly in Natal and Zululand in 1929 to 1932.—Proceedings Academy of Natural Sci- ences Philadelphia 86:405—514, 53 figs. 1937. Zoological results of the third De Schauensee Siamese expedition. Part 7. Fishes obtained in 1936.—Proceedings Academy of Natural Sciences Philadelphia 89:125—264. . 1944. Fishes obtained in the New Hebrides by Dr. Edward L. Jackson. Proceedings Academy of Natural Sciences Philadelphia 96:155—199. Gilchrist, J. D. E, & W. W. Thompson. 1909. Descrip- VOLUME 109, NUMBER 1 tions of fishes from the coast of Natal (Part 2). —Annals South African Museum 6(3):213—279. Gloerfelt-Tarp, T., & P. J. Kailola. 1984. Trawled fishes of southern Indonesia and northwestern Austra- lia. Australian Development Assistance Bureau, Australia, Directorate General of Fisheries, In- donesia, German Agency for Technical Cooper- ation, Federal Republic of Germany, 406 pp. Giinther, A. 1860. Catalogue of the Acanthopterygian fishes in the collection of the British Museum. 2. Squamipinnes, Cirrhitidae, Triglidae, Trachini- dae, Polynemidae, Sphraenidae, Trichiuridae, Scombridae, Carangidae, Xiphiidae. London, xxi + 548 pp. 1872. 56. Notice of some species of fishes from the Philippine Islands—Annals and Mag- azine of Natural History Series 4, 10:397—399. . 1880. Report on the shore fishes. In Zoology of the voyage of HMS “Challenger’”’.—Chal- lenger Reports, Zoology 1(6):1—82, 32 pls. Herre, A. W. 1953. Check list of Philippine fishes. Fish & Wildlife Service, U. S. Department of Interior, Research Report 20:977 pp. Houttuyn, M. 1782. Beschryving van eenige Japanse visschen, en andere zee-schepzelen. Verh. Holl. Maatsch. Wet. Haarlem 20 (2):311—350. Hubbs, C. L., & K. EK Lagler. 1958. Fishes of the Great Lakes Region.—Cranbrook Institute of Science Bulletin 26:213 pp. Hughes, D. R. 1981. Development and Organization of the Posterior Field of Ctenoid Scales in the Pla- tycephalidae——Copeia 1981(3):596—606. . 1985. Scale morphology and relationships of the flatheads (Pisces: Platycephalidae). Unpub- lished M.Sc. thesis, University of Sydney, Syd- ney, Australia. Jones, S., & M. Kumaran. 1980. Fishes of the Lacca- dive Archipelago. Mathrubhumi Press, Cochin, India, 760 pp., 603 figs. Jordan, D. S., & B. W. Evermann. 1903. Notes on a collection of fishes from the Island of Formo- sa.—Proceedings U.S. National Museum 25(1289):315-368, 29 figs. , & C. Hubbs. 1925. Record of fishes obtained by David Starr Jordan in Japan, 1922.—Memoirs Carnegie Museum 10(2):93-346. —, & C. W. Metz. 1913. A catalog of the fishes known from the waters of Korea.—Memoirs Carnegie Museum 6(1):1—65, Pls. 1—10. , & R. E. Richardson. 1908. A review of the flatheads, gurnards, and other mail-cheeked fish- es of the waters of Japan.—Proceedings of the U.S. National Museum 33(1581):629—670, 9 figs. , 9. Tanaka, & J. O. Snyder. 1913. A catalogue 33 of the fishes of Japan.—Journal College Science, Imperial University 23(1):497 pp., 369 figs. , & W. FE Thompson. 1914. Record of the fishes obtained in Japan in 1911.—Memoirs Carnegie Museum 6(4):205—213, 37 figs., pls. 24-92. Knapp, L. W. 1984. Family Platycephalidae. In W. Fischer & G. Bianchi, eds., FAO species iden- tification sheets for fishery purposes. Western Indian Ocean fishing area 51. FAO, Rome, vol. 3, no pagination. 1986. Family No. 155: Platycephalidae. In M. M. Smith & P. C. Heemstra, eds., Smiths’ sea fishes. Macmillan, Johannesburg, South Af- rica, 482—486, 12 figs., pls. 29-30. Kyushin, K., K. Amaoka, K. Nakaya, H. Ida, Y. Tan- ino, & T. Senta. 1982. Fishes of the South Chi- na Sea. Japan Marine Fishery Resource Re- search Center, Tokyo, 333 pp., 291 pls. Masuda, H., K. Amaoka, C. Araga, T. Uyeno, & T. Yoshino (eds.). 1984. The fishes of the Japa- nese Archipelago, 437 pp., 274 figs., 370 pls. , C. Araga, & T. Yoshino. 1975. Coastal fishes of southern Japan. Tokai University Press, To- kyo, 379 pp., 11 figs., 1445 pls. Matsubara, K., & A. Ochiai. 1955. A revision of the Japanese fishes of the family Platycephalidae (the flatheads)—Memoirs College Agriculture, Kyoto University 68: 110 pp., 33 figs., 3 pls. Munro, I. S. R. 1955. The marine and fresh water fishes of Ceylon. Department of External Af- fairs, Canberra, 349 pp., 56 pls. . 1967. The fishes of New Guinea. Department of Agriculture, Stock and Fisheries, Port Mores- by, 650 pp., 84 pls., 23 figs. Sainsbury, K., P. J. Kailola, & G. G. Leyland. 1985. Continental shelf fishes of northern and north- western Australia. Clouston & Hall and Peter Pownall Fisheries Information Services, Can- berra, 375 pp. Sauvage, H. E. 1875. Fishes. Jn Alfred Grandidier, Histoire physique, naturelle et politique de Madagascar, 1887-91. Paris, vol. 16, 543 pp., 61 pls., 525 figs. Smith, J. L. B. 1950. The sea fishes of Southern Af- rica. Central News Agency, Ltd., South Africa, 550 pp., 103 pls. Tilesius von Tilenau, W. G. 1812. “‘Atlas zur Reise um de Welt’? unternommen auf Behalf Seiner Kaiserlichen Majistae Alexander des Ersten auf den Schiffen NADESDA und NEVA unter dem Commande des Captains von Krusenstern. At- las. St. Petersburg, pl. 59, fig. 2. Tomiyama, I., & T: Abe. 1963. Pisces and Cyclosto- mata Jn Encyclopaedia Zoologica. Hokuryukan, Tokio, 342 pp, 1020 pls. Whitley, G. 1940. The Nomenclator Zoologicus and some new fish names.—Australian Naturalist 10(8):242—243. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):34-38. 1996 New locality records of freshwater decapods from México (Crustacea: Atyidae, Cambaridae, and Palaemonidae) Gabino A. Rodriguez-Almaraz and Ernesto Campos (GAR-A) Facultad de Ciencias Biolégicas, Universidad Aut6noma de Nuevo Leon, A.P. 105-E Ciudad Universitaria, San Nicolas de los Garza, Nuevo Leon, México; (EC) Facultad de Ciencias, Universidad Aut6énoma de Baja California, A.P. 2300, Ensenada, Baja California 22800, México Abstract.—Collections in the Mexican states of Nayarit, Nuevo Leon, Ta- maulipas, San Luis Potosi and Veracruz, provided significant new distribution records or data for seven freshwater decapods species. Information on each species is presented, including restricted synonymy, previously known distri- bution, new localities, and notes on the significance of the new data. Villalobos-Hiriart et al. (1993) reported that Mexican freshwater decapods com- prised about 132 species. Of these, many are known only from the type locality and, in several cases, knowledge on habitat is scarce or non-existent. Rapid changes in habitat conditions, introduction of non-na- tive species, and decline in abundance of some elements of the original crustacean decapod fauna in northern México (see Contreras-Balderas 1991; Rodriguez-Al- maraz & Campos 1994) prompted us to up- date the diversity, ranges, and habitat of the 22 species reported from that region. Col- lections in selected freshwater habitats in the states of Nayarit, Nuevo Leén, Tamau- lipas, San Luis Potosi, and Veracruz pro- duced significant new distribution data for seven freshwater decapods species one Atyidae, one Cambaridae and five Palae- monidae). Abbreviations used are: UANL, Arthropoda Collection, Facultad de Cien- cias Biologicas, Universidad Autonoma de Nuevo Leé6n; UABC, Invertebrate Collec- tion, Facultad de Ciencias, Universidad Au- tonoma de Baja California; IBUNAM-EM, Crustacean Collection, Instituto de Biolo- gia, Universidad Nacional Aut6noma de México. For each species listed, a restricted synonymy is provided and includes original description or redescription, changes in ge- neric assignment, and known distribution. Family Atyidae Potimirim mexicana (De Saussure, 1857) Potimirim mexicana.—Villalobos-Figueroa, 1959:295-313, pl. VI, figs. 29-38, pl. VII, figs. 39-50, pl. VIII, figs. 51-54, pl. IX, figs. 55, 56, map. 2, 3.—1982:217, 223.—Hart, 1961:67, 76.—Smalley, 1963: 177, 179, 181.—Villalobos-Hiriart et al., 1993:281. Known distribution.—Meéxico. This spe- cies is found near the sea in the hydrolog- ical system of the Gulf of México slope, from Rio Soto la Marina basin, Tamaulipas, to Rio Grijalva basin, Tabasco, and rivers emptying in Laguna del Carmen, Campe- che. Central America. Honduras, Costa Rica. Antilles. Cuba, Jamaica and Puerto Rico (Villalobos-Figueroa 1982). Material examined.—21 females, Estero de Canala, Guayabitos, Nayarit, 21°05'N, 105°15'W, 1 Jun 1995 (UANL). 8 males, 13 females, Camino a las Varas, Nayarit, 21°10’N, 105°14'’W, 2 Jun 1995. 4 males, Rio Coy (Rio Choy) [Rio Panuco basin], 28 km south of Valles city, San Luis Potosi, 21°51'N, 98°56'W, 27 Aug 1994 (UANL). 3 males (topotypes, IBUNAM-EM uncat.), VOLUME 109, NUMBER 1 M. A. Quevedo Park, Playa Norte, Vera- cruz, Veracruz, 19°14’N, 96°08'W, 1958. Remarks.—Our specimens agree with Villalobos-Figueroa’s (1959) description except that we found up to 15 uncinuli on the appendix interna and, like in P. poti- mirim (Miller), a plumose seta close to it. Even so, morphometric and shape features recorded by Villalobos-Figueroa allow easy separation of these two species. This record represents the farthest inland finding of P. mexicana in the Gulf of México slope, ap- proximately 250 km upstream in the Rio Panuco-Rio Coy basin, and is the first re- port of this species in the Mexican Pacific slope. Family Cambaridae Procambarus roberti Villalobos-Figueroa & Hobbs, 1974 Procambarus (Pennides) roberti Villalo- bos-Figueroa & Hobbs, 1974:8, fig. 2.— Hobbs, 1989:74. Known distribution and habitat.—Ditch from La Media Luna, 4.8 miles (7.7 km) south of Rio Verde (on highway to Pedro Montoya) and 2.5 miles (4 km) west on dirt road to Mina El] Refugio, San Luis Potosi (type locality). In streams. Material examined.—1 male I, 3 fe- males, 6 juvenile males, 7 juvenile females, El Venado, San Luis Potosi (151 northwest of San Luis Potosi city and 105 km south- west of Matehuala city, San Luis Potosi), 22°58'N, 101°04’W, Feb 1994 (UANL). Remarks.—The habitat of Procambarus roberti at El Venado is a spring-pool of ap- proximately 20 m? with clear water and a rocky bottom. Only scarce grasses were ob- served in the margin of the pool amongst which the crayfishes were collected. Ostra- cods of the genus Darwinula and Cypron- odopsis were in the sediment of the bottle where the crayfishes were fixed; however, the entocytherid symbiont, Ankylocythere barbouri Villalobos-Figueroa & Hobbs, 1974, was not found. It is not clear how P. roberti moved 160 km northwest from La 35 Media Luna to El Venado since no super- ficial aquatic drainage system connects these localities. An hypogean dispersion or human introduction are two possible expla- nations. Family Palaemonidae Macrobrachium acanthurus (Wiegmann, 1836) Palaemon acanthurus Wiegmann, 1836: 150. Macrobrachium acanthurus, Hedgpeth, 1949:30, figs. la, 2, 5.—Holthuis, 1952: 45-53, pl. 8, 9, figs. a, b.—Villalobos- Figueroa, 1982:217, 224.—Williams, 1984:66—-68, figs. 44, 45.—Markham et al., 1990:419. Known distribution and habitat.—From North Carolina, U.S.A. to Brazil (Holthuis 1952; Williams 1984). In México it ranges from southern Tamaulipas to Quintana Roo (Markham et al. 1990). The species lives in fresh or sometimes brackish water, and gen- erally is not found far inland (Holthuis 1952). Material examined.—3 males, Rio Ra- mos [San Juan basin], municipality of Al- lende, Nuevo Le6én, 25°16’N, 100°00’W, 15 Apr 1992 (UANL). Remarks.—This is the first record of M. acanthurus in the state of Nuevo Leon and represents the farthest inland finding of this species in México, approximately 360 km upstream in the Rio Bravo-Rio San Juan ba- sin. Macrobrachium hobbsi Nates-Rodriguez & Villalobos-Hiriart, 1990 Macrobrachium hobbsi Nates-Rodriguez & Villalobos-Hiriart, 1990:7, Fig. 3.—Vil- lalobos-Hiriart et al., 1993:285. Know distribution.—Los Tuxtlas area, southern Veracruz; Arriaga and Tonala, Chiapas; Rio Ostuta, Oaxaca, and Rio Mur- ga near Petatlan, Guerrero; presumably to 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Nayarit (Nates-Rodriguez & Villalobos- Hiriart 1990; Villalobos-Hiriart 1993). Material examined.—4 males, 1 female, 20 juveniles, Rio Coy (=Rio Choy), 28 km south of Ciudad Valles, San Luis Potosi, Apr, Jun 1994 (UANL); 1 male, 1 female, Rio Huichihuayan, 70 km south of Ciudad Valles, San Luis Potosi, 21°32’N, 98°56'W, Apr 1994 (UANL); 7 juveniles, small pool, km 160 on the highway Tuxpam-Tampico, (30 km south of Panuco [Ozuluama]), Ve- racruz, 21°51’N, 97°48'W, Nov 1993 (UANL); 3 juveniles, small pool, km 240 on highway Poza Rica-Tuxpam, Veracruz, 20°45'N, 97°31'W, Nov 1993 (UANL). Remarks.—The presence of M. hobbsi in northern Veracruz and San Luis Potosi rep- resents a northern range extension of ap- proximately 850 km. It is remarkable that juveniles of M. hobbsi were found in the same habitat with Palaemonetes mexicanus Strenth, at Rio Coy, San Luis Potosi. This does not agree with Strenth’s (1976) con- clusion that juveniles of Macrobrachium competitively exclude Palaemonetes. Macrobrachium olfersi (Wiegmann, 1836) Palaemon olfersii Wiegmann, 1836:150. Macrobrachium olfersi.—Holthuis, 1952: 95-103, pl. 24, pl. 25, figs. a, b—Hedg- peth, 1949:35, figs. 1d, 4, 5.—Villalobos- Figueroa, 1967:167—171.—Villalobos- Hiriart, 1993:285. Known distribution.—Western Atlantic slope. Lower Cape Fear River near South- port, North Carolina; Florida; Louisiana; Texas; and southern Veracruz, México to Santa Catarina, Brazil (Holthuis 1952). Eastern Pacific slope. From Chiapas to Nayarit, México (Villalobos-Hiriart et al. 1993»). Material examined.— | male, 3 juveniles, Rio Limon, 8 km north of Ciudad Mante, Tamaulipas, 22°49'N, 98°56’W, 12 Jun 1994 (UANL). Remarks.—The identification of M. ol- fersi was possible by comparing the adult male above noted with the description and figures provided by Holthius (1952). The shrimps were collected in a shallow water area (15 cm depth) of the Rio Lim6n, amongst aquatic vegetation. This records represent the farthest inland finding of this species in México, approximately 205 km upstream in the Rio Guayalejo-Rio Limon basin. Palaemonetes kadiakensis Rathbun, 1902 Palaemonetes kadiakensis Rathbun, 1902: 93.—Strenth, 1976:2.—1994:91. Palaemonetes (Palaemonetes) kadiakensis, Holthuis, 1949:92.—1952:212, pl. 51, figs. k—n, pl. 52, figs. a, b—Smalley, 1964: 231.—Flemming, 1969:444.—Villalobos- Figueroa & Hobbs, 1974:15, fig. 8b. Known distribution.—This species is widespread in fresh-water habitats of Cen- tral U.S.A., including the shores of lakes Ontario, Erie, and Michigan, the Mississip- pi basin, and the basin of several rivers emptying east and west of the Mississippi river in the Gulf of México (Holthuis 1952). The only Mexican record is that of Creaser (1932) for the state of Nuevo Leon, northeast México (Strenth 1976). Material examined.—4 males, 5 females (1 ovigerous), Rio Sabinas Hidalgo, Valle- cillos, Nuevo Leén, 26°39'N, 99°59'W, 20 Apr 1979 (UANL). Tamaulipas: 21 males, 15 females, 13 Mar 1981 (UABC); 1 male, 3 ovigerous females, 22 Oct 1981; 1 male, 4 ovigerous females, 22 Nov 1981; 1 male, 6 females, 30 Jun 1982, Rio Alamo, Cd. Mier, 26°25'34”N, 99°06'41"W; 1 male, 4 females (2 ovigerous), 23 Oct 1983, Presa Falcon, Miguel Aleman, 26°39'N, 99°12'W; 1 male, 2 females, 30 Mar 1983; 1 male, 4 females, 14 Mar 1987; Rio Bravo, Mata- moros 25°50'N, 97°24'W (UANL). Remarks.—Based on the present record we believe that the material reported by Creaser (1932) from Nuevo Leon came from the Rio Alamo-Rio Salado basin (northern Nuevo Le6én), which empties into the Rio Bravo del Norte (Rio Grande) in VOLUME 109, NUMBER 1 the state of Tamaulipas. Previous collecting efforts in Central Nuevo Leon (Rio San Juan basin) failed to produce specimens of P. kadiakensis. The material of the Rio Alamo (13 Mar 1981) was collected in shallow (<I m depth) and clear-greenish water, the bottom was muddy and the aquatic vegetation in- cluded Chara sp. Additional information on the habitat is noted in Ruiz-Campos et al. (1985). Palaemonetes mexicanus Strenth, 1976 Palaemonetes mexicanus Strenth, 1976:7, fig. 3.—1994:291. Known distribution and habitat.—Small spring-fed river, 15 miles (24 km) west of Ciudad Valles, San Luis Potosi (type local- ity). Material examined.—7 males, 20 fe- males (5 ovigerous), Rio Coy, 28 km south of Ciudad Valles, San Luis-Potosi, Apr, Jun 1994 (UANL). Remarks.—This is apparently the first record of P. mexicanus since its original de- scription by Strenth (1976). The new local- ity 1s about 50 km southeast of the type locality. Hobbs & Hobbs (1989) reported P. mexicanus to Mante river (Ciudad Mante, Tamaulipas); however, Strenth (1994) pointed out that Hobbs & Hobbs’ speci- mens are of P. hobbsi Strenth, 1994. Five ovigerous females (cl. 11.4—18.5 mm) were carrying 13 to 31 eggs having a diameter of 0.89-1.12 mm (X = 0.99) by 1S=14) om C& = 125), Acknowledgments Financial support for a short-term visit to the Crustacean Collection, Instituto de Biol- ogia, Universidad Nacional Aut6noma de México (UNAM) was provided by Direc- cion de Intercambio Académico, UNAM to GAR-A. Our great appreciation is given to J. E Fitzpatrick Jr., H. H. Hobbs III, N. E. Strenth and J. L. Villalobos-Hiriart for their criticism and helpful comments on the 37 manuscript. EC is a fellow of the ““Progra- ma de Estimulo al personal Académico 94/ 95”’ of the Universidad Autonoma de Baja California (UABC). This is contribution number 4 of the program “‘Crustaceos De- capodos (formerly Cambarinos) del Norte de México” of the UABC and Universidad Autonoma de Nuevo Leon. Literature Cited Contreras-Balderas, S. 1991. Conservation of Mexi- can freshwater fishes: some protected sites and species and recent federal legislation. Pp. 191— 197 in W. L. Minckley and J. E. Deacon, eds., Battle against extinction: native fish manage- ment in the American West. University of Ari- zona Press, Tucson, Arizona. De Saussure, H. 1857. Diagnoses de quelques Crus- tacés nouveaux de 1’Amérique tropical.—Re- vue et Magasin de Zoologie pure et appliquée 2(9):501—S05. Creaser, E. P. 1932. The decapod crustaceans of Wis- consin.—Transactions of the Wisconsin Acad- emy of Sciences, Arts and Letters 27:321—338. Flemming, L. E. 1969. Use of male external genitalic details as taxonomic characters in some species of Palaemonetes (Decapoda, Palaemonidae).— Proceedings of the Biological Society of Wash- ington 82:443—452. Hart, C. W. 1961. The freshwater shrimps (Atyidae and Palaemonidae) of Jamaica, W.I. with a dis- cussion of their relation to the ancient geogra- phy of the Western Caribbean area.—Proceed- ings of the Academy of Natural Sciences of Philadelphia 113(4):61—80. Hedgpeth, J. W. 1949. The North American species of Machrobrachium (River shrimps).—Texas Journal of Science 1:28-38. Hobbs, H. H. Jr. 1989. An illustrated checklist of the American crayfishes (Decapoda: Astacidae, Cambaridae, and Parastacidae).—Smithsonian Contribution to Zoology 480:1—235. , & H. H. Hobbs III. 1989. New locality re- cords for two poorly known mexican freshwater shrimps (Decapoda, Palaeomonidae).—Crusta- ceana 57(2):220—222. Holthuis, L. B. 1949. Note on the Palaemonetes (Crustacea Decapoda) found in the United States-of America.—Proceedings of the Kon- inklinjke Nederlandse Akademie van Weten- schappen 52:87—95. . 1952. The subfamily Palaemoninae. A gen- eral revision of the Palaemonidae (Crustacea Decapoda Natantia) of the America, I].—Allan Hancock Foundation Occasional Papers 12:1— 396. 38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Markham, J. C., EK E. Donath-Hernandez, J. L. Villa- lobos-Hiriart, & A. Canta Diaz-Barriga. 1990. Notes on the shallow-water marine crustacea of the Caribbean coast of Quintana Roo, Méxi- co.—Anales del Instituto de Biologia, Univer- sidad Nacional Aut6noma de México, Serie Zoologia 61(3):405—446. Nates-Rodriguez, J. C., & J. L. Villalobos-Hiriart. 1990. Dos nuevas especies de camarones de agua dulce del género Macrobrachium Bate, (Crustacea, Decapoda, Palaemonidae), de la vertiente occidental del Pacifico.—Anales del Instituto de Biologia, Universidad Nacional Au- tonoma de México, Serie Zoologia, 61(1):1—11. Rathbun, M. J. 1902. Description of new Decapod Crustaceans from the West coast of North America.—Proceeding of the United States Na- tional Museum 24:885—905. Rodriguez-Almaraz G. A., & E. Campos. 1994. Dis- tribution and status of the crayfishes (Cambar- idae) of Nuevo Leon, México.—Journal of Crustacean Biology 14(4):729-735. Ruiz-Campos, G., M. Torres-Morales and S. Contrer- as-Balderas. 1985. Peces del Rio Alamo, Sub- cuenca del Bravo, México. II. Estructura y din- amica de la comunidad ictica.—Instituto de In- vestigaciones Cientificas, Universidad Auténo- ma de Nuevo Leon (México) 2:51—75. Smalley, A. E. 1963. The genus Potimirim in Central America (Crustacea, Atyidae).—Revista de Biologia Tropical 11(2):177-183. 1964. A new Palaemonetes from México (Decapoda: Palaemonidae).—Crustaceana 6: 229-232. Strenth, N. E. 1976. A review of the systematics and zoogeography of the freshwater species of Pa- laemonetes Heller of North America (Crustacea: Decapoda).—Smithsonian Contribution to Zo- ology 228:1—28. . 1994. A new species of Palaemonetes (Crus- tacea: Decapoda: Palaemonidae) from north- eastern México.—Proceedings of the Biological Society of Washington 107:291—295. Villalobos-Figueroa, A. 1959. Contribuci6n al cono- cimiento de los Atyidae de México. II. (Crus- tacea, Decapoda). Estudio de algunas especies del género Potimirim (Ortmannia), con descrip- cion de una especie nueva en Brasil.—Anales del Instituto de Biologia, Universidad Nacional Auto6noma de México 30:269-330. . 1967. Estudio de los Palaemonidos de Méx- ico I. Macrobrachium acanthochirus una nueva especie del SE de México.—Anales del Insti- tuto-de Biologia, Universidad Nacional Auton- oma de México, Serie Zoologia, 26(1/2):167— 174. . 1982. Decapoda. Pp. 215-239 in S. H. Hurl- bert and A. Villalobos-Figueroa, eds., Aquatic biota of México, Central America and the West Indies. Aquatic Biota-San Diego State Univer- sity Foundation, San Diego California. , & H. H. Hobbs, Jr. 1974. Three new crusta- ceans from La Media Luna, San Luis Potosi, México.—Smithsonian Contribution to Zoology 174:1-18. Villalobos-Hiriart, J. L., A. Canti Diaz-Barriga, & E. Lira-Fernandez. 1993. Los Crustaceos de Agua Dulce de México.—Revista de la Sociedad Mexicana de Historia Natural 44:267—290. Wiegmann, A. FE A. 1836. Beschreibung einiger neuen Crustaceen des Berliner Museums aus Mexiko und Brasilien.—Archiv fiir Naturges- chichte 2(1):145—-151. Williams, A. B. 1984. Shrimps, lobsters and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington D.C., 550 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):39-43. 1996 Neocrangon zacae (Chace, 1937) synonymized with N. resima (Rathbun, 1902), and compared with N. communis (Rathbun, 1899) (Decapoda: Caridea: Crangonidae) Mary K. Wicksten Department of Biology, Texas A&M University, College Station, Texas 77843, U.S.A. Abstract.—Features previously used to distinguish between Neocrangon za- cae and N. resima overlap greatly. The rostrum can be low and narrow, slightly elevated and deep, or high with a plate with or without teeth. The width of the hand of the first pereopod also varies, as does the distal width of the antennal scale. Similar variability exists in the shape of the rostrum of N. communis, but this species has a prominent carina on the fifth abdominal somite and has broader hands of the first pereopod and a more horizontal dactyl. Usually, adult N. resima are smaller than N. communis. Neocrangon zacae is considered to be a junior synonym of N. resima. A redescription of Neocrangon resima is presented. Crangonid shrimp of the genus Neocran- gon Zarenkov, 1965, are characterized as follows: two median carapace spines, with- out submedian spines between the middor- sal line and hepatic spine, the branchioste- gal spine not very strong or flared sideways, the gastric region not depressed below the general level of the carapace, one or two spines at the distal end of the merus of first pereopod, second pereopods subchelate and subequal in length to other pereopods, fourth and fifth pereopods with normal dac- tyls; not broadened; lateral surface of the abdomen not sculptured. Four species have been reported from the northeastern Pacific: N. communis (Rathbun, 1899), N. resima (Rathbun, 1902), N. abyssorum (Rathbun, 1902) and N. zacae (Chace, 1937). Of these, N. abyssorum is the most distinct, having enlarged eyes, a bent antennular pe- duncle, a long and narrow rostrum and a thin exoskeleton. It generally occurs on the lower continental slope, but has been re- ported as shallow as 97 m (Butler 1980). Colleagues conducting environmental impact surveys had difficulty distinguishing between N. communis, N. resima and N. za- cae. Neocrangon resima has been recog- nized by its elevated rostrum with a ventral plate (Rathbun 1904:fig. 65). However, Rathbun noted that the development of the rostrum seemed to be dependent on age, with specimens smaller than 20 mm not showing this feature. Schmitt (1921) quoted Rathbun as stating that the rostrum of N. communis also could be variable, with some specimens regenerating a narrow rostrum with an elevated tip. The illustration of the rostrum given by Schmitt (1921:fig. 64) shows a bilobed rostrum with a small tooth, while that by Green & Butler (1988) shows a rostrum ending in a vertical plate. The rostrum of N. zacae was described as as- cending at a slight angle “which varies somewhat’’ (Chace 1937). The hands of the first pereopod of N. re- sima were reported by Schmitt (1921) to be shorter and with a more longitudinal margin than in N. communis. The hands of the first pereopod of N. zacae were said by Chace (1937) to have a “longer palm”’ than those of N. resima, as well as a more longitudinal 40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dactyl. However, Green & Butler (1988) did not indicate any difference between the pro- portions of the palm of their specimen of N. resima and that of any other species. Schmitt quoted Rathbun as saying that N. resima had a “‘slight’’ median carina on the fifth abdominal somite while N. communis had a “blunt” carina. Green & Butler (1988) illustrated a specimen of N. resima with a “‘moderate’’ median dorsal carina, but noted that the carapace and abdominal sulci bore a short pubescence, as Butler (1980) reported for N. communis. Neocran- gon zacae lacked a carina on the fifth ab- dominal somite (Chace 1937). The ranges and habitats of the three spe- cies have been reported to overlap in the eastern Pacific. According to Butler (1980), N. communis ranges from the Bering Sea to San Diego, California at 16-1537 m on mud. Green & Butler (1988) reported N. re- sima from mud or mixed mud and sand bot- toms from Alice Arm, British Columbia to San Domingo Point, Baja California; Schmitt (1921) reported the depth range as 28-487 m. Neocrangon zacae has been re- ported from Monterey Bay, California to north of Gorgona Island, Colombia (al- though most reports come from California, U.S.A. and Baja California, Mexico) at depths from shore to 572 m. It has been reported from muddy or rocky bottoms. In southern California and Baja California, N. zacae usually was collected at shallower depths than N. communis (Wicksten 1980). Neocrangon communis and N. resima have been reported to be collected at the same trawling station (Schmitt 1921; Green & Butler 1988). Considering the overlap in range, depth and morphological features reported for N. communis, N. zacae and N. resima. I ex- amined specimens to determine what con- sistent differences could be found between them. Comparison of these specimens in- dicate that only two of these species are dis- tinct. Materials and Methods I examined 179 specimens identified as N. communis, N. resima or N. zacae from Santa Monica Bay and Monterey Bay, Cal- ifornia and Cook Inlet and the northeastern Gulf of Alaska. One hundred fifty two spec- imens from Santa Monica Bay were loaned by Ann Dalkey of the Hyperion Treatment Plant, Los Angeles County Environmental Monitoring Division; the specimens from Monterey Bay and Alaska were loaned by the California Academy of Sciences (CAS 014038, 020155, 020163 and 020155). I also examined the six syntypes of N. resi- ma, collected off San Diego, California, from the National Museum of Natural His- tory, Smithsonian Institution (USNM 25246). For each specimen, I noted total length, shape (low, deep, with plate, with or with- out teeth) and elevation of the rostrum, presence or absence of a noticeable dorsal carina of the fifth abdominal somite, and shape of the hand of the first pereopod. These data were tabulated and compared with the original descriptions of the species. Results The most striking difference among the shrimp examined was that all specimens from the Gulf of Alaska and Cook Inlet had a pronounced and blunt dorsal carina on the dorsal surface of the fifth abdominal somite (Fig. 1f). None of those from California had more than a vague trace of a dorsal carina (Fig. Ic). The Alaskan specimens were larger, ranging from 37.1—61.1 mm in total length. The specimens from California ranged from 19.5—50.6 mm in total length, but most individuals (59) were between 35— 40 mm long. The shape of the rostrum was low and narrow in 87 specimens from California and 13 Alaskan specimens, elevated but without a plate in 52 specimens from Cal- ifornia and eight from Alaska, and elevated and with a plate in 14 specimens from Santa Monica Bay, three from Monterey VOLUME 109, NUMBER 1 Al f Fig. 1. Neocrangon resima (Rathbun, 1902). a, 10 shapes of the rostrum; b, subchela of first pereopod; c, dorsal surface of fifth abdominal somite in frontal view. Neocrangon communis. d, three shapes of the rostrum; e, subchela of first pereopod; f, dorsal surface of fifth abdominal somite in frontal view. 42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bay and one from Alaska. In other speci- mens, the rostrum was broken off. In the collection from Santa Monica Bay, speci- mens with an elevated rostrum with a plate comprised about 9% of the total collection. Some of the shapes of the rostrum are shown in Fig. la and Id. It was difficult to determine the difference between “‘elevated and deep” and “elevated with a plate’’; moreover, the number of teeth Gf any) also varied. Most of the rostra did not have teeth except for a pointed apex. Among the syn- types of N. resima, the shape of the rostrum varied: two had a very low rostrum, one had a short but upraised rostrum and three had a moderately upraised rostrum. It is noteworthy that none of the syntypes had a raised rostrum with a plate, as illustrated by Rathbun (1904:fig. 65b). Most of the specimens from Alaska had a broader hand and also a more horizontal dactyl of the first pereopod than those from California (Fig. le). In general, the speci- mens from California had a more elongated hand with a more vertical dactyl (Fig. 1b), similar to that illustrated by Chace (1937: fig. 9d) for N. zacae. Discussion In comparing the syntypes of N. resima, the other specimens and the descriptions, the presence or absence of a carina on the dorsal surface of the fifth abdominal somite was the most reliable distinguishing feature for recognition of species. Such a carina is present in N. communis. Neocrangon com- munis also may have a broader hand of the first pereopod, about 2.7—3X long as wide, although some specimens may have a hand as much as 3.7X long as wide (Butler 1980). The hand has a dactyl that closes somewhat horizontally. The specimens from Alaska could be confidently identified as N. communis. The specimen illustrated by Green & Butler (1988) as N. resima probably also belongs to N. communis. Ex- cept for the variation in the shape of the rostrum, the specimens fit well the descrip- tion of NV. communis given by Butler (1980, as Crangon communis). The shape of the rostrum is unreliable as a diagnostic character of species of Neo- crangon. Among the specimens from Mon- terey Bay and Santa Monica Bay, it varied from low to high, shallow to deep, and with or without a plate, which sometimes had teeth. Variation also occurred among the syntypes of N. resima and the specimens of N. communis. Rathbun (1904) stated that the develop- ment of the rostral plate was dependent on age, ““specimens 20 mm long show no ev- idence of it.”” However, in two specimens with a total length of 22—28 mm, there was an elevated rostrum; another had a rostrum with a well-developed plate. Because my samples contained only one specimen less than 20 mm long, it is difficult to say whether or not the development is related to age or size. The smallest specimen, 19.5 mm long, had a low rostrum without a plate. The features of the fifth abdominal so- mite and first pereopod are indistinguish- able between N. resima and N. zacae. The ranges as reported for the two species, their habitat and depth also overlap. In the ab- sence of any other distinguishing features, Neocrangon zacae (Chace, 1937) must be considered to be a junior synonym of N. resima (Rathbun, 1902). It is unfortunate that the species was illustrated in Rathbun’s work from an atypical specimen having a raised rostrum and a ventral plate, for this shape of the rostrum occurs in less than 10% of the population. A redescription of N. resima, including new information on variability within the species, follows. Neocrangon resima (Rathbun, 1902) Crangon resima Rathbun, 1902: 889.— Rathbun 1904:124, fig. 65. Crago resima.—Schmitt 1921:96, fig. 64. Crago zacae Chace, 1937:136, text-fig. 9. Neocrangon zacae.—Kuris & Carlton 1977:554.—Wicksten 1980:39. VOLUME 109, NUMBER 1 Neocrangon resima.—Kuris & Carlton 1977:554. Redescription. (Modified from Chace 1937).—Rostrum usually narrow, with pro- nounced dorsal sulcus and blunt apex, vari- able in length, not reaching end of cornea of eye to exceeding entire eye. Rostrum as- cending, usually at slight angle but as much as 45° in some specimens; ventral plate, with or without teeth, may be present. Carapace about 0.25 length of abdo- men. Gastric region not depressed below general outline of carapace. Two spines in dorsal midline, posterior one longer and arising in front of middle of carapace. Strong hepatic spine in line with anterior dorsal spine. Anterior margin with antennal and branchiostegal spines. Eye large and black. Abdominal somites with few small setae along ventral surface but without obvious pubescence. Somites 1 and 2 with elevated posterior portions, elevations preceded by broad, shallow transverse sulci. Somites 1— 4 without carinae. Pleura of first to third somites with rounded or squared margins, fourth pleuron with minute posteroventral spine, fifth pleuron with posterolateral point. Fifth somite occasionally with ob- scure median carina. Sixth somite approxi- mately 1.6X length of fifth, bearing pair dorsal longitudinal carinae separated by median sulcus and bounded by shallower lateral sulci. Telson longer than sixth so- mite, with slight median sulcus, bearing 2 pairs dorso-lateral spines, apex blunt. Antennular peduncle with 3 segments, second 2-3 length of third, stylocerite short. Antennular flagella exceeding anten- nal scale. Basicerite of antenna with lateral spine. Spine of antennal scale exceeding blade, antennal scale about 3.5xX long as wide. Third maxilliped long and slender, ex- tending slightly beyond antennular flagella, exopod with lash. First pereopod subche- late, merus with spine at outer angle of dis- tal end, palm of subchela 3—3.3X long as 43 wide, dactyl closing obliquely. Second per- eopods slender, shorter than first, chelate. Third pereopods slender, extending well be- yond third maxillipeds, fourth and fifth per- eopods shorter; all with simple, slender dac- tyls. Uropods equal to or exceeding length of telson. Color in life translucent, mottled finely with greenish-brown and scarlet on dorsal and lateral surfaces; ventral surface white, antennae banded with scarlet and white, uropods and telson mottled. Literature Cited Butler, T; H. 1980. Shrimps of the Pacific coast of Canada.—Canadian Bulletin of Fisheries and Aquatic Sciences 202:1—280. Chace, E A. Jr. 1937. The Templeton Crocker Expe- dition. VII. Caridean decapod Crustacea from the Gulf of California and the west coast of Lower California.—Zoologica 22:109-138. Green, G., & T. H. Butler. 1988. Range extensions of three caridean shrimps to British Columbia wa- ters.—Royal British Columbia Museum Contri- butions to Natural Science 8:1—7. Kuris, A. M., & J. T. Carlton. 1977. Description of a new species, Crangon handi, and new genus, Lissocrangon, of crangonid shrimps (Crustacea: Caridea) from the California coast, with notes on adaptation in body shape and coloration.— Biological Bulletin 153:540—-559. Rathbun, M. J. 1899. List of Crustacea known to oc- cur on and near the Pribilof Islands. Pp. 555— 557 in D.S. Jordan et al., ed., The fur seals and fur seal islands of the north Pacific Ocean, Washington, D.C., Pt. 3. 1902. Descriptions of new decapod crusta- ceans from the west coast of North America— Proceedings of the U.S. National Museum 24: 885-905. . 1904. Decapod crustaceans of the northwest coast of North America.—Harriman Alaska Ex- pedition, Series 10:1—210. Schmitt, W. L. 1921. The marine decapod Crustacea of California.—University of California Publi- cations in Zoology 23:1—470. Wicksten, M. K. 1980. Range extensions of four spe- cies of crangonid shrimps in the eastern Pacific Ocean (Decapoda: Crangonidae).—Bulletin of the Southern California Academy of Sciences 79:38-41. Zarenkov, N. A. 1965. Revision of the genera Cran- gon Fabricius and Sclerocrangon G.O. Sars (Decapoda, Crustacea).—Zoologicheskii Zhur- nal 44:1761—1775. (In Russian.) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):44—52. 1996 Daldorfia Rathbun, 1904 (Crustacea: Decapoda) from the Neogene of Japan Hiroaki Karasawa and Hisayoshi Kato (H. Karasawa) Mizunami Fossil Museum, Yamanouchi, Akeyo, Mizunami, Gifu 509-61, Japan; (H. Kato) Natural History Museum and Institute, Chiba, 955-2 Aobacho, Chuoku, Chiba 260, Japan Abstract.—Daldorfia nagashimai, new species, is described from the Hi- gashimorogata Formation (Upper Miocene - Lower Pliocene), Miyazaki Group of Miyazaki Prefecture, Japan. This species is characterized by a large carapace with irregular, large, granulose tubercles dorsally and without a deep, eroded area behind a longitudinally hexagonal mesogastric lobe. Another specimen representing a second unnamed species is recorded from the Aoso Formation (Upper Miocene) of Miyagi Prefecture, Japan. These species represent the first records of Neogene decapod crustaceans from Japan and extend the geologic range of the genus Daldorfia to the Late Miocene age. The subfamily Parthenopinae is a small group including eight Recent genera, Daira de Haan, 1833, Dairoides Stebbing, 1920, Daldorfia Rathbun, 1904, Leiolambrus A. Milne Edwards, 1878, Parthenope Weber, 1795, Solenolambrus Stimpson, 1871, Thy- rolambrus Rathbun, 1894, and Tutankha- men Rathbun, 1925. The fossil record of Parthenope Weber, 1795, is robust in Ce- nozoic deposits throughout the world (Glaessner 1969). Daira is recorded from the Eocene-Miocene of Europe, the Plio- cene of Fiji, and the Miocene of Japan (Glaessner 1969; Miller 1984; Miiller & Collins 1991; Karasawa 1993). The only known fossil Leiolambrus was described from the Upper Eocene Bartonian of En- gland (Quayle & Collins 1981) and Tut- ankhamen also has been reported from the Oligocene Gambier Limestone of Australia (Jenkins 1985). The fossil records of the other four Recent genera are unknown throughout the world, but the extinct, Me- solambrus declinatus Miiller & Collins, 1991, similar to Thyrolambrus Rathbun, 1894, was described from the Szépvdélgy Formation (Upper Eocene) of Hungary; Glaessner (1969) recognised the Upper Eo- cene, Phrynolambrus Bittner, 1893 as the junior synonym of the subgenus Pseudo- lambrus Paulson, 1875, in Parthenope, but subsequently Guinot (1979) separated Phrynolambrus from Pseudolambrus and suggested that Phrynolambrus has a close affinity with Daira and Dairoides. The purpose of this paper is to describe a new species and a related, but unnamed species of Daldorfia from the Neogene de- posits of Japan. The materials were collect- ed from a road cut (Loc. MYZ-4 of Kara- sawa 1993, 31°56’48"N, 131°16'46’E) at Akatani, Uranona, Takaoka-cho, Higashi- morogata-gun, Miyazaki Prefecture (Fig. 1A). Siltstone of the Aya Member of the Higashimorogata Formation, Miyazaki Group (Upper Miocene to Upper Pleisto- cene) is exposed at this locality (Tomida 1991; Karasawa 1993). This formation is assigned to Zones N.17b—18 (latest Mio- cene-earliest Pliocene) of Blow’s scale of planktonic foraminifera (Suzuki 1987). Daldorfia nagashimai, new species, 1s based upon the holotype and two paratype specimens included within calcareous nod- ules. Karasawa (1993) reported two species of crabs from this locality. The decapod as- VOLUME 109, NUMBER 1 45 Sinkansen Line Tohoku aia ee TOMIYA ARE | Yoshidagawa River yd Hage -bearing Meare rshins X ccality Bay C) Sendai i) ob Fossil-bearing locality o~ mays wi S —/ Oyodogawa \\ River Takaoka The Sea of Hyuga Fig. 1. Map of Japan showing the fossil-bearing localities. semblage is dominated by Carcinoplax prisca Imaizumi. Two specimens of Linu- parus sp. aff. L. trigonus (von Siebold) have been found also. Tomida (1991) re- ported some molluscs, Perotrochus sp., Bathybembix sp., Hindsia sp. and Acila submirabilis Makiyama from the locality. Decapods collected from the locality sug- 46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gest an environment within the lower sub- littoral zone on a muddy to sandy bottom (Karasawa 1993). A large manus of the second species, Daldorfia sp., was obtained from the Aoso Formation at the exposed road-cut south of Matsugai, Tomiya Town, Miyagi Prefecture (38°20'22’N, 140°55’56"E) (Fig. 1B). The Aoso Formation consists mainly of cross- laminated, poorly consolidated coarse- grained sandstone with remarkable key beds of tuff layer (K7) intercalated within the lower part and a pumiceous tuff layer underlying the conglomerate bed (K8) in the middle part of the Formation (Kitamura et al. 1983). The decapod fossil was ob- tained from this lithology beneath the K8 beds. Fujiwara (1992) discussed the Late Miocene molluscan assemblages of the re- gion. He discriminated the Serripes-Miya and Glycymeris-Dosinia assemblages with- in the K8 beds, and regarded the inhabitants to be subneritic to mesoneritic, respectively. According to Fujiwara (1992) and Saito & Fujiwara (1994), the middle to the upper part of the Aoso Formation lying several meters above the K7 tuff bed is correlated to Blow’s N17b Zone. Therefore, the geo- logic age of the fossil bearing horizon is assignable to the latest Miocene. From an- other locality of the Aoso Formation, a small number of fingers referable to the families Callianassidae and Parthenopidae were collected by the same person who dis- covered the material described below. Section Heterotremata Guinot, 1977 Superfamily Parthenopoidea MacLeay, 1838 Family Parthenopidae MacLeay, 1838 Subfamily Parthenopinae MacLeay, 1838 Genus Daldorfia Rathbun, 1904 Type species.—Cancer horridus Linnae- us, 1758, by monotypy (ICZN Opinion 1582); Recent, Indo-West Pacific. Geologic range.—Late Miocene—Re- cent. Daldorfia nagashimai, new species Figs. 2—4 Material. KMNH IvP 300,022, holo- type (Kitakyushu Museum and Institute of Natural History, 6, Nishihonmachi 3-cho- me, Yahatahigashiku, Kitakyushu, 805 Ja- pan), coll. H. Nagashima, 1994; MFM83058, 83059, 2 paratypes (Mizunami Fossil Mu- seum, Yamanouchi, Akeyo, Mizunami, Gifu, 509-61 Japan), coll. S. Tomida, 1993. Diagnosis.—Large parthenopid; dorsal carapace covered with irregular, large gran- ulose tubercles; mesogastric lobe large, lon- gitudinally hexagonal, without deep, eroded area behind it; major cheliped like Daldor- fia horrida, \arge, long, with stout fingers. Description.—A large Daldorfia; cara- pace appears to be broadly pentagonal in outline, width approximately 1.4 times length. Front broadly triangular, down- turned, with shallow, ovate dorsal hollow. Orbits small, subovate, directed anterolat- erally. Anterolateral margin strongly con- vex, bearing irregular, large tubercles; deep notch defined lateral termination of from cervical groove. Posterolateral margin strongly convex. Posterior margin obscured by poor preservation. Dorsal surface with inflated, tuberculate regions separated by shallow, moderately well-defined grooves. Protogastric lobes strongly convex, with sharply pointed tu- bercles on highest parts, deep depressions between gastric and hepatic lobes. Meso- gastric lobe with large, granulose tubercles, longitudinally hexagonal, gently convex; narrow anterior mesogastric process with tubercles longitudinally arranged; deep de- pression on either side of mesogastric lobe. Cardiac lobe broken. Highest part of strong- ly convex hepatic regions with sharply pointed tubercles. Branchial lobes uneven, with irregular, large, granulose tubercles and small setal pits. Intestinal region de- pressed, with irregular tubercles. Dactylus and propodus of right major cheliped (paratype) preserved, but tips of both fingers, and proximal end and mesial VOLUME 109, NUMBER 1 47 Fig. 2. Daldorfia nagashimai, new species, holotype, X1.0; dorsal view; showing central and left side of carapace, parts of merus and carpus of left cheliped. surface of propodus broken. Fingers with a wide gape. Dactylus stout, with irregular, granulose tubercles; occlusal surface miss- ing. Imperfect fixed finger with a broad, flattened, molariform tooth on occlusal sur- face. Palm long, covered with longitudinal rows of irregular, conical, granulose tuber- cles decreasing in size proximally; greatest distal width about 1.5 times proximal width; tubercles large on dorsal and lateral surfaces, rather small on ventrolateral sur- face; ventral margin with conical, granulose tubercles; four longitudinal rows of spines present on ventromesial surface; two bro- ken bases of large mesial spines present, others missing. Carpus and merus of left cheliped (holotype) with granulose tuber- cles, merus bearing acute spines on dorsal margin. Pereiopods and ventral aspects unknown. Derivation of name.—The species name honors H. Nagashima, who collected the holotype. Remarks.—Daldorfia comprises nine Re- cent species from the Indo-Pacific and East Atlantic Oceans. Of these, Daldorfia na- gashimai most closely resembles D. horri- da (Linnaeus 1758) and D. rathbuni (de Man 1902), from the Indo-West Pacific Oceans, but the new species has a moder- ately uneven, not eroded dorsal surface with irregular, granulose tubercles. This new 48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. species has a large, longitudinally hexago- nal mesogastric lobe with large, granulose tubercles, but D. horrida and D. rathbuni each have a small, rounded mesogastric lobe. Absence of a deep, eroded area behind the mesogastric lobe in the new species dis- tinguishes it from D. horrida and D. rath- buni. There is, in the general features of the major cheliped, considerable similarity be- tween D. nagashimai, and D. horrida: the latter bears two large, conical, distal tuber- cles, one on the lateral surface and the other on the dorsal surface of the propodus, but in D. nagashimai, the distal ends of the lat- eral and dorsal surfaces are covered with irregular, conical tubercles. Absence of spines on the mesial surface of the dactylus readily distinguishes D. nagashimai from D. horrida. Recent D. horrida and D. rathbuni differ from most other parthenopine crabs, includ- ing the other species of Daldorfia (.e., D. investigatoris (Alcock 1895), D. spinosis- sima (A. Milne Edwards 1862)), in having both fingers of the major cheliped with large blunt molariform denticles on the proximal occlusal surfaces. Zipser & Vermeij (1978) observed that the Recent Daldorfia horrida uses the occlu- sal surfaces with molariform teeth of the ma- Daldorfia nagashimai, new species, paratype, X1.0. Dorsal view of carapace. jor cheliped to crush gastropod shells (.e., Cerithium, Cypraea, Drupa, Cantharus, Vasum) and to feed on hermit crabs. Ng & Rodriguez (1986) described detail the den- tition on both chelipeds of D. horrida. There is, on the occlusal surface of the major chela, a great similarity between the fossil species and Recent D. horrida. Thus, D. nagashimai appears to have acquired the crushing be- havior of feeding on gastropods and hermit crabs at least by the Late Miocene. Daldorfia sp. Fig. 5 Material. SSME 13320 (Sendai Sci- ence Museum, 4-1, Forest Park, Dainohara, Aobaku, Sendai 981). 1 specimen, coll. Y. Takaizumi, 1991. Description.—A large propodus of left minor cheliped, oval in cross section, cov- ered on every surface except for fixed fin- ger with large, conical tubercles. Tubercles vary in size, up to | cm in diameter. Lateral surface has flattened tubercles, dorsal bor- der bears rather pointed ones. A hook- shaped projection with which carpus artic- ulates on dorsal border situated near prox- imal articulation. Fixed finger short, about Y; of total length, slightly curved inward, VOLUME 109, NUMBER 1 Fig. 4. Daldorfia nagashimai, new species, paratype, X0.93. Right major cheliped: mesial; d, ventral view. 49 a, dorsal; b, lateral; c, 50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. stout with apex faintly curved downward. Occlusal surface with 3 blunt, subconical teeth. A shallow, short longitudinal furrow on apex below occlusal edge. Remarks.—In general shape, the present chela most closely resembles the minor chela of D. horrida, but differs in lacking granulose tubercles on the lateral surface Daldorfia sp., X0.9. Propodus of left major cheliped: a, dorsal; b, lateral; c, mesial view. and spines on the ventral margin. The mi- nor chela of D. nagashimai being as yet unknown, precludes comparison with D. sp. Acknowledgments We thank R. M. Feldmann (Kent State University) and J. S. H. Collins (London) VOLUME 109, NUMBER 1 for critically reading our manuscript, and H. Nagashima (Fukuoka City, Fukuoka), S. Tomida (Chukyo Gakuin University, Gifu), Y. Takaizumi (Sendai City, Miyagi) for of- fering their fossil decapod specimens. We also thank O. Fujiwara (Tono Geoscience Center, Gifu), Y. Okazaki (Kitakyushu Mu- seum and Institute of Natural History, Fu- kuoka), T. Komai (Natural History Museum and Institute, Cheba), and Y. Okumura (Mi- zunami Fossil Museum, Gifu), who offered much valuable advice. Literature Cited Alcock, A. 1895. The Brachyura Oxyrhyncha: mate- rials for a carcinological fauna of India, no. 1.— Journal of the Asiatic Society of Bengal 65: 157-291. Bittner, A. 1893. Decapoden des pannonischen Ter- tiars.—Sitzungsberichte der kais. Akademie der Wissenschaften in Wien, Mathematisch-Natur- wissenschaftiche Klasse 102:10—37. Fujiwara, O. 1992. A new species of Halicardia (Bi- valvia) from the Late Miocene of Miyagi Pre- fecture, northeast Honshu, Japan.—Saito Ho-on Kai Museum, Research Bulletin 60:9—20. Glaessner, M. EF 1969. Part R. Arthropoda. Pp. R400— 456 in R. C. Moore, ed., Treatise on invertebrate paleontology. The Geological Society of Amer- ica and the University of Kansas. Guinot, D. 1977. Propositions pour une nouvelle clas- sification des Crustacés, Décapodes, Brachy- oures.—Comptes-rendus hebdomadaires des sé- ances de 1’ Académie des Sciences, Paris, series D 285:1049-1052. . 1979. Données nouvelles sur la morphologie, la Phylogenése et la taxonomie des Crustacés . Décapodes Brachyoures.—Mémoires du Mu- séum National d’Histoire Naturelle, série A. Zoologie 112:354 pp. Haan, W. de 1833 (1833-1850). Crustacea. Pp. i-xvii, i-XXxi, 1x—xvi, 1-243, pls. A-J, L-Q, 1-55 in P. EF von Siebold, Fauna Japonica sive Descrip- tio Animalium, quae in Itinere per Japoniam, Jussu et Ausiciis Superiorum, qui Summun in India Batava Imperium Tenent, Suscepo, Annis 1823-1830 Collegit, Notis, Observationibus et Adumbrationibus Illustravit. Ludguni-Batavo- rum. Jenkins, R. J. E 1985. Fossil spider crabs from Aus- tralia——Special Publication of South Australia Department of Mines and Energy 5:145—165. Karasawa, H. 1993. Cenozoic decapod Crustacea from southwest Japan.—Bulletin of the Mizun- ami Fossil Museum 20:1—93. 51 Kitamura, N., A. Ozawa, & H. Nakagawa. 1983. Ge- ology of the Yoshioka district. Quadrangle Se- ries, scale 1:50,000. Geological Survey of Ja- pan. 50 pp. Gin Japanese with English abstract). Linnaeus, C. 1758. Systema Naturae per Regma Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Syn- onymis, Locis. Edition 10, 1:1ii + 824. Hol- miae. MacLeay, W. S. 1838. On the brachyurous decapod Crustacea brought from the Cape by Dr. Smith. Pp. 53-71 in Illustrations of the Annulosa of South Africa; being a Portion of the Objects of Natural History Chiefly Collected during an Ex- pedition into the Interior of South Africa, under the Direction of Dr. Andrew Smith, in the Years 1834, 1835, and 1836; Fitted out by “The Cape of Good Hope Association for Exploring Cen- tral Africa.”’ London. Man, M. de. 1902. Ergebnisse einer zoologischen For- schungsreise in den Molukken und Borneo, im Auftrange der Senckenbergischen Naturforsch. Gesellschaft ausgefiihrt von Dr. Willy Ktiken- thal. Teil. 2, Reiseergebon., Bd. 3, Heft. 3.— Abhundlungen der Senckenbergischen Natur- forschenden Gesellschaft 85:467—-929. Milne Edwards, A. 1862. Faune carcinologique de Vile de la Réunion, Annexe F de |’ ouvrage in- titulé: Notes sur l’ile de la Réunion par L. Mail- lard. Pp. Fl—16, plates XVII-XIX in L. Mail- land, Notes sur l’ile de la Réunion (Bourbon). Paris, Dentu. . 1878 (1873-1881). Etudes sur les Xiphosures et les Crustacés de la région Méxicaine. Pp. 1— 368 in Mission scientifique au Méxique et dans l’ Amérique centrale, Recherches Zoologiques pour servir a l’histoire de la faune de l’ Amé- rique centrale et du Méxique, 5. Miiller, P. 1984. Decapod Crustacea from the Baden- ian.—Geologica Hungarica, Series Palaeonto- logica 42:317 pp. , & J. S. H. Collins. 1991. Late Eocene coral- associated decapods (Crustacea) from Hunga- ry.—Contributions to Tertiary and Quaternary Geology 28:47-92. Ng, P. K. L., & G. Rodriguez. 1986. New records of Mimilambrus wileyi Williams, 1979 (Crustacea: Decapoda: Brachyura), with notes on the sys- tematics of the Mimilambridae Williams, 1979, and Parthenopidae MacLeay, 1838, sensu Guin- ot. 1978.—Proceedings of the Biological Soci- ety of Washington 99:88—99. Paulson, O. 1875. Podophthalmata and Edriophthal- mata (Cumacea). Studies on Crustacea of the Red Sea with notes regarding other seas. Part I. xiv + 144 pp. Kiev (in Russian). Quayle, W. J., & J. S. H. Collins. 1981. New Eocene 52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON crabs from the Hampshire Basin.—Palaeontol- ogy 24:733-758. Rathbun, M. J. 1894. Descriptions of a new genus and four new species of crabs from the Antillean region.—Proceedings of the United States Na- tional Museum 17:83-86. 1904. Some changes in crustacean nomen- clature.—Proceedings of the Biological Society of Washington 17:169-172. . 1925. The spider crabs of America.—United States National Museum Bulletin 129:xx + 613 Pp. Saito, T. & O. Fujiwara. 1994. Tertiary planktonic fo- raminiferal biostratigraphy near Sendai district. Palaeontological Society of Japan, Abstracts of the 1994 Annual Meeting, p. 37 (in Japanese). Stebbing, T. R. R. 1920. South African Crustacea (Part X of S. A. Crustacea, for the Marine In- vestigations of South Africa).—Annals of South African Museum 17:231—272. Stimpson, W. 1871. Preliminary report on the Crus- tacea dredged in the Gulf Stream in the Straits of Florida, by L. E de Pourtales, Assist. U.S. Coast Survey.—Bulletin of the Museum of Comparative Zodlogy at Harvard College 2: 109-160. Suzuki, H. 1987. Stratigraphical study of Miyazaki Group.—Contributions from the Institute of Ge- ology and Paleontology, Tohoku University 90: 1—24 (In Japanese with English abstract.). Tomida, S. 1991. Pleurotomariid fossil from the Mi1- yazaki Group, Kyushu, Japan.—Bulletin of the Mizunami Fossil Museum 18:111-118. Weber, E 1795. Nomenclator entomologicus secun- dum Entomologiam Systematicam ill. Fabricii adjectis recens detectis et varietatibus. viii + 171 pp. Chilonii and Hamburg. Zipser, E., & G. J. Vermeij. 1978. Crushing behavior of tropical and temperate crabs.—Journal of Ex- perimental Marine Biology and Ecology 39: 135-150. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):53-69. 1996 Systematics and distribution of the genus Calocarides (Crustacea: Decapoda: Axiidae) Brian Kensley Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—The nine currently recognized species of Calocarides are re- viewed, and two new species are described: C. capensis from the south-east Atlantic Ocean off the Cape of Good Hope, and C. macphersoni from the Atlantic Ocean off Namibia. Species of the genus are particularly characterized by the possession of dentate median, submedian, and lateral carinae of the carapace, weakly pigmented or unpigmented eyes, and robust, quite heavily sclerotized, tuberculate and/or spinose first pereopods. The approximately 100 described species of Axiidae show their greatest diversity in shallow tropical and subtropical seas, with several genera confined to the tropics, e.g., Coralaxius Kensley & Gore, 1982, Parax- iopsis De Man, 1905, Paraxius Bate, 1888, Scytoleptus Gerstaecker, 1856. A few gen- era, however, have colonized colder and deeper waters and show considerable diver- sity and broad geographic range, e.g., Cal- ocarides Wollebaek, 1908, and Eiconaxius Bate, 1888, the latter an inquiline of hex- actinellid sponges. Calocarides is geo- graphically widespread in continental shelf- Slope regions, and contains cold-water in- habiting, soft-bottom burrowing species, some of which may be associated with zones of upwelling. The genus is reviewed here, two new species are described, and a number of taxonomic and _ distributional problems noted, relating to species previ- ously included in the genus. Abbreviations: cl—carapace length; HUMZ—Laboratory of Marine Zoology, Hokkaido University; ICMB—Instituto de Ciencias del Mar, Barcelona; MNHN—Mu- séum National d’Histoire Naturelle, Paris; SAM—South African Museum, Cape Town; UBZM—Zoological Museum, Uni- versity of Bergen; USNM—National Mu- seum of Natural History, Smithsonian Institution. Systematic Account Family Axiidae Genus Calocarides Wollebaek, 1908 Calocaris (Calocarides) Wollebaek, 1908: 3, 23s Axiopsis (Calocarides).—De Man, 1925:1, 2, 6, 67, 71.—Balss, 1957:1579. Calocarides.—Bouvier, 1940:97.—Sakai & de Saint Laurent, 1989:4, 78.—Poore, 1994:98. Euconaxius {laps. cal. for Eiconaxius Bate, 1888] Trybom, 1904:384. Type species.—By subsequent designa- tion by Sakai & de Saint Laurent, 1989, Eu- conaxius coronatus Trybom, 1904. Type lo- cality: Skagerrak, North Sea, 230-500 m. Gender: Masculine. Diagnosis.—Gonochoristic. Carapace having small supraocular spine present, as part of lateral rostral series; post-cervical carina low, rounded, barely discernible in some species, lacking spines; rostrum at same level as anterior carapace, margins armed; median carina dentate; submedian carina dentate; lateral carina dentate. Eye reaching to about midlength of ros- trum, cornea weakly pigmented or unpig- 54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Branchial formula for Calocarides coron- atus. r = reduced. Maxillipeds Pereopods 1 2 3 1 D 3 4 5 Exopod 1 1 1 — — — — — Epipod 1 1 1 1 1 1 — Podobranch —_—_ — r r r r— — Arthrobranch — — 2 2.) 2 2) 2. — Pleurobranch — — — Setobranch — — 1 1 1 1 1 1 mented; eyestalk longer than eye. Antennal scaphocerite elongate. Maxilla 2, scaphog- nathite having spinulose flagellum. Maxil- lipeds 1—3 with exopods and epipods. Pe- reopod 1, chelipeds asymmetrical, some- what sexually dimorphic. Propodi and dac- tyli of pereopods 3-5 slender, not expanded, dactyli simple. Pleopodal rami slender elongate; appendix interna present on pleopods 2—5. Pleopod | ¢ absent; pleo- pod 1 @ uniramous, of 2 articles. Pleopod 2 3 with appendix masculina of single lobe. Uropod with lateral ramus with su- ture. Telson longer than wide, or at least as long as wide, with dorsal spines. Branchial formula: see Table 1. Species: Calocarides capensis, new species. Off Ta- ble Bay, South Africa, 1280 m. Calocarides coronatus (Trybom, 1904). Swedish and Norwegian fjords, North Sea, 80—700 m. Calocarides lev (Zarenkov, 1989). Gulf of California, 2000—2036 m. Calocarides longispinis (McArdle, 1901). Bay of Bengal, 549 m. Calocarides macphersoni, new species. Off Namibia, 300-480 m. Calocarides quinqueseriatus (Rathbun, 1902). Off mouth of Columbia River, Or- egon, to southern California, 292—2013 m. Calocarides rostriserratus (Andrade & Baez, 1977). Off Chile, 320-400 m; off Peru, Gulf of Panama, 576-825 m. Calocarides rudolfi (Zarenkov, 1989). Off Angola, 490-510 m. Calocarides soyoi (Yokoya, 1933). Off northern Japan, 138—270 m. Calocarides spinulicauda (Rathbun, 1902). Off Bodega Head, California, 113 m; Pu- get Sound, 70—256 m; British Columbia, 91-128 m. Calocarides vigila Sakai, 1992. New Zea- land, 74—146 m. Taxonomic remarks.—The species Cal- ocaris lev Zarenkov, 1989, described from a hydrothermal vent at 2000—2036 m in the Gulf of California, is almost certainly a spe- cies of Calocarides. The proportions and spination of the first chelipeds bear more resemblance to C. rostriserratus from Chile, Peru, and Panama, than to C. quin- queseriatus from southern California. Giv- en that this species was taken from greater depth than either of the two abovemen- tioned species, its relative isolation of the Gulf of California, and the lack of males, it seems prudent to await further material be- fore deciding on the status of this species. Calocaris rudolfi Zarenkov, 1989, from 490-510 m off Angola is likewise a species of Calocarides, and bears some resem- blance to C. macphersoni (see Remarks un- der C. macphersoni). Although Sakai & de Saint Laurent (1989) tentatively placed Axiopsis tenuicor- nis De Man, 1905, Axius habereri Balss, 1913, Axius armatus Smith, 1881, and Ax- iopsis werribee Poore & Griffin, 1979, within Calocarides, none of these possess a majority of the features that define the genus. The specific identity of Calastacus quin- queseriatus and Axius spinulicauda amu- rensis, both recorded from Japan by Kob- jakova (1937) will remain uncertain until the material can be located and examined. Ecological and biological notes.—Little information exists for species of Calocari- des, except for C. coronatus and C. quin- queseriatus. Both of these species are known to burrow into soft bottom sedi- ments. In the case of C. coronatus, these sediments are usually mud with a high VOLUME 109, NUMBER 1 (60%) silt content and a temperature of 6°— 7.5°C. Calocarides coronatus overlaps in bottom distribution with the calocarid shrimp Calocaris macandreae, but domi- nates in deeper water (Brattegard 1966). For all the records of C. quinqueseriatus, the bottom sediments were recorded as ei- ther yellow or green mud, with tempera- tures ranging from 13°—16°C. Even less is known of the prey organisms of species of Calocarides. Calocarides macphersoni from Namibia was taken from the gut-contents of two bottom feeders, viz. the scorpaenid fish Helicolenus sp., and the ray Raja confundens, again suggesting that the species burrows in soft sediments. Regarding reproduction and fecundity, C. coronatus produces a about 65 eggs/female, with an egg diameter 15% of carapace length (n = 2). Calocarides quinqueseriatus produces about 180 eggs/female, with an egg diameter of about 7% of carapace length (n = 4). These egg sizes are in the high range for free-living axiids, suggesting that a fairly advanced larva hatches, having a rel- atively short planktonic larval lifespan. Spe- cies which produce thousands of eggs per female generally have much smaller eggs, e.g., Oxyrhynchaxius caespitosa with 16,000 eggs/female, egg diameter 1.3% of carapace length; Axiopsis serratifrons with 4000 eggs/ female, egg diameter 2.7% of carapace length. The commensal species of Eiconax- ius by contrast, produce far fewer large eggs, 8—30 eggs/female, with an egg diameter 16— 38% of carapace length. Elofsson (1959) re- corded a 10 mm stage II larva (Gurney 1942) of C. coronatus from 600 m. Distribution.—From the depth ranges of the 11 species of Calocarides (see list of species), it is clear that they are continental Shelf/slope dwellers, probably with a pref- erence for soft-sediment bottoms into which they can burrow. In several cases, the shelf/ slope habitat is also characterized by up- welling of deeper water, e.g., off the Cape of Good Hope and Namibia, off Pacific North America, off Chile, and in localized areas off Norway. 55) The widespread geographical distribution of the species of Calocarides, in the Atlan- tic, Indian, and Pacific oceans (Fig. 1), along with the most northerly (C. corona- tus) and most southerly (C. vigila) records for the Axiidae, suggests a considerable age for the genus. If, as has been suggested, the axiid-like shrimps evolved from Tethyan ancestors (Kensley 1994), shallow-water precursors of Calocarides must have spread with tectonic plate movements and the opening of the present-day oceanic basins, at the same time penetrating to varying depths the waters of the continental shelves. The direction of this invasion from shallow to deeper shelf waters may be reflected in that some eye-pigment is retained in some species (e.g., C. soyoi) while other species completely lack eye-pigment (e.g., C. ca- pensis). Calocarides capensis, new species Fig. 2 Calastacus longispinis.—Stebbing, 1910: 367, non McArdle, 1901. Calocaris (Calastacus) longispinis.—De Man, 1925:8 (part), non McArdle, 1901.—Barnard, 1950:503, fig. 93 d-f. Calocaris longispinis.—Kensley, 1981:30, non McArdle, 1901. Calocarides coronatus.—Sakai & de Saint Laurent, 1989:79, non Trybom, 1904. Material examined.—Holotype, SAM- A940, ovigerous 2 cl 19.0 mm (carapace damaged), R/V Pieter Faure 40 miles N79°E of Table Mountain, Cape of Good Hope, South Africa, 1280 m. Diagnosis.—Carapace surface minutely pitted, especially posteriorly; rostrum with 5—6 lateral teeth; median carina well marked, unarmed; submedian carina bear- ing | or 2 spines; lateral carina with 3 or 4 spines. Abdominal pleura ventrally round- ed, pleuron 2 widest. Telson with 1 pair of small spines dorsally, 3—4 small mobile spines at posterolateral corner, posterior margin gently convex between posterolat- eral and median spines. 56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rostriserratus rudolfie macphersonie capensis e Fig. 1. Distribution of species of Calocarides. Maxilliped 3, merus with 4 spines on posterior margin; carpus with single small distal spine. Pereopod 1, larger cheliped, is- chium with single strong distal spine; merus with strong spine at midlength of posterior margin, few small tubercles in proximal half, anterior margin with 5 distal spines; carpus with few low tubercles on anterior margin and few small tubercles on lateral surface; propodal palm slightly longer than fingers, outer surface with tiny scattered acute tubercles and strong ridge near pos- terior margin reaching almost to tip of fixed finger, anterior margin with about 11 strong spines, cutting margin of finger finely toothed, with large proximal tubercle; dac- tylus with few tubercles on anterior margin, cutting edge finely toothed, with proximal tubercle. Smaller cheliped, ischium with strong distal spine; merus with strong spine at midlength of posterior margin, anterior margin with 3 distal spines; carpus with few tubercles on anterior margin and lateral sur- face; propodal palm longer than fingers, lat- eral surface with few scattered small tuber- cles and ridge near posterior margin, ante- rior margin with about 11 spines; fixed fin- ger and dactylus as in larger cheliped. Pereopod 2, ischium with few small spines and strong distal spine on posterior margin; merus with 3 widely spaced spines on pos- terior margin. Pleopod 1 consisting of sin- gle slender elongate ramus. Pleopods 2—5 each with appendix interna on mesial mar- gin of endopod. Lateral uropodal ramus with single small spine on outer margin; su- ture bearing 4—8 irregular spines; mesial ra- mus with 4 spines on middorsal ridge. Remarks.—See ‘Remarks’ after Calocar- ides macphersoni. Etymology.—tThe specific name derives from the type locality, the Cape of Good Hope. Calocarides coronatus (Trybom, 1904) Euconaxius coronatus Trybom, 1904:384— 390, pl. 20, figs. 1-10, 13, 14, pl. 21, figs. 1-8. VOLUME 109, NUMBER 1 7 Fig. 2. Calocarides capensis, new species: A, Carapace in lateral view, scale = 5 mm; B, Anterior carapace in dorsal view; C, Abdomen in lateral view; D, Telson in dorsal view; E, Pleopod 1, female; E Pleopod 2, female; G, Pereopod 1, smaller cheliped; H, Pereopod 1, larger cheliped; I, Maxilliped 3; J, Pereopod 2. 58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Calocaris (Calocarides) coronatus.—Wol- leback 908-35 ee loo: Axiopsis (Calocarides) coronatus.—De Mans 1925:6 3677 aile Calocarides coronatus.—Bjorck, 1913:7.— Balss, 1925:209.—Poulsen, 1940:208, 216, fig. 4.—Bouvier, 1940:97.—Chris- tiansen, 1955:1-4; 1972:40, fig. 45.— Elofsson, 1959:3—9, figs. 1-20.—Bratte- gard, 1966:45-52.—Sakai & de Saint Laurent, 1989:4, 80, 101, fig. 20. Euconaxius crassipes Trybom, 1904:390— Sl, , ZO, iiss, Wl, WZ. Calocaris (Calocarides) crassipes.—Wol- lebaek., 1908:35 5-135 17, 235 ple 1-7 Axiopsis (Calocarides) crassipes.—De Man, 1925:6, 67, 71. Material examined.—UBZM 50400, 6 cl 14.8 mm, Sorfjord, Norway, 60°11’N, 6°34'E, 589-600 m.—UBZM 49598, 4 ¢ cl 13.5, 13.9, 15.5, 15.5 mm, Kvinnherad- fjord, Norway, 59°56'N, 5°45’E, 512 m.— UBZM 49595, ¢ cl 14.0 mm, Kvinnherad- fjord, Norway, 59°59'N, 5°53’E, 660-670 m.—UBZM 50461, ovigerous @ cl 14.4 mm, Osafjord, Norway, 60°40'N, 6°55’E, 300.—UBZM 49589, Alfjord, Norway, 59°41'N, 5°34’E, 450-465 m.—UBZM, ovigerous © cl 16.1 mm, Fusafjord, Nor- way, 60°12’N, 5°45’E, 240 m.—UBZM 49591, 3 cl 15.1, 2 cl 15.2 mm, Bomla- fjord, Norway, 59°40'N, 5°22’E, 360 m. Distribution.—From Malangenfjord, northern Norway, to southwestern Sweden, 80-700 m. Diagnosis.—Carapace surface smooth. Rostrum with 4 pairs of lateral teeth, con- tinuous with lateral carina bearing 3 teeth; median carina with | tooth anterior to tu- bercle; submedian carina bearing 2 teeth. Abdominal pleuron 1 ventrally narrowly rounded; pleuron 2 broadly rounded; pleura 3-5 rounded, with | or 2 small teeth on anterior margin. Telson with 5 or 6 lateral teeth, 2 mobile spines at posterolateral an- gle; posterior margin evenly convex, with median tooth; 1 or 2 small spines proxi- modorsally on rounded ridge. Antennal scaphocerite reaching distal margin of peduncle article 4. Maxilliped 3, merus with 4 spines on posterior margin, 3 more distal spines large; carpus with small distal spine on posterior margin. Pereopod 1, larger cheliped: ischium with row of acute tubercles on posterior margin, distal- most a large spine; merus with posterior surface expanding distally, mesial and lat- eral margin defined by row of small acute tubercles, mesial margin with strong spine at about midlength, anterior margin bearing 1 strong spine and few blunt tubercles; car- pus bearing small tubercles dorsodistally; fingers slightly more than half length of propodal palm, latter with lateral and most of mesial surface tuberculate, fixed finger with triangular tooth at about midlength of cutting edge; fingers widely gaping, dacty- lus downcurved, strongly tuberculate, with 1 or 2 stronger tubercles on cutting edge. Smaller cheliped: ischium, merus, and car- pus as in larger cheliped; propodal palm with lateral and mesial surface tuberculate, tubercles along anterior margin forming spines, fingers about % length of palm, fixed finger with blunt proximal triangular tooth followed by regularly spaced small teeth; dactylus with row of spines along an- terior margin, cutting edge with regularly spaced small teeth. Pereopod 2, ischium and basis each with single posterodistal spine; merus with 3 evenly spaced spines on posterior margin. Lateral uropodal ra- mus with about 18 spines along suture, 2 small distal spines on lateral margin; mesial ramus with single distolateral spine, dorsal ridge bearing 5 or 6 spines. Calocarides macphersoni, new species Biss 3 Calastacus longispinis.—Macpherson, 1983:45, fig. 26; 1991:405, non Mc- Ardle, 1901. Material examined.—Holotype, USNM 243561, 6 cl 20.4 mm, R/V Benguela II sta 66, off Namibia, 20°55’S, 12°23’E, 475— 480 m, 9 Sep 1980.—Paratypes, USNM VOLUME 109, NUMBER 1 Fig. 3. in dorsal view; C, Pereopod 2; D, Maxilliped 3; E, Pereopod 1, larger cheliped; EK Pereopod 1, smaller cheliped. 243562, 3 cl 17.8 mm, R/V Benguela II sta 66, off Namibia, 20°55’S, 12°23’E, 475— 480 m, 9 Sep 1980.—Paratype, ICMB, R/ V Benguela II sta P-37, 3 cl 21.5 mm, off Namibia, 21°33’S, 12°39’E, 380-390 m, from stomach of scorpaenid fish Helicolen- 59 Calocarides macphersoni, new species: A, Lateral view, scale = 5 mm; B, Telson and right uropod us Ssp.—Paratypes, ICMB, R/V Benguela II sta P-66, 2 5 cl 20.3 mm, one damaged, off Namibia, 20°55’S, 12°23’E, 475-480 m.—Paratypes, ICMB, R/V Benguela IV sta P-95, 2 6 cl 21.0 mm, 21.2 mn, off Namibia, 20°42’S, 12°14’E, 410-417 m, 60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON one specimen from stomach of ray Raja confundens, one from stomach of scorpaen- id fish Helicolenus sp. Diagnosis.—Rostrum with 2-3 pairs of lateral teeth; median carina with single tooth anterior to tubercle, entire posterior to tu- bercle; submedian carina poorly defined, with 1 or 2 teeth; lateral carina with 3—5 teeth posterior to supraocular. Carapace sur- face smooth. First abdominal pleuron ven- trally narrowly rounded, pleura 2—6 broadly rounded, small spine on anterior margin of pleura 3—6. Telson with 5 small fixed spines in posterior half of lateral margin, 3—4 small mobile spines mesial to posteriormost lateral spine; posterior margin gently convex be- tween lateral spine and median spine; dorsal surface with one pair of small fixed spines. Maxilliped 3, posterior margin of ischi- um unarmed; merus with 3-4 strong distal spines and 2—3 tiny more proximal spines on posterior margin; carpus with small pos- terodistal spine. Pereopod 1: larger chela, ischium with row of small denticles and single strong spine along posteromesial margin, small distal spine on anterior mar- gin; merus with posteromesial surface tu- berculate, having single strong tooth at about midlength, anterior margin with about 5 small spines and several small tu- bercles; outer and upper surface of carpus evenly tuberculate; propodal palm about /, longer than fingers, entire surface evenly and fairly densely tuberculate, posterior margin with tuberculate carina extending onto fixed finger, anterior margin with row of somewhat larger tubercles; fingers prox- imally widely gaping, distally meeting, fixed finger with proximal half of cutting edge having few low rounded tubercles, distal half straight, bearing numerous small rounded teeth; surface of dactylus tubercu- late, cutting edge as in fixed propodal fin- ger. Smaller chela, ischium with few small spines and single strong spine on posterior margin, single distal spine on anterior mar- gin; merus with row of small tubercles along posteromesial margin with | or 2 strong spines at about midlength, row of small tubercles along posterolateral margin, anterior margin with row of irregularly large and small spines and tubercles; outer and upper surface of carpus fairly densely tuberculate; propodal palm about Y; longer than fingers, posterolateral carina feebly tu- berculate, extending almost to apex of fixed finger, outer and upper surfaces tuberculate, anterior margin with double row of large and small spines and tubercles; cutting mar- gins of fixed propodal finger and dactylus only slightly excavate proximally, remain- der of margin bearing row of small evenly- spaced teeth. Pereopod 2, posterior margin with row of small spines and single strong distal tooth; posterior margin of merus with 2—4 spines. Lateral uropodal ramus with 2 or 3 small distal spines on lateral margin, single mobile spine mesial to distalmost, transverse suture armed with row of 13—15 small spines; mesial ramus with single dis- tal spine, dorsal ridge with irregular row of about 8 small spines. Remarks.—From McArdle’s (1901) de- scription, and Alcock & McArdle’s illustra- tions (1902, pl. 57, figs. 2, 2a), and De Man’s (1925:118) key, the Bay of Bengal species Calastacus longispinis lacks spines on the anterior median carina, has five spines on the free lateral margins of the ros- trum, and three spines on the submedian ca- rina. The antennal scaphocerite reaches al- most to the end of peduncle article 4. The single female lacked both chelipeds of pe- reopod 1. Given that the Namibian material has a strong spine anterior to the tubercle on the median carina, and that the antennal scaphocerite reaches only to the distal third of peduncle article 4, it seems unlikely that the Atlantic and Indian Ocean specimens are the same species, especially considering the considerable geographical separation. The species described above as Calocar- ides capensis, differs from C. macphersoni in having more slender and less granular chelipeds of pereopod 1, with relatively more elongate spines on the anterior margin of the propodal palm; in having fewer but stronger distal spines on the anterior margin VOLUME 109, NUMBER 1 of the merus of pereopod 1; in having far fewer spines on the suture of the lateral ra- mus of the uropod; and in having a more elongate telson. Major differences between C. coronatus. and C. macphersoni are seen in the size (cl ovigerous @ 13.5—-16.1 mm vs. cl ovigerous 2 17.8-25.0 mm resp.), and in the more spinose upper margin of the merus in C. macphersoni. The species described as Axius (Neaxius) laevis by Bouvier (1915) from 698 m off Cape Bojador, Spanish Sahara, West Africa, (holotype, MNHN, @ cl 5.1 mm) was syn- onymized with C. coronatus by Sakai & de Saint Laurent (1989:81, fig. 21). The latter authors illustrated the single damaged spec- imen of C. laevis, the caption of their illus- tration erroneously referring to Eiconaxius laevis. Given the damaged condition of the specimen, and the fact that it lacks spines on the submedian carina, has an antennal scaphocerite shorter than in C. coronatus, has pereopod 1 lacking the fine tubercula- tion of the propodus and the anterior spi- nation of the merus of C. coronatus, and a telson longer and narrower than in C. co- ronatus, it seems unlikely that the West Af- rican species is Synonymous with C. coron- atus, or indeed, that it is a species of Cal- ocarides. From the figure of Calocarides rudolfi (Zarenkov, 1989, fig. 3) from An- gola, several differences in spination sug- gests that it differs from C. macphersoni. In C. rudolfi, the merus of maxilliped 3 bears more elongate and numerous spines, and the merus of both chelae of pereopod 1 has larger and more numerous anterodistal and posterodistal spines than in the Namibian species; the carpus of pereopod 2 bears a posterodistal spine, and the telson bears two pairs of fixed dorsal spines, while C. mac- phersoni lacks a carpal spine on pereopod 2 and the telson bears a single pair of fixed spines. Posterolateral mobile spines on the telson were not illustrated for the Angolan species; C. macphersoni bears four such spines on each side. Etymology.—The species is named for 61 carcinologist Dr. Enrique Macpherson of Barcelona, Spain, who made the material from Namibia available for study. Calocarides quinqueseriatus (Rathbun, 1902) Figs. 4, 5 Calastacus quinqueseriatus Rathbun, 1902: 887.—Schmitt, 1921:113, fig. 76.— Balss, 1925:209. Calocaris (Calastacus) quinqueseriata.— De Man, 1925:8, 118. Calocarides quinqueseriatus.—Sakai & de Saint Laurent, 1989:79, 103. Material examined.—Syntypes, USNM 25240, 9 3 cl 15.5—22.7, 2 ovigerous @ cl 15.4, 18.2, 3 2 cl 17.5-19.4, 19.4, R/V AI- batross sta 3196, San Luis Obispo Bay, California, green mud, 366 m.—USNM 28321, 5 6, cl 13.5—21.6, 1 ovigerous 2 cl 16.9 mm, 9 8 cl 15.1—20.5, R/V Albatross sta 2909, Santa Barbara Channel, Califor- nia, 375 m, green mud.—USNM 152525, 22 3 cl 18.9-26.0, 9 ovigerous 2 20.0- 23.3, 11 & cl 21.0—23.0, R/V Albatross sta 4436, San Miguel Island, California, 483— 496 m, green mud.—USNM 28325, 16 6, cl 11.3-18.0, 1 ovigerous @ cl 24.0, 7 2 cl 10.2-19.2, R/V Albatross sta 3195, San Luis Obispo Bay, California, 461 m, green mud.—USNM 28236, 5 6 cl 14.8—24.0, 4 2 cl 13.0—22.5, R/V Albatross sta 3198, off Point Conception, California, 508 m, green mud.—USNM 28327, 7 63 cl 12.9-20.5, 3 ovigerous ¢ cl 19.0—20.0, 6 2 cl 10.3—19.5, R/V Albatross sta 3199, Santa Barbara Channel, California, 426 m, green mud.— USNM 28232, ¢ cl 18.0, 2 2 cl 21.0, 21.9 R/V Albatross sta 3187, off Point Sur, Cal- ifornia, 545 m, brown and grey ooze.— WSINME233205 6 cl 20 3592 cl 7 2RIN Albatross sta 2892, off Point Conception, California, 520 m, yellow mud.—USNM 28528, 2 6, cl 18.1, 24.1, R/V Albatross sta 3200, Santa Barbara Channel, Califor- nia, 485 m, green mud.—USNM 28319, 1 3d cl 19.0, 1 2 cl 12.1, R/V Albatross sta 2891, off Point Conception, California, 426 62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Calocarides quinqueseriatus: A, Lateral view, scale = 10 mm; B, Pereopod 1, larger cheliped; C, Pereopod 1, smaller cheliped; D, Telson and right uropod in dorsal view; E, Pleopod 2, male; EK Pleopod 3, male; G, Pleopod 1, female; H, Pleopod 2, female. m, brown ooze.—USNM 120119, 2 cl 18.2, R/V Commando sta, mouth of Colum- bia River, Oregon, 732 m.—USNM 28324, 6 cl 17.5, 2 cl 15.0, R/V Albatross sta 3193, off San Simeon Bay, California, 293 m, green mud.—USNM 28329, ovigerous 2 cl 19.5, R/V Albatross sta 3201, Santa Barbara Channel, California, 512 m.— USNM 28322, ¢ cl 17.4, R/V Albatross sta 2979, off Anacapa Island, California, 710 m, green mud.—USNM 53021, 2 cl 16.2, R/V Albatross sta 4425, off San Nicolas Is- land, California, 2013 m, green mud, fine sand, globigerina ooze.—USNM 155735, 6 cl 20.5, R/V Albatross sta 4387, off San Diego, California, 1830 m, green mud.— USNM 243350, ovigerous 2 damaged, R/ V Commando sta, SW mouth of Columbia River, Oregon, 1556 m.—USNM 243351, 3 cl 18.4, R/V Commando sta, SW mouth VOLUME 109, NUMBER 1 A B E pa Fig. 5. L Calocarides quinqueseriatus, variation in rostral and anterior carapace spination: A, Syntype, ovi- gerous female, San Luis Obispo Bay, California, cl 15.4 mm; B, Syntype, ovigerous female, San Luis Obispo Bay, California, cl 18.2 mm; C, Ovigerous female, San Miquel Is., California, cl 22.0 mm; D, Ovigerous female, San Miguel Is., California, cl 23.3; E, Female, Baja California, cl 12.0 mm; FE Female, Baja California, 20.7 mm; G, Male, Columbia River mouth, Oregon, 18.2 mm; H, Syntype, male, San Luis Obispo Bay, California, cl 19.1 mm; I, Syntype, male, San Luis Obispo Bay, California, cl 20.9 mm; J, Male, San Miguel Is., California, 24.2 mm; K, Male, San Miguel Is., California, cl 26.0 mm; L, Male, Columbia River, Oregon, cl 27.2 mm. of Columbia River, Oregon, 1373 m.— USNM 243352, 2 damaged, R/V Comman- do sta, SW mouth of Columbia River, Or- egon, 1556 m.—USNM 243353, ¢ cl 17.3, 2 2 damaged, R/V Commando sta, SW mouth of Columbia River, Oregon, 1922 m.—USNM 243354, 2 damaged, R/V Commando sta, SW mouth of Columbia River, Oregon, 1556 m.—USNM 155737, 2 cl 20.0, R/V Albatross sta 5675, San Cristobal Bay, Baja California, 520 m.— USNM 155736, 2 ¢ cl 19.8—21.0, R/V AI- batross sta 4433, off Santa Rosa Island, California, 445 m, green mud.—USNM 243355, 216) Cl G15; 2) 2 cl N3:8—15205 R/ V Albatross sta 4351, NE of Point Loma, California, 774 m, soft green mud. Diagnosis.—Carapace smooth; 5—6 pairs rostral teeth including small supraocular; median carina with 1—4 spines anterior to tubercle, 1—2 posterior; 1—5 spines on sub- median carina; 3—7 spines on lateral carina. Abdominal pleuron | with ventral spine usually present; pleuron 2 ventrally round- ed; pleura 3—6 rounded with spine on an- terior margin. Telson with lateral margin 64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON having 5—6 small spines; | pair spines on dorsal surface. Maxilliped 3, ischium with about 10 small spines on posterior margin; merus with 5—6 small proximal spines, 3 large dis- tal spines on posterior margin. Pereopod 1, larger (broader) cheliped: ischium with 1 strong distal and several small proximal spines on posterior margin; merus with in- ner and outer row of tiny tubercles flanking posterior row of about 17 strong spines, outer surface distally with few scattered tiny tubercles, upper (anterior) margin with irregular row of spines, distal spines largest; carpus with outer surface and distal margin bearing scattered acute tubercles; upper propodal palm subequal in length to fingers, posterior margin of palm and fixed finger a denticulate carina, upper (anterior) margin with about 14 strong spines, mesial and lat- eral surfaces of palm fairly densely granu- lar/tuberculate, fixed finger with broad tu- bercle on proximal cutting edge, distally denticulate; dactylus cutting edge with proximal tubercle, distally denticulate, up- per margin with several proximal spines. Pereopod 1, smaller (narrower) cheliped: is- chium, merus, and carpus as in larger che- liped; upper propodal palm subequal in length to fingers, tuberculation as in larger cheliped, cutting edge of fixed finger with few small tubercles proximally, remainder of edge finely denticulate; dactylus, cutting edge finely denticulate, with about 2 spines on upper margin. Pereopod 2, ischium with 6 small spines on posterior margin; merus with 5 small proximal spines, single strong distal spine. Pleopod 1 in female a slender elongate ramus; pleopod 2 in male with rod-shaped appendix masculina bearing dis- tal setae, slightly shorter than appendix in- terna; pleopods 2—5 in female, 3—5 in male with appendix interna at about proximal third of mesial margin of endopod. Uropod with lateral ramus having 2 small distal spines on lateral margin, single mobile sub- marginal spine; about 14 spines along su- ture; mesial ramus with single distolateral spine, 4 spines on dorsal surface. Size ranges.—Male, maximum carapace length 26.0 mm; ovigerous females, cara- pace length 15.4—24.0 mm. Remarks.—There is considerable varia- tion in the spination of the anterior carapace (Fig. 5) and the chelipeds of pereopod 1 of this species, often made more marked either by injuries to the chelae that have healed, or by loss and regeneration of the chelipeds, resulting in a range of cheliped proportions and armature. The majority of specimens have been taken from soft bottom sediments, especial- ly from green mud. The species recorded as Calastacus quin- queseriatus from 1150 m off Japan (Kob- jakova 1937:142, pl. 2, fig. 8) appears to have the first pereopod chelae relatively shorter, more robust, and less granular than in true C. quinqueseriatus. It thus seems un- likely that the Japanese species is conspe- cific with the California/Oregon species. Calocarides rostriserratus Andrade & Baez, 1977 ica Calastacus rostriserratus Andrade & Baez, IST VLES, We. I Calocarides quinqueseriatus.—Sakai & de Saint Laurent, 1989:78, 79, 103, non Rathbun, 1902. Material examined.—USNM 173366, 4 6 cl 153 mm, 153 5mm tos enim we0 mm, 2 @ cl 14.5 mm, 15.9 mm, off Tum- bes, Peru, 3°33.8’'S, 81°01.4’E, 576 m, 28 Aug 1979.—USNM 243392, ¢ cl 16.1 mm, R/V Gilliss sta GS-22, Gulf of Panama, 7T28'N, 7912 W;, 825 m, ts Janeio7es Diagnosis.—Carapace smooth; rostrum with 5—6 pairs of lateral teeth including small supraocular; 2 median carina teeth plus tubercle; 4—5 teeth on submedian ca- rina; 4—6 teeth on lateral carina. Abdominal pleura 2—5 ventrally rounded. Telson with 2-3 pairs small lateral spines, 2 small distal mobile spines at posterolateral angle, 2 pairs spines on dorsal surface. Maxilliped 3, ischium with 5 small prox- VOLUME 109, NUMBER 1 Fig. 6. 65 wi, : S X ' Calocarides rostriserratus: A, Lateral view, scale = 5 mm; B, Anterior carapace in dorsal view; C, Telson and right uropod in dorsal view; D, Pereopod 1, larger cheliped; E, Pereopod 1, smaller cheliped; EF Pleopod 1, female; G, Pleopod 2, female; H, Pleopod 2, male; I, Pleopod 3, male. imal teeth on posterior margin, distal mar- gin entire; merus with 2 strong distal teeth on posterior margin. Pereopod 1, larger (broader) cheliped, ischium with about 5 small proximal and single strong distal Spine on posterior margin; merus with 5 distal spines on upper (anterior) margin, posterior margin with single row of spines of varying length, those at about midlength longest; carpus with several (6—7) small spines on upper surface; upper propodal palm 1.2 times length of fingers, bearing row of 8 strong spines, lower border con- sisting of flattened distally narrowed ridge having dentate margins, medial and lateral surfaces bearing moderately dense scattered conical tubercles, cutting edge of fixed fin- ger having rounded proximal cusp; dactylus with upper margin bearing about 8 strong spines, cutting edge having rounded proxi- mal cusps, distally finely denticulate. Pereo- pod 1, smaller (narrower) cheliped, ischium 66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with 2—3 small proximal and single strong distal spine on posterior margin; merus with 5—6 strong distal spines on anterior (upper) margin, single row of spines along posterior (lower) margin, those at about midlength longest; carpus with 8-10 spines on upper surface; upper propodal palm about 1.2 times length of fingers, bearing irregular row of about 12 spines, lower border formed by flattened ridge with dentate mar- gins, inner and outer surfaces bearing scat- tered acute conical tubercles; dactylus with several small spines on upper surface, cut- ting edge more or less evenly denticulate. Pereopod 2, basis with single spine on pos- terior surface; ischium with proximal and one or 2 stronger distal spine on posterior margin; merus with 3 or 4 spines in mid- region of posterior margin. Pleopod 1 in fe- male a slender elongate ramus; pleopod 2 in male with rod-shaped appendix mascu- lina bearing distal setae, noticeably shorter than appendix interna; pleopods 2—5 in fe- male, 3—5 in male with appendix interna at about proximal third of mesial margin of endopod. Uropod with lateral ramus having 2—3 small lateral spines plus distal submar- ginal mobile spine; about 13 spines along suture; mesial ramus with single distal mar- ginal spine, 4—5 spines on upper surface. Remarks.—In spite of Sakai & de Saint Laurent’s (1989:80) statement that C. ros- triserratus 1s probably a synonym of C. quinqueseriatus, Calocarides rostriserratus differs from C. quinqueseriatus, its geo- graphically nearest congener, in several eas- ily seen features: in having two pairs of dorsal telsonic spines (one pair in C. quin- queseriatus), and two or three lateral tel- sonic spines (five or six in C. quinqueser- iatus); and in having the chelae of pereopod 1 broader and less elongate, and with spines only in the distal half of the anterior margin of the merus (full anterior margin of merus bearing spines in C. quinqueseriatus). Calocarides soyoi (Yokoya, 1933) Axius soyoit Yokoya, 1933:49, fig. 25.— Horikoshi et al., 1982:30, 33, 38, 39, 120, 145, 168. Axiopsis (Axiopsis) soyoi.—Sakai, 1987: 303. Calocarides soyoi.—Sakai & de Saint Lau- rent, 1989:4, 83, 103.—Kensley & Ko- mai, 1992:81, fig. 1. Material examined.—USNM 243563, 2 2 cl 12.0 mm, 15.0 mm, ovigerous ¢ cl 17.9 mm, off Fukushima Prefecture, Japan, 37°04'N, 141°31.7’E, 270 m.—USNM 243564, 3 cl 14.2 mm, ovigerous ¢ cl 18.8 mm, 2 cl 11.8 mm, off Hachinohe, Aomori Prefecture, Japan —HUMZ-C990, 3 @ cl 11.3 mm, 14.1 mm, 13.0 mm, off Hachi- nohe, Aomori Prefecture, Japan, 42°52.6’N, 145°22.2'E, 152 m.—HUMZ-C994, 2 el 16.9 mm, off Fukushima Prefecture, Japan, 37°17.3'N, 141°21.4’E, 141 m—HUMZ- C998, 3 6 cl 12.4 mm, 15.0 mm, 15.3 mm, off Fukushima Prefecture, Japan, 37°04’N, 141°31.7’E, 270 m. Remarks.—This species was recently re- described and figured by Kensley & Komai (1992). Calocarides spinulicauda (Rathbun, 1902) Fig. 7 Axius spinulicauda Rathbun, 1902:886; 1904:149, fig. 90. Axiopsis spinulicauda.—Schmitt, 1921:111, fig. 74.—Butler, 1961:60, figs. 1, 2, pl. 1. Axiopsis (Axiopsis) spinulicauda.—De Man, 1925:6, 67, 69. Acanthaxius spinulicaudus.—Sakai & de Saint Laurent, 1989:4, 66, 103. Material examined.—Holotype, USNM 25239, 2 cl 15.5 mm, R/V Albatross sta 3172, off Bodega Head, California, 113 m. Diagnosis.—Carapace surface smooth; rostrum with 5 (left) and 6 (right) lateral teeth, including supraocular; median carina barely reaching onto base of rostrum, with 3 spines anterior of tubercle, entire poste- rior to tubercle; postcervical median carina obsolete except close to posterior margin of carapace; submedian carina short, entire; lateral carina entire posterior to supraocular VOLUME 109, NUMBER 1 67 Fig. 7. C, Anterior carapace in dorsal view; D, Telson and right uropod in dorsal view. spine of rostral series. First abdominal pleu- ron ventrally narrowed apically rounded with tiny denticle; pleura 2—6 broadly rounded. Telson with lateral margin bearing larger fixed anterior spine, 4-5 small more posterior fixed spines; posterior margin evenly convex; submedian longitudinal ridge bearing 3 fixed spines, pair of fixed anterior spines mesial to longitudinal ridge present. Maxilliped 3, ischium with 1 or 2 tiny denticles on posterior margin; merus with 3 Calocarides spinulicauda: A, Carapace in lateral view, scale = 5 mm; B, Abdomen in lateral view; distal spines on posterior margin. Pereopod 1, both chelipeds missing. Pereopod 2, me- rus with 2 or 3 well separated spines on posterior margin. Lateral uropodal ramus with 7 small distal spines on lateral margin, transverse suture with row of 11—12 small spines; mesial ramus with 2 strong spines on lateral margin, dorsal ridge with 7 or 8 small fixed spines. Remarks.—Rathbun (1902), in her de- scription of A. spinulicauda, noted that the chelae of the first pereopod were missing. 68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Butler (1961) figured both chela of pereo- pod 1, from material from British Colum- bia. While the lateral and mesial faces of the propodal palm do not appear to be tu- berculate, as is characteristic of Calocari- des, the spination of the dactyli, propodi, carpi, and meri are characteristic of this ge- nus. Calocarides spinulicauda, however, disagrees with Calocarides in lacking spi- nation on the lateral and submedian carinae. Sakai & de Saint Laurent (1989) placed this species in their new genus Acanthaxius, which is characterized by a tendency to having a double cornea, and a lack of pleu- robranchs, neither of which are seen in C. spinulicauda. With only the female holo- type available for examination, the nature of the male pleopods and the first pereopods being uncertain, unambiguous generic placement is almost impossible. Axius spinulicauda amurensis Kobjako- va, 1937, (see also Derjugin & Kobjakova 1935:142), described from the Sea of Japan, while clearly not a subspecies of the Cali- fornian Calocarides spinulicauda Rathbun, 1902, is difficult to place generically, given the single small figure and the abbreviated description. Acknowledgments I thank Dr. Enrique Macpherson of the Instituto Ciencias del Mar, Barcelona, Mrs. Michelle van der Merwe of the South Af- rican Museum, and Dr. A. Fosshagen of the Zoological Museum, Bergen, for the loan of material. The paper was considerably im- proved by the comments and suggestions of Dr. Rafael Lemaitre and anonymous re- viewers, to whom I am very grateful. Literature Cited Alcock, A., & A. EF McArdle. 1902. Illustrations of the zoology of the Royal Indian Marine Survey ship Investigator, under the command of Com- mander T. H. Heming, RN. Crustacea, part 10, plates 56-67. Calcutta: Office of the Superin- tendent of Government Printing, India. Andrade, H., & P. Baez. 1977. Calastacus rostriser- ratus n. sp. (Crustacea, Decapoda, Macrura, Ax- iidae).—Anales del Musec de Historia Natural de Valparaiso 10:65—67. Balss, H. 1913. Diagnosen neuer ostasiatischer Ma- cruren.—Zoologischen Anzeiger 62(5):234— 25% . 1925. Macrura der Deutschen Tiefsee-Expe- dition. 1. Palinura, Astacura und Thalassinid- ea.—Wissenschaftliche Ergebnisse der Deutsch- en Tiefsee-Expedition auf dem Dampfer “Val- divia” 1898-1899, 20(4):189-216. . 1957. Decapoda.—Dr. H. G. Bronn’s Klassen und Ordnungen des Tierreichs 5(1)7(12):1505— 1672. Bamard, K. H. 1950. Descriptive catalogue of South Af- rican decapod Crustacea (Crabs and Shrimps).— Annals of the South African Museum 38:1—837. Bate, C. S. 1888. Report on the Crustacea Macrura collected by H.M.S. Challenger during the years 1873—76.—Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873-76, Zoology 24:i-xc, 1-942. Bjorck, W. 1913. Decapoden aus dem Kattegatt und aus dem Skagerak.—Arkiv for Zoologi 8(3):1— 12. Bouvier, E. L. 1915. Thalassinidés nouveaux capturés au large des cétes Soudanaises par le ““Talis- man.’—Bulletin du Muséum National d’His- toire Naturelle, Paris 21(6):182—-185. 1940. Décapodes Marcheurs.—Faune de France 37:1—399. Brattegard, T. 1966. Ecological and biological notes on Calocarides coronatus (Crustacea Thalassi- nidea).—Sarsia 24:45—52. Butler, T. H. 1961. Records of decapod Crustacea from British Columbia.—Canadian Journal of Zoology 39:59-62. Christiansen, E. N. 1955. Notes on Calocarides co- ronatus (TIrybom) (Crustacea Decapoda).—As- tarte [1]12:1—5. Derjugin, K. M., & S. I. Kobjakova. 1935. Zur De- kapodenfauna des Japanischen Meeres.—Zool- ogischer Anzeiger 112(5/6):141—-147. Elofsson, R. 1959. A new decapod larva referred to Calocarides coronatus (Trybom).—Universite- tet 1 Bergen, Arbok 1959, Naturvitenskapelig rekke 7:1—10. Gerstaecker, A. 1856. Carcinologische Beitrage —Ar- chiv fiir Naturgeschichte 22(1):101—-162. Gurney, R. 1942. Larvae of Decapod Crustacea. Lon- don, Ray Society. 306 pp. Horikoshi, M., T. Fujita, M. Takeda, M. Okiyama, S. Ohta, E. Tsuchida, & M. Yamamoto. 1982. Preliminary compilation of the results obtained by the “Survey of Continental Shelf Bordering Japan” carried out on board the S/S S6y6-Maru during 1923-1930. In M. Horikoshi, ed., Report for a part of the result of Grant-in-Aid for Co- operative Research (A), Ministry of Education, VOLUME 109, NUMBER 1 Science and Culture: “Studies on Off-shore and Deep-sea Faunas in the West Pacific and Indian Oceans” (Proj. no. 00534033). Ocean Research Institute, University of Tokyo, 252 pp. Kensley, B. 1981. On the zoogeography of southern African decapod Crustacea, with a distributional checklist of the species.—Smithsonian Contri- butions to Zoology 338:1—64. . 1994. The genus Coralaxius redefined, with the description of two new species (Crustacea: Decapoda: Axiidae).—Journal of Natural His- tory 28:813-828. , & R. H. Gore. 1982. Coralaxius abelei, new genus and new species (Crustacea: Decapoda: Thalassinidea: Axiidae): a coral inhabiting shrimp from the Florida Keys and the western Caribbean Sea.—Proceedings of the Biological Society of Washington 103(3):558—572. , & T. Komai. 1992. Redescription of Calocar- ides soyoi (Yokoya, 1933) from Japan (Crusta- cea: Decapoda: Axiidae).—Proceedings of the Biological Society of Washington 105(1):81—85. Kobjakova, S. I. 1937. Systematische Uebersicht der Dekapoden aus dem Ochotskischen und Japan- ischen Meere.—15:93—154. [In Russian, with German summary. ] Macpherson, E. 1983. Crustaceos Decapodos captu- rados en las costas de Namibia.—Resultados Expediciones Cientificas (Supl. Inv. Pesq.) 11: 3-79. 1991. Biogeography and community struc- ture of the decapod crustacean fauna off Na- mibia (Southeast Atlantic)—Journal of Crus- tacean Biology 11(3):401-415. McArdle, A. F 1901. Natural history notes from the Royal Indian Marine Survey Ship ‘Investiga- tor’, Commander T. H. Heming, R. N., com- manding. Series 3, No. 5. An account of the trawling operations during the surveying-season of 1900—1901.—Annals and Magazine of Nat- ural History (7)8:517—526. Man, J. G. de. 1905. Diagnoses of new species of ma- crurous decapod Crustacea from the “‘Siboga-Ex- pedition”.—Tijdschrift der Nederlandsche Dier- kundige Vereeniging (2)9(3/4):587-614. . 1925. The Decapoda of the Siboga-Expedi- tion. Part VI. The Axiidae collected by the Si- boga-Expedition.—Siboga-Expeditie monogra- phie 39a5:1—127. Poore, G. C. B. 1994. A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to families and genera.—Memoirs of the Mu- seum of Victoria 54:79-120. , & D. J. G. Griffin. 1979. The Thalassinidea (Crustacea: Decapoda) of Australia.—Records of the Australian Museum 32(6):217—321. 69 Poulsen, E. M. 1940. On the occurrence of the Thal- assinidea in Danish waters.—Videnskabelige Medelelser, Copenhagen 104:207—239. Rathbun, M. J. 1902. Descriptions of new decapod crustaceans from the west coast of North Amer- ica.—Proceedings of the United States National Museum, 24(1272):885—905. . 1904. Decapod crustaceans of the northwest coast of North America.—Harriman Alaska Ex- pedition 10:1—210. Sakai, K. 1987. Two new Thalassinidea (Crustacea: Decapoda) from Japan, with the biogeograph- ical distribution of the Japanese Thalassinid- ea.—Bulletin of Marine Science 41(2):296—308. 1992. Axiid collections of the Zoological Museum, Copenhagen, with the description of one new genus and six new species (Axiidae, Thalassinidea, Crustacea).—Zoologica Scripta 21(2):157-180. , & M. de Saint Laurent. 1989. A check list of Axiidae (Decapoda, Crustacea, Thalassinidea, Anomura), with remarks and in addition descrip- tions of one new subfamily, eleven new genera and two new species.—Naturalists 3:1—104. Schmitt, W. L. 1921. The Marine Decapod Crustacea of California, with Special Reference to the Decapod Crustacea Collected by the United States Bureau of Fisheries Steamer “‘Albatross”’ in connection with the Biological Survey of San Francisco Bay during the Years 1912-1913.— University of California Publications in Zoolo- gy, 23:1-470, pls. 1-50, 165 text figures. Smith, S. I. 1881. Preliminary notice of the Crustacea dredged, in 64 to 325 fathoms, off the south coast of New England, by the United States Fish Commission in 1880.—Proceedings of the United States National Museum 3:413—452. Stebbing, T. R. R. 1910. General catalogue of South African Crustacea (Part V of S. A. Crustacea, for the Marine Investigations in South Afri- ca).—Annals of the South African Museum 6(4):281-593. Trybom, E 1904. Two new species of the genus Eu- conaxius.—Arkiv for Zoologi 1(4):383-393. Wollebaek, A. 1908. Remarks on decapod crustaceans of the North Atlantic and the Norwegian Fiords (I and II).—Bergens Museums Aarbog 12:1—74. Yokoya, Y. 1933. On the distribution of decapod crus- taceans inhabiting the continental shelf around Japan, chiefly based upon the materials collect- ed by S. S. S6y6-Maru, during the years 1923— 1930.—Journal of the College of Agriculture, Tokyo Imperial University 12(1):1—226. Zarenkov, N. A. 1989. Three new species of Calo- caris Bell (Decapoda Axiidae).—Zoologiskii Jhurnal 68(11):24—30. [In Russian, with English summary. | PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):70—-74. 1996 A new species of the axiid shrimp genus Acanthaxius from the Caribbean (Crustacea: Decapoda: Thalassinidea) Brian Kensley Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Acanthaxius kirkmilleri is described from slope depths (420-440 m) in the Caribbean Sea off Anguilla, and is the first unambiguous record of the genus in the Atlantic. The species is characterized by a slender rostrum longer than the eyestalks, highly spinose and relatively slender chelipeds of pereopod 1, and seven spines on the suture of the lateral uropodal ramus. A single female specimen of a distinctive axiid shrimp was collected by the R/V John Elliott Pillsbury in 1969, off Anguilla. The specimen is described here, as part of a se- ries of papers documenting the diversity of the Axiidae in the western Atlantic, and is deposited in the collections of the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). Family Axiidae Genus Acanthaxius Sakai & de Saint Laurent, 1989 Acanthaxius Sakai & de Saint Laurent, 1989:4, 12, 14, 66.—Poore, 1994:98 [key].—Sakai, 1994:192. Type species.—By original designation, Axiopsis (Axiopsis) pilocheira Sakai, 1987. Type locality: off Honshu Island, Japan, 360 m. Gender: Masculine. Diagnosis.—Gonochoristic. Rostrum at same level as anterior carapace, narrow, lat- eral margins dentate; carapace glabrous to faintly rugose; with median, submedian and lateral carinae dentate; post-cervical carina absent. Eye with cornea pigmented; eye- stalk relatively elongate. Antennal scapho- cerite (acicle) just reaching distal margin of peduncle article 4, slightly arcuate. Maxil- liped 3, exopod not clearly bent; ischial crest well developed. Pereopod 1, chelipeds asymmetrical, with spines on anterior (up- per) margin of propodal palm and dactylus. See Table 1 for branchial formula. Pleopo- dal rami slender; pleopod 1 in female uni- ramous; pleopods 2—5 with appendix inter- na. Lateral uropodal ramus with transverse suture. Telson longer than wide, with dorsal submedian fixed spines (Table 1). Remarks.—The definition of this genus contains some uncertainties, as the type species, A. pilocheira Sakai, 1987, is known only from the holotypic female. Im- portant characters of pleopods 1 and 2 of the male are is thus unknown, although the absence of pleopod 1 was deduced from other species placed in the genus. Three of the species assigned to this genus by Sakai & de Saint Laurent (1989) and Sakai (1994), A. caespitosa (Squires, 1979), A. hirsutimana (Boesch & Smalley, 1972), and A. spinosissimus (Rathbun, 1906), do not fit the generic definition, in that all three have pleopod 1 of the male present, and the car- apace is covered with granules, both fea- tures more characteristic of Oxyrhynchaxius Parisi, 1917. The species A. spinulicauda (Rathbun, 1902), also included in Acan- thaxius Sakai & de Saint Laurent, 1989, is clearly a species of Calocarides Wollebaek, 1908. The use of the post-cervical carina, present in some species, absent in others, also needs reassessment as a diagnostic character. The advanced characters of the genus VOLUME 109, NUMBER 1 Table 1.—Branchial formula for Acanthaxius pil- ocheira. r = reduced. Maxillipeds Pereopods 1 2 3 1 2 3 4 5 Exopod Epipod 1 1 Podobranch — r Arthrobranch — — Pleurobranch — — Setobranch —_—- — would appear to be the relatively slender and dentate rostrum, with its basal pair of spines in a supraorbital position; the form of the chelipeds of pereopod 1, and es- pecially that of the smaller chela with the slender dentate fingers being 1.5—2.0 times longer than the propodal palm; the presence of spines on the upper margins of the dac- tylus and propodus of pereopod | chelae; the absence of pleurobranchs; and the pres- ence of epipods on pereopods 1-4. Species: Acanthaxius amakusana (Miyake & Sakai, 1967). Amakusa Island, Kyushu, Japan, 20—40 m. Acanthaxius kirkmilleri, new species. Off Anguilla, 421-439 m. Acanthaxius miyazakiensis (Yokoya, 1933). Southern Miyazaki-ken, Japan, 137 m; Philippines, 136—210 m. Acanthaxius pilocheira (Sakai, 1987). Off Honshu Island, Japan, 360 m. Acanthaxius polyacantha (Miyake & Sakai, 1967). East China Sea, 118 m. Acanthaxius polychaetes Sakai, 1994. Off Great Barrier Reef, Australia, 260 m. Acanthaxius kirkmilleri, new species lias, I, 2 Material examined.—Holotype, USNM 243492, 2 carapace length 12.0 mm, R/V, John Elliott Pillsbury sta P-984, off An- guilla, 18°26.4’N, 63°12.6'W, 421-439 m, brown mud bottom, 22 Jul 1969. Diagnosis.—Female: Carapace glabrous; 71 rostrum slender, with 2 pairs of strong dor- sal teeth (apex missing); median carina Starting at rostral base, with 2 teeth anterior to tubercle, 1 posterior; submedian carina with 4 teeth; lateral carina with single strong tooth posterior to basal rostral tooth; postcervical carina poorly defined, most marked anteriorly. Abdominal pleuron 1 ventrally narrowed; pleuron 2 broad, ven- trally truncate; pleura 3—5 with tiny denticle on anterior margin, anteroventrally round- ed, posteroventrally slightly angled. Telson with lateral margin having single anterior tooth, 2 mobile posterolateral spines, pos- terior margin convex, with median tooth; dorsal surface with 2 pairs of teeth (poster- 1ormost tooth on right side doubled). Antennal scaphocerite (acicle) slender, acute, reaching to distal margin of peduncle article 4. Maxilliped 3, ischium with 3 teeth on posterior margin; merus with 5 teeth in- creasing in length distally on posterior mar- gin. Pereopod 1, larger chela, ischium with 4 teeth on posterior margin; merus with 7 teeth and several smaller denticles on pos- terior margin, 5 strong distal teeth on an- terior margin; carpus with 3 strong teeth on anterior margin, several smaller denticles on lateral surface, including strong submar- ginal tooth; fingers subequal in length to propodal palm, propodus strongly setose on upper and lower surfaces, with 4 strong teeth on anterior margin, row of 12 sub- marginal teeth on ventrolateral surface, lat- eral surface of palm with many small acute denticles; fixed finger with cutting edge bearing numerous small rounded teeth; dac- tylus bearing 5 strong teeth on upper mar- gin, cutting edge bearing numerous small rounded teeth. Smaller chela, ischium, me- rus, and carpus as in larger chela; fingers about % longer than propodal palm, latter setose, with 4 strong teeth on upper margin, row of about 10 submarginal teeth ventral- ly, lateral surface with several small acute denticles, fixed finger with cutting edge straight, bearing numerous small serrations; dactyl bearing 4 strong teeth on upper mar- gin, cutting edge straight, bearing numerous 72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Anterior carapace in dorsal view; C, abdomen in lateral view; D, telson and right uropod in dorsal view. Scale = 2 mm. small serrations. Pereopod 2, anterior and posterior margins of all articles setose; is- chium with single strong posterodistal tooth; merus with 3 strong teeth on poste- rior margin; carpus with single strong tooth at midlength of anterior margin. Pereopod 3, merus with 3 teeth on posterior margin; propodus with 5 small clumps of short spines on posterior margin. Pereopod 4, merus with single small posterodistal tooth; dactylus setose; propodus with 4 small clumps of tiny spines on posterior margin. Acanthaxius kirkmilleri, new species, holotype (USNM 243492). A, Carapace in lateral view; B, Pereopod 5, propodus with distolateral clus- ter of setae; dactylus setose. Pleopod 1 a single slender ramus, distally flagelliform. Pleopods 2—5 with slender appendix interna articulating at about proximal third of en- dopod. Uropod with lateral ramus having 2 teeth on lateral margin, mobile spine at an- gle of suture, 6 spines along suture; mesial ramus with 2 teeth on lateral margin, 4 teeth on dorsal ridge including distal marginal tooth. Remarks.—The major differences be- VOLUME 109, NUMBER 1 73 G Fig. 2. Acanthaxius kirkmilleri, new species, holotype (USNM 243492). A, Pereopod 1, larger cheliped; B, Pereopod 1, smaller cheliped; C, Pereopod 2; D, Pereopod 3; E, Pereopod 4; EK Pereopod 5; G, Pleopod 1; H, Pleopod 2. Scale (A-F) = 2 mm. tween this western Atlantic species and the holotype of A. pilocheira (USNM 231418) lie in the lateral ramus of the uropod (lateral margin more spinose than A. kirkmilleri), the generally more spinose and slightly more robust chelipeds of pereopod 1 in A. pilocheira, and the more elongate telson (1.5 times longer than basal width) in A. kirkmilleri (1.25 times longer than basal width in A. pilocheira). The differences between the present spe- cies and the other Japanese congeners are easily discerned: A. miyazakiensis is a far more setose species, especially on the che- lipeds and carapace; A. polyacantha pos- sesses a markedly tuberculate carapace, and relatively more robust chelipeds; A. amak- usana has a rostrum shorter than the eye- stalks, and a relatively broader telson. The Australian species A. polychaetes is more 74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON setose, especially on the carapace and ab- domen, and possesses squatter and more se- tose chelipeds of pereopod 1 Etymology.—The species is named for Dr. Kirk D. Miller, longtime friend and companion. Acknowledgments I am grateful to Dr. Rafael Lemaitre for reading the manuscript, and making useful suggestions for its improvement. Literature Cited Boesch, D. F, & A. E. Smalley. 1972. A new axiid (Decapoda, Thalassinidea) from the northern Gulf of Mexico and tropical Atlantic.—Bulletin of Marine Science 22(1):45—52. Miyake, S., & K. Sakai. 1967. Two new species of Axiidae (Thalassinidea, Crustacea) from the East China Sea.—Journal of the Faculty of Ag- riculture, Kyushu University 14(2):303-310. Parisi, B. 1917. I Decapodi giapponesi del Museo di Milano. V. Galatheidea e Reptantia.—Atti della Societa Italiana di Scienze Naturali 56:1—24. Poore, G. C. B. 1994. A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to the families and genera—Memoirs of the Museum of Victoria 54:79—120. Rathbun, M. J. 1902. Descriptions of new decapod crustaceans from the west coast of North Amer- ica.—Proceedings of the United States National Museum 24(1272):885—905. . 1906. The Brachyura and Macrura of the Ha- waiian Islands.—U.S. Fish Commission Bulle- tin for 1903, Part 3:827—930. Sakai, K. 1987. Two new Thalassinidea (Crustacea: Decapoda) from Japan, with the biogeograph- ical distribution of the Japanese Thalassinid- ea.—Bulletin of Marine Science 41(2):296—308. . 1994. Eleven species of Australian Axtidae (Crustacea: Decapoda: Thalassinidea) with de- scriptions of one new genus and five new spe- cies.—The Beagle, Records of the Museums and Art Galleries of the Northern Territory 11: 175-202. , & M. de Saint Laurent. 1989. A check list of Axiidae (Decapoda, Crustacea, Thalassinid- ea, Anomura), with remarks and in addition de- scriptions of one new subfamily, eleven new genera and two new species.—Naturalists 3:1— 104. Squires, H. J. 1979. Axiopsis caespitosa (Thalassinid- ea, Axiidae), a new species from the Pacific coast of Colombia.—Canadian Journal of Zo- ology 57(8):1584—1591. Wollebaek, A. 1908. Remarks on decapod crustaceans of the North Atlantic and the Norwegian Fiords (I & IL).—Bergens Museums Aarbog 12:1-—74. Yokoya, Y. 1933. On the distribution of decapod crus- taceans inhabiting the continental shelf around Japan, chiefly based upon the materials collect- ed by S. S. Sd6y6-Maru, during the year 1923— 1930.—Journal of the College of Agriculture, Tokyo Imperial University 12(1):1—226. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):75—90. 1996 A new genus for Anapagrides sensu De Saint Laurent-Dechancé, 1966 (Decapoda: Anomura: Paguridae) and descriptions of four new species Patsy A. McLaughlin and Janet Haig (PAMcL) Shannon Point Marine Center, Western Washington, 1900 Shannon Point Road, Anacortes, Washington 98221-4042, U.S.A.; (JH) Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007, U.S.A. (deceased). Abstract.—Anapagrides De Saint Laurent-Dechancé, 1966, as originally di- agnosed by its author, has recently been found not to agree with several es- sential characters of its type species, Eupagurus (Spiropagurus) facetus Melin, 1939. Since Anapagrides necessarily is restricted to the taxon exemplified by its type species, the new genus, Laurentia, is proposed for the four undescribed species previously referred to the former genus. These species are now fully described and the systematic relationship of Anapagrides sensu stricto to the new genus is discussed. Ina study of the hermit crabs of Maluku, Indonesia, collected during the Alpha Helix Expedition of 1975, to northern Australian and eastern Indonesian waters, Haig & Ball (1988) recognized and illustrated a species, referable to the genus Anapagrides as de- fined by De Saint Laurent-Dechancé (1966). Haig & Ball (1988) declined to de- scribe or name their taxon as a new species in deference to Michéle de Saint Laurent, who had indicated, at the time of her orig- inal generic diagnosis (De Saint Laurent- Dechancé 1966) and subsequently (De Saint Laurent 1968), that three new IndoPacific species of Anapagrides re- mained to be described. Although De Saint Laurent-Dechancé (1966) based her diag- nosis of Anapagrides on these undescribed taxa, she designated the nominal species Eupagurus (Spiropagurus) facetus Melin, 1939, as the type of the genus. McLaughlin & Sandberg (1995) recently completed a review of three of Melin’s (1939) species, including Anapagrides fa- cetus. These authors found that Melin’s spe- cies differed from De Saint Laurent-De- chancé’s (1966) generic diagnosis to such a significant extent, that a major emendation of Anapagrides was required. Anapagrides sp. of Haig & Ball (1988), while agreeing with Anapagrides sensu De Saint Laurent, can no longer be assigned to Anapagrides as emended. In addition to Haig & Ball’s (1988) Maluku specimen, we have now been able to examine De Saint Laurent’s undescribed species. These are described herein as new species in Laurentia, new ge- nus. Specimens for this review are from the Dutch Siboga, U.S. Philippine Albatross, U.S., and Australian/Indonesian Alpha He- lix expeditions. The holotype of Haig & Ball’s species has been deposited in the In- donesian National Institute of Oceanogra- phy, Jakarta (NIOJ). Specimens from the Albatross Expedition have been returned to the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM), and the Siboga material to the Zoologisch Museum, Amsterdam (ZMA). One measurement, shield length (SL), mea- sured from the tip of the rostrum to the mid- point of the posterior margin of the shield, provides an indication of animal size. 76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Laurentia, new genus Anapagrides De Saint Laurent-Dechancé, 1966:262 (in part).—Miyake, 1978:142 (in part), see remarks. Anapagrides: De Saint Laurent, 1968: 1115.—Haig & Ball, 1988:177; not An- apagrides De Saint Laurent-Dechancé, 1966. Type species.—Laurentia albatrossae, new species. Gender: feminine. Diagnosis.—Eleven pairs of trichobran- chiate gills. Rostrum narrowly triangular, produced beyond bases of ocular acicles. Ocular acicles slender. Antennular pedun- cles with elongate ultimate segment often provided with | or more long setae at distal margin. Antennal peduncle with supernu- merary segmentation. Maxillule with exter- nal lobe of endopod somewhat produced, not recurved. Crista dentata of third max- illiped without accessory tooth. Sternite of third maxillipeds unarmed, but with shal- low median depression. Chelipeds unequal or subequal, right appreciably stouter. Am- bulatory legs with elongate dactyls; carpi with dorsodistal spine. Fourth pereopods with single row of scales in propodal rasp. Sternite of fifth pereopods developed as sin- gle small subovate or subquadrate lobe. Coxa of left fifth pereopod in males with moderately long or long, sometimes coiled, sexual tube (Fig. 1E, F) provided with ter- minal tuft of stiff setae; right fifth coxa with gonopore; 3 uniramous or unequally bira- mous unpaired left pleopods. Females with paired gonopores; no paired pleopods, un- paired left pleopods on somites 2—5. Uro- pods markedly asymmetrical. Telson with transverse suture only weakly indicated; posterior lobes usually asymmetrical, ter- minal margins oblique; posterolateral mar- gins delineated at least on left. Etymology.—This genus is named in honor of the noted French carcinologist, Michele de Saint Laurent. Remarks.—Miyake (1978) included An- apagrides sensu lato among the hermit crab genera reported from Japan. Although he cited the gill structure as being phyllobran- chiate, the remainder of his generic diag- nosis was based on that of De Saint Lau- rent-Dechancé (1966); A. facetus was listed as the only species. Miyake (1978) gave no indication that he had any personal knowl- edge of the taxon. Laurentia albatrossae, new species Figs. 1, 2, 3A, B Holotype.—é (SL = 3.2 mm), Philip- pine Islands, Sulu Archipelago, 5.5 mi NW of Jolo Light, 06°09'N, 120°58’E, Albatross station 5141, 153 m; 15 Feb 1908; USNM ASI LOR Paratype.—Ovigerous @° (SL = 1.97 mm), ? Banda, 9-36 m; ZMA Crust.: De.201763. Description.—Shield (Fig. 1A) slightly longer than broad; anterior margin between rostrum and lateral projections concave; an- terolateral margins sloping; posterior mar- gin truncate; dorsal surface glabrous. Ros- trum triangular, well developed, reaching nearly half length of ocular acicles, termi- nating subacutely. Lateral projections well developed, triangular, with submarginal spine. Ocular peduncles (including corneae) approximately ¥; shield length; corneae slightly dilated. Ocular acicles subtriangu- lar, with submarginal spine. Antennular peduncles, when fully extend- ed, overreaching ocular peduncles by %4 to nearly entire length of ultimate segment; ul- timate segment with 1 or 2 long setae on dor- sodistal margin; penultimate segment with | or 2 short setae; basal segment with statocyst region expanded laterally and dorsoventrally flattened, with distal hook-like process; dor- solateral margin with small distal spine. Antennal peduncles overreaching ocular peduncles by approximately % length of ul- timate segment. Fifth and fourth segments with few long setae. Third segment un- armed or with small ventrodistal spinule. Second segment with dorsolateral distal an- gle produced, terminating in acute spine and with small secondary spine on mesial VOLUME 109, NUMBER 1 c= = ines — = D aS oo Cir (Cs +4 D-F poner an B -+———H1,6 77 Fig. 1. Laurentia albatrossae, new species. Holotype, USNM 275922: A, shield and cephalic appendages; B, dactyl of second right pereopod (mesial view); C, dactyl, propodus and carpus of left fourth pereopod (lateral view); D, anterior lobe of sternite of third pereopods; E, coxae and sternite of fifth pereopods (ventral view); E coxa of left fifth pereopod and sexual tube (dorsal view); G, telson. Scales equal 2.0 mm (A, B), 1.0 mm (C, E-G), and 0.5 mm (D). margin distally; dorsomesial distal angle with prominent acute spine. First segment sometimes with spinule at dorsolateral dis- tal angle; small spine on ventrolateral mar- gin distally. Antennal acicle reaching to base of cornea or slightly beyond; termi- nating in acute spine and with long setae on mesial margin. Antennal flagellum with 2 or 3 long and sometimes | or 2 shorter setae every | to 3 articles. 78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Laurentia albatrossae, new species. Holotype, USNM 275922: A, right cheliped (setae omitted); B, left cheliped (setae omitted); C, right second pereopod (lateral view); D, left second pereopod (lateral view); E, dactyl of left second pereopod (mesial view); EF left third pereopod (lateral view); G, dactyl of left third pereopod (mesial view). Scale equals 2.0 mm. VOLUME 109, NUMBER 1 Fig. 3. Chelipeds of species of Laurentia, new genus. L. albatrossae, new species: holotype, USNM 275922: A, left cheliped; B, right cheliped. ?L. albatrossae: 6, ZMA: C, right cheliped; D, left cheliped. L. senticosa, new species: holotype, ZMA: E, right cheliped. Scales equal 1.0 mm (A, B); 0.5 mm (C); and 0.25 mm (D, E). 80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Crista dentata with 10 to 15 regularly- spaced, small teeth. Right cheliped [Figs. 2A (setae omitted), 3B] moderately elongate, rather slender. Dactyl approximately ¥% length of palm; cutting edge with 1 small calcareous tooth proximally and 1 prominent calcareous tooth near mid-margin, row of very small calcareous teeth distally; terminating in very small corneous claw; dorsomesial margin not clearly delimited; dorsal surface convex, armed with numerous, scattered small spines or very fine spinules extending onto mesial face dorsally, and partially ob- scured by long setae; ventral surface with scattered long setae. Palm only slightly shorter than carpus; dorsomesial margin with row of slender, sometimes irregular, acute spines and long setae, dorsal surface convex, armed with scattered small spines and fine spinules practically obscured by long setae, dorsolateral margin with long setae proximally, but with small spines or fine spinules distally and on fixed finger; dorsal surface of fixed finger with scattered fine spinules also obscured by long setae, cutting edge with 1 calcareous tooth in proximal half, very small calcareous teeth distally, terminating in small weakly calci- fied claw. Carpus slightly longer than me- rus; dorsomesial margin with short, trans- verse rows of long setae proximally and more distal row of moderately slender, acute spines, dorsal surface with 3 very ir- regular rows of small spinules, decreasing in size laterally, dorsolateral margin not de- limited; mesial face with transverse rows of long setae, lateral and ventral surfaces also with transverse rows of long setae, but few- er in number. Merus with transverse rows of setae on dorsal surface, extending onto lateral face dorsally; ventrolateral margin with 1 prominent and 1 smaller spine dis- tally, ventromesial margin with 1 prominent spine at distal angle, 1 spine at midlength, and | transverse setal row and smaller spine proximally. Ischium with row of transverse setal tufts adjacent to mesial margin ven- trally. Coxa with prominent spine on ven- trolateral distal angle and second, smaller spine on ventromesial margin in distal half. Left cheliped [Figs. 2B (setae omitted), 3A] (missing in paratype) slender, not much shorter than right. Dactyl approximately % longer than palm; cutting edge with row of very small corneous teeth, terminating in corneous claw; dorsal surface with irregu- lar, nearly double row of very small spi- nules in midline, partially obscured by long setae, dorsomesial margin rounded and armed with scattered tiny spinules. Palm about % length of carpus, dorsomesial mar- gin with 3 acute spines, 2 proximally sep- arated by broad space from medianly placed third spine, dorsal surface with nu- merous tiny spinules generally obscured by long setae and 3 slightly stronger spines proximally on weakly delimited dorsolat- eral margin; fixed finger with dorsal surface similarly armed with tiny spinules and long dense setae, cutting edge with row of very small calcareous teeth interspersed with corneous teeth, terminating in tiny corneous claw; ventral surfaces all with weakly de- fined transverse rows of long setae. Carpus approximately as long as merus; dorsome- sial margin with row of slender, acute spines, dorsolateral margin with slightly smaller row of spines, both partially ob- scured by long setae; lateral and mesial fac- es with transverse rows of long setae, ven- trolateral margin with small spine distally. Merus with long setae on dorsal, lateral and mesial surfaces; ventrolateral margin with 3 prominent acute spines and transverse rows of long setae; ventromesial margin with 1 spine near distal angle and 2 rather widely- spaced spines in proximal half. Ischium with long setae on ventral margin. Second and third pereopods (Figs. 1B, 2C-G) (right third missing in holotype, both second pereopods missing in paratype) generally similar from left to right. Dactyls slightly longer than propodi, slender, in dor- sal view slightly twisted, terminating in long, slender corneous claws; in lateral view curved ventrally; dorsal margins each with a row of stiff setae, mesial faces with VOLUME 109, NUMBER 1 long setae, ventral margins also with long setae and with 5 to 8 corneous spines in distal half to 7%. Propodi with 1 or 2 cor- neous spines on ventrodistal margin, long setae dorsally and ventrally. Carpi each with 1 spine on dorsal surface adjacent to dorsodistal angle (third) and 1 additional spine on dorsal surface proximally (sec- ond). Meri each with 1 spine on ventral Margin in distal third (second) or unarmed (third), dorsal margins with tufts of long se- tae. Ischia unarmed. Fourth pereopod (Fig. 1C) without preungual process; carpus with small spine at dorsodistal margin. Sternite of third pereopod (Fig. 1D) with small, sub- ovate or subquadrate anterior lobe armed distally with 1 small spine. Sternite of fifth pereopods (Fig. 1E) semisubovate, with long distal setae. Male pleopods unequally biramous; ex- opod well developed, endopod reduced. Telson (Fig. 1G) with posterior lobes strongly asymmetrical, each with prominent spine at outer angle (largest on left) and 1 or 2 additional spines on oblique terminal margin. Affinities.—Laurentia albatrossae most closely resembles L. senticosa, new species in having the dorsal surfaces of the chelae armed with numerous tiny spines; however, the right chela of L. albatrossae is more elongate and slender than that of L. senti- cosa. The dorsomesial margin of the palm of the right cheliped of L. albatrossae car- ries a row of moderate to strong spines that are lacking in L. senticosa. Other distin- guishing characters are pointed out under L. senticosa. Etymology.—Named for the U.S. Fish Commission Steamer Albatross. Distribution.—Philippines, Sulu Archi- pelago and Banda, Indonesia; 5 to 53 m. Remarks.—In her original report on An- apagrides sensu lato, De Saint Laurent- Dechancé (1966:262) commented that a species collected in the Indo-Pacific by the Albatross and Siboga was very near Melin’s (1939) species [Eupagurus (Spiropagurus) facetus| and that her genus Anapagrides 81 was proposed for the former and two other new species from the same region. Of the specimens from the three Albatross stations initially considered by De Saint Laurent, only one, from station 5141, actually rep- resents Laurentia. One specimen, from Si- boga station 99 [Sulu Archipelago, 6°7.5'N, 120°26'E], appears to be one of the speci- mens specifically referred to by De Saint Laurent-Dechancé (1966). It is a male (SL = 1.79 mm; ZMA Crust.: De.201764) that closely resembles L. albatrossae, but is now almost entirely lacking calcification, and morphological details are difficult to in- terpret. Therefore, we tentatively assign this specimen (Fig. 3C, D) to L. albatrossae, but do not consider it a paratype. The female paratype presumably from the Siboga Ex- pedition carries only the label *“*? Banda, 5— 20 fms” in what appears to be De Saint Laurent’s hand writing. Laurentia sibogae, new species Fig. 4 Holotype.—&, ovigerous (SL = 1.5 mm), Sailus Ketjil, Paternoster Island, “‘Si- boga’”’ station 37, dredged to 18 m, 30, 31 Mar 1899, ZMA Crust.: De.201762. Description.—Shield (Fig. 4A) slightly longer than broad; anterior margin between rostrum and lateral projections distinctly concave; anterolateral margins terraced; posterior margin roundly truncate; dorsal surface glabrous. Rostrum well developed, appreciably overreaching lateral projec- tions, triangular, terminating rather bluntly. Lateral projections prominently developed, obtusely triangular, with strong marginal or submarginal spine. Ocular peduncles (including corneae) ap- proximately %4 length of shield, moderately stout, corneae only slightly dilated. Ocular acicles narrowly triangular, terminating acutely and with small submarginal spine. Antennular peduncles elongate, when fully extended, overreaching distal margin of cornea by approximately % length of ul- timate segment. Ultimate segment with few 82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Laurentia sibogae, new species. Holotype, ZMA Crust.: De.201762: A, shield and cephalic append- ages; B, right cheliped; C, left cheliped; D, right second pereopod (lateral view); E, dactyl of right second pereopod (mesial view); E left third pereopod (lateral view); G, dactyl of left third pereopod (mesial view); H, anterior lobe of sternite of third pereopods; I, coxae and sternite of fifth pereopods; J, telson. Scales equal 1.0 mm (A-G) and 0.5 mm (H-J). VOLUME 109, NUMBER 1 short setae on ventral surface. Penultimate segment glabrous. Basal segment with 1 small acute spine- on dorsolateral margin distally. Antennal peduncles overreaching ocular peduncles by % to % length of ultimate seg- ment. Fifth and fourth segments with scat- tered, moderately long setae. Third segment with very small spinule on ventrodistal margin. Second segment with dorsolateral distal angle produced, terminating in simple or bifid spine; dorsomesial distal angle with moderately strong spine. First segment with 1 acute spine ventrolateral margin distally. Antennal acicle moderately short, not reaching beyond proximal margin of ulti- mate peduncular segment. Antennal flagel- lum with 1 or 2 quite long and 1-4 shorter setae practically every article. Third maxilliped with 7 or 8 small teeth on crista dentata; proximal 3 or 4 more prominent. - Right cheliped (Fig. 4B) with chela mod- erately slender. Dactyl only slightly longer than palm; dorsomesial margin with 3 low spinulose protuberances, dorsal surface with few scattered very small spinules and sparse tufts of long setae; cutting edge with 2 widely spaced large calcareous teeth in proximal *%4 and few small calcareous teeth interspersed with corneous teeth distally. Palm with 4 widely-spaced spines or spi- nulose tubercles on dorsomesial margin, dorsal surface convex, apparently complete- ly unarmed, but with scattered long setae, dorsolateral margin not delimited on palm, but marked by tufts of long, stiff setae on fixed finger, dorsal surface of fixed finger also with numerous low protuberances and tufts of long, stiff setae; cutting edge with 2 large and several smaller calcareous teeth; ventral surfaces of dactyl, palm, and fixed finger all with tufts of long setae. Carpus approximately as long as palm; dorsomesial margin with 2 widely spaced, strong spines and proximal transverse row of long stiff setae, dorsal surface with few transverse rows of long, stiff setae, dorsolateral margin not delimited, lateral and ventral surfaces 83 with scattered long setae, 1 acute spinule at ventrolateral distal angle. Merus approxi- mately as long as carpus; dorsodistal mar- gin with long setae, and few scattered long setae on dorsal surface; ventrolateral mar- gin with 2 spines in distal half, ventrome- sial margin with small spine at distal angle. Ischium unarmed, but with few long setae. Left cheliped (Fig. 4C) with moderately long slender chela; dactyl and fixed finger lacking distinct hiatus, and not noticeably curved ventrally. Dactyl slightly longer than palm; dorsal and ventral surfaces un- armed, but with numerous tufts of long se- tae. Palm somewhat elevated in midline, perhaps minutely serrate and with long se- tae, dorsomesial and dorsolateral faces slop- ing, unarmed but with scattered long setae. Carpus approximately % longer than palm, covered dorsally, mesially and to lesser ex- tent laterally and ventrally with long setae; dorsomesial distal angle with acute spine, dorsolateral margin with row of 3 acute, rather widely-spaced spines and long setae; ventrolateral distal angle with acute spine. Merus approximately equaling length of carpus; dorsal and ventral surfaces with scattered long setae; ventrolateral margin with 2 acute spines distally, ventromesial margin with 1 acute spine at distal angle. Ischium with long setae on ventral margin. Ambulatory legs (Fig. 4D-—G) generally similar from left to right; dactyls slender, “4 to % longer than propodi; terminating in long slender corneous claws; dorsal mar- gins each with row of long, corneous, spine-like bristles in distal 74 and long setae, ventral margins each with row of corneous spines (6 to 8) in distal %4 or slightly more and additional long setae; mesial and lateral faces with scattered setae. Propodi with 1 corneous spinule at least on left second and third; dorsal and ventral surfaces all with tufts of long setae. Carpi each with small almost transparent spine at dorsodistal mar- gin, few long setae on dorsal and ventral margins. Meri with long setae dorsally and ventrally. Ischia also with long setae on ventral margins. Fourth pereopod without 84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON preungual process at base of claw; carpus without dorsodistal spine. Sternite of third pereopods (Fig. 4H) with anterior lobe roundly subquadrate, anterior margin with long setae and | spine. Sternite of fifth per- eopods (Fig. 41) subovate, with median tuft of long setae. Telson (Fig. 4J) with transverse suture in- dicated in part by tuft of setae (left side); terminal margins oblique, each with 4 spines, stronger on left and/or at outer an- gles. Affinities —Of the four species of Lau- rentia currently recognized, L. sibogae ap- pears most closely related to L. balli new species. In both species the dorsomesial margin of the right chela is similarly armed with a row of spines; however, the dactyl of the right chela has only a few low pro- tuberances in L. sibogae, whereas in L. balli this margin is provided with a row of spines; the dorsal surfaces of the palm and fixed finger have a few small spines. Ad- ditionally, the dorsodistal margin of the car- pus of the right cheliped is unarmed in L. sibogae, but carries a prominent median spine in L. balli. Etymology.—Named for the Dutch re- search vessel “‘Siboga.”’ Distribution.—At present, known only from the type locality; 18 m. Laurentia balli, new species lay, S) Anapagrides sp.: Haig & Ball, 1988:177, Fig. 8. Holotype.—é (SL = 2.0 mm), off Pulau, Saparua, Indonesia, 3°37.9'S, 128°38.6— 39'E, Alpha Helix Saparua station 3, 29 Mar 1975, coll. E. Ball, NIOJ. Description.—Shield (Fig. 5A) approxi- mately as long as broad; anterior margin be- tween rostrum and lateral projections con- cave; anterolateral margins sloping; poste- rior margin roundly truncate. Rostrum tri- angular, produced beyond bases of ocular acicles, terminating subacutely. Lateral pro- jections well developed, triangular, termi- nating acutely and with marginal or sub- marginal spinule. Ocular peduncles approximately equaling length of shield, cylindrical, with corneae slightly dilated. Ocular acicles narrowly tri- angular, with small marginal or submarginal spine. Antennular peduncles (when extended) overreaching corneae by % to % length of ultimate segment. Ultimate segment with 1 or 2 long setae on dorsodistal margin. Pen- ultimate segment with few scattered setae. Basal segment with statocyst area broad and dorsoventrally flattened, with acute, curved spine on lateral margin. Antennal peduncles equaling or slightly overreaching distal margins of corneae. Fifth and fourth segments with few scat- tered setae. Third segment with acute spine on ventral margin. Second segment with dorsolateral distal angle produced, termi- nating in acute spine, lateral margin with few setae; dorsomesial distal angle with moderately strong spine, mesial and lateral surfaces with few long setae. First segment with spinule on dorsolateral distal angle and 1 acute spine ventrolaterally. Antennal aci- cle reaching beyond base of fifth peduncu- lar segment, slightly arcuate, terminating in small spine and 1 to 3 long setae. Antennal flagellum moderately short, with 1 or 2 long and O to 4 short setae every article. Crista dentata of third maxilliped with 10 to 13 small teeth, most proximal 3 largest. Right cheliped with dactyl slightly short- er than palm; dorsomesial margin with row of very small, moderately widely-spaced spinules, surfaces all with numerous long setae; cutting edge with 2 prominent cal- careous teeth in proximal half, row of small calcareous teeth distally; terminating in small corneous claw. Palm (Fig. 5B) slight- ly shorter than carpus; dorsomesial margin with row of 4 slender spines, dorsolateral margin not delimited, dorsal surface with scattered setae and few spinules laterally, more numerous on fixed finger; lateral face with short transverse rows of long setae; cutting edge of fixed finger with 2 promi- VOLUME 109, NUMBER 1 Fig. 5. 85 A Laurentia balli, new species. Holotype, NIOJ: A, shield and cephalic appendages; B, chela of right cheliped; C, chela of left cheliped; D, carpus of second right pereopod (mesial view); E, anterior lobe of sternite of third pereopods. Scale equals 1.0 mm (A—D) and 0.5 mm (E). nent and 2 smaller calcareous teeth proxi- mally, row of small calcareous teeth distal- ly. Carpus slightly longer than merus; dor- somesial margin with 2 widely-spaced acute spines distally and 2 spinulose pro- tuberances proximally, 1 additional spine on distal margin medially; ventrolateral margin with acute spine distally; all sur- faces with scattered tufts of long setae. Me- rus subtriangular; dorsal margin with tufts of moderately long setae; ventrolateral dis- tal angle with acute spine and tuft of long 86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON setae. Ischium with scattered long setae dorsally and ventrally. Left chela (Fig. 5C) with distinct hiatus between dactyl and fixed finger. Dactyl slightly longer than palm; unarmed but with long setae on mesial face; cutting edge with row of small corneous teeth, terminating in small corneous claw. Palm with dorsal sur- face slightly elevated in midline, with few scattered moderately short setae mesially and laterally, dorsolateral margin with long setae, more numerous on fixed finger; cut- ting edge of fixed finger with row of small calcareous teeth. Carpus slightly longer than palm, approximately equaling length of merus; dorsolateral margin with row of 3 widely-spaced spines, dorsomesial margin with 1 strong spine distally and 3 short, transverse ridges and long setae in proximal *2z, extending onto mesial face dorsally; lat- eral face with scattered somewhat shorter setae. Merus subtriangular; dorsal margin with row of moderately long setae; ventro- mesial margin with long setae and 1 acute spine at distal angle; ventrolateral margin row of long setae and 2 prominent spines in distal half. Ischium damaged in holotype. Ambulatory legs with dactyls at least 1% times longer than propodi; in lateral view, Straight (second left missing in holotype) or curved (third); dorsal surfaces each with row of long, moderately stiff setae; mesial and lateral faces each with few scattered se- tae; ventral margins each with 6 corneous spines in distal *%4. Propodi with scattered setae dorsally and ventrally; ventrodistal margins each with corneous spine. Carpi each with small spine on dorsal surface near distal angle, second (Fig. 5D) with addi- tional small spine in proximal half, long se- tae dorsally and to lesser extent ventrally. Meri and ischia with long setae dorsally and ventrally. Fourth pereopod with unarmed carpus. Sternite of third pereopods with an- terior lobe (Fig. 5E) roundly subquadrate, with 4 small marginal spines. Sternite of fifth pereopods subquadrate, with long mar- ginal setae. Telson (now missing from holotype) with transverse suture not distinct; posterior lobes markedly asymmetrical, with terminal margins oblique, each with 3 or 4 spines, strongest at outer angle, lateral margin of left with narrow corneous plate. Affinities.—As previously mentioned, L. balli appears most closely allied to L. si- bogae. In addition to the characters already cited that separate the two species, there are differences in the armature of the anterior lobe of the sternite of the third pereopods and the asymmetry of the telsonal lobes. In L. balli, the anterior lobe of the sternite car- ries 4 marginal spines; only 1 is present in L. sibogae. The posterior lobes of the telson are strongly asymmetrical in L. balli, but very weakly so in L. sibogae. Etymology.—lIt gives us great pleasure to name this species for Eldon E. Ball, in rec- ognition of his work on Pacific hermit crabs. Distribution.—Saparua and possibly Banda, Indonesia. Remarks.—The figures of ““Anapagrides sp.’ (Haig & Ball 1988) are generally di- agrammatic, and some do not accurately de- pict certain characters. Specifically, the dor- solateral distal angle of the second antennal peduncular segment is illustrated as a sim- ple spine; however, there actually is a sec- ond spine on the mesiodistal margin; the first peduncular segment has a small spinule at the dorsolateral distal angle. The dactyl, fixed finger, and dorsolateral surface of the palm of the right cheliped appear unarmed in Haig & Ball’s Fig. 8B. As may be seen in our figure (Fig. 5B), the dactyl has a few small spines on the dorsomesial margin and there are a few small spines laterally on the palm; on the fixed finger, a few spinules are present on the dorsal surface proximally and on the dorsolateral margin. The right second pereopod appears to have only a dorsodistal spine on the carpus in Haig & Ball’s fig. 8E. In lateral view, the typical second spine on the proximal half of the dorsal surface is not readily observable; however, in mesial view (Fig. 5D) it is quite obvious. VOLUME 109, NUMBER 1 One 3 (SL = 2.32 mm, ZMA Crust.: De.201765) lacking chelipeds and ambula- tory legs, from “Siboga”’ station ““Banda,”’ is tentatively assigned to L. balli. It shares with the holotype the strongly asymmetrical telsonal lobes, and similarly armed anterior lobe of the sternite of the third pereopods. However, as variation among species of Laurentia is not known, we are not consid- ering this Banda specimen a paratype. Laurentia senticosa, new species Figs. 3E, 6 Holotype.—é (SL = 1.79 mm), Seram Ceram, Indonesia, 02°28.5'S, 131°3.3’E, Si- boga station 166, 118 m, 22 Aug. 1899, ZMA Crust.: De.201760a. Paratypes.—1 3,1 @ (SL = 1.28, 1.40 mm), Seram Ceram, Indonesia, 02°28.5’S, 131°3.3'E, Siboga station 166, 118 m, 22 Aug 1899, ZMA Crust.: De.201760b. Description.—Shield (Fig. 6A) slightly to considerably longer than broad; anterior margin between rostrum and lateral projec- tions concave; anterolateral margins sloping or terraced; posterior margin truncate. Ros- trum triangular, well developed, reaching approximately half length of ocular acicles, terminating bluntly or subacutely. Lateral projections well developed, triangular, with marginal or submarginal spine. Ocular peduncles subcylindrical, approx- imately *% shield length; corneae dilated. Ocular acicles subtriangular, with small submarginal spine. Antennular peduncles overreaching ocu- lar peduncles by about 7; length of ultimate segment; ultimate segment with 1 or 2 long, simple or plumose setae on dorsodistal mar- gin; penultimate segment glabrous; basal segment with statocyst region expanded lat- erally and dorsoventrally flattened, dorso- lateral margin with long slender spine dis- tally. Antennal peduncles overreaching ocular peduncles by %—%4 length of ultimate seg- ment. Fifth and fourth segments with few long setae. Third segment unarmed or with 87 small ventrodistal spinule. Second segment with dorsolateral distal angle produced, ter- minating in acute spine and sometimes with prominent slender spine. First segment with small spine on ventrolateral margin distally. Antennal acicle reaching to base of cornea or slightly beyond, terminating in acute spine and with long setae on mesial margin. Antennal flagellum missing. Crista dentata with 5—10 large or irreg- ularly-sized teeth, largest 3 or 4 often prox- imal. Right cheliped (Figs. 3E, 6B) moderately long and stout; with hiatus between dactyl and fixed finger. Dactyl approximately *% length of palm; cutting edge with 2 widely- spaced calcareous teeth in proximal %, row of very small calcareous teeth distally; ter- minating in very small corneous claw; dor- somesial margin with row of very small spines; dorsal surface convex, with scat- tered long setae, midline with irregular rows of very small spinules; ventral and mesial surfaces also with scattered long se- tae. Palm as long or slightly longer than carpus; dorsomesial margin not delimited, dorsal surface convex, armed with scat- tered, very fine spinules extending onto me- sial face and few scattered long setae, dor- solateral margin with few long setae but otherwise not delimited proximally, with fine spinules distally and on fixed finger; dorsal surface of fixed finger with scattered fine spinules and long setae; cutting edge with 3 broad calcareous teeth, terminating in small corneous claw. Carpus approxi- mately equal to length of merus; dorsome- sial margin with row of moderately slender, acute spines, dorsal surface with 2 to 4 ir- regular longitudinal rows of small spinules, dorsolateral margin not delimited; mesial face with transverse rows of long setae, lat- eral and ventral surfaces also with scattered long setae. Merus with transverse rows of setae on dorsal margin, extending onto lat- eral face dorsally; ventrolateral margin with 2 prominent spines distally, ventromesial margin with 1 prominent spine at distal an- gle, 1 additional spine originating from se- 88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Laurentia senticosa, new species. Holotype, ZMA Crust.: De.201760a: A, shield and cephalic ap- pendages; B, right cheliped; C, left cheliped; D, right second pereopod (lateral view); E, left third pereopod (lateral view); F anterior lobe of sternite of third pereopods; G, telson. Scale equals 1.0 mm (A—E) and 0.5 mm (E G). VOLUME 109, NUMBER 1 tal protuberance at midlength, and | trans- verse setal row and smaller spine proxi- mally. Ischium with setae mesially and ven- trally. Coxa sometimes with prominent spine on ventrolateral distal angle. Left cheliped (Fig. 6C) slender; not ap- preciably shorter than right but much less robust. Dactyl approximately equal to or slightly longer than palm; cutting edge with row of very small corneous teeth, terminat- ing in corneous claw; dorsal surface con- vex, with 4 or 5 spinules and tufts of long setae in midline, low protuberances and long setae on dorsomesial margin. Palm about *% length of carpus; dorsal surface convex, armed with irregular longitudinal rows of tiny spinules, fewer on dorsomesial side and on fixed finger, dorsolateral margin with row of slender acute spines; palm and fixed finger both with scattered long setae; cutting edge of fixed finger with row of widely-spaced corneous teeth, terminating in small corneous claw; ventral surfaces all with weakly defined short transverse rows of long setae. Carpus approximately as long as merus; dorsolateral margin with row of slender, acute spines, dorsomesial margin also with row of spines strongest distally, both rows partially obscured by long setae; mesial and ventral faces with transverse rows of long setae; lateral face with scat- tered long setae, ventrolateral margin with acute spine distally. Merus with long setae on dorsal, lateral and ventral surfaces; ven- trolateral margin with 2 prominent acute spines distally, continued as row of smaller spinules on lateral face ventrally; ventro- mesial margin with 1 spine near distal an- gle. Ischium with long setae on ventral mar- gin. Coxa with prominent spine at ventro- lateral distal angle. Second and third pereopods (Fig. 6D, E) (holotype with only third left remaining) generally similar from left to right. Dactyls slightly to considerably longer than propo- di, slender; in dorsal view, slightly twisted; in lateral view, curved ventrally; terminat- ing in long, slender corneous claws; dorsal margins each with a row of stiff setae, me- 89 sial faces with long setae, ventral margins also with long setae and 5—8 long corneous spines in middle half to %. Propodi with 1 or 2 corneous spines on ventrodistal mar- gin, long setae dorsally and ventrally. Carpi each with 1 spine on dorsal surface adjacent to dorsodistal angle (third) and 1 additional spine on dorsal surface proximally (sec- ond). Meri each with | spine on ventral Margin in distal third (second) or unarmed (third), dorsal margins with tufts of long se- tae. Ischia unarmed. Fourth pereopod with small preungual process at base of claw; carpus unarmed. Sternite of third pereopods with small, subovate or subtriangular ante- rior lobe (Fig. 6F) usually with 1 or 2 mar- ginal spines. Telson (Fig. 6G) with posterior lobes only slightly asymmetrical, each outer an- gle acutely developed and 1 or 2 slight pro- tuberances and tufts of setae on oblique ter- minal margins. Affinities.—In having the dorsal surfaces of the palms of both chelipeds armed with spinules, L. senticosa closely resembles L. albatrossae. However, the two species are immediately distinguished by the presence in the latter species of a row of distinct spines on the dorsomesial margin of the palm of the right chela and the more nu- merous, smaller and more regular teeth of the crista dentata. The telsons of the two species are also distinctly different. Etymology.—From the Latin senticosus meaning full of thorns, and referring to the spinulose dorsal surfaces of the chelae of this species. Distribution.—Currently known only from the type locality, Seram Ceram, I[n- donesia. Discussion As previously indicated, Melin’s (1939) Eupagurus (Spiropagurus) facetus {also cit- ed by Melin as Eupagurus (Catapagurus) facetus], not only was originally designated as the type species of Anapagrides, but, as the only nominal species assigned to the ge- 90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nus, must be considered the type species whether or not it agrees with the generic diagnosis given by De Saint Laurent-De- chancé (1966) (L. B. Holthuis, pers. comm.). As emended by McLaughlin & Sandberg (1995), Anapagrides sensu stricto is characterized as follows: 1) phyllobran- chiate gills; 2) males with short sexual tube on right fifth coxa; 3) prominent accessory tooth on the crista dentata; 4) females with single gonopore on coxa of left third pereo- pod; 5) coxa of the male third left pereopod frequently with female-type gonopore. In contrast, Laurentia has: 1) trichobranchiate gills; 2) males with long sexual tube arising from the coxa of left fifth pereopod and provided with a terminal tuft of stiff setae; 3) no accessory tooth on the crista dentata; 4) females with paired gonopores; 5) males without female-type genital pore on coxa of left third pereopod. All of the species presently recognized in Laurentia are morphologically quite simi- lar. Among the materials reviewed initially by M. de Saint Laurent, is an additional lot from the Siboga Expedition, which is as- signable, at least in part to Laurentia; how- ever, none of the specimens are in suffi- ciently good condition now to permit us to assign them with confidence to any of the described taxa. Siboga station 260 contains the bodies of at least five specimens of Lau- rentia, but also representatives of at least two other genera. Of the remaining append- ages, some most probably belong to the Laurentia specimens, but none can be matched with certitude to any of the bodies. Acknowledgements We are deeply indebted to Michele de Saint Laurent for having made the Alba- tross and Siboga specimens available for study. The photographs are the work of E. J. McGeorge. This is a scientific contribu- tion from the Shannon Point Marine Center. Literature Cited Haig, J.. & E. E. Ball. 1988. Hermit crabs from north- ern Australian and eastern Indonesian waters (Crustacea Decapoda: Anomura: Paguroidea) collected during the 1975 Alpha Helix Expedi- tion.—Records of the Australian Museum (1988) 40:151—-196. McLaughlin, P. A., & L. Sandberg. 1995. Redescrip- tions of Gustaf Melin’s 1939 ““Eupagurus (Pa- gurillus)”’ exiguus, ““Eupagurus (Catapagu- rus)” vallatus, and “Eupagurus (Spiropagu- rus)” facetus (Decapoda: Anomura: Paguridae) based on the type material—Journal of Crus- tacean Biology, 15(3):569-587. Melin, G. 1939. Paguriden und Galatheiden von Prof. Dr. Sixten Bocks Expedition nach den Bonin- Inseln 1914.—Kungliga Svenska Vetenskapsak- ademiens Handlingar (3)18(2):1-119. Miyake, S. 1978. The crustacean Anomura of Sagami Bay: 1—200 (English), 1-161 (Japanese). To- kyo: Hoikusha Publishing Co. Saint Laurent-Dechancé, M. de. 1966. Remarques sur la classification de la famille des Paguridae et sur la position systématique d’/ridopagurus de Saint Laurent. Diagnose d’Anapagrides gen. nov.—Bulletin du Muséum National d’ Histoire Naturelle (2) 38(3):257—265. Saint Laurent, M. de. 1968. Révision des genres Ca- tapaguroides et Cestopagurus et description de quatre genres nouveaux. I. Catapaguroides A. Milne Edwards et Bouvier et Decaphyllus nov. gen. (Crustacés Décapodes Paguridae) (suite). Bulletin du Muséum National d’ Histoire Natu- relle (2) 39(6) (1967):1100-1119. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):91—96. 1996 Rediscovery of Cymodocella algoense from South Africa (Crustacea: Isopoda: Sphaeromatidae) Brian Kensley and Mary Bursey (BK) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (MB) East London Museum, PO. Box 11021, Southernwood 5213, South Africa Abstract.—The tiny sphaeromatid isopod Cymodocella algoense, which is characterized by a bulbous tubular posterior pleotelson, is rediscovered 120 years after having been described from a single specimen from Algoa Bay. The new material was collected from the delta of the Nahoon River on the east coast of South Africa. Stebbing (1875) described a single tiny sphaeromatid isopod taken from the sand and fragments in a collection of sponges and gorgonians sent to him from Algoa Bay, South Africa. The species has not been recorded since. In 1910, Stebbing cast some doubt on the accuracy of the original local- ity data. Barnard (1940), in his list of Cy- modocella species from South Africa, men- tions C. algoensis with the annotation “‘spe- cies dubia, juv.”’ thereby casting uncertainty on its identity. In the course of an ecological survey of the benthos of the flood-tidal delta of the Nahoon River estuary, East London, on the east coast of South Africa, seven intertidal and seven subtidal sites were sampled by portable hydraulic suction sampler. Seven tiny adult female sphaeromatids were col- lected at two of the sites. On examination they proved to be the long forgotten Cy- modocella algoense, which is here rede- scribed. The Nahoon River estuary is about 200 km north-east of the type locality, Al- goa Bay. Three specimens have been de- posited in the National Museum of Natural History, Smithsonian Institution (USNM 253240), two in the South African Muse- um, Cape Town, and two in the East Lon- don Museum, South Africa. Family Sphaeromatidae Subfamily Dynameninae Cymodocella algoense (Stebbing, 1875) Figs. 1-3 Sphaeroma algoense Stebbing, 1875:187, alle Ilex, ike, She Cymodocella algoensis.—Stebbing, 1905: 30; 1910:430.—Barnard, 1940:492 (spe- cies dubia).—Bruce, 1995:14. Material.—5 non-ovigerous @, total length 2.9-4.0 mm, dissected 2 3.3 mm; Nahoon River estuary flood-tidal delta, East London, South Africa, 1.6 m at bottom of channel at mouth, low spring tide, coarse sand, coll. M. Bursey, 22 Feb 1993.—2 non-ovigerous 2, total length 3.0-4.0 mm; Nahoon River estuary flood-tidal delta, East London, South Africa, 2 m on steep sloping bank 500 m from mouth, low spring tide, sand, coll. M. Bursey, 8 Feb 1993. Description.—Female: Body dorsally strongly arched; surface of cephalon, per- eon, and pleon smooth. Epistome roughly triangular, distally rounded, lateral margins slightly convex, leading to short, rounded, diverging ‘legs’. Pleotelson reaching well beyond uropodal rami, consisting of broad, inflated basal half having 2 obscure, sub- median protuberances, posterior half tubu- 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Cymodocella algoense: A, Female in dorsal view, scale = 1 mm; B, Posterior pleon in ventral view; C, Female in lateral view; D, Epistome; E, Antennule; EK Antenna; G, Mandible; H, Grinding surface of man- dibular molar; I, Mandibular palp; J, Maxilliped; K, Maxilla 1; L, Maxilla 2. VOLUME 109, NUMBER 1 93 Fig. 2. Cymodocella algoense: A-G, Pereopods 1-7 respectively. 94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. lar, apically slightly narrowed with circular foramen. Antennular peduncle with robust basal article twice length of article 2, flanking and reaching anteriorly beyond epistome; article 3 slightly shorter and about half width of article 2; flagellum of 9 articles, single aesthetasc present on articles 3-8. Antennal peduncle of 5 articles, articles 1 and 2 short, subequal; articles 3—5 increas- ing in length distally; flagellum of 11 arti- cles. Mandible with sclerotized incisor hav- ing 3 cusps; lacinia mobilis with 2 cusps; 2 Cymodocella algoense: AE, Pleopods 1—5 respectively. short plus 2 more elongate fringed spines in spine-row; molar truncate, grinding sur- face subcircular, ridged, with marginal se- tae; palp with 4 fringed setae distally on article 2, 8 stout fringed setae on article 3. Maxilla 1, outer ramus with 5 broad simple and 4 denticulate distal spines; inner ramus with 4 circumplumose setae. Maxilla 2, in- ner ramus with about 12 mesiodistal cir- cumplumose setae; outer ramus with 4 elongate denticulate spines distally on each lobe. Maxilliped, endite distally having 7 stout, basally circumplumose and distally VOLUME 109, NUMBER 1 rounded setae, with single coupling hook on mesial margin; articles 2—5 of palp dis- tomesially setose, articles 2 and 3 mesiodis- tally somewhat lobed. Pereopod 1 more ro- bust but shorter than following pereopods; basis having pile of short stiff setules on dorsal surface; ischium with several simple setae on dorsal surface; merus with single stout anterodistal and posterodistal fringed seta; carpus triangular, with very short free anterior margin, with stout posterodistal fringed seta; propodus with 3 stout fringed setae on posterior surface; dactylus having bilobed accessory spine at base of unguis. Pereopod 2 more slender than following legs; basis with pile of short stiff spinules on anterior surface; ischium with 3 elongate simple setae on anterior surface; merus with 2 simple anterodistal setae; carpus rectan- gular, *% length of propodus. Pereopods 3— 7 essentially similar, pereopod 7 slightly more slender than preceding legs; basis and ischium with pile of short spinules on an- terior surface; ischium with 4—6 elongate simple setae on anterior surface; merus much shorter than ischium, with 2—4 simple setae on rounded anterodistal lobe; carpus with single bifringed seta distally, except in pereopod 7 which has 6 bifringed setae; an- terior surfaces of merus, carpus, and prop- odus having numerous short scales/spi- nules; dactylus with bifid accessory spine at base of unguis. Pleopods 1—3 each with elliptical exopod bearing plumose marginal setae; endopod triangular, subequal in length to exopod in pleopod 1, reaching well beyond exopod in pleopods 2 and 3, bearing plumose marginal setae. Pleopod 4, both rami pleated, exopod with single short distal seta. Pleopod 5, both rami pleated, exopod with transverse suture in distal half, armed with 3 spinulose lobes. Uropod with exopod freely articulating under endopod, elliptical, considerably shorter than endo- pod, latter distally rounded. Male: unknown. Remarks.—Bruce (1995), in a revision of the tube-tailed sphaeromatids, demonstrated that the genus Cymodocella was polyphy- 95 letic, defined a second genus, Diclidocella, to accommodate some of the species, moved two southern African species to Js- chyromene, and removed C. hawaiiensis Bruce, 1994, from the group altogether. The genus Cymodocella Pfeffer, 1887, currently contains four species (see Bruce 1995:5). The type species, C. tubicauda Pfeffer, 1887, was redescribed by Brandt & Wagele (1989). Cymodocella algoensis can now be included in this group, agreeing well with the revised diagnosis provided by Bruce (1995). Although Stebbing’s descrip- tion of C. algoense is limited, the figure of the whole animal in dorsal view shows the distinctively shaped convex-sided pleotel- sonic tube. The remaining four South Af- rican species are of uncertain generic status, and require redescription, given the lack of detail in existing descriptions. Acknowledgments The first author thanks Leon Engelbrecht, Robson Ntsabo, and Deon Smit for helping to operate the hydraulic pump during sam- pling. Dr. Niel Bruce of the Zoological Mu- seum, Copenhagen, commented on the manuscript, and generously shared his un- published work on the tube-tailed sphaero- matids, which greatly improved this short contribution. We are also grateful to Ms. Marilyn Schotte for reading a draft of the paper. Literature Cited Barnard, K. H. 1940. Contributions to the crustacean fauna of South Africa. XII. Further additions to the Tanaidacea, Isopoda, and Amphipoda, to- gether with keys for the identification of the hitherto recorded marine and fresh-water spe- cies.—Annals of the South African Museum 32(5):381-543. Brandt, A., & J.-W. Wagele. 1989. Redescriptions of Cymodocella tubicauda Pfeffer, 1887, and Ex- osphaeroma gigas (Leach, 1818) (Crustacea, Is- opoda, Sphaeromatidae).—Antarctic Science 1: 205-214. Bruce, N. L. 1994. New records of marine isopod crustaceans (Sphaeromatidae) from Hawaiian waters.—Crustaceana 67(3):381—386. 96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1995. The taxonomy and phylogeny of tube- tailed sphaeromatid isopods (Crustacea) with descriptions of new species and a new genus from southern Australia—Ophelia 43:127—180. Harrison, K., & J. P. Ellis. 1991. The genera of the Sphaeromatidae (Crustacea: Isopoda): a key and distribution list.—Invertebrate Taxonomy 5: 915-952. Pfeffer, G. 1887. Die Krebse von Stid-Georgien nach der Ausbeute der Deutschen Station 1882— 1883.—Jahrbuch der Hamburgischen Wissen- schaftlichen Anstalten 4:43—150. Stebbing, T. R. R. 1875. On some new exotic sessile- eyed crustaceans.—Annals and Magazine of Natural History (4)15:184—-188. . 1905. On the Isopoda. Pages 1—64, in W. A. Herdman, Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Ma- naar, Supplementary Report 23. 1910. General catalogue of South African Crustacea.—Annals of the South African Mu- seum 6(4):281—593. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):97—102. 1996 Spelaeomysis villalobosi, a new species of mysidacean from northeastern México (Crustacea: Mysidacea) Maria Elena Garcia-Garza, Gabino A. Rodriguez-Almaraz, and Thomas E. Bowman (MEG-G, GAR-A) Facultad de Ciencias Biol6égicas, Universidad Autonoma de Nuevo Leon, Ap. postal 105-E Cd. Universitaria, San Nicolas de los Garza, Nuevo Leon, México; (TEB) Department of Invertebrate Zoology, NHB-163, Smithsonian Institution, Washington, D.C. 20560, U.S.A. (deceased) Abstract.—Spelaeomysis villalobosi, a new species of mysid is described from subterranean habitats in Nuevo Leén, México. This is the third known species of Spelaeomysis from México. Spelaeomysis villalobosi is similar to Spelaeomysis quinterensis (Villalobos-Figueroa, 1951) previously described from caves in Tamaulipas and San Luis Potosi, but differs in the rounded shape of the anterolateral lobe of the eyestalk, in the 4 cusped incisor of the mandible, the presence of three spines on the dactylus of the maxilliped, and three claws on the dactylus of pereopod 1; the exopod of pleopod 4 is 2-segmented; the margin of the telson is convex. A key to the known species of Spelaeomysis is given. Resumen.—Spelaeomysis villalobosi, es una nueva especie de misidaceo hi- pogeo encontrada en el Estado de Nuevo Leon, México. Esta especie es la tercera conocida de Spelaeomysis en México. Spelaeomysis villalobosi presenta parecido morfolégico con Spelaeomysis quinterensis Villalobos-Figueroa, 1951 ubicada en cuevas de Tamaulipas y San Luis Potosi, pero existen diferencias notorias entre ambas especies, principalmente con respecto a la forma de los pedunculos oculares, la dentici6n del proceso incisivo de la mandibula, la pre- sencia de espinas en el dactilo del maxilipedio, el nimero de ufas en el dactilo del pereiopodo 1, ple6podo 4 con su exopodo con 2 segmentos en vez de 4, y la forma y ornamentacion del tels6n y el urdpodo. Se incluye una clave para las especies de Spelaeomysis conocidas. The freshwater mysids of México are poorly known. Antromysis cenotensis, A. reddelli, Spelaeomysis olivae and S. quin- terensis have been found in hypogean hab- itats (Bowman 1982). Taphromysis louisi- anae is the first freshwater epigean mysi- dacean recorded from México (Garcia-Gar- za et al. 1992). Spelaeomysis quinterensis was recorded from caves in Tamaulipas and San Luis Potosi (Villalobos-Figueroa 1951; Reddell 1981), and S. olivae is known from caves in Oaxaca (Bowman, 1973), both species have disjunct distributions. Family Lepidomysidae Clarke, 1961 Genus Spelaeomysis Caroli, 1924 Spelaeomysis villalobosi, new species Figs. 1-18 Material examined.—Holotype male, UANL (Universidad Autonoma de Nuevo Leon) CCRMYOO1, 1 female paratype (UANL CCRMY002) from a draw-well, Rancho Monte Carmelo, Cerralvo, Nuevo Le6n, 100 Km Northeast of Monterrey, Nuevo Leon, about 26°04'N, 99°37'W, 21 Oct 1989, leg. Maria Elena Garcia-Garza, 1 male paratype (UANL CCRMY003) from 98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-7. Spelaeomysis villallalobosi n. sp., Male holotype: 1, Male, lateral; 2, Eyestalks; 3, Antenna 1; 4, Antenna 2; 5, Telson; 6, Mandible; 7, Mandible palp. a small spring near the Pilon River, Mon- temorelos, Nuevo Leon, about 25°11'N, 99°48'W, 17 May 1990, leg. M. Valdez- Marroquin. 3 male paratypes (UANL CCRMY004) from Chorrera Cave, High- way Linares-Galeana, Linares, Nuevo Leén, 24°46’N, 99°37'W, 4 Apr 1994, leg. Carlos Cavazos-Camacho. Description.—Length of holotype 7 mm. Caparace with triangular rostrum (Fig. 2) VOLUME 109, NUMBER 1 99 Figs. 8-11. Spelaeomysis villalobosi n. sp. Male holotype: 8, Maxilla 1; 9, Maxilla 2; 10, Maxilliped; 11, Pereopod 1. not covering eyestalks; anterolateral lobe of wider than long, produced anterolaterally caparace reaching to midlength of eye- into pointed lobe. stalks. Eyestalk without ommatidia slightly Pleuron of pereonite 7 produced poste- 100 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 12-18. Spelaeomysis villalobosi n. sp. Male holotype: 12, Pereopod 7; 13, Pleopod 1; 14, Pleopod 2; 15, Pleopod 3; 16, Pleopod 4; 17, Pleopod 5; 18, Uropod. riorly into lobe. Telson linguiform (Fig. 5), as long as wide at base; posterior 0.4 armed, with 24 marginal spines; apex with 2 long spines flanked by 2 short spines. Antenna 1 peduncle (Fig. 3), segment 1 widest, about 1.1 times longer than segment 2; segment 3, shortest 0.95 X as long as seg- ment 2; segment 1, with 4 plumose setae at distomedial corner and 5 naked setae at dis- tolateral corner; segment 2, with row of plumose and naked setae on medial margin and 2 small setae on distolateral corner; segment 3 with naked and plumose setae on medial margin and small seta on distal part of lateral the margin; distal lobe as wide as long; bearing 3 small apical setae. Flagel- lum with 45 segments. Antenna 2 (Fig. 4), scale 2.5X longer than wide, reaching to first third of last segment of peduncle, fla- gellum broken. VOLUME 109, NUMBER 1 101 Table 1.—Differences between Spelaeomysis quinterensis and S. villalobosi. S. quinterensis S. villalobosi Rounded 1.3X Narrows at anterior 3 strong teeth Eyestalk anterolateral lobe Telson length/width Telson shape Mandibular incisor Mandibular palp Mandibular palp segment 2/seg- ment 3 Maxilla 2 endopod 1.8X Maxilliped, merus of endopod 4 setae Pereopod 1 merus 2 setae Pereopod 1 dactyl 1 terminal claw Pleopod 4 exopod 4 segments Uropod: exopod/endopod 1.25X Left Mandible (Fig. 6), incisor with 4 conspicuous cusps; lacinia mobilis bearing 5 denticles; spine row with 6 naked setae; molar process armed with minute setae; palp (Fig. 7), segment 22.6 longer than segment 3, both segments with stout barbed setae on distal part of outer margin. Maxilla | (Fig. 8), inner lobe with 4 pec- tinate terminal setae and 3 naked subter- minal setae; 3 small setae on outer margin and one long setae on proximal inner mar- gin. Outer lobe with 14 long apical setae of different types; inner margin. with row of fine hairs extending onto surface. Maxilla 2 (Fig. 9), segment | shorter than segment 2, produced medially to midlength of segment 2; segment 2 with 7 setae on distomedial margin; exopod 3X longer than wide, with about 15 long plumose marginal setae. Maxilliped (Fig. 10), merus of the en- dopod with 7 plumose setae, carpus with 7 plumose setae. Propodus, distal outer mar- gin with 2 long plumose setae and 3 smaller naked setae; dactylus with 3 spines and 4 naked setae in distal part. Pereopod 1 (Fig. 11), endopod is robust; merus with 6 naked setae; outer margin propodus with a long setae and 3 smaller setae; dactylus ending in 3 claws. Pereopds 2-7 as in Fig. 12. Pleopod 1 (Fig. 13), endopod with 2 long Segment 1 longer than segment 2, not produced distomedially Pointed 1.0 Margins evenly convex 4 strong teeth With slightly concave margins 2.6X Segment 1 shorter than segment 2, produced distomedially 7 setae 6 setae 3 terminal claws 2 segments 1.30 naked setae on medial margin and 5 long naked distal setae; exopod proximal seg- ment with 4 short and 3 longer naked me- dial setae; distal segment with 7 long naked setae. Pleopod 2 (Fig. 14), endopod elon- gate, 2.8 times longer than width; segment 3 of exopod about half length of segment 2, setae on segment 2 short. Pleopod 3 (Fig. 15), endopod not reach- ing distal part of exopod segment 1, with 4 small setae on inner margin; segment 2 of exopod with a subterminal seta. Pleopod 4 (Fig. 16), exopod with 2 segments. Pleopod 5 (Fig. 17), exopod subequal in length to endopod; segment 2 about 1.25X longer than segment 3, bent laterally at midlength, with 5 small proximolateral setae, 2 longer subterminal setae, and a terminal seta as long as 3rd segment. Uropod protopod (Fig. 18), medial lobe with numerous curved spines and a long tooth; proximal part with 2 slender setae. Endopod reaching oblique suture of exo- pod. Exopod, proximal segment with 15 spines of almost equal size. Etymology.—Named in memory of Dr. Alejandro Villalobos-Figueroa, distin- guished Mexican carcinologist. Remarks.—Spelaeomysis villalobosi 1s the third species of the genus known for México. Spelaeomysis quinterensis (Villa- lobos-Figueroa, 1951) and S. villalobosi, 102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON new species, have similar distribution in northeastern México; and S. olivae (Bow- man, 1973) is known only from Oaxaca state in southern México. Spelaeomysis vil- lalobosi is similar to S$. quinterensis; the principal differences are listed in Table 1. The seven known species of Spelaeomysis may be distinguished by the following key. Key to the Species of Spelaeomysis 1. Eyes with small anteriolateral area of OMNIA GIA ss A er ee esate erates 2 — Eyes without visual elements ......... 3} 2. Apical spine of telson about 2X length of 2 flanking spines .......... S. serratus (Fage, 1924) — Apical spine of telson more than 6X length of 2 flanking spine ...S. cardisomae Bowman 1973 3. Eyestalks fused medially, forming single DUALS ce eee cer ras Sein. Reiter S. longipes (Pillai & Mariamma, 1963) =eeEVestalksisecpalate = aaa nee 4 4. Eyestalks produced anterolaterally into Subthiancular lobes eens 5) — Eyestalks subquadrilangular, not pro- duced anterolaterally ................ 6 5. Pleopod 4 with L/W 4 segmented exo- pod; telson about 1.2; lateral margins slightly concave near midlength .S. quinterensis (Villalobos-Figueroa, 1951) — Pleopod 4 with L/W 2 segmented exo- pod; telson about 1.0; lateral margins evenly convex ........ S. villalobosi n. sp. 6. Telson L/W 1.5, with about 20 marginal spinal fox aoa S. nuniezi Bacescu & Orghidan, 1971 — Telson L/W 1.1—1.2, with about 40 mar- ginal spines ..... S. olivae Bowman, 1973 Acknowledgments We are grateful to Dr. Elva Escobar-Bri- ones of the Instituto de Ciencias del Mar y Limnologia, UNAM, México for her com- mentaries and criticism during the elabora- tion of the original draft. Besides, we sin- cerely thank two anonymous reviewers for their comments on this manuscript. Part of this work was carried out in Washington by the second author during a Short-term Vis- itor Award from the Smithsonian Institu- tion’s Office of Fellowships and Grants. Literature Cited Bacescu M., & T. Orghidan. 1971. Antromysis cuban- ica n. sp. et Spelaeomysis nuniezi n. sp. mysis cavernicoles nouvelles de Cuba.—Revue Rou- maine de Biologie, série de Zoologie 16:225— 231. Bowman, T. E. 1973. Two new American species of Spelaeomysis (Crustacea: Mysidacea) from a Mexican cave and land crab burrows.—Asso- ciation for Mexican Cave Studies, Bulletin 5: 13-20. . 1982. Mysidacea. Pp. 201—202 in S. H. Hurl- bert & A. Villalobos-Figueroa, eds., Aquatic Bi- ota of México, Central America and the West Indies. San Diego State University Foundation, San Diego, California, xv + 529 pp. Caroli, E. 1924. Su di un Misidaceo Cavernicolo (Spelaeomysis bottazii n. g., n. sp.) di Terra d’ Otranto.—Atti dell Accademia Nazionale die Lincei Rendiconti, Classe die Scienze Fisiche Matematiche Naturali 33:512—513. Clarke, D. 1961. Proposal of a new name, Lepido- mysis, for the preoccupied mysidacean generic name Lepidops Zimmer, 1927.—Crustaceana 2: 251-252. Fage, L. 1924. Sur un type nouveau de Mysidacé des eaux souterraines de l’isle de Zanzibar—Comp- tes Rendus Hebdomadaires des Séances de 1’? Académie des Sciences, Paris 178:2127—2129. Garcia-Garza, M. E., G. A. Rodriguez-Almaraz, & T. E. Bowman. 1992. Distribution of the opossum shrimp Taphromysis louisianae (Mysidacea), including new far inland records.—Journal of Crustacean Biology 12:101—103. Pillai, N. K., & T. Mariamma. 1963. On the discovery of the primitive mysidacean family Lepidomy- sidae in India.—Current Science, Bangalore 32: 219-220. Reddell, J. R. 1981. A review of cavernicole fauna of México, Guatemala and Belize.—Bulletin of the Texas Memorial Museum 27:1—327. Villalobos-Figueroa, A. 1951. Un nuevo Misidaceo de las grutas de Quintero en el Estado de Tamau- lipas.—Anales del Instituto de Biologia, Univ- ersidad Nacional Aut6noma de México 22:191— 218. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):103—111. 1996 A new copepod species from California, U.S.A.: Hesperodiaptomus californiensis (Crustacea: Copepoda: Calanoida: Diaptomidae) Megan Scanlin and Janet W. Reid (MS) College Station Box 3505, The College of William and Mary, Williamsburg, Virginia 23186-3505, U.S.A.; (JWR*) Research Associate, Department of Invertebrate Zoology/MRC- 163, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. (*Corresponding author) Abstract.—Hesperodiaptomus californiensis, new species (Copepoda: Cal- anoida: Diaptomidae) was collected from vernal pools in Lassen County, north- ern California, U.S.A. It differs from its three most morphologically similar congeners, Hesperodiaptomus schefferi, Hesperodiaptomus victoriaensis and Hesperodiaptomus kiseri in having both leg 5 endopods of the male relatively long, the left lateral protrusion on the genital double somite of the female directed dorsally, and in other details. Co-occurring diaptomids were Hesper- odiaptomus novemdecimus and Leptodiaptomus tyrrelli. A new species of diaptomid copepod was collected in Lassen County, California, as part of a study of vernal pools (described by King et al. 1996). The new copepod oc- curred in three pools and at the time of col- lection was abundant in every pool. Wilson (1959) listed 16 species of Hes- perodiaptomus in North America; since then no new species has been recognized from this continent. However, the taxon Hesperodiaptomus arcticus (Marsh, 1920) s.l. may be composed of several cryptic species. Boileau & Hebert (1988) and Bo- ileau (1991) found genetic differences be- tween different North American popula- tions of H. arcticus, which were accompa- nied by subtle morphological differences. On the other hand, Stepanova (in Borutskii et al. 1991) rejected efforts by Streletskaya (1983, 1986) to split Siberian H. arcticus into three taxa: H. koolensis Streletskaya, 1983, H. judayi Streletskaya, 1986 and H. kurenkovi Streletskaya, 1986, because of the supposedly minor degree of morpholog- ical differences. Some other members of Hesperodiapto- mus are so similar morphologically that the females cannot be reliably assigned to a particular species with present knowledge. These possible species complexes are H. breweri (Wilson, 1958a), H. eiseni (Lillje- borg in Guerne & Richard, 1889), and H. arcticus; aS well as H. kiseri (Kincaid, 1953) and H. victoriaensis (Reed, 1958). The high degree of morphological similar- ity between H. kiseri and H. victoriaensis creates some difficulty in establishing the species from California as a new taxon, be- cause it shares several characters with both these species. Hesperodiaptomus species and other diaptomid copepods are characterized clas- sically by the features of the fifth legs of the males and females, the right antennule of the male and setation of the left anten- nule of the male and both antennules of the female, and features of the somites, partic- ularly the shape of the thoracic “wings” (expansions of the sixth thoracic somite). Pending eventual revision of the genus Hes- perodiaptomus and the possible discovery of additional morphological and genetic characteristics, we describe the new species using these traditional discriminators. 104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The copepod specimens were originally fixed in buffered formalin and transferred to 70% ethanol for long term storage. Field sampling methods were described by King et al. (1996). Descriptions were made from whole specimens in glycerin and/or lactic acid; dissected specimens were mounted ei- ther in CMC-10 or in commercial polyvinyl] lactophenol with a little Chlorazol Black E added. All measurements were made from specimens in glycerin. Specimens were de- posited in the collections of the United States National Museum of Natural History, Smithsonian Institution (USNM) and the Natural History Museum of Los Angeles County, California (LACM). Family Diaptomidae G. O. Sars Genus Hesperodiaptomus Light, 1938 Hesperodiaptomus californiensis, new species Figs. 1-3 Type material.—Holotype 6, USNM 264057; allotype 2, USNM 264058; para- types: 3 dd 3 2, each dissected on slide in PVL or in CMC-10, and 464 dd 292 + copepodids, USNM 264059; 463 dd 22 + copepodids, LACM 95-50.1; all from California, Lassen County, pool #41, 40°37'N, 121°03'W, immediately adjacent to west side of Route 44, 1.4 km north of Bogart Safety Rest Area/Ranger Station, 2 m tow of wet meadow edge with plankton net, sample 215. Additional, non-type material.—372 33 22 + copepodids, California, Lassen County, pool #42, 40°32'N, 121°0’W, 30 m west of Route 44, just south of railroad track crossing, 2 m tow of wet meadow lake edge with plankton net, sample 218, USNM 264060. 894 ¢6d5 22, pool #42 bottom, sample 217, LACM 95-50.3. 41 36 @Q, pool #41, sample 214, USNM 264061. 400+ dd 2°, pool #42 surface, sample 216, LACM 95-50.4. 28 dd 2°, Califor- nia, Lassen County, pool #43 (Long Lake), 40°31'N, 120°59'W, 100 m west of Route 44, about 0.36 km south of pool #41 (about 1.5 km south of intersection with Route A21), 2 m tow of wet meadow lake bottom with plankton net, sample 220, USNM 264062. 10 dd 2 2, pool #43 (Long Lake), sample 219, LACM 95-50.2. All collected 23 April 1992 by J. L. King, D. Gluesen- kamp and J. Tritt. Undissected specimens preserved in 70% ethanol. Co-occurring species (determined by JWR).—Pool #41: Hesperodiaptomus nov- emdecimus (Wilson, 1953), Leptodiaptomus tyrrelli (Poppe, 1888), Acanthocyclops car- olinianus (Yeatman, 1944), Diacyclops crassicaudis var. brachycercus (Kiefer, 1927), Microcyclops rubellus (Lilljeborg, 1901), Bryocamptus washingtonensis Wil- son, 1958b; pool #42: H. novemdecimus, L. tyrrelli, A. carolinianus, Acanthocyclops vernalis (Fischer, 1853) s.1., Diacyclops na- vus (Herrick, 1882); pool #43 (Long Lake): H. novemdecimus, L. tyrrelli, A. carolini- anus, B. washingtonensis. Male.—Length (mm) of holotype 2.21; of ten paratypes, mean = 2.14, median = 2.18, range = 1.93—2.32. Pediger 5 (sixth thoracic somite) (Fig. la), lateral ““wings” symmetrical, each with two sensilla. Uro- some (Fig. la) symmetrical except urosom- ites 1 and 4 slightly produced laterally and posteriorly. Urosomites 2—4 and caudal rami with pore-canals, middle dorsal pore- canal of urosomite 4 displaced slightly to left of dorsal midline. Inner sides of caudal rami with long hairs, rest of ramal surfaces with short sparse hairs. Rostral points (Fig. 1b) short, acute. Antennules (Fig. la, c, d) reaching mid- dle of urosomite 2. Right antennule with two setae on segment 6; segment 8 with one seta and one spine; segment 10 with one seta and one stout spine reaching middle of segment 12; segment 11 with one seta and one stout spine, reaching distal end of seg- ment 13; segment 13 with one stout spine reaching past middle of segment 14; seg- ment 14 without spine; segment 15 with small acute process pointing distally; seg- ment 16 with two setae. Segment 23 (Fig. 1d), process at distal end straight, reaching VOLUME 109, NUMBER 1 105 <= < SX = KK KE DLE 100 pm Fig. 1. Hesperodiaptomus californiensis, new species, male: a, Habitus, dorsal; b, Rostral points (indicated by arrows); c, Right antennule, segments 6-17; d, Right antennule, segments 23-25; e, Leg 2 endopod, segments 1 and 2, posterior. 106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ag we AVIS SOIC as Fig. 2. Hesperodiaptomus californiensis, new species, male: a, Leg 5, posterior; b, Left leg 5 exopod, posterior; c, Left leg 5 exopod and endopod, anterior; d, Left leg 5 endopod, posterior; e, Right leg 5 endopod, anterior. Both scales = 50 pm. end of last segment of antennule. Setation of left antennule as in female. Leg 2 endopod segment 2 (Fig. le) with- out defined Schmeil’s organ, but with small transverse ridge, more or less developed on different specimens. Left leg 5 (Fig. 2a—d): Coxa, small pro- cess tipped with sensillum near outer pos- terodistal margin. Basis with lateral seta. Exopod segment 1 longer than segment 2, with haired pad on inner margin, hairs thick and as long as about half width of endopod. Exopod 2, tip forming stout, blunt, smooth lateral process; also with serrate digital pro- cess. Inner margin of exopod 2 with two pads, proximal pad haired, proximal hairs of this pad thinner and shorter than distal hairs; distal pad covered with rows of teeth and few tiny hairs near tip of exopod. Pos- terior surface of exopod 2 with longitudinal groove. Endopod reaching midlength of ex- opod 2, of one segment, narrowing at distal %. Tip of endopod narrowing in acute pro- cess; also with five small subterminal spines and group of subterminal hairs. Right leg (Fig. 2a, e): Coxa with small posterodistal process ending in sensillum. Basis with lateral seta, shallow longitudinal VOLUME 109, NUMBER 1 R <A » << > > TEOMA = C é AMM iam, a en Fig. 3. 107 100 pm Hesperodiaptomus californiensis, new species, female: a, Habitus, dorsal; b, Rostral points (indicated by arrows); c, Antennule, segments 1—9; d, Leg 5, anterior; e, Tip of leg 5 endopod, posterior. groove on posterior surface, and rounded, unornamented protrusion on middle of in- ner margin; length of protrusion equal to width of endopod, its width equal to % width of endopod. Outer margin of exopod 1 with large, distally directed conical pro- cess on distolateral corner. Exopod 2 almost twice length of exopod 1, with shallow lon- gitudinal groove along distal % of inner margin; lateral spine at distal % of exopod 2, % thickness of endopod and ¥% length of exopod 2, straight, finely denticulate. Ter- minal claw tapering gradually from en- larged base, twice length of exopods 1 and 2 combined, with row of teeth extending from tip of claw almost to base. Right en- dopod without suture on posterior surface, suture visible on anterior surface where en- 108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dopod narrows slightly at midlength; en- dopod tapering to blunt point; with 3—5 spines near outer distal margin, and subter- minal group of small hairs. Female.—Length (mm) of allotype 2.39; of ten paratypes, mean = 2.20, median = 2.22, range = 1.95—2.33. Prosome (Fig. 3a) symmetrical, thoracic wings symmetrical, each with two sensilla located on lateral and posterior margins. Genital double somite extended laterally in asymmetrical conical projections each tipped with sensillum, left projection directed dorsally, right projection laterally. Caudal rami haired in pattern sim- ilar to that of male. Rostral points (Fig. 3b) low, rounded. Antennule (Fig. 3a, c) reaching mid- length of genital double somite. Segments 1—8 or 9 each with few tiny hairs irregularly scattered on posterior surface. Seta on seg- ment | long, reaching midlength of segment 4. Appendages per segment as follows (Ro- man numerals = segment, Arabic numerals = number of setae, a = aesthetasc, sp = spine): I(1+a), W@Gta), Il(1+a), IV), V(i+a), VIC), Vil(i+a), VIII(1+sp), IX(2+a), X(1), XI(2), X11 +sp+a), XIIIC), XIV(1+a), XV(1), XVI(1+a), XVIIC1), XVIII(1), XIX(1+a), XX(1), XXI(), XXII(2), XXII(2), XXTV(5+a). Leg 2 (not figured) of all females ex- amined without trace of Schmeil’s organ or transverse ridge on endopod 2. Leg 5 (Fig. 3d, e): Coxa with posterior lateral protrusion ending in sensillum. Basis with lateral seta reaching % of length of ex- opod 1. Endopod two-segmented, distal segment twice length of proximal segment. Endopod 2 with two setae at tip, tip pro- truding in acute point between these setae, and 3—5 subterminal teeth surrounding tip of endopod. Exopod 2 a little longer than exopod 1, claw with inner margin toothed. Exopod 2 with small articulated spine lat- eral to outer margin of exopod 3, reaching from slightly beyond middle to end of ex- opod 3. Exopod 3 distinct from exopod 2. Inner spiniform seta of exopod 3 serrate on one or both margins, longer and stouter than outer, naked spiniform seta. Type locality.—Pool #41, 40°37'N, 121°03'W, Lassen County, California. Etymology.—The species name is given after the State of California, in which the type locality is located. Habitat description—The three pools (41, 42 and 43—Long Lake) are relatively large for vernal pools (400 xX 400 m, 300 x 300 m and 800 X 200 m respectively at time of sampling), shallow (maximum depths 0.15—0.6 m), clear, and covered 70— 90% with aquatic grasses. The electrical conductivity was low (30, 60 and 10 wMHO), water temperature 16, 16 and 23 C, pH 7.7, 7.7 and 8.2, alkalinity 20, 28 and 8 ppm, and total dissolved solids 10, 30 and 0 ppm respectively. Discussion and comparisons.—Hespero- diaptomus californiensis differs from H. schefferi (Wilson, 1953) in the male leg 5: the conical process on the coxa of the right leg is larger; the inner protrusion on the ba- sis of the right leg of H. californiensis is rounded, not quadrate as in H. schefferi; and the endopods of both left and right legs are longer in H. californiensis. In leg 5 of the female, H. californiensis has the endo- pod extended in a terminal point. The gen- ital double somite in H. schefferi has very slight lateral projections. Hesperodiaptomus victoriaensis 1s mor- phologically closest to H. californiensis. The male of H. californiensis differs from H. victoriaensis in having the right leg 5 with longer endopod, and in having the in- ner process on the basis larger and placed at about midlength rather than at the inner proximal corner. In the female of H. victo- riaensis, the lateral projections of the gen- ital double somite are slightly asymmetri- cal, the left projection being directed pos- terolaterally rather than dorsally. Both H. schefferi and H. victoriaensis possess an acute process on the distal mar- gin of segment 16 of the right antennule of the male. Such a process is lacking in H. californiensis. VOLUME 109, NUMBER 1 The differences between H. kiseri and H. californiensis are primarily in leg 5 of the male: in H. californiensis there is no prox- imal protrusion on the inner margin of the right basis, and only one protrusion on the middle of the inner margin. There is no dis- tally placed spiniform projection on the in- ner margin of the right exopod 1. The small protrusion on the posterior surface of the right exopod 2, present in H. kiseri, is lack- ing in A. californiensis. The claw is smoothly tapered in H. californiensis, with- out angles as in H. kiseri. The lateral pro- jections of the genital double somite of the female are asymmetrical in H. californien- sis, Symmetrical in H. kiseri. The thoracic wings are not expanded laterally in H. cal- iforniensis. In A. californiensis, the left lateral pro- jection on the female double somite is dor- sally directed. The second (middle) uro- somite is relatively long. On leg 5 of the male, the endopodites are relatively longer than in other similar species. In the most recent diagnosis of the genus Hesperodiaptomus, Borutskii et al. (1992) Stated that the left antennule of the male is like that of the female. However, H. hirsu- tus (Wilson, 1953) always, and H. schefferi sometimes, display sexual dimorphism in setation on some segments (Wilson 1953). The observed setation pattern of H. califor- niensis was constant, similar in both sexes, and the most common pattern found in the genus (cf. Wilson 1953). Species of the genus Hesperodiaptomus are supposed to lack Schmeil’s organ (Wil- son 1953). Schmeil’s organ (Schmeil 1896) is a small protuberance of unknown func- tion, which in diaptomids if present is lo- cated on the posterior surface of leg 2 en- dopod 2. If the small transverse ridge in this position which was noted in some of the males of H. californiensis examined, does correspond to Schmeil’s organ, this is the first reported occurrence of this structure in the genus. Most species of Hesperodiaptomus occur in northwestern North America. A few spe- 109 cies are found in eastern Canada and the U.S.A., and the ranges of three species ex- tend into Siberia (Borutskii et al. 1991). The species that are morphologically clos- est to H. californiensis occur well north of its range. Hesperodiaptomus kiseri is found in the State of Washington and in Saskatch- ewan (Wilson 1959). Hesperodiaptomus schefferi is found in the Pribilof Islands, Alaska, and the northern Rocky Mountain States (Wilson 1959). Hesperodiaptomus victoriaensis has been collected in the mountains of southwestern Alberta (Ander- son 1967, 1971), near Churchill, Manitoba (Hebert 1985; Boileau & Hebert 1988), and on Victoria Island, N.W.T. (Reed 1958). Two other species of diaptomids, Lepto- diaptomus tyrrelli and Hesperodiaptomus novemdecimus, occurred in large numbers together with H. californiensis in all three pools sampled. Many instances of co-oc- curring diaptomid species have been re- ported and have been much discussed in the published literature, including reviews by Cole (1961) and Hutchinson (1967). Of 34 examples of co-occurrence listed by Cole (1961), 10 involved congeneric species. Hutchinson (1967) observed that co-occur- ring diaptomid species tend to be of differ- ent sizes and suggested that these size dif- ferences may indicate non-overlapping feeding niches, and that differences in feed- ing would likely be found to be related to structural differences. Several co-occurrences of species of Hesperodiaptomus have been reported. Wilson (1953) reported H. wardi (Wilson, 1953) together with H. novemdecimus. An- derson (1971) collected H. shoshone (S. A. Forbes, 1893) with H. arcticus, and H. shoshone with H. victoriaensis. Hammer & Sawchyn (1968) found H. arcticus in the same pond as H. kiseri, although the inci- dences were separated by a five-day inter- val. These authors noted distinct size dif- ferences between the two species, but pos- tulated mutual exclusion between them. An especially interesting example was given by Anderson (1967), who found H. shoshone 110 with H. victoriaensis and L. tyrrelli together in a pond in Alberta. There, H. shoshone individuals averaged 1.6 times longer than H. victoriaensis and preyed actively upon the much smaller L. tyrrelli. Anderson (1967) described the enlarged clawlike se- tae of the maxilliped of H. shoshone, which were apparently an adaptation for this pred- atory activity. The corresponding setae of H. victoriaensis were found to be unmodi- fied, and this species apparently did not pre- date upon the smaller diaptomid. In the case of the Lassen County Hesperodiaptomus pair, the setae of the mouthparts and max- illiped of both species are unmodified. Hes- perodiaptomus novemdecimus averaged larger than H. californiensis in all three ponds, and the two could be reliably sorted because of this size difference. In accor- dance with Hutchinson’s (1967) hypothesis, we suggest that differences between these species in feeding niches are likely to exist, although we have made no direct observa- tions. Acknowledgments This article was prepared during the se- nior author’s participation in the Mentor- ship Program of the Thomas Jefferson High School for Science and Technology, Alex- andria, Virginia. We thank the collector, Ja- mie L. King, University of California, Da- vis for providing habitat information and a draft of the article by King et al. Jamie King and two anonymous reviewers made valu- able suggestions on an earlier draft. Literature Cited Anderson, R. S. 1967. Diaptomid copepods from two mountain ponds in Alberta.—Canadian Journal of Zoology 45:1043-1047. . 1971. Crustacean plankton of 146 alpine and subalpine lakes and ponds in western Canada.— Journal of the Fisheries Research Board of Can- ada 28:311-321. Boileau, M. G. 1991. A genetic determination of cryptic species (Copepoda: Calanoida) and their postglacial biogeography in North America. Zoological Journal of the Linnean Society 102: 375-396. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON , & PD. N. Hebert. 1988. Genetic differenti- ation of freshwater pond copepods at Arctic sites—Hydrobiologia 167/168:393—400. Borutskii, E. V., L. A. Stepanova, & M.S. Kos. 1991. Revision of the Calanoida of Freshwaters of the USSR. Zoological Institute, USSR Academy of Sciences, Saint Petersburg, 503 pp. (in Russian; abstract in English). Cole, G. A. 1961. Some calanoid copepods from Ar- izona with notes on congeneric occurrences of Diaptomus species.—Limnology and Oceanog- raphy 6(4):432—442. Fischer, S. 1853. Beitrage zur Kenntnis der in der Umgegend von St. Petersburg sich findenden Cyklopiden; (Fortsetzung).—Bulletin de la So- ciété Impériale des Naturalistes de Moscou 26: 74-100 + plates II, III. Forbes, S. A. 1893. A preliminary report on the aquatic invertebrate fauna of the Yellowstone National Park, Wyoming, and of the Flathead region of Montana.—Bulletin of the United States Fish Commission for 1891:207—258 + plates XXX VII- XL. Guerne, J. de, & J. Richard. 1889. Révision des Cal- anides d’eau douce.—Mémoires de la Société Zoologique de France 2:53—181 + plates I-IV. Hammer, U. T., & W. W. Sawchyn. 1968. Seasonal succession and congeneric associations of Diap- tomus spp. (Copepoda) in some Saskatchewan ponds.—Limnology and Oceanography 13(3): 476-484. Hebert, P D. N. 1985. The ecology of the dominant copepod species at a low arctic site —Canadian Journal of Zoology 63:1138-1147. Herrick, C. L. 1882. Cyclopidae of Minnesota with notes on other copepods.—Report of the Geo- logical and Natural History Survey of Minne- sota 10:221—223 + plates I-VII. Hutchinson, G. E. 1967. A treatise on limnology. II. John Wiley & Sons, Inc., New York, 1115 pp. Kiefer, E 1927. Freilebende Stisswasser-Copepoden aus Nordamerika.—Zoologischer Anzeiger 72: 262-268. Kincaid, T. 1953. A contribution to the taxonomy and distribution of the American fresh water calan- oid crustacea. The Calliostoma Company, Se- attle, Washington, 73 pp. + plates 1—5. King, J. L., M. A. Simovich, & R. C. Brusca. 1996. Endemism, species richness, and ecology of crustacean assemblages in northern California vernal pools.—Hydrobiologia (in press). Light, S. E 1938. New subgenera and species of Diap- tomid copepods from the inland waters of Cal- ifornia and Nevada.—University of California Publications in Zoology 43:67—78. Lilljeborg, W. 1901. Synopsis specierum huc usque in Suecia observatorum generis Cyclopis.—Kon- VOLUME 109, NUMBER 1 glige Svenska Vetenskaps-Akademiens Han- dlingar 35:3-118 + Plates I-VI. Marsh, C. D. 1920. The fresh water Copepoda of the Canadian Arctic Expedition 1913—18.—Report of the Canadian Arctic Expedition 1913-18, 7(Crustacea), Part J:3—25. Poppe, S. A. 1888. Diagnoses de deux espéces nou- velles du genre Diaptomus Westwood.—Bulle- tin de la Société Zoologique de France 13:158— 159. Reed, E. B. 1958. Two new species of Diaptomus from arctic and subarctic Canada (Calanoida, Copepoda).—Canadian Journal of Zoology 36: 663-670. Schmeil, O. 1896. Deutschlands freilebende Siisswas- ser-Copepoden. III Teil: Centropagidae. Erwin Nagele Verlag, Stuttgart, 143 pp. Streletskaya, E. Y. 1983. The “‘eiseni” group of the genus Hesperodiaptomus (Copepoda, Calanoi- da) and a new species H. koolensis from the Chukot Peninsula.—Zoologicheskii Zhurnal 62: 1474-1480 (in Russian). 1986. Toward a revision of the Beringian freshwater calanoid crustaceans (Copepoda, 111 Calanoida). Pp. 64—99 in Biogeography of the Beringian sector of the Subarctic: Proceedings of the Tenth All-Union Symposium, Vladivos- tok, 1986 (in Russian) (not seen; cited in Bo- rutskii et al. 1991). Wilson, M. S. 1953. New and inadequately known North American species of the genus Diapto- mus.—Smithsonian Miscellaneous Collections 122:1—30. . 1958a. New records and species of calanoid copepods from Saskatchewan and Louisiana.— Canadian Journal of Zoology 36:489-497. 1958b. North American harpacticoid cope- pods. 4. Diagnoses of new species of fresh-wa- ter Canthocamptidae and Cletodidae (genus Huntemannia).—Proceedings of the Biological Society of Washington 71:43-48. . 1959. Free-living Copepoda: Calanoida. Pp. 738-794 in W. T. Edmondson, ed., Ward and Whipple’s fresh-water biology. Second edition. John Wiley & Sons, Inc., New York, 1248 pp. Yeatman, H. C. 1944. American cyclopoid copepods of the viridis-vernalis group (including a de- scription of Cyclops carolinianus n. sp.).— American Midland Naturalist 32:1—90. PROCEEDINGS OF THE BICLOGICAL SOCIETY OF WASHINGTON 109(1):112-117. 1996 Orecturus amplus, a new species (Copepoda: Siphonostomatoida: Asterocheridae) from an alcyonacean in New Caledonia Arthur G. Humes Boston University Marine Program, Marine Biological Laboratory, Woods Hole, Massachusetts 02543, U.S.A. Abstract.—A new species of siphonostomatoid copepod, Orecturus amplus, is described from New Caledonia, where it is associated with the alcyonacean Siphonogorgia variabilis Hickson. The new species may be distinguished from its six congeners by its large size and by detailed features of its external anatomy. Copepods are frequent associates of Al- cyonacea (Humes 1990). Species of the or- der Poecilostomatoida are by far the most common copepod associates (nearly 180 species) of these cnidarians. Those of other orders are much less frequently found with soft corals (Cyclopoida, 4 species; Harpac- ticoida, 1 species; and Siphonostomatoida, 3 species). Three poecilostomatoids have been recorded from the alconacean Siphon- ogorgia. In Madagascar, Acanthomolgus hians (Humes & Ho 1968) lives with Si- phonogorgia pichoni Verseveldt (host orig- inally reported as S. pendula Studer) and Acanthomolgus longispinifer (Humes & Ho 1968) with S. pichoni (see Humes & Stock 1973). In the Moluccas (at Pulau Gomumu, south of Obi), Acanthomolgus brevifurca Humes, 1990, is associated with S. varia- bilis Hickson. The relatively small number of siphon- ostomatoid copepods reported to date from soft corals probably reflects the lack of at- tention so far given to these associations. In this paper, a siphonostomatoid belonging to the genus Orecturus, living with Siphono- gorgia in New Caledonia, is described. Materials and Methods The host alcyonacean colony was isolat- ed immediately after collection in a plastic bag containing sea water. Sufficient 95% ethanol was later added to make approxi- mately a 5% solution. After 2—3 hours, the soft coral was gently rinsed, and the wash water poured through a fine net (approxi- mately 120 holes per 2.5 cm, each hole ap- proximately 225 wm square). The copepods were then recovered from the sediment re- tained. The copepods were measured and studied in lactic acid. The length of the body does not include the setae on the caudal rami. Dissections were prepared using the wood- en slide method described by Humes & Gooding (1964). All drawings were made with the aid of a camera lucida. Order Siphonostomatoida Thorell, 1859 Family Asterocheridae Giesbrecht, 1899 Genus Orecturus Humes, 1992 Orecturus amplus, new species Figs. la—g, 2a—i, 3a—i Type material—T 2°, 9 36 from the alcyonacean Siphonogorgia variabilis Hickson, in 30 m, outside Récif Mtere, near Nouméa, New Caledonia, 22°20'40"S, 166°13'55"E, 23 Jul 1971. Holotype & (USNM 274207), allotype 6 (USNM 274208), and 11 paratypes (4 22,7 dd) (USNM 274209) deposited in the National Museum of Natural History, Smithsonian Institution, Washington. Remaining para- types in the collection of the author. VOLUME 109, NUMBER 1 Fig. 1. 113 Za” — Zp aa —— ee Orecturus amplus, new species. Female. a, body, dorsal (scale A); b, body, lateral (A); c, urosome, dorsal (B); d, genital area, dorsal (D); e, anal somite and caudal ramus, dorsal (C); f, egg, ventral (B); g, rostral area, ventral (B). A, = antennule, A, = antenna, S = siphon. Female.—Body (Fig. la, b) with moder- ately broad prosome. Length 1.54 mm (1.50—1.56 mm) and greatest width 0.80 mm (0.78—0.88 mm), based on 7 specimens in lactic acid. Greatest dorsoventral thickness 0.55 mm. Somite bearing leg 1 fused with cephalosome. Somite bearing leg 3 with pos- teriorly extended, narrowly rounded epi- mera. Somite bearing leg 4 narrow, with rounded epimera, overlapped dorsally by ter- gum of preceding somite. Ratio of length to width of prosome 1.38:1. Ratio of length of prosome to that of urosome 2.05:1. Somite bearing leg 5 (Fig. Ic) 120 x 264 um, indented laterally, with scalelike spines laterally. Genital double-somite broader 114 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SSS SS rs Fig. 2. Orecturus amplus, new species. Female. a, antennule, postero-outer (scale C); b, antenna, outer (C); c, siphon, ventral (B); d, mandible, anterior (C); e, maxillule, anterior (C); f, maxilla, posterior (C); g, maxilliped, posterior (C); h, leg 1 and intercoxal plate, anterior (E); i, leg 2 and intercoxal plate, anterior (E). than long, 117 wm long in midline, 160 pm long including posterolateral spiniform pro- cesses, 231 ym wide; ratio 0.51:1, taking length at midline. Genital areas located dor- solaterally near indentation. Each genital area with 2 small setae (Fig. 1d). Two post- genital somites from anterior to posterior 55 x 172 wm (length including posterolateral processes) and 180 X 160 pm, longer so- mite with small scalelike spines (continuing on dorsal and ventral surfaces, but not shown in Fig. Ic, e). Caudal ramus (Fig. le) 60 X 75 pm, wider than long, ratio 0.8:1. Dorsal seta 70 VOLUME 109, NUMBER 1 115 Fig. 3. Orecturus amplus, new species. Female. a, leg 3 and intercoxal plate, posterior (scale E); b, leg 4 and intercoxal plate, anterior (E); c, leg 5, ventral (C). Male. d, body, dorsal (A); e, somites of legs 3 and 4, dorsal (B); f, urosome, dorsal (E); g, antennule, dorsal (C); h, leg 5, ventral (D); genital somite showing leg 6, ventral (D). um, with elongate pedicel. Outer lateral seta 250 ym, outermost terminal seta 275 14m, innermost terminal seta 340 wm, and 2 median terminal setae 460 wm (outer) and 625 wm (inner) swollen proximally. All se- tae with long lateral setules. Outer margin of ramus with few scalelike spines and pos- tero-inner corner with group of slender se- tules. Dorsal surface of body without visible sensilla. Entire egg sac not seen, but isolated egg oval (Fig. 1f), 226 179 wm. Rostrum (Fig. 1g) weak. Antennule (Fig. 116 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2a) 585 wm long, 17-segmented. Lengths of its segments (measured along their posterior nonsetiferous margins): 39 (78 wm along its anterior margin), 26, 83, 26, 25, 4, 21, 34, 36, 36, 38, 38, 39, 44, 26, 23, and 13 wm, respectively. Formula for armature: 2, 2, 10, 2a Ole 252 ae aeeleaesthetascs 2, 3, and 5 + | aesthetasc. Enlarged seta on third segment 74 pm long. Certain setae on segments 1—4 with lateral setules. An- tenna (Fig. 2b) with elongate, 1-segmented exopod 57 X 10 wm, bearing short inner seta and long terminal barbed seta 95 pm; both sides of exopod with setules. Endopod with first segment unarmed, second seg- ment with 3 short setae and long terminal seta 120 wm long. Siphon (Fig. 2c) 935 wm long, reaching almost to ventral edge of intercoxal plate of leg 4. Mandible (Fig. 2d), maxillule (Fig. 2e), and maxilla (Fig. 2f) resembling those of congeners (see Humes, 1992, 1993, 1994). Maxilliped (Fig. 2g) with basis lacking in- ner seta but having small spinules along outer margin. Endopod 3-segmented, armed with 2,2,1; terminal claw 120 wm. Legs 1-4 (Figs. 2h, 1, 3a, b) segmented and armed as follows (Roman numerals in- dicating spines, Arabic numerals represent- ing setae): P, coxa O-1 basis 1-I exp I-1; I-1; II,I,5 enp 0-1; 0-2; 1,5 P, coxa 0-1 basis 1-Oexp I-1; I-1; III,1,4 enp O-1; 0-2; 1,1,1,3 P; coxa O-1 basis 1-Oexp I-1; I-1; I,II,3 enp 0-1; 0-2; 1,1,3 P, coxa O-1 basis 1-O exp I-1; I-1; I,II,3 enp O-1; 0-2; 1,1,2 Leg 1 with inner barbed spine on basis 52 wm. Leg 5 (Fig. 3c) placed ventrally (as shown in Fig. Ic). Suboval free segment 107 X 70 pm, ratio 1.53:1. Five setae from outer to inner 75, 65, 36, 25, and 27 wm. Two innermost setae stout, almost spine- like, and lacking setules. Other setae with lateral setules. Few scalelike spines along outer edge of free segment. Adjacent dorsal seta, here located ventrally, 52 wm with lat- eral setules. Leg 6 represented by 2 small setae on genital area (Fig. 1d). Color of living specimens red, eye red. Male.—Body (Fig. 3d) with prosome more slender than in female. Length 0.98 mm (0.94—1.02 mm) and greatest width 0.43 mm (0.40—0.45 mm), based on 9 spec- imens in lactic acid. Greatest dorsoventral thickness 0.33 mm. Somite bearing leg 4 with sharply pointed epimera (Fig. 3c). Ra- tio of length to width of prosome 1.69:1. Ratio of length of prosome to that of uro- some 2.46:1. Somite bearing leg 5 (Fig. 3f) 73 X 127 wm including processes (length at midline 62 wm). Genital somite 60 X 86 pm in- cluding leg 6 (length at midline 65 pm). Three postgenital somites from anterior to posterior 42 X 112 wm (length at midline 26 pm), 26 X 104 wm (length at midline 16 pm), and 104 X 112 pm. Caudal ramus 39 X 52 wm, ratio 0.75:1, resembling that of female. Dosal surface of body without visible sensilla. Urosome with scalelike spines as in female. Rostrum as in female. Antennule (Fig. 3g) 13-segmented, with segments 8 and 9 clearly separated (in other species these segments tending to be fused). Lengths of segments (measured along their posterior nonsetiferous margins): 21 (52 wm along anterior margin), 22, 42, 23, 13, 9, 14, 26, 48, 29, 57, 52, and 48 wm, respectively. Formula for armature: 2, 2, 10 + 3 aesthet- ascs, 2 +> | aesthetasc, 6; 1p 1B 25 leet + 1 aesthetasc, and 5 + 1 aesthetasc. Aesthetascs on segments 3, 4, and 10 (these present in all males examined) very hya- line. Antenna as in female. Siphon, mouthparts, and legs 1-4 like those of female. Leg 5 (Fig. 3h) with free segment 50 X 32 wm, ratio 1.56:1, otherwise placed ven- trally and similar to that of female. VOLUME 109, NUMBER 1 Leg 6 (Fig. 31) posteroventral flap on genital somite bearing 2 slender setae. Color of living specimens as in female. Etymology.—The specific epithet am- plus, Latin meaning large, alludes to the rel- atively large size of this species compared to congeners. Remarks.—Orecturus amplus may be distinguished by its size. None of its six congeners exceeds a length (in the female) of 1.25 mm, with their average length of only 1.11 mm. Selected features for further differentiation are as follows. In O. braca- tus (Stock & Kleeton 1963) the genital dou- ble-somite is rectangular. In O. excavatus Humes, 1989, the outer side of segment 1 of the exopod of leg 1 is excavated and the posterior part of the claw of the maxilliped is swollen. In O. finitimus Humes, 1993, the enlarged seta on segment 3 of the antenna is longer than the segment and the basis of the claw of the male maxilliped has a low prominence. In O. forticulus Humes, 1993, the prosome is broad, the maxilla has an unusually stout claw, the somite bearing leg 3 has sharply pointed epimera, and the first segment of the exopod of leg 1 has an outer thornlike process. In O. grandisetiger Hu- mes, 1992, the prosome is broad, the en- larged seta on segment 3 of the antennule is longer than the segment, the somite bear- ing leg 3 has sharply pointed epimera, and the basis of the male maxilliped has a weak inner protuberance and lacks spinules. In O. sakalavicus Humes, 1994, the antennule lacks an enlarged seta, the somite bearing leg 3 has sharply pointed epimera, and the basis of the male maxilliped has an inner lobe and lacks spinules. The outer spinules on the basis of the maxilliped seen in the new species are found only in O. forticulus. The color of living colonies of the soft 117 coral host, Siphonogorgia variabilis, was as follows: slender creamy white stems, small- er branches and twigs pale yellow, polyps lavender blue to red. The red color of Or- ecturus amplus more closely resembles that of the polyps than that of other parts of the colony, suggesting that the copepods may live on or in the polyps. Literature Cited Humes, A. G. 1989. Acontiophorus excavatus, a new species (Copepoda: Siphonostomatoida) asso- ciated with the soft coral Dendronephthya (A\- cyonacea) in the Indo-Pacific.—Proceedings of the Biological Society of Washington 102:916— 923. 1990. Synopsis of lichomolgid copepods (Poecilostomatoida) associated with soft corals (Alcyonacea) in the tropical Indo-Pacific.— Zoologische Verhandelingen 266:1—201. . 1992. Copepoda associated with the thorny coral Antipathes (Antipatharia) in the Indo-Pa- cific.—Journal of Natural History 26:709-744. 1993. Copepoda associated with gorgona- ceans (Cnidaria) in the Indo-Pacific.—Bulletin of Marine Science 53:1078—1098. 1994. Copepoda associated with octocorals in northwestern Madagascar, including Orectu- rus sakalavicus n. sp. from the telestacean Coe- logorgia palmosa.—Transactions of the Amer- ican Microscopical Society 113:117—126. , & R. U. Gooding. 1964. A method for study- ing the external anatomy of copepods.—Crus- taceana 6:238-240. , & J.-S. Ho. 1968. Cyclopoid copepods of the genus Lichomolgus associated with octocorals of the families Xeniidae, Nidaliidae, and Teles- tidae in Madagascar.—Proceedings of the Bio- logical Society of Washington 81:693—750. , & J. H. Stock. 1973. A revision of the family Lichomolgidae, cyclopoid copepods mainly as- sociated with marine invertebrates —Smithson- ian Contributions to Zoology 127:1—368. Stock, J. H., & G. Kleeton. 1963. Copépodes associés aux invertébrés des cdtes du Roussillon 3.— Acontiophorus bracatus n. sp. un cyclopoide si- phonostome associé aux six octocoralliaires.— Vie et Milieu 14:551—560. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):118—137. 1996 Records of Enchytraeidae (Annelida: Oligochaeta) from west Florida. 1. Mesenchytraeus, Cognettia, Bryodrilus, Hemienchytraeus, Henlea and Buchholzia Brenda Healy Department of Zoology, University College, Belfield, Dublin 4, Republic of Ireland Abstract.—Records of eleven species of Enchytraeidae belonging to six gen- era are given for terrestrial habitats in Florida west of Talahassee. All are new records for the southern part of North America. A revised diagnosis of Mes- enchytraeus 1s provided and a new species, Mesenchytraeus hamiltoni, is de- scribed which is characterised by the occurrence of secondary pharyngeal glands in V and a very short spermathecal ectal duct. Four species of Cogettia occurred, including two new species: Cognettia floridae which belongs to the group of species with three pairs of primary and two pairs of secondary pha- ryngeal glands and male organs in the normal position, and a species for which the description is incomplete. Bryodrilus novaescotiae, Hemienchytraeus ste- phensoni and H. bifurcatus are fully described and the occurrence of the wide- spread species Henlea perpusilla, H. ventriculosa and Buchholzia fallax is re- corded. The species recorded and described in this paper occurred in four series of collec- tions of Enchytraeidae made in West Flor- ida (west of Talahassee) during March 1984, April-May 1987, March 1988 and December 1993. Collection procedures and the range of habitats sampled have been de- scribed in a previous paper that also con- tains comments on faunal composition and ecology (Healy 1989). The enchytraeid fau- na was found to be rich but patchily dis- tributed. Among the 70—75 taxa recognised were at least 24 species believed to be new to science, a further 10 species new to North America and 18 already known from the continent but new to Florida. The large number of new species and new regional records is explained by the fact that there have been few studies of enchytraeids in North America and almost none in the Southeast. The only enchytraeids previous- ly recorded from Florida are two species of Grania from the marine sublittoral on the Atlantic coast (Kennedy 1966, Coates & Erséus 1985), Stephensoniella marina from the marine littoral on Virginia Key (Coates 1983) and the freshwater species Barbidri- lus paucisetosus from the Appalachicola River (Loden & Locy 1980). There are no records of terrestrial species. This paper records details of 11 terrestrial species belonging to six of the 13 genera found in West Florida. Most of the previ- ously known species were originally de- scribed from European populations, thus some morphological differences are to be expected in North American specimens. When Florida specimens depart from the original descriptions or those of European material as given by Nielsen & Christensen (1959), these differences are noted. A full description of Bryodrilus novaescotiae is given because the original description was based on a small number of fixed specimens only. Hemienchytraeus stephensoni and H. bifurcatus are also fully described, extend- ing the known range of variability in these widespread species. VOLUME 109, NUMBER 1 Materials and Methods Samples were collected from a wide range of habitats, particularly moist or wet humus in woodlands or swamps, and soil from capillary zones at the edge of water bodies such as bayheads, ponds and streams. Enchytraeids were absent or in poor condition in dry soils and rare in sub- merged substrates where they were gener- ally replaced by Tubificidae. The species described in this paper occurred at nine lo- calities. Worms were extracted from soil samples using the wet funnel method which uses light and heat from light bulbs to expel worms from samples into water-filled fun- nels (O’Connor 1955, Healy & Rota 1992). Some specimens of all species were exam- ined live and measurements of live worms or their organs were made on worms lightly compressed under a cover slip. Other ma- terial was fixed in 70% ethanol or Bouin’s fixative and stored in 70% ethanol. Most Specimens were stained in paracarmine and whole-mounted in Canada Balsam but a few unstained mounts were also prepared. Type specimens and other material have been deposited in the United States Nation- al Museum of Natural History, Washington D.C. (USNM) and a few specimens are in the Royal Ontario Museum (Invertebrate Zoology), Toronto (ROMIZ). Genus Mesenchytraeus Eisen, 1878 Mesenchytraeus Eisen, 1878:67. Analycus Levinsen, 1884:230. Mesenchytraeus Eisen 1879:10, 1904:14— 17; Michaelsen 1887:369—372, 1889:15— 16; Cejka 1914:5; Welch 1920:42—44; Cernosvitov 1937a:278; Nielsen & Chris- tensen 1959:30-31. Type species.—Mesenchytraeus primae- vus Eisen, 1878. Following the listing of Eisen’s species M. primaevus, M. mirabilis and M. falci- forrnis (Eisen 1878) as species dubia by Nielsen & Christensen (1959), Brinkhurst 119 & Jamieson (1971) designated Analycus Levinsen, 1884 as the correct name for the genus on grounds of priority. However, Pip- er et al. (1982) re-examined Ejisen’s mate- rial and concluded that the specimens were undoubtedly members of Mesenchytraeus although their poor condition precluded the distinction of specific characters. As the original descriptions were deemed ade- quate, they therefore restored the name Mesenchytraeus designating M. primaevus as type species. Diagnosis.—Medium to large worms, usually white but sometimes yellow due to colored coelomocytes or black when the epidermis is pigmented. Setae sigmoid, dis- tinctly nodulated or with slight swellings. Head pore at or near the apex of the pro- stomium. Clitellum well developed, some- times extends over several segments. Paired pharyngeal glands on septa at 4/5 and 5/6, secondary pharyngeal glands, when pres- ent, ventral in two to five segments. Tran- sition between esophagus and intestine gradual. Peptonephridia absent. Esophagus and intestine without diverticula or append- ages. Dorsal vessel originates in, or more usually behind, clitellum. Coelomocytes of one type, nucleate, rather small for the size of the worm, round, oval or spindle-shaped. Nephridia with poorly developed interstitial tissue, allows the coils of the canal to be seen clearly, preseptale consisting of a ne- phrostome on a short or long neck, post- septale with two or three lobes. Brain rather short, often broader than long, its anterior border indented or cleft. Paired seminal vesicles usually present, sometimes extend- ing within the egg sac into post-clitellar segments. Sperm funnel small, barrel- shaped or cylindrical. Sperm duct often en- larged to form an atrium just before it en- ters penial bulb. Atrial glands (prostates) and penial glands present or absent. Egg sac present, often extends through several seg- ments behind clitellum. Spermathecae sim- ple or with diverticula, free or fused to esophagus. Remarks.—The above diagnosis is a 120 modified version of those given by Eisen (1878, 1879, 1904), and Nielsen & Chris- tensen (1959), and takes into account more recent descriptions. Nodulated setae were considered diagnostic by Nielsen & Chris- tensen but were not mentioned by Eisen, either in his generic diagnoses or in any of his detailed species descriptions. Subse- quent reviews of the genus (e.g., Michael- sen 1887, 1889; Cejka 1914, Welch 1920, Cernosvitov 1937a) similarly omitted men- tion of the character. Re-examination of the material described in Eisen’s 1878 paper by Piper et al. (1982) confirmed the presence of noduli in all three species and it must be assumed that the character was overlooked by early workers. The nodulus may be very weak, however, and may be represented by little more than a slight swelling, as in the new species described below, making it dif- ficult to recognise for an inexperienced worker. A nodulus is unusual in enchy- traeids and is present in only one other ge- nus. However, reliance should not be placed on this character when constructing or us- ing keys to genera. Other modifications of existing diagnoses concern the postseptal regions of the nephridia, which Nielsen & Christensen describe as bilobed but which frequently have three lobes, and the egg sac, which Eisen describes as single and median but which may extend backward as two distinct sacs. Eisen (1904) described the penial bulb as a muscular cushion con- taining radiating muscular strands, which in some species are powerful and usually con- tain internal glands (penial glands), but the bulb structure is variable and the whole structure may be quite compact. Although not always present, an atrium, formed by expansion of the sperm duct at the entrance to the penial bulb, sometimes with attached glands, is unique in the Enchytraeidae. None of the characters proposed by Eisen (1878, 1879) as grounds for erecting the new genus has proved to be useful; these are: the presence of sperm balls or sper- matophores in the coelom (not recorded in any other enchytraeid genus but only pres- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ent in some Mesenchytraeus species), brain deeply divided in front (not unique to Mes- enchytraeus, indentation sometimes shal- low), and an unusually short vas deferens (usually short but not always exceptionally so). Cardiac glands (Herzk6rper of Mi- chaelsen), mentioned by early workers, are internal structures in the dorsal vessel, not easily detected in whole mounts. In prac- tice, the most reliable character that distin- guishes Mesenchytraeus is the nephridium with its free nephrostome (also found in some other genera), poorly developed in- terstitial tissue and lobed postseptale. In live worms, the nephridia have a very dis- tinctive appearance that makes Mesenchy- traeus specimens instantly recognisable. Distribution.—Eisen (1904) described 18 species of Mesenchytraeus from western parts of North America (mainly Alaska and California) and a further ten species have been described from western regions by other workers; in contrast, only three spe- cies are recorded from the eastern part of the continent (Tynen 1975). The genus also appears to be well represented in Siberia (Nurminen 1973a, Piper et al. 1982, Timm & Popchenko 1978, Timm 1994) while only eleven species are recognised in Eu- rope. Very few species have been found in the tropics. The genus thus appears to have its present centre of distribution in northern regions of the western Nearctic and eastern Palearctic. The poor representation in West Florida, where only one species was at all frequent, supports this view. North America, South America, Green- land, Europe, Spitzbergen, Siberia, Japan, Antarctic, ?Africa. Mesenchytraeus hamiltoni, new species Fig. 1 Material examined.—Holotype: USNM 170722 whole mounted specimen from edge of “‘sewage pond’’, University of West Florida, Pensacola, collected by author, De- cember 1993. Paratypes from the type lo- VOLUME 109, NUMBER 1 Fig. 1. 121 Mesenchytraeus hamiltoni, new species. A, anterior segments, dorsal view; B, dorsal setal bundle; C, brain; D, spermatheca of live worm, contracted; E, spermatheca of live worm, expanded; E coelomocytes; G, dorsal clitellar gland cells and clear interspaces; H, sperm funnel, live specimen; I, sperm funnel, fixed specimen; J, male pore and associated structures; K, nephridium. am ampulla, at atrium, br brain, dv dorsal vessel, ed ectal duct, hp head pore, mp male pore, pb penial bulb, phb pharyngeal bulb, pph primary pharyngeal gland, pr prostate, sd sperm duct, sp spermatheca, sph secondary pharyngeal gland. cality: USNM 170723—170726, two stained and two unstained whole mounts, one with clitellum partly removed. Other material: seven whole mounts in author’s collection. Approximately 16 live mature specimens examined. Type locality.—Wet or saturated peaty sand with moss and plant roots at edge of a pond receiving clean water from a sewage treatment plant, water level constant. pH 4.5-5.3. Etymology.—The species is dedicated to Professor Paul Hamilton of the University of West Florida who introduced me to many interesting and rewarding localities. Description.—Medium to large, white, yellowish or pink worms; live specimens 18-30 mm, 0.7 mm in diameter. Fixed specimens often much contracted, mature individuals measuring 8-22 mm with a prominent clitellum up to 1.2 mm in di- ameter. Segments in mature specimens 55— 86 (X = 66, SD 7.9, n = 11), 55-75 in submature ones. Setae sigmoid with nodu- lus poorly developed or absent, without en- tal hook (Fig. 1B), 108—120 pm in the pre- clitellar region, 120-144 «wm in posterior segments. Lateral bundles with three or four setae, ventral bundles with 3—12 in precli- tellar segments, 3—11 behind the clitellum, setae about equal in length within bundle. Setae absent dorsally in XII and XIII, oc- casionally present ventrally in XII, reduced in number or absent ventrally in XIII. Cu- 122 taneous glands in two or three rows at about the level of the setae, or scarcely visible. Clitellum extends from “XI to % or all of XIII with numerous small gland cells con- taining coarse granules, irregularly scat- tered with clear interspaces, that mask in- ternal organs in live as well as stained spec- imens (Fig. 1G). Gland cells may be almost continuous ventrally in fully mature indi- viduals. Head pore at, or near, tip of pro- stomium. Primary pharyngeal glands on 4/5 and 5/6 free dorsally, either unlobed or with two or three lobes; secondary glands, each with several lobes, situated ventrally in V, VI, VII and usually in VIII (Fig. 1A). A few specimens had either one or two glands in IX and one individual possessed a lobed gland unilaterally in X (Fig. 1A). Two small bulbs on dorsal posterior border of the phar- ynx in IV (Fig. 1A). Chloragocytes small, brown, present from IV, form a dense layer from VIII, 12—16 cells across the intestine in the preclitellar region in mounted speci- mens. Esophagus merges with intestine without increase in diameter, intestine en- larges behind origin of dorsal vessel. Coe- lomocytes 14-30 wm, oval or spindle- shaped, with rather coarse granules and a weakly staining nucleus (Fig. 1F). Dorsal vessel originates in XI—XIII with dilatations in VII—XII and conspicuous branches in II and III (Fig. 1A); bifurcation anterior to brain. Blood faintly or distinctly red. Brain about as broad or slightly broader than long, about 80 wm in length, truncated or slightly indented posteriorly, deeply indent- ed on its anterior border (Fig. 1C). Nephrid- ial nephrostome elongated, on a long neck, postseptale with two or three lobes; efferent duct arises mid-ventrally (Fig. 1K). First nephridia at 6/7, occasionally at 5/6. Testes large and spongy in mature worms, appear brown by transmitted light in living specimens. Seminal vesicles paired, confined to XI or extend into X. Sperm funnels small, barrel-shaped, 120— 160 ym in length, about 1.5 times as long as wide, with a raised collar about as wide PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON as the funnel in fixed specimens, appearing slightly narrower in live worms (Fig. 1H, I). Sperm duct 14—16 wm wide, about ten times length of funnel, loosely coiled in XII. Male openings irregular or semicircu- lar slits. Each sperm duct widens distally to form a thick-walled, fusiform atrium that opens centrally into penial bulb; three large, slightly lobed glands attached to bulb (Fig. 1J). Two egg sacs develop asymmetrically on 11/12 as pouches, each containing a string of oocytes and eggs, reaching to XVIII on one side. Ovarian tissue diffuse, extending back in egg sacs which develop before sperm funnels and ducts. Up to five mature eggs present at a time. Spermathe- cae simple, without diverticula, each con- sists of an elongated ampulla that tapers en- tally to merge with esophagus in V and ec- tally to a short ectal duct, only a little longer than wide, scarcely longer than thickness of body wall (Fig. 1D, E). Ampullae expand and contract and occasionally pulsate. When contracted, wall becomes rugose or folded internally (Fig. 1D). Only small amounts of sperm, arranged lengthwise, in ampulla. Spermathecae develop early and are fully formed in specimens only half the final size and in which there are no signs of male organs or oocytes. Remarks.—Among the ten or so de- scribed species of Mesenchytraeus with spermathecae attached to the esophagus in V, but which lack diverticula, M. hamiltoni is unique in the shortness of the ectal duct of its spermatheca and in the presence of secondary pharyngeal glands in V. These features are deemed sufficient to character- ise the species although details of structures associated with the male opening, often an important diagnostic character in the genus, have not been described for most of the species with which it might be compared. Mesenchytraeus hamiltoni most resembles M. glandulosus (Levinsen, 1884) which has five pairs of secondary pharyngeal glands but has a spermathecal ectal duct equal in length to one third of the ampulla, lacks secondary pharyngeal glands in V and has VOLUME 109, NUMBER 1 a nephridial funnel without a neck. The number of pharyngeal glands was variable for individuals of the same size and stage of maturity. Although the species was plentiful at the type locality and at several other sites, ma- ture specimens were rare. None was found in March 1984, only four in April-May 1987, three in March 1988 and seven in De- cember 1993. A low proportion of adults in populations is common in the genus (per- sonal observation). Habitat.—Mesenchytraeus hamiltoni was common in the Pensacola region and was sometimes the dominant enchytraeid in wet substrates. It occurred in a variety of habitats including woodland leaf litter in valleys and swamps, Sphagnum and pitcher plant (Sarracenia) bogs, and saturated soils at the edge of water bodies where the high- est densities occurred. At the latter sites they were often accompanied by Tubifici- dae. The pH range was 3.9—5.8. Distribution.—In and around the campus of the University of West Florida, Pensa- cola; Blackwater Forest, at the edge of a swamp; Avalon Peninsula, edge of a pitcher plant (Sarracenia) bog; Lakewood, among Sphagnum and grassroots at edge of lake. Not found in the east of the study area. Mesenchytraeus sp. Material examined.—Seven live, imma- ture specimens. Remarks.—A somewhat yellowish spe- cies was present at several hardwood forest sites but no fully mature specimens were available for a definitive identification. Specimens had two pairs of primary and two pairs of secondary pharyngeal glands and spermathecae communicating with the esophagus in V, each with a pair of elon- gated diverticula. The species resembles M. beumeri (Michaelsen, 1886) in general ap- pearance. Habitat and distribution.—In hardwood leaf litter. Occasional at the University of West Florida, Pensacola and in Torreya 123 State Park; common at Tall Timbers Forest Research Station. Genus Cognettia Nielsen & Christensen, 1959 Cognettia species are recognised by sig- moid setae, without nodulus, in bundles of two or three; nephridia with preseptale con- sisting of funnel only and an antero-ventral or mid-ventral origin of the efferent duct; absence of peptonephridia; absence of esophageal and intestinal diverticula; dorsal vessel originating in or behind clitellum; a simple penial bulb and a simple spermathe- ca, not attached to the oesophagus. Testes, Ovaries and male pores may be displaced forward by up to four segments. Distribution.—The only record of Cog- nettia in North America is for C. glandu- losa from Montreal, Canada (Nurminen 1973b). The genus has proved so far to be characteristic of wet, acid soils in cold or cold-temperate lands and members are usu- ally the dominant enchytraeids in tundra and coniferous forests of northern Europe. However, the discovery of four species in Florida shows that Cognettia species can survive in a warm climate while the pres- ence of two new species suggests that North America, like Europe, may have its own en- demic species. Canada, Greenland, Europe, Spitzbergen, Siberia, Japan, Ecuador, Antarctic. Cognettia floridae, new species Fig. 2 Material examined.—Holotype: USNM 170727, stained whole mount, Edward Ball Nature Trail, University of West Florida, Pensacola, a hardwood, bayhead swamp, collected by the author April 1987. Para- types from the type locality and other sites in the University of West Florida: USNM 170728—170732, two stained and two un- stained whole mounts. Other material: ROMIZ 13242-13243, stained whole mounts; 11 whole mounts in the author’s collection; 42 live specimens examined. 124 Fig. 2. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 100 pm Cognettia floridae, new species. A, anterior segments, lateral view; B, dorsal clitellar gland cells and clear interspaces; C, brain; D, spermatheca of live worm; E, spermatheca of fixed specimens; E nephridium; G, coelomocytes; H, setae; I, sperm funnel, live worm. am ampulla, br brain, dv dorsal blood vessel, ec ectal glands, ed ectal duct, mu muscle; pph primary pharyngeal gland, sg secondary pharyngeal gland, sp spermatheca. Type locality.—In moist, wet or saturated sandy humus under Taxodium and _ hard- wood trees in a partly tidal, freshwater bay- head swamp; some areas subject to periodic flooding. pH 3.9—4.7. Etymology.—Named after the State of Florida, genitive case. Description.—Live specimens greyish- white, 7—9 mm, fixed specimens 5—9 mm, diameter 0.5—0.52 mm, 0.56 mm at clitel- lum. Segments (29)39—46 (X 40.9, SD = 3.6, n = 23). Setae without ental hook (Fig. 2H), 80—96 pm in preclitellar region, 80— 104 wm in posterior segments, three per bundle in all regions, occasionally only two, setae absent in XII. Cutaneous gland cells small, in numerous rows or more or less scattered. Clitellum prominent in live worms, extending over XII—XIII with squarish gland cells arranged in transverse rows. In stained mounts, the gland cells, which are packed with coarse granules, are irregularly scattered with clear interspaces, absent mid-ventrally (Fig. 2B). Head pore at O/1. Three pairs of primary pharyngeal glands, all free dorsally, without ventral lobes, and two pairs of compact secondary glands, situated ventrally in V and VI (Fig. 2A). Esophagus merges gradually with in- testine from 6/7. Chloragocytes present from IV, forming a dense layer from VI, 10—12 across intestine in preclitellar region in compressed specimens. Dorsal vessel arises at 13/14, occasionally in XIV or XV, anterior vessel bifurcates at level of 0/1. Blood colorless in specimens from drier habitats, faintly or distinctly red in those from wet substrates. Coelomocytes round or oval, finely granular with a prominent nucleus (Fig. 2G), 14—32 wm long, about a quarter of length of setae. Nephridial ne- phrostome on a short neck, long efferent duct arises antero-ventrally or mid-ventrally VOLUME 109, NUMBER 1 (Fig. 2F). First nephridium at 6/7. Brain 160—200 pm long, 1.5—2 times longer than wide, deeply incised posteriorly (Fig. 2C). Seminal vesicle unpaired, dorsal, con- fined to XI. Sperm funnels small, cylindri- cal, in live worms about 100 pm long, four or five times longer than wide, equal to about 4—'%3 diameter of live worm (Fig. 21). In fixed specimens, length usually about 4 of worm diameter. Collar slightly narrower than funnel. Sperm duct long and narrow, diameter 6 pm, coiled in XII, opens through a compact penial bulb, greatest diameter about 90 wm, which occupies % to Y; of the diameter of clitellum in mounted speci- mens. One, or usually two, eggs present at a time. Spermathecae confined to V. Ectal ducts thick-walled, about four times length of ampullae, each with a mass of fused, elongate cells on anterior face, near opening (Fig. 2D, E). Ampulla dorsal to oesophagus in V, thin-walled, 35—50 pm long, 1.2 times longer than wide, with most of sperm ar- ranged lengthwise and extending into prox- imal part of ectal duct in most specimens (are221D: EB). Remarks.—Cognettia floridae belongs to a group of species that comprises C. cog- nettii (Issel, 1905), C. lapponica Nurminen, 1965, C. hibernica Healy, 1975 and C. zic- sit D6zsa-Farkas, 1989 all of which repro- duce sexually and have reproductive organs in the normal position for Enchytraeidae. It resembles C. lapponica, C. hibernica and C. zicsii in having three pairs of primary pharyngeal glands and two pairs of second- aries and spermathecae confined to V but it is unique in the asymmetrical ectal swell- ings on the spermathecal ducts. Habitat.—The species was common in and around the campus of the University of West Florida and plentiful at two other lo- calities. It was found in a range of sub- strates including sandy peat and peaty sand, Sphagnum, coniferous and broad-leaved leaf litter, finely divided humus and among roots of aquatic plants at the edge of water bodies. All sites were wet, saturated or flooded, with pH levels 3.9—4.5, among the 125 lowest for the region. Mature individuals were common. Distribution.—Pensacola, campus of the University of West Florida; Avalon Penin- sula, pitcher plant (Sarracenia) bog; Lake- wood, edge of a lake. Not found in the east- ern part of the study area. Cognettia sp. Fig. 3 Material examined.—Two live mature specimens collected in April 1984 were available for the following description but both were damaged during examination and no further mature individuals could be found. The remaining material consists of one stained, whole-mounted, submature specimen, USNM 170732, three stained whole mounts of immature specimens USNM 170733-170735 and ten whole mounts of immatures in the author’s collec- tion. Approximately 30 live, immature in- dividuals were also examined. Description.—Medium sized, rather slen- der worms, creamy white, internal organs partly masked by abundant coelomocytes in live worms. Length of live, mature speci- mens 10—12 mm; live, non-fragmenting im- matures reach 20 mm. Maximum length of fixed immature specimens 12 mm, diameter 0.28—0.37 mm. Segments 50 and 58 in the mature specimens but reach 64 in immatu- res. Setae without ental hook, 68—84 pm in all regions, three ventrally, three or some- times two laterally. Setae absent from IX or X in mature individuals. Clitellum only slightly raised, extends over IX—%2X (over X—%2X]I in submature specimen), gland cells irregularly scattered or partly in transverse rows (Fig. 3D). Male pores in X or XI. Cu- taneous glands small, in numerous rows or scarcely visible. Head pore at, or just an- terior to O/1. Five pairs of unlobed, primary pharyn- geal glands in large individuals but fifth pair often small, developing unilaterally or absent (Fig. 3G). Only four pairs in mature worms. Anterior glands may be united dor- 126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. glands; E, preclitellar nephridium; EF postclitellar nephridium; G, anterior segments, lateral view. am ampulla, br brain, ed ectal duct, eg ectal gland, mu muscle, ph pharyngeal gland, nc ventral nerve cord. sally while posterior ones are usually free. Secondary pharyngeal glands absent. Esophagus merges gradually with intestine. Chloragocytes present in a dense layer from behind last pair of pharyngeal glands i.e., in VIII or LX, 10—14 across the intestine in compressed specimens. Dorsal vessel from XIV or XV, anterior bifurcation just behind 0/1. Blood colorless or faintly red. Coelom- ocytes 20—30 wm, round or oval with a sharp outline, packed with fine granules that make them appear brown by transmitted light in living worms (Fig. 3B). They are Cognettia sp. A, spermatheca, live worm; B, coelomocytes; C, sperm funnel, live worm; D, clitellar usually very abundant, often so densely packed as to mask internal organs. Nephrid- ia with free nephrostome and elongate pre- septale, efferent duct long, arising antero- ventrally or mid-ventrally (Fig. 3E, F). Brain about 1.5 times longer than broad with a truncate or sinuous posterior border. Seminal vesicle present, unpaired. Sperm funnels cylindrical, about four times longer than wide, each with narrower, scarcely raised, collar (Fig. 3C). Sperm ducts long and narrow, opening at compact penial bulbs. Spermathecal ectal ducts long, slen- VOLUME 109, NUMBER 1 der, without swellings, leading to ovoid am- pullae in VII (Fig. 3A). One large ectal gland anterior to each spermathecal open- ing. Remarks.—The above description is in- complete owing to the shortage of mature specimens but the distinctive appearance of live individuals made them easily recognis- able in samples, even as immatures, and there seems little doubt that this is a new species. Many worms were regenerating an- terior or posterior segments and the species obviously reproduces principally by frag- mentation; sexual individuals were rare in the population. The species is close to C. sphagnetorum (Vejdovsky, 1878) which re- produces in the same way and which also has four or five pairs of primary pharyngeal glands and no secondary glands. Cognettia sphagnetorum, however, has a chamber containing a ring of sperm at the junction between the ectal duct of the spermatheca and the ampulla and more anterior male openings (in VIII or IX). Other Cognettia with displaced reproductive organs, namely C. glandulosa (Michaelsen, 1888), C. paxi (Moszynski, 1938) and C. anomala (Cer- nosvitov, 1928), all have secondary pharyn- geal glands. While there are no important diagnostic differences between immature forms of C. sphagnetorum and the present species, the abundant coelomocytes with sharp outline and densely granular cyto- plasm give the Florida species a distinctive appearance. Habitat and distribution.—University of West Florida, Pensacola, in the drier parts of a bayhead swamp and in an adjoining ravine with hardwoods. Common in moist, but not wet, leaf litter and humus, pH 3.6— 4.5, above the zone subject to periodic flooding. Cognettia ?sphagnetorum (Vejdovsky, 1878) Pachydrilus sphagnetorum Veydovsky 1878: 304. Cognettia sphagnetorum Nielsen & Chris- 127 tensen 1959:42-43, figs. 28, 29; Kas- przak 1986:124-125, figs. 332-334; Chalupsky 1992:142, fig. 10. Material examined.—Two stained, whole- mounted immature specimens in the author’s collection. Three live immature specimens examined. Remarks.—The absence of mature indi- viduals makes a definitive identification im- possible, but specimens resembled C. sphagnetorum from Ireland in general ap- pearance although they had only four pairs of pharyngeal glands. Nielsen & Christen- sen (1959) give five pairs for Danish ma- terial, except in mature individuals, but I have found in Irish populations that the fifth pair is often absent in fragmenting individ- uals. Fragmentation is the normal method of reproduction in C. sphagnetorum and sexual individuals are always rare. Material from Florida included one specimen with a developing clitellum and one which ap- peared to be a juvenile, with only 20 seg- ments, although according to Christensen (1959) the species does not seem to be ca- pable of producing viable eggs. Habitat and distribution.—Lakewood, among grass roots and leaf litter at the edge of a lake, pH 4.24.4. Europe, Iceland, Spitzbergen, Antarctic. (Some of these re- cords are based on immatures only.) Cognettia ?glandulosa (Michaelsen, 1888) Pachydrilus sphagnetorum var. glandulosus Michaelsen, 1888:483. Marionina glandulosa Issel 1905:455—456, fig. 3. Enchytraeoides glandulosus Cernosvitov 1928:16—-17, Pl. 1, figs. 10, 11. Cognettia glandulosa Nielsen & Christen- sen 1959:43—-44, fig. 30; Kasprzak 1986: 125-126, figs. 335-337. Material examined.—USNM 170736— 170737, two stained, whole-mounted im- mature specimens; two whole mounts in the author’s collection; five live, immature in- dividuals examined. 128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 100 um Fig. 4. Bryodrilus novaescotiae. A, anterior segments, dorsal view; B, spermatheca, live worm; C, sper- mathecal pore, surface view; D, setal bundle; E, preclitellar nephridium; FE postclitellar nephridium; G, brain; H, coelomocytes; I, clitellar glands, dorsal view; J, sperm funnel, live worm; K, male pore, ventral view. am ampulla, br brain, ch chloragocytes, dv dorsal blood vessel, ed ectal duct, ec ectal glands, en ental duct, eg esophageal gland, sp spermatheca. Remarks.—The distinctive feature of this species is the presence of five pairs of pri- mary and five (occasionally four) pairs of secondary pharyngeal glands. Immature specimens are commonly identified on the basis of this character alone. The sperma- theca is also distinctive. No sexual individ- uals were found, however, during this sur- vey, thus a positive identification is not pos- sible. Specimens resembled C. glandulosa from Ireland in general appearance. The presence of regenerating fragments indicat- ed that individuals were reproducing by fragmentation. Habitat and distribution.—Ponce de Leon, leaf litter on floodbank of stream is- suing from limestone, pH of litter 5.34. Canada, Greenland, Iceland, Europe, Sibe- ria, Antarctic. Genus Bryodrilus Ude, 1892 Diagnostic characters of the genus are sigmoid or curved setae, without nodulus, of different sizes within a bundle; four esophageal diverticula in VI; peptonephri- dia and intestinal diverticula absent; dorsal vessel originates within or behind the cli- tellum; coelomocytes of uniform size and shape; preseptale of nephridium consists of funnel only, efferent duct arises anteriorly, near the septum (“‘Henlea type’’) or medi- ally; compact penial bulb; simple sperma- thecae whose ental ducts usually unite be- fore communicating with the dorsal wall of the esophagus. Distribution.—So far confined to the northern Holarctic, the majority of species being recorded from North America and Si- beria. Alaska, Canada, Greenland, Europe, Siberia. Bryodrilus novaescotiae Bell, 1962 Fig. 4 Bryodrilus novaescotiae Bell, 1962:169— 171, Pl. iv, figs. 1-9. VOLUME 109, NUMBER 1 Material examined.—Three stained whole mounts USNM 170746—170748; three stained whole mounts ROMIZ 13241, 13244-13245, 17 whole mounts in the au- thor’s collection. Approximately 30 live specimens examined. Type material not lo- cated. Description of new material.—Medium to large species, white or yellowish due to color of chloragogen tissue and abundant coelomocytes. Length variable; live speci- mens 10-20 mm, reach 30 mm _ when stretched; fixed specimens 8-16 mm, di- ameter 0.7—0.8 mm, slightly wider at clitel- lum. Segments (37)45—64 (X = 54.2, SD = 5.5, n = 19). Setae sigmoid, without ental hook, three or four in lateral bundles, five or six (occasionally seven) in anterior ven- tral bundles, four to six ventrally behind cli- tellum (Fig. 4D). Setae diminish in size from outside of bundle to mid-line and measure 50—100 pm in preclitellar region, 112-128 wm in posterior segments. Cuta- neous gland cells in about ten rows anteri- orly, more noticeable at level of setae, eight or nine rows per segment behind clitellum. Clitellum extends over XII and 4—% XIII, only slightly raised, with numerous small, coarsely granular, irregularly scattered gland cells and polygonal interspaces (Fig. 41). Ventrally, gland cells nearly continu- ous. Head pore a transverse slit just anterior to O/1. Three pairs of lobed pharyngeal glands, all free dorsally and with ventral lobes (Fig. 4A). Four esophageal glands in VI, closely applied to the esophagus, situated dorso-lat- erally and ventro-laterally, each with five to seven regular lobes (Fig. 4A), apparently solid, no internal canals seen. Chloragocy- tes small, average 10—15 in number across intestine in compressed specimens, form a dense layer from 6/7 at point where gut widens gradually. Coelomocytes numerous, round or oval, finely granular, nucleus not visible in live worms but distinct when stained, diameter (16)20—42 pm (Fig. 4H). In live individuals, coelomocytes appear brown by transmitted light. Dorsal vessel 129 originates at 11/12 or sometimes 12/13, with segmental dilatations in VI—XI, ante- rior bifurcation at 0/1. Blood colorless. Ne- phridia with small, free nephrostome and elongate preseptale, postseptale with long, narrow efferent duct that arises antero-ven- trally in preclitellar region, mid-ventrally in posterior segments (Fig. 4E, F). First ne- phridium at 6/7. Brain usually about 1.5 times longer than wide but sometimes shorter with a sinuous posterior border and straight or indented anterior border (Fig. 4G). Testes small, oval and compact with a smooth outline. Seminal vesicle unpaired, confined to XI. Sperm funnels cylindrical or taper distally, each with a distinct collar of about same width as funnel (Fig. 4J), length in live specimens 150—180 p.m, three or four times longer than wide, up to five times longer when stretched, equal to %4 to ¥% of diameter of worm. In mounted speci- mens, funnel has a smooth outline and is usually bent into a semicircle. Sperms duct long and narrow, coiled in XII, about 6 wm in diameter; each opens through a large but compact penial bulb, about 60 pm in lon- gitudinal diameter but sometimes smaller. In fixed specimens, bulb occupies about % of diameter of clitellum. Male openings semicircular slits (Fig. 4K). One egg pres- ent at a time. Spermathecae large, more or less confined to V (Fig. 4B). Ectal duct of spermatheca composed of two parts roughly equal in length, a stout, thick-walled section communicating with one side of ampulla, and a large, bulbous, asymmetrical mass of cells surrounding a narrow chamber that opens to the outside through a straight or curved slit at 4/5, surrounded by a protrud- ing ring of large cells forming a rosette at the surface up to 190 wm across (Fig. 4C). Ampulla thin-walled, roughly pear-shaped in live specimens but contracting to become more spherical when fixed. Ampullae taper entally, ental ducts usually unite just before communicating with esophagus at 5/6 or in anterior part of VI (Fig. 4B), but occasion- ally communicate separately. Ampullae and 130 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ental ducts contain abundant sperm, mostly arranged lengthwise. Remarks.—The distinctive character of B. novaescotiae is its spermathecae. The large swellings near the openings of the ec- tal ducts are not present in any other mem- ber of the genus. Bell (1962) also consid- ered the solid esophageal glands to be unique as they are hollow in other species. His specimens differed chiefly in the form and dimensions of the spermathecae, each of which had a large, hemispherical cham- ber in the ectal bulb, a smaller “‘spherical”’ ampulla and a longer common ental duct. He mentioned two ectal glands in addition to the mass of fused cells around the orifice, but these are not shown in his figure (Bell 1962, fig. 3). Bell also noted a hemispher- ical chamber within the penial bulb which could not be detected in mounted speci- mens from Florida. Other differences in Bell’s specimens are a shorter brain, about half as long as wide, irregular-shaped coe- lomocytes (possibly resulting from poor fixation) and nephridia with a large presep- tale and an efferent duct that arises termi- nally but is folded back to give the impres- sion of leaving subterminally or even near the septum. This latter character cannot be considered significant since in sectioned material it can be difficult to tell whether the distal portion of the nephridial duct is folded inside or outside the sheath of the postseptale. Bell did not mention lobes on the esophageal glands, which are not pres- ent in other known species of Bryodrilus, the dimensions of the sperm funnel, or the presence of a seminal vesicle. Habitat and distribution.—University of West Florida, Pensacola, in drier parts of a hardwood swamp and an adjoining wooded ravine, mainly above the level of periodic flooding. Soil a moist, spongy humus with numerous fine roots, pH 3.5-4.5. Canada (Nova Scotia). Genus Hemienchytraeus Cernosvitov, 1934 Hemienchytraeus species possess a me- dian, unpaired, bifurcated peptonephridium arising from the dorsal wall of the pharynx, which is unique in the Enchytraeidae. Other useful characters are setae in pairs; absence of esophageal or intestinal diverticula; ne- phridia with large preseptale enclosing the nephrostome; long sperm ducts usually coiled into a spirals; small, compact penial bulbs and free spermathecae each of which has an ectal duct without glands at the opening and a simple ampulla without di- verticula. Distribution.—Hemienchytraeus species were the most abundant and widespread en- chytraeids in West Florida, especially in moist or wet habitats. Preliminary sampling in Peninsular Florida (unpublished) indi- cates that the same is true there. The genus is mainly tropical; only one species is re- corded from Europe and is uncommon. Eu- rope, India, Japan, Africa, South America. Hemienchytraeus stephensoni (Cognetti, 1927) Fig. 5, A-I Enchytraeus cavicola Stephensen, 1924 (non Joseph 1880):127—129, Pl. 6, figs. 1-4. Enchytraeus stephensoni Cognetti, 1927:4. Enchytraeus myrmecophilus Cernosvitov, 1930a:85—89, figs. 1-9. Enchytraeus rangoonensis Stephensen, 19317 INV 79s tiene Hemienchytraeus stephensoni (E. cavicola = E. stephensoni + E. myrmecophilus + E. rangoonensis) Cernosvitov, 1934:298— 304, figs. 1-11. Hemienchytraeus stephensoni Cernosvitov 1939:92—-93, figs. 43-49; Christoffersen 1979:40—-46, figs. 1-23; Nakamura 1984: 32-33, Fig. 1B, C, F; Dozsa-Farkas 1989:200—202, figs. 24-33. Material examined.—Five stained, whole mounts, USNM_ 170741-170745, four stained, whole mounts ROMIZ 13239— 13240, 13246-13247; 30 stained, whole mounts in the author’s collection. Approx- imately 50 live specimens examined. Description of new material.—Small to VOLUME 109, NUMBER — 131 Fig, 5. D, preclitellar nephridium; E, postclitellar nephridium; EF brain; G, anterior setal bundle; H, posterior setal bundle; I, coelomocytes. Hemienchytraeus bifurcatus. J, peptonephridium, K, sperm funnel, live worm; L, spermatheca. Buchholzia fallax. M, esophageal gland in IV. pb primary branch, sb secondary branch, tb tertiary branch. medium-sized worms, live specimens rather transparent, with a prominent clitellar re- gion due to presence of large eggs. Live specimens 8-10 mm, fixed specimens 6—9 mm, diameter 0.2—0.3 mm, 0.24—0.38 mm at clitellum. Segments 36—48 (X = 44.1, SD = 2.7, n = 29). Setae two per bundle, oc- casionally three, with weak ental hook. Se- tae absent in XII. In anterior region setae straight or slightly curved and rather small, 32—40 pm long, more curved, thicker and longer in posterior half of worm, 50-62 wm in terminal segments (Fig. 5G, H). Cuta- neous gland cells small and inconspicuous, in about four rows per segment. Prostomi- um with numerous dark-staining gland cells. Clitellum over XIH—“XIII, gland cells small, arranged in more or less regular transverse rows, especially in ventral region and on anterior and posterior borders of cli- tellum. Glands may be absent mid-ventral- ly. Head pore near tip of prostomium. Some Hemienchytraeus stephensoni. A, peptonephridium; B, sperm funnel, live. worm; C, spermatheca; specimens were in mucus tubes encrusted with particles of organic matter. Three pairs of pharyngeal glands, all united dorsally and with ventral lobes. Pep- tonephridium arises from dorsal side of pharynx in III, proximal portion of variable length, thick-walled with a coiled or sinu- ous lumen, primary and secondary branch- es narrower but also with coiled lumina (Fig. 5A). Each secondary branch divides into three to five thin-walled, finger-like processes. Entire peptonephridium confined to IV. Chloragocytes from V, form a dense layer from VII, 11—13 cells across intestine in preclitellar region in compressed speci- mens. Transition between esophagus and intestine gradual with enlargement behind pharyngeal glands. In all specimens col- lected from the Pensacola region in 1993, a section of the intestine behind the clitellum, usually between XIV—XVI and XXVIII-— XXX, was enlarged and brown in colour. In 132 other specimens, the brown region was re- duced or absent. Coelomocytes sparse, of various sizes and shapes, but mainly round, 8-18 wm, almost hyaline in live individu- als, faintly granular with a distinct nucleus in fixed specimens (Fig. 51). Dorsal vessel originates in XIII, usually with isolated, yellow, dendritic cells on surface of vessel (seen in live worms); anterior bifurcation at 0/1. Blood colorless. Nephridia with large, ovoid preseptale that encloses a dorsally di- rected nephrostome, postseptale of variable shape, usually ovoid in preclitellar seg- ments where duct arises mid-ventrally and elongated in posterior segments where duct is sub-terminal or terminal (Fig. 5D, E). First nephridium at 6/7. Brain about 1.5 times longer than wide, deeply indented or even cleft on its anterior border, posterior border sinuous or straight (Fig. 5F). Testes compact. Small, paired seminal vesicles usually present in XI. Sperm fun- nels funnel-shaped, somewhat flattened, 190—360 pm long, 40—112 wm wide, some- what longer than diameter of worm, gen- erally six to eight times longer than wide in live specimens, with collar about same width or slightly narrower than rest of fun- nel (Fig. 5B). Funnels taper to spirally coiled ducts 5—6 wm in diameter, which open at small, compact penial bulbs, lon- gitudinal diameter 54—82 ym. Male pores curved slits. One to three large eggs present at a time, contained in an egg sac that ex- tends to 14/15 or rarely 15/16. Spermathe- cae with long, thick-walled ectal ducts, 12— 18 wm in diameter, outer surface rough, which swell to form sperm-containing chambers in V or VI, continue as smooth, thin-walled tubes, and terminate in thin- walled, cylindrical ampullae in VI-IX (Fig. 5C). Length of spermathecae variable; am- pullae may be twice or several times longer than wide, swollen and occupying up to 2% segments when mature. Remarks.—Hemienchytraeus stephen- soni 1S a very variable species, hence the number of nominal species synonymised by Cernosvitov (1934). The variability has PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON been commented on by Cernosvitov (1939), who noted differences in the length of the basal part of the peptonephridium and in the form and complexity of the septal (pharyn- geal) glands in South American specimens. Christoffersen (1979) distinguished four types among Brazilian material from a sin- gle locality that differed in body length and width, setal length, pharyngeal glands, form of the peptonephridium, origin of dorsal vessel, number and shape of nephridia, form of the penial bulbs, length of sperm funnels and length of spermathecae and am- pullae. He concluded that H. stephensoni is a complex of cryptic species. Two extreme forms were recognised among West Florida material that differed in lengths of setae, sperm funnels, spermathecal ampullae and the proximal section of the peptonephri- dium and in the form of the pharyngeal glands, but intermediates were also found. Size and segment number and dimensions of setae and sperm funnel are within the range of specimens from South America (Christoffersen 1979, Do6zsa-Farkas 1989) and Japan (Nakamura 1984), but specimens differ in having both secondary and tertiary branches to the peptonephridium and in the presence of seminal vesicles. No individu- als were found with more than two second- ary branches on each side of the peptone- phridium, as in material from Lake Titicaca (Cernosvitov 1939) and Ecuador (D6zsa- Farkas 1989). In spite of Christoffersen’s ability to recognise distinct types within a region, and the differences between mate- rial from Florida and South America, I be- lieve H. stephensoni to be one variable spe- cies with a wide ecological tolerance and a wide geographic range. Habitat and distribution.—Hemienchy- traeus. stephensoni was by far the most common enchytraeid in the region and was recorded from all localities and most kinds of inland habitat. Pensacola, woodland, swamp, edge of a pond, floodbank of a river and boggy area, Avalon Peninsula, pitcher plant (Sarracen- ia) bog; Ponce de Leon, woodland at edge VOLUME 109, NUMBER 1 of a stream; Lakewood, edge of a lake; Marianna, woodland on limestone and bank of a river; Torreya State park, woodland on limestone; Port St. Joseph, edge of roadside ditch. pH range 3.6—7.2. Mainly tropical. South America (Argen- tina, Paraguay, Brazil, Bolivia, Ecuador), India, Burma, Central Africa, Japan. Re- corded from almost every conceivable ter- restrial habitat including woodland, grass- land, tree bark, anthills, bat guano and the marine littoral zone. New for North Amer- ica. Hemienchytraeus bifurcatus Nielsen & Christensen, 1959 Fig. 5J—L Hemienchytraeus bifurcatus Nielsen & Christensen 1959:45, figs. 23-27. Hemienchytraeus bifurcatus Nakamura 1984:31-32, fig. 1A, D, E.; Kasprzak 1986:178, figs. 639-643. Material examined.—Three stained, whole mounted specimens USNM 170746— 170748; one stained whole mount ROMIZ 13248; seven stained whole mounts in the author’s collection. Approximately 28 live specimens examined. Description of new material.—Medium to small worms, live specimens 4—6 mm, transparent when viewed microscopically, with a prominent clitellar region due to presence of rather large eggs. Fixed length 4—6 mm, diameter 0.3—0.4 mm, 0.3—0.45 mm at clitellum. Segments (25)29—44 (X = 35.2, SD = 4.9, n = 20). Setae straight or slightly curved with a weak ental hook, two per bundle throughout, absent in XII, 32— 58 m. Cutaneous glands small, indistinct. Clitellum over XII-“XIII with small gland cells arranged in more or less transverse rows or sometimes irregularly, smaller or absent in mid-ventral region. Head pore near tip of prostomium. Some specimens were in mucus tubes with adhering debris and soil. Three pairs of pharyngeal glands, all lobed and united dorsally. Peptonephridium 133 arises from mid dorsal region of pharynx in Ill, proximal section of variable length, branching dichotomously into two primary and four secondary branches (Fig. 5J). Proximal part of unpaired section hollow and contractile in live specimens, distal sec- tion and primary branches stout with a coiled or sinuous lumen, secondary branch- es thin-walled with a wide lumen. Entire peptonephridium confined to IV. Chlora- gocytes from V, forming a dense layer from VII. Esophageal-intestinal transition gradu- al, gut expands behind pharyngeal glands. Coelomocytes sparse, of various shapes and sizes, dominant type round or oval, 10-14 zm, nucleated, granular. Dorsal vessel orig- inates in XIII or XIV, anterior bifurcation at 0/1. Blood colorless. Nephridial presep- tale large, postseptale egg-shaped with pos- tero-ventral efferent duct in preclitellar re- gion, more elongated with terminal or sub- terminal duct behind clitellum. Nephridia starting at 4/5 or 5/6, absent X—XVI. Brain rectangular, about 1.5 times longer than wide, deeply indented or even cleft on its anterior border, more or less truncated pos- teriorly. Small, paired seminal vesicles usually present, sometimes only developed on one side. Sperm funnels funnel-shaped, three or four times longer than broad in live worms, 100—150 pm in length, equal to % or % di- ameter of worm, with a collar equal to or slightly narrower than rest of funnel (Fig. 5K). Funnels taper to long, spirally-coiled ducts in XII, open at small, compact penial bulbs, about 32 wm across. One to three large eggs present at a time, egg sac ex- tending to XIV. Spermathecae long, each consists of a fairly stout, thick-walled ectal duct, without glands at the orifice, duct swells to form a sperm-containing chamber in V or VI and then extends as a thin-walled tube to an ovoid or spherical ampulla situ- ated in any of segments VI-IX, usually in VII (Fig. 5L). Remarks.—This species, like the previ- ous one, is very variable, especially in size, form of the nephridia and lengths of sper- 134 mathecae and sperm funnels. The different dimensions of the reproductive organs are not related to stage of maturity, for individ- uals with sperm in the ampulla may have long or short spermathecae and either short or long sperm funnels. Danish specimens (Nielsen & Christensen 1959) differed in having a brain indented posteriorly, ne- phridia with terminal efferent ducts, coe- lomocytes with refringent granules, no sem- inal vesicle and a sperm funnel only 2-3 times longer than wide. A longer sperm funnel was also reported in Japanese spec- imens (Nakamura 1984). The species is similar in general appear- ance to HA. stephensoni but is generally smaller and has somewhat more granular coelomocytes, more noticeable in live spec- imens. The two species often occurred to- gether and although the form of the pepto- nephridium is quite different, this was sometimes difficult to see clearly. Mature individuals could be reliably distinguished by the greater dimensions of the sperma- thecal ampulla and sperm funnel in H. ste- phensoni, as noted by Nakamura (1984). Habitat and distribution.—Not as abun- dant, nor as widespread as H. stephensoni. Pensacola, woodland leaf litter at the Uni- versity of West Florida, grass cuttings at Scenic Hills golf course; Avalon Peninsula, pitcher plant (Sarracenia) bog; Ponce de Leon, woodland soil and leaf litter; Marian- na, woodland soil and leaf litter on lime- stone. pH range 3.9—6.9. Denmark, France, Japan, India. New for North America. Genus Henlea Michaelsen, 1889 The principal diagnostic characters of the genus are straight or slightly curved setae, usually of unequal size within a bundle, the outer ones longer, esophagus expands abruptly into the intestine (with the possible exception of the sub-genus Hepatogaster), esophageal appendages present, intestinal diverticula present or absent, a preclitellar origin of the dorsal vessel, usually in VIII or [X, and nephridia with free nephrostome PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and anterior origin of the efferent duct, of- ten described as “‘Henlea type’ but not unique to that genus. Cosmopolitan: E and W of North Amer- ica, S America, Europe, Siberia, Africa, Antarctic. Henlea perpusilla Friend, 1911 aug. Cernosvitov 1937b Henlea perpusilla Friend, 1911:466—467. Henleanella perpusilla Friend, 1913a:89. Henlea bisetosa Friend, 1914:135. Henlea inusitata Friend, 1913a:83—84. Henlea minima Friend, 1913a:84. Henlea nivea Cernosvitov, 1930b:88, figs. 3-6. Henlea perpusilla Friend 1913b:270—271, figs. 34-35; Cernosvitov 1937a:194—196, figs. 2-5; Nielsen & Christensen 1959: 58-59, figs. 46, 47; Kasprzak 1986:256— 257, figs. 992-996. Material examined.—Two stained, whole mounts USNM_ 170749-170750; two stained whole mounts in the author’s col- lection. Live specimens examined, 27. Remarks.—The large number of syn- onyms that are now recognised reflects a wide variation in size and morphology (Cernosvitov 1941, Nielsen & Christensen 1959), which is partly explained by the ex- istence of different cytotypes (Nielsen & Christensen 1959). Size of the live speci- mens (6-8 mm) and segment number (30— 33) of Florida material are within the range of European specimens. Habitat and distribution.—Common in Marianna State Park, deciduous woodland on limestone, sandy humus, sandy alluvial deposits and humus with limestone frag- ments; Tall Timbers Research Station, live oak forest, dark, sandy loam and leaf litter. pH range 5.5—7.1. Absent from the west of the study area. Canada (Cornwallis Is., Prince of Wales Is., Devon Is., Rocky Mountains), Greenland, Europe, Lebanon, Siberia, Japan, Antarctic, Bolivia. VOLUME 109, NUMBER 1 Henlea ventriculosa d’Udekem, 1854 Enchytraeus ventriculosus d’Udekem, 1854: 863, figs. 1, 4, 6-9. Henlea multispinosa Friend, 1913a:85. H. (Udekemiana) ventriculosa Cernosvitov, 1930b:75. Henlea groenlandica Cernosvitov, 1929: 146, figs. 1, 2. H. (Udekemiana) groenlandica Cernosvi- tov, 1930b:75. Fridericia stewarti Stephenson, 1909:109. Henlea ventriculosa Michaelsen, 1889:31— 32, 1900:69-70; Nielsen & Christensen 1959:62, figs. 50, 53, 57; Kasprzak 1986: 264-265, figs. 1028-1030. Material examined.—Two stained, whole mounts USNM_ 170751-170752; three stained, whole mounts in the author’s col- lection. Five live specimens examined. Remarks.—Size, 5—8 mm when fixed, and segment number (38—44) of Florida in- dividuals are within the range of European specimens but the spermathecae have short- er ectal ducts, about equal to the length of the ampullae, while each ental duct is lon- ger than the combined lengths of ampulla and ectal duct. In addition, there are several small glands at the ectal opening. Danish specimens lacked these glands, the ectal duct was longer and the ental duct shorter (Nielsen & Christensen 1959). In spite of these differences, the species could be iden- tified with confidence by the characteristic gut diverticula. Habitat and distribution.—Marianna State Park, deciduous woodland on lime- stone, rotting wood, leaf litter, humus with limestone fragments. pH range 5.3—6.7. Canada (Cornwallis Is., Prince of Wales Is.), USA (Massachusetts, N. Carolina), Eu- rope, Siberia, Tibet, Japan, New Zealand, S. America. Genus Buchholzia Michaelsen, 1887 The genus is recognised by the following unique combination of characters: sigmoid 135 setae, decreasing in size within bundles to- ward the dorsal and ventral midlines of the body; hollow esophageal appendages in IV; transition between esophagus and intestine abrupt with one or two dorsal diverticula at 7/8; anteclitellar origin of the dorsal vessel in the region of the intestinal diverticula; and small, hyaline, anucleate coelomocytes in addition to the normal kind. So far, confined to the northern hemi- sphere but not recorded from Asia. Canada, Europe, N. Africa. Buchholzia fallax Michaelsen, 1887 (Fig. 5M) Buchholzia fallax Michaelsen, 1887:374— 376, pl. 21, fig. 4A—E. Buchholzia fallax Beddard, 1895:334—335; Michaelsen 1900:72-73; Cernosvitov 1928:9; Ude 1929:57-58; Nielsen & Christensen 1959:65, figs. 59-61; Wilcke 1967:72, fig. 6A, B; Kasprzak 1986:117— 118, figs. 294—298, table 8. Material examined.—Two stained, whole mounts USNM 170753—170754; ten whole mounts in the author’s collection. Live specimens examined, 18. Remarks.—Distinguished from B. appen- diculata (Buchholz, 1862), which was re- corded from Montreal by Nurminen (1973b), by the presence of one, rather than two, intestinal diverticula and only three in- stead of four pairs of pharyngeal glands, and by having male organs in the usual po- sition for enchytraeids. All specimens ex- amined are small, 4-7 mm compared with 12-14 mm for specimens from Denmark (Nielsen & Christensen 1959). Segments 30—38 (Danish material 37—42). A seminal vesicle is usually present, stated to be ab- sent by Nielsen & Christensen and not men- tioned by Michaelsen (1887). The pulsating esophageal appendages in IV are of a dif- ferent shape from those figured by Nielsen & Christensen (1959:143, fig. 60); they are elongate with partial cross walls dividing the cavity into three compartments (Fig. 5M), rather than rosette-like. The small, 136 secondary cavities in the spermathecal am- pulla, figured by Michaelsen, could not be seen and have not been mentioned by other authors. Some specimens were inside mu- cus tubes encrusted with rings of soil par- ticles. Habitat and distribution—Common in Marianna State Park, deciduous woodland on limestone. A few in a similar habitat in Torreya State Park. pH range 5.5—7.1. Ab- sent from the west of the study area. Eu- rope, N. Africa. New for North America. Acknowledgments Support for this work was provided by the Mary Ball Washington Foundation at the University of West Florida, Pensacola. I am grateful to members of the Department of Biology for useful information on the re- gion and in particular to Paul Hamilton for his help in locating the wide range of hab- itats sampled. The friendly hospitality of the Department was much appreciated. Literature Cited Brinkhurst, R. O., & B. G. M. Jamieson. 1971. Aquat- ic Oligochaeta of the world. Oliver and Boyd, Edinburgh, 860 pp. Beddard, E. F 1895. A monograph of the order of the Oligochaeta. Clarendon Press, Oxford. Bell, W. A. 1962. Enchytraeids (Oligochaeta) from various parts of the world.—Transactions of the American Microscopical Society 81:158—177. Buchholz, R. 1862. Beitrage zur Anatomie der Gat- tung Enchytraeus.—Schriften Physikalisch- 6konomischen Gesellschaft zu K6nigsberg 3: 93-132. Cejka, B. 1914. Die Oligochaeten der Russischen in den Jahren 1900-1903 unternommenen Nord- polarexpedition. III. Uber neue Mesenchy- traeus-Arten. IV. Verzeichnis der wahrend der Expedition gefunden Oligochaeten-Arten.— Mémoires de |1’Académie Impériale des Sci- ences de Petrograd, 8th series, 29:1—31, plates I-IV. Cernosvitov, L. 1928. Die Oligochaeten der Karpath- en.—Zoologische Jahrbiicher, Abteilung Syste- matik, Okologie und Geographie der Tiere 55: 1-28. 1929. Communication preliminaire sur les oligochetes récoltées par M. P. Remy pendant la croisiere arctique effectuée par le ““Pourquoi- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pas?” en 1926 sous la direction du Dr. J.-B. Charcot.—Bulletin du Muséum national d’ His- toire naturelle 2 144-149. . 1930a. Eine neue Enchytraeiden-Art aus dem Ameinenhaufen Argentiniens.—Zoologischer Anzeiger 88:85—89. . 1930b. Sur quelques oligochétes de la région arctique et des Iles Faeroer—Annales des Sci- ences naturelles, Zoologie, Paris 14:65—110. . 1934. Zur Kenntnis der Enchytraeiden. I1.— Zoologischer Anzeiger 105:295—305. . 1937a. System der Enchytraeiden.—Bulletin de |’ Association russe pour les recherches scientifiques 4 Prague 5:263—294. . 1937b. Zur Kenntnis der Enchytraeiden. II Revision der Friendschen Enchytraeidentyp- en.—Zoologischer Anzeiger 117:191—205. . 1939. Report of the Percy Sladen Trust Ex- pedition to Lake Titicaca, IV. Oligochaeta.— Transactions of the Linnaean Society 1:81—116. . 1941. Revision of Friend’s types and descrip- tions of British Oligochaeta——Proceedings of the Zoological Society of London 111:237—280. Chalupsky, J. 1992. Terrestrial Enchytraeidae (Oli- gochaeta) and Parergodrilidae (Polychaeta) from Sweden, with description of a new enchy- traeid species.—Zoologica Scripta 21:133—150. Christensen, B. 1959. Asexual reproduction in the En- chytraeidae (Olig.).—Nature 184:1159-1160. Christoffersen, M. L. 1979. Some enchytraeids (Oli- gochaeta) from Sierra do Mar, Sao Paulo, Bra- zil.—Boletim de Zoologia da Universidade do Sao Paulo 4:39-52. Coates, K. A. 1983. A contribution to the taxonomy of the Enchytraeidae (Oligochaeta). Review of Stephensoniella, with new species records.— Proceedings of the Biological Society of Wash- ington 96:411-419. , & C. Erséus. 1985. Marine enchytraeids (Oli- gochaeta) of the coastal northwest Atlantic (northern and mid U.S.A).—Zoologica Scripta 14:103-116. Cognetti, L. 1927. Lumbricidi dei Carpazi.—Bolleti- no dei Musei e Laboratorii di Zoologia e di An- atomia comparata della R. Universita di Genova (Ser. 2.) 7:18. Dozsa-Farkas, K. 1989. Neue Enchytraeiden-Arten (Oligochaeta) aus Ekuador—Acta Zoologica Hungarica 35:191—203. Eisen, G. 1878. Redogorelse for Oligochaeter, sam- lade under de Svenska expeditionerna till Ark- tiska trakter—Ofversigt af Kunglinga Veten- skaps-Akademiens Forhandlingar 3:63—79. 1879. On the Oligochaeta collected during the Swedish Expeditions to the Arctic regions in the years 1870, 1875 and 1876.—Kunglinga Svenska Vetenskaps-Akademiens Forhandligar 15:1-49, plates 1-16. VOLUME 109, NUMBER 1 1904. Enchytraeidae. Harriman Alaska Ex- pedition. Alaska XII. Doubleday, Page and Co., New York, 123 pp., pl. E-XX. Friend. H. 1911. New British Henleas.—The Zoolo- gist, (Ser. 4) 15:464—468. . 1913a. A key to British Henleas.—The Zo- ologist, (Ser. 4) 17:81—91. . 1913b. British enchytraeids V. Species new to science.—Journal of the Royal microscopical Society 255-271. . 1914. British enchytraeids VI. New species and revised list—Journal of the Royal micro- scopical Society 128-154. Healy, B. 1975. A description of five new species of Enchytraeidae (Oligochaeta) from Ireland.— Zoological Journal of the Linnaean Society 56: 315-326. . 1989. Preliminary report on the Enchytraei- dae (Oligochaeta) of West Florida.—Hydrobiol- ogia 180:47-56. , & E. Rota. 1992. Methods for collecting En- chytraeidae during expeditions.—Soil Biology and Biochemistry 24:1279—-1281. Issel, R. 1905. Oligocheti della fauna italiana I. En- chytreidi di Val Pellice.—Zoologische Jahr- bticher 22:451—476. Kasprzak, K. 1986. Skposzczety wodne i glebowe, II. Rodzina: Wazonkowce (Enchytraeidae). Pan- stwowe Wydawnictwo Naukowe, 366 pp. Kennedy, C. R. 1966. A taxonomic revision of the genus Grania (Oligochaeta: Enchytraeidae).— Journal of Zoology 148:399—407. Levinsen, G. M. R. 1884. Systematisk-geographisk Oversigt over de nordiske Annulata, Gephyrea, Chaetognathi, og Balanoglossi.—Videnskabeli- ge Meddeleser fra dansk naturhistorisk Foren- ing 1 Kgébenhaven (1883):92—350. Loden, M. S., & S. M. Locy. 1980. Barbidrilus pau- cisetous, new genus, new species (Oligochaeta: Enchytraeidae), from eastern America.—Pro- ceedings of the Biological Society of Washing- ton 93:1173-1176. Michaelsen, W. 1886. Uber Chylusgefasssysteme bei Enchytraeiden.—Archiv fiir mikroscopische Anatomie 28:294. 1887. Enchytraeiden-Studien.—Archiv fiir mikroscopische Anatomie 30:366—376, pl. 21. 1888. Beitrage zur Kenntnis der deutschen Enchytraeidenfauna.—Archiv fiir mikroskopis- che Anatomie, Bonn 31:483—498. . 1889. Synopsis der Enchytraeiden.—Abhan- dlungen und Verhandlungen des Naturwissen- schaftlichen Vereins in Hamburg 11:3-—59, plate. . 1900. Oligochaeta. Das Tierreich, 10. Lief, Friedlander und Sohn, Berlin. 575 pp. Moszynski, A. 1938. Marionina paxi, ein neuer Bor- stenwurm von den Haberwiesen.—Pax. Beitra- 137 ge zur Biologie des Glatzer Schneeberge 4:343— 345. Nakamura, Y. 1984. Enchytraeids in Japan (3).—Eda- phalogia 31:31—34. Nielsen C. O., & B. Christensen. 1959. The Enchy- traeidae, critical revision and taxonomy of Eu- ropean species.—Natura jutlandica 8—9:1—160. Nurminen, M. 1965. Enchytraeids (Oligochaeta) from northern Norway and western Lapland.—An- nales Zoologici fennici 2:11—12. . 1973a. Distribution of northern enchytraeids (Oligochaeta).—Annales Zoologici fennici 10: 483-486. . 1973b. Enchytraeidae (Oligochaeta) from the vicinity of Montreal, Canada.—Annales Zool- ogici fennici 10:399—402. O’Connor, FE B. 1955. Extraction of worms from a coniferous forest soil—Nature, London 175: 815-816. Piper, S. R., S. E MacLean, & B. Christensen. 1982. Enchytraeidae (Oligochaeta) from taiga and tundra habitats of north eastern U.S.S.R.—Ca- nadian Journal of Zoology 60:2594—2609. Stephenson, J. 1909. Report on a collection of smaller Oligochaeta, made by Capt. EK H. Stewart, J. M. S., in Tibet.—Record of the Indian Museum, Calcutta 3:105—114. . 1924. Oligochaeta of Siju Cave, Garo Hills, Assam.—Record of the Indian Museum 26: 127-135, pl. 6. 1931. Descriptions of Indian Oligochaeta IIl.—Record of the Indian Museum 33:173—202. Timm, T. 1994. Propappidae and aquatic Enchytraei- dae (Oligochaeta) from the farthest southeast of Russia.—Hydrobiologia 278:67—78. , & V. Popchenko. 1978. The aquatic Oligo- chaeta of the Murmansk region. In Seasonal phenomena in freshwater biology.—Hydrobio- logical researches of the Limnological Station, Institute of Zoology and Botany, Academy of Sciences of the Estonian SSR:71—132. Tynen, M. J. 1975. A checklist and bibliography of the North American Enchytraeidae (Annelida: Oligochaeta).—Syllogeus 9:2-14. Ude, H. 1892. Ein neues Enchytraeiden-Genus.— Zoologischer Anzeiger 15:344—345. d’Udekem, J. 1854. Description d’une nouvelle es- péece d’Enchytraeus.—Bulletin de 1’ Académie royale scientifique de Belgique 21:583—864. Vejdovsky, E 1878. Zur Anatomie und Systematik der Enchytraeidae.—Sitzungsberichte der K. Boh- mischen Gesellschaft der Wissenschaften, Prag 1877:294—304. Welch, P. S. 1920. The genera of the Enchytraeidae (Oligochaeta).—Transactions of the American Microscopical Society 39:25—50. Wilcke, D. E. 1967. Oligochaeta. Jn P. Brohner et al., eds., Die Tierwelt Mitteleuropas, 1 (7). Quelle und Meyer, Leipsig, 161 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):138-142. 1996 Artacama valparaisiensis, a new species of Terebellidae (Annelida: Polychaeta) from subtidal soft bottoms of Valparaiso Bay, Chile Nicolas Rozbaczylo and Marco A. Méndez Departamento de Ecologia, Facultad de Ciencias Bioldgicas, P. Universidad Catélica de Chile, Casilla 114-D, Santiago, Chile Abstract.—A new species of terebellid, Artacama valparaisiensis, is de- scribed from Valparaiso Bay, Central Chile. The species closely resembles A. crassa Hartman, 1967, from the South Shetland Islands, Antarctica, but can be distinguished on the basis of the following morphological criteria: shape and number of proboscidial papillae, size and number of the branchial filaments, relative size of lobe at the upper edge of uncinal ridges in abdominal parapodia and presence of constrictions in the limbate setae. Systematics of the Terebellidae was re- viewed recently by Holthe (1986). Accord- ing to his review, Artacama Malmgren, 1866, with eight valid species is the only genus in subfamily Artacaminae. McHugh (1995) has carried out a cladistic analysis of the Terebellidae in which she shows that Artacama is the sister taxon of a large clade within the Amphitritinae. The Artacaminae has therefore been synonymized with the Amphitritinae, which is diagnosed by the presence of double rows of uncini on pos- terior thoracic segments. The character that identified the Artacaminae, a peristomium modified on its ventral side to form a con- spicuous proboscis-like organ adorned with papillae, is considered an autapomorphy for the genus Artacama (McHugh 1995). Only two species of Artacama, A. crassa Hart- man, 1967 and A. proboscidea Malmgren, 1866, have been reported from southeastern Pacific, in antarctic and subantarctic waters (Rozbaczylo 1985). From 1978 to 1980 an extensive research program on benthic communities at Valparaiso Bay, Central Chile, was carried out by Dr. José Stuardo (presently at Universidad de Concepcion) and Dr. Héctor Andrade (presently at A & A Tecnolab S.A., Divisi6n Ambiental). Three areas with different particle size (sand, sandy-mud and sandy-silt), organic matter and other biochemical components were sampled, from 34 to 59 meters depth. Samples from these areas have been studied by Stuardo et al. (1981) regarding their bio- chemical and granulometric composition. They showed that among the three sampled areas there were differences in proteins, lip- ids and organic matter; the sandy-mud area showed the highest values and sandy area the lowest values. Among the abundant polychaetes collected during that program numerous individuals of Artacama were found. These differed from the other known species reported for the genus, and conse- quently are described as a new species. Specimens of the new species were com- pared with the holotype of A. crassa de- posited in the National Museum of Natural History, Washington, D.C. (USNM 55569) and a specimen of A. proboscidea from Western Canadian Arctic (USNM 41132) identified by E. & C. Berkeley. Type and paratype specimens of the new species are deposited in the National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. (USNM); Sala de Sistematica, Departamento de Ecologia, VOLUME 109, NUMBER 1 Pontificia Universidad Cat6élica de Chile, Santiago (SSUC); and Museo de Zoologia, Universidad de Concepcién (UCCC). Materials and Methods Polychaetes were collected at Valparaiso Bay, Central Chile. Samples were taken monthly, from November 1979 to October 1980, with a 0.1 m* Van Veen dredge at three areas with different particle size (sand, sandy-mud and sandy-silt), from 34 to 59 meters depth. Polychaetes were fixed in 10% formalin and preserved in 70% eth- anol. Figures were prepared by means of a drawing tube on a Wild M-5 microscope. Scanning electron microscopic (SEM) ob- servations and photographs were obtained using a JEOL JSM-25SII microscope. Artacama valparaisiensis, new species lanes, Il, ZW Material examined.—Central Chile: Val- paraiso Bay, off Punta Osas, ca. 32°59'20'S, 71°33'56"W, 44 m, H. Andrade, coll., 9 Jul 1979, holotype (USNM_ 170005), and 14 paratypes (USNM 170006); 4 Sep 1979, 8 paratypes (UCCC 23169-23176), and 14 paratypes (SSUC 6719). Description.—Holotype incomplete with 52 setigers, measuring about 60 mm long, width about 9 mm at thorax. Body (Fig. 1a) thick anteriorly and colorless in alcohol, with first 8-11 segments enlarged, tapers posteriorly to a narrow pygidial end. Total length of one of the largest complete spec- imens, is 104 mm including proboscis, width 12 mm at thorax and 4 mm at ab- domen, and consists of 102 setigerous seg- ments; 17 thoracic setigers and 85 abdom- inal setigers. Large, ovoid and papillose proboscis (Fig. la) extends forward from buccal segment below mouth. Prostomium (Fig. 1b) is small folded, bilobed structure, with small oral aperture immediately below. Eyes absent. Small, horseshoe-shaped ten- tacular lobe on dorsal side of peristomium, with a dorsal indentation and numerous ten- 139 tacles, short and clubbed, most of which are broken. Proboscis covered overall with nu- merous, conical and minute, papillae (Fig. Ic) arranged in about ninety longitudinal rows. Lateral lappets absent on segments 2— 4. Three pairs of long filiform branchiae on segments 2—4; each branchia is a tuft of ap- proximately 50 equal-length filaments aris- ing from basal stump. Nephridiopores, in form of short tubes, number five pairs; the best developed are on segment 3 below sec- ond pair of branchiae, in line with the more posterior notopodia; smaller ones are on segments 6, 7, 8 and 9, between notopodia and uncinal ridges, postero-ventral to no- topodia. Ventral glandular pads present on first 10 setigers. Thorax with fixed number of setigers; seventeen bundles of notosetae Starting on segment 4; uncini first present from the second setiger (fifth segment); oc- cur in single rows on first six uncinal ridg- es, then in double rows from setigers 8 to 17, oriented “fang to fang.’’ Thoracic no- topodia (Fig. 1d) with triangular lamellae, postsetal lobe larger than presetal; 40—50 long, pointed setae decreasing in length from dorsal to ventral part of notopodia, lat- erally winged, with one or two constrictions (Fig. le). Thoracic neuropodia with avicu- lar uncini with a large fang surmounted by four or five alternating rows of many small teeth (Fig. 2a, b). Abdomen with numerous segments bearing parapodia (Fig. 1g, h) with flaplike tori and dorsally large, subcir- cular membrane which increase in size to- wards the posterior end; uncini in single rows restricted to ventral margin; abdomi- nal uncini resemble thoracic but main fang is longer and thinner and have six or seven alternating rows of teeth (Figs. If, 2c, d). Posterior end tapers to terminal pygidium (Fig. 11), with terminal anus, with crenulate sides. Geographical distribution—Known only from Valparaiso Bay, Chile. Remarks.—Artacama_ valparaisiensis new species resembles, most closely, A. crassa Hartman, 1967 and A. proboscidea Malmgren, 1866. However A. valparaisien- 140 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SQ noe! 009 99 rong aiyase riron nares» X N S ns N N N iN (J Lede { <i Fig. 1. Artacama valparaisiensis new species (USNM 170005). a, entire animal in left lateral view; b, prostomium in dorsal view with the oral aperture immediately below; c, proboscidial papillae; d, thoracic par- apodium, right side anterior view; e, limbate thoracic notoseta from fifth setiger, in lateral view showing con- strictions; f, abdominal uncinus in frontal and lateral view, from tenth abdominal parapodium; g, second abdom- inal parapodium, right side anterior view, with large foliaceous dorsal lobe and uncinal torus; h, tenth abdominal parapodium, right side anterior view, with large foliaceous dorsal lobe and uncinal torus; i, posterior end, in lateral view. Scales = 1 cm for a; 1 mm for b, g, h, i; 0.5 mm for c, d; 0.01 mm for f; 0.1 mm for e. VOLUME 109, NUMBER 1 141 Fig. 2. Artacama valparaisiensis new species, SEM micrographs: a, thoracic uncini in lateral view from the first uncinigerous segment (fifth thoracic segment); b, detail of the upper part of an uncinus of thoracic segment 11 (first segment with double rows of uncini); c, abdominal uncini in lateral view from the fourth abdominal segment; d, detail of the upper part of an uncinus of the fourth abdominal segment. Scales = 0.01 mm for a—d. sis differs from A. crassa in that papillae of the proboscis are more abundant, conical, and smaller; the number of branchial fila- ments is higher (ca. 50 in each branchiae) and each filament is longer and thinner; presence of constrictions in the limbate se- tae; thoracic notopodia with postsetal la- mellae longer than presetal lobe, while in A. crassa both lobes are short; abdominal parapodia with the uncinigerous tori longer and the dorsal membrane comparatively smaller. On the other hand, A. proboscidea shows fewer branchial filaments, shorter and thicker than those of A. valparaisiensis. Papillae of the proboscis of A. proboscidea are bigger and fewer than in A. valparais- iensis. Dorsal membrane of abdominal par- apodia of A. valparaisiensis is bigger than those of A. proboscidea. Absence of con- strictions in the limbate setae of A. probos- cidea. McIntosh (1885) described A. chal- lengeriae from several localities off Ker- guelen Islands, 46—202 m; Hartman (1959) considers this species similar to A. probos- cidea. Description of A. challengeriae is so general and vague that it is difficult to com- pare it with A. valparaisiensis. Neverthe- less, from figures (Pl. LI. fig. 6; PI. 142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON XXVIIIA. figs. 23a, b, 24, 25) in McIntosh (1885) can be established that A. challen- geriae differs from A. valparaisiensis at least in that the former has proboscidial pa- pillae bigger an fewer; edges of limbate se- tae are entire, without constrictions; ventral margin of uncini presents an ondulating outline while in A. valparaisiensis has a uniform and slight convexity the same as in A. crassa and A. proboscidea. Etymology.—tThe specific name valpa- raisiensis refers to the type locality, Val- paraiso Bay. Ecological remarks.—Artacama_ valpa- raisiensis Was more abundant between May and September, with densities of 100 indiv./ m? in May and 250 indiv./m? in September. The species showed the highest density in the sandy-mud area (30—250 indiv./m?), the second one was the sandy-silt area (5—100 indiv./m7); at the sandy area the species was extremely rare. In the sandy area few spec- imens of small size were found only in May and September while in the sandy-mud and sandy-silt areas numerous specimens of dif- ferent size were found throughout the sam- ple period. This pattern of distribution ap- pears to be correlated with the different or- ganic matter content of sediments in the sites sampled (Stuardo et al. 1981). In the population studied it was possible to distin- guish adult females by the presence of eggs in the coelomic cavity. Maximum egg size was 165 wm diameter. Acknowledgments We thank Dr. Jose Stuardo, Departamen- to de Oceanologia, Universidad de Concep- cion, who made specimens available for sudy and Dr. Héctor Andrade, A & A Tec- nolab S.A., Divisi6n Ambiental who fur- nished information about collecting. We specially thank Clara Yafiez for her skill and patience in producing the illustrations used in this paper and José Morillas for the scanning electron photomicrographs. We thank Dr. Kristian Fauchald of the National Museum of Natural History, Washington, D.C., for the loan of specimens for com- parison. The manuscript was greatly im- proved by the valuable comments and sug- gestions provided by Dr. Christopher J. Glasby and an anonymous reviewer who also provided valuable bibliography. Literature Cited Hartman, O. 1959. Catalogue of the polychaetous an- nelids of the World, part If.—Allan Hancock Foundation Publications Occasional Paper 23: 355-628. 1967. Polychaeta annelids collected by the USNS Eltanin and Staten Island Cruises, chief- ly from Antarctic Seas.—Allan Hancock Mono- graphs in Marine Biology 2:1—387. 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. Malmgren, A. J. 1866. Nordiska Hafs-Anunlater.— Ofversigt af Konglia Vetenskaps-Akademiens Forhandlingar, Stockholm 22:355—410, pls. 18— Dor McHugh, D. 1995. Phylogenetic analysis of the Am- phitritinae (Polychaeta: Terebellidae).—Zoolog- ical Journal of the Linnean Society (in press). McIntosh, W. C. 1885. Report on the Annelida Poly- chaeta collected by H.M.S. Challenger during the Years 1873-1876.—Jn Report on the Sci- entific Results of the Voyage of H.M.S. Chal- lenger during the Years 1873 under the Com- mand of Captain George S. Nares, R.N., ER.S. and the Late Captain Frank Tourle Thomson, R.N., Zoology, 12(34):1-554, plates 1-55, and 1A—39A, 1 map. Rozbaczylo, N. 1985. Los Anélidos Poliquetos de Chile. Indice sinonimico y distribucion geo- grafica de especies.—Monografias Bioldgicas 3:1—-284. Stuardo, J.. M. A. Soto, H. Andrade, & R. Aguilar. 1981. Caracteristicas granulométricas y com- ponentes bioquimicos de los sedimentos de tres estaciones submareales de Valparaiso.—Revista de Biologia Marina, Instituto de Oceanologia, Universidad de Valparaiso, 17(2):171—-196. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):143-149. 1996 Review of Hermilepidonotus Uschakov, 1974, and two species of polynoid polychaetes (Lepidonotinae) Marian H. Pettibone Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—The genus Hermilepidonotus Uschakov, 1974, and type species Lepidonotus robustus Moore, 1905, are reviewed, and compared with the close- ly related Lepidonotus helotypus (Grube, 1877), based on examination of types and available new material. Under Lepidonotus helotypus (Grube, 1877), from China, Seidler (1924:56) in- cluded Lepidonotus robustus Moore, 1905, from the Gulf of Alaska. This was followed by Annekova (1937), from the North Japan Sea, by Hartman (1938), from Alaska, and by Uschakov (1950, 1955), from the Okhotsk Sea. On further study, L. robustus was considered to be distinct from L. hel- otypus, by Hartman (1948) and by Uschak- ov (1974). Based mainly on a characteristic pharynx, the latter author referred L. ro- bustus to the new genus Hermilepidonotus. The close similarity of these two Pacific Ocean boreal species is the basis for adding L. helotypus to Hermilepidonotus. The two species are more fully described herein and some new records are presented. In addition to the specimens deposited in the National Museum of Natural History, Smithsonian Institution (USNM), types and additional material were obtained on loan or in ex- change from the following sources: the for- mer British Museum (Natural History), London (BMNH), now renamed the Natural History Museum, London, through J. D. George and A. I. Muir; National Science Museum, Tokyo (NSMT), through M. Ima- jima; Zoological Institute Academy of Sci- ence, Leningrad (ZIASL), through P. V. Us- chakov; Zoologisches Museum, Berlin (ZMB), through G. Hartwich; and Zoolo- gisches Staatsmuseum, Hamburg, through G. Hartmann-Schroder. Family Polynoidae Kinberg, 1856 Subfamily Lepidonotinae Willey, 1902 Genus Hermilepidonotus Uschakov, 1974, emended Type species.—Lepidonotus robustus Moore, 1905, by monotypy and original designation. For: H. robustus (Moore, 1905) and H. helotypus (Grube, 1877), new combination. Diagnosis.—Body elongate-oval, with 26 segments. Prostomium lepidonotoid, bi- lobed, with 3 antennae and 2 palps; cera- tophore of median antenna in anterior notch, lateral antennae inserted terminally on anterior extensions of prostomium, on same level as median antenna; 2 pairs of eyes on posterior half of prostomium. First or tentacular segment not distinct dorsally; tentaculophores lateral to prostomium, each with aciculum, few setae on inner side, and pair of dorsal and ventral tentacular cirri, similar to antennae. Elytra and large ely- trophores 12 pairs, on segments 2, 4, 5, 7, O) Mil, 13, WS; 15 WD, Zl, ZB. aly levexe, suboval, leaving middorsum uncovered, without fringes of papillae, thick, soft, with micropapillae on (H. robustus) or with mi- crotubercles and conical macropapillae (1. helotpyus). Dorsal cirri on non-elytrigerous segments, with cylindrical cirrophores and smooth styles; dorsal tubercles bulbous in- flated areas. Second or buccal segment with first pair of elytrophores, biramous para- 144 podia, and long ventral buccal cirri lateral to ventral mouth. Eversible muscular phar- ynx with 2 pairs of chitinous hooked jaws and border papillae; [about 15 pairs (13-17, in H. helotypus) or border papillae plus nu- merous branched, threadlike papillae, in H. robustus)|. Parapodia biramous, with small conical notopodia on anterodorsal sides of large neuropodia; neuropodia with subequal presetal and postsetal lobes, with smaller, truncate supraacicular parts and longer, di- agonally truncate subacicular parts. Noto- setae slender, with spinous rows, short, with blunt tips and longer, with fine tips. Neu- rosetae stout, with long spinous regions and long, bare, entire tips. Ventral cirri short, subulate. Nephridial papillae beginning on segment 8. Pygidium with anal ridge and pair of anal cirri. Hermilepidonotus robustus (Moore, 1905) Fig. 1 Lepidonotus robustus Moore, 1905:544, pl. 36:figs. 32—35.—Hartman, 1948:12. Lepidonotus helotypus.—Seidler, 1924:56 (part)—Annenkova, 1937:145, pl. 1:fig. 1, pl. 3:fig. 14—Hartman, 1938:109.— Uschakov, 1950:157; 1955:128, fig. 19, A-E; 1965:107, fig. 19, A-E.—Chlebov- itsch, 1961:165. Not Grube, 1877. Hermilepidonotus robustus.—Uschakov, 1974:458, figs. A-H (synonymy); 1982: 112, pl. 33, 1-8.—Buzhinskaja, 1985:82. Material examined.—Alaska: Shelikov Strait, 88-119 m, from hermit crab, Alba- tross sta 4291, 15 Aug 1903, holotype of L. robustus (USNM 5523). Virgin Bay, T. Kincaid, collector, 1 specimen (USNM 32311). Coal Harbor, Unga Island, 1872, W. H. Dall, collector, 1 specimen (USNM 18652). Kodiak Island, W. J. Fisher, collec- tor, 1 specimen (USNM 18867). Aleutian Islands, R. C. McGregor, collector, 1 spec- imen (USNM 18904). Canoe & Cold Bays, 27-73 m, Sep & Oct 1940, 4 specimens (USNM 21318-21320; ZIASL, ident. by Hartman, 1948). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bering Sea: West Black Hill, 18 m, 27 Jul 1957, Weber, collector, 1 specimen (USNM 32310). Okhotsk Sea: 16 Aug 1947, P. V. Us- chakov, collector, 1 specimen (USNM 43574, from ZIASL, as L. helotypus by Us- chakov, 1950). Description.—Length of holotype 37 mm, width with setae 17 mm, 26 segments. Lengths of specimens examined 13-40 mm, widths 7-16 mm. Uschakov (1974) report- ed lengths to 65 mm, widths with setae to 20 mm. Body elongate-oval, flattened dor- soventrally. Segments multiringed (wrin- kled), with pigmented transverse bands. Large oval elytra leaving middorsum un- covered, without fringes of papillae. Elytra rather thick, soft, of gelatinoid texture, with pigmented network on median and poste- rior areas, colorless on anterior and lateral areas; surfaces with slightly raised, bulbous, white spots with small “‘papillae” (thick- ened cuticle?) in center and scattered mi- cropapillae (Fig. 1J; Moore 1905, pl. 36:fig. 32; Uschakov 1974, figs. C, D; 1982, pl. 33,3). Dorsal cirri on non-elytrigerous par- apodia with cylindrical cirrophores, bulbous basally, on posterodorsal sides of notopo- dia, with styles long, cylindrical, with fila- mentous tips, and extending beyond neu- rosetae; dorsal tubercles forming slightly inflated areas (Fig. 1G; Moore 1905, pl. 36: fig. 35; Uschakov 1974, fig. B; 1982, pl. 33, 2») Bilobed prostomium with stout cerato- phore of median antenna in anterior notch, style rather short, with filamentous tip; lat- eral antennae with ceratophores formed of anterior extensions of prostomium, equal in length and lateral to ceratophore of median antenna, with styles slightly shorter; palps stout, tapering, slightly longer than anten- nae; anterior pair of eyes in region of great- est width of prostomium, larger than pos- terolateral pair; tentaculophores with few (O-7) slender setae on inner sides; dorsal and ventral tentacular cirri similar to me- dian antenna (Fig. 1A; Uschakov 1974, fig. Ne NSIS, foils 32, 1). VOLUME 109, NUMBER 1 145 Fig. 1. Hermilepidonotus robustus, A, D-J, holotype of Lepidonotus robustus (USNM 5523); B, C, pharynx from specimen from Aleutian Is. (USNM 18904): A, Dorsal view of anterior end, posterior pair of eyes partially hidden by segment II; style of left dorsal tentacular cirrus missing; setae lacking on right tentaculophore; B, Muscular pharynx removed from body and cut open midventrally and flattened, showing left and right ventral jaws on outside and dorsal pair of jaws in middle, and distal pharyngeal papillae; C, Pharyngeal branched papillae; D, Right elytrigerous parapodium from segment II, anterior view, acicula dotted; E, neuroseta from same; F Right middle elytrigerous parapodium, anterior view, acicula dotted; G, Right middle cirrigerus para- podium, posterior view; H, Short and long notosetae from same; I, Short lower and longer upper neurosetae from same; J, Right middle elytron, with detail of micropapillae and pigmented network. Scales = 1.0 mm for A, B; 0.2 mm for C:; 1.0 mm for D, E G; 0.1 mm for E, H, I; 1.0 mm for J. 146 Segment II without nuchal fold, with large bulbous elytrophores, biramous para- podia, and long ventral buccal cirri, lateral to ventral mouth, similar to tentacular cirri (Fig. 1D; Uschakov 1974, fig. A; 1982, pl. 33, 1). Notosetae similar to following seg- ments; neurosetae more slender than fol- lowing, with long spinous regions and long tapered bare tips (Fig. 1E). Eversible mus- cular pharynx with 2 pairs of large hooked jaws and numerous border papillae enclos- ing numerous branched, threadlike papillae, forming thick brush (Fig. 1B, C; Uschakov 1974, figs. EK G, H; 1982, pl. 33, 6-8). Biramous parapodia with small subconi- cal notopodia on anterodorsal bases of larg- er neuropodia; smaller supraacicular part of neuropodia truncate and larger subacicular part diagonally truncate (Fig. 1K G; Moore 1905, pl. 36:fig. 35; Uschakov 1975, fig. B; 1982, pl. 33, 2). Notosetae forming close tuft, slender, with close transverse rows of fine teeth, some shorter, with blunt tips, and some longer, tapering to slender tips and ex- tending beyond tips of neuropodia (Fig. 1F H; Moore, 1905 pl. 36:figs. 34, 35). Neu- rosetae stout, dark golden color, forming dense bundles arranged in horizontal rows; 2 supraacicular and 7 subacicular (Moore 1905), with long spinous regions (13-20 rows), Supraacicular longer than subacicular ones, all with long, slightly hooked,