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BRIAN ROBBINS ASSOCIATE EDITORS Classical Languages Invertebrates GEORGE C. STEYSKAL THOMAS E. BOWMAN STEPHEN D. CAIRNS Plants Vertebrates Davip B. LELLINGER RICHARD P. VARI Insects RosBertT D. GORDON All correspondence should be addressed to the Biological Society of Washington, Smithsonian Institution Washington, D.C. 20560 ALLEN PRegss INC. LAWRENCE, KANSAS 66044 OFFICERS AND COUNCIL of the BIOLOGICAL SOCIETY OF WASHINGTON FOR 1988-1989 OFFICERS President KRISTIAN FAUCHALD President-Elect LESLIE W. KNAPP Secretary G. DAVID JOHNSON Treasurer DON E. WILSON COUNCIL Elected Members GARY R. GRAVES MEREDITH L. JONES W. RONALD HEYER RAYMOND B. MANNING W. DUANE HOPE WAYNE N. MATHIS TABLE OF CONTENTS Volume 101 Adkison, Daniel L. Pseudione parviramus and Aporobopyrus collardi, two new species of Bopyridae (isopodasEpicaridea) sromiytives Gullit ie Vc xc 0 eee eerie Alvarifio, Angeles. Pandea cybeles, a new medusa from the Sargasso Sea (Coelenterata: Antho- TMEGUSAS Pare dae) ae SA PR ae re Nea ae eee eee Baba, Keiji and David K. Camp. Two species of galatheid crustaceans (Decapoda: Anomura) new to Florida, Munida spinifrons Henderson, and Munidopsis kucki, mew SPeCves .....--cc-eceeeeeeeeenee Barnard, J. L. and James Darwin Thomas. Vadosiapus copacabanus, a new genus and species of Exoedicerotidae from Brazil (Crustacea, Armphipoda) .iie..e.--eeeecccseceeeeeeeececceeceeeeeeeececeeeceeeeeeeennnnneeeennenneennnney Barnard, J. L. and James Darwin Thomas. Ipanemidae, new family, Ipanema talpa, new genus and species, from the surf zone of Brazil (Crustacea: Amphipoda: Haustorioidea) -...... Bayer, Frederick M. Mirostenella articulata, a remarkable new genus and species of primnoid octocoral (Gorgonacea) with umcalcified axial MODES nine eeeeeeeeeeeeeeeeeeeeeeeeeeeeneeeeeeeeuneeeeeeenuneseneennnneeeneeennneee Bayer, Frederick M. and Jeffrey Stefani. A new species of Chrysogorgia (Octocorallia: Gorgonacea) from New Caledonia, with descritions of some other species from the western Pacific Birdsong, Ray S. Robinsichthys arrowsmithensis, a new genus and species of deep-dwelling gobiid fishctrom' the:western Caribbean 22. ee ee ee eee Bishop, Gale A. A new crab, Seosus wadei, from the Late Cretaceous Coon Creek Formation, TO mTOMG So uty VAT SSISS Up PO ss a Bae a ee Bishop, Gale A. New fossil crabs, Plagiophthalmus izetti, Latheticocarcinus shapiroi, and Sagit- tiformosus carabus (Crustacea, Decapoda), from the Western Interior Cretaceous, U.S.A. ........... Bleeker, J. and van der Spoel. Diacria picola and Diacria maculata, two new pteropod mollusc species from the Atlantic amd Pacific OC@ATS ce ceeccteecccecceeeeseecceeeeeceseetemneneceeceeneeenunneseceeeeeetununeneeees Bowman, Thomas E. Nitokra sphaeromata, a new harpacticoid copepod crustacean associated with the wood-boring isopod, Sphaeroma peruvianum, in Costa Rica oe ccccececeeeeeeeeeeeeeeeeeeeeeeeeeeee Bowman, Thomas E. and Thomas M. Iliffe. Tulumella unidens, a new genus and species of thermosbaenacean crustacean from the Yucatan Peninsula, M@X1CO oo... ccececcscsssseeeeeeeeeeeceeseeeeseeeeeeeee Bruce, A. J. Synalpheus dorae, a new commensal alpheid shrimp from the Australian Northwest Bruce, Niel L. Aega /eptonica, a new species of aegid crustacean from the tropical western Atlantic, with notes on Rocinela oculata Harger and Rocinela kapala, new Species eee eeeeeee eee Bruce, Niel L. Hadromastax merga, a new genus and species of marine isopod crustacean (Lim- noriidae) from southeastern Australia, with discussion on the status of the families Keuphyliidae eI STE Vit S11 AAG as ks A Sa a A CEE ORR OE De IO ic eee eC ee Bruce, Niel L. and Elizabeth B. Harrison-Nelson. New records of fish parasitic marine isopod crustaceans (Cymothoidae, subfamily Anilocrinae) from the Indo-west Pacific ...0...000.cc- cece Buden, Donald W. Geographic variation and probable sources of the northern mockingbird in thesBalvarmiavsl's] aris inset es Se i a ete ee ee Bullock, Robert C. The systematics of the southern hemisphere chiton genera Radsia Gray, 1847, and Sypharochiton Thiele, 1893 (Mollusca: Polyplacophora: Chitomidae) .......-cccecccceeeseeeeeeeeeeeeee Bullock, Robert C. Notes on some Rhyssoplax from the Pacific Ocean (Mollusca: Polyplacophora: @ ini tomid ae) ps sere ce Na ia: ae SSA Oe ne a ee cel aie ae Cairns, Stephen D. Cryptotrochus, new genus and two new species of deep-water corals (Scler- actinia:jurbinoliumae) iat see a Ce ore tall a ac Calder, Dale R. Turritopsoides brehmeri, a new genus and species of athecate hydroid from Belize (Eiydrozoasi@lavidac) |. 2 = ae 8 uc ee ee Camp, David K. and Richard W. Heard. Booralana tricarinata, a new species of isopod from the WestermyAtlantic/Oceani (Crustacea: Isopoda:|Cirolanidac) = Campos, Martha R. and Gilberto Rodriguez. Notes on the freshwater crabs of the genus Moritschus Pretzmann, 1965 (Crustacea: Decapoda: Pseudothelphusidae) with description of M. narinnen- Srswimom-southerme Colom biaies ye. a a 8 a0 eke Ne ld eters seg Ae Castro, Ricardo M. C. Semaprochilodus varii, a new species of prochilodontid fish (Ostariophysi: Characiformes) from the Marowijne River, Surinam 576-584 102-108 414-422 366-374 614-621 251-256 257-279 438-443 72-78 375-381 60-66 171-175 221-226 843-852 95-101 346-353 585-602 475-486 280-299 682-692 709-716 229-233 603-613 640-643 503-508 Clark, Janice and J. L. Barnard. Tonocote introflexidus, a new species of marine amphipod from Argentina (Crustacea: Gammaridea: Amphipoda) cece eeeenneceenennnneennennnnneennune Clark, Paul F. and Bella S. Galil. Redescriptions of Tetralia cavimana Heller, 1861 and Trapezia cymodoce (Herbst, 1799) first stage zoeas with implications for classification within the super- family Xanthoidea (Crustacea: Brachyura) 2... ..cccceccecceeeceeeceveeeeeeteteneeeeeeneeeeeeeeeeeseeeneenneeeesenentusunuunnnnnneneeeeeeeeeeeeeee Coates, Kathryn A. and Robert J. Diaz. Description of Guaranidrilus oregonensis (Oligochaeta: Enchytraeidae) from North America, with additional comments on the genus oe Cocroft, Reginald B. and W. Ronald Heyer. Notes on the frog genus Thoropa (Amphibia: Lep- todactylidae) with a description of a new species (Thoropa SAXatilis) 0... cece eens Cressey, Roger and Coiln Simpfendorfer. Pseudopandarus australis, a new species of pandarid CopepodstromyAustraliamyst ark) 2255265 2 8 ee oe ee eee Cutler, Edward B. and Norma J. Cutler. A revision of the genus Themiste (Sipuncula) ...................... Deets, Gregory B. and Ju-shey Ho. Phylogenetic analysis of the Eudactylinidae (Crustacea: Co- pepoda: Siphonostomatoida), with descriptions of two NEW QeEMeTa cece eceeeececeeeeeeeeeeeeeeeeeeeeetteene Eakin, Richard R. A new species of Pogonophryne (Pices: Artedidraconidae) from the South Shetland bislandssAmtarcticaass Sessa seins lan SO ee ew as PS es the en Ernst, Carl H. Redescriptions of two Chinese Cuora (Reptilia: Testudines: Emydidae) ..................... Ernst, Carl H. Cuora mccordi, a new Chinese box turtle from Guangxi Province 0.0.0 Erséus, Christer. Deep-sea Tubificidae (Oligochaeta) from the Gulf of M@X1CO oi. .c.ccccccccceseeeeeeeeeeeee Erséus, Christer and Michael R. Milligan. Limnodriloides faxatus and Doliodrilus puertoricensis, new Limnodriloidinae (Oligochaeta: Tubificidae) from Puerto Rico eee Erséus, Christer. Taxonomic revision of the Phallodrilus rectisetosus complex (Oligochaeta: Tubi- FY 1 ELS) eee NE ML AS Ory snes Paine Oe crea Bo oy Pall a Ee ul ha te Ferraris, Carl J., Jr. Relationships of the neotropical catfish genus Neumuroglanis, with a descrip- tion of a new species (Osteichthys: Siluriformes: Pimelodidae) 2c ccececeeeeeeeseeeeseeneeeeeeeeeeeeeeeee Felminger, Abraham. Parastephos esterlyi, a new species of copepod (Stephidae: Calanoida: Crus- tacea) piromuSany Dies opBaya Californias eee ee Gable, Michael F., Eric A. Lazo-Wasem, and Adam J. Baldinger. A description of the pigmented and non-stygobiontic females of Podobothrus bermudensis Barnard & Clark, 1985 (Crustacea: Ary pba ol tse a Ate cl eh ©) i Pa I a ie BN Sn oe Sen I Oe Gali, Frank, Albert Schwartz, and Alejandro Suarez. A new subspecies of Leiocephalus personatus (Sauriaslouanidae) from blartit 2 eee os Oe weet ioe i ee ee Ghosh, H. C. and Raymond B. Manning. Types of stomatopod crustaceans in the Zoological Surpyeyya@ team chi ieee sacs casera es eh ere alee es leet heel ye a dle aoe Goodman, Steven M. Patterns of geographic variation in the Arabian Warbler Sylvia leucomelaena CASES ERS willie) petite Sik cde es Sede nS. RMN Maho villas eel Ba Send nalts ot a ark Od, Hershler, Robert and Leslie Hubricht. Notes on Antroselates Hubricht, 1963, and Antrobia Hu- bricht, 1971 (Gastropoda: Hyrobiicae) oo. c ccc ccccccceeeeceseeeestnnneennnnneeneeeeeceeeseseeeeseeeeneeeneeeeesnttsnstevennnnnnnneeneeseeeeeese Hyer, W. Ronald. A notable collection of Cycloramphus (Amphibia: Leptodactylidae) from Bahia, Brazil, with a description of a new species (Cyclorammphus miguel) i -.....-ccececeeeeeeeeeeeeee eee eeeeeeeeen een Hobbs, Horton H., Jr., and William L. Pflieger. Cambarus (Erebicambarus) maculatus, a new crayfish (Decapoda: Cambaridae) from the Meramec River Basin of Missouri .....- Hobbs, Horton H., Jr. and Henry W. Robison. The crayfish subgenus Girardiella (Decapoda: Cambaridae) in Arkansas, with the descriptions of two new species and a key to the members of the gracilis group in the genus PrOCQMDArus -...:c.cccccsccccccveveveeesseevvcveeeseesesesteneseeeeeesesesssneseseeseesesesnnneeseveeeeesensnnneeeeee Hogans, W. E. Pennella makaira, new species (Copepoda: Pennellidae) from the Atlantic Blue Marlin, Makaira nigricans in the Caribbean Sea ooiii..........ccscssccccccscsssssessecsssessessesssnnnsesseeseeessesnnsseseseeeeestsnneseseeseeesseee Holt, Perry C. The correct name of Ceratodrilus orphiorhysis Holt, 1960 (Annelida: Branchiob- (LEN Ninh il) fe Resear mata ies lena eared YP theedaa Lua te ne See Any Deal Nie Holt, Perry C. Four new species of Cambarincolids (Clitellata: Branchiodellida) from the south- eastern United States with a redescription of Oedipodrilus macbaini (Holt, 1955) Hope, W. Duane. A review of the marine nematode genera Platycoma and Proplatycoma, with a description of Proplatycoma fleurdelis (Enoplida: Leptosomatidae) ee eeceeceeeeeeeeeeeeeeeeeeeeeeeee Hope, W. Duane. Syringonomus dactylatus, a new species of bathyal marine nematode (Enoplida: Leptosomatidae) and a supplementary description of Syringonomus typicus Hope and Murphy, 1S GS ete eee ecient siete betes Peels, Pea nea Perk Palak: 2 AUN AEE oo oe NR Leek baile! 8 pelt Humes, Arthur G. Bythocheres prominulus, a new genus and species (Copepoda: Siphonosto- matoida) from deep-water cold seeps at the West Florida escarpment i eeeeeeeee eee 354-365 853-860 773-783 209-220 340-345 741-766 317-339 434-437 155-161 466-470 67-71 11-14 784-793 509-516 309-313 146-150 1-3 653-661 898-911 730-740 151-154 644-652 391-413 15-19 308 794-808 693-706 717-729 568-575 Humes, Arthur G. Hyalopontius boxshalli, new species (Copepoda: Siphonostomatoida), from a deep-sea hydrothermal vent at the Galapagos Rift Se —e————————— James, Samuel W. Diplocardia hulberti and D. rugosa, new earthworms (Annelida: Oligochaeta: IMegascolecidae))tromy Kan Sas) === ssn Jones, Meredith L. and Stephen L. Gardiner. Evidence for a transient digestive tract in Vesti- TOV TA CULT AA sa A EE A 2 EAE IRL EEE ae Keppner, Edwin J. Thoonchus longisetosus and Oxyonchus striatus, new species of free-living marine nematodes (Nematoda: Enoplida) from northwest Florida, U.S.A. Kornicker, Louis S. Mydocopine ostracoda of the Alaskan continental shelf Kropp, Roy K. The status of Cryptochirus hongkongensis Shen, 1936 (Brachyura: Cryptochi- TLC.) see a I ea a tS a ee ae eV ee ree eee Kropp, Roy K. The status of Cryptochirus coralliodytes Heller and Lithoscaptus paradoxus Milne Edwards (Brachyura?€ryptochiridae) 20 ee eee Kritsky, D. C., W. A. Boeger, and V. E. Thatcher. Neotropical Monogenea. II. Rhinoxenus, new genus (Dactylogyridae: Ancyrocephalinae) with descriptions of three new species from the nasal Cavities of Amazonian Characoidcaye= en Lemaitre, Rafael and Robert H. Gore. Redescription, ecological observations, and distribution of the caridean shrimp Plesionika escatilis (Stimpson, 1860) (Decapoda: Pandalidae) _... Manning, Raymond B. Notes on albuneid crabs. (Crustacea: Decapoda: Albuneidae) from the Centrallicast:Coasty@ LK lo ric ais Ae ee soe tal sh ENS Mes ee Be OR cea ee ala es ee Manning, Raymond B. The status of Callianassa hartmeyeri Schmitt, 1935, with the description of Corallianassa xutha from the west coast of America (Crustacea, Decapoda, Thalassinidea) Mathis, Wayne N. and Jin Zuyin. A review of the Asian species of the genus Lamproscatella Hendeli(DipteravE phy dri ac) see eS ha ea ee ee ee eee eee Miura, Tomoyuki. Poecilochaetus koshikiensis, a new polychaete species from Shimo-Koshiki MS Leary Gh 2) ea py eA hs TE SN tad Ea OEE ue Ws Pin ah gE IE er Monterrosa, Oscar E. Heterocarpus cutressi, new species, and Plesionika macropoda Chace, 1939: two caridean shrimps of the family Pandalidae (Crustacea: Decapoda) from Puerto Rico and ste WSS 25 Vata eS Learn Se a Sa ci Sah Morgan, Gary S., Omar J. Linares, and Clayton E. Ray. New species of fossil vampire bats (Mammalia: Chiroptera: Desmodontidae) from Florida and Venezuela o.........cccccceccseccceeeeeceeeeeeeeeeeeees Nakamura, K. and C. Allan Child. Pycnogonida of the western Pacific islands IV. On some species drommbthre aR yakeyis] earn cl S32 es aie kaon Ae a eae yee ise dane nce Nakamura, Koichiro and C. Allan Child. Pycnogonida of the western Pacific Islands V. A collection Loni daKey ICON eA WO) AY CONV) vim oy ony SeeHOOVO LY cee ee Ng, Peter K. L. Allopotamon, a new genus for the freshwater crab Potamon (Potamonautes) tambelanensis Rathbun, 1905 (Crustacea: Decapoda: Potamidae) from the Tambelan Islands Ohwada, Takashi. Streptospinigera alternocirrus, a new species of Polychaeta (Syllidae: Eusyl- INGSAYS) \paeenetsmeah eo lon Mra eU rare ant ANGER DAS Ne Wye MOLOM MP tora Wao tO ti Oe) Ue nen SRLEt Rater oe cat mae Na aca «ee Peters, Esther C., Stephen D. Cairns, Michael E. Q. Pilson, John W. Wells, Walter C. Jaap, Judith C. Lang, Carol E. (Cummings) Vasleski, and Lauren St. Pierre Gollahon. Nomenclature and biology of Astrangia poculata (=A. danae, =A. astreiformis) (Cnidaria: Anthozoa) ............. Pettibone, Marian H. New species and new records of scaled polychaetes (Polychaeta: Polynoidae) from hydrothermal vents of the northwest Pacific Explorer and Juan de Fuca ridges .................. Reid, Janet W. Copepoda (Crustacea) from a seasonally flooded marsh in Rock Creek Stream Mallews Barkan lar yilarid 5 seo 0 200 aes Peay Pie ek ph Dla i canals nadie ena aaa lt Reid, Janet W. Records of cyclopoid and harpacticoid copepods (Crustacea) from a spring in AGH IVI0T:4K0) 01) DM Castanea Muna ya Bee MAMIE Vege onay, SMe nary woe Sir ere oe ee The ee ah Robinson, Harold. Studies in the Lepidaploa complex (Vernonieae: Asteraceae) IV. The new genus, TECSSINGIGNEN US 5 BOE BSUS a pote Areas Obese g 0 al, eet ee a ga A ee Sai ane tea ig Robinson, Harold. Studies in the Lepidaploa complex (Vernonieae: Asteraceae) V. The new genus Gary Solace aso Soe ANA a 8 WN RL OR ed al a le AOL cco Ne EN oho 32 Uh ras al dey aes A A Rr ec Rozbaczylo, N. and J. C. Castilla. A new species of polychaete, Scolelepis anakenae (Polychaeta: Spionidae) from Easter Island, South Pacific Ocean, with ecological comments Sohn, I. G. Darwinulocopina (Crustacea: Podocopa), a new suborder proposed for nonmarine Paleozoic to Holocene Ostracoda 825-831 300-307 423-433 183-191 549-567 866-871 872-882 87-94 382-390 626-632 883-889 540-548 671-675 633-639 912-928 662-670 809-816 861-865 83-86 234-250 192-208 31-38 314-316 929-951 952-958 959-965 767-772 817-824 Sig, Jiirgen and Richard W. Heard. Tanaidacea (Crustacea: Peracarida) of the Gulf of Mexico. V. The family Pseudotanaidae from less than 200 meters, with the description of Pseudotanais mexikolpos, n. sp., and a key to the known genera and species of the world eee Springer, Victor G. Rotuma lewisi, new genus and species of fish from the southwest Pacific (Gobroidersex ents tira cl ae) ee ee aa ee eee ee aU alate ps Ae eee eee ea Starnes, Wayne C. and Robert E. Jenkins. A new cyprinid fish of the genus Phoxinus (Pisces: Cypriniformes) from the Tennessee River drainage with comments on relationships and bio- OYTO NT et ee eee ee ee Steadman, David W. A new species of Porphyrio (Aves: Rallidae) from archeological sites in the INVA UU SA SoS Mca 11 Cl S paseo ee 2 NO ee Steadman, David W., Susan E. Schubel, and Dominique Pahlavan. A new subspecies and new records of Papasula abbotti (Aves: Sulidae) from archeological sites in the tropical Pacific ....... Thomas, James Darwin and J. L. Barnard. Elasmopus balkomanus, a new species from the Florida Key se(Grustace ase rai poli oda) ea a EI ees el ala wot SER Tourtellotte, Gary and Henry Kritzler. Scionella papillosa, a new species of polychaete (Polychaeta: Terebellidae) from the southwest Florida continental shelf ccc cccccccccsccccecsssessessstetstncccsscsseceseeeeeeeeesee Turner, Richard L. and Cathleen M. Norlund. Labral morphology in heart urchins of the genus Brissopsis (Echinodermata: Spatangoida), with an illustrated revised key to western Atlantic SPIE CIC S paar een ee aA a IM IRE ec Cs A BO Tae lle Si Nl ose Van Syoc, Robert J. Description of Membranobalanus robinae, a new species of sponge barnacle (Cirripedia, Archeobalanidae) from Baja California, with a key to the genus . ee Villalba, Antonio and José M. Viétez. Polychaetous annelids from the intertidal rocky substratum of a polluted area of the Ria de Pontevedra (Galicia, Spain). 2. Taxonomic aspects with de- SCIP EL ONO fap B1L SURAT LCL TALT CCl aN eg S [Meet a Ne Warén, Anders. Neopilina goesi, a new Caribbean monoplacophoran mollusk dredged in 1869 ... Wasmer, Robert A. The parasitic isopod Holophryxus acanthephyrae Stephensen (Epicaridea: Dajidae) from the Subantarctic South Pacific, with notes on its synonym and host .................... Weems, Robert E. Paleocene turtles from the Aquia and Brightseat formations, with a discussion of their bearing on sea turtle evolution and phylogeny cece eeeeeeeeeeeeennneeeenenetennnnn Weitzman, Stanley H. and Richard P. Vari. Miniaturization in South American freshwater fishes; ANGONSTAVIC WHAT GNIS CUS STC Meee te ea a eee A GIR Ge ee Wicksten, Mary K. and Matilde Mendez G. New records for Ogyrides alphaerostris and a new species, Ogyrides tarazonai (Crustacea: Ogyrididae), from the eastern Pacific Ocean _. Wilson, Robin S. Synonymy of the genus Nectoneanthes Imajima, 1972, with Neanthes Kinberg, NS GO7(Bolychae tacwiNeret chica e) yee ee Berets Ma escheslscgetacendene eee espe tacpsct nce! Wongratana, Thosaporn. Leiognathus pan, a new ponyfish (Pisces: Leiognathidae) from Thailand, with comments On Thai letogmath ids cece cceecceeeeeeeeeeseesensennnnnnnnnneneceseneeeeeeeeeeneeeeeneeeseeseetnuusnnnnnnneneneeeeeeeeeeeee 39-59 530-539 517-529 162-170 487-495 838-842 79-82 890-897 832-837 176-182 676-68 | 20-30 109-145 444-465 622-625 4-10 496-502 INDEX TO NEW TAXA VOLUME 101 (New taxa are indicated in italics, new combinations designated n.c.) COLENTERATA Hydroza Pandea Cybele «sito. 5 00 ace a uh ioleee Peas dd cee 48 SE Bia sa ae 103 MUTKILODS O1d CS ea ee eee ee 229 brehmeri 229 GhirySOgOrgiavadiiiel eo e se Peeyla L e A eEl oe Db aes ite eee tea ere ee 260 bracteata 263 calypso 265 CHU) SOLS Pisa eye 0M el GN By La fae a a UN los RSD A nea at lb SN ee 267 CLO CER DSO LENG SS ie AL Lh IN he IOS A AS ae UN de NAN eA OR 269 SCUTALD LT ETUS cs) ICEL Se ELM eB Sa Sel SL aah Na A RUE er LER A 270 CCHYDEOLFOCTUS set 5 AE Net AWN dere) Eg AR ON oe hn ehh ean Oe aL hae oI St es 709 carolinensis... 711 VETS ach Nh Nyt Vt MS A ae re le Mc eu me Se Pd oe a A 712 WATT O SCC TCU U Ge See ie I PI Be Nal nl oe AGM a gli sll sa V2 Ale ee a ee ADS 251 CTIA CUL TOS 0) hatiaed es an BIS atl NGS Oa ais aa NS Ue Al ea ce IONS cea RL ee ROO 252 PLATYHELMINTHES Trematoda 88 89 92 88 ORV OMCIUSTS TTL S as Oa ole CU BLSNINTES ee UD He ade ger te Oe ened ite RICO Oe eer ees dea tere ae 186 Proplatycoma fleurdelis ......... 695 Syringonomus dactylatus 722 MA OOMCHUS HON SISCLOS TES en WA ele alteh ice NOt ee pen) te 0 1 ames vedic MOvaeMENEE ar ole omnhen Rinne ee 183 MOLLUSCA Monoplacophora INGO pIlinascOes islet c 8 cee Ten eee sas Aen Nir: fly RE a cata IN atl seg ea LAC alae So kas Sed Ss 676 Gastropoda LACT AGIA CEL CE CaS cee hee UNI Nao ND uals Nespas WONG ENN nine ep 61 piccola 60 ANNELIDA Polychaeta Bepidonotopodium) piscesae = i ah ae IN eee a ee 193 ENOTES 2P4 121711610 | ree Re A PURO O NCTE Cee ee rae yy Sl ee 176 INeanthestlatipodaim:c.. a ue) leno. en gee eR Uk ear ene 9 maligna tae 2 a i een I 2 ST OI ee 9 Opisthotrochopodserssra701 li ffecie mese etn arama aa 203 ROECILOGH AC TU SHKOSILEKCLE TiS 1s wrmatbesats arene ester te veo s EN ee Sete el ll ores ol Hi erie EN Ne SClOn el ay 7110S cx nmeaeaae tees eaten es ere aie ae rN We odleca ey sede eet Oe pus reer nes al Sa es ee SCOlETE DiS 71 Ke 17 cc Peseta eee eee seer eal et re ere eA NO StreptospinigeraraltennOcinis ae ene ee eae ee ee eee BYE YG EU NBN LON AY SUA sks ee ae ed PE oc AO eR Diplocandiak)iee Dent tee ress scene eestor mre tebdites AO a ea Mees Br Bn eos me aoe et Pe a LAS rugosa Dol Od nilus#aUertOni Cer SiS en ee oh rue ne nee acon ieee SPs ES AV NE ee Guarani a rilUsyore co crisis se esse eer ates etre ee PE ede ee aN ee pe ee MEWTATVO A Tall OVAL SS wf Acca LES pees ten Pa oe one I a lo es Sh ena ok ae HAT OAT SKCO 11S 111 CLI1 Sees ese ea RRs ll eae Nr Soe cy ee se ee ene FILO LCS GUUS wees dl elie Elie SN AV ANE, adept Pre oA en NC Gs vn nl ale AL EECA Camibarincol anh ob ly eee eee MIMO Seay ERIN BH 2 oe ha ne elegy ee «Si aes PRN Ee ENTS OATUUSECAHTON ATT11LS RRR a MNES etl ans De sea Ned ella 2S OR gree hu IEP MeEdipoOdalUseareSO Sal His ese ep AE AON by RL AE ears ee ne een ede SA tO rill SH iV ESC Ce se NTN al APTA A NIN IL eT N ts VAAAN A Na ef he AOS ARTHROPODA Pycnogonida FAM ODLOGAG YIU SHEN CLS:S 14:5 eeseraee eer Maceo eel es Coeee Nt aereresvaltee een aMiee Mea aS Go LAL Ra Re a he ee Galli palleMme ge ytd CHS wees MA eee MON acta eae gr eee pee Dea a ae UR oS es tn 8 er ha lhe NCR AGC PILOT CNet est rN eli aS Ry OPIN eAR eS iy eed Vee RAIN 2 NR eter aa U6) 0X0) 21 1/1) ee tambelanense n.c....... Aporobopyrus collardi........... Bathnyleberi SHO Crit CErts tS ea ee ea leit lanl Ad ls ol fe ede el elt ear hoow.c nahn, SOA ye tenes OO rel ATV AYE CCU TUL Ca a ah A NAO SI sh es Ne cea te Al lpetae ana UENO DAE BV TROCILCH 2 Stesmn esta g Moder LLU GR aN A eat IE Seattle eet a ett ieee nae mee eh i dD UTI OF YUL IUL 11 S eae a RE) pe Se En ety hl SL a al ety i ys eR Cambarus (Erebicambarus) maculatus... (COLI OSS OT IULS posceed crore aa eta tI ta ah eat cineca ERR AS CLI VACHS LS comers te le ceca Le er oe ee na een cere etter ate EN UE EY IE, Goralllianassaecy/t/1apestaene eins a ull nie estes. RR ole alee ON tay Se al sr ed DAUR VTA TE OE OTN A rete his ese is art nn reo es EIR SE NU gg Na wc al Mee kv Diacyclops palustris FS] US TUNG POSH! AO FY 11 1 AS eee ae aa oe DEI QHA ITO IN TESA Gosek eR er LSP ca OS eR UPR TO Se SOA oe IFAS COMO GAT POULS CULM CS S Ui ao a a ee 2 aed er Oe RRR Heterocladius LDV S'S CL CS as tn an ar hE eRe ct el yD see, NT MeLsy all @PPOMMENAS WO TS HCL Lip St a acta eens A Ms SUE BTR AU ETSI EAN TEV AUT AIDA ceed Peer Re en tu ae aur a ares A A WN a shapiroi Membranobalanus robinae INNO TIES CIV FLCUETUFLC 1S US memes recat eee a ee Pe cial i ee Munidopsis kucki.................. Nitokra sphaeromata OpyriGdes LAP AZONAL 2.2.2.2: 622 Parastephos esterlyi 309 Perineal 77a a ea er ee Nr ene 15 Plagiophthalmus izetti 375 Plesionika escatilis 1. Co ... cece 383 Procambarus (Girardiella) ferrugineus......... 391 TOUTS oO ee TN EN A eh eR La I et lor oN asd 398 PSEWATOME PIVEN GTTIUS ee a eae ce cee ee ee eee ee 576 Pseuid@ par ars: Quester Quis csc 020 eS ee oe UN ee a 340 Pseudotanais (Pseudotamais) ext Ol ..cceececccccccsseeessvcceccecseesssvsenececseeeesvesnneceeeeeeesensensececeeeeeeenumnnnececeenensunneeeeeeeene 48 ROGINE AV Kap ala Mea a EE RE To a NR RA IO an cl 100 IS STEELS ONIILOSULS oie Meh Lege Sete go VND TOLLE np 379 CON ADUS HE SEC ela) as in 2a tg a Alla Ae ea iat au Peel Near oe eae ea 379 Scleroconehiavcdip lance. ie ie see I i A UR ee a 560 SYP) ON YS ease Oe eA eat rat eR ee ee eee VT TYH(O17 (2 RAIN Seal co CAL Areas AA en NCL OR Lies near ae enPcaO DE ase ei analrenenrCrWcles Huei ey V2 Tonocote introflexidus 357 TOTAITT UI Toh cas a eI et es ay ett MEW shes Notas ain de Ste Wm ene UL ON 221 UNITAS ee esas re Pee Maia ee ea HS aod sega ae Netea eh GLI re foe, ced Me ae oo 221 VAAOSIGp Us Wee Be: culled ce A TANI OVNIER Depa ae ans hi | Pees BY Lt OWL Sie Be 2 A es Ve ee ee 369 COP ACAD GIUUS se BGR gE EN NE AP Me Nee aN EAL Ae le Le a au aes O RNa i et A 370 Insecta amiproscatellay(eamproscatella) | ely cl ieee ee eee eae rc eee 543 SETULC Gc te Reet eth i SUN A eel i hE I Mg tate el Os Una UO a ee 544 LED CLONES TS 2 it Se eto Bg US I AN aE ss ee Die nn ge atte tO I 547 CHORDATA Pices TES LO PM ACIS 7711 ee cee ar LUCIE Secchi 8 occ le a ol 496 IN@mATUTO SATIS 7 ts CEA LEA LTS ee Oe I asa yc rl a 510 Phoxinus tennesseensis 518 Pogonophryme Cewitty ccc cecceeeeeeeeeseecesvsneeccevcveeeeeeeeeeeeeeeeee 434 HERO ELT YU CASA an ee Sete Sele sates tna aa aul ORNL LAR ce eR Ra 531 TOMOES Ty sl Se eee SL 9 ee AI a eA A A LU elt Dad 1S eT 535 TRODU TESTE Hata) Siz ian eeN iti 2G OU ATE SSI SEPA ee I ei ge Ue Re la ard a 438 arrowsmithensis 439 Sema pnochalOauss yer yep eee ee Ve CNMI Te lal a er cee seen dade Ea NOTES i a a el 504 Cycloramphus migueli 151 BUG OO PAGS CONCALTLES scenes cca ark NT HUT fe UN Ua ULI hd telat caisen i ON and ales De A 210 Reptilia Gata ple ray) ffiesc ce ae Ce UE eI a Pa eh te ca ee ce a 125 QUOT ANTC CONT a ae! ee SEAS oe Nile UN ca rel ene SO NLS eee cl ee ear ae pee ree ee 466 Wollochelys:coatesisch ccskc so Wa an ee OUI ASS aie oe ee Oeil lic rece 132 Helocephalus PeErsOnatus, C/attoproSOp ore a tees Mia i iiaeaniee teller ni atlelect ssa lee enna 1 Osteopygis roundsi 120 RLANCLOCNEL YS) ire Bes 18 LN AES ENC CU ha eR ate DUP ade cd stays Wve aan 116 SLVOLED soc 3 PS stil a MN USI a Ny aL to Te aca Oo Orca ee A San 116 Aves Bapasula.abbotti costellog:. 2.25 oily i ya eg ge cle al de 490 BOnphyrio Paep ae nals ENS es SO INE are a an ee sae oe 163 Mammalia DESTINO GUSTOKCH ACO ADIOS meester NEI ae hat ar ae eR et Rg tn 913 CL CLC ULL Co Grater ae ee ren Ne OE ae mare dc ett lal OO a ieee Se SS 920 PLANTAE TTT AS sas I A Sa a ae te Dn ee 959 pseudascaricida 959 GQ) SOL eer a a I SD nr ee etre ee a ea ; 956 (for seven new name combinations see pp. 956-957) JESUP SG UOU AA IS SS ee eet oe ee U2 veo een 939 ISI NT AIHA ce a cee eS A A ne 945 (QlicaCen Tal is) eee es ees PE eats aN ea Nase eres eat OE A eo cas Ce a a Ia 949 (for 100 new name combinations see pp. 940-950) bs} fi ‘ La ne 1s awres VOLUME 101 NUMBER 1 29 APRIL 1988 ISSN 0006-324X THE BIOLOGICAL SOCIETY OF WASHINGTON 1986-1987 Officers President: Austin B. Williams Secretary: C. W. Hart, Jr. President-elect: Kristian Fauchald Treasurer: Don E. Wilson Elected Council Stephen D. Cairns Richard P. Vari Mason E. Hale Stanley H. Weitzman Robert P. Higgins Donald R. Whitehead Custodian of Publications: David L. Pawson PROCEEDINGS Editor: Brian Robbins Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Stephen D. Cairns Plants: David B. Lellinger Insects: Robert D. Gordon Vertebrates: Richard P. Vari Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $15.00 include subscription to the Proceedings of the Biological Society of Washington. Library subscriptions to the Proceedings are: $25.00 within the U.S.A., $30.00 elsewhere. The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued sporadically) are available. Correspondence dealing with membership and subscriptions should be sent to The Treasurer, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, National Museum of Natural History, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 1-3 A NEW SUBSPECIES OF LEIOCEPHALUS PERSONATUS (SAURIA: IGUANIDAE) FROM HAITI Frank Gali, Albert Schwartz, and Alejandro Suarez Abstract. — A new isolated subspecies of Leiocephalus personatus is described from near the southern coast of the Haitian Tiburon Peninsula. Comparisons are made with its most closely related geographical congener, L. p. personatus. Schwartz (1967) recorded one female Leiocephalus personatus Cope from the town of Aquin, Département du Sud, Haiti, which he questionably assigned to the subspecies L. p. personatus. Aquin lies on the southern coast of the Haitian Tiburon Peninsula, and the locality is separated from the nearest population of L. personatus by the inter- vening Massif de la Hotte. Much additional material, collected by Richard Thomas, from the Aquin area allows us to clarify the status of the population of Aquin L. per- sonatus, which is distinct from L. p. per- sonatus, and for which we propose the name. Leiocephalus personatus elattoprosopon, new subspecies Holotype. —USNM 197370, adult male, ca. 1 km inland, basal portion of the Morne Dubois “‘peninsula,” Dépt. de Sud, Haiti, one of series collected 20 Jul 1971 by native collectors; original number ASFS V26037. Paratypes.—(All from Haiti, Dept. du Sud): ASFS V26036, same data as holo- type; CM 83051-60, MCZ 156222-31, MNHNSD 221-30, USNM_ 197383-91, Aquin, 12-13 Jul 1971, native collectors; ASFS V25793, ca. 11 km SE Vieux Bourg d’Aquin, 14 Jul 1971, native collectors; ASFS V26030-31, beach area at base of Morne Dubois “‘peninsula,” 20 Jul 1971, R. Thomas; ASFS V2605 1-621, basal area at the base of the Morne Dubois “peninsula,” 21 Jul 1971, native collectors. Associated specimen. —ASFS V25728, Grosse Caye, south of Aquin, Dépt. du Sud, Haiti. Diagnosis.—A subspecies of Leiocepha- lus personatus characterized by pale gray middorsal region with pale buff lateral stripes; ventral color pale green with very pale green (nearly white) spotting; mask present but restricted and usually dorsal to ear opening; chin and anterior throat dark gray rather than black; sides red with pale gray-green centers to scales; median dorsal scales between occiput and vent 40-63 (x = 50.1 + 1.7 = twice standard error of mean), median rows of dorsal scales 19—40 (28.6 + 1.6). Distribution. —Département du Sud, Hai- ti, from the vicinity of Vieux Bourg d’Aquin northward to Aquin and onto the Morne Dubois ‘“‘peninsula’’; presumably also Grosse Caye off this southern coast. Description of holotype. — Adult male with following measurements and counts (all measurements in millimeters): SVL (snout— vent length) 80, tail length 105; dorsal crest scales occiput-axilla 16, dorsal crest scales on trunk 26; one-half midbody scales 22; subdigital fourth toe tricarinate scales 22/ 22: loreals 4; 4 prefrontal scales, row in- complete; 7/8 supraocular scales; supraor- bital semicircles incomplete; parietal scales in contact. Dorsal ground color pale gray, dorsolat- eral stripes pale buff in life but faded in preserved specimen; sides red with pale gray- green centers to scales, becoming greener on lower sides, with gray to pale stripes which 7) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON may be very indistinct; mask dark brown with dark gray in loreal region, mask re- stricted and ending dorsal to ear opening; top of head brown with gray supraocular scales; upper surface of hindlimbs dark green with pale green spotting, forelimbs pale gray- green with pale green spotting; chin and an- terior throat dark gray rather than black, followed by pale gray scales randomly smudging balance of throat; ventral ground color pale green (gray-green laterally) with very pale green (nearly white) spotting; tail brown above and light brown below. Variation.—The series of 55 specimens of L. p. elattoprosopon has the following scale counts: dorsal crest scales occiput-vent 40-63 (X = 49.8 + 1.7); dorsal crest scales occiput-axilla 16-27 (21.4); dorsal crest scales on trunk 19-40 (28.6 + 1.6); tricar- inate scales 15—26 (21.4); loreals 4-17 (5.7); supraoculars 6/6 (30 specimens), 6/5 (1), 5/5 (2), 7/6 (10), 7/7 (6), 7/8 (4), 6/8 (2); semicircles more often complete (95%), and parietals usually in contact (91%). The pre- frontal row varies between 3 and 5 scales (Mo = 3), the median head shields are 4—8 (Mo = 5), and the frontoparietal row is more often complete than not (49 of 55 speci- mens). The largest male has a SVL of 83 and is a topotype, and the largest females measure 60 SVL and are from Aquin. The series of specimens was described in life as follows. Males have a dorsal pale gray ground color; some adult males have pale buff dorsolateral stripes obvious, others lack this feature. All males have 2 to 4 dark brown nuchal bars, more obvious in some than in others. Sides have red scales with other pale scales gray-green centered, the scales be- coming greener on the lower sides, and lim- ited ventrolaterally by gray to pale stripes which may be very indistinct. Mask dark brown with dark gray in loreal region; mask present but restricted and usually dorsal to ear opening. In some specimens the mask extends further posteriorly than in others. The ventral ground color is pale green (gray- green laterally) with very pale green (nearly white) spotting. The throats are green and ventrals are 40-64 (x = 50.1 = 1.7), and median dorsal trunk scales are 19-40 (28.6 + 1.6). In L. p. personatus these counts are 47-61 (53.5 + 1.0) and 24—40 (32.4 + 1.0), respectively. Remarks. —Leiocephalus p. elattoproso- pon is similar to L. p. personatus in having a head scutellation formula of 3-5-4 (pre- frontals-medians-frontoparietals). Both subspecies have masks, but they differ in that the mask in L. p. elattoprosopon usually is restricted dorsal to the ear opening, unlike that of L. p. personatus in which the mask extends half way to the forelimb insertion, including the eyes and the loreal region, and as far anteriorly as the tip of the snout. The distribution of L. p. elattoprosopon as currently known is restricted to the Morne Dubois “peninsula,” and north to Aquin and Vieux Bourg d’Aquin, a distance of about 0.7 km. It is intriguing that Ameiva chrysolaema evulsa Schwartz (1973) and L. p. elattopro- sopon occur in much the same general area, although the former species occurs (as far as known) only on Grosse Caye, whereas L. p. elattoprosopon occurs on the adjacent mainland, as well as on Grosse Caye. Both lizards represent southern outlier popula- tions of species that are limited, on the Ti- buron Peninsula, to the northern coast (Leiocephalus; 25 km) or extend only a short distance along the coast west of Port-au- Prince (Ameiva; 85 km). Intervening areas, which have been intensively sampled, seem not to harbor populations of either lizard. It seems that the Aquin-Morne Dubois ‘““‘peninsula” region has a peculiar relictual fauna that was once more widespread, or continuous with relatives farther to the east and north. Etymology. — The name elattoprosopon is from the Greek elatton (less) and prosopon (face, mask), in reference to the reduced dark mask in this subspecies in contrast to that in L. p. personatus. The name is used as an appositional noun. VOLUME 101, NUMBER 1 Specimens examined. —(Other than L. p. elattoprosopon; all are L. p. personatus from Haiti): Dépt. du Sud, Jeremie (MCZ 3615, two syntypes); Les Roseaux (MCZ 74626- 39); 16 km E Baradéres (USNM 80773-66); Grand Boucan (USNM 80774-87); Pres- qu’ile de Baradéres, vicinity of Grand Bou- can (ASFS V26282-338); Petit Trou de Nippes (USNM 80788-98); 1.7 km SE Anse a Veau (ASFS V42829); 2.7 km S Anse a Veau, 46 m (ASFS V43187-88); Ti Anse, ca. 9.5 km NW Miragoane (ASFS V26212- 17); 9.7 km SW Miragoane (MCZ 25433); Dépt. de l’?Ouest: Léogane (MCZ 13834- 37); 3.2 km SW, less than 1.6 km E Leéogane, +122 m(ASFS V8317-—22); Bayeux (USNM 75906); L’>Acul (USNM 72607-12); just W Grand-Goave (ASFS V36552-68, ASFS V45471, ASFS V45513, ASFS V45528). Acknowledgments We wish to thank C. Butterfield, D. A. Daniels, J. W. Norton, W. W. Sommer, R. Thomas, and T. M. Thurmond for assis- tance in the field. Specimens in the Museum of Comparative Zoology (MCZ) and the Na- tional Museum of Natural History (USNM) were borrowed by Schwartz for his 1967 paper, and data from them have been used 3 in the meristic sections of the present paper. For these previous loans we are grateful to Ernest E. Williams, the late Doris M. Coch- ran, and the late James A. Peters. The ho- lotype and paratypes of L. p. elattoprosopon have been deposited in the cited collections as well as in the Carnegie Museum of Nat- ural History (CM), the Museo Nacional de Historia Natural, Santo Domingo (MNHNSD), and the Albert Schwartz Field Series (ASFS). Specimens in the ASFS were in part collected under NSF grant B-023603. Literature Cited Schwartz, A. 1967. The Leiocephalus (Lacertilia, Iguanidae) of Hispaniola. II. The Leiocephalus personatus complex.— Tulane Studies in Zool- ogy 14(1):1-53. . 1973. A new subspecies of Ameiva chryso- laema (Sauria, Teiidae) from Haiti.— Herpeto- logica 29(2):101-105. (FG) 156 South Melrose Drive, Miami Springs, Florida 33166; (AS) 10000 South West 84th Street, Miami, Florida 33173; (AS) 7645 West 16th Court, Hialeah, Flor- ida 33104. Send reprint requests to A. Schwartz. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 4-10 SYNONYMY OF THE GENUS NECTONEANTHES IMAJIMA, 1972, WITH NEANTHES KINBERG, 1866 (POLYCHAETA: NEREIDIDAE) Robin S. Wilson Abstract. —Nectoneanthes oxypoda (Marenzeller, 1879) and N. alatopalpis (Wesenberg-Lund, 1949) are synonymized with Neanthes succinea Leuckart, 1847. Nectoneanthes Imajima, 1972, with type species Nereis (Alitta) oxypoda Marenzeller, thus becomes a junior synonym of Neanthes Kinberg, 1866. An undescribed species of Neanthes is recognized. Nectoneanthes vimai (Izuka, 1912) is once more referred to Neanthes, and Nectoneanthes latipoda Paik, 1973 and N. multignatha Wu, Sun & Yang, 1981 are referred to Neanthes for the first time. The genus Nectoneanthes was erected by Imajima (1972) for two species of Neanthes Kinberg, 1866, which lack falcigers in all neuropodia: N. oxypoda (Marenzeller, 1879) (type species by original designation) and N. ijimai (Izuka, 1912). Three additional species have subsequently been placed in the genus: Nectoneanthes latipoda Paik, 1973; N. multignatha Wu, Sun & Yang, 1981; and N. alatopalpis (Wesenberg-Lund, 1949) which was transferred from Neanthes to Nectoneanthes by Wu, Sun & Yang, 1981. In a previous paper (Wilson 1984) I ex- amined all Australian material of N. oxy- poda and synonymized these records with Neanthes succinea (Leuckart, 1847). I have now been able to examine type material of Nereis (Alitta) oxypoda Marenzeller and Nereis alatopalpis Wesenberg-Lund, which are here shown to be junior synonyms of Neanthes succinea. Nectoneanthes Imaji- ma, 1972 thus falls into synonymy with Neanthes Kinberg, 1866. Loss and breakage of atokous setae occurs prior to metamorphosis in epitokous nere- idids (Clark 1961), hence absence or ap- parent absence of neuropodial falcigers should not be accepted as a valid taxonomic character unless undamaged specimens en- compassing a wide size range and thus age are available for study. The specimens ex- amined in this study and all literature rec- ords represent large specimens with ex- panded parapodial lobes which are probably approaching epitoky. The three remaining species that have been referred to the genus Nectoneanthes and an undescribed form previously con- fused with N. oxypoda appear to be valid species and are here referred to the genus Neanthes. This determination can only be confirmed when atokous material of these species becomes available for study; how- ever, it has not been possible to obtain ma- terial of these three oriental species. These taxa are discussed below. All species discussed here are adequately figured in the original descriptions or in sub- sequent publications; these figures are re- ferred to in the relevant descriptions below and no new figures are presented here. The specimens examined in this paper are deposited in the following institutions: Brit- ish Museum (Natural History), London (BMNH); National Science Museum, To- kyo (NSMT); Zoological Museum, Copen- hagan (ZMK); Naturhistorisches Museum, Vienna (ZMV) and National Museum of Natural History, Smithsonian Institution, Washington (USNM). VOLUME 101, NUMBER 1 Neanthes Kinberg, 1866 Neanthes Kinberg, 1866. Type species: N. vaalii Kinberg, 1866 (designated by Hart- man, 1959:249).—Hartman, 1940:219 (generic definition emended); 1959:249 (for additional synonymies). Nectoneanthes Imajima, 1972. Type species: Nereis (Alitta) oxypoda Marenzeller, 1879 (by original designation). New synonymy. Definition. —Eversible pharynx with con- ical paragnaths on both rings. Four pairs of tentacular cirri. Parapodia biramous. No- tosetae homogomph spinigers; neurosetae including homogomph and heterogomph spinigers (emended after Fauchald 1977:89). Remarks. —As a consequence of the new synonymies made in this paper, several species previously placed in Nectoneanthes Imajima have been referred to Neanthes Kinberg. These species are not presently known to possess neuropodial falcigers and thus violate the widely used definition of Neanthes provided by Fauchald (1977). Wilson (1984:221) however noted that three species of Neanthes belonging to Group I in Fauchald’s (1972) subdivision of the ge- nus are already known to lack neuropodial falcigers: N. agnesiae (Augener, 1918); N. chingrighattensis (Fauvel, 1932) and N. mexicana Fauchald, 1972. In view of the incomplete material presently available of these species, and the loss and replacement of setae which is known to occur in ageing nereidids, the presence of neuropodial fal- cigers should no longer be included in the generic definition of Neanthes and the ge- neric definition of Fauchald (above) is emended accordingly. Additional comments on the synonymy of Nectoneanthes with Neanthes are includ- ed in the Neanthes succinea Remarks sec- tion (below). Neanthes succinea (Leuckart, 1847) Nereis succinea Leuckart, 1847:154, pl. 2, figs. 9, 11. B) Nereis (Alitta) oxypoda Marenzeller, 1879: 120-122, pl. 2, fig. 3. New synonymy. Nereis oxypoda. —Monro, 1934:362-363 (in part, specimen collected by Chen). Nereis (Neanthes) oxypoda. —\banez, 1972: 24-26, fig. 1. Nereis alatopalpis Wesenberg-Lund, 1949: 281-283, figs. 15-17. New synonymy. Nectoneanthes alatopalpis.—Wu, Sun & Yang, 1981:152-153, fig. 94A—-F; 1985: 168-170, fig. 94A—-F. Neanthes succinea. —Wilson, 1984:218- 221, fig. 4A-F (includes additional syn- onymies). Material examined.—JAPAN: (‘“‘wahr- scheinlich bei Yokohama”? Marenzeller, 1879:122) coll. Roretz 1876, 2 syntypes of Nereis oxypoda, ZMV 2140. Tokyo Bay, coll. Doderlein, 1 specimen of Nereis oxy- poda, ZMV 781. CHINA: Amoy, coll. T. Y. Chen, No. 27, 1 specimen of Nereis oxypoda, BMNH ZK 1933.3.2.40. IRAN: Gulf of Oman, Sta 72C, 12 km von Yask, coll. G. Thorson, 20.4.1937, ho- lotype (by monotypy) of Nereis alatopalpis, ZMK. Description. —Two syntypes of Nereis oxypoda: larger syntype with pharynx partly extended, consisting of anterior fragment of 169 setigers, about 260 mm long, 9 mm wide anteriorly (excluding parapodia), de- creasing to 7 mm wide in middle setigers and 3.5 mm wide in posteriormost setigers; smaller syntype with pharynx fully everted, consisting of anterior fragment of 103 se- tigers, about 100 mm long, 5 mm wide an- teriorly decreasing to 2.5 mm wide in pos- teriormost setigers. Color pale yellow. Prostomium dorsoventrally flattened, with 1 pair of triangular palps with very small palpostyles, | pair of antennae about 4 as long as prostomium. Four pairs of tenta- cular cirri, longest extending back 3-4 se- tigers, with cylindrical cirrophores basally; cirrophores of 2 dorsal pairs longer than ventral pairs, forming '3—'4 of length of cir- 6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rus. Jaws amber, with 11-12 small lateral teeth on smaller syntype; larger syntype with darker, stouter jaws with indistinct teeth. Pharynx with pale amber paragnaths, point- ed cones on area VI, low domes on all other areas. Larger syntype with many maxillary ring paragnaths indistinct or lost, area I = O; Il = 3-4; III = 4; IV = about 6; V = 0; VI = 10 in circular group on left, 0 on right side; VII-VIII = about 50 small paragnaths in 2 irregular rows. Smaller syntype with I = 0; II = 14/16; WI = 5; IV = 14/17; V = 1; VI = 10, 10; VU-VIII = 56 in 2 irregular rows. Notopodia and neuropodia each with 3 triangular lobes throughout, posterior parapodia with dorsal notopodial lobe be- coming greatly expanded and enclosing dor- sal cirrus in large notch (Marenzeller 1879: pl. 2, fig. 3). Dorsal and ventral cirri ap- proximately equal to length of neuropodial lobes throughout. Both syntypes with most setae shed or broken; few notopodial hom- ogomph spinigers present and homogomph and heterogomph setae with appendages missing remain in many neuropodia. Specimen from Japan (ZMV 781) con- sisting of anterior fragment of 81 setigers, about 100 mm long, 6 mm wide anteriorly; only 25 anterior setigers in good condition. Agrees with description of syntypes except as follows: pharynx everted, area I = 4 in longitudinal series; II = 19/21; HIIl= 6; IV = NG/175 Ve — 32 VAI a2 ViVi = 77. in 2—3 rows, including single evenly spaced anterior row of larger cones. Setae of ante- rior setigers mostly intact, homogomph spi- nigers in notopodia, homogomph and het- erogomph spinigers in both neuropodial fascicles. Specimen from China (BMNH ZK 1933.3.2.40) immature female epitoke with coelomic gametes, entire but regenerating posteriorly, 80 setigers, about 75 mm long, 5 mm wide (part of material examined by Monro 1934 and Wilson 1984). This spec- imen also agrees with the above descrip- tions and apparently lacks neuropodial fal- cigers. Holotype of Nereis alatopalpis (ZMK) in- complete posteriorly, broken into 3 pieces: an anterior fragment with pharynx everted, 46 setigers, 25 mm long, 2.5 mm wide; frag- ment of 16 setigers, 7 mm long and fragment of 10 setigers, 4 mm long. Prostomium strongly flattened, triangular, with small palpostyles, antennae 4 as long as prosto- mium; 4 pairs of tentacular cirri, longest extending back 4 setigers, 2 dorsal pairs with cirrophores accounting for about 4 of length of cirri. Jaws translucent amber, smooth distally, with 5—6 indistinct teeth basally. Paragnaths pale amber domes, arranged as follows: area I = 1; I] = 11/9; II] = 0; IV = 8/9; V = 0; VI = 8/11 in circular groups; VII-VIUI = about 32 in 2 widely spaced irregular rows. Notopodia and neuropodia each with 3 acutely triangular lobes, dorsal and ventral cirri about as long as parapodial lobes. Dorsal notopodial lobe greatly ex- panded posteriorly, enclosing dorsal cirrus in large notch (Wesenberg-Lund 1949:fig. 16a-d). Notosetae homogomph spinigers, neurosetae homogomph and heterogomph spinigers in both fascicles. Remarks. —Marenzeller’s original de- scription of Nereis oxypoda was very de- tailed and agrees with that of the two syn- types re-examined in this study. There are no significant differences in the form of the paragnaths and parapodia between this ma- terial and epitokes and sub-epitokes of Neanthes succinea (Leuckart) as described by Pettibone (1963:165) and Wilson (1984: 218). Nereis alatopalpis Wesenberg-Lund was also found to be identical to previous descriptions of N. succinea. Nereis oxypoda and N. alatopalpis are thus formally syn- onymized with Neanthes succinea and Nec- toneanthes Imajima, 1972 (of which N. oxy- poda is the type species) becomes a junior synonym of Neanthes Kinberg, 1866. Imajima (1972) gave absence of neuro- podial falcigers and expansion of the dorsal notopodial lobe as distinguishing characters of Nectoneanthes; however, these are char- acters associated with the development of VOLUME 101, NUMBER 1 epitoky. All material previously referred to Nectoneanthes represents large (100-300 mm long) specimens and it is probable that even the “‘atokes”’ described by Izuka (1912) and Imajima (1972) are in fact epitokes at varying stages of development. Several smaller specimens of N. succinea and the holotype of N. alatopalpis examined here apparently lack neuropodial falcigers; how- ever, in the absence of a detailed study of the ontogeny of N. succinea this character alone is insufficient to form the basis of a new taxon. Other nereidid species can apparently achieve sexual maturity in either their sec- ond or third year (e.g., Hediste diversicolor (Miller) and Nereis pelagica Linnaeus as indicated by Olive & Garwood (1981), Met- tam et al. (1982), and Moller (1985)), hence sexual maturity and epitokal modifications may also occur in N. succinea at different sizes and ages. It is possible that different populations differ in this respect: e.g., Reish (1977) noted that a population of N. suc- cinea introduced into the Salton Sea, Cali- fornia, differed from nearby natural popu- lations only in reaching a larger size. This paper does not address the possi- bility that sibling species may be present within what is now known as Neanthes suc- cinea. Examples already exist of widely dis- tributed nereidid species complexes which are morphologically similar but exhibit dis- tinct reproductive patterns (reviewed by Smith 1958): Hediste! diversicolor (Miller)/ Neanthes japonica (Izuka)/N. limnicola (Johnston) and Platynereis dumerilii (Au- douin & Milne Edwards)/P. megalops (Ver- rill)/P. massiliensis (Moquin-Tandon). Neanthes succinea may also be shown to represent such a complex; however, large numbers of specimens and knowledge of life histories of populations from widespread ' Hediste is currently held to be a monotypic genus (Hartmann-Schroder 1971, Fauchald 1977); a review of this group is required since obviously closely related species are presently placed in separate genera. localities will be needed to detect sibling species if they exist. The present study shows that the taxa Nectoneanthes, N. oxypoda, and N. alato- palpis (recorded only from within the range of Neanthes succinea) can no longer be re- tained. Distribution. —Neanthes succinea is widely distributed in the Northern and Southern Hemispheres (Pettibone 1963, Day 1967). It is likely that some of the more recent records are the result of introductions by humans, both intentional and accidental (Smith 1963, Reish 1977, Proskurina 1979). Neanthes sp. Nereis oxypoda. —Okuda, 1933:247, pl. 13, figs. f-h (not Marenzeller 1879). Nereis oxypoda var.—Monro, 1934:362- 363 (in part, 2 specimens collected by Ping) (not Marenzeller 1879). Nereis (Neanthes) oxypoda. —Treadwell, 1936:268 (not Marenzeller 1879). Nectoneanthes oxypoda.—Imajima, 1972: 113-117, figs. 35-37.—Paik, 1972:135, fig. 4a—h; 1973:82, figs. 1k, 1, 2, table 1 (not Marenzeller 1879). Neanthes (Nectoneanthes) oxypoda var.— Wilson, 1984:220-221 (not Marenzeller 1879). Material examined.—JAPAN: Kojima Bay, Okayama Prefecture, coll. K. Izuka, 17 Dec 1906, 3 specimens labelled Necto- neanthes oxypoda, NSMT 5290-5292. CHINA: Amoy, Outer Harbour, coll. Prof. C. Ping, 6 Apr 1923, No. 3, 2 speci- mens identified by Monro, 1934 as Nereis oxypoda var., BMNH ZK 1926.4.27.22-23. Amoy, No. 27, coll. T. Y. Chen, | gravid female identified as Nereis (Neanthes) oxy- poda by Treadwell, 1936, USNM 22240. Description. —Six specimens: smallest 104 setigers, 80 mm long, 4.5 mm wide (anterior fragment); largest 160 setigers, 260 mm long, 6 mm wide (complete). Prostomium flat- tened, wider than long, | pair of triangular palps with small palpostyles, 1 pair of an- 8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tennae about 4 as long as prostomium. Four pairs of tentacular cirri, longest extending back to setiger 3, 2 dorsal pairs with short indistinct cirrophores. Two pairs of dark red subdermal eyes. Jaws translucent amber with 6-12 distinct lateral teeth. Paragnaths amber, consisting of pointed cones on areas VI and maxillary ring, low domes else- where, arranged as follows: area I = 2-4 in longitudinal series; If = 16-30; III = 4-15 in triangular patch; IV = 24-32; VI = 8-13 in circular group; V—VIII = continuous band of 100-200 paragnaths, 6-8 rows deep dor- sally and ventrally, tapering to 2-3 rows deep laterally (Imajima 1972:fig. 35b, c). Noto- podia and neuropodia each with 3 acutely triangular lobes throughout and with dorsal and ventral cirri of about equal length to that of each parapodial lobe. Parapodial lobes of similar proportions throughout, ex- cepting dorsal notopodial lobe greatly ex- panded posteriorly, enclosing dorsal cirrus in large notch (Imajima 1972:fig. 35d—h). Notosetae homogomph spinigers, neurose- tae homogomph and heterogomph spinigers in dorsal and ventral fascicles. Anal cirri extending back over posteriormost 12-13 setigers. Remarks.—\majima (1972) apparently did not examine the syntypes of Nereis (Alit- ta) oxypoda Marenzeller, 1879, in desig- nating that species as the type species of Nectoneanthes. Imajima’s detailed descrip- tion of new material from several Japanese localities agrees with that given above for Neanthes sp. but differs significantly from Neanthes succinea in the arrangement of paragnaths on the oral ring: N. succinea has a ventral band of up to about 80 paragnaths tapering to a narrower band laterally and absent dorsally; Neanthes sp. has 100-200 oral ring paragnaths forming a continuous band 6-8 rows deep dorsally and ventrally and tapering to 2-3 rows laterally, exactly as figured by Imajima (1972:fig. 35b, c). In all other respects these specimens agree with the description of large specimens of N. suc- cinea but in view of the substantial differ- ences noted above this material is here con- sidered to represent a separate species. Neanthes sp. is not formally described and named here since all specimens examined are large and lack neuropodial falcigers; as with N. succinea (see above), smaller spec- imens of this form may be found to possess neuropodial falcigers but in any case it is desirable that a formal description of this taxon be based on a wider size range of ma- terial than is currently available. The above discussion shows that Imaji- ma (1972) misidentified his specimens as Nereis (Alitta) oxypoda Marenzeller, 1879, when designating that species as the type species of Nectoneanthes. Under Article 70 (b) of the International Code of Zoological Nomenclature (3rd edition), the case must then be referred to the International Com- mission on Zoological Nomenclature to designate a type species for the genus Nec- toneanthes; in the interest of stability I shall be proposing to the Commission that Nereis (Alitta) oxypoda Marenzeller, 1879, be so designated. Distribution. —Recorded from China, Ja- pan and Korea. Neanthes ijimai (Izuka, 1912) Nereis yimai Izuka, 1912:174-176, pl. 2, fig. 1, pl. 19, figs. 1-9. Neanthes ijimai.—Imajima and Hartman, 1964:144-145. Nectoneanthes ijimai. —Imajima, 1972: 117.—Wu, Sun & Yang 1981:146-148, fig. 9OA-H; 1985:161-163, fig. OA—H. — Srinivasa Rao & Sarma, 1982:447, figs. G5 Us Nectoneanthes imajimai (sic.).— Wilson, 1984:220 (demonic typographical error for Nectoneanthes ijimai). Remarks. —Izuka’s original description referred to a very large specimen (390 mm long for 215 segments); however, the only subsequent reports of new material relate to smaller specimens (40 mm long by Srini- vasa Rao & Sarma 1982; up to 12 mm long, 3.5 mm wide for 25 anterior setigers by Wu et al. 1981). These smaller specimens were VOLUME 101, NUMBER 1 reported to lack neuropodial falcigers; how- ever, as noted above in the Neanthes suc- cinea Remarks section, this character alone should not form the basis of a new genus. Neanthes ijimai is distinguished from other similar species discussed here by the ab- sence of paragnaths on areas I, III and V of the pharynx. Distribution. —Recorded from China, Ja- pan and India. Neanthes latipoda (Paik, 1973), new combination Nectoneanthes latipoda Paik, 1973:81-84, figs. 1, 2. Remarks. —The description of Neanthes latipoda by Paik (1973) was apparently based on larger specimens (163 mm long for 127 segments, 15 mm wide). As with the species of Nectoneanthes discussed above, the dis- tinguishing characters of N. latipoda can probably be attributed to the large size and approaching epitoky of the specimens ex- amined. The description and figures by Paik show that N. latipoda is very similar to ep- itokes of N. succinea but differs in the ar- rangement of paragnaths: the band of para- gnaths of the maxillary ring are continuous through the dorsal region as a single scat- tered band. As noted above, the maxillary ring band is not continuous dorsally in N. succinea. Neanthes latipoda appears to be intermediate between Neanthes sp. (above) and Neanthes succinea and is retained here as a Separate species. Distribution. —Korea. Neanthes multignatha (Wu, Sun & Yang, 1981), new combination Nectoneanthes multignatha Wu, Sun & Yang, 1981:148-149, fig. 91A—H; 1985: 163-164, fig. 91A—H. Remarks. —The figures of N. multignatha in the original description by Wu, Sun & Yang 1981 indicate that this species is very close to N. succinea, perhaps differing in the reduced numbers of paragnaths on areas III and IV of the pharynx. Distribution. —China. Acknowledgments I wish to thank the following persons for providing loan material: K. Fauchald (USNM), D. George and A. Muir (BMNBH), M. Imajima (NSMT), J. Kirkegaard (ZMK), and E. Kritscher (ZMV). I also thank K. Fauchald for encouragement and assistance with the literature, and M. Harvey, P. Hutchings, M. Pettibone, and G. Poore for their constructive comments on the manu- script. Literature Cited Augener, H. 1918. Polychaeta. Jn W. Michaelsen, ed., Beitage zur Kenntnis des Meeresfauna West- Africas 2(2):67-625. Clark, R. B. 1961. The origin and formation of the heteronereis. — Biological Reviews of the Cam- bridge Philosophical Society 36:199-236. Day, J. H. 1967. A monograph on the Polychaeta of southern Africa. Part 1. Errantia. British Mu- seum (Natural History) London, 458 pp. Fauchald, K. 1972. Benthic polychaetous annelids from deep water off western Mexico and adja- cent areas in the eastern Pacific Ocean.— Allan Hancock Foundation Monographs in Marine Biology 7:1—575. . 1977. The polychaete worms. Definitions and keys to orders, families and genera.—Los An- geles County Museum of Natural History Sci- ence Series 28:1-190. Fauvel, P. 1932. Annelida Polychaeta of the Indian Museum, Calcutta.—Memoirs of the Indian Museum 12:1-262. Hartman, O. 1940. Polychaetous annelids Part II. Chrysopetalidae to Goniadidae.—Allan Han- cock Pacific Expeditions 7:173-286. . 1959. Catalogue of the polychaetous annelids of the world Part I.—Allan Hancock Founda- tion Occasional Paper 23:1-353. Hartmann-Schr6éder, G. 1971. Annelida, Borsten- wurmer, Polychaeta.—Tierwelt Deutschlands 58:1-594. Ibanez, M. 1972. Notas sobre algunas especies de Anélidos Poliquetos nuevas para las costas de Espana, con especial referencia a Nereis (Neanthes) oxypoda Marenzeller y Onuphis (Nothria) geophiliformis Moore.—Boletin Real Sociedad Espanola de Historia Natural (Seccion Biologica) 70:23-31. 10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Imajima, M. 1972. Review of the annelid worms of the family Nereidae of Japan with descriptions of five new species or subspecies.— Bulletin of the National Science Museum, Tokyo 15:37- S33}, , & O. Hartman. 1964. The polychaetous an- nelids of Japan.—Allan Hancock Foundation Publication Occasional Paper 26:1—452. Izuka, A. 1912. The errantiate Polychaeta of Japan. — Journal of the College of Science, Imperial Uni- versity, Tokyo 30:1—262. Kinberg, J. G. H. 1866. Annulata nova.— Kungliga Svenska Vetenskapsakademiens, Stockholm. Ofversigt af Fordhandlingar 22:167-179, 238- 258. Leuckart, R.S. 1847. Verzeichnis der zur Fauna Hel- goland’s gehérenden wirbellosen Seethiere. Pp. 136-170 in H. Frey and R. Leuckart, eds., Bei- trage zur Kenntnis wirbelloser Thiere mit be- sonderer Berticksichtigung der Fauna des Nord- deutschen Meeres. Braunschwieg, F. Vieweg and Sohn. Marenzeller, E. von. 1879. Siidjapanische Anneliden I.—Denkschrift fiir Akadademie der Wissen- schaften Wien 41:109-152. Mettam, C., V. Santhanam & M.S. C. Havard. 1982. The oogenic cycle of Nereis diversicolor under natural conditions.—Journal of the Marine Bi- ological Association of the United Kingdom 62: 637-645. Moller, P. 1985. Production and abundance of ju- venile Nereis diversicolor, and oogenic cycle of adults in shallow waters of western Sweden. — Journal of the Marine Biological Association of the United Kingdom 65:603-617. Monro, C.C. A. 1934. Ona collection of Polychaeta from the coast of China.— Annals and Magazine of Natural History (10) 13:353-380. Okuda, S. 1933. Some polychaete annelids used as bait in the Inland Sea.—Annotationes Zoolog- icae Japonenses 14:243-253. Olive, P. J. W.,& P.R. Garwood. 1981. Gametogenic cycle and population structure of Nereis (He- diste) diversicolor and Nereis (Nereis) pelagica from north-east England.—Journal of the Ma- rine Biological Association of the United King- dom 61:193-213. Paik, E.-I. 1972. The polychaetous annelids in Korea (I).—Bulletin of the Korean Fisheries Society 5: 128-136. 1973. The polychaetous annelids in Korea (II). Description of Nectoneanthes latipoda, sp. nov.—Bulletin of the Korean Fisheries Society 6:8 1-84. Pettibone, M. H. 1963. Marine polychaete worms of the New England region. 1. Aphroditidae through Trochochaetidae. — Bulletin of the United States National Museum 227:1-356. Proskurina, Y.S. 1979. The state of introduced species in the Aral Sea and the prospects for their spread. — Gidrobiologicheskii Zhurnal 15(3):37- 41 [in Russian]. Translated in Hydrobiological Journal 15(3):30-34, 1979 [1980]. Reish, D. J. 1977. The role of life history studies in polychaete systematics. — Allan Hancock Foun- dation, Olga Hartman Memorial Volume, pp. 461-476. Los Angeles. Smith, R. I. 1958. On reproductive pattern as a spe- cific characteristic among nereid polychaetes. — Systematic Zoology 7:60-73. . 1963. On the occurrence of Nereis (Neanthes) succinea at the Kristenburg Zoological Station, Sweden, and its recent northward spread. — Ar- kiv for Zoologie 15:437-441. Srinivasa Rao, D., & D. V. Rama Sarma. 1982. New polychaete records from Indian waters. —Jour- nal of the Bombay Natural History Society 79: 445-450. Treadwell, A. L. 1936. Polychaetous annelids from Amoy, China.— Proceedings of the United States National Museum 83(2984):26 1-279. Wesenberg-Lund, E. 1949. Polychaetes of the Iranian Gulf. Jn K. Jessen and R. Sparck, eds., Danish scientific investigations in Iran 4:247—400. Wilson, R. S. 1984. Neanthes (Polychaeta: Nereidi- dae) from Victoria with descriptions of two new species. — Proceedings of the Royal Society of Victoria 96:209-226. Wu, Boaling, Ruiping Sun, & Deijang Yang. 1981. The Nereidae (Polychaetous annelids) of the Chinese Coast.—Institute of Oceanography, Academia Sinica Beijing [in Chinese, English summary]. English translation published by China Ocean Press, Beijing, and Springer-Ver- lag, Berlin, vi + 234 pp., 1985. Museum of Victoria, 285-321 Russell Street, Melbourne 3000, Victoria, Austra- lia. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 11-14 LIMNODRILOIDES FAXATUS AND DOLIODRILUS PUERTORICENSIS, NEW LIMNODRILOIDINAE (OLIGOCHAETA: TUBIFICIDAE) FROM PUERTO RICO Christer Erséus and Michael R. Milligan Abstract. — Limnodriloides faxatus, new species, with long, heavily muscular atrial ducts, dorsal spermathecal pores and torch-shaped sperm bundles in the spermathecal ampullae, and Doliodrilus puertoricensis, new species, with atrial ducts that are only moderately modified (by comparison with its congeners) and minute spermathecae, are described from the west coast of Puerto Rico. A few marine Tubificidae (subfamilies Rhyacodrilinae, Phallodrilinae, and Tubi- ficinae) were recently described from Aguadilla and Mayagtiez Bays in Puerto Rico by Milligan (1986) from material col- lected by Dr. A. Stoner for the Center for Energy and Environmental Research, Uni- versity of Puerto Rico. A new member of the Limnodriloidinae from the same col- lection, Limnodriloides faxatus, is de- scribed in the present paper together with Doliodrilus puertoricensis, another new species from the west coast of Puerto Rico. The material of the latter, which was placed at the first author’s disposal by Dr. M. L. Jones at the National Museum of Natural History (USNM), Washington, D.C., orig- inates from a benthic community study in Laguna Joyuda (near Mayaguez) conducted by Mr. R. L. Castro at the Department of Marine Sciences, University of Puerto Rico. The anterior end of one of the specimens of D. puertoricensis was sectioned and stained in Heidenhain’s hematoxylin and eosin; the remaining material was stained in paracarmine or Grenacher’s alcohol bo- rax Carmine and mounted whole in Canada balsam. The types series are deposited at the USNM. Limnodriloides faxatus, new species Fig. 1 Holotype. -USNM mounted specimen. Type locality. —Mayaguez Bay, | mile off- shore, 18°15'48”N, 67°12'05”W, 11.5 m, silt and clay with pockets of detritus (coll. A. Stoner, 14 May 1985). Paratypes. -USNM 101461, 2 speci- mens from Aguadilla Bay, 18°29'03’N, 67°10'44”W, 16m, mud and shell fragments (coll. A. Stoner, 18 Apr 1985). Other material examined (authors’ col- lection).—Six specimens: | from type lo- cality, 4 from locality of paratypes, and | from another station in Aguadilla Bay, 38 m, coarse sand and shell (coll. A. Stoner, 18 Apr 1985). Description. —Length (only | complete specimen) 7.1 mm, 52 segments. Width at XI (compressed, whole-mounted worms), 0.17-0.26 mm. Clitellum extending over XI-XII in 1 paratype, poorly developed in other specimens. Setae (Fig. 1A) bifid, 35- 50 um long, 1.5—2 wm thick, with upper tooth slightly thinner and shorter than low- er. Setae 2-3 per bundle anteriorly, absent from X—XI, 2 per bundle in postclitellar seg- 101460, whole- 12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. ments. Male pores paired in line with ven- tral setae in posterior part of XI. Spermathe- cal pores in line with dorsal setae in middle of X. Pharyngeal glands in IV-V. Pair of slen- der oesophageal diverticula present ante- riorly in IX. Male genitalia (Fig. 1B) paired. Vas deferens 16-22 um wide, probably somewhat longer than atrium (not observed in its full length in available material), en- tering atrium apically. Atrial ampulla 40- 75 um long, 28-50 um wide, ventrally bear- ing conspicuous prostatic pad; latter causing wall of ampulla to bulge considerably. Pros- tate gland large and lobed. Atrial duct slen- der (about 280 um long in holotype), entally narrow and without granulation, in middle part with granulated inner epithelium and thin outer lining, ectally without granula- tion but with up to 10 wm thick muscular lining. Atrial duct terminating in large pa- pilla at inner end of voluminous copulatory sac. Spermathecae (Fig. iB, s) oblong, 100- 145 um long, 35-55 um wide, with short (sometimes indistinct) ducts and thin-walled ampullae. In postcopulatory individuals, each ampulla with 1-2 torch-shaped, very Limnodriloides faxatus: A, Free-hand drawing of seta; B, Lateral view of genitalia in segments X— XI. (aa = atrial ampulla, ad = atrial duct, mf = male funnel, pp = pseudopenial papilla, ppa = prostatic pad, pr = prostate gland, s = spermatheca, vd = vas deferens.) characteristic bundles of sperm, latter partly embedded in amorphous mass of secretion. Remarks. — The dorsal spermathecae, the long atrial ducts, and the absence of sper- mathecal setae render L. faxatus a member of the agnes-group within Limnodriloides (cf. Erseus 1982). The new species is easily distinguished from the other forms within this group by its wide and heavily muscular atrial ducts (ducts narrow and with thin muscular lining in the other species), and the characteristic shape of the sperm bun- dles in the spermathecae. Etymology.—The name faxatus means ‘with torch,’ or ‘flame,’ and refers to the appearance of the sperm bundles in the spermathecae. Distribution and habitat.—Known only from the west coast of Puerto Rico. Subtid- al, muddy or shelly sediments, 11.5-38 m depth. Doliodrilus puertoricensis, new species Fig. 2 Holotype. -USNM mounted specimen. Type locality.—Laguna Joyuda, an en- 101462, whole- VOLUME 101, NUMBER 1 Fig. 2. Doliodrilus puertoricensis: A, Ventral view of segments X—XI, showing size and location of sper- mathecae and male ducts; B, Free-hand drawing of seta; C, Spermatheca; D, Male efferent duct. (mp = male pore, sp = spermathecal pore; other abbreviations as in Fig. 1.) closed brackish-water lagoon (salinities fluctuating between 8 and 44%) located 9 km S of Mayagiiez, W Puerto Rico, 1—2.5 m, mud (material coll. R. L. Castro on sev- eral occasions between 9 Nov 1978 and 10 Jul 1979, holotype on 10 Jan 1979). Paratypes.—USNM_ 101463-101468, 1 sectioned and 5 whole-mounted specimens from type locality. Other material. —USNM_ 101469- 101485, 17 whole-mounted specimens, in- cluding immature and partially mature worms, from type locality. Description.—Length (fixed specimens) 6.1-10.7 mm, 44-65 segments. Diameter at XI (whole-mounted, somewhat compressed worms) 0.32-0.44 mm. Clitellum extending over XI—XII. Setae (Fig. 2B) bifid, with up- per tooth thinner than, but about as long as, lower. Bifids 40-60 um long. 1.5—2.5 wm thick, (1)2—4 per bundle anteriorly, (1)2-3 per bundle in postclitellar segments. Ventral setae absent from XI. Male pores (Fig. 2A, mp) paired in line with ventral setae, pos- terior to middle of XI. Spermathecal pores (Fig. 2A, sp) paired in line with ventral se- tae, anteriorly in X. Pharyngeal glands in III-V. Oesophagus dilated, thick-walled and with reticulate blood-plexus, in whole length of IX. Male ducts and spermathecae small in relation to body size (cf. Fig. 2A). Male genitalia (Fig. 2D) paired. Vas deferens 7-10 um wide, about as long as atrium, joining apical end 14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of latter. Atrium slender, tubular, about 165 um long, with thin but distinct outer lining of muscles. Both ental part of atrial ampulla and ectal duct narrow (10-19 um wide), with thin inner epithelium; middle part of atrium (ectal part of ampulla) conspicuously wider (about 25 um), containing cone-shaped prostatic pad. Atrial duct shorter than am- pulla, somewhat granulated in its middle part, ectally slightly dilated and opening to exterior through simple pore. Prostate gland small, lobed. Spermathecae (Fig. 2C) very small, with 24—27 um long, 18-19 um wide, ducts, and more or less spherical, 20-30 um wide ampullae; ducts ectally dilated in fully mature specimens. Sperm as a small, curved bundle in each spermathecal ampulla. Remarks. — This is the third species in the genus Doliodrilus Erséus, 1984, which ac- commodates limnodriloidines with (1) modified oesophagus in IX (but without proper anteroventral diverticula in this seg- ment), (2) slender atria with a distinct pros- tatic pad in most ectal part of each atrial ampulla, (3) simple atrial ducts terminating in simple male pores, although the most ectal part of the ducts may be dilated, sharp- ly bent, or modified into a posteriorly di- rected blind sac, and (4) a similar dilation of the most ectal part of the spermathecal ducts; cf. definition of genus given by Erséus (1985). Doliodrilus puertoricensis is with regard to the atrial ducts the least modified species of the genus; in D. tener Erséus, 1984 (from Hong Kong) there is an abrupt bend about two-thirds down the length of the atrial ducts and in D. diverticulatus Erséus, 1985 (from Saudi Arabia) this ‘bend’ is further elabo- rated into a blind sac. The new species is further distinguished from the other two by its very small, roundish spermathecal am- pullae, and its fewer setae (up to 5 or 6, occasionally even 7, setae per bundle an- teriorly in the others), and from D. diver- ticulatus by its lack of an unpaired, dorsal oesophageal diverticulum in segment IX. Distribution and habitat.—Known only from the type locality, west coast of Puerto Rico. Subtidal muds, subject to fluctuating salinities, 1-2.5 m depth. The species co- occurs with Thalassodrilides gurwitschi (Hrabé) and T. belli (Cook); cf. Erséus (1981). Acknowledgments We are indebted to Dr. M. L. Jones (USNM) for placing the Laguna Joyuda specimens at the first author’s disposal, to Ms. Charlene D. Long (Salem, Massachu- setts) for environmental data on Laguna Jo- yuda, and to Ms. Barbro Lofnertz and Mrs. Aino Falck-Wahlstrom for technical assis- tance. Literature Cited Erséus, C. 1981. Taxonomy of the marine genus Thalassodrilides (Oligochaeta: Tubificidae).— Transactions of the American Microscopical Society 100:333-344. . 1982. Taxonomic revision of the marine ge- nus Limnodriloides (Oligochaeta: Tubifici- dae).— Verhandlungen des naturwissenschaft- ichen Vereins in Hamburg (Neue Folge) 25:207— BH . 1984. The marine Tubificidae (Oligochaeta) of Hong Kong and southern China. — Asian Ma- rine Biology 1:135-175. 1985. Annelida of Saudi Arabia. Marine Tubificidae (Oligochaeta) of the Arabian Gulf coast of Saudi Arabia.— Fauna of Saudi Arabia 6 (1984):130-154. Milligan, M. R. 1987. Marine Tubificidae (Oligo- chaeta) from Puerto Rico with descriptions of two new species, Tubificoides aguadillensis and Heterodrilus paucifascus.— Proceedings of the Biological Society of Washington 100(3):480- 489. (CE) Swedish Museum of Natural His- tory, Stockholm, and (postal address:) De- partment of Zoology, University of Gote- borg, Box 25059, S-400 31 Goteborg, Sweden; (MRM) Mote Marine Laboratory, 1600 City Island Park, Sarasota, Florida S350 Ts PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 15-19 PENNELLA MAKAIRA, NEW SPECIES (COPEPODA: PENNELLIDAE) FROM THE ATLANTIC BLUE MARLIN, MAKAITRA NIGRICANS, IN THE CARIBBEAN SEA W. E. Hogans Abstract.—A new species of pennellid copepod, Pennella makaira, is de- scribed. Collected from the flesh of Makaira nigricans off Puerto Rico, P. makaira belongs to the group of species in the genus Pennella which are less than 50 mm in total length. It differs from its congeners in the size and ar- rangement of cephalothoracic papillae, and the structure and segmentation of the first and second antennae. During an examination of the parasitic copepod genus Pennella Oken, 1815, in the collection of the United States National Museum of Natural History, three speci- mens of a previously undescribed species were discovered. This new species, P. ma- kaira is described and illustrated herein. Methods The cephalothorax of each specimen was cleaned with 20% warm NaOH. After clean- ing, one each of the first and second anten- nae and swimming legs were removed from the cephalothorax of two specimens and cleared in 85% lactic acid to reveal struc- tural details. Figures were drawn with the aid of a camera lucida or drawing tube under phase-contrast microscopy (Photozeiss). Pennella makaira, new species Figs. 1-6 Description (based on 3 specimens).— Pennellidae: cephalothorax (Fig. 2a, b) sub- spherical, with rounded anterior end slight- ly excavate at center in dorsal view. Surface of anterior end completely covered by pa- pillae (antennary processes). Papillae large, clavate to clublike, branched at margins; be- coming smaller and spherical to knoblike at center. Junction of posterior end of cepha- lothorax and anterior of neck with 2 lateral horn holdfasts. Holdfast horns cylindrical, unbranched. Neck short, stout, cylindrical, expanded at junction with trunk. Trunk cy- lindrical, slightly transversely ridged, three- quarters total length, curved slightly ven- trally in all specimens. Oviducts on ventral surface at junction of posterior end of trunk and anterior of abdomen. Abdomen dor- soventrally compressed, posterior extrem- ity bifid, rounded. Ventral surface of ab- domen covered by 2 rows of plumelike processes; plumes single or secondarily branched (Fig. 3). Egg strings (Fig. 4) in- complete, portions remaining cylindrical, filiform. Eggs uniseriate. Mean total length of specimens 28 mm (27-29 mm). First an- tennae (Fig. 5) 3-segmented, setose. Basal segment subquadrangular. Second segment cylindrical, with at least 22 setae (some probably missing in all specimens). Ter- minal segment three-quarters length of sec- ond segment, with apical armature of 12 setae; 9 long, flagelliform; 3 short, spini- form. First antennae on dorsal surface of cephalothorax, near anterior margin. Sec- ond antennae (Fig. 6) immediately anterior to first, 2 segmented, chelate, with large re- curved claw closing on triangular spur on distal margin. Single, stout seta at base of 16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | GR. rm Fig. 1. Pennella makaira: trunk, lateral; cephalo- thorax, dorsal (holotype). claw. Basal segment subquadrangular. Coxa of second antennae attached to subquadran- gular scleritorized plate with single trian- gular pigment spot. Mouth pennellid, bur- ied between papillae on anterior end of cephalothorax. Maxillae, maxillipeds not discernible. Four pairs of swimming legs on ventral surface close to posterior margin of cephalothorax. First 2 pairs biramous, third and fourth pairs uniramous. Armament formula as follows: Endopod Exopod 1 D) 1 2 Leg | 1-0 7 le S31 Leg 2 1-0 i] 1-1 Sl Leg 3 — — 0-0 ?,? Leg 4 — — 0-0 4,1 Male and pre-metamorphosis female un- known. Host. —Makaira nigricans Lacépéde (At- lantic blue marlin). Site of infestation. —Branchiostigeal area: cephalothorax, lateral horns and neck of parasite embedded in flesh. Locality. —San Juan Canyon, Puerto Rico. Holotype. —USNM 111554. Paratype. —USNM 222997. Discussion. —During the examination of more than 200 specimens of Pennella in the USNM, three features were found which could be used for separating species of the genus: 1) Overall size of the mature female. Wilson (1917) used total length of the par- asite (excluding egg strings) to separate the known species into two groups, those longer than 100 mm and those less than 50 mm. This is, from my examinations, a valid char- acter for general sorting of species; P. ma- Kaira is in the less-than-50 mm category. 2) The number of segments of the first and second antennae and the structure of the terminal segment of the second antennae is consistent for each species. 3) In small species, the size, shape and arrangement of cephalothoracic papillae is similar among NO © eS L> &IR¥ ce S Od: OS A) yy, S, artic NE TTA rer, Me anil high Figs. 2-6. Pennella makaira: 2a, Anterior end of paratype, ventral; 2b, Same, dorsal; 3, Plumules from abdominal brush; 4, Egg string (incomplete); 5, First antenna, dorsal; 6, Terminal segment of second antenna, dorsal. PROCEEDINGS OF — 8 Table 1.—Comparisons of antennae structure of Pennella makaira to similar species in the genus Pen- nella. Second antennae number of segments First antennae number of Species segments NW FN WH W W . biloboa elegans . exocoetl longicauda makaira n. sp. platycephalus robusta . Sagitta Ge] ge) AS) aches) gach as) WN WW * * No available information. a Based on two specimens identified by C. B. Wilson from Exocoetus volitans (USNM 112252). > Based on one specimen identified by the author from Histrio histrio (BMNH 1985, 471). specimens collected from the same host species or individuals of host species. Pennella makaira is the smallest species of Pennella yet recorded from a scombroid fish. One other slightly larger species from scombrids, P. biloboa Kirtisinghe, 1932, was reported from the sailfish, [stiophorous brevirostris collected off Sri Lanka. The in- adequate description of this species pre- cludes any detailed comparisons. I have been unable to obtain the single type specimen for comparative purposes. From the limited figures 1n the original description, P. biloboa appears to be a slimmer, more delicate par- asite than P. makaira. Gnanamuthu (1957) described four species of Pennella from flyingfishes off the coast of India. None of THE BIOLOGICAL SOCIETY OF WASHINGTON the species are adequately described. I have not been able to secure type material for any of these species: P. elegans, P. longicauda, P. platycephalus, and P. robusta. They could, judging from the limited figures, be speci- mens ofa single species differing only in age. All four species have total lengths less than P. makaira. Two other small species, P. exocoeti (Holten, 1802), and P. sagitta (Lin- naeus) appear to be valid. They have been well described previously and figured and can be distinguished from the new species as follows: P. exocoeti is found only on flyingfishes of genus Exocoetus. It has a 4-segmented first antenna, whereas the first antenna of P. makaira has only three seg- ments. In addition, the distal spur of the terminal segment of the second antenna of P. exocoeti is much larger and more sharply pointed than in P. makaira. Pennella sagitta is found only on the sargassum fish Histrio histrio; it is a very delicate parasite with no spherical cephalothoracic papillae. One species which approaches P. makaira in to- tal length, P. diodontis Oken, Chamisso, & Eysenhardt, 1821, is found, supposedly, only on diodontid porcupine fishes. The validity of the species has been questioned frequent- ly (see Wilson 1917). It has never been ad- equately described. Although most of the above-listed species are of uncertain valid- ity, a comparison of antenna structure and papillae form for each (compiled from the original descriptions) with those exhibited by P. makaira is shown in Tables 1 and 2, respectively. Table 2.—Comparison of the structure of papillae of the cephalothorax of Pennella makaira to similar species in the genus Pennella. Species Papillae shape Papillae arrangement Papillae size P. biloboa spherical to clavate, unbranched partially covering variable P. elegans spherical to clavate, unbranched partially covering similar P. exocoeti club-like to spherical, unbranched partially covering variable P. longicauda clavate, unbranched partially covering variable P. makaira n. sp. tubiform to spherical, bifid irregular branching completely covering variable P. platycephalus spherical clavate to spherical, branching partially covering similar P. robusta clavate to spherical, branching completely covering variable P. sagitta tubiform, irregular branching completely covering similar VOLUME 101, NUMBER 1 Acknowledgments I thank Dr. Roger Cressey for reviewing this paper and for the loan of material used during this study from the Smithsonian In- stitution, Dr. Paul Illg for collecting the par- asites, Dr. Z. Kabata for providing infor- mation on the appendage structure in Pennella, Dr. M. Deb (ZSI) for searching for the types of Gnanamuthu’s specimens, and B. Fullerton for preparing the type- script. 19 Literature Cited Gnanamuthu, C. P. 1957. Lernaeid copepods para- sitic on flyingfish.— Parasitology 47(1—2):119- 125. Kirtisinghe, P. 1932. Two new parasitic copepods from Ceylon.—Parasitology 24(4):548-551. Wilson, C. B. 1917. North American parasitic co- pepods belonging to the family Lernaeidae, with a revision of the entire family.— Proceedings of the United States National Museum 53:1-149. Huntsman Marine Laboratory, St. An- drews, New Brunswick EOG 2X0, Canada. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 20-30 THE PARASITIC ISOPOD HOLOPHRYXUS ACANTHEPHYRAE STEPHENSEN (EPICARIDEA: DAJIDAE) FROM THE SUBANTARCTIC SOUTH PACIFIC, WITH NOTES ON ITS SYNONYMY AND HOST Robert A. Wasmer Abstract. —The dajid isopod Holophryxus acanthephyrae Stephensen, 191 2a, is newly reported from the subantarctic South Pacific. A female of the species, and the male recovered from her marsupium, collected attached to the carapace of a specimen of the oplophorid shrimp Acanthephyra pelagica (Risso) are described. It is concluded that Jsophryxus concavus Schultz, 1977, the type species of the genus Jsophryxus, recorded from the antarctic and subantarctic South Pacific, is a junior synonym of Holophryxus acanthephyrae. The three other known species of Jsophryxus, I. quadratohumerale Schultz, 1978, I. poly- andrus Schultz, 1978, and IJ. septapodus Schultz, 1978, are transferred to Hol- ophryxus. Physical damage caused to the host shrimp by H. acanthephyrae is discussed. A list of the known species of Holophryxus, their distributions, and known host species is presented. During a study of the pelagic caridean shrimps belonging to the family Oplophori- dae collected by the USNS Eltanin from the antarctic and subantarctic South Pacific during the U.S. Antarctic Research Pro- gram (Wasmer 1986), I found an oviger- ous female dajid isopod loose among spec- imens of the oplophorid Acanthephyra pelagica (Risso) collected at station 1723 of Eltanin cruise 24. A note in the sample bot- tle indicated that the isopod was collected attached to one of the shrimps in the sam- ple, and a male specimen of A. pelagica in the bottle was found to have marks on the dorsal surface of its carapace which ap- peared to have been made by the mandibles and pereopods of the isopod. An additional male specimen of A. pelagica from Eltanin cruise 19, station 1480 was noted to have marks on its carapace similar to those on the shrimp from station 1723 of cruise 24, although no isopod was present in the sam- ple bottle when it was obtained from the Smithsonian Oceanographic Sorting Center (SOSC). I tentatively identified the female isopod (and the male found enclosed in her mar- supium) from station 1723 as a member of the genus Holophryxus Richardson, based on the redescription by Butler (1964) of H. alaskensis Richardson, 1905 (the type species of the genus) and my examination, 1966, of the type specimens of H. califor- niensis Richardson, 1908 (synonymized with H. alaskensis by Butler in 1964) and of H. giardi Richardson, 1908. The exam- ination of the types of these latter two species was made in conjunction with the identifi- cation of several dajids from shrimps col- lected off the coast of Oregon by oceano- graphic vessels of Oregon State University. Of the five previously recognized species of Holophryxus, only one has been recorded from an oplophorid shrimp. Stephensen (1912a) described H. acanthephyrae based on a mature female collected off Greenland VOLUME 101, NUMBER 1 attached to the carapace on an oplophorid he identified as Acanthephyra purpurea, but which he later concluded (1912b) was 4. multispina, a synonym of A. pelagica (see Wasmer 1986). The recent redescription of the female and the first description of the male of H. acanthephyrae by Jones & Smal- don (1986), published while the current work was in an early draft stage, strengthened my conclusion that the female and male isopod from Eltanin station 1723 are H. acanthe- phyrae Stephensen. The dajid genus Isophryxus was estab- lished by Schultz (1977), with the type species J. concavus. The description was based on the holotype female and the al- lotype male found enclosed in her marsu- pium collected by a midwater trawl at sta- tion 1480 of Eltanin cruise 19. Eight additional specimens (six females and two males) of the species were recorded (Schultz 1977) from six midwater trawl stations oc- cupied by the E/tanin during cruises 11, 15, and 19 in the antarctic and subantarctic South Pacific. The host species of none of these specimens was known to Schultz. Schultz pointed out that I. concavus is sim- ilar to the type species of the genus Holo- phryxus, from which it otherwise differs by having the pereonal segments indicated on the dorsal surface of the female. Schultz (1978) subsequently described three addi- tional species of Isophryxus (I. quadrato- humerale, I. polyandrus, and I. septapodus) from specimens collected by midwater trawls during E/tanin cruises 24, 25, and 35 in the antarctic and subantarctic South Pa- cific. Again, the host species is not known for these three species. The purpose of this paper is to describe the female and male Holophryxus acanthe- phyrae from Eltanin station 1723, and to propose, based on examination of the per- tinent types, that Jsophryxus concavus Schultz is a synonym of Holophryxus acan- thephyrae Stephensen. Evidence of physical damage caused to the host shrimp by H. acanthephyrae is also discussed. Lastly, a 21 list of the known species of Holophryxus, their distributions, and known host species is presented. Suborder Epicaridea Family Dajidae Holophryxus acanthephyrae Stephensen Fig. 1A—D Holophryxus acanthephyrae Stephensen, 1912a:112, figs. 13, 15-21.—Jones & Smaldon, 1986:303, figs. 1-6. TIsophryxus concavus Schultz, 1977:93, figs. 17A-17I, 18A—18H, 180, 18P. Material. —Ovigerous female (22.7 mm long and 10.7 mm wide), and male (2.3 mm long, 1.1 mm wide) from marsupium of fe- male, collected attached to a male specimen of the oplophorid shrimp Acanthephyra pe- lagica (Risso), cl. 26.0 mm, from USNS E/- tanin cruise 24, sta 1723 (18 Jul 1966, 40°01'S, 149°57'W to 40°05’S, 149°55’W, 880 m), USNM 233550. Description. — Adult female: Body sym- metrical, oblong-ovate, dorsal surface con- vex, ventral surface strongly concave ante- riorly. Color in alcohol light yellow. Dorsal prominence of cephalon slightly bilobed, not visible from below. Cephalic ridge narrow in dorsal view, anterior margin sinuous. Eyes absent. True segmentation of pereon evident only in lateral view, indicated by posterior 4 pairs of 5 coxal plates; first pair of coxal plates fused with cephalic ridge, forming anterior boundary of oral area; fifth pair small, sep- arated from fourth by distinct space. Dorsal surface of pereon with 3 shallow transverse furrows in integument behind rounded prominence of cephalon. Pleon continuous with but narrower than pereon, tapering posteriorly to rounded ex- tremity, unsegmented, lacking pleopods and uropods. Dorsal surface of pleon deeply ex- cavated on both sides of midline near junc- tion with pereon. Oral area bounded anteriorly and later- ally by flattened flange-like cephalic ridge. 22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Holophryxus acanthephyrae, Eltanin station 1723: A, Dorsal view, mature female; B, Ventral view, anterior part of pereon, mature female, x = pit organ; C, Ventral view, posterior part of pereon and _pleon, mature female; D, Dorsal view, male. Cephalic ridge with pair of narrow, slit-like pores or pit organs (Fig. 1B, x) about half- way from midline to lateral edge. First and second pairs of antennae unsegmented and flattened, with elongate lobe-like peduncles surrounding oral area. Tips of mandibles styliform, visible in central part of oral area; remaining mouthparts not visible without dissection. Five pairs of short pereopods present (pereopods 6 & 7 lacking), posterior 4 pairs arising from bases near separate cox- al plates. Pereopods prehensile, all of sim- ilar size and shape, with dactyls strongly recurved and clawlike. Five pairs of incubatory plates (ooste- gites) arising from bases of pereopods; only first and fifth pairs visible, covering other 3 pairs. Fifth pair of plates largest, extending to posterior end of pleon and forming largest part of marsupium, medial edges overlap- ping slightly; posterior margin of each end- ing in flattened crest just before pleon, each fringed with long spines (12 on left, 13 on right) above posterior end of marsupium. Male: Elongate, symmetrical, dorsal sur- face slightly convex, ventral surface con- cave. Color in alcohol yellowish white. Cephalon large, rounded anteriorly, com- pletely fused with pereonite 1, lateral mar- gin slightly indented posteriorly, indicating anterior limit of first pereonite. Eyes absent. First antennae short, composed of 3 articles; VOLUME 101, NUMBER 1 second antennae somewhat longer (com- posed of 3 articles?), directed posteriorly; neither pair visible in dorsal view. Mandi- ble tips projecting from oral area. Pereonites 2—6 distinctly separated from each other, pereonite 7 almost completely fused with pleon; lateral margins of second through sixth rounded, lateral margins of seventh conical. Seven pairs of prehensile pereopods present. Pleon conical, slightly longer than pereo- nites 1-6 combined, longer than wide, un- segmented, tapering posteriorly to narrowly rounded extremity. Pleopods and uropods absent. Remarks. — Based on my comparison of the female and male dajid specimens. and their host (Acanthephyra pelagica) from El- tanin station 1723 with the female holotype of Holophryxus acanthephyrae Stephensen, and its type host (Acanthephyra pelagica) obtained on loan from the Universitetets Zoologiske Museum in Copenhagen, as well as the description of the species given by Jones. & Smaldon (1986), it is concluded that these specimens are identical with H. acanthephyrae. There appear to be no significant differ- ences between the holotype of H. acanthe- phyrae and the specimen from the E/tanin oplophorid. Both specimens are ovigerous females of comparable size and shape. The morphological details visible in dorsal, ven- tral, and lateral views of the two specimens are in very good agreement, although there is a minor difference in the number of spines on the posterior marginal crests of the fifth incubatory plates of the two specimens (14 on each in the case of the holotype, but only 12 and 13 on the specimen from the Eltan- in). The female described here is also in es- sential agreement with the female H. acan- thephyrae redescribed and illustrated by Jones & Smaldon (1986), although that specimen is smaller and somewhat more de- formed. The cephalic pores, first pointed out by Jones & Smaldon (1986), are also 23 easily seen on the female described here. I was unable to see the pores on the holotype of H. acanthephyrae or on other members of the genus examined in the course of this study; perhaps the opening to the pore, which may be a sense organ of some type, can be contracted or closed off by the isopod so that it would be especially hard to see on some preserved specimens. It is unfortunate that Stephensen did not have a male specimen of H. acanthephyrae with which the present male specimen can be compared. It is very likely that his ho- lotype female has a male in her marsupium; several of the previously recorded males of other Holophryxus species have been taken from ovigerous females on which the me- dial margins of the fifth incubatory plates are close together or from females which have recently emptied marsupia (Rustad 1935, Butler 1964, Jones & Smaldon 1986). The male described here is very similar to the male described by Jones & Smaldon (1986). The shape of the lateral margins of the seventh pereonites differ in the two spec- imens, being more conical in the present specimen. However, as pointed out by Jones & Smaldon (1986), there are few diagnostic characters separating the males known for the different species of Holophryxus, as well as evidence of some variation in body form between males of the same species. The fact that the holotype of H. acanthe- phyrae and one of the four specimens of the species reported by Jones & Smaldon (1986), as well as the present female, were collected on specimens of the oplophorid shrimp Acanthephyra pelagica is strong evidence that the dajid from the E/tanin material is Holophryxus acanthephyrae. Each of the five species of Holophryxus for which the host is known occurs on only a single species of pelagic decapod shrimp (see below). Holophryxus acanthephyrae has been re- corded previously from the type locality in Davis Strait west of Greenland (60°07’N, 48°26'W) and from the Western Ap- proaches and Bay of Biscay (Stephensen 24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1912a, Jones & Smaldon 1986). The present record from the subantarctic South Pacific thus considerably extends the known dis- tribution of the species. This new record is not totally unexpected when the wide dis- tribution of its host in the North Atlantic from about 13°N northward, the Mediter- ranean, the South Atlantic from 24°S south- ward, and the Indo-Pacific between 32°S and 57°S (Wasmer 1986) is considered. Discussion Notes on synonymy. —Comparison of the holotype and allotype of [sophryxus con- cavus Schultz in the National Museum of Natural History with the holotype of Holo- phryxus acanthephyrae and with the female and male dajids from E/tanin station 1723 identified herein as H. acanthephyrae, has convinced me that Schultz (1977) mistak- enly separated Jsophryxus from Holo- phryxus and that Isophryxus concavus and Holophryxus acanthephyrae are in fact the same species. By the rule of priority, [so- phryxus concavus should therefore be con- sidered a junior synonym of Holophryxus acanthephyrae and Isophryxus becomes a junior synonym of Holophryxus. Schultz (1977) appears to have over- looked Butler’s 1964 redescription of Holo- phryxus alaskensis Richardson, 1905, the type species of the genus. Butler pointed out that folds in the dorsal integument of the pereon of several specimens of that species give the impression of segmentation, but that the folds are not related to segmenta- tion. Richardson did not mention well de- fined transverse folds in her description or show them in her figures of H. alaskensis; however, the specimens were apparently somewhat stout and irregularly shaped when preserved (Butler 1964), and for that reason the folds are probably not so evident. Folds or furrows are also present on the dorsal integument of the pereon of Hypodajus georgiensis Nierstrasz & Brender 4 Brandis (1931:212, figs. 106-108), synonymized with Holophryxus alaskensis by Butler (1964). Anterior dorsal integumentary fur- rows of the pereon are present to some ex- tent on all other known species of Holo- phryxus and have been noted in descriptions or are visible on figures of those species, as indicated in the following citations: H. giardi Richardson, 1908:fig. 1; H. richardi, Ste- phensen, 191 2a:fig. 11; Rustad, 1935:16, fig. 4: H. acanthephyrae Stephensen, 1912a:113, fig. 15; Jones & Smaldon, 1986:309, fig. 2A and 2B; and H. fusiformis Shiino, 1937:188, fig. LA and 1B. The flattened posterior marginal crests armed with long fringing spines on the fifth incubatory plates of the holotype and para- type females of J. concavus (not mentioned in Schultz’s description of the species but visible in his figs. 17A and 18B) also indi- cate that these specimens belong to the ge- nus Holophryxus. Such crests are apparent- ly present on mature females of the other species of the genus. Although Richardson (1905) does not specifically mention the crests and spines in her diagnosis of the ge- nus or in her description of H. alaskensis, her fig. 8c seems to indicate the presence of spines on the posterior part of the fifth in- cubatory plates. Butler (1964), in his rede- scription of H. alaskensis, mentions the presence of such fringed crests on the fifth incubatory plates on mature females of the species, although his fig. 1c curiously does not show them. The crests are also drawn and described by Nierstrasz & Brender a Brandis (1931:212, fig. 108) in their descrip- tion of Hypodajus georgiensis. Richardson (1908) does not mention the fringed crests in her description of Holophryxus giardi and they do not show on her fig. 1, but in my examination of the syntypes I found that they are in fact present on the mature female but folded downward and under the pos- terior edge of the fifth incubatory plates so as to be hidden from view; this is the result of the specimen being somewhat deformed when preserved. For the other species in the genus, the following citations show the pres- VOLUME 101, NUMBER 1 Fig. 2. Acanthephyra pelagica: A, Type host to Holophryxus acanthephyrae Stephensen, from Tyjalfe station 322, view of posterior right carapace; B, Posterior right carapace edge of specimen from E/tanin station 1723; C, Posterior right carapace edge of specimen from E/tanin station 1480; D, Posterior dorsal surface of carapace of specimen from Eltanin station 1480, showing marks caused by mandibles and pereopods of isopod; E, Posterior dorsal surface of carapace of specimen from E/tanin station 1723, showing marks caused by mandibles and pereopods of isopod. ence of this morphological feature: H. rich- ardi, Stephensen, 1912a:109, 110, figs. 9, 11; Rustad, 1935:26, fig. 14; H. acanthe- phyrae Stephensen, 1912a:115, figs. 15, 17; Jones & Smaldon, 1986:309, figs. 2C and 2D; and H. fusiformis Shiino, 1937:190, figs. 38}, (Ce The allotype male and other males re- covered by Schultz (1977) from females of I. concavus show no apparent morpholog- ical differences between the male H. acan- thephyrae described here and that described by Jones and Smaldon (1986). The host of Jsophryxus concavus was un- known to Schultz (1977) because the iso- pods apparently fell off the hosts at the time of collection. The presence of a male spec- imen of the oplophorid Acanthephyra pe- lagica (carapace length 26.5 mm), USNM 233552, from Eltanin station 1480 (the type locality station for I. concavus) with marks from the pereopods and mandibles of an 26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON isopod on the posterior dorsal surface of its carapace (Wasmer 1986), strongly suggests that this shrimp may have been the host of the holotype of J. concavus. The marks on the shrimp (Fig. 2D) are similar to those described and illustrated by Stephensen (1912a) on the type host of Holophryxus acanthephyrae and to those on the specimen of A. pelagica (Fig. 2E) which was the host of the dajid identified as H. acanthephyrae from Eltanin station 1723; this latter shrimp is on deposit in the National Museum (USNM 233551). Although it is difficult to gauge accurately the size of the isopod responsible for the marks on the shrimp from E/tanin station 1480, the marks are not inconsistent with an isopod 21.6 mm long (the length of the holotype of J. concavus). The holotype of H. acanthephyrae is 22 mm long (Stephen- sen 1912a), and the specimen from E/tanin station 1723 is 22.7 mm long; the marks on the carapaces of these latter two specimens are in the same size range as those on the presumed host specimen of J. concavus. An attempt was made to identify addi- tional host specimens of A. pelagica from the six E/tanin stations listed by Schultz (1977) where the paratypes of J. concavus were collected. This attempt was not suc- cessful; none of the 14 specimens of A. pe- lagica from five of the stations have any marks indicating they served as hosts to iso- pods, and no specimens of A. pelagica were reported from the sixth station. However, if the specimen of A. pelagica from Eltanin station 1480 is assumed to be the type host of I. concavus, then the evidence becomes stronger that the specimens described as J. concavus are in reality H. acanthephyrae. In addition to the undue significance at- tached by Schultz (1977) to the presence of the dorsal integumentary folds on the pere- on of the genus /sophryxus, several other, more minor, erroneous interpretations of morphology are present in his diagnosis of the genus and in the description of J. con- cavus. These misinterpretations involve the supposed absence of both pairs of antennae in the females, the presence of “flattened mouthparts,” and the misnumbering of the lateral edges of the pereonites (coxal plates). The two pairs of antennae on females of the genus Holophryxus have presented dif- ficulties of interpretation for most previous investigators, probably due to the rather un- usual appearance of these appendages and to the lack of sufficient numbers of speci- mens for adequate dissection. Nierstrasz & Brender a Brandis (1931) and Shiino (1937) incorrectly considered both pairs of anten- nae to be entirely absent and interpreted them as the maxillipeds. Richardson (1905) considered both antennae to be rudimen- tary and articulated. Butler (1964) consid- ered them to be rudimentary and flattened, with the first pair being questionably artic- ulated; he also called a part of the peduncle of the second antennae the maxilliped, an error pointed out by Coyle & Mueller (1981). Stephensen (1912a) considered both pairs of antennae to be present but questionably articulated. Rustad (1935), with access to a number of specimens of H. richardi, was able to carry out detailed dissections and preparations with hot sodium hydroxide and pyrogallic acid stain to investigate the struc- ture of the two pairs of antennae, as well as the mouthparts. Of the mouthparts, only the tips of the mandibles and part of the maxillipeds were normally visible without dissection. The two pairs of antennae were described by Rustad as being more or less deformed and “cushiony,” and encircling the oral cone; even though they occasionally appeared to be articulated, the special pre- parative techniques used showed them to be unjointed. Comparison of Schultz’s (1977) fig. 17B of the anterior ventral part of J. concavus with fig. 16 of Stephensen (191 2a) and figs. 2 and 7 of Rustad (1935) leads to the con- clusion that what Schultz refers to as flat- tened mouthparts on his specimens repre- sent the first and second pairs of antennae of Stephensen and Rustad. VOLUME 101, NUMBER 1 In at least several (and perhaps all) of the species of Holophryxus, the first pair of cox- al plates on adult females is coalesced with the cephalic ridge and the posterior four pairs of coxal plates are generally separate and evident as rounded or acute projections lat- eral to pereopods 2-5 (see Richardson 1908: 690, fig. 1, for H. giardi; Rustad 1935:9, fig. 2, for H. richardi; Shiino 1937:188, fig. 1B, C, for H. fusiformis; Stephensen 1912a:113, llr, fig. 16, and Jones & Smaldon 1986: 309, fig. 2C for H. acanthephyrae). Figure 1B herein, showing the anterior ventral re- gion of the specimen of H. acanthephyrae from El/tanin station 1723, is in general agreement with fig. 16 of Stephensen (191 2a) and fig. 2C of Jones & Smaldon (1986) and shows the first coxal plates to be fused with the cephalic ridge, with the first pereopods arising from near the posterolateral part of the ridge. The lateral edges of pereonal seg- ments I-IV of Schultz (1977) should there- fore more correctly be referred to as coxal plates II-V. The three additional species of /so- phryxus (I. quadratohumerale, I. polyan- drus, and I. septapodus described by Schultz (1978) from Eltanin cruises 11, 15, and 19 need to be briefly considered here. The pres- ence of weak anterior furrows or folds sug- gesting segmentation of the pereon, the presence of fringing spines on the posterior margin of the fifth oostegites, and the form of the males clearly indicate that J. quad- ratohumerale and I. polyandrus represent species of Holophryxus. The situation with I. septapodus is perhaps less clear-cut than that with the other described species of Jso- pDhryxus. Rustad (1935), Butler (1964), and espe- cially Coyle & Mueller (1981) have shown that the development of the juvenile female stage into the mature Holophryxus female involves loss of pleopods, uropods, abdom- inal segmentation, a change in the mandi- bles from the sagittate form of the crypto- niscid to a rasplike organ capable of drilling into the carapace, and a reduction of pereo- 27 pod 7 (with only its coxal plate remaining); in the final changes to the adult form, coxal plate 7 is lost, oostegites 1-5 are formed, the body region between pereopods 5 and 6 lengthens, and pereopod 6 is finally lost and its coxal plate reduced to a small bump or papilla which may persist at the posterior part of the pereon (as illustrated in Nier- strasz & Brender a Brandis 1913, Shiino 1937, and Butler 1964). Breeding does not occur until the marsupium is formed (Coyle & Mueller 1981), a condition that is prob- ably not complete until the fringing spines are present on the posterior margins of the fifth oostegites. Contrary to Schultz’s statement (1978:83) about the absence of spines on the posterior margin of the marsupium of J. septapodus, my examination of the holotype of this species showed them to be present. Based on the above information, I suggest that the gravid female holotype of J. septapodus is a specimen of Holophryxus on which the fringing spines are present on the posterior marsupial margin, but which still has ves- tiges of the seventh pereopods and the cor- responding seventh coxal plates, represent- ed by the posterior two pairs of processes on the posterior ventral part of the pereon (see Schultz 1978:fig. 11 A), as well as joint- ed sixth pereopods and the corresponding sixth coxal plates, represented by the more anterior two pairs of processes on the pos- terior ventral part of the pereon. Whether these posterior appendages are retained much beyond maturity in H. sep- tapodus (Schultz) can be determined only when additional mature specimens come to hand. Rustad (1935:16-17, figs. 3, 6) has shown that at least in H. richardi there is some individual variation in regard to the stage of development at which the sixth pe- reopods finally disappear (being absent on a specimen of 11.2 mm total length, but still present on another of about 12.9 mm total length); in both these cases, fringing spines are already present on the fifth oostegites. He also suggests (Rustad 1935:12) that the 28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON entire genus Holophryxus is possibly excep- tional among the Dajidae in having the oos- tegites developed before the sixth pereopods have completely disappeared. It is significant to note that Schultz (1978) considered the first pereonal segment to be completely fused with the cephalon in the latter three Jsophryxus species. Damage to host.—The holotype of H. acanthephyrae was collected and photo- graphed attached to the posterior surface of the carapace of its host, with its cephalon directed posteriorly (Stephensen 191 2a:fig. 13); one of the four specimens of the species reported by Jones & Smaldon (1986:fig. 1) was collected attached to its host in the same position. Three other species of Holo- phryxus (H. alaskensis, H. giardi, and H. richardi) have been found similarly at- tached to their hosts (Richardson 1908:figs. 2, 4; Stephensen 191 2a:fig. 13; Rustad 1935; Butler 1964, 1980:fig. 7B). The pereopods and mandibles of the isopod leave marks on the carapace of the host as the isopod grows, moves, and feeds (Stephensen 191 2a: figs. 14, 18). The mandibles perforate the carapace to the epithelial and connective tissues underneath, but the clawlike dactyls of the pereopods do not completely perfo- rate the carapace (Rustad 1935). Figure 2E shows the marks made by the pereopods and mandibles of the specimen of H. acanthephyrae collected attached to the specimen of Acanthephyra pelagica from Eltanin station 1723; Fig. 2D shows similar marks on the specimen of 4. pelagica from Eltanin station 1480, the presumed host to I. concavus. The holes left by the mandibles tend to be larger than those of the pereopods and surrounded by a large ring of darkly stained tissue. The marks present on both shrimps indicate that the isopods progres- sively moved posteriorly on the carapaces as the isopods increased in size. Coyle & Mueller (1981) provided com- plete descriptions of the larval and juvenile stages of Holophryxus alaskensis. The epi- carid stage of H. alaskensis uses stage V of the copepod Euchaeta elongata as an inter- mediate host, on which it metamorphoses through the microniscid stage to the cryp- toniscid stage, which then seeks out the shrimp final host (Pasiphaea pacifica). The cryptoniscid crawls beneath the carapace of the shrimp and attaches to the body wall in the branchial chamber above the gills, where it metamorphoses to the juvenile stage. It then moves out of the branchial chamber, up the side of the first abdominal somite and onto the dorsal surface of the carapace, where it attaches and assumes the mature adult form. Few details of the life cycle of H. acan- thephyrae are known, although Stephensen (1912a) described an epicarid larval stage from the marsupium of the holotype. From marks and damage (described below) to the carapaces of the two host specimens of A. pelagica at hand, and to the type host of H. acanthephyrae (which accompanied the ho- lotype when it was obtained on loan from the Universitetets Zoologiske Museum), I suggest that the cryptoniscid stage of H. acanthephyrae also attaches inside the bran- chial chamber of the host. The juvenile fe- male then apparently burrows through the branchial region of the carapace and gains access directly to the outer surface of the carapace, from whence it moves dorsally, begins to feed, and develops into the mature female. While I was examining the type host of H. acanthephyrae, I noticed an oblong, de- pressed scar on the posterior part of the right branchial region of the shrimp (Fig. 2A). The same area is visible in Stephensen’s (1912a:fig. 13) photograph as a darkened area, although over the years the darkened color has apparently become much less ev- ident. The two E/tanin specimens of A. pe- lagica which served as hosts for H. acan- thephyrae show evidence of damage to the right branchial region of their carapaces (Fig. 2B, C), as if something ate or burrowed through at these locations. The damage is contiguous with the posterior edge of the VOLUME 101, NUMBER 1 carapaces of these specimens and the edges of the wounds are rounded and partly healed, as if they are of some age. A small male specimen of 4. pelagica, carapace length 11.5 mm, from station 1204 of Eltanin cruise 14 has two relatively fresh, unhealed holes in the left branchial region of its carapace. Schultz (1977) recorded two small (2.2 and 2.5 mm long) female dajids from the same Eltanin station; the speci- mens were not given a formal species name at the time, but were identified simply as dajid species. Since the specimens bear some resemblance to the immature female of H. giardi described and illustrated by Richard- son (1908), I suggest that these small dajids may represent a species of Holophryxus, and that it is probable, based on the damage to the specimen of 4. pelagica from station 1204, that they represent juvenile females of H. acanthephyrae which had only re- cently burrowed from the branchial cham- ber of the host shrimp to the outside of the carapace, from where they were dislodged upon collection or subsequent handling. Distributions and hosts of Holophryxus species. —Eight species of Holophryxus are currently recognized. The final host species is known for only five of the eight, and the intermediate host is known for only one of these five. Each of the five species for which the final host is known apparently uses a single species of pelagic decapod shrimp as a host. It is expected that the distributions of the various Holophryxus species would therefore be limited to those of their single host shrimps. The currently recognized Holophryxus species, their distributions, and known hosts are as follows (in chronological order); 1. H. alaskensis Richardson, 1905. North- eastern North Pacific, Santa Barbara Channel, California to Prince William Sound, Alaska. Intermediate host, stage V of the copepod Euchaeta elongata; fi- nal host Pasiphaea pacifica Rathbun. 2. H. giardi Richardson, 1908. Northwest- 29 ern North Pacific, off Bering Island (54°48’N, 164°54’E). Final host, Ben- theogennema borealis (Rathbun). 3. H. richardi Koehler, 1911. Central North Atlantic (33°41'N, 36°55’W) north to Davis Strait west of Greenland (66°21'N, 57°04’W), in Denmark Strait between Iceland and Greenland, south of Iceland, to west coast of Norway, and to south of England. Final host, Sergestes arcticus Kroyer. 4. H. acanthephyrae Stephensen, 191 2a. North Atlantic from Davis Strait (60°07'N, 48°26'W) west of Greenland and to south of England, and subantarc- tic and antarctic waters of South Pacific. Final host, Acanthephyra pelagica (Ris- SO). 5. H. fusiformis Shiino, 1937. Northwest- ern North Pacific off east coast of Japan. Final host, Sergia prehensilis (Bate). 6. H. quadratohumerale (Schultz, 1978). Subantarctic waters of southeastern South Pacific. Final host not known. 7. H. polyandrus (Schultz, 1978). Subant- arctic waters of southeastern South Pa- cific. Final host not known. 8. H. septapodus (Schultz, 1978). Subant- arctic waters of southeastern South Pa- cific. Final host not known. Acknowledgments This work was supported in part by Con- tract PC-206882 from the Cooperative Sys- tematics Research Program of the Smith- sonian Oceanographic Sorting Center, funded by grant DPP-7920835 from the National Science Foundation, Division of Polar Programs, B. J. Landrum, Principal Investigator. Iam indebted to Torben Wolff of the Universitetets Zoologiske Museum, Copenhagen for the loan of the holotype of Holophryxus acanthephyrae and its host, and to Thomas E. Bowman of the National Museum of Natural History for the loan, in 1966, of the holotype of H. californiensis and the syntypes H. giardi, and for the more 30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON recent opportunity to examine the types of Isophryxus concavus, I. quadratohumerale, I. polyandrus, and I. septapodus, and to reexamine the syntypes of H. giardi at the National Museum of Natural History. I es- pecially thank Thomas E. Bowman, Mal- colm Jones, of Plymouth Polytechnic, and Anthony Futcher, of Columbia Union Col- lege, for reading an early draft of the manu- script. Literature Cited Butler, T. H. 1964. Redescription of the parasitic isopod Holophryxus alaskensis Richardson, and a note on its synonymy.—Journal of the Fish- eries Board of Canada 21(5):971-976. 1980. Shrimps of the Pacific coast of Cana- da.— Canadian Bulletin of Fisheries and Aquat- ic Sciences 202:1-280. Coyle, K. O., & G. J. Mueller. 1981. Larval and juvenile stages of the isopod Holophryxus alas- kensis (Epicarida, Dajidae), parasitic on deca- pods.—Canadian Journal of Fisheries and Aquatic Sciences 38(11):1438-1443. Jones, M. B., & G. Smaldon. 1986. On the genus Holophryxus Ysopods: Epicaridea), with de- scription of the male and redescription of the female of Holophryxus acanthephyrae. —Jour- nal of the Marine Biological Association of the United Kingdom 66(2):303-314. Koehler, R. 1911. Isopodes nouveaux de la famille des Dajides provenant des campagnes de la Princesse Alice. — Bulletin de |’ Institut Océanog- raphique de Monaco 196:1-34. Nierstrasz, H. F., & G. A. Brender a Brandis. 1931. Papers from Dr. Th. Mortensen’s Pacific Ex- pedition 1914-16. 57. Epicaridea II.— Viden- skabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn 91:147-226. Richardson, H. 1905. Isopods of the Alaska Salmon Investigation.— Bulletin of the United States Bureau of Fisheries 24:209-221. . 1908. On some isopods of the family Dajidae from the northwest Pacific Ocean, with descrip- tions of a new genus and two new species.— Proceedings of the United States National Mu- seum 33:689-696. Rustad, D. 1935. Notes on Holophryxus richardi Koehler (?) (Fam. Dajidae).— Bergens Museum Arbok 1934, Naturvidenskapelig Rekke 13:1- 31. Schultz, G. A. 1977. Bathypelagic isopod Crustacea from the Antarctic and Southern Seas. — Biology of the Antarctic Seas V, Antarctic Research Se- ries 23:69-128. 1978. More planktonic isopod crustaceans from Subantarctic and Antarctic Seas. — Biology of the Antarctic Seas VII, Antarctic Research Series 27:69-89. Shiino, S. M. 1937. Holophryxus fusiformis, a new species of Dajidae, Epicaridea.— Annotationes Zoologicae Japonenses 16(3):188-191. Stephensen, K. 1912a. Report on the Malacostraca collected by the ““Tjalfe’’-Expedition, under the direction of Cand. Mag. Ad. S. Jensen, especially at W. Greenland. — Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 1 Kjoben- havn 64:57-134. . 1912b. Corrections to the paper on the Mala- costraca from the “Tjalfe’’-Expedition.— Vi- denskabelige Meddelelser fra Dansk Naturhis- torisk Forening i Kjobenhavn 64:329-330. Wasmer, R. A. 1986. Pelagic shrimps of the fam- ily Oplophoridae (Crustacea: Decapoda) from the Pacific sector of the Southern Ocean: USNS Eltanin Cruises 10, 11, 14-16, 19-21, 24, and 25.—Biology of the Antarctic Seas XVII, Ant- arctic Research Series 44:29-68. Biology Department, Columbia Union College, Takoma Park, Maryland 20912. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 31-38 COPEPODA (CRUSTACEA) FROM A SEASONALLY FLOODED MARSH IN ROCK CREEK STREAM VALLEY PARK, MARYLAND Janet W. Reid Abstract. — Species of Copepoda (Crustacea) collected from a seasonally flooded marsh in Rock Creek Stream Valley Park, Maryland, included the cyclopoids Acanthocyclops vernalis (Fischer), Diacyclops bicuspidatus thomasi (S. A. Forbes), and Diacyclops palustris, new species, and the harpacticoid Attheyella (Mra- zekiella) americana (Herrick). The female of D. palustris is described; the species seems most closely related to D. haueri (Kiefer). Diacyclops clandestinus Yeatman is renamed D. yeatmani. A key to North American species of Dia- cyclops is presented. Some specimens of A. americana differ slightly from extant descriptions in details of the anal operculum and setation of the third swimming leg. Mr. William B. Yeaman of the National Park Service brought a sample of water and detritus from a marsh in Rock Creek Stream Valley Park, Maryland, to Dr. Thomas E. Bowman of the Department of Invertebrate Zoology for identification of the aquatic fau- na. The sample contained four species of Copepoda: Acanthocyclops vernalis (Fisch- er, 1853), 23 22, 3 66, 2 copepodites; Dia- cyclops bicuspidatus thomasi (S. A. Forbes, 1882), 1 6; Attheyella (Mrazekiella) amer- icana (Herrick, 1884), 7 92, 1 6; and 2 °° of a previously undescribed species of Diacy- clops. This species is described below as D. palustris. Variations of the population of A. americana from extant descriptions are also noted below. The collection was made on 23 Aug 1986 in a small marsh located in Rock Creek Stream Valley Park, Unit 3, Montgomery County, Maryland, on the west side of the park opposite Puller Drive, Kensington; about 39°01'41’N, 77°05'27”W. The marsh is shaded by broad-leaved deciduous hard- wood trees such as red maple (Acer rubrum) and black willow (Salix nigra). The area is in the floodplain of Rock Creek; surface water usually persists from November through May during years of normal or above normal rainfall. Standing water dis- appears in summer and early fall, although soils remain saturated. Any heavy rain which occurs during this period will briefly return the area to its standing water conditions. The first heavy rains following autumn leaf drop typically refill the wetland. The marsh is classified as a Palustrine System, Class FO (forested), Subclass 1 (broad-leaved de- ciduous), Water Regime E (seasonally flooded, saturated) (National Wetlands In- ventory, Fish and Wildlife Service, U.S. De- partment of the Interior; Cowardin et al. 1979). During the unusually dry summer of 1986, standing water was absent for long periods, though soils remained saturated; the collection was made after a rain. Family Cyclopidae Sars, 1913 Diacyclops palustris, new species Figs. 1-19 Material examined. —1 ?, holotype, dis- sected on | slide, National Museum of Nat- ural History—USNM 232186. 1 9°, para- type, dissected on | slide, USNM 232187. Female.—Length of holotype (excluding 32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Diacyclops palustris, new species, female holotype: 1, Habitus, dorsal, omitting pores on somites (both next innermost terminal setae on caudal rami broken); 2, Posterior 2 pedigers and genital segment, left lateral; 3, Urosome, dorsal; 4, Genital segment, ventral. Scale a, Fig. 1; Scale b, Figs. 2-4; scales = 100 um. caudal setae) 1140 wm; length of paratype 1130 wm. Posterior 2 pedigers (Figs. 1, 2) not markedly expanded laterally, each with small tubercle on posterolateral margin; posterior margins of urosomites with ser- rate hyaline membranes, except anal somite which has fine spinules along posteroventral margin. All somites (Figs. 2-4) with tiny pits; pattern of these pits different in holo- type and paratype. Genital segment (Figs. 2-4) expanded anteriorly, slightly longer than broad; seminal receptacle with poste- rior part little expanded. Caudal ramus (Fig. 3) 4.5-6 x longer than broad, with lateral row of 5 spinules about %4 distance from base of ramus. Lateral seta of ramus in- serted at distance from base of ramus equal to about 65% of length of ramus. Both lon- gest apical setae of holotype broken; lengths of setae of paratype as follows: lateral 40 VOLUME 101, NUMBER 1 33 Figs. 5-14. Diacyclops palustris, new species, female holotype: 5, Antennule, ventral; 6, Antennule, articles 1 and 2, dorsal; 7, Antenna, anterior; 8, Antenna, article 1, posterior; 9, Labrum; 10, Mandible; 11, Maxillula; 12, Maxilla; 13, Maxilliped, posterior; 14, Maxilliped, anterior. Scale = 100 um. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 101, NUMBER I 35 Table 1.—Proportions of some structures of Diacyclops palustris, new species, compared to those of D. haueri (Kiefer, 1931a). Abbreviations: CR, caudal ramus; P4 enp3, article 3 of endopod of leg 4; P5, leg 5. Structure D. palustris D. haueri CR, ratio of length : breadth 4.5-6:1 8-10:1 CR, ratio of length of next outermost terminal seta to length of outermost 5.0:1 5.81:1 terminal seta CR, ratio of next innermost terminal seta to length of outermost terminal seta 8.1:1 9.66:1 P4 enp3, ratio of length : breadth 2.26:1 2.53-3.0:1 P4 enp3, ratio of length of endopod to length of inner terminal spine 1.34:1 1.86:1 P5, ratio of length of terminal spine to length of article 2 1.7:1 um; dorsal 63 um; innermost to outermost apical setae 65, 420, 260 and 52 um re- spectively; thus innermost apical seta about 1.25x longer than outermost apical seta. All caudal setae finely plumose. Antennule (Figs. 5, 6) of 17 articles, reaching midlength of prosomite 2; with slender esthetascs on articles 12, 16 and 17; tew small pits on dorsal surface of article 1; and 2 terminal articles with narrow, smooth hyaline membrane. Antenna article | (Figs. 7, 8) with 3 groups of spinules on posterior surface, | group of 4 spinules on anterior surface, and 3 spinules near base on lateral margin; paratype with some pits on anterior surface of articles 3 and 4. Labrum and re- maining mouthparts as in Figs. 9-14. Swim- ming legs 1—4 (Figs. 15-17) with rami of 3 articles and spine formula 2,3,3,3. Leg 1, seta on medial expansion of basipod 2 reaching distal margin of endopod article 2. Legs 2 and 3 essentially similar. Basal la- mellae of all legs without armament, but with smooth protrusions each side, those of leg 1 most and those of leg 4 least developed. Terminal article of endopod of leg 4 2.26 x longer than broad; inner terminal spine 1.40 longer than outer terminal spine. Leg 5 (Fig. 18) of 2 free articles, inner —_ Figs. 15-21. Asi spine of article 2 1.7 longer than article, outer seta nearly 3 x longer than article. Left leg 5 of paratype (Fig. 19) with 3 subequal setae; right leg 5 as in holotype. Leg 6 (Figs. 2—4) consisting of 2 spinules and | seta. Etymology. —Named for the marsh hab- itat. Remarks. —Among North American species of Diacyclops, D. palustris most re- sembles D. haueri, with which it shares a slender leg 4 terminal endopod article hav- ing the inner terminal spine longer than the outer. However, comparison of the Rock Creek specimens with the original descrip- tions of D. haueri (Kiefer, 193la, b) and with specimens from a pond in Wooster, Ohio, collected by Mr. Andrew Weaver on 23 Feb 1948 and lent by Dr. Harry C. Yeat- man, revealed several differences. The cau- dal rami of D. palustris are relatively short, and there exist other differences in propor- tion between the two species (Table 1). The posterior expansion of the seminal recepta- cle of D. palustris is less developed than in D. haueri (Fig. 20). The anal operculum of D. haueri is unusually convex and strongly thickened (Kiefer 1931b:fig. 23). Setules of the maxilliped of D. palustris (Fig. 13) are sparse, while most setae of the maxilliped 15-18, Diacyclops palustris, new species, female holotype: 15, Leg 1; 16, Leg 2; 17, Leg 4 (setules of most setae of swimming legs omitted for clarity); 18, Leg 5. Fig. 19, D. palustris, female paratype, left leg 5. Figs. 20-21, Diacyclops haueri (Kiefer, 1931a), females from Wooster, Ohio: 20, genital segment, ventral; 21, maxilliped. Scale a, Figs. 19, 20; Scale b, Figs. 15-18, 21; scales = 100 um. 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of D. haueri bear many, closely set setules (Fig. 21). The seta on the medial expansion of the basipod 2 of leg 1 of D. haueri reaches the apex of endopod article 3. The inner terminal spine of endopod article 3 of leg 4 of D. palustris is straight, while that of D. haueri is bent outwards at midlength. Diacyclops palustris is the fifteenth species and subspecies of the genus recorded from North America (Dussart & Defaye 1985). Since the widely used keys of Yeatman (1958) and of Pennak (1978) include only 11 and 10 species respectively, it seems use- ful to include an updated key. The formu- lation of this key owes much to the exten- sive discussions of variations within North American populations of several members of the genus Diacyclops by Yeatman (1944). Dussart and Defaye (1985) suggested that in order to avoid confusion with D. /an- guidoides clandestinus (Kiefer, 1926), D. clandestinus Yeatman, 1964 be renamed. I concur and propose the new name D. yeat- mani, used below. The name of course hon- ors Dr. Harry C. Yeatman for his many contributions to copepod systematics. Key to Females of Species of Diacyclops Recorded from North America 1. Antennule with 11 articles — Antennule with 12 articles — Antennule with 16 or 17 articles 5 2. Lateral seta of caudal ramus in- serted at about midlength of ramus D. nanus (G. O. Sars, 1863) — Lateral seta of caudal ramus in- serted at distal *4to*4oframus 3 3. Leg 4 endopod 3 with outer ter- minal spine and inner terminal seta ... D. jeanneli putei (Yeatman, 1943) — Leg 4 endopod 3 with 2 terminal SPMINVCS srsie spec ac heen amare aa ... D. languidoides (Lilljeborg, 1901) 4. Caudal rami 4-5 longer than LOA) fic. ae 3092 ts od tera Meas RN ... D. crassicaudis (G. O. Sars, 1863) — Caudal rami 3.1-3.6 x longer than broad NO), ISIE We 13. . Antennule of 17 articles D. crassicaudis brachycercus (Kiefer, 1927) Antennule of 16 articles Fe ee D. languidus (G. O. Sars, 1863) . Leg 4 endopod 3, outer terminal spine longer than inner terminal SPINE OL Stas secur: Us eee 7 Leg 4 endopod 3, inner terminal spine or seta longer than outer ter- Mmunalespine-. 2.4 seoe oe ee 9 Seta of article 2 of leg 5 about 2 x length Of spine wae .em Pee 8 Seta and spine of article 2 of leg 5 subequal .. D. navus (Herrick, 1882) Outer terminal spine of endopod 3 of leg 4 about 1.5 length of inner terminal spine; endopod 3 of leg 4 2-3 x longer than broad ... Ae ea D. bicuspidatus (Claus, 1857) Outer terminal spine of endopod 3 of leg 4 about 2 x length of inner terminal spine; endopod 3 of leg 4 3-4.2 x longer than broad D. bicuspidatus thomasi (S. A. Forbes, 1882) . Leg 4 endopod 3, outer margin with seta Leg 4 endopod 3, outer margin with spine ....D. nearcticus (Kiefer, 1934) Leg 4 endopod 3 with 2 terminal spines Leg 4 endopod 3 with outer spine and inner seta 2 Teel D. jeanneli (Chappuis, 1929) Leg 4 endopod 3 stout, about 1.5- 1.7x longer than broad ........ WZ Leg 4 endopod 3 slender, about 2.2—-2.5x longer than broad .... 13 Innermost terminal caudal seta shorter than outermost; caudal rami 5—7 x longer than broad ... eget a D. bisetosus (Rehberg, 1880) Innermost terminal caudal seta longer than outermost; caudal rami 3-—4.5 x longer than broad EN Ra D. yeatmani, new name Caudal rami 8-10 longer than VOLUME 101, NUMBER 1 broad; leg 4 endopod 3, length of endopod about 1.9 x length of in- ner terminal spine Sate eNO ME Te D. haueri (Kiefer, 193 1a) — Caudal rami 4.5-6x longer than broad; leg 4 endopod 3, length of endopod about 1.3 x length of in- ner terminal spine eR SPs ait Es Laaekes D. palustris, new species Cee Family Canthocamptidae Sars, 1906 Attheyella (Mrazekiella) americana (Herrick, 1884) Material examined.—1 2, dissected on 2 slides; 5 2°, 2 66, alcohol-preserved; USNM 232188. Remarks. —The Rock Creek Park popu- lation varies little from those from North Carolina and Wisconsin, exhaustively de- scribed by Coker (1934). Two Rock Creek females bear two setae rather than one on the inner margin of leg 3 exopod article 2; Coker noted similar variability in the se- tation of endopod article 2 of this leg in females. The anal opercula of Rock Creek females bear 10-15 teeth, while those of the two males bear 12 and 14 teeth; Coker men- tioned variations in this character also, his specimens from North Carolina having 12- 18 “‘spinules”, and those from Wisconsin having 20-24 “‘spinules”’. Acknowledgments For his gift of specimens and description of the habitat, I thank Mr. William B. Yea- man of the National Park Service; my thanks also go to Dr. Harry C. Yeatman for his kindness in lending specimens and provid- ing information on variations in Diacyclops haueri. Literature Cited Chappuis, P. A. 1929. Copépodes cavernicoles de Amérique du Nord (note préliminaire). — Bul- letin de la Société des Sciences de Cluj 4:51—57. 37 Claus, C. 1857. Weitere Mitteilungen tiber die ein- heimischen Cyclopiden.—Archiv fiir Natur- geschichten 23:205—211, pl. 11. Coker, R. E. 1934. Contribution to knowledge of North American freshwater harpacticoid cope- pod Crustacea. — Journal of the Elisha Mitchell Scientific Society 50:75-141, pls. 1-15. Cowardin, L. M., V. Carter, F. C. Golet, & E. T. LaRoe. 1979. Classification of wetlands and deepwater habitats of the United States.— Biological Ser- vices Program, Fish and Wildlife Service, U.S. Department of the Interior FWS/OBS-79/31. v + 103 pp. Dussart, B. H., & D. Defaye. 1985. Répertoire mon- dial des copépodes cyclopoides. Editions du C. N. R. S., Paris. 236 pp. Fischer, S. 1853. Beitraége zur Kenntniss der in der Umgegend von St.-Petersburg sich findenden Cyklopiden (Fortsetzung).— Bulletin de la So- ciété Impériale des Naturalistes de Moscou 26: 74-100, pls. I, II. Forbes, S. A. 1882. On some Entomostraca of Lake Michigan and adjacent waters. — American Nat- uralist 16:640—650, pl. IX. Herrick, C. L. 1882. Papers on the Crustacea of the fresh-waters of Minnesota. I. Cyclopidae of Minnesota with notes on other Copepoda. — Re- ports of the Geological and Natural History Sur- vey of Minnesota 10:219-—233, pls. I-VII. . 1884. A final report on the Crustacea of Min- nesota included in the Orders Cladocera and Copepoda. Geological and Natural History Sur- vey of Minnesota. Johnson, Smith & Harrison, Minneapolis. 191 pp., 22 pls. Kiefer, F. 1926. Uber einige Krebse aus der Wasser- leitung von Ofingen.—Schriften des Vereins fiir die Geschichte und Naturgeschichte der Baar und der angrenzenden Landsteile in Donaue- schingen 16:273-283. . 1927. Freilebende Siisswasser-Copepoden aus Nordamerika. — Zoologischer Anzeiger 72:262- 268. 193la. Kurze Diagnosen neuer Siisswasser- Copepoden.— Zoologischer Anzeiger 94:219- 224. 1931b. Zur Kenntnis der freilebenden Siiss- wassercopepoden, insbesondere der Cyclopiden Nordamerikas. — Zoologische Jahrbuch flr Sys- tematik 61:579-620. 1934. Neue Ruderfusskrebse aus Nordamer- ika.— Zoologischer Anzeiger 107:269-271. Lilljeborg, W. 1901. Synopsis specierum huc usque in Suecia observatorum generis Cyclopis.— Kongliga Svenska Vetenskapsakademiens Handlingar 35:1-118, pls. 1-6. Pennak, R. W. 1978. Fresh-water invertebrates of the United States. Second Edition. John Wiley & Sons, New York. 803 pp. 38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rehberg, H. 1880. Beitrag zur Kenniniss der freile- benden Stisswasser-Copepoden. — Abhandlung- en herausgegeben vom Naturwissenschaftlichen Verein zu Bremen 6:533-554, pl. VI. Sar, G. O. 1863. Oversigt af de indenlandske Fersk- vandscopepoder, af Stud.—Forhandlinger, Vi- denskabs-Selskabet I Christiana, Aar 1862:212— 262. Yeatman, H.C. 1943. Rare cyclopoid copepods from wells in North Carolina.—Journal of the Elisha Mitchell Scientific Society 59:27-36. 1944. American cyclopoid copepods of the viridis-vernalis group, (including a description of Cyclops carolinianus n. sp.).— American Midland Naturalist 32:1—90. . 1958. Cyclopoida. Pp. 795-815 in W. T. Ed- mondson, ed., Fresh-water biology, second edi- tion, John Wiley and Sons, Inc., New York. 1964. A new cavernicolous cyclopoid cope- pod from Tennessee and IIlinois.—Journal of the Tennessee Academy of Science 39:95-98. NHB-163, Department of Invertebrate Zoology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 39-59 TANAIDACEA (CRUSTACEA: PERACARIDA) OF THE GULF OF MEXICO. V. THE FAMILY PSEUDOTANAIDAE FROM LESS THAN 200 METERS, WITH THE DESCRIPTION OF PSEUDOTANAIS MEXIKOLPOS, N. SP. AND A KEY TO THE KNOWN GENERA AND SPECIES OF THE WORLD Jurgen Sieg and Richard W. Heard Abstract.—No published records exist for pseudotanaid tanaidaceans in the Gulf of Mexico. In this study three species, /ungentitanais primitivus (Sieg, 1973), Pseudotanais mortenseni Sieg, 1973, and P. mexikolpos, n. sp. are re- ported from the Gulf or immediately adjacent waters of the Florida Keys. TIungentitanais primitivus and P. mortenseni were previously known only from their type localities, both at San Thomas Harbor in the West Indies. Pseudo- tanais mexikolpos is described from a single location in the West Flower Garden Banks off Texas. It most closely resembles two Mediterranean species, P. medi- terraneus G. O. Sars, 1882, and P. unicus Sieg, 1977. It can be separated from P. mediterraneus by having a much longer disto-sternal seta on the propodus of peraeopods 4—6 and from P. unicus by its shorter cephalothorax and distinctly larger peraeopod 3. A key to the five known genera and 25 described species of the family is included and taxonomic problems concerning several species are briefly discussed. There are no published records of the tan- aidacean family Pseudotanaidae for the Gulf of Mexico. Of the 24 described species only three, Jungentitanais primitivus (Sieg, 1973), Pseudotanais kurchatovi Kudinova-Paster- nak & Pasternak, 1978, and P. mortenseni Sieg, 1973, have been reported from the Caribbean region. During the past 10 years, three species of pseudotanaids were col- lected from less than 200 m in conjunction with several environmental baseline studies conducted in the Gulf of Mexico and the immediately adjacent waters of the Florida Keys. These specimens were made available to us for study and are the subject of this report. The family Pseudotanaidae was original- ly established by Sieg (1973, 1977) for those species having a marsupium formed by a single pair of sheet-like oostegites. As in all other tanaidaceans except the Tanaidae, the eggs develop between the oostegites. The Tanaidae are the only other family with one pair of oostegites. This group is unique, however, in having the oostegites trans- formed into sack-like structures, termed ““ovisacs’” by Lang (1960), in which the eggs are incubated (see Sieg 1980, 1984; Johnson & Attramadal 1982). Other characters of the Pseudotanaidae given by Sieg (1977) were the: (1) reduced size of the first two peraeonites, (2) maxilliped with basis com- pletely fused, (3) first antenna of females with 3 or 4 segments, and (4) mandibles with pars molaris reduced. Males are known only for the genus Pseu- dotanais G. O. Sars, 1882. They are rarely collected and are distinctly dimorphic, with 40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a body shape similar to the males of Lep- tognathia and Typhlotanais. The pleon is strongly enlarged and distinctly longer than one-third of the total body length (rostrum to tip of telson); whereas in females the pleon is approximately one-fifth of the total body length (Fig. 1: Pseudotanais). Each pleonite contains large bundles of dorso-ventral musculature to move the strongly devel- oped pleopods which are characteristic of all males, even for those species in which the females lack pleopods. The first antenna has 7 segments with the first 2 dorsoven- trally depressed and probably functioning as a unit. The third peduncle segment is distinctly smaller and movable. The four flagellar segments each bear aesthetascs dis- tally (segments 1-3 with a cluster of aesthe- tascs and segment 4 with a single terminal aesthetasc). The second antenna is like that of the female. The mouthparts are lacking except for the maxilliped, which is slightly smaller than that of the female. Unlike the female the lateral margins of the maxilliped are fused with the carapace fold. The che- lipeds are slightly larger, and the peraeopods and uropods appear more slender than in the female. In a monograph of the family, Sieg (1977) grouped the known species into two subfamilies, the Cryptocopinae and the Pseudotanainae. The Cryptocopinae is rep- resented by four genera: Cryptocope G. O. Sars, 1882; Cryptocopoides Sieg, 1977; Iun- gentitanais Sieg, 1977; and Paraiungenti- tanais Sieg, 1977. The subfamily Pseudo- tanainae has only the type genus, but contains most of the family’s described species. Sieg (1977) also divided the genus Pseudotanais into the subgenera Pseudota- nais, s.s. and Akanthinotanais Sieg, 1977. Since 1977 three species of Pseudotanais, two species of Cryptocopoides, and two species of Cryptocope have been described: Pseudotanais guillei Shiino, 1978, from the Kerguelen Islands; P. kurchatovi Kudinova- Pasternak & Pasternak, 1978, from deep water in the Caribbean, and P. siegi Kudi- nova-Pasternak, 1985, from the North At- lantic; Cryptocopoides rostralis Tzareva, 1982, which was transferred to the genus Typhlotanoides Sieg, 1983 (Sieg 1986a:102) and Cryptocopoides rotundata Tzareva, 1982 (which was synonymized with the neuter of Mirandotanais vorax Kussakin & Tzareva, 1974 by Sieg 1986a:138-139). Within Cryptocope Kudinova-Pasternak (1982) described C. longa and C. vitjazi, both from the deep waters of the Mediter- ranean Sea. Finally, Kudinova-Pasternak & Pasternak (1978:187-188) mentioned a specimen of Cryptocope sp. from deep water in the Caribbean Sea, but with reservations they later reassigned it to the genus Lepto- gnathia G. O. Sars, 1882 (Kudinova-Pas- ternak & Pasternak 1981:117). A key to the known genera and all the described species of the family is presented here. For bibliographic, taxonomic, and zoogeographic information see Sieg (1983). For terminology refer to Sieg (1977, 1980). The following abbreviations are used in the key and figures: A.1 = first antenna, A.2 = second antenna, L = labrum, Md(r) = right mandible, Md(l) = left mandible, Mx.1 = first maxilla, Mx.2 = second max- illa, La = labium (paragnaths), Mxp = max- illiped, Epi = epignath (=maxillipedal epi- podite), Che = cheliped, P.1-P.6 = peraeopod | to peraeopod 6, Pl.1—P1.5 = pleopod 1 to pleopod 5, Plt = pleotelson, and Uro = uropod. Key to the Genera and Species of Pseudotanaidae 1. A.1 7-segmented (Fig. 1: Pseudo- LONRQIS) Ae FR ee eel ee males — A.14- or 3-segmented ......... 2 2. A.1 4-segmented (Fig. 1: subfam- ily. Cry. ptoOcopinac) mena aaa 3 — A.1 3-segmented (Fig. 1: subfam- ily Pseudotanainae) ............ 6 3. Pleopods reduced, without setae (Fig. 1: Pl.3); Mx.1 with 9 terminal Spines) .4e Cryptocope abbreviata (G. O. Sars, 1868) VOLUME 101, NUMBER 1 Pleopods well developed and with setae; Mx.1 with 5 short or 9 longer terminal spines . Cheliped slender, with carpus and propodus elongate (Fig. 1: Che.1, Che.2); A.1 elongate; pleopodal endopodite with one distal inner seta and exopodite with pilose proximal seta (Fig. 1: Pl.1) Cheliped (Fig. 1: Che.3) and A.1 stout; pleopodal exopodite and en- dopodite each only with some dis- tal outer setae (Fig. 1: Pl.2) 2 Ea AN Cryptocopoides arcticus (Hansen, 1913) . Without eyes (Fig. 1); cheliped with carpus 4 times as long as broad (Fig. 1: Che.2); Mx.1 with 9 well developed terminal spines .... Paraiungentitanais longidigitatus (Kudinova-Pasternak, 1975) With eyes (Fig. 1); cheliped with carpus 5 times as long as broad (Fig. 1: Che.1); Mx.1 with 5 small terminal spines Iungentitanais primitivus (Sieg, 1973) . Carpal spines of P.2—P.6 of typical shape (Fig. 1: 1 subgenus Akan- thinotanais) Carpus of P.2—P.6 with 1 trans- formed blade-like spine (Fig. 1: 2 subgenus Pseudotanais s. str.) ... 14 . With eyes; pars molaris always POUCH ee We Mate Li hiee, 8 Without eyes; pars molaris point- ed or ending in several blunt pro- cesses . Merus of P.4—P.6 with 2 short spine-like setae of equal size .... 9 Merus of P.4—P.6 with 1 short and 1 long spine-like seta . Uropodal exopodite nearly as long as endopodite _... Pseudotanais gerlachi Sieg, 1977 Uropodal exopodite only slightly longer than first segment of endo- 11. 1S 41 Pseudotanais malayensis Sieg, 1977 podite . Cheliped with carpus elongate (at least 2.5 times longer than broad) and elongate chela (Fig. 1: 4); is- chium of P.3—P.6 with | long seta Carpus of cheliped stout (less than twice as long as broad) and chela of typical shape (Fig. 1: 3); ischium of P.3-P.6 with short seta ...... _...Pseudotanais guillei Shiino, 1978 Peraeonites 4 and 5 relatively small, broader than long, lateral MAneinS CONVekeee een ee Pseudotanais mortenseni Sieg, 1977 Peraeonites 4 and 5 large, at least as long as broad, lateral margins straighter, with a protuberance at articulation of peraeopods 2 hhaaiabeti in seus pment ae Pseudotanais siegi Kudinova-Pasternak, 1985 Cheliped with carpus elongate (about 3.5 times longer than broad) and elongate chela (Fig. 1: 4); pars molaris with several blunt pro- cesses (Fig. 1: 9) 1 SEALE E Chuan atu Pseudotanais longipes Hansen, 1913 Carpus of cheliped stout (less than twice as long as broad) and of typ- ical shape (Fig. 1: 3); pars molaris pointed: (Figs 1slD) ieee ane 13 Sternal border of propodus in P.4— P.6 with 2 distal spine-like setae, longer than claw; propodus 3 times as long as claw .. Pseudotanais gaussi Vanhoffen, 1914 Sternal border of propodus in P.4— P.6 with 1 distal spine-like seta, shorter than claw; propodus twice as long as claw Pseudotanais similis Sieg, 1977 . With eyes; pars molaris ending in blunt processes (Fig. 1: 10) or sim- ple point (Fig. 1: 12) Without eyes; pars molaris of vari- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 42 sjeue,opnesd snyediosoj *q luasuayjoW “gq SISUBAIPE|PW “gq snyedioso} “qd 4js's sleuejopnasd a sejayooioew “qd snjejnoo “q sedipuo| “dq a SIBPUBJOUIWJUeYY IpjOlySuepsou “q snsojesi6uo| “d SIUIJJe “d 8 Z z Sliejow sied aePuleUuUBeBUOPNSAS, seplodov0,dhin NS Seurdoo00};dAl9O VOLUME 101, NUMBER 1 15. 17. 18. 20. — Fig. 1. able shape (broad with several spine-like setae; thin with several pointed or blunt processes; bifur- cate or simple point (Fig. 1: 6-8, 11-12) Pars molaris ending in several blunt spine-like processes (Fig. 1: LO) AE ALAR! Pseudotanais oculatus Hansen, 1913 (syn. Paratanais nanaimoensis Fee, 1927) Pars molaris pointed (Fig. 1: 12) 16 . Cephalothorax longer than broad; P.3 distinctly smaller than P.2 .. Nena Pseudotanais unicus Sieg, 1977 Cephalothorax broader than long, P.3 as large as P.2 Propodus of P.4—P.6 distally with one long sternal seta, distinctly longer than claw ae Pseudotanais mexikolpos, n. sp. Propodus of P.4—P.6 with one ster- nal distal seta, about as long as claw SO resiuehi is Pseudotanais mediterraneus G. O. Sars, 1882 20 Pleopods present Pleopods absent . Chela with a gap between fixed fin- ger and articulation of dactylus (Fig. 1: 5); pars molaris pointed (Fig. 1: 12) ..Pseudotanais forcipatus (Lilljeborg, 1864) Cheliped without gap between fixed finger and articulation of dactylus (Fig. 1:3); pars molaris with 3 blunt processes .... Pseudotanais lilljeborgi G. O. Sars, 1882 Chela with a gap between fixed fin- ger and articulation of dactylus (Fig. 1: 5); pars molaris pointed (Fig. 1: 12) Chela without gap between fixed Die MD AB. 24. DS 43 finger and articulation of dactylus (Fig. 1:3); pars molaris not pointed 22 Exopodite of uropod at most as long as first segment of endopo- dite; endite of maxilliped com- pletely fused medially, without notch or setae ... Pseudotanais jonesi Sieg, 1977 Exopodite of uropod distincly longer than first segment of en- dopodite; endite of maxilliped with 1 pair of small setae on each side of the disto-medial notch ... Pseudotanais abyssi Hansen, 1913 Pars molaris bifid (Fig. 1: 11) ... Pseudotanais macrocheles G. O. Sars, 1882 Pars molaris broad or tapered, but always with several pointed pro- cesses (Fig. 1: 6-8) Pars molaris broad, with 3 small distal setae in addition to pointed processes (Fig. 1: 6) _.. Pseudotanais affinis (Hansen, 1887) Pars molaris tapered, without dis- tal setae Endopodite and exopodite of uro- pod nearly equal in length (pars molaris tapered and with several pointed processes of nearly equal SIZE) M5 et eie Pseudotanais vitjazi Kudinova-Pasternak, 1966 Exopodite of uropod distinctly shorter than endopodite Pars molaris with 1 process dis- tinctly longer than others (Fig. 1: 7); carpus of P.4—P.6 with | long sternal seta (as long as propodus) Pseudotanais longisetosus Sieg, 1977 Pars molaris with all processes of nearly equal size (Fig. 1: 8); carpus of P.4—P.6 with short sternal seta 75) Major characteristics of the different pseudotanaid subfamiliar taxa. 44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pseudotanais nordenskioldi Siegeal9i7/7/ Remarks. —Two species recently referred to the genus Cryptocope, C. longa Kudi- nova-Pasternak, 1982, and C. vitjazi Ku- dinova-Pasternak, 1982, are not included in the key, since they both appear to be most closely related to Leptognathia (Leptogna- thiinae). Both species differ distinctly from the described species of Cryptocope in the overall shape of the body and chela, the morphology of the pars incisiva, and the setation of the pleopods. The pars incisiva in Cryptocope is deeply incised while those of Leptognathia spp., C. longa and C. vitjazi are only slightly crenulated (see Sieg 1986b; Kudinova-Pasternak 1982:figs. 3, 4). The pleopodal exopodite and endopodite of Cryptocope only have distal setae (see Sieg 1977), whereas those of C. /onga and C. vitjazi like Leptognathia spp. also each bear one proximal seta (see Kudinova-Pasternak 1982:figs. 3, 4). Based on these observa- tions, we tentatively transfer C. Jonga and C. vitjazi to the genus Leptognathia, as di- agnosed by Sieg (1986b). Pseudotanais borceai Baéescu, 1960, is not included in the key because of apparent in- consistencies in the original and only de- scription. The type material has been lost (Bacescu, pers. comm.), and at present it is not possible to determine its taxonomic sta- tus. Additional specimens from the type lo- cality are needed to clarify this problem. We also excluded P. kurchatovi Kudi- nova-Pasternak & Pasternak, 1978 from the key because it does not belong to Pseudo- tanais. Our reasons are based on the fol- lowing criteria. In their original description Kudinova-Pasternak & Pasternak (1978: 188-190, fig. 5) described and illustrated the antenna 1 with the second segment short and annular as well as the third segment being distinctly longer. In all other known species of Pseudotanais the second segment is nearly as long as the third. Other mor- ohological differences that make the sys- tematic position of P. kurchatovi uncertain are: (1) the shape of the cheliped (especially that of the chela), (2) the proportion of pe- raeonites (peraeonites | and 2 atypically long and peraeonites 4 and 5 atypically broad), (3) the shape of dactylus and its terminal spine on P.4—P.6 (in all other species of Pseudotanais a short claw is present while in P. kurchatovi the dactylus and spine are unfused with a combined length nearly equal to that of propodus), and (4) the shape of the pars molaris 1s more reminiscent of species belonging to the Typhlotanaidae rather than to the genus Pseudotanais. It also should be mentioned that at least in P. nordenskioldi Sieg, 1977, P. longise- tosus Sieg, 1977, and P. oculatus Hansen, 1913, the figures of the pars molaris given by Sieg (1977) are misleading, since there are no distinct spines or setae as illustrated (see Figs. 43, 46, 49). In these species the pars molaris ends in several spine-like teeth, not spines or setae. Subfamily Cryptocopinae Sieg, 1977 Iungentitanais primitivus (Sieg, 1973) Figs. 2-4, 12 Material. — FLORIDA MIDDLE GROUNDS: Sta 151, 28°32'13’N, 84°18'40’W, 25-28 Jun 1979, 31.3—33.0 m, coral reef, dive, 1 female, Marine Science Consortium (MESC) Cat. No. 6157- 10495.—Same station, 15-19 Oct 1978, 1 female, MESC Cat. No. 6157-10499 and 2 females, Cat. No. 6157-10500.—Sta 481, 29°30'52”"N, 84°18'59"W, 6-8 Oct 1978, 28.5—33.0 m, coral reef, dive, 1 female, MESC Cat. No. 6157-10496.—Same sta- tion, 14-18 Oct 1978, 1 female, MESC Cat. No. 6157-10498.—Sta 247, 28°32'16’N, 84°18'36’W, 26-30 Jan 1979, 31.3-33.0 m, coral reef, dive/grab, 1 female, MESC 281261. FLORIDA KEYS: Pumpkin Creek, Key Largo, 25°19’N, 80°16’W, no depth infor- mation, 2 Jun 1981, 2 females, Invertebrate Zoology Collection of the Gulf Coast Re- Fig. 2. Iungentitanais primitivus (Sieg, 1973), female. 46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.03 mm Fig. 3. Jungentitanais primitivus (Sieg, 1973), female. VOLUME 101, NUMBER 1 Fig. 4. Jungentitanais primitivus (Sieg, 1973), female. 47 48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON search Laboratory (GCRL) Museum, Cat. No. I 86-1127.—Shark Channel, between lower Saddlebunch Keys and Big Coppitt Key, no depth information, 5 Jan 1982, 24°36'’N, 81°39'W, 2 females dissected in Sieg Coll. and 1 female (lost). Remarks. —Tungentitanais primitivus 1s characterized by a slender cheliped and a 4-segmented antenna 1. The presence of well developed eye-lobes containing clearly pig- mented visual elements distinguishes the species from its closest relative Paraiungen- titanais longigigitatus (Kudinova-Paster- nak, 1975). The mouthparts of two specimens were dissected to supplement the original de- scription. As noted by Sieg (1977:12) both mandibles of the holotype, the only speci- men then available, were lost during dis- section. The mandibles (Fig. 3) do not differ markedly from those of Paraiungentitanais longidigitatus. Each mandible is well de- veloped and has a broad pars molaris. The surrounding wall of the crushing area is slightly indented and has 1 tooth-like pro- cess. The distal margin of the pars incisiva of the right mandible is crenulated. The tip is bifid, and the lower tooth probably rep- resents the fused lacinia mobilis. On the left mandible the lacinia mobilis is small and not fused with the pars incisiva. Distribution (Fig. 12).—The holotype was collected from the harbor of Saint Thomas, West Indies, at a depth of 18.2—27.3 m. Our records from the Florida Keys and Florida Middle Grounds indicate that the species may have a broad distribution throughout the tropical waters of the Gulf of Mexico and the Caribbean (about 18-33 m). Pseudotanais G. O. Sars, 1882 Remarks.—In our material each of the two subgenera established by Sieg (1977) is represented by a single species. The sub- genus Akanthinotanais is characterized by having long meral spines on P.4—P.6 while Pseudotanais, s.s. has one transformed blade-like spine. Pseudotanais mortenseni Sieg, 1977 belongs to the former subgenus and Pseudotanais mexikolpos, n. sp. to the latter. Pseudotanais (Akanthinotanais) mortenseni Sieg, 1977 Figs. 5-7, 12 Material.—FLORIDA KEYS: Pumpkin Creek, Key Largo, 25°19'N, 80°16'W, no depth information, 2 Jun 1981, 1 female, dissected in Sieg Coll.—Shark Channel, be- tween lower Saddlebunch Keys and Big Coppitt Key, 24°36'N, 81°39’W, no depth information, | Jan 1982, 1 female, dissected in Sieg Coll. and 1 female (lost). Remarks. —Pseudotanais mortenseni is distinguished from the other members of the subgenus Akanthinotanais by the fol- lowing combination of characters: (1) pres- ence of eyes, (2) a pointed pars molaris, (3) the shape of the cheliped, and (4) the ar- mament of the merus (see key and Figs. 5— i): Distribution (Fig. 12).—This is the first report of this species since its original de- scription from the harbor of Saint Thomas, West Indies. Like Jungentitanais primitivus (Sieg, 1973), it may have a broad distri- bution in the tropical waters of the Gulf of Mexico and the Caribbean. Pseudotanais (Pseudotanais) mexikolpos, new species Figs. 8-11, 12 Synonymy. —cf. Paratanais sp. A., Anon- ymous (Texas A&M University), 1978:772. Material. — Holotype: 1 female, National Museum of Natural History, USNM 231765; off Texas coast, East Flower Gar- den Bank, 72 m, Sta 902, 27°54'36.64’N, 93°32'53.27”W. Paratypes: | females, dis- sected in Sieg Coll., Sta 899-4-SED, | neu- ter, Sta 896-6b-SED, Invertebrate Zoology Collection of the Gulf Coast Research Lab- oratory Museum Cat. No.; 1 female, Sta Fig. 5. Pseudotanais mortenseni Sieg, 1977. 50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.05 mm Fig. 6. Pseudotanais mortenseni Sieg, 1977. 52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 896-3-4, USNM 231766; | neuter, Sta 897- 8-3, USNM 231768; 1 neuter, Sta 896-6b- 4, USNM 231769: all from type locality. Additional material. —2 females of Texas coast, Stetson Bank, 28°09'N, 94°16'W, lost in slide preparation. Type locality.—East Flower Garden Bank, off Texas coast, 27°54'36.64’N, 93°34'53.27”W, 72 m. Etymology. —The first part of the name is taken from Mexico while the second part is of Greek origin (kolpos = bosom). Description of female (paratype). — Length of adult female reaching about 1.5 mm, about 4.2 times longer than broad (Fig. 8). Cephalothorax: Short, 1.1 times broader than long; eye-lobes well developed and with visual elements; rostral area convex, ante- rior 14 smallest, broadest posteriorly and corners rounded; no setae. Peraeonites: Lateral margins of all perae- onites rounded in dorsal view, bearing no setae, spines, or peaks. First to third pe- raeonites small, first 6.9 times, second 5.5 times, and third 3.5 times broader than long; fourth and fifth peraeonites largest, fourth only 1.7 times and fifth 1.8 times broader than long; sixth peraeonite broadest poste- riorly, margins strongly convex, about 2.3 times broader than long. Antenna | (Fig. 8): Three-segmented. First segment 3.9 times longer than broad, with 1 seta at midlength and one seta, | setule, and 3 feathered hairs distally. Second 2.3 times longer than broad, with 2 distal setae. Third segment elongate, nearly 3.9 times longer than broad, tip with | aesthetasc, 2 normal setae, and 4 setae having bifid tips. Antenna 2 (Fig. 8): Six-segmented. First segment small, partly fused with cephalo- thorax, and unarmed. Second stout, as long as broad, outer border with | spine-like dis- tal seta. Third segment also short, but 1.5 Fig. 8. times longer than broad, outer border with 1 spine-like distal seta. Fourth segment elongate, bent sternally, 4.6 times longer than broad, with 3 setae and | feathered hair distally. Fifth nearly 2.9 times longer than broad, | distal seta. Sixth segment very small, conical, with 2 short and 3 long setae. Labium (Fig. 9): Hood-shaped, covered with fine setules. Mandibles (Fig. 9): Not strongly calcified, but well developed and of typical shape. Pars molaris reduced, small, thin, with pointed tip. Left mandible with slightly crenulated pars incisiva, lacinia mobilis well developed, with 1 large and 4 small teeth. Right mandible with crenulated pars inci- siva, lacinia mobilis fused to pars incisiva, represented only by well-developed edge. Labium (Fig. 9): Consisting of 1 lobe, out- er lobe totally lacking; inner lobe deeply in- cised at middle. Maxilla 1 (Fig. 9): Endite bearing circle of 9 spines. One-segmented palp as long as endite, with 2 terminal setae. Maxilla 2 (Fig. 9): Of typical shape, oval, lacking setae. Maxilliped (Fig. 9): Well developed, with- out coxae. Basis fused medially, with a small seta near articulation of palpus; inner lobe of normal size, totally fused, distal margin undulate (wave-like), with 2 rows of 4 set- ules. Palpus 4-segmented; distal margin of first segment slanted towards inner border, trapezoidal, 1.3 times longer than broad, unarmed; second segment also trapezoidal, but inner border longer than outer, inner border with 2 setae; third segment elongate, about 2.4 times longer than broad, inner border with 3 strong setae. Fourth about 2.6 times longer than broad, outer border with 1 seta, inner border with 4 strong setae. Epignath (Fig. 9): Of typical shape; elon- gate, tip rounded and unarmed. Pseudotanais mexikolpos, n. sp., female, paratype. 54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON L Md; Md \ Epi Mx. 2 Fig. 9. Pseudotanais mexikolpos, n. sp., female, paratype. 56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 11. Cheliped (Fig. 8): Well developed, of typ- ical shape, chela with small gap. Side-piece of normal size, articulating with basis be- hind a distal conjunction. Basis stout, about 1.6 times longer than broad, unarmed. Me- rus small, triangular, with | midsternal seta. Carpus relatively short, 1.6 times longer than broad, tergal border with | small proximal and | small distal seta; sternal border with 1 short and | long seta at midlength. Propo- dus with fixed finger elongate, nearly 3 times longer than broad, with ““comb”’ at base of dactylus consisting of five small setae, with 2 groups of setules nearby; fixed finger with 3 tergal setae, tip not markedly sclerotized, sternal border with | small seta; with 1 small seta close to articulation of dactylus. Dac- tylus straight, with | small proximal seta. Peraeopod | (Fig. 10): Slender; coxa not fused with peraeonite, with 1 seta. Basis 5.4 times longer than broad, unarmed. Ischium annular, with | tergal seta. Merus nearly 1.5 times longer than broad, with 1 tergal setule. Carpus 2.4 times longer than broad, with 1 distal rostral seta. Propodus elongate, five times longer than broad, with 1 distal tergal setule. Dactylus and terminal spine not fused, combined length nearly equal to that of propodus. Peraeopod 2 (Fig. 10): Shorter than P.1; Pit Pseudotanais mexikolpos, n. sp., female, paratype. coxa not fused with peraeonite, having 1 seta. Basis slightly bent sternally, 4.6 times longer than broad, sternal border with 1 feathered hair at midlength. Ischium an- nular, with 1 tergal setule. Merus slightly over 2 times longer than broad, tergal bor- der with | rostral and 1 caudal seta distally. Carpus shorter, 1.8 times longer than broad, tergal border with | strong blade-like spine reaching about 0.6 times length of propo- dus, with 1 additional caudal seta. Propodus about 4.6 times longer than broad, with 1 tergal spine-like seta distally. Dactylus and terminal spine unfused, short, combined length reaching 0.45 times that of propodus. Peraeopod 3 (Fig. 10): Similar to P.2, ex- cept propodus 2.4 times longer than broad, therefore carpal blade-like spine appearing smaller than in P.2, but in reality of same size. Peraeopod 4 (Fig. 10): Somewhat stouter than P.1—P.3. Coxa fused to peraeonite, no setae. Basis nearly 3.6 times as long as broad, tergal border with 2 feathered hairs, sternal border with 1 proximal setule. Ischium an- nular, with 2 tergal setae. Merus bent ster- nally, 1.5 times longer than broad, tergal border with 1 rostral and 1 caudal spine- like seta distally. Carpus elongate, about 3 times longer than broad, tergal border dis- VOLUME 101, NUMBER 1 -) ound! 6 ENS Jamaica Di im J . SOlNuy Caribbean Sea Fig. 12. Distribution of the pseudotanaid species in the Caribbean and the Gulf of Mexico. @ Jungentitanais primitivus (Sieg, 1973). Hl Pseudotanais mortenseni Sieg, 1977. & Pseudotanais mexikolpos, n. sp. tally with 1 rostral blade-like spine and 1 spine, sternal border distally with 1 caudal seta. Propodus slightly longer than carpus, 4.2 times longer than broad, tergal border with | rostral and | caudal spine-like setae distally, sternal border with | spine-like seta, which is distinctly longer than terminal seg- ment. Dactylus and terminal spine fused to claw. Peraeopod 5 (Fig. 10): Similar to P.4. Ba- sis unarmed, dactylus bearing 1 additional seta, and sternal border of propodus with 3 to 4 tubercles. Peraeopod 6 (Fig. 10): Similar to P.4 and P.5, except propodus bearing distally 1 ad- ditional small seta on sternal border. Pleopods (Fig. 11): All 5 pairs of pleopods similar. Basis small, slightly longer than broad, unarmed. Exopodite 1-segmented, with 6 setae on outer border. Endopodite l-segmented, with 1 small seta appearing articulated to inner border, outer border with 4 setae, none appearing pinnate. Pleotelson (Fig. 11): Of typical shape, not elongate, 1.4 times broader than long; cau- dal point protuberant, with 2 small medial setae; | additional seta at each caudal cor- ner. Uropods (Fig. 11): Biramous. Basis short, as long as broad, unarmed. Endopodite 2-segmented; first segment 2.1 times as long as broad, with 2 feathered hairs and | seta distally; second segment as long as first, but more slender, about 2 times longer than broad, 2 small, 3 long, and | feathered hair at tip. Exopodite 2-segmented, reaching slightly more than *%4 length of endopodite; first segment 2.5 times longer than broad, 1 distal seta; second more slender, 3.8 times longer than broad, with | long and | short seta at tip. Remarks.—This species appears to be most closely related to P. unicus Sieg, 1977 and P. mediterraneus G. O. Sars, 1882, both from the Mediterranean. Pseudotanais mexikolpos can be separated from P. unicus by its short cephalothorax and peraeopod 3, which is distinctly smaller in P. unicus. It is distinguished from P. mediterraneus by having a much longer disto-sternal seta on propodus of P.4—P.6. Distribution (Fig. 12).—Pseudotanais mexikolpos is presently known from the type locality, in the vicinity of Golum Lake, a hypersaline seep, on the East Flower Gar- den Bank, and from Stetson Bank, both off Texas coast. Acknowledgments Sara LeCroy, through Applied Biology Inc., provided the specimens from the Flor- 58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ida Keys (Pumpkin Creek and Shark Chan- nel). Loan of the material from the Florida Middle Ground was arranged by Mike Dar- deau, Museum Curator, Marine Environ- mental Sciences Consortium, Dauphin Is- land, Alabama. Specimens from the East Flower Garden Bank were provided by Eric N. Powell, Department of Oceanography, Texas A&M University. The specimens from the Florida Middle Ground and the East Flower Garden Bank were collected with support provided by the U. S. Bureau of Land Management under Contract Nos. AA551-CT8-35 and AA550-CT7-15, re- spectively. Ruth Shulman provided critical assistance in the preparation of the manu- script and M. Schilling inked the pencil drawings. This project was also supported by the Deutsche Forschungsgemeinschaft. Literature Cited Bacescu, M. 1960. Citera animale neucunescute inca in Marea si descrierea unor Malacostracei noi (Elaphognathia monodi n. sp. si Pontotanais borcei n. g. n. sp.) provenind din apele pontici prebosforice.—Studii si Cercetari di Biologie (Biologie Animale) 12(2):107—124. Fee, A.R. 1927. The Isopoda of Departure Bay and vicinity with description of new species, varia- tions and color notes.—Contributions to Ca- nadian Biologie and Fisheries 3:15—46. Hansen, H. J. 1887. Oversigt over de paa Dijmpha- togtet indsamlede krebsdyr. Pp. 183-286, in Lueken, C. F. (ed.), Diympha togets zoologisk- botaniske udbytte. Kopenhagen. 1913. Crustacea Malacostraca II.— Danish Ingolf-Expedition 3(3):1-145. Johnson, S. B., & Y. G. Attramadal. 1982. A func- tional-morphological model of Tanais cavolinii Milne-Edwards (Crustacea, Tanaidacea) adapt- ed to a tubicolous life-strategy.—Sarsia 67:29- 42. Kudinova-Pasternak, R. K. 1966. Tanaidacea Crus- tacea of the Pacific ultra abyssals.—Zoologi- cheskii Zhurnal 45:516-535. 1975. Tanaidacea (Crustacea, Malacostraca) from the Atlantic sector of Antarctic and Sub- antarctic.—Trudy Instituta Okeanologii 103: 194-228. . 1982. Deep sea Tanaidacea (Crustacea, Mal- acostraca) from Mediterranean Sea. — Trudy In- stituta Okeanologii 117:151-162. 1985. Tanaidacea (Crustacea, Malacostraca) collected on the summit and the foot of Great Meteor seamount.—Trudy Instituta Okeanol- ogii 120:52-64. —., & F. Pasternak. 1978. Deep-sea Tanaidacea (Crustacea, Malacostraca) collected in the Ca- ribbean Sea and Puerto Rico Trench during the 16th cruise of R/V ‘Akademic Kurchatov’ and the resemblance between the fauna of deep-sea Tanaidacea of the Caribbean region and the Pa- cific.—Trudy Instituta Okeanologii 113:178- 197. ,& . 1981. Tanaidacea collected by the Soviet Antarctic Expedition.— Trudy Instituta Okeanologii 115:108-125. Lang, K. 1960. The genus Oosaccus Richardson and the brood-pouch of some tanaids.— Arkiv for Zoologi (2)13:77-80. Lilljeborg, W. 1864. Bidrag till kaennedomen om de inom Sverige och Norrige foerkommande Crustaceer af Isopodernas underordning och Tanaidernas familj. Inbjudningsskrifter Uni- versitetet Uppsala, 31 pp. Sars, G. O. 1868. Undersogelser over Christianiaf- jorden dybvandsfauna.—Nytt Magasin for Naturvidenskapene 16:305-—362. 1882. Rivision af gruppen: Isopoda Cheli- fera.—Archiv for Mathematik og Naturviden- skab 7:1-54. Shino, S. M. 1978. Tanaidacea collected by French scientists on board the survey ship ‘Marion Du- fresne’ in the region around the Kerguelen Is- lands in 1972, ’74, ’75, ’76.—Scientific Report Shima Marineland 5:1-122. Sieg, J. 1973. Ein Beitrag zum Natiirlichen System der Dikonophora Lang. Dissertation, Christian- Albrechts Universitat Kiel, 298 pp. 1976. Zum natiirlichen System der Dikono- phora Lang (Crustacea, Tanaidacea).—Zeit- schrift fur Zoologische Systematik und Evolu- tionsforschung 14:177-198. . 1977. Taxonomische Monographie der Fam- ilie Pseudotanaidae (Crustacea, Tanaidacea). — Mitteilungen aus dem Zoologischen Museum in Berlin 53:3-109. . 1980. Taxonomische Monographie der Tan- aidae Dana, 1849 (Crustacea: Tanaidacea).— Abhandlungen der Senckenbergisch Natur- forschenden Gesellschaft 537:1—267. 1983. Tanaidacea.—Crustaceorum Catalo- gus 6:1-552. 1984. Neuere Erkenntnisse zum natirlichen System der Tanaidacea. Eine phylogenetische Studie. — Zoologica (Stuttgart) 136:1-132. . 1986a. Tanaidacea (Crustacea) from the Ant- arctic and Subantarctic. 1. Material collected from Tierra del Fuego, Isla de los Estados, and VOLUME 101, NUMBER 1 along the west coast of the Antarctic Peninsu- la.— Antarctic Research Series 45:1-180. 1986b. Tanaidacea (Crustacea) von der An- tarktis und Subantarktis. IJ. Tanaidacea gesam- melt von Dr. J.-W. Wagele wahrend der Deutschen Antarktis Expedition 1983.—Mit- teilungen aus dem Zoologischen Museum der Universitat Kiel 2(4):1-80. Texas A&M University. 1978. South Texas topo- graphic features study. A final report to the U.S. Department of the Interior Bureau of Land Management Outer Continental Shelf Office New Orleans, Louisiana, Contract No. AA 550-CT- 6-18. College Station, Texas 775 pp. Tzareva, L. A. 1982. Doplonenie k faune kleschne- nosnich osslikov (Crustacea, Tanaidacea) schelfovich son antarktiki 1 subantarktiki. Pp. 59 40-61, in Kavanov, A. I. (eds.), Fauna i Ras- predelenie Rakoobranich Notalnich i Antark- titscheskich vod. Akademia Nauk SSSR, Vla- divostok. Vanhoffen, E. 1914. Die Isopoden der Deutschen Sidpolar Expedition 1901-—1903.— Deutsche Suidpolar Expedition 15:449-598. (JS) Universitat Osnaburck, Abt. Vechta/ FB 13, DriverstraBe 22, D-2848 Vechta, Federal Republic of Germany; (RWH) In- vertebrate Zoology Section, Gulf Coast Re- search Laboratory, Ocean Springs, Missis- sippi 39564. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 60-66 DIACRIA PICCOLA AND DIACRIA MACULATA: TWO NEW PTEROPOD MOLLUSC SPECIES FROM THE ATLANTIC AND PACIFIC OCEANS J. Bleeker and S. van der Spoel Abstract. —A few specimens of Diacria from the northwestern Pacific Ocean were described as intermediates between D. trispinosa forma trispinosa and D. rampali (cf. Van der Spoel 1982). New records from this Diacria form from the Pacific and Atlantic prove that these ‘intermediates’ are a common phe- nomenon, not an incidental occurrence. The co-occurrence of this Diacria form with other related species of the genus made it acceptable that a good species is involved, for which the name Diacria maculata is proposed. In one of the Northwestern Pacific Ocean sediment samples very small Dia- cria trispinosa-like specimens were found. Though it is known that the western Pacific representatives of D. trispinosa may be small (Rampal 1975), the present material, twice as small as the smallest previously known specimen, is consid- ered to represent a species new to science for which the name D. piccola is proposed. Diacria piccola, new species Figs. 2-4 Material.—Locality data of D. piccola with numbers of specimens in parentheses: Holotype USNM 276782 and paratype USNM 859098, both from Albatross sta 5236, Philippines, off Magabao Is., E. Min- danao, 8°50'45’N, 126°26'52”E, 494 fms, fine gray sand, 12 feet Agassiz beam trawl, 11 May 1908 (2). Description. —The specimens collected from the sediment are subfossils or Recent specimens which have been buried in the sediment for a long period, as evidenced by decalcification. No color patterns remain visible. In the holotype the caudal spine is broken off below the closing septum, the width near the septum (> 15% of shell width) is relatively large compared to other species (<15% of shell width). The lateral spines are worn off. The dorsal side shows 3 ribs; the central one is especially broad and flat- tened. The lateral sides between the closing septum and the lateral spines are nearly straight and meet at an angle of 66°. The dorsal lip is curled dorsally. The ventral lip is slightly curved ventrally. The 2 lateral ribs on the ventral side are indistinct. Mea- surements of the holotype: shell width 2.52 mm, shell length 3.32 mm, length between lateral spines and closing septum 1.60 mm, distance between lateral spines and closing septum 2.00 mm, distance between lateral spines and centre of upper lip 1.80 mm, width of shell aperture 1.08 mm, height of shell aperture 0.49 mm, angle of lateral sides 66°. The paratype is slightly smaller than the holotype and damaged at the upper shell lip; it is filled with sediment. Type material. —Deposited at the Smith- sonian Institution. The holotype USNM 276782 and one paratype USNM 859098 are both from A/batross sta 5236, 494 fms, fine gray sand. Type locality. — Philippines, off Magabao Is., E. Mindanao, 8°50’'45”N, 126°26'52’E. Etymology.—The small size and resem- VOLUME 101, NUMBER 1 blance to a small pipe is reflected in the Italian word piccolo. It is here treated as a neo-Latin adjective. Diacria maculata, new species Figs. 4, 5, 8, 9 Material. —Locality data of D. maculata with numbers of specimens in parentheses; geographic positions between square brack- ets are added by the present authors: Ho- lotype USNM 283052 and paratypes USNM 859099 (8), both from: Albatross sta 5470, Philippines, Lagonboy Gulf, E. Luzon, 13°37'30’N, 123°41'09”E, 560 fms (esti- mate), 18 Jun 1909 (9). USNM 859100: Fish Hawk sta 1154, off Martha’s Vineyard, 39°55'31”N, 70°39'W, 193 fms (1). —USNM 38491: Albatross sta 2222, S of Martha’s Vineyard, 39°03'15’N, 70°50'45’W, 1537 fms (2).—USNM 859101, Albatross sta 2760, off Bahia, Bra- zil, 12°07'00"S, 37°17'00”W, 1019 fms, ooze (1). —USNM 859102, Albatross sta 5580, Darvel Bay, Borneo, 23 m, W of Sibutu, 4°52'45"N, 119°06'45”E, 162 fms, brown sand and coral (2).—USNM 258143: Albatross sta 5238, Philippines, off Pt. Lambajon, E. Mindanao, 7°34'45’N, 126°38'15”E, 380 fms, green mud (4).— USNM 274767: Albatross sta 5394, Phil- ippines, off Dumurug Masbate, 12°00'30’N, 124°05'36”E, 153 fms, green mud (5).— USNM 276003: Albatross sta 5265, Phil- ippines, off Matocot Pt., W. Luzon, 13°41'15”N, 120°00'50’E, 135 fms, sand and mud (1).—USNM 859103, Albatross sta 5236 Philippines, off Magabao Is., E. Min- danao, 8°50’'45’N, 126°26'52”E, 494 fms, fine gray sand (4).—USNM 281794: Alba- tross sta 5392, Philippines, off Adyagan Is., E. Masbata, 12°13'15”N, 124°05'03”E, 135 fms, green mud and sand. (5).—USNM 284465: Albatross sta 5314, China Sea, off Pratas Is., 21°41’N, 116°46’E, 122 fms, sand and broken shells (1). —USNM 284819: Al- batross sta 5313, China Sea, off Pratas Is., 21°30'N, 116°49’E, 150 fms, sand (2).— USNM 288113: Albatross sta 5425, Phil- 61 Fig. 1. ferred to. A, Shell length; B, Length between lateral spines and closing septum; C, Shell width; D, Distance between lateral spines and closing septum; E, Distance between lateral spines and center of upper lip; F, Height of shell aperture; G, Width of shell aperture. Diacria shell showing measurements re- ippines, Jolo Sea off Cagayan Is., 9°37'45’N, 121°11'E, 495 fms, gray sand, mud, coral and sand (9).—USNM 289069: Albatross sta 5453, Philippines, Albay Gulf, off Legaspi light, E. Luzon, 13°12'N, 123°49'18’E, 146 fms (1). —USNM 289259: Albatross sta 5429 Philippines, off Fondeado Is., E. Palawan, 9°41'30”N, 118°50'22”E, 766 fms, green mud. (1 + cf. 2).—USNM 289640: Alba- tross sta 5459, Philippines, Albay Gulf, E. Luzon, 13°10’21”N, 123°59’54”E, 201 fms (2).—USNM 289802: Albatross sta 5582, Borneo, off Si Amil Is., 4°19'54’N, 118°58'38”E, 890 fms, gray mud and fine sand (1).—USNM 334804: [21°15'N, 158°W] Albatross sta 3908, Hawaiian Is., S. coast Oahu Is., 304—308 fms, fine white sand and mud. (1).—USNM 859104 Albatross sta 2644, off Cape Florida, 25°40’00’N, 80°00'00”"W, 193 fms, gray sand (1).— USNM 357699: Eolis sta 118, Miami, Flor- ida, off Govt. cut NE, 30 fms (1). —-USNM 859105: [25°35’N, 80°05'’W] Eolis sta 153, off Foway light, Fla., SE 3,5 mi. (1); USNM 859106: [25°35'N, 80°05'W] Eolis sta 174, 62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 2-6. Diacria piccola: 2, Holotype, dorsal view; 3, Holotype, ventral view; 4, Paratype, ventral view. Diacria maculata: 5, Holotype, dorsal view; 6, Holotype, ventral view. off Foway light, Florida, ENE, 58 fms (1).— USNM 859107: [25°30'N, 80°11’W] Eolis sta 302, off Sand Key, Florida, 100 fms. (1).—USNM 859108: [25°30'N, 80°11'’W] Eolis sta 324, off Sand Key, Florida, 100 fms (11).—USNM 859109 Eolis sta 329, off Sambo Reef, Florida, 135 fms (8).—USNM 859110: [25°35’N, 80°05'W] Eolis sta 340, off Foway light, Florida, 209 fms. (3).— USNM 429577: Johnson Smithsonian Exp. Sta 32, 18°25'50"N, 67°14'55”W 200-280 fms (1). —-USNM 859115: Johnson Smith- sonian Exp. sta 93, 18°38'00’N, 65°09'30"W, 350-400 fms (2).—USNM 859111: Johnson Smithsonian Exp. sta 25, 18°32'15”N, 66°22'10”W, 240-300 fms (1). —USNM 859112: State University of Iowa Exp. sta 25, off Pelican Is., Barbados, 80 fms, coarse sand (1).—USNM 859113: [13°57'S, 59°33'W] State University of lowa Exp. sta 54, off cable station, Barbados, 33 fms rocky (1). -USNM 859114 coarse sand (1); 59°33’'W] (1).—USNM 859114 Alba- tross sta 2711, New Jersey, 38°59’00’N, VOLUME 101, NUMBER 1 63 Figs. 7-10. 7, Diacria piccola x D. rampali, ventral view; 8, Diacria piccola x D. trispinosa, ventral view; 9, Diacria maculata from the Pacific Ocean; 10, Diacria maculata from the Atlantic Ocean. 70°07'00"W, 1544 fms globigerina ooze (1). Description.—The type series was col- lected from the sediment but no decalcifi- cation has occurred, and the color pattern is well preserved so that very recently sed- imented specimens are represented. In the holotype the caudal spine is broken off be- low the closing septum, the width at the closing septum shows similarity to that in Diacria trispinosa, for example. The lateral spines are straight and sharp, slightly di- rected caudally. The dorsal side shows 3 moderately developed ribs. The central rib is most elevated in the caudal part, the lat- eral ones in the middle. The lateral sides between lateral spines and closing septum are nearly straight and form an angle of 109°. Growth lines are evident on both sides. The color pattern on the dorsal side consists of a continuous band along upper and lateral shell margins; a separate patch of color is found on the caudal half of the median dor- sal rib. The upper lip is curved only slightly dorsally. The ribs on the ventral side are very indistinct, the ventral aperture lip curves prominently ventrally. The color 64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pattern on the ventral side also forms a con- tinuous band along upper and lateral shell margins, and a roughly quadrangular color patch is found on the upper half of the ven- tral side, as well as a patch on the narrowing caudal part of the ventral side. Sizes in the holotype are: shell width 8.80 mm, shell length 7.36 mm, length between lateral spines and closing septum 3.20 mm, dis- tance between lateral spines and closing sep- tum 4.80 mm, distance between lateral spines and center of upper lip 5.68 mm, width of shell aperture 3.20 mm, height of shell 0.72 mm, aperture angle between lat- eral sides 109°. Type material. — Deposited at the Smith- sonian Institution. The holotype USNM 283052 and 8 paratypes USNM 859099 are all from Albatross sta 5470, 560 fms (esti- mate). Type locality.—Philippines, Lagonboy Gulf, E. Luzon, 13°37'30’N, 123°41'09’E. Etymology. —The color spots (=macula- ta) on both sides give this species its name. Discussion. —Sediment samples are used throughout as the forms described as new in this paper are not recorded from the plankton. In the large collections of plank- ton samples studied from the areas con- cerned no comparable specimens were found so that it is not excluded that the new species are fossil or subfossils. The measurements given in the descrip- tions and table are indicated in Fig. 1. Diacria piccola is distinct from all related species by its small size and narrow angle between the lateral sides. D. maculata is distinct from all other species by its color pattern. Diacriais composed of the following taxa: the Diacria trispinosa group: Diacria trispi- nosa forma trispinosa (De Blainville, 1821), between 40°N and 40°S in all oceans. Diacria trispinosa forma atlantica Du- pont, 1979, between 40°N and 70°N in the Atlantic Ocean. Diacria trispinosa forma atlantica Du- pont, 1979 upwelling type (cf. Hilgersom & Van der Spoel 1988) in upwelling area off NW Africa. Diacria rampali Dupont, 1979, between 30°N and 30°S in all oceans. Diacria major (Boas, 1886), in Central waters of all oceans. Diacria maculata, n. sp., in NW bound- ary currents of Atlantic and Pacific Oceans. Diacria piccola, n. sp., W tropical Pacific near Philippines. The Diacria quadridentata group (cf. Van Leyen & Van der Spoel 1982): Diacria quadridentata (De Blainville, 1821). Indo-Pacific between 30°N and 30°S. Diacria costata Pfeffer, 1879. Central . water form (cf. Van Leyen & Van der Spoel 1982) of the Pacific. Diacria costata Pfeffer, 1879. Equatorial water form (cf. Van Leyen & Van der Spoel 1982) of Indian and Pacific Oceans. Diacria danae Van der Spoel, Equatorial waters of all oceans. Diacria schmidti schmidti Van der Spoel, 1971. Eastern Tropical Pacific. Diacria schmidti occidentalis Van Leyen & Van der Spoel, 1982. Western Tropical Pacific near Philippines. Diacria erythra erythra Van der Spoel, 1971. Red Sea and Western Indian Ocean. Diacria erythra crassa Van der Spoel, 1971. Red Sea. Variation and distribution. —Diacria pi- cola is known currently only from near the Philippines, thus any conclusion on its dis- tribution is excluded. Further, the two spec- imens available cannot give an indication of variation. In the sample with the type material, however, two specimens also were found that were somewhat larger and re- semble in shape D. rampali (Fig. 7) or D. trispinosa (Fig. 8). This may indicate that interbreeding of D. piccola with related species is still possible. Deformation or ab- normal growth are not expected to generate shells of the type described here as all aber- rant shells found show either asymmetry or 1968. VOLUME 101, NUMBER 1 65 60. 80 100 120 140 160 E 180 160 140 120 100 80 W. es [eee eee ee N 80 Banta ho 70 = 60 | it 50 + 40 + Miso ; = + SS 20 ae 10 ke 0 10 20 30 H 40 50 60 Ae 70 e cent ananny oes { ; 80 | SSS Se See eee ee eee eee ee ee (See 6 8 | Ss Fig. 11. different aberrations in different growth re- gions of the shell, which is not the case in D. piccola. Rampal (1975) described the size varia- tion of Diacria trispinosa in the Pacific Ocean, but though a tendency towards smaller sizes is clear, it is not acceptable that D. piccola forms the extreme of a size vari- ation of one of the two mentioned species. The gap between the size range of D. piccola and the other species is too large and size variation will be binomial with a separate maximum for D. piccola. Diacria maculata is found in the western North Pacific and in the western North At- lantic oceans (Fig. 11). Though the distri- bution of this species probably is not com- pletely known, it is evident that it does not occur along the eastern borders of the oceans. An endemic occurrence in western bound- ary waters is not known for other species, though a few (distant) neritic taxa have their range in these water masses. Probably D. Distribution of Diacria maculata (closed circles) and D. piccola (open circles). Table 1.—Morphometric data (minimum—maxi- mum) in mm based on the present material and lit- erature data showing the unique position of D. picola and the more intermediate position of D. maculata. Shell Shell Aperture Angle of Species width length height lateral sides D. major' 7.0-11.0 7.5-9.0 0.5-1.0 — D. t. dark 7.0-10.0 6.0-8.0 — — type? D. macu- 9.8 7.4 0.9 100-110° lata? D. t. atlan- 6.5-12.0 6.0-9.5 0.4—0.8 — tica! D. major 6.5-8.0 7.2-8.4 0.5-0.9 100-110° D. macu- lata 5.5-8.8 5.0-7.2 0.5-0.8 100-110° D. t. tris- pinosa' 5.5-9.0 5.0-6.5 0.5-0.9 — D. t. tris- pinosa 5.6-7.8 5.4-6.8 0.5-0.8 80-90° D. rampali' —5.5-8.5. 5.0-7.0 0.5-0.8 — D. rampali 5.5-6.9 5.5-6.6 0.5-0.7 80—90° D. piccola 2.0-2.4 2.5-2.8 0.24 65-80° ' After Dupont 1979. ? After Hilgerson & Van der Spoel 1988. 3 After Van der Spoel 1982. 66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON maculata is a neritic taxon in the Diacria trispinosa group of oceanic species. Atlantic and Pacific specimens do not differ in color pattern and shape, the Pacific ones are only slightly larger than the Atlantic ones, as can be seen in Table 1. In shape D. maculata is closest to D. ma- jor and in shape to D. rampali, but many characters, like the closing membrane in the caudal spine, make it look like a D. trispi- nosa, So that it was originally considered an intermediate between both (Van der Spoel 1982). Acknowledgments Dr. C. F. E. Roper and M. Sweeney kindly provided the material on which this pub- lication is based and commented on the manuscript. Literature Cited Dupont, L. 1979. Note on variation in Diacria Gray, 1847 with descriptions of a species new to sci- ence, Diacria rampali nov. spec., and a form new to science, Diacria trispinosa forma atlan- tica nov. forma.— Malacologia 18:37—52. Hilgersom, P., & S. van der Spoel. 1988. East west variation in Diacria off Northwest Africa.— Malacological Review [in press]. Rampal, J. 1975. Les Thécosome (Mollusques pé- lagiques). Systématique et évolution. Ecologie et biogéographie mediterranéennes. — Thése Université Provance Marseille CNRS.A.0.- 11932, 485 pp. Van der Spoel, S. 1982. Intermediates between Dia- cria trispinosa and D. rampali (Mollusca, Ptero- poda).— Basteria 46:121-123. Institute of Taxonomic Zoology, Univer- sity of Amsterdam, Plantage Middenlaan 53, P.O. Box 20125, 1OOOHC Amsterdam, The Netherlands. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 67-71 DEEP-SEA TUBIFICIDAE (OLIGOCHAETA) FROM THE GULF OF MEXICO Christer Erséus Abstract. — Four bathyal species of the subfamily Phallodrilinae (Phallodrilus constrictus, n. sp., P. grasslei Erséus, 1984, Bathydrilus connexus, n. sp., and B. longiatriatus Erséus, 1983) are reported from the northern part of the Gulf of Mexico. Phallodrilus constrictus is characterized by bifid penial setae (three per bundle), and heavily muscular atria and copulatory sacs, the latter enclosing pendant penes. Bathydrilus connexus, which belongs to the ‘B. adriaticus group,’ has more or less single-pointed setae in postclitellar segments, and blunt, some- what chisel-shaped penial setae (two, occasionally three, per bundle). Marine oligochaetes collected during a study of the northern Gulf of Mexico con- tinental slope were sent to the author for identification from LGL Ecological Re- search Associates, Inc. (Bryan, Texas). The material contained four bathyal species be- longing to the subfamily Phallodrilinae, of which two are new to science. The taxon- omy of these species is treated here. Other tubificids found are Limnodriloides mono- thecus Cook, 1974 (Limnodriloidinae) and a species of Tubificoides (Tubificinae), but they occurred only at stations of about 350 m depth. The latter of these two will be treated by the author elsewhere. The specimens studied here were collect- ed by boxcore sampling along two transects, south of Louisiana and south of western Florida, respectively. Some worms were stained with paracarmine by the author; all individuals were then mounted whole in Canada balsam. The material is deposited at the National Museum of Natural History (USNM), Smithsonian Institution, Wash- ington, D.C. Phallodrilus Pierantoni, 1902 (Generic definition: see Erséus 1984a) Phallodrilus constrictus, new species Figs. 1A—C Holotype. —USNM 113701, from S of W Florida, 28°16'42”N, 86°15'06”W, 625 m, 16 Apr 1984. Paratypes.—USNM_ 114501-114502, 2 specimens: | from 28°07'05’N, 86°19'15’W, 860 m, 18 May 1985; 1 from 28°21'45’N, 86°48'05”W, 852 m, 20 May 1985. Description. —One paratype 3.2 mm long, 20 segments (other specimens not com- plete). Width at XI (compressed, whole- mounted specimens) 0.14—0.17 mm. Cli- tellum extending over !/2X—'2XII. Somatic setae (Fig. 1A) bifid, with upper tooth thin- ner and slightly longer than lower, at least anteriorly. These setae 28-47 um long, about 1-1.5 wm thick, 3-4 per bundle anteriorly, 2 per bundle in postclitellar segments. Peni- al setae (Fig. 1B; C, ps) slender, bifid, with upper tooth thinner and shorter than lower, about 45-50 wm long, 2 um thick, 3 per bundle. Male pores paired ventrally and posteriorly in XI. Spermathecal pores paired in line with ventral setae, anteriorly in X. Pharyngeal glands in IV—VI. Male geni- talia (Fig. 1C) paired. Vas deferens not ob- served. Atrium oval, 53-63 um long, 33- 37 wm wide, with 3-8 wm thick lining of muscles, and ciliated and somewhat gran- ulated inner epithelium. From ectal end of atrium short, narrow duct leading into oval, heavily muscular copulatory sac, 47-49 wm long, 30-40 um wide. Penis present, narrow and pendant within copulatory sac. Ante- 68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (oe) Fig. 1. A-C, Phallodrilus constrictus, n. sp.: A, Free-hand drawing of somatic seta; B, Free-hand drawing of penial seta; C, Somewhat ventral view of spermathecae and one of male ducts in segments X—XI (note: septum between X and XI not observed). D-E, Phallodrilus grasslei Erséus: D, Penial setae; E, Spermatheca. Abbre- viations: a atrium; cs copulatory sac; p penis; pr | anterior prostate gland; pr 2 posterior prostate gland; ps penial seta; s spermatheca. rior prostate gland attached to apex of atrium. Posterior prostate attached to ectal end of atrium, near exit of narrow duct. Spermathecae (Fig. 1C, s) cylindrical, 82- 94 um long, 21-28 wm wide, consisting of short, partly hollow ducts, and slender am- pullae, all with 2—4 um thick lining of mus- cles; sperm in small compartments within ampullae. Remarks. —Phallodrilus constrictus is named for the distinct ‘constriction’ on the male duct between the atrium proper and the muscular copulatory sac. The species bears resemblance to P. vu/nus Erséus, 1983 and P. cristolatus Erséus, 1983, two bathyal species from the NE Atlantic which also have bifid penial setae and penial structures (latter termed pseudopenes by Erséus 1983). [Note that the illustrations of P. vu/nus and P. cristolatus were transposed in the original paper; the legend of Erséus’ (1983) fig. 1 actually refers to the drawing of fig. 2 and vice versa.] Phallodrilus contrictus is, how- ever, easily distinguished from both of these by its very conspicuous, muscular copula- tory sacs and by the morphology of its sper- mathecae. Another close relative of the new species is probably P. davisi Erséus, 1984, known from the continental shelf off Mas- sachusetts (Erseus 1984c). It has well de- veloped copulatory sacs and penes, as well as compartmented spermathecae. The male ducts and the spermathecae of P. davisi are, however, not as muscular as those of P. con- strictus, and the species lacks penial setae. Distribution and habitat. —Northern Gulf of Mexico. Upper continental slope sedi- ment, 625-860 m. Phallodrilus grasslei Erséus, 1984 Fig. 1D, E Phallodrilus grasslei Erséus, 1984b:101-103, figs. 1-3. Type material.—USNM 96493-96495, holotype and 2 paratypes from 1789 m depth, S of Massachusetts, NW Atlantic (see Erseus 1984b). VOLUME 101, NUMBER 1 New material examined. -USNM 113702, 1 specimen from S of Louisiana, 28°04'18’N, 86°34'24’W, 1330 m, 17 Apr 1984. Remarks. —This species was previously known only from off Massachusetts. The new individual conforms well to the original description. For instance it has sigmoid, hooked penial setae, 2 per bundle (Fig. 1D), which is the most striking feature of P. gras- slei. The spermathecal ampullae of the orig- inal material had thick walls (Erseus 1984b: fig. 1). The spermathecae of the new ma- terial are, however, more thin-walled (Fig. 1E). Distribution and habitat. —Northern Gulf of Mexico (new record), S of Massachusetts. Continental slope sediments, known from 1330-1789 m depth. Bathydrilus Cook, 1970 (Generic definition: see Erséus 1981, and modification by Erséus 1983) Bathydrilus connexus, new species Fig. 2A—C Holotype. —USNM 113703, from S of W Florida, 28°09'36’"N, 86°25’00"W, 845 m, 17 Apr 1984. Paratypes. -USNM 113704-113710 and 114503-114505, 10 specimens from S of W Florida and Louisiana: 1 from 26°57'48’N, 89°31'00”W, 2490 m, 28 Nov 1983; 2 from 26°58'00’N, 89°31'48”W, 2467 m, 29 Nov 1983; 1 from 26°56'54’N, 89°36'12”W, 2377 m, 13 Apr 1984; 2 from 28°16'42’N, 86°15'06’”W, 625 m, 16 Apr 1984; 1 from 28°00'24"N, 86°38'48’"W, 2853 m, 18 Apr 1984; 2 from 28°14'50’N, 86°09'47’W, 618 m, 16 May 1985; and 1 from 28°00'10’N, 86°38'43"W, 2902 m, 21 May 1985. Description. — Length (2 complete worms) 7.8-12.9 mm, 43-55 segments. Width at XI (compressed, whole-mounited specimens) 0.23-—0.39 mm. Epidermal glands not ob- served. Clitellum extending over '2X—XII. Somatic setae (Fig. 2A, B) 45-75 um long, 2.5-3 um thick, 2—3(4) per bundle ante- 69 riorly, 2(3) per bundle in postclitellar seg- ments. In segments II—VIII, setae bifid, with upper tooth thinner and shorter than lower (Fig. 2A). From IX, setae sharply single- pointed or with very much reduced upper tooth (Fig. 2B). Penial setae (Fig. 2C, ps) straight or somewhat curved, (3) per bundle, 60-85 um long, entally S—6 um wide (ectally Narrower), with blunt, somewhat chisel- shaped tips directed towards and located near midventral line. Male pores paired in line with ventral somatic setae, posteriorly in XI. Spermathecal pores paired in lateral lines, in anteriormost part of X. In several specimens, male and spermathecal pores el- evated on bulbous protuberances. Pharyngeal glands in IV—VII. Male gen- italia (Fig. 2C) paired. Vas deferens 7-12 um wide, slightly longer than atrium, en- tering latter somewhat ectal to middle, to- gether with anterior prostate gland. Atrium spindle-shaped, 100-175 um long, 50-56 um wide at middle, with very thin outer lining of muscles and ciliated inner epithe- lium. Middle part of atrium densely gran- ulated. Posterior prostate glands attached to apical, inner end of atrium. Ectally, atrium terminating in simple pseudopenis. Sper- mathecae (Fig. 2C, s) with short, indistinct ducts and large, sacciform ampullae; latter with a few large roundish ‘spermatozeug- mata.’ Remarks.—This species is named con- nexus (Latin meaning ‘linked together’ or ‘bordering upon’), because of its close re- lationship with a whole group of, largely shallow-water, species of Bathydrilus [B. adriaticus (Hrabe, 1971), B. litoreus Baker, 1983, and others] characterized by more or less erect, spindle-shaped atria, bisetal or trisetal penial bundles, and sacciform sper- mathecae with sperm arranged in ‘sper- matozeugmata.’ Bathydrilus connexus is in fact very similar to B. adriaticus, but is re- garded as a separate species because of the detailed morphology of its setae. Sharply single-pointed postclitellar setae (Fig. 2B), are not known from any other member of 70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ie 100 um al Fig. 2. A-C, Bathydrilus connexus, n. sp.. A, Free-hand drawing of anterior seta; B, Free-hand drawing of postclitellar setae; C, Lateral view of spermatheca and male duct in segments X—XI. D, Bathydrilus longiatriatus Erséus: spermathecae in segments [X—X. Abbreviations: a atrium; pp pseudopenis; pr | anterior prostate gland; pr 2 posterior prostate gland; ps penial seta; s spermatheca; sb sperm bundle; sz ‘spermatozeugma’; vd vas deferens. this group. Moreover, the penial setae of the new species are blunt, those of B. adriaticus are pointed. Distribution and habitat. —Northern Gulf of Mexico. Continental slope sediments, known from 618-2902 m depth. Bathydrilus longiatriatus Erséus, 1983 Fig. 2D Bathydrilus longiatriatus Erséus, 1983:143- 144, fig. 10. Type material. —USNM 72988, holotype from about 2850 m depth, N of Surinam (see Erséus 1983). New material examined. -—USNM 113717-113722 and 114506, 7 specimens from S of W Florida and Louisiana: 3 from 26°58'00"N, 89°31'48"W, 2467 m, 29 Nov 1983; 1 from 27°28'24"N, 89°46'48’W, 1390 m, 13 Apr 1984; 1 from 28°09'36’N, 86°25'00"”W, 845 m, 17 Apr 1984; | from 27°27'41"N, 89°47'19”W, 1506 m, 15 Nov VOLUME 101, NUMBER 1 1984: 1 from 28°35'22’N, 86°46'26’W, 622 m, 13 May 1985. Remarks. — Bathydrilus longiatriatus was originally described on the basis of a single specimen from off Surinam. Very long, muscular atria (about 300-400 um long in new material), extending to the end of seg- ment XII, characterize the species. The ho- lotype was precopulatory, but in some of the new individuals the spermathecae con- tain an amorphous granular mass with a few bundles of sperm (Fig. 2D). Some of the worms from the Gulf of Mexico have fine particles adhering to the cuticle. Distribution and habitat. —Northern Gulf of Mexico (new record) and N of Surinam. Continental slope sediments, known from 622-2953 m depth. Acknowledgments The LGL specimens reported on in this publication were collected through funding by the U.S. Department of Interior, Min- erals Management Service, Gulf of Mexico Regional OCS Office under Contract Num- ber 14-12-0001-30046 and 14-12-0001- 30212. I am indebted to Dr. Linda H. Pequegnat (LGL Ecological Research Asso- ciates, Inc.) for placing the oligochaete ma- terial at my disposal, and to Ms. Barbro Lofnertz and Mrs. Aino Falck-Wahlstrom for technical assistance. Literature Cited Baker, H.R. 1983. New species of Bathydrilus Cook (Oligochaeta; Tubificidae) from British Colum- bia.—Canadian Journal of Zoology 61:2162- 2167. Cook, D. G. 1970. Bathyal and abyssal Tubificidae Tal (Annelida, Oligochaeta) from the Gay Head— Bermuda transect, with descriptions of new gen- era and species.— Deep-Sea Research 17:973- 981. 1974. The systematics and distribution of marine Tubificidae (Annelida: Oligochaeta) in the Bahia de San Quintin, Baja California, with descriptions of five new species. — Bulletin of the Southern California Academy of Sciences 73: 126-140. Erséus, C. 1981. Taxonomic studies of Phallodrilinae (Oligochaeta, Tubificidae) from the Great Bar- rier Reef and the Comoro Islands with descrip- tions of ten new species and one new genus. — Zoologica Scripta 10:15-31. . 1983. Deep-sea Phallodrilus and Bathydrilus (Oligochaeta, Tubificidae) from the Atlantic Ocean, with descriptions of ten new species. — Cahiers de Biologie Marine 24:125-146. 1984a. Taxonomy of some species of Phal- lodrilus (Oligochaeta: Tubificidae) from the Northwest Atlantic, with description of four species. — Proceedings of the Biological Society of Washington 97:812-826. . 1984b. New and little-known species of deep- sea Tubificidae (Oligochaeta) from the North- west Atlantic. — Zoologica Scripta 13:101-106. 1984c. Taxonomy of some species of Phal- lodrilus (Oligochaeta: Tubificidae) from the Northwest Atlantic, with description of four new species. — Proceedings of the Biological Society of Washington 97:812-826. Hrabe, S. 1971. On new marine Tubificidae of the Adriatic Sea.—Scripta Facultatum Scientiarum Naturalis Ujep Brunensis, Biologia 3, 1:215- 226. Pierantoni, U. 1902. Due nouvi generi di Oligocheti Marini rinvenuti nel Golfo di Napoli.—Bollet- tino della Societa di Naturalistii Napoli 16:113- 117. Swedish Museum of Natural History, Stockholm, and (postal address:) Depart- ment of Zoology, University of Goteborg, Box 25059, S-400 31 Goteborg, Sweden. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 72-78 A NEW CRAB, SEORSUS WADET, FROM THE LATE CRETACEOUS COON CREEK FORMATION, UNION COUNTY, MISSISSIPPI Gale A. Bishop Abstract.—A fossil dakoticancroid crab, Seorsus wadei, new genus and species, is described from the early Maastrichtian Coon Creek Formation of Union County, Mississippi. This crab belongs to the Dakoticancer Assemblage pre- viously described from the Blue Springs locality. The only known specimens consist of a nearly complete, crushed, carapace steinkern and a partial carapace steinkern. Seorsus differs from other members of the Dakoticancridae in car- apace proportions, shape and ornamentation. Collection of the Blue Springs locality (Bishop 1983) on 10 Mar 1984, resulted in the discovery of a new fossil crab. The Blue Springs locality is situated geographically in the Mississippi Embayment, an ancient arm of the Cretaceous Tethys Seaway that oc- cupied much of present Mississippi, Ala- bama, and western Tennessee. Two speci- mens were collected from the sandy claystone shelf sediments of the Coon Creek Formation of early Maastrichtian Age (Rus- sell et al. 1983). Correlation with other North American Cretaceous rocks indicates an ab- solute age of approximately 67 million years before present. Both specimens are internal casts of the carapace and are preserved as steinkerns. Systematic Paleontology Order Decapoda Latreille, 1803 Superfamily Dakoticancroidea Rathbun, 1917 Family Dakoticancridae Rathbun, 1917 Seorsus, new genus Type species. —The type species of Seor- sus 18 Seorsus wadei, n. sp. Diagnosis. —Carapace of moderate size, longer than wide (L/W = 1.10), widest at anterior 13; lateral margins distinctively convergent posteriorly; grooves broad, moderately defined; areolations very tumid, especially epibranchial lobes; cardiac region with small central tubercle; metabranchial region with subtle transverse and submar- ginal ridges. Claws and legs unknown. Etymology. —‘‘Seorsus”’ is derived from the Latin; apart, separate, severed, to in- dicate the carapace shape which sets this taxon apart from the other dakoticancroid genera and also the fragmented or severed nature of both specimens. The gender is masculine. Occurrence. —Seorsus is known from a single species based on two carapace spec- imens from the Coon Creek Formation, ear- ly Maastrichtian, of Union County, Missis- SIppl. Seorsus wadei, new species Fig. 1A—F Diagnosis. —Carapace longer than wide (L/W = 1.10), widest at anterior 14; lateral margins distinctively convergent posterior- ly. Grooves and areolations similar to those in other members of family. Carapace evenly granulate. Description. —Carapace 1.10 times longer (23.3 mm) than wide (21.1 mm), widest at anterior '4; front relatively straight; antero- lateral margin broadly rounded; lateral mar- gins relatively straight, converging poste- VOLUME 101, NUMBER 1 riorly; posterolateral margins tightly rounded; hind margin straight. Carapace re- gions tumid but poorly differentiated be- cause of broad shallow grooves. Cervical groove sinuous, posteriorly reflexed around large mesogastric region; hepatic groove and longitudinal gastric grooves poorly defined but present; branchiocardiac and cervical grooves define epimeral peninsula; anterior branchial groove very shallow; posterior marginal groove sharply defined. Cephalic arch differentiated into large mesobranchi- al, intermediate protogastric, and small he- patic regions each raised into a granulate tumid areolation. Scapular arch differen- tiated into sagittal ridge consisting of car- diac and intestinal regions; and branchial regions consisting of raised epibranchial ridge, a smaller mesobranchial ridge, and metabranchial area. Cardiac region with low media! boss. Hind margin raised into a pos- terior rim. Transverse, low submarginal bosses on posterior of metabranchial re- gions. Orbits large and broad occupying 48% of carapace width. Carapace regions evenly granulated except for more coarsely gran- ulated highest parts of hepatic and epibran- chial regions. Legs very granulate, decreas- ing in size posteriorly, P; very small. Sternum and abdomen poorly exposed. Ab- domen narrow (male) and abdominal so- mite A, with longitudinal ridge and trans- verse ridge. Comparison. — Seorsus wadei is an enig- matic fossil. As with many fossil crabs rep- resented by unique or only a few specimens, diagnostic characteristics necessary to con- fidently assign the species or genus to a higher taxonomic unit are often lacking. Assign- ment in such cases can only be made on the basis of circumstantial evidence such as general similarities in carapace shape, car- apace morphology, and carapace ornamen- tation. Although admittedly not the best possible criteria, assignment to a family seems better than to such taxa as “Family Indeterminate,” or not describing them at all. Once such a taxon Is described it is avail- 73 able for scientific discussion and possible reassignment as other carcinologists eval- uate the data. Seorsus wadei has a generalized carapace lacking clearly definitive characteristics. Based solely on shape, Seorsus is most sim- ilar to trapezohedral members of the Family Geryonidae but differs from them by lack- ing the dentate front and the large orbits. Seorsus also resembles some members of the Family Goneplacidae in its trapezohe- dral shape but differs from them in having clearly divided orbits. The carapace areo- lation of Seorsus resembles closely that of members of the Family Dakoticancridae. If Seorsus is viewed obliquely from the front, this similarity is enhanced and the trape- zohedral carapace outline becomes quite similar to that of Dakoticancer Rathbun, 1917. The carapace ornamentation of Seor- sus wadei consists of low, intermittent gran- ulation similar to that of Dakoticancer, Avi- telmessus Rathbun, 1923, and Tetracarcinus Weller, 1905, 1907. Because of these sim- ilarities, Seorsus is assigned to the Family Dakoticancridae. Seorsus wadei differs from other dakoti- cancroid crabs by its trapezohedral shape. Seorsus wadei is more arched and much smaller than Avite/messus grapsoideus Rathbun, 1923, but larger than Tetracarci- nus subquadratus Weller, 1905, and trape- zohedral rather than quadrate. Seorsus wa- dei most resembles Dakoticancer in areolation and ornamentation. Seorsus wadei is trapezohedral whereas D. australis Rathbun, 1935, is nearly rectangular. Seor- sus wadei is most similar to D. overanus Rathbun, 1917, which is slightly trapezo- hedral in carapace outline, but differs by being much more trapezohedral, having a relatively larger cephalic arch, and nearly lacking the prominent epimeral peninsulas of D. overanus. Variation. —Because this species is rep- resented by only two specimens, variation is undefinable. The mode of preservation as a steinkern preserves basic body shape and, 74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 101, NUMBER 1 because of compression of the exterior mold onto the interior mold, some of the carapace ornamentation. Description of a new species based on two specimens is justified by sam- pling (a collection of about 1500 fossil deca- pods) which indicates that this species is such a minor taxon in the Blue Springs Da- koticancer Assemblage that additional spec- imens may not be collected for some time. Such low abundances in fossil and recent assemblages necessitate the description of new taxa based on few specimens when dis- covery of additional specimens cannot rea- sonably be expected for some time. Type locality.—The only known speci- mens, the holotype (GSCM 1693) and para- type (GSCM 1694), were collected from the Coon Creek Formation in Union County, Mississippi, in the NE'’4, NW'4, SE'4, sec- tion 9, T. 8S., R. 4E. The holotype is de- posited in the collection of the GSC Mu- seum (Georgia Southern College, Statesboro, Georgia 30460-8161). Etymology.—The trivial name “‘wadei”’ honors the stratigrapher and paleontologist, Bruce Wade, whose pioneering efforts built the foundation for subsequent studies of Cretaceous fossils of the Mississippi Em- bayment. Remarks.—The specimens collected are phosphatic steinkerns (internal molds) that are grayish orange (10 YR 714) to yellowish gray (5 Y 712) in color. The carapaces are broken by compression and show two sizes (0.5 mm and 1.2 mm) of open burrows within them. The crabs’ legs have been bro- ken away by weathering but were partly rep- licated by dental impression wax for study. — Fig. 1. 75 This is the typical mode of preservation at the Blue Springs locality. The Dakoticancer australis Assemblage The Blue Springs Dakoticancer Assem- blage (Bishop 1983 fig. 3, table 1) consists of 11 decapods and is dominated by a few taxa (D. australis (49.0%), Protocallianassa mortoni (Pilsbry, 1901) (26.9%), and Ho- ploparia tennesseensis Rathbun, 1926 (10.1%)) and eight other taxa present in very small numbers (Bishop 1985). Seorsus wad- ei constitutes less than 0.1% of it. The as- semblage is thought to represent a preserved community fraction (Bishop 1981) ofa Cre- taceous decapod-dominated community that periodically inhabited the Cretaceous sea bottom (Bishop 1986a). The Superfamily Dakoticancroidea and Family Dakoticancridae were erected by Rathbun (1917:385) to contain the crab Da- koticancer overanus described from the Western Interior Cretaceous (South Dako- ta). In 1935 Rathbun described D. overanus australis and reassigned Tetracarcinus subquadratus to the Family Dakoticancri- dae. Glaessner (1960:46) remarked on similarities and reassigned Avitelmessus grapsoideus to Dakoticancridae. This clas- sification is the currently accepted model (Glaessner 1969:R491). Bishop (1983:426) raised D. overanus australis to species level rank. Rathbun (1937:26) described D. ols- soni, but it should be assigned to another genus. The current constitution of the Superfamily Dakoticancroidea is presented below with diagnoses and _ illustrations (Fig. 1). North American Cretaceous dakoticancroid crabs. A-F, Seorsus wadei, n. sp.: A-D, Holotype (GSCM 1693) in dorsal, ventral, anterior, and left lateral views, x 2.0; E-F, Comparative dorsal views of Holotype and Paratype (GSCM 1694). G-M, Comparative views of other dakoticancroid taxa: G, Tetracarcinus subquad- ratus Weller, 1905, carapace (GAB 37-1113); K, Same, right claw, x 2.0; H, Dakoticancer overanus Rathbun, 1917, carapace (GAB 4-2006, x 1.5); L, Same, right claw (GAB 4-B4, x 2.0); I, Dakoticancer australis Rathbun, 1935, carapace (USNM 73840, x 1.5); M, Same, right claw (GAB 37-1094, x2.0); J. Avitelmessus grapsoideus Rathbun, 1923, carapace (USNM 25411, x 2.5). 76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Superfamily Dakoticancroidea Rathbun, 1917 nom. correct. Glaessner (1969, R491) (pro Dakoticancroideae Rathbun, 1917) Diagnosis. —‘‘Carapace rectangular to transversely ovoid, front narrow; orbits well developed, median part of cardiac groove weak, branchiocardiac groove strong, no lineae; third maxillipeds elongate; sternum of female without longitudinal grooves, fifth pereiopods subdorsal, small.’’ (Glaessner 1969:R491). Family Dakoticancridae Rathbun, 1917 Diagnosis. —Characters of superfamily. Genus Dakoticancer Rathbun, 1917 Diagnosis. —*‘Carapace rectangular to transversely ovoid, wider than long, front narrow, orbits well developed, bilobed; me- dian part of cardiac groove weak, gastric regions hardly separated from cardiac-in- testinal region, branchiocardiac groove well developed, pleural sutures on carapace sides; genital openings on coxae, female on third leg and male on fifth; fifth legs much re- duced. Chelae equal.’’ (Bishop 1983:424). Dakoticancer overanus Rathbun, 1917 Fig. 1H, L Diagnosis. —Carapace moderate size, slightly wider than long, widest across epi- branchial regions, sides sinuous, slightly convergent posteriorly, well-differentiated by grooves, cervical groove prominent; or- namented by granules over areolations; pos- terior shelf narrow, rim upturned. Claws equal, much longer than high. Dakoticancer australis Rathbun, 1935 Fig. 11, M Diagnosis. —*‘Carapace large, slightly longer than wide, widest across branchial and hepatic regions (sides nearly parallel), well differentiated by grooves, ornamented by granules over entire surface. Claws sim- ilar, equal, short, stout, and crested; carpal articulation very oblique; fingers short, downturned.” (Bishop 1983:426). non Dakoticancer olssoni Rathbun, 1937 Remarks. —Rathbun assigned this Cre- taceous crab to Dakoticancer with no com- parison. The specimen in question (USNM 495104) was examined by me on 26 Aug 1977 at which time I made the note, ““This is not a Dakoticancer.” The specimen has probable affinity to the Necrocarcinids, pos- sibly being assignable to Protonecrocarcinus Forster, 1968, or to Cyphonotus Bell, 1863. Genus Tetracarcinus Weller, 1905 Tetracarcinus subquadratus Weller, 1905 Fig. 1G, K Diagnosis. —Carapace small, subquad- rate, widest across epibranchial region, length nearly equals width; sides sinuous; areolations low, granulate, cardiac region large; weakly differentiated by smooth grooves, particularly peripherally, on ce- phalic arch, and deeper on scapular arch. Claws equal, inflated proximally, fingers small. Seorsus, new genus Seorsus wadei, new species Fig. 1A-F Genus Avitelmessus Rathbun, 1923 Avitelmessus grapsoideus Rathbun, 1923 Fig. 1J Diagnosis. —Carapace very large, flat, cir- cular, widest at midlength anterolateral margin spinose. Grooves poorly developed except for those delimiting sagittal ridge and epimeral peninsula; sagittal ridge well-dif- ferentiated, epibranchial somewhat tumid, other regions not well-differentiated; regions relatively level, especially peripherally. Claws equal, short, crested, granulate; fin- gers large, downturned, fluted, and toothed. VOLUME 101, NUMBER 1 Biogeography Dakoticancroid crabs are confined to the latest Cretaceous rocks (Santonian-Maas- trichtian) of North America on the shallow shelves of the Tethys Sea. The taxa are more or less endemic but with boundaries of their distributions somewhat overlapping. 7 etra- carcinus subquadratus is abundant on the Northern Atlantic Coastal Plain and in the Mississippi Embayment, with but one rec- ord in the Western Interior Seaway. Avitel- messus grapsoideus is found on the South- ern Atlantic Coastal Plain and is abundant in the Mississippi Embayment where it oc- curs in the dominant Avite/messus Assem- blage in thin layers over large areas. The unusually large size of this crab, its abun- dance, and its occurrence in laminated shales and calcareous sandstones are correlated with yet undefined environmental condi- tions. Dakoticancer australis is the predom- inant decapod in the D. australis Assem- blage of Northern Mississippi (Bishop 1983), and Seorsus wadei is a minor element in it. Dakoticancer overanus is the predominant decapod in the repeated D. overanus Assem- blages of the middle Western Interior Sea- way (Bishop 1981). Little has been substan- tiated about evolutionary development of the dakoticancroid crabs. It is apparent, however, that Jetracarcinus is the oldest known dakoticancroid, with abundant San- tonian—Campanian records on the Northern Atlantic Coastal Plain, and that Dakotican- cer probably evolved from Tetracarcinus and migrated into the Western Interior dur- ing the Late Campanian and into the Mis- sissippi Embayment during the Maastrich- tian (Bishop 1986b:129). Acknowledgments The specimen described was collected with the direct support of the Society of the Sigma Xi (Lisa Juray, Grant-in-Aid) and the indirect support of Georgia Southern Col- lege through its Faculty Research Commit- Vy tee. Special thanks go to Mr. Ralph Harris of Blue Springs for acting as guide and to the students who accompanied me on this trip. The manuscript was read and strength- ened by criticism from an anonymous re- viewer. Darkroom assistance was provided by Jeanne Amos, and the manuscript was typed by Mrs. Donna Cain and Mrs. Judith McQuaig. Literature Cited Bishop, G. A. 1981. Occurrence and fossilization of the Dakoticancer Assemblage, Upper Creta- ceous Pierre Shale, South Dakota. Pp. 383-413 in J. Gray, P. F. Boucot, and W. B. N. Berry, eds., Communities of the past. Hutchinson Ross Publishing Co., Stroudsburg, Pennsylvania. . 1983. Fossil decapod Crustacea from the Late Cretaceous Coon Creek Formation, Union County, Mississippi.—Journal of Crustacean Biology 3(3):417-430. 1985. A new crab, Prehepatus harrisi (Crus- tacea, Decapoda) from the Coon Creek and Prai- rie Bluff Formations, Union County, Mississip- pi.—Journal of Paleontology 59(5):1028-1032. 1986a. Taphonomy of the North American Decapods.— Journal of Crustacean Biology 6(3): 326-355. 1986b. Occurrence, preservation, and bio- geography of the Cretaceous crabs of North America. Pp. 111-142 in R. H. Gore and K. L. Heck, eds., Crustacean issues 4: Crustacea bio- geography. Balkema Press, Rotterdam, The Netherlands. Glaessner, M. F. 1960. The fossil decapod Crustacea of New Zealand and the evolution of the Order Decapoda.—New Zealand Geological Survey Paleontoiogical Bulletin 31:1-63. 1969. Decapoda. Pp. R399-R533 in R. C. Moore, ed., Treatise on invertebrate paleontol- ogy, Part R, Arthropoda 4, volume 2. Univer- sity of Kansas Press (Lawrence) and the Geo- logical Society of America. Rathbun, M. J. 1917. New species of South Dakota Cretaceous crabs. — Proceedings of the U.S. Na- tional Museum 52(2182):385-391. 1923. Decapod crustaceans from the Upper Cretaceous of North Carolina. — North Carolina Geological and Economic Survey 5:403-408. 1926. Decapoda. Jn B. Wade, the fauna of the Ripley Formation on Coon Creek Tennes- see.— U.S. Geological Survey Professional Pa- per 137:184-191. 1935. Fossil Crustacea of the Atlantic and 78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Gulf Coastal Plain.—Geological Society of Weller, S. 1905. The fauna of the Cliffwood (N.J.) America, Special Paper 2:1—160. clays.—Journal of Geology 13(4):324—337. —. 1937. Cretaceous and Tertiary crabs from ———. 1907. A report on the Cretaceous Paleon- Panama and Colombia.—Journal of Paleontol- tology of New Jersey. — Geological Survey of New ogy 11(1):26-28. Jersey, Paleontology Series 4:846-853. Russell, E. E., D. M. Keady, E. A. Mancini, & C. C. Smith. 1983. Upper Cretaceous lithostratig- Institute of Arthropodology and Parasi- raphy and biostratigraphy in northeast Missis- sippi, southwest Tennessee and northwest Al- tology, Department of Geology and Geog- abama, shelf chalks and coastal clastics. Spring ! aphy, Georgia Southern College, States- Field Trip, April 7-9, 1983.—Gulf Coast Sec- boro, Georgia 30460-8149. tion, Society of Economic Paleontologists and Mineralogists, 72 pp. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 79-82 SCIONELLA PAPILLOSA, A NEW SPECIES OF POLYCHAETE (POLYCHAETA: TEREBELLIDAE) FROM THE SOUTHWEST FLORIDA CONTINENTAL SHELF Gary Tourtellotte and Henry Kritzler Abstract. —Scionella papillosa, a new species of terebellid polychaete from the southwest Florida continental shelf is described. It more closely resembles the only other Atlantic species, S. /ornensis Pearson, 1969, than the Pacific species, S. japonica Moore, 1903, S. estevanica Berkeley & Berkeley, 1942, and S. vinogradovi (Ushakov), 1955, in lacking a ridge on segment 4, but it differs from all four in having densely packed small digitiform papillae on the ven- trolateral surfaces of the second segment. From collections made in December, 1983 on the southwest Florida continental shelf by Environmental Science and Engi- neering, and LGL Ecological Research As- sociates under a contract with the Minerals Management Service, U.S. Department of the Interior, there were segregated a number of specimens of an unidentifiable terebellid polychaete whose closest affinities seemed to be with the genus Scionella Moore, 1903. This genus had hitherto been known only from the cold waters of the North Pacific and Scotland. Family Terebellidae Subfamily Amphitritinae Scionella Moore, 1903 Type species. —Scionella japonica Moore, 1903, by monotypy. Material examined. —North Atlantic Ocean, Gulf of Mexico, 26°17'4.5’N, 082°19'57”W, 16 m, 6 Dec 1983 (Environ- mental Science and Engineering, Inc., Tam- pa, Florida and LGL Ecological Research Associates, Inc., collectors). Holotype, USNM 99376, paratype USNM 99377, nu- merous other specimens in the collections of the senior author. Description. —Holotype, complete speci- men with 20 thoracic and about 65 abdom- inal segments, 17 mm long and | mm wide. Body cylindrical in section. No distinct ven- tral glandular scutes. Fused prostomium and peristomium forming a collar surrounding terminal mouth. Numerous tentacles aris- ing from dorsal and lateral surfaces of collar. Eyespots lacking. First segment bearing small lateral lappets. Second segment hav- ing large, translucent lateral lappets, ven- trolateral surfaces of which covered with densely packed, small digitiform papillae (Fig. 1a). Second segment also with median, V-shaped, opaque pad accentuating ap- pearance of lateral lappets of this and fol- lowing segments. From anterior dorsal edge of second segment, one pair of bottle-brush branchiae arising, each consisting of stout, ringed stalk bearing on its distal *4 a cluster of filaments arranged in whorls, too densely packed to reveal manner of insertion (Fig. 1b). All specimens having one small and one large branchia, smaller being about 3 size of larger. In holotype, larger equal in length to first 7 segments. Third segment bearing largest lateral lap- pets. Smallest on fourth, on which appear- ing first notosetae, continuing through seg- ment 20. In anterior setigers, notopodia very 80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. D, Abdominal uncinigerous pinnule; E, Thoracic uncinus. short, each bearing 6—7 smooth-tipped bi- limbate setae. Limbs on one side slightly wider than on opposite (Fig. lc). In poste- rior thorax, notopodia somewhat more prominent, although still no more than short rounded protuberances bearing up to 12 similar setae. Avicular uncini, none of which having elongate basal stalks, as in Pista, borne in single rows of 25-30 on segments 5-10 and in double rows of 10-20, inter- locking face to face, on segments 1 1-20. Ab- dominal uncini occurring in single rows, having prominent suspensory ligaments and borne on well-developed, flap-like pinnules (Fig. 1d). Throughout, uncini having several rows of denticles forming a crown above main fang (dental formula MF:8:9:10:~) (Fig. le). Terminal anus surrounded by 4 short, rounded protuberances. Scionella papillosa, holotype. A, Anterior end, ventral view; B, Anterior end, dorsal view; C, Notoseta; Etymology.—The specific name, papil- losa, derived from Latin, refers to the patches of digitiform papillae on the ventrolateral surfaces of the second segment. Remarks. —The most useful treatment of the genus Scionella is that of Pearson (1969). In his key (pp. 513-514) the four species then known fall into two groups, those with a dorsal ridge on segment 4 (S. japonica Moore, 1903, S. estevanica Berkeley & Berkeley, 1942, and S. vinogradovi (Usha- kov, 1955)), and those without (then only S. lornensis Pearson, 1969). Perhaps some significance may be attached to the fact that the first group occurs only in the Pacific, whereas the second has been found only in Atlantic waters. Scionella papillosa, also an Atlantic species and lacking a dorsal ridge on the fourth segment, may be grouped with VOLUME 101, NUMBER 1 S. lornensis. Scionella papillosa, however, differs from the latter in having lateral lap- pets on the first 4 segments; in that the lat- eral lappets on the second segment are less prominent than those on the third, a con- dition which is reversed in S. /ornensis; in that it lacks well-developed ventral glan- dular scutes; and in that none of its gill fil- aments are dichotomously branched. It is not possible to state how S. papillosa compares with the others with respect to nephridia. This character which was given much weight by Pearson and was a principal basis for classification by Hessle (1917) has not been useful in dealing with the generally very small polychaetes from the Gulf of Mexico offshore continental shelf. Certain- ly, S. papillosa does not have the large bot- tle-shaped nephridial papillae on the fourth segment, as depicted by Hartman (1969) for a California specimen of S. japonica. The number of uncini per segment, much larger in S. japonica (up to 160 thoracic and 65 abdominal) than in S. papillosa, may not be a significant difference, the numbers pos- sibly being a function of the size of the spec- imens. The collar-like fused prostomium and peristomium of S. papillosa bears little re- semblance to the trefoil-shaped structure in S. japonica, as described by Moore and fig- ured by Hartman (1969:635, fig. 1) or, pre- sumably, to these structures in S. esteva- nica. Were it not for the dorsal ridge on the fourth segment, S. estevanica would easily fit in with the S. papillosa, S. lornensis group, for its gills also arise from segment 2 and, apparently, its lateral lappets lack the alate configuration and increasingly more dorsal insertion which set off the other two species so plainly. In any case, S. papillosa is distinguished from all the others by the patches of papillae on the second segment. Accordingly, Pearson’s key may be up- dated as follows: 81 Key to the Species of Scionella Moore, 1903 la. Segment 4 with a prominent Gorsalenidgememn 3 Ib. Segment 4 without aridge.. 2 2a (1b). Lateral lappets on the first three segments S. lornensis Pearson 2b (1b). Lateral lappets on the first four segments ....S. papillosa, n. sp. 3a (la). Gills carried on segment4 . 4 3b (la). Gills carried on segment 2 .. Ry Meester: S. estevanica Berkeley & Berkeley 4a (3a). Edges of segment 4 lappets Crenulaten.) eee See ane bans S. vinogradovi (Ushakov) 4b (3a). Edges of segment 4 lappets smooth ..... S. japonica Moore Ecology and distribution. — Scionella pap- illosa has been dredged from relatively shal- low depths in the subtropical waters of the Gulf of Mexico, whereas the other four species have been found in greater depths in the cold waters off California, Japan, Western Canada, the Sea of Okhotsk, and Scotland. It lives in mucous tubes adorned with fine sand grains, buried in medium to fine, mainly calcareous, sediments. The sa- linity of the overlying water was 35.1%o and the dissolved oxygen 7.3 mg/liter. It is as- sociated with other polychaetes: Cirropho- rus sp., Axiothella mucosa, Mediomastus californiensis, Exogone dispar, Pseudover- milia sp.; the amphipods Ampelisca sp., Ti- ron tropakis, and Eudeuenopus honduranus; the cumaceans Oxyurostylis smithi, and Cy- claspis spp.; the decapod Lucifer faxoni; and the cephalochordate Branchiostoma cari- baeum. Acknowledgments The authors are indebted to Pat Hutch- ings and Marian Pettibone for reviewing the manuscript, and to Kristian Fauchald for 82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON his assistance and encouragement. The specimens were collected under U.S. De- partment of the Interior Minerals Manage- ment Service Contract No. 14-12-001- 30071. Literature Cited Berkeley, E., & C. Berkeley. 1942. North Pacific Poly- chaeta, chiefly from the west coast of Vancouver Island, Alaska, and Bering Sea.— Canadian Journal of Research 20:183-208, 6 figs. 1952. Annelida Polychaeta sedentaria. Ca- nadian Pacific Fauna. — Fisheries Research Board of Canada 9b(2):1-139. Hartman, O. 1969. Atlas of the sedenteriate poly- chaetous annelids from California. Allan Han- cock Foundation, University of Southern Cal- ifornia, Los Angeles, California, 812 pp. Hessle, C. 1917. Ziir Kenntnis der terebellomorphen Polychaeten.—Zoologiska Bidrag fran Uppsala 5:39-258. Moore, J. P. 1903. Polychaeta from the coastal slope of Japan and from Kamchatka and Bering Sea. — Proceedings of the Academy of Natural Sci- ences, Philadelphia 55:401—490, pls. 23-27. Pearson, T. H. 1969. Scionella lornensis sp. nov., a new terebellid (Polychaeta: Annelida) from the west coast of Scotland with notes on the genus Scionella Moore, and a key to the genera of the Terebellidae recorded from European waters. — Journal of Natural History 3:509-516. Ushakov, P. 1955. (Polychaetous annelids of the far eastern seas of the USSR.) [In Russian.]— Aka- demiya Nauk SSSR, Keys to the Fauna of the USSR 56:1-433 (translated 1965 by the Israel Program for Scientific Translations, Jerusalem). (GT) Environmental Science and Engi- neering, Inc., P.O. Box 23601, Tampa, Flor- ida, 33623-3601; (HK) Department of Oceanography, Florida State University, Marine Laboratory, Route 1, Box 219A, Sopchoppy, Florida 32358. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 83-86 STREPTOSPINIGERA ALTERNOCIRRUS, A NEW SPECIES OF POLYCHAETA (SYLLIDAE: EUSYLLINAE) Takashi Ohwada Abstract. —A new species of Eusyllinae, Streptospinigera alternocirrus, from Aburatsubo Bay on the Pacific coast of Central Japan is described. Although itis most closely related to S. heteroseta Kudenov in possessing enlarged acicula, composite spinigers, and both superior enlarged and capillary simple setae, it is distinguished from the latter in the morphology and arrangement of the dorsal cirri. Streptospinigera Kudenov, 1983, was de- scribed as a genus closely related to Strep- tosyllis Webster & Benedict, 1884, in having enlarged anterior setae, but differing from the latter in possessing composite spinigers and superior capillary simple setae. During the course of my study on the settlement and recruitment of polychaete larvae (Ohwada 1986) numerous tiny syllids were collected. Some of them fit well in the genus Streptospinigera, but differ from S. hetero- seta Kudenov, the only known species in the genus, in their morphology and arrange- ment of the dorsal cirri. They are described below as a new species. Type specimens are deposited in the Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. (USNM), British Museum (Natural Histo- ry), London (BM(NH)), and National Sci- ence Museum, Tokyo (NSMT). Streptospinigera alternocirrus, new species Bigs. 2 Material examined. — Misaki, Kanagawa, Japan: Aburatsubo Bay (35°09'18’N, 139°36'55”E), 2 m, clean medium sand, coll. T. Ohwada, 19 Nov 1984, holotype (USNM 01494); 22 Oct 1984, paratype (USNM 01495); 25 Jan 1985, paratype (BM(NH)ZB 1986.695); 20 May 1985, paratype (NSMT- Pol. P 236). Material examined represents a selection of material available. Description. — Holotype, sexually mature, 1.37 mm long, 0.23 mm wide without para- podia at middle widest part of body, with 32 setigers. Three paratypes 1.09 to 1.75 mm long, 0.10 to 0.17 mm wide with 26 to 33 setigers. Body generally lacking pigmen- tation in alcohol preservation. Prostomium wider than long, with palps large, fused basally, directed anteroventral- ly, laterally incised, distally rounded, lack- ing palpal cirri (Fig. la, b). Three antennae smooth, clavate; with long median antenna in middle of prostomium, lateral antennae lateral to small ocular spots (Fig. la). Four lenticular eyes in trapezoidal arrangement (in holotype, anterior right eye as 2 separate spots, in paratype (USNM 01495), anterior left eye as 2 separate spots), and 2 small ocular spots lateral to bases of lateral an- tennae (Fig. la, b). Peristomium a complete ring dorsally and laterally, with 2 pairs of smooth clavate tentacular cirri; dorsal pair slightly longer than ventral pair. Pharynx completely everted in paratype (USNM 01495), unarmed, thickly sclero- tized, distal border surrounded by 10 pa- pillae proximal to smooth anterior end of sclerotized part (Fig. 1b), extending to about middle part of setiger 3, almost as long as proventriculus. Proventriculus barrel- shaped, about twice as long as wide, ex- 84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Streptospinigera alternocirrus: a, c, holotype (USNM 01494), b, paratype (USNM 01495), c-f, paratype (BM(NH)ZB 1986.695), g, paratype (NSMT-Pol. P 236): a, Anterior half, dorsal view, pharynx partially everted; b, Dorsal view, anterior end with pharynx everted, antennae not shown (positions dotted); c, Parapodium from setiger 3, dorsal view; d, Same, anterior view; e, Parapodium from setiger 20, anterior view; f, Parapodium from setiger 25, anterior view; g, Posterior end, dorsal view, long right anal cirrus missing. tending from setiger 7 through 10 in holo- Parapodia of setigers | to 5 large, distally type whose pharynx partially everted (Fig. truncate (Fig. lc, d). Parapodia from setiger la), with about 30 transverse rows ofmuscle 6 smaller and conical (Fig. la, e, f). Dorsal points, also arranged in 2 opposite, diagonal cirri of setiger 1 long, resembling tentacular planes. cirri but slightly longer; those of setigers 2, VOLUME 101, NUMBER 1 3, 5 smooth, cirriform (Fig. la, c, d). In setiger 4 dorsal cirri longer, with 2 articu- lations (Fig. la). Dorsal cirri of setiger 6 with 4 rather large articles (Fig. 1a). Dorsal cirri of setigers 7 and 8 smooth, thick, sub- ulate, thereafter as long as setae with 4-6 large articles (Fig. 1f) on odd-numbered se- tigers and smooth, thick, subulate, extend- ing beyond parapodial lobes (Fig. le) on even-numbered setigers to end of body (Fig. la, g). Each article containing dense yellow- ish-brown granules. Ventral cirri smooth throughout, short, blunt, inflated on setigers 1-3 (Fig. 1d), thereafter becoming elongate to setiger 7, then subulate to posterior end (Fig. le, f). Neuroacicula of setigers 2—5 twice as thick as followings, with dorsally curved tips (Fig. 2b-e, h). Those from setigers 1 and 6, to end of body slender, distally beak-shaped (Fig. 2a, f, g). Notoacicula very slender, needle-like (Fig. 21) present in all setigers; those in few anterior setigers especially thin. Superior simple seta present in all seti- gers. Those of setigers 1-5 distally falcate, rather short, as long as shafts of compound spinigers; those of setigers 2—5 enlarged (Figs. Ic, d, 2j). From setiger 6 to end of body, stout simple seta replaced by long slender capillary seta, slightly curved dorsally and finely serrated on lower curved side (Figs. le, f, 2k) (sometimes transitional forms between two types of simple setae on setiger 6). Setigers 1-5 each with 10-11 thick com- posite setae including 6-9 falcigers having short unidentate blades (Fig. 21, n) and 2—4 spinigers having long blades (Fig. 2m), all with saw-tooth cutting margins, lacking dis- tal hoods and sheaths; those of setigers 2— 5 thicker than those of setiger 1. Shaft tips smooth with rather pointed superior branch and rounded inferior branch; cup-shaped socket for blade between two branches (Fig. 20). Setigers 6 to end of body each with 6-9 slender composite setae including 2-3 su- perior spinigers having very long finely ser- 85 0.02 mm n q k Fig. 2. Streptospinigera alternocirrus, paratype (BM(NH)ZB 1986.695):a—g, Neuroacicula, dorsal views: a, Setiger 1; b, Setiger 2; c, Setiger 3; d, Setiger 4; e, Setiger 5; f, Setiger 6; g, Setiger 20; h, Neuroaciculum, setiger 3, anterior view; 1, Notoaciculum, setiger 20, anterior view; J, Enlarged superior simple seta, setiger 4, posterior view; k, Superior capillary seta, setiger 10, posterior view; 1, Medial composite falciger, setiger 1, anterior view; m, Upper composite spiniger, setiger 1, anterior view; n, Medial composite falciger, setiger 5, posterior view; 0, Shaft tip of same, dorsoposterior view; p, Superior composite spiniger, setiger 15, pos- terior view; q, Medial composite spiniger, setiger 16, posterior view; r, Inferior composite falciger, setiger 22, posterior view. rated blades of similar length (Fig. 2p), 2- 4 medial spinigers having long minutely ser- rated blades (Fig. 2q) and 2—4 inferior fal- cigers having rather short unidentate blades with minutely serrated cutting margins (Fig. 2r) becoming shorter ventrally (Fig. 1f); all lacking distal hoods and sheaths. Medial spinigers sometimes with basal spur (Fig. 86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2q). Shaft tips smooth, similar to those of setigers 1-5, but socket for blade generally inconspicuous, not recognized in superior spinigers. Inferior simple seta absent throughout body. Pygidium very small with 3 smooth anal cirri including short filiform midventral and 2 long lateral ones (Fig. 1g). In holotype, gametes present from setiger 10 to end of body. Remarks. —Streptospinigera alternocir- rus agrees with S. heteroseta in having en- larged acicula in the anterior region, com- posite spinigers and both enlarged and capillary superior simple setae. Streptospi- nigera alternocirrus, however, differs from S. heteroseta in having not only articulate dorsal cirri but also subulate dorsal cirri af- ter setiger 7. In addition, while articulate dorsal cirri become long only on setigers 6, 9, 11, 13, 16, 18 in S. heteroseta, long ar- ticulate dorsal cirri occur on setiger 6 and, alternately with smooth thick subulate dor- sal cirri from setiger 9 to end of body in S. alternocirrus. Dorsal cirri of setiger 4 are short, smooth, subulate to cylindrical in S. heteroseta, and rather long with two articles in S. alternocirrus. Streptosyllis pettiboneae Perkins, 1980, agrees with Streptospinigera alternocirrus in having similar arrangement of articulate and smooth dorsal cirri; S. pettiboneae, how- ever, has only composite falcigers and stout superior simple setae. Ecology. —Streptospinigera alternocirrus occurs abundantly in clean medium to fine sands. Epitokes were occasionally found. In these epitokes, four lenticular eyes are en- larged, and several very long natatory setae develop superior to usual capillary simple seta on the segments swollen with repro- ductive material. Larval settlement oc- curred mainly from April to June in the type locality, and the highest density of 25,000 inds./m? was recorded during this settling season (Ohwada 1986). The adult popula- tion disappeared soon after the breeding season. Etymology.—The specific name, alter- nocirrus, derives from the Latin, alterno, meaning alternate, and cirrus meaning curls; it is used as a noun in apposition. Acknowledgments I am indebted to the staff of the Misaki Marine Biological Station, University of Tokyo for their assistance in the sampling. Dr. Marian Pettibone, Smithsonian Insti- tution, kindly reviewed the manuscript and offered invaluable suggestions. Literature Cited Kudenov, J..D. 1983. ‘Streptospinigera heteroseta, a new genus and species of Eusyllinae (Polychaeta: Syllidae) from the western shelf of Florida.— Proceedings of the Biological Society of Wash- ington 96:84—-88. Ohwada, T. 1986. The importance of settling period in the determination of the distribution of poly- chaetous annelids in the shallow-water sediment bottom. Dr.Sc. thesis, University of Tokyo, 111 pp. Perkins, T.E. 1980. Syllidae (Polychaeta), principally from Florida, with descriptions of a new genus and twenty-one new species.—Proceedings of the Biological Society of Washington 93:1080- 1172. Webster, H. E. & J. E. Benedict. 1884. The Annelida Chaetopoda from Provincetown and Wellfleet, Massachusetts.—Annual Report of the Com- missioner for Fish and Fisheries for 1881:699— 747, 8 pls. Ocean Research Institute, University of Tokyo, 15-1, 1-Chome, Minamidai, Na- kano-ku, Tokyo, 164 Japan. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 87-94 NEOTROPICAL MONOGENEA. 11. RHINOXENUS, NEW GENUS (DACTYLOGYRIDAE: ANCYROCEPHALINAE) WITH DESCRIPTIONS OF THREE NEW SPECIES FROM THE NASAL CAVITIES OF AMAZONIAN CHARACOIDEA D. C. Kritsky, W. A. Boeger, and V. E. Thatcher Abstract. — Rhinoxenus, new genus is proposed for ancyrocephalines char- acterized by 1) possessing a pair of dorsal haptoral spikes which replace or are modifications of the dorsal anchor/bar complex; 2) lacking a dorsal bar; and 3) having hook pair 2 situated on two bilateral lobes of the trunk. Species of Rhinoxenus infest the nasal cavities of Neotropical charcoid fishes. Three new species are described: R. piranhus, new species, from Serrasalmus nattereri (Kner), R. arietinus, new species, from Schizodon fasciatum Agassiz and Rhy- tiodus argenteofuscus Kner, and R. nyttus, new species from S. fasicatum. Two undescribed species of Rhinoxenus are reported from S. fasciatum and Hydro- lycus scomberoides (Cuvier), respectively. Relatively few species of Dactylogyridae have been recorded from nasal cavities of freshwater fishes. In North America, Rogers (1967) described Aplodiscus nasalis (Dac- tylogyrinae) from the nose of the Alabama hogsucker, Hypentelium etowanum (Cato- stomidae). Dechtiar (1969) described two species of Dactylogyrinae, Pellucidhaptor catostomi and P. nasalis, from the nasal cavities of Catostomus catostomus and C. commersoni (Catostomidae), respectively, in Ontario. Recently, Klassen and Beverley- Burton (1985) reported Ligictaluridus mon- ticellii (Ancyrocephalinae) from the nose of Ictalurus nebulosus (Ictaluridae) in Ontario. The latter species had previously been re- ported from the nasal cavities of North American Ictaluridae imported into Europe (Cognetti de Martiis 1924, Prost 1973). Ad- ditional records of Dactylogyridae from the nose of freshwater fishes exist from Europe, the USSR, and Asia (Smirnova et al. 1964, Strelkow & Ha Ky 1964, Gussev and Strijak 1972). At present there are no reports of dactylogyrids from the nares of Neotropical or Ethiopian freshwater fish. Recently, we examined the nasal cavities of a few species of freshwater fish collected from the Manaus area in Brazil. These pre- liminary collections from four species of Characoidea, one species of Siluriformes and one clupeid host have revealed several new monogenean species and genera, suggesting that the nose of fishes may be a specific site of infestation for an unique fauna in the Neotropical Region. The present study in- cludes the description of dactylogyrids com- prising a new genus from the characoid hosts. Materials and Methods Fish were collected by net from the en- virons of Manaus, Amazonas, Brazil, dur- ing 1983 and 1984. The host brain and spi- nal cord were destroyed by pithing, after which the nasal cavities of each fish were washed with a strong current of 1:4000 for- malin solution. After this washing, the nasal cavity was disrupted with a probe and again washed with the formalin solution. The washings were collected and examined with a dissecting microscope; parasites were re- moved with a fine probe and fixed in a 3% formalin solution. Preparation of helminths 88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON for study followed the procedures outlined by Kritsky, Thatcher & Boeger (1986). Mea- surements, all in micrometers, were made with the aid of a filar micrometer, except length of the cirrus was approximated using a Minerva curvimeter on camera lucida drawings. Type specimens were deposited in the helminth collections of the Instituto Nacional de Pesquisas da Amazonia (INPA), the U.S. National Museum (USNM), and the University of Nebraska State Museum (HWML). Rhinoxenus, new genus Diagnosis. —Dactylogyridae, Ancyroce- phalinae. Body robust, divisible into ce- phalic region, trunk, haptor (peduncle ab- sent). Tegument thin, smooth. Cephalic lobes, head organs, cephalic glands present. Four eyes. Mouth subterminal, midventral; pharynx muscular, glandular; esophagus short; intestinal caeca (2) confluent poste- rior to gonads, lacking diverticulae. Genital pore midventral. Gonads intercaecal, over- lapping; testis dorsal to ovary. Vas deferens looping left intestinal caecum; seminal ves- icle a dilation of vas deferens, frequently with constriction near midlength; copula- tory complex comprising coiled cirrus with counterclockwise rings, accessory piece ba- sally articulated to cirrus by proximal pro- jection lying within rings of cirrus. Vagina sinistral in anterior trunk; seminal recep- tacle present. Vitellaria well developed, coextensive with caeca. Haptor armed with ventral pair of anchors, ventral bar, 6 pairs of hooks with ancyrocephaline distribution (Mizelle 1936) (hook pair 2 removed from haptor, lying in lateral or ventrolateral lobes on posterior half of trunk), one pair of dorsal spike-like structures which replace or rep- resent modifications of dorsal anchors. Hooks with proximally inflated shank. Na- sal parasites of Neotropical characoid fish- es. Type species, host, and locality. —Rhi- noxenus piranhus, n. sp. from Serrasalmus nattereri (Kner), Ilha Marchantaria, Rio So- lim6es, near Manaus, Amazonas, Brazil. Other species. — Rhinoxenus arietinus, n. sp. from Schizodon fasciatum Agassiz (type) and Rhytiodus argenteofuscus Kner (an ap- parent accidental host); Rhinoxenus nyttus, n. sp. from Schizodon fasciatum; R. spp. from Schizodon fasciatum and Hydrolycus scomberoides (Cuvier) (USNM 79264, 79265, respectively). Etymology.—The generic name is from Greek (rhin/o = nose + xen/o = guest). Rhinoxenus piranhus, new species Figs. 1-8 Host. —Serrasalmus nattereri (Kner), the piranha caju, Serrasalmidae. Locality. —Iha Marchantaria, Rio Soli- moes near Manaus, Amazonas, Brazil (type) 21 Sep 1983, 15 Aug 1984, 14 Sep 1984, 25 and 26 Nov 1984; Furo do Catalao, near Manaus, Amazonas, Brazil 27 Nov 1984. Type specimens. —Holotype, INPA PA287-1; paratypes, INPA PA287-2 to PA287-11, USNM 79180, 79181, HWML 23307. Description (based on 79 specimens; 20 measured). — Body 758 (582-1014) long, fu- siform, slightly flattened dorsoventrally; greatest width 165 (132-206) in posterior trunk. Cephalic lobes poorly developed, 2 terminal, 2 bilateral; 2 pairs of head organs lying in cephalic lobes and adjacent cephalic area; cephalic glands unicellular, bilateral, posterolateral to esophagus. Eyes usually equidistant, members of anterior pair smaller than those of posterior pair; con- spicuous lens associated with posterior eyes; eye granules ovate, small; accessory gran- ules absent. Pharynx ovate, with distal suck- er-like modification, 34 (27-39) in diame- ter. Haptor trapezoidal, 113 (93-136) wide, 111 (86-135) long. Ventral anchor 131 (120- 137) long, with superficial root modified into ventromedial protuberances serving as ar- ticulation point for bar, evenly curved shaft, point with saucer-like termination; base 25 VOLUME 101, NUMBER 1 (20-32) wide. Anchor filament conspicu- ous, basal. Dorsal haptoral spike 116 (97- 130) long, with small bulbous proximal end, tapered distal point; each termination with conspicuous cap of tissue. Ventral bar 52 (46-55) long, flattened, with small terminal protuberances. Hook pair 2—24 (23-25) long, lying in small lobe on ventral surface of pos- terior trunk, stout, with heavy point, erect thumb, inflated proximal 7% of shank. Hap- toral hooks 27 (24-29) long, elongate, del- icate, with curved point, erect thumb, shank slightly inflated proximally. FH loop ex- tended to near beginning of shank dilation. Cirrus a coil of about 2'4 rings; base with bilaterally sclerotized lobes, ring diameter 28 (25-31), cirrus length 175. Accessory piece spathulate distally, with lateral rod- like thickening, proximal projection ex- tending along distal one-half cirral ring serving as cirral guide; length of distal spath- ulate portion 51 (45-55). Testis elongate ovate, 126 (115-144) long, 36 (34—37) wide; seminal vesicle with medial constriction; prostatic reservoirs not observed. Ovary elongate, with varying diameter, greatest width 34 (23-48), length 183 (137-202); oviduct short; ootype, uterus not observed; genital pore posterior to intestinal bifurca- tion; vagina with distal double cavity from which simple tube arises; seminal receptacle a dilation of vaginal tube, lying diagonally to left of body midline. Egg lacking fila- ments, with proximally thickened shell. Remarks. —Rhinoxenus piranhus is the type species for the genus. The specific name is derived from the common or local name of the host. Rhinoxenus arietinus, new species Figs. 9-17 Hosts. — Schizodon fasciatum Agassiz, the aracu pintado, Anostomidae (type), and Rhytiodus argenteofuscus Kner, the aracu pau de negro, Anostomidae. Locality.—UWha Marchantaria, Rio Soli- moes, near Manaus, Amazonas, Brazil (type) 89 S. fasciatum, 25 Nov 1983; aquarium, INPA, Manaus, Amazonas, Brazil (R. ar- genteofuscus, 8 Feb 1984. Type specimens.—Holotype, INPA PA288-1; paratypes, INPA PA288-2 to PA288-4, USNM 79183, 79184, HWML 23309, 23310. Description (based on 17 specimens).— Body flattened dorsoventrally, with lateral margins of trunk curled ventrally (Fig. 10); length 395 (310-501) greatest width 169 (133-202) in posterior trunk at level of go- nads. Cephalic area with 2 poorly developed terminal lobes; head organs conspicuous, usually 2 pairs; cephalic glands indistinct, posterolateral to pharynx. Eyes subequal; members of anterior pair approximately twice the distance apart of members of pos- terior pair; eye granules ovate, small; ac- cessory granules absent or few in cephalic region. Pharynx spherical, 23 (20-27) in di- ameter. Haptor subquadrate, with 2 large posteriorly directed pads shaped as horns of a ram; pads dense staining, apparently glandular; haptor 126 (111-138) wide, 108 (89-131) long. Ventral anchor 106 (88-115) long, robust, with large base, well-devel- oped roots, evenly curved shaft and point, obtuse point tip; base 54 (50-61) wide. An- chor filament well developed. Dorsal hap- toral spike 110 (89-126) long, variable in width, robust, with distal acute tip, proxi- mal and distal tissue caps. Ventral bar 43 (41-46) long, subrectangular, with slightly expanded ends. Hook pair 2 lying on lateral lobe of trunk just posterior to body mid- length, with heavy point, erect thumb, prox- imal %3 of shank inflated. Haptoral hooks elongate, delicate, with blunt thumb, prox- imal 34 of shank dilated. FH loop extending to near level of termination of shank infla- tion. Cirrus a coil of about 1! rings, base with large sclerotized lobes, ring diameter 29 (26-33), cirrus length 107-110. Acces- sory piece basally articulated to cirral base, proximally follows first cirral ring, with or- nate termination; length of terminal portion 32 (27-36). Testis ovate, 80 (78-83) long, 90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-8. Rhinoxenus piranhus: 1, Holotype (ventral), 2, Vagina (dorsal); 3, Copulatory complex (dorsal); 4, Ventral bar; 5, Dorsal haptoral spike; 6, Hook pair 2; 7, Haptoral hook; 8, Ventral anchor. All figures are draw to the 30-micrometer scale except Fig. 1 (200-micrometer). VOLUME 101, NUMBER 1 91 Figs. 9-17. Rhinoxenus arietinus: 9, Holotype (ventral); 10, Ventrolateral illustration showing relationship of haptoral anchors, pads, and spike, and bilateral body lobes; 11, 12, Copulatory complexes; 13, Ventral bar; 14, Hook pair 2; 15, Haptoral hook; 16, Dorsal haptoral spike; 17, Ventral anchor; Figs. 11-17 are drawn to the 30-micrometer scale; Fig. 9 to the 100-micrometer scale. 36 (29-43) wide, seminal vesicle with in- conspicuous medial constriction, one pros- tatic reservoir lying to right of cirral base. Ovary 85 (81-88) long, 21 (18-23) wide, usually an elongate sigmoid organ which may appear subovate in contracted speci- mens as a result of margins between curves being closely appressed; oviduct short; ootype, uterus not observed; genital pore posterior to or at level of intestinal bifur- cation; vagina sclerotized, tubular, with dis- tal loop, expanding proximally to large pyr- 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 18-24. Rhinoxenus nyttus: 18, Holotype (ventral); 19, Hook pair 2; 20, Haptoral hook; 21, Copulatory complex; 22, Dorsal haptoral spike; 23, Ventral bar; 24, Ventral anchor. All drawings are to the 30-micrometer scale except Fig. 18 (100-micrometer). iform seminal receptacle. Vitellaria dense, coextensive with gut. Remarks. —This species is unique among the Dactylogyridae by possessing two pos- terior lobes on the haptor. The lobes are apparently glandular and serve in attach- ment within the nasal cavity of the host. The specific name is from Latin (arietinus = like a ram’s head) and is based on the dorsal haptoral lobes which impart a ram’s head appearance to the haptor. Rhinoxenus nyttus, new species Figs. 18-24 Host. —Schizodon fasciatum Agassiz, the aracu pintado, Anostomidae. Locality. —Uha Marchantaria, Rio Soli- moes, near Manaus, Amazonas, Brazil 25 Nov 1983. Type specimens.—Holotype, INPA PA 286-1; paratypes, USNM 79182, HWML 23308. VOLUME 101, NUMBER 1 Description (based on 3 specimens).— Body foliform; length 334 (299-358), great- est width 92 (88-96) at various points along trunk. Cephalic lobes poorly developed or absent; 4 head organs in anterolateral ce- phalic region; cephalic glands inconspic- uous, lying posterolateral to pharynx. Mem- bers of posterior pair of eyes larger, closer together than those of anterior pair; eye granules ovate, small; accessory granules few or absent in cephalic area and anterior trunk. Pharynx subspherical, 24—25 in diameter; gut obscured by vitellaria. Haptor subrec- tangular, 81 (70-93) long, 66 (65-67) wide. Ventral anchor lacking roots, base with 2 superficial projections articulating to ven- tral bar, shaft curved, point undulated; an- chor 76 (71-80) long, base 25 (24—26) wide. Anchor filament well developed. Dorsal haptoral spike 59 (57-60) long, delicate, with distal hollow point, proximal and distal caps. Ventral bar subtrapezoidal, with short pos- terolateral protuberances; bar 41 (39-42) long. Hook pair 2-18 (16-20) long, lying on 2 bilateral lobes on posterior trunk, with delicate point, erect blunt thumb, shank di- lated along proximal half. Haptoral hooks 21 (20-22) long, delicate, with fine point, blunt thumb, slightly dilated proximal shank; FH loop extending to near beginning of dilated portion of shank. Cirrus com- prising a coil of 10-15 rings, simple base; ring diameter 35 (33-39); cirrus 1300 long. Accessory piece basally articulated to cirrus, proxima!ly twisted and lying within cirral rings, distally serving as a cirrus guide; ac- cessory piece 29-30 long. Testis, vas def- erens, prostatic reservoirs not observed; seminal vesicle pyriform. Ovary elongate, fusiform, 15-16 wide, 54-55 long; oviduct short; ootype, uterus not observed; vagina comprising an elongate coiled tube expand- ing into large fusiform seminal receptacle; vitellaria dense, scattered throughout trunk. Remarks. —Based on the morphology of the haptoral bar, the dorsal haptoral spikes, and the ventral anchor base, Rhinoxenus nyttus 1s most closely related to the type 93 species, R. piranhus. The coiled cirrus and vaginal tube, each comprising numerous rings, and the acute and doubly bent anchor point in R. nyttus serve to differentiate these species. The specific name is a neologism suggested by the Greek (Vyss/o = to prick) and refers to the presumed action of the dorsal haptoral spike. Discussion Although the morphology of the internal organ systems of Rhinoxenus species is not unique among the Dactylogyridae, the ge- nus is characterized by some outstanding features not found among members of other genera in the family. Most notable among these are: 1) the modification of the dorsal anchors into paired haptoral spikes which lack an articulating bar and 2) the removal of hook pair 2 from the haptor to bilateral lobes on the trunk. In a few species of Dac- tylogyridae (some Cosmetocleithrum spp., among others), hook pair 1 may be located on the peduncle, but this condition is not considered as significant as that shown by species of Rhinoxenus. In Cosmetocleith- rum, hook pair | remains in close proximity to the haptor and probably functions sim- ilarly to that of other dactylogyrids with normal hook distributions. Modifications of the haptors of Rhinoxenus species ap- parently relate to the needs for attachment determined by the physical surface of the host’s nares. However, collection tech- niques precluded a determination of how the parasites utilize their highly modified haptors at specific attachment sites within the nasal cavity. Determination of haptoral and hook 2 function in Rhinoxenus species will most likely depend on observations of living specimens on their hosts. The uniqueness of Rhinoxenus precludes determination of closely related genera, even though the genus clearly is a member of the Ancyrocephalinae (sensu Yamaguti 1963) based on anatomy of the internal organ sys- tems (digestive and reproductive). It is un- 94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON likely that it belongs in the Dactylogyrinae, which includes most North American par- asites from nasal cavities of fishes (Aplo- discus, Pellucidhaptor), since the anchor/bar complex is ventral (dorsal in Dactylogyri- nae), 4A hooks are lacking, and species of Rhinoxenus parasitize characoid fishes (pri- marily Cyprinidae in Dactylogyrinae). Acknowledgments This study was supported in part by the Max Plank Institute, Plon, Germany, dur- ing collection of fish hosts and the Conselho Nacional de Desenvolvimento Cientifico e Tecnologico through a study grant (20.0115/ 84) to WAB. Literature Cited Cognetti de Martiis, L. 1924. Nuovo gyrodactylide parassita nella cavita olfattiva di Amiurus catus L.-—Belletino Societa Naturalisti Napoli 36, Ser. II, 16:76-81. Dechtiar, A. O. 1969. Two new species of monoge- netic trematodes (Trematoda: Monogenea) from nasal cavities of catostomid fishes.— Journal of the Fisheries Research Board of Canada 26:865— 869. Gussev, A. V., & O. I. Strijak. 1972. New species of Pellucidhaptor (Monogenea) from Europe.— Parazitologiya 6:555-557. Klassen, G. J., & M. Beverley-Burton. 1985. Ligic- taluridus Beverley-Burton, 1984 (Monogenea: Ancyrocephalidae) from catfishes (Siluriformes: Ictaluridae) in North America with redescrip- tions of the type species, Ligictaluridus pricei (Mueller, 1936), and three others.—Canadian Journal of Zoology 63:715-727. Kritsky, D. C., V. E. Thatcher, & W. A. Boeger. 1986. Neotropical Monogenea. 8. Revision of Uro- cleidoides (Dactylogyridae, Ancyrocephali- nae). — Proceedings of the Helminthological So- ciety of Washington 53:1-37. Mizelle, J.D. 1936. New species of trematodes from the gills of Illinois fishes.— American Midland Naturalist 17:785-806. Prost, M. 1973. Fish Monogenoidea of Poland. II. Parasites of Ictalurus nebulosus (Lesueur). Re- vision of genera Cleidodiscus Mueller, 1934 and Urocleidus Mueller, 1934.— Acta Parasitologica Polonica 21:315-326. Rogers, W. A. 1967. Studies on Dactylogyrinae (Monogenea) with descriptions of 24 new species of Dactylogyrus, 5 new species of Pellucidhaptor, and the proposal of Aplodiscus gen. n.—Journal of Parasitology 53:501-524. Smirnova, K. V., Yu. A. Strelkov, V. A. Timofeeva, & S.S.Shul’man. 1964. The nasal cavities of teleost fish as a localization for parsites.—Zool- ogicheskii Zhurnal 43:1649-1658. Strelkov, Yu. A, & Ha Ky. 1964. A new instance of an unusual localization of the genus Dactylo- gyrus of the Monogenoidea in nasal cavities of fish. —Zoologicheskii Zhurnal 43:1236-1238. Yamaguti, S. 1963. Systema Helminthum. IV. Mon- ogenea and Aspidocotylea. Interscience Pub- lishers, Inc., New York. 699 pp. (DCK) Department of Allied Health Professions and Idaho Museum of Natural History, Idaho State University, Pocatello, Idaho 83209; (WAB) Instituto Nacional de Pesquisas da Amazonia, Manaus, Amazo- nas, Brazil, and Department of Biological Sciences, Idaho State University, Pocatello, Idaho 83209; (VET) Instituto Nacional de Pesquisas da Amazonia, Manaus, Amazo- nas, Brazil. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 95-101 AEGA LEPTONICA, A NEW SPECIES OF AEGID ISOPOD CRUSTACEAN FROM THE TROPICAL WESTERN ATLANTIC, WITH NOTES ON ROCINELA OCULATA HARGER AND ROCINELA KAPALA, NEW SPECIES Niel L. Bruce Abstract. —Aega leptonica, n. sp. is described and figured. The status and distribution of Rocinela oculata is reviewed and a new name, Rocinela kapala, is provided for Australian material previously identified as R. oculata. Remarks are given on the presence of the mandibular molar process in aegids, and the mouthparts of the family are rediagnosed. The Aegidae of the United States have been little studied, the only major work since Richardson’s (1905) monograph being that of Brusca (1983) on the East Pacific Aega species. The only recent new record of At- lantic aegids was of Aega monophthalma Johnston (Treat 1980). While examining collections at the Smithsonian I came across a new species of Aega, and also material identified as Roci- nela oculata. This Atlantic material, iden- tified as R. oculata, proved not to belong to that species, but also led to a re-evaluation of the status of Australian material identi- fied as R. oculata, which is here described as a new species. The following abbrevia- tions are used: AM-—Australian Museum; NSW—New South Wales, Australia; USNM—USS. National Museum of Natural History, Smithsonian Institution; QM— Queensland Museum. Mouthpart Morphology of the Aegidae Some confusion over the presence or ab- sence of the mandibular molar process in the Aegidae has appeared in recent litera- ture. Brusca (1983), in his diagnosis to the family, stated “mandible without lacinia mobilis, spine row, or molar process.”’ This was later repeated by Brusca and Iverson (1985), and Bowman (1986) stated that the aegid mandible lacked the molar process. This seems to be due to a misinterpretation of Brusca’s (1983) figs. That the family Ae- gidae possess a molar process has been am- ply demonstrated, most notably by Hansen (1890) who figured it for Barybrotes agilis (plate 9, fig. 3f), Aega psora (plate 9, fig. 4c, d) and Rocinela danmoniensis (plate 10, fig. 1b-e). The figs. for Aega vigilans. A. laevis, A. lethrina, A. coroo, A. beri, Rocinela ocu- lata and Alitropus typus given by Bruce (1983) all show a molar process on the man- dible. The molar process of Aega vigilans has small spines on the anterior margin. Aega leptonica described herein retains a very re- duced spine row, also noted in other aegids by Hansen (1890). There are several other aspects of mouth- part morphology frequently not noted. The mandible palp has 4 articles, the proximal one often fused to the second, also figured by Hansen (1890). The maxilla is made up of two articles, the large one usually termed lateral lobe, to which is attached the very much smaller medial lobe (there is never a basal lobe with two distinct terminal lobes as figured by Brusca (1983), this appearing to be an artifact caused by folding under a slide coverslip). The maxilla is essentially the same as that of the Cymothoidae. The PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 101, NUMBER 1 maxillule is nearly always figured without a medial lobe, but in Aega /eptonica this small lobe is present. The maxilliped of Aega has an article often not figured. This article is the coxa, the ar- ticle lateral to it is the epipod as figured and identified by Hansen (1890), and not part of a divided coxa. In males the epipod may be small, or with a few marginal setae; in females the maxilliped develops large lam- inar lobes, one from the basis and one from the coxa. The mouthparts of the Aegidae are here rediagnosed: Mandible incisor narrow, mo- lar process present, lacinia mobilis and spine row (usually) absent; palp of 3 or 4 articles, articles 1 and 2 often coalesced. Maxillule slender, styliform with terminal spines. Maxilla with 2 articles, small distomedial lobe joined to larger lateral lobe, each lobe with 2 or more apical spines. Maxilliped with endite and epipod; palp with 3-5 ar- ticles, at least articles 3 and 4 with large hooked spines. Aega leptonica, new species Figs. 1, 2 Material.—Female Holotype, 28.5 mm, off Tortugas, Florida, 30 Jul 1932, 1048 m, Fish Hawk sta 65-32 (USNM 227124). Type locality. —Off Dry Tortugas, Flori- da, U.S.A., ca. 24°00'N, 83°00'W. Description. — Body about 3 times as long as wide. Cephalon with median rostrum; eyes large, united, medially occupying about 0.25 length of cephalon. Pereonites 4—7 with partial transverse impressed line. Coxae be- coming more acute posteriorly; all coxae with oblique carina. Pleonites all visible; pleonites 1—4 each with horizontal carina. —_— Fig. 1. 97 Pleotelson slightly wider than long, lateral Margins convex, converging to narrow apex; each margin with 6 large spines and mar- ginal setae; apex with subrectangular exci- sion, lateral margins of which diverge. Antennule peduncle article 3 longest, ex- ceeding combined lengths of articles 1 and 2; peduncle article 4 fused to 3; flagellum with 17 articles, extending to middle of pe- reonite 1. Antenna peduncles articles 1-3 short; 4 and 5 about equal in length, each longer more than twice as long as article 3; flagellum with 27 articles, extending to an- terior to pereonite 3. Frontal lamina approximately pentago- nal, posterior part stem-like; anterior mar- gins slightly convex. Mandible incisor trun- cate; molar process prominent, vestigial spine row present; palp 4 articled, article 1 short, fused to article 2, distolateral margin of article 3 and most of lateral margin of article 4 with stiff setae. Maxillule with 1 broad-based triangular spine and 8 slender spines. Maxilla with 2 gently curved spines on endopod, exopod with 3 large curved spines, 1 curved and | straight spine. Max- illiped palp medial margins with 2 slender spines at distal angle of article 2, 5 spines and 4 setae on article 3, 4 large recurved spines on article 4, and 2 curves spines and 2 setae on article 5; endite with 4 setae. Pereopods all slender. Pereopods 1-3 with slender propodus and elongate dactylus which exceeds length of propodus; 2 small spines on posterior margin of carpus, | small spine on posterior margin of merus, | small spine on anterodistal angle of ischium. Pe- reopods 2 and 3 with more numerous and larger spines than pereopod 1. Pereopod 7 basis with evenly spaced setae along an- teromedial margin; posterior margins of is- Aega leptonica, holotype: A, Lateral view; B, Cephalon; C, Mandible; D, Mandible, detail; E, Clypeal region; F, maxilla and detail; G, maxillule and detail; H, Antennule; I, Antenna; J, Antennal flagellum, distal articles; K, Maxilliped; L, Maxilliped endite; M, Maxilliped palp; N, Pleotelson and uropods. Scale bar represents 5 mm. 98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pees \\e z Y \ ees K G Fig. 2. Aega leptonica, holotype: A, Pereopod 1; B, Pereopod 2; C, Pereopod 7; D, Left uropod, ventral view; E, Right uropod, endopod apex, dorsal view; F-I, Pleopods 1-3,.5 respectively. chium and merus indented, each indenta- Pleopods with only endopod of pleopod tion with spines as well as spines at distal 5 naked; endopods of pleopods 3 and 4 with angles; carpus with | spine on posterior setae only on distomedial margin; endopods margin; propodus with 3 spines on posterior 3-5 with small acute process on mediodistal margin. angle. Peduncle of pleopod 1 with 9 cou- VOLUME 101, NUMBER 1 99 Fig. 3. Rocinela oculata, holotype. A, Dorsal view of cephalon; B, Propodal palm, pereopod 1; C, Propodal palm of peropod 2; D, Lateral margins of right uropodal rami. pling hooks decreasing to 7 on peduncle 4. Uropods extending to apex of pleotelson, rami about equal in length. Exopod lateral margin convex, with 15—16 marginal spines and continuous marginal setae, medial mar- gin with 5 spines and continuous marginal setae. Endopod lateral margin sinuate, with continuous marginal setae and three spines; medial margin with continuous marginal setae and 8 spines; apices of both rami bifid. Male. —Not known. Color.—On collection label: “Pink on posterior three somites .. . eyes chestnut.” Remarks. —The shape of the pleotelson, medially united eyes, and slender anterior pereopods readily identify this species. Aega leptonica is most similar to those species which have a prominently excised pleotel- son apex: Aega beri Bruce, and Aega quad- ratisinus Richardson. Characters common to all three species include antennule, an- tennal, clypeal, pereopodal, pleopodal and uropod morphology. Aega leptonica differs from those two species in having medially united eyes, and more slender anterior pe- reopods. The similar species Aega excisa Schioedte & Meinert, 1879, differs in having smoothly convex pleotelson lateral margins, and the pletoelson excision is V-shaped, not rect- angular. Etymology.—The specific epithet is de- rived from combining the Greek words /ep- tos (slender, thin) and onychos (talon or claw). Rocinela oculata Harger Fig. 3 Rocinela oculata Harger, 1883:97, pl. II, figs. 2-2a, pl. IV, fig. 1.—Ruichardson, 1898:9; 1900:219; 1901:523; 1904:34, 35; 1905:191, fig. 195.—Schultz, 1969:199, fig. 314. Not Rocinela oculata.—Bruce, 1983:778, figs. 15, 16 (=Rocinela kapala, new species). Material. —Holotype, immature (13.3 mm), off Georgia, U.S.A. (MCZ 3910). Remarks.—The single specimen is too fragile to allow dissection, but additional figures of appendages in situ are given. 100 Material in the Smithsonian collections from the Caribbean and Florida identified as R. oculata (USNM 7513, USNM 90440) are not that species, the morphology and spination of pereopods | to 3 being incom- patible with Harger’s holotype. This mate- rial appears to be an undescribed species of Rocinela. Distribution. —Known only from the type locality, off Georgia, U.S.A. Rocinela kapala, new species Rocinela oculata.—Bruce, 1983:778, figs. 15, 16 (not R. oculata Harger, 1883). Material.—HOLOTYPE male (41.5 mm), east of Clarence River mouth, NSW, Aus- tralia, 29°25.2’S, 153°49.5’E, 12 Oct 1975, 450 m, coll. NSW State Fisheries on F.R.V. Kapala (AM P31740) (specimen figured by Bruce 1983). PARATYPES. 4 males, 2 fe- males, same data as holotype (AM P31705). Other paratypes detailed by Bruce (1983) AM P17950, P21014, P31704, QM W 10464. New records.—Male (36.0 mm), female (48.0 mm), off Sydney, NSW, 33°47’S, 151°58’E, 6 Dec 1972, 765 m (AM P37143); male (48.0 mm), off Sydney, NSW, 33°42’S, 151°57'E, 25 Sep 1984, 632 m (AM P37142); female (41.0 mm) off Clarence River, NSW, 29°46'S, 153°42’E, 26 Apr 1978, 405 m (AM P37141); all coll. NSW State Fisheries, FRV Kapala. Remarks.—This material was described in detail by Bruce (1983) and identified as R. oculata. Under remarks Bruce (1983) commented that specificity of Australian and western Atlantic material would only be confirmed by examination of adult speci- mens of R. oculata from off the U.S.A. Re-examination of the holotype of R. oculata showed that the two populations cannot be considered conspecific, and the Australian species is here named as new. Rocinela kapala can easily be distin- guished from R. oculata by: shorter, ante- riorly rounded rostrum; pereopods 1 to 3 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with palm less produced, posterior margin convex; pereopods 4-7 less spinose; per- eonites 4 to 7 become elongated in adults. Etymology. — Named after the New South Wales Fisheries vessel, FRV Kapala. Acknowledgments This initial part of this study was carried out while in receipt of a Smithsonian Post- doctoral Fellowship (1984) and completed while in receipt of a Queen’s Fellowship in Marine Science. The Australian Museum is gratefully acknowledged for provision of fa- cilities. Literature Cited Bowman, T. E. 1986. Tridentella recava, a new iso- pod from tilefish burrows in the New York Bight (Flabellifera: Tridentellidae).—Proceedings of the Biological Society of Washington 99:269- D3. Bruce, N.L. 1983. Aegidae (Isopoda: Crustacea) from Australia with descriptions of three new species.—Journal of Natural History 17:757- 788. Brusca, R.C. 1983. A monograph on the isopod fam- ily Aegidae in the tropical eastern Pacific. 1. The genus Aega.—Allan Hancock Monographs in Marine Biology, Allan Hancock Foundation 12: 1-39. — .,&E. W. Iverson. 1985. A guide to the marine isopod Crustacea of Pacific Costa Rica.—Re- vista de Biologia Tropical 33, supplemento 7: 1-77. Hansen, H. J. 1890. Cirolanidae et familiae nonnul- lae propinquae Musei Hauniensis.—Kongeligt Videnskabernes Selskab Skrifter, 6, Raekke, Naturvidenskabelig og Mathematisk Afdelung 5:237-426. Harger, O. 1883. Reports on the results of the dredg- ing, under the supervision of Alexander Agassiz, on the East Coast of the United States, during the summer of 1880, by the U.S. Coast Survey Steamer ‘Blake’, Commander J. P. Bartlett, U.S.N., commanding.—Bulletin of the Museum of Comparative Zoology, Harvard 11:91-104. Richardson, H. 1898. Description of four new species of Rocinela, with a synopsis of the genus. — Pro- ceedings of the American Philosophical Society 37:8-17. . 1900. Synopsis of North-American inverte- brates. VIII. The Isopoda.— American Natural- ist 34:207—230, 295-309. VOLUME 101, NUMBER 1 . 1901. Key to the isopods of the Atlantic coast of North America, with description of new and little known species. — Proceedings of the United States National Museum 23:493-597. 1904. Isopod collected in Japan in the year 1900 by the U.S. Fish Commission Steamer ‘Al- batross,’ and in the year 1881 by the U.S:S. Palos.— Proceedings of the United States Na- tional Museum 27:32-89. . 1905. A monograph on the isopods of North America.— Bulletin of the United States Mu- seum 54:1-727. Schioedte, J. C., & F. Meinert. 1879. Symbolae ad monographium cymothaorum crustaceorum isopodum familiae I. Aegidae.— Naturhistorisk Tidskrift, Series 3, 12:319-414, pls. 7-13. 101 Schultz, G. A. 1969. How to know the marine isopod crustaceans. Dubuque, Iowa, Wm. C. Brown Co., 359 pp. Treat, S.-A.F. 1980. New record of Aega monoph- thalma Johnston (Isopoda: Flabellifera: Aegi- dae) in the tropical western Atlantic. — Bulletin of Marine Science 30:912-914. Division of Invertebrate Zoology, Aus- tralian Museum. Present Address: Queens- land Museum, P.O. Box 300, South Bris- bane, QLD 4101, Australia. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 102-108 PANDEA CYBELES, A NEW MEDUSA FROM THE SARGASSO SEA (COELENTERATA: ANTHOMEDUSAE: PANDEIDAE) Angeles Alvarino Abstract.—A new medusa is described and illustrated. It is compared to related ‘species in the genus, Pandea conica (Quoy & Gaimard, 1827) and Pandea rubra Bigelow, 1913. It differs from those species in the proportions of the umbrella and its conical process, number of ribs on the umbrella, number of marginal tentacles, ocelli on the basal bulb of tentacles, and the large size of the stomach. The diagnostic characteristics of these species and Pandea cybeles are compiled in a table. Information is included on the distribution of the species throughout the world. The genus Pandea Lesson, 1843, includes two valid species, Pandea conica (Quoy & Gaimard, 1827) and Pandea rubra (Big- elow, 1913. Mayer (1910) recognized five species of Pandeidae: Pandea conica (Quoy & Gaimard, 1827), P. saltoria (Sars, 1835), P. minima von Lendenfeld, 1884, P. vio- lacea Agassiz & Mayer, 1899, and P. maasi Maas, 1904. According to Kramp (1965) P. maasi is a synonym of Euphysa flammea (Linko, 1905); P. minima and P. violacea are ju- venile stages of indeterminable Tiaridae, placing also P. violacea under Merga vio- lacea (Agassiz & Mayer, 1899); and P. sal- toria 1s a species of Aglantha. During the Sargasso Sea Biowat cruise on the R/V Knorr in Apr 1975, plankton col- lections were obtained, and some specimens of Medusae, Siphonophora, and Cteno- phora were kindly sent to me by Michael Latz (Department of Biological Sciences, University of California, Santa Barbara) for identification. The medusae included three specimens of a new species of Pandea, de- scribed below. Pandea cybeles, new species Figs. 1, 2 Material. —NE Sargasso Sea, Biowat Cruise of R/V Knorr, Apr 1975, from tows with Tucker '2 m net with 333 wm mesh: Sta 240, 34°53.22'N, 70°06.01'W, ca. 100 m depth, 20 Apr 1975, holotype, USNM 77473.—Sta 231, 33°55.74'N, 69°59.11'W, ca. | m depth, 19 Apr 1975, paratype, USNM 77474. A third specimen from Sta 230, 33°55.74'N, 69°59.11'W, ca. | m depth, 19 Apr 1975, was left at the University of California, Santa Barbara. Description. —Umbrella_ bell-shaped, slightly higher than wide (heights 18, 20 and 25 mm, widths 14.5, 16.3 and 20 mm re- spectively), with conical apical process about 5 mm long (measurements included in total height) in largest medusa (Fig. 1). Exum- brella with ridges reaching from tip of apical process to each marginal tentacle, alternat- ing with grooves running from tip of apical process to spaces between tentacles at edge of umbrella. Crest of each ridge with thin whitish band, probably formed by rows of nematocysts. The four wide radial canals with rough margins. Circular or ring canal simple, about half width of radial canals, with smooth margins (Fig. 1). Centripetal canals lacking. Velum narrow. Marginal tentacles about 40, 10 at each space between perradii, long, of same size, hollow. Base of tentacle with thick conical laterally compressed bulb or spur clasping margin of umbrella. Abaxial spur of each tentacle having | red ocellus. VOLUME 101, NUMBER 1 Fig. 1. Pandea cybeles, habitus. Stomach large, completely filling sub- umbrellar cavity, attached for about '4 of its length to subumbrella perradii. Mouth with 4 perradial lips deeply and complexly folded, with crenulated edges, reaching border of umbrella. Gonads extending over interradial and perradial zones, completely covering stom- ach, forming irregular network of sinuous ridges and pits, latter corresponding to in- ternal ovular formations (Fig. 2B). Color of stomach, mouth, gonads, and border of umbrella pinkish, with light pur- ple and violet tones. Thick jelly umbrella and conical apical process, crystal clear with violet tones, revealing darker tones of stom- ach and gonads within. Etymology.—Named after Cybele, god- dess of nature. Remarks. — Differential morphological characteristics of Pandea cybeles, P. conica, and P. rubra are given in Table 1. 103 Fig. 2. Pandea cybeles: A, Detail of bulbs in mar- ginal tentacles; B, Detail of internal ovular formations in gonads. Pandea cybeles differs from P. conica in the proportion of length to width of the um- brella, the number of ribs, the number of marginal tentacles, and the size of the stom- ach. Pandea cybeles differs from P. rubra in having a conical apical process of the um- brella, and ribs and nematocyst tracks on the umbrella, all of which structures are ab- sent in P. rubra. The circular canal is broad- er in P. rubra than in P. cybeles. The number of marginal tentacles is different in the two species, and P. rubra does not have ocelli on the basal bulb of the tentacles. The stomach of P. rubra reaches to half the length of the subumbrellar cavity; in P. conica it occupies about 4 or '2 of the length of the subumbrellar cavity, whereas in P. cybeles it fills completely the subumbrellar cavity, reaching the margins of the um- brella. Differences between Pandea rubra and Pandea cybeles are obvious and do not re- quire further discussion. However, Pandea 104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Principal differential characteristics of the species of Pandea and the new species. Characteristics Pandea conica (Quoy & Gaimard, 1827) Pandea rubra Bigelow, 1913 Pandea cybeles, n. sp. Umbrella Umbrella size Velum Apical conical process Ribs and ridges Radial canals Circular canal Marginal tentacles Stomach Mouth Gonads Bell barrel-shaped, about twice as high as wide, jelly fairly thick, with apical process 20-30 mm height, 10-15 mm wide Narrow Conical at summit, with apical ectodermal thickening 16 or in the 20’s. Exum- brellar nematocyst ribs 4 broad, smooth or jag- ged Narrower than radial ca- nals, smooth outlines 16 or in the 20’s, smooth, hollow, with conical laterally com- pressed basal bulb, without well developed abaxial spur, with one abaxial ocellus. No ru- dimentary marginal tentacles Large, pyramidal, almost filling upper '4 of sub- umbrellar cavity, at- tached about % of length to subumbrellar perradii 4 perradial lips with folded crenulate edges On entire interradial walls of stomach, forming coarse mesh- work of ridges and pits, surrounding the stomach Bell-shaped, as high or slightly higher than wide, with rounded summit, without apical process. Thin soft walls 30-40 mm height up to 75 mm Narrow Not present No ribs or exumbrellar nematocyst tracks 4 broad with wavy or jagged outlines Broad, with smooth out- lines 18—24 of various sizes, hollow, smooth, with large conical basal bulb, not laterally compressed, with dis- tinct abaxial spur clasping margin of um- brella. No ocelli. No rudimentary marginal tentacles Large, with broad base, about half-height of subumbrellar cavity, attached to subumbrel- lar perradii for about +, of its length 4 lips with folded crenu- late margins Very fine meshwork of pits, interradially on stomach, close-meshed irregular network of ridges with pits be- tween them Bell-shaped, slightly higher than wide, of thick jelly, with thick apical conical process 18-25 mm height and 14.5-20 mm wide Narrow Present, long conical ectodermal thickening at top of umbrella Present 40 exumbrellar ribs with nematocyst band at crest edge, extending from tip of conical process to edge of umbrella 4 broad with jagged outline Narrower than radial canals, about half width, smooth out- lines About 40, 10 from perradial to perradial, hollow, long, with conical laterally compressed basal bulb, with abaxial spur and ocellus. No rudimentary marginal tentacles Large, filling completely subum- brellar cavity, attached '4 of its length to subumbrellar perra- dii 4 perradial lips with complexly folded crenulate edges reaching border of umbrella Extended over interradial and perradial zones covering com- pletely stomach, forming net- work of ridges and oval pits VOLUME 101, NUMBER 1 Table 1.—Continued. Pandea conica (Quoy & Gaimard, 1827) Characteristics Stomach and gonads red- dish, brownish or yel- lowish. Mouth lips reddish or pink, tenta- cles milky yellow, ocel- li red or reddish brown. Subumbrella colorless Color Distribution Atlantic, Mediterranean Pandea rubra Bigelow, 1913 Subumbrella, stomach, mouth, gonads and marginal tentacles deep brownish red or chocolate Bermuda, NW Pacific, 105 Pandea cybeles, n. sp. Stomach, mouth, gonads and border of umbrella pink with light purple and violet tones Sargasso Sea Bering Sea, British Co- lumbia. Probably in- habiting deep waters conica and Pandea cybeles are more closely related, as both have a long conical process at the top of the umbrella. Therefore, a dis- cussion on the descriptions of Pandea co- nica published by various authors will en- lighten the separation of the species. There is some disagreement among the various authors on the number of marginal tentacles, as well as in the proportion of height and width of the umbrella. It appears in some instances that authors were includ- ing under Pandea conica specimens belong- ing to different species. However, all authors basically agree, when defining Pandea conica, that the height of the umbrella is about twice its width, the manubrium extends only along the upper '4 or half of the subumbrellar cavity, and the number of ribs on exumbrella and marginal tentacles is in the 20’s, about 24. A chronological review of descriptions by various authors follows. Quoy & Gaimard (1827) first described the medusa as Dianaea conica, collected near the Strait of Gibraltar, as body elongated conically pointed at the top, “tentacles small, filamentous, in the 20’s, with reddish spot at their base. The umbrella presented as many striae as tentacles. Manubrium with 4 small short arms, pink in color, the rest of the medusa transparent.” Their illustra- tion of the medusa shows size proportions of umbrella, stomach and tentacles. Maas (1904) found abundant specimens of Pandea conica at Monaco Bay, and those had 8, 16, or 20 tentacles. Mayer (1910) described Pandea conica as *‘barrel-shape sides bluntly pointed, 21 mm high and 10 mm wide, with 8 to 24 well developed longitudinal rib-like ridges along exumbrella and equal number of marginal tentacles with abaxial ectodermal ocelli. Stomach wide and short with 4 folded lips having sinuous margins.”’ The colors of go- nads, tentacles, and ocelli were brownish red, yellowish-milky and dark purple, re- spectively. Mayer’s illustration (1910:117) shows short additional ridges extending up to '4 from the border of the umbrella, and the conical top is missing. Either this is an illustration of a newly-born medusa or it is not P. conica. Vanhoffen (1911) indicated P. conica 12.5 mm high and 10 mm wide with conical pro- cess of 2 mm, 4 radial canals and ring canal. The largest specimens had 19 or 20 tenta- cles. Other characteristics agreed with de- scriptions by other authors. Hartlaub (1913) also indicated the um- brella extending into a conical process at the top, being 21 mm high and 10 mm wide, with 24 marginal tentacles, and the stomach 106 extending along the upper 3 of the sub- umbrellar cavity. Gonads and color agree with previous descriptions. Figure 286 on page 339 of Hartlaub 1913 illustrates his description. Browne (1916) referred to Pandea juve- nile specimens obtained at the Chagos Ar- chipelago, which could be related to Pandea conica. The smallest was about 2.5 mm in diameter. Uchida (1927) discussed the character- istics of Pandea conica and P. rubra. He identified and described specimens ob- tained at Misaki, Japan, as Pandea conica, with ‘“‘bell somewhat prismatic, with trun- cated top, 30 mm high and 17 mm wide; exumbrella with 44 longitudinal ridges. 4 Radial canals, wide, jagged, widest in the lower half. Ring canal narrower. Tentacles 44, all of similar length, hollow. Manubrium wide and short, nearly filling the upper half of bell cavity, fused with the upper part of subumbrella.”” The rest of the descriptive part agrees with other authors, in gonads, color, etc. He explained, ““Ocellus could not be found in preserved specimens.” The truncate top of the umbrella of Uchi- da’s specimens indicates that those speci- mens apparently do not belong to Pandea conica (see fig. 38, page 214 of Uchida 1927). Ranson (1936) discussed the character- istics of species of Pandea, mainly referring to P. conica and P. rubra. The collections Ranson analyzed included abundant spec- imens or various sizes, enabling him to de- termine series of development of the me- dusa. A table on page 83 compiles the characteristics of P. conica as described by various authors. Analysis of data in that table suggests that under P. conica were in- cluded some specimens belonging to other species. Russell (1953) described P. conica with higher than wide umbrella, conical summit, 16 to 24 longitudinal exumbrellar nema- tocyst ribs and corresponding number of marginal tentacles; jelly fairly thick. Velum PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON narrow. Stomach pyramidal filling upper half of subumbrellar cavity, etc. Kramp (1961) described P. conica “up to 21 mm high, 10 mm wide, with a conical apex terminating with a peculiar patch of thickened ectoderm; exumbrella with lon- gitudinal ribs and ridges. Manubrium about half as long as bell cavity, with short mouth tube and folded lips. Radial canals fairly narrow, smooth. 16-24 tentacles with lat- erally compressed basal bulbs, with abaxial ocellus.” Radial canals in P. conica as described by Kramp (1961) do not agree with those in P. cybeles, which are wide with jagged outlines. According to Kramp (1961) the Pandea conica of Bigelow (1918) found between Chesapeake Bay and Bermuda is possibly a new species. The main anatomical features used to dis- tinguish species of Pandea are: shape, di- mensions, and characteristics of umbrella; shape and dimensions of stomach; charac- teristics of canals; number and character- istics of tentacles. In Pandea conica according to most au- thors, the height of the umbrella is almost twice its width. The stomach occupies the upper '4 or probably half of the subum- brellar cavity. Ribs and tentacles are in the 20’s, usually up to 24. In Pandea cybeles, width and height of umbrella are rather similar, only slightly higher than wide. The stomach is large, fill- ing completely the subumbrellar cavity, ex- tending to the edge of the umbrella. Ribs and marginal tentacles number 40. Differences between P. conica and P. cy- beles are clear when comparing the illustra- tions by Quoy & Gaimard (1827), Hartlaub (1913), Russell (1953), and Kramp (1959, 1965), with those of P. cybeles in the present work. Distribution.—The medusa is a mero- planktonic stage. Specimens of Pandea, in- cluding Pandea cybeles have been mainly obtained far offshore at oceanic localities. VOLUME 101, NUMBER 1 Therefore, the medusae experience wide oceanic distribution, enjoying a long-lived pelagic medusoid stage. Kramp (1959) stat- ed that the large size of the medusae may indicate a long pelagic life, which is advan- tageous for increasing dispersion of the pop- ulation, with the opportunity to be trans- ported by currents to regions distant from the normal habitat of the species. Kramp (1959) also suggested that in Pandea conica, wide oceanic distribution is due to the fact that its hydroid is attached to the shell of the pteropod Cleodora cuspidata. Segura (1984) found specimens of Pandea sp. at several locations northwest of the Ga- lapagos Islands, off Ecuador and Peru. They were juvenile specimens of some species of Pandea, with umbrella 1.0 to 3.0 mm high and 1.0 to 2.0 mm wide, with no apical projection, umbrella of thick mesoglea, ex- umbrella with longitudinal rows of nema- tocyst tracks in same number as marginal tentacles. Velum narrow. Stomach large, filling almost completely the subumbrellar cavity. Radial canals wide, circular canal narrow. Marginal tentacles up to 24, ten- tacular basal bulbs conical, laterally com- pressed. No ocelli at basal bulb of tentacles. Pandea conica has been observed near Bermuda (Bigelow 1918, 1938), China (Chiu 1954), Gibraltar (Hartlaub 1913), Tristan de Cunha (Haeckel 1879), Alboran Sea, Tir- rhenian Sea, Sidra Sea, Messina, Aegean Sea, Marmora Sea, Strait of Gibraltar (Kramp 1924), Japan and Philippine Islands (Kramp 1928), Gulf of Guinea (Kramp 1955), West Africa and off Argentina (Kramp 1957), Sargasso Sea, west of Spain, east of Azores (Kramp 1959), East Africa, Ceylon, East Australia, New Zealand, Vietnam (Kramp 1968), Strait of Gibraltar (Quoy & Gaimard 1827), Mediterranean Sea (Ranson 1936), Japan (Uchida 1927), Agulhas Current (Vanhoffen 1911), China Seas (Zhang 1979). Pandea rubra has been recorded from British Columbia and Puget Sound (Arai & Brinckmann-Voss 1980), Gulf of Alaska, 107 Aleutians, Southeast Kamchatka, Sea of Okhotsk, San Francisco (Bigelow 1913), Bermuda (Bigelow 1938), West Ireland (Kramp 1920), west of British Isles (Kramp 1929, Russell 1953), south and southwest of Iceland (Kramp 1926), South Africa, Weddell Sea, Antarctic (Kramp 1957a, b), Northwest Pacific, Ceylon, Northeast Pa- cific (Kramp 1965), USSR Far East waters (Naumov 1956), Cochin, Malabar-Trivan- drum coastal waters (Vannucci et al.1970). Acknowledgments I would like to express my appreciation to Michael Latz from the University of Cal- ifornia Santa Barbara, for providing me with the plankton material from the Sargasso Sea, and to Frank D. Ferrari of the Smithsonian Oceanographic Sorting Center for the sorted Hydromedusae from the Sargasso Eel U. Maine collections sent to me for analysis, in order to obtain more specimens of Pan- dea. 1 am grateful to Thomas E. Bowman for his kind advice and careful editorial as- sistance. Thanks are also due to Debra Lo- sey (Librarian at Southwest Fisheries Cen- ter) for providing me with the literature I have needed for this work, and to John F. Carr and John R. Hunter of the SWFC for reading the manuscript. Literature Cited Arai, M. N., & A. Brinckmann-Voss. 1980. Hydro- medusae of the British Columbia and Puget Sound.—Canadian Bulletin of Fisheries and Aquatic Sciences (204):1—-192. Bigelow, H. B. 1913. Medusae and Siphonophorae collected by the U.S. Fisheries Steamer AL- BATROSS in the Northwestern Pacific 1906.— Proceedings of the U.S. National Museum 14: 1-119. 1918. Some Medusae and Siphonophorae from the Western Atlantic. — Bulletin of the Mu- seum of Comparative Zoology at Harvard Col- lege 63:365-442. 1938. Plankton of the Bermuda Oceano- graphic Expeditions. VIII Medusae taken during 108 the years 1929 and 1930.— Zoologica (New Y ork) 23(5):99-189. Browne, E.T. 1916. Medusae from the Indian Ocean. The Percy Sladen Trust Expedition. — Transac- tions of the Linnean Society of London (Zool- ogy), series 2, 17:169-210. Chiu, S. T. 1954. Studies on the Medusae fauna of the South-eastern China coast with notes on their geographical distribution. — Acta Zoologica Sin- ica 6(1):49-57. Haeckel, E. 1879. Das System der Medusen. Erster Theil einer Monographie der Medusen. Pp. 1- 360. Jena. Hartlaub, V. 1913. Craspedote Medusen. Teil I, Lief. 3, Family Tiaridae.—Nordisches Plankton (17)12:237-363. Kramp, P. L. 1920. Anthomedusae and Leptome- dusae.— Report of the Scientific Research of the MICHAEL SARS North Atlantic Deep Sea Ex- pedition 3(2):1-24. : 1924. Medusae.—Report on the Danish Oceanographic Expeditions 1908-1910 to the Mediterranean and Adjacent seas (8)2, Biol. K1: 1-40 1926. Medusae II. Anthomedusae.— Danish Ingolf Expedition 5(10):1-102. .1928. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-1916. XLIII Hydromedusae. I Anthomedusae. — Videnskabelige Meddelelser Dansk Naturhistorisk Forening i Kgbenhavn 85: 27-64. . 1955. The Medusae of the tropical west coast of Africa.—Atlantide Report 3:239-324. 1957a. Hydromedusae of the DISCOVERY Collections.— Discovery Reports 29:1-128. 1957b. Medusae.—British-Australian-New Zealand Antarctic Research Expedition 1929- 31, (B)6:151-164. 1959. The Hydromedusae of the Atlantic Ocean and adjacent waters. — Dana Reports (46): 1-283. 1961. Synopsis of the Medusae of the World.—Journal of the Marine Biological As- sociation of the United Kingdom 40:1-469. 1965. The Hydromedusae of the Pacific and Indian Oceans.— Dana Reports (63):1-162. 1968. The Hydromedusae of the Pacific and Indian Oceans.— Dana Reports (72):1—200. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Maas, O. 1904. Méduses provenant des Campagnes des yachts HIRONDELLE et PRINCESS ALI- CE.—Résultats des Campagnes Scientifiques, Monaco 28:1-71. Mayer, A. G. 1910. Medusae of the World.—Car- negie Institution of Washington Publication 109: 1-735. Naumov, D. V. 1956. Medusae in the Far-eastern waters of the Soviet Union.—Akademia Nauk SSSR. Zoologicheskii Institut. Trudy Problem- nykh i Tematisheskikh Soveshchsnii 6:34-41. [In Russian] Quoy, J.R.C., & J. P. Gaimard. 1827. Observations zoologiques faites 4 bord de 1ASTROLABE, en mai 1826, dans le détroit de Gibraltar. — Annales des Sciences Naturelles, Paris 10:1—21, 172-193. Ranson, G. 1936. Méduses provenant des Cam- pagnes du Prince Albert I de Monaco.—Résul- tats des Campagnes Scientifiques accomplies sur son yacht par Albert I, Prince Souverain de Monaco 92:1—245. Russell, F.S. 1953. The Medusae of the Brtitish Isles. Cambridge University Press, p. 1-530. Segura, L. 1984. Morfologia, sistematica y zoogeo- grafia de las medusas (Cnidaria, Hydrozoa, Scy- phozoa) del Pacifico Tropical Oriental.—Pu- blicacion Especial del Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Au- tonoma de México (UNAM) 8:1-320. Uchida, T. 1927. Studies on Japanese Hydromedu- sae. I. Anthomedusae.—Journal of the Faculty of Sciences Imperial University of Tokyo, Sec- tion 4, Zoology 1(3):145-241. Vanhoffen, E. 1911. Die Anthomedusen und Lep- tomedusen des Deutschen Tiefsee Expedition 1898-1899.— Wissenschaften Ergebnisse VAL- DIVIA 19(5):191-233. Vannucci M., V. Santhakumari, & E. P. dos Santos. 1970. The ecology of Hydromedusae from Cochin area.— Marine Biology 7:49-58. Zhang, J. 1979. A preliminary analysis of the hydro- medusae fauna of the China Sea areas.—Acta Oceanologica Sinica 1(1):127—237. NOAA, NMEFS, Southwest Fisheries Center, P.O. Box 271, La Jolla, California 92038. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 109-145 PALEOCENE TURTLES FROM THE AQUIA AND BRIGHTSEAT FORMATIONS, WITH A DISCUSSION OF THEIR BEARING ON SEA TURTLE EVOLUTION AND PHYLOGENY Robert E. Weems Abstract.—The Piscataway Member of the Aquia Formation (upper Paleo- cene: Thanetian) has yielded remains of six species of turtles: Aspideretes vir- ginianus (Clark), Planetochelys savoiei (n. gen., n. sp.), Osteopygis roundsi (n. sp.), Dollochelys coatesi (n. sp.), Catapleura ruhoffi (n. sp.), and Allopleuron insularis (Cope). The Brightseat Formation (lower Paleocene: Danian) has yield- ed three taxa: Taphrosphys sulcatus (Leidy), Agomphus sp., and Osteopygis emarginatus Cope. These faunas, when compared with Late Cretaceous and early Eocene marine turtle faunas, provide valuable insights into the evolu- tionary history of sea turtles from Late Cretaceous through Early Tertiary time. No catastrophic terminal Cretaceous extinction event among sea turtles is indicated by this succession. Rather, strong decline in the late Campanian is followed by modest recovery in the Thanetian and Ypresian. This decline and renaissance closely matches the global pattern of oceanic cooling and warming in Late Cretaceous—early Tertiary time. Turtle remains have been reported from the Aquia Formation of the Pamunkey Group since 1895, but mostly from isolated fragments (for example, see Clark & Martin 1901). Not until Lynn (1929) described a fairly complete carapace of ““Amyda’’ vir- giniana was there sufficient material ade- quate to characterize even one species. Even so, Lynn did not demarcate the suture boundaries on this specimen and the pub- lished illustration does not show them clear- ly. Since the publication of Lynn’s paper, more chelonian material has turned up spo- radically both in the Aquia Formation and in the underlying Brightseat Formation, but it has not been described. This new mate- rial, although far from ideal, is still sufficient to expand greatly our knowledge of the di- versity and taxonomy of the turtles of Pa- leocene age and sheds new light on their anatomy. Age of the Aquia and Brightseat Forma- tions. —The Aquia formerly was considered to be a lower Eocene unit (Clark & Martin 1901), but Loeblich & Tappan (1957) dem- onstrated that it should properly be consid- ered part of the upper Paleocene column on the basis of its contained planktonic Fora- minifera. Greater refinement in the strati- graphic position of this unit has been achieved by Gibson and others (1980), whose work indicated that the entire Aquia lies within calcareous nannoplankton zones NP5 through NP9. The Aquia is divided into two members (Clark & Martin 1901), a lower member named the Piscataway which lies within NP5 through NP8, and an upper member named the Paspotansa which falls entirely within NP9. This division im- plies that the Piscataway accumulated 60 to 57 Ma and the Paspotansa accumulated 57 to 55 Ma (Fig. 1). Both members of the Aquia belong within the Thanetian Stage of the Paleocene (Hardenbol & Berggren 1978). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 110 5 51 ‘(0861 ‘Te 19 WOSqId “[O6] UIE 7 YreIO Joye) errs, pue purlAreypy ur dnoip Aoyunwed 94} SuIsLIdu0d suONeULIO} pue siaquidu ay) jo sosuevlosy "| “314 N ise) + i i <— — |< | | | Ay | \ | ' | | ! | | I I | | —=) {| Ss — |< | | Fig. 25. Diagram showing the two major evolutionary trends in the plastrons of advanced sea turtles. Primitively the plastral elements are squarish and only slightly reduced, as illustrated by the Chelospharginae (bottom). In more advanced dermochelyoid forms, there is a tendency for the plastral elements to become more reduced by compression (shown by arrows) toward two axes (dashed lines) oriented in an anteroposterior direction, leaving a very broad medial plastral fontanelle (upper left). In cheloniid and toxochelyid turtles, the plastral elements tend to become compressed toward a single axis oriented transversely across the midline of the shell (upper right), leaving broad anterolateral and posterolateral plastral fontanelles. In the living Chelonia mydas, both types of compression seem to occur, but this is a highly derived condition, established in the Late Tertiary, atypical of other cheloniids. Because the entire superfamily to which many more species of Cretaceous and early the Dermochelyidae belong is so thin-shelled Tertiary dermochelyids remain to be de- and fontanellized, well-preserved speci- scribed and that any phylogeny of this fam- mens are rare. It therefore seems likely that ily made at the present time will be neces- VOLUME 101, NUMBER 1 sarily sketchy and very incomplete. Similarly, the details of the phylogeny of the family Cheloniidae are still sketchy and in- complete due to lack of described material at many horizons. Nevertheless, at the subfamily and higher taxonomic levels, a cladistic phylogeny can be drawn relating all of the various known families and subfamilies of sea turtles on the basis of shared derived characteristics (see Fig. 26). This proposed phylogeny 1) entirely re- moves occurrences of apparently advanced cheloniids from the Cretaceous and 2) im- plies that the Dermochelyidae occur as far back in time as unequivocal members of the Toxochelyidae. Kirgizemys, which was de- scribed by Nessov & Khozatskiy (1978) as a Lower Cretaceous toxochelyid, is ex- tremely primitive and shows only a few de- rived toxochelyid traits in its shell, such as developing lateral plastral fontanelles and a tendency toward anteroposterior constric- tion of the plastron toward the midline. The familial assignment is quite possibly cor- rect, but without cranial material it is im- possible to be certain based on so few ob- servable derived toxochelyid character states. Although the Dermochelyidae, as here defined, seem to stand in contrast to the Toxochelyidae and Cheloniidae, they do show some affinities with the Protostegidae, principally in the strongly reduced carapace, the proportions of elements in the shoulder girdle, and the morphology and histology of the humerus. Therefore, it is reasonable to associate these two families within the su- perfamily Dermochelioidea as an entirely separate line of sea turtle evolution from the Toxochelyidae and Cheloniidae, which can be associated together as the superfam- ily Chelonioidea. The four constituent fam- ilies of these two superfamilies are the same four recognized by Gaffney (1975b) as con- taining all Cretaceous and Cenozoic sea tur- tles. The taxonomic organization of these various families and subfamilies, and the described genera which can be assigned con- 141 Table 2.—Taxonomy of the sea turtles, showing su- perfamily, family, and subfamily levels of classification and their organization. Below the subfamily level, phy- logenetic pathways are not indicated and genera are listed seriatim. Superfamily Chelonioidea Family Plesiochelyidae Plesiochelys, Portlandemys Family Toxochelyidae Subfamily Toxochelyinae Dollochelys, ?Kirgizemys, Portochelys, Thino- chelys, Toxochelys Subfamily Lophocheliinae Ctenochelys, Lophochelys, Neurochelys, Peritre- sius, Prionochelys Family Cheloniidae Subfamily Osteopyginae Erquelinnesia, Osteopygis, Rhetechelys Subfamily Eocheloniinae Argillochelys, Catapleura, Eochelone, Glariche- lys, Puppigerus, Tasbacka Subfamily Syllominae ?Bryochelys, Kurobechelys, Syllomus Subfamily Cheloniinae Caretta, Carolinochelys, Chelonia, Eretmoche- lys, Lepidochelys, Procolpochelys Superfamily Dermochelyoidea Family Protostegidae Subfamily Chelospharginae Calcarichelys, Chelosphargis, Rhinochelys Subfamily Protostegidae Archelon, Protostega Family Dermochelyidae Subfamily Desmatochelyinae Corsochelys, Desmatochelys Subfamily Allopleuroninae Allopleuron, Eosphargis, Glyptochelone, Proto- sphargis Subfamily Dermochelyinae Cosmochelys, Dermochelys, Psephophorus fidently to them, are summarized in Ta- ble 2. Once Desmotochelys and Corsochelys have been removed from the Chelonioidea, the major evolutionary radiation of the Cheloniidae cannot be found before the be- ginning of the Tertiary. Persistently gener- alized sea turtles, possibly similar to Thi- nochelys and Portochelys, appear to have given rise to a primitive cheloniid, Cata- pleura arkansaw, by the Late Cretaceous 142 (0) © ac} > o i (3) 49) on ® a Toxochelyinae Lophocheliinae Osteopyginae Eocheloniinae Syllominae Cheloniinae Chelospharginae Protosteginae Desmatochelyinae Allopleuroninae Dermochelyinae Fig. 26. Cladogram showing the relationships and key shared derived characters of the known families and subfamilies of sea turtles. Numbers refer to the following character states or character complexes: 1) Palatine and internal carotid arteries equal in diameter, basisphenoidal trabeculae lie close together; foramina anterius canalis carotici interni lie close together and are separated by relatively thin bar of bone; dorsum sellae high and separated from sella turcica and carotid foramina by prominent bone surface; posterior part of sella turcica not concealed by overhanging dorsum sel- lae (Gaffney 1975b, 1984). 2) Palatine artery and can- alis caroticus lateralis larger than internal carotid artery and canalis caroticus internus; front limb with elongate digits and metacarpals forming a flipper with elements oriented in one plane (Gaffney 1975b, 1984). 3) Hy- oplastra and hypoplastra reduced by constriction anteroposteriorly toward a transverse midline axis (see Fig. 25). 4) Neurals serrated, forming mid-dorsal keel, peripherals moderately to strongly serrated, connection between pygal and suprapygal very narrow or broken, hyopiastron, hypoplastron, and xiphiplastron with conspicuous ventral keel knobs, epineural elements present in advanced genera (Zangerl 1953). 5) Second- ary palate present, postnuchal fontanelles lost at least in adult stages. 6) Prefrontal has an extended posterior process, nearly or entirely excluding frontal from orbit, fissura ethmoidalis deep, palate flat with low tomial ridge, vomer has straight contact with premaxillaries, enlarged complete secondary palate organized differ- ently from that in the Cheloniinae (Fastovsky 1985). 7) Secondary palate completely developed, rear limbs markedly modified for swimming rather than walking. 8) Secondary palate lost, jaw margins develop pseu- dodont crenulations, deltopectoral crest of humerus moved far distally toward the middle of the humerus. 9) Much of dermal bone layer absent in carapace, re- sulting in costal ribs that are free of overlying bone; front flippers very long in relation to body; large radial crest of humerus displaced distally, humerus tends to develop vascularized chondroepiphyses; braincase partially cartilaginous in adult (Wieland 1906, Zangerl 1953, Gaffney 1984, Rhodin 1985). 10) Pointed and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Schmidt 1944). But only later, in the late Paleocene, did the major radiation of che- loniids begin. Late Cretaceous (Santonian through Campanian) radiations of sea tur- tles did occur, but these only involved the families Toxochelyidae, Protostegidae, and Dermochelyidae. This major radiation end- ed with the Campanian, however, and no protostegids of any kind are known to have survived into the Maastrichtian. The toxo- chelyines are known only from a single form in the Maastrichtian (Toxochelys weeksi), and Dollochelys is the only known Paleo- cene form. Similarly, the lophochelyines are represented only by Peritresius in the Maas- trichtian and a single fragmentary form from the Danian (Wood 1973). In contrast, a modest radiation of the Allopleurines oc- curred in the Maastrichtian and the osteo- pygines make their first appearance. In gen- eral, though, both the Maastrichtian and Danian appear to have been times of rela- tively low sea turtle diversity. This general pattern of decline reverses in the Thanetian with the beginning of the cheloniid radia- tion. In the Ypresian (early Eocene), sea tur- tles reached their peak of Cenozoic diver- sity. Although lophochelyines make their last appearance in the form of Neurochelys (Moody 1980a), and osteopygines in the form of Erquelinnesia (Zangerl 1971), most of this assemblage consists of eochelyines and allopleurines. The later Tertiary history of the sea turtles so far is too poorly known to indicate exact relationships or to get a feeling for detailed changes in diversity. — decurved beak; nasal elements lost; costals very greatly reduced, extending no farther down the axis of the rib than a distance equal to their width (Zangerl 1953). 11) Hyoplastra and hypoplastra reduced by constric- tion laterally toward two parallel anteroposterior axes (see Fig. 25). 12) Nuchal elongated to project forward over neck region, neurals longer than wide. 13) Nuchal element deeply emarginated, neurals as wide or wider than long. 14) Normal shell elements (reduced) over- lain by mosaic of suturally interconnected dermal bones. VOLUME 101, NUMBER 1 # of species 100 90 70 60 50 40 143 Deep Ocean Paleotemperature O16) 20 (0) Megayears Fig. 27. Diversity in sea turtle families in comparison with the oxygen isotope derived oceanic paleotem- perature curve for the northern mid-latitudes (temperature curve adapted from Douglas and Woodruff 1981). Many factors could explain the strong correlation between ocean temperature and sea turtle diversity, including (1) increased areal extent of shallow sea habitat during warm spells, (2) more efficient physiology during warm spells, or (3) better egg viability during warm spells. Cen = Cenomanian, Tur = Turonian, Con = Coniacian, San = Santonian, Cam = Campanian, Maa = Maastrichtian, Dan = Danian, Tha = Thanetian, Ypr = Ypresian, Lut = Lutetian, Bar = Bartonian, Pri = Priabonian, Rup = Rupelian, Cha = Chatian, Aqu = Aquitanian, Bur = Burdigalian, L = Langhian, S = Serravalian, Tor = Tortonian, M = Messinian, Z = Zanclian, P = Piacenzian, Q = Quaternary undifferentiated. Generally, however, there has been decline since the early Eocene to the present diver- sity of five species worldwide. Sea Turtles and the Cretaceous—Tertiary Transition As indicated by the previous discussion, sea turtles seem to have had two periods of exceptional radiation and diversity, once in the Campanian and once in the Ypresian. In between those radiations, both the Maas- trichtian and the Danian represent a time of exceptionally low diversity. No major ex- tinction among sea turtles can be localized at the Maastrichtian—Danian boundary. Rather, the most dramatic time of extinc- tion occurs near the end of the Campanian. This pattern of evolutionary expansion and contraction does not seem to be strongly affected either by the recently suggested me- teoric impact event at the Maastrichtian— Danian boundary (Alvarez et al. 1980) or by the more gradual series of climatic and geologic changes suggested by Officer & Drake (1983), but does match nicely the changes in the estimated average oceanic temperatures for the mid-northern latitudes through the Late Cretaceous and early Ter- tiary (Fig. 27). The correlation is so close that the diversity of sea turtles seems to have been controlled either directly by av- erage ocean temperatures around the world or by the same factors that controlled such temperatures, rather than by short-term dramatic events that may have occurred sporadically throughout geologic time. If 144 extraterrestrial causes for this extinction are to be invoked, only a large interstellar dust and debris cloud which reduced the inten- sity of sunlight over an interval of millions of years would fit the pattern observed. As a generally quiet period in sea turtle history, the Maastrichtian—Danian interval was a time of continuity and stability, marked by neither the radical changes nor great innovations that would occur later in Tertiary. Thus, at a time when many other animal and plant groups have been de- scribed as being in a period of massive change or decline, the sea turtles appear to have been essentially unaffected by any great changes taking place around them. As the early Tertiary global warming trend unfold- ed, the sea turtles concurrently underwent a new evolutionary radiation, undimin- ished in their evolutionary potential by the Cretaceous—Tertiary transition. Acknowledgments I would like to thank Nicholas Hotton III and Donald Baird for their very helpful comments and criticisms on this work. Opinions expressed are those of the author, and do not necessarily reflect those of the reviewers. Literature Cited Alvarez, L. W., W. Alvarez, F. Asaro, & H. V. Michel. 1980. Extraterrestrial cause for the Cretaceous- Tetiary extinction.—Science 208(4448):1095- 1108. Baird, D. 1964. A fossil sea-turtle from New Jersey. — New Jersey State Museum Investigations 1:1- 26. Bennet, R. R., & G. G. Collins. 1952. Brightseat For- mation, a new name for sediments of Paleocene age in Maryland. — Washington Academy of Sci- ence Journal 42(4):114—-116. Carpenter, K. 1981. Preneural in the evolution of Trionyx.—Copeia 198 1(2):456-457. Clark, W. B. 1895. Contributions to the Eocene fauna of the Middle Atlantic Slope.—Johns Hopkins University Circular 15(4):3-6. 1896. 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Comparative chondro-os- seous development and growth of marine tur- tles.—Copeia 1985(3):752-771. Schmidt, K. P. 1944. Two new thalassemyd turtles from the Cretaceous of Arkansas.— Field Mu- seum of Natural History, Geology Series 8(1 1): 63-74. Shuvalov, V. F., & V. M. Chkhikvadze. 1979. O stratigraficheskom 1 sistematicheskom poloz- henii nekotorykh presnovodnykh cherepalh iz novykh melovykh mestonakhozhdeniy Mon- goli in R. Barsbold, ed., Fauna mezozoya i kay- nozoya Mongolii, sovmestnaya Sovetsko-Mon- gol’ skaya Palenotologicheskaya Ekspeditisiya. — Trudy 8:58-76, 123, 125-126, 149. Wieland, G. R. 1905. Structure of the Upper Cre- taceous turtles of New Jersey.— American Jour- nal of Science 27:112-132. 1906. The osteology of Protostega.—Mem- oires of the Carnegie Museum 2(7):279-298. Wiman, C. 1930. Fossile Schildkréten aus China.— Paleontologica Sinica, series C 6(3):1-56. Wood, R.C. 1973. Fossil marine turtle remains from the Paleogene of the Congo.— Musée Royal de VAfrique Centrale, Annales, Serie in-8° (Sci- ences Geologiques) 75:1—28. Yeh Hsiang-Kuei [also spelled Ye Xiang-kui]. 1963. Fossil turtles of China.—Paleontologica Sinica (150), new series C, 18:1-111. . 1973. Fossil chelonians from Wuérho (Urho). In Reports of the Paleontological Expedition to the Xinjiang; (2) the pterosaurian fauna from Wueérho (Urho).— Academia Sinica, Institute of Vertebrate Paleontology and Paleoanthropology Memoir 11:8-11. Zangerl, R. 1953. The vertebrate fauna of the Selma Formation of Alabama. Part 4, The turtles of the family Toxochelyidae.— Fieldiana, Geolog- ical Memoirs 3(4):137-—277. 1971. Two toxochelyid sea turtles from the Landenian sands of Erquelinnes (Hainaut), of Belgium.—Institut Royal des Sciences Natu- relles de Belgique, Memoire 169:1-32. Mail Stop 928, U.S. Geological Survey, Reston, Virginia 22092. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 146-150 A DESCRIPTION OF THE PIGMENTED AND NON-STYGOBIONTIC FEMALES OF PODOBOTHRUS BERMUDENSIS BARNARD & CLARK, 1985 (CRUSTACEA: AMPHIPODA: DULICHIIDAE) Michael F. Gable, Eric A. Lazo-Wasem, and Adam J. Baldinger Abstract. —The females of Podobothrus bermudensis (Amphipoda: Dulichi- idae) are described. Three females and two males, all from the shallow subtidal zone, are compared with each other and with the single cave-dwelling male used by Barnard & Clark (1985) to establish the monotypic genus. Most sig- nificantly, all females are pigmented and all males are non-pigmented. The mandibular palp is judged to be the most useful character for distinguishing Podobothrus from the closely related Podocerus. Podobothrus bermudensis Barnard & Clark, 1985, was described as a sea-cave amphipod species (and genus) from a male specimen collected from Green Bay Cave, Bermuda. Examinations of collections at the Yale Peabody Museum (YPM), made in Bermuda in 1985, and of United States Na- tional Museum (USNM) collections, made by M. L. Jones in the early 1980’s, have uncovered five additional specimens of this new species including three females. Be- cause these animals exhibit important mor- phological and habitat differences from the type specimen, we believe that descriptions of the specimens and remarks on their dis- tribution will provide valuable information for amphipod taxonomists and stygobiol- ogists. Description. — Female: Body pigmented, most notably on abdomen and thoracic areas (Fig. 1). Pereonites dorsally smooth except for a slight mucronation on pereonite 2, spine groups on distoposterior corners of pereonites 2—4, 6 and pleonite 1. Pleonites 1-3 with slight dorsal carinations. Uroso- mites as in males. Coxae reduced, subrectangular, distinctly larger than those of males, coxae 1—4 touch- ing, coxae 5—7 overlapping. Head and eyes as in males, except eyes pigmented, ap- pearing red in alcohol (Fig. 1). Antenna 1, 118% body length, ratio to peduncle of antenna 2, 1:1; flagellum 50% of peduncular length, 5-articulate; accessory flagellum, 1-articulate, as in male. Antenna 2, proportion of peduncular article 4 to 5, 1:1.1, flagellum missing. Mouthparts as in male except: lower lip with facial tubercles extending from internal margins of anterior lobes; medial margin of inner plate of maxilla 2 less setose, apical setae, plumose; setae of mandibular palp, distinctly pinnate (Fig. 2). Gnathopod | similar to that of male with exception of dactyl, which in female lacks inner marginal spination, but possesses se- tae and a distal annulation (Fig. 2). Gnatho- pod 2 similar to that of male, but more weakly subchelate, palmar margin of article 6 finely serrated, dactyl distally annulated; pigment spots along the length of the ap- pendage (Fig. 2). Pereopod 7 (others miss- ing) with pigment spots along its length. Pleopods as in male, except peduncular segments pigmented (Fig. 1). Uropods as in male, but uropod 3 lacking inner setule. Tel- son as in male. Material examined.—YPM 8269: Ferry VOLUME 101, NUMBER 1 147 Z Fig. 1. Podobothrus bermudensis, ovigerous female, 2.2 mm. YPM No. 8269: Top, Lateral view of body pigmentation (4x); Middle, Close-up of pigmented eye and adjacent pigment granules (40 x); Bottom, Lateral view of pigmentation on pleon and peduncular segments of pleopods (10 x). 148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ere \ Fig. 2. Podobothrus bermudensis, female, 1.9 mm. USNM Acc. No. 346847. Legend: major body parts marked by abbreviations beginning with uppercase letters; enlargements of gnathopod articles marked by a lowercase “‘d.”” Abbreviations: Gn, gnathopod; Md, palp of mandible; d, dactyl. Reach, St. George’s, Bermuda, Cove W of cent to west side of dock at BBS, M. L. BBS, M. F. Gable, 28 May 1985, 1 oviger- ous female, 2.2 mm; Ferry Reach, St. George’s, Bermuda, Cove W of BBS, M. F. Gable, 28 May 1985, 1 male, 1.7 mm. USNM Acc. No. 346847: Ferry Reach, Ber- muda, adjacent to west side of dock at BBS, M. L. Jones, 2 Sep 1981, Cohen net through Thalassia, 2 females, 1.8 mm and 1.9 mm; Ferry Reach, St. George’s, Bermuda, adja- Jones, 2 Sep 1983, 1 male, 1.8 mm. Remarks. — Barnard & Clark (1985) men- tioned a close resemblance between Podo- bothrus and the genus Podocerus. An un- determined species of Podocerus does occur in Bermuda. Because antennae and limbs of the two genera are often broken or miss- ing in preserved specimens, we indeed found difficulty in distinguishing the two. Many VOLUME 101, NUMBER 1 characteristics of damaged specimens (e.g., spine lengths of the uropods) are too sub- jective for one not already versed in telling the two genera apart. We found that the best feature for quickly and decisively distin- guishing Podocerus sp. from Podobothrus bermudensis 1s the third article of the man- dibular palp, clearly visible without dissec- tion, long, slender, and highly setose in the latter species, but somewhat short, stout, and with only a few short apical setae in the former species. Differences between male and female P. bermudensis were mentioned in our de- scription, but emphasis should be given to several notable differences. The last two ar- ticles of female gnathopod 2 differ signifi- cantly from those of the males, not unex- pectedly. (An interesting dimorphism between right and left second gnathopods for the only male specimen with both gnathopods intact involves distinct differ- ences in size and robustness of the append- ages.) The pinnate setation on the mandib- ular palp is not shown by Barnard & Clark (1985), but we found this type of setation on both female and male specimens. The two most noteworthy differences, however, are the coxal plates and pigmentation. The coxal plates of all females are larger than those of our male specimens and of the spec- imen described by Barnard & Clark. Their male holotype, from a cave, is without pig- mentation; our three females all have var- ious degrees of pigmentation, including pig- mented eyes, and our two males have none. All five of our specimens are from the shal- low subtidal habitat, and pigmentation in this species, from the few specimens at hand, therefore appears to be sex-linked and to have nothing to do with habitat. Such knowledge might be useful for stygobiolo- gists examining cave-related anatomical and morphological losses (e.g., pigmentation) to trace the evolutionary route of freshwater cavernicoles from marine ancestors (e.g., Stock 1986). The habitats occupied by P. bermudensis 149 support the statements made by Maddocks & Iliffe (1986) and Stock (1986) that certain island marine-cave and open-water taxa are often identical. Because of the great em- phasis currently being placed on anchiha- line and marine-cave fauna, care must be exercised in ascribing cavernicolous attri- butes to new species discovered in caves for those geographic areas and taxa, such as Bermuda and its amphipods, where shal- low-water surveys are virtually non-exis- tent. We have evidence to indicate that oth- er cave amphipods described this decade from Bermuda may also be found with reg- ularity in shallow-water marine habitats. Gable & Lazo-Wasem (1987) address this situation briefly in their review of the Ber- muda caprellid amphipods. Further shal- low-water surveys may indeed change the views on endemism (considerably higher in cave fauna than among littoral species) ex- pressed by some for Bermuda (e.g., Iliffe, Hart & Manning 1983). Acknowledgments The authors wish to thank Dr. J. L. Bar- nard (USNM) for the loan of material and Dr. P. Willenz (YPM) for photographic as- sistance. This study has been supported in part by a Connecticut State University Re- search Grant to the first author. This paper is contribution No. 1165 from the Bermuda Biological Station for Research. Literature Cited Barnard, J. L., & J. Clark. 1985. A new sea-cave amphipod from Bermuda (Dulichiidae).—Pro- ceedings of the Biological Society of Washington 98:1048-1053. Gable, M. F., & E. A. Lazo-Wasem. 1987. The ca- prellids (Amphipoda: Caprellida) of Bermuda, a survey of specimens collected from 1876-1987: including cave inhabitants and the description of Deutella aspiducha, new species.— Proceed- ings of the Biological of Washington 100(3):629- 639. lliffe, T. M., C. W. Hart, Jr., & R. B. Manning. 1983. 150 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Biogeography and the caves of Bermuda.—Na- ture 302:141-142. Maddocks, R. F., & T. M. Iliffe. 1986. Podocopid Ostracoda of Bermudian caves.—Stygologia 2: 26-75. Stock, J. H. 1986. Two new amphipod crustaceans of the genus Bahadzia from ‘blue holes’ in the Bahamas and some remarks on the origin of the insular stygofaunas of the Atlantic.—Journal of Natural History 20:921-933. (MFG & AJB) Department of Biology, Eastern Connecticut State University, Willimantic, Connecticut 06226-2295: (EAL-W) Division of Invertebrate Zoology, Peabody Museum of Natural History, Yale University, 170 Whitney Avenue, P.O. Box 6666, New Haven, Connecticut 06511- 8161. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 151-154 A NOTABLE COLLECTION OF CYCLORAMPHUS (AMPHIBIA: LEPTODACTYLIDAE) FROM BAHIA, BRAZIL, WITH A DESCRIPTION OF A NEW SPECIES (CYCLORAMPHUS MIGUELT) W. Ronald Heyer Abstract.— Members of the frog genus Cycloramphus were unknown north of the Rio Doce, Espirito Santo, Brazil until 1986, when two species of the genus were collected in southern Bahia, Brazil. The first, Cycloramphus fuli- ginosus, was previously known from the states of Espirito Santo and Rio de Janeiro. The second species is new and described as Cycloramphus migueli. Hypotheses are made to be tested by new data. Heyer & Maxson (1983:356) predicted that members of the frog genus Cycloram- phus did not occur north of the Rio Doce within the Atlantic Forest Morphoclimatic Domain. The ecology of most Cycloram- phus species is closely associated with small, fast-flowing mountain streams; topographic maps indicated that there were no areas north of the Rio Doce with enough topo- graphic relief to provide suitable habitat. Prior to 1986, no Cycloramphus had been collected north of the Rio Doce. However, in October of 1986, Miguel T. Rodrigues collected two species of Cycloramphus from Fazenda Unacau and Rio de Una, Bahia, and saw one individual of a third (distinct, possibly new) species. Twelve specimens of Cycloramphus fuliginosus were collected along small ('2-1'2 m wide), fast flowing, clear streams with rocky bottoms. Most specimens were encountered in the forest, and a few where the forest was replaced by cacao plantations. Dr. Rodrigues also col- lected, among rocks about 100 m from a stream, a single specimen of Cycloramphus representing a new species, proposed herein as: Cycloramphus migueli, new species Fig. | Holotype.—MZUSP 63450, adult male, from Brazil: Bahia; Sao José (do Macuco), Fazenda Unacau, 15°09’S, 39°18'W; Miguel T. Rodrigues, 13 Oct 1986. Diagnosis.—The webless species of Cy- cloramphus are bolitoglossus, carvalhoi, ca- tarinensis, diringshofeni, eleutherodactylus, granulosus, migueli, stejnegeri, and valae. The toes are free of web and fringe and the dorsum is smooth in C. migueli; the toes are ridged or fringed and the dorsum warty- granular in catarinensis, granulosus, and valae. The leg is shorter in C. migueli (e.g., foot length 36% SVL) than in diringshofeni (average foot length 50% SVL in males) or eleutherodactylus (average foot length 48% in males). Cycloramphus migueli is most similar in appearance to bolitoglossus, car- valhoi, and stejnegeri (this grouping was pre- viously recognized as the genus Craspedog- lossa [see Heyer 1983, for discussion]). The venter and posterior thigh surfaces of C. migueli are dark with small, distinct, irreg- ular light spots; the venter and posterior thigh surfaces of C. carvalhoi are uniform 152 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. brown; and the throat, chin, and posterior thigh surfaces of C. stejnegeri are dark brown with distinct light spots, while the belly is mostly unpigmented. In addition, C. mig- ueli is smaller (male SVL 42 mm) than either carvalhoi (male SVL 59-62 mm) or stejne- geri (male SVL 45-47 mm). The ventral and posterior thigh patterns of C. migueli and bolitoglossus are similar. The two species differ in size (C. migueli male 42 mm SVL, bolitoglossus males 29-34 mm SVL) and C. migueli has a distinct fold, from the pos- terior corner of the eye to the shoulder, which is lacking in C. bolitoglossus. Description of holotype.—Snout nearly round from above, slightly obtuse in profile; canthus rostralis indistinct; loreal convexly obtuse in cross section; tympanum hidden; vomerine teeth in obtuse series posterior to and between choanae; no vocal slits or vocal sac; first finger just longer than second; fin- gers lacking fringe or web; outer metacarpal tubercle large, square with rounded corners; inner metacarpal tubercie large, oblong, separated from subarticular tubercle of thumb by less than diameter of subarticular tubercle; thumbs lacking asperities; dorsal and ventral textures smooth; fold from pos- terior corner of eye to shoulder; inguinal Dorsal and ventral views of holotype of Cycloramphus migueli. gland large, circular, diameter greater than ¥3 length of thigh; toe tips rounded, not ex- panded; toes without fringe or web; subar- ticular tubercles under toes weakly devel- oped; outer metatarsal tubercle large, rounded, elongate, about '2 size of oblong inner metatarsal tubercle; no tarsal or meta- tarsal folds; posterior surface of tarsus and sole of foot smooth. SVL 42.1 mm, head length 16.1 mm, head width 19.2 mm, eye-nostril (middle of opening) distance 3.0 mm, thigh length 17.3 mm, shank length 14.0 mm, foot length 15.1 mm. Dorsum with a dark, finely lichenous pat- tern of black and brown; brown interorbital band; face mostly black with three indis- tinct, slanted stripes from eye to edge of upper lip, most anterior stripe directed for- ward from eye, other two directed poste- riorly; flank same color as dorsum; inguinal gland uniformly dark; upper limb surfaces barred black and brown; tips of digits white; venter black with small distinct white dots; under-surfaces of hands and feet gray with all tubercles and digit tips conspicuously white; posterior surface of thigh dark brown with small, irregular, distinct, light spots. Etymology. —Named for Dr. Miguel T. VOLUME 101, NUMBER 1 Rodrigues, who has made significant col- lections of frogs throughout much of Brazil. Distribution. —Known only from the type locality (Fig. 2). Discussion The occurrence of Cycloramphus fuligi- nosus in the State of Bahia is surprising for two reasons. First, no member of any stream associated species group of Cycloramphus was expected north of the Rio Doce (Heyer & Maxson 1983:356). However, small, swift, clear streams obviously occur north of the Rio Doce, comparable in habitat to those found south of the river. Intensive collecting now will have to be done to ascertain the northern distributional limits of Cyclo- ramphus in the Atlantic Forest system. Sec- ond, the occurrence of C. fuliginosus is in itself surprising. All other members of the same species group have much smaller geo- graphic ranges (see Heyer 1983, and Heyer & Maxson 1983) than C. fuliginosus. As cur- rently understood, C. fuliginosus is not con- tinuously distributed throughout its range; rather it is known from three small areas in the State of Rio de Janeiro (in or near the city of Rio de Janeiro, Corcovado, and the Serra da Mangaratiba [Heyer 1983, fig. 38]); one locality in Espirito Santo (Santa Tere- sa); and now two nearby localities in south- ern Bahia (Fig. 2). There is no morpholog- ical differentiation among the samples from the states of Rio de Janeiro, Espirito Santo, or Bahia. In fact, all the Bahia and Espirito Santo individuals have the distinct tripar- tite odontoids on the lower jaw pointed out by A. Lutz (1929:10) as characterizing C. fuliginosus. As this characteristic often ap- pears late in the ontogeny of C. fuliginosus, being found consistently only in large spec- imens (Lutz 1929:10), I did not include the character in my previous work (Heyer 1983). Additional data confirm Lutz’s observation that C. fuliginosus is the only species of Cy- cloramphus to have bipartite or tripartite mandibular odontoids. Thus, it would ap- 153 Fig. 2. Distribution of Cycloramphus migueli (tri- angle), C. fuliginosus (dots and triangle), and C. stejne- geri (Square) in eastern Brazil. Cycloramphus stejnegeri geographically is the closest member of the species clus- ter to which C. migueli belongs. State initials, BA = Bahia, ES = Espirito Santo, MG = Minas Gerais, RJ = Rio de Janeiro. pear that C. fuliginosus either (1) has until relatively recently had a continuous distri- bution but now Is divided into relictual pop- ulations, or (2) has been separated for a longer period of time but has not undergone morphological differentiation characteristic of other species of Cycloramphus isolated for similar periods of time. Estimates of ge- netic relatedness among the Rio de Janeiro, Espirito Santo, and Bahia populations are needed to resolve the issue. Cycloramphus migueli has a morphology indicative of a fossorial existence and, along with its presumed closest relatives (bolito- glossus, carvalhoi, and stejnegeri) most like- ly has some form of direct development (Heyer & Crombie 1979). These four species are allopatric, each with a small geographic range (bolitoglossus occurs in the Serra do Mar in the states of Parana and Santa Ca- tarina, carvalhoi in the Serra da Manti- queira, and stejnegeri in the Organ Moun- 154 tains, Fig. 2 and see Heyer 1983, figs. 30, 42). Assuming that C. migueli is closely re- lated to bolitoglossus, carvalhoi, and stejne- geri, it is not surprising that a population found in Bahia would represent a new species, nor that a member of this cluster would occur in the Atlantic Forests north of the Rio Doce as species with direct de- velopment should be able to occur through- out the Atlantic Forest Domain. From the same localities in Bahia, Dr. Rodrigues collected species of Crossodac- tylus and Hylodes, two genera typically found in the same streams as Cycloramphus south of the Rio Doce. Notable by its absence from these collections is a representative of the genus Thoropa, which elsewhere is found in the same types of habitats as Cyclo- ramphus. Acknowledgments Dr. Miguel T. Rodrigues (Instituto de Biociéncias, Universidade de Sao Paulo) generously turned the Bahia Cycloramphus materials over to me for study and also pro- vided his ecological notes for the specimens. Reginald B. Cocroft and Ronald I. Crombie (Smithsonian Institution) reviewed the manuscript. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Research for this paper was supported in. part by the Museu de Zoologia da Univer- sidade de Sao Paulo (MZUSP) and the Smithsonian Institution’s IESP Neotropical Lowland Research Program. Literature Cited Heyer, W.R. 1983. Variation and systematics of frogs of the genus Cycloramphus (Amphibia, Lepto- dactylidae).— Arquivos de Zoologia 30:235-339. ——,&R.I. Crombie. 1979. Natural history notes on Craspedoglossa stejnegeri and Thoropa pet- ropolitana (Amphibia: Salientia, Leptodactyli- dae).—Journal of the Washington Academy of Sciences 69:17-20. , & L. R. Maxson. 1983. Relationships, zoo- geography, and speciation mechanisms of frogs of the genus Cycloramphus (Amphibia, Lepto- dactylidae).— Arquivos de Zoologia 30:341-373. Lutz, A. 1929. Taxonomia e biologia do genero Cy- cloramphus.—Memorias do Instituto Oswaldo Cruz 22:5-16 (Portuguese), 17—25 (English) + 5 plates. Department of Vertebrate Zoology, Am- phibians and Reptiles, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 155-161 REDESCRIPTIONS OF TWO CHINESE CUORA (REPTILIA: TESTUDINES: EMYDIDAE) Carl H. Ernst Abstract. — Validity of the turtle Cuora pani Song, 1984, from Shaanxi Prov- ince, China, which has become available to American and European scientists via the pet trade, has been questioned. It has been confused with Cuora yun- nanensis Boulenger, 1906, a poorly known species not seen since the type series was collected from two highland sites in Yunnan Province, China. Major dif- ferences exist, and the two species are redescribed and illustrated. In the last year several taxa of Chinese box turtles, family Emydidae, have become available to the scientific community of North America and Europe through the pet trade. These turtles have all been mistak- enly referred to as Cuora yunnanensis (Bou- lenger 1906), a highland species from Yun- nan Province, China, and one that has not been collected since the type series. Most of the imported turtles are the recently named, but poorly described Cuora pani Song, 1984. The original descriptions of both C. yun- nanensis and C. pani are vague and suggest many similarities that do not exist. Cuora pani was described in Chinese with a brief English summary, thus contributing to the confusion of non-Chinese readers. To clear- ly differentiate these two taxa, I offer the following descriptions and illustrations. Methods and materials. —Straight-line measurements of each specimen were taken with dial calipers accurate to 0.1 mm of the greatest carapace length, carapace width and depth of the level of the seam between ver- tebrals 2 and 3, marginal width (the differ- ence between the carapacial width and the width across the pleurals taken between the points of juncture of the marginals and pleu- rals at the level of the seam between ver- tebrals 2 and 3), greatest plastron length, greatest width and length of both plastral lobes, greatest bridge length, greatest width and length of vertebrals 1 and 2 and pleural 2, and the medial seam length and greatest width of all plastral scutes. Careful notes and drawings were made of head, neck, limb, carapacial, plastral, and bridge patterns. Colors were recorded from living turtles and color transparencies. Shell proportions are expressed as ratios of one measurement to another. Sixteen ratios proved useful (ab- breviations used in the text are given in parentheses): width/length of cervical scute (W/L CS), width/length of first vertebral (W/L Ist V), width/length of second ver- tebral (W/L 2nd V), width/length of second pleural (W/L 2nd Pl), marginal width/car- apacial width (MW/CW), marginal width/ carapacial length (MW/CL), carapacial width/carapacial length (CW/CL), carapa- cial depth/carapacial length (D/CL), cara- pace depth/carapacial width (D/CW), plas- tral length/carapacial length (PL/CL), bridge length/plastral length (B/PL), bridge length/ carapacial length (B/CL), length of anterior plastral lobe/plastral length (APL/PL), width of anterior plastral lobe/plastral length (APW/PL), length of posterior plastral lobe/ plastral length (PPL/PL), and width of pos- terior plastral lobe/plastral length (PPW/ PL). The number of rows of large scales at the lateral edge of the antibrachium between the claw of digit V and the first horizontal skin fold proximal to the elbow (presented in text as foreleg scale rows) was recorded. Specimens from the following collections 156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Carapace and plastron of Cuora yunnanensis. VOLUME 101, NUMBER 1 were examined (abbreviations used in the text are given in parentheses): British Mu- seum of Natural History (BMNH), William H. Randel, Hatboro, Pennsylvania, person- al collection (WHR), National Museum of Natural History, Smithsonian Institution (USNM). Cuora yunnanensis (Boulenger, 1906) Figs. 1, 2 Cyclemys yunnanensis Boulenger, 1906:567. Cuora yunnanensis Smith, 1931:88. Syntypes. —-BMNH 1946.1.22.97 (female) from Tongchuan Fu, Yunnan Province, China, and 1946.1.22.98-99, 1946.1.23.1- 3 (4 males, | female) from Yunnan Fu, Yun- nan Province, China; John Graham. Specimens examined. —BMNH 1946.1.- 22.1-3; 1946.1.23.98-99. Diagnosis.—A brown species of Cuora with a flattened, tricarinate carapace, a pos- teriorly notched plastron with a complete interanal seam, the plastral seams narrowly marked with dark pigment, brown skin, a light postorbital stripe, and large yellow or orange blotches on the chin and throat. Description. —Carapace length to 140 mm (males 126, females 140, measurements of largest specimen taken from Boulenger 1906), elliptical, flattened (D/CL, 0.33-0.39, X = 0.352; D/CW, 0.49-0.51, x = 0.502; CW/CL 0.65-0.77, x = 0.704); widest at level of marginals 8—9, highest at posterior of vertebral 3 (posterior of vertebral 2 in one specimen). Sides straight with no in- dentation at the bridge. Marginals flared, those most posterior slightly serrated, the first anterior marginals widest, and those over the bridge narrowest (MW 11.0-16.6 mm, x = 13.2; MW/CW 0.18-0.215, < = 0.202; MW/CL, 0.13-0.17, x = 0.143). There is a slight notch between the twelfth marginals. Cervical scute longer than wide (W/L CS 0.76-0.89, X = 0.79). Each ver- tebral scute wider than long, but much nar- Sy7/ rower than the pleural scutes. The first and fifth vertebrals are flared, but the first does not extend to the seams separating margin- als 1 and 2 on each side. General texture of carapacial scutes smooth in older individ- uals, but granulated with low, rounded ru- gosities in those up to 88 mm CL. Three longitudinal keels are present; that extend- ing medially along the vertebrals is highest, while the lateral two, which extend along the dorsal third of the pleural scutes on each side, are lower and may be almost non- existent in larger individuals. Carapace in preserved specimens is olive-brown to chestnut-brown with narrow black seams and some yellow along the pleural-marginal border. Plastral length to 115.5 mm (males 115.5, females 62.4, but largest female reported by Boulenger (1906) not examined). PL/CL 0.92-0.94, x = 0.928; hinged between the pectoral and abdominal scutes, slightly up- turned anteriorly and posteriorly notched. The posterior plastral lobe is longer and wider than the anterior lobe (APL/PL 0.42- 0.45; x = 0.438; PPL/PL 0.56-0.58, x = 0.568; APW/PL 0.45-0.49; x = 0.480; PPW/PL 0.51-0.54, * = 0.526). Lateral margins of the femoral and anal scutes curved gradually toward the midline. Bridge moderate in length with a small axillary and a small inguinal scale; B/PL 0.28-0.31, x = 0.297; B/CL 0.26-0.29, X = 0.276. The av- erage plastral formula is Abd. > Pect. > An. > Gul. > Fem. > Hum. The interanal seam is complete. Plastron light brown with dark brown or black seams. A dark bar is present on the bridge, and the undersides of the marginals are yellow with dark seams and some faded dark pigment. The head is narrow, the snout pointed and only slightly projecting, and the upper jaw is slightly hooked. Coloration is olive to brown. A narrow yellow black-bordered stripe extends backward from the eye above the tympanum and onto the neck, and a second, similar stripe extends posteriorly from the corner of the mouth, passes below PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 158 hth Reig ae eRe. oe Head of Cuora yunnanensis. Fig. 2. VOLUME 101, NUMBER 1 the tympanum and onto the neck. The chin and throat are marked with large yellow to orange blotches. The jaws are yellow to tan. The neck is brown with two narrow yellow or orange, black-bordered, longitudinal stripes on each side. The digits are fully webbed. The forelegs are brown with longitudinal stripes of yel- low or orange spots. There are several large, band-like, transverse scales on the anterior surface of each foreleg; foreleg scale rows 9- 11, x = 10. The hindlegs are brown with a narrow outer fringe of small yellow scales. The tail is brown with two black-bordered, yellow dorsal stripes. Males have concave plastra, and longer tails with the vent beyond the carapacial rim; females have flat plastra and shorter tails with the vent beneath the posterior marginals. The female carapace is more domed than that of the male. Cuora pani Song 1984 Fig. 3 Holotype. —Shaanxi Institute of Zoology 80170 (adult male); Al/otype.—SIZ 80171 (adult female), both from Xujiaba, Shaanxi Province, China; 17 Jun 1981, Song Ming- Tao. Specimens examined. —WHR 1-11 (live; 5 males, 6 females) USNM 4 males). Diagnosis. —A species of Cuora with red- dish-brown vertebrals on an olive to dark brown, carinate carapace, a posteriorly notched plastron with a complete inter- anal seam, a plastral pattern of large, often separated, black blotches, yellow skin, a yel- low postorbital stripe, and an unmarked chin and throat. Description. —Carapace length to 156.5 mm (males 120 from Song 1984; females 156.5), oval, flattened. (D/CL 0.30-0.39, X = 0.352; D/CW, 0.35-0.50, x = 0.467; CW/CL 0.64-0.79, x = 0.729); widest at level of marginal 8, highest near seam sep- arating vertebrals 2 and 3. Sides straight with no indentation at the bridge. Marginals 159 flared, those most posterior slightly serrat- ed, the first anterior marginals widest, those along the sides may be slightly upturned (MW 9.2-18.8 mm, x = 12.7; MW/CW 0.16-0.24, x = 0.197; MW/CL 0.12-0.17, X = 0.143). There is a slight notch between the twelfth marginals. Cervical scute longer than wide (W/L CS 0.35-0.91, x = 0.710). Each vertebral scute wider than long, but narrower than the pleural scutes. The first and fifth vertebrals are flared. General tex- ture of the carapacial scutes smooth in older individuals, but granulated with low, rounded rugosites in those to 84 mm CL. A low keel extends medially along the ver- tebrals; no lateral keels are present. Cara- pace in live individuals is olive-brown with chestnut-brown to reddish-brown pigment on the vertebrals, and occasionally as spots on the pleurals, and with a narrow yellow rim. Plastral length to 146.5 mm (males 84.0, but PL not reported for large male in Song 1984; females 146.5); PL/CL 0.91-0.97, X = 0.932; hinged between the pectoral and abdominal scutes, slightly upturned ante- riorly and notched posteriorly. The poste- rior plastral lobe is longer and wider than the anterior lobe (APL/PL 0.40-0.46, x = 0.423; PPL/PL 0.54-0.58, x = 0.560; APW/ PL 0.48-0.54, x = 0.502; PPW/PL 0.51- 0.60, xX = 0.551). Lateral margins of the fem- oral and anal scutes curve gradually toward the midline. Bridge moderate in length with a small axillary and a small inguinal scale; B/PL 0.27-0.35, X = 0.311; B/CL 0.25-0.34, X = 0.290. The average plastral formula is An. > Pect. > Abd. > Gul. > Fem. > Hum., but much variation exists in the 15 specimens examined, 4 had the average for- mula, 5 were An. > Abd. > Pect. > Gul. > Hum. > Fem.; 3 each were An. > Abd. > Pect. > Gul. > Fem. > Hum. and Pect. > An. > Abd. > Gul. > Fem. > Hum. The interanal seam is complete. Plastron yellow with large, triangular-shaped black blotches which may touch, but are often separated. A black bar is present on the bridge, and 160 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON * * - - area, Fig. 3. Carapace and plastron of Cuora pani. VOLUME 101, NUMBER 1 the undersides of the marginals are yellow, with some black pigment on those at the bridge. The head is narrow, the snout is pointed but nonprojecting, and the upper jaw is slightly hooked. Coloration is lemon-yel- low. A narrow, black-bordered stripe ex- tends posteriorly from the eye onto the neck, and a second similar stripe runs backward from the corner of the mouth through the tympanum and onto the neck. Both of these stripes almost disappear in older individ- uals. The iris in life is greenish-yellow. The jaws, chin and throat are immaculate yel- low. The neck is greenish-yellow dorsally and laterally, but yellow ventrally; narrow often broken, yellow stripes may occur on the sides or dorsal surface. The digits are fully webbed. The limbs are olive on the outer surface but yellow to orange beneath. There are several large, band-like, transverse scales on the anterior surface of each foreleg; foreleg scale rows 10-15, x = 11.7. The tail is olive with two black-bordered, yellow-dorsal stripes. 161 Sexual dimorphism as in C. yunnanensis. Acknowledgments I wish to thank William H. Randell for calling my attention to this problem, and for allowing me to examine his live speci- mens of Cuora pani. George R. Zug com- mented on an early draft of the manuscript, and Evelyn M. Ernst helped in the exami- nation of the specimens. Literature Cited Boulenger, G. A. 1906. Descriptions of new reptiles from Yunnan.—Annals and Magazine of Nat- ural History (7)17:567-568. Smith, M. A. 1931. The fauna of British India, in- cluding Ceylon and Burma. Reptilia and Am- phibia. Volume I. Loricata, Testudines. Lon- don, Taylor and Francis. 185 pp. Song, M-T. 1984. A new species of the turtle genus Cuora (Testudoformes: Testudinidae).— Acta Zootaxonomica Sinica 9:330-332. Department of Biology, George Mason University, Fairfax, Virginia 22030. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 162-170 A NEW SPECIES OF PORPHYRIO (AVES: RALLIDAE) FROM ARCHEOLOGICAL SITES IN THE MARQUESAS ISLANDS David W. Steadman Abstract.—A new species of gallinule, Porphyrio paepae, is described from bones found in archeological sites on Hiva Oa and Tahuata, Marquesas Islands. This gallinule represents the only species of Porphyrio in the Pacific east of Samoa, Tonga, and Niue, where P. porphyrio occurs today. The occurrence of a species of Porphyrio in the Marquesas Islands, at the northeastern edge of Polynesia, suggests that congeners may have occurred through much of eastern Polynesia in prehistoric times. Bones from Polynesian archeological sites have shown that the avifaunas of individual islands were much richer in species before human colonization. Here I describe a new species of gallinule of the genus Porphyrio from archeological sites on Hiva Oa and Tahuata islands in the Marquesas. The sites on Hiva Oa were described by Bellwood (1972, 1979:331-337) and Sinoto (1979). The site on Tahuata was excavated recently by B. Rolett, whose studies are in progress. The extinct landbirds from sites on both islands are reviewed briefly by Steadman (in press) and include new species of parrots (Steadman & Zarriello, 1987). Materials and methods.—Comparative skeletons are from the Division of Birds, USNM. Aside from those of Porphyrio (see Tables 1—4), skeletons of Gallirallus philip- pensis, Gallirallus owstoni, Gallicrex cine- rea, Gallinula chloropus, and Fulica atra, as well as fossils of undescribed species of Gal- lirallus from the Marquesas and elsewhere, were examined to determine generic rela- tionships. Abbreviations: BPBM = Bernice P. Bishop Museum; USNM = National Mu- seum of Natural History, Smithsonian In- stitution; yr B.P. = corrected radiocarbon years before present. Osteological termi- nology usually follows that of Baumel et al. (1979). Measurements were taken with dial calipers with 0.05 mm increments, rounded to the nearest 0.1 mm. Systematic Paleontology Class Aves Order Gruiformes Family Rallidae Genus Porphyrio Following Olson (1973:409), I regard the genus Porphyrio to include “Porphyrula”’ and ‘“‘Notornis.” The fossil material is re- ferred to Porphyrio rather than to other large rails (Gallirallus, Gallicrex, Gallinula, Fu- lica) because of the following characters. Carpometacarpus—processus pisiformis extends farther cranially; fovea carpalis cau- dalis deeper; great distance from processus pisiformis to departure of os metacarpi ma- jus and os metacarpi minus; short distal symphysis of os metacarpi majus and os metacarpi minus. Synsacrum—stout over- all; strongly bent ventrally in caudal por- tion. Femur—more obtuse angle formed at the junction of the impressiones obturato- riae and trochanter femoris; similar size and position of the impressiones iliotrochanter- ia and linea intermuscularis caudalis; me- dial margin of the condylus medialis ori- ented roughly parallel to the shaft rather than diagonal. Tibiotarsus—deep sulcus VOLUME 101, NUMBER 1 ee 2g eee Woe Cad Mays 7 A B C Fig. 1. 163 D E F The femur of Porphyrio in cranial (A-C) and caudal (D-F) aspects. A, D, P. martinica, USNM 501122, male; B, E, P. paepae, holotype, BPBM 165649, unsexed; C, F, P. porphyrio, USNM 488868, male. Scale bar = 10 mm. musculo fibularis; sharp cranio-medial and cranio-lateral margins of the shaft; deep in- cisura intercondylaris. Tarsometatarsus— extremely thin proximo-medial portion of shaft; more distinct fossa metatarsi. Pedal phalanx — more elongate. Porphyrio paepae, new species Figs. 1-4 Holotype. — Femur lacking only the me- dial portion of facies articularis acetabular- is, BPBM 165649, from Hanatekua Shelter No. 2 Site (MH3-12), square E2-F2, 100- 140 cm, Hiva Oa, Marquesas Islands, 19 Dec 1967, P. Bellwood. (This site has been referred to as Hanatukua, with an earlier Bishop Museum site designation of MH- 11.) Paratypes. —Hiva Oa, 1967, P. Bellwood: Pelvis—BPBM 165656, Hanatekua Shelter No. 2 Site (MH3-12), square D1, 80-100 cm. Femur (tentatively referred)—BPBM 168539, Hanatekua Beach Site (MH3-13, formerly designated MH-12), test pit 24, 40+ cm. Four tibiotarsi—BPBM 165651, Hanatekua Shelter No. 2 Site (MH3-12), square E1-H1, layer 10 (section collapse); BPBM 165652, Hanatekua Beach Site (MH3-13), test pit 2, layer 4; BPBM 167496, Hanatekua Beach Site (MH3-13), test pit 22, layer 4, 40+ cm; BPBM 165654, Ha- naui Site (MH-22), test pit 8. Two tarso- metatarsi—BPBM 165650, Hanatekua Shelter No. 2 Site (MH3-12), square E2-F2, 100-150 cm; BPBM 165653, Hanatekua Beach Site (MH3-13), test pit 23, 30+ cm. Tahuata, Hanamiai Site (MT1), 1985, B. Rolett: Carpometacarpus—BPBM 166427, square L11, layer 137. Synsacrum—BPBM 166432, square L10, layer G/H16 (240-250 cm). Four femora—BPBM 166434, square 112, layer G/H14 (230-240 cm); BPBM 166430, square L11, layer 136; BPBM 166426, square L1l, layer 137; BPBM 166424, square K9, layer F10 (180+ cm). 164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A B C D E F Fig. 2. The tibiotarsus of Porphyrio in cranial (A—C) and caudal (D-F) aspects. A, D, P. martinica, USNM 501122, male; B, E, P. paepae, BPBM 165651, unsexed; C, F, P. porphyrio, USNM 488868, male. Scale bar = 10 mm. Two tibiotarsi—BPBM 166428, square M13, layer H31 (255-260 cm); BPBM 166433, square N13, layer H33 (260-265 cm). Two tarsometatarsi—BPBM 166429, square K11, layer G/H16 (220-230 cm); BPBM 166425, square K 10, layer G13 (220- 230 cm). Diagnosis. —Smaller than any species of Porphyrio except the ‘“‘Porphyrula’’ group (P. martinica, P. alleni, P. flavirostris) (Ya- bles 1-4). Carpometacarpus: short; os meta- carpi minus and os metacarpi majus fused proximally for a greater distance. Femur: shaft broader than in P. martinica; shaft straighter than in P. porphyrio, P. polioce- phalus, or P. pulverulentus; sulcus patellaris broader than in P. martinica; impressio lig- amentum cruciati caudalis deeper than in P. martinica; in medial aspect, condylus medialis joins caudal surface of the shaft nearly perpendicularly (more obtuse angle in all others); in lateral aspect, the proximal margin of crista trochanteris less rounded than in P. porphyrio. Tibiotarsus: shaft rel- atively shallower than in P. porphyrio; in- cisura intercondylaris relatively broader and deeper than in all others; proximal portion of condylus lateralis more rounded and joins the shaft more perpendicularly (an obtuse angle is formed in all others). Etymology. —From the Marquesan word paepae, which is a prehistoric rectangular stone platform that served as a base for var- ious dwellings (Bellwood 1979:333 and fig. 12.3). Like the extinct gallinule described here, the paepae is no longer an active part of Marquesan culture, although its remains are scattered on the islands. Remarks.—No skeletons were available for Porphyrio alleni of Africa or P. flaviros- tris of South America. The former is about the size of P. martinica, while the latter is smaller than P. martinica. Discussion The chronology of the Hanatekua Shelter No. 2 Site is poorly defined, although the bones of Porphyrio paepae from this site may be from pre-cultural levels (Y. H. Sin- VOLUME 101, NUMBER 1 165 A B C D E F Fig. 3. The tarsometatarsus of Porphyrio in dorsal (A-C) and plantar (D-F) aspects. A, D, P. martinica, USNM 501122, male; B, E, P. paepae, BPBM 165650, unsexed; C, F, P. porphyrio, USNM 488868, male. Scale bar = 10 mm. A B Cc D E F Fig. 4. The carpometacarpus of Porphyrio in dorsal (A—C) and ventral (D-F) aspects. A, D, P. martinica, USNM 501122, male; B, E, P. paepae, BPBM 166427, unsexed; C, F, P. porphyrio, USNM 488868, male. Scale bar = 10 mm. 166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Measurements (in mm) of the carpometacarpus of Porphyrio, with mean, range, and sample size. F = female. M = male. U = sex unknown. Total length Proximal width Proximal depth Least depth of shaft Porphyrio paepae 29.3 6.8 3.6 Del Tahuata, Marquesas (U) 1 1 1 1 BPBM 166427 Porphyrio porphyrio 45.2 9.5 4.8 3.0 Bechuanaland (M) 1 1 1 1 USNM 488868 Porphyrio porphyrio — 10.5 4.9 = New Caledonia (M) 1 1 USNM 561547 Porphyrio porphyrio 47.1 10.0 4.8 3.6 New Caledonia (F) 47.0-47.2 9.9-10.0 4.8 3.5-3.7 USNM 561549, 561551 2 2 2 2 Porphyrio porphyrio 52.4 10.8 5.0 3.9 Captive (M) USNM 52.0-52.9 10.8-10.9 4.9-5.2 3.9 555504, 557529 2 2 2 2 Porphyrio poliocephalus 43.5 9.5 4.4 2.8 Thailand (F) 1 1 1 1 USNM 343212 Porphyrio poliocephalus 54.5 11.5 5.3 3.6 Captive (M) USNM 491890 1 1 1 1 Porphyrio poliocephalus 43.5 10.0 4.3 3.4 Captive (F) USNM 347372 1 1 1 1 Porphyrio pulverulentus 47.0 10.0 4.7 3.4 Philippines (M, U) 46.5—47.4 9.9-10.2 4.6-4.8 3.3-3.5 USNM 200817, 226035 DD 2 2 2 Porphyrio martinica 32.6 7.1 3.4 2.1 Florida (M) 31.6-33.1 6.8-7.5 3.2-3.5 2.0—2.2 USNM 501122, 501126, 5 5 5 5 501128, 501129, 501131 Porphyrio martinica 30.8 6.6 3.2 Del Florida (F) USNM 30.6-31.3 6.3-6.9 3.0-3.4 2.0-2.3 501123-501125, 501130 4 4 4 4 oto, pers. comm.). Kirch (1986:25) reported a radiocarbon date of 1930 + 80 yr B.P. for Hanatekua Shelter No. 2, although I am not sure how this date relates stratigraphically to the fossils of P. paepae. The chronology of the Hanamiai Site on Tahuata is cur- rently being resolved and lies mainly in the range of 1500 to 800 yr B.P. (B. Rolett, pers. comm.). All bones of P. paepae from Ha- namiai are associated with Polynesian ar- tifacts from the lowest cultural levels. The only other large rails from the Mar- quesas are extinct, undescribed, flightless species of Gallirallus from Ua Huka, Nuku Hiva, Hiva Oa, and Tahuata. The wing ele- ments in these species are much smaller than those in any species of Porphyrio, including P. paepae. Based upon the size and mor- phology of the carpometacarpus, P. paepae had somewhat reduced powers of flight compared to congeners, yet probably was not flightless. A crude indication of volancy is presented in Table 5 by the ratio of the total lengths of the carpometacarpus and femur (the only wing or leg bones known in P. paepae for which the total length could VOLUME 101, NUMBER 1 167 Table 2.— Measurements (in mm) of the femur of Porphyrio, with mean, range, and sample size. F = female. M = male. U = sex unknown. Porphyrio paepae Hiva Oa, Marquesas (U) Holotype BPBM 165649 Porphyrio paepae Tahuata, Marquesas (U) BPBM 166430, 166434 Porphyrio porphyrio New Caledonia (M) USNM 561547 Porphyrio porphyrio New Caledonia (F) USNM 561549, 561551 Porphyrio porphyrio Bechuanaland (M) USNM 488868 Porphyrio porphyrio Captive (M) USNM 555504, 557529 Porphyrio poliocephalus Thailand (F) USNM 343212 Porphyrio poliocephalus Captive (M) USNM 318498, 344474, 491890 Porphyrio poliocephalus Captive (F) USNM 289454, 321151, 347372 Porphyrio poliocephalus Captive (U) USNM 19761, 227119 Porphyrio pulverulentus Philippines (M) USNM 226035, 292297 Porphyrio pulverulentus Philippines (F) USNM 292296 Porphyrio martinica Florida (M) USNM 501122, 501126, 501128, 501129, 501131 Porphyrio martinica Florida, Louisiana, Dominican Republic (F) USNM 288205, 499438, 501123-501125, 501130 Total length 52.5 72.6 72.4-72.8 2 74.5 1 76.8 73.8-79.8 a 68.8 1 76.2 62.7-83.7 3 67.0 64.6-70.7 3 69.0 61.2-76.8 2 78.2 77.9-78.4 2 75.4 1 54.2 52.8-55.4 4 50.7 47.7-52.4 6 Depth of proximal end Well 1 11.2 11.0-11.4 2 9.3 11.2 11.0-11.4 2 10.8 1 Least width of shaft Least depth of shaft Width of distal end 13.8 12.6 12.3-12.9 2 13.5 13.3-13.7 2 11.7 1 13.2 10.8-14.6 3 11.3 10.9-11.8 3 12.0 10.8-13.3 a) 13.8 13.4-14.2 ey 13.4 1 Depth of condylus medialis 7.0 1 10.0 9.6 9.6-9.7 2 11.3 11.2-11.4 2 9.8 9.4 8.1-10.8 2 11.1 10.5-11.7 ” 10.8 168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.—Measurements (in mm) of the tibiotarsus of Porphyrio, with mean, range, and sample size. F = female. M = male. U = sex unknown. PE = proximal end. PS = pons supratendineus. Width of Height of _—_ Height of Depth of Depth of Least width shaft at Least depth condylus condylus condylus condylus of shaft PE of PS of shaft Distal width medialis lateralis medialis lateralis Porphyrio paepae 4.0 6.4 3.4 9.8 5.8 8.2 9.6 8.9+ Hiva Oa (U) BPBM 3.7-4.2 6.1-6.7 3.3-3.5 1 1 1 1 1 165651, 165652, 3 yD) 4 165654, 167496 Porphyrio paepae — 5.8 3.4 8.6+ 5.4+ 7.3 8.5+ 8.3 Tahuata (U) BPBM 5.7-6.0 3.4 1 1 1 1 1 1664228, 166433 2, 2 Porphyrio porphyrio 5.1 7.2 3.8 10.2 6.0 9.1 10.5 10.2 Bechuanaland (M) 1 1 1 1 1 1 1 1 USNM 488868 Porphyrio porphyrio 6.0 8.1 4.8 11.1 7.0 9.8 11.6 11.3 New Caledonia (M) 1 1 1 1 1 Line 1 1 USNM 561547 Porphyrio porphyrio 5.6 7.6 4.3 10.4 6.4 9.2 11.0 10.4 New Caledonia (F) 5.4-5.9 7.4-7.8 4.3 10.3-10.5 6.3-6.4 9.2-9.3 10.8-11.1 10.2-10.5 USNM 561549, 561551 2 D, 2 yD, 2 2 2} 2 Porphyrio porphyrio — 7.7 — 10.6 6.3+ 9.9 10.9+ 10.6+ Tikopia, Solomon Is. 1 1 1 1 1 1 BPBM 166227, 166263 Porphyrio porphyrio 5.7 8.6 4.6 11.6 6.8 10.2 12.0 11.2 Captive (M) 5.5-5.9 8.3-8.9 2 11.5-11.8 6.8-6.9 10.2-10.3 11.9-12.1 10.9-11.5 USNM 555504, 557529 2 D yp) D 2) D} 2 Porphyrio poliocephalus 4.4 6.6 3.5 9.2 5.6 8.2 9.9 9.2 Thailand (F) 4.3-4.6 6.4-6.7 3.4-3.6 9.1-9.2 5.3-5.9 8.0-8.3 9.6-10.2 2 USNM 343212, 343213 D) 2 2 2 D D) 2 Porphyrio poliocephalus 5.2 7.6 4.0 10.6 6.2 9.0 11.1 10.0 Captive (M) 4.5-5.8 6.8-8.5 3.44.6 9.1-12.0 5.3—7.1 7.8-10.3 9.2-13.0 8.6-11.3 USNM 344474, 491890 2 2 2 yD) 2 2 2 2 Porphyrio poliocephalus 4.8 6.8 4.0 9.7 6.0 8.4 10.0 9.4 Captive (F) 4.7-4.9 6.5-7.1 3.6-4.3 1 2 8.1-8.6 1 9.3-9.4 USNM 321151, 347372 D 2 2 2 2) Porphyrio pulverulentus 5.5 7.8 4.2 11.2 6.5 10.0 11.4 10.7 Philippines (M, U) 5.4-5.6 7.7-7.9 4.1-4.4 2 6.3-6.7 9.9-10.1 11.0-11.8 10.5-10.9 USNM 200817, 226035 2 2 2 2 2 2, 2 Porphyrio martinica 3.5 5.0 Del Toll 4.3 6.2 7.0 6.9 Florida (M) 3.3-3.7 4.8-5.4 2.6-2.9 6.9-7.5 4.1-4.7 6.1-6.4 6.9-7.3 6.8—7.1 USNM 501122, 501126, 5 5 5 5 5 5 5 5 501128, 501129, 501131 Porphyrio martinica 3.4 4.9 2.6 6.8 4.2 6.0 6.7 6.6 Florida, Louisiana, 2.8-3.6 4.7-5.1 2.5-2.8 6.3-7.5 3.9-4.4 5.8-6.3 6.1-7.0 6.2-7.1 Dominican Republic (F) 6 6 6 6 6 6 6 6 USNM 288205, 499438, 501123-501125, 501130 be determined). Although the value of this fossils are of unknown sex. Note also that ratio is less in P. paepae than in other con- the skeletons of captive individuals of P. geners, the difference is small and might be porphyrio and P. poliocephalus have higher of little significance, especially because the ratios than those of wild individuals. For VOLUME 101, NUMBER 1 169 Table 4.—Measurements (in mm) of the tarsometatarsus of Porphyrio, with mean, range, and sample size. F = female. M = male. U = sex unknown. Minimum width ofMinimum depth of shaft Porphyrio paepae 4.9 Hiva Oa, Marquesas (U) 1 BPBM 165650 Porphyrio paepae = Tahuata, Marquesas (U) BPBM 166425 Porphyrio porphyrio 5.0 Bechuanaland (M) 1 USNM 488868 Porphyrio porphyrio — New Caledonia (M) USNM 561547 Porphyrio porphyrio 5.5 New Caledonia (F) 5.4-5.6 USNM 561549, 561551 2 Porphyrio porphyrio — Tikopia, Solomon Is. (U) BPBM 166228, 166230 Porphyrio porphyrio 5.9 Captive (M) 5.7-6.1 USNM 555504, 557529 2 Porphyrio poliocephalus 4.6 Thailand (F) USNM 343212 1 Porphyrio poliocephalus 6.5 Captive (M) USNM 491890 1 Porphyrio poliocephalus 5.2 Captive (F) USNM 347372 1 Porphyrio pulverulentus 5.5 Philippines (M, U) 5.1-5.9 USNM 200817, 226035 y) Porphyrio martinica 3.8 Florida (M) 3.7-3.9 USNM 501122, 501126, 5 501128, 501129, 501131 Porphyrio martinica 3.6 Florida (F) 3.2-3.9 USNM 501123-501125, 4 501130 shaft Proximal width Width of hypotarsus 2.9 _ _ 1 — 10.4 5.1 1 1 3.6 10.9 5.4 1 1 1 3.9 11.8 5.8 1 1 1 3.6 10.9 5.4 3.6-3.7 10.8-11.0 5.3-5.5 2 Dp; yy 3.7 — 5.6 1 1 4.2 12.4 5.8 4.1-4.2 12.3-12.5 5.8-5.9 yy) 2 2 3D 9.9 4.7 1 1 1 4.3 12.4 6.3 1 1 1 3.3 10.4 5.0 1 1 1 3.8 11.6 6.0 3.74.0 11.5-11.6 6.0-6. 1 2 2 2 2.5 7.6 4.0 2.3-2.6 7.3-7.9 3.84.3 5 4 4 2.5 7.0 3.6 2.4-2.6 6.7—-7.2 3.4-3.7 4 4 4 this and other reasons, I do not consider the captive individuals to be an absolutely ac- curate representation of either the quanti- tative (Tables 1—5) or qualitative characters of P. porphyrio or P. poliocephalus. The di- agnosis 1s based only upon wild individuals. In all of the tables, the values for P. mar- tinica provide the best assessment of the intraspecific variation of wild populations. Balouet & Olson (in press) have described a flightless, extinct species of Porphyrio from late Holocene fossils from New Caledonia. This new species was much larger even than P. porphyrio, which is the only other con- gener in the Pacific, other than the large, flightless species on New Zealand (“‘Notor- nis’’) and the historically extinct P. albus of Lord Howe Island, which was similar in size 170 Table 5.—Ratio of total lengths: carpometacarpus divided by femur, expressed as a percentage. From data in Tables | and 2. Porphyrio paepae 56.7 (—) Tahuata, Marquesas (U) 1 BPBM 166427, 166434 Porphyrio paepae 55.8 Tahuata and Hiva Oa, Marquesas (U) 1 BPBM 166427, 165649 Porphyrio porphyrio 60.7 Bechuanaland (M) 1 USNM 488868 Porphyrio porphyrio 64.8 New Caledonia (F) 64.6-65.1 USNM 561549, 561551 2 Porphyrio porphyrio 68.4 Captive (M) 66.3-70.5 USNM 555504, 557529 2 Porphyrio poliocephalus 63.2 Thailand (F) USNM 343212 1 Porphyrio poliocephalus 66.3 Captive (M) USNM 491890 1 Porphyrio pulverulentus 60.4 Philippines (M) 1 USNM 226035 Porphyrio martinica 60.6 Florida (4M, 4F) 58.4-62.7 USNM 501122-501126, 8 501128, 501130, 501131 to P. porphyrio but had smaller wings. In the Pacific, P. porphyrio occurs from the New Guinean region east through Mela- nesia to the western Polynesian islands of Tonga, Samoa, and Niue; it has not been recorded, either living or as a fossil, from eastern Polynesia. Thus P. paepae repre- sents an eastward range extension of 3200 km for the genus. Because Porphyrio once inhabited the Marquesas Islands, at the northeast corner of Polynesia, it is likely that fossils of these gallinules will be dis- covered elsewhere in eastern Polynesia. Acknowledgments For access to fossil specimens, I thank T. Han, C. Kishinami, B. Rolett, and Y. H. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sinoto. Comparative skeletons were made available by J. P. Angle, J. P. Dean, and S. L. Olson. Curatorial assistance was provid- ed by D. Pahlavan, S. E. Schubel, and M. C. Zarriello. B. Rolett kindly provided un- published data about his recent excavations at Hanamiai. The research was supported by National Science Foundation grant BSR- 8607535. N. G. Miller, S. L. Olson, D. Pah- lavan, and S. E. Schubel commented on the manuscript. This is contribution number 542 of the New York State Museum and Science Service. Literature Cited Balouet, J. C., & S. L. Olson. [In press]. Fossil birds from late Quaternary deposits in New Caledo- nia.—Smithsonian Contributions to Zoology. Baumel, J. J., A. S. King, A. M. Lukas, J. E. Breazile, & H. E. Evans (eds.). 1979. Nomina Anatom- ica Avium. Academic Press, London, 664 pp. Bellwood, P. 1972. A settlement pattern survey, Ha- natekua Valley, Hiva Oa, Marquesas Islands. — Pacific Anthropological Records 17:1-S0. . 1979. Man’s Conquest of the Pacific. Oxford University Press, New York, 462 pp. Kirch, P. V. 1986. Rethinking East Polynesian pre- history.—Journal of the Polynesian Society 95: 9-40. Olson, S. L. 1973. A classification of the Rallidae.— Wilson Bulletin 85:381-416. Sinoto, Y. S. 1979. The Marquesas. Pp. 110-134 in J. D. Jennings, ed., The prehistory of Polynesia. Harvard University Press, Cambridge, Massa- chusetts. Steadman, D. W. [In press]. Fossil birds and bio- geography in Polynesia.— Acta XIX Congressus Internationalis Ornithologicus. —, & M.C. Zarriello. 1987. Two new species of parrots (Aves: Psittacidae) from archeological sites in the Marquesas Islands. — Proceedings of the Biological Society of Washington 100(3):518- 528. Biological Survey, New York State Mu- seum, The State Education Department, Al- bany, New York 12230. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 171-175 NITOKRA SPHAEROMATA, A NEW HARPACTICOID COPEPOD CRUSTACEAN ASSOCIATED WITH THE WOOD-BORING ISOPOD, SPHAEROMA PERUVIANUM, IN COSTA RICA Thomas E. Bowman Abstract.— Nitokra sphaeromata is described from the pleopods of the marine flabelliferan isopod Sphaeroma peruvianum, collected from burrows in man- groves in the Gulf of Nicoya, Costa Rica. It is the fourth species of Nitokra, a genus with about 40 species, known to have commensal relationships. Other crustaceans associated with species of Sphaeroma, Iais spp. (Isopoda: Asellota), and Microsyssistria (Ostracoda: Entocytheridae) are listed. The commonly used spelling Nitocra is replaced by Boeck’s correct original spelling Nitokra. During an investigation of the effect of the wood-boring isopod, Sphaeroma peru- vianum Richardson, 1910, on red man- groves of Pacific Costa Rica by Richard C. Brusca and Diane Perry, a harpacticoid co- pepod was found on the pleopods of the isopod. Specimens were sent to me for iden- tification, and they proved to be the new species of Nitokra described below. Nitokra sphaeromata, new species Figs. 1-16 Material. —Costa Rica: Gulf of Nicoya, Punta Morales (ca. 10°04’N, 84°58'W), from pleopods of Sphaeroma peruvianum col- lected from its burrows in red mangroves: holotype 2 USNM 234096; paratypes 4 9, 2 6, 1 juv. USNM 234097. Leg. Richard C. Brusca and Diane Perry, Feb 1984. Etymology. —Sphaeromata = “of Sphae- roma,” the host genus. Description. —Lengths of 4 2, 1.22, 1.24, 1.25, 1.26 mm; of 2 6, 1.10, 1.16 mm. Pro- some (head + pedigers 1-4) subequal in length to urosome (pediger 5 thru caudal rami). Greatest body width at level of head. Rostrum oval, short. Urosomites 2—5 with submarginal row of spinules limited to lat- eral surface, except on anal somite where row is marginal and continues dorsally. Anal operculum with marginal spinules. Caudal ramus about as long as wide, without spi- nules; medial seta and inner of 2 lateral seta subequal in length; outer lateral seta half as long; dorsal seta reduced to short spine at base of inner terminal seta. Antenna | of 2 with row of setules on anterior margin of segment 1; in 6 with 2 serrate spines on segment 4 and 2 clublike setae on penultimate segment. Antenna 2, outer seta of exopod longer than others. Mandibular palp, segment 1 with seta on produced distomedial corner; segment 2 with 1 marginal and 5 terminal setae. Max- illa 1 inner lobe with 3 terminal spines and 1 seta on medial margin; middle lobe with 3 terminal setae; outer lobe with 5 terminal setae and 2 long setae on basal swelling. Maxilla 2 proximal endite produced into curved process with flaring tip armed with close-set setules; middle endite with 2 ter- minal setae; distal endite produced into claw with 2 slender basal setae and | basal seta with broadened tip having concave terminal margin. Maxilliped segment | with 3 setae on posterior margin and | at anterodistal corner. All rami of swimming legs 1—4 3-merous. Leg 1 exopod segments (Re 1-3) subequal 172 in length; Re 1-3 with 1-1-5 outer and ter- minal spines (Se and St) and O-1-0 inner setae (Si); endopod segments (Ri 1-3) with no Se, 1-1-0 Si, and 3 terminal setae. Ri 1 about 2 x as long as Ri 2 and Ri 3 combined and slightly longer than entire exopod. Inner spine on 2nd basipod modified to dumbbell shape in é. Re 1-3 of legs 2-4 with 1-1-3 Se, those of leg 2 enlarged, 1 St, and 0-1-3 Si. Ri 1- 3 of legs 2-4 with 1-1-3, 1-1-3, and 1-1-4 Si, and 1 St. Leg 5 of 2, proximal segment as long as broad, with 5 distal setae; next-to-outer- most seta slightly more than 2x length of segment, other setae subequal, as long as segment. Distal segment slightly longer than proximal segment, 2 x as long as wide, with 5 distal setae having proportionate lengths (medial to lateral) 24:20:10:18:28 = 100. Leg 5 of 6, proximal segment very short, with 4 distal setae; 3 medial setae about 3 Xx as long as segment, lateral seta about as long as segment. Distal segment about 2 x as long as wide, with 5 setae having proportionate lengths (medial to lateral) 14:13:38:5:30 = 100. Leg 6 of 6 a tiny lobe armed with 2 setae; medial seta nearly 2 x length of lateral seta. Comparisons.—In Wells (1976) Nitokra sphaeromata keys to N. mediterranea Brian (1928) and N. reunionensis Bozik (1969), from Italy and Réunion Island (western In- dian Ocean) respectively. These two species, however, are clearly distinct from N. sphae- romata. Nitokra mediterranea is much smaller, 0.4 mm in length. In legs 2-4 the endopod is shorter than the first two exopod segments PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON combined, whereas in N. sphaeromata the endopod of legs 2-4 is nearly as long as the exopod. In leg 1 the third exopod segment has four rather than five spines. In the @ leg 5 proximal segment the two lateral setae are subequal, and the distal segment has six rather than five setae. In the é leg 5 proximal segment the two lateral setae are subequal. Finally, the anal operculum of N. mediter- ranea 1s unarmed. Nitokra reunionensis is also smaller than N. sphaeromata, 2 ca. 0.75 mm, 30.70 mm. It has a dorsal spine row on the preanal segment, and the spines of the anal oper- culum reach beyond the posterior margin of the anal segment. The rami of leg 1 are subequal in length. The 2 leg 5 proximal segment has four rather than five setae. The 6 leg 5 proximal segment has two rather than four setae, and the distal segment has six rather than five setae. There are also im- portant differences in the armament of max- illa 1 and 2. From the Pacific coast of the Americas the only Nitokra’s known are the two new taxa described from Monterey Bay, Cali- fornia, by Lang (1965), N. spinipes armata and N. affinis f. californica, and N. lacustris columbianus Reid (1988) from Bahia So- lano, Colombia (6°17'N, 77°25'W). In both California forms the subterminal row of spines on the urosomites extends complete- ly around them, whereas in N. sphaeromata the row is confined to the lateral surface. Also in both the endopod of leg 4 reaches barely beyond the proximal margin of the second exopod segment, and the ¢ leg 6 has three rather than two setae. In Nitocra la- custris the exopod of leg | is about as long =) Figs. 1-16. Nitokra sphaeromata: 1, Female, lateral; 2, Female, dorsal; 3, Anal segment and caudal rami of female, dorsal; 4, Antenna 1, female, setae of distal segments omitted; 5, Antenna 1, male; 6, Antenna 2; 7, Mandible; 8, Maxilla 1; 9, Maxilla 2; 10, Maxilliped; 11, Leg 1, arrow points to endopod segment 3 in lateral view, dashed line leads to modified spine on 2nd basipod of male; 12, Leg 2; 13, Leg 3; 14, Leg 4; 15, Leg 5, female; 16, Legs 5 and 6, male. 174 as the endopod, whereas in N. sphaeromata the exopod is shorter than the Ist segment of the endopod. Host relationship.—Nothing is known about the relationship of Nitokra sphaero- mata with its host, Sphaeroma peruvianum, except that it occurs on the pleopods of the isopod. The copepods were discovered on preserved Sphaeroma’s long after the field work in Costa Rica had been completed. Other crustaceans are associated with species of Sphaeroma. The asellote isopod Tais californica (Richardson) occurs on Sphaeroma quoyanum Milne Edwards (=S. pentadon Richardson) (Menzies & Barnard 1951, Hurley 1956, Rotramel 1972), and on S. terebrans Bate, as I. singaporensis Men- zies & Barnard (Menzies & Barnard 1951, Pillai 1961). Jais pubescens (Dana) is as- sociated with S. walkeri Stebbing (Stebbing 1917, Barnard 1920, 1955) and S. annan- dalei Stebbing (Barnard 1955). The recently introduced population of S. walkeri in San Diego Bay, California, is not infested with Tais (Carlton & Iverson 1981). The collec- tions of the Division of Crustacea, Smith- sonian Institution, contain associated spec- imens of Sphaeroma peruvianum and Tais californica from the Gulf of Guayaquil, Ec- uador, donated in 1977 by Billy A. F. Ham- mond, then of the University of North Car- olina, Wilmington. Two species of the entocytherid ostracode genus Microsyssitria Hart, Nair, & Hart are associated with Sphaeroma terebrans: M. indica Hart, Nair, & Hart (1967) in Kerala State, India, and M. nhlabane Hart & Clark (1984) from Lake Nhlabane, South Africa. Nitokra sphaeromata is the fourth species of Nitokra known to have commensal re- lationships. Nitokra bdellurae lives in the egg capsules of Bdellura propinqua and B. candida, flatworms which live upon the car- apace of the horseshoe crab Limulus (Lid- dell 1912). Nitokra divaricata lives in the gill chambers of the crayfishes Astacus flu- viatilis (see Chappuis 1926) and A. Jlepto- dactylus (see Jakubisiak 1939). Nitokra me- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dusaea was found on the exumbrellar surface of an unidentified species of the scyphozoan genus Aurelia (Humes 1953). Nitokra or Nitocra? The correct spelling of the harpacticoid genus is Nitokra, Boeck’s (1865) original spelling. Later authors, except Brady & Robertson (1873) have spelled the genus Ni- tocra. The first to use the latter spelling ap- pears to have been Giesbrecht (1881), who did so without comment. It can be pre- sumed that Giesbrecht considered Nitocra the correct Latin form, the Greek letter kap- pa being transliterated as “‘c,”’ as given today in ICZN Appendix B. However, ICZN Ar- ticle 11(b)(v) states that the letter ““k”? may be used in scientific names, hence Nitokra Boek, 1865 is clearly available. Nitocra is an “incorrect subsequent spelling” and not available [ICZN Article 33(c)]. Nitokra can- not be suppressed as an unused senior syn- onym [ICZN Article 79(c)], since Nitocra is not available and therefore not a synonym. The choices available under the ICZN are two: (1) Return to the correct original spell- ing Nitokra, even though it has not been used for more than a century, and Nitocra, with about 40 species, has appeared in the zoological literature several hundred times. (2) Ask the Commission to use its plenary power to suppress Nitokra and validate Ni- tocra, in accordance with usage. I have elected the first choice because the matter seems too trivial to submit to the Commission. Although well known among copepodologists, Nitocra is scarcely a fa- miliar name among other zoologists. I am not replacing a long-standing name but re- turning to the original spelling, following the ICZN. Acknowledgments I am grateful to Richard C. Brusca for sending me specimens of the new harpac- ticoid, to Frederick M. Bayer for a discus- sion of the correct spelling of the genus, and VOLUME 101, NUMBER 1 to Janet W. Reid for reviewing the manu- script. Literature Cited Barnard, K.H. 1920. Contributions to the crustacean fauna of South Africa. 6. Further additions to the list of the marine Isopoda.— Annals of the South African Museum 17:319-438. 1955. Additions to the fauna-list of South African Crustacea and Pycnogonida.— Annals of the South African Museum 43:1-107. Boeck, A. 1865. Oversigt over de ved Norges Kyster iagttagne Copepoder henhgrende til Calani- dernes, Cyclopidernes og Harpactidernes Fam- ilier.— Forhandlinger i Videnskabs-Selskabet i Christiana, Aar 1864:226-282. Bozik, B. 1969. Copépodes Harpacticoides de la Ré- union.— Bulletin du Muséum National d’His- toire Naturelle, Paris (2)41(4):867-882. Brady, G. S., & D. Robertson. 1873. Contributions to the study of the Entomostraca. VIII. On ma- rine Copepoda taken in the west of Ireland. — Annals and Magazine of Natural History (4) 12: 126-142. Brian, A. 1928. Ricerche faunistiche nella isole ital- iane dell’Egeo. I copepodi bentonici marini.— Archivio Zoologico Italiano 12(3—4):293-343. Carlton, J. T., & E. W. Iverson. 1981. Biogeography and natural history of Sphaeroma walkeri Steb- bing (Crustacea: Isopoda) and its introduction to San Diego Bay, California.—Journal of Nat- ural History 15:31-48. Chappuis, P. A. 1926. Harpacticiden aus der Kie- menhohle de Flusskrebses. — Archiv fur Hydro- biologie 17(3):515-520. Giesbrecht, W. 1881. Vorlaufige Mittheilung aus ein- er Arbeit tiber die freilebenden Copepoden des Kieler Hafens.— Zoologischer Anzeiger 4:254— 258. Hart, C. W., Jr., & J. Clark. 1984. A new commensal ostracod of the genus Microsyssitria from South Africa (Ostracoda: Entocytheridae: Microsys- sitrinae). — Proceedings of the Biological Society of Washington 97(1):217—220. —., N. Balakrishnan Nair, & D. G. Hart. 1967. A new ostracod (Ostracoda: Entocytheridae) commensal on a wood-boring marine isopod from India.—Notulae Naturae, Academy of Natural Sciences of Philadelphia 405:1-11. 175 Humes, A. G. 1953. Two new semiparasitic harpac- _ticoid copepods from the coast of New Hamp- shire.—Journal of the Washington Academy of Sciences 43(11):360-373. Hurley, D. E. 1956. The New Zealand species of Jais (Crustacea Isopoda).— Transactions of the Roy- al Society of New Zealand 83(4):715-719. Jakubisiak, S. 1939. Nitocrella divaricata (Chappuis), komensal raka.—Archiwum Hydrobiologii i Rybactwa 12:117-121. Lang, K. 1965. Copepoda Harpacticoidea from the Californian Pacific Coast.—Kungliga Svenska Vetenskapsakademiens Handlingar 10(2):1—560. Liddell, J. A. 1912. Nitocrameira bdellurae, nov. gen. et sp., a copepod of the family Canthocampti- dae, parasitic in the egg cases of Bdellura.— Journal of the Linnean Society of London, Zo- ology 32:87-94. Menzies, R. J., & J. L. Barnard. 1951. The isopodan genus Jais (Crustacea).— Bulletin of the South- ern California Academy of Sciences 50(3):136- 151. Pillai, N. K. 1961. Wood-boring Crustacea of India. Government of India Press, Simla, viii + 61 pp. Reid, J. W. 1988. Cyclopoid and harpacticoid co- pepods (Crustacea from Mexico, Guatemala and Colombia).— Transactions of the American Mi- croscopical Society 107(2). Richardson, H. 1910. Report on isopods from Peru, collected by Dr. R. E. Coker.— Proceedings of the United States National Museum 38(1729): 79-85. Rotramel, G. 1972. Jais californica and Sphaeroma quoyanum, two symbiotic isopods introduced to California (Isopoda, Janiridae and Sphaero- matidae). —Crustaceana, supplement 3:193-197. Stebbing, T.R.R. 1917. The Malacostraca of Durban Bay.—Annals of the Durban Museum 1:435- 450. Wells, J. B. J. 1976. Keys to aid in the identification of marine harpacticoid copepods., Aberdeen University Press, Aberdeen, U. K. 215 pp. Department of Invertebrate Zoology, Na- tional Museum of Natural History, NHB- 163, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 176-182 POLYCHAETOUS ANNELIDS FROM THE INTERTIDAL ROCKY SUBSTRATUM OF A POLLUTED AREA OF THE RIA DE PONTEVEDRA (GALICIA, SPAIN) 2. TAXONOMIC ASPECTS WITH THE DESCRIPTION OF LUGIA ATLANTICA, N. SP. Antonio Villalba and José M. Viéitez Abstract. —Some taxonomic aspects derived from a study of the taxocoenosis of polychaetes inhabiting the intertidal zone of an area in the ria de Pontevedra (NW of Spain), affected by industrial wastes, are presented. A new species, Lugia atlantica (Phyllodocidae), is described, and detailed descriptions of spec- imens of the genus Micronereis Claparéde (Nereididae) and family Capitellidae with dubious taxonomic positions, are offered. In a previous report (Villalba & Viéitez 1985), the taxocoenosis of polychaetous an- nelids inhabiting the intertidal zone of an area in the ria de Pontevedra was studied, and the effects of industrial wastes from Kraftpaper and chlorine-soda factories on these components of the benthic fauna were discussed. Among the sampled material, some specimens with interesting taxonomic implications were found. These are de- scribed and discussed below. Three sampling stations were established on the southern side of the estuary (Fig. 1). The studied area, sampling stations, mate- rial and methods are described elsewhere (Villalba & Viéitez 1985). Family Phyllodocidae Williams, 1851 Genus Lugia Quatrefages, 1865 Lugia atlantica, new species Fig. 2 Lugia sp. Villalba & Viéitez, 1985:376. Phyllodocidae ind. Villalba & Viéitez, 1985: 373-376. Material examined.—Sta M (42°24'N, 05°00’W), lowest intertidal band, holotype and paratype. Sta A, intertidal band of Ulva rigida and Fucus vesiculosus, paratype. Sta B, intertidal band of U. rigida and F. vesi- culosus, paratype. All specimens deposited in the collection of the Zoology Department of the University of Alcala de Henares, Ma- drid. Description of holotype. —Elongate body, tapering anteriorly and posteriorly, with constriction on segment 10, at level of end of inverted proboscis. Segments 25, 1.8 mm long and 0.4 mm wide. Prostomium round, slightly bilobed, about as wide as long; nar- rower anterior end with 4 cirriform anten- nae of equal length, slightly shorter than prostomium (Fig. 2A). Without median an- tennae or nuchal papilla. One pair of lensed eyes on posterior part of prostomium. Pro- boscis inverted; when observed through body wall it consisted of wider region with papillae and perhaps narrower bare region. Segment | dorsally distinct from prosto- mium, with pair of tentacular cirri tapering gradually distally, longer than antennae. Segment 2 with pair of similar dorsal ten- tacular cirri but twice as long; setal lobes with aciculum and setae; pair of ventral cirri shorter than dorsal tentacular ones, but longer than following ventral cirri; similar in shape and size to remaining dorsal cirri. 1 N T lar fe lag il se S—= a S= Che, entacular formula S N SK { ig 2A). VOLUME 101, NUMBER 1 Parapodia uniramous; from segment 3 consisting of lanceolate dorsal cirrus, seti- gerous lobe with aciculum and setae, and shorter, oval ventral cirrus. Setal lobes with 2 rounded presetal lobes of equal size. Acic- ulum with blunt tip and very slender exten- sion protruding from setal lobe (Fig. 2D). Setae of 2 kinds: compound setae variable in number per parapodium, from 3 setae in 2nd segment to 9 in middle region of body. Compound setae consisting of smooth shaft with swollen joint bearing bundle of very small spines and 2 larger unequal spines; blades minutely serrated, with obliquely striated surface (Fig. 2B); simple setae, sin- gle one in parapodia of segments 20 to 23, with very finely serrated distal tip (Fig. 2C). Parapodia of posterior 2 segments very re- duced, without setae, perhaps area of active growth. Anal cirri lacking. Color in formalin light brown with darker spots. Paratypes.—Segments from 25 to 28. One specimen with proboscis slightly everted, showing portion covered with fleshy pa- pillae. Number of posterior segments bear- ing simple setae from 2 to 6. All of paratypes with 2 lanceolate anal cirri, slightly longer than dorsal cirri of middle region. Remarks. —Quatrefages (1866:152) in- cluded Eteone aurantiaca Schmarda, 1861, and E. pterophora Ehlers, 1864, in his new genus Lugia. Bergstrom (1914:184) selected Ehlers’s species as the type species and in- cluded a diagnosis for the genus. Bergstrom (1914:201) also retained Schmarda’s species in Eteone. In her Catalogue, Hartman (1959: 154) incorrectly listed Schmarda’s species as the type species of Lugia; this was fol- lowed by Fauchald (1977:49). Uschakov (1958:204) added Lugia rarica from the Kurile-Kamchatka trench but later (1972:121, 1974:116), referred it to Mys- tides rarica, based on the lack of dorsal cirri on segment 3. Uschakov (1972:116, 1974:113) added Lugia abyssicola from abyssal bottoms off Japan and the Pacific Ocean off California. N77 Fig. 1. the Peninsula of Placeres, Pl (with two sampling sta- tions) and Mogor, M (the third sampling station); (2) Detail of the Peninsula of Placeres showing the location of two sampling stations, A and B, and the industrial complex, F. The location of the sewer is marked by the arrow. (Dique = Dike). (1) Map of the ria de Pontevedra showing Campoy & Alquézar (1982:123) and Alos, Campoy & Pereira (1982:143) added Lugia incognita from the Iberian Peninsula. Lugia atlantica differs from L. ptero- phora, which is characterized by setae with smooth blades (without serrations and stria- tions), lacking simple setae, and by the pres- ence of a very elongate prostomium (Ehlers 1864:pl. 6, figs. 22-25; Fauvel 1923:fig. 631). There are ecological and morphological differences between L. atlantica and L. abyssicola, the later inhabiting the abyssal zone and lacking eyes and simple setae (Us- chakov 1972, 1974:figs. 1-5). Lugia atlantica differs from L. incognita, the latter having much shorter and rounded tentacular cirri with rounded tips, dorsal cirri, lacking simple setae and with fewer compound setae per parapodium (Campoy & Alquézar 1982:fig. 2A—F). Etymology.—The name atlantica is pro- 178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Lugia atlantica holotype. A, Dorsal view of anterior region; B, Compound setae; C, Simple setae from posterior parapodium; D, Aciculum. VOLUME 101, NUMBER 1 179 lhe 1444 Fig. 3. Micronereis sp. A, Pharynx of male with jaws and buccal cavity (Cb); B, Compound homogomph setae; C, Shaft of compound setae showing articulation. Family Nereididae Johnston, 1845 posed because this is the first time speci- : Genus Micronereis Claparéde, 1863 mens of the genus Lugia have been found in the Atlantic Ocean. The others have been Micronereis sp. described from the Mediterranean Sea (L. Figs. 3, 4A. pterophora, L. incognita) and the Sea of Ja- Material examined. —Sta M. Lowest in- pan and the Pacific Ocean (L. abyssicola). _ tertidal band, 1 male specimen. 180 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. A, Micronereis sp. Dorsal view of anterior end showing jaws, one of right tentacular cirri detached by manipulation; B, Capitellidae indet., thoracic region showing genital hook between setigers 9 and 10 (arrow); C, Capitella capitata (?), fragment of abdominal region with bud-like expansion; D, Capitella capitata (?), abdominal fragment, enlarged. Description. — Body flat, 2 mm long and 0.6 mm wide excluding setae, with 16 se- tigerous segments. Prostomium broad, slightly bilobed, distinct from peristomium, with 2 pairs of eyes in trapezoidal arrange- ment, anterior pair slightly larger and lat- erally located along maximum width, pos- terior pair located in posterior portion of prostomium (Fig. 4A). Paired lobes located anteroventrally, called palps after Paxton (1983). Tentacular cirri 4 pairs with broad bases, gradually tapering distally. Anterior 2 pairs lateral to prostomium, of similar length; posterior 2 pairs slightly longer, lat- eral to peristomium, one pair anterior and ventral, other pair posterior and dorsal (Fig. 4A). Pharynx with 2 yellow jaws extending from peristomium to setiger 2. Jaws of male consisting of 2 regions: base with 3 teeth, broadest of them with 8 denticles. Base con- tinuing distally into slender prolongation ending in 2 round expansions and sharp tip; buccal cavity with circle of many rows of very small paragnaths (Fig. 3A). Setigers 1 and 2 with uniramous para- podia, consisting of setigerous lobe, acicu- lum, fascicle of setae, and ventral cirrus; dorsal cirrus lacking. Parapodia of following setigers biramous, with divergent rami. VOLUME 101, NUMBER 1 Dorsal and ventral cirri similar in length. Acicula sharp-pointed with straight tips. Compound setae with homogomph articu- lations, with joint slightly crenulate; smooth shafts without transverse septa and long blades slightly serrate, sharp-pointed. Each neuropodium of setiger 3 with pair of cop- ulatory hooks. Anal cirri lacking. Remarks. —The taxonomic status of this genus has been studied and changed in dif- ferent reviews (Reish 1961, Banse 1977, Buzhinskaya 1981, Paxton 1983). Our spec- imen cannot be included with conviction in any of the valid species that Paxton (1983) considered. The specimen differs from M7- cronereis nanaimoensis Berkeley & Berke- ley, 1953, M. halei Hartman, 1954, M. eni- wetokensis Reish, 1961, and M. bansei (Hartmann-Schroder 1979) because all of them have shortened type of jaws. Our spec- imen also differs from M. ochotensis Buz- hinskaya, 1981, with different kind of jaws, shafts of compound setae with septa and blades with much more conspicuous den- ticles. It differs from M. minuta (Knox & Cameron 1970), with accessory parapodial cirri and simple setae, lacking copulatory hooks, and blades of the compound setae very coarsely serrated. It differs from M. piccola Paxton, 1983, in having ocular spots on the segments, compound falcigers, and shafts with septa. Our specimen does not completely agree with M. variegata Claparéde, 1863. The main difference resides in the shape of the jaws, this character being the most useful for the diagnosis of this species (Amoureux et al. 1978). Paxton (1983) could not ex- amine the holotype of the species. Based on specimens that she considered similar to the original, however, she offered a drawing of the jaws of a male, which differ from our specimen. Other characters established by this author, those related to number of se- tigers, length, and setae, are similar to our specimen. In spite of these differences, we consider that one specimen is not enough to establish a new taxon. 181 _ Family Capitellidae Grube, 1862 Capitellidae indet. Fig. 4B Material examined.—Sta A, intertidal band of Ulva rigida, complete specimen and one without abdominal region. Description. —The external appearance agrees with Capitella capitata (Fabricius 1780), except for copulatory apparatus lo- cated between setigers 9 and 10, instead of 8 and 9 (Fig. 4B). Setigers 1 to 6 with cap- illary setae exclusively. Parapodial rami of segment 7 with hooded hooks, except right ventral ramus with capillary setae. Setiger 8 with hooded hooks exclusively. Setiger 9 and 10 with hooded hooks in ventral rami and genital hooks in dorsal. Remarks. —We have not found in the lit- erature any genus of this family with the copulatory apparatus between setigers 9 and 10. These specimens could lead to the es- tablishment of a new genus, but we prefer to wait for new findings of specimens with this character, especially from unpolluted areas. Genus Capitella Blainville, 1828 Capitella capitata (Fabricius 1780) Fig. 4C, D Material examined.—Sta A, intertidal band of Ulva rigida, fragment of the abdom- inal region of a specimen belonging very likely to Capitella capitata. It bears a lateral segmented bud-like expansion with hooded hooks in every segment, showing the typical arrangement for the species. Remarks.—We have not found any ref- erence in the literature pointing out the pos- sibility of budding in this family. Using high levels of heavy metals in culture, Reish (1974, 1977) obtained larvae of this species with bifurcated abdomens. These abnormal specimens, however, never passed the 8-segmented metatrochophore stage. The fragment we are reporting belongs to an adult specimen. We do not discount the idea that the bud-like expansion may be a kind of 182 anomaly caused by the industrial pollution of the area. Acknowledgments We are grateful to Dr. G. Bellan (Station Marine d’Endoume, Marseille) for his com- ments throughout the preparation of this paper. Literature Cited Alos, C., A. Campoy, & F. Pereira. 1982. Contri- bucion al estudio de los anélidos poliquetos en- dobiontes de esponjas.— Actas del II Simposio Ibérico de Estudios del Bentos Marino 3:139- 157. Amoureux, L., F. Rullier, & L. Fishelson. 1978. Sys- tematique et ecologie d’annelides polychétes de la presqu’il du Sinai. — Israel Journal of Zoology 27:57-163. Banse, K. 1977. Anew subfamily, Notophycinae (Po- lychaeta: Nereididae), for Micronereis Cla- paréde and Quadricirra new genus. Pp. 115- 140 in D. J. Reish, and K. Fauchald, eds., Es- says on polychaetous annelids in memory of Dr. Olga Hartman. Allan Hancock Foundation, University of Southern California, Los Angeles. Bergstrom, E. 1914. Zur Systematik der Polychae- tenfamilie der Phyllodociden. — Zoologiska Bid- rag fran Uppsala 3:37-224. Buzhinskaya, G. N. 1981. On taxonomic status of the genus Micronereis Claparéde, 1863 (Poly- chaeta, Nereidae). — Zoologischesky Zhurnal 60: 1256-1261. [In Russian, English summary]. Campoy, A., & E. Alquézar. 1982. Anélidos Poli- quetos de las formaciones de Dendropoma pe- traeum (Monterosato) de las costas del Sureste de Espana.— Actas del II Simposio Ibérico de Estudios del Bentos Marino 3:121—137. Ehlers, E. 1864. Die Borstenwiirmer (Annelida Chae- topoda), vol. 1. Wilhelm Engelman, Leipzig, 268 pp. Fauchald, K. 1977. The Polychaete worms. Defini- tions and keys to the orders, families and gen- era.— Natural History Museum of Los Angeles County Science Series 28:1-188. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fauvel, P. 1923. Polychétes Errantes.—Faune de France 5:1-488. Hartman, O. 1959. Catalogue of the polychaetous annelids of the world.— Allan Hancock Foun- dation Publications, Occasional Paper 23 (1)2: 1-628. Paxton, H. 1983. Revision of the genus Micronereis (Polychaeta: Nereididae: Notophycinae).—Rec- ords of the Australian Museum 35(1):1-18. Quatrefages, A. de. 1866 (1865). Histoire naturelle des Annelés marins et d’eau douce. Annélides et Gephyriens.— Librairie Encyclopédique de Roret, Paris 2(1):1-336. Reish, D. J. 1961. A new species of Micronereis (An- nelida, Polychaeta) from the Marshall Islands.— Pacific Science 15:273-277. . 1974. Induction of abnormal polychaete lar- vae by heavy metals.—Marine Pollution Bul- letin 5(8):125-126. . 1977. Effects of chromium on the life history of Capitella capitata (Annelida: Polychaeta). Pp. 199-207 in F. J. Vernberg et al., eds., Physio- logical responses of Marine Biota to pollutants. Academic Press, New York. Uschakov, P. V. 1958. Two new species of poly- chaetes. belonging to the family PhyHodocidae from the abysses of the Kurile-Kamchatka Trench.— Trudy Instituta Okeanologii 27:204— 207. [In Russian] . 1972. Polychaetes. Pp. 1-271 in Fauna of the U.S.S.R. Academy of Sciences of the U.S.S.R., Zoological Institute, I. [In Russian, English translation, Israel Program for Scientific Trans- lations, Jerusalem, 1974:1—259.} Villalba, A., & J. M. Viéitez. 1985. Estudio de la fauna de anélidos poliquetos del sustrato rocoso intermareal de una zona contaminada de la ria de Pontevedra (Galicia). I. Resultados bioce- noticos.—Cahiers de Biologie Marine 26:359- 377. Departamento de Biologia Animal, Uni- versidad de Alcala de Henares, 28071 Al- cala de Henares, Madrid, Spain. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 183-191 THOONCHUS LONGISETOSUS AND OX YONCHUS STRIATUS, NEW SPECIES OF FREE-LIVING MARINE NEMATODES (NEMATODA: ENOPLIDA) FROM NORTHWEST FLORIDA, U.S.A. Edwin J. Keppner Abstract. — Two new species of free-living marine nematodes from the genera Thoonchus and Oxyonchus (Nematoda: Enoplida) are described from sediments off the Gulf of Mexico and St. Andrew Bay, Bay County, Florida, U.S.A. Thoonchus longisetosus is unique in the presence of distinct labial setae rather than labial papillae, in the size and shape of the amphid, the position of the excretory pore, and in the shape of the gubernaculum. Oxyonchus striatus is unique in the presence of distinct transverse striations of the cuticle and in the size and shape of the spicules and gubernaculum. Keys are provided to the species of each genus. The two new species described here were recovered from sediment in a bayou and lagoon off St. Andrew Bay, and from sedi- ment adjacent to a rock outcrop in the Gulf of Mexico, Bay County, Florida. Sediment samples were obtained from the bay sites with a 4.5 cm diameter corer to a depth of 10 cm in the sediment. Sediment samples were obtained from the Gulf of Mexico with SCUBA equipment, and a 3.5 cm diameter corer to a depth of 5-10 cm. Nematodes were extracted by repeated washing of the sediment in seawater, allow- ing the heavier particles to settle for 15 sec and decanting the supernatant water and suspended material to a second container. The suspended material from four washings was allowed to settle for 15—20 min, and the supernatant water was decanted. Nema- todes were removed alive from the remain- ing sediment, fixed in hot alcohol-formalin- acetic acid for 24 h, cleared in glycerine, and mounted in anhydrous glycerine on Cobb slides. Measurements were made with a cali- brated ocular micrometer. All measure- ments are given in wm unless othewise in- dicated, and the mean is followed by the range in parentheses. The taxonomic hier- archy is that of Gerlach & Riemann (1974). Enchelidiidae Filipjev, 1918 Thoonchinae Gerlach & Riemann, 1974 Thoonchus Cobb, 1920 Thoonchus longisetosus, new species Figs. 1-5, 10, 11 Diagnosis. —One male from Gulf of Mex- ico. Body slender; cuticle thin, smooth. Head with circle of 6 long slender labial setae and single circle of 10 (6 + 4) cephalic setae, 4 submedian setae about *% length of other 6. Cervical setae long, slender; present from immediately posterior to amphid to level of nerve ring. Somatic setae not observed; cau- dal setae present. Stoma large with heavily cuticularized walls and 3 teeth, right sub- ventral tooth largest; size and position of left subventral and dorsal teeth about equal. About 6 complete irregular circles of small round denticles present anterior to dorsal and left subventral teeth. Amphid posterior to base of stoma; wide with thickened lateral margins. Excretory pore posterior to base of 184 stoma. Esophagus long, muscular; not ex- panded at junction with intestine. Tail con- ical, then cylindrical. Spinneret and caudal glands present; caudal gland cell bodies ex- tend anterior to cloaca. Female unknown. Male (n = 1): Length 2.62 mm, width at midbody 32. Head diameter 21 at level of cephalic setae. Labial setae 7 long, cephalic setae 12 and 8 long. Stoma 21 long, 19 wide at level of denticles. Amphid 18 wide. Esophagus 778 long; excretory pore 30 and nerve ring 202 from anterior end. Tail 86 long, 33 wide at cloaca. Two midventral areas of thickened cuticle 102 and 150 an- terior to cloaca; each with central depres- sion and small central sensilla. One pair small, subventral papillae present 78 ante- rior to cloaca. Single midventral, cuticular elevation with pore 22 anterior to cloaca. Two pairs subventral setae immediately an- terior to cloaca. Spicules 35 long, arcuate with curved tip. Corpus of gubernaculum surrounding spicules; gubernacular apoph- ysis 8 long, dorsally directed. a = 81.9; b = 3.37; c = 30.5. Type specimen. — Holotype male, USNM VHS. Type locality.—Sediments consisting of coarse sand, shell, and sponge spicules in Gulf of Mexico adjacent to exposed rock outcrop Bay County, Florida (30°12'45’N, 86°02'10'’W). Etymology.—The specific name is from the Latin /ongus for long and seta for bristle in reference to the long labial setae of this species. Remarks.—The genus Thoonchus con- tains three species, 7. ferox Cobb, 1920, T. inermis Gerlach, 1953, and 7. giganticus Belogurova, Belogurov & Khamula, 1986. Cobb (1920) stated that 7. ferox has labial papillae and the amphids are faint. He fig- ured the amphids as small and located over the base of the stoma. Males of T. ferox have a single elevated precloacal supplementary organ and a pair of subventral papillae im- mediately anterior to the cloaca. Spicules are arcuate, the corpus of the gubernaculum PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON does not surround the spicules, and the gu- bernaculum has a thin proximal extension parallel to the spicules. Chitwood (1960) described 7. ferox as having labial papillae, an excretory pore an- terior to the cephalic setae, and amphids located over the base of the stoma. Males have at least two subventral precloacal pap- illoid supplements and the spicules and gu- bernaculum are as described by Cobb (1920). Gerlach (1953) described the male, and Gerlach (1954) described the female of T. inermis. Thoonchus inermis has labial pa- pillae, the excretory pore is at the base of the stoma, amphids are located over the middle of the stoma, and the right subven- tral tooth is largest. Males do not have pre- cloacal supplements or papillae and the gu- bernaculum has a pointed apophysis. Thoonchus longisetosus can be distin- guished from 7. ferox by the presence of long labial setae rather than labial papillae, the amphid is posterior to the stoma and is large with thickened lateral margins rather than thin-walled and over the stoma, and the excretory pore is posterior to the stoma rather than anterior to the cephalic setae. The male genital apparatus of 7. /ongise- tosus differs from that of T. ferox in that the gubernaculum surrounds the spicules and the gubernaculum has a broad apophysis rather than a thin proximal extension par- allel to the spicules. Thoonchus longisetosus can be distin- guished from 7. inermis by the presence of long labial setae rather than papillae, the amphids are large and posterior to the sto- ma rather than over the middle of the sto- ma, and the excretory pore is posterior to the stoma rather than at the base of the stoma. The male genital apparatus of T. /on- gisetosus differs from that of 7. inermis in the presence of precloacal supplements and papillae, in the presence of a gubernaculum that surrounds the spicules, and the guber- naculum has a broad blunt apophysis rather than a pointed apophysis. The three species of Thoonchus discussed VOLUME 101, NUMBER 1 185 Figs. 1-5. Thoonchus longisetosus: 1, Male, head, right lateral view; 2, Male, head, left lateral view; 3, Male, posterior end, right lateral view; 4, Male, right spicule and gubernaculum, lateral view; 5, Male, precloacal supplements, right lateral view (anterior, top; posterior, bottom). Scales in um. 186 above are quite similar in the presence of the oncholaimoid structure of the stoma. The stoma is 1.1—1.5 times as long as broad and the teeth are short, broad, and originate from a wide base at the posterior end of the stoma. The spicules are short and arcuate. The gubernaculum has an expanded corpus and an apophysis. Cobb (1920) and Chit- wood (1960) emphasized the oncholaimoid shape of the stoma as characteristic of the genus Thoonchus. Wieser (1953) stated in reference to the genera of Thoonchinae (Thoonchus and Ditlevsenella Filipjev, 1925) “buccal cavity oncholaimoid i.e. the two subventral teeth strongly developed and with a broad base.” Belogurova et al. (1986) described a male nematode from the Tatar Strait as Thoon- chus giganticus. They placed this species in Thoonchus based on the presence of three teeth in the stoma and the absence of sup- plements in the male. Examination of the drawings of 7. giganticus reveals that the shape of the stoma and teeth differ signifi- cantly from the oncholaimoid type of stoma and teeth of the other species in the genus. The stoma is long and narrow (2.5 times as long as broad). The subventral teeth are fig- ured as almost equal, narrow, and extend from a narrow base at the posterior end of the stoma to almost the anterior end of the stoma. The dorsal tooth is smaller and slightly posterior to the anterior terminus of the subventrals. The spicules of 7. gi- ganticus are elongate (chord = 558; arc = 360) and thin. The gubernaculum is a thin plate without apophysis or extension. The precloacal organs are papilloid. The shape of the stoma, teeth, and male genital apparatus deviate significantly from that of Thoonchus. The shape of the these parts are more similar to those of certain genera in the Enchelidiinae rather than Thoonchinae. Based on the information available, it is not prudent to emend the genus Thoonchus to include this species. It is also difficult to place this species within the existing genera in the Enchelidiinae or PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON erect a new genus for it. Therefore, Thoon- chus giganticus Belogurova, Belogurov, & Khamula, 1986, is considered of doubtful generic assignment. The following key differentiates the three species of Thoonchus. Ile Labial sensillae papilliform; am- phid over stoma; excretory pore at level of base of stoma or an- terior to cephalic setae - Labial sensillae setiform; amphid posterior to stoma; excretory pore posterior to stoma Bite AONE RO as T. longisetosus n. sp. Excretory pore anterior to ce- phalic setae; male gubernaculum with thin proximal extension parallel to spicules; precloacal supplements and papillae present en he ee T. ferox Cobb, 1920 — Excretory pore at base of stoma; male gubernaculum with pointed apophysis; precloacal supple- ments and papillae absent Se TE © tetee T. inermis Gerlach, 1953 2(1). Enoplidae Dujardin, 1845 Oxyonchinae De Coninck, 1965 Oxyonchus Filipjev, 1927 Oxyonchus striatus, new species Figs. 6-10, 13, 14 Diagnosis. —One male, one female, and one juvenile. Body long, slender. Cuticle thick with distinct transverse striations be- ginning at base of head and extending length of body. Head with 3 low lips, 6 labial setae, and single circle of 10 (6 + 4) cephalic setae; 4 submedian setae about 45% length of oth- er 6 in male, 36% of other 6 in female. Ce- phalic organ not observed. Cephalic capsule thin, distinctly light refractive. Cervical se- tae of variable length, numerous to level of nerve ring; longest setae in dorsal, ventral, and lateral rows. Somatic setae short, sparse; caudal setae variable in length, numerous. Stoma with 2 large subventral teeth, left VOLUME 101, NUMBER 1 subventral longer, reaching just anterior to anterior margin of mandibles; dorsal tooth not observed. Mandibles complex, well-de- veloped; each with two thick arched rods converging medially. Each rod with small tooth (denticle) at base of transverse bar. Rods connected by broad transverse bar with claws. A series of refractile rods or tubules extending radially from oral surface of each mandible, terminating at about posterior margin of transverse bar. Amphid near base of head, oblong with thickened anterior margin. Esophagus vesiculate-crenulate; ex- cretory pore not observed. Tail cylindrical for first two-thirds, then narrower; tip slightly expanded. Caudal glands and spin- neret present. Male (n = 1): Length 3.41 mm, width at midbody 43. Head 38 long, 34 wide at level of cephalic setae. Labial setae 6 long; ce- phalic setae 33 and 15 long. Amphid 7 wide. Subventral teeth 26 and 23 long. Esophagus 590 long; nerve ring 135 from anterior end. Tail 115 long, 40 wide at cloaca. Single, midventral, tubular supplement 22 long, lo- cated 74 anterior to cloaca. One pair sub- ventral, papillae 61 posterior to cloaca. Spicules 140 long, capitulum funnel-shaped then narrowed, then expanded and gradu- ally narrowed to flattened tip. Gubernacu- lum bipartite, each lateral piece a narrow tube 28 long, apophysis absent. a = 79.3; b = 5.78; c = 29.7. Female (n = 1): Length 4.44 mm, width 54 at midbody. Head 37 long, 37 wide at level of cephalic setae. Cephalic setae 30 and 11 long. Amphid 7 wide. Subventral teeth 30 and 28 long. Esophagus 683 long; nerve ring 150 from anterior end. Tail 112 long, 45 wide at anus. Reproductive system di- delphic, amphidelphic, ovaries reflexed. Vulva 2.74 mm from anterior end. Two eggs present, 120 and 144 long. a = 82.2; b = 6.50; c = 39.6; V = 62%. Type specimens. — Holotype male, USNM 77134; allotype female, USNM 77135; paratype juvenile, USNM 77136. Type locality. — Holotype male from non- 187 vegetated sediment consisting of fine sand and detritus from mouth of Freshwater Bay- ou off St. Andrew Bay, Bay County, Florida (85°39'00"W, 30°07'30’N). Allotype female from nonvegetated sediment consisting of fine sand and detritus from Grand Lagoon off St. Andrew Bay, Bay County, Florida (85°43'15”W, 30°07'50’'N). Etymology. — From the Latin stria for fur- row in reference to the transverse striations in the cuticle of this species. Remarks. —The genus Oxyonchus is characterized by the presence of two large subventral teeth in the stoma that extend to the anterior end of the mandibles. The dor- sal tooth is reduced or absent. Mandibles are arch-shaped, well developed, and den- ticles can be present on the inner surface of the mandibular plate. Until now all species for which males are known have arcuate, relatively short spicules (1-1.9 cloacal di- ameters). De Coninck & Stekhoven (1933) provid- ed a key to the species of Oxyonchus and considered O. stateni Allgen, 1930 as in- sufficiently characterized and possibly a synonym of O. australis (De Man, 1904) Filipjev, 1927. Wieser (1953) revised the genus Oxyonchus and provided a key to the species. He considered O. statenia synonym of O. australis. Species added to the genus Oxyonchus since 1953 are: O. subantarcti- cus Mawson, 1958: O. culcitatus Wieser, 1959; and O. ditlevseni Inglis, 1964. Allgen (1959) described five new species of Oxyonchus; O. brachysetosus, O. crassi- colis, O. macrodon, O. notodentatus, and O. parastateni. He disagreed with the position of Wieser (1953) regarding the synonymy of O. stateni and O. australis but did not provide additional figures or information to support the validity of O. stateni. The de- scription, figures, and absence of figures for the new species of Oxyonchus provided by Allgen (1959) are insufficiently adequate to distinguish the species from others and to place these species in a key. Therefore, O. brachysetosus, O. crassicolis, O. macrodon, 188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 6-10. Oxyonchus striatus: 6, Male, head, left lateral view; 7, Male, precloacal supplement and guber- naculum, right lateral view; 8, Male, posterior end, right lateral view; 9, Female, head, left sublateral view; 10, Female tail, lateral view. Scales in um. VOLUME 101, NUMBER 1 Figs. 11-14. Figs. 11-12. Thoonchus longisetosus: 11, Male, head, lateral view; 12, Male posterior end, lateral view; Scale bars = 10 um. Figs. 13-14. Oxyonchus striatus: 13, Male, head, lateral view; Scale bar = 10 um; 14, Male, posterior end, lateral view; Scale bar = 20 um. 190 O. notodentatus, and O. parastateni are con- sidered species dubia. Oxyonchus striatus, n. sp. differs from all other species of the genus in the presence of the distinctly striated cuticle, the length and shape of the spicules, and the shape of the gubernaculum. The following key to the species of Oxy- onchus is to a great extent based on the key developed by Wieser (1953). Wieser (1953) stated in the introductory remarks to his key that “‘as to the species described there are some problems which can not be cleared up from the records in the literature.’ The statement remains applicable, and some of the species included in the genus do not appear to belong in Oxyonchus. An exam- ination of the type specimens for each species would be required in order to complete a thorough analysis of the genus. Key to Species of Oxyonchus Filipjev, 1927 ils Cuticle with transverse stria- tions; spicules long, slender, not arcuate, 3.5 cloacal diameters long; gubernaculum without apophysis O. striatus, n. sp. - Cuticle without transverse stria- tions; spicules arcuate, broad, 1- 1.9 cloacal diameters long; gu- bernaculum with or without ADOPHYSIS 82 bce ee ee 2, 2(1). Tail 9-10 cloacal diameters long Pea Ges O. dubius (Filipjev, 1918) De Coninck & Stekhoven, 1933 - Tail 7 cloacal diameters long or LESS rece ee a rene Oy ate MA 3 3(2). Longer cephalic setae 0.8 head diameter or less in length .... 4 - Longer cephalic setae 1 head di- ameter or more in length .... 8 4(3). Precloacal supplement absent in INVA C ee oh cr elk oh ae eae eats 5 - Precloacal supplement present Dinelanall © Vere a ac ee acne oa 6 5(4). 6(4). 7(6). 11(8). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Tail 7 cloacal diameters long; spicules 1.9 cloacal diameters long; gubernaculum - with apophysis ... O. problematicus Filipjev, 1946 Tail 4.2 cloacal diameters long; spicules | cloacal diameter long; gubernaculum without apoph- VSIS ie a cee eto aed O. pachylabiatus Stekhoven, 1946 Mandibles each with 14-15 denticles; cephalic capsule 45— S07 melongeee ase O. acantholaimus (Saveljev, 1912) Filipjev, 1927 Mandibles with about 6 denti- cles; cephalic capsule less than 45 um long Spicules 1.5 cloacal diameters in length O. hamatus (Steiner, 1916) Filipjev, 1927 Spicules 1.0 cloacal diameter in length O. australis (De Man, 1904) Filipjev, 1927 Tail 4-4.25 anal diameters in length Longer cephalic setae 2.7 head diameters in length Sa NS O. culcitatus Wieser, 1959 Longer cephalic setae about 1.0 head diameter in length Mandibles each with 15-20 denticles; male precloacal sup- plement 2-—2.5 cloacal diame- ters from cloaca O. dentatus (Ditlevsen, 1918) Filipjev, 1947 Mandibles each with about 4 denticles; male precloacal sup- plement 3.8—4.2 cloacal diam- eters from cloaca 5 Re coke O. polaris Filipjev, 1927 Precloacal supplement about 2.2 cloacal diameters from cloaca; gubernacular apophysis direct- 10 VOLUME 101, NUMBER 1 ed caudally Se rich. AES O. ditlevseni Inglis, 1964 - Precloacal supplement about 3.7 cloacal diameters from cloaca; gubernacular apophysis direct- ed dorsally . O. subantarcticus Mawson, 1958 Acknowledgments Sincere appreciation is expressed to Dr. W. Duane Hope of the National Museum of Natural History, Smithsonian Institu- tion, for reviewing the manuscript, exam- ining the specimens, and directing attention to the complex structures in the specimens. Appreciation is also expressed to Mr. & Mrs. William Fable, U.S. National Marine Fish- eries Service, Panama City, Florida for col- lecting the sediment samples from the Gulf of Mexico. Literature Cited Allgen, C. 1959. Free-living marine nematodes.— Further Zoological Results of the Swedish Ant- arctic Expedition 5:1-293. Belogurova, L. S., I. O. Belogurov, & E. S. Khamula. 191 1986. New species of Enchelidiidae nematodes from Tatar Strait.—Biologiya Morya 12:25-30. Chitwood, B. G. 1960. A preliminary contribution on the marine nemas (Adenophorea) of north- ern California.— Transactions of the American Microscopical Society 79:347-384. Cobb, N. A. 1920. One hundred new nemas.—Con- tributions to a Science of Nematology 9:217- 343. De Coninck, L. A., & J. H. Schuurmans-Stekhoven. 1933. The free-living marine nemas of the Bel- gian coast II.—Mémoirs du Musée Royale d’Historie Naturelle Belgique 58:1-163. Gerlach, S. A. 1953. Die Nematoden Besiedelung des Sandestrandes und des Kiistengrundwassers an der italienischen Kiiste. I, Systematischer Teil. — Archivio Zoologica Italiano 37:517-640. . 1954. Les Nematodes marins libres des eaux souterraines littorales d’esposende (Portugal). — Vie et Milieu 4:83-94. , & F. Riemann. 1974. The Bremerhaven checklist of aquatic nematodes. — VerOffentlich- ungen des Instituts fur Meeresforschung in Bre- merhaven, Supplement 4:1-735. Weiser, W. 1953. Free-living marine nematodes I. Enoploidea.—Acta Universitatis Lundensis; Medica, Mathematica, Scientiae Rerum Natur- alium (N. F. 2) 49:1-155. 306 Hibiscus Avenue, Panama City Beach, Florida 32407. PROC. BIOL. SOC. WASH. 101(1), 1988, pp. 192-208 NEW SPECIES AND NEW RECORDS OF SCALED POLYCHAETES (POLYCHAETA: POLY NOIDAE) FROM HYDROTHERMAL VENTS OF THE NORTHEAST PACIFIC EXPLORER AND JUAN DE FUCA RIDGES Marian H. Pettibone Abstract. —The joint Canadian-American Seamount Expedition in the North- east Pacific uncovered additional polynoid polychaetes associated with hydro- thermal vents of the Explorer and Juan de Fuca Ridges. They included five species, two new, in three subfamilies of Polynoidae: Lepidonotopodinae: Lep- idonotopodium piscesae, new species; Macellicephalinae: Levensteiniella kin- caidi Pettibone: Branchinotogluminae: Branchinotogluma grasslei Pettibone, P. sandersi Pettibone, Opisthotrochopodus tunnicliffeae, new species. The di- agnosis of Opisthotrochopodus is emended. Habitats of some species are dis- cussed. Among the unusual marine life discov- ered in August 1983 by the joint Canadian American Seamount Expedition (CASM) in the Northeast Pacific (Tunnicliffe & Juniper 1983, Harman 1984, Canadian American Seamount Expedition 1985) and later stud- ies on the Axial Seamount of the Juan de Fuca Ridge (Tunnicliffe et al. 1985) and the Explorer Ridge (Tunnicliffe et al. 1986), were five species of polychaete worms belonging to three subfamilies of Polynoidae. The subfamilies were described earlier by me from hydrothermal vents of the Tropical East Pacific: the Galapagos Rift and 21°N (Pettibone 1983, 1984, 1985a, b), and three of the five species were previously described from these areas: Levensteiniella kincaidi, Macellicephalinae; Branchinotogluma grasslei and B. sandersi, Branchinotoglu- minae. The remaining two are new species: Lepidonotopodium piscesae, Lepidonoto- podinae, and Opisthotrochopodus tunniclif- feae, Branchinotogluminae. The polynoids were collected by the Ca- nadian submersible DSRV Pisces IV in 1983, 1984, and 1986, and by DSRV Alvin in 1984, and sent to me for study by Verena Tunnicliffe. Additional specimens collected by DSRV Alvin in 1984 were provided by Meredith L. Jones. The specimens were ob- tained from four vent areas: Explorer Ridge (SO°N); Endeavour segment of the Juan de Fuca Ridge (48°N); Axial Seamount of the Juan de Fuca Ridge (46°N); and Southern Juan de Fuca Ridge (45°N). The Axial Seamount of the Juan de Fuca Ridge was described by Tunnicliffe & Ju- niper (1983), Harmon (1984), Canadian American Seamount Expedition (1985), and Tunnicliffe, Juniper, & de Burgh (1985). Axial Seamount is a volcanic feature formed by hydrothermal activity, 46° off the U.S.- Canadian border. The seamount is crossed by a fissure of about 1510 meters, within which is a system of vents with water tem- peratures up to 35°C. Dense thickets of ves- timentiferan tubes overgrow the major vents and consist of deposits of sulfide-forming chimneys. Large quantities of mucus bind the tubes together and provide habitats for many organisms. Chemosynthetic bacteria are found in the mucous strands. The vents vary from diffuse flow through bacterial mats to large structures (over 2 meters high) built VOLUME 101, NUMBER 1 by the vestimentiferan tube worms. The m1- nor vents are surrounded by bacterial mats, small prone vestimentiferans, and limpets. Colonial protozoans (folliculine ciliates) and bacterial mats cover most surfaces of the active fissures. Associated animals of the Axial Sea- mount, in addition to the vestimentiferan Ridgeia piscesae Jones, with its large, soft collared tubes, were two species of alvinellid polychaetes: Paralvinella palmiformis Des- bruyéres & Laubier, the large ““palm-worm”’ that wraps the caudal end around the Rid- geia tubes with the anterior end held up- right; and P. pandorae Desbruyéres & Lau- bier, the smaller “‘pandora-worm,” that forms mucus-lined tubes on the bases of the vestimentiferans. The ampharetid 4Amphis- amytha galapagensis Zottoli was also found living in small tubes of mineral fragments on the periphery of the vents. Two new polynoid polychaetes were tentatively iden- tified as species A and B by D. Weston; they were examined and referred herein to Lep- idonotopodium piscesae, n. sp. (species A) and Branchinotogluma grasslei Pettibone, 1985 (species B). The larger white polynoids (=L. piscesae) were observed crawling on vestimentiferan tubes and rocks and grazing on bacterial and protozoan mats; its white coloration was due to the filamentous bac- teria attached to the elytra. The smaller red polynoids (=B. grasslei) were found closer to the water flow than the larger white species. The extensive hydrothermal vents of the Explorer Ridge (SO°N, 130°W, 1820 m) were described by Tunnicliffe et al. (1986). This is the most northern of the recently explored vent areas of the eastern Pacific. The vents are formed of large sulfide mounds project- ing 2 to 20 meters in height through thick grey hydrothermal sediment. The mounds are covered with active chimneys and vent grey ‘smoke’ which 1s a fluid rich in hydro- gen-sulfide and of a high temperature (25-— 310°C). Vestimentiferan tubes were the ma- jor habitat for other species; limpets were 193 particularly abundant. Three polynoid species were listed (Pettibone, personal communication): Branchinotogluma sp. (=B. grasslei, B. sandersi Pettibone, 1985), Lepidonotopodium sp. (=L. piscesae, n. sp.), and Levensteiniella sp. (=L. kincaidi Petti- bone, 1985). Holotype, paratypes, and additional specimens are deposited in the National Museum of Natural History, Smithsonian Institution (USNM). Paratypes and repre- sentative specimens are deposited in the National Museum of Natural Sciences, Ot- tawa, Canada (NMCA). Family Polynoidae Subfamily Lepidonotopodiinae Pettibone, 1983 Lepidonotopodium Pettibone, 1983 The genus includes three previously de- scribed species: L. fimbriatum Pettibone, 1983, from off western Mexico at 21°N; L. riftense Pettibone, 1984, and L. williamsae Pettibone, 1984, both from the Galapagos Rift. A new species from the Explorer and Juan de Fuca Ridge systems is added. Lepidonotopodium piscesae, new species Figs. 1-4 Large white polynoid, Tunnicliffe, Juniper, & de Burgh, 1985:454, 455, 459, fig. 4 (bottom left). [Axial Seamount] Lepidonotopodium sp., Tunnicliffe et al., 1986:407. [Explorer Ridge] Lepidonotopodium, n. sp. (A), Desbruyéres et al., 1985:114. [Juan de Fuca, sulfide rocks] Material. —Hydrothermal vents of Northeast Pacific, dives by DSRV Pisces IV (P) and DSRV Alvin (A): EXPLORER RIDGE, 49°45'N, 130°16’W, 1812-1823 m: P1494-600/605, Gulati and Gusher vents, 1 Jul 1984, 14 paratypes (8 small) (USNM 102883); P1497-611, Crab vent, 4 Jul 1984, 7 paratypes (USNM 102886); P1505-617, Buster Thruster vent, 19 Aug 1984, 8 para- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 194 VOLUME 101, NUMBER 1 types (USNM_ 102888).—ENDEAVOUR SEGMENT, JUAN DE FUCA RIDGE, 4T°ST'N, 129°06'W, 2208 m, A1452-621/ 641, 7 Sep 1984, 7 paratypes (USNM 102889).—AXIAL SEAMOUNT, JUAN DE FUCA RIDGE, 45°56-59'N, 130°01- 03’W, 1546-1592 m: A1411, 16 Jul 1984, 2 young paratypes (USNM 102882); A1413, 18 Jul 1984, holotype (USNM 102878), 3 paratypes (USNM 102881), 23 paratypes (USNM 102887), 3 paratypes (NMCA1987- 0449); P1327-639, 17 Aug 1983, Taylor’s vent, paratype (USNM 102884); P1721-628, 17 Jul 1986, Embley’s Inferno, 3 paratypes (USNM 102879); P1728-632, 29 Jul 1986, Demon vent, 3 small paratypes (USNM 102880). SOUTHERN JUAN DE FUCA RIDGE, 24 Oct 1984, 44°39'N, 130°22'W, 2200 m: A1455-1B-616, 4 paratypes (USNM 102890); A1463-7B-615, paratype (USNM 102885). Measurements. — Holotype from Endea- vour Segment, Juan de Fuca Ridge, 28 mm long, 13 wide with setae, with 26 segments, last one small; additional paratypes 13-29 mm long, 8-12 mm wide, with 24—26 seg- ments. Adult paratypes from Explorer Ridge 10-23 mm long, 6-11 mm wide, with 24— 26 segments: young paratype 4 mm long, 3 mm wide, with 18 segments, last one small. Adult paratypes from Axial Seamount, Juan de Fuca Ridge, 11-28 mm long, 6-14 mm wide, with 24—28 segments, last one minute; young paratype 9 mm long, 7 mm wide, with 22 segments. Paratypes from Southern Juan de Fuca vent 19-29 mm long, 9-13 mm wide, with 25—26 segments. Description. — Body sturdy, elongate-oval, rounded anteriorly and posteriorly, flat- — Fig. 1. 195 tened ventrally and arched dorsally; color brownish to tan. Elytra 11 pairs, on seg- ments 2, 4, 5, alternate segments to 21; with dorsal cirri on posterior segments. Elytra large, overlapping, covering dorsum, thick, stiff, opaque, attached eccentrically on prominent elytrophores. First pair oval, with anterolateral notch (for passage of dorsal tentacular cirrus), following ones subreni- form, posterior pair (11th) subtriangular (Fig. 1A—D). Elytral surface thickly covered with opaque, rounded to conical microtu- bercles, especially thick on somewhat raised transverse areas in middle of elytra and along posterior and lateral borders, and with scat- tered globular and filiform micropapillae; both tubercles and papillae with bacterial “hairs” attached (Fig. 1A—D). Dorsal cirri on segments lacking elytra with large cylin- drical cirrophores on posterior sides of no- topodia, with styles long, tapered, extending beyond tips of neurosetae; dorsal tubercles on cirrigerous segment elongate, inflated (Fig. 2C). Surfaces of elytrophores, dorsal tubercles, and anterior sides of parapodia with numerous ciliated ridges (Figs. 1E, 2A— (©). Prostomium bilobed, with anterior lobes prominent, tapered, with rather long frontal filaments; wider on posterior half, whitish, with pair of tannish or dark spots, appearing as “‘eyes’’; median antenna with large cylin- drical ceratophore in anterior notch with style short tapered, about length of ceratop- hore; palps stout, smooth, tapering, about 1’ time length of prostomium (Fig. 1E). First or tentacular segment fused to prosto- mium, not visible dorsally; tentaculophores lateral to prostomium, without setae, each Lepidonotopodium piscesae, A-F, holotype; G, H, paratype, USNM 102890: A, Left Ist elytron, segment 2, with detail of microtubercles, micropapillae, and bacterial “hairs”; B, Left 2nd elytron, segment 4; C, Right middle elytron, with detail of A; D, Left 11th elytron, segment 21; E, Anterior end, dorsal view, palp, dorsal and ventral tentacular cirri of right side smaller, regenerating; F, Same, ventral view, right parapodium of segment 2 not shown; G, Extended pharynx, frontal view, showing border papillae and dark red jaws; H, Dorsal jaws removed. Scales = 2.0 mm for A—D; 1.0 mm for E-G; 0.5 mm for H. 196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON en ee See ee ee D : F Fig. 2. Lepidonotopodium piscesae, holotype: A, right middle elytragerous parapodium, anterior view, with detail of neuropodial papillae with bacterial “hairs”; B, Same, posterior view; C, Right middle cirrigerous parapodium, posterior view; D, Notosetae; E, Supraacicular neuroseta; F, Upper and lower subacicular neu- rosetae, with detail of spinous rows. Scales = 1.0 mm for A—C; 0.1 mm for D-F. with pair of tentacular cirri, subequal in prominent cirrophores lateral to ventral length, slightly shorter than palps; without mouth, with styles similar to tentacular cir- distinct facial tubercle (Fig. 1E, F). ri, larger than following ventral cirri (Fig. Second or buccal segment with first pair 1E, F). Ventral mouth enclosed in fleshy of large elytrophores, biramous parapodia, upper, lateral and posterior lips medial to and ventral buccal cirri attached basally on parapodia of segments | and 2 (Fig. 1F). VOLUME 101, NUMBER 1 11 12 14 15 197 E Fig. 3. Lepidonotopodium piscesae, paratypes, A, USNM 102887; B-E, USNM 102889: A, Ventral view of left side of segments 11-15, showing segmental papillae; B, Left neuropodium from segment 12, anterior view, separated from notopodium, neuroaciculum dotted; C, Same, posterior view; D, Supraacicular neurosetae from same; E, Upper and lower subacicular neurosetae from same. Scales = 1.0 mm for A; 0.5 mm for B, C; 0.1 mm for D, E. Large muscular eversible pharynx with 7 pairs of border papillae and 2 pairs of strong, hooked, dark red jaws; dorsal row of pa- pillae subequal in size, with middle one only slightly larger; ventral row with lateral pair and 3 medial papillae smaller, with next to lateral pair elongate and tapered (Fig. 1G). Jaws serrated on cutting edge, with up to 12 or so teeth (Fig. 1G, H). Biramous parapodia with shorter noto- podia on anterodorsal sides of longer neu- ropodia (Fig. 2A—C). Notopodium large, rounded, with projecting acicular lobe on posterior side, hidden by very numerous notosetae, and nearly enclosed by well-de- veloped, large, flaring bract, attached to acicular lobe on posterior side; lower thin- ner part of bract bordered with papillae with attached bacterial “hairs” (Figs. 2A, B, 4A). Neuropodium diagonally truncate, with shallow notch on posterior lower side and deep notch on posterior upper side; upper part of presetal acicular lobe projecting be- yond shorter postsetal lobe; distal borders of neuropodial lobes papillate with attached bacterial “‘hairs’”’ (Figs. 2A—C, 3B, C). No- tosetae very numerous, forming radiating bundles, varying in length from short to long, much stouter than neurosetae, distal part with 2 rows of spines and blunt tapered tips, most with numerous attached bacterial “hairs” (Figs. 2D, 4B). Neurosetae numer- ous, forming fan-shaped bundles. Upper, supraacicular neurosetae few (4 or so) em- anating from upper posterior notch of neu- ropodium, stouter than lower neurosetae, with 2 rows of prominent, lateral spines and bare tapered tips (Figs. 2E, 3D). Very nu- 198 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON LEEBS Fig. 4. Lepidonotopodium piscesae, paratype, USNM 102889: A, Left notopodium and elytrophore from segment 13, separated from neuropodium, posterior view, notoaciculum dotted, with detail of papillae on ventral part of bract; B, Short (left) and long (right) notosetae from same, with bacterial ‘“‘hairs”; C, Left neuropodium from segment 13 anterior view, neuroaciculum dotted; D, Same, posterior view; E, Supraacicular neuroseta from same, with detail of fine spinous rows; F, Subacicular neuroseta from same, with detail of spinous rows. Scales = 0.5 mm for A, C, D; 0.1 mm for B, E, F. merous subacicular neurosetae with slightly hooked tips and fine spinous rows on cutting edge (Figs. 2F, 3E, 4F). On 7 paratypes, neu- ropodia of segment 13 differing by presence of only 1—2 long, stout, reddish sabre-like supraacicular neurosetae, appearing smooth but with longitudinal rows of fine spines (Fig. 4C-E), with greater development of upper fimbriated bract (compare Figs. 3B— D and 4C-E). Ventral cirri attached on mid- dle of neuropodia, tapering to slender tips, and extending to tips of neuropodia (Figs. 2A-C, 3B, C). About half of adult paratypes with 5 pairs of elongate ventral papillae on segments 1 1 to 15, attached on middle bases of neuro- podia, and extending to near bases of ven- tral cirri or beyond (Figs. 3A—C, 4C, D). Pygidium bulbous dorsal lobe between parapodia of posterior few smaller seg- ments, with or without pair of anal cirri. Etymology. —The species is named for the Canadian submersible DSRV Pisces IV. Distribution. —Hydrothermal vents of Northeast Pacific, in 1546 to 2208 meters. Biology. —Lepidcnotopodium piscesae lives on the periphery of the vents, crawling on rocks and vestimentiferan tubes, grazing VOLUME 101, NUMBER 1 on protozoan and bacterial mats. One spec- imen was observed on time-lapse photog- raphy crawling to the top of a tube and grop- ing around inside (V. Tunnicliffe, in litt.). The elytra are often coated with bacterial filaments, giving them a white color. Lepidonotopodium piscesae is most closely related to L. williamsae from the Galapagos Rift. The two species differ from one another as follows: L. piscesae has 5 pairs of elongate segmental papillae on segments 11-15, in- stead of 4 pairs on segments 1 2—15; a phar- ynx with 7 pairs of unequal-sized border papillae, instead of subequal papillae, jaws with up to 12 basal teeth, instead of 7, and has 1-2 elongate sabre-like supraacicular neurosetae on segment 13 of a few speci- mens, instead of lacking them entirely. Subfamily Macellicephalinae Hartmann-Schroder, 1971, emended Pettibone, 1976 Genus Levensteiniella Pettibone, 1985 Levensteiniella kincaidi Pettibone Fig. 5 Levensteiniella kincaidi Pettibone, 1985b: 741, figs. 1-3. Levensteiniella sp. Tunnicliffe et al., 1986: 407. [Explorer Ridge] Material. —Hydrothermal vents of Northeast Pacific, dives by DSRV Pisces IV (P) and DSRV Alvin (A): EXPLORER RIDGE, 49°45'N, 130°16’W, 1818-1823 m: P1494-604/605, 1 Jul 1984, 5 specimens (USNM 102869), 1 specimen (NMCA 1987- 0451); P1505-602, 19 Aug 1984, Busted Thruster vent, | specimen (USNM 102867).—ENDEAVOUR SEGMENT, JUAN DE FUCA RIDGE, 47°57'N, 129°04-06'W, 2208-2213 m A1418, 24 Jul 1984, 1 specimen (USNM 102868); A1447- 614, 3 Sep 1984, 2 specimens (USNM 102870); A1452-621, 7 Sep 1984, Lt. Obo vent, 1 specimen (USNM 102871).—AX- IAL SEAMOUNT, JUAN DE FUCA RIDGE, 45°56—-57'N, 130°01'W, 1546-1553 m;A1411, 16 Jul 1984, residue from animal 199 container, young specimen (USNM 102866); A1413, 18 Jul 1984, young spec- imen (USNM 102872). Remarks. —The 12 specimens agree with previously described specimens from the Galapagos Rift and 21°N vent sites. Some specimens are larger than previously re- ported (12 mm long, 6 mm wide, with 25 segments). The largest specimen from Ex- plorer Ridge is 16 mm long, 9 mm wide, for 25 segments, with the last one small; additional adult specimens are 12-15 mm long, 7-9 mm wide, for 25 segments: a small specimen is 7 mm long, 4 mm wide, for 23 segments, with the last 3 small. The largest specimen from the Endeavour Segment is 16 mm long, 9 mm wide, for 25 segments; a small specimen is 6 mm long, 5 mm wide, for 22 segments. Two young specimens from Axial Seamount are 1.2 mm long, 1.2 mm wide, for 11 segments. Most of the elytra were missing. On a small specimen from Endeavour Segment, Juan de Fuca Ridge, three pairs of elytra remained on segments 2, 4, and 5; in ad- dition to the usual filiform papillae on the surface, some papillae on the posterior and lateral borders were enlarged basally (Fig. 5A, B). Four of the larger specimens had long ventral papillae on segments 11 and 12, sometimes directed dorsally between the parapodia. Distribution. —Hydrothermal vents of Tropical East Pacific: Galapagos Rift and East Pacific Rise at 21°N, in 2457-2633 m:; Northeast Pacific: Explorer Ridge, Endea- vour Segment and Axial Seamount, Juan de Fuca Ridge, in 1546-2213 m. Subfamily Branchinotogluminae Pettibone, 1985 Branchinotogluma Pettibone, 1985 Branchinotogluma grasslei Pettibone Red polynoid polychaete, Tunnicliffe & Ju- niper, 1983:966. [Axial Seamount, Juan de Fuca Ridge] Branchinotogluma_ grasslei Pettibone, a A Fig. 5. \ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON JPN B Levensteiniella kincaidi, small specimen (6 < 5 mm, with 22 segments), USNM 102870: A, Left Ist elytron, segment 2, with detail of border papillae; B, Right 2nd elytron, segment 4, with detail of border and surface papillae. Scale = 0.5 mm. 1985a:457, figs. 5, 6.—Newman, 1985 (list). Branchinotogluma sp., Tunnicliffe et al., 1986:407. [Explorer Ridge] Material. —Hydrothermal vents of Northeast Pacific, dives by DSRV Pisces IV (P) and DSRV Alvin (A): EXPLORER RIDGE, 49°45'N, 130°16'W, 1818 m: P1494, 1 Jul 1984, 1 specimen (USNM 102864); P1495-606, 2 Jul 1984, Lunch Hour vent, 2 specimens (USNM 102861).— ENDEAVOUR SEGMENT, JUAN DE FUCA RIDGE, 47°57'N, 129°04—06’W, 2208 m: A1419, 25 Jul 1984, 13 specimens (USNM 102859), 2 specimens (NMCA 1987-0452); A1452.-21, Lt. Obo vent, 7 Sep 1984, 4 specimens (USNM 102865).—AX- IAL SEAMOUNT, JUAN DE FUCA RIDGE, 45°59'N, 130°03’W, 1592 m: P1327, 17 Aug 1983, Taylor’s vent, 2 spec- imens (USNM 102860); P1728-634, 29 Jul 1986, Demon vent no. 1, 6 specimens (USNM 102863).—SOUTHERN JUAN DE FUCA RIDGE, A1410, 15 Jul 1984, 45°13'N, 130°09'W, 2380 m, on chaleopy- rite and sphalerite, 1 specimen (USNM 102862). Remarks.—The 31 specimens from the four vent areas agree with the specimens previously described from the Galapagos Rift and 21°N vent sites. Specimens from the Explorer Ridge are 14-23 mm long, 7— 10 mm wide with setae and have 21 seg- ments. From Endeavour Segment, larger specimens are 13-31 mm long, 6-12 mm wide, and have 21 segments; small speci- mens are 5—7 mm long, 3—4 mm wide, and have 18-20 segments. From Axial Sea- mount, the larger specimens are 12-26 mm long, 6-11 mm wide and have 21 segments; the small specimen is 5 mm long, 3 mm wide, with 19 segments. Specimen from Southern Juan de Fuca Ridge is 25 mm long, 12 mm wide, and has 21 segments. Distribution. —Hydrothermal vents of Tropical East Pacific: Galapagos Rift and East Pacific Rise at 21°N, in 2482-2633 m; Northeast Pacific: Explorer Ridge, Endea- vour Segment and Axial Seamount, Juan de Fuca Ridge, and Southern Juan de Fuca Ridge, in 1495-2380 m. Biology. —Branchinotogluma grasslei has been found living on rocks and the soft ves- timentiferan tubes of Ridgeia piscesae Jones in the vicinity of very hot vents and high VOLUME 101, NUMBER 1 hydrogen-sulfide levels. They are bright red and very difficult to see in the shimmering hot water of the vents (V. Tunnicliffe, in litt.). Branchinotogluma sandersi Pettibone Branchinotogluma sandersi Pettibone, 1985a:453, figs. 3, 4.—Newman, 1985: 232 (list). Branchinotogluma sp., Tunnicliffe et al., 1986:407. [Explorer Ridge] Material. —Hydrothermal vents of Northeast Pacific, dives by DSRV Pisces IV (P) and DSRV Alvin (A): EXPLORER RIDGE, 49°45'N, 130°16’W, 1818 m, P1494, 1 Jul 1984, 2 specimens (USNM 102857).—ENDEAVOUR SEGMENT, JUAN DE FUCA RIDGE, 47°57'N, 129°04--06'W, 2208 m: A1419, 25 Jul 1984, 3 specimens (USNM 102853), 1 specimen (NMCA 1987-0453); A1452-621, Lt. Obo vent, 7 Sep 1984, 3 specimens (USNM 102856).—AXIAL SEAMOUNT, JUAN DE FUCA RIDGE, 45°59’'N, 130°03’W, 1592 m: P1327-640, 17 Aug 1983, Taylor’s vent, | specimen (USNM 102854): P1728- 634, 29 Jul 1986, Demon vent no. 1 spec- imen (USNM 102855). Remarks. —The 11 specimens from three vent areas agree with the previously de- scribed specimens from the Galapagos Rift and 21°N vent sites. Two specimens from Explorer Ridge are 8—12 mm long, 4-6 mm wide with setae, for 21 segments. Larger specimens from Endeavour Segment mea- sure 12-15 mm long, 6-8 mm wide, for 21 segments; smaller specimens are 8-9 mm long, 5-6 mm wide, for 21 segments, with the posterior 3 segments not fully devel- oped. A large specimen from Axial Sea- mount is 21 mm long, 11 mm wide, for 21 segments. One specimen from Endeavour Segment (USNM 102856) was observed with groups of very long white filaments emerging from the body on the left side between parapodia 201 of segments 12-13 and 13-14, possibly in- dicative of a parasite. Distribution. —Hydrothermal vents of Tropical East Pacific: Galapagos Rift and East Pacific Rise at 21°N, in 2451-2633 m; Northeast Pacific: Explorer Ridge, Endea- vour Segment and Axial Seamount, Juan de Fuca Ridge, in 1592-2208 m. Branchinotogluma sp. B Branchinotogluma sp. B, Pettibone, 1985a: 466. Material. —Explorer Ridge, 49°45.6'N, 130°16.1'W, 1818 m, P1494, 1 Jul 1984, 2 young specimens (USNM 102858). Remarks. — The two young specimens, 4— 6 mm long, 2-3 mm wide with setae, and having 16-17 segments, may be juveniles of B. grasslei, B. sandersi, or Opisthotro- chopodus tunnicliffeae; all three species were found at the same station. Opisthotrochopodus Pettibone, 1985, emended Type species. —Opisthotrochopodus alvi- nus Pettibone, 1985a; by original designa- tion. Gender: masculine. Opisthotrochopodus is emended to in- clude the new species O. tunnicliffeae. Diagnosis.—Body short, with 21 seg- ments, first achaetous. Elytra 10 pairs, on Segments 25 495.) spose els. lispeli/eeand 19. Dorsal cirri with short cirrophores and long styles, and dorsal tubercles, in line with elytrophores, on segments 3, 6, 8, 10, 12, 14, 16, 18, 20, and 21. Arborescent bran- chiae 4 per segment, attached to lateral sides of elytrophores, dorsal tubercles, and dorsal sides of notopodia, beginning on segment 3 and continuing to segment 17 or 18. Pro- stomium bilobed with triangular anterior lobes bearing frontal filaments, with cera- tophore of median antenna in anterior notch, with paired ventral palps, without lateral antennae or eyes. First or tentacular seg- ment not visible dorsally; tentaculophores 202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Opisthotrochopodus tunnicliffeae, A-H, holotype; I, paratype, USNM 102874: A, Anterior end, dorsal view; B, Same, ventral view; C, Left elytragerous parapodium of segment 2, anterior view, acicula dotted; D, Left 1st elytron, segment 2; E, Left 2nd elytron, segment 4; F, Left 8th elytron, segment 15; G, Left 9th elytron, segment 17; H, Left 10th elytron, segment 19; I, Dorsal view left side of segments 8 (cirrigerous) and 9 (elytra- gerous), showing only base of dorsal cirrus. nB, notopodial bract. Scales = 1.0 mm for A, B, I; 1.0 mm for C; 2.0 mm for D-H. VOLUME 101, NUMBER 1 lateral to prostomium, each with dorsal and ventral tentacular cirri, without setae. Sec- ond or buccal segment with first pair of ely- tra, biramous parapodia, and ventral or buccal cirri attached to basal parts of para- podia lateral to mouth; styles longer than following ventral cirri. Parapodia biramous, with notopodia shorter than neuropodia. Notopodia of elytragerous segments 2-17 with prominent rounded bracts enclosing conical acicular lobes and notosetae, or only on segment 2. Neuropodia with longer, flat- tened, conical presetal and shorter, rounded postsetal lobes. Notoseta stouter than neu- rosetae, straight, acicular, smooth or with spines. Neurosetae long, slender, finely spi- nous, with slightly hooked tips. Ventral cirri short, tapered. Posterior 4 segments (18-21) compressed; parapodia directed posteriorly and greatly modified, with expanded deli- cate lamellae; elongated cylindrical noto- podia fused with dorsal cirrophores and dis- tal styles on segments 20 and 21; with unique neuropodial wheel organs on segment 20, including stout acicular neurosetae. Pharynx with 5 border papillae, 3 dorsal and 2 ven- tral; 2 pairs of dorsal and ventral hooked jaws, each with row of numerous, minute teeth along edge. Segmental ventral papillae or lamellae on segments 12-17. Pygidium with or without pair of anal cirri. Opisthotrochopodus tunnicliffeae, new species Figs. 6-9 Material. —Hydrothermal vents of Northeast Pacific, dives by DSRV Pisces IV (P) and DSRV Alvin (A): EXPLORER RIDGE, 49°45'N, 130°16’W, 1818 m: P1494, 1 Jul 1984, paratype (NMCA 1987- 0450): P1495-606, 2 Jul 1984, Lunch Hour vent, paratype (USNM 102877).—AXIAL SEAMOUNT, JUAN DE FUCA RIDGE, 16 Jul 1984, 45°57'N, 130°01’W, 1553 m, A1411, residue from animal container, ho- lotype (USNM 102873), 4 paratypes, 2 young (USNM 102874-6). 203 Description. —Length of holotype 25 mm, width with setae 11 mm, segments 21. Length of largest paratype (USNM 102874) 30 mm, width 12 mm, segments 21. Two smaller paratypes 15-17 mm long, 7-9 mm wide, with 21 segments. Two young para- types (USNM 102876) 2—2.5 mm long, 2- 2.5 mm wide, with 14-17 segments, last ones minute. Body flattened ventrally, arched dorsally, slightly tapered anteriorly and posteriorly, with parapodia longer than body width; posterior few segments (18-21) compressed, with parapodia modified (Fig. 8B, C). Elytra large, overlapping, covering dorsum; round to oval, posterior pair small- er, elongate-oval, enclosing posterior mod- ified segments (Fig. 6 D-H). Elytra smooth, stiff, somewhat vaulted around place of at- tachment to elytrophores. Elytrophores large, projecting posteriorly, with arbores- cent branchiae attached on lateral sides (Figs. 61, 7B, 8C, D). Dorsal tubercles elongate, projecting posteriorly with arborescent branchiae on distal part (Figs. 61, 7A, 8C). Prostomium bilobed, without eyes or lat- eral antennae; anterior lobes subtriangular, with rather long frontal filaments; median antenna with bulbous ceratophore in ante- rior notch, subulate style with long slender tip, shorter than tentacular cirri; palps stout, tapered, smooth (Fig. 6A, B). First or ten- tacular segment forming upper and lateral lips of mouth; tentaculopores lateral to pro- stomium, without setae, each with pair of tentacular cirri, dorsal tentacular cirrus longer than palp, ventral one shorter than dorsal (Fig. 6A, B). Second or buccal seg- ment forming posterior lip of mouth, bear- ing biramous parapodia and first pair of ely- trophores; notopodia with hood or bract (nB) on anterodorsal side encircling small bundle of notosetae and conical acicular lobe; ven- tral buccal cirri similar to tentacular cirri, longer than following ventral cirri (Fig. 6A— C). Muscular pharynx observed cut open, showing usual pattern in genus: 5 papillae around opening, 3 dorsal and 2 ventral; 2 pairs of amber-colored jaws minutely den- 204 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \\ SS 7 Fig. 7. Opisthotrochopodus tunnicliffeae, holotype: A, Left middle cirrigerous parapodium, posterior view; B, Left middle elytragerous parapodium, anterior view; C, Left middle neuropodium, anterior view, notopodium cut off, neuroaciculum dotted; D, Notosetae from same; E, Supraacicular neurosetae from same, with detail magnified; F, Subacicular neuroseta from same, with detail magnified, one view of tip turned. Scales = 1.0 mm for A-C; 0.1 mm for D, F. ticled on inner border (see Pettibone 1985a: notopodium rounded, with projecting, ta- fig. 8H, I). pered acicular lobe and radiating bundle of Biramous parapodia of segments 3-17 numerous notosetae; neuropodium with similar, with smaller notopodium on _ presetal lobe long, subconical, with pro- anterodorsal side of larger neuropodium; jecting acicular lobe; postsetal lobe shorter, VOLUME 101, NUMBER 1 205 Fig. 8. Opisthotrochopodus tunnicliffeae, holotype: A, Ventral view of segments 12-17, showing ventral papillae on segments 12-15 and ventral lamellae on segments 16 and 17, parapodia incompletely shown; B, Ventral view of posterior end, including segments 17—21 and pygidium, wheel organ of segment 20 withdrawn on right side; C, Dorsal view of same; D, Left elytragerous parapodium from segment 17, anterior view; E, Right cirrigerous parapodium from segment 18, posterior view; F, Right elytragerous parapodium from segment 19, anterior view, acicula dotted. nL, notopodial lamella. Scales = 1.0 mm for A-C; 1.0 mm for D-F. rounded; lobes widely separated on dorsal side, with projecting small lobe on dorsal base (hidden from view by anterior noto- podium), with fan-shaped bundle of very numerous neurosetae (Figs. 7A—C, 8D). No- tosetae stouter than neurosetae, long, straight, tapered, acicular, with 2 rows of spines; shorter notosetae mostly smooth (Fig. 7D). Neurosetae long, slender, slightly wid- er basally, with slightly hooked tips; su- 206 praacicular neurosetae with widely spaced spines extending to near tips, finely spinous distally (Fig. 7E): subacicular neurosetae finely spinous (Fig. 7F). Dorsal cirrus on cirrigerous segments on posterodorsal side of notopodium, with cylindrical cirrophore and long style extending far beyond setae; dorsal tubercle extending laterally, with ar- borescent branchia distally; additional branchia on dorsal base of notopodium (Figs. 61, 7A). Elytragerous segments with large, bulbous elytrophore extending pos- terolaterally, with large arborescent bran- chia on lateral base and additional branchia on dorsal base of notopodium (Figs. 7B, 8C, D). Ciliated bands on dorsum, extending onto elytrophores and dorsal tubercles, as well as on anterior and posterior sides of parapodia (Figs. 6A, I, 7A, B). Ventral cirri on middle of neuropodia tapering to slender tips (Fig. 7A, B). Cirrigerous parapodium of segment 18 much smaller than preceding parapodium, with long dorsal cirrus projecting posterior- ly and single, small arborescent branchia projecting from dorsal tubercle (Fig. 8B—E). Segment 19 with smaller elytrophores ap- proaching medially, usually without bran- chiae (small one on under side of left ely- trophore of holotype), with small parapodia wedged between adjacent parapodia and hidden from view dorsally; notopodium with delicate lamella (nL) on dorsal side; notosetae stout, acicular, with 2 rows of spines; neurosetae slender, minutely spi- nous, with capillary tips (Fig. 8B, C, F). Segment 20 with parapodia greatly mod- ified. Neuropodium enlarged (correspond- ing to wheel organ of O. alvinus), inflated, with projecting, conical acicular lobe, with neurosetae projecting from wide opening and ventral cirrus attached near base (Figs. 8B, C, 9A, B, F). Neurosetae consisting of up to 4 stout, acicular, harpoon setae (Fig. 9E) and long slender setae with 2 rows of long spines on basal half, tapering to slen- der, bare tips (Fig. 9D). Neuropodium on holotype inflated and extended on left side PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON but retracted and wrinkled on right side (Figs. 8B, C, 9A, F); largest paratype (USNM 102874) with modified neuropodia extend- ed on both sides. Notopodium with acicular lobe fused to cirrophore of dorsal cirrus, with style extending posteriorly; basal part of notopodium compressed against inflated neuropodium and hidden from view until separated, with row of about 13 short, spi- nous notosetae enclosed in delicate lamella (nL) (Fig. 9A—C). Small paratype of 17 x 9 mm (USNM 102877) with both neuropodia inflated, each with 2—3 harpoon neurosetae. Slightly smaller paratype of 15 : SA IN SS SS \ \ N N \ \ N Q N \ \ \ Fig. 4. Booralana tricarinata, holotype male: A-E, Pleopods 1-5; F, Right uropod; G, Penes. Scale = 1 mm for A-F, 3 mm for G. VOLUME 101, NUMBER 3 Fig. 5. Booralana tricarinata: A-C, Left uropods of 47, 41, and 37 mm males, respectively, from Puerto Rico; D, Right uropod, adult (?) male from British Virgin Islands (fish stomach); E, Endopod, right uropod, adult (?) male from Puerto Rico (fish stomach); F, Left uropod, adult 47 mm female from Puerto Rico; G, Left uropod, adult 42 mm female from Grand Bahama Island. Scale = 2 mm. hamian males, including the holotype, the distolateral margin of the endopod is entire and has a dense fringe of setae; whereas An- tillean males have several sharp tubercles along the distolateral margin and a thin fringe of setae, similar to the condition seen on females from both areas and on smaller subadult males from the Bahamas. More- over, the tubercles on the distolateral mar- gin are more strongly developed on females from Puerto Rico than on females of similar size from the Bakamas. The appendix masculina on Bahamian males is relatively shorter than that of males from the Antilles, although there is an ex- ception in this case, too. The appendix mas- culina on each Bahamian male does not reach the distal end of the inner ramus of the second pleopod, whereas it does reach the distal end of the ramus on each of the two intact males from off the north coast of Puerto Rico and on two of the three frag- mented males from elsewhere in the An- tilles. However, on one of the fragmented males from Anegada, British Virgin Islands, the appendix masculina is short, not reach- ing the end of the ramus. The populations from the Bahamas and the Antillean Islands may be diverging, or the species may be quite variable. We may be seeing in our limited material phenotypic effects of great water depth or temperature differences between the localities represent- ed. Nevertheless, the absence of a dense se- 612 tal fringe on the endopods of uropods of apparently adult males from the Antilles is worrisome. For now, we have chosen to conservatively consider the populations as one variable species. When more material becomes available, a further investigation can be made of their differences. Affinities. — Booralana tricarinata can be distinguished immediately from B. bathy- nella and B. wundurra by the presence and configuration of carinae on the dorsal sur- face of the pleotelson. Booralana tricarinata has one longitudinal carina on the midline of the pleotelson and a pair of submedian carinae. Booralana bathynella lacks carinae on the pleotelson; B. wundurra has a pair of submedial carinae but lacks one on the midline. The shape of the pleotelson of B. tricarinata is most similar to that of B. ba- thynella; in both species the lateral margins of the pleotelson are convex, sloping evenly to a broad, truncate posterior margin. In B. wundurra the posterior margin is relatively much narrower than in the other two species, and it is upturned in males. Finally, the eyes of preserved specimens of B. tricarinata are brown or black, but the eyes of B. wundurra are red (Bruce 1986:136). The posterolateral notch in the enlarged lateral margin of the third pleonite may be a reliable generic character, although the notch is more weakly developed in B. wun- durra than in the other two species (N. Bruce, pers. comm.). In specimens of B. tricarinata the basal segment of the uropod becomes inserted in the notch when the pleotelson is flexed ventrally. Functionally, this meshing of the pleotelson and pleonite appears to be a locking mechanism used for leverage when the isopod is clinging to a fish carcass. Etymology. —The specific name, tricari- nata, refers to the three prominent carinae on the dorsal surface of the pleotelson. Type locality.—Atlantic Ocean, North- west Providence Channel, about 3 km southeast of Lucaya, Grand Bahama Island (approximately 26°32’N, 78°33’W), 200-250 m depth. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Distribution and ecology. — Booralana tri- carinata is known from outer-shelf and up- per-slope depths off the Little Bahama Bank and the Antilles Islands; 110-610 m depths. There was no depth recorded with the sample from Smithsonian-Bredin Station 74-A-58 at Redonda Island, but the female was taken at the anchorage there. The col- lection data includes the entry “‘(dip net?).” We have found that the collection was made in depths shallower than those at the col- lection sites of other material examined, at the surface, with light and dip net. The limited data indicate that B. tricar- inata is an epibenthic scavenger, because most of the specimens came from baited mollusk or fish traps. Another isopod cap- tured with B. tricarinata in baited mollusk traps off Grand Bahama Island is Cirolana minuta Hansen, 1890 (see Kensley & Schotte 1987:233-236). Acknowledgments We thank William G. Lyons, Jack Wors- fold, Robert Quigley, and Steven I. Can- dileri, who collected the original series of specimens and made them available to us. Raymond B. Manning arranged for the loan of specimens housed at the National Mu- seum of Natural History, Smithsonian In- stitution. William Lyons and Thomas H. Perkins made valuable suggestions for im- provement of the manuscript, and Llyn French improved figure layout. Special thanks go to Niel L. Bruce for deferring de- scription of B. tricarinata to us and for his constructive comments on the manuscript. Literature Cited Bruce, N. L. 1981. The Cirolanidae (Crustacea: Iso- poda) of Australia: New species and a new genus from southeastern Australia.—Records of the Australian Museum 33(13):644-672, figs. 1-15. . 1985. Calyptolana hancocki, a new genus and species of marine isopod (Cirolanidae) from Aruba, Netherlands Antilles, with a synopsis of Cirolanidae known from the Caribbean and Gulf VOLUME 101, NUMBER 3 of Mexico. —Journal of Crustacean Biology 5(4): 716, figs. 1-3. 1986. Cirolanidae (Crustacea: Isopoda) of Australia.— Records of the Australian Museum, Supplement 6:1—239, figs. 1-152. Kensley, B. 1987. Further records of marine isopod crustaceans from the Caribbean.— Proceedings of the Biological Society of Washington 100(3): 559-577, figs. 1-11. —., & M. Schotte. 1987. New records of isopod Crustacea from the Caribbean, the Florida Keys, and the Bahamas.— Proceedings of the Biolog- 613 ical Society of Washington 100(1):216—247, figs. 1-21. (DKC) Florida Department of Natural Resources, Bureau of Marine Research, 100 Eighth Avenue, S.E., St. Petersburg, Florida 33701-5095; (RWH) Invertebrate Zoology Section, Gulf Coast Research Laboratory, P.O. Box 7000, Ocean Springs, Mississippi 39564. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 614-621 IPANEMIDAE, NEW FAMILY, [PANEMA TALPA, NEW GENUS AND SPECIES, FROM THE SURF ZONE OF BRAZIL (CRUSTACEA: AMPHIPODA: HAUSTORIOIDEA) J. L. Barnard and James Darwin Thomas Abstract.—A new family of haustorioid Amphipoda resembling the Phox- ocephalopsidae and Urothoidae has been discovered in sands of the surf zone off Rio de Janeiro, Brazil. Like the recently described Cheidae, the new species- genus-family combines various characters of haustorioids in a unique manner and bears one unprecedented apomorphic character: an alate, uncleft telson. The new taxon has the following characters of Urothoidae: antenna 2, head, and distal parts of pereopods but has a distinctive antenna 1, coxae 1-2, man- dibular palp, telson and epimera 1-2. Ipanema has the following characters of Phoxocephalopsidae: coxae, mandibles, and uropod | but has distinctive an- tennae 1—2, epimera 2-3, pereopods 5-7 and telson. The taxon has some char- acters found mostly or only in Urohaustoriidae, such as epimera 2-3, but differs in many ways from that family, such as: head, antennae 1-2, parts of mandibles and maxillipeds, maxilla 1, parts of pereopods 3—7, epimeron 1, and especially uropods 1-2. The Ipanemidae are created for Ipanema talpa. The unusual combination of anten- nae 1-2 with mouthparts, uropods 1—2, head shape and pereopods precludes its assign- ment to any existing family group. It bears an apomorphic telson not heretofore de- scribed from the superfamily Haustorioidea except in the otherwise remote Urohaus- toriidae. Ipanema combines characters of Phoxocephalopsidae, Urothoidae and Uro- haustoriidae. For example, antenna 2 and to a certain extent antenna | have characters of Urothoidae, whereas coxae 1-4 and uro- pod 1 are like Phoxocephalopsidae while the epimera have some characters of Uro- haustoriidae. Corrections to literature.—We have re- evaluated the cephalic cheek of Phoxoceph- alopsidae to be well developed in contrast to the opinion of Barnard & Drummond (1982). Ipanemidae, new family Type genus. — Ipanema, new genus. Gen- der feminine. Etymology.—A beach of Rio de Janeiro, Praia de Ipanema. Diagnosis. —Rostrum weak, broad; cheek strong. Antenna | of neither haustoriid nor urothoid form, peduncle short, no articles elongate, each thick, article 1 with weak se- tation, article 2 furnished with dense row of spines, article 3 about 0.67 as long as article 1, poorly armed; no geniculations present; flagellum longer than peduncle and heavily armed with aesthetascs; accessory flagellum 2-articulate. Antenna 2 of urothoid form, articles 4 and 5 slender, with long lines of spines not organized into ranks, posterior margins lacking glass-spines (typical of Uro- haustoriidae), ordinary setae and bulbar se- tae extremely sparse, flagellum very short, 3-articulate. Epistome and upper lip fused VOLUME 101, NUMBER 3 together, lower margin rounded. Right and left mandibles alike, with narrow but stubby and simple incisors lacking teeth laciniae mobiles alike, rakers absent, molar of me- dium size (compared to Urothoidae), weak- ly triturative, choppers weak to absent; palp 3-articulate, article 3 with urohaustoriid se- tal-spine distribution (versus urothoid setal distribution), spines hooked but unawned. Lower lip with mandibular lobes well de- veloped. Maxilla 1 with biarticulate palp, inner plate small and spout-like, with | large seta. Inner plate of maxilla 2 without oblique facial seta row, with one medial seta. Max- illipeds with unexpanded bases and no baler lobes, with rather small inner plate but nor- mally enlarged and spinose outer plate; palp 4-articulate, article 2 expanded, article 4 rectangular, with 2 thin nail-setae. Coxae 1-2 minute and hidden by coxa 3, coxa 4 dominant, weakly excavate behind, coxa 3 adz-shaped; coxae 5—6 with comma- shaped posterior lobe. Gnathopods feeble, subsimilar, simple (gnathopod 2 with mi- nute palm), article 3 short. Article 5 of pe- reopods 3-4 slender, not lobate; dactyls of pereopods 3-7 well developed; pereopod 5 of weakly haustorious form, article 2 ex- panded, articles 4-5 weakly expanded and with few facial spines; article 2 of pereopods 5 and 7 diverse, with that of pereopod 6 intermediate in form; no pereopod with un- derslung articulation. Gills on coxae 2-5 only, 6—7 apparently without gills because of long forward reach of beating pleopods, gill 2 (on coxa 2) spear-shaped, others de- creasing in size, subrectangular sacs. Pleopods like urohaustoriids, thus pe- duncles wider than long, pleopod 3 inferior; peduncles as long as wide. Epimeron | fully developed, small, with 1 seta, epimera 2-3 equally dominant, all epimera with pos- terodorsal “‘hip.’’ Urosomites ordinary. Rami of uropod | styliform, naked; of uro- pod 2 rod-like and spinose; uropod 3 of ordinary haustorioid-phoxocephalid kind, neither ramus dominant, article 2 on outer ramus small and poorly setose. 615 Telson very short, much wider than long, essentially entire, each side with alate lobe projecting upward. Relationship. — Differing from the Phox- ocephalopsidae in the completely distinc- tive antennae 1-2, of which antenna 1 has a heavily spinose (versus setose) article 2 and the accessory flagellum is poorly de- veloped; of which antenna 2 has slender ar- ticles 4-5 bearing large facial spines in lines rather than small spines in ranks and on which the posterior margins have almost no long setae, almost no bulbar setae and no glass spines. Also differing from Phoxo- cephalopsidae in the thin article 5 of pe- reopods 3-4; the diversity of article 2 on pereopods 5-7, with article 2 on pereopod 5 constricted apically; thin apical articles of pereopods 5-7; the presence of a long com- ma-shaped posterior lobe on coxae 5-6; the equal size of epimera 2-3 and the hips on epimera 1-3; the strange telson (versus or- dinary and cleft); and the reduced inner plate of maxilla 1 bearing a single seta. Differing from Urothoidae in the short articles of the peduncle on antenna 1, with article 2 heavily spinose, the long primary flagellum of antenna 1; the kind and distri- bution of setal-spines on article 3 of the mandibular palp (versus regular setae and presence of an A-seta in Urothoidae); the tiny coxae 1-2; the hipped shapes of epim- era 1—3; and the more delicate mandible. Phoxocephalopsidae and Urothoidae ap- pear to be more closely related than pre- viously perceived: both have a well devel- oped lateral cephalic cheek, styliform rami on uropods 1-2 and many similarities in mandibles, maxillae, and maxillipeds. Phoxocephalopsids differ from urothoids principally in the antennae, in which articles 1-3 of antenna | are short, not geniculate, with huge setal patch on article 2; article 4 of antenna 2 is widely expanded and bears 3 kinds of posterior armaments, including glass spines; article 5 also is expanded and bears 2 kinds of armaments posteriorly; the mandibular palp bears hooked setal-spines PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 616 ‘ \ ©) ) Serr >, iN BY °K HE VOLUME 101, NUMBER 3 and the dactyl of the maxilliped is less elon- gate and less unguiform. Differing from the Cheidae in the large molar and lack of significant rostrum, nor- mal uropod 2, untoothed incisors and sim- ple gnathopods. Differing from the complex Pontoporei- idae in the shortened peduncles of the pleo- pods, the short article 1 of antenna | and the lack of long setae (distinct from spines) on pereopods 5-7; also differing from all pontoporeiids except Priscillina in the spi- nose antenna 2. Differing from the Haustoriidae in the 4-articulate (versus 3-) palp of the maxil- liped, the presence of only spines (no flex- ible setae) on pereopods 5-7, the hidden coxae 1-2, large mandibular lobes of the lower lip, the presence of only a few stiff spines (no long flexible setae) on mandib- ular palp article 3, the unexpanded article 4 of antenna 2, the poor setation on the inner plates of maxillae 1-2 and the ordi- nary, non-enlarged outer plate of maxilla 2. Ipanema, new genus Type species. —Ipanema talpa, n. sp. Diagnosis. —With the characters of the family. Ipanema talpa, new species Figs. 1-4 Etymology. —L., talpa, mole. Description (of holotype male “‘s” 2.57 mm).—Rostrum obsolescent; eyes medi- um, ommatidia few, interspersed in alcohol with faint purplish pigment, each with an- teromedial ovate ganglion. Antenna | ar- ticle 1 with sparse penicillate setules, article 3 almost naked, accessory flagellum 3-ar- —_ Fig. 1. 617 ticulate, primary flagellum long, with 8 ar- ticles, formula of aesthetascs on articles 1— 8 = 2-6-5-4-4-1-1-x (vestigial). Article 4 of antenna 2 with anterolateral line of 14 spines mostly alternating long and short spines pointing in alternate directions, with 3 an- terodistal spines; posterior margin and face with 3-4 bulbar setules, posterodistal corner with line of 4 spines and seta and 2 marginal setae; article 5 with anterolateral line of 8 spines mixed long and short, posterior mar- gin mostly naked, posterodistal corner and apex with 5 long setae, 2 bulbar setae; fla- gellum with 2 articles. Upper lip and epistome forming large subcircle from anterior view. Right and left mandibles identical, incisor extended, mod- erately narrow, simple; lacinia mobilis lin- guiform; rakers absent; molar large, weakly triturative, with weak chopper region, no seta; mandible organized so that when pressed into flat plane palp extending in odd direction (see illustration of labral-mandib- ular complex), article 1 short, article 2 with 1—2 small midlateral setae, article 3 with 5 mixed-size apical spines in haustoriid for- mula of 2-2-1. Lower lip without cones. Maxilla 1 with small spout-like inner plate bearing one long seta, outer plate with 9 simple spines, palp feeble, 2-articulate, slen- der, reaching less than two-third along outer plate, with 2 huge apical setae. Inner plate of maxilla 2 narrower and shorter than outer plate, without oblique facial seta row, with one medial seta, outer plate with 2 apico- lateral setae. Inner plate of maxilliped rather small and very broad (relative to other haus- torioids), with one medial seta, one apical seta and 2 penicillate spines; outer plate spi- nose medially, naked apically; palp huge, article 2 expanded and medially setose, ar- Ipanema talpa, holotype male “‘s” 2.57 mm. Capital letters in figures refer to parts; lower case letters to left of capital letters refer to specimens and to the right refer to adjectives as described below: A, antenna; B, body; C, coxa; E, epimeron; G, gnathopod; H, head; J, prebuccal; L, labium; M, mandible; P, pereopod; Q, spine; R, uropod; S, maxilliped; T, telson; U, upper lip; V, palp; W, pleon; a, anterior; d, dorsal, t, left. 618 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Ipanema talpa, unattributed figures = holotype male ‘“‘s” 2.57 mm; p = male “p” 2.98 mm. Letter codes, see Fig. 1. VOLUME 101, NUMBER 3 619 Fig. 3. Ipanema talpa, unattributed figures = holotype male “s” 2.57 mm; p = male “‘p” 2.98 mm. Letter codes, see Fig. 1. 620 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ny” AU \\ QVM Wt Fig. 4. Ipanema talpa, holotype male “‘s’’ 2.57 mm. Letter codes, see Fig. 1. ticle 3 curved and fitting hump of article 2, dactyl not unguiform but not stubby, rect- angular, armed with 2 huge apical setal- spines. Coxae 1-2 very small, hidden by coxa 3, coxa 1 extended forward into point armed with stiff setal-spine. Article 2 of gnatho- pods 1-2 and pereopods 3—4 poorly setose; article 5 of gnathopod 1 broader and more strongly setose than on gnathopod 2, gnathopod 1 simple, gnathopod 2 scarcely subchelate, with shorter dactyl than gnatho- pod 1, with minute non-skid scales on prop- odus (see enlarged illustration, Fig. 1). Pereopods 3-4 alike, article 4 with naked margins, one long seta at each apex, article 5 narrow but with weak posterodistal lobe armed with 3 spines; many spines of all pe- reopods with lines of bosses and apical bi- fidation formed by enlarged boss, some multibossed at apex; article 6 slender, spine formula = 3-3-1; dactyl long and straight, with inner boss near apex. Article 2 of pe- reopod 5 grotesque, expanded and lobate proximally, tapering sharply at apex, of pe- reopod 7 almost evenly ovatorectangular and bearing about 7 long posterior setae, of pereopod 6 intermediate between 5 and 7; dactyls elongate, like those of pereopods 3- 4. Pereonites 6-7 and pleonites 1-3 with lateral ridge. Epimera 1-3 similar, with hipped posterodistal corner, weakly exca- vate posterior margin bearing setule, round- ed ventral margin with 2 long facial setae VOLUME 101, NUMBER 3 on epimera 2-3, epimeron 2 also with mid- posterior facial seta. Pleopods with 2 cou- pling hooks, outer rami about 1.5 times as long as inner, articles on outer rami = 9-8- 7, on inner rami = 6-6-5. Urosome ordinary. Uropod 1 with short peduncle bearing ventrolateral spine, ven- trodistal spine, distomedial spine, rami sty- liform and naked, outer very long and ex- ceeding apex of uropod 2, inner much shortened. Uropod 2 with short peduncle and apical spine on each side; rami rod-like, outer with one apical and one apicolateral spine, inner with 2 apical and one apico- medial spine. Uropod 3 with short peduncle bearing 2 apical spines on each side, rami elongate, inner almost as long as outer, well setose, outer with small second article bear- ing 2 long setae. Telson 2.4 times as wide as long, with lateral wings bearing 3 setules each. Female.—None. All specimens are males, with large penial processes. Illustrations. —Mandible and maxilla 1 enlarged equally. Holotype. -USNM 195181, male “‘s”’ 2.57 mm. Type locality.—Brazil, Rio de Janeiro, Praia Vermelha, Rio Orca Beach, 7 May 1985, coarse sand, 4 m, with numerous small echinoids and platyischnopid amphipods, coll. J. D. Thomas. 621 Material.—From type locality, USNM 196184, male “‘p’”’ 2.98 mm, male “‘q” 2.95 mm, male “‘r’’ 2.10 mm and 4 other males. Distribution. —Brazil, Rio de Janeiro, outer surf zone, 4 m. Acknowledgments We thank Dr. David Challinor, Assistant Secretary for Science of the Smithsonian In- stitution, for providing funds to travel to Brazil. The second author also was sup- ported by NSF Grant BSR-8515168. We thank Linda B. Lutz of Vicksburg, Missis- sippi for inking our illustrations. Janice Clark assisted us in the laboratory. Dr. Glo- ria Alonso kindly offered us valuable sug- gestions for improvements. Literature Cited Barnard, J. L., & M. M. Drummond. 1982. Gam- maridean Amphipoda of Australia, Part V: Su- perfamily Haustorioidea.—Smithsonian Con- tributions to Zoology 360:1-148. (JLB) Department of Invertebrate Zool- ogy, NHB-163, Smithsonian Institution, Washington, D.C. 20560; (JDT) Reef Foun- dation, P.O. Box 569, Big Pine Key, Florida 33043. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 622-625 NEW RECORDS FOR OGYRIDES ALPHAEROSTRIS AND A NEW SPECIES, OGYRIDES TARAZONAI (CRUSTACEA: OGYRIDIDAE), FROM THE EASTERN PACIFIC OCEAN Mary K. Wicksten and Matilde Mendez G. Abstract.—A new species, Ogyrides tarazonai, related to O. hayi, is described from Nicaragua and Peru. This shallow-water species has only one spine on the dorsal midline of the carapace. Specimens of O. alphaerostris (Kingsley) have been found off southern California and western Mexico. Species of Ogyrides (Caridea: Ogyrididae) are burrowing shrimps of warm-temperate and tropical waters. In the eastern Pacific, the genus is reported from specimens taken off western Mexico (Wicksten 1983, Car- vacho & Olson 1984). Examination of ma- terial from California, Mexico, Nicaragua, and Peru led to the determination that two species are present in the eastern Pacific. The specimens discussed herein are depos- ited in the collections of the Allan Hancock Foundation, University of Southern Cali- fornia (AHF), the California Academy of Sciences (CAS), the Instituto del Mar del Peru (IMARPE), and the Estacion Mazatlan (EMU). We thank Michel Hendrickx, Es- tacion Mazatlan, for allowing us to examine specimens from off Sinaloa. The illustra- tions are by Debbie Meier, Texas A&M University. Ogyrides tarazonai, new species Fig. 1 Description. —Rostrum triangular, acute, setose; slightly longer than width at base and longer than extracorneal teeth. Extracorneal teeth rounded, infracorneal teeth shorter than extracorneal but rectangular. Ptery- gostomial angle projected but rounded. Car- apace carinate along midline posterior to rostrum, setose, bearing | strong movable spine directed forward. Eyestalks reaching beyond end of anten- nular peduncles, thickened at base, curving slightly mesad toward narrowed midlength, then broadening distally toward slightly ex- panded but small terminal cornea. First an- tennular article about equal to second when measured from base, second article 2 as long as broad, third 0.5 x second. Styloce- rite with 2 strong spines, superior spine al- most reaching end of first segment of an- tennular peduncle, inferior spine surpassing it. Squamous part of scaphocerite broad, lanceolate, with tiny lateral spine, exceeding second segment of antennular peduncle. Carpocerite as long as or slightly surpassing antennular peduncle. Distoinferior margin of basicerite bearing 2 small acute spines. Third maxilliped exceeding antennules. Ratio of article lengths 10:6:2. First chelipeds 0.4 x length of third max- illipeds, symmetrical. Ischium 0.6 x merus, bearing rounded protrusion on inferior margin. Merus 3x long as broad. Carpus 4x long as broad distally, with distal end 1.8 x as broad as proximal and bearing 1 stout spine or knob on lateral margin. Chela about as broad as carpus, but not as long, with fingers 0.7 x length of palm. Carpal articles of second legs 4, with length ratio of 10:4:3:4 progressing distally. Third legs with ischium unarmed, shorter than merus. Merus 3x as long as broad, bearing large spine subterminally on infe- rior margin. Carpus 0.7 X merus, broadened VOLUME 101, NUMBER 3 623 imm Fig. 1. Ogyrides tarazonai, paratype, female: A, Lateral view of front; b, Telson and uropods; c, Frontal region in dorsal view; d, Fourth pereopod; e, First pereopod; f, Second pleopod; g, Fifth pereopod; h, Third pereopod. Third maxillipeds, second pereopods and thelycum-like structure broken or missing in this specimen. distally and bearing many long setiferous bristles. Propodus stout, 1.7 x long as broad, with setose margin, bases of setae forming serrate edge. Dactyl spatulate, shorter than propodus. Fourth legs slender, ischium 0.6 x merus. Merus 5 x as long as broad, bearing long setae. Carpus 0.6 X merus. Propodus shorter than carpus, tapered distally. Both carpus and propodus bearing long setae. Dactyl curved, spatulate, 0.5 propodus. Fifth legs very slender, ischium about equal to merus, merus 6 X as long as broad, carpus 624 shorter than propodus, propodus and dactyl of approximately equal length. Propodus and dactyl setose, dactyl spatulate. Thelycum-like structure of female narrow and elongate, lying between coxae of fourth legs and ventral to sternal plates, anteriorly reaching bases of third legs, anterior margin with V-shaped cleft, lateral margins nearly straight, posterior margins apparently at- tached to coxae of legs and to sternum. Cox- ae of fifth legs bearing lobes joining as low, continuous process reaching from leg to leg. Second pleopod with appendix interna on endopod. Telson 1.8 as long as broad, tip sub- acute, part distal to posterolateral spines representing 0.4 of total telson length. Lat- eral margins bearing broad, low rounded projection just anterior to middle. Outer spines of posterolateral pair short, inner spines longer. Low ridges on dorsal surface of telson. Two pairs of dorsal spines, difh- cult to see, well separated. Both uropod rami exceeding telson. Type material. —Holotype: female, total length in millimeters 19.4; Ventanilla (11°50’S), Peru, shallow water, 40 m from high tide line, 18 Mar 1984, Juan Tarazona, IMARPE.— Paratype: female, ovigerous, total length 27.7; about 15 km S of San Juan del Sur, Pacific coast of Nicaragua, 27 Jan 1974, A. J. Ferreira, CAS 044016. Remarks.—Only one other species of Ogyrides has only a single dorsal spine on the carapace. Ogyrides hayi Williams, from the western Atlantic, shares this feature with O. tarazonai, as well as also having four segments in the carpus of the second pereo- pod. However, O. hayi has a spine on the ischium of the third leg. The spines on its stylocerite do not reach or surpass the first segment of the antennular peduncle. Of the two specimens of the new species, the holotype is more intact. We deposited the holotype at the Instituto del Mar del Peru, hoping to illustrate it later. Unfortu- nately, neither of us was able to return to Callao to draw this specimen, nor did we PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON trust the postal service to send the holotype to us safely. We therefore illustrated what we could from the paratype, which is broken and damaged on the ventral surface. The description in the text is of the holotype. Etymology. —The specific epithet honors Juan Tarazona, collector of the new species and dedicated benthic ecologist. Ogyrides alphaerostris (Kingsley) Ogyris alphaerostris Kingsley, 1880:420, pl. 14, fig. 7. Ogyrides alphaerostris.—Williams, 1981: 144; 1984:107, fig. 74.—Carvacho & Ol- son, 1984:66, figs. 3-4. (See Williams, 1981, for a fuller synonymy.) Material. —Off Huntington Beach, Cali- fornia (33°38'30”N, 118°03’00”W), 26-28 m, sand, 20 Apr 1940, Velero III sta 1127- 40, 1 specimen, AHF. Remarks. —The specimen from off Hun- tington Beach agrees with the description of O. alphaerostris as given by Williams (1984). The number of dorsal spines on the cara- pace of this species is variable, ranging from three to 14. The specimen from California bears nine carapace spines. We have been unable to reexamine the specimens of Ogyrides mentioned in the monograph by Wicksten (1983). These probably belong to O. alphaerostris, which we have identified in grab samples from off the coast of Sinaloa, Mexico. The latter rec- ords will be published elsewhere in a report on benthic decapod crustaceans of Sinaloa (M. E. Hendrickx, pers. comm.). Literature Cited Carvacho, A., & R. Olson. 1984. Nuevos registros para la fauna carcinologica del noreste de Méx- ico y descripcion de una nueva especie: Eualus subtilis n. sp. (Crustacea: Decapoda: Natan- tia). Southwestern Naturalist 29(1):59-71. Kingsley, J. S. 1880. On a collection of Crustacea from Virginia, North Carolina, and Florida, with a revision of the genera of Crangonidae and Pa- laemonidae.— Proceedings of the Academy of Natural Sciences of Philadelphia 31:383-—427. VOLUME 101, NUMBER 3 Wicksten, M. K. 1983. A monograph on the shallow water caridean shrimps of the Gulf of California, Mexico.— Allan Hancock Monographs in Ma- rine Biology 13:1-59. Williams, A. B. 1981. Western Atlantic shrimps of the caridean shrimp genus Ogyrides.—Journal of Crustacean Biology 1(1):143-147. 1984. Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine 625 to Florida. Washington, D.C., Smithsonian In- stitution Press, 550 pp. (MKW) Department of Biology, Texas A&M University, College Station, Texas 77843; (MMG) Fondo de Promocion de Ex- portaciones no Tradicionales, Casilla Postal 205, Tumbes, Peru. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 626-632 NOTES ON ALBUNEID CRABS (CRUSTACEA: DECAPODA: ALBUNEIDAE) FROM THE CENTRAL EAST COAST OF FLORIDA Raymond B. Manning Abstract.—Three species are reported: Albunea paretii Guérin-Meéneville, 1853, Lepidopa benedicti Schmitt, 1935, and L. websteri Benedict, 1903. The latter species is recorded from southeast Florida for the first time. Study material includes rare males of both species of Lepidopa. A key to western Atlantic albuneids is presented. Until now, one species of the albuneid genus Lepidopa Stimpson, 1858, L. bene- dicti Schmitt, 1935, had been reported from the southeast coast of Florida (Holthuis 1960, Efford 1971). Of the six Atlantic species of the genus, only two, L. benedicti and L. websteri Benedict, 1903, had been reported from localities in Florida (Efford 1971, Gore et al. 1978, Abele & Kim 1986), although a larva, tentatively identified with a third species, L. richmondi Benedict, 1903, was taken in the plankton east of Jupiter Inlet, Martin County, on the Atlantic coast of Florida (Gore & Van Dover 1981). In 1982 and 1983, collections made with a yabby pump in the vicinity of Fort Pierce, Florida, yielded material of both L. bene- dicti and L. websteri at one locality on the Atlantic coast just north of the St. Lucie Inlet (St. Lucie site of Gore et al. 1978). There are few records for members of A/- bunea from the central east coast of Florida, although this is well within the known range of the two western Atlantic species. Material available for study included a specimen from the Indian River at Fort Pierce. East American Species of Albuneidae The east American albuneids comprise ten species in four genera: A/bunea Weber, 1795 (two species), Blepharipoda Randall, 1840 (one species), Lepidopa Stimpson, 1858 (six species), and Zygopa Holthuis, 1960 (one species). In all, the family now includes seven genera; Efford & Haig (1968) provided a key to the genera. The two species of A/bunea Weber (1795: 94), A. gibbesii Stimpson (1859:78) and A. paretii Guérin-Meéneville (1853:48), were treated by Williams (1984) and Abele & Kim (1986). Both have wide ranges in the Amer- icas, from the Carolinas to Brazil, and A. paretii also occurs off West Africa. The single east American species of Ble- pharipoda Randall (1840:130), B. doelloi Schmitt (1942:2), is known only from Ar- gentina. Efford (1971) reviewed the American species of Lepidopa Stimpson (1858:230), and recorded five species from the Atlantic coast of the Americas. Since then a sixth east American species, L. dexterae Abele & Efford, 1972, has been described from Pan- ama. Efford (1971:99) recognized four species groups in the genus: Myops, bene- dicti, venusta, and californica. The first three of these are represented in the western At- lantic. The following species of Lepidopa are known from the western Atlantic: L. benedicti Schmitt (1935:210); south- eastern and northwestern Florida to south- east of Veracruz, Mexico, in the Gulf of Mexico. L. dexterae Abele & Efford (1972:502); Panama. VOLUME 101, NUMBER 3 L. distincta Corréa (1968:77); Dominican Republic and Brazil. L. richmondi Benedict (1903:895); West Indies and Caribbean Sea to Brazil. L. venusta Stimpson (1859:79); West In- dies to Brazil. L. websteri Benedict (1903:892); Atlantic coast of southeastern United States. The genus Zygopa was established by Holthuis (1960:21) for Z. michaelis Hol- thuis (1960:22), from Curacao. Since its original description, Z. michaelis has been recorded from off Palm Beach and Miami on the east coast of Florida (Gore & Becker 1977), and from the Florida west coast (Reames et al. 1982). The east American albuneids can be dis- tinguished by means of the following key. 1. Carapace with 4 lateral spines [Ble- pharipoda| B. doelloi Schmitt — Carapace with | lateral spine .... 2 2. Carapace lacking median spine or angled projection on anterior mar- gin. Eyes very small, fused together (Zyzopal gee Z. michaelis Holthuis — Carapace with median spine or an- gled projection on anterior margin. EVeSiseparaten guy. 7ii se nuke 3 3. Eyes narrow, elongate, with distinct cornea. Lateral spine of carapace be- low linea anomurica [4/bunea] ... 4 — Eyes (eye plates) broad, oval or squarish, lacking distinct cornea. Lateral spine of carapace above lin- ea anomurica [Lepidopa] 4. Dactyli of second and third pereo- pods with blunt lobe basally on an- terior border .... A. gibbesii Stimpson — Dactylus of second pereopod with broad spur, of third pereopod with acute, slender spur at base of ante- PLO TADOLG CIB: ius ae craigs ck AE ETS PUN Fo. A. paretii Guérin-Meéneville 5. Eye plates distinctly squarish, flat- tened anteriorly, anterior edge with teeth or denticles [benedicti Group] — Eye plates round or oval, anterior edge smooth 6. Groove along posterior edge of car- apace continuous, not interrupted in middle of posterior margin [sub- rostral spine present] ............ Be ate a er L. richmondi Benedict — Groove along posterior edge of car- apace interrupted medially ....... 7 7. Subrostral spine present ......... De ae aS: L. dexterae Abele & Efford — Subrostral spine absent .......... L. benedicti Schmitt 8. Antennal flagellum with 7 articles. Carpus of third maxilliped extend- ing Over propodus, overlapping dac- tylus. Subrostral spine absent [myops Group]. ce. L. distincta Corréa — Antennal flagellum with 8 articles. Carpus of third maxilliped not ex- tending to dactylus. Subrostral spine present [venusta Group] 9. Width of emargination in edge of second abdominal somite no greater than length of this somite along midline. Anterior margin of cara- pace extending in almost straight line between base of orbit and anterolat- eral projection ... L. websteri Benedict — Width of emargination in anterior edge of second abdominal somite greater than length of this somite along midline. Anterior margin of carapace with distinct anterior bulge between base of orbit and anterolat- eral projection ... L. venusta Stimpson ey Almost all of the material reported here was taken at the same site (St. Lucie, At- lantic, of Manning collections; see Manning & Felder, in press). Locality data are as fol- lows: Atlantic Ocean, Florida, Martin County, about | mile north of St. Lucie In- let, 27°10.9'N, 80°09.5'W, shallow sand flat between emergent sabellariid reef and sand beach. Most of the specimens were taken with a yabby pump, a suction device used to sample infaunal organisms; some were 628 Fig. 1. mm, sta FP-82-2: Carapace. Lepidopa benedicti Schmitt, female, cl 5.0 taken with a garden rake, used on sand at the edge of the wave line. The lepidopas were disturbed by the rake and could be seen re-burying themselves. Specimens have been deposited in the In- dian River Coastal Zone Museum, Harbor Branch Oceanographic Institution, Fort Pierce, Florida (IRCZM), and the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). Albunea paretii Guérin-Meéneville, 1853 Albunea paretii. —Gore & Becker, 1977:220 [no material].— Young, 1978:177 [South Carolina].— Williams, 1984:249, fig. 183 [original citation and other references giv- en here].— Fox & Ruppert, 1985:52, 122, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 128, 259, 286 [South Carolina].— Abele & Kim, 1986:x, 38, 427, 428, 748, figs. d-g on p. 429 [Florida].—Chace et al., 1986:338, fig. on pl. 112 [Bermuda]. Albunea paretti.—Pequegnat, 1975:45 [no localities]. Material. — Florida, St. Lucie County, In- dian River, island west of Coon Island, Fort Pierce Inlet; intertidal, with yabby pump; 6 Mar 1985; W. D. Lee and S. Petry; 1 male (USNM). Size. —Carapace length of male 10.9 mm. Remarks.—There are other representa- tives of this species from off Jacksonville Beach, off Daytona Beach, and off Cape Canaveral in the collections of the National Museum of Natural History. The species has a wide range in the Atlantic, from North Carolina to Texas, the West Indies to Brazil, and off West Africa (Holthuis & Manning 1970, Williams 1984). Lepidopa benedicti Schmitt, 1935 Fig. 1 Lepidopa scutellata.—Benedict, 1903:891, 894, fig. 6 [Pensacola and Morris Cut, Miami; not L. scutellata Fabricius, 1793]. Lepidopa benedicti Schmitt, 1935:210 [Re- placement name for L. scutellata sensu Benedict, 1903].—Holthuis, 1960:31, 32, 33, 34, 35, fig. 5 [several records, from east coast of Florida to southwest coast of Texas; lectotype selected].—Efford, 1971:76, figs. la, 2e, 3a, 4e, p, q, 5a, 6e, n, 7a [Florida to Mexico].—Gore & Beck- er, 1977:220 [no material].—Gore et al.,1978:231, 247 [central east Florida]. — Gore & Van Dover, 1981:1016, 1030, 1033 [southeastern and central eastern Florida; no material].—Abele & Kim, 1986:x, 38, 427, 428, 752, fig. h on p. 429 [Florida]. Material. —RBM stations: FP-82-2, 13 Jul 1982, R. B Manning and C. W. Hart, Jr., yabby pump, | female. FP-82-7, 16 Jul 1982, R. B. Manning, yabby pump, | male [taken VOLUME 101, NUMBER 3 together with L. websteri]. FP-83-6, 13 Jul 1983, R. B. and L. K. Manning, W. D. Lee, rake, 1 male, 2 females [taken together with L. websteri]. All specimens USNM. Vero Beach, 100 meters south of South Beach Park, along shore, in colony of Emer- ita, 4 Mar 1973, 1 female (IRCZM 89: 514).—Fort Pierce Inlet, St. Lucie County, worm reef, 6 Feb 1974, 1 female (IRCZM 89:2558).— Walton Rocks, St. Lucie Coun- ty, worm reef, LES, LB, MGR leg., 19 Jul 1974, 1 female IRCZM 89:2604). Size. —Carapace lengths of 2 males, 3 mm and 3.5 mm, of 6 females, 4.5 to 17 mm. Efford (1971) reported that the largest spec- imen known was a female 22 mm long; he studied only 2 males, the largest measuring 4 mm. Remarks. —Eye pigment is present in all of these specimens, two lots of which were taken together with L. websteri. Apparently these specimens are from the northernmost locality known for this species on the east coast of Florida, and they dem- onstrate that the species is fairly common off the Florida east coast. The specimens in the collections of the Indian River Coastal Zone Museum may have been the basis for the comment by Gore & Van Dover (1981:1016), ““The sec- ond continental species, L. benedicti Schmitt, 1935, occurs in the Gulf of Mexico and along the southeastern and central east- ern Florida coastline (... Gore, unpub- lished).” Lepidopa websteri Benedict, 1903 Figs. 2, 3 Lepidopa websteri Benedict, 1903:891, 892, fig. 3 [Fort Macon, North Carolina]. —Ef- ford, 1971:91, figs. 1b, k, 2j, 3r, 4h, m, 5g, 1, 60 [North Carolina, South Carolina, west Florida, Mississippi].—Gore & Becker, 1977:220 [no material].— Young, 1978:177 [South Carolina].— Williams, 1984:250, fig. 184 [additional records from Sapelo Island, Georgia, and Petit 629 Fig. 2. Lepidopa websteri Benedict, female, cl 5.0 mm, sta FP-83-3: a, Carapace; b, Second abdominal somite. Bois Island, Mississippi].—Fox & Rup- pert, 1985:39, 114, 122, 129, 187, 196, 259, 286 [South Carolina].—Abele & Kim, 1986: x, 38, 427, 428, 752, fig. 1 on p. 429 [Florida]. Material.— RBM stations: FP-82-7, 16 Jul 1982, R. B. Manning, yabby pump, | female [taken with L. benedicti]. FP-83-3, 8 Jul 1983, R. B.and L. K. Manning, yabby pump, 630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Lepidopa websteri Benedict, female, cl 9.0 mm, sta FP-84-4: Carapace. 1 female. FP-83-6, 13 Jul 1983, R. B. and L. K. Manning, W. D. Lee, rake, 1 male [taken with L. benedicti]. FP-84-4, 10 Jul 1984, R. B. and L. K. Manning, D. L. Feld- er, and W. D. Lee, yabby pump, | female. Specimen from FP-82-7 deposited in IRCZM, remainder deposited in USNM. Size. —Carapace length of male, 4 mm, of 3 females, 5 to 9 mm. Williams (1984) reported a specimen 19 mm long, and Ef- ford (1971) reported a male 6 mm long. Remarks. —The male and the smallest fe- male (Fig. 2) differ from the two larger spec- imens (Fig. 3) in having a distinct elongate patch of dark pigment in each eye; this pig- ment is completely absent in larger speci- mens of this species. Efford (1971:61) noted that eye pigment was always present in ju- veniles. Until now this species was not known south of Sapelo Island, Georgia, on the At- lantic coast of the United States (Williams 1984). Like some other Carolinian species generally considered to be restricted to the Carolinas and Georgia on the Atlantic coast, b Fig. 4. Lepidopa venusta Stimpson, female, cl 6.5 mm, Trinidad (USNM 141353): Carapace. e.g., Callichirus major (Say, 1818) (see Man- ning & Felder 1986) and Pinnixa cristata Rathbun, 1900 (see Manning & Felder, in press), this species appears to be not un- common in the central Florida east coast. Efford (1971:91) noted that “‘the easiest character separating venusta from websteri” is the presence in L. venusta of a bulge or obtuse projection on the anterior margin of the carapace, between the ocular sinus and the anterolateral spine. This is a subtle but VOLUME 101, NUMBER 3 distinctive character, and as I can find no published figure of the carapace of L. ve- nusta in which this feature is well illustrat- ed, I provide here a figure of L. venusta (Fig. 4). Curiously, Efford (1971), in his revision of Lepidopa, gave no figure of the carapace of L. venusta. The only recent figure of the carapace of this species was given by Gomes (1965); in it the bulges are not well marked. Acknowledgments This study was carried out at the Smith- sonian Marine Station at Link Port, Fort Pierce, Florida, and is contribution number 199 from that laboratory. The help of W. D. Lee in the field is gratefully acknowl- edged. I thank Paula Mikkelsen for allowing me to study material of Lepidopa in the collection of the Indian River Coastal Zone Museum, Harbor Branch Oceanographic Institution, and Richard Heard, Gulf Coast Research Laboratory, for reading a draft of the manuscript. The figures were prepared by my wife Lilly. Literature Cited Abele, L. G., & I. E. Efford. 1972. A new species of Lepidopa, L. dexterae, (Anomura, Albuneidae), from the Caribbean coast of Panama.—Pro- ceedings of the Biological Society of Washington 84:501-506. —, & W. Kim. 1986. An illustrated guide to the marine decapod crustaceans of Florida. —Flor- ida Department of Environmental Regulation, Technical Series 8(1), parts 1&2, 760 pp. Benedict, J. E. 1903. Revision of the Crustacea of the genus Lepidopa.—Proceedings of the United States National Museum 26(1337):889-895. Chace, F. A., Jr., J. J. McDermott, P. A. McLaughlin, & R. B. Manning. 1986. Order Decapoda (shrimps, lobsters and crabs). Pp. 312-358 in W. Sterrer, ed., Marine fauna and flora of Ber- muda. John Wiley & Sons, New York. Corréa, M. M. G. 1968. Descricao de uma espécie nova do género “‘Lepidopa” Stimpson, e sua ocorréncia no litoral brasileiro (Decapoda, Al- buneidae).— Revista Brasileira de Biologia 28(1): 77-86. Efford, I. E. 1971. The species of sand crabs in the genus Lepidopa (Decapoda: Albuneidae).— Zoologischer Anzeiger 186(1/2):59-102. 631 ——,, & J. Haig. 1968. Two new genera and three new species of crabs (Decapoda: Anomura: Al- buneidae) from Australia.— Australian Journal of Zoology 16:897-914. Fox, R. S., & E. E. Ruppert. 1985. Shallow-water marine benthic macroinvertebrates of South Carolina: Species identification, community composition and symbiotic associations. — The Belle W. Baruch Library in Marine Science 14: 330 pp. Gomes, M. M. de Almeida. 1965. Redescricaéo de ‘‘Lepidopa venusta’”’ (Stimpson) e sua ocorrén- cia no litoral sul do Brasil (Decapoda, Albu- neidae).— Revista Brasileira de Biologia 25(1): 97-103. Gore, R. H., & L. J. Becker. 1977. Zygopa michaelis Holthuis, 1960: A first record and range exten- sion to the continental United States (Decapoda Anomura, Albuneidae).—Crustaceana 33(2): 219-221, pl. 1. , L. E. Scotto, & L. J. Becker. 1978. Com- munity composition, stability, and trophic par- titioning in decapod crustaceans inhabiting some subtropical sabellariid worm reefs. Studies on decapod Crustacea from the Indian River region of Florida, IV.— Bulletin of Marine Science 28(2): 221-248. —., & C. L. Van Dover. 1981. Larval develop- ment in the laboratory of Lepidopa richmondi Benedict, 1903, with notes on larvae of Amer- ican species in the genus (Anomura: Albunei- dae). 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[In press.] The Pinnixa cristata complex in the western Atlantic, with descriptions of two 632 new species (Crustacea, Decapoda, Pinnother- idae).—Smithsonian Contributions to Zoology. Pequegnat, L. H. 1975. List of catalogued inverte- brate species in the Texas A&M Systematic Col- lection of Marine Organisms. Mimeographed. Texas A&M University. 109 pp. Randall, J. W. 1840. Catalogue of the Crustacea brought by Thomas Nuttall and J. K. Town- send, from the west coast of North America and the Sandwich Islands, with descriptions of such species as are apparently new, among which are included several of different localities, previ- ously existing in the collection of the Acade- my.—Journal of the Academy of Natural Sci- ences of Philadelphia 8:106-147, pl. 3-7. Reames, R. C., R. W. Heard, & T. S. Hopkins. 1982. Records of Zygopa michaelis Holthuis, 1960 (Decapoda: Anomura: Albuneidae) from the Gulf of Mexico.—Gulf Research Reports 7(2): 171-173. Schmitt, W. L. 1935. Crustacea Macrura and Ano- mura of Porto Rico and the Virgin Islands. — Scientific Survey of Porto Rico and the Virgin Islands 15(2):125—227. 1942. A new species of sand bug, Blephari- poda doelloi, from Argentina.—Smithsonian Miscellaneous Collections 101(18):1-10, pl. 1. Stimpson, W. 1858. Crustacea Anomura. Prodromus desriptionis animalium evertabratorum, quae PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON in Expeditione ad Oceanum Pacificam Septen- tronialem, a Republica Federata missa, Cad- waladaro Ringgold et Johanne Rodgers Duci- bus, observavit et descripsit.—Proceedings of the Academy of Natural Sciences, Philadelphia 1858:225—252. 1859. Notes on North American Crustacea, I.—Annals of the Lyceum of Natural History, New York 7:49-93, pl. 1 [pp. 1-47 on separate]. Weber, F. 1795. Nomenclator entomologicus secun- dum Entomologiam Systematicam ill. Fabricii adjectis speciebus recens detectis et varietatibus. vili + 171 pp. Chilonii et Hamburgii. 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. xviii + 550 pp. Young, A. M. 1978. Superorder Eucarida, Order De- ’ capoda. Pp. 171-185 in R. G. Zingmark, ed., An annotated checklist of the biota of the coastal zone of South Carolina. University of South Carolina Press, Columbia. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 633-639 HETEROCARPUS CUTRESST, NEW SPECIES, AND PLESIONIKA MACROPODA CHACE, 1939: TWO CARIDEAN SHRIMPS OF THE FAMILY PANDALIDAE (CRUSTACEA: DECAPODA) FROM PUERTO RICO AND THE U.S. VIRGIN ISLANDS Oscar E. Monterrosa Abstract.—Two species of deep-water (320-777 m) pandalid shrimps (Crus- tacea: Decapoda: Caridea) are reported. Previously undescribed, Heterocarpus cutressi, n. sp., has two longitudinal lateral carinae with the dorsalmost origi- nating anteriorly at the antennal spine; the abdomen has a blunt median ridge on the third, fourth, and fifth somites. Plesionika macropoda Chace, 1939 has the dorsal surface of the rostrum smooth except for eight to 10 teeth on the basal crest; the dactyls of the walking legs are armed with five teeth along the posterior margin. Specimens of both species were collected in baited, wire- mesh traps. In this paper, a prelude to a more com- plete report on deep-water shrimps collect- ed in the waters around Puerto Rico and the U.S. Virgin Islands, I describe one species of pandalid and redescribe another. The former is in the genus Heterocarpus A. Milne-Edwards, 1881, consisting of over 20 species and subspecies and characterized by a dorsal and at least one longitudinal lateral carinae (one exception) and with the second pair of pereopods unequal and dissimilar. The other belongs to the genus Plesionika Bate, 1888, with over 60 species, and char- acterized by a carapace without lateral ca- rinae and an abdomen that is unarmed dor- sally. A redescription of Plesionika macropoda Chace, 1939 was decided upon due to the lack of figures in the original de- scription. For a thorough report on the Pan- dalidae, the reader is referred to Chace (1985). Holotype and paratypes of Heterocarpus cutressi are deposited in the National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. (USNM); para- types are also deposited in the Museum of Natural History, Paris, France (MNHP); and in the Marine Invertebrate Museum of the University of Puerto Rico at the Depart- ment of Marine Sciences Field Station, Mayaguez, Puerto Rico (UPR). Heterocarpus cutressi, new species Figs. 1-3 Material.— Holotype: USNM 234162, 1 6, southwest of Puerto Rico, 17°51.7’N, 67°15.4'W, 777 m, 22 Dec 1983, coll. D. Hensley, 37.4 mm carapace length. Paratypes: USNM 234163, 1 3, 1 2 (ovig.), south of Puerto Rico, 17°54.3'N, 66°51.0'W, 500 m, 28 Mar 1985, coll. O. Monterrosa; UPR, 2 6, St. Thomas, U.S. Virgin Islands, May 1983, coll. M. Brandon; UPR, | 8, north-northwest of Buck Island, St. Croix, U.S. Virgin Islands, 550-650 m, Jun—Jul 1985, coll. I. Clavijo. Size of paratypes (carapace length): male, 23.8-37.3 mm (n = 3); female, 36.1 mm; ovigerous female, 34.2 mm. All specimens were captured in baited wire-mesh fish traps. Description. —Rostrum somewhat longer than carapace in smaller specimens, slightly shorter than carapace in larger ones, di- Fig. 1. x 1.8). rected slightly dorsad, with lateral carina extending over posterior 743-% of length; armed with 10 or 11 dorsal teeth extending to anterior end, including 5-7 postorbital teeth, armed ventrally with 6—10 teeth. Car- apace with median postrostral ridge nearly complete; with 2 lateral carinae, dorsalmost directly in line with antennal spine and ex- tending to posterior margin, lateral carina reaching 7 to 34 carapace length; with slight depression found at midlength, between ca- rinae; branchiostegal spine slightly over- reaching antennal spine (Fig. 1). Abdomen (Fig. 2A) with Ist and 2nd so- mites rounded dorsally, 3rd somite with distinct middorsal carinae, somites 4 and 5 slightly carinate middorsally, somite 6 with subparallel ridges partially subdivided by a middorsal sulcus. Pleuron 4 with small pos- teroventral tooth, pleuron 5 posteroven- trally produced into elongate spine. Telson 13 length of 6th somite, with shallow mid- dorsal concavity reaching almost to poste- rior tip; 5 pairs of dorsolateral spinules, pos- teriormost pair situated dorsolateral to base of long, lateral spine of 2 apical pairs. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Heterocarpus cutressi, male holotype from South Puerto Rico, carapace length 37.4 mm (Magnification: Eye subpyriform, maximum diameter about 4 of carapace length; ocellus lacking. Antennular peduncle (Fig. 2B) with sty- locerite acute, reaching at least to midlength of external margin of 2nd segment of pe- duncle. Antennal scale (Fig. 2C) with blade slight- ly overreaching distolateral spine, distal margin broadly rounded. Mouthparts (Fig. 2D-I) as illustrated. Mandibular palp (Fig. 2D) with basal article lobate at inner distal angle, 2nd article (Fig. 2E) broad, about '2 length of terminal ar- ticle. Third maxilliped overreaching anten- nal scale by *% length of terminal article, armed terminally with small corneous spine, penultimate segment about 7% length ter- minal segment. Pereopods with well-formed epipods on 4 anterior pairs. First pereopod slightly overreaching antennal scale; fingers small (Fig. 3D), less than 1 mm in length, with single apical spine on each finger. Second pair of pereopods with shorter one (Fig. 3C) reaching distal “—’% of antennal scale, chela subequal to carpus in length, fingers with VOLUME 101, NUMBER 3 GEN 635 Figs. 2. Heterocarpus cutressi, male holotype from South Puerto Rico: A, Posterior part of abdomen, dorsal aspect including telson and uropods; B, Left antennule, dorsal aspect; C, Left antennal scale, dorsal aspect; D, Right mandible; E, Right mandibular palp, reverse aspect; F, Left 1st maxilla; G, Left 2nd maxilla; H, Left Ist maxilliped; I, Left 2nd maxilliped (Magnifications: A, x 3.2; B, C, D, F, G, H, I, x65.; E, x 13.5). entire cutting edges, subequal to palm in length, carpus comprising 6 articles, is- chium carinate along posterior margin; longer 2nd pereopod (Fig. 3E) overreaching antennal scale by lengths of chela and distal % of carpus, latter with 22—25 articles. Third pereopod (Fig. 3D) extending beyond an- tennal scale by lengths of dactyl, propodus and distal 4 to ¥% of carpus; dactyl (Fig. 3E) % length of propodus, with 5 spines on pos- terior margin; carpus, in addition to row of spines along posterior margin, with 1 or 2 spines on lateral margin at 3—% its length; 3 distal segments, combined, slightly longer than carapace. Fourth pereopod overreach- ing antennal scale by lengths of dactyl and propodus; dactyl as in 3rd pereopod; | or 2 spines projecting posterolaterally from car- pus; 3 distal segments, combined, slightly longer than carapace. Fifth pereopod over- reaching antennal scale by length of dactyl and 74—%4 of propodus; dactyl as in 3rd and 4th pereopods; carpus with accessory spine projecting midlaterally; 3 distal segments, combined, 1,.-1% as long as carapace. Endopod of first pleopod of male (Fig. 3F) nearly '2 length of exopod, distally broad- ened with mesial margin strongly sinuous, bearing minute hooks, without obvious notch or sinus. Appendix masculina of 2nd pleopod (Fig. 3G) armed with over 20 spines of varying lengths on anteromesial and dis- tal margins (Fig. 3H). Exopod of 3rd pleo- pod about % as long as carapace. Mesial 636 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Heterocarpus cutressi, A-C, F-H, male holotype from South Puerto Rico; D, E, male paratype from southwest Puerto Rico: A, Chela left 1st pereopod; B, Left 2nd pereopod; C, Right, same; D, Left 3rd, 4th, 5th pereopod; E, Dactyl left 3rd pereopod; F, Right 1st pleopod; G, Right 2nd pleopod showing appendices masculinae and internae; H, Right appendices masculinae and internae, mesial aspect (Magnifications: D, x 3.2; B, C, F, G, <6.7; E, x 12.4; A, x25; H, <37.5). branch of uropod slightly overreaching tel- son proper; lateral branch slightly longer. Coloration. —Integument orange-red ex- cept for white areas on midlateral surface and dorsal posterior margin of carapace, lat- eral surface of each abdominal somite, base of uropods, telson, antennal scale and basi- cerite. Ova red-brown. Habitat.—A sediment sample for one of the trapping stations indicated a mud bot- tom. Remarks. —Of the 21 species and sub- species within Heterocarpus, only H. cu- tressi and the Indo-Pacific H. woodmasoni Alcock, 1901 have a carapace with the dor- salmost of two lateral carinae originating anteriorly at the antennal spine (i.e., directly in line). The West Indian species is easily distinguished from H. woodmasoni by the absence of the median tooth near the mid- length of the 3rd abdominal somite of the latter species (Chace 1985). Etymology. —It is a pleasure to name this species for Prof. Charles E. Cutress who in- spired this investigation. His experience in- cludes 22 years of dedicated teaching of ma- VOLUME 101, NUMBER 3 637 Fig. 4. Plesionika macropoda, female holotype from West Puerto Rico, carapace length 21.3 mm (Magni- fication: x 4.9). rine invertebrate systematics and biology in the Department of Marine Sciences, Uni- versity of Puerto Rico at Mayaguez. Plesionika macropoda Chace, 1939 Figs. 4, 5 Plesionika macropoda. Chace, 1939:37.— Springer & Bullis,1956:12.—Bullis & Thompson, 1965:8. Material examined. —USNM, 1 4, 5 2 (3 ovig.), off Fredericksted, St. Croix, U.S. Vir- gin Islands, 320 m, 27 Aug 1982, coll. W. Tobias; USNM, 3 4, 1 2, north of Mona Island, Puerto Rico, 18°14.3'’N, 67°52.7'W, 500 m, 10 Mar 1984, coll. O. Monterrosa; UPR, 5 2 (2 ovig.), south Puerto Rico, 17°54.3'N, 66°51.0'W, 500 m, 28 Mar 1985, coll. O. Monterrosa; UPR, 2 2 (1 ovig.), north St. Thomas, U.S. Virgin Islands, 550- 650 m, Jun-Jul 1985, coll. I. Clavijo; MNHP, 5 4, 2 2, south Puerto Rico, 17°53.6’N, 66°55.1’W, 475 m, 7 Oct 1985, R/V Chapman, NMFS, cruise no. 507, sta 28; UPR, 1 2, south Puerto Rico, 17°53.2'N, 66°54.6’W, 615 m, 11 Oct 1985, R/V Chap- man, NMFS, cruise no. 507, sta 34. Size of material examined (carapace length): males, 16.4—20.8 mm (n = 9); fe- males, 9.5-21.3 mm (n = 10); ovigerous females, 20.6-22.1 mm (n = 6). All specimens were captured in baited, wire-mesh fish traps, except 1 2 from R/V Chapman, sta 34, which was collected in a bottom trawl. Geographic range.—Collections of Ple- sionika macropoda have been few but the species appears to be distributed throughout the Greater and Lesser Antilles (Cuba, Chace 1939; Gulf of Mexico, Springer & Bullis 1956; Guadeloupe, Crosnier, pers. comm.; and Puerto Rico and the U.S. Virgin Is- lands). Bathymetric range. —Specimens have been collected between 320-650 m. Remarks.— Apart from Plesionika mac- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Plesionika macropoda, A—O, female holotype; P, Q, male paratype from West Puerto Rico: A, Left eye and orbit; B, Telson and uropods, dorsal aspect; C, Left antennule, mesiodorsal aspect; D, Left antennal scale, dorsal aspect; E, Left mandible; F, Left 1st maxilla; G, Left 2nd maxilla; H, Left 1st maxilliped; I, Left 2nd maxilliped; J, Distal end of left 3rd maxilliped; K, Chela of left 1st pereopod; L, Left 2nd pereopod; M, Dactyl and distal end of propodus of left 4th pereopod; N, Tegumental scale from posterodorsal area of carapace; O, Same, lateral aspect; P, Endopod of left 1st pleopod; Q, Left appendices masculinae and internae, mesial aspect (Magnifications: A, B, D, x3.2; C, E-I, L, x 6.4; P, x 13.2; K, M, Q, x25; J, x 165; N, O, x 323). ropoda Chace, 1939, other western Atlan- tic members of the genus having a rostrum that is dorsally smooth (excluding the basal crest) include P. polyacanthomerus Pequeg- nat, 1970 and P. martia (A. Milne-Edwards, 1883). Plesionika polyacanthomerus is eas- ily differentiated by the presence of a broad curved ridge on the lateral surface of the carapace (Pequegnat 1970). Plesionika mar- tia on the other hand, has a rostrum with fewer dorsal teeth at the basal crest (5—9 vs. 8-10 for P. macropoda) and considerably more ventral teeth (34-56 vs. 9-14 respec- tively) (Pequegnat 1970; Crosnier & Forest 1973, fig. 63d). In addition, dactyls of the walking legs of Plesionika martia are armed with a single terminal tooth (A. Milne-Ed- wards 1883, pl. 21), dactyls of P. macropoda have 5 teeth. To my knowledge, Plesionika edwardsii (Brandt, 1851) and P. macropoda are the only two species within the genus to have multiple teeth along the posterior margin of the dactyls, P. edwardsii having 4 (Chace VOLUME 101, NUMBER 3 1985, fig. 26e) and P. macropoda with 5 (includes terminal tooth in both cases). Fea- tures distinguishing P. macropoda from P. edwardsii respectively, include longer walk- ing legs, a rostrum with fewer dorsal teeth (8-10 vs. 28-34), color of the eggs (red- brown vs. blue), and well-formed versus vestigal epipods. Acknowledgments I am grateful to Dr. Fenner A. Chace, Zoologist Emeritus, National Museum of Natural History, for his invaluable and un- selfish assistance throughout the course of this study. In addition, Dr. Alain Crosnier of the Division of Crustacea, Museum of Natural History in Paris, contributed im- portant suggestions to the revision of the manuscript. My sincere thanks to the Smithsonian Institution, especially to members of the Division of Crustacea, and to Prof. C. Cutress, Mrs. B. Cutress, and Dr. R. Appeldoorn of the University of Puerto Rico at Mayaguez, who provided helpful comments and assistance. This investigation was funded by a Smithsonian Institution Graduate Student Fellowship and a grant to Dr. R. Appel- doorn from the Puerto Rico Sea Grant Pro- gram (PD-19). Literature Cited Bullis, H. R., Jr., & J. R. Thompson. 1965. Collec- tions by the exploratory fishing vessels Oregon, 639 Silver Bay, Combat, and Pelican made during 1956 to 1960 in the southwestern North Atlan- tic.—United States Fish and Wildlife Service Special Scientific Report— Fisheries 510:1—130. Chace, F. A., Jr. 1939. Preliminary descriptions of one new genus and seventeen new species of decapod and stomatopod Crustacea in Reports on the scientific results of the First Atlantis Ex- pedition to the West Indies, under the joint aus- pices of the University of Havana and Harvard University. — Memorias de la Sociedad Cubana de Historia Natural 13(1):31—54. . 1985. The caridean shrimps (Crustacea: De- capoda) of the A/batross Philippine Expedition, 1907-1910, Part 3: Families Thalassocarididae and Pandalidae.—Smithsonian Contributions to Zoology 411: iv + 1-143, 62 figs. Crosnier, A., & J. Forest. 1973. Les crevettes pro- fondes de |’Atlantique oriental tropical. —Faune Tropicale (O.R.S.T.O.M.) 19:1—409, 121 figs. Milne-Edwards, A. 1883. Recueil de figures de crus- taces nouveaux ou peu connus, 3 pp., 44 pls. Paris. Pequegnat, L.H. 1970. Contributions on the biology of the Gulf of Mexico, 4: Deep-sea caridean shrimps with descriptions of six new species. — Texas A&M University Oceanographic Studies, 1:59-123, 17 figs. Springer, S., & H. R. Bullis, Jr. 1956. Collections by the Oregon in the Gulf of Mexico. [U.S.] Fish and Wildlife Service, Special Scientific Report— Fisheries 196:134 pp. Department of Marine Sciences, Univer- sity of Puerto Rico, P.O. Box 5000, Maya- guez, Puerto Rico 00709-5000. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 640-643 NOTES ON THE FRESHWATER CRABS OF THE GENUS MORITSCHUS PRETZMANN, 1965 (CRUSTACEA: DECAPODA: PSEUDOTHELPHUSIDAE) WITH DESCRIPTION OF M. NARINNENSIS FROM SOUTHERN COLOMBIA Martha R. Campos and Gilberto Rodriguez Abstract. — Moritschus narinnensis, n. sp. adds a second species to a genus of pseudothelphusid crabs previously considered to be monotypic. The distri- bution of the genus appears to be confined to the high mountains of the Pacific slope of the Western Cordillera of Colombia and Ecuador, in areas neighboring the boundary between them, but their closest affinities are with Hypolobocera henrici from the Amazonian drainage of Ecuador and Peru. The generic di- agnosis is redefined to accommodate the new species and a key is given to separate the two taxa. In 1897, Mary J. Rathbun, following the system of genera of freshwater crabs then in use, described a sample of small crabs collected by H. Deyrolle near Quito, Ec- uador, which she named Pseudothelphusa ecuadorensis. Almost 70 years later, Pretz- mann (1965) created the monotypic sub- genus Moritschus of the genus Hypolobo- cera, to receive this species. In his monograph of the Pseudothelphusidae, Ro- driguez (1982) gave generic status to Mor- itschus and mentioned the similarity of its gonopod to that of Hypolobocera henrici, a species from the Amazonian drainage of Ecucador and Pert. The species is fairly common around Quito and has been col- lected repeatedly in that area. Recent inten- sive collections in southern Colombia have revealed the presence there of a second, un- described species of Moritschus. The material is deposited at the Museum of Natural History, Universidad Nacional de Colombia, Bogota (ICN-MHN), and in the reference collection of the Instituto Venezolano de Investigaciones Cientificas (IVIC). Tribe Hypolobocerini Pretzmann, 1971 Genus Moritschus Pretzmann, 1965 Moritschus ecuadorensis (Rathbun, 1897) Material examined. — Alluriquin, affluent of Rio Toachi, SE of Santo Domingo de los Colorados, Pichincha Province, Ecuador; 31 Nov 1980; H. Diaz; 12 males, the largest cb. 22.0 mm, cl. 13.9 mm (IVIC). Remarks.—These specimens agree well with the description and illustration of the species given by Rathbun (1897) and Ro- driguez (1982). The border of the marginal lobe of the gonopod has a band of very tiny spinules, barely visible with the stereoscop- ic microscope at a magnification of 500 times. The 5th abdominal sternites of the males are deeply invaginated to form gon- opodial receptacles where the apices of the gonopods are lodged. However, the mature males usually carry the gonopods outside the abdominal fossa, protruding on each side of the folded abdomen. The large gonopo- dial receptacles and the extrusion of the gonopods are also characteristic of Hypo- lobocera henrici; in other pseudothelphu- VOLUME 101, NUMBER 3 sids the gonopodial receptacles are shallow- er and the gonopods are folded inside the abdominal fossa. The known distribution of the species covers the Pacific slope of the Western Cor- dillera of Eucador, between the Guaylla- bamba and Toachi rivers, tributaries of the Esmeraldas River, at altitudes between 880 and 2740 m above sea level. Moritschus narinnensis, new species Fig. | Material examined.—Quebrada Taibai, Vereda Piedra Verde, Inspeccion Buena Vista, Municipio Barbacoas, 1140 m above sea level, Narino Department, Colombia; 16 Aug 1987; G. Arango; 1 male holotype, cb. 25.2 mm, cl. 15.5 mm, 12 male para- types, the largest cb. 24.4 mm, cl. 15.0, 16 female paratypes, the largest cb. 24.2 mm and cl. 15.3 mm (ICN-MHN No. CR 0605). Description. —The cervical groove is straight and shallow, it reaches the margin of the carapace. The anterolateral margin has a conspicuous notch behind the orbit; it does not meet the outer orbital angle, but curves upward above the orbital margin; it is smooth, except for a faint crenulation near its midlength. The postfrontal lobes are ill defined, their presence being indicated only by 2 small scars. The median groove is ob- solescent. The surface of the carapace be- hind the front is moderately inclined ante- riorly and towards the mid-line. The upper border of the front is arched and bilobed in dorsal view, bearing a row of tubercles; the lower margin is strongly sinuous in frontal view and conspicuously marginated; it lies a little in front of the upper one. The front between the upper and lower margins is high. The surface of the carapace is smooth, cov- ered by papillae not visible to the naked eye; for the most part, the regions are clearly defined. The palm of the larger cheliped is inflated. The fingers have a strong gape between them; the mobile finger is strongly arched. The walking legs are slender and elongate, the 641 longest being the third pair (total length greater than the breadth of carapace); in this pair the merus is 3.2 times longer than wide. The exopod of the third maxilliped is 0.32 the length of the ischium of the endognath. The orifice of the efferent branchial channel is Open. The male gonopod is slender and strongly arched laterally; the lateral lobe, which is elongate and moderately prominent, occu- pies the distal half of the appendage; the apex is strongly elongate in the meso-lateral plane and strongly arched, the caudal end produced in a strong fingerlike process di- rected proximally. The elongate process over the field of spines, which in all species of Hypoloberini has 2 rudimentary papillae, in this species is entire, with its distal margin bordered with minute spines. In addition to the spines surrounding the spermatic ap- erture, other small closely-set ones cover the lateral margin of the apex and the fingerlike process. Etymology.—The species is named after the Narino Department in Southern Co- lombia. Remarks.—The locality of the present species is on the Pacific slope of the Western Cordillera of Colombia, which is a contin- uation of the Western Cordillera of Ecua- dor, but 170 km to the North of the area frequented by Moritschus ecuadorensis, in the basin of the Paita River. The species of Moritschus are among the smallest within the Pseudothelphusidae. /. narinnensis can be easily distinguished from M. ecuadorensis by details of the male gon- opod given in the key below. Key to the Species of the Genus Moritschus 1. Gonopod with caudal end produced in short beak. Elongate process over field of spines with 2 rudimentary papillae directed laterally. Lateral margin of apex unarmed eae ire, TERS Ceti: bat M. ecuadorensis — Gonopod with caudal end pro- 642 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Moritschus narinnensis, holotype, ICN 0605: a—e, Left gonopod (a, Caudal view; b, Same lateral view; c, Apex in caudal view; d, Apex in mesial view; e, Apex in distal view); f, Third maxilliped, left; g, Aperture of efferent channel; h, Larger chela; i, Detail of anterolateral margin of carapace at the outer orbital angle. duced in strong fingerlike process di- rected proximally. Elongate process over field of spines entire, its distal margin bordered with minute spines. Lateral margin of apex and fingerlike process covered with small closely- SCUSPINES® hire. See M. narinnensis Since the gonopod type found in Mo- ritschus somewhat resembles the gonopod of some Hypolobocera, particularly H. hen- rici, a revised definition of Moritschus is given below to distinguish between the gen- era and to accommodate the new species described above. VOLUME 101, NUMBER 3 Moritschus Pretzmann, 1965 Pseudothelphusid crabs of small size (car- apace breadth usually less than 26 mm). The anterolateral margin of the carapace does not meet the outer orbital angle, but curves upward above the orbital margin. The ex- ognath of the third maxilliped is less than half the length of the lateral margin of the ischium of the endognath. The male gono- pod is slender and strongly arched laterally; the lateral lobe occupies the distal half of the appendage; it is elongate and moderately prominent; the apex is strongly elongate in the meso-lateral plane and strongly arched; its caudal end is produced in a beak or strong fingerlike process directed proximally. Literature Cited Pretzmann, G. 1965. Vorlaufiger Bericht uber die Familie Pseudothelphusidae.—Anzeiger der 643 Osterreichischen Akademie der Wissenschaften Mathematische Naturwissenschaftliche Klasse, Jahrgang 1965 1:1-11. Rathbun, M. J. 1897. Description de nouvelles es- péces de Crabes d’eau douce appartenant aux collections du Muséum d’Histoire naturelle de Paris.—Bulletin du Muséum National d’His- toire Naturelle de Paris 3(2):58-61. Rodriguez, G. 1982. Les Crabes d’eau douce d’Amé- rique.-Famille des Pseudothelphusidae. — Faune Tropicale 22:1—223. (MRC) Universidad Nacional, Instituto de Ciencias Naturales, Apartado Aéreo 7495, Bogota, Colombia; (GR) Centro de Ecologia, Instituto Venezolano de Investi- gaciones Cientificas, Apartado 21827, Ca- racas 1020-A, Venezuela. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 644-652 CAMBARUS (EREBICAMBARUS) MACULATUS, A NEW CRAYFISH (DECAPODA: CAMBARIDAE) FROM THE MERAMEC RIVER BASIN OF MISSOURI Horton H. Hobbs, Jr., and William L. Pflieger Abstract. — Cambarus (Erebicambarus) maculatus is described from the Mer- amec River and its tributaries in eastern Missouri. Its speckled color pattern, which is unique in the genus Cambarus, prompted the vernacular name, “‘Freck- led Crayfish.” It may be distinguished from its closest relatives, C. (E.) hubbsi Creaser (1931) and C. (E.) rusticiformis Rhoades (1944), by the color pattern and by the tapering central projection of the first pleopod of the first form male. So long as the crayfish described here re- tains its color pattern, it is one of the most easily recognized of any of the American species. Only one other, Procambarus (Gi- rardiella) tulanei Penn (1953), an inhabitant of the Ouachita River Basin in Arkansas and Louisiana, exhibits a similar freckled pattern, but the chelae of the males of that species bear a heavy beard on the mesial surface of the palm; moreover, the pleura of members of both sexes are broadly rounded rather than being acute as they are in this crayfish from Missouri. Closely allied to the new species from the Meramec River Basin, and occupying con- tiguous, if not overlapping, ranges are the troglobitic Cambarus (Erebicambarus) hu- brichti Hobbs (1952) and a probable eco- logical equivalent, C. (E£.) hubbsi Creaser (1931), which seems to be limited to the White and St. Francis watersheds of Mis- souri and Arkansas. Cambarus (Erebicambarus) maculatus, new species Figs. 1, 2 Cambarus hubbsi. — Williams 1954:908, fig. 233 [in part]. Freckled Crayfish, Pflieger, 1987a:23; 1987b:7. Diagnosis.—Body pigmented; eyes well developed. Rostrum, with thickened lateral carinae, broad basally, tapering gradually to base of acumen where studded with paired, conspicuous, dorsally projecting, corneous tubercles; apex of acumen with similar, dor- sally directed, strong tubercle. Postorbital ridges thickened and capped at anterior ex- tremity with prominent, corneous, some- times spiniform tubercle. Suborbital angle obtuse, usually rounded. Cervical spines represented by rounded, inconspicuous tu- bercles. Areola of adults 3.8 to 5.9 times as long as wide and comprising 37.0 to 40.6% of entire length of carapace (43.8 to 49.0% of postorbital carapace length), and bearing 3 to 6 punctations across narrowest part. Chela with single row of 9 to 12 (usually 10) often poorly defined tubercles on mesial surface of palm, remainder of which punc- tate. Fingers with tubercles restricted to op- posable margins; fixed member with third tubercle from base distinctly larger than others on either finger (size difference not evident in regenerated chelae); both fingers with well defined dorsomedian longitudinal ridges. Prominent subacute hook on is- chium of third pereiopod distinctly over- reaching basioischial articulation, and op- posed by tubercle on basis. Pleura of second through fifth abdominal segments tapering to acute or subacute apices. First pleopod VOLUME 101, NUMBER 3 645 Fig. 1. Cambarus (Erebicambarus) maculatus (all from holotype except c and e which are from paratypic male, form II, and g, m which are from allotype): a, Lateral view of carapace; b, c, Mesial view of first pleopod; d, Epistome; e, f, Lateral view of first pleopod; g, Annulus ventralis and adjacent sternites; h, Basal podomeres of third, fourth, and fifth pereiopods; i, Antennal scale; j, Caudal view of first pleopods; k, Dorsal view of carapace; 1, Dorsolateral view of abdomen; m, n, Dorsal view of distal podomeres of cheliped. 646 of first form male terminating in two ele- ments: gently curved, scythelike, tapering central projection with apex, lacking sub- apical notch, directed caudoproximally; and mesial process inflated and directed cau- dolaterally with apex situated lateral to apex of central projection although reflexed somewhat mesially. Color tan to olive tan with darker brown to almost black spots. First pleopods present in female. Holotypic male, form I. —Cephalothorax subovate in cross section; carapace rather strongly depressed (Fig. la, k), its greatest width decidedly broader than height at cau- dodorsal margin of cervical groove (20.1 and 13.0 mm). Rostrum with conspicuously thickened, elevated, and tapering lateral ca- rinae studded with obliquely set, dorsally disposed, corneous tubercles, latter clearly marking base of acumen which, bearing prominent corneous, dorsally directed tip, reaching about midlength of ultimate pod- omere of antennular peduncle. Subrostral ridges rather weak and evident in dorsal as- pect along slightly less than basal third of rostrum. Suborbital angle obtuse. Postor- bital ridges weak except anteriorly where bearing upturned, corneous tip. Branchio- stegal spine obsolete. Areola 5 times as long as broad, with 4 or 5 punctations across narrowest part; length of areola 38.7% of total length of carapace (46.5% of postor- bital carapace length). Cervical spine rep- resented by inconspicuous tubercle no larg- er than others nearby. Carapace densely punctate dorsally, becoming granulate to tu- berculate laterally, especially from area of cervical tubercle anteriorly. Abdomen shorter than carapace (33.7 and 37.7 mm); pleura (Fig. 11) tapering ventral- ly, all acute. Cephalic section of telson with lateral margins comparatively strongly con- verging posteriorly and bearing single prominent, fixed spine in each caudolateral corner; caudal section narrow and appear- ing remarkably small. Proximal podomere of uropod lacking spines on both lobes; dis- tally subtruncate mesial ramus with poorly PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON developed median keel ending in small pre- marginal spine. Cephalomedian lobe of epistome (Fig. 1d) broadly rounded with small cephalomedian projection, margin somewhat thickened, and central part of lobe elevated ventrally; main body with distinct fovea, and epistomal zy- goma rather weakly arched. Ventral surface of proximal podomere of antennule with very small tubercle at about base of distal fourth and slightly lateral to median line of podomere. Antennal peduncle devoid of spines and tubercles; flagellum broken, but probably reaching fifth or sixth abdominal tergum. Antennal scale (Fig. 11) about 2.5 . times as long as broad with mesial and lat- eral margins subparallel for more than half length, distomesial margin of lamellar area tapering to join prominent distolateral spine which attaining about same level anteriorly as apex of rostrum. Mesial half of ischium of third maxilliped with linear clusters of tubercles bearing long, stiff setae; setae scat- tered over lateral half short and inconspic- uous; distolateral extremity of podomere not produced. Right chela (Fig. 1n) about 2.5 times as long as wide, moderately depressed; palm, with length of mesial margin subequal to greatest width of podomere and bearing row of 10 very low tubercles; remainder of palm densely punctate and lacking tubercles other than usual ones associated with articulation of adjacent podomeres. Both fingers with low but clearly defined dorsal and ventral median longitudinal ridges. Arched fixed finger rather smaller than those of some of its congeners and with row of 9 (10 on left) tubercles, third from base largest, rather evenly distributed along opposable margin; usual additional large tubercle situated be- low row just proximal to distal 3 tubercles of row; finger otherwise unremarkable. Typ- ically heavy dactyl with row of 11 (12 on left) tubercles on opposable margin and off- set larger one slightly below row and im- mediately distal to gap between third and fourth (fourth and fifth on left) tubercles VOLUME 101, NUMBER 3 from base; gap lying almost opposite largest tubercle on opposable margin of fixed fin- ger. Minute denticles abundant on oppos- able margins of fingers of chelae of many crayfishes sparse and largely limited to sin- gle fine, short row extending between dis- talmost tubercle and corneous tip of finger. Mesial surface of dactyl lacking tubercles, even basally; podomere entirely punctate except for those on opposable margin. Carpus of cheliped longer than broad (13.4 and 9.5 mm), with moderately deep, oblique, longitudinal furrow dorsally flanked by punctations; mesial surface with prominent mesiodistally directed spine and single (2 on left) small tubercle near proximomesial base; ventral surface polished and with sin- gle tubercle situated on distal margin mesial to ventrolateral articular knob. Merus with 2 small dorsodistal tubercles; ventral sur- face with mesial row of 7 (left with 9) cor- neous tipped spines and 2 (1 on left) rep- resenting lateral row of other crayfishes; lateral and mesial surfaces sparsely punc- tate. Basioischial podomere with row of 4 poorly developed tubercles ventromesially. Hook on ischium of third pereiopod only (Fig. 1h), acute, overreaching basioischial articulation, and opposed by tubercle, flanked by setiferous punctations, on basis. Coxa of fourth pereiopod with prominent caudomesial boss, caudal face of which in almost same plane as remainder of caudal surface of podomere. Boss absent from coxa of fifth pereiopod. First pleopods (Fig. 1b, f, }) reaching coxae of third pereiopods, symmetrical, and with small gap between their bases. See “‘Diagnosis”’ for description of terminal elements. Allotypic female. —Excluding secondary sexual characteristics, differing in only few minor respects from holotype: telson and uropods decidedly hirsute; mesial lobe of proximal podomere of uropod produced caudally in spine; tubercle on ventral sur- face of basal podomere of antennule situ- ated slightly more proximally; chela (Fig. lm) proportionately narrower, about 2.7 647 Table 1.—Measurements (mm) of Cambarus (E.) maculatus. Holo- Allo- Morpho type type type Carapace: Entire length Sled 36.0 32.0 Postorbital length 31.4 29.0 26.6 Width 20.1 18.9 17.2 Height 13.0 12.7 11.2 Areola: Width 2.9 2.4 2.1 Length 14.6 14.2 11.5 Rostrum: Width S72 5.2 5.0 Length 6.3 7.0 5.4 Right chela: Length, palm mesial margin 14.2 13.2 10.5 Palm width 14.5 11.9 11.1 Length, lateral margin 37.1 32.6 25.6 Dactyl length 18.3 16.6 13.2 Abdomen: Width 17.6 17.7 13.5 Length 33.7 31.6 DST times as long as wide; mesial margin of palm with row of 11 ill-defined tubercles. Fixed finger with row of 13 (left with 12, proxi- malmost lacking) tubercles along opposable margin, fifth from base largest; opposable margin of dactyl with row of 13 (left with 14) tubercles and offset 1 following gap; ven- tral surface of merus of cheliped with mesial row of 8 tubercles (9 on left) and lateral row represented by only 1 tubercle. (See Table 1) Annulus ventralis (Fig. 1g) firmly fused to sternite XIII, irregularly ovate with great- er axis lying transversely, about 1.7 times as broad as long; cephalomedian trough moderately deep but not flanked by con- spicuous cephalolateral prominences; trans- verse sulcus well defined with tongue ex- tending dextrally, disappearing under high, longitudinally creased caudal wall. Postan- nular sclerite only slightly narrower than an- nulus and only little more than half as long. 648 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Z/ Fig. 2. Dorsolateral view of first form male of Cambarus (E.) maculatus from Huzzah Creek, Crawford County, Missouri. Morphotypic male, form II.—Differing from holotype in following respects: apical spines on postorbital ridges directed ceph- alolaterally and only slightly dorsally; telson with less convergent margins, and anterior section with pair of movable spines mesial to fixed ones in caudolateral corners; mesial lobe of basal podomere of uropod with prominent caudally projecting spine; mesial margin of palm of chela with row of 14 tubercles (left with 10); opposable margin of fixed finger with row of 8 tubercles (left with 10); opposable margin of dactyl with row of 11 tubercles and offset one (left with 10 and 1); row of minute denticles on op- posable margins of both fingers longer and more conspicuous; ornamentation of more basal podomeres within ranges noted in ho- lotype; hooks on ischia of third pereiopod much less well developed, not reaching bas- ioischial articulation, but small opposing tubercle on basis clearly evident. First pleo- pods (like Fig. lc, e) differing from that of holotype chiefly in structure of central pro- jection which much more massive, com- paratively shorter, and non-corneous. (See Table 1.) Color notes (Fig. 2).—Carapace yellowish tan to pale orange tan with very dark brown to black spots, largest of which subcircular and occurring dorsolaterally over bran- chiostegites and in posterior part of areola. Dorsal surface of abdomen, telson, and uro- pods with yellow or tan base infused with brown, thus yellowish to orange brown, with dark spots similar to, but smaller than, those on carapace. Background color of chelipeds often much paler, sometimes almost cream, and, except ventrally, bearing dark brown to black spots of varying sizes and shapes; only few of those on dorsal part of carpus and merus as large as those on branchio- stegites. Second through fifth pereiopods similarly colored but with spots small and more sparse. Size.—The largest specimen available is a first form male from Huzzah Creek in Crawford County having a carapace length of 44.4 (postorbital carapace length 37.2) mm. The smallest first form male, from Washington County, has corresponding lengths of 24.6 and 20.4 mm. The smallest ovigerous female available has a carapace length of 32 mm (postorbital length 27.8 VOLUME 101, NUMBER 3 mm), the largest, 36 and 30.6 mm, respec- tively. Type locality. —WHazel Creek at Route C, 8 miles (12.8 km) northeast of Courtois, Washington County, Missouri (T. 36N, R. 1W, SE'4 Sec. 24). The specimens collected there were found beneath rocks in the plunge pool just below the concrete road crossing. At low water this pool measured 14 m in width and 8.4 m in length, with a maximum depth of 1 m. At this locality, C. (E.) mac- ulatus occurred in association with Orco- nectes (Procericambarus) punctimanus (Creaser 1933), O. (P.) luteus (Creaser, 1933) and O. (P.) medius (Faxon, 1884). The latter species comprised 73.2% of our collections, while C. (E.) maculatus comprised 5.5%, the least common of the four species. Disposition of types. —The holotype, al- lotype, and morphotype (USNM 219292, 219293, and 219280, respectively) are de- posited in the National Museum of Natural History, Smithsonian Institution. Of the paratypes, 3éI, 2éII, 12, and 1j2 are in the collection of Ronald D. Oesch; the remain- ing ones are in the National Museum of Natural History. Range and specimens examined. —This new crayfish appears to be restricted to the Meramec River and its tributaries in eastern Missouri (Fig. 3) where it has been found in the following localities: Crawford Coun- ty: (1) Huzzah Creek at end of Rte E (T. 38N, R. 3W, Sec. 11), 141, 4 Mar 1977, W. L. Pflieger, A. Buchanan; (2) Meramec Riv- er at Cooks Station (T. 36N, R. SW, NE%4 Sec. 6), 22, 12 Oct 1984, WLP, 1éI, 1¢II, 18 Jul 1985, WLP, 141, 6 May 1986, WLP; (3) Meramec River at Hwy 8, 8.75 mi (14 km) W of Steelville (T. 37N, R. 5W, SW'4 Sec. 6) 29, 1j9, 25 Aug 1968, J. E. Cooper, M. R. Cooper, 3éI, 2éII, 24 Oct 1985, WLP, 3611, 22, 5 May 1986, WLP, 161, 16 Apr 1987, R. D. Oesch; (4) Huzzah Creek (T. 37N, R. 2W, Sec. 6), 341, 12, 23 Oct 1986, WLP; (5) Trib of Huzzah Creek 1.5 mi (2.4 km) N of Dillard, 14II, 10 Sep 1955, Nell Crenshaw, John Crenshaw; (6) Huzzah 649 Creek below Hwy 8, 14], 1¢II, 12, 133, 9 May 1987, RDO. Franklin County: (7) Meramec River at Meramec State Park (T. 40N, R. 2W, Sec. 13), 161, 4 Mar 1977, WLP, AB. Washington County: (8) Type locality, 5éI, 22, 18 Oct 1984, WLP, 1éI, 22, 5 May 1986, WLP, 1éI, 22, 4j6, 25 Mar 1985, WLP; (9) Big River at Hwy 21, 3 mi N of Caledonia (T. 36N, R. 2E, Sec. 25), 1j2, 25 Aug 1968, JEC, MRC, 12, 1j¢, 17 Oct 1984, WLP; (10) Mill Creek at Rte N (T. 36N, R. 3E, NE Seew22)) 2615" 25° Mar 19855 WEP: (i) Fourche Renault Creek at Rte AA, 1 mi W of Summer Lake (T. 38N, R. 1E, SW Sec 34), 161, 22 Oct 1986, WLP; (12) Mine Bre- ton Creek at Hwy 8, 16], 11 Apr 1984, RDO; (13) Cedar Creek at Hwy 32, 1.5 airmi (2.4 km) ESE of Caledonia, 1j¢, 18 Aug 1948, ABW, ABL, 12, 21 Jun 1987, RDO; (14) Big River, 8.6 mi (13.8 km) S of Potosi, 1éII, 236, 18 Aug 1948, ABW, ABL; (15) Mineral Fork at W Hwy 57 bridge, 1¢II, 7 Jul 1979, J. F. Payne; (16) Big River at Hwy DN MAO I We IES We Variations. — Barring variations that have almost certainly resulted from injury, re- generation, or abrasion that often appear to be commensurate with the length of the pe- riod since the most recent molt, our speci- mens exhibit remarkable uniformity in both mensural and meristic qualities. Such ratios as length of areola/total length of body yielded a mean of 38.6 + 1.653; length of the areola/postorbital carapace length, 46.3 + 1.125; and areola length/areola width, 4.9 + 0.597. Moreover, ratios involving the length of the chela, its maximum width, and length of the palm, even when regenerated chelae were considered, show little varia- tion. Similarly the differences in the num- bers of tubercles on the body and the var- ious podomeres of the chelipeds are comparatively small, hardly exceeding the variations pointed out in the descriptions of the primary types. Perhaps the most con- spicuous variation occurs in the relative de- velopment and numbers of tubercles on the regenerated chelipeds. Such chelae can be 650 Fig. 3. recognized by the absence ofan enlargement of the third, fourth, or fifth tubercle from the base on the opposable margin of the fixed finger. While it is likely that such a difference does not exist in a cheliped that was lost and regenerated in the earlier in- stars, should a large tubercle be wanting on the opposable margin of that finger, one may be sure that the appendage is a regenerated one, and is likely to bear a larger number of tubercles on that and the corresponding margin of the dactyl. Ecological notes.—Cambarus (E.) mac- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Distribution of Cambarus (E.) maculatus (black circles) and Cambarus (E.) hubbsi (open circles). ulatus is largely restricted to a deeply dis- sected region of the eastern Ozarks known as the Coutois Hills. The bedrocks of this region consist principally of limestones and dolomites of the Gasconade and Potosi for- mations. Chert gravel weathered from these rocks is the principal bottom type in the streams. Much of the region is sparsely pop- ulated and is within the boundaries of the Mark Twain National Forest. The streams are very clear, with permanent flow main- tained by numerous springs. C. (E.) mac- ulatus occurs in streams ranging in size from VOLUME 101, NUMBER 3 order 3 to 7, but is most abundant in those of order 4 or 5. It is usually found in pools, at depths of 20 cm or more, beneath rocks that are well seated in gravel. Its presence is often revealed by small piles of gravel pushed out of the shallow cavities that it excavates. The habitat and habits of this species are not notably different from those of C. (E.) hubbsi. In 12 collections made by one of us (WLP), five other species of crayfishes, all belonging to the genus Orconectes occurred. The species and number of collections were: O. (Procericambarus) luteus, 12; O. (P.) punc- timanus, 9; O. (P.) medius, 8; O. (Billecam- barus) harrisonii (Faxon 1884), 4; and O. (P.) hylas (Faxon 1890), 2. Life history notes.—In our collections from the months of October, March, and April, 18 of 20 males were Form I, while 5 of 6 males collected during the months of July to September were Form II. Three ovigerous females occurred in collections made on 5 May 1986. These females mea- sured 32, 34, and 36 mm carapace length and carried 67, 63, and 127 eggs, respec- tively. The eggs were about 1.8 mm in di- ameter and were grey to nearly black in col- or. Relationships.—Cambarus (Erebicam- barus) maculatus has its closest affinities with C. (E.) hubbsi and C. (E.) rusticiformis Rhoades (1944). Differing from their other congeners, these three crayfishes with strongly depressed bodies possess rostra, the lateral carinae of which end abruptly at the base of the acumen and usually bear mar- ginal tubercles at their apices. The carinae are thickened and diverge at their bases. All three have rather broad, moderately densely punctate areolae. The rather weak tubercles along the mesial margin of the palms of their chelae are largely confined to a single row. All three exhibit a striking similarity in their abdominal pleura, which taper to acute an- gles (apices often rounded) that, when the abdomen is extended, are directed almost ventrally. Similarly structured pleura, which may be found to be associated with adap- 651 tations to lotic habitats, occur in the Asta- cidae and in members of the cambarid gen- era Cambaroides and the monotypic Barbicambarus cornutus Faxon (1884), all of which are at least primarily stream dwell- ers. The “‘freckled’”’ color pattern alone serves to distinguish this crayfish from C. (E.) hubbsi and C. (E.) rusticiformis, but in pre- served specimens in which the pigment has completely faded, distinguishing between them becomes more difficult. First form males of the new species are unique among them in that the central projection of the first pleopod tapers to an acute tip, whereas in the other two the comparatively broad distal part of the element bears a subapical notch. Except for the color patterns, we have been unable to discover a single character that will serve consistently to distinguish the second form males, females, and juveniles of the three, although the obliquely set mar- ginal spines of the rostrum that project dor- sally rather than anterodorsally can usually be relied upon to recognize most members of C. (E.) maculatus. The recognition of the allopatric (with respect to the other two) C. (E.) rusticiformis, which is restricted to streams east of the Mississippi River (where it is largely, if not entirely, restricted to the Cumberland Basin, and thus their ranges are not even contiguous) presents no prob- lem. Although Cambarus (E.) hubbsi is re- stricted largely, if not entirely, to the White and St. Francis watersheds of Missouri and Arkansas, there is a Form I male of this species at the National Museum of Natural History (USNM 219315) which was re- portedly collected from the Meramec River east of Eureka, Missouri, in 1934. If this record is valid, the range of C. (E.) hubbsi overlaps that of C. (E.) maculatus, and rec- ognition of the identity of bleached second form males and females of the two, at least sometimes, might be problematical. Recent collections at this locality, however, includ- ed specimens of only the latter species. There can be little doubt that before the 652 ancestral Erebicambarus stock crossed the Mississippi River (Hobbs 1969:110) to be- come established in the Meramec, St. Fran- cis, and White river basins, it had already acquired a mien shared by the segment of the stock that was left behind and that gave rise to the modern C. (E.) rusticiformis. In our view then, the close resemblance exist- ing between the latter, C. (E.) maculatus, and C. (E.) hubbsi, has resulted from a re- tention of features that have been little al- tered during the course of their long-time, continuous occupancy of similar ecological niches rather than from convergence in the three undergoing similar adaptations inde- pendently. Of the three Erebicambarus oc- curring west of the Mississippi River (Ozark Region), only C. (E.) hubrichti exhibits fea- tures that appear to us to be markedly dif- ferent from those we believe to have been characteristic of the pioneering ancestor, and we suggest that these are associated with its ancestors becoming adapted to a spelean en- vironment. At least two features of C. (E.) maculatus, unique in the subgenus Erebicambarus, sug- gest that this crayfish is farther removed from the ancestral stock than is C. ( E.) hubbsi. These are the speckled color pattern and the tapered central projection of the first pleopod in first form males. Acknowledgments Appreciation is extended to Ronald D. Oesch, who is associated with the Cooper- ating Schools of the St. Louis Suburban Area, for lending us specimens of C. (E£.) macu- latus from his collection and to other col- lectors mentioned under “Range and spec- imens examined’? who have donated specimens to the Smithsonian Institution. For their helpful comments on the manu- script, acknowledgment is also made with thanks to C. W. Hart, Jr., of the National Museum of Natural History, and to Douglas G. Smith, of the University of Massachu- setts. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Creaser, E.P. 1931. Three new crayfishes from Pueb- la and Missouri. — Occasional Papers of the Mu- seum of Zoology, University of Michigan 224: 1-10, 17 figs. 1933. Descriptions of some new and poorly known species of North American crayfishes. — Occasional Papers of the Museum of Zoology, University of Michigan 275:1-21, 2 pls. Faxon, W. 1884. Descriptions of new species of Cam- barus, to which is added a synonymical list of the known species of Cambarus and Astacus. — Proceedings of the American Academy of Arts and Sciences 20:107-158. 1890. Notes on North American crayfishes, family Astacidae.— Proceedings of the United States National Museum 12(785):619-634. ; Hobbs, H. H., Jr. 1952. A new albinistic crayfish of the genus Cambarus from southern Missouri with a key to the albinistic species of the genus (Decapoda, Astacidae).— American Midland Naturalist 48(3):689-693, 8 figs. 1969. On the distribution and phylogeny of the crayfish genus Cambarus. In P. C. Holt, R. L. Hoffman, and C. W. Hart, Jr., eds., The dis- tributional history of the biota of the southern Appalachians, Part I: Invertebrates. — Virginia Polytechnic Institute, Research Division Mono- graph 1:93-178, 20 figs. Penn, G. H. 1953. A new crawfish of the genus Pro- cambarus from Louisiana and Arkansas (De- capoda, Astacidae).—Journal of the Washing- ton Academy of Sciences 43(5):163-166, 12 figs. Rhoades, R. 1944. The crayfishes of Kentucky, with notes on variation, distribution and descriptions of new species and subspecies. — American Mid- land Naturalist 31(1):111-149, 10 figs. Pflieger, W. L. 1987a. An introduction to the crayfish of Missouri.— Missouri Conservationist 48(4): 17-31, illustrated in color. . 1987b. Reprint of 1987a with pagination 1- 15. Williams, A. B. 1954. Speciation and distribution of the crayfishes of the Ozark Plateaus and Ouachi- ta Provinces.— University of Kansas Science Bulletin 36(12):803-918, 253 figs. (HHH) Department of Invertebrate Zo- ology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560; (WLP) Fish and Wildlife Re- search Center, Missouri Department of Conservation, 1110 College Avenue, Co- lumbia, Missouri 65201. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 653-661 TYPES OF STOMATOPOD CRUSTACEANS IN THE ZOOLOGICAL SURVEY OF INDIA H. C. Ghosh and Raymond B. Manning Abstract.—A list of the types of stomatopod crustaceans in the Zoological Survey of India is presented. This historically important collection includes type specimens of 42 species, including primary types (holotype, lectotype, or syntype) of 36 species. The collection of types of stomatopod crustaceans held by the Zoological Survey of India forms comprises the most impor- tant collection of types of Indo-West Pacific species in the world, with no less than 42 species, or about 10% of the known sto- matopods, represented in the collection by type specimens. The collection is based largely on material reported on by James Wood-Mason and Stanley W. Kemp be- tween 1875 and 1921, supplemented by a few additions since then. Kemp prepared the first major regional monograph of the stomatopods, and his work remains today as an essential tool for anyone studying the group. Species are first listed alphabetically un- der their original combination. Then species are listed by current names, cross-refer- enced to the original combination. The following) information is included under each species represented in the col- lection: catalogue number, location, date, collector or source of material, and number of specimens. Condition of material is good unless otherwise specified. For some localities, coordinates or alter- nate spellings, taken from gazetteers of the U.S. Board on Geographic Names, have been added; these always are given in square brackets. Where possible we have added station data from “Biological collections of the R.I.M.S. “‘Jnvestigator.”’ List of stations 1884-1913.” Pages 1-35. Indian Museum, Calcutta. 1914. Station data that we have added also is given in square brackets. Under each species, the material is listed in the same order as listed by Kemp (1913). We thank B. K. Tikader, former Director of the Zoological Survey of India, for his permission to undertake the preparation of this catalogue. STOMATOPOD TYPES IN THE ZOOLOGICAL SURVEY OF INDIA, CALCUTTA (Listed alphabetically under original combination) Chorisquilla andamanica Manning, 1975 Chorisquilla andamanica Manning, 1975: 258, fig. 3 [=Chorisquilla excavata (Miers, 1880)]. Holotype. —9177/6, off Andaman Islands [12°30'N, 92°45’E], India, 20 fm (=37 m), Investigator, 1 male. Paratypes. —9177/6, off Andaman Is- lands, 20 fm (=37 m), Jnvestigator, 4 fe- males (one specimen broken in middle).— 9850-3/6, off Little Andaman Island [10°45’N, 92°30’E], 10.5 fm (=19 m), Jn- vestigator, 1 male, 3 females.—9846-9/6, off Table Island [14°12’N, 93°22’E], Coco Group, Andaman Islands, 5 fm (=9 m), Jn- vestigator, 1 male, 3 females. Clorida decorata Wood-Mason, 1875 Clorida decorata Wood-Mason, 1875:231. Holotype. —3086/5, Andaman Islands [12°30'N, 92°45’E], India, J. Wood-Mason, 1 female (delicate, broken). 654 Remarks. —Kemp (1913) gave Port Blair as the type locality. Coronis spinosa Wood-Mason, 1875 Coronis spinosa Wood-Mason, 1875:232 [=Heterosquilla tricarinata (Claus, 1871)]. Lectotype (selected by Holthuis 1967: 11).—3052/5, Port Blair [11°36’N, 92°45’E], Andaman Islands, India, J. Wood-Mason, 1 female (very delicate, dactylus missing). Paralectotype. — 3053/5, Dunedin [45°52’S, 170°30’E], New Zealand, Capt. F. W. Hut- ton, | female. Remarks.—This is a cold water species known to be common in New Zealand; the specimen from the Andaman Islands may have been labelled incorrectly. Gonodactylus chiragra var. platysoma Wood-Mason, 1895 Gonodactylus chiragra var. platysoma Wood-Mason, 1895:11, pl. 3, figs. 3-9) [=Gonodactylus platysoma Wood-Ma- son, 1895]. Lectotype. —3033/5, Society Islands, Otago Museum, | female. Paralectotypes. —3039/5, Mauritius [20°18’S, 57°35'E], purchased, 2 females. Remarks. — Although Kemp (1913:163) considered three specimens from Port Blair, Andamans, to be the types, Wood-Mason did not mention these specimens or mate- rial from the Andamans in his original ac- count. The female from the Society Islands is here selected as the lectotype. Gonodactylus choprai Manning, 1967 Gonodactylus choprai Manning, 1967:16, fig. 6. Paratypes.—C 1334/2, Indian Ocean, off Somali coast, 9°36’N, 51°01’E, 78-82 m, 16 Dec 1964, International Indian Ocean Ex- pedition, Anton Bruun sta 9-444, exchange from Smithsonian Institution, 1 male, 1 fe- male. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Gonodactylus furcicaudatus Miers, 1880 Gonodactylus furcicaudatus Miers, 1880: 124, pl. 3, figs. 13-16 [=Mesacturus fur- cicaudatus (Miers, 1880)]. Syntype. —7225/10, type locality un- known, H.M.S. Alert, exchange from British Museum (Natural History), 1 male. Gonodactylus glyptocercus Wood-Mason, 1875 Gonodactylus_ glyptocercus Wood-Mason, 1875:232 [=Haptosquilla glyptocercus (Wood-Mason, 1875)]. Holotype. —3038/5, Nicobar Islands [8°00'N, 93°30’E], India, S. Kurz, 1 female. Gonodactylus lanchesteri Manning, 1967 Gonodactylus lanchesteri Manning, 1967:11, fig. 4. Paratype.—C 1337/2, Junghi Bay, Ibo Archipelago [12°20’S, 40°36’E], Mozam- bique, 5—18 fm (=9-33 m), A. Patience, ex- change from Smithsonian Institution, 1 fe- male. Gonodactylus nefandus Kemp, 1911 Gonodactylus nefandus Kemp, 1911:93 [=Haptosquilla nefanda (Kemp, 1911)]. Syntypes. —3045/5, Andaman Islands [12°30'N, 92°45’E], India, J. Wood-Mason, 6 males, 16 females.—3040/5, same data, 1 female.—9025/6, same data, 20 fm (=37 m), /nvestigator, 1 female.—2825-6/7, Port Blair [11°36’N, 92°45’E], Andaman Islands, G. H. Booley, 2 males.— 2546/3, Cheduba [Cheduba Island = 18°48’N, 93°38’E], Ara- kan coast, Burma, 7 fm (=13 m), Jnvesti- gator, 3 females. Gonodactylus proximus Kemp, 1915 Gonodactylus proximus Kemp, 1915:183, pl. 1, figs. 9, 10 [=Haptosquilla proxima (Kemp, 1915)]. VOLUME 101, NUMBER 3 Syntypes. —9296/10, Port Galera [Puerto Galera = 13°30’N, 120°57’E], Min- doro, Philippine Islands, Philippine Bureau of Science, | male, 1 female. Lophosquilla tiwarii Blumstein, 1974 Squilla costata.—Kemp, 1913:84 [speci- men from Burmese coast only]. [Not Squilla costata De Haan, 1844.] Squilla sp. prox. costata. —Tiwari & Biswas, 1952:354, fig. 2. Lophosquilla tiwarii Blumstein, 1974:123, fig. 8. Paratype. —C 332/1, 4 miles NNE of Ka- busa Island [Kabosa Island = 12°49'N, 97°53'E], Mergui Archipelago, Burma, /n- vestigator [? sta 550, 33 fm (=60 m), 22 Oct 1913], 1 female. Remarks. — This specimen from the Mer- gui Archipelago is a paratype of Blumstein’s species, as she referred to the accounts of both Kemp and Tiwari & Biswas in her orig- inal description. Lysiosquilla insignis Kemp, 1911 Lysiosquilla insignis Kemp, 1911:94 [=Het- erosquilloides insignis (Kemp, 1911)]. Holotype. — 7568/10, off North Andaman Island, India, 14°27’N, 93°50’E, 235 fm (=430 m), 19 Apr 1907, Investigator sta 375, 1 male (condition delicate, dactylus missing). Remarks.—Kemp (1913) gave 50°7’N, 79°7'E as the coordinates for the Jnvesti- gator station at which this species was tak- en. His coordinates apparently are in error. Lysiosquilla maculata var. sulcirostris Kemp, 1913 Lysiosquilla maculata var. sulcirostris Kemp, 1913:116, pl. 8, figs. 92, 93 [=Ly- siosquilla sulcirostris Kemp, 1913]. Holotype. —7564/10, Andaman Islands [12°30’N, 92°45’E], Homfray and Tytler, 1 male. 655 Lysiosquilla multifasciata Wood-Mason, 1895 Lysiosquilla multifasciata Wood-Mason, 1895:1, pl. 1, figs. 4-7 [=Acanthosquilla multifasciata (Wood-Mason, 1895)]. Holotype. —3342/7, Bombay [18°58’N, 72°50'E], India, Bombay Natural History Society, 1 female (both dactyli missing). Manningia andamanensis Ghosh, 1975 Manningia andamanensis Ghosh, 1975:33, fig. 1. Holotype.—C 1268/2, Aerial Bay Rock Reef, Diglipur, North Andaman Island [13°15’N, 92°55’E], India, 22 Feb 1970, A. G. K. Menon, 1 female. Odontodactylus southwelli Kemp, 1911 Odontodactylus southwelli Kemp, 1911:94 [=Odontodactylus brevirostris (Miers, 1884)]. Syntypes. —3526/10, Andaman Islands [12°30’N, 92°45’E], India, /nvestigator, | fe- male.— 3502/10, Andaman Islands, 53 fm (=97 m), Investigator, 1 male, 1 female.— 9751/6, off Cinque Island, Andaman Is- lands, 20 fm (=37 m), Jnvestigator, 1 fe- male.—9747/6, off Interview Island [12°55'N, 92°43’E], Andaman Islands, /n- vestigator, 1 male.—7313/10, N. Cheval Paar [Cheval Bank = 08°40’N, 79°46’E], Ceylon [=Sri Lanka], 1 female. Remarks. — Although Kemp (1913) con- sidered the specimens from lot 3526/10 as the types, in his original description he did not specify a holotype. All of the specimens assigned to this species by him in 1913 ap- pear to be syntypes. Oratosquilla hindustanica Manning, 1978 Oratosquilla hindustanica Manning, 1978: IS, ERs I NSC: Holotype. —7037-41/10, Tuticorin [8°47'N, 78°08'E], Madras, Gulf of Manaar, South India, J. Hornell leg., 1 male. 656 Paratypes. —ex. 7037—41/10, same data, 12 males, 17 females. Remarks. —These specimens also are paralectotypes of Squilla oratoria var. per- pensa Kemp, 1911 (see Manning 1978: 15,16). Oratosquilla pentadactyla Manning, 1978 Oratosquilla pentadactyla Manning, 1978: 19, fig. 10. Paratype.—8107/10, Kilakarai [9°14’S, 78°47'E], Ramnad District, India, from fishermen’s nets, 17 Feb 1913, S.W. Kemp, 1 male. Oratosquilla subtilis Manning, 1978 Oratosquilla subtilis Manning, 1978:33, fig. 19. Holotype. —7536/10, off Vizagapatam coast [Visakhapatnam = 7°42’N, 83°18’E], Madras, India, 20 fm (=37 m), /nvestigator, 1 female. Paratypes. —ex. 320/1, off Kabusa Island [Kabosa Island = 12°49'N, 97°53’E], lower Burma, 25-35 fm (=46—-64 m), Investigator, 1 male, 1 female. Remarks.—The holotype is also a syn- type of Squilla gonypetes Kemp, 1911. Squilla annandalei Kemp, 1911 Squilla annandalei Kemp, 1911:99 [=Har- piosquilla annandalei (Kemp, 1911)]. Syntypes. —7563/10, Gulf of Martaban, Burma, 14°46’N, 95°52’E, 61 fm (=112 m), 7 Mar 1901, Investigator sta 328, 1 fe- male.—1748/10, same area, 14°26'N, 96°23’E, 67 fm (=123 m), /nvestigator [sta 226, 24 Mar 1897], 1 male.— 1749/10, same area, 14°38'12”N, 96°24'30’E, 53 fm (=97 m), /nvestigator [sta 225, 14°38’'15’N, 96°24'30”E, 24 Mar 1897], 1 female. Remarks. -- As Kemp (1911) did not des- ignate a holotype in the original description of this species, but did mention that he had PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON studied four specimens, all of the four spec- imens reported by him in 1913 must be considered to be syntypes, in spite of his suggestion in 1913 that the type was the female from 7563/10. Squilla bengalensis Tiwari & Biswas, 1952 Squilla bengalensis Tiwari & Biswas, 1952: 352, fig. 1b, c [=Cloridopsis bengalensis (Tiwari & Biswas, 1952)]. Holotype. —C 3013/1, Salt Lakes, Lower Bengal, India, T. N. Poddar, 1 male. Paratype. —C 3014/1, Piali River, Uttar- bhag, Lower Bengal, India, 23 May 1934, S.L. Hora, 1 male. Squilla boopis Kemp, 1911 Squilla boopis Kemp, 1911:97 [=Busquilla quadraticauda (Fukuda, 1911)]. Holotype.—1710/10, Gulf of Martaban, Burma, 14°26'N, 96°23’E, 67 fm (=123 m), Investigator [sta 226, 24 Mar 1897], 1 fe- male. Squilla bombayensis Chhapgar & Sane, 1967 Squilla bombayensis Chhapgar & Sane, 1967:1, fig. 1 [=Clorida bombayensis (Chhapgar & Sane, 1967)]. Holotype. —C 4621/1, Bombay [18°58’N, 72°50'E], India, intertidal water, 19 Apr 1963, S. R. Sane, 1 female. Squilla brasiliensis Calman, 1917 Squilla brasiliensis Calman, 1917:139, figs. 1-3. Paratype. —C 243/1, Atlantic Ocean, near Cabo Frio, Brazil, 22°56’S, 41°34’W, 40 fm (=73 m), 2 May 1913, Terra Nova sta 42, exchange from British Museum (Natural History), 1 male. VOLUME 101, NUMBER 3 Squilla denticauda Chhapgar & Sane, 1967 Squilla denticauda Chhapgar & Sane, 1967: 4, fig. 2 [=Clorida denticauda (Chhapgar & Sane, 1967)]. Holotype. —C 4622/1, Bombay [18°58'N, 72°50’E], India, intertidal water, 17 Jan 1963, B. F. Chhapgar, 1 male. Squilla foveolata Wood-Mason, 1895 Squilla foveolata Wood-Mason, 1895:2, pl. 2, fig. 1 [=Dictyosquilla foveolata (Wood- Mason, 1895)]. Syntypes. —3332/9, Hong Kong [Hong Kong Island = 22°15'N, 114°11’E], G. Den- nys, | male, 4 females. Squilla gilesi Kemp, 1911 Squilla gilesi Kemp, 1911:95 [=Lenisquilla gilesi (Kemp, 1911)]. Syntypes. —7514—-18/10, off Madras coast, India, 14°18'15”N, 80°18'30”E, 88-110 fm (=161-—201 m), Jnvestigator [sta 154, 3 Jan 1894], 1 male, 3 females.—488/10, Madras coast, India, /nvestigator, 1 male.—5800/9, off Orissa coast, India, 19°49'N, 86°43’E, Investigator, 1 female.—3077/5, northeast Bay of Bengal, 20°18’N, 90°50’E, 65 fm (=119 m), Investigator, 4 males.—7519/10, Persian Gulf, 26°20'30”N, 54°52’30”E, 35 fm (=64 m), /nvestigator [sta 345, 20 Oct 1904], 1 female. Squilla gonypetes Kemp, 1911 Squilla gonypetes Kemp, 1911:96 [=Ora- tosquilla gonypetes (Kemp, 1911)]. Lectotype (selected by Manning 1978: 13).—3359/7, off Cheduba [Cheduba Island = 18°48'N, 93°38’E], Arakan coast, Burma, 7 fm (=13 m), Jnvestigator, 1 male. Paralectotypes. —3476/10, off Andaman Islands [12°30'N, 92°45’E], India, 60 fm (=110 m), /nvestigator, 1 male, 1 female.— 4421/10, Persian Gulf, 26°24’N, 56°02’E, 657 47 fm (=86 m), /nvestigator [? sta 296, 26°4'N, 56°2’E, 12 Apr 1902], 1 female. Remarks.—The latitude for this station may have been cited erroneously by Kemp. Squilla hieroglyphica Kemp, 1911 Squilla hieroglyphica Kemp, 1911:96 [=Al/- ima hieroglyphica (Kemp, 1911)]. Holotype. —7327/10, type locality un- known, | female. Squilla holoschista Kemp, 1911 Squilla holoschista Kemp, 1911:97 [=Ora- tosquilla holoschista (Kemp, 1911)]. Syntypes. —7926/9, Sandheads, Ganges delta, India, A. J. Milner, 1 male.—7301/ 10, Puri [19°48'N, 85°51’E], Orissa coast, India, N. Annandale, F. H. Gravely, and S. W. Kemp, 6 males, 2 females.—3147/5, Madras, Madras Museum, 7 males, 12 fe- males. —7445-—50/10, Cuddalore [11°45’N, 79°45'E] and Porto Novo [11°29’N, 79°46’E], south India, T. H. Hill, 7 males, 13 females.— 3062/5, Ceylon [=Sri Lanka], Colombo Museum, | male, 2 females (spec- imens completely fragmented). Remarks.— Although Kemp (1913) re- ported that the large lot from Madras (3147/ 5) were types, apparently all of the speci- mens reported by him are syntypes. In his original account (1911:98) he remarked that ‘In all, several hundreds of specimens have been examined.” Squilla interrupta Kemp, 1911 Squilla interrupta Kemp, 1911:98 [=Ora- tosquilla interrupta (Kemp, 1911)]. Syntypes. —9827—-8/6, Hong Kong [Hong Kong Island = 22°15’'N, 114°11’E], Hong Kong Museum, | male, 1 female. —3329/9, same locality, G. Dennys, 3 males, 3 fe- males.—4743-5/9, Singapore? [1°17'N, 103°51'E], Malaysia, Raffles Museum, 3 males. — 3095/5, Camorta [Camorta Island 658 = 8°08'N, 93°30’E], Nicobar Islands, India, F. Stoliczka, 1 female.—3094/5, Akyab [20°09'N, 92°54’E], Arakan coast, Burma, W. Dodgson, | female.—7982-91/9, Sand- heads, Hughli delta, India, Commissioners of H. M. Pilot Brigs., 5 males, 5 females. — 1284-6/7, same data, 2 males, 1 female.— 7783/10, Ghapa Natta, Mutlah River, near Calcutta, India, S. B. Nath, 1 male. —7268- 74/10, mouth of River Hughli, India, J. Munro, 5 males, 5 females.—3080/5, off Mutlah Light, Hughli River, India, G. M. Giles, 1 male.—8043/10, Puri [19°48'N, 85°51’E], Orissa coast, India, F. H.Gravely, 1 male, 1 female.—3335-7/7, off Ganjam coast, Madras, India, 7 fm (=13 m), Jn- vestigator, 3 males.—5830/9, off Vizaga- patam coast, Madras, India, 7.5—9 fm (=14— 16 m), /nvestigator [? sta 87, 10 Feb 1890], 1 male.—7557/6, 7559/9, Bombay [18°58'N, 72°50’E], India, Bombay Natural History Society, 4 males, 14 females.— 7272-8/10, Panvel Creek, Bombay, India, J. Caunter, 2 males, 4 females.— 7004/10, Karachi? [24°52’N, 67°03’E], Pakistan, Ka- rachi Museum, | male, 1 female.— 3047/5, 3067/5, Arabian Sea, Karachi Museum and A. O. Hume, 4 males, 4 females. — 647/10, Persian Gulf, F. H. Townsend, 1 female.— 7282-3/10, Persian Gulf, F. H. Townsend, 1 female.—7282-3/10, Persian Gulf, 29°20’N, 48°47’E, 13 fm (=24 m), Investi- gator [sta 352, 19 Nov 1905], 2 males.— 7279-8 1/10, same area, 28°59'N, 50°5’E, 25 fm (=46 m), Investigator [sta 349, 8 Oct 1905], 2 males, 1 female. Remarks. —Kemp (1911:99) remarked that he had examined “‘very numerous spec- imens.” Apparently all of the specimens listed by him in 1913 are syntypes, although in 1913 he considered the specimens from lot 7982-91/9 as types. Squilla investigatoris Lloyd, 1907 Squilla investigatoris Lloyd, 1907:7, 10 [=Natosquilla investigatoris (Lloyd, 1907)]. Syntypes. —5354—65/10, Arabian Sea, off PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the southeast coast of Arabia, 15°8'30’N, 51°52'15”E, 110 fm (=201 m), sandy mud, Investigator sta 364, 27 Mar 1906, 9 males, 5 females. Squilla lirata Kemp & Chopra, 1921 Squilla lirata Kemp & Chopra, 1921:303, figs. 3, 4 [=Lophosquilla lirata (Kemp & Chopra, 1921)]. Syntypes. —C 306/1, Singapore [1°17’N, 103°51'E], Malaysia, Capt. Hutcheson leg., 1 male, 1 female. Squilla mauritiana Kemp, 1913 Squilla mauritiana Kemp, 1913:66 [in syn- onymy], 68 [=Oratosquilla mauritiana (Kemp, 1913)]. Syntypes. —4269/4, Mauritius [20°18’S, 57°35'E], purchased, 1 male, | female (bad- ly mutilated). Squilla merguiensis Tiwari & Biswas, 1952 Squilla merguiensis Tiwari & Biswas, 1952: 350, fig. la [=Clorida merguiensis (Tiwari & Biswas, 1952)]. Holotype. —C 302/1, 4 miles NNE of Ka- busa Island, Burma, 33 fm (=60 m), Jn- vestigator [sta 550, 12°52'N, 97°54’E, 22 Oct 1913], 1 male. Squilla mikado Kemp & Chopra, 1921 Squilla mikado Kemp & Chopra, 1921:301, fig. 2 [=Kempina mikado (Kemp & Cho- pra, 1921)]. Holotype. —7685/10, Misaki [Misaki Harbor = 33°23’N, 132°07’E], Shikoku Is- land, Japan, A. Owston leg., | male. Paratype.—C 304/1, Misaki, Japan, K. Aoki leg., 1 male. Squilla oratoria var. perpensa Kemp, 1911 Squilla oratoria var. perpensa Kemp, 1911: 98 [=Oratosquilla perpensa (Kemp, 1911)]. VOLUME 101, NUMBER 3 Lectotype (selected by Manning, 1978: 23).—ex. 4851-72/9, Hong Kong [Hong Kong Island = 22°15’N, 114°11’E], G. Den- nys leg., 1 female. Paralectotypes. —4851-72/9, 6728/10, data same, 13 males, 16 females.—9817— 20/6, Hong Kong, Hong Kong Museum, 1 male, 5 females.—8179/7, Mergui Archi- pelago, Burma, museum collector, 1 fe- male.—5776-80/9, off Irrawaddy delta, Burma, 15°20’N, 94°55’E, 20 fm (=37 m), Investigator [sta 60, 23 Nov 1889], 2 males, 1 female.—5769/9, 6729/10, Sandheads, Hughli delta, India, A. J. Milner, 2 fe- males.— 7266-—7/10, Madras coast, India, 1 male, 1 female.—4520/10, Persian Gulf, 26°20'N, 53°54’W, 53 fm (=124 m), Jn- vestigator [sta 292, 2 Nov 1901], 1 female. Remarks. —Manning (1978:23) pointed out that Kemp’s (1913) material of this species included representatives of at least two other species, O. hindustanica Man- ning, 1978, and O. solicitans Manning, 1978. Apparently all of the specimens listed by Kemp (1913) were syntypes of S. oratoria var. perpensa; in his original description of this species (1911:98) he remarked that he had seen ““Very numerous specimens.” Squilla scorpio var. immaculata Kemp, 1913 Squilla scorpio var. immaculata Kemp, 1913:45, pl. 2, fig. 31 [=Cloridopsis im- maculata (Kemp, 1913)]. Syntypes. —3090/5, E of Terribles [19°24'N, 93°18’E], Arakan coast, Burma, 13 fm (=24 m), Investigator, 1 male.—1317/ 7, Bay of Bengal, V. Ball, 1 male.—3093/5, Calcutta [22°32’N, 88°22’'E], India, J. Wood- Mason and V. Ball, 6 males, 5 females. — 7531/10, Pratapnagar, Calcutta, India, B. L. Chaudhuri, 1 male.—7784/10,Ghaga Nata, Mutlah River, near Calcutta, India, brack- ish water, S. B. Nath, 14 males, 7 females. — 7532/10, Saugor Island, mouth of Hughli River, India, J. Munro, | male.— 1407-9/ 7, Hughli estuary, India, museum collector, 1 male, 1 female.—3091/5, Port Canning, 659 Ganges delta, India, R. D’Cruz, 1 female. — 7533/10, Karachi [24°52’N, 67°03’E], Pa- kistan, 1 male.—7528/10, locality un- known, 2 males, 4 females. Squilla stridulans Wood-Mason, 1894 Squilla stridulans Wood-Mason, in Alcock, 1894:409 [=Kempina stridulans (Wood- Mason, 1894)]. Syntypes. —7035-40/9, Masulipatam coast, Madras, India, 15°56’20’N, 81°26'10’E, 95 fm (=174 m), Investigator [sta 120, 24 Dec 1890], 6 females. —6935- 6/9, same area, 15°56’50”N, 80°30’30’E, 240 fm (=439 m), Investigator [sta 119, 24 Dec 1890], 2 females. Remarks.—In his original description, Wood-Mason (in Alcock 1894:411) listed material from two Investigator stations: sta 119, 95 fm, and sta 120, 240 fm. Specimens from these stations are syntypes, in spite of Kemp’s indication (1913:80) that two spec- imens from the Orissa coast of India were types. Squilla supplex Wood-Mason, 1875 Squilla supplex Wood-Mason, 1875:232 [=Alimopsis supplex (Wood-Mason, 1875)]. Holotype. — 3048/5, Bombay [18°58’N, 72°50’E], India, F. Stolizcka, 1 male (deli- cate). Squilla tenuispinis Wood-Mason, 1891 Squilla tenuispinis Wood-Mason, in Wood- Mason & Alcock 1891:271 [=Squilloides tenuispinis (Wood-Mason, 1891)]. Syntypes.—5801/9, off Ganjam coast, Madras, India, 18°40’N, 84°46’E, 98-102 fm (=179-187 m), Investigator [sta 96, 18°30’N, 4 Mar 1890], 1 male.—3081/5, off Cheduba [Cheduba Island = 18°48'N, 93°38’E], Arakan coast, Burma, Jnvesti- gator, | female. Remarks.—In the original description, Wood-Mason (in Wood-Mason & Alcock 1891:272) mentioned two specimens taken 660 by the Jnvestigator: one from sta 96 in 90 to 100 fm and one from off Cheduba [Is- land], Arakan coast of Burma, in about the same depth. Of the specimens listed by Kemp (1913:49), these two specimens are syntypes; Kemp listed them as types. The latitude given by Kemp for the ma- terial from sta 96 is incorrect by 10 minutes. Squilla wood-masoni Kemp, 1911 Squilla wood-masoni Kemp, 1911:99 [=Or- atosquilla woodmasoni (Kemp, 1911)]. Syntypes. —7513/10, Port Jackson [33°51’S, 151°15’E], New South Wales, Australia, Australian Museum, | female. — 7294/10, Pondicherry, India, purchased, 1 female.—3098/5, Madras’ [13°05’N, 80°17’E], India, 3 males, 2 females. —3078/ 5, 3087/5, Madras, India, Madras Museum and purchased, 2 males.—7295/10, off Madras coast, India, 14°4’57”N, 80°20'50’E, 23 fm (=42 m), Investigator [sta 158, 10 Jan 1894], 1 female.—7297-8/10, Puri [19°48'N, 85°51’E], Orissa coast, India, N. Annandale, F. H. Gravely, and J. Caunter, 9 males, 3 females.—7992/9, Sandheads, Ganges delta, India, W. M. Daly, 1 female. Remarks. —In his original account, Kemp (1911:99) stated that he had seen ‘“‘Numer- ous examples.”’ Apparently all of the spec- imens listed by him in 1913 are syntypes. Types of Stomatopoda in the Zoological Survey of India, Listed by Current Name Acanthosquilla multifasciata (Wood-Ma- son, 1895) See Lysiosquilla multifasciata Alima hieroglyphica (Kemp, 1911) See Squilla hieroglyphica Alimopsis supplex (Wood-Mason, 1875) See Squilla supplex Busquilla quadraticauda (Fukuda, 1911) See Squilla boopis Chorisquilla excavata (Miers, 1880) See Chorisquilla andamanica Clorida bombayensis (Chhapgar & Sane, 1967) See Squilla bombayensis PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Clorida decorata Wood-Mason, 1875 Clorida denticauda (Chhapgar & Sane, 1967) See Squilla denticauda Clorida merguiensis (Tiwari & Biswas, 1952) See Squilla merguiensis Cloridopsis bengalensis (Tiwari & Biswas, 1952) See Squilla bengalensis Cloridopsis immaculata (Kemp, 1913) See Squilla scorpio var. immaculata Dictyosquilla foveolata (Wood-Mason, 1895) See Squilla foveolata Gonodactylus choprai Manning, 1967 Gonodactylus lanchesteri Manning, 1967 Gonodactylus platysoma Wood-Mason, - 1895 See Gonodactylus chiragra var. platyso- ma Haptosquilla glyptocercus (Wood-Mason, 1875) See Gonodactylus glyptocercus Haptosquilla nefanda (Kemp, 1911) See Gonodactylus nefandus Haptosquilla proxima (Kemp, 1911) See Gonodactylus proximus Harpiosquilla annandalei (Kemp, 1911) See Squilla annandalei Heterosquilla tricarinata (Claus, 1871) See Coronis spinosa Heterosquilloides insignis (Kemp, 1911) See Lysiosquilla insignis Kempina mikado (Kemp & Chopra, 1921) See Squilla mikado Kempina stridulans (Wood-Mason, 1894) See Squilla stridulans Lenisquilla gilesi (Kemp, 1911) See Squilla gilesi Lophosquilla lirata (Kemp & Chopra, 1921) See Squilla lirata Lophosquilla tiwarii Blumstein, 1974 Lysiosquilla sulcirostris (Kemp, 1913) See Lysiosquilla maculata var. sulcirostris Manningia andamanensis Ghosh, 1975 Mesacturus furcicaudatus (Miers, 1880) See Gonodactylus furcicaudatus Natosquilla investigatoris (Lloyd, 1907) See Squilla investigatoris Odontodactylus brevirostris (Miers, 1884) See Odontodactylus southwelli VOLUME 101, NUMBER 3 Oratosquilla gonypetes (Kemp, 1911) See Squilla gonypetes Oratosquilla hindustanica Manning, 1978 Oratosquilla holoschista (Kemp, 1911) See Squilla holoschista Oratosquilla interrupta (Kemp, 1911) See Squilla interrupta Oratosquilla mauritiana (Kemp, 1913) See Squilla mauritiana Oratosquilla pentadactyla Manning, 1978 Oratosquilla perpensa (Kemp, 1911) See Squilla oratoria var. perpensa Oratosquilla subtilis Manning, 1978 Oratosquilla woodmasoni (Kemp, 1911) See Squilla woodmasoni Squilla brasiliensis Calman, 1917 Squilloides tenuispinis (Wood-Mason, 1894) See Squilla tenuispinis Literature Cited Alcock, A. 1894. On the results of the deep-sea dredg- ing during the season 1890-9 1 (concluded). Nat- ural history notes from H.M. Indian Marine Survey Steamer ‘Investigator,’ Commander R. F. Hoskyn, R.N., late commanding, series 2, no. 1.—Annals and Magazine of Natural History (6)13:4C0-411. Blumstein, R. 1974. Stomatopod crustaceans from the Gulf of Tonkin with the description of new species. —Crustaceana 26(2):113-126. Calman, W. T. 1917. Stomatopoda, Cumacea, Phyl- locarida, and Cladocera. Crustacea, Part [IV.— British Antarctic Terra Nova Expedition, 1910, Natural History Report, Zoology 3(5):137-162. Chhapgar, B. F., & S.R. Sane. 1967. Two new species of Squilla (Stomatopoda) from Bombay. — Crus- taceana 12(1):1-8. Ghosh, H. C. 1975. A new species of Manningia (Stomatopoda, Gonodactylidae) from the An- daman Islands.—Crustaceana 28(1):33-36. Holthuis, L. B. 1967. Fam. Lysiosquillidae et Bathy- squillidae: Stomatopoda I., in H.-E. Gruner and L. B. Holthuis, eds., Crustaceorum Catalogus 1: 1-28. The Hague, W. Junk. Kemp, S. 1911. Preliminary descriptions of new species and varieties of Crustacea Stomatopoda in the Indian Museum.—Records of the Indian Museum 6(2):93-100. 1913. An account of the Crustacea Stoma- topoda of the Indo-Pacific region, based on the collection in the Indian Museum.— Memoirs of the Indian Museum 4:1—217, pls. 1-10. 661 1915. Ona collection of stomatopod Crus- tacea from the Philippine Islands. — Philippine Journal of Science (D)10(3):169-187, pl. 1. ——., & B. Chopra. 1921. Notes on Stomatopo- da.— Records of the Indian Museum 22(4):297-— 311. Lloyd, R. E. 1907. Contributions to the fauna of the Arabian Sea, with descriptions of new fishes and Crustacea.— Records of the Indian Museum 1: 1-12. Manning, R. B. 1967. Notes on the demanii section of genus Gonodactylus Berthold with descrip- tions of three new species (Crustacea: Stoma- topoda).— Proceedings of the United States Na- tional Museum 123(2618):1-27. . 1975. Two new species of the Indo-West Pa- cific genus Chorisquilla (Crustacea, Stomato- poda), with notes on C. excavata (Miers). — Pro- ceedings of the Biological Society of Washington 88:253-261. 1978. Further observations on Oratosquilla, with accounts of two new genera and nine new species (Crustacea: Stomatopoda: Squillidae).— Smithsonian Contributions to Zoology 272:iii + 44 pp. Miers, E. J. 1880. On the Squillidae.—Annals and Magazine of Natural History (5)5:1-30, 108- 127, pls. 1-3. 1884. Crustacea. Report of the zoological collections made in the Indo-Pacific Ocean dur- ing the voyage of H.MLS. “‘Alert,” 188 1-82:178- 322, 513-575, pls. 18-35, 46-52. B.M., Lon- don. Tiwari, K. K., & S. Biswas. 1952. On two new species of the genus Squilla Fabr., with notes on other stomatopods on the collections of the Zoological Survey of India.—Records of the Indian Mu- seum 49(3-4):349-363. Wood-Mason, J. 1875. On new or little-known crus- taceans.— Proceedings of the Asiatic Society of Bengal 1875:230-232. . 1895. Figures and descriptions of nine species of Squillidae from the collection in the Indian Museum, pp. 1-11, pls. 1-4. Calcutta: Indian Museum. —, &A. Alcock. 1891. Note on the result of the last season’s deep-sea dredging. Natural history notes from H.M. Indian Marine Survey Steamer ‘Investigator,’ Commander R. F. Hoskyn, R.N., commading.— Annals and Magazine of Natural History (6):7:258-272. (HCG) Zoological Survey of India, 27 Ja- waharlal Nehru Road, Calcutta 700 016, India; (RBM) Department of Invertebrate Zoology, National Museum of Natural His- tory, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 662-670 PYCNOGONIDA OF THE WESTERN PACIFIC ISLANDS IV. ON SOME SPECIES FROM THE RYUKYU ISLANDS K. Nakamura and C. Allan Child Abstract.—A small collection of Pycnogonida from the southern Ryukyus and miscellaneous specimens from Okinawa are recorded with their zooge- ography. One new species, Callipallene tridens, is described and figured and 11 known species are recorded for the first time in the Ryukyus. A high incidence of juveniles and larvae suggests that the months of collection, May and June, may be the peak of the breeding and hatching season in these islands. Only two species of pycnogonids were known from the Ryukyu Islands prior to this report. The specimens reported here were taken in the Sakishima Group of the southern Ryukyus and from Okinawa. The first pycnogonid described from Okinawa was Pycnogonum benokianum Ohshima (1935:137-139), from Benoki (=Benochi, 26°48’N, 128°14’E). The second species re- ported from the Ryukyus was Endeis mollis (Carpenter), also by Ohshima (1935:139), taken at Ishigaki Island in the Sakishima Group south of Okinawa. Specimens of E. mollis are reported from the Sakishima Group, but P. benokianum has not been recorded since the type specimens were de- scribed. Hedgpeth (1949:304—397, fig. 49) subsequently redescribed this species from Ohshima’s notes and a preserved type spec- imen in his paper on collections taken in Japan by the Albatross. Utinomi (1971:327) also repeated Ohshima’s record in his list of Japanese pycnogonids. There are now 13 species known from the Ryukyus with one of them, Callipallene tri- dens, described herein as new. At least two and probably three other species are rep- resented in these collections but the ques- tionable specimens are juveniles or larvae and do not have characters sufficiently de- veloped for recognition or description. The known Ryukyu Islands species have their strongest faunal affinity with the Phil- ippines where six of the species are also known, suggesting some form of transport by shallow northward-flowing currents. Three of the species are also found in Japan while three are found to the south in In- donesia and beyond. There are also three species found in the Marshall, Hawaii, and Samoa Island groups while a more distant distribution occurs for two species recently described from Aldabra Atoll in the Indian Ocean. Two other widely distributed species are known from the western Indian Ocean while one additional species, Pigrogromitus timsanus, 1s known to be pantropical. The distribution of most of these species is so incompletely known that their true distri- bution can not be suggested at this time. They are all shallow-water species. Family Phoxichilidiidae Genus Anoplodactylus Wilson, 1878 Anoplodactylus pectinus Hedgpeth Anoplodactylus pectinus Hedgpeth, 1948: 234-236, fig. 34.—Child, 1982a:372-—373 [literature]; 1988a:20. Material examined. —Sakishima: Funau- ki Bay, Iriomote Island, 5 m, 7 Jun 1987; 1 6 with eggs, 2 2, 1 juv. Southwest Ishigaki Harbor, Ishigaki Island, 24 m, 10 Jun 1987; 1 3. VOLUME 101, NUMBER 3 Distribution.—This species was first de- scribed from specimens taken in the Florida Keys and is now known from many local- ities in Florida and the Caribbean. It has also been taken in Madagascar and more recently in the northern Philippines. Its oc- currence in the southern Ryukyus consti- tutes only a small range extension from the Philippines, but the range of collections in which the species appears suggests that it may be another pantropical species. All known depths of capture are shallow, the deepest being 27 meters. Remarks. —The moderately large size (leg span about 12 mm) of this species in rela- tion to other tropical members of this genus, its slender appearance, and the fairly long cribriform cement gland slit of the male along with the serrate major heel spine in both sexes all serve to make this an easily identified species. Anoplodactylus perforatus Nakamura & Child Anoplodactylus perforatus Nakamura & Child, 1982:288-291, fig. 3; 1983:49; (in press). Material examined. —Sakishima: south- west Ishigaki Harbor, Ishigaki Island, 8-12 m, 4 Jun 1987; 1 4, 1 2. Distribution. —This species is often taken in large numbers per capture effort in west- ern Sagami Bay, Japan, while another sim- ilar large aggregation was recently reported from Western Samoa. These 2 Ryukyu specimens serve to help bridge the wide dis- tributional gap between Japan and Samoa while contributing nothing new to the known depth range of 7-113 meters. Remarks. —This is an easily recognized species, at least in the male, by its many cement gland pores (17 to 25) per femur and the long slender oviger having an incon- gruously tiny terminal segment. Both sexes have a very short propodal lamina and long chelae fingers lacking teeth. 663 Anoplodactylus, species indeterminate Material examined. —Sakishima: Fu- naura Bay, Iriomote Island, night, at sur- face, 10 May 1986; 2 juv, 1 larva. Same locality, depth, 3 Jun 1986; 4 juv, 5 larvae. Funauki Bay, Iriomote Island, 5 m, 7 Jun 1987; 1 larva. Iriomote Island, 11-20 m, 7 Jun 1987; 1 2? juv. Southwest Ishigaki Har- bor, Ishigaki Island, 2-4 m, 10 Jun 1987: 1 Q. Remarks. —There are at least two species represented by these females and young but none of them is sufficiently developed or was taken with adult males to permit iden- tification. The relatively large number of Anoplodactylus young in relation to the total number of pycnogonids taken during col- lecting in the Sakishima Islands group is possibly significant in suggesting that May and June represent the breeding-hatching season for the genus. Family Endeidae Genus Endeis Philippi, 1843 Endeis nodosa Hilton Endeis (Phoxichilus) nodosa Hilton, 1942: 47-48, fig. 4. Endeis nodosa.—Stock, 1968:59 [key].— Child, 1982b:275—277, fig. 2g—i. Material examined. —Okinawa: Naha Harbor, on breakwater, 1.5 m, 27 Jun 1987; 1 subadult. Distribution.—This species is rare be- cause it is found on islands seldom or never exposed to collecting for marine microin- vertebrates. Its type locality is Hawaii and it has subsequently been taken at Enewetak Atoll in the Marshall Islands. Okinawa is only the third known locality where it has been found and it is apparently not common in any of these places. All known collecting depths are littoral or very shallow waters. Remarks.—This species is fairly easily distinguished from others of the genus by the presence of a large node or low conical tubercle on the midventral femur and for 664 the many diverticula or blind pockets of the central intestinal branches in the legs. The integument is usually opaque so that the gut branches and diverticula are easily seen. There are sometimes many more finger-like diverticula on the intestines than are figured by Child (1982b:276, fig. 2h). Endeis mollis (Carpenter) Phoxichilus mollis Carpenter, 1904:182- 183, figs. 1-7. Endeis mollis.—Utinomi, 1971:327 [liter- ature].—Stock, 1975a:1083-1085; 1975b: 76.—Child, 1979:66.—Nakamura & Child, 1983:41.—Stock, 1986:440.— Child, 1988a:20-21. Material examined. —Sakishima: Fu- naura Bay, Iriomote Island, 3 Jun 1986; 2 juv. Same locality and date, night, at sur- face; 1 juv. Distribution. —This is a pantropical-tem- perate species found as far north as Sagami Bay, Japan. Ohshima reported it from the Sakishima Group at Ishigaki Island (Oh- shima 1935:139), the only pycnogonid found in the southern Ryukyus, but it could also be expected to inhabit Okinawa. It is a shal- low-water species, having been taken from the shoreline down to 60 meters. Remarks. —This species does not have the small side pockets along its gut diverticula in each leg nor does it have the conspicuous conical ventral tubercle on the femorae as does E. nodosa. Family Ammotheidae Genus Eurycyde Schiddte, 1857 Eurycyde, species indeterminate Material examined. —Sakishima: Irio- mote Island, 11—20 m, 7 Jun 1987; 1 juv. Remarks. — This juvenile specimen is in- determinable although it has several char- acters reminiscent of Eurycyde setosa Child (1988a:8-10, fig. 3), from the Philippines. This recently described species has 7 large spines at the ocular tubercle apex, but the abdomen of the unique type is missing. The PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Iriomote specimen has 4—5 large spines at the ocular apex and several similar spines on the curved abdomen. The lateral pro- cesses are also closely spaced as in E. setosa. The proboscis of this juvenile has not yet divided into the characteristic 2 segments and the specimen otherwise has no visible identifying characters. Family Callipallenidae Genus Callipallene Flynn, 1929 Callipallene novaezealandiae (Thomson) Pallene novae-zealandiae Thomson, 1884: 246-247, pl. 14, figs. 1-4. Callipallene novaezealandiae. —Child, 1983: ° 708 [literature]. Material examined. —Sakishima: Fu- naura Bay, Iriomote Island, night, at sur- face, 3 Jun 1986; 2 3, 1 juv. Southwest Ishi- gaki Harbor, Ishigaki Island, 3 m, 4 Jun 1987; 4 6 with eggs, 2 6, 4 2, 8 juv, 15 larvae. Same locality, 2-4 m, 10 Jun 1987; 2 4, 3 2, 7 juv, | larva. Distribution. —This species is known from eastern Africa, Australia, Japan, New Zea- land, and from Palau and Enewetak Atoll in Micronesia, all from shallow localities. This series of specimens extends its distri- bution to an intermediate locality between Japan and islands to the south and east. Remarks. —These specimens, particular- ly the males, have the same variation in Oviger denticulate spine counts per segment as seen in the Micronesian specimens, but otherwise agree with these specimens in di- agnostic characters. Callipallene tridens, new species Fig. 1 Material examined. —Sakishima: south- west Ishigaki Harbor, Ishigaki Island, 3 m, 4 Jun 1987; 1 6 with eggs (holotype, USNM 234430), 5 6, 5 2, 2 juv (paratypes, USNM 234431). Description. —Size very tiny, leg span only 3.6 mm. Trunk fairly elongate, segmenta- tion lines faint, incomplete. Lateral pro- VOLUME 101, NUMBER 3 cesses no longer than wide, distally sepa- rated by their own diameter or less, glabrous. Neck very short, without parallel sides, ex- pansion before chelifores no wider than long. Ocular tubercle very broad-based, only half as tall as basal width, eyes large, slightly pigmented, lateral papillae present, not prominent. Proboscis short, 1.5 times long- er than diameter, with low rounded ven- trodistal bulges, constricted distally just posterior to flat lips. Abdomen very short, less than length of 4th lateral processes, dis- tally rounded, armed with 2 short lateral setae. Chelifore scape moderately slender, 3 times longer than its diameter, armed with 3 dorso- and laterodistal setae longer than scape diameter. Chela palm subtriangular, with 10-11 ectal and lateral setae longer than palm diameter. Fingers longer than palm, carried anaxially, slender, overlap strongly at tips, armed with 4—5 long sharp teeth on immovable finger and 6 similar teeth on movable finger. Movable finger longer and more curved than immovable finger. Oviger short, 4th segment only slightly longer than 3rd, 0.6 as long as 5th which is armed with rounded distal apophysis bear- ing 2-3 short setae. Strigilis segments each shorter than last, armed with 1-2 ectal short setae distally and dimorphic denticulate spines endally in the formula 5:5:5:6. Prox- imal spines with many tiny lateral serra- tions, distal serrations slightly larger; distal spines with tiny serrations on proximal side, long finger-like serrations on distal side of spines. Egg size very large, 4 times larger than 5th segment diameter. Legs fairly short, slightly inflated, cement gland tubes or pores not evident. Second and 3rd coxae armed with very long ven- trodistal setae, each 2—3 times longer than segment diameter. Femur with slight ven- trodistal bend, armed with several distal and lateral setae not as long as segment diam- eter. Tibiae slightly shorter than femur, Ist tibia armed with few lateral and distal setae shorter than segment diameter, 2nd tibia 665 with more setae, some longer than segment diameter. Tarsus very short, armed with dorsal and ventral setae, some longer than segment diameter. Propodus slightly inflat- ed, straight, without heel but with 4 stout heel spines and 7-8 sole spines, very short distal lamina, and few lateral and distal se- tae. Claw moderately curved, half propodal length, with very short auxiliary claws shaped like tridents bearing broad pointed spatulate median spine flanked by 1 or 2 tiny pointed lateral serrations. Auxiliaries slightly inflated, curved in lateral view. Female paratype: propodal claw longer, 0.7 propodus length, more curved distally. Trunk and legs slightly larger than those of male, oviger reduced, without distal apoph- ysis on 5th segment. Measurements (holotype in mm).— Trunk length, 0.6; trunk width (across 2nd lateral processes), 0.29; proboscis length, 0.2; ab- domen length, 0.05; third leg, coxa 1, 0.08; coxa 2, 0.19; coxa 3, 0.1; femur, 0.33; tibia 1, 0.29; tibia 2, 0.31; tarsus, 0.04; propodus, 0.22; claw, 0.11. Distribution. —Known only from the type locality, Ishigaki Harbor, Ishigaki Island, in the southern Ryukyus, in 3 meters. Etymology.—The name tridens (Latin: forked with 3 tines) refers to the 3-lobed “trident” auxiliary claws of this species. Remarks. — The presence of short trident- like auxiliary claws is unique among known pycnogonid species. Auxiliary claws are used as a usually reliable diagnostic character among pycnogonid genera and some have them while some do not. They are usually simple curved claws, but some species have tiny endal setules or spinules. These are most often found among Callipallene species and these trident-like claws add another form to the seemingly endless variety of characters. This new species does not conform in oth- er ways to most Callipallene species; the chelae are atypical in having very slender fingers placed anaxially on the palm and bearing long slender teeth, the ocular tu- bercle is broader based and is shorter than 666 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Callipallene tridens, holotype male: A, Trunk, dorsal view; B, Trunk, lateral view; C, Third leg; D, Distal segments of third leg with auxiliary claw enlarged; E, Chela; F, Oviger with single egg attached; G, Oviger distal and proximal denticulate spines, enlarged; H, Oviger terminal segment, distal spine enlarged. VOLUME 101, NUMBER 3 in almost any other species known, and the propodus has a short distal sole lamina, a character unknown to this genus but com- mon among species of the genus Anoplo- dactylus. These atypical and unique char- acters are sufficient to permit description of this as a new species and they continue to expand the diversity of characters found among the Pycnogonida. Genus Cheilopallene Stock, 1955 Cheilopallene hirta Child Cheilopallene hirta Child, 1988b:63-66, fig. 5. Material examined. —Sakishima: Fu- naura Bay, Iriomote Island, night, at sur- face, 3 Jun 1986; 1 4, 1 2. Funauki Bay, Iriomote Island, 5 m, 7 Jun 1987; 1 Q. Southwest Ishigaki Harbor, Ishigaki Island, 5 m, 4 Jun 1987; 1 juv. Distribution. —This species is known only from Aldabra Atoll, western Indian Ocean, its type locality, in 0-21 meters. The Saki- shima specimens greatly extend this distri- bution eastward into the Pacific and north- ward to the southern Ryukyus in similarly shallow waters. Remarks. — There is remarkably little dif- ference between these specimens and the type series. The male fourth oviger segment is very slightly longer than with the Aldabra specimens and there are the usual 1—2 spine differences in the denticulate spine counts among specimens. The four stout spines next to the chela immovable finger of the type are reduced to three with several additional slender setae on the adjacent raised pad, but the specimens agree exactly otherwise, in- cluding the hirsute distal proboscis. Genus Pigrogromitus Calman, 1927 Pigrogromitus timsanus Calman Pigrogromitus timsanus Calman, 1927:408- 410, fig. 104a-f. — Child, 1982a:367-368; 1988a:21 [literature]. Material examined.—Okinawa: Maeda 667 Misaki, reef flat, on Lybia tesellata [Xan- thidae: Crustacea], 15 Nov 1985; 1 juv. Distribution.—This species has a pan- tropical but scattered distribution in many varied localities. It has been taken in Hawaii and is known from the Philippines, so it is not surprising to find it in Okinawan waters. It inhabits shallow depths, the deepest cap- ture being 108 meters. Remarks.—This is an easily recognized species in a monotypic genus. It superfi- cially looks like several species of the genus Pycnogonum in habitus, but has chelifores with small functional chelae, and has an Oviger without denticulate spines but with a terminal claw having a proximal spine in a forcipulate arrangement with the large claw. Live specimens of this species have a uniform whitish-tan coloration. Genus Propallene Schimkewitsch, 1909 Propallene curtipalpus Child Propallene curtipalpus Child, 1988a:21-—23, fig. 9. Material examined. —Sakishima: Irio- mote Island, 11-20 m, 7 Jun 1987; 2 4, 1 2, 2 juv. Distribution. —This species was originally described from material taken in the Visa- yan Islands, central Philippines, from depths of 2 meters or less. The presence of the species in the southern Ryukyus is not sur- prising considering the supposed corridor of pycnogonid genera thought to inhabit is- lands from New Zealand north along the western Pacific archipelagoes as far as Japan (Child 1983:713). The corridor hypothesis was originally proposed to include the pre- dominantly Antarctic genus Austrodecus which has several species found from New Zealand to Japan. It now appears that the corridor should include other genera, among them Propallene, with the majority of its species found from the Indian Ocean (in- cluding South Africa) and Australia north along the corridor to Japan. The genus Sci- piolus with representatives from South Af- 668 rica, Indonesia, and Japan, and the genus Hemichela having species known from Aus- tralia, Indonesia, and the Philippines, should possibly be included as members of this cor- ridor fauna. There are several endemic gen- era known to the East Indies, but none of the above genera nor any families are con- fined to the western Pacific corridor as en- demics although the majority of their species are known from parts of the corridor and Indian Ocean-Australian localities. Our knowledge of the pycnogonids of this vast region remains very fragmentary and the distribution of any of these genera may be greatly expanded to more distant parts of the Pacific in future collecting. The known depth at which this species has been taken is now extended to 20 meters by the Ryukyu records. It has not been taken on Okinawa Island. Remarks. —This is the only known species in a genus with very look-alike members to have a reduced number of cement gland tubes (4-5) arranged on the femur (1) and both tibiae (1-2 each), and a very reduced palp which is less than half the proboscis length. Where specimens are sympatric, they can not be differentiated to species with only females in hand. The males with palps and cement glands bear the only characters with which to separate any two species. Necks have some variation in length among a large suite of females. The only instance of sym- patry in this genus, to our knowledge, is in P. longiceps and P. saengeri, both of which appear in Sagami Bay collections (Naka- mura & Child 1983:61-62). Propallene longiceps (BOhm) Pallene longiceps Bohm, 1879:59-60. Propallene longiceps. —Nakamura & Child, 1983:61-62 [literature].—Kim & Hong, 1986:41.—Hong & Kim, 1987:158. Material examined. —Okinawa: Heanza Shima, 30 Apr 1987; 11 6 with eggs. Distribution. —This species was believed PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON to be endemic to the main Japanese Islands until it was reported from the Korean coast by Kim & Hong (1986:41). This record is the first for the species from the Ryukyus. Further collecting in these many islands will possibly yield other endemics from the main islands to the north. Remarks.—One of the most common pycnogonids in Sagami Bay, this species has more than 10 cement gland tubes per leg which are usually confined to the ventral femur and has a palp almost as long as the proboscis with a constriction in the second or longest segment. Family Nymphonidae Genus Nymphon Fabricius, 1794 Nymphon diabolus Child Nymphon diabolus Child, 1988a:23-25, fig. 10; 1988b:75. Material examined. —Sakishima: Fu- naura Bay, Iriomote Island, night, at sur- face, 3 Jun 1986; 1 2. Distribution. — This species was described from specimens taken in Negros Oriental Province, central Philippines, and was sub- sequently found on Aldabra Atoll, Indian Ocean. All collecting depths are very shal- low. Its distribution is extended northward to the Ryukyus by the above record. This is the first known record of the species being taken in a night plankton haul at the surface. Remarks.—This specimen is very much like the adult female listed under other ma- terial from the Philippines. The species is notable for its reduced anterior size (chelae, proboscis and palps) in relation to its trunk, for the “horns” at the ocular tubercle apex, for the short main and auxiliary claws as related to the length of the propodus, and for the slender graceful chelae having long fingers bearing from 9 to 12 long pointed teeth. This female has the marked horns of the Philippine specimen, but there are slight variations in the tarsus-propodus ratio and the oviger denticulate spine count. VOLUME 101, NUMBER 3 Nymphon, species indeterminate Material examined. —Sakishima: Irio- mote Island, 11-20 m, 7 Jun 1987; 1 juv. Remarks.—This juvenile is close to N. micronesicum Child, but is insufficiently de- veloped for determination. It has the tiny lateral tubercles on the ocular tubercle, but without horns, and has short auxiliaries and a main claw with rugosities. The bifurcate chelae teeth are on much more slender fin- gers which strongly overlap at the tips, un- like those of the type specimen. The ovigers are not sufficiently developed to permit comparison. Family Austrodecidae Genus Austrodecus Hodgson, 1907 Austrodecus tubiferum Stock Austrodecus gordonae (part).—Stock, 1954: 153, fig. 76e. Austrodecus tubiferum Stock, 1957:75-77, fig. 43. Material examined.—Okinawa: Horse- shoe Cliffs, 1 km WNW of Onna Village, 73.2 m, coll. R. F. Bolland, 28 Sep 1981; 1g. Distribution. — This species is only known from off Okinose in Sagami Bay, Japan, at 183 meters (type locality) and the above specimen extends this distribution south to the Ryukyus in shallower water. Remarks.—This female conforms very well to the figures given for the type (a male), except that the median trunk tubercles and those of the first coxae are smaller, the ovi- ger 1s very similar, and the palp has the deep cleft in its terminal segment although there is apparently no indication of a suture line beyond the cleft as shown for the type (Stock 1957:76, fig. 43c). Acknowledgments We wish to thank Mr. Hiroaki Akishige, captain of the Kakuyo Maru, and the crew of the vessel for their many kindnesses to 669 the first author (K.N.) in collecting speci- mens, Mr. Nobuhiro Ooshiro of the Yae- yama Branch of the Okinawa Prefectural Fisheries Experimental Station for pycno- gonids from his night tow samples, and Mr. Norikazu Shikatani of the Department of Marine Science, University of the Ryukyus, for bringing specimens from sortings of Fishery Bureau material to the attention of one of us (K.N.). All specimens are deposited in the Na- tional Museum of Natural History, Smith- sonian Institution, under the catalog num- bers of the old U. S. National Museum system. Literature Cited Bohm, R. 1879. Ueber zwei neue von Dr. Hilgendorf in Japan gesammelte Pycnogoniden. —Sitzungs- berichte der Gesellschaft Naturforschender Freunde zu Berlin 1879 (4):53-60, figs. ac. Calman, W.T. 1927. 28. Report on the Pycnogonida. Zoological Results of the Cambridge Expedition to the Suez Canal, 1924.—Transactions of the Zoological Society of London 22(3):403-410, 3 figs. Carpenter, G. H. 1904. Report on the Pantopoda collected by Prof. Herdman at Ceylon in 1902.— Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, 2, Supplementary Report XIII:181-—184, 1 pl. Child, C. A. 1979. Pycnogonida of the Isthmus of Panama and the Coasts of Middle America. — Smithsonian Contributions to Zoology, 293:1- 86, 25 figs. 1982a. Pycnogonida from Carrie Bow Cay, Belize. Jn K. Rutzler and I. G. Macintyre, eds., The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, 1: Structure and Communi- ties. —Smithsonian Contributions to the Marine Sciences 12:355—380, figs. 163-167. 1982b. Pycnogonida of the Western Pacific Islands I. The Marshall Islands.— Proceedings of the Biological Society of Washington 95(2): 270-281, 3 figs. 1983. Pycnogonida of the Western Pacific Islands II. Guam and the Palau Islands. —Pro- ceedings of the Biological Society of Washington 96(4):698-714, 5 figs. 1988a. Pycnogonida of the Western Pacific Islands III. Recent Smithsonian-Philippine Ex- peditions.—Smithsonian Contributions to Zo- ology 468:1—32, 12 figs. 670 1988b. Pycnogonida from Aldabra Atoll.— Bulletin of the Biological Society of Washington 8:45-78, 9 figs. Hedgpeth, J.W. 1948. The Pycnogonida of the West- ern North Atlantic and the Caribbean.—Pro- ceedings of the United States National Museum 97(3216):157—342, 49 figs., 3 charts. 1949. Report on the Pycnogonida collected by the Albatross in Japanese waters in 1900 and 1906.—Proceedings of the United States Na- tional Museum 98(3231):233-321, 33 figs., 1 chart. Hilton, W. A. 1942. Pycnogonids from Hawaii.— Occasional Papers of the Bernice P. Bishop Mu- seum 17:43-55, 10 figs. Hong, J. S., & I.H. Kim. 1987. Korean Pycnogonids chiefly based on the collections of the Korea Ocean Research and Development Institute. — Korean Journal of Systematic Zoology 3(2):137— 164, 17 figs. Kim, I. H., & J. S. Hong. 1986. Korean shallow- water pycnogonids based on the collections of the Korea Ocean Research and Development Institute.—Korean Journal of Systematic Zo- ology 2(2):35-52, 9 figs. Nakamura, K., & C. A. Child. 1982. Three new species of Pycnogonida from Sagami Bay, Japan.— Pro- ceedings of the Biological Society of Washington 95(2):282-291, 4 figs. SS & . 1983. Shallow-water Pycnogonida from the Izu Peninsula, Japan.— Smithsonian Contributions to Zoology 386:1-71, 21 figs. ——., & (in press). Pycnogonida of the Western Pacific Islands V. A Collection by the Kakuyo Maru from Samoa. Proceedings of the Biological Society of Washington. Ohshima, H. 1935. On a sea spider inhabiting the Okinawa region. — Dobutsugaku Zasshi 47:137— 139. (Zoological Society of Japan, in Japanese.) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Stock, J. H. 1954. Pycnogonida from the Indo-West Pacific, Australian and New Zealand waters. — Videnskabelige Meddelelser fra Dansk Natur- historisk Forening (Kobenhaven) 116:1-168, 81 figs. 1957. The pycnogonid family Austrodeci- dae.— Beaufortia 6(68):1-81, 43 figs. . 1968. Pycnogonida collected by the Galathea and Anton Bruun in the Indian and Pacific oceans. — Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening (Kobenhaven) 131:7- 65, 22 figs. . 1975. Pycnogonida from the continental shelf, slope, and deep sea of the tropical Atlantic and East Pacific. Biological results of the University of Miami deep-sea expeditions, 108.— Bulletin of Marine Science, 24(4):957—-1092, 59 figs. 1986. Pycnogonida from the Caribbean and the Straits of Florida. Biological results of the University of Miami deep-sea expeditions. — Bulletin of Marine Science, 38(3):399-441, 15 figs. Thomson, G. M. 1884. On the New Zealand Pyc- nogonida, with descriptions of new species. — Transactions & Proceedings of the New Zealand Institute 16(1883):242—248, 3 pls. Utinomi, H. 1971. Records of Pycnogonida from shallow waters of Japan.— Publications of the Seto Marine Biological Laboratory, 18(5):317— 347. (KN) Ocean Research Institute, Univer- sity of Tokyo, 1-15-1 Minamidai, Nakano- ku, Tokyo 164, Japan; (CAC) Department of Invertebrate Zoology (Crustacea), Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 671-675 POECILOCHAETUS KOSHIKIENSIS, A NEW POLY CHAETE SPECIES FROM SHIMO-KOSHIKI ISLAND, JAPAN Tomoyuki Miura Abstract. —A new species of polychaetous annelid, Poecilochaetus koshikien- sis, collected at a depth of 200 m off Shimo-Koshiki Island, Kyushu, southern Japan, is described. The new species is characterized by posterior notopodial hooks and basally fused anal cirri. The genus Poecilochaetus Claparéde, 1875 consists of 16 nominal species (Pilato & Cantone 1976, Read 1986). Their main di- agnostic characters include the epidermal smoothness, the distribution of flask-shaped (ampullaceous) postsetal lobes (dorsal and ventral cirri), the length and number of nu- chal organs, and the occurrence and distri- bution of several types of setae. Despite the small number of species and the large in- terspecific variability in morphology, some difficulties in the identification of species arise from the insufficient descriptions of most species, their intraspecific variability (Milligan & Gilbert 1984) and the confused terminology of the setae. For example, “‘plu- mose setae”’ called by Hartman (1939) cor- responds to “‘pinnate setae” by Milligan & Gilbert (1984), while “‘plumose setae” by the latter refers to “‘spinose setae” by the former. In this study, the terminology of setae by Read (1986, Table 2) is chiefly fol- lowed. The worms are very fragile and com- plete specimens are rarely collected, even from shallow bottoms. The extensive mor- phology and habitat were described only on few particular species, such as Poecilochae- tus serpens Claparéde from England by AI- len (1904), P. australis Nonato from Brazil by Nonato (1963), and P. johnsoni Hartman from Florida by Taylor (1966). Only a single species, Poecilochaetus ja- ponicus Kitamori, 1965, has been reported from Japanese coasts. In this study, a new species of Poecilochaetus is described based on a complete female and two additional mature specimens collected in Japan. The types are deposited in the National Museum of Natural History, Smithsonian Institution (USNM), and the National Science Mu- seum, Tokyo (NSMT). The specimens were fixed in 10% for- malin-seawater and preserved in 70% al- cohol. Parapodia of the holotype were ob- served by a scanning electron microscope after dehydration in ethyl alcohol and crit- ical drying in CO). Family Poecilochaetidae Hannerz, 1956 Poecilochaetus Claparéde, in Ehlers, 1875 Poecilochaetus koshikiensis, new species Figs. 1-3 Material. —Japan, off Shimo-Koshiki Is- land, Kagoshima Prefecture, 31°40.0'N, 129°40.7’E, 200 m, muddy sand, 4 Jun 1987, Nansei-Maru Cruise 1987-TRS5, holotype, complete female (USNM 104126), para- type, incomplete male (USNM 104127), paratype, incomplete female (NSMT-Pol. P-249). Types. —Holotype consisting of 68 seti- gers, 43 mm long, 1.5 mm wide without parapodia, 2.6 mm with parapodia. Mature types with gametes in coelom. Description. —Prostomium small, round- ed, with 2 pairs of eyes, anterior left eye PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. view of setigers 9, with chitinized plate, and 10; d, Posterior end, dorsal view. consisting of several pigment spots (in ho- lotype, Fig. 1a). Nuchal organ trilobed, mid- dle lobe extending posteriorly to setiger 4, flattened laterally with ciliated margin, lat- eral lobes small and discoid (Fig. la) and tentacular palps missing. Facial tubercle present in front of ventral mouth and di- rected anteriorly (Fig. 1b). Stout setae of setiger | directed forward, forming cephalic cage; parapodial postsetal lobes of anterior 3 setigers directed more or less forward (Fig. la, b). Weakly chitinized middorsal elevat- ed plate present on setiger 9 (Fig. Ic). Py- gidium with pair of cylindrical dorsal cirri and pair of short ventral cirri fused in single Y-shaped structure (Fig. 1d). Posterior seg- ments well marked by stout notopodial spines. Parapodia all biramous. First parapo- dium with very short notopodial and very long neuropodial postsetal lobes (Fig. 2a). Notopodial postsetal lobes longer than neu- ropodial ones on parapodia 2-6, both cy- lindrical (Fig. 2c). Notopodial postsetal lobes of setiger 5 longer than those of setigers 3, 4, and 6. Both postsetal lobes of parapodia 7-13 flask-shaped (ampullaceous) with swollen tips (Fig. 2d). Succeeding parapodia Poecilochaetus koshikiensis, holotype: a, Anterior end, dorsal view; b, Same, ventral view; c, Dorsal on middle of body with long conical post- setal lobes (Fig. 2e, f). Far posterior post- setal lobes again cylindrical but smaller, notopodial ones with many conical tuber- cles (Fig. 2h, 1). Numerous papillae present on anteroventral sides of anterior parapodia of setigers 1, 2, 6—8 (Fig. 2a, c, d). Interramal parapodial sensory organs present on all se- tigers except for setigers 6-9. A well-devel- oped sensory organ cup-shaped with cilia in distal hole (Fig. 2b). Sensory organs varying in size and sometimes withdrawn and de- tected only by their cilia (Fig. 2g). Sensory organs of anterior 5 setigers well developed, those of others smaller. Branchiae absent. First parapodium with stout cephalic se- tae forming cephalic cage, very long, surface covered with numerous short spines (Figs. la, b, 2a, 3c, d). Parapodia 2 and 3 with slender hispid capillaries, spiral setae (membranous capillary) and 3 or 4 neuro- podial falcate spines (Fig. 2c). Falcate spines curved posteriorly to body axis, distal tip smooth in SEM observation, but finely hir- sute in LM observation (Fig. 3a, b). Spiral setae with spirally fringed pectinate margin, present on all parapodia except for first se- tiger, their margins with long stout teeth on VOLUME 101, NUMBER 3 673 3 " \y Fig. 2. Poecilochaetus koshikiensis: a, Parapodium 1, anterior view; b, Sensory organ below notopodial postsetal lobe of same; c, Parapodium 2; d, Parapodium 7; e, Parapodium 14; f, Parapodium 17; g, Sensory organ of same; h, Parapodium 54; i, Parapodium 61. 674 Way N 3 5 y a Ye N y% é , N27 A Ze N iT Z Va \ Y fh Bee! J Hy \ Z YA 4 We N Y UY y Wh ie N Y f 5s y Wie N Z lke Wy x Zo / i MY y, ay \ Z / N pe, Lig : y \ % d jie of i N ae / pill J 7, x oY / fig ay \ GY fae eo 0 ah el N y H } Y seh N: % foe | fe of Vee / / G ey Vee is ae | yy j i Hf i H i i | / i : i [ y ! ! i | ai ew : ae | i 5 : i ™~ c SR | b \ ves t i a () a yA ' We ee Vn - y ; | | | ii | NY | A aig WR 4 ii We i i Ny “ é | NG y H eae | I. X i H i ! | k | : Li ' PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Poecilochaetus koshikiensis: a, Neuropodial falcate spine from setiger 2; b, Enlarged tip of same; c, Section of cephalic seta from setiger 1 in lateral view; d, Same, convex side; e, Spiral seta; f, Distal part of same; g, Proximal part of saine; h, Pappose seta; i, Aristate seta; j, Slender hispid seta; k, Posterior notopodial hook. proximal part and fine teeth on distal part (Fig. 3e-g). Parapodia 4-16 with slender hispid capillaries and spiral setae (Fig. 2d, e). Parapodia 17-50 with pappose setae covered with long, fine flexible spinules on distal part and stout spinules on proximal part (Figs. 2f, 3h). Aristate setae first present on parapodia 48, unidentate with hoods covered with numerous short spines (Figs. 2h, 31). Posterior parapodia with slender hispid setae (Fig. 3j). Stout smooth noto- podial hooks forming prominent dorsal shield in posterior 10 segments (Figs. 1d, 2i, 3k). Remarks.—In the genus Poecilochaetus, seven species have flask-shaped cirri on VOLUME 101, NUMBER 3 parapodia 7—13. Of these seven species, two species and a subspecies, P. serpens Allen, 1904, P. serpens honiarae Gibbs, 1971, and P. tropicus Okuda, 1935, 1937, have branchiae, differing easily from the abranchiate new species. P. johnsoni Hart- man, 1939 and P. australis Nonato, 1963 differ from P. koshikiensis in lacking stout notopodial hooks which make remarkable dorsal shields of several preanal segments in the new species. P. fauchaldi Pilato & Cantone, 1976 differs from the new species in lacking aristate spines and interramal sensory organs on segments posterior to se- tiger 5. P. paratropicus Gallardo, 1968 has well-developed lateral lobes on the nuchal organ, while P. koshikiensis has only small discoidal ones. P. japonicus Kitamori, 1965, the only species recorded from Japan, and known only from anterior fragments, has a very short middle lobe on the nuchal organ, reaching setiger 2, compared to the new species with middle lobe extending back to setiger 4. Etymology.—The species name is de- rived from the type locality, Shimo-Koshiki Island. Literature Cited Allen, E. J. 1904. The anatomy of Poecilochaetus Claparéde.— Quarterly Journal of Microscopi- cal Science, N.S. London 48:79-151, 1 fig., 6 pls. Ehlers, E. 1875. Beitrage zur Kenntnis der Vertical- verbreitung der Borstenwiirmer im Meere.— Zeitschrift fiir Wissenschaftliche Zoologie, Leip- zig 25:1-102, 4 pls. Gallardo, V. A. 1968. Polychaeta from the Bay of Nha Trang, South Viet Nam. Scientific Results of Marine Investigations of the South China Sea and the Gulf of Thailand 1959-1961.—Naga Report 4(3):35-279, 59 pls. 675 Gibbs, P. 1971. The polychaete fauna of the Solomon Islands.— Bulletin of the British Museum (Nat- ural History), Zoology, London 21(5):99-211, 17 figs. Hannerz, L. 1956. Larval development of the poly- chaete families Spionidae Sars, Disomidae Mes- nil, and Poecilochaetidae n. fam. in Gullmar Fjord (Sweden).—Zoologiska Bidrag fran Upp- sala 31:1-204, 57 figs. Hartman, O. 1939. New species of polychaetous an- nelids from southern California.— Allan Han- cock Pacific Expedition 7(2):157—-172, 2 pls. Kitamori, R. 1965. Two new species of rare families, Disomidae and Paralacydonidae (Annelida: Polychaeta). — Bulletin of Tokai Regional Fish- eries Research Laboratory (44):41—44, 2 figs. Milligan, M. R., & K. M. Gilbert. 1984. Chapter 9, Poecilochaetidae. Pp. 1-7 in J. M. Uebelacker & P. G. Johnson, eds., Taxonomic guide to the polychaetes of the northern Gulf of Mexico, Volume 2. Barry A. Vittor and Associates, Mo- bile, Alabama. Nonato, E. 1963. Poecilochaetus australis n. sp. (An- nelida, Polychaeta).—Neotropica 9(28):17—26, 15 figs. Okuda, S. 1935. Poecilochaetus tropicus n. sp., a re- markable sedentary polychaete from the South Seas. — Proceedings of the Imperial Academy of Japan 11:289-291, 2 figs. 1937. Polychaetous annelids from the Palau Islands and adjacent waters, the South Sea Is- lands.— Bulletin of the Biogeographical Society of Japan 7(12):257-315, 59 figs. Pilato, G., & G. Cantone. 1976. Nuove specie de Poecilochaetus e considerazioni sulla famiglia dei Poecilochaetidae (Annelida, Polychaeta). — Animalia, Catania 3(1/3):29-63. Read, C. B. 1986. New deep-sea Poecilochaetidae (Polychaeta: Spionida) from New Zealand.— Journal of Natural History 20(2):399-414, 36 figs. Taylor, J. L. 1966. A Pacific polychaete in South- eastern United States. — Quarterly Journal of the Florida Academy of Sciences 29(1):21-26. Faculty of Fisheries, Kagoshima Univer- sity, 4-50-20 Shimoarata, Kagoshima, 890 Japan. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 676-681 NEOPILINA GOEST, A NEW CARIBBEAN MONOPLACOPHORAN MOLLUSK DREDGED IN 1869 Anders Waren Abstract.—A new monoplacophoran species, Neopilina goesi, dredged NW of the Virgin Islands, from the upper part of the continental slope, is described. This new species resembles N. oligotropha Rokop and N. zografi (Dautzenberg & Fischer), but has a more marginal apex than the former and a finer sculpture than the latter species. Axel Theodor Goés (1835-1897) was a Swedish physician and zoologist whose main zoological interest was Foraminifera. Dur- ing 1862-1870 he was employed at the gar- rison at St. Barthelemy, Leeward Islands, then a Swedish colony. In 1866 and 1869 Goés dredged around St. Barthelemy at depths down to 400 fathoms (720 m). This yielded rich material of different groups of animals. A very brief description of the dredgings was given by Goés (1882:8). These collections were given to the Swedish Mu- seum of Natural History in 1870 (Theel 1916:205) and formed the base for several reports on Caribbean invertebrates. The molluscs, however, were only roughly sort- ed to species by N. Odhner and never pub- lished upon, although he labelled many species with manuscript names. A consid- erable part of the molluscan species col- lected by Goés is undescribed and will prob- ably remain so until better material becomes available, because most of it consists of bro- ken juveniles extracted from shell gravel. The monoplacophoran, here described as a new species, had been labelled ““Acmea sp.,”’ by Odhner, an irony of fate, as it was Odhner who named the class Monoplacophora (in Wenz 1940; not Wenz in Knight 1952, as often is quoted). Neopilina Lemche, 1957 Neopilina goesi, new species Figs. 1-3, 6-11 Type material. —Holotype, Swedish Mu- seum of Natural History type collection no. 3635a, the shell figured in Figs. 2, 6, 7. Para- type, type collection no. 3635b, the shell figured in Figs. 1, 3, 8-11. Type locality. —‘‘Windward off the east- ern keys of the Virgin Islands, 200-300 fath- oms (360-540 m),” upper continental slope of the western Caribbean. Distribution. —Known only from the type locality and type material. Description. —Shell very small, colorless, fragile, regularly ovate and depressed, with apex reaching slightly outside margin. Api- cal region consisting of central part without sculpture, but rather rough and with signs of wear, 0.13 x 0.16 mm (Figs. 3, 6); sur- rounded by smooth area, diameter 0.3 mm, sculptured only by numerous pores or pits and 2 or 3 incomplete anterior concentric ridges. Outside this area more regular sculp- ture of concentric and radial ridges starting, giving surface distinctly reticulated appear- ance, with distinct tubercules at intersec- tions of concentric and radial ridges. Con- centric ridges somewhat more close-set and irregular at edge of shell, but very regular at central part of shell. Shell interior very smooth with no muscle scars visible and only some traces of radial sculpture appar- ent by transparency. Shape regularly convex with apex distinctly overhanging anterior ventral margin. Dimensions: Holotype 1.79 x 1.40 x 0.56 mm. Paratype 1.54 x 1.21 x 0.46mm. Remarks. —The two specimens are small- er than the maximum dimensions known for any other monoplacophoran species VOLUME 101, NUMBER 3 (smallest species: Laevipilina hyalina (McLean, 1979), 2.28 mm), but the more crowded concentric radial sculpture at the edge indicates that the specimens are ap- proaching full-grown size. The inside of the shell does not show any muscle scars, although the condition of the shell is good enough to have shown them if they had been present. This agrees with oth- er recent monoplacophorans (Wingstrand 1985:47). The reason for considering this limpet a monoplacophoran is the similarity in shell characters to N. oligotropha Rokop, 1972, and N. zografi (Dautzenberg & Fi- scher, 1896), which from anatomy and shell structure, respectively, are known to belong to this group. A part of the shell is covered by tracks or shallow furrows about 1 wm wide, similar to those made by beetles under the bark of trees (Figs. 9-10). These furrows occur on the area equipped with pores and on the early part of the area with adult sculpture. They appear to be too regular to be caused by corrosion, but their origin is unknown. The presence of three different sculptural areas on the shell, in accordance with the growth lines, indicates that N. goesi passes through three different ontogenetic growth stages in its life history. I cannot, however, presently correlate these with the different phases in the shell development of gastro- pods or lamellibranchs with planktotrophic larvae. Nor is it possible to conclude that the bowl-shaped initial part which lacks pores is a larval shell although this seems possible. Wingstrand (1985) concluded that the old description by Lemche & Wing- strand (1959) ofa spirally coiled larval shell was erroneous. He supported this on a re- port by Menzies (1968) where a young mol- lusc supposed to be a monoplacophoran was figured with a large bulbous larval shell. However, the identification of Menzies’s specimen may be questioned. It closely re- sembles the young of many Lepetellacea Dall, 1881 (Archaeogastropoda), but his photo indistinctly shows a prismatic struc- 677 ture of the shell, not present in Lepetellacea. Wingstrand concluded that it probably rep- resented a monoplacophoran and that this larval shell is later rejected, possibly as in Patella (Linné, 1758). Menzies’s figure is, however, too poor to allow any conclusions about whether the larval shell is spirally coiled as in the Docoglossa, where it con- sists of half a whorl, or simply bowl-shaped as assumed by Wingstrand. I have seen the process of larval shell re- jection in Patella coerulea (Linné, 1758) (Figs. 12-14). Here a narrow zone around the larval and part of the first postlarval shell is dissolved, evidently actively (Warén unpubl. data, see also Gardner 1986), after which the larval shell falls off at the slightest touch. The place where the larval shell has been attached is marked by a distinct scar with a large pore, now closed. These obser- vations agree with Smith’s (1935) descrip- tion of the larval development of Patella, based on sectioned material. His fig. 29b shows a small apical chamber, cut off from the main part of the body by a septum and connected to the body by a narrow string of tissue. Behind the scar is a second impres- sion from the overhanging part of the larval shell. A very similar mode of rejection of the larval shell has also been observed in the Lepetellidae (Warén, unpubl. data). The absence of a pronounced scar from the larval shell (compare Figs. 3, 5 and 14), the shape of the apex, and the presence of three well-marked apical zones seem to ar- gue against Wingstrand’s hypothesis about rejection of the larval shell, but it is possible that there occur different modes of larval development among monoplacophorans. Pores like those obvious in Figs. 6—9 have not previously been reported from any re- cent monoplacophoran. I have, however, seen them also in N. zografi, although they are less obvious in that species, probably because the shells available of N. zografi were less well preserved. These pores are most numerous in the smooth area around the scar of the protoconch, but occur less 678 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | ies 22osse abe ists beaenette Menterisiinert St ee OOP Tapes ¥ VOLUME 101, NUMBER 3 679 Figs. 6-11. Neopilina goesi, details of sculpture: 6, 7, Apical area of holotype, side view, showing bowl- shaped apex; 8, Apical area of paratype, showing distribution of pores; 9, Detail of pores; 10, Transition from part with pores to adult sculpture; 11, Adult sculpture on central part of shell. Scale lines: Fig. 6, 100 um; Figs. 7-11, 10 wm. Arrows indicate transition between different sculptural zones. —_— Figs. 1-5. Neopilina goesi and N. zografi: 1, N. goesi, paratype, maximum diameter 1.54 mm; 2, N. goesi, holotype, lateral view, length 1.79 mm; 3, N. goesi, paratype, apical area; 4, N. zografi, paratype, in Muséum National d’Histoire Naturelle, Paris, maximum diameter 3.9 mm; 5, N. zografi, apical area of specimen in Fig. 4. Scale lines 50 um. Arrows indicate the different sculptural zones. 680 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 12-14. Patella coerulea (L.), from Calvi, Corsica, taken at the pier of Station de Recherche Sousmarines et Océanographiques: 12, Juvenile shell, diameter 0.8 mm; 13, Larval shell (smooth) and part of initial teleconch ready to fall off; 14, Larval shell lost. In the center of the calcareous plug that seals off apex, a closed pore can be seen. This pore is the last connection to the larval shell and may in some way assist in the active dissolution of the zone of breakage. Scale lines: Fig. 13, 30 um, Fig. 14, 20 um. densely all over the shell. I did not study them in detail and their significance is not known. Pores are known in the Silurian ge- nus 7ryblidium Lindstr6m, 1880 (Erben et al. 1968). In that genus they branch inwards, something that could not be examined in N. goesi because of lack of material. There are two known species of Mono- placophora that resemble N. goesi in shell morphology, viz. N. zografi, from the bathy- al zone of the Azores, and N. oligotropha, from abyssal depths north of the Hawaiian Islands. Neopilina oligotropha differs from N. goe- si in having the apex placed well behind the anterior margin of the shell. Other differ- ences are impossible to discern because of the poor illustrations of that species, except that N. oligotropha evidently has a coarser sculpture; N. goesi has 28 concentric ridges behind the apex at a size of 1.79 mm, while N. oligotropha has 32 at a size of 3.0 mm. Neopilina zografi differs mainly in having a blunter apex, the sides of which form a wider angle in dorsal view and perhaps also by reaching a larger size, 3.9 mm. The systematic position of N. goesi within Monoplacophora is uncertain, as no soft VOLUME 101, NUMBER 3 parts are known. From the shell, it is ob- vious that it does not belong to Laevipilina McLean, 1979, whose species have a rather smooth shell, with a low but dense nodular sculpture, caused by the large prisms that build up the shell. Prisms are also obvious in Vema Clarke & Menzies, 1959, where Laevipilina was described as a subgenus. Monoplacophorus Moskalev, Starobogatov, & Filatova, 1983, may be related to the present new species, but the description of the shell and the illustrations do not allow any conclusions. Therefore, I have followed Rokop (1972) and Bouchet, McLean, & Warén (1983) and included this small, strongly sculptured species in Neopilina. Acknowledgments I want to thank Dr. Philippe Bouchet, Dr. James H. McLean, Dr. H. Mutvei, and Prof. K. G. Wingstrand for discussions about this species and Dr. Bouchet, also, for preparing some of the SEM photos at ““Centre de mi- croscopie du CNRS, Paris.” Literature Cited Bouchet, P., J. H. McLean, & A. Warén. 1983. Mono- placophorans in the North Atlantic.—Ocean- ologica Acta 6:117-118. Dautzenberg, P., & H. Fischer. 1896. Dragages ef- fectuées par I’Hirondelle et Princesse-Alice. Mollusques Gasteropodes.—Mémoires de la Société Zoologique de France 9:395—498. Clarke, A. H., & R. J. Menzies. 1959. Neopilina (Verma) ewingi, a second new living species of the Paleozoic class Monoplacophora. — Science 129:1026-1027. Erben, H. K., G. Flajs, & A. Siehl. 1968. Uber die Schalenstruktur von Monoplacophoren.—Ab- handlungen der Matematisch-Naturwissen- schaftlichen Klasse der Akademie der Wissen- schaften und der Literatur, Mainz 1968(1):1—- 24. Gardner, J. P. A. 1986. A method for the investi- gation of the shell structure of newly settled lim- pets.—Journal of Molluscan Studies 52:35-37. Goés, T. A. 1882. On the reticularian Rhizopoda of 681 the Caribbean Sea.—Kongliga Svenska Veten- skaps-Akademiens Handlingar 19(4):1-151. Knight, J. B. 1952. Primitive fossil gastropods and their bearing on gastropod classification. — Smithsonian Miscellaneous Collections 117(13): 1-56. Lemche, H. 1957. A new living deep sea mollusc of the Cambro-Devonian class Monoplaco- phora.— Nature, London 179:413-416. ——, & K. G. Wingstrand. 1959. The anatomy of Neopilina galatheae Lemche, 1957.—Galathea Report 3:9-71. Lindstr6m, G. 1880. Fragmenta Silurica e dono Car- oli Henrici Wegelin (N.P. Angelin). 15. Hol- miae. Linné, C. von. 1758. Systema Naturae. 10th Edition. Uppsala. 822 pp. McLean, J.H. 1979. Anew monoplacophoran limpet from the continental shelf off southern Califor- nia. — Contributions in Science, Natural History Museum of Los Angeles County 307:1-19. Menzies, R. J. 1968. New species of Neopilina of the Cambro-Devonian class Monoplacophora from the Milne-Edwards Deep of the Peru-Chile Trench, R/V Anton Bruun. — Proceedings of the Symposium on Mollusca of the Marine Biolog- ical Association of India 3:1-19. Moskalevy, L. I., Z. A. Starobogatov, & Z. A. Filatova. 1983. New data on the abyssal Monoplaco- phora from the Pacific and South Atlantic oceans. —Zoologicheski Zhurnal 112:981-995. Rokop, R. J. 1972. A new species of monoplaco- phoran from the abyssal North Pacific. — Veliger 15:91-95. Smith, F. G. W. 1935. The development of Patella vulgata. —Philosophical Transactions of the Royal Society of London 225:95-125. Theel, H. 1916. Evertebratafdelningen 1841-1916. In Naturhistoriska Riksmuseets Historia. Kungliga Vetenskapsakademien. Stockholm. Almavist & Wiksell. 290 pp. Wingstrand, K.G. 1985. On the anatomy and rela- tionships of recent Monoplacophora.—Gala- thea Report 16:1—94. Wenz, W. 1940. Ursprung und friihe Stammesge- schichte der Gastropoden.— Archiv ftir Mollus- kenkunde 72:1-109. Department of Invertebrate Zoology, Swedish Museum of Natural History, Box 50007, S-104 05 Stockholm, Sweden. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 682-692 NOTES ON SOME RHYSSOPLAX FROM THE PACIFIC OCEAN (MOLLUSCA: POLYPLACOPHORA: CHITONIDAE) Robert C. Bullock Abstract. Taxonomic comments are presented for the following Rhyssoplax from the Pacific Islands: R. discolor (Souverbie, 1866), R. ectypa (Rochebrune, 1884), and R. rhynchota (Rochebrune, 1884) from New Caledonia; R. spino- setata (Bergenhayn, 1930), from the Fiji Islands; R. whitleyi Iredale & Hull, 1932, from the Cook Islands; R. perviridis (Carpenter, 1865) from the Society Islands; Rhyssoplax sp. from the New Hebrides east to the Samoan Islands; and the widespread R. pulcherrima (Sowerby, 1842), which ranges from South- east Asia to New Caledonia. The latter species is conspecific with R. excellens Iredale & Hull, 1926, C. excellens capricornensis Ashby, 1928, and R. nier- strasziana Kaas, 1957. Rhyssoplax ectypa, which has been considered a junior synonym of R. discolor for nearly a century, is recognized as a separate species. Chiton (Clathropleura) pacificus Thiele, 1910, is a junior synonym of R. per- viridis. Thiele’s (1910) published description and figure of the type specimen of R. rhynchota were erroneously based on R. crawfordi (Sykes) from South Africa. Species of the polyplacophoran genus Rhyssoplax Thiele, 1893, are sporadically represented among malacological collec- tions from the islands of the central Pacific Ocean, and their taxonomy is poorly under- stood. The predominant reason for this sit- uation is the inadequate collections avail- able for study. Large, well preserved collections from all major island groups will be required before sound taxonomic con- clusions can be reached. Study of many Pacific Rhyssoplax is also perplexing due to the inadequate published work of the past. The works of Rochebrune (1884 and other papers) especially have proved to be troublesome. Pilsbry (1893: 151), in a discussion of Rochebrune’s use of the controversial name Gymnoplax, cor- rectly observed the outcome of Roche- brune’s efforts: “‘It should be noted that the irrepressible Rochebrune still uses Gym- noplax in a generic sense, in order presum- ably, to disguise his species;—an unneces- sary precaution, for his diagnoses of Chitons generally defy identification of either genus or species.” Iredale & Hull (1932:158) com- mented on the problems associated with type material in the Muséum National d’His- toire Naturelle in Paris: ““Rochebrune ran riot among these shells in the Paris Mu- seum. Using a classification of his own which has defied interpretation, he lost, mislaid and transferred labels and specimens, de- scribing the types of former workers under different genera and then renaming his own new species until there is no confidence in any specimens of this class in that Mu- seum.”’ Bullock (1972) noted that some of this material is present in the Dautzenberg collection at the Institut Royal des Sciences Naturelles de Belgique in Brussels. The zoogeography and phylogenetic re- lationships of Pacific Island Rhyssoplax will prove to be of great interest. Preliminary examination of the distributional patterns indicate that a few species, for example R. VOLUME 101, NUMBER 3 pulcherrima (Sowerby, 1842), are broadly distributed across several island groups, while a larger numer of species appear to be endemic to a single island group. Examples of the latter group include: R. discolor (Sou- verbie, 1866), known only from New Cal- edonia; R. whitleyi Iredale & Hull, 1932, found only in the Cook Islands; and R. spi- nosetata (Bergenhayn, 1930), reported only from the Fiji Islands. Major conclusions, however, must be made only after the tax- onomy of the group is better known. Materials and Methods Specimens were examined from the ma- jor collections of the U.S. and Europe listed below. Additional material was received from the Western Australian Museum and the Australian Museum. Girdle scales of se- lected specimens were prepared for scan- ning electron microscopy (SEM) following procedures described by Bullock (1985). Samples were mounted on aluminum stubs using double coated tape, and coated with carbon and gold/palladium in a Denton DV- 502 vacuum evaporator. All work was done using an ISI MSM-3 SEM located in the Department of Zoology at the University of Rhode Island. The following abbreviations are used: AMS, Australian Museum, Sydney; ANSP, Academy of Natural Sciences of Philadel- phia; BMNH, British Museum (Natural History), London; ICZN, International Code of Zoological Nomenclature; IRSN, Institut Royal des Sciences Naturelles de Belgique, Brussels; MCZ, Museum of Comparative Zoology, Harvard University, Cambridge; MHNB, Muséum d’Histoire Naturelle de Bordeaux; MNHNP, Muséum National d’Histoire Naturelle, Paris; NRS, Naturhis- toriska Riksmuseet, Stockholm; USNM, National Museum of Natural History, Washington, D.C.; WAM, Western Austra- lian Museum, Perth; ZMA, Zoodlogisch Mu- seum, Amsterdam; ZMHU, Zoologischen Museum, Humboldt-Universitat, Berlin; 683 ZMK, Universitetets Zoologiske Museum, Copenhagen; ZMU, Uppsala Universitets Zoologiska Museum, Uppsala. Genus Rhyssoplax Thiele, 1893 Rhyssoplax Thiele, 1893:368. Type species, Chiton affinis Issel, 1869, by declaration of the ICZN, Opinion 951. Remarks.—In general, the name Rhys- soplax has been used for small to moderate- sized, non-New World polyplacophorans that are otherwise similar in shell and girdle scale morphology to the genus Chiton from the New World. Employment of Rhysso- plax at the generic level is not accepted by all polyplacophoran workers. Malacologists from Australia and New Zealand for the most part have championed this use, while others, forexample Kaas & Van Belle (1981), have used Rhyssoplax as a subgenus of Chiton Linnaeus, 1758. Substantial radular differences exist between Chiton and Rhyssoplax: in Chiton the distal edge of the centro-lateral tooth is perpendicular to the longitudinal axis of the animal, while in all Rhyssoplax the distal edge is parallel to the longitudinal axis. Furthermore, the denticle cap of the major lateral tooth of Rhyssoplax is rather short with a circular black tab dis- tally on the anterior surface. In Chiton the denticle cap tends to be elongate and the black tab is usually elongate. It is advisable to consider these groups as separate genera within the subfamily Chitoninae. Rhyssoplax discolor (Souverbie, 1866) Fig. 13 Chiton discolor Souverbie [in Souverbie & Montrouzier], 1866:252, pl. 9, figs. 1, la (ins. Art, l’Archipel calédonien; lectotype, selected by Strack [1986], in MHNB). Chiton tuberculosus Souverbie [in Souver- bie & Montrouzier], 1866:251, pl. 9, fig. 3 (ins. Art, l’Archipel calédonien; holo- type in MHNB). Chiton (Rhyssoplax) discolor Souverbie.— 684 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-9. Shells of Pacific Rhyssoplax: 1, Rhyssoplax perviridis Carpenter, Arue, Society Islands, 11 mm [curled] (USNM 672794); 2, Holotype of Chiton perviridis Carpenter, Tahiti, Society Islands, 1 1 mm(MCZ 141125); 3, Rhyssoplax ectypa (Rochebrune), New Caledonia, 20.5 mm (BMNH); 4, Syntype of Lepidopleurus ecyptus Rochebrune, 20 mm (MNHNP); 5, Chiton rhynchotus ‘Rochebrune’ Thiele [=R. crawfordi (Sykes)], 9.5 mm VOLUME 101, NUMBER 3 685 Figs. 10-15. (Rochebrune), W end, Ricaudy Reef, Noumea, New Caledonia, 291 x (ANSP 271045); 11, Rhyssoplax pul- cherrima (Sowerby), 3 mi [4.8 km] S of Airport Beach, Phuket Is., Thailand, 81 x (ANSP 286765); 12, Rhyssoplax spinosetata (Bergenhayn), Korolevu, Viti Levu, Fiji Islands, 95x (ANSP 236891); 13, Rhyssoplax discolor (Souverbie), near radio station, Baie Ouemo, Noumea, New Caledonia, 46 x (ANSP 271200); 14, Rhyssoplax ectypa (Rochebrune), 0-4 ft [0-1.2 m], Thio, New Caledonia, 107 x (ANSP 238361); 15, Rhyssoplax sp., E side Scanning electron micrographs of girdle scales of Pacific Rhyssoplax: 10, Rhyssoplax rhynchota of Vailele Bay, Upolu Is., Western Samoa, 169 x (ANSP 198086). Strack, 1986:195, fig. 4a, b [description; types fig?d.]. Remarks. — Rhyssoplax discolor is a com- mon species that varies considerably in col- or. Risbec (1946), who reported on the anat- omy of this species, noted that on the rocks at Noumea R. discolor is second in abun- dance only to Acanthopleura gemmata (Blainville, 1825). Pilsbry (1894) erroneously believed that Lepidopleurus ectypus Rochebrune is con- specific with this species, and all subsequent authors have accepted this conclusion. However, an examination of syntype ma- terial of L. ectypus [IRSN and NMHNP] (Fig. 4) revealed that it is distinct and not at all closely related to R. discolor (see re- marks under R. ectypa). — The New Caledonian R. discolor is not very similar to R. coryphea (Hedley & Hull, 1912) from Norfolk Island, as Iredale & Hull (1932) suggested. Rhyssoplax coryphea is more highly elevated, more elongate, and has longitudinal ribs which only partly ex- tend across the valve. The girdle scales of R. discolor (Fig. 13) characteristically ex- hibit a broad shelf across the apical surface and have many apical pustules; in R. co- ryphea this shelf is nearly lacking and the pustules are limited to a narrow region at the apex. Locality records. —New Caledonia: Koe Reef, 2 mi [3.2 km] SSE Touho; Touho Bay; Koe, near Touho; Thio; Yate (all ANSP); Noumea Harbour (AMS); Port Ngéa, Ma- genta, Noumea (ANSP); Baie des Isoles, [curled] (MNHNP); 6, Rhyssoplax rhynchotus (Rochebrune), barrier reef, 5 mi [8 km] WSW Gatope Is., Voh, New Caledonia, 9.5 mm (ANSP 269722); 7, Rhyssoplax spinosetata (Bergenhayn), N shore of Port Ellington, NNE Viti Levu, Fiji Islands, 17.5 mm (MCZ 279505); 8, Rhyssoplax whitleyi Iredale & Hull, 0.25 mi [0.4 km] N of Matavera, NE Rarotonga, S Cook Islands, 11 mm (MCZ 252808); 9, Rhyssoplax sp., E side of Vailele Bay, Upolu Is., Western Samoa, 7 mm (ANSP 198086). 686 Ouemo, Magenta, Noumea (AMS); S side, Baie de Citron; Baie de Citron, Noumea, in 1-6 ft [0.3-1.8 m] (both ANSP); Ile Cesar, Anse Vata, Noumea (AMS); mouth of stream entering Anse Vata Bay, Noumea, in 2-6 ft [0.6—1.8 m]; Tonghouen Bay, 45 km N of Noumea; Plage de Poe, Bourail; S side, Gatope Is., Voh; SW side, Isle Mouac, Poume; 2 mi [3.2 km] N of Poume (all ANSP). Rhyssoplax ectypa (Rochebrune, 1884) Figs. 3, 4, 14 Lepidopleurus ectypus Rochebrune, 1884:37 (Nouvelle Calédonie; syntypes in IRSN and MNHNP). Description. — Animal medium in size, up to 38 mm in length, 19 mm in width. Valves subcarinate, angle about 90°. Anterior valve straight to convex; post-mucral slope of posterior valve abrupt, concave at first, con- vex ventrally. Mucro moderately sharp, often worn, centrally located on posterior valve. Jugal region smooth. Central areas with numerous, fine, slightly irregular, longitudinal ribs, up to 28 per side; dorsal edge somewhat sloping, ventral edge more abrupt; first few dorsal ribs not reaching preceding valve. Lateral triangle sharply elevated, with 3-6 nodular, radiating ribs; nodes on posterior rib longitudinally elon- gate. Terminal areas with similar radial sculpture, up to 25 on anterior and posterior valves. New radial ribs begin anew between existing ribs. Shell color grayish white with a few darker speckles at jugum or, more commonly, cream orange or reddish brown. Girdle very light orange or alternately band- ed white and light orange. Interior of valves white to dull greenish white. Insertion plates: Intermediate valves with a single slit per side; insertion teeth well grooved, but not deeply pectinate. Anterior valve with 10 teeth; posterior valve with 13 teeth. Hypostracum: Jugal tract in central depression with numerous transverse slits PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and continuous over callus as small holes in posteriordepression. Primary slit-ray with longitudinally elongate slits on callus, more circular, smaller slits in posterior depres- sion. Secondary slit-ray absent. Girdle scales: Moderate in size, rather ovate, with reticular sculpture predominate except near apex where large, fairly close- packed nodules are arranged in rows. Apical shelf lacking (Fig. 14). Remarks. — Rhyssoplax ectypa is distin- guished from R. discolor by its more sharply defined lateral triangle, more abrupt post- mucral slope on the posterior valve, nar- rower ribs of the central area which are strongest near the lateral triangle, and by the fact that as the animal grows, additional ribs of the lateral triangle and end valves begin separately between existing ones, not as bifurcations of previous ribs. The girdle scales of the two species greatly differ (Figs. 13, 14). In R. ectypa the scales are ovate rather than roundly triangular and the char- acteristic apical shelf of R. discolor is lack- ing. The apical pustules of R. ectypa are linearly arranged, not random as in R. dis- color. Certain structural aspects, such as the sharply defined lateral triangle and the man- ner by which new ribs are added, indicate that R. ectypa is most closely related to the more northern R. komaiana (Is. & Iw. Taki, 1929), which may be differentiated by its more inflated girdle scales and more pro- nounced ribs of the central areas of the in- termediate valves. Rhyssoplax ectypa has not been recog- nized as a valid taxonomic entity since its original description by Rochebrune (1884). At least five species of Rhyssoplax are now reported from New Caledonia: Rhyssoplax subassimilis (Souverbie, 1866), known only by the holotype (Strack 1986); the abundant R. discolor (Souverbie); and the uncommon R. ectypa, R. rhynchota (Rochebrune, 1884), and R. pulcherrima (Sowerby, 1842). Gym- noplax ludoviciae Rochebrune, 1884 [syn- type IRSN], stated to be from New Cale- donia, is Chiton (Chondroplax) granosus VOLUME 101, NUMBER 3 Frembly, 1827, from the Pacific coast of South America. Gymnoplax alphonsinae Rochebrune, 1884 [syntypes IRSN and MNHNP], also stated to be from New Cal- edonia, is actually Rhyssoplax aerea (Reeve, 1847) from New Zealand. Distribution. —Rhyssoplax ectypa is known only from New Caledonia. Locality records. —New Caledonia: (BMNH, IRSN, MNHNP); 0-4 ft [0-1.2 ml], Thio, New Caledonia (ANSP). Rhyssoplax sp. Figs. 9, 15 Remarks. —The collections of the ANSP and USNM include a few examples of small Rhyssoplax that possibly are juveniles of one or more species. The shell sculpture is reminiscent of that of R. ectypa from New Caledonia. The girdle scales from the dif- ferent localities vary in the size of the apical pustules, the linearity of these pustules, the existence of small riblets that extend from the pustular region ventrally to the reticular sculpture, and the presence in some cases of broader ribs. I figure an example from the Samoan Islands (Fig. 9) and a girdle scale from it (Fig. 15). Locality records. —New Hebrides: rocky reef flat, ESE of Inyeung Is., Aneityum (USNM).—Tonga Islands: Niuafoou Is. (USNM).— Hoorn Islands: over the outside fringing reef, NW of Mua, Alofi (USNM).— Samoan Islands: Asili, Tutuila (USNM); Pago Pago, Tutuila (USNM); east side of Vailele Bay, Upolu Is. (ANSP). Rhyssoplax pulcherrima (Sowerby, 1842) Fig. 11 Chiton pulcherrimus Sowerby, 1842:103 (Gindulman, ins. Bohol, Philippinarum; type in BMNH).—Leloup, 1952:34, pl. 1, fig. 2; pl. 4, fig. 1; text-fig. 13 [girdle ele- ments and esthetes fig’d.]. Ischnochiton pulcherrimus (Sowerby).— Pilsbry, 1893:130, pl. 27, fig. 47, 48. Chiton reticulatus Nierstrasz, 1905:81, pl. 687 2, fig. 36; pl. 7, figs. 195-199 (Insel San- guisiapo, 12 m; Saleyer; syntypes in ZMA). Non Reeve, 1847, nec Dupuis, 1918. Rhyssoplax excellens Iredale & Hull, 1926: 181, pl. 19, figs. 22, 27, 40 (Darnley Is- land, Torres Strait; type in Macleay Mu- seum, Sydney) [description].—Iredale & Hull, 1927:116, pl. 13, figs. 22, 27, 40.— Mackay, 1934:150, pl. 16 Chiton (Rhyssoplax) excellens capricornen- sis Ashby, 1928:169, pl. 12, figs. 1, 13 (Capricorn Group; location of type un- known). Rhyssoplax nierstrasziana Kaas, 1957:85 (new name for C. reticulatus Nierstrasz, 1905, non Reeve, 1847). Remarks. — Rhyssoplax pulcherrima dif- fers from all other Rhyssoplax, and all other Chitoninae, by its cancellate sculpture of the central areas. This species has been rede- scribed on a number of occasions because it seems that authors have been unwilling to recognize its broad distribution. Austra- lian workers in particular have maintained that their specimens represented a species different from the one in the Philippine Is- lands. Pilsbry (1893) considered this species to be an Ischnochiton, but an examination of the radula, especially the denticle cap of the major lateral tooth, indicates its chitonid affinities. A study of the shell morphology of the Siboga Expedition material revealed that Chiton reticulatus Nierstrasz, 1905, is this species, and its renaming by Kaas (1957) was unjustified. Rhyssoplax komaiana (Is. & Iw. Taki, 1929), which occurs sympatri- cally with R. pulcherrima in at least Malay- sia, the Philippines, and possibly south to the Admiralty Islands (ANSP 182204), dif- fers by its rounded valves and lack of can- cellate sculpture. The prominent central ribs on the girdle scales of R. pulcherrima (Fig. 11) easily distinguish this species from other carinate Rhyssoplax. Distribution. —A tropical species, Rhys- 688 soplax pulcherrima occurs from Thailand to the Philippine Islands and south to Queens- land, Australia, the New Hebrides, and New Caledonia. Locality records. —Thailand: Pulau Tan- ga, Butang Group (USNM); coral rubble reef, 3 mi [4.8 km] S of Airport Beach, Phuket Is., Andaman Sea (ANSP); Koh Chang, Gulf of Siam (ZMK).— Malaysia: Pulau Siburu, N of Sipora, SW of Sumatra (USNM); Pi- sang Is. (IRSN); N shore of Toba Is., N end of Aru, Molucca Is. [5°21’S, 136°27'E] (WAM).--Philippine Islands: Gindulman, Bohol Is. (BMNH); rock shore at Cabcaben, SE Bataan, Luzon Is. (ANSP).— New Heb- rides: Anelgauhat, Aneityum Is., on Turbo marmoratus L. (ANSP).—New Caledonia: 4-7 ft [1.2—2.1 m], barrier reef, 5 mi [8 km] WSW of Gatope Is., Voh, SW New Cale- donia (ANSP). Rhyssoplax spinosetata (Bergenhayn, 1930) Figs. 7, 12 Chiton spinosetatus Bergenhayn, 1930:24, pl. 1, figs. 17-28; pl. 2, figs. 29-31 (Viti Levu, Fiji Islands; type in NRS). Description.— Animal medium to small in size, reaching a length of 22 mm, a width of 14 mm. Valves fairly carinate, angle about 100°. Anterior valve straight to convex; post- mucral slope of posterior valve straight to slightly concave. Mucro rather blunt, slight- ly anteriorly acentric on posterior valve. Ju- gal region smooth; central area with up to 20 well formed longitudinal ribs per side, usually all reaching preceding valve; dorsal edge of each rib somewhat sloping, lateral edge abrupt. Lateral triangle with 5-8 ra- diating, occasionally bifurcating rows of low, broad nodules. Terminal areas with similar radial sculpture, up to 41 rows on anterior valve, 29 on posterior valve. Shell color variable. Some specimens dark brown with scattered lighter, maculated areas and dark girdle of yellowish white scales irregularly PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON banded with dark brown. Other specimens much lighter, usually yellowish green mac- ulated with small greenish brown markings, or yellowish white with similar maculations with lateral areas orange; girdle alternately banded yellowish white and dull green. In- terior of valves blue or white with bluish green to dull green streak extending from mucro along slope of posterior depression. Insertion plates: Intermediate valves with a single slit per side, occasionally doubly slit; insertion teeth highly grooved, not deeply pectinate. Anterior valve with 9-12 teeth; posterior valve with 11-15 teeth. Hypostracum: Much of central depres- sion with irregular thin grooves and slits. Primary slit-ray well developed, with prom- inent diagonal slits. Secondary slit-ray lack- ing. Girdle scales: Moderately large, rather smooth; fine reticular sculpture ventrally; central area smooth with obsolete stria- tions; apical area sharply defined by close- packed pustules of moderate size. Remarks. —A comparison of Bergen- hayn’s type specimen of R. spinosetata with additional specimens collected from the Fiji Islands has firmly established the identity of R. spinosetata, although slight differences exist between populations. Rhyssoplax spi- nosetata appears to be related most closely to two New Zealand species, R. stangeri (Reeve, 1847) and R. canaliculata (Quoy & Gaimard, 1835). Distribution. —Rhyssoplax spinosetata is apparently endemic to the Fiji Islands. Locality records. — Fiji Islands: Kaba, Viti Levu (ZMU); N shore of Nananu-i-ra Is., 3 mi [4.8 km] N of Port Ellington, NNE Viti Levu (ANSP, MCZ); fringe reef, Korolevu, Viti Levu (ANSP, MCZ). Rhyssoplax perviridis (Carpenter, 1865) Figs. 1; 2 Chiton (Lophyrus) perviridis Carpenter, 1865:511 (Central Pacific [type locality VOLUME 101, NUMBER 3 here restricted to Tahiti, Society Islands]; holotype MCZ 141125). Chiton (Clathropleura) pacificus Thiele, 1910:93, pl. 10, figs. 1-4 (Huahine and Anaa [Society Is.]; type in ZMHU). Description.—Animal rather small, at- taining a length of 13 mm, a width of 8.5 mm. Valves subcarinate, angle about 105°. Anterior valve straight; post-mucral slope of posterior valve concave. Mucro moder- ately sharp, anteriorly acentric on posterior valve. Jugal region smooth. Central areas with up to 17 longitudinal ribs, the more dorsal ones not reaching preceding valve; dorsal surface of each rib sloping, ventral edge abrupt. Lateral triangle elevated, with 4—6 weak, low-noduled ribs. Terminal areas with similar radial sculpture; ribs obsolete. Shell color usually yellowish white; lateral edge of each longitudinal rib translucent brown or dark green. Small, irregular green- ish brown flecks on lateral triangle; larger splotches along anterior edge of lateral tri- angle and occasionally along posterior edge. Some immature specimens smoother, to- tally green. Girdle alternately banded light blue-green and yellowish white. Interior of valves white. Insertion plates: Intermediate valves with a single slit per side; insertion teeth highly grooved, moderately pectinate. Anterior valve with 10-11 teeth; posterior valve with 11-12 teeth. Hypostracum: Jugal tract of central depression with scattered transverse slits; entire central depression nearly transparent. Primary slit-ray with prominent slits on cal- lus, a row of irregular small holes in pos- terior depression. Girdle scales: Rather large, roundly tri- angular, moderately inflated; reticular sculpture ventrally; 14-17 broad ribs on central area, thinning near apex; small num- ber of close-packed pustules at apex; apical shelf barely evident. Remarks. —The unique type of R. pervir- 689 idis (Fig. 2), formerly in the Pease collection, is in the Museum of Comparative Zoology. Although the label and the original descrip- tion state the locality to be “Central Pacif- ic,” Pease (1872) mentioned that the spec- imen was from Tahiti. The type locality is therefore restricted to Tahiti, Society Is- lands. Thiele (1910), who was no doubt un- aware of the identity of Carpenter’s species, described Chiton pacificus on the basis of specimens from Huahine and Anaa, also in the Society Islands; an examination of Thiele’s type indicates it is conspecific with R. perviridis. Rhyssoplax perviridis differs from the Rhyssoplax of neighboring island groups, but appears to be most closely related to R. /ins- leyi Burghardt, 1973, from the Hawaiian Islands and R. whitleyi Iredale & Hull, 1932, from the Cook Islands. Rhyssoplax linsleyi has more even radial sculpture and the lon- gitudinal ribs of the central areas are some- what better formed. Rhyssoplax perviridis is easily differentiated from R. whitleyi by lacking the stronger longitudinal ribs of the central area and the low, but prominent, nodules of the lateral triangle. The girdle scales of all three species are quite similar, except that those of R. /ins/eyi have thin ribs in the central area while those of R. pervir- idis and R. whitleyi have broad ribs. Distribution. —Rhyssoplax perviridis is known only from the Society and Tubuai Islands. Locality records. —Society Islands: (MCZ, USNM); W of Motu Tapu, Bora Bora Is.; Arue, Tahiti; E side of Taunoa Pass, Tahiti (all USNM); fringe reef, opposite pass, Pu- neavia, Tahiti (ANSP, MCZ); Huahine; Anaa (both ZMHU).—Tubuai Islands: W of wharf, Moerai, Rurutu (USNM). Rhyssoplax whitleyi Iredale & Hull, 1932 Fig. 8 [?] Chiton (Clathropleura) alphonsinae ‘Rochebrune’ Thiele, 1910:93 [in part]. Non Rochebrune, 1884. 690 Rhyssoplax whitleyi Iredale & Hull, 1932: 145, pl. 9, figs. 15, 16 (Rarotonga, Cook Islands; type in AMS [not seen]) [descrip- tion]. Remarks. — Rhyssoplax whitleyi is distin- guished from other Rhyssoplax from the central Pacific by its yellowish brown and blue coloration and by the prominent low nodules on the lateral triangle. Its girdle scales are moderately large, roundly trian- gular with reticular sculpture evident ven- trally and at sides; there are 8-10 broad ribs on the central area and moderate-sized pus- tules on a flattened apical area. Rhyssoplax whitleyi differs from R. perviridis by having better formed longitudinal ribs on the cen- tral areas. In R. spinosetata from the Fiji Islands, the nodules are less pronounced and the scales (Fig. 12) very weakly striated, not distinctly striated as in R. whitleyi. Also, the apical pustules of the girdle scales of R. spi- nosetata are sharply limited to the apex, and not converging into the central area ribs as in R. whitleyi and R. perviridis. Distribution. —Rhyssoplax whitleyi has only been recorded from the Cook Islands. Locality records. —Cook Islands: Avatiu Harbor to Motu Toa, Rarotonga (USNM); 0.25 mi [0.4 km] N of Matavera, NE Rar- otonga (ANSP, MCZ); Koromiri Is., SE Rarotonga (ANSP); Avaavaroa Passage, S Rarotonga (ANSP); fringe reef off Aroa Creek, SW Rarotonga (ANSP). Subgenus Anthochiton Thiele, 1893 Anthochiton Thiele, 1893:377. Type species by monotypy, Anthochiton tulipa (Quoy & Gaimard, 1835). Remarks.—The name Anthochiton has been used for Rhyssoplax-like species that have a smooth lateral triangle and end valves. It appears useful at this time to use this name to denote a separate lineage with- in the genus Rhyssoplax that is character- ized by these features. The taxonomic status of nomina associated with Rhyssoplax, such PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON as Anthochiton, Tegulaplax Iredale & Hull, 1926, Delicatoplax Iredale & Hull, 1926, and Mucrosquama Iredale & Hull, 1926, must await detailed radular and anatomical studies. Rhyssoplax (Anthochiton) rhynchota (Rochebrune, 1884) Figs. 6, 10 Gymnoplax rhynchotus Rochebrune, 1884: 39 (Nouvelle Calédonie; type not found in the MNHNP).—Pilsbry, 1894:100. Rhyssoplax rhynchotus (Rochebrune). — Risbec, 1946:163, fig. 18 (anatomy, es- thetes, radula). Non Chiton (Clathropleura) rhynchotus ‘Rochebrune’ Thiele, 1910:92, pl. 9, figs. 52-55 [=Rhyssoplax crawfordi (Sykes, 1898)]. Description. —Animal rather small, reaching a length of 9.5 mm, a width of 5.5 mm. Valves subcarinate, angle about 90°. Mucro rather blunt, anteriorly acentric on posterior valve. Jugal region smooth. Cen- tral areas with up to 12 well formed longi- tudinal ribs per side. Lateral triangle smooth, often somewhat inflated. Terminal areas smooth. Shell color tan, yellowish green, or light orange, often with irregular darker markings. Girdle similarly colored. Interior of valves white. Insertion plates: Intermediate valves with a single slit per side; insertion teeth ap- pearing smooth but obsoletely grooved. Hypostracum: Jugal tract with numerous, faint transverse grooves. Primary slit-ray with many longitudinally elongate slits. Girdle scales: Moderate in size, ovate to rectangular; reticular sculpture ventrally, about 21 broad, close-packed ribs on central and apical areas (Fig. 10). Remarks. —Rhyssoplax rhynchota is characterized by its small size, strong lon- gitudinal ribs of the central areas, and slight- ly inflated lateral triangle. The specimen marked “‘type”’ in the MNHNP is not this species. This latter specimen was figured and VOLUME 101, NUMBER 3 described by Thiele (1910) as an example of Rochebrune’s species, but an examina- tion of the valves (Fig. 5), which clearly have a concave lateral triangle, and the girdle scales reveal it to be a young example of R. crawfordi (Sykes, 1898) from South Africa. The nearly smooth insertion teeth of R. rhynchota are unusual for Chitoninae. A check of the radular proved it to be typically chitonid. Distribution. —Rhyssoplax rhynchota oc- curs from the Bismarck Archipelago south to New Caledonia. Locality records. —Bismarck Archipela- go: New Ireland (BMNH).—New Caledo- nia: (MNHNP); Oubatche (AMS); N tip of Santa Marie (Isle Ngéa), Magenta, Noumea; 0-3 ft [0O-0.9 m], W end, Ricaudy Reef, Noumea; 4—7 ft [1.2—2.1 m], barrier reef, 5 mi [8 km] WSW Gatope Is., Voh (all ANSP). Acknowledgments For assistance during my visits to their institutions and for the loan of specimens, I gratefully acknowledge: K. Boss and R. Turner (MCZ), H. Coomans (ZMA), T. Karling and C. Holmquist (NRS), R. Kilias (ZMHU), J. Knudsen (ZMK), E. Leloup (IRSN), B. Métivier and A.-M. Testud (MNHNP), G. Davis and R. Robertson (ANSP), J. Rosewater, C. Roper, and H. Rehder (USNM), J. Taylor, K. Way, and J. Peake (BMNH), W. Ponder (AMS), B. Wil- son and L. Joll (WAM). Although in general collections of Pacific island Rhyssoplax are meagre, I especially wish to acknowledge the collections that were made possible by the efforts of G. and M. Cline, V. Orr Maes, and R. Ostheimer (ANSP) and H. Rehder (USNM); without this material the present study would have been impossible. Literature Cited Ashby, E. 1928. Notes on a collection of chitons (Polyplacophora) from the Capricorn Group, Queensland.— Transactions and Proceedings of 691 the Royal Society of South Australia 52:167- 173, pl. 12 [in part]. Bergenhayn, J. R. M. 1930. Die Loricaten von Prof. Dr. Sixten Bocks Pazifik-Expedition 1917-1918 mit spezieller Beriicksichtigung der Perinotum- bildungen und der Schalenstruktur. —G6teborgs Kungliga Vetenskaps-och Vitterhets-Samhilles Handlingar, femte F6ljden, (B) 1 (12), 52 pp., 3 pls. Bullock, R. C. 1972. Notes on the genus Chiton in the western Indian Ocean (Mollusca: Polypla- cophora).— Occasional Papers on Mollusks, Museum of Comparative Zoology, Harvard University 3:237-251, pl. 44. . 1985. The Stenoplax limaciformis (Sowerby, 1832) species complex in the New World (Mol- lusca: Polyplacophora: Ischnochitonidae).— Ve- liger 27:291-307, 24 figs. Carpenter, P. P. 1865. Description of two species of Chitonidae, from the collection of W. Harper Pease, Esq.— Proceedings of the Zoological So- ciety of London for 1865:511-512. ICZN [International Commission on Zoological No- menclature]. 1971. Opinion 951. Rhyssoplax Thiele, 1893 (Amphineura): designation of a type species under the plenary powers.—Bulletin of Zoological Nomenclature 28:18-19. Iredale, T., & A. F. B. Hull. 1926. A monograph of the Australian loricates. VI.—Australian Zool- ogist 4:164-185, pls. 18-20. ——,, & . 1927. A monograph of Australian loricates. Sydney, 168 pp., 21 pls. , & 1932. The loricates of the Neo- zelanic region. — Australian Zoologist 7:119-164, pls. 7-10. Kaas, P. 1957. Notes on Loricata. 5. On some preoc- cupied names.—Basteria 21:83-87. ,& R.A. Van Belle. 1981. Catalogue of living chitons (Mollusca: Polyplacophora). Rotter- dam, W. Backhuys, 144 pp. Leloup, E. 1952. Polyplacophores de l’Océan Indien et des cdtes de l’Indochine francaise.—Meé- moires de |’Institut Royal des Sciences Natu- relles de Belgique, Deuxiéme Série, Fasc. 47, 69 pp., 6 pls. Mackay, J. 1934. Notes onarare loricate, Rhyssoplax excellens Iredale and Hull.—Memoires of the National Museum, Victoria 8:150—153, pl. 16. Nierstrasz, H. 1905. Die Chitonen der Siboga-Ex- pedition. —Siboga-Expeditie Monographie 48:1-— 114, 8 pls. Pease, W. H. 1872. Polynesian Chitonidae.—Amer- ican Journal of Conchology 7:194-195. Pilsbry, H. A. 1892-1894. Polyplacophora.— Manual of Conchology 14:1—128 [1892], 129-350 [1893]; 15:1-133 [1894]. Risbec, J. 1946. Etudes anatomiques sur les amphi- 692 neures de la Nouvelle-Calédonie.—Journal de la Société des Océanistes 2:129-190, pl. 2. Rochebrune, A. T. de. 1884. Diagnoses d’espéces nouvelles de la famille des Chitonidae (Deux- iéme supplément).— Bulletin de Société Philo- matique de Paris (7) 8:32-39. Souverbie, M., & R. P. Montrouzier. 1866. Descrip- tions d’espéces nouvelles de l’Archipel calédo- nien.—Journal de Conchyliologie 14:248-261, pl. 9. Sowerby, G. B. 1842. Descriptions of four species of the genus Chiton, brought by H. Cuming, Esq. from the Philippine Islands. — Proceedings of the Zoological Society of London for 1841:103-104. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Strack, H. L. 1986. Notes on New Caledonian chi- tons. I. The species described by St.-M. Sou- verbie. — Basteria 50:193-198, 5 figs. Thiele, J. 1893. Polyplacophoren. Pp. 353-401 and pls. 30-32 in F. H. Troschel, Das Gebiss der Schnecken, volume 2. Berlin, Nicolaische Ver- lagsbuchhandlung. 1910. Revision des Systems der Chitonen, II.—Zoologica 22:71—132, pls. 7-10. Department of Zoology, University of Rhode Island, Kingston, Rhode Island 02881. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 693-706 A REVIEW OF THE MARINE NEMATODE GENERA PLATYCOMA AND PROPLATYCOMA, WITH A DESCRIPTION OF PROPLATYCOMA FLEURDELIS (ENOPLIDA: LEPTOSOMATIDAE) W. Duane Hope Abstract. —The taxonomic histories of the genera Platycoma Cobb, 1894 and Proplatycoma Platonova, 1976 are reviewed. Proplatycoma fleurdelis, n. sp. from Barbados West Indies, is described, and it is the first record of this genus in the western North Atlantic. The structure of the buccal capsule, amphidial flaps, spicula and gubernacula are discussed, and the genera Proplatycoma and Platycoma are redefined. An artificial key to the males of the species of Platyco- ma and Proplatycoma is provided. The genus Platycoma was proposed by Cobb (1894:399) to receive a single new species, P. cephalata and named for the “Two flat hairs of unequal length that grow from the inner margin of the anterior border of each ...” amphid of the males (Cobb 1894:400). Platonova (1976:138) and Platt & Warwick (1983:178) have identified these structures as setae. Platycoma africana (Gerlach, 1959) Gerlach, 1962, P. curiosa (Gerlach, 1955) Gerlach, 1962, and P. sud- africana Inglis, 1966 were subsequently added to the genus. Males of species whose descriptions were subsequent to those of the type species lack the pair of “‘hairs” or “‘se- tae’? emerging from each amphid, but all possess what is presumed to be a modifi- cation of this character. The amphid in males of P. africana is covered by a flap that is broad at its posterior end and tapered an- teriorly to a blunt, central point. The am- phidial flap in males of P. curiosa and P. sudafricana is tapered to form a central, an- teriorly directed lobe, and each has a dorsal and ventral lobe as well. The dorsal and ventral lobes in P. curiosa are highly branched, whereas in P. sudafricana they are not branched, but sloped posteriorly, giving the flap the appearance of an arrow- head. Platonova (1976:139) proposed the genus Proplatycoma to which she transferred P. curiosa, P. africana, and P. sudafricana, with the latter designated as the type species. In the diagnosis of Proplatycoma, Platonova (1976:139) states that the amphids of the males are elongated anteriorly, sometimes with unique processes, and that cuticular, labial outgrowths are absent around the buccal aperture. Although Platonova (1976: 138) did not give a formal diagnosis for Platycoma, it is stated in her key to the gen- era of Platycominae that there are two flat setae situated near each amphid, and cutic- ular, labial outgrowths are present. Plato- nova (1976:137) also proposed a new subfamily, Platycominae, for the genera Platycoma, Proplatycoma, Platycomopsis and Pilosinema. A collection of marine nematodes from Barbados, West Indies, deposited at the Na- tional Museum of Natural History, Smith- sonian Institution (USNM), contains two males, three females and three juveniles of Proplatycoma fleurdelis, n. sp. These spec- imens most closely resemble those of P. sud- africana, known only from males. The syn- types of P. sudafricana were studied for the purpose of making a comparison between members of the two species. The morpho- 694 Fig. 1. of amphidial flap of Proplatycoma fleurdelis: A, Length of flap; B, Minimum (anterior) width of flap; C, Max- imum (posterior) width of flap; D, Length of anterior lobe; E, Length of dorsal and ventral lobes; F, Distance from oral surface of head to posterior margin of flap. Diagram showing location of measurements metric and meristic data for P. sudafricana presented in the discussion of P. fleurdelis are from original observations. These specimens provided an opportu- nity to make detailed observations on the structure of the amphidial flap, the cuticu- lar, labial outgrowths, and the spicula and gubernaculum. Platycoma and Proplaty- coma are redefined on the basis of these observations. Materials and methods. —Specimens were fixed in formalin and mounted in anhy- drous glycerine between coverslips mount- ed on Cobb aluminum frames. Morpho- metric data were obtained by measuring camera lucida drawings and photographs of the specimens with a Sumagraphics Digi- tizing Pad. Morphometric and meristic data in the text are given first for the male ho- lotype (USNM 77129), followed by that for the male paratype (USNM 77130). Some measurements were not possible for the male PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON paratype; in those cases it is stated that the data is for the holotype. Similarly, for fe- males the data is given for USNM 77131 followed by that for USNM 77132, and for juveniles the data is given for USNM 77133, USNM 77170, and USNM 77171, in that order. Where only two items of data are given for the juveniles, it is that for USNM 77170 that has been omitted, because the position of the head of that specimen made certain measurements impossible. Mea- surements of the male amphid were made as shown in Fig. |. For each set of data, the mean + one standard deviation is given in parentheses. Statistics are not calculated for the juveniles because the specimens are most likely at different stages of development. After measurements were made on male specimen USNM 77130, it was removed from its slide mount, rehydrated, and freeze dried for scanning electron microscopy by a method described elsewhere (Hope 1982: 2). The specimen was precoated with car- bon, coated with gold/palladium, and ex- amined with an Hitachi S-570 SEM. Platycoma Cobb, 1894 Platycoma Cobb, 1894:399.—Inglis, 1966: 85.—Gerlach & Riemann, 1974:482.— Platonova, 1976:137. Type species. —Platycoma cephalata Cobb, 1894. Diagnosis. —Leptosomatidae. Body slightly tapered from mid region toward each end; cu- ticle, at least in head region, with very fine transverse striae. Head region of males con- stricted at level of amphid, then cylindrical to level of cephalic sensilla; anterior end rounded, truncate or slightly concave. Head region of female uniformly tapered, without constriction. Amphid cyathiform; cuticle at posterior rim of amphidial aperture flap- like in males and extended anteriorly over amphidial aperture; each amphidial flap with 2 parallel, tapered lobes. Amphidial flap absent in females. Oral aperture surrounded by 3 microla- VOLUME 101, NUMBER 3 bia, each microlabium triangular in profile with apex anteriorly directed. Odontia, on- chia or other forms of buccal armament ab- sent. Anterior end of esophagus attached to cephalic cuticle; oral aperture dilated by la- bial retrodilator muscles in anterior end of esophagus. Subventral esophageal gland duct orifices open into buccal capsule at base of subventral microlabia. Spicula typical in structure and position, and with ventrolat- eral wedge- or rod-shaped accessory struc- ture; gubernaculum with small, paired, dor- sally directed apophyses. Proplatycoma Platonova, 1976 Proplatycoma Platonova, 1976:139. Type species. — Proplatycoma sudafricana (Inglis, 1966) Platonova, 1976. Diagnosis. —Leptosomatidae. Body slightly tapered from mid region toward each end; cu- ticle with very fine transverse striae. Head region of males constricted at level of am- phid, then cylindrical to level of cephalic sensilla; anterior end rounded to slightly concave. Head region of female uniformly tapered. Cephalic capsule present or absent. Amphid cyathiform; cuticle at posterior rim of amphidial aperture in males flap-like and extended anteriorly over amphidial aper- ture. Amphidial flap tapered to single, an- teriorly directed point without dorsal and ventral lobes, or flap with single, central, anteriorly directed lobe and with dorsal and ventral lobes; amphidial flap absent in fe- males. Oral aperture surrounded by 3 microla- bia, each microlabium triangular in profile with apex anteriorly directed. Odontia, on- chia or other forms of buccal armament ab- sent. Anterior end of esophagus attached to cephalic cuticle; oral aperture dilated by la- bial retrodilator muscles in anterior end of esophagus. Subventral esophageal gland duct orifices open into buccal capsule at base of subventral microlabia. Spicula typical in structure and position; wedge- or rod-shaped accessory structure ventrolateral to spicula; 695 gubernaculum with paired caudally directed apophyses. Caudal glands absent. Proplatycoma fleurdelis, new species Figs. 1-5 Holotype. —Male, USNM 77129. Paratypes. —Male, USNM 77130; two fe- males (USNM 77131 and 77132); and three juveniles (USNM 77133, 77170, and 77171). Type locality.—Barbados, West Indies. The type specimens were collected by Dr. Bjorn Urhammer in Jan and May 1968. Un- fortunately, the type specimens came from a composite sample from the top 20 cm of sediment at high- and low-water marks at Skeete’s Bay, the top 10 cm of sediment at the edge of Green Pond, and from the top 20 cm of sediment at the low-water mark in Bottom Bay. Therefore, it is not possible to identify the exact location(s) from which this material was obtained. Description. — Males. — Body a b Cc length USNM 77129 128 6.6 52 8.135 mm USNM 77130 142 6.9 60 9.672 mm Body diameter uniform except near an- terior and posterior ends; midbody diam- eter 64 and 68 (66 + 3) um; body width at base of esophagus 62 um (holotype); at level of nerve ring 55 um (holotype); at level of cephalic sensilla 25 and 21 (23 + 3) um. Head region (Figs. 2A, C; 5A) constricted at level of amphid, cylindrical to level of cephalic sensilla, then rounded anteriorly from level of cephalic sensilla. Oral surface of head slightly concave to rounded. Striae of cuticle (Fig. 2B) very shallow; periodicity of striae 400 nm. Head with 6 papilliform, inner labial sensilla (Figs. 2B; 5A). Single circle of 6 setiform, outer labial sensilla and 4 setiform cephalic sensilla (Figs. 2A; 5A), 8 um from oral surface of head in both specimens; outer labial sensilla of ho- lotype 20-25 (22 + 3) um and cephalic sen- 696 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Proplatycoma fleurdelis: A, SEM micrograph of lateral surface of head of male paratype, USNM 77130; scale equals 10 um; B, SEM micrograph of oral surface of male paratype, USNM 77130; scale equals 3 um; C, Photomicrograph of head of holotype male USNM 77129 in optical sagittal section; scale equals 10 um; D, Photomicrograph of left amphidial flap of male holotype USNM 77129; scale equals 10 um; E, Photomi- crograph of right amphidial flap of holotype; scale equals 10 um. Abbreviations.— AA, amphidial aperture; AF, amphidial flap; AS, amphidial gland secretion; CC, cephalic capsule; DM, dorsal microlabium; DO, level of dorsal esophageal gland orifice; LP, inner labial papillae; MG, microlabial groove; RM, retrodilator muscles of buccal aperture; SM, subventral microlabium; SO, subventral esophageal gland orifice; VL, ventral lobe of amphidial flap. silla of holotype 20-22 (21 + 1) um long. Cervical region of holotype with 21 sensilla on right side of body, 20 on left. Length of cervical sensilla 14-19 (16 + 2) wm. Ce- phalic capsule (Figs. 2C; 5A) 3 wm long on both specimens, visible only on dorsal and ventral sides of head when head viewed in sagittal, optical section. Cuticle at posterior rim of amphidial ap- erture extended anteriorly, flap-like over aperture; amphidial flap (Figs. 2A, D, E; 5A) tri-lobed and with shape of fleur-de-lis. Sur- VOLUME 101, NUMBER 3 Fig. 3. Proplatycoma fleurdelis: A, SEM micrograph um; B, SEM micrograph of ventral, precloacal region of male paratype USNM 77130; scale equals 10 um; C, SEM micrograph of a postero-lateral view of the posterior ventromedian supplement of paratype USNM 77130; scale equals 5 um; D, Same as C, but in ventral view; scale equals 1 wm. Abbreviations.— AS, anterior ventro- median supplement; CS, subventral caudal seta; MD, depression produced by the contraction of copulatory muscles; PS, posterior ventromedian supplement; SS, subventral, setiform supplements. Dhak Dames of tail of paratype male USNM 77130; scale equals 20 face of flap striated; lobes nonstriated and and ventral lobes arched with rounded ends more electron reflective than flap (Fig. 2A). directed posteriorly. Right and left amphid- Middle, anteriorly-directed lobe tapered to _ ial flaps of holotype 7 um long; width of flap point with tip often bent outward; dorsal at its anterior end 4 wm on right and left 698 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Proplatycoma fleurdelis, photomicrographs of female paratype USNM 77131: A, Head in optical sagittal section, scale equals 10 um; B, Right lateral surface of head; scale equals 10 wm; C, Optical sagittal section of tail; scale equals 40 um. Abbreviations.— AA, amphidial aperture; AC, amphidial fovea; CC, cephalic capsule; DM, dorsal microlabium; RM, retrodilator muscles of the buccal aperture. VOLUME 101, NUMBER 3 Fig. 5. Proplatycoma fleurdelis: A, Lateral view of head of holotype, USNM 77129; scale equals 10 um; B, Lateral view of tail of holotype, USNM 77129; scale equals 20 um; C, Lateral view of head region of female paratype, USNM 77131; scale equals 10 um. sides and width of flap at its posterior end 9 um on right and left sides; anterior lobe of left flap 8 um long; dorsal and ventral lobes of right and left amphidial flaps 7 um long. Distance from oral surface of head to posterior margin of right and left flaps 24 and 26 (25 + 1) um, respectively. Amphidi- al glands well developed; posterior end of 700 right gland of holotype 687 wm from oral surface of head. Oral aperture (Fig. 2B) triradiate and sur- rounded by 3 microlabia (Figs. 2A, B, C; 5A). Microlabia triangular with apex di- rected anteriorly, and each set off from head by microlabial groove (Fig. 2B). Buccal cap- sule triradiate and dilated by 3 obliquely oriented, labial retrodilator muscles (Figs. 2C; 5A), 1 in each sector at anterior end of esophagus. Buccal armament absent. Ori- fice of dorsal esophageal gland (Figs. 2C; 5A) in lumen of esophagus, 18 and 20 (19 + 1) um posterior from oral surface. Ori- fices of right and left subventral esophageal glands on mesal surface of right and left microlabia, respectively (Fig. 2B). Esopha- gus 1240 and 1400 (1320 + 114) um long, cylindrical, slightly tapered anteriorly, and posteriorly crenate. Renette absent. Somatic sensilla uncommon between level of nerve ring and near level of cloacal vent. Right lateral chord of holotype with 48 dor- solateral loxometanemes of type I with cau- dal filaments, and 28 without caudal fila- ments; in left lateral chord 58 with caudal filaments and 29 without. Hypodermal gland cells present in hypodermal chords, each gland cell opening on surface of cuticle at dorsolateral or ventrolateral margin of chord. Testes paired and opposed, each on left of gut. Seminal vesicle highly convoluted. Ejaculatory duct enveloped with muscle cells, each cell oblique with dorsal margin anterior to ventral margin. Spicula (Fig. 5B) paired, curved ventrally; right spiculum of holotype 73 and left spiculum 76 (75 + 2) um long, measured on chord. Proximal end of each spiculum bluntly rounded without distinct capitulum; diameter of proximal third uniform, distal end tapered. Ventral surface of distal two-thirds of each spiculum recessed. Gubernaculum (Fig. 5B) with paired apophyses directed caudally; each apophysis keel-shaped, straight; right apophysis in holotype 18 and left 13 (16 + 4) um long. Wedge-shaped accessory struc- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ture (Fig. 5B) present in recessed area on ventrolateral surface of spiculum; proximal end narrow; distal end of right accessory structure in holotype 9 and distal end of left 11 (10 + 1) wm wide and with 13 fine trans- verse striae on right and with 12 on left side of body. Copulatory muscles of body wall anterior to spicula (Fig. 5B) oblique and nearly in continuous sheets. Setiform, subventral supplements (Figs. 3A; 5B) 18-24 (20 + 2) um long and in series of 4 and 4 on right side of body and 3 and 4 on left; anterior supplements 41 and 49 um anterior to cloacal vent on right side of body and 35 and 36 (40 + 6) um on left; posterior subventral supplements approxi- mately adanal. Posterior ventromedian supplement (Figs. 3A, B, C, D; 5B) disc- shaped and pedunculate, on raised surface of body 63 and 79 (71 + 11) um anterior to cloacal vent; center of disc with slightly raised, longitudinally oriented ridge (Fig. 3D); posterior end of ridge expanded and with central depression (sensory receptor ?). Anterior ventromedian supplement (Fig. 3A, B) with 4 and 3 pairs of setiform sen- silla, 1 member of each pair on right and left sides respectively of sagittal plane; sen- silla of paratype 3-5 (4 + 1) um long; pos- terior pair 112 and 108 (111 + 2) wm from cloacal vent, and anterior pair 127 and 118 (123 + 6) um from cloacal vent. Dorsolat- eral, lateral and ventrolateral setiform sen- silla sparsely distributed precaudally to near anterior limit of copulatory muscle. Tail 158 and 160 (159 + 1) wm long with terminal, conical spike (Figs. 3A; 5B). Paired, stout, subventral setiform sensilla (Figs. 3A; 5B), 8 and 9 (9 + 1) um long, present on tail 67 and 81 (74 + 10) um posterior from cloacal vent, or 45 and 51 (48 + 4)% of tail length. One subdorsal se- tiform sensilla present on each side of tail near base of spike; additional subdorsal cau- dal setiform sensilla sparse or absent. Clo- acal body diameter 59 and 48 (53 + 8) um. Caudal glands absent. Females. — VOLUME 101, NUMBER 3 Body a b c Vv length USNM 77131 133 85 86 63% 10.619 mm USNM 77132 107 8.0 60 65% #£7.718 mm Body diameter at midlength 80 and 72 (76 + 6) um; at level of base of esophagus 70 and 68 (69 + 1) um; at level of nerve ring 65 and 63 (64 + 1) um; at level of cephalic sensilla 29 and 28 (29 + 1) um. Head (Figs. 4A; 5C) uniformly tapered from cephalic sensilla to oral surface, then trun- cate to slightly concave. Head with 6 papilliform, inner labial sen- silla (Fig. 5B). Single circle of 6 setiform, outer labial sensilla and 4 setiform, sub- median, cephalic sensilla (Fig. 5C), 11 and 13 (12 + 1) wm from oral surface. Outer labial sensilla 13-17 and 15-16 (15 + 2) um long, and 4 submedian cephalic sensilla 1 5— 17 and 16-18 (17 + 1) um long. Cervical region with 22 and 19 (21 + 1) sensilla on right side of neck and 20 and 17 (19 + 2) on left. Cervical sensilla 3-14 (9 + 4) and 4-14 (9 + 4) um long. Cephalic capsule 5 um long on both specimens. Amphid (Figs. 4B; 5C) cyathiform; aperture transversely oval, 16 and 15 (16 + 0.4) um from oral surface; amphidial fovea 9 and 8 (9 + 1) um long and 7 and 6 (7 + 0.4) um wide. Oral aperture triradiate and surrounded by 3 microlabia (Fig. 4A; SC). Microlabia set off from head by microlabial grooves. Oral surface of head concave with contrac- tion of retrodilator muscles (Figs. 4A; 5C). Buccal armature absent. Orifice of dorsal esophageal gland (Fig. 5C) 21 and 19 (20 + 1) um from oral surface. Orifices of sub- ventral glands on mesal surface of subven- tral microlabia. Esophagus 1255 and 967 (1111 + 204) um long, cylindrical, slightly tapered anteriorly, and posteriorly crenate. Renette absent. Somatic sensilla sparse. Hypodermal gland cells present in hypodermal chord, each gland cell opening on surface of cuticle at dorsolateral and ventrolateral margin of 701 chord. Hypodermal chord in vulvar region without specialized gland cells. Reproduc- tive system amphidelphic and antidro- mous. Gonads ventral to gut; anterior ovary on left side of oviduct, posterior on right. Tail (Fig. 4C) tapered dorsally and ven- trally; length 124 and 128 (126 + 3) um with 1 subdorsal sensillum and 2-4 sensilla at base of caudal spine on each side of body. Anal body diameter 59 and 60 (60 + 1) um. Caudal glands absent. Juveniles. — Body a b c length USNM 77133 60.3 4.8 Dies 3.498 mm USNM 77170 92.0 5.1 31.5 5.683 mm USNM T7171 133.0 7.3 54.0 8.268 mm Similar to females. Midbody diameter 58, 62, and 62 wm; body diameter at base of esophagus 61, 61, and 62; at level of nerve ring 59, 52, and 56; at level of cephalic sen- silla 20, 22, and 23 um. Head truncate to slightly concave on oral surface. Cephalic sensilla 11-14, 12-13, and 13-17 um long and 10 and 8 um from oral surface of head. Amphid cyathiform; aperture transversely oval; anterior rim of aperture 13 um from oral surface of head in each of 2 specimens; amphidial fovea longitudinally ovoid, 9 by 6 um in each of 2 specimens. Cephalic cap- sule 2.5 and 3.0 um long. Distance from oral surface to orifice of dorsal esophageal gland 19 and 16 wm. Esophagus 730, 1108, and 1135 um long. Cervical region with 15, 18, and 24 sensilla on right side; 10, 20, and 22 on left; cervical sensilla 6-14, 5-14, and 6— 15 um long. Tail tapered on dorsal and ventral sur- faces; length 127, 180, and 153 um; terminal spike present. Each side of tail with 1 or 2 subdorsal setiform sensilla. Anal body di- ameter 47, 49, and 54 um. Caudal glands absent. Diagnosis. —Proplatycoma fleurdelis is most similar to P. sudafricana, with the am- 702 phidial flap of the males of each species hav- ing a median lobe and unbranched, dorsal and ventral lobes. The males of the new species differ from those of P. sudafricana in that the dorsal and ventral lobes of each amphidial flap are arched with their re- spective tips pointed dorso- and ventrocau- dally so that the overall shape of the am- phidial flap resembles the fleur-de-lis; the values for ‘“‘a” and “‘b” are 135 + 10 and 7 + 0.2 respectively; the distances from the cloacal vent to the posterior and anterior ventromedian supplements are 71 + 11 and 117 + 29 um respectively; the number of setiform subventral supplements are 3 or 4 on the right side of the body and 4 on the left. In P. sudafricana the dorsal and ventral branches respectively are sloped dorso- and ventrocaudally so that the overall shape of the amphidial flap resembles an arrowhead; the values of “‘a”’ and “‘b”’ for the only whole syntype specimen are 174 and 8 respective- ly; the distances from the cloacal vent to the posterior and anterior ventromedian sup- plements are 115 + 17 wm and 185 + 23 wm respectively; the number of setiform subventral supplements are 2 on the right and 2 on the left sides of the body. Etymology.—The specific epithet is a noun derived from fleurdelis (middle French) for flower of the lily, which denotes the shape of the amphidial flap. Discussion. —SEM observations reveal that the base of the amphidial flap in males of P. fleurdelis is covered with striated exo- cuticle, whereas the anterior and dorsal and ventral lobes are without striations and have the same relatively high electron-reflective appearance as the cephalic and cervical sen- silla (Fig. 2A). This may be due to the fact that both the sensilla and lobes of the am- phidial flaps are extensions of the meso- or endocuticle without the external covering of the exocuticle. Even if this is so, it does not necessarily imply that the branches of the amphidial flaps are derived from sensory sensilla, because, unlike the sensilla, each does not arise from a socket, and there is PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON no evidence from light microscopy ofa nerve passing into them. Although SEM obser- vations have not been made on the amphid- ial flaps of the males of the other species of Proplatycoma, or those of Platycoma ceph- alata, it is assumed that the flaps are ho- mologous. The only difference between them is assumed to be the number, shape and position of the lobes. The amorphic sub- stance at the edge of the amphidial aperture (Fig. 2A) is assumed to be a secretion of the amphidial gland. The 3 projections around the oral aper- ture are here interpreted to be microlabia, homologous with the microlabia among members of Leptosomatidae (Hope 1982: 12). Like microlabia, they are set off from the head region by a groove, the microlabial groove, to the base of which is attached the labial retrodilator muscle for the buccal cap- sule. It was not possible to determine if this muscle is attached to an apodeme, as it is in Deontostoma (Hope 1982:4). Also, as in other leptosomatids, there is an orifice of a subventral esophageal gland duct on the mesal surface of each subventral microla- bium. However, in other members of Lep- tosomatidae, the subventral gland duct ori- fice opens into the mandibular groove on the mesal surface of the microlabium (Hope 1982:8). This groove separates the micro- labium from the mandibular ridge, the latter being absent in members of Proplatycoma. The microlabia of P. fleurdelis differ from those of Deontostomatinae (Leptosomati- dae) in that they are triangular in profile, and tapered to a relatively thin anterior edge (Fig. 2A, B). When the labial retrodilator muscles are relaxed, the microlabia project anteriorly beyond the anterior surface of the head (Fig. 2A, B), but when these muscles are contracted, the buccal cavity is dilated and the microlabia are pulled posteriorly so that they no longer protrude beyond the oral surface (Figs. 2C; 4A; 5A, C). In this con- dition, microlabia may appear tooth-like, especially when the head is viewed laterally in optical longitudinal section. This inter- VOLUME 101, NUMBER 3 pretation applies to females as well. It is also apparent that complete contraction of the labial retrodilator muscles causes the oral surface of the head to become concave (Figs. 4A; 5C). An examination of the syntypes of P. sud- africana during this study has revealed the presence of microlabia similar to those of P. fleurdelis and a complete absence of buc- cal armament. This is in contrast to the in- terpretations of Inglis (1966:83) who does not mention the presence of lips or micro- labia and states that the buccal cavity has “*.. . three small wholly cuticular onchia on each sector ....’ A head remains on only one of the two syntypes of this species, and in that specimen the microlabia are pulled posteriorly into the buccal cavity. It is con- cluded that Inglis (1966:83) erroneously identified the microlabia as onchia. The structure and function of the labial region among members of P. sudafricana is iden- tical to those among members of P. fleur- delis. It is further postulated that the buccal ap- erture is surrounded by 3 microlabia, and the buccal cavity is without odontia or on- chia in Proplatycoma curiosa and P. afri- cana, as well as in Platycoma cephalata. Thus, in the illustrations of the male heads in the original description of P. curiosa and P. africana, the microlabia have probably been pulled posteriorly into the buccal cap- sule (Gerlach 1955:254; 1959:361). The il- lustration of the male head in the original description of Platycoma cephalata is with- out sufficient detail to allow conclusions concerning the structure of the labial re- gion (Cobb 1894:400). However, in the illus- trations of the head of a male of this spe- cies published by Bresslau & Schuurmans Stekhoven (1940:table II, figs. 6a, b, c), the tips of the microlabia are level with the oral surface of the head, whereas in the illustra- tions ofa male of the same species published by Platt & Warwick (1983:179), the micro- labia extend anterior to the oral surface. Even though the projections are interpreted 703 in both publications as being teeth, they are undoubtedly microlabia, and the difference in the degree to which the microlabia are extended is attributable to the degree by which the labial retrodilator muscles are contracted. This applies to males and fe- males. The foregoing are relevant with regard to the diagnoses of Platycoma and Proplaty- coma. Although the former genus was not redefined by Platonova (1976:139), she characterizes Proplatycoma as follows: Nematodes characterized by sharp sexual dimorphism in structure of amphids. In fe- males amphids typically cyathiform but slightly elongated in longitudinal direction. In males anterior end of amphids elongated anteriorly and sometimes with unique pro- cesses. Cuticular labial outgrowths absent. Cervical setae may be numerous and may be scattered over the preneural region in groups or singly. Tail wide, terminally acic- ular. Spicules small, blunt, and reduced. Gubernaculum significantly longer than spicules, curved in the middle, with large blunt dorsal process. The cuticular labial outgrowths of the above diagnosis are the microlabia of this study. Because it has been demonstrated that specimens of P. fleurdelis and P. sudafricana do have microlabia, and that microlabia probably occur as well in P. curiosa and P. africana, Platycoma and _ Proplatycoma cannot be separated on the basis of this character. Platonova’s (1976:140) interpretation of the spicula and gubernacula in her diagnosis of Proplatycoma is probably based upon an earlier erroneous interpretation by Gerlach (1959:361; 1962:85) who regarded the straight, wedge-shaped structure ventrolat- eral to the spicula as being the spicula, and the spicula as being the gubernaculum with a caudal apophysis. This interpretation is in contrast to that made by Inglis (1966:83) in his original description of P. sudafricana and in the present study of P. fleurdelis. As may be seen in Fig. 5B, the structure of the spicu- 704 la and gubernacula and the number and ar- rangement of the muscles associated with them are similar to those found in other leptosomatid species, especially in those cases where the gubernaculum possess a caudal apophysis. This interpretation for the structure and arrangement of the spicula and gubernacula corresponds to that made by Platt & Warwick (1983:178) in their rede- scription of Platycoma cephalata. The only difference that exists between males of Platycoma and Proplatycoma is that in the former the apophyses of the gubernaculum are smaller and directed dorsad, whereas in males of all species of Proplatycoma the apophyses are larger and directed caudad. The remainder of the characteristics giv- en by Platonova (1976:139) for Proplaty- coma also apply to Platycoma. Further- more, the presence or absence of the cuticular labial outgrowths is the only char- acter by which Platonova (1976:138) sep- arates Platycoma and Proplatycoma in her key. Therefore, the two genera cannot be distinguished from one another as they are defined by Platonova. In the new definitions given here, the males of the two genera may be distinguished by differences in 1.) the amphidial flap, which is comprised of two, anteriorly-directed, parallel lobes in the case of Platycoma, and by a single, anteriorly- directed lobe, with or without dorsal and ventral lobes, in Proplatycoma; and 2.) the apophyses of the gubernaculum, which are smaller and dorsally directed in Platycoma, and larger and caudally directed in Pro- platycoma. There are no known characters by which the females of the two genera can be separated. The subfamily Platycominae is retained, but it is recognized that at present it cannot be defined by a single synapomorphic char- acter. Further study of members of Platy- comopsis Ditlevsen, 1926 and Pilosinema Platonova, 1976 is needed. The following key to the species of Pla- tycoma and Proplatycoma is for the iden- tification of males only, because adult fe- males of P. curiosa and P. sudafricana have PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON not been described, and females of other species of the genera are not sufficiently well known. Artificial Key to the Males of the Species of Platycoma and Proplatycoma 1. Each amphidial flap tapered to a single anteriorly directed tip with- out dorsal and ventral lobes, or with a single, central, pointed lobe, and with dorsal and ventral lobes. — Apophysis of gubernaculum cau- dally directed Proplatycoma 2 — Each amphidial flap with 2 parallel, anteriorly directed lobes; each lobe tapered anteriorly, without dorsal and ventral lobes; apophysis of gu- bernaculum dorsally directed conte Platycoma cephalata Cobb, 1894 2. Amphidial flap with single central and dorsal and ventral lobes — Amphidial flap tapered to single, anteriorly-directed tip without dor- sal and ventral lobes Ny Lee eters P. africana (Gerlach, 1959) 3. Dorsal and ventral lobes not branched — Dorsal and ventral lobes branched LORY ae eee P. curiosa (Gerlach, 1955) 4. Amphidial flap arrowhead-shaped; dorsal and ventral lobes sloped pos- teriorly; 2 subventral supplements on each side of body NR area ir « P. sudafricana Inglis, 1966 — Amphidial flap with shape of fleur- de-lis; dorsal and ventral lobes arched; 3 or 4 subventral supple- ments on each side of body gies on hus et aaa P. fleurdelis, n. sp. eee ee ew ew Ce ee Distribution and ecology.—The original description of specimens of Platycoma cephalata was based on material extracted from sandy sediment collected in the Bay of Naples, Italy (Cobb 1894:399). This species was redescribed by Southern (1914: 33) from specimens obtained from a habitat of sand and shells at a depth of 44 m in VOLUME 101, NUMBER 3 Clew Bay, Ireland, and by Bresslau & Schuurmans Stekhoven (1940:13) from specimens, including females, inhabiting Polygordius-sediment collected at Helgo- land in the North Sea. Proplatycoma africana was originally col- lected in coarse, intertidal sand on the Insel Abd el-Kuri, Gulf of Aden (Gerlach 1959: 360). Subsequently, it was found in medium sand in the intertidal zone of Fadiffollu Atoll, Maldive Islands (Gerlach 1962:85); coral sand in the littoral and supralittoral zones of Sarso Island, Red Sea (Gerlach 1967a: 15; 1967b:20); coarse sand between the low and mean water levels on the coast of India (Bay of Bengal) near Waltair (Rao & Gan- apati 1968a:40; 1968b:97); fine to coarse sand between low and mean water levels on the coast of India (Bay of Bengal) near Puri and Konarak, Orissa (Nagabhushanam & Rao 1969:77; Rao 1969:94); medium sand between low and mid-tide levels on Pudi- madaka Beach, Andhra Pradesh, India (Rao 1970:112). Proplatycoma curiosa was originally col- lected from the intertidal zone on the coast of El Salvador near San Salvador (Gerlach 1955:253), and subsequently found in coarse sand on the coast of Brazil near Victoria and Macaé (Gerlach 1957:463). Proplatycoma sudafricana is known only from its type habitat and locality, which is sand of the surf zone near Durban, Republic of South Africa (Inglis 1966:83). In summary, the known distribution of P. cephalata, the only species of the genus, is the eastern North Atlantic and Mediter- ranean Seas, roughly between 40°N (Bay of Naples) and 54°N (Helgoland), and its depth range is to at least 44 m. Members of the genus Proplatycoma, on the other hand, range from about 20°N, as in the case of P. africana (Rao 1969:89) to almost 30°S, as in P. sudafricana (Inglis 1966:83). The known depth distribution for all members of Proplatycoma does not extend below the intertidal zone. It appears, therefore, that differences between Platycoma and Pro- platycoma are to be found in geographic 705 distribution and, possibly, habitat distri- bution. Members of Platycoma are temper- ate and are known to range into subtidal depths, whereas members of Proplatycoma are tropical to subtropical and limited to the intertidal zone; and John Lambshead of the British Museum (Natural History) for the loan of type specimens. Acknowledgments I gratefully thank Ms. Abbie Yorkoff for her technical assistance; Bob Higgins, Ar- mand Maggenti, and Richard O’Grady for reviewing the manuscript; the staff of the SEM Laboratory for taking the scanning electron micrographs. Literature Cited Bresslau, E., & J. H. Schuurmans Stekhoven. 1940. Marine freilebende Nematoda aus der Nord- see.— Bruxelles, Musée Histoire Naturelle de Belgique. 74 pp. Cobb, N. A. 1894. Tricoma and other new nematode genera. — Proceedings of the Linnean Society of New South Wales 8(2):389-421. Gerlach, S. A. 1955. Zur Kenntnis der freilebenden marinen Nematoden von San Salvador. —Zeit- schrift fiir Wissenschaftliche Zoologie 158:249- 303. 1957. Die Nematodenfauna des Sand- strandes an der Kiiste von Mittelbrasilien (Bra- silianische Meeres-Nematoden IV).—Mittei- lungen aus dem Zoologischen Museum in Berlin 33:411-463. 1959. Drei neue Nematoden aus dem Kius- tengrundwasser der Insel Abd el-Kuri (Golf von Aden).— Zoologischer Anzeiger 163:360-364. . 1962. Freilebende Meeresnematoden von den Malediven.—Kieler Meeresforschungen 18(1): 81-108. 1967a. Die Fauna des Kiistengrundwassers am Strand der Insel Sarso (Rotes Meer).—Me- teor-Forschungsergebnisse (D) 2:7-18. . 1967b. Freilebende Meeres-Nematoden von den Sarso-Inseln (Rotes Meer).— Meteor-For- schungsergebnisse (D) 2:19-43. , & F. Reimann. 1974. The Bremerhaven checklist of aquatic nematodes.— Ver6ffentli- chungen des Instituts fiir Meeresforschung in Bremerhaven, Supplement 4, part 2:405-736. Hope, W. D. 1982. Structure of head and stoma in the marine nematode genus Deontostoma (En- oplida: Leptosomatidae).—Smithsonian Con- tributions to Zoology 353:1—22. 706 Inglis, W.G. 1966. Marine nematodes from Durban, South Africa.— Bulletin of the British Museum (Natural History) Zoology 14(4):81-106. Nagabhushanam, A. K., & G.C. Rao. 1969. Prelim- inary observations on a collection of shore-fau- na of the Orissa coast, India.— Proceedings of the Zoological Society of Calcutta 22:67-82. Platonova, T. A. 1976. Nizshie Enoplida (Svobod- nozhivushchie Morskie Nematody) morei SSSR. In T. A. Platonova & V. V. Gal’tsova. Issle- dovanie Fauny Morei. XV:X XIII. Nematody i ikh Rol’v Meiobentose. (Translated to English. Lower Enoplida of the seas of the Soviet Union. Pp. 1-214 inT. A. Platonova & V. V. Gal’tsova. Nematodes and their role in the meiobenthos. — Studies on Marine Fauna 15(23):1-366. Amer- ind Publishing Co., New Delhi.) Platt, H. M., & R. Warwick. 1983. Freeliving Marine Nematodes. Part I. British Enoplids. Published for the Linnean Society of London and The Es- tuarine and Brackish-water Sciences Associa- tion. London, Cambridge University Press. 307 pp. Rao, G. C. 1969. The marine interstitial fauna in- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON habiting the beach sands of Orissa coast. —Jour- nal of the Zoological Society of India 21:89- 104. . 1970. Three new interstitial gastrotrichs from Andhra coast, India.—Cahiers de Biologie Ma- rine 11(1):109-120. —., & P. N. Ganapati. 1968a. Some new inter- stitial gastrotrichs from the beach sands of Wal- tair coast.— Proceedings of the Indian Academy of Science (B) 67:35-53. , & 1968b. The interstitial fauna in- habiting the beach sands of Waltair coast. — Pro- ceedings of the National Institute of Science of India (B) 34:82-125. Southern, R. 1914. Nemathelmia, Kinorhyncha and Chaetognatha (Clare Island Survey, Part 54).— Proceedings of the Royal Irish Academy 31(54): 1-80. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(3), 1988, pp. 707-708 INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE The following Opinions were published on 25 March 1988 in vol. 45, part 1 of the Bulletin of Zoological Nomenclature: Opinion No. 1464 Calcarina calcar d’Orbigny, 1839 (currently Pararotalia calcar; Protista, For- aminiferida): specific name conserved. 1465 Laomedea gracilis Sars, 1850 (currently Clytia gracilis; Cnidaria, Hydrozoa): specific name conserved. 1466 Terebratula triangulus Valenciennes, 1819, 7. catulloi Pictet, 1867 and T. janitor Pictet, 1867 (Brachiopoda, Articulata): specific names con- served. 1467 Criopus Poli, 1791 and Criopoderma Poli, 1795 (Brachiopoda): suppressed. 1468 Orbicula Cuvier, 1798 (Brachiopoda, Inarticulata): suppressed. 1469 Crania tuberculata Nilsson, 1826 (Brachiopoda): conserved. 1470 _ SINUITIDAE Dall, 1913, MACLURITIDAE Carpenter, 1861 and EUMPHALIDAE de Koninck, 1881 (Gastropoda, Archaeogastropoda): conserved. 1471 Aplysia (originally Laplysia) viridis Montagu, 1804 (Mollusca, Gastropoda): specific name conserved. 1472 Cyclaxyra Broun, 1893 (Insecta, Coleoptera): conserved. 1473 Tetropium Kirby, 1837 (Insecta, Coleoptera): conserved. 1474 Tropiphorus Schonherr, 1842 (Insecta, Coleoptera): conserved. 1475 Dexia Meigen, 1826 (Insecta, Diptera): Musca rustica Fabricius, 1775 des- ignated as the type species. 1476 Agromyza Fallén, 1810 (Insecta, Diptera): Agromyza reptans Fallén, 1823 designated as the type species. 1477 Napomyza Westwood, 1840 (Insecta, Diptera): conserved. 1478 Lycaena mirza Plotz, 1880 (currently Azanus mirza; Insecta, Lepidoptera): specific name conserved. 1479 Antispila Hiibner [1825] (Insecta, Lepidoptera): Antispila stadtmuellerella Hubner [1825] designated as type species. 1480 Apanteles ornigis Weed, 1887 (currently Pholetesor ornigis; Insecta, Hyme- noptera): specific name conserved. 1481 Siphamia Weber, 1909 and S. permutata Klausewitz, 1966 (Osteichthyes, Perciformes): conserved. 1482 Heteroclonium bicolor Cope, 1896 (currently Bachia bicolor; Reptilia, Squa- mata): specific name conserved. 1483 Rhabdodon Matheron, 1869 (Reptilia, Ornithischia): conserved. Direction No. 122 Bubo Dumeril, 1806 and Surnia Duméril, 1806 (Aves): Official List entries completed. 708 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE The following applications, and one general article, were published on 25 March 1988 in vol. 45, part 1 of the Bulletin of Zoological Nomenclature. Comment or advice on these applications is invited for publication in the Bulletin and should be sent to the Executive Secretary, ICZN, @ British Museum (Natural History), Crom- well Road, London SW7 5BD, U.K. General Article An appraisal of the Zoology of C. S. Rafinesque. Case No. 2609 Madrepora limax Esper, 1797 (currently Herpolitha limax) and Fungia tal- pina Lamarck, 1801 (currently Polyphyllia talpina; both Cnidaria, Anthozoa): proposed conservation of the specific names. 2610 Coenobita Latreille, 1829 (Crustacea, Decapoda): proposed conservation 2613 Sphaeroma hookeri Leach, 1814 (currently Lekanesphaera hookeri; Crus- tacea, Isopoda): proposed conservation of the specific name. 2607 Hydrobius Leach, 1815 and Berosus Leach, 1817 (Insecta, Coleoptera): con- firmation of type species. 2481 Elachista Treitschke, 1833 (Insecta, Lepidoptera): proposed conservation, and confirmation of type species designation. 2617 Colias alfacariensis Ribbe, 1905 (Insecta, Lepidoptera): proposed availability as a senior synonym of ‘“‘Colias australis Verity, 1911.” 2411 Ludita Nagy, 1967 (Insecta, Hymenoptera): proposed designation of Tiphia villosa Fabricius, 1793 as type species. 2608 Vespa triangulum Fabricius, 1775 (currently Philanthus triangulum, Insecta, Hymenoptera): proposed conservation of the specific name. 2598 Ictiobus Rafinesque, 1820 (Osteichthyes, Cypriniformes): proposed conser- vation. 2556 Hydrolycus Miller & Troschel, 1844 (Osteichthyes, Cypriniformes): confir- mation proposée de Hydrocyon scomberoides Cuvier, 1819 comme espéce-type. 2621 Ascalabotes gigas Bocage, 1875 (currently Tarentola gigas; Reptilia, Squa- mata): proposed conservation of the specific name. 2605 Euryotis brantsii A. Smith, 1834 (currently Parotomys brantsii; Mammalia, Rodentia): proposed conservation of the specific name. ee . y iw ta ‘ea¥ ee ea ee INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (botany, zoology, and paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with a summary in an alternate language when appropriate. Submission of manuscripts. —Submit manuscripts to the Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History NHB-108, Smithsonian Insti- tution, Washington, D.C. 20560. Review. —One of the Society’s aims is to give its members an opportunity for prompt pub- lication of their shorter contributions. 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Academic Press, New York. Figures and tables with their legends and headings should be self-explanatory, not requiring reference to the text. Indicate figure and table placement in pencil in the margin of the manu- script. Plan illustrations in proportions that will efficiently use space on the type bed of the Proceedings. Original illustrations should not exceed 15 x 24 inches. Figures requiring solid black backgrounds should be indicated as such when the manuscript is submitted, but should not be masked. CONTENTS Geographic variation and probable sources of the northern mockingbird in the Bahama Islands Donald W. Buden A new subspecies and new records of Papasula abbotti (Aves: Sulidae) from archeological sites in the tropical Pacific David W. Steadman, Susan E. Schubel, and Dominique Pahlavan Leiognathus pan, a new ponyfish (Pisces: Leiognathidae) from Thailand, with comments on Thai leiognathids Thosaporn Wongratana Semaprochilodus varii, a new species of prochilodontid fish (Ostariophysi: Characiformes) from the Marowijne River, Surinam Ricardo M. C. Castro Relationships of the neotropical catfish genus Nemuroglanis, with a description of a new species (Osteichthys: Siluriformes: Pimelodidae) Carl J. Ferraris, Jr. A new cyprinid fish of the genus Phoxinus (Pisces: Cypriniformes) from the Tennessee River drainage with comments on relationships and biogeography Wayne C. Starnes and Robert E. Jenkins Rotuma lewisi, new genus and species of fish from the southwest Pacific (Gobioidei, Xenisth- midae) Victor G. Springer A review of the Asian species of the genus Lamproscatella Hendel (Diptera: Ephydridae) Wayne N. Mathis and Jin Zuyin Myodocopine ostracoda of the Alaskan continental shelf Louis S. Kornicker Bythocheres prominulus, a new genus and species (Copepoda: Siphonostomatoida) from deep- water cold seeps at the West Florida escarpment Arthur G. Humes Pseudione parviramus and Aporobopyrus collardi, two new species of Bopyridae (Isopoda: Epicaridea) from the Gulf of Mexico Daniel L. Adkison New records of fish parasitic marine isopod crustaceans (Cymothoidae, subfamily Anilocrinae) from the Indo-west Pacific Niel L. Bruce and Elizabeth B. Harrison-Nelson Booralana tricarinata, a new species of isopod from the western Atlantic Ocean (Crustacea: Isopoda: Cirolanidae) David K. Camp and Richard W. Heard Ipanemidae, new family, Ipanema talpa, new genus and species, from the surf zone of Brazil (Crustacea: Amphipoda: Haustorioidea) J. L. Barnard and James Darwin Thomas New records for Ogyrides alphaerostris and a new species, Ogyrides tarazonai (Crustacea: Ogyrididae), from the eastern Pacific Ocean Mary K. Wicksten and Matilde Mendez G. Notes on albuneid crabs (Crustacea: Decapoda: Albuneidae) from the central east coast of Florida Raymond B. Manning Heterocarpus cutressi, new species, and Plesionika macropoda Chace, 1939: two caridean shrimps of the family Pandalidae (Crustacea: Decapoda) from Puerto Rico and the U.S. Virgin Islands Oscar E. Monterrosa Notes on the freshwater crabs of the genus Moritschus Pretzmann, 1965 (Crustacea: Decapoda: Pseudothelphusidae) with description of M. narinnensis from southern Colombia Martha R. Campos and Gilberto Rodriguez Cambarus (Erebicambarus) maculatus, a new crayfish (Decapoda: Cambaridae) from the Mer- amec River Basin of Missouri Horton H. Hobbs, Jr., and William L. Pflieger Types of stomatopod crustaceans in the Zoological Survey of India H. C. Ghosh and Raymond B. Manning Pycnogonida of the western Pacific islands IV. On some species from the Ryukyu Islands K. Nakamura and C. Allan Child Poecilochaetus koshikiensis, a new polychaete species from Shimo-Koshiki Island, Japan Tomoyuki Miura Neopilina goesi, a new Caribbean monoplacophoran mollusk dredged in 1869 Anders Warén Notes on some Rhyssoplax from the Pacific Ocean (Mollusca: Polyplacophora: Chitonidae) Robert C. Bullock A review of the marine nematode genera Platycoma and Proplatycoma, with a description of Proplatycoma fleurdelis (Enoplida: Leptosomatidae) W. Duane Hope International Commission on Zoological Nomenclature: Opinions and Applications 475 487 496 503 509 SET) 530 540 549 568 576 585 603 614 622 626 633 640 644 653 662 671 676 682 693 707 MNGS pwd hbo, VOLUME 101 NUMBER 4 7 DECEMBER 1988 ISSN 0006-324X THE BIOLOGICAL SOCIETY OF WASHINGTON 1988-1989 Officers President: Kristian Fauchald Secretary: G. David Johnson President-elect: Leslie W. Knapp Treasurer: Don E. Wilson Elected Council Gary R. Graves Meredith L. Jones W. Ronald Heyer Raymond B. Manning W. Duane Hope Wayne N. Mathis Custodian of Publications: David L. Pawson PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Stephen D. Cairns Plants: David B. Lellinger Insects: Robert D. Gordon Vertebrates: Richard P. Vari Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $15.00 include subscription to the Proceedings of the Biological Society of Washington. Library subscriptions to the Proceedings are: $25.00 within the U.S.A., $30.00 elsewhere. The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued sporadically) are available. Correspondence dealing with membership and subscriptions should be sent to The Treasurer, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, National Museum of Natural History, Washington, D.C. 20560. THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 709-716 CRYPTOTROCHUS, NEW GENUS AND TWO NEW SPECIES OF DEEP-WATER CORALS (SCLERACTINIA: TURBINOLIINAE) Stephen D. Cairns Abstract.—A new genus in the Turbinoliinae is described: Cryptotrochus, characterized by having an imperforate theca, a papillose columella, and well- developed P2. A partial key to the 21 recognized genera of Turbinoliinae shows it to be similar to Kionotrochus, but differing in corallum shape and in having well-developed P2. Two widely geographically separated new species are de- scribed in this genus: C. carolinensis, known only from 320-338 m off North Carolina, and C. javanus, known only from 585 m in the Java Sea. Species of the subfamily Turbinoliinae have small, solitary, unattached coralla that are completely invested by the polyp. This complete investiture of the corallum by soft tissue allows for deep intercostal furrows from the calice to the base, which is char- acteristic of the subfamily. They are among the smallest of the Scleractinia; one species, Oryzotrochus stephensoni Wells, 1959, has an adult calicular diameter of 1.5 mm, and the genus is so named for its resemblance in size and shape to a grain of rice. There are approximately 118 species of Turbinoliinae attributed to 21 genera, eight of which are exclusively fossil genera. Species in the subfamily are known from the Upper Cretaceous to the Recent and are worldwide in distribution (including the Antarctic), the Recent species recorded from depths of 9- 835 m. Some species of Turbinoliinae are known to be interstitial in habitat. Species of Turbinoliinae are most common in the Australian-New Zealand and Indonesian regions, especially in the Miocene; only five species are described from the Atlantic Ocean. While studying the deep-water coral col- lection of the R/V Gosnold at the National Museum of Natural History (USNM) in 1977, I discovered a species of Turbinoli- inae from off North Carolina (Gosnold sta. 1811) that was very different from any yet described from the Atlantic but quite sim- ilar to an Indo-West Pacific species (1.e., No- tocyathus conicus (Alcock, 1902)). Thinking that this single lot of specimens might be improperly labelled, I placed it aside. Sev- eral years later I discovered a damaged basal part of the same species from Gosnold sta. 1841, not far from Gosnold sta. 1811. More recently, I diagnosed two specimens collected from the Java Sea by the R/V Gal- athea as having the same generic attributes. In reviewing all genera of the Turbinoliinae in preparation for a revision of the deep- water Scleractinia of the Philippine Islands (Cairns, in prep.), I determined that the Gosnold and Galathea samples are two new species and represent a new genus, de- scribed herein as Cryptotrochus. A key is provided for the imperforate paliferous gen- era of Turbinoliinae (9 of 21 genera in the subfamily) to help distinguish the new ge- nus. Order Scleractinia Bourne, 1900 Suborder Caryophylliina Vaughan and Wells, 1943 Family Caryophylliidae Dana, 1846 Subfamily Turbinoliinae Milne Edwards and Haime, 1848 Cryptotrochus, new genus Diagnosis.—Solitary, conical, free coral- lum with pointed base. Imperforate theca. 710 Well-developed, discrete P2. Columella pa- pillose. Ahermatypic. Discussion.—Among the 21 genera of Turbinoliinae, nine can be characterized as having pali and an imperforate theca (see Key). Of these nine, only two genera have pali restricted to the second-cycle septa (P2): Kionotrochus and Cryptotrochus. Although the type species of Kionotrochus, K. suteri Dennant, 1906, was originally described as lacking pali, and Wells (1956) also defined the genus as lacking pali, Squires (1960) cor- rectly noted that K. suteri does, in fact, have small, rod-like P2 that merge almost indis- tinguishably with the columella (Fig. 13). This is the basic difference between the two genera: Kionotrochus has indistinct rod-like P2, whereas Cryptotrochus has discrete la- mellar P2. Another important difference is corallum shape: Cryptotrochus has a conical corallum with a pointed base; Kionotrochus has a bowl-shaped corallum with a rounded base (Fig. 14), often the result of transverse division (Squires 1964). Based on corallum shape, it is unlikely that Cryptotrochus re- produces by transverse division. Other less important differences are that species of Cryptotrochus have highly exsert S1—2 and four cycles of septa, wheras K. suteri has moderately exsert septa and only three cycles of septa. Kionotrochus was divided into two subgenera by Squires (1960): the nominal subgenus and K. (Cylindrophyllia). Cylin- drophyllia is considered to be a junior syn- onym of Peponocyathus (Cairns, in prep.), resulting in Kionotrochus s. str. being mono- typic. It is endemic to northeastern New Zealand, 48-241 m (Squires & Keyes 1967). Although not apparent from the key, Cryptotrochus is very similar to Notocy- athus, being similar in size, corallum shape, septal exsertness, costae, and columella. The only point of difference concerns the pali: Cryptotrochus has six P2, whereas Notocy- athus has 12 P3, arranged in 6 V-shaped pairs, the Pl and P2 of Notocyathus usually being suppressed in the adult stage. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Two species are assigned to Cryptotro- chus, C. carolinensis and C. javanus; how- ever, two more undescribed species are known from off New Caledonia and the Chesterfield Islands (H. Zibrowius, pers. comm. and examination of specimens). One differs from C. javanus in having a more slender corallum, having only one row of granules per costa, and having less promi- nent pali. It is known from a similar depth range: 555-680 m. The second undescribed species differs from C. javanus in having prominent lateral granules on costal edges, S3 almost as large as S2, and highly granular septa. It is known only from 223-345 m. Etymology.—Cryptotrochus is a combi- nation of ‘“‘kryptos” (Greek: hidden, con- cealed) and “trochos’’ (Greek: wheel), a common sufhx of turbinoliid generic names, together alluding to the inconspicuous pres- ence of these small corals, which are only now being described from the Atlantic. The gender is masculine. Type species.—Cryptotrochus carolinen- sis, nN. sp., here designated. Distribution. — Currently known only from off North Carolina and Java Sea, 320-585 m. Also New Caldonia and Chesterfield Is- lands, 223-680 m (H. Zibrowius, pers. comm.). Partial Key to the Genera of Turbinoliinae with Emphasis on the Imperforate Paliferous Genera (+ denotes exclusively fossil genera) — . Corallum perforate + Turbinolia Lamarck, 1816; Cono- cyathus Orbigny, 1849; Trema- totrochus Tenison-Woods, 1879; + Bothrophoria Felix, 1909 1’. Corallum imperforate Columella absent . +Dominicotrochus Wells, 1937 2’. Columella present 3. Pali absent Sphenotrochus Milne Edwards and VOLUME 101, NUMBER 4 a 6’. 8. Haime, 1848; Platytrochus Milne Edwards and Haime, 1848; + Koilotrochus Tenison- Woods, 1878; Holcotrochus Dennant, 1902; +Sphenotro- chopsis Alloiteau and Tessier, 1958; Oryzotrochus Wells, 1959; Wellsotrochus Squires, 1960 Melealiepresentmerte sn ce ee 4 Onlvae2spresent a eee 5 FS DTeSeMt ante: come, eee eee 6 . Pali before all but last cycle of sep- ta (usually P1-—3), but P1l—2 sup- pressed in adult Notocyathus ... 8 P2 fused to columella, obscure; corallum bowl-shaped with ROUNGECIDASE see es ee: Best t Kionotrochus Dennant, 1906 P2 independent and prominent; corallum conical with pointed base see eee Cryptotrochus, n. gen. Septa correspond to costae; trans- verse division lacking ree +Coronocyathus Alloiteau and Tessier, 1958 Septa alternate with costae; trans- verse division present Corallum discoidal; pali in one crown of 12 elements; columella spongy Dunocyathus Tenison-Woods, 1878 . Corallum cuneiform; pali in 2 crowns of 10-12 elements; colu- melaspapulloseme nn ae lee. Idiotrochus Wells, 1936 Eighteen primary septa bet + Monticyathus Alloiteau and Tessier, 1958 Typical hexameral symmetry (SHES 2 SBES4)) seas oe 9 P3 fused to one another in V-shaped pairs; P2 usually sup- pressed in adult; septa indepen- dent; corallum conical, with pointed base Notocyathus Tenison-Woods, 1880 . P3 fused to intermediate P2; P2 711 well developed; higher cycle septa fuse with lower cycle septa; cor- allum of variable shape, but never with a pointed base ........... 10 Corallum relatively large (adult calicular diameter over 10 mm), cuneiform; P1 well developed .. Tropidocyathus Milne Edwards and Haime, 1848 Corallum small (adult calicular diameter less than 8 mm) and of variable shape (e.g., cylindrical, hemispherical); Pl often poorly developed .. Peponocyathus Gravier, 1915 (=Cylindrophyllia Yabe and Eguchi, 1937) HO; 10’. Cryptotrochus carolinensis, new species Figs. 1-9 Types.—Holotype: Gosnold sta. 1811, USNM 46914.—Paratypes: Gosnold sta. 1811 (8) USNM 46915; Gosnold sta. 1841 (1) USNM 81082, 33°38.5'N, 76°29.3'’W, 338 m. Type locality: Gosnold sta. 1811: 33°00.5'N, 77°16.2’W (off Cape Fear, North Carolina), 320 m. Description.—Corallum conical, with pointed basal angle of 55°-65°, the corallum becoming cylindrical at height of about 5.5 mm. No evidence of asexual regeneration from basal fragments. Largest specimen (holotype) 6.4 mm in calicular diameter and 7.7 mm tall; calice circular. Cl1—2 measures 0.16—0.17 mm wide at calicular edge; C3- 4 narrower, about 0.10 mm wide. Costae separated by intercostal furrows 0.17—0.21 mm wide and about 0.55 mm deep at ca- licular edge. Each costa bears a unilinear row of outward projecting triangular teeth, each about 0.14 mm in basal width and height, producing a serrate costal edge. Smaller cylindrical granules, 0.09 mm tall and 0.045 mm in diameter, adorn the lateral edges of costae. Costae arranged in typical turbinoliid fashion (Fig. 2): 12 Cl-—2 origi- 712 nate at the base, each C2 bifurcating into 2 C3 very close to the base (0.3 mm from epicenter), and each C3 bifurcating into 2 C4 about 1.5 mm from base epicenter, the original C2—3 continuing beyond points of bifurcation. At level of C3—4 bifurcation, the 12 Cl—2 abruptly decrease in width to accommodate the doubled number of cos- tae. Theca about 0.15 mm thick; corallum white. Septa hexamerally arranged in 4 cycles according to the septal formula: S1>S2=2S3>S4. S1 highly exsert (up to 1.4 mm) and extend about 0.8 distance to col- umella. S1-3 have straight, vertical inner edges and rounded upper edges. S2 less exsert (about 1.1 mm) and extend slightly over '2 distance to columella. Each S2 bordered in- ternally by a large, flat palus (P2), each about 0.60 mm deep and 0.3 mm thick. S3 slightly less exsert than S2 but extend an equal dis- tance to columella, at which point their low- er inner edges create a porous fusion with the P2. S4 slightly less exsert than S3 and extend toward columella only about '4 the distance, at which point their entire inner edges create a porous fusion with the ad- jacent S3. Septal faces covered by low (about 0.07 mm tall) triangular granules aligned in rows perpendicular to septal edges (Figs. 7— 9). Granulation of pali coarser, with blunt granules up to 0.15 mm tall. Fossa very shallow to nonextant, con- taining the 6 P2 and columella. Rounded upper edges of P2 extend above calicular edge but below the upper edges of the S4; inner edges of pali fused to columella. Col- umella composed of 7-9 tuberculate pa- pillae, fused among themselves and to sur- rounding pali. Upper edges of columellar papillae stand above upper edges of pali, equally as high as the S4. Pali and colu- mellar elements morphologically discrete. Discussion.—C. carolinensis is distin- guished from C. javanus, the only other species in the genus, by three characters: 1) the inner edges of the higher-cycle septa of C. carolinensis are fused to the lower-cycle PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON septa or pali (S4 to S3, S3 to P2); septa of C. javanus are independent, 2) its costae bear a uniserial row of spines; costae of C. jJavanus bear up to three rows of granules per costa at the calicular margin, and 3) its pali have rounded upper edges that do not rise above the level of the columella; the pali of C. javanus have slender, pointed up- per edges that rise well above the columella. Etymology.— This species is named car- olinensis for North Carolina, its type-local- ity. Distribution.— Known only from off Cape Fear, North Carolina, 320-338 m. Cryptotrochus javanus, new species Figs. 10-12 Types.— Holotype: Galathea sta. 490, Zoologisk Museum, Copenhagen.—Para- type: Galathea sta. 490 (1) Zoologisk Mu- seum, Copenhagen. Type locality: 5°25’S, 117°03’E (eastern Java Sea), 585 m. Description.—Corallum conical, with a pointed basal angle of 75°, the corallum be- coming cylindrical at a height of about 6.0 mm. No evidence of asexual regeneration from basal fragments. Holotype 9.2 mm in calicular diameter and 10.0 mm tall; calice circular. Costae equal in width near calic- ular edge, about 0.32 mm wide, separated by deep intercostal furrows about 0.18 mm wide and 0.60 mm deep. Basally, each costa bears one row of coarse teeth; several mm from the base epicenter, each costa bears 2 rows of teeth; and near the calicular edge, the C1—2 each bear 3 rows of outward-pro- jecting coarse teeth. Costal arrangement identical to that of C. carolinensis. Corallum white. Septa hexamerally arranged in 4 cycles according to the formula: S1>S2>S3>S4. S1 highly exsert (up to 1.5 mm) and extend about 0.8 distance to columella. S1—2 have straight, vertical inner edges and rounded upper edges. S2 slightly less exsert and ex- tend about 0.7 distance to columella. Each S2 bordered internally by a tall, flattened, VOLUME 101, NUMBER 4 M3 Figs. 1-6. Cryptotrochus carolinensis (1, 3-4, Holotype, USNM 46914; 2, 5-6, Paratype from Gosnold sta. 1811, USNM 46915): 1, Lateral view of holotype, x 7.1; 2, Corallum base showing typical costal bifurcations, x 18; 3-4, Calicular views of holotype, x 7.8; 5-6, Costae, showing large triangular outward projecting teeth and smaller cylindrical lateral granules, x56, x 202, respectively. 714 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 7-9. Cryptotrochus carolinensis, paratype from Gosnold sta. 1811, USNM 46915: 7, Stereo view of broken corallum exposing septocostal face, aligned septal granules, palus, and cross section of theca, x20; 8, Oblique view of septocostal face, x 34; 9, Enlargement of septal granules, x 260. VOLUME 101, NUMBER 4 715 Figs. 10-14. (10-12, Holotype of Cryptotrochus javanus) 10, Stereo view of calice, x 5.9; 11, oblique view of calice showing relative septal exsertness, 6.1; 12, Lateral view of corallum showing costal pattern, x 6.6. (13-14, Kionotrochus suteri, topotypic specimen from off Cuvier Island, New Zealand, 38 fm (=70 m), USNM 78586) 13-14, Calicular and lateral views, x 12.8, x 14.6, respectively. 716 sharp palus (P2) having a vertical inner edge. S3 less exsert than S2 and extend about 0.6 distance to columella; their inner edges are slightly sinuous and not fused to any other septa. S4 less exsert than S3 and extend about 0.3 distance to columella; their inner edges also free, not fused to adjacent septa. Upper septal faces smooth, without granulation, but lower septal faces sparsely covered by low pointed granules. Fossa moderately deep, containing the 6 P2 and columella. Upper edges of pali ex- tend just above calicular edge. Columella composed of 2 papillae, terminating below level of pali. Discussion.—Comparisons to C. caroli- nensis are made in the account of that species. Etymology.—This species is named ja- vanus for the Java Sea, in reference to its type-locality. Distribution.— Known only from the type- locality. Acknowledgments I would like to thank H. Zibrowius (Sta- tion Marine d’Endoume, Marseille) for commenting on an early draft of the manu- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON script, and K. W. Petersen (Zoologisk Mu- seum, Copenhagen) for permission to ex- amine and publish on the Galathea specimens. Literature Cited Cairns, S. D. (in prep). A revision of the deep-water Scleractinia of the Philippine Islands and ad- jacent waters. Part 1. Fungiacyathidae, Micra- baciidae, Turbinoliinae, Guyniidae, and Fla- bellidae.—Smithsonian Contributions to Zoology. Squires, D. F. 1960. The scleractinian genus Kiono- trochus and Cylindrophyllia.—Records of the Dominion Museum 3(4):283-288. 1964. New stony corals (Scleractinia) from northeastern New Zealand.—Records of the Auckland Institute Museum 6(1):1-9. —., & I. W. Keyes. 1967. The marine fauna of New Zealand: scleractinian corals.—New Zea- land Department of Scientific and Industrial Re- search Bulletin 185: 46 pp., 6 pls. Wells, J. W. 1956. Scleractinia. Pp. F328—-F444 in R. C. Moore, ed., Treatise on invertebrate paleon- tology, Part F, Coelenterata. University of Kan- sas Press, Lawrence, Kansas. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 717-729 SYRINGONOMUS DACTYLATUS, A NEW SPECIES OF BATHYAL MARINE NEMATODE (ENOPLIDA: LEPTOSOMATIDAE) AND A SUPPLEMENTARY DESCRIPTION OF SYRINGONOMUS TYPICUS HOPE AND MURPHY, 1969 W. Duane Hope Abstract.—The taxonomic history of the genus Syringonomus is reviewed, and the genus is redefined. A supplementary description of the males of Syringo- nomus typicus is given, and Syringonomus dactylatus is described. The structure of the amphidial flap of the males is compared and contrasted between both species of Syringonomus, and between species of Syringonomus, Platycoma and Proplatycoma. The geographic distribution and ecology of Syringonomus is discussed and compared with that of Platycoma and Proplatycoma. The genus Syringonomus has, until now, contained a single species, S. typicus Hope & Murphy, 1969. The description of the type species was based upon specimens col- lected in the North Atlantic at a depth of 3806 m, which remains the only record of its occurrence. The report that S. typicus had been collected from near Recife, Brazil (Bongers 1983:855) is based upon misiden- tified specimens of the new species, Syrin- gonomus dactylatus. Syringonomus was assigned by Hope & Murphy (1969) to the subfamily Leptoso- matinae (Leptosomatidae). However, the sexually dimorphic males of Syringonomus are unique among members of this subfam- ily in that the neck 1s slightly narrowed and the cuticle is thickened between the amphid and cephalic sensilla. Also, the amphids were originally described as having the shape of an “inverted lyre,’ with a fringed border, and with or without a tapered process di- rected posteriorly from the center of the an- terior edge of the lyre. An amphid by this description, in fact, is not known among other marine nematodes. The males of Syringonomus dactylatus, which in other regards resemble the males of S. typicus, have a lobed flap, obvious even in light microscopic observations, that covers the amphidial aperture. This obser- vation made it necessary to question the original interpretation of the amphid of S. typicus. Consequently, the amphid was studied with the SEM, and the results are included in the following supplementary de- scription of the type species. Materials and methods.-The original de- scription of the males of S. typicus was based upon the holotype (USNM 39489) and three paratypes (USNM 39490-39492) (USNM = National Museum of Natural History). Of the three paratypes, one has been sec- tioned for histological examination, and another is a young male that was moulting when fixed and is in poor condition. There- fore, of the original type material, only the holotype (USNM 39489) and one paratype (USNM 39490) were used in this redescrip- tion of S. typicus. However, three additional males (USNM 77178-77180) have been in- cluded in this redescription. They were not part of the original type series, but were sort- ed, for the purposes of this study, from the same samples as those from which the type material had originally been obtained. One 718 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Morphometric and meristic data for males of Syringonomus typicus. USNM number 39489 39490 77178 77179 77180 Midbody diameter 83 95 — 77 100 Body width at base of esophagus 70 80 — 74 82 Body width at level of nerve ring 69 74 — 70 74 Head width at level of cephalic sensilla 32 30 D2) 33 28 Length of cephalic capsule on dorsal and ven- 5/3 5/4 —/— 6/5 3/2 tral sides of head, d/v Distance from oral surface to outer labial and 17 15 13 16 15 cephalic sensilla Length of outer labial sensilla 3,3,4,4 3,3 3,3,3,4 3,4,4,4,5 4,5 Length of cephalic sensilla 5,5,5 5,6 4,4,4,5 5,5,6,6 5,6 Number cervical sensilla anterior end to nerve 2/3 1/2 —/— 4/2 5/4 ring; right/left Length of amphidial flap, r/1 3/3 3/5 —/3 6/— 5/6 Anterior width of amphidial flap; r/1 7/7 7/7 —/4 5/— 5/5 Posterior width of amphidial flap; r/I 10/10 9/11 —/9 15/9 8/9 Length of amphidial lobes; r/1 5/5 6/5 5/4 —/3 5/6 Distance from oral surface to posterior margin 27/31 27/31 —/24 32/30 32/31 of flap; r/1 Distance from oral surface to posterior end of 677/706 762/700 —/— 840/849 800/792 amphidial gland (% of esophagus length); r/1 (92/96) (107/98) —/— (96/97) (91/90) Distance from oral surface to orifice of dorsal 43 31 — 39 34 esophageal gland Distance from oral surface to nerve ring 300 305 — 368 344 Length of esophagus 736 715 = 872 879 Position of testes relative to gut (v = ventral; | v/l v/l —/— V1 v/\ = left); anterior/posterior Length of spicula; r/I 80/77 80/73 —/— —/72 $2/55 Length of lateral accessory component; r/1 14/13 —/— —/— 15/16 11/13 Cloacal body diameter 79 83 — 88 85 Tail length 142 138 — 143 132 ce —’”’ indicates data that were not obtainable. of these three topotype males (USNM 77178) was critical point dried for SEM by a method described elsewhere (Hope 1982: 2). The technique differs from the original in that a graded series of acetone was used for dehydration, and amyl acetate was ex- cluded from the process. The specimen was precoated with carbon, coated with gold/ palladium, and examined with an Hitachi S-570 scanning electron microscope. The description of S. dactylatus is based upon specimens collected with an epiben- thic sled at station 167, cruise 31 of Atlantis IT, and donated to the National Museum of Natural History by the Woods Hole Ocean- ographic Institution. All specimens of both species were fixed in 4% formalin in sea water. Those em- ployed in light microscopic observations are mounted in anhydrous glycerin between coverslips on Cobb aluminum frames. Mor- phometric data were obtained by measure- ments from camera lucida drawings or elec- tron micrographs. Morphometric and meristic data for each specimen of S. typicus are given in Table 1, and for each specimen of S. dactylatus in Table 2. In some cases, several data are given for one character, such as the length of each of several outer labial or cephalic sensilla, which are separated from one another by commas. The range, mean, and plus or minus one standard de- VOLUME 101, NUMBER 4 719 Table 2.—Morphometric and meristic data for Syringonomus dactylatus. Males Females Juv USNM number 77172 77173 77174 TiS Wii Midbody diameter 87 84 124 103 76 Body width at base of esophagus 78 79 88 82 68 Body width at level of nerve ring 70 Vi 70 73 59 Head width at level of cephalic sensilla 30 30 30 28 29 Length of cephalic capsule on dorsal and ventral NA —/3 6/4 6/5 4/5 sides of head, d/v Distance from oral surface to outer labial and 23 14 13 15 15 cephalic sensilla Length of outer labial sensilla DMB Sr DDD DB} 2,3,3 2,3,3,3 3,3 Length of outer cephalic sensilla 3,3,3 3,3,3,3 4,5,5 3,4,5 4,4,5 Number of cervical sensilla anterior end to nerve 5/3 3/4 5/3 6/5 7/6 ring; right/left Length of cervical sensilla DES Dp 2,3,3,4 2,4 3,5 Length of amphidial flap; r/1 5/4 4/3 NA NA NA Anterior width of amphidial flap; r/1 5/5 7/6 NA NA NA Posterior width of amphidial flap; r/I 10/11 11/11 NA NA NA Length of anterior amphidial lobes; r/] 8.8/8,8 8,8/8,8 NA NA NA Length of dorsal amphidial lobes; r/1 9/9 8/6 NA NA NA Length of ventral amphidial lobes; r/1 7/8 Wi NA NA NA Distance from oral surface to posterior margin of flap 39/29 30/30 17/18 21/20 20 (males); to amphidial aperture (females); r/1 Distance from oral surface to posterior end of 846/884 707/740 NA NA NA amphidial gland (% of esophagus length); r/1 (86/90) (73/76) Distance from oral surface to orifice of dorsal 31 23 26 21 24 esophageal gland Distance from oral surface to nerve ring 331 333 295 307 235 Length of esophagus 988 970 851 943 673 Distance from oral surface to excretory pore 119 148 NA NA NA Position of gonads relative to gut (r = right; v = v/r r/r v/v v/v NA ventral); anterior/posterior Length of spicula on arc; r/I 93/84 79/69 NA NA NA Length of lateral accessory component; r/1 14/15 14/— NA NA NA Cloacal and anal body diameters 79 79 83 75 63 Tail length 121 123 120 103 82 NA indicates data that are not applicable; ‘“‘“—’’ indicates data that were not obtainable. viation of the data are given in the text. The points between which the various measure- ments of the amphidial flaps and lobes were made is depicted in Fig. 1A and B. Syringonomus Hope & Murphy, 1969 Syringonomus Hope & Murphy, 1969:511. Type species.—Syringonomus typicus Hope & Murphy, 1969:512. Emended diagnosis. —Leptosomatidae. Body elongate and spindle-shaped. Anterior end bluntly rounded. Neck slightly reduced in width at level of amphid, slightly tapered from amphid to level of cephalic sensilla, and head rounded anteriorly from level of cephalic sensilla. Tail bluntly conical. Cu- ticle with fine transverse striae, at least pos- terior to amphid. Cuticle of neck in males thickened between level of amphids and ce- phalic sensilla; not thickened in females. Cephalic capsule present or absent in males; present in females. Amphidial aperture of males wide, slit-like, and covered with lobed 720 amphidial flap; cephalic cuticle fringed at edge of amphidial flap and outside edge of lobes. Amphid of females cyathiform with small, pore-like aperture; amphidial flap ab- sent. Amphidial glands well developed in males, apparently absent in females. Lips short, flap-like, directed anteriorly and not set off by microlabial groove. Buccal capsule unarmed. Anterior end of esophagus at- tached to cephalic cuticle; cephalic cuticle at anterior end of esophagus not thickened. Oral aperture dilated by retrodilator mus- cles in anterior end of esophagus. Dorsal esophageal gland orifice in lumen of esoph- agus; subventral esophageal gland orifices apparently at anterior end of buccal capsule. Dorsolateral and ventrolateral orthometa- nemes present; some with, others without, caudal filaments. Spicula with paired, ven- trolateral, accessory structures. Gubernac- ulum absent and ventromedian supple- ments absent. Caudal glands well developed, and extended anteriorly beyond level of clo- acal and anal vents. Syringonomus typicus Hope & Murphy, 1969 Figs. 1A; 2A—C Nec Syringonomus typicus sensu Bongers, 1983:855. Material examined. — Holotype. —Male, USNM 39489. Paratype. —Male, USNM 39490. Topotypes. —Males, USNM 77178, 77179 and 77180. Type locality.—Sediment from epiben- thic trawl collected between 39°37.0’N, 66°47.0’W and 39°37.5'N, 66°44.0’W at 3806 m on 24 Aug 1966. Although males, USNM 77178 thru 77180 are not paratype specimens, they are from the type locality. Supplementary description of males. — Total a b Cc length USNM 39489 65.9 13 37.9 5.377 mm PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON USNM 39490 65.3 8.7 44.9 6.206 mm USNM VIMY) 89.4 7.8 47.9 6.844 mm USNM 77180 58.2 6.6 44.1 5.824 mm Body elongate, gradually tapered from midbody to level of nerve ring and anal vent. Midbody diameter 77-100 (89 + 11) um; body width at base of esophagus 70-82 (77 + 5.5) um, at level of nerve ring 69-74 (72 + 2.6) um, and at level of cephalic sensilla 22-33 (29 + 4.4) um. Cervical region slight- ly constricted at level of amphid, then slightly tapered to level of cephalic sensilla. Head (Fig. 2A) rounded anteriorly. Cuticle thickened at level of amphid. External sur- face of cuticle posteriorly from amphid with very shallow, transverse striae (Fig. 2A, C); periodicity of striae 300 nm. Cephalic cap- sule apparent when head viewed in optical sagittal section; length of cephalic capsule 3-6 (5 + 1.3) and 2—5 (4 + 1.3) um on dorsal and ventral sides of head respectively. Inner labial papilliform sensilla not ob- served. Head with single circle of 6 setiform, outer labial sensilla and 4 setiform cephalic sensilla (Fig. 2A), 13-17 (15.2 + 1.5) wm from oral surface of head. Outer labial sen- silla 3-5 (3.7 + 0.7) wm long; cephalic sen- silla 4-6 (5.1 + 0.7) um long. Cervical re- gion between cephalic sensilla and nerve ring with 1-5 (3 + 1.8) setiform sensilla on right side, and 2—4 (2.8 + 1.0) on left; length of cervical sensilla ca. 4 wm. Cuticle at posterior rim of amphidial ap- erture extended anteriorly, flap-like over aperture (Figs. 1A; 2A—C); anterior end of each flap with paired, anteriorly directed, finger-like lobes (Fig. 2A—C), tapered and rounded at tip. Length of flap (Fig. 1A) on right side of head 3-6 (4.3 + 1.5) wm, and 3-6 (4.3 + 1.5) on left. Width of anterior end of flap on right side of head 5-7 (6.0 + 1.2), and on left side 4-7 (5.8 + 1.5) um; width at posterior end of flap on right side VOLUME 101, NUMBER 4 Fig. 1. 72M Diagram showing location of measurements of the amphidial flap of 1A.) Syringonomus typicus and 1B.) S. dactylatus A. Length of flap. B. Minimum (anterior) width of flap. C. Maximum (posterior) width of flap. D. Length of central anteriorly directed lobes. E. Length of dorsal and ventral lobes. F. Distance from oral surface of head to posterior margin of flap. of head 8-15 (10.5 + 3.1), and on left 9- 11 (9.6 + 0.9) um; length of lobes on right side of head 5—6 (5.3 + 0.5), and on left 3- 6 (4.9 + 0.9) um. Distance from oral surface of head to posterior margin of flap on right side of head 27-32 (29.5 + 2.9), and on left 24-31 (29.4 + 3.1) um. Cephalic cuticle with fringed border (Fig. 2B) at edges of amphidi- al flaps and lobes. Amphidial glands well developed; distance from oral surface of head to posterior end of right amphidial gland (% of esophagus length) 677 (92%) to 840 (107%) (770 + 70 um; 97 + 7%), and to posterior end of left amphidial gland 700 (90%) to 849 (98%) (762 + 72 wm; 95 + 4%). Edges of amphidial aperture dorsal and ventral from flap and lobes often with pre- sumed secretion of amphidial gland (Fig. 2A, B). Buccal aperture (Fig. 2C) triradiate and surrounded by 3 anteriorly directed, oral flaps (Fig. 2A, C). Anterior to posterior length of each flap greatest midway between corners of oral aperture; flaps not set off from head by groove. Buccal armature ab- sent. Orifice of dorsal esophageal gland in cuticular wall of esophageal lumen 31-43 (37 + 5.3) wm from oral surface of head. Subventral esophageal glands apparently with orifices at anterior end of buccal cap- sule. Esophagus cylindrical, slightly narrower anteriorly; external surface of esophagus smooth. Distance from oral surface of head to anterior edge of nerve ring 300-368 (329 + 33) wm. Length of esophagus 715-879 (801 + 87) um. Renette not observed. Orthometanemes present at dorsal and ventral margins of lateral hypodermal chords; caudal filaments present in most metanemes, absent in others. Hypodermal glands absent. Testes paired and opposed; testes on ven- tral or left side of gut. Spicula paired, curved ventrally; right spicula 52-80 (71 + 16.2) and left 55-77 (70 + 8.6) um long, mea- sured on arc; capitulum not distinct. Lateral accessory component at distal end of each spiculum; right accessory component | 1-15 (Bs==o2 5) -andileft ISG nC seeeale7))eeem long. Gubernaculum not apparent. Dorso- ventral copulatory muscles on each side of body separated from one another. Precau- dal, subventral, setiform sensilla present; not distinguishable from somatic setae. Ventro- median supplement absent. Tail bluntly conical, 132-143 (139 + 5) um long, with subdorsal and subventral se- 722 tiform sensilla. Cloacal body diameter 79- 88 (84 + 3.9). Cuticle of tail terminus with median, crescent-shaped lamella penetrated by spinneret. Cell bodies of each of 3 caudal glands extend anteriorly beyond cloacal vent. Syringonomus dactylatus, new species Figs. 1B; 3A—-C; 4A, B Syringonomus typicus sensu Bongers, 1983: 855. Material examined. — Holotype. —Male, USNM 77172. Paratypes. —Male, USNM 77173; fe- males USNM 77174 and 77175; juvenile USNM 77176. Type locality. —Approximately 52 km off coast of Recife, Brazil, 7°58.0’—7°50.0’'S; 34°17.0'W at a depth of 943-1007 m. Col- lected 20 Feb 1967. Description. — Males.- Total a b Cc length USNM VUNT2 76.3 6.7 54.3 6.613 mm USNM 77173 78.1 6.7 53.1 6.528 mm Body elongate, slightly tapered from mid- body to level of nerve ring and anal vent. Midbody diameter 84—87 (85.5 + 2.1) um; body width at base of esophagus 78-79 (78.5 + 0.7) um, at level of nerve ring 70-77 (73.5 + 4.9) um, and at level of cephalic sensilla 30 um in both specimens. Neck region (Fig. 3A) slightly constricted and cuticle thick- ened between level of amphid and cephalic sensilla. Thickness of cuticle posterior to amphids 4 um, and between amphid and cephalic sensilla 7 wm in both specimens. Head rounded anteriorly. Cuticle posterior from head with fine transverse striae; pe- riodicity of striae ca. 550 wm. Cephalic cap- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sule absent in holotype (Fig. 3A), apparent only on ventral side in optical sagittal sec- tion of male paratype; length 3 um. Papilliform, inner labial sensilla not ob- served. Outer labial sensilla and cephalic sensilla setiform (Fig. 3A) and in single cir- cle of ten, 14-23 (18.5 + 6.4) um from oral surface. Outer labial sensilla 2—3 (2.4 + 0.5) um long; all cephalic sensilla in both spec- imens 3 wm long. Cervical region between amphid and nerve ring with 3-5 setiform, sensory sensilla on right side of body, and 3-4 on left (3.8 + 1.0); length of cervical sensilla 2—3 (2.3 + 0.5) wm. Cuticle at posterior rim of amphidial ap- erture extended anteriorly, flap-like over aperture; amphidial flap (Figs. 1B; 3A) with 2 central, anteriorly directed lobes and 1 dorsal and | ventral lobe; dorsal and ventral lobes directed obliquely towards anterior; all lobes tapered slightly and distally round- ed. Edge of aperture extended dorsally and ventrally beyond posterior end of flap. Ce- phalic cuticle apparently with fringe at edge of flap and lobes. Length of flap (Fig. 1B) 4-5 (4.5 + 0.7) um on right side of head and 3-4 (3.5 + 0.7) um on left; width of posterior end of flap 10-11 (10.5 + 0.7) wm on right, and 11 wm on left in both speci- mens; width at anterior end of flap 5-7 (6 + 1.4) wm on right and 5-6 (5.5 + 0.7) um on left; length of anterior lobes 8 um on both sides of both specimens; length of dorsal lobes 8-9 (8.5 + 0.7) on right side and 6— 9 (7.5 + 2.1) um on left side; and length of ventral lobes 7 um on both specimens on right side and 7-8 (7.5 + 0.7) um on left side. Distance from oral surface of head to posterior margin of flap 30-39 (34.5 + 6.4) um on right side, and 29-30 (29.5 + 0.7) um on left side. Amphidial glands (Fig. 4A) well developed; distance from oral surface of head to posterior end of gland (in wm and % of esophagus length) on right side 707 (73%) to 846 wm (86%) (777 + 98 um; 80 + 9%) and on left 740 (76%) to 884 um (90%) (812 + 102 wm; 83 + 10%). VOLUME 101, NUMBER 4 sei 723 Fig. 2. Syringonomus typicus. Scanning electron micrographs of male topotype, USNM 77178. A. Left side of head; scale equals 10 wm. B. Left amphidial flap; scale equals 5 um. C. Anterior view of head and left side of cervical region; scale equals 10 wm. Abbreviations.— AF, amphidial flap; AL, amphidial lobe; AS, amphidial secretion; CB, fringed border of amphidial aperture; OF, oral flap; TS, transverse striae. Buccal aperture triradiate; microlabia or oral flaps not evident in light microscopic observations (Fig. 3A). Buccal capsule tri- radiate and dilated by 3 obliquely oriented, labial retrodilator muscles (Fig. 3A), one in each sector of esophagus. Buccal armature absent. Orifice of dorsal esophageal gland (Fig. 3A) in lumen of esophagus 23-31 (27 + 5.7) um from oral surface of head. Sub- ventral esophageal glands (Fig. 3A) appar- ently with orifices at anterior end of buccal capsule. Esophagus cylindrical, slightly narrower anteriorly; exterior surface of esophagus smooth. Distance from oral surface of head to anterior edge of nerve ring 331-333 (332 + 1) um. Length of esophagus 970-988 (979 + 13) wm. Renette present and excretory 724 pore 119-148 (134 + 20.5) um from oral surface of head. Dorso- and ventrolateral orthometa- nemes present; some with caudal filaments, others without. Hypodermal gland cells ab- sent. Testes paired and opposed; testes on ven- tral or right of gut. Seminal vesicle straight without convolutions. Spicula (Fig. 3B) paired, curved ventrally; right spicula 79-— 93 (86 + 10) and left 69-84 (77 + 11) wm long, measured on arc; capitulum well de- veloped. Lateral accessory component (Fig. 3B) at distal end of each spiculum; right accessory component 14 um long on both specimens, and left 15 wm on only specimen measured; distal end thick-walled and nar- row; proximal end thin-walled and slightly flared. Gubernaculum not apparent. Dor- soventral copulatory muscles on each side of body (Fig. 3B) separated. Precaudal, subventral, setiform sensilla present (Fig. 3B); not distinguishable from other somatic setae. Length of setiform sup- plements | to 3 um on both specimens. Ven- tromedian supplement absent. Tail (Fig. 3B) bluntly conical, 121-123 (122 + 1) um long, with dorsolateral, sub- ventral and subterminal setiform sensilla. Cloacal body diameter 79 wm in both spec- imens. Cuticle on ventral surface of tail of holotype, 85 um posterior from anal vent, penetrated by hypodermis without evidence of setae or glands (Fig. 3B); not observed in paratype. Cuticle of tail terminus (Fig. 3B) with median, crescent-shaped lamella pen- etrated by spinneret. Cell bodies of each of 3 caudal glands extend anteriorly beyond cloacal vent (Fig. 3B). Females. — Body a b c Vv length USNM 77174 42 6 43 62% 5.156 mm USNM 77175 56 6 56 64% 5.791 mm Midbody diameter 103-124 (114 + 15) um; body width at base of esophagus 82-— PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 88 (85 + 4) um, at level of nerve ring 70— 73 (71.5 + 2) um, and at level of cephalic sensilla 28-30 (29 + 1) wm. Neck region uniformly tapered; head region (Fig. 3C) slightly constricted at level of amphid, but cuticle not thickened. Head rounded ante- riorly (Fig. 3C). Striae of cuticle shallow, periodicity ca. 550 nm. Cephalic capsule (Fig. 3C) apparent only in optical sagittal section of head; length of cephalic capsule on dorsal side of head 6 um in both speci- mens, and length on ventral side 4—5 (4.5 ae (0), 7/)) (Mao, Papilliform, inner labial sensilla not ob- served. Six outer labial and 4 cephalic se- tiform sensilla in single circle of ten (Fig. 3C), 13 to 15 (14 + 1.4) wm from oral sur- face of head. Outer labial sensilla 2-3 (2.7 + 0.5) um long; cephalic sensilla 3-5 (4.3 + 0.8) wm long. Cervical region (Fig. 4A) between amphid and nerve ring with 5 or 6 setiform sensilla on right side of body and with 3-5 (4 + 1.4) on left; length of cervical sensilla 2—4 (3 + 0.9) um. Amphid (Fig. 3C) cyathiform with pore-like aperture 17-21 (19 + 2.8) um from oral surface on right side of head, and 18-20 (19 + 1.4) um on left side. Amphidial flap absent; amphidial glands not evident. Buccal cavity and anterior end of esoph- agus as in males. Distance from oral surface of head to orifice of dorsal esophageal gland duct (Fig. 3C) 21-26 (23.5 + 3.5) um; to anterior edge of nerve ring 295-307 (301 + 8.5) um. Length of esophagus 851-943 (897 + 65) wm. Renette not observed. Somatic sensilla uncommon between level of nerve ring and near level of cloacal vent. Dorso- and ventrolateral orthometanemes present; some with caudal filaments, others without. Hypodermal gland cells absent. Gonoducts opposed and ovaries anti- dromus. Anterior and posterior gonads ven- tral to gut in both specimens; both ovaries on right side of oviduct in both specimens. Tail (Fig. 4B) bluntly conical, 103-120 (112 + 12) wm long. Anal body diameter 75-83 (79 + 5.7) um. Cuticle of tail terminus with median, crescent-shaped lamella penetrated a SA Bas =a WW ‘Xs Cc Fig. 3. Syringonomus dactylatus. A. Illustrated lateral view of the head of the holotype, USNM 77172; scale equals 10 um. B. Illustrated lateral view of the tail of the holotype, USNM 77172; lateral accessory component 1s Situated between 4 arrow heads at distal end of spiculum; scale equals 20 um. C. Illustrated lateral view of the head of the female paratype USNM 77175; scale equals 10 um. Abbreviations.—AA, amphidial aperture; AF, amphidial flap; AG, amphidial gland; AL, amphidial lobe; CC, cephalic capsule; CG, caudal gland; CL, crescent-shaped layer of cuticle; DD, duct of dorsal esophageal gland; DG, dorsal esophageal gland; RM, retro- dilator muscle; SG, subventral esophageal gland. Ds 726 te Z| CES. / PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Syringonomus dactylatus. A. Photomicrograph of the cervical region of the holotype, USNM PUN To unlabeled arrows point to amphidial gland; scale equals 100um. B. Photomicrograph of the posterior body region and tail of the paratype USNM 77175; scale equals 100 um. Abbreviations. — AV, anal vent; CG, caudal gland; NR, nerve ring. by spinneret. Cell bodies of each of 3 caudal glands (Fig. 4B) extend anterior beyond anal vent. Juvenile. — Body a b Cc length USNM 77176 52 6 49 3.984 mm Midbody diameter 76 um; body width at base of esophagus 68 um, at level of nerve ring 59 um, and at level of cephalic sensilla 29 um. Neck region uniformly tapered; head region not constricted nor cuticle thickened at level of amphid. Cuticle with fine trans- verse Striae; periodicity of striae ca. 500 nm. Cephalic capsule apparent on dorsal and ventral sides of head in optical sagittal sec- tion; length of cephalic capsule on dorsal side 4 um and on ventral side 5 um. Papilliform, inner labial sensilla not ob- served. Six outer labial and 4 cephalic se- tiform sensilla in single circle of ten, 15 wm from oral surface of head. Outer labial sen- silla 3 wm long; cephalic sensilla 4 to 5 (4.5 + 0.7) um long. Right cervical region be- tween amphid and nerve ring with 7 seti- form sensilla, and left with 6; cervical sen- sila 3 to 5 (4 + 1.4) wm long. Amphid cyathiform with pore-like aperture, 20 wm VOLUME 101, NUMBER 4 posterior from oral surface of head; am- phidial flap absent. Amphidial glands not observed. Buccal cavity and anterior end of esoph- agus as in adults. Distance from oral surface of head to orifice of dorsal esophageal gland duct 24 um; to anterior edge of nerve ring 235 um; to posterior end of esophagus 673 um. Renette not observed. Dorso- and ventrolateral orthometa- nemes present, with and without caudal fil- aments. Hypodermal gland cells absent. Tail bluntly conical, 82 wm long. Anal body di- ameter 63 um. Cuticle of tail terminus with median, crescent-shaped lamella penetrated by spinneret. Cell bodies of each of 3 caudal glands extend anterior beyond anal vent. Diagnosis.—The males of S. dactylatus may be distinguished from the males of the only other species of the genus, S. typicus, by the presence of a dorsal and a ventral lobe on each amphidial flap. The females of S. dactylatus have a body length of 5.156-5.791 (5.474 + 0.449) mm and the esophagus is 851-943 (897 + 65) um. The corresponding values for females of S. typicus are 3.320—4.940 (4.230 + 0.510) mm and 556-717 (664 + 45) um. Although these values are likely to overlap if obtained from larger populations, they are the only characters known at present by which the females may be distinguished from one another. Etymology. —The specific epithet is Greek, masculine gender, for finger, refer- ring to the finger-like processes of the am- phidial flap. Discussion.— As noted in the introduc- tion of this work, it is stated in the original description of S. typicus that the amphid has the shape of an inverted lyre, with or without a posteriorly directed central pro- cess. It is apparent from the present study that the inverted lyre corresponds to the outline of the paired lobes of the flap, and the posteriorly directed process corresponds to the space between the lobes when they 727 are separated. The “process” is obscure when the lobes are together. Thus, the description of an inverted lyre-shaped structure was a misinterpretation of what has been dem- onstrated in this study to be a bilobed am- phidial flap. The misinterpretation of the amphidial flap in the original description had made it difficult to relate the structure of the amphid to that of any other taxon, especially in Lep- tosomatidae. From the new interpretation, it is apparent that the amphidial flaps of the males of Syringonomus resemble those found in males of Platycoma and Proplaty- coma, (Platycominae: Leptosomatidae) in position, presence of lobes, and in that they occur in males only. In particular, the am- phidial flap in males of S. typicus, with its paired, anteriorly directed lobes, is similar to the amphidial flap in the males of Platy- coma cephalata Cobb, 1894. Also, the am- phidial flaps of males of S. dactylatus are similar to those of Proplatycoma sudafri- cana Inglis, 1966 and P. fleurdelis Hope, 1988 in having unbranched dorsal and ven- tral lobes in addition to those that are cen- tral. There are obvious differences as well. The ends of the dorsal and ventral lobes of the flaps of P. sudafricana and P. fleurdelis are directed posteriorly, whereas they are di- rected anteriorly in the case of S. dactylatus. Also, each flap in the males of each species of Proplatycoma has a single, anteriorly di- rected, central lobe, instead of the paired, central lobes that occur in the males of both species of Syringonomus and males of Platycoma cephalata. Another difference apparent from the SEM observations of S. typicus and P. fleurdelis is that the lobes of flaps in the former species have electron re- flective qualities similar to those of the sur- rounding cephalic and cervical cuticle. By contrast, the lobes in males of P. fleurdelis are more electron reflective than is the cu- ticle of the surrounding head and neck re- gions (Hope 1988). Finally, there is no evi- 728 dence that the amphidial flaps are framed by a fringe on the cephalic cuticle, as is the case in the males of both species of Syrin- gonomus. For obvious phylogenetic reasons, it is important to determine if the amphidial flap of Syringonomus is homologous with that of Platycoma and Proplatycoma. Its simi- larity among all three genera in position, general structure, and occurrence in males only is convincing evidence that they are homologous. The differences concerned with the number, position and shape of the lobes of the amphidial flaps within Proplatycoma, er between P. cephalata on the one hand and Proplatycoma on the other, are similar to the differences between either genus and Syringonomus. Therefore, differences of this nature are not regarded as evidence against homology. The fringe bordering the am- phidial flap in males of both species of Syr- ingonomus is a feature of the adjacent cu- ticle of the neck region, and so is not concerned with the question of the homol- ogy of the flap itself. Finally, because a dif- ference in electron reflective properties has been observed between males of S. typicus and males of P. fleurdelis only, it is at best a weak argument against homology. There- fore, it is concluded from the available evi- dence that the amphidial flap of Platycoma, Proplatycoma, and Syringonomus are ho- mologous. A lobed amphidial flap is not known to occur in any other taxon within the Order Enoplida, and so it is interpreted to be an apomorphic character within the family Leptosomatidae. By this character, it is as- sumed that Platycoma, Proplatycoma, and Syringonomus share a common ancestor not known to be shared by any other taxon. Males of Proplatycoma fleurdelis Hope, 1988, P. sudafricana Inglis, 1966, and both species of Syringonomus have prominent amphidial glands that extend to near the posterior end of the esophagus. According to Bongers (1983), the amphidial glands are enlarged in males of Leptosomatum, and it PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON may be that this is a unique (synapomor- phic) character state indicative of a mono- phyletic relationship among the genera that share this character. However, taxonomic changes are deferred until after a more ex- haustive morphological and phylogenetic analysis of Leptosomatidae. Distribution and ecology. —The localities from which specimens of S. typicus and S. dactylatus have been collected are limited to their respective type localities. Therefore, representatives of this genus are at this time known to inhabit only the western North and South Atlantic, although members of the former species were collected at tem- perate latitudes, whereas specimens of the latter were collected at tropical latitudes. This is in contrast to the much wider dis- tribution known for representatives of the genera Platycoma and Proplatycoma (Hope 1988). However, the apparently limited geographic distribution for the genus Sy- ringonomus in comparison to Platycoma and Proplatycoma is likely to be a function of the relatively inaccessible depths inhab- ited by representatives of the genus Syrin- gonomus. All known specimens of S. typicus have been collected at 3806 m, and S. dactylatus at 943-1007 m. This is in contrast to all nominal species of Proplatycoma whose known depth distributions do not extend below the intertidal zone, and the maxi- mum known depth for Platycoma cephalata Cobb, 1894, the only nominal species of the genus, is 44 m (Hope 1988). Therefore, from available data it appears that the species of Platycoma and Proplatycoma are limited to very shallow depths, whereas known species of Syringonomus are inhabitants of bathyal sediments. A review of the literature has revealed that, where data is available, specimens of Platycoma and Proplatycoma inhabit fine to coarse, sandy sediment (Hope 1988). There is no published data on the sediment from the sites at which S. typicus and S. dactylatus were collected. However, accord- VOLUME 101, NUMBER 4 ing to George Hampson of the Woods Hole Oceanographic Institution (personal com- munication), the sediment from station 167 of Atlantis ITs cruise 31 is silty mud. It is likely that the sample from which S. typicus was sorted also contained substantial amounts of mud given that it was collected at abyssal depths. However, the sediment from which the latter species was sorted also contained tests of the foraminiferan Rhab- damina abyssorum. These tests were con- structed of fine particles of sand, so sand also must have been present in this sample. Although definitive data is lacking, it ap- pears that specimens of Syringonomus in- habit sediments muddier than those inhab- ited by specimens of Platycoma and Proplatycoma. It is very possible that S. typ- icus does not in its natural environment in- habit tests of Rhabdamina abyssorum, but may have done so upon being sorted live from the sediment. Acknowledgments I thank Abbie Yorkoff for her technical assistance; Tom Bongers, Corinne Erhlich, Armand Maggenti, and Richard O’Grady for their valuable comments on the manu- 729 script; and the SEM laboratory for taking the scanning electron micrographs. I also thank George Hampson of the Woods Hole Oceanographic Institution for providing in- formation concerning the sediment collect- ed at station 167 of Atlantis ITs cruise 31. Literature Cited Bongers, T. 1983. Revision of the genus Leptoso- matum Bastian, 1865 (Nematoda: Leptosoma- tidae).--Proceedings of the Biological Society of Washington, 96(4):807-855. Hope, W. D. 1982. Structure of head and stoma in the marine nematode genus Deontostoma (En- oplida: Leptosomatidae).--Smithsonian Contri- butions to Zoology 353:1—22. . 1988. A review of the nematode genera Pla- tycoma and Proplatycoma, with a description of Proplatycoma fleurdelis, (Enoplida: Leptoso- matidae).— Proceedings of the Biological Soci- ety of Washington, 101(3):693-706. ——., & D.G. Murphy. 1969. Syringonomus typ- icus new genus, new species (Enoplida: Lepto- somatidae) a marine nematode inhabiting are- naceous tubes.— Proceedings of the Biological Society of Washington 82:511-518. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 730-740 NOTES ON ANTROSELATES HUBRICHT, 1963 AND ANTROBIA HUBRICHT, 1971 (GASTROPODA: HY DROBIIDAE) Robert Hershler and Leslie Hubricht Abstract. — Antroselates Hubricht, 1963 and Antrobia Hubricht, 1971, mono- typic cavesnail genera from eastern United States, are redescribed. The presence of a spermathecal duct in the female reproductive system of these snails in- dicates that, contrary to earlier opinions, they belong to the Hydrobiidae: Lit- toridininae. Both genera are well-differentiated members of the Cochliopina Morrison, 1946 group, characterized by possession of trochoidal shells and simple penes. The monotypic genera Antroselates and Antrobia are poorly known members of the North American aquatic cavesnail (Gas- tropoda: Hydrobiidae) fauna. Initial de- scriptions of these unusual-shelled snails lacked critical details from soft-part mor- phology and speculations on their affinities have varied considerably. Antroselates, for instance, was initially considered a hydro- biid (Hubricht 1963:138), but also has been placed in the Micromelaniidae (Taylor 1966: 171, Burch 1982:3). During the course of an ongoing review of North American cav- esnails, we examined what may be the only alcohol-preserved material available for these genera and determined that, contrary to all previous assertions, both are well-dif- ferentiated members of the Cochliopina group of Hydrobiidae: Littoridininae (Her- shler 1985). Redescriptions of Antroselates and Antrobia follow. Antroselates Hubricht, 1963 Antroselates Hubricht, 1963:138. Type species, Antroselates spiralis Hubricht, 1963, by monotypy and original desig- nation. Diagnosis.—Shell small, solid, trochoid- low conic, with 4.5 rounded whorls sculp- tured with numerous elevated spiral lines. Protoconch tilted or depressed, with punc- tate surface lined by spiral grooves. Aper- ture enlarged, with expanded outer lip. Um- bilicus narrowly to broadly open. Operculum paucispiral, with rapidly ex- panding whorls. Animal blind, unpigment- ed. Basal cusps of central radular teeth emerging from lateral angles. Cephalic ten- tacles without hypertrophied ciliary tufts. Ctenidium broad and elongate, with ca. 25 filaments. Intestine with coil in pallial roof. Penis simple, flagellate, non-glandular; sur- face of distal portion striated. Females ovip- arous. Spermathecal duct present, connect- ing posteriorly with albumen gland and anteriorly with capsule gland. Oviduct opening into ventral surface of albumen gland. Bursa copulatrix large, positioned partly posterior to albumen gland. Remarks. — Placement of this genus in the Micromelaniidae was based on the incorrect observation that the central radular teeth lacked basal cusps (Hubricht 1963:138). We assign the genus to the Hydrobiidae: Lit- toridininae on the basis of possession of a spermathecal duct in the female reproduc- tive system (see Davis et al. 1982, Hershler 1985). The globose shell and simple penis indicate assignment to the informal Coch- liopina group (Hershler 1985). Affinity with Antrobia is indicated by similarity of pro- toconch sculpture and duct arrangements in the pallial oviduct complex (including VOLUME 101, NUMBER 4 unique connection between spermathecal duct and albumen gland). These snails are, however, separable by differences in shell, penial and radular morphology. Antroselates spiralis Hubricht Figs. 1, 2, 3a—c, 4 Antroselates spiralis Hubricht, 1963:138- 139, pl. 8, figs. a, b; (Holotype FMNH 116916).—Burch 1982:108, fig. 108. Material examined.—Kentucky. Ed- mondson County: Echo River Spring, Mammoth Cave National Park, USNM 673526 (paratypes), USNM 673527 (para- types); River Styx, Mammoth Cave, Mam- moth Cave National Park, UF 40624 (al- cohol series [unrelaxed]). Description. —Shell (Fig. 1) 4.3-5.2 mm in height, slightly taller than wide. Whorls shouldered adapically, sutures slightly in- dented. Body whorl enlarged, spire rela- tively small. Shell white-clear, translucent, with amber periostracum. Surface typically eroded with small holes. Protoconch surface slightly pitted; spiral grooves (Fig. 1d) fairly broad and slightly depressed. Spiral lines of irregular height crossing strong growth lines on teleoconch (Fig. 1b, c). Aperture ellip- soidal, longer than wide, rounded above and somewhat angled below, and tilted relative to coiling axis, with anterior side advanced. Inner lip somewhat thickened and reflected below, broadly adnate to body whorl above. Outer lip thin. Operculum (Fig. 2a) thin, amber-colored, with pronounced growth lines. Animal of three whorls (Fig. 1b). Proxi- mal portions of tentacles without any ex- ternally visible vestiges of eyespots. Gen- eralized formulae of radular teeth (Fig. 4): central, 5(6)-1-5(6)/3-3; lateral, 4(5)-1-4(5); inner marginal, 15—18; outer marginal, |2— 16. Central teeth trapezoidal; basal cusps small, central cusps only slightly enlarged. Lateral teeth with enlarged, hoe-like central cusps (Fig. 4b). Stomach slightly longer than style sac. Posterior edge of stomach without 731 caecal chamber. Intestinal loop filling pos- terior half of pallial roof (Fig. 2b). Osphra- dium filling 25-33% of ctenidium length. Testis (Ts, Fig. 2b) massive, covering stomach and extending from posterior edge of prostate gland (overlapping most of stomach) almost to posterior end of animal. Branches of testis bearing numerous grape- like lobes and draining to narrow, thin vas efferens. Seminal vesicle consisting of a few thickened coils ventral to anterior testis. Prostate gland (Fig. 2d) thickened, near-cir- cular in outline; almost entire length posi- tioned in pallial roof. Vas deferens entering and exiting antero-ventral portion of gland; anterior portion of duct having thickened coil. Penis emerging from behind right ce- phalic tentacle, large relative to head (Fig. 2c), with a tight clockwise coil. Penis (Fig. 3a) unciliated, vermiform, with short distal- most section tapering gently. Vas deferens having thickened coils in base of penis; duct undulating in penis. Distal half of structure striated (Fig. 3a), with surface consisting of broad ridges alternating with narrow fur- rows (Fig. 3b). Terminus (Fig. 3c) thick- ened, smooth. Opening of vas deferens ter- minal, simple. Ovary a simple lobed mass filling about one half whorl behind posterior edge of stomach. Albumen and capsule glands (Ag, Cg, Fig. 2e) about equal in length; capsule gland bipartite (posterior section much longer). Oviduct (Ov) with single, thickened coil on left side of albumen gland. Kidney- shaped bursa copulatrix pressed against and slightly overlapping posterior edge of al- bumen gland. Bursa duct narrow (Dbu). Seminal receptacle (Sr) small, almost duct- less, pear-shaped, positioned on left side of albumen gland and slightly overlapping bursa. Oviduct without connection to al- bumen gland. Communication to latter achieved by short duct emerging from sper- mathecal duct (Dsd, Fig. 2f) slightly anterior to point where oviduct joins seminal recep- tacle duct. This unique arrangement implies that oocytes travel (anteriorly) a short dis- 732 Fig. 1. height, 4.95 mm); b, Dorsal aspect, showing well-developed spiral lines (shell height, 3.12 mm); c, Apical whorls (bar = 0.43 um); d, Protoconch (bar = 176 um). tance through the spermethecal duct before entering duct to albumen gland. Spermathe- cal duct (Sd) fairly wide, pressed against capsule gland. Spermathecal duct entering anterior capsule gland; genital aperture (Ga) simple, terminal. Distribution.— Found in a few caves and springs in vicinity of Mammoth Cave Na- tional Park, Kentucky, as well as in a cave stream in adjacent Indiana (Fig. 5; Hubricht 1963). Snails “found on the undersides of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SEM micrographs of shells of Antroselates spiralis Hubricht, USNM 673527 (paratypes): a, (Shell large stones in running water’ (Hubricht 1963:139). The species was not found by the senior author during a 1987 trip to the Mammoth Cave area. Antrobia Hubricht, 1971 Antrobia Hubricht, 1971:95. Type species, Antrobia culveri Hubricht 1971, by monotypy and original designation. Diagnosis.—Shell small, low conical, VOLUME 101, NUMBER 4 733 a Ts 1.0 mm 1.0 mm Epc Fig. 2. Morphology of Antroselates spiralis Hubricht, UF 40624: a, Dorsal view of operculum; b, Dorsal view of male (head—foot removed); c, Dorsal view of head and penis; d, Right lateral view of prostate gland; e, Left lateral view of pallial oviduct complex; f, Left lateral view of bursa copulatrix, seminal receptacle (rotated to expose underlying structures), and associated ducts. Only the anterior portion of the albumen gland is shown. Ag = albumen gland; Avd = anterior vas deferens; Bu = bursa copulatrix; Cg = capsule gland; Dbu = duct from bursa copulatrix; Dg = digestive gland; Dov = duct connecting oviduct and spermathecal duct; Epc = posterior end of pallial cavity; Ga = genital aperture; In = intestine; Jnc = juncture between anterior vas deferens and “neck”; Ov = oviduct; Pr = prostate gland; Pvd = posterior vas deferens; Sd = spermathecal duct; Sr = seminal receptacle; Sts = style sac; Ts = testis. 734 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. SEM micrographs of critical point dried penes of Antroselates spiralis Hubricht, UF 40624 (a—c) and Antrobia culveri Hubricht, LH coll. (d): a, Dorsal view of penis (bar = 37 wm); b, Close-up of striated surface (bar = 213 wm); c, Close-up of penial tip, showing simple terminal opening (bar = 25 um); d, Dorsal view of penis showing irregular ciliation (bar = 136 um). VOLUME 101, NUMBER 4 broadly umbilicate, with 3.5 well-rounded whorls having indented sutures. Protoconch and teleoconch lined with regularly spaced shallow spiral grooves. Aperture simple, sub- circular. Operculum multispiral, with slow- ly expanding whorls. Animal blind, unpig- 735 fi | 7) i) Fig. 4. Radula of Antroselates spiralis Hubricht, UF 40624: a, Centrals (bar = 17.6 um); b, Laterals (bar = 12 wm); c, Inner marginals (bar = 13.6 wm); d, Outer marginals (bar = 15 um). mented. Cusps numerous on all radular teeth; central teeth broadly trapezoidal with elongate lateral angles. Cephalic tentacles without hypertrophied ciliary tufts. Ctenid- ium reduced, consisting of ca. eight small filaments. Penis simple, non-glandular, dag- 736 Fig. 5. Antrobia culveri Hubricht. ger-like. Females oviparous. Spermathecal duct present, connecting posteriorly with al- bumen gland and anteriorly with capsule gland. Bursa copulatrix positioned partly posterior to albumen gland. Remarks.—Burch (1982:3) placed this genus in the Lithoglyphinae, presumably based on its low-conical “‘lithoglyphine-like”’ shell. Given the presence of a spermathecal duct in the female reproductive system (ab- sent from lithoglyphines; Thompson 1984), these snails are more appropriately placed in the Hydrobiidae: Littoridininae. Antro- bia is distinguished from other members of the Cochliopina group by its radular mor- phology (similar to that of Fontigens Pils- bry, 1933) and teleoconch sculpture. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 4 Antrobia culveri e Antroselates spiralis Drainage map of eastern United States showing distributions of Antroselates spiralis Hubricht and Antrobia culveri Hubricht Figs. 3d, 6-8 Materials examined.—Missouri. Taney County: stream in Tumbling Creek Cave, 4.5 mi NE of Protem, FMNH 164171 (Ho- lotype), FMNH 164170 (Paratypes); Leslie Hubricht coll. 38780 (alcohol series [re- laxed]). Description. —Shell (Fig. 6) 1.9—2.2 mm high, very slightly taller than wide. Sutures indented. Shell white-clear, translucent, with amber periostracum. Surface typically part- ly covered by white deposits. Protoconch (Fig. 6d, e) with 1.0-1.25 whorls, having a slightly pitted surface with well-spaced, spi- ral grooves. Grooves sometimes ending af- VOLUME 101, NUMBER 4 V3 Fig. 6. SEM micrographs of shells of Antrobia culveri Hubricht, FMNH 164170 (paratypes): a, b (Shell heights, 1.9, 2.0 mm, respectively); c, Close-up showing spiral grooves (bar = 0.43 um); d, e, Views of apex (bars = 176 um). ter first teleoconch whorl (erosion?). Teleo- conch roughened with collabral growth lines. Aperture longer than wide, near-planar, ad- nate to a small portion of the body whorl above, and slightly tilted relative to coiling axis (adapical side advanced; Fig. 6b). Inner lip reflected, somewhat thickened; outer lip thin. Operculum (Fig. 7a) thin, amber. Animal of two whorls. Proximal portion of tentacles without any externally obvious eyespot vestiges. Generalized formula of radular teeth (Fig. 8): central, 6(7)-1-6(7)/ 3-3; lateral, 16-17; inner marginal, 29; outer marginal, 21. Cusp morphology very sim- ilar for all tooth types: note absence of en- larged central cusps on centrals and laterals. Basal portion of central teeth well excavated between lateral angles and basal process. Stomach considerably longer than style sac. Posterior stomach edge without caecal chamber. Intestinal loop undulating gently in pallial roof. Osphradium about 33% of ctenidium length. Testis a single, unlobed mass, filling one- halt whorl of animal and partly overlapping the posterior stomach. No vas efferens not- 738 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.5 mm Ag Epc Fig. 7. Morphology of Antrobia culveri Hubricht, LH coll.: a, Dorsal view of operculum; b, Dorsal view of penis showing course of vas deferens (dotted line); c, Left lateral view of pallial oviduct complex; d, Left lateral view of bursa copulatrix, seminal receptacle (rotated to expose underlying structures), and associated ducts. Ag = albumen gland; Bu = bursa copulatrix; Cg = capsule gland; Dbu = duct from bursa copulatrix; Dov = duct between oviduct and spermathecal duct; Epc = posterior end of pallial cavity; Ga = genital aperture; Ov = oviduct; Sd = spermathecal duct; Sr = seminal receptacle. ed; seminal vesicle consisting of a few thick- ened coils entering anterior edge of testis. Prostate gland small, with 50% of length in pallial roof. Vas deferens entering and ex- iting central area of prostate. Penis (Fig. 7b) small, with tight clockwise coil, well-ta- pered. Surface irregularly ciliated (Fig. 3d), with cover particularly heavy distally. Dis- tal end of penis undifferentiated; terminal pore simple. Vas deferens undulating slight- ly within penis. Ovary an unlobed mass filling a relatively small portion of digestive gland posterior to the stomach. Albumen gland slightly longer than capsule gland (Ag, Cg, Fig. 7c). Capsule gland bipartite with posterior section much larger. Oviduct (Ov) with single coil on left side of albumen gland. Bursa (Bu) sac-like, with a narrow duct (Dbu, Fig. 7d). Seminal receptacle (Sr) small, positioned partly an- terior to bursa. Oviduct without connection to albumen gland. Communication to latter achieved by short duct emerging from sper- mathecal duct (Dsd, Fig. 2f) anterior to point where oviduct joins seminal receptacle duct. Spermathecal duct (Sd) moderately wide, pressed against capsule gland and entering gland near anterior terminus. Genital ap- erture (Ga) simple, terminal. Distribution.—Endemic to underground VOLUME 101, NUMBER 4 739 Fig. 8. Radula of Antrobia culveri Hubricht, LH coll.: a, Centrals (bar = 7.5 um); b, Laterals (bar = 7.5 um); c, Inner marginals (bar = 6 um); d, Outer marginals (bar = 3.8 um). stream of Tumbling Creek Cave in SE Mis- Acknowledgments souri (Fig. 5). Snails are currently uncom- We thank Drs. A. Solem (FMNH) and F. mon in the cave (pers. comm. 1986, T. J. G. Thompson (UF) for loan of specimens Aley, Ozark Underground Laboratory). under their care. The National Park Service 740 (Mammoth Cave National Park) provided a collecting permit and field assistance dur- ing Hershler’s visit to Mammoth Cave. Molly Kelly Ryan and Cathy Flamer pre- pared the map, and Paul Greenhall provid- ed other assistance with plate preparations. An anonymous reviewer provided useful criticism of the manuscript. Literature Cited Burch, J. B. 1982. Freshwater snails (Mollusca: Gas- tropoda) of North America.—U.S. Environ- mental Protection Agency, Contract No. 68-03- 1280, EPA-600/3-82-026, 294 pp. Davis, G. M., M. Mazurkiewicz, & M. Mandracchia. 1982. Spurwinkia: morphology, systematics, and ecology of a new genus of North American marshland Hydrobiidae (Mollusca: Hydrobi- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON idae).— Proceedings of the Academy of Natural Sciences of Philadelphia 134:143-177. Hershler, R. 1985. Systematic revision of the Hy- drobiidae (Gastropoda: Rissoacea) of the Cua- tro Cienegas Basin, Coahuila, Mexico. — Mala- cologia 26:31-123. Hubricht, L. 1963. New species of Hydrobiidae.— Nautilus 76:138-140. . 1971. New Hydrobiidae from Ozark caves. — Nautilus 84:93-96. Taylor, D. W. 1966. A remarkable snail fauna from Coahuila, Mexico. — Veliger 9:152—228. Thompson, F.G. 1984. North American freshwater snail genera of the hydrobiid subfamily Litho- glyphinae. — Malacologia 25:109-141. (RH) NHB STOP 118, Department of In- vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (LH) 4026 35th Street, Meridian, Mississippi 39301. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 741-766 A REVISION OF THE GENUS THEMISTE (SIPUNCULA) Edward B. Cutler and Norma J. Cutler Abstract.—The 31 putative species of the sipunculan genus Themiste and the morphological characters used to differentiate them are critically reviewed. The monograph of Stephen & Edmonds (1972) is used as a starting point and all changes made in the intervening years are reiterated. All available type material was studied and new collections of Hawaiian and Californian material are used to analyze within-deme variation. Four characters are determined to be useful at the species level and three at the subgeneric level. A key to and a discussion of each of the 10 remaining species (plus one reduced to subspecies) with the newly designated synonyms are presented. A brief statement of the known distribution and an overall summary of the zoogeography of the two subgenera (largely non-overlapping) are given. This is a continuation of our series in which we reevaluate the species of sipun- culan worms (e.g., Cutler & Cutler 1985a, b, 1986, 1987). With this work we complete our survey of the genera in the Class Si- punculida. The monograph of Stephen & Edmonds (1972) is the starting place for this work (25 species names). In that same work there were three names listed as species in- quirendae or incertae sedis that subsequent- ly were considered members of this genus and are addressed herein. The one species and the subgenera erected and the two species transferred into this genus since that time are also included (Table 1). The genus Themiste was erected by Gray in 1828, but this name dropped from sight for over a century. The name Dendrosto- mum (Grube & Oersted, 1858) took its place until 1964 when Stephen resurrected Gray’s name. The genus has been placed in its own family, Themistidae of the order Golfingi- iformes by Cutler & Gibbs (1985). While no formal subgenera exist in Ste- phen & Edmonds (1972), there were six groups that Edmonds later (1980) converted into three subgenera. These subgenera were considered in Gibbs & Cutler (1987), and the one with four retractors (Stephensonum) was determined to be a junior synonym of the genus name Golfingia. One of its two species (Themiste pinnifolia Keferstein, 1865) was regarded as nomen dubium and the other (7. stephensoni Stephen, 1942) was a mixture of Golfingia capensis and T. la- geniformis (see below). The remaining two subgenera (based on differences in contrac- tile vessel villi or tubules) are used here. We do question the presumed homologous na- ture of these tubular extensions. They are very different in size, distribution, and number. It seems likely that these elabo- rations resulted from two different evolu- tionary events (homoplasy). The type (10 mm trunk), and only spec- imen of Themiste spinifera, Sluiter, 1902 is not in the museum collections (ZMUA) with the other Siboga material. Certain aspects of the figures and description make us doubt the generic affinity, and, with no way to ver- ify this, we reduce its status to that of in- certae sedis. Whenever possible we have obtained type material to verify the original descriptions. In several cases we have made detailed ob- servations on series of recently collected in- dividuals to better evaluate the traditionally used morphological characters. Collecting 742 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Species considered and proposed taxonomic changes. Present name Proposed name Subgenus Themiste Themiste alutacea (Grube & Oersted, 1858) Themiste blanda (Selenka et al, 1883) Themiste dyscrita (Fisher, 1952) Themiste hennahi Gray, 1828 Themiste hexadactyla (Sato, 1930) Themiste lissa (Fisher, 1952) Themiste orbiniensis (de Quatrefages, 1865) Themiste perimeces (Fisher, 1928) Themiste petricola (Amor, 1964) Themiste pyroides (Chamberlin, 1920) Dendrostomum ramosum Quatrefages, 1865 Sipunculus rapa Quatrefages, 1865 Themiste rosacea (Amor, 1964) Themiste schmitti (Fisher, 1952) Themiste spinifera (Sluiter, 1902) Themiste zostericola (Chamberlin, 1920) no change no change no change no change Themiste pyroides Themiste hennahi Themiste alutacea Themiste hennahi Themiste alutacea no change Themiste hennahi Thimiste hennahi Themiste alutacea Themiste hennahi incertae sedis Themiste hennahi Subgenus Lagenopsis Themiste cymodoceae (Edmonds, 1956) Themiste dehamata (Kesteven, 1903) Themiste elliptica (Sato, 1934) Themiste fisheri (Amor, 1964) Themiste fusca (Edmonds, 1960) Themiste glauca (Lanchester, 1905) Themiste huttoni (Benham, 1904) Themiste lageniformis (Baird, 1868) Themiste minor (Ikeda, 1904) Themiste pyriformis (Lanchester, 1905) Themiste robertsoni (Stephen & Robertson, 1952) Themiste tropica (Sato, 1935) Themiste variospinosa Edmonds, 1980 no change no change Themiste dehamata Themiste dehamata Themiste minor minor Themiste lageniformis Themiste minor huttoni no change Themiste minor minor Themiste lageniformis Themiste lageniformis Themiste lageniformis no change Subgenus stephensonum (now void) Themiste pinnifolia (Keferstein, 1865) Themiste stephensoni (Stephen, 1942) trips to Hawaii and California have greatly facilitated this effort. The opportunity to see the differences in the habitat of these ani- mals and to observe living material is in- valuable. We first discuss the morphological char- acters in light of our recent analyses. Next a key to all the species we consider valid and a section where each of these species is discussed; this includes a synonymy, a dis- cussion of newly added junior synonyms, and asummary of their distribution. Within nomen dubium Golfingia capensis each subgenus the species are grouped ac- cording to whether or not hooks are present. A short zoogeographical summary is also presented. The following abbreviations are used in this text for the museums from which we borrowed material: Bernice P. Bishop Mu- seum, Homolulu (BPBM); British Museum (Natural History) (BMNH); Museum of Comparative Zoology, Harvard (MCZH); Museum fur Naturkunde der Humboldt- Universitat zu Berlin (MNHU); Royal Scot- VOLUME 101, NUMBER 4 tish Museum, Edinburgh (RSME); Santa Barbara Museum of Natural History (SBNH); University Zoological Museum, Cambridge (UZMC); National Museum of Natural History, Washington, D.C. (USNM); Zoologisk Museum, Copenhagen (UZMK); Zoological Institute, Tohoku University, Sendai (ZITU); Zoological Mu- seum, Hamburg (ZMUH); Zoology Mu- seum, University of Tokyo (ZMUT). Morphological Characters Historically the following characters have been used to differentiate Themiste species. Earlier descriptions were not always based on a good understanding of possible vari- ation within a population (species). In this section we evaluate these characters in light of our experience with large sample sizes. It must be emphasized that if the specimens are not relaxed prior to fixation the mea- surements of diverse parts can be mislead- ingly different. For instance, in the Hawai- ian 7. lageniformis population we had access to one collection with completely retracted introverts and another which was preserved with introverts extended. The length of the introverts did vary considerably (Fig. 1), but this is partly an artifact of preservation tech- nique. Introverts of retracted specimens were measured by dissecting the body wall and determining the distance from the ne- phridiopores to the base of the tentacles. We have observed other differences that are not of the classical type but worth men- tioning. The niche concept is particularly applicable in the Californian species; we found one (7. hennahi) lives in unconsoli- dated sediments (various mixtures of silt/ sand/gravel), and a second (7. dyscrita) lives in discrete holes made by other animals in shales or sandstones. These also have dif- ferent sporozoan parasites in their intestine (F. Hochberg, pers. comm.) and respond very differently to anesthesia (7. hennahi is much more likely to extend its introvert in response to refrigeration or menthol). The 743 70 e @ - RELAXED e O - CONTRACTED INTROVERT LENGTH AS % OF TRUNK TRUNK LENGTH (in mm) Fig. 1. Introvert length of 7. /ageniformis to show how such measurements can differ between relaxed and contracted worms. As in other genera, the relative length of this body part becomes shorter with age (in linear regression b = —1.2 and — 0.27). third species (7. pyroides) lives in fissures/ crevices in granitic rock, is much more eu- rybathyal, and does not relax easily. A sim- ilar case (ill-defined but different niches) seems to exist for the Australian complex (S. J. Edmonds, pers. comm.). Karyology may be an attribute that bi- ologists can use to differentiate species. A more detailed statement will be published separately, but, based on a preliminary analysis of three species, it appears that the members of this genus have ten pairs of chromosomes. The apparent similarities preclude using this as a diagnostic character at the species level. Information about de- velopmental pathways and parasites might be useful, but at this time the data base is far from complete. 1. Contractile vessel villi. —In sipunculans a closed, fluid filled, tubular system extends into the tentacles from one or two vessels running along the esophagus functioning as a circulatory system helping in the exchange of gases between the external and internal 744 = 4 ‘a ! NM t WN N01 22. SQK Fig. 2. Contractile vessel villi from a 5 mm T. /a- geniformis showing bifurcations in a small specimen and the increasing frequency of bifurcations toward the posterior end (bottom of figure). Scale line is 1 mm. environments. It may also be a turgor pres- sure system not unlike the polian vessels in echinoderms aiding in the expansion of the tentacular crown. In five genera, digitiform villi along this vessel increase the volume and the surface area for internal gas ex- change. In this genus are two rather different configurations, and this character is used to separate the two subgenera (Edmonds 1980). The more common pattern (found in other genera) of many (more than 100) short villi is found in T. (Lagenopsis). In T. (Themiste) a unique design exists: few (usually 8-14) long threadlike extensions off the posterior quarter of the vessel, often with a corkscrew or beads-on-a-string shape. A question that has been raised about the shorter villi is whether or not the individual units bifurcate; a few species have been al- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON leged to lack any bifurcating villi. Our ex- amination of recently collected Hawaiian 7. lageniformis, ranging in size from 2-33 mm, shows that while more bifurcation exists in larger worms, all those over 3 mm show at least some evidence of bifurcation (Fig. 2). One must look carefully in smaller worms. In addition to an ontogenetic change, we have observed an anterior/posterior gradi- ent. The branching complexity (and size of villi) increase towards the posterior end— sometimes more than three terminal branches can be seen. In the subgenus 7. (Themiste) none of the tubular extensions (villi seems inappro- priate here) bifurcates in the same manner. In some larger specimens a complex branching occurs near the base (see Fisher 1952 pl. 30, fig. 3). The variable, occasion- ally used in species descriptions, is the num- ber, e.g., more than 10 vs. 10 or less. The exact number of tubules is difficult to de- termine, and our data suggest that smaller worms (under 30 mm) have less than 10 while larger worms (same population) usu- ally have more than 10 tubules. To use this difference as a species specific character is unwise. Only a few references to anastomosing ““networks”’ of the vessel around the esoph- agus exist, but our examination shows this feature to be age dependent, thus rarely de- veloped in smaller worms (see 7. pyroides section for more detail). Therefore, its use as a taxonomic character is limited. 2. Introvert hooks.—In both subgenera some species carry dark, horny, chitinoid hooks scattered on the introvert. In some genera hooks appear to be deciduous, but that does not seem to be the case here. How- ever, we know very little about the ontogeny of those species lacking hooks as adults. The one such species we do have information on is 7. lageniformis. Awati & Pradhan (1935) reported hooks on early stages and Pilger (1987) reported hooks in a five day specimen. Also, Williams (1977:38) ob- served that 13 day old Hawaiian specimens VOLUME 101, NUMBER 4 (about 0.5 mm) have “‘.. . several rows of hooks.”’ We saw no hooks on our 2-3 mm worms from Hawaii. One record suggests the reverse pattern exists in one species that has hooks as adults: Fisher’s (1952) discussion of 7. pyroides in- cludes the observation that smaller worms have fewer hooks (no hooks in those less than 18 mm, 4-18 hooks in two 18 mm worms, 50 hooks in a 24 mm worm). One must remember that Fisher’s measurements include both the trunk and the introvert, the later being about ' the total. Our recent collections from the same area (Carmel and Monterey) included only a few small worms and all of these have hooks (30-80 hooks in worms with a total length of 15—21 mm; trunks from 10-14 mm). In the 7. blanda we collected in Japan only one with a 2 mm trunk has no hooks; those 3 mm and larger carried them. These data suggest that all Themiste species, as early juveniles, have hooks which are soon lost. While some species never re- place these, other species do so in later stages of ontogeny. This hypothesis needs more testing, but Rice (1967) did observe one row of hooks in one month old specimens of 7. pyroides, the same species Fisher was refer- ring to above. She has also recorded (Rice 1975b) the presence of a row of hooks at one month in 7. alutacea. Nevertheless, a presence/absence dichotomy in adult worms exists and therefore, this character can be helpful to the systematist. If these early ju- venile hooks are real and are arranged in rings, we may have reason to reverse our earlier assumptions about the polarity of this character in our cladistic analyses, 1.e., this may be the plesiomorphic not the apo- morphic state. The adult hooks are often large, sparse, and arrayed over a wide band on the prox- imal part ('‘4-%) of the introvert (Fig. 3). Descriptions sometimes include comments on their number, size, and arrangement. As shown above, the number changes with trunk size. We originally thought that the 745 amount of introvert covered by hooks (most animals have less than 45% covered, but in others the hooked region seems to cover more than 50% of the introvert) as geneti- cally determined. Observations from our field work in California strongly suggest that those animals living in areas of high energy (open rocky intertidal) have fewer hooks covering a smaller area with many more broken hooks than other members of the same species at more protected or subtidal depths (e.g., 7. pyroides). It now seems clear that, within limits, this trait 's partly an en- vironmentally determined feature. The ar- rangement is not constant within a species or species specific, generally being scattered and oriented posteriorly (one exception is T. variospinosa). Hook size also changes with trunk size: In the Japanese 7. blanda pop- ulation we analyzed, a 5 mm worm had hooks about 50 um tall, an 8 mm worm had 75 um hooks, and an 11 mm worm had hooks about 100 um tall. The upper limit in this genus is about 450 um in large adults. 3. Tentacular crown. — The diverse inter- pretations of this feature have been one of the most troublesome aspects of Themiste. The dendritic branching pattern is unique within Sipuncula and makes the identifi- cation of these worms to the generic level easy (if the tentacles are extended). How- ever, the question of how many tentacular ““stems”’ exist has led to some unfortunate confusion. Perhaps the classic (but not only) case is the holotype of 7. hennahi. Gray (1828) saw five main stems; Stephen (1964) reported only four main branches; but, when Rice and Stephen (1970) reexamined the same worm, they reported six tentacular stems. The lack of consistency in language de- scribing these tentacles has compounded the confusion. We are defining the terms as fol- lows: The four structures arising from the oral disc are the stems; these stems divide into branches (primary, secondary, etc.); the final subdivisions or terminal units are ten- tacules. Finally, one tentacle is the entire 746 Fig. 3. array of subunits beginning with one stem. While this is somewhat arbitrary, it is con- sistent with several earlier authors (Fisher 1952, Edmonds 1980). The basic plan in this genus is four stems, but most of the problems arise when the first dichotomy appears close to the base of a stem. During the ontogeny of these worms different species exhibit different patterns vis-a-vis the timing of the “splitting.”’ Two adult patterns result—either 4 or 6 (occa- sionally 5) putative stems. Those animals with 5 or 6 may have a developmental path- way that involves an early post-larval sub- division of the dorsal pair. At times the dor- sal stems will be longer/larger than the ventrals. Our work with the Hawaiian and Californian populations supports the hy- pothesis that as these worms increase in size a corresponding increase in the complexity of the tentacular crown occurs, more branches and more tentacules (Fig. 4). Therefore, the number of apparent tentac- ular stems varies and should not be consid- ered species specific. A separate issue relates to the branching pattern that has been asserted to be either dendritic, pinnate, or palmate by different authors, sometimes for the same species. These adjectives have been used with min- imal precision and suggest differences that PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Introvert hooks showing different sizes and orientation among species: A. 7. minor minor; B. T. pyroides; C. T. variospinosa. Scale lines all equal 200 um. do not exist. In the subgenus Lagenopsis the four stems divide into smaller branches rel- atively few times (1-3), depending on their age/size (Fig. 4). On all of these branches as well as the main stems, terminal tentacules are given off along the entire length from near the branch points to the tip. The length of these tentacules exhibits some consisten- cy within a species. We measured the lon- gest ones in T. lageniformis, 2—4 mm; short- er in 7. cymodoceae, 1-2 mm; and shortest in 7. dehamata, 0.5-1 mm). The trunk lengths of the measured specimens, in order, are 10-30 mm, 79-85 mm, and 45-68 mm. In the subgenus 7hemiste each stem branches dichotomously several times with discrete internodes. In small individuals (less than 15 mm) the dendritic crown exists but with fewer branches and shorter internodes (see Fig. 4 and Fisher 1952 for more detail). The terminal tentacules are present as clus- ters near the tips of the terminal branches. Tentacules are widely spaced (what Fisher (1952) illustrated and called processes) along the branches. In small worms the relative size of the parts gives the crown a more Lagenopsis-like appearance. In general, the complexities and variations in the branch- ing patterns mitigate against using it for taxonomic purposes at the species level. The exception (helpful in particular species com- VOLUME 101, NUMBER 4 747 D Fig. 4. Left dorsal tentacle from two Themiste species showing size-related changes in branching pattern in the two subgenera: A-C are T. (Themiste) hennahi trunk lengths are 25, 80 and 140 mm. D-F are T. (Lagenopsis) lageniformis; trunk lengths are 6, 16 and 30 mm. Scale line is 1 mm. plexes), is unequal length of dorsal vs. ven- other genera the relative anterior/posterior tral stems and whether or not the ventral relationship of these openings has been re- stems split at the same time as dorsals. ported and sometimes used to differentiate 4. Anus-nephridia relationship.—As in species. The language is often general; these 748 NEPHRIDIOPORES (% OF TRUNK LENGTH POSTERIOR TO ANUS) !0 30 50 70 90 TRUNK LENGTH (in mm) Fig.5. Distance nephridiopores are posterior to anus as percentage of trunk length. Circles are T. lagenifor- mis, squares are T. dyscrita, triangles are T. hennahi. This distance decreases with size but with much over- lap among species. are at the same level, nearly the same level, or the nephridiopores open posterior to the anus. In our study of the Hawaiian 7. /a- geniformis population, the nephridia are al- most always posterior to the anus, com- monly at a distance equal to 3—7% of the trunk length, occasionally up to 12-15%, but a few at the same level (Fig. 5). In one Californian population of 7. hennahi this distance is usually only 0-1% but in a few small worms it is 5%. In a population of 7. dyscrita this distance is 1—4%. Fisher (1952) weighted this heavily in the California com- plex to distinguish his 7. /issa from its neighbors, but our measurements of Fish- er’s material show that only in the holotype is this distance 10% while in four of the paratypes this value ranged from 4—6%. Our conclusion is that since this distance is small, not easy to measure precisely, and, most importantly, since there can be much variation within demes, it has very little taxonomic significance and should not be used. 5. Trunk size and shape. —While this kind of information is often reported imprecisely (large vs. small or slender vs. stout), it may have some meaning. Based on our obser- vations there do appear to be two size classes for adult worms. Those that generally in- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON habit soft substrata commonly have trunks exceeding 60 mm (some over 200 mm) while those living in rock or coral rarely exceed 30 mm. However, the north Pacific 7. py- roides may reach 150 mm. Most of the smaller worms have a similar pyriform shape. Shape of preserved material is partly dependent on the animal’s unique set of cir- cumstances prior to fixation (physical mi- crohabitat and anesthesia, if any). However, worms living in unconsolidated sediments are cylindrical and elongate while those from rock/fissures/coral have stouter and more pyriform bodies. When these occupy holes made by boring bivalves the body is shaped by the mold chosen by the juvenile worm (Fig. 6). This shape is difficult to quantify and therefore difficult to communicate without ambiguity. Our observations on living, freshly-collected material lead us to conclude that trunk shape is partly envi- ronmentally determined. Nevertheless, this character (especially as an indicator of niche) can be used by the taxonomist. 6. Fixing muscle attachments.—Small threadlike muscles connect some part of the digestive tract to the body wall. The general pattern in Themiste is three muscles labeled Fl, F2, F3. In the subgenus 7Themiste, F1 anchors the esophagus in the area of the long contractile vessel villi to the body wall of the mid-trunk, while in Lagenopsis it an- chors the esophagus near the posterior part of the trunk where the esophagus turns for- ward and may take the form of several threads or a membrane. F2 is generally on the posterior esophagus or first gut coil while F3 is more often on the last intestinal coil or the rectum. The precise insertion of these muscles has been asserted to be constant and taxonomically meaningful (Fisher 1952 compares T. hexadactyla to T. pyroides or T. dyscrita to T. zostericola). However, Fisher did illustrate variations and Foster (1974:856) working with several hundred 7. dyscrita from one deme commented on the novel and abnormal arrangement of the F2, other than that described by Fisher. Our VOLUME 101, NUMBER 4 749 Fig. 6. Comparison of fully contracted (A, C, and E) with extended (B, D, and F) body shape in three pairs of living worms from different Californian microhabitats: A-B. 7. dyscrita from boring mollusc (pholid) holes in shale at Santa Barbara; C-D. Same species from burrows in sandstone at Santa Cruz; E-F. 7. hennahi from coarse sand at Santa Barbara. 750 RETRACTOR ORIGINS AS % OF TRUNK LENGTH TRUNK LENGTH (in mm) Fig. 7. Introvert retractor muscle origins on body wall as percentage of trunk length towards the posterior end in three species. Circles are T. lageniformis, squares are T. dyscrita, triangles are T. hennahi. This is not a size related character (all values for b (linear regression) are less than 0.1). Own analysis shows variation within pop- ulations (some lacking the full complement or the insertion of F2 being at different points and sometimes branched with several points of attachment). Therefore, as this variation also overlaps among species, it can only be used in a general manner. 7. Retractor muscle origins. —In adults one pair of muscles functions to retract the in- trovert into the trunk. These muscles have their origins on the body wall on either side of the ventral nerve cord and insert in the cephalic region. The anterior/posterior lo- cation of these origins has sometimes been used by systematists to describe a species using phrases like “in the middle third”’ or “‘in the last quarter.’ One must also deter- mine whether the author is talking about the whole body (including the introvert) or just the trunk when giving measurements. In order to standardize these data we record it as a percentage of the trunk length using the nephridiopores to mark the anterior border of the trunk. Our analysis shows that the vast majority of individuals have their retractor origins from 60-80% of the dis- tance towards the posterior end. In two Cal- ifornia populations that we examined (7. dyscrita and T. hennahi), there is a smaller PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 35 i) a NUMBER OF INTESTINAL COILS a 10 30 50 70 90 TRUNK LENGTH (in mm) Fig. 8. Number of intestinal coils showing varying increases in number with increase in size for the same species as Fig. 7. The positive slopes (b in the linear regression) are 0.5, 0.16 and 0.09. size related shift in the point of origin than that noted in other genera (Fig. 7). As this figure shows 7. dyscrita is generally more anterior than 7. hennahi, which is more an- terior than 7. Jageniformis. The linear regression is a very poor fit because of much overlap, especially in smaller worms, and so much scatter of the data points. 8. Number of intestinal coils. — As in other genera the number of coils is directly cor- related with the size of the worm: com- monly 12 to 24 coils (Fig. 8). One apparent exception to this is: some larger individuals (over 200 mm) in the California population of T. hennahi may have more than 80 coils. Four species seem to have irregular, loosely wound gut coils; however, since we find much within-deme variation and overlap among species, this is not a useful character. 9. Pigmentation on tentacles and intro- vert. —Some species have dark bluish pig- ment in a collar-like band around the in- trovert and others have dark purple/blue pigmented patches on the tentacles (Fig. 9). The presence or absence of the dark collar on the introvert is consistent. Pigment on the branches is inconsistent, but its presence or absence on the tentacules of certain species is reliable and helpful to the system- atist. 10. Papillae size and shape.—As with VOLUME 101, NUMBER 4 Fig. 9. Distal introverts of two species; T. dyscrita on left with pigmented collar region and pale tentacules; T. hennahi on the right with unpigmented collar and dark tentacules. other genera, these worms have glandular secretory papillae on the introvert and trunk. These are more numerous and larger near both ends of the trunk. While some authors have used the size and/or shape of these bodies to distinguish species we do not con- cur. The morphology of these papillae is simply too variable to be useful. 11. Spindle muscle.—As in other sipun- culans, a thread-like muscle runs through the center of the gut coil, connected along its length to help anchor the intestinal coils. It originates anteriorly in all genera but does not insert on the body wall posteriorly in this genus. Some authors have described it as terminating on the rectum in certain species. After carefully analyzing this char- acter, we conclude that all Themiste species are similar: after originating from the body wall anterior to the anus it appears to be- come part of the rectal wall, is difficult to trace but can be seen posteriorly attached to the caecum before entering the gut coil. Summary. — Of these characters, the type of contractile vessel villi, branching pattern of the tentacular crown, and the origin and insertion of Fl are useful at the subgeneric level only. The presence or absence of hooks, trunk size and shape, pigmented collar on the introvert, and tentacule pigmentation can be useful at the species level. The re- maining characters exhibit too much with- in-deme variation or among species overlap to have value for the systematist. Key to Themiste Species 1. Contractile vessel with numerous (more than 40) short digitiform villi (subgenus Lagenopsis) ........... 2 — Contractile vessel with few (less than 20) long thread-like tubular exten- sions (subgenus Themiste) ....... 6 2. Introvert without hooks ......... 3 — Introvert bearing dark scattered HOOKS yoanae aes wens See eee 5 3. Small (trunk <35 mm); introvert with purple “collar”; lives in hard SUDSiTates: eee ee T. lageniformis V2 — Large (trunk commonly >40 mm); unpigmented introvert “‘collar’’; lives in sand/mud 4. Trunk pyriform; undivided tentacle stems of equal length and less than 4% of trunk length .... 7. cymodoceae — Trunk elongate, cylindrical; undi- vided tentacle stems longer (> 4.5% of trunk) and of unequal length (dor- sal pair longer than ventrals) Me 2 aos ee T. dehamata 5. Introvert hooks oriented posterior- ly; 50-100 um tall T. minor — Introvert hooks point in all direc- tions; 30—400 um tall 6. Introvert without hooks ......... 7 — Introvert bearing dark scattered hooks’ 5: eee eee suc ne diay pres 8 7. Introvert collar purple; tentacules white; lives in rocks ....... T. dyscrita — Introvert collar unpigmented; ten- tacules with pigment spots; lives in unconsolidated sediments ........ 8. Introvert collar unpigmented; ten- tacules with pigment spots; Atlantic Ocean T. alutacea — Introvert collar purple; tentacules unpigmented; Pacific Ocean 9. Tentacular crown appears to have 6 stems; many subdivisions of branches, tentacules mainly at distal tips T. pyroides — Tentacular crown with 4 stems each dividing once to give 8 branches; tentacules appear all along branch AAS A wre aad Peta ter ear tinea tr Oe T. blanda Taxonomic Section Genus Themiste Gray, 1828 Type species.—Themiste hennahi Gray, 1828 by monotypy. Diagnosis. — Introvert shorter than trunk; body wall with continuous muscle layers; oral disc carries tentacles basically sur- rounding mouth but extending with growth along margins of branching stemlike out- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON growths of oral disc; with or without hooks; two introvert retractor muscles; contractile vessel with villi or tubules; spindle muscle not attached posteriorly; two nephridia; species small to large sized (adults 4—400 mm). Subgenus Themiste (Themiste) Edmonds, 1980 Diagnosis. — Contractile tubules (villi) few (less than 20), long and threadlike. Within this subgenus one finds two sub- sets: those with and those without hooks as adults. The two species lacking hooks live in the eastern Pacific Ocean. The three species that bear hooks are similar to one another. Two of these live in the mid lati- tudes of the northern Pacific Ocean, and one lives in the western Atlantic. Those without hooks: Themiste dyscrita (Fisher, 1952) Dendrostomum dyscritum Fisher, 1952:417— 419, fig. 87 B-E; pl. 30, fig. 3, pl. 34. Themiste dyscrita.—Stephen & Edmonds, 1972:199.—Oglesby, 1968:155-177.— Foster, 1974:856.—Rice, 1980:493—494. Material examined. —USNM, Fisher’s California material (holotype, 21221; and 21538, 27680, 27682) and Rice’s Oregon specimens (35935); SBNH, Hollister Ranch (42496M); our 1987 Santa Barbara and Santa Cruz, California material. Discussion. —Purple pigment on the smooth “‘collar” region just behind the ten- tacles but lacking pigment on the tentacules distinguishes this species from 7. hennahi. The nephridiopores commonly open 2—3% of the trunk length posterior to the anus. Fixing muscle F2 is more often (but not always) found on the rectum and the re- tractors commonly originate at 65-75% of the distance to the posterior end of the trunk. Another difference from 7. hennahi is its subspherical trunk. 7. dyscrita’s ecology is also different; it lives in burrows made by VOLUME 101, NUMBER 4 other animals, commonly pholid bivalves, in intertidal shales or sandstones. It does not appear to coexist with the hooked 7. pyroides, which may live in the same bay but occupies a different niche. The body col- or ranges from pale cream in small worms to dark brown in larger ones. Distribution. —The U.S. west coast from Oregon to southern California at intertidal depths (one record at 18 m). Themiste hennahi Gray, 1828 Themiste hennahi Gray, 1828:8, pl. 6, figs. 4, 4a.— Baird, 1868:98.—Stephen, 1964: 458; 1967:92—94.— Rice & Stephen, 1970: 53-56.—Amor, 1970:495-504; 1975:21- 23.—Stephen & Edmonds, 1972:201- 203.—Tarifeno, 1975:251-—266; 1976:29- 37.—Tarifeno & Rojas, 1978:118-119. Dendrostomum lissum Fisher, 1952:419- 422, pl. 35. Themiste lissa.—Stephen & Edmonds, 1972: 206.—(Not Cutler & Cutler, 1979a:966- 967). Dendrostoma mytheca Chamberlin, 1920: 30. Dendrostoma perimeces Fisher, 1928:196—- 198, pl. 6, figs. 3, 3a, pl. 7, fig. 1, pl. 8, figs. 2, 2a; 1952:415-417.—MacGinitie, 1935:63 1-682. Themiste perimeces.—Stephen & Ed- monds, 1972:207-—209. Dendrostoma peruvianum Collin, 1892:179— 180.—Fischer, 1914b:12-13. Dendrostomum peruvianum. —Wesenberg- Lund, 1955:12-13. Dendrostomum ramosum Quatrefages, 1865:629.—Leroy, 1936:425. Themiste ramosa.—Baird, 1868:98.—Ste- phen & Edmonds, 1972:340 (in list of in- certae sedis, etc.).—Saiz Salinas, 1984: 184-185. Sipunculus (Aedematosomum) rapa Qua- trefages, 1865:627.—Saiz Salinas, 1984: 172-173. Phascolosoma rapa. —Baird, 1868:86. Dendrostomum schmitti Fisher, 1952:422, fig. 87F. 753 Themiste schmitti.i—Amor, 1970:499.— Stephen & Edmonds, 1972:212. Dendrostoma zostericolum Chamberlin, 1920:30.— Peebles & Fox, 1933:201. Dendrostomum zostericolum. — Fisher, 1952:411-415, pl. 30, fig. 1, text-fig. 87A, pls. 31—32.—Gross, 1954:403-423. Themiste zostericola. —Stephen & Ed- monds, 1972:213-214.—Rice, 1980:493. Material examined.—BMNH, paralec- totype (1965.16.2/5); RSME, two Peru specimens identified by Stephen (1965.27.2); our own 1987 Santa Barbara and Monterey Bay, California material. 7. /issa: USNM, Fisher’s holotype (21222) and (27671). T. perimeces: USNM, Holotype (19615), para- type (21547), and other California material (26446, 27612); SBNH (42493, 42494, 47660); specimens at Moss Landing Marine Lab. D. peruvianum: MNHU, Collin’s syn- types (Nr. 2087); ZMUH, Fischer’s from Chile (V6184). D. ramosum and D. rapa: MNHN, Quatrefages’ types (V25). T. schmitti: USNM, Fisher’s holotype (21216), paratypes (21217). 7. zostericola: MCZH, type (2181; this is a dried up bag of skin and of no value); SBNH (47662); USNM, Fisher’s Baja material; many Californian specimens collected for us by Pacific Bio- Marine. Discussion. — Themiste hennahi is an eastern Pacific taxon inhabiting intertidal gravely to silty sand (including mud flats, eel grass beds and between and under turn- able boulders). Trunks are commonly 25- 100 mm long (occasionally up to 200 mm, rarely larger) with a slender pyriform to cy- lindrical shape sometimes with a nipple-like posterior end. The introvert retractors com- monly originate at 50-60% of the trunk length (Fig. 7) and the introvert is without a pigmented collar. The careful redescrip- tion by Rice and Stephen (1970) is well il- lustrated. Tarifeno (1975) gives detailed ecological and behavioral information. The tentacular crown is asymmetrical, with the dorsal pair being longer and splitting sooner than the ventral pair giving the appearance 754 of six stems. The nephridiopores open O- 10% of the trunk length posterior to the anus, in larger worms this distance is usually less than 2%. The 18 syntypes of Collins’ T. peruvian- um (reduced to junior synonym by Stephen 1964) all had pyriform trunks with the pos- terior end coming to a blunt point. The spin- dle muscle is very strong, and in a few this seems to be contracted along the rectum giving the latter an accordion pleated look. This population allegedly has protuber- ances on the esophagus. However, we see only a few bumps, which easily rub off and appear to be agglutinated gametes adhering to the esophageal wall. Quatrefages’ (1865) two species (7. rapa and 7. ramosum) were considered incertae sedis until the reconsideration by Saiz-Sa- linas (1984) wherein he determined them to be synonyms of this species. When Rice & Stephen (1970:56) rede- scribed this species, they made the following statement: “Themiste hennahi also shows many similarities to the species described as Dendrostomum zostericolum Chamber- lin 1919 and Dendrostomum schmitti Fish- er 1952. The possibility that these may all represent a single species remains to be de- termined by future studies.”’ We are unable to find any truly meaningful characters that would serve to differentiate these species. Therefore, we reluctantly agree with their suggestion and place the familiar 7. zoster- icolain synonomy. The apparent gap in dis- tribution (Peru to Baja California) may be an artifact of undercollection in Central America, but we discarded the possibilty of subspecific rank. The small ones look dif- ferent from the large ones, but, when one fills the gap with middle sized worms, the continuum becomes clear. Fisher’s 7. perimeces was originally dif- ferentiated from 7. zostericola by having more gut coils, fixing muscles usually ab- sent, papillae of two or three sizes, and fewer contractile vessel villi. Our examination of Fisher’s material (some not previously dis- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sected) and other California specimens demonstrated that this dichotomy is artifi- cial. At least one fixing muscle is present in almost every worm. There are usually 8-14 contractile vessel tubules, but one 9 mm worm has only four, so this, as well as both papillae size and number of gut coils, is somewhat age dependent. Variation exists and the worms bearing this name are simply large T. hennahi. Rice (1980) in the most recent general treatment of the California fauna did not include 7. perimeces. Themiste lissa is not well known (our to- tal sample size is 19) and the putative dif- ferences from 7. hennahi are subtle. Fisher (1952:422) comments on the close relation- ship between this taxon and his new 7. schmitti (now considered a synonym of 7. hennahi): “*... possibly they represent the extremes of geographic variation of one species.” We concur. In his key Fisher used the location of the gonads “on the surface of the retractors” vs. “‘on body wall just back of origins” to separate these two taxa. In our analysis of the 7. hennahi/peruvian- um and T. lissa material, we found the go- nads to be located on the body wall, on the retractors, and on combinations of this in different worms but showing significant within-deme variation. When Fisher asserted that T. /issa differed from the other Californian species (he did not mention 7. hennahi, which had not been “‘rediscovered”’ at that time), he put consid- erable weight on the distance between the anus and the nephridiopores—he gave no measurements but stated that the distance was “conspicuous.” In the holotype (with a 20 mm trunk) the distance is 10% of the trunk length, but in six of Fisher’s paratypes (7-11 mm trunks) it varies from 4—7% (in one worm these are at the same level). In two recently collected specimens the dis- tance was 12% in an 8 mm worm and 5% in an 11 mm worm. As noted in the intro- duction, this is not a species specific char- acter. Our present construct includes worms of VOLUME 101, NUMBER 4 very different sizes from what may appear to be rather different habitats. The proba- bility exists that some live in more optimal niches than others and that these can achieve a larger size. If a trend exists, it seems to be that the higher latitudes are better suited to this genotype. The smallest worms come from the Gulf of California, and this pop- ulation may be isolated with restricted gene flow. A reexamination of the two T. lissa from Durban (Cutler & Cutler, 1979a) showed them to be Antillesoma antillarum. Distribution. —Chile and Peru, Gulf of California and California; unconsolidated intertidal or shallow subtidal sediments. Quatrefages’ 7. ramosa specimens are la- beled, “Qf. du Bresil M. Pissis,’’ but we are hesitant to accept this as a valid location as there have been no 20th century confir- mations outside the eastern Pacific. Those with hooks: Themiste alutacea (Grube & Oersted, 1858) Dendrostomum alutaceum Grube & Oerst- ed, 1858:118.— Quatrefages, 1865:630.— Diesing, 1859:765.—Keferstein, 1865: 438.—Selenka et al., 1883:84-85.— Fischer, 1895:18; 1922:18.—Gerould, 1913:417-418.—Leroy, 1936:425. Themiste alutacea. —Baird, 1868:98.—Ste- phen & Edmonds, 1972: 196-197.—Cut- ler, 1973:162-164.—Cutler & Cutler, 1979b:105-106.—Rice, 1975a:37-47, 1975b:147. Sipunculus (Phymosomum) orbiniensis de Quatrefages, 1865:622. Phascolosoma orbiniense. —Baird, 1868: 93.—Stephen & Edmonds, 1972:339 (in list of incertae sedis etc.). Themiste (Themiste) orbiniensis. —Saiz-Sa- linas, 1984:124—132. Themiste blandum [sic].—Murina, 1968: 423. Dendrostomum petricolum Amor, 463-467, pl. 3, 5 figs. 1964: 755 Themiste petricola.—Stephen & Edmonds, 1972:209.—Amor, 1975:185-192. Dendrostomum rosaceum Amor, 1964:459, pls. 1, 2 and 3. Themiste rosacea.—Stephen & Edmonds, 1972:211-212. Material examined.—MNHU, Grube’s type (1033) plus some of Selenka’s from Brazil; other material from North Carolina (Cutler, 1973) and M. Rice’s Florida ma- terial. T. orbiniensis: MNHN, Quatrefages’ material (V25). 7. rosacea and T. petricola: USNM, paratypes (30995, 30996). T. blan- da: ZIAS, Murina’s Gulf of Mexico speci- mens. Discussion.—This warm water, western Atlantic species inhabits hard substrates (spaces in coral, oyster beds or soft rock), is small (trunk rarely over 25 mm, commonly less than 15), and exhibits indirect devel- opment. As an early juvenile it has a single ring of introvert hooks, but these are soon lost and replaced by scattered adult hooks. We have sufficient data on this species (Rice 1975b) to assert that early in a worm’s life four tentacular stems exist; two of these often subdivide near the base to give the appear- ance of six (sometimes five) in many adult worms. This is different than the four di- viding into eight seen in 7. blanda. The subsequent branching of these primary units is limited in the smaller worms, and the thin tentacules (with pigment spots) are located all along each branch. The hooks in this species can be numerous (over 100 in one 5 mm worm) but not as large as in other members of this subgenus (75—150 um). This species has lecithotrophic pelagic larvae, 1.e., indirect development (Rice 1975b). Themiste rosacea and T. petricola were described from Argentina, and, while we have not been able to obtain additional ma- terial, our examination of the literature and the paratypes convinces us that these three putative taxa are not biologically distinct. The presumed differences are in the number of tentacles, the number of hooks, the size, 756 shape and color of the trunk, and the po- sition of the fixing muscles. As noted in the Introduction, these characters are too vari- able to be meaningful. While gene flow may be restricted and subspecific status might be appropriate, we cannot discern consistent and meaningful morphological differences. Our examination of Murina’s Gulf of Mexico worms convinced us that they be- long to this taxon. The hooks in this pop- ulation are larger, measuring up to 300 um. Saiz-Salinas (1984) presented a detailed re- description of Quatrefages’ T. orbiniensis and a comparison to others in this group. Towards the end of his discussion, he noted that a future comparative study of speci- mens may result in the synonymizing of Quatrefages’ name. When Amor examined this material in 1969, she left a note in the bottle that she thought it was T. alutacea, and our analysis of all available information brings us to the same conclusion. Distribution.—Western Atlantic Ocean from North Carolina (34°N) to Argentina (42°S) at depths less than 30 m. Themiste blanda (Selenka, deMan, & Bulow 1883) Dendrostomum blandum Selenka et al., 1883:85-86, pl. 1, fig. 9, pl. 11, figs. 159- 162.—Selenka, 1885:14.—Ikeda, 1904: 53-55; 1924:30-31.—Ostroumov, 1909: 319-324.—(Not Fischer, 1922:18-19)— Sato, 1930:24—28; 1937:162; 1939:412.— Fisher, 1952:405. Themiste blanda. —(Not Murina, 1968: 423)—Stephen & Edmonds, 1972:197.— Cutler, Cutler & Nishikawa, 1984:285. Material examined. —MNHU, type and paratype (962, 1034); Our 1979 Japanese material. Discussion. —In Cutler et al. (1984:285) we noted “. . . what has been called JT. blan- da may be simply small 7. hexadactyla.” Our current studies cast some doubt on that hypothesis and we now consider 7. hex- adactyla to be a junior synonym of T. py- roides. While acknowledging the very sim- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ilar nature of this taxon and the other two hooked members of this subgenus, we retain T. blanda as a distinct species for the pres- ent. It differs from 7. pyroides in two ways: 1. Tentacular crown—this array is simpler with four tentacular stems, each dividing only once, and it has longer tentacules along each branch giving each branch a somewhat pinnate appearance. 2. The upper limit of trunk size seems to be about 25 mm (vs. 200 mm). The similarities are marked and some of the alleged differences referred to by earlier authors (such as fixing muscle ar- rangements) are not constant enough to be meaningful. The similarities to 7. alutacea are also marked, but it differs by having four stems dividing symmetrically into eight, lacking pigment spots on its stouter tenta- cules, and by living in cool northern Pacific water vs. warmer western Atlantic water. Distribution. —Intertidal in Honshu and Hokkaido, Japan. Themiste pyroides (Chamberlin, 1920) Dendrostoma pyroides Chamberlin, 1920: 31.—Fisher, 1952:406-409. Themistse pyroides. —Stephen & Edmonds, 1972:210.—Rice, 1967:143-171; 1980: 492-—493.—Frank, 1983:22. Dendrostoma blandum.— Fischer, 1922:18— 19. Dendrostoma petraeum Fisher, 1928:195— 196. Dendrostoma hexadactylum Sato, 1930: 28-33, pl. 4, figs. 20-24, text-figs. 13-15; 1937:162-163; 1939:412.—Okuda, 1946: 224. Dendrostomum hexadactylum. — Fisher, 1952:410-411, pl. 30, fig. 2. Themiste hexadactyla. —Stephen & Ed- monds, 1972:203-204.— Cutler & Cutler, 1981:75.—Cutler, Cutler & Nishikawa, 1984:285-286. Material examined. —MCZH, Chamber- lin’s type (2.182) and paratype (2.183); USNM, several of Fisher’s California spec- imens (21503, 21548), a few from Vancou- ver, B.C. identified by M. Rice (96094); our VOLUME 101, NUMBER 4 1987 Carmel and Monterey, California specimens. 7. hexadactyla: ZITU, type (G26) plus other Sato material (1-4, 1-5, 4-11, 5-11, 5-15; one now at USNM, 59994): USNM, Fisher’s California speci- mens (21504, 27672). Discussion.— This taxon is similar to the two previous species but does have a pyri- form shape with a bluntly pointed posterior end regardless of size (Fig. 10). The mor- phological distinction we now afhirm is; four primary tentacular stems divide close to their base into 2, 3, or 4 branches, each then subdividing, as the worm grows, into small- er branches giving off unpigmented tenta- cules mostly at the distal ends. In specimens the same size as a given 7. blanda or T. aluiacea, the crown of T. pyroides is much more voluminous with shorter terminal tentacules. Whereas many specimens from central California have trunks 20-40 mm long, some worms from the more northern latitudes (British Columbia and Hokkaido) may exceed 100 mm. The fixing muscle placement is not as constant as Fisher (1952) implies nor is the anastomosing network of contractile vessel branches as clearcut or consistent. In 12-14 mm worms only 8 or 10 long tubules are at the terminal end of the vessel, while in a 23 mm worm one sees branches off the main vessel adhering to the sides of the esophagus. In a 40 mm worm these branches are more elaborate and sur- round the esophagus. This network be- comes much more complex in an 80 mm worm, much like that shown in Fisher’s (1952) plate 29, fig. 3. Distribution. —Honshu and Hokkaido, Japan and west coast of North America from southern Alaska to Baja California at inter- tidal depths. Subgenus 7Themiste (Lagenopsis) Edmonds, 1980 Diagnosis. —Contractile vessel villi many (more than 100), short and digitiform. Two subsets exist in this taxon, those species with and without introvert hooks. Within the hookless triad is one common 757 Fig. 10. Themiste pyroides of different sizes show- ing typical pointed pyriform trunks, hooks on intro- verts and voluminous tentacular crowns. circum-subtropical inhabitant of hard sub- strates, T. /ageniformis, which has a long list of junior synonyms. The other two species seem to be endemic to Australia liv- ing in soft substrates. The hooked subset also has two taxa endemic to Australia and one more widely spread subtropical taxon. Those without hooks: Themiste cymodoceae (Edmonds, 1956) Dendrostomum cymodoceae Edmonds, 1956:299-301, pl. 2, fig. 1, text-figs. 15- 16; 1957:55-63.—Akesson, 1958:147- ND Ike Themiste cymodoceae. —Stephen & Ed- monds, 1972:197—198.—Edmonds, 1980: 38-40. Material examined.—Three specimens (79-85 mm trunks) from the type locality from collections of S. J. Edmonds. Discussion. —This species is similar to 7. dehamata in many ways (e.g., pigmented 758 tentacules but unpigmented collar), and we have reservations about its distinct status. However, S. J. Edmonds (pers. comm.) has collected and observed these in the living state and feels strongly that they are differ- ent species. The differences we can agree on are not sharp or easily quantified. The trunk in T. cymodoceae is flask shaped or pyri- form (length 50-90 mm but less than five times the width) whereas 7. dehamata is slender and elongate (length at least 10 times width). The second morphological differ- ence has to do with the tentacular crown. In T. cymodoceae the tentacular stems are shorter (2.5—4% of trunk length) and are of equal size (giving it a bushy appearance). In T. dehamata the stems are longer overall (4.5-7% of trunk length) and in each worm the dorsal stems are longer than the ventrals by 25-67% (Edmonds describes this as plu- mose). The niches of these two species may differ; however, they have both been col- lected from among the roots in intertidal eel grass beds (Edmonds, 1980). Distribution. —South Australia; intertidal unconsolidated sediments. Themiste dehamata (Kesteven, 1903) Dendrostoma dehamatum Kesteven, 1903: 69-73, pl. 7, fig. 7. Dendrostomum dehamatum. —Edmonds, 1956:296. Themiste dehamata.—Stephen & Ed- monds, 1972:198-199.—Edmonds, 1980: 34-36. Dendrostoma ellipticum Sato, 1934:20-22, pl. 1, fig. 10, text-figs. 22-25; 1939:411. Themiste elliptica.—Stephen & Edmonds, 1972:199-200.—Cutler & Cutler, 1981: 74-75; Cutler, Cutler & Nishikawa, 1984: 283. Dendrostomum fisheri Amor, 1964:467— 469, pl. 3 and 4. Themiste fisheri.—Stephen & Edmonds, 1972:200. Material examined. —Four specimens from near the type locality from the collec- tions of S. J. Edmonds. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Discussion. — This taxon was recently re- described by Edmonds (1980). Whereas the written description resembles 7. lagenifor- mis (see preceding section for comparison to 7. cymodoceae), it is distinct in several ways. Themiste dehamata is more cylin- drical, elongate, larger (commonly over 40 and up to 170 mm), and lives in sand or mud habitats. Sato’s (1934) D. ellipticum was based on a single 64 mm worm, which cannot now be located. Although Sato did not comment on the contractile vessel villi branching (and his drawing is not sufficiently detailed), Fisher (1952) asserted that they were un- branched, and, therefore, it was distinct from similar species. Sato differentiated it from others with short villi on the basis of pa- pillae shape (elongate elliptical in this and roundish in others including 7. dehamata). In Stephen & Edmonds’ key (1972) they were unable to differentiate these two species. As suggested in Cutler & Cutler (1981) there now seems no justification for retaining this species name. Edmonds (1980:35) first suggested that T. fisheri is very similar to this species. The only “significant” difference, according to him, is the absence of a rectal caecum in 7. fisheri; however, Amor stressed the different number of tentacular stems. Amor’s species rests on a single 100 mm specimen, which we have tried unsuccessfully to obtain for examination. Having only this one worm with only these minor differences, we here- by reduce it to a junior synonym. Distribution. —New South Wales and South Australia; intertidal unconsolidated sediments. Because of our synonymies, one record is from Japan and one from Argen- tina. Since these latter specimens cannot be examined, we would caution against in- cluding them in zoogeographical analyses. Themiste lageniformis Baird, 1868 Themiste lageniformis Baird, 1868:98-99, pl. 10, fig. 3.—Rice & Stephen, 1970:66-— 67.—Stephen & Edmonds, 1972:205- VOLUME 101, NUMBER 4 206.—Cutler, 1977a:147.— Williams, 1977:1-206.—Cutler & Cutler, 1979a: 966.—Edmonds, 1980:41-42.—Pilger, 1982:143-156.—Cutler, Cutler, & Nishi- kawa, 1984:283-284. Dendrostoma signifer Selenka et al., 1883: 86-87 (part).—Sluiter, 1886:515; 1891: 115; 1902:19.—Fischer, 1895:17; 1914a: 72-74; 1914b:10-11 (part); 1919:282-283 (part); 1922:19 (part); 1923:22.—Auge- ner, 1903:300—301.—Ikeda, 1904:56- 57.—Hammerstein, 1915:2—3.—Grave- ley, 1927:87.—Awati & Pradhan, 1935: 102-113; 1936:114-131.—Leroy, 1942: 41-43. Dendrostomum signifer.—Edmonds, 1956: 297.—Wesenberg-Lund, 1959a:198-199; (Not 1959b:213; 1963:129-130). Dendrostoma minor. —Chin, 1947:100. Dendrostoma robertsoni Stephen & Rob- ertson, 1952:438—439, pl. 1, figs. 3-4. Dendrostomum robertsoni. —Wesenberg- Lund, 1963:130. Themiste robertsoni. —Stephen & Ed- monds, 1972:210-—211.—Stephen & Cut- ler, 1969:116.—Cutler & Cutler, 1979a: 967-968. Dendrostoma stephensoni Stephen, 1942: 252-253 (part). NOT Themiste stephensoni. —Cutler, 1977b: 154. Dendrostoma tropicum Sato, 1935:313-315, pl. 4, fig. 15, text-fig. 11; 1939:411. Dendrostomum tropicum. — Wesenberg- Lund, 1963:131-132. Themiste tropica.—Stephen & Edmonds, 1972:213.—Cutler, 1977b:154.—Cutler & Cutler, 1981:76—77.— Murina, 1981:16. Phascolosoma glaucum Lanchester, 190Sa: 82, TOIL, 1s, 3. Golfingia (Thysanocardia) glauca. —Ste- phen & Edmonds, 1972:123.—Cutler, 1977b:152. Themiste glauca. —Gibbs, Cutler, & Cutler, 1983:302. Phascolosoma_ pyriformis Lanchester, 1905b:39, pl. 2, fig. 5.—Sato, 1939:404— 406. 759 Golfingia pyriformis.—Murina, 1964:261; 1967:43. Golfingia (Thysanocardia) lanchesteri pro Phascolosoma_ pyriformis Lanchester, 1905b (pre-occupied by Phascolosoma pyriforme Danielssen in Théel 1875): Ste- phen & Edmonds, 1972:124. Themiste pyriformis. —Gibbs, Cutler, & Cutler, 1983:302. Material examined.—BMNH, syntypes (165.25.9/10); BPBM, Hawaiian collections (N101, 1973.205); Naval Oceans Systems Center, Kailua (at BPBM) and our 1984 Hawaiian worms plus Sta. 87 from Ft. Pierce, FL in the collections of M. Rice. D. signifer. ZMUH, Type (V2052); MNHU (384, 982). T. robertsoni: RSME, type (1958.23.74); UZMK, Wesenberg-Lund’s; our own from Indian Ocean. D. stephensoni: RSME, type (1958.23.34) plus (1958.23.35, 37, 38; 1966.1.10). 7. tropicum: Galathea Sta. 256 in our collections; the type cannot be located. G. glauca: UZMC, type; UZMK, Galathea Sta. 630, 631; G. pyriformis: UZMC, type material; ZITU, Sato’s from Formosa (3-13). Discussion. —This is the most common, most studied, and most widely distributed species in the genus. Baird’s original de- scription (1868) of this species was over- looked by Selenka and others for almost a century. Williams (1972, 1977) and Wil- liams & Margolis (1974) have contributed to an understanding of its burrowing habits and its development (indirect with pelagic larvae). Pilger (1987) has demonstrated that this species may also be parthenogenic. The nature of this species has been fairly clear over the years with two exceptions. Several authors confused the issue by including some animals with introvert hooks. These are now considered to be 7. minor huttoni, a taxon remarkably similar to this one except bear- ing hooks. Wesenberg-Lund (1963) report- ed the presence of two or three rows of hooks in two worms from Tristan de Cunha. We have not been able to locate these specimens and are reluctant to assign them to this ge- 760 nus because of the hook arrangement and her lack of comment on the tentacular crown. The alleged number of tentacular stems: 4, 5, 6 or 8 has also added to the confusion (see Morphological Characters). When Stephen & Robertson (1952) de- scribed 7. robertsoni, they differentiated it from this species on the basis of number of tentacular stems plus a few other points such as number of fixing muscles, length of re- tractors, number of contractile vessel villi, etc. That model was followed by a few other authors, but, as pointed out by Cutler & Cutler (1979a), the differences are subjec- tive. Our current reexamination of this ma- terial indicates that these particular char- acters are within the range found in one 7. lageniformis population. While the holotype and some of Stephen’s (1942) material of 7. stephensoni are Gol- fingia capensis (Gibbs & Cutler 1987, Cutler & Cutler 1987), four specimens are 7. /a- geniformis. The original foundation for T. tropica was one 14 mm worm (Sato 1935). He did not compare it to this species since he incor- rectly said it (D. signifer at that time) had long contractile vessel villi. Wesenberg-Lund (1959b) reported two worms from 90-110 m in the northern Gulf of Guinea. We could not find these in any museum collections, and, since these are the only records from this part of the Atlantic, we are skeptical. A careful reading of her description suggests that these are more probably two Thysan- ocardia catharinae. She later (1963) record- ed three small (S-10 mm) worms from Capetown. There were a few puzzling as- pects to her paper including rings of hooks and an apparent contradiction (key vs. text) about whether or not the villi were branched, a matter now known to be size dependent, not species specific. Without access to these worms we cannot reassign them with con- fidence, but they are in the 7. minor group. Cutler’s (1977b) and Murina’s (1981) worms were all less than 20 mm long, and nothing PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON would exclude them from our present con- cept of 7. lageniformis (i.e., some un- branched contractile vessel villi in small specimens). Now it is clear that 7. tropica should be considered a junior synonym. The confusion created by Lanchester’s (1905a,b) two species (Phascolosoma glauca and P. pyriformis) has been addressed in Gibbs et al. (1983) when they revised the golfingiid subgenus now genus) 7hysano- cardia. At that time it was recognized that these two taxa belong in 7hemiste, but they were not assigned to any different species in anticipation of the present work. All of Lanchester’s specimens and some of Sato’s (1939) T. pyriformis were highly contracted so that the anus and nephridiopores were withdrawn (both said “‘on the introvert” — a feature also noted by Murina (1964)). In some of our strongly retracted recently col- lected Hawaii material, the anus and ne- phridiopores were withdrawn, giving the false impression of being on the introvert (see section on Morphological Characters). Since we now find no significant differences, we submerge both names under 7. /ageni- formis. Themiste lageniformis is one of the small- er species (rarely more than 35 mm) which lacks hooks (after early juvenile stages), usu- ally lives in soft rock, old coral or mussel beds, has a bluish band or collar on the introvert (Sometimes very pale), and long tentacules. Distribution. —Well established in the western Pacific (from Japan to Australia and Hawaii) and throughout the Indian Ocean. It has been recorded from South and South- west Africa, Cuba, and Florida. Species with hooks: Themiste minor minor (Ikeda, 1904) Dendrostoma minor Ikeda, 1904:57—59, text-figs. 16, 92-95.—Sato, 1939:411.— Stephen, 1942:252.—(Not Chin, 1947: 100). VOLUME 101, NUMBER 4 Dendrostomum minor. —Wesenberg-Lund, 1963:128-129. Themiste minor. —Stephen & Cutler, 1969: 116.—Stephen & Edmonds, 1972:207.— Cutler, 1977a:147.—Cutler & Cutler, 1979a:967; 1980:206; 1981:75-76.— Cutler, Cutler, & Nishikawa, 1984:284— 285. Dendrostoma signifer. — Fischer, 1914b:10- 11 (part); 1919:282-283 (part); 1922:19 (part). Dendrostomum fuscum Edmonds, 165-167, pl. 3, figs. 7-9. Themiste fusca. —Stephen & Edmonds, 1972:200-201.—Edmonds, 1980:40-41. Golfingia coriacea. —Murina, 1972:298. 1960: Material examined. —ZMUT, Ikeda’s type material was located but is completely dried out; RSME, Stephen and Cutler’s South African worms (1966.1.9); our 1979 Japanese material from near the type lo- cality (neotype named in Cutler and Cutler, 1981; part deposited at USNM). 7. fuscum: two Australian worms from the collections of S. J. Edmonds. Discussion. —Except for the presence of introvert hooks (generally 50-200 um tall) and a more loosely wound gut coil, this tax- on is ecologically and morphologically sim- ilar to 7. lageniformis. The animals are small, i.e., trunk length usually less than 20 mm and we have seen 4 mm worms which are sexually mature. Edmonds’ (1960) differentiated 7. fusca from 7. minor on the basis of presumed but vague differences in hook and papillae structure, attachment of esophagus and the arrangement of the contractile vessel villi. In 1980 Edmonds did say that it was hard to distinguish it from some Japanese 7. mi- nor but that the contractile vessel and fixing muscles in the Japanese worms seem sim- pler. From our analysis of the literature and the animals, we conclude that the differ- ences Edmonds reported probably result from Ikeda’s narrowly constructed descrip- 761 tion but are not biologically meaningful, i.e., this amount of variation does exist within demes (see section on Morphological Char- acters above). Murina’s 1972 record of a single 7 mm worm which she named Golfingia coriacea was attributed to this species by Edmonds (1980:29), and we concur. Chin’s 1947 rec- ord includes the statement that there were no hooks on his material; we herein move Chin’s record to 7. lageniformis. Distribution. —Japan, China, South Aus- tralia, southern Africa, and one record from southern Argentina. The latter record (Cut- ler and Cutler, 1980) is a 4 mm worm of uncertain affinity. It seems clear that this is a cool temperate, intertidal, and shallow subtidal species. The few deep water records are based on few, very small specimens and may represent either anomalies or confu- sion in station data. The Cutler (19774) rec- ord at 4510 m should, therefore, be ignored in zoogeographical analyses. Themiste minor huttoni (Benham, 1904) Phascolosoma huttoni Benham, 1903:177-— 184. Dendrostoma huttoni Benham, 1904:306- 307. Dendrostomum huttoni.—Edmonds, 1960: 164-165. Themiste huttoni.—Stephen & Edmonds, 1972:204.—Cutler, 1977a:146.—Ed- monds, 1980:36-38. Dendrostoma signifer Selenka et al., 1883: 86-87 (part).—Augener, 1903:337.— Fischer, 1919:282-—283 (part); 1926:206. Material Examined. —RSME, 1958.23. 124; two from the collections of S. J. Ed- monds. Discussion. —In the years 1903-1904 Benham published one faunal index and two publications in which the name “‘huttoni” appears. The formal new species description was in the 1904 paper. This name is pre- 762 sented in a confusing context along with Si- punculus aeneus Baird, 1868 and S. /utu- lentus Hutton, 1879. The former has been synonymized under Siphonosoma australe (Edmonds 1961; Rice and Stephen 1970) and the latter placed on the list of incertae sedis (Stephen & Edmonds 1972). Themiste minor huttoni is here consid- ered a subspecies of a more cosmopolitan taxon with restricted gene flow between the two populations. The similarities to the nominate form are substantial. The differ- ences we can identify are: Larger (trunk length may reach 55 mm), more hooks, and the hook bearing region extends over more (55-75 vs. 25-35%) of the introvert. These are not clear distinctions and may represent only a more optimal set of conditions and less wave action, inducing or permitting a different expression of the genotype (e.g., larger size and more hooks produced or few- er rubbed off, like the situation seen in the Californian/Japanese 7. pyroides). This hy- pothesis needs testing. Distribution. — Australia, New Zealand and Chatham Is.; intertidal, hard substrates. Themiste variospinosa Edmonds, 1980 Themiste variospinosa Edmonds, 1980:42- 43, figs. 62, 68, 69. Material examined.—A specimen from the collections of S. J. Edmonds from the type locality. Discussion. —The six worms upon which this species is based have many ecological and morphological similarities to 7. minor huttoni. The principal difference lies in the nature of the introvert hooks (Fig. 3). They exhibit much greater variation in size on any given worm (30-400 um tall) and are pointing in all directions, not just poste- riorly. While this may be only a localized variation of the surrounding population of T. minor huttoni, we propose no change in its status at this time. We hope that addi- tional material will be collected to more firmly test the validity of this taxon. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Distribution. —Queensland, Australia, in- tertidal coral. Zoogeographical Summary The most striking feature is how non- overlapping the two subgenera are. The five Themiste (Themiste) species live in the western Atlantic, eastern Pacific and Jap- anese waters. Three of the five 7. (Lagen- opsis) are endemic to Australia/New Zea- land, the fourth lives in the western Pacific Ocean and South Africa while the fifth is circum-tropical/subtropical (but absent in the eastern Pacific). Overlap of subgenera exists on Honshu (central Japan) where T. (T.) blanda and T. (T.) pyroides are found along with 7. (L.) minor. Another case is the Caribbean (and east coast of Florida) where 7. (L.) lageniformis and T. (T.) alu- tacea coexist. Southern Argentina may be a third such place, but at this time the data base is not large enough to use with confi- dence. Secondly, like in many other genera, there is One very widespread species (the circum- tropical 7. /Jageniformis) while most others have rather restricted distributions. Finally, this genus is absent from the North Atlantic above Cape Hatteras on one side and West Africa on the other. Acknowledgments The encouragement and shared frustra- tions as well as the exchange of ideas and specimens with S. J. Edmonds, Adelaide, have contributed greatly to the completion of this work. Our work in Hawaii was ac- complished with the assistance of B. Burch, Bishop Museum and M. Hadfield, Kewalo Marine Laboratory, University of Hawai. The field work in California was dependant on the generous cooperation of F. Hoch- berg, Santa Barbara, J. Nybakken, Moss Landing, and M. Saffo, Santa Cruz. L. Cut- ler, Palo Alto, assisted in the lab and field work in Hawaii and California while J. Sil- verstein, Seattle, provided valuable assis- VOLUME 101, NUMBER 4 tance in California and was responsible for obtaining the karyotypes. J. Swartwout, Utica; G. McDonald, Santa Cruz; and Anne Darling, Clinton, assisted with the artwork. Financial support was provided by the Na- tional Science Foundation (grant BSR 83- 14301 and BSR 86-15315). The cooperation of the following persons and institutions in the loan of reference ma- terial and/or providing access to their col- lections was essential to the completion of this project and greatly appreciated: R. Sims (BMNH); H. Levi (MCZH); J. Renaud- Mornant (MNHN); G. Hartwich (MNHU); S. Chambers (RSME); M. Rice (USNM),; J. Kirkegaard (UZMK); Zoological Institute Tohoku University, Sendai; M. Dzwillo (ZMUH); Zoological Museum University of Tokyo. Literature Cited Akesson, B. 1958. 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The fauna of Akkeshi Bay XV. Ge- phyrea.— Contributions of the Akkeshi Marine Biological Laboratory 43:221—225. Ostroumov, A. 1909. Sur les gephyriens du nord de la mer du Japan.—Ezhegodnik Zoologischeskii Muzei 14:319-321. Peebles, F., & D. L. Fox. 1933. The structure, func- tions and general reactions of the marine sipun- culid worm Dendrostoma zostericola. — Bulletin of the Scripps Institution of Oceanography 3(9): 201-224. Pilger, J. F. 1982. Ultrastructure of the tentacles of Themiste lageniformis (Sipuncula).—Zoomor- phology 100:143-156. 1987. Reproductive biology and develop- ment of Themiste lageniformis, a parthenogenic sipunculan.— Bulletin of Marine Science 41(1): 59-67. Quatrefages, A. 1865. Gephyrea Inermia.— Histoire naturelle des Annelés marins et d’eau douce, Paris 2:599-632. Rice, M. E. 1967. A comparative study of the de- velopment of Phascolosoma agassizii, Golfingia pugettensis, and Themiste pyroides with a dis- cussion of developmental patterns in the Sipun- cula.— Ophelia 4:143-171. 1975a. Survey of the Sipuncula of the coral and beach rock communities of the Caribbean 765 Sea. Pp. 35-41 in M. E. Rice and M. Todorovic, eds., Proceedings of the International Sympo- sium of Sipuncula and Echiura, volume 1. Naucno Delo Press, Belgrade. 1975b. Observations on the development of six species of Caribbean Sipuncula with a review of development in the phylum. Pp. 141-160 in M.E. Rice and M. Todorovic, eds., Proceedings of the International Symposium of Sipuncula and Echiura, volume |. Nauéno Delo Press, Bel- grade. 1980. Sipuncula and Echiura. Pp. 490-498 in R. H. Morris, D. P. Abbott, and E. C. Had- erlie, eds., Intertidal Invertebrates of California. Stanford University Press, Stanford. ——, & A.C. Stephen. 1970. The type specimens of Sipuncula and Echiura described by J. E. Gray and W. Baird in the collections of the British Museum of Natural History.—Bulletin of the British Museum of Natural History 20(2):49— V2; Saiz-Salinas, J. I. 1984. Los tipos de Sipunculidos de deQuatrefages del Museo de Paris.— Universi- dad del Pais Vasco, Bilbao, 1-258 pp. Sato, H. 1930. Report on the biological survey of Mutsu Bay. 15. Sipunculoidea.—Science Re- ports. Series 4 Biology 5:1—40. 1934. Report on the Sipunculoidea, Echiu- roidea, and Priapuloidea collected by the Soyo- Maru Expedition of 1922—1930.—Science Re- ports of the Tohoku University Fourth Series (Biology) 9:1—32. 1935. Sipunculoidea and Echiuroidea of the West Caroline Islands. — Tohoku Daigaku, Sen- dai, Japan, Scientific Reports series 4(10):299- 329. 1937. Echiuroidea, Sipunculoidea, and Pri- apulaidea obtained in northeast Honshu, Ja- pan.— Saito Ho-on Kai Museum, Research Bul- letin 12:137-176. . 1939. Studies on the Echiuridea, sipunculids, and priapulids of Japan.—Science Reports of Tohoku Imperial University 14(4):339-459. Selenka, E. 1885. Report on the Gephyrea collected by H.M.S. Challenger during 1873-76.— Report of scientific results of voyage of Challenger Zo- ology 13(36):1-25. , J. G. de Man, & C. Bulow. 1883. Die Sipun- culiden, eine systematische Monographie. — Semper, Reisen in Archipel Phillippinen 2, 4: 1-131. Sluiter, C. P. 1886. Beitrage zu der kenntnis der Ge- phyreen aus dem Malayischen Archipel.—Na- tuurkundig Tijdschrift voor Nederlandisch-In- die 45:472-517. 1891. Die Evertebraten aus der Sammlung des Koniglichen naturwissenschaftlichen Ne- reins in Nederlandisch-Indien in Batavia. — Na- 766 tuurkundig Tijdschrift voor Nederlandisch-In- die 50:102-123. 1902. Die Sipunculiden und Echiuriden der “Siboga” Expedition, nebst Zusammenstellung der Uberdies aus den indischen Archipel be- kannten Arten.—Siboga-Expeditie, Monogra- phie 25:1-53. Stephen, A. C. 1942. The South African Intertidal zone and its relation to ocean currents. Notes on the intertidal sipunculids of Cape Province and Natal.— Annals of the Natal Museum 10(2): 245-256. 1964. A revision of the classification of the phylum Sipuncula.—Annals and Magazine of Natural History 7(13):457—-462. 1967. Notes on a collection of sipunculids from Peru.—Exposes annuels de Biochemie Medicale 21:91-94. —, & E. B. Cutler. 1969. Ona collection of Si- puncula, Echiura, and Priapulida from South African waters.— Transactions of the Royal So- ciety of South Africa 38(2):111-121. ,&8S.J. Edmonds. 1972. The phyla Sipuncula and Echiura. London, Trustees British Museum (Nat. Hist.), 528 pp. ——.,, & J. D. Robertson. 1952. A preliminary re- port on the Echiuridae and Sipunculidae of Zan- zibar.— Proceedings of the Royal Society of Edinburgh 64(4):426-444. Tarifeno, E., & M. Rojas. 1978. El phylum Sipuncula en Chile.—Studies on Neotropical Fauna and Environment 13:93-134. Tarifeno, E.S. 1975. Burrowing pattern and its vari- ation with different kinds of substratum in the Chilean sipunculan Themiste hennahi Gray 1828 (syn. Dendrostomum pervianum Collin 1892). Pp. 251-266 in M. E. Rice and M. Todorovic, eds., Proceedings of the International Sympo- sium of Sipuncula and Echiura, volume 1. Naucno Delo Press, Belgrade. 1976. Resistance to environmental stress of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON temperature and salinity in the Chilean sipun- culan Themiste hennahi Gray 1828 (syn. Den- drostomum peruvianum Collin 1892). Pp. 29- 38 in M. E. Rice and M. Todorovic, eds., Pro- ceedings of the International Symposium of Si- puncula and Echiura, volume 2. Naucno Delo Press, Belgrade. Théel, H. 1875. Etudes sur les géphyriens inermes des mers de la Scandinavie, du Spitzberg et du Groenland.—Bihang till Kungliga Svenska Ve- tenskaps-Akademiens Handlingar 3(6):1—30. Wesenberg-Lund, E.. 1955. Reports of Lund Univer- sity Chile Expedition, 1948-49. 19. Gephyrea from Chile.—Lunds Universitets Arsskrift 51(10):1-23. 1959a. Sipunculoidea and Echiuroidea from Tropical West Africa.— Atlantide Report 5:177- 210. 1959b. Sipunculoidea and Echiuroidea.— Resultats Scientifiques des Campagnes de la “Calypso” 4:207-217. 1963. South African Sipunculids and Echiu- roids from coastal waters. — Videnskabelige Meddelelser Danske fra Naturhistorisk Foren- ing 1 Kjobenhavn 125:101-146. Williams, J. A. 1972. Development of a rock bur- rowing sipunculid inhabiting stony coral.— American Zoologist 12(4):723. . 1977. Functional development in four species of the Sipuncula.— University of Hawaii, dis- sertation: 1-218. , & S. V. Margolis. 1974. Sipunculid burrows in coral reefs: evidence for chemical and me- chanical excavation. — Pacific Science 28(4):357— 359. Biology Department, Utica College of Syracuse University, Utica, New York 13502. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 767-772 A NEW SPECIES OF POLYCHAETE, SCOLELEPIS ANAKENAE (POLYCHAETA: SPIONIDAE) FROM EASTER ISLAND, SOUTH PACIFIC OCEAN, WITH ECOLOGICAL COMMENTS N. Rozbaczylo and J. C. Castilla Abstract.—A new species of Spionidae, Scolelepis anakenae from Easter Is- land, is described. Scolelepis anakenae, which closely resembles Scolelepis chi- lensis (Hartmann-Schréder) from the coast of Chile was the only polychaete species found in sand at Anakena beach during a macrofaunistic study. Com- ments about its distribution and population density in the intertidal zone are included. Easter Island, Rapa Nui in the native lan- guage, is one of the most isolated points in the southeast Pacific Ocean (27°07’S, 109°22’W) and the easternmost outpost of the Indo-Pacific Region. It lies 2230 miles from South America; the nearest land to the west 1s the uninhabited atoll Ducie Island, 1250 miles away, and Pitcairn Island of the Tuamotu Archipelago, 290 miles farther west. Its marine fauna is of the highest bio- geographic and evolutionary interest be- cause of the remarkable degree of endemism of its shore fauna at the specific level con- sidering that the island is not only very young (2.5 million years), but rather small, with an area of approximately 118 km? (New- man & Foster 1983). Recently, Castilla & Rozbaczylo (1987) prepared a bibliographic review on the lit- toral marine invertebrate fauna of Easter Island. The polychaete fauna is only par- tially known. The papers of Chamberlin (1919), Fauvel (1936), and Kohn & Lloyd (1973) include most of the records from the island. About 60 species of polychaetes are known to occur intertidally on Easter Is- land. Most of them inhabit rocky shores, since the coastline of the island is formed by volcanic rocks with just a few sandy beaches. Hitherto, no research has been done on the polychaete fauna of the island’s sandy beaches. On Anakena, one of the two prin- cipal sandy beaches on the island, the mac- rofauna was sampled quantitatively along a perpendicular sea transect. Among the spec- imens collected, a new and undescribed species of spionid of the genus Scolelepis was found. This increases the known species of spionid polychaetes on Easter Island to three, since Prionospio (Minuspio) cirrifera (Wirén) and Tripolydora spinosa Wood- wick, were the only species previously known (Rozbaczylo 1985). The new species is described and ecological comments on its distribution and population density are in- cluded. Type specimens have been deposited in the Museo Nacional de Historia Natural, Santiago (MNHN); Sala de Sistematica, La- boratorio de Zoologia, Pontificia Univer- sidad Catolica de Chile, Santiago (SSUC); the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM); and in the personal reference col- lection of the first author (NR). Materials and methods.—Samples were collected on 16 Oct 1982 at Anakena beach (27°04'S, 109°20’W) during an expedition sponsored by the Tinker Foundation and the Pontificia Universidad Catolica de Chile through the Project “The Chilean Oceanic Islands: Our Knowledge and Future Needs” 768 (Las Islas Oceanicas Chilenas: Nuestro Conocimiento y Necesidades Futuras). Two transects perpendicular to the sea front were laid on the eastern portion of the beach, 22 and 28 m long, extending from the top part of the berm down to the lower intertidal limit at low tide. The profile of the beach was determined according to the method of Emery (1961) and its slope was 1:20 m. Low tide (0.23 cm) occurred at 10.30 AM. Sam- pling stations were set every 4 meters along the transects. The samples, 0.1 m7’, were taken at each station using a square iron sampler of 31.6 cm, open at both ends, and introduced to a depth of 20 cm in the sand. The sand was removed with a spade and sieved through a 1.5 mm mesh screen. Sam- ples were washed in sea water and the mac- rofauna retained in the mesh was fixed in 10% formalin, then transferred to 70% al- cohol; polychaetes were preserved in 70% ethanol with glycerine added. No analysis of the sand was made. The density of poly- chaetes at each level was expressed as num- bers per square meter, by extrapolating from the data obtained with the sampler. Scan- ning Electron Microscope (SEM) observa- tions were made with a JEOL-25 II. Pho- tographs and drawings of the new species were executed by the first author. Figures were prepared by means of a drawing tube on a Wild M-5 microscope. Scolelepis anakenae, new species Figs. 1, 2 Material examined.—Easter Island, An- akena Bay, 27°04'S, 109°20'W, along a tran- sect in the intertidal zone, 16 Oct 1982, J. C. Castilla, coll. (aumerous specimens); 7 Mar 1984, J. C. Castilla, coll., holotype (MNHM 80009); paratype (MNHN 80010); 62 specimens (SSUC 6481); 41 specimens (USNM 104937). Diagnosis.— Body wider anteriorly, flat- tened dorsoventrally, with short segments; posterior region subcylindrical, with longer segments. Prostomium pointed anteriorly, extending posteriorly as cephalic ridge to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON anterior margin of setiger one; 2 pairs of eyes. Peristomium with low lateral wings partially surrounding prostomium and forming ventral ruffled surface; palps ex- tending posteriorly to setiger 20-30. Para- podia biramous throughout, with ciliated sensory organs between rami. Cirriform branchiae beginning on setiger 2, partially fused with notopodial lamellae, with sub- terminal enlargements forming knobbed tips. Anterior neuropodial lamellae entire and rounded up to setiger 34—40; in poste- rior segments completely divided into elon- gated, thickened interramal lamellae and conical ventral lobe. Anterior notosetae and neurosetae all winged capillaries; bidentate neuropodial hooded hooks from setiger 35— 37 to end of body; without notopodial hooks. Pygidium forming thick cushionlike pad. Description.—Color in alcohol light tan or pale brownish. One of largest specimens 36 mm long, 2 mm wide, with 120 setigers; one of smallest individuals 18 mm long, 1.2 mm wide, with 86 setigers; maximum body width reached approximately at setiger 20. Body distinctly divided into 2 regions: fu- siform anterior region with segments rather flattened and rectangular in cross section; posterior region of numerous more or less rounded segments; segments from anterior region shorter than those from posterior re- gion; change from one region to other marked by appearance of neuropodial hooded hooks and modifications in para- podial structure and setal distribution. Prostomium pointed anteriorly, wider subterminally, and extending posteriorly as cephalic ridge to anterior margin of setiger 1 (Fig. la). Two pairs of dark eyes in nearly straight transverse row, located on each side of thickened part of prostomial ridge, just behind slight depression at beginning of ridge; dorsal pair more or less rounded, ven- tral pair oval. Peristomium well developed, with low lateral wings, partially encompassing pro- stomium and forming ventral ruffled sur- face (Fig. 1b). Short, eversible, somewhat VOLUME 101, NUMBER 4 769 Fig. 1. Scolelepis anakenae: a, Anterior end, dorsal view, palps omitted; b, Same, lateral view; c, Setiger 1, anterior view; d, Setiger 2, anterior view; e, Capillary notoseta from anterior row of setiger 7 (third from top): f, Setiger 3, anterior view; g, Setiger 14, anterior view; h, Setiger 32, anterior view; i, Setiger 36, posterior view; j, Setiger 76, posterior view; k, Setiger 114, posterior view; 1, Neuropodial hooded hook from posterior para- podium, lateral view; m, Same, frontal view; n, Posterior end of body, dorsal view. Scale under b applies to a— d, f-k, n. lobulated proboscideal region. Palps thick- ened basally, tapering gradually, with con- spicuous basal sheath or palpophore; when extended posteriorly palps reaching setiger 20-30. Parapodia biramous throughout. First se- tiger smallest, although well developed, with lamellae of almost same size; notopodial lamellae more or less triangular while neu- ropodial more rounded (Fig. Ic). All para- 770 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VLE 2 Fig. 2. Scolelepis anakenae, scanning electron micrographs: a, Anterior parapodium showing ciliated sensory organs between rami (arrow); b, Anterior end, dorsal view, showing ciliated bands across dorsum (arrow). VOLUME 101, NUMBER 4 podia with ciliated sensory organs between rami, more conspicuous in specimens pre- served in formalin than when transferred to alcohol (Fig. 2a, arrow). Branchiae beginning on setiger 2, fused to notopodial lamellae, being free only at their tips, with distinct subterminal enlarge- ments forming knobbed tips (Fig. 1d). Bran- chial pairs connected across dorsum by nar- row ciliated ridge, rather difficult to see with light microscopy but evident with SEM (Fig. 2b, arrow). Notopodial lamellae fused to branchiae except distally in more anterior setigers, becoming more separated poste- riorly and retaining only basal fusions in far posterior setigers (Fig. 1k); free margin of notopodial lamellae folded. Anterior notosetae and neurosetae all winged capillaries, similar in structure, with shafts bearing fine granulations along inter- nal striations and transparent sheaths with fine striations (Fig. le). Notosetae arranged in 2 rows in anterior setigers and single rows more posteriorly. Each row in anterior se- tigers forming fan-shaped fascicle of 13-15 setae per ramus, with 3-4 longer ones in superior position; setae of each row almost of same length, being shorter in anterior row than in posterior row. Median and posterior setigers with only 4—7 notosetae, with striat- ed shafts but without granulations. Postsetal neuropodial lamellae more or less rounded on first 5—6 setigers (Fig. 1c— f), being just slightly longer than wide, in- creasing in size and becoming broadly rounded in following setigers (Fig. 1g). Neu- ropodial lamellae entire up to setigers 31- 40, then developing slight notch (Fig. 1h) and becoming deeper in following few se- tigers, and then becoming completely di- vided into rounded dorsal and conical ven- tral lobes (Fig. 11). Neuropodial lobes becoming more prolonged vertically and further separated forming elongated thick- ened interramal lamellae and small conical ventral lobe (Fig. 1j, k). Neurosetae on an- terior setigers all capillaries in 2 transverse rows, similar in arrangement and structure 771 to notosetae, numbering about 12 setae per row with 3-4 slender setae, slightly sepa- rated from remainder, appearing like small ventral fascicle. Neuropodial hooded hooks beginning on setiger 35-37 and continuing up to end of body. Neuropodial fascicles in single rows, typically with 5—8 hooded hooks and 2-4 capillary setae in dorsal and ventral position, dorsal ones with striated shafts but without granulations, ventral ones with shafts coarsely granulated. Hooded hooks bidentate, with apical tooth formed by 2 incompletely separated teeth fused by ridge (Fig. 1l—m). Pygidium with ventral cushion (Fig. 1n), with single achaetous preanal segment. Anus dorsal, surrounded by crenulate margin. Distribution. — Known only from type lo- cality, Anakena Bay (27°04’S, 109°20’W), north coast of Easter Island, intertidal zone, sandy beach. Etymology.—The specific epithet is taken from the type locality, Anakena Bay where, according to tradition, King Hotu Matu’a and his followers from some region located in the east or northeast of Polynesia arrived as settlers on the island. Remarks. —Scolelepis anakenae is closely related to S. chilensis (Hartmann-Schroder, 1962) from the coast of Chile in that the apical teeth of the hooded hooks are fused by a ridge, as clearly shown by Blake (1983, fig. 2K—L). It differs from S. chilensis in hav- ing conspicuous ciliated sensory organs be- tween rami in all parapodia and by the total absence of notopodial hooded hooks. In S. chilensis, neuropodial hooded hooks begin on setiger 25—32, while in S. anakenae they begin more posteriorly, around setiger 35- 37. Further, the SEM reveals significant dif- ferences in the ciliated bands across the dor- sum between S. anakenae, which are nar- row, and S. chilensis with wide and dense bands. In S. anakenae, the branchiae show conspicuous subterminal enlargements forming knobbed tips; this character has not been mentioned for S. chilensis either in Hartmann-Schroder’s original description V2 or by Blake (1983) in a recent revision of the species. Ecological notes.—Scolelepis anakenae was the only polychaete found in the sandy beach transects at Anakena during this study. According to its macrofaunal components, the beach can be characterized by three fringes. The upper fringe, extending 8-10 m down from the berm, is very poor in mac- rofauna with only a single species of isopod and S. anakenae in very low densities, up to 10/m? towards the lower portion of the fringe. The middle fringe extends down for 8 m more and S. anakenae is the only species present, with maximum densities ranging between 460-180/m/7 at the lower border, and 260-—70/m? at the upper border. The lower fringe encompasses the infralittoral sector of the beach, characterized by several species, including gastropods, sipunculans, and hippidean crustaceans; S. anakenae was found in low densities of about 20/m7?. Acknowledgments The authors acknowledge financial sup- port by the Tinker Foundation and Ponti- ficia Universidad Catolica de Chile, through the Project ““The Chilean Oceanic Islands: Our Knowledge and Future Needs” (Las Is- las Oceanicas Chilenas: Nuestro Conoci- miento y Necesidades Futuras). Marco Mendez helped to collect specimens of Sco- lelepis chilensis (Hartmann-Schroder) from the type locality in Huasco, Chile. We thank Dr. Marian H. Pettibone for reading and making numerous helpful suggestions on the manuscript. Literature Cited Blake, J. A. 1983. Polychaetes of the family Spion- idae from South America, Antarctica and ad- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON jacent seas and islands. Pp. 205-287 in L. Kor- nicker, ed., Biology of the Antarctic Seas XII. Antarctic Research Series, volume 39. Ameri- can Geophysical Union, Washington, D.C. Castilla, J. C., & N. Rozbaczylo. 1987. Invertebrados marinos de Isla de Pascua y Sala y Gomez. Pp. 181-215 in J. C. Castilla, ed., Islas Oceanicas Chilenas: Conocimiento Cientifico y Necesi- dades de Investigacion Ediciones Universidad Catolica de Chile. Chamberlin, R. V. 1919. The Annelida Polychaeta. — Memoirs of the Museum of Comparative Zo- ology at Harvard College 48:1-493. Emery, K. O. 1961. A simple method of measuring beach profiles.— Limnology and Oceanography 6:90-93. Fauvel, P. 1936. Sur quelques annélides polychétes de I’Ile de Paques.—Bulletin du Muséum Na- tional D’Histoire Naturelle, Série 2, 8:257—259. Hartmann-Schroder, G. 1962. Die Polychaeten des Eulitorals. Pp. 57-167 in G. Hartmann-Schro- der & G. Hartmann, eds., Zur Kenntnis des Eu- litorals der chilenischen Pazifikkiiste und der argentinischen Kite Siidpatagoniens unter be- sonderer Bericksichtigung der Polychaeten und Ostracoden.— Mitteilungen aus dem Hambur- gischen Zoologischen Museum und Institut, Supplement 60. Kohn, A. J., & M. C. Lloyd. 1973. Marine polychaete annelids of Easter Island.—Internationale Re- vue der Gesamten Hydrobiologie 58(5):69 1-712. Newman, W. A., & B. A. Foster. 1983. The Rapanu- aian faunal district (Easter and Sala y Gomez): In search of ancient archipelagos. — Bulletin of Marine Science 33(3):633-644. Rozbaczylo, N. 1985. Los anélidos poliquetos de Chile. Indice sinonimico y distribucion geo- grafica de especies. — Monografias Biologicas 3: 1-284. Departamento de Ecologia, Facultad de Ciencias Biologicas, Pontificia Universidad Catolica de Chile, Casilla 114-D, Santiago, Chile. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 773-783 DESCRIPTION OF GUARANIDRILUS OREGONENSIS (OLIGOCHAETA: ENCHYTRAEIDAE) FROM NORTH AMERICA, WITH ADDITIONAL COMMENTS ON THE GENUS Kathryn A. Coates and Robert J. Diaz Abstract. —Guaranidrilus oregonensis, n. sp., is described from specimens found in the Columbia River near Miller Sands, Oregon, and its relationships to other nominal species of Guaranidrilus are briefly considered. This is the first record of the genus from North America. Examination of type material of G. glandulosus, type species of the genus, and of G. rarus, showed that the former does not have esophageal appendages in VI, like G. oregonensis, and that the former two species are quite distinct. Guaranidrilus sawayai is a prob- able synonym of G. rarus. The new species is closely related to G. glandulosus, and two other species without esophageal appendages. A comparative table allows quick distinction of species of Guaranidrilus without oesophageal ap- pendages. The genus Guaranidrilus was erected by Cernosvitov (1937a, b) for three enchy- traeid species from Argentina, G. glandu- losus, G. rarus and G. fridericoides, the latter of which is a nomen nudum, and, provi- sionally, for G. columbianus (Michaelsen, 1913) from Colombia. Brinkhurst & Jamie- son (1971) subsequently designated G. glan- dulosus as the type species. Some diagnostic characteristics of Guaranidrilus noted by Cernosvitov (1937a, b) were occurrence of two setae in each setal bundle, compact ap- pendages on the dorsal side of the esophagus in VI, a large pair of intestinal diverticula originating at the transition from oesopha- gus to intestine, coiled vasa deferentia, and adiverticulate spermathecae not commu- nicating with the intestine. Guaranidrilus lamottei was described by Omodeo (1958) from Ivory Coast, but none of the nine specimens examined was ma- ture. Omodeo expressed some ambivalence about whether the specimens should be at- tributed to Guaranidrilus or to a new genus. The systematic position of this species is still uncertain (Righi 1973, Healy 1979) and no type material is locatable. Seven other species of Guaranidrilus have since been described from Brazil: G. sa- wayai Righi, 1973, G. mboi Righi, 1975, G. athecatus Christoffersen, 1977, G. atlanti- cus Christoffersen, 1977, G. joanae Chris- toffersen, 1977, G. oiepe Righi, 1974, and G. finni Christoffersen, 1977. The latter two species lack esophageal appendages in VI. In the most recent review of Guaranidrilus, Healy (1979) described G. europeus, the first species from Europe, which also is without esophageal appendages in VI. She also de- scribed G. cernosvitovi Healy, 1979, based on undescribed material collected by Cer- nosvitov (1937b) (see G. fridericoides no- men nudum). This review did not cover the recently described South American species. Healy (1979) examined Cernosvitov’s original material of Guaranidrilus, located at the British Museum (Natural History) (BMNH), and syntypes of G. columbianus in the Zoologisches Museum, Hamburg. Af- ter her examinations, Healy questioned the 774 validity of G. rarus, suggesting that it could not be distinguished reliably from G. glan- dulosus. The quality of the syntype material of G. columbianus did not allow Healy to confirm all recorded anatomical details, but characteristics such as the presence of bi- lobed gut diverticula and form of the ne- phridia indicated to her that it belongs in Guaranidrilus, and, probably, is distinct from the other species, although presently not fully described. In a phylogenetic study (Coates 1987a) it was found that both bisetate setal bundles and free adiverticulate spermathecae are plesiomorphies of Enchytraeidae. As well, coiled vasa deferentia is a synapomorphy of all or most of Enchytraeidae, although lost in more recent lineages (Coates 1987b, in press a). Within Guaranidrilus, gut diver- ticula have been lost one or more times and extra-lineal hybridization possibly compli- cates the genealogy of the lineage (Coates, in press b). Recent collections made in the Columbia River near Miller Sands, Oregon, yielded the first specimens of Guaranidrilus for North America. Material and Methods Type specimens of Guaranidrilus glan- dulosus, G. rarus, G. europeus, and G. cer- nosvitovi were borrowed from the BMNH. Material of the new species was received from benthic collections made by the United States Army Corps of Engineers, from a field study to test the feasibility of developing marsh habitats on dredged material in the Columbia River estuary, Oregon (see Clair- ain et al. 1978 for details). All of the new material was stained in alcoholic borax car- mine, dehydrated through an ethanol to xy- lene series then mounted whole in Canada balsam. Type material of the new species has been deposited at the National Museum of Natural History, Smithsonian Institution (USNM) and at the Royal Ontario Museum, Department of Invertebrate Zoology (ROMIZ). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Guaranidrilus Cernosvitov, 1937 Guaranidrilus Cernosvitov, 1937a:148- 149.— 1937b:282—283. Healy, 1979:7-8. Type species.—Guaranidrilus glandulo- sus Cernosvitov, 1937. Discussion.—Nominal species of Guar- anidrilus are small to medium-sized terres- trial and freshwater enchytraeids with sim- ple-pointed, anodulate, straight or slightly curved setae. These occur in 4 bisetate bun- dles per segment, except laterally in XII of mature specimens and rarely in some pre- clitellar segments. The head pore is anterior on the prostomium, and other coelomic pores are absent. Secondary pharyngeal gland lobes may be present. The species have compact nephridia with large anteseptal parts including some of the efferent canal. Their seminal vesicles are unpaired and dorsal, or apparently absent, and vasa defer- entia are usually coiled. Vasa deferentia ter- minate in very small or no penial bulbs. Spermathecae, if present, are paired, free, with thick-walled ectal ducts and usually a sperm-containing dilation in V. The saccate ampulla may extend through several seg- ments. The genus is primarily characterized by a pair of hollow, dorso-lateral gut diver- ticula originating near 7/8 or, rarely, origi- nating in X. The diverticula sometimes are constricted, forming anterior and posterior lobes. These diverticula have been lost in some species. Overall, the nominal species of Guaran- idrilus are relatively underived within the lineage of Enchytraeidae. Only the presence of gut diverticula has been determined to be a synapomorphy of the genus (Coates, in press b). Gut diverticula at the esophago- intestinal transition (usually about VIII) also occur in the enchytraeid genera Buchholzia, Enchylea, Punahenlea, Henlea, and Aspi- dodrilus. The evolutionary origins of their diverticula are apparently all independent of those in nominal species of Guaranidrilus and, at least, some diverticular forms are reported to be distinguishable by their gen- VOLUME 101, NUMBER 4 eral appearances (Healy 1979). One group of Guaranidrilus is further characterized by esophageal appendages in VI. The species of this group also have relatively large sem- inal vesicles, whereas these are apparently absent or small in other nominal species of Guaranidrilus. Specimens of several nominal species of Guaranidrilus with esophageal appendages, namely G. atlanticus, G. athecatus, G. joan- ae, and G. mboi, were not available (G. Righi & M. L. Christoffersen, pers. comm.) during the period of this research. Examination of specimens is required in order to complete a revision of the genus both because of taxo- nomic problems (see below) and descriptive inconsistencies. Each of these species does have some easily recognized specific char- acteristic: G. atlanticus has single esopha- geal diverticula in VI; G. athecatus lacks spermathecae; G. joanae has setae with very broad ental ends in anteriormost and pos- teriormost segments; and G. mboi has a pair of gut diverticula in X, rather than at 7/8 and is without pharyngeal glands at 6/7. The summarized status of the nominal species of Guaranidrilus as a result of the conclusions drawn by Righi (1973), Chris- toffersen (1977), Healy (1979), Coates (1987a, in press a, b), and the taxonomic studies herein 1s: group 1: without esophageal appendages, with small or no seminal vesicles: G. glandulosus Cernosvitov, 1937 G. oiepe Righi, 1974 (?=G. glandulo- SUS) G. europeus Healy, 1979 G. oregonensis new species; group 2: with esophageal appendages, in VI, rarely absent; seminal vesicles large, rarely absent: G. rarus Cernosvitov, 1937 (G. sawayai Righi, 1973 = G. rarus) (see fol- lowing) G. cernosvitovi Healy, 1979 G. mboi Righi, 1975 G. joanae Christoffersen, 1977 WHS G. athecatus Christoffersen, 1977 G. atlanticus Christoffersen, 1977, with several described forms G. finni Christoffersen, 1977; other species: G. columbianus (Michaelsen, 1913) sp. dubia, incompletely described G. lamottei Omodeo, 1958 gen. dubia, incompletely described, based on immature specimens. New Species Description and Remarks on Other Species A. Species without esophageal appendages in VI Guaranidrilus oregonensis, new species ais seale2 Material examined.—Columbia River, Miller Sands, Oregon, about 46°15.00’'N at 123°37.50'W. Collected VIII.1980 by E. J. Clairain and C. Newling. Holotype (USNM 118242) and 6 paratypes (USNM 118243- 118245; and ROMIZ 11226-1228). Etymology. —Named for the type locali- ty. Description. — Fixed, whole-mounted specimens 3.0 to 4.5 mm long, 0.19 to 0.24 mm [xX = 0.22, s = 0.02, n = 5] wide at clitellum (specimens slightly compressed) (Fig. 1A); with 26 to 34 segments. Head pore dorsal on anterior third of prostomi- um. Setae 2 per bundle, absent entirely from XII, straight with ental hook. Preclitellar ventral setae 24 to 33 um [X = 29,s =4,n = 4] long (Fig. 1B, b); setae from clitellum to posterior 4 or 5 segments 28 to 41 um [xX = 35, s = 5, n = 5] long; posterior setae 52 to 78 um [X = 62, s = 9, n = 6] long (Fig. 1B, a); more or less equal laterally and ven- trally. Clitellum (Fig. 1A, c) extending over ¥> XII to % XIII. Gland distribution: irreg- ular dorsally, disrupted by large hyaline cells; in transverse rows ventro-laterally; reduced but complete ventrally between male pores. Cutaneous glands inconspicuous, in | to 4 regular transverse rows per segment, if only 776 Fig. 1. ments through segment XVI; B, Setae from posterior segments, XX VIII or X XIX, and b. anterior segments III and IV; C, Nephridium at 7/8; D, Sperm funnels, vasa deferentia and male pore, ventro-lateral view; E, Spermatheca, dorso-lateral view; Abbreviations: c, clitellum; d, gut diverticulum; f, female pore; m, male pore; p, pharyngeal glands; s, seminal vesicle; v, dorsal blood vessel. 1 row, mid-segmental. Spermathecal pore small, in 4/5 in line with lateral setae; male pore small, in line with ventral setae, on a small papilla. With sensory or adhesive pa- pillae on pygidium. Jnternal.— Posterior margin of brain deeply incised. Dorsal an- terior blood vessel bifurcating in prosto- mium, originating in X to XII (Fig. 1A, v). Pharngeal glands (Fig. 1A, p) united dor- sally at 4/5, 5/6 and 6/7; ventral lobes in V and VI; small, secondary glands antero-ven- tral in V and VI. Bilobed gut diverticula (Fig. 1A, d) communicating with gut in pos- terior half of VII, posterior lobes of diver- ticula extending into VIII; inner walls of diverticula ciliate. Without other gut diver- ticula or appendages. Chloragocytes golden- brown, most obvious in preclitellar seg- ments. Lymphocytes nucleate, egg-shaped. Preclitellar nephridia on 6/7 through 8/9 (Fig. 1C); preseptal part half or more of en- tire nephridium; efferent duct terminal; in- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A-E, Guaranidrilus oregonensis, n. sp., whole-mounted specimens. A, Lateral view of anterior seg- terstitial tissue well-developed. Postclitellar nephridia of same shape as preclitellar, usu- ally only | or 2 pairs. Seminal vesicle small (Fig. 1A, s) extending into X. Testes com- pact, paired on posterior of 10/11. Sperm funnels cylindrical, canal eccentric; granu- lar; funnels about 61 wm [n = 3, s = 22] long and 42 um [n = 3, s = 7] wide; collar subequal funnel width (Fig. 1D). Vasa de- ferentia long, narrow, coiled in XII (Fig. 1D); with very slight widening at penial pore. Male pore simple (Fig. 1A, m), without pen- ial bulb, but with glandular, slightly en- larged, epidermal cells adjacent. Ovaries paired ventrally on posterior of 11/12. Fe- male funnel (Fig. 1A, f) on 12/13, incon- spicuous; pore in or just posterior to 12/13. Eggs extending posteriad into XV, in an egg sac, usually only 1 egg large. Spermathecal ectal ducts (Fig. 1E) thick-walled, without associated glands, with a small, sperm-con- taining widening in V; ampulla thin-walled, VOLUME 101, NUMBER 4 oregonensis n.sp. glandulosus Cernos. body length (mm) 3.0-4.5 3-7 number of segments 26-34 = setal length anterior 24-33 =33 (pm) posterior 52-78 +54 gut diverticula bilobed bilobed pharyngeal glands —_* Ge or So, lobes 8 spermathecae scale equals 7/8 0.1 mm vil seminal vesicle anterior absent ia oiepe Righi europeus Healy finni Christoffersen 3.5-8.5 3-6 8-14 31-34 23-29 41-56 = 20 24-32 =57 =37 48-61 unilobed bilobed bilobed > i} Od Vi \) absent small/absent anterior & posterior Fig. 2. Tabular comparison of Guaranidrilus oregonensis n. sp. and the 4 other nominal species of Guar- anidrilus without esophageal appendages. irregular shaped, saclike, extending to VI or VII; sperm free in ampulla. Habitat and distribution.—The Miller Sands area in the Columbia River, Oregon, is an active site for the disposal of dredged material. Snag Isiand, on which the oligo- chaetes were found, was a natural marsh used as a reference area. The tidal range in the freshwater marsh habitat of the worms was 1.9 m. Guaranidrilus specimens were collected in the high marsh where the sed- iments were a matrix of rhizomes and sandy mud (61% silt-clay) with 4.5% volatile sol- ids. The dominant plants were spike-rush (Eleocharis palustris), Lyngby’s sedge (Car- ex lyngbei), and tufted hairgrass (Des- champsia caespiotosa) (Clairain et al. 1978). The benthos in this habitat was domi- nated numerically by oligochaetes, 93% of the total, with 84% of them being tubificids, 14% enchytraeids, and 2% lumbriculids. Other dominant taxa were the amphipod Anisogammarus confervicolus, sphaeriid clam, Corbicula fluminea, and chironomid and chrysomelid insect larvae (Clairain et al. 1978). In general the oligochaete fauna of the Miller Sands area is diverse with many unusual tubificid species (Brinkhurst & Diaz 1985). Remarks.—A few characteristics distin- guish Guaranidrilus oregonensis and G. glandulosus. Small differences exist between the species in location of the origin of the dorsal blood vessel, in distribution of cli- tellar gland cells, and in form of pharyngeal glands (Fig. 2). Guaranidrilus glandulosus also is reported to lack a seminal vesicle whereas one is present, although small, in G. oregonensis. A further reason for recognition here of a distinct species is the apparent geographic separation of the respective populations of 778 100 pm Fig. 3. Guaranidrilus glandulosus, BMNH 1949.3.1.951, dorsal sagittal section through V and VI showing ectal ducts of spermathecae. Abbreviations: b, body wall; e, spermathecal ectal ducts; p, pharyngeal glands (dorsal lobe at 5/6); se, septum. Guaranidrilus glandulosus and G. oregon- ensis. Broad dispersion of some earthworm species by human activities are reported (Sims & Gerard 1985), but there is no ex- isting evidence for this having occurred with Guaranidrilus species. There are few rec- ords of the genus outside of South America (Europe in Healy 1979; and Florida, Healy, pers. comm.) and few of the species are known to have large ranges of distribution. That of G. rarus, including Argentinian and Brazilian localities, is the broadest if the proposed synonymy of G. sawayai is valid. It is of concern that descriptions of both Guaranidrilus oregonensis and G. glandu- losus were based on small numbers of spec- imens and that both South and North America are poorly explored for enchy- traeids. As well, there is no clear indication of the conditions required for speciation within enchytraeid lineages (Coates 1987b). Long periods of geographic separation need not guarantee the development of repro- ductive isolation mechanisms nor evolution in unique directions, unconstrained by ear- lier, shared historical events. Guaranidrilus oregonensis is compared to G. glandulosus and the three other nominal PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON species of Guaranidrilus without esophageal appendages, G. europeus, G. oiepe, and G. finii, in Figure 2. Guaranidrilus glandulosus Cernosvitov, 1937 Figs. 2, 3 Guaranidrilus glandulosus Cernosvitov, 1937a:149-151, figs. 1-9.— 1937b:282.— Christoffersen, 1977:table 1.—Healy, 1979:8, table 1. Material examined. —Syntypes, BMNH 1949.3.1.949-950, transverse and tangen- tial sections; BMNH 1949.3.1.951-952, sagittal sections; BMNH 1949.3.1.953, whole-mounted specimens; collected at Loreto, Misiones, Argentina, 18.XI.1931. Habitat and distribution. —In rich humic soul and in the detritus along a small stream. Loreto, near Santa Ana, Misiones, Argen- tina. Remarks.—The specimens or parts of specimens catalogued as BMNH 1949.3.1.952 and 953 were not suitable for observations of internal structures. In the complete description Guaranidri- lus glandulosus, Cernosvitov (1937a, p. 151) stated that compact esophageal appendages ““peptonéphridies” are located dorso-later- ally in VI. Such structures were not found in any of BMNH 1949.3.1.949-951. Even though the ducts of the spermathecae follow a circuitous route into segment VI (Fig. 3, e) sections of these would not be confused with esophageal appendages as the cellular struc- tures of these organs are distinctive (see Figs. 3, 4). Cernosvitov (1937b) recognized two oth- er species of Guaranidrilus in the material obtained from Argentina, but only Guar- anidrilus rarus was described by him (Cer- nosvitov 1937a) at the same time as G. glandulosus; G. cernosvitovi was not de- scribed until some years later by Healy (1979). Both G. rarus and G. cernosvitovi are reported to have esophageal append- ages, as did all the type specimens of them VOLUME 101, NUMBER 4 779 Fig. 4. A-B, Guaranidrilus rarus, BMNH 1949.3.1.955. A, Dorsal sagittal section of segments I through VII showing esophageal appendages; B, Enlargement of esophageal appendages, sagittal section ventral to dorsal blood vessel. Abbreviations: a, esophageal appendages; d, gut diverticulum; e, spermathecal ectal ducts; 0, esophagus; p, pharyngeal glands; v, dorsal blood vessel. examined here (see below). Cernosvitov (1937a, fig. 9, pt) clearly showed esophageal appendages in his illustration of G. glan- dulosus and it seems possible that he con- fused two species, but the type material that is still usable is of one species, without such appendages. Guaranidrilus oiepe Righi, 1974 Fig: 2 Guaranidrilus oiepe Righi, 1974:140-141, figs. 32-40.—1975:144.—1981:427- 428.—Christoffersen, 1977:table 1. Habitat and distribution. —In decompos- ing wood covered with grasses, in a tem- porary pond. Mato Grosso, Rio de Janeiro, and Minas Gerais, Brazil. Remarks.—Specimens of Guaranidrilus oiepe with sperm in the spermathecae have never been found, possibly indicating that the specimens seen were not mature or that the species is parthenogenetic. This species was distinguished from G. glandulosus by the shape and distribution of pharyngeal glands and by the shape of the intestinal diverticula (Righi 1974) (Fig. 2). Good rec- ords of intraspecific variation for either of these characters are not available. Re-ex- amination of G. olepe is required to further substantiate its specific distinction from G. glandulosus. Type and other material is not 780 currently available for loan outside the country of its origin (Righi, pers. comm.). Guaranidrilus europeus Healy, 1979 Fig. 2 Guaranidrilus europeus Healy, 1979:1 1-13, figs. 2a-i, 3a-d. Material examined. — Paratype, BMNH 1978.39.2, whole-mounted; collected at le Corneau near Arcachon, France, 21.X.1977, by B. Healy. Habitat and distribution. —Wet, marshy pasture at the edge of a small pond, pasture soil, and in moist sandy loam. Near Arca- chon, southwest France and Roncesvalles, Spanish Pyrenees. Remarks. —For the purpose of compari- son with the South American species, some characteristics are noted from the type ma- terial in addition to the description made by Healy (1979). The setae of anterior seg- ments were notably shorter than the setae of posteriormost segments (Fig. 2); nephrid- ia were present from 6/7; and secondary pharyngeal glands were present in V and VI (Fig. 2). No scattered glands on the wall of the spermathecal ectal duct, as reported by Healy, were obvious in the fixed material but cells of the walls of the duct were coarse- ly granular. Gland cells of the clitellum were irregularly interspersed with hyaline cells in dorso-lateral regions. Penial bulbs were no more than glandular areas in the epidermis surrounding the male pores. Guaranidrilus europeus was described as lacking a seminal vesicle but evidence of a very small seminal vesicle (Fig. 2) was found in the one specimen examined here. The form of the pharyngeal glands, spermathe- ca, and possibly the gut diverticula still dis- tinguish it from the other species without esophageal appendages (Fig. 2). Guaranidrilus finni Christoffersen, 1977 Fig. 2 Guaranidrilus finni Christoffersen, 1977: 188-190, figs. 1-12, table 1. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Habitat and distribution.—Banks of a small brook. Evangelista de Souza, Serra do Mar range, Sao Paulo, Brazil. Remarks.—This species is distinctively larger than the other species without esoph- ageal appendages (Fig. 2). The sperm fun- nels and spermathecae are correspondingly proportioned. Guaranidrilus finni is also unique in having encapsulated sperm bun- dles in the spermathecae (Fig. 2). Phylogenetic analyses (Coates 198b, in press) indicate that Guaranidrilus finni is closely related to the larger species of Guar- anidrilus with esophageal appendages rather than to other nominal species of Guarani- drilus lacking such appendages. In G. finni the absence of esophageal appendages is a derived character state. The opinion of Christoffersen (1977) that G. finni and G. atlanticus, in particular, had a close affinity also 1s supported. B. Some species with esophageal appendages in VI Guaranidrilus rarus Cernosvitov, 1937 Fig. 4 Guaranidrilus rarus Cernosvitov, 1937a: 151-153, figs. 10-15.—1937b:282.— Christoffersen, 1977:table 1.—Healy, 1979:14—-15, table 1. ?Guaranidrilus sawayai Righi, 1973:470- 472, figs. 1-9.—1975 [as G. rarus]:143- 144.—1981:428, figs. 1-2.—Christoffer- sen, 1977:table 1. Material examined. —Syntypes, BMNH 1949.3.1.955, tangentially sectioned, BMNH 1949.3.1.954, whole-mounted; col- lected at Loreto, Misiones, Argentina, 27.XI1.1931, by L. Cernosvitov. Habitat and distribution. — Under the bark of an old tree, in a tree-growing bromeliad; in a periodically flooded stream bed near Rio Parana; near Rio Capiroa. Loreto near Santa Ana, Misione, Argentina; Serra do Cipo, Minas Gerais, and Bataguassu, Mato Grosso, Brazil. Remarks. —The whole-mounted type VOLUME 101, NUMBER 4 781 Fig. 5. Guaranidrilus cernosvitovi, BMNH 1949.3.1.960.1A, sagittal lateral section showing prostomium to segment VII, with expansion of ectal duct of one spermatheca and one esophageal appendage. Abbreviations: a, esophageal appendages; e, spermathecal ectal ducts. BMNH 1949.3.1.954 was unidentifiable and, if an oligochaete, immature. The original description of Guaranidrilus rarus (Cernosvitov 1937a) indicated that the esophageal appendages in VI were ventral. Examination of the type material has shown this to be incorrect, and the esophageal ap- pendages are dorso-lateral, as usual for the genus (Fig. 4A, B, a). Righi (1973, 1981) already may have recognized this error as he did not indicate any difference between G. rarus and G. sawayai for that character. Righi (1973, 1975, 1981) has compared Guaranidrilus rarus to G. sawayai in detail. He most recently (1981) suggested that some of the characters he originally used to dis- tinguish these, in particular the form of the brain and the form of the intestinal diver- ticula, were not good specific characters as presently determined. Righi (1973) de- scribed G. sawayai as possessing small, rounded penial bulbs. Bulbs were not ap- parent in his figures of the species, although a few small structures once were indicated around the male pore (Righi 1973, fig. 9). Penial gland development in sexually ma- ture specimens may show enough variation in the species so that the state recorded for G. sawayai is not different from their ab- sence as recorded for G. rarus. As Healy (1979) pointed out, the description of G. rarus was based on very few specimens. Other characters that Righi (1981) iden- tified as having states distinguishing Guar- anidrilus rarus from G. sawayai were sec- ondary pharyngeal glands and the position of the most anterior pair of nephridia. As G. sawayai, the sectioned type material of G. rarus has secondary glands in V and VI and the first pair of nephridia on 6/7. Righi also indicated that the intestinal diverticula in G. sawayai extended across the dorsal surface of the gut. Such a dorsal continuity could not be seen clearly in any available material of G. rarus (Fig. 4A, d) but this too could fall within the range of intraspecific variation recognized for other species of Guaranidrilus (Christoffersen 1977). Observations of available material strongly suggest that Guaranidrilus sawayai is a junior synonym of G. rarus. Examina- tion of type material of G. sawayai ulti- mately is required to decide the status of the synonymy. 782 Data available in the literature regarding the lengths of setae (Righi 1973) for Guar- anidrilus sawayai, 3 wm anteriorly and 15 um posteriorly, are very likely incorrect be- cause the cuticle of the body wall is often thicker in an enchytraeid than the former measurement. Guaranidrilus cernosvitovi Healy, 1979 Fig. 5 Guaranidrilus fridericoides Cernosvitov, 1937b:282, nomen nudum. Guaranidrilus cernosvitovi Healy, 1979:10— 11, fig. la-f, table 1. Material examined. —Paratypes, BMNH 1949.3.1.960a—, 2 sagitally sectioned spec- imens on three slides, BMNH 1949.3.1.957/ 8, whole-mounted specimen; collected at Loreto, Misiones, Argentina, 12.VI.1932. Remarks.—The accession numbers for the holotype and other paratypes are, re- spectively, BMNH_ 1949.3.1.956 and 1949.3.1.959, 96la—b and 962-967 (E. G. Easton, pers. comm.). Accession numbers for this species were not correctly indicated in Healy (1979). Guaranidrilus cernosvitovi is one of the larger nominal species of Guaranidrilus, be- longing in a lineage with G. finni and G. atlanticus and several other South Ameri- can species (Coates, in press D). It shares with these the character of ventral epider- mal copulatory pads or papillae, occurring primarily in the clitellar region. As other nominal species of Guaranidrilus and “‘low- er enchytraeids,” G. cernosvitovi retains a sperm-containing expansion on each of the spermathecal ectal ducts just ental to the spermathecal pore (Fig. 5, e). These appar- ently were overlooked by Healy (1979). The esophageal appendages of this species are associated with large blood sinuses (Fig. 5, a). Acknowledgments We are indebted to Buddy Clairain and Charlie Newling of the U.S. Army Corps of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Engineers Waterways Station, Vicksburg, Mississippi, for collection of the material and permission to publish this report. This paper is Contribution No. 1435 of the Viriginia Institute of Marine Science. Literature Cited Brinkhurst, R. O., & R. J. Diaz. 1985. Isochaetides columbiensis, new species (Oligochaeta: Tubi- ficidae) from the Columbia River, Oregon.— Proceedings of the Biological Society of Wash- ington 98:949-953. Brinkhurst, R. O., & B.G. M. Jamieson. 1971. Aquatic Oligochaeta of the World. Edinburgh, Oliver and Boyd. 860 pp. Cernosvitov, L. 1937a. Notes sur les Oligochaeta (Naididées et Enchytraeidées) de l’Argentine. — Anales del Museo Argentino de Ciéncias Nat- urales 39:135-157. . 1937b. System der Enchytraeiden.— Bulletin de l’Association pour les Russe Recherches Scientifiques 4 Prague 5:262-295. Christoffersen, M. L. 1977. New species of Guar- anidrilus Cernosv. (Enchytraeidae, Oligochaeta) from Serra do Mar, Sao Paulo, Brazil. — Studies on Neotropical Fauna and Environment 12:187— 206. Clairain, E. J., R. A. Cole, R. J. Diaz, A. W. Ford, R. T. Huffman, L. J. Hunt, & B. R. Wells. 1978. Habitat development field investigations Miller Sands marsh and upland development site, Co- lumbia River, Oregon. Summary Report, Dredged Material Research Programme Tech- nical Report D-77-38:1-72, U.S. Army Corps of Engineers Waterways Station, Vicksburg, Mississippi. Coates, K. A. 1987a. Phylogenetics of some Enchy- traeidae (Annelida: Oligochaeta): A preliminary investigation of relationships to the Haplotax- idae. Hydrobiologia 155:91-106. 1987b. Phylogenetic analysis of some En- chytraeidae (Annelida: Oligochaeta): Parsimony analysis of structural characters. Unpublished Ph.D. dissertation. University of Victoria, Vic- toria, British Columbia. 392 pp. —. (in press a). Phylogeny and origins of Enchy- traeidae (Annelida: Oligochaeta). Hydrobiolo- gia, 1989. . (in press b). Preliminary investigations of hy- bridization/reticulate evolution in Guaranidri- lus (Oligochaeta: Enchytraeidae). Hydrobiolo- gia, 1989. Healy, B. 1979. Review of the genus Guaranidrilus (Oligochaeta, Enchytraeidae) with the descrip- tion of two new species. — Bulletin of the British Museum of Natural History (Zoology) 37:7-15. VOLUME 101, NUMBER 4 Michaelsen, W. 1913. Die Oligochaeten Columbias. — Mémoires de la Société Neuchateloise des Sci- ences Naturelles 5:202—253. Omodeo, P. 1958. La reserve naturelle intégrale du Mont Nimba. IV. 1. Oligochétes. — Institut fon- damental d’Afrique noire. Mémoires. 53:9-17. Righi, G. 1973. Sobre trés espécies brasileiras de En- chytraeidae (Oligochaeta).— Boletim de Zoolo- gia e Biologia Marinha n.s. 30:469-482. 1974. Notas sobre os Oligochaeta, Enchy- traeidae do Brazil.— Papéis Avulsos de Zoologia 28:127-145. 1975. Algumas Enchytraeidae (Oligochaeta) brasileiras. —Ciéncia e Cultura 27:143-150. 1981. Notas sobre Enchytraeidae (Oligo- chaeta) brasileiras.— Revista Brasil de Biologia 41:427-430. 783 Sims, R. W., & B. M. Gerard. 1985. Earthworms. /n D. M. Kermack and R. S. K. Barnes, eds., Syn- opses of the British Fauna. #31, Cambridge Uni- versity Press, Cambridge. 171 pp. (KAC) Department of Invertebrate Zo- ology, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, Canada MSS 2C6; (RJD) Virginia Institute of Marine Science, School of Marine Science, College of Wil- liam and Mary, Gloucester Point, Virginia 23062. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 784-793 TAXONOMIC REVISION OF THE PHALLODRILUS RECTISETOSUS COMPLEX (OLIGOCHAETA: TUBIFICIDAE) Christer Erséus Abstract. —A complex of small marine tubificids previously regarded as one variable species is revised, partly on the basis of new material from Belize, Barbados, Florida, Hawaii, Great Barrier Reef (Australia), Fiji and Aldabra atoll (Indian Ocean). Four different species are recognized: Phallodrilus recti- setosus Erséus, 1979, s.s., P. deminutius Erséus, 1979, P. molestus, new species >) and P. heronensis Erséus, 1981. They are distinguished from each other by differences in the number of (both somatic and penial) setae and in the mor- phology of the spermathecae. All four species exhibit wide geographical dis- tribution in the warmer parts of the world; P. rectisetosus and P. molestus, at least, appear to be circumtropical. A complex of closely related intertidal and subtidal tubificids, belonging to the marine genus Phallodrilus Pierantoni, 1902, was described by Erséus (1979) as a polytypic species, P. rectisetosus. The worms were characterized by their (1) small body size, (2) numerous, straight and sharply single- pointed penial setae arranged in tight bun- dles, (3) small, spindle-shaped atria, each with two compact prostate glands, and (4) pear-shaped, somewhat bipartite sper- mathecal ampullae. Material from Italy and France was regarded as the nominate sub- species, while specimens from Bermuda and Florida, having fewer setae in the somatic bundles than those from Europe, were as- signed to a separate taxon, P. rectisetosus deminutius Erséus, 1979. Later, a third sub- species, P. rectisetosus heronensis Erséus, 1981, was established for material from Australia’s Great Barrier Reef, separated from the other subspecies on dimensional differences in penial setae and atria. Pecu- liarities in the appearance of its spermathe- cae were, however, also noted (Erséus 1981: 16). The distinction between the nominate P. rectisetosus and the subspecies heronensis appeared difficult to uphold, when some- what intermediate specimens were found in Saudi Arabia, and it was therefore suggested (Erséus 1985) that the whole group should be regarded as one taxonomic entity (P. rec- tisetosus), until additional material from other parts of the world and non-morpho- logical data become available. It was sus- pected (op. cit.) that more than one biolog- ical species may be involved, but it was found difficult to express the morphological variation observed in a further taxonomic division. Subsequently, additional material of this complex has become available, and it is the basis for the present study. Firstly, material from Belize in Central America, consisting of two similar but distinguishable forms, led to a re-appraisal of the original material of P. rectisetosus deminutius. Upon re-exam- ination the latter proved to contain the same two distinct forms, now concluded to be separate species. Secondly, specimens from Barbados, the Gulf of Mexico, Hawaii, Fiji, Australia, and Aldabra (Indian Ocean)— either collected by the author or received from other sources—further supported the view that more than one species is involved VOLUME 101, NUMBER 4 in the rectisetosus complex. The more than 300 individuals thus available (including the old material), could be separated into four groups, the members of each being slightly but consistently different morphologically from those of any other group, even over great geographical distances. The four forms are now regarded as different species, and they are diagnosed in the present paper. Except when otherwise stated below, the new material was collected by the author while visiting: (1) the Carrie Bow Cay field station on the Barrier Reef off Belize (op- erated by the Smithsonian Institution) (1985), (2) the Bellairs Research Institute of McGill University in Barbados (1979), (3) the Lizard Island Research Station in the Great Barrier Reef (1982), and (4) the Mana Island Resort in Fiji (1982); or by Mr. D. Davis (University of Hawaii) and the au- thor, during the latter’s stay at (5) the De- partment of Zoology in Honolulu (1987). The live worms were sorted from sieved washings of sand samples, and then fixed in Bouin’s fluid. Material was also received from the Gulf of Mexico (courtesy M. R. Milligan), and the Aldabra Atoll (Seychelles) in the Indian Ocean (K. Fauchald, Smithsonian Institu- tion). The type series and other material of P. rectisetosus deminutius were re-examined during a visit to the National Museum of Natural History (USNM), Smithsonian In- stitution, Washington, D.C. One worm from Fiji was sectioned and stained in Azan. The other specimens were stained in paracarmine and mounted whole in Canada balsam. Types and other refer- ence material have been deposited in the USNM, the Australian Museum in Sydney (AMS), and the British Museum (Natural History) in London (BMNH). The penial setae (in segment XI) were counted in all available specimens. Often the bundle on one side of the worm did not contain the same number of setae as that of the other side. For the statistics presented below, n thus refers to the total number of 785 bundles studied, i.e., twice the number of specimens. Systematics General description of members of the P. rectisetosus complex. —Small, somewhat transparent tubificids, only a few mm long. Prostomium typically phallodriline: gener- ally round, set off from peristomium. Cli- tellum extending over most posterior part of segment X, whole of XI and most of XII. Somatic setae bifid, with upper teeth thinner and often shorter than lower, and with sub- dental ligament. Bifids few per bundle, an- teriorly up to four in P. rectisetosus s.s., but only two to three in other species, poste- riorly one to two per bundle. Penial setae (ventral setae of XI) straight and short, sharply single-pointed, with tips often somewhat curved. In most species penials generally more than five per bundle (usually about three per bundle in P. molestus, n. sp.), arranged very tightly together, parallel or in somewhat fan-shaped formation with- in bundle. Bundles median to male pores; latter in line with ventral somatic setae in posterior part of XI. Spermathecal pores usually in line with ventral somatic setae anteriorly in X, near intersegmental furrow IX/X. Pharyngeal glands well developed in IV— VI. Male genitalia (cf. Figs. 1D, 2C, F) paired. Vas deferens longer than atrium, entering apical end of latter. Atrium spin- dle-shaped, more or less erect, bearing two compact prostate glands, anterior one at- tached closer to entrance of vas deferens than posterior one. Outer lining of atrium very thin, inner epithelium ciliated and often granulated. Male pore inconspicuous; penis absent. Spermathecae with sperm in loose bundle or random mass. Key to the Four Species within the Complex 1. Bifid setae fewer per bundle poste- riorly than anteriorly (Penial setae generally more than 5 per bundle) 2 786 — Bifid setae almost invariably 2 per bundle throughout body (Penial se- tae generally 3-5, only occasionally more than 5 per bundle) ..... molestus 2. Spermathecal ampullae _pear- shaped, gradually widening towards inner part and thus not clearly set off from duct (Figs. 2C, F), and with sperm in large bundle or mass (Atria at least about 45 um long) — Spermathecal ampulla small, some- what spherical, clearly set off from long duct, and with a few loosely arranged spermatozoa (Fig. 3). (Atria at most about 40 um long) ....... SALOME Da NAD bets EA BLE Mine A) SG heronensis 3. Bifid setae 2—3(4) per bundle ante- riorly, 2 per bundle in postclitellar segments rectisetosus — Bifid setae 2 (occasionally 3) per bundle anteriorly, 1 (2) per bundle in postclitellar segments ...deminutius Phallodrilus rectisetosus Erséus, 1979, sensu stricto Fig. 1A Phallodrilus rectisetosus rectisetosus Er- seus, 1979:190-191, fig. 6.—Erséus 1981: table I. Phallodrilus rectisetosus. —Erséus 1985:136- 137, figs. 3-4. Type material.—In Swedish Museum of Natural History (3086-3087): seven speci- mens from Ischia, Italy (see Erséus 1979). Other material. —Re-examined: speci- mens from Italy, France and Saudi Arabia in author’s collection (see Erséus 1979, 1985).—New material: USNM 104941, one specimen from Paiko Beach, Maunalua Bay, Oahu, Hawaii, barely subtidal mixed sand (2 Apr 1987). Diagnosis. —Length 3.2—-11.9 mm, 38-63 segments. Bifids 2—3(4) per bundle ante- riorly, 2 per bundle in postclitellar seg- ments. Penial setae 25-50 um long, gener- ally about 7-9 per bundle (total range 4-15, x = 7.9, SD = 1.9, n= 68). Atria 45-90 um PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON long. Spermathecal ampullae pear-shaped, with sperm in large bundle or mass. Description of specimen from Hawaii. — Length and segment number unknown (worm not complete). Width at XI 0.23 mm. Bifids 2-3 per bundle anteriorly, 2 per bun- dle in postclitellar segments. Penial setae (Fig. 1A) 30-35 um long, 8 per bundle. Atria about 60 um long. Spermathecae about 75 um long, with sperm in ampullae. Variation in number of penial setae per bundle (Fig. 4).—Italy (n = 58): range 4-11, x = 7.7, SD = 1.4.—France (n = 2): range 4-7, x = 5.5, but worm not fully mature. — Saudi Arabia (n = 6): range 6-15, x = 10.3, SD = 3.3.—Hawaii (n = 2): 8 (both bun- dles). Distribution and habitat.—Hawaii (new record), Italy, Atlantic coast of France and Saudi Arabia (Fig. 5). Intertidal and barely subtidal sand. Phallodrilus deminutius Erséus, 1979 Figs. 2A—C Phallodrilus rectisetosus deminutius (part) Erséus, 1979:191-192, fig. 7.—Erséus 1981:table I. Type material re-examined. —USNM 56200-56202, six specimens from Ferry Reach, Bermuda (see Erséus 1979). Other material re-examined. —USNM 56223, two specimens from Blue Hole area, Bermuda, coll. by M. L. Jones (see Erséus 1979).—Author’s collection: 12 specimens from Bermuda (11 from type locality; 1 from Whale Bone Bay, intertidal). Note that sev- eral other individuals from Bermuda, pre- viously assigned to “P. rectisetosus demi- nutius’ now are identified as P. molestus, n. sp., below. New material. —USNM 104134, one specimen from N of inner end of dock at Carrie Bow Cay, Barrier Reef off Belize, 0.5 m, patch of poorly sorted medium to coarse sand (6 Nov 1985). USNM 104133, 104135, two specimens from N of dock, Carrie Bow Cay, 1 m, fine to medium sand and rubble (10 Nov 1985). USNM 104132, one spec- VOLUME 101, NUMBER 4 imen from Bellairs Reef, St. James, Bar- bados, 13-14 m, poorly sorted coarse sand (29 Oct 1979).—Author’s collection: two specimens from 50 m N of Carrie Bow Cay, 0.7 m, rubble bed with coarse to medium sand beneath (8 and 24 Nov 1985); two specimens from immediately W of N tip of Carrie Bow Cay, 0.5 m, patch of medium sand amongst scattered rocks and Thalassia (8 Nov 1985); one specimen from SW of S tip of Carrie Bow Cay, edge of reef, near beginning of Thalassia bed, 2 m, poorly sorted sand (12 Nov 1985); and two spec- imens (coll. H. R. Baker) from near dock of Carrie Bow Cay, 1.5 m, poorly sorted coral sands with algal debris, roots and Halimeda flakes (25 Apr 1982). Diagnosis. —Length 3.4-5.9 mm, 36-44 segments. Bifids 2(3) per bundle anteriorly, 1(2) per bundle in postclitellar segments. Penial setae 30-46 wm long, generally 5-8 per bundle (total range 4-11, x = 6.7, SD = 1.5, n = 60). Atria 60-85 um long. Sper- mathecal ampullae pear-shaped, with sperm in large bundle or mass. Description of material from Belize and Barbados. — Length 3.5—5.8 mm, 36-43 seg- ments. Width at XI 0.14—0.27 mm. Bifids (Fig. 2A) 2(3) per bundle in II-VIII, 1 (very occasionally 2) per bundle thereafter, 28—37 um long, about 2 um thick. Penial setae (Figs. 2B, C, ps) 35—40 um long, about 2 um thick, 4-8 per bundle. Male genitalia shown in Fig. 2C. Vas deferens about 9 um wide. Atrium 60-85 wm long, 23-35 wm wide. Sperma- thecae (Fig. 2C, s) 80-145 um long, con- sisting of short ducts and pear-shaped, thin- walled ampullae; latter narrower ectally than entally, maximally 35-70 um wide; sperm as thick bundle or large mass. Variation in number of penial setae per bundle (Fig. 4).—Bermuda (n = 38): range 4-11, x = 7.0, SD = 1.7.—Belize (n = 20): range 4-8, x = 6.2, SD = 1.1.—Barbados (n = 2): range 7-8, x = 7.5. Remarks.—The single individual from Fowey Rocks, Miami, Florida, that was identified as “‘P. rectisetosus deminutius” by Erséus (1979) is actually a specimen of P. 787 Fig. 1. penial setae. B—D, Phallodrilus molestus, from Hawaii: B, Somatic seta; C, Penial setae; D, Lateral view of spermatheca and male genitalia in segments X—XI. Ab- breviations: a atrium, pr | anterior prostate gland, pr 2 posterior prostate gland, ps penial setae, s sperma- theca, vd vas deferens. A, Phallodrilus rectisetosus, from Hawaii, molestus. However, although P. deminutius co-occurs with P. molestus in many places (even at the type locality of the former), the holotype and paratypes of it proved all, upon re-examination, to be the same species. The key characteristic separating this species from all the others within the com- plex is the low number of setae; with few exceptions the postclitellar bundles are uni- setal. It is otherwise very similar to P. rec- tisetosus, both in terms of number of penial setae and in spermathecal morphology. Distribution and habitat.—Belize, Bar- bados (both new records) and Bermuda (Fig. 5); apparently present throughout the Ca- ribbean area. Intertidal and subtidal sand, down to at least about 14 m depth. Phallodrilus molestus, new species Figs. 1B—D, 2D-F Phallodrilus rectisetosus deminutius (part) Erséus, 1979:191-192.—Erséus 198 1:ta- ble I. 788 A 10 ym D | 10 ym E 20pm PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. A—C, Phallodrilus deminutius, from Belize: A, Somatic seta; B, Penial setae; C, Lateral view of spermathecae and male genitalia in segments X—XI. D-F, Phallodrilus molestus, from Belize: D, Somatic setae; E, Penial setae; F, Lateral view of spermatheca and male genitalia in segments X—XI. Abbreviations: a atrium, pr 1 anterior prostate gland, pr 2 posterior prostate gland, ps penial setae, s spermatheca, vd vas deferens. Holotype. —USNM_ 118352, whole- mounted specimen from outer end of dock at Carrie Bow Cay, Barrier Reef off Belize, 1.5 m, poorly sorted, somewhat muddy, medium to coarse sand (6 Nov 1985). Paratypes. -USNM 118353 -118356, four specimens from type locality. Other material re-examined (author’s collection).— Nine specimens from Ber- muda (Erséus 1979:191): two from Ferry Reach (type locality of P. deminutius), three from North Rock, one from Tobacco Bay, one from Harrington Sound, and two from Bailey’s Bay. One specimen from Fowey Rocks, Miami (Erséus 1979). Other material.—USNM 118357, one specimen from inner end of bay at river mouth, River Bay, NE Barbados, low water mark, coarse sand (27 Oct 1979).—USNM 118358-118361, four specimens from Pai- ko Beach, Maunalua Bay, Oahu, Hawaii, barely subtidal mixed sand (2 Apr 1987).— AMS W202443, one specimen from beach at northern side of Palfrey Island, S of Liz- ard Island, Great Barrier Reef, Australia, lower intertidal, gravel with silt and fine sand (15 Nov 1982).—BMNH _ 1987.5.1/2, 1987.5.3, two specimens from Mana Island, off Lautoka, Viti Levu, Fiji (5 Dec 1982): one from flat W of Mana Island Resort, subtidal off beach rock, 0.5 m, fine coralline sand with lots of gravel and coral rubble; and one from rocks at W end of North Beach (Mana Island), tide pool in lower intertidal, stones and pebbles with some muddy sand beneath.—Author’s collection: 20 speci- VOLUME 101, NUMBER 4 mens from type locality; 177 specimens from 32 other sampling stations in the reef areas near Carrie Bow Cay, Belize (Nov 1985); six specimens from Oahu, Hawaii, four from Paiko Beach (see above), one from off Keehi Lagoon, Honolulu, about 70 m, medium sand (Nov 1986; coll. D. Davis), and 1 from Kawaikui Beach Park, Maunalua Bay (9 Sep 1987; coll. D. Davis).—M. R. Milligan Col- lection: 1 specimen from about 4 km off mouth of Crystal River, W Florida (Gulf of Mexico), 28°58.7'N, 82°48.7'W, 4 m, coarse sand and shells (Nov 1984; coll. M. R. Mil- ligan). Etymology. —The species name molestus is Latin for “troublesome,” referring to the previous difficulties in separating this form from P. deminutius. Diagnosis. —Length 3.0-6.8 mm, 30-43 segments. Bifids almost invariably 2 per bundle, throughout body. Penial setae 28— 47 um long, generally 3—5 per bundle (total range 2-7, x = 3.1, SD = 0.7, n = 444). Atria 40-80 um long. Spermathecal am- pullae pear-shaped, with sperm in large bundle or mass. Description of material from Belize (in- cluding type material) and Barbados.— Length 3.7—-5.9 mm, 35-42 segments. Width at XI 0.19-0.24 mm. Bifids (Fig. 2D) 2 per bundle throughout (very occasionally | per ‘bundle’), 26—42 wm long, 1.5—2.5 wm thick. Penial setae (Figs. 2E, F, ps) 35-46 um long, about 2 wm thick, generally 3, but some- times 2, 4 or even 5, per bundle. Male gen- italia shown in Fig. 2F. Vas deferens 7-14 um wide. Atrium 45-80 um long, 20—30 um wide. Spermathecae (Fig. 2F, s) 130-210 um long, consisting of short ducts and pear- shaped, thin-walled ampullae; latter nar- rower ectally than entally, maximally 25- 95 um wide; sperm as thick bundle or large mass. Description of material from Hawaii, Great Barrier Reef and Fiji.—Length 3.2- 5.2 mm, 36—42 segments. Width at XI 0.20- 0.35 mm. Bifids (Fig. 1B) 30-42 um long, 1.5—2.5 um thick, 2 (anteriorly sometimes ~ only 1) per bundle throughout. Penial setae 789 (Fig. 1C) 28-42 um long, 1.5-2 um thick, generally 4—5 (total range 3-6) per bundle. Male genitalia shown in Fig. 1D. Vas def- erens 5—7 um wide. Atrium 40-60 um long, 15-23 um wide. Spermathecae (Fig. 1D, s) 60-90 um long, consisting of short ducts and pear-shaped, thin-walled ampullae; lat- ter narrower ectally than entally, maximally 28-42 um wide; sperm as large mass, or absent (lumen then filled with a round mass of secretion, as shown in Fig. 1D). Variation in number of penial setae per bundle (Fig. 4).— Belize (n = 394): range 2- 5, x = 3.0, SD = 0.6.— Bermuda (n = 18): range 2—5 (one bundle with 7), x = 4.1, SD = 1.1.—Miami (n = 2): range 4-5, x = 4.5.— Gulf of Mexico (n = 2): 2 (both bundles). — Barbados (n = 2): 3 (both bundles).—Ha- wall (n = 20): range 3-6, x = 4.4, SD = 0.7.—Great Barrier Reef (n = 2), range 3- 4, x = 3.5.—Fiyi (n = 4): range 4-5, x = 4.3, SD = 0.4. Remarks. —Some of the specimens from Bermuda and Florida were previously (Er- seus 1979) identified as P. rectisetosus de- minutius, but they have bisetal bundles of bifids throughout the body, and a low num- ber of penial setae, the two features unique for the form now recognized as a separate species, P. molestus. Distribution and habitat.—Belize, Bar- bados, Florida, Bermuda, Hawaii, Great Barrier Reef, Fiji (Fig. 5). Lower intertidal and subtidal sands, down to at least 70 m depth. Phallodrilus molestus is one of the most common tubificids on the Barrier Reef off Belize. Phallodrilus heronensis Erséus, 1981 Fig. 3 Phallodrilus rectisetosus heronensis Erséus, 1981:15-17, figs. 1-3, table I. Type material. —In Queensland Museum (G 12544-12551): eight specimens from Heron Island, Great Barrier Reef, Australia (see Erséus 1981). Fig. 3. Phallodrilus heronensis, spermathecae: A, Specimen from Fiji; B, Specimen from Aldabra. New material.—BMNH 1987.5.4/5 and 1987.5.6, one sectioned and two whole- mounted specimens from reef flat off ““Pic- nic Point,’ Mana Island, off Lautoka, Viti Levu, Fiji, 0.5 m, coarse gravelly sand (7 Dec 1982).—USNM 104129-104131, three specimens from Aldabra Atoll (Seychelles), coll. K. Fauchald: one from Basin Lebine, 0.5 m, from Caulerpa sp. clump (13 Mar 1985); and two from in front of laboratory, in Thalassodendron (Apr 1983). Diagnosis. —Length 2.3-4.6 mm, 33-39 segments. Bifids 2—3 per bundle anteriorly, 2 per bundle in postclitellar segments. Pe- nial setae 20-35 um long, 4—7 (occasionally up to 11) per bundle (x = 6.4, SD = 1.6, n = 30). Atria very small, only 25—40 um long. Spermathecae club-shaped, with long ducts and small, somewhat spherical ampullae; latter containing a few loosely arranged spermatozoa. Description of material from Fiji. — Length, 2.7-2.9 mm, 35-37 segments. Width at XI 0.26-0.27 mm. Bifids 30-35 um long, 1.5—2 wm thick, 2-3 per bundle anteriorly, 2 per bundle in postclitellar seg- ments. Penial setae about 25-30 um long, 4—5(6) per bundle. Atrium about 30 um long, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON about 10 wm wide. Spermathecae (Fig. 3A) 55-70 um long, maximally 17—26 um wide, with sperm as loose bundle in ampullae. Description of material from Aldabra. — Length 2.3-3.5 mm, 33-37 segments. Width at XI 0.10-0.18 mm. Bifids 25-37 um long, 1-2 um thick, 2-3 per bundle anteriorly, 2 per bundle in postclitellar segments. Penial setae 20-25 um long, 7(9) per bundle. Atrium about 25-30 um long, 11-15 wm wide. Spermathecae (Fig. 3B) only 40-47 um long, 10-20 um wide, with a few sper- matozoa in ampullae. Variation in number of penial setae per bundle (Fig. 4).— Heron Island, Great Bar- rier Reef (n = 18): range 5-11, xX = 6.7, SD = 1.6.—Fiji (n = 6): range 4-6, x = 4.8, SD = 0.7.— Aldabra (n = 6): range 7-9, X = 7.3, SD = 0.7. Remarks. —The very small atria and spermathecal ampullae are diagnostic for P. heronensis, and separate it from the other species of the complex. Distribution and habitat.—Aldabra, Fiji (new records) and Great Barrier Reef (Fig. 5). Intertidal and barely subtidal sand, down to at least 0.5 m depth. Discussion The species complex considered here is a homogeneous group of marine tubificids, with a few variable and somewhat confusing characters that made the original classifi- cation difficult (Erseus 1979, 1981, 1985). The new specimens from Florida, Belize, Barbados, Hawaii, Fiji, Australia and Al- dabra, however, have revealed a pattern in the variation indicating the presence of four distinct species. Three characters have shown to be particularly useful. First, the bifid (somatic) setae, which morphologically are more or less identical in the four forms (Figs. 1B, 2A, D), are up to 3(4) per bundle anteriorly, diminishing to 2 per bundle posteriorly, in P. rectisetosus and P. heronensis, whereas the bundles are VOLUME 101, NUMBER 4 (almost invariably) bisetal anteriorly and unisetal posteriorly in P. deminutius. Such a posterior decline in setal numbers does not characterize P. molestus, which has bi- setal bundles throughout the body. Second- ly, the latter species is also unique in its very low number of penial setae, in fact it is the only species in the group that in most cases can be separated solely on the basis of this number. The other three species have very wide, overlapping ranges for the numbers of penial setae (see Fig. 4), the means being very close to each other. Thirdly, although variable in shape, the spermathecal ampul- lae of P. rectisetosus, P. deminutius and P. molestus are large (relative to those of P. heronensis) and pear-shaped, sometimes with a tendency towards a bipartition (cf. Erséus 1979:fig. 6), and they are not always clearly set off from the ducts. In P. hero- nensis, however, the ampullae are very small, roundish and clearly set off from the (relatively long) ducts (Fig. 3). Some of these differences may appear slight and not, per se, interspecific. For in- stance, a variation in the number of somatic setae between one and two, or two and three, per bundle may be considered intraspecific in other tubificid species. However, in the present study, which covers material from 13 different areas of the world (Fig. 5), each individual could rather easily be assigned to one of the four distinct forms; no inter- mediates were found. The forms are not even fully separated geographically, thus refuting the alternative of them being geographical races. As illustrated in Fig. 5, two forms are => Fig. 4. Variation in the number of penial setae per bundle, in various populations of the Phallodrilus rec- tisetosus complex. A, P. rectisetosus from Italy (A1), France (A2), Saudi Arabia (A3), and Hawaii (A4); B, P. deminutius from Bermuda (B1), Belize (B2), and Barbados (B3); C, P. molestus, from Belize (C1), Ber- muda (C2), Miami (C3), Gulf of Mexico (C4), Barbados (C5), Hawaii (C6), Great Barrier Reef (C7), and Fiji (C8); D, P. heronensis from Great Barrier Reef (D1), Fiji (D2), and Aldabra (D3). (ieee | A2 A3 0 oe ; I Al 0 10 0 ali Baoan Bl "4 fle an B 3 200 100 (C4 0 ial F Re ‘ C2 14 tee C3 2 co | C4 a4 Fa C5 vi 0 aL C6 a cy 3 ak ie C8 gs Uv 5. a oe3 Borge bt al. D2 KS a | 791 gaa lcs ue a me 23) fh SG Bo 9). SO Sh We GI We ss Number of penial setae per bundle 792 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON @ P rectisetosus * P deminutius e P molestus n.sp. A P heronensis Fig. 5. species from the same area are encircled. sympatric (in various combinations) in 5 of the 13 areas studied. The records suggest, however, that the Caribbean P. deminutius and the Indo-Pacific P. heronensis may be geographically more restricted than the ap- parently circumtropical P. rectisetosus and P. molestus. Although these four species without doubt are closely related, they do not necessarily constitute a strictly monophyletic group. Their simple male ducts and penial setae do not appear to be apomorphic traits, but rather less derived structures possibly sim- ilar to those of an ancestral phallodriline; the male genitalia of most other members of the subfamily appear to be derived from the biprostate (i.e., unmodified) atria of Phallodrilus. This implies that the rectise- tosus group, as well as the genus Phallodrilus as a whole, is likely to be paraphyletic. The recently described P. duplex Erséus, 1987, from the Mediterranean coast of Israel, most probably belongs to the same lineage. It has bisetal bundles throughout body, except for the penial ones which are trisetal, and it thus appears to be very closely related to P. mo- lestus; it is distinguished from the latter by its somewhat more slender atria and the modified tips of its penial setae (see Erséus 1987). Particularly the latter feature makes Geographical distribution of the species within the Phallodrilus rectisetosus complex. Records of two it a slightly more derived species than the four taxa treated in the present paper. Acknowledgments I am indebted to Dr. K. Ruetzler (USNM and Carrie Bow Cay field station), Dr. F. Sander (former director, Bellairs Research Institute), and Dr. B. Goldman (former di- rector, Lizard Island Research Station), for working facilities at their respective labo- ratories; to Dr. L. Zann (Suva) for assistance in planning my work and arranging a re- search permit in Fiji; to Dr. K. Fauchald (USNM), Drs. H. Spero and M. Gustavson (Barbados), Dr. F. Doujak (Canberra), and Mr. D. Davis (now in San Francisco), for most valuable assistance in the field work; to Dr. R. O. Brinkhurst (Sidney, Canada), Mr. M. R. Milligan (Sarasota, FL), and Dr. K. Fauchald, for making material in their collections available for study; to Ms. B. Lofnertz and Mrs. A. Falck-Wahlstrom (Goteborg) for skillful technical assistance; to the Smithsonian’s Walter Rathbone Ba- con Scholarship Fund, and the Swedish Natural Science Research Council, for fi- nancial support; and to R. O. Brinkhurst for critical review of the manuscript. This paper is contribution number 244 of the Smith- VOLUME 101, NUMBER 4 793 sonian Institution’s Caribbean Coral Reef MK. 1985. Annelida of Saudi Arabia. Marine Ecosystems Program. It is also a contribu- Tubificidae (Oligochaeta) of the Arabian Gulf tion from the Bellairs Research Institute of Coast of Saudi Arabia.— Fauna of Saudi Arabia : g : 6(1984):130-154. McGill University, Barbados. 1987. A new species of Phallodrilus and rec- ords of two other marine Tubificidae (Oligo- Literature Cited chaeta) from the Mediterranean coast of Is- rael.— Israel Journal of Zoology 33(1984/85):73— Erséus, C. 1979. Taxonomic revision of the marine 78. genus Phallodrilus Pierantoni (Oligochaeta, Tubificidae), with descriptions of thirteen new : species. — Zoologica Scripta 8:187—208. Zoo-tax, Swedish Museum of Natural 1981. Taxonomic studies of Phallodrilinae History, Stockholm, and (postal address:) (Oligochaeta, Tubificidae) from the Great Bar- Department of Zoology, University of Go- rier Reef and the Comoro Islands with descrip- teborg, Box 25059, S-400 31 Géteborg, tions of ten new species and one new genus. — Sweden Zoologica Scripta 10:15-31. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 794-808 FOUR NEW SPECIES OF CAMBARINCOLIDS (CLITELLATA: BRANCHIOBDELLIDA) FROM THE SOUTHEASTERN UNITED STATES WITH A REDESCRIPTION OF OEDIPODRILUS MACBAINI (HOLT, 1955) Perry C. Holt Abstract. — Four new species of cambarincolid branchiobdellidans, Cambar- incola bobbi from Virginia, Ellisodrilus carronamus and Oedipodrilus anisog- nathus from Tennessee, and Sathodrilus rivigeae from Arkansas are newly described. In addition, Oedipodrilus macbaini (Holt, 1955) is redescribed. Re- lationships and distributions are discussed. When the branchiobdellidan collections belonging to the Virginia Polytechnic Insti- tute and State University were given to the National Museum of Natural History of the Smithsonian Institution in the Spring of 1986, I retained in my possession as a loan certain collections from the southeastern United States, including all the material from Tennessee, which I knew to contain a number of new species. This opportunity is taken to describe four of these species and to more fully describe Oedipodrilus mac- baini (Holt, 1955). The region composed of the southern Ap- palachians and associated uplands (the Piedmont and the Interior Low Plateaus) and the Ozark Mountains contains the most diverse branchiobdellidan fauna known for any similarly sized region in the world and may include the original home of the order. An understanding of the relationships and history of the branchiobdellidans awaits a fuller description of this fauna. Some discussion of generalities and ge- neric and familial diagnoses may be found in Holt (1986). Herein, the intent is merely to describe new species with minimal em- phasis placed on any other questions. Cambarincola bobbi, new species Fig. | Type specimens.—Holotype, USNM 101496, four paratypes, USNM 101497- 101499, taken on Cambarus bartonii bar- tonil (Fabricius) from a medium-sized stream in Tom’s Brook (a town) 5.7 miles south of Strasburg, Shenandoah County, Virginia, by Marvin L. Bobb and Perry C. Holt, 22 Jul 1948. Diagnosis. —Small- to medium-sized worms (holotype 2.2 mm in length); lips obscurely lobed; no oral papillae; dorsal ridges weakly developed; jaws subequal in size, dorsal one slightly larger than ventral one, dark in color, dental formula 5/5; bursa about 4 to *%4 body diameter in length, elongate ovoid, atrial fold thin, penial sheath about 4 total length, penis membraneous; spermiducal gland large, length about twice its diameter, no deferent lobes, flexed an- tero-ventrad at its mid-point; prostate about ¥, length, *%% diameter of spermiducal gland, differentiated, with large clear ental bulb; spermatheca with prominent ectal duct, bulb ovoid to lanceolate (in optical section), length slightly less than diameter of body. VOLUME 101, NUMBER 4 795 Fig. 1. Cambarincola bobbi, holotype. a, Lateral view of entire animal; b, Lateral view of reproductive systems; c, Optical section through bursa and penis; d, Lateral view of jaws. Abbreviations: af, atrial fold; b, bursa; ba, bursal atrium; ejd, ejaculatory duct; p, penis; pr, prostate; ps, penial sheath; sb, spermathecal bulb; sd, spermathecal duct; sg, spermiducal gland. Etymology.—For Dr. Marvin L. Bobb, friend, fellow student, and for a season, field companion. Description. —The five type specimens of Cambarincola bobbi have the following mean dimensions: total length, 2.0 mm; greatest diameter, 0.3 mm; head length, 0.3 mm; head diameter, 0.2 mm; diameter, seg- ment I, 0.2 mm; diameter, sucker, 0.3 mm. In external appearance, C. bobbi is un- remarkable. The lips have four dorsal, two ventral, indistinct lobes. Oral papillae are not detectable, yet there are slight, low un- dulations of the inner margin of the mouth. The posterior fourth of the head is demar- cated by a shallow external sulcus and a prominent pharyngeal one. Supernumerary muscles of the dorsal portions of the ante- rior prosomites of the body segments are poorly developed, thus in profile the body outline is smooth: there are no dorsal ridges. The sucker is prominent and slightly greater in diameter than segment I. The length of the jaws is about 1, that of the head; hence, of usual comparative size. The animals available for this study are all strongly bent ventrad. The exact distribu- tion of the teeth, consequently, is difficult to ascertain in most specimens. There is a prominent apical tooth on each jaw with 796 two small lateral ones flanking it on each side, producing a 5/5 dental formula. The spermiducal gland is large and re- flexed to some extent, often to a right angle, at its mid-length. It has no deferent lobes. The slender prostate is about '2 the length of the spermiducal gland and is “‘differen- tiated,” that is, composed of cells less dense- ly granular than those of the spermiducal gland. A large, clear ental bulb is present. The bursa is elongate ovoid in shape. The penis is distinctly less muscular than is usual among members of the genus, consisting of an ovoid sac, presumably protrusible, with a few strands (? muscular) traversing it. The ejaculatory duct is of the usual type, a short muscular tube. The spermathecal duct is about '2 the body diameter in length; the bulb is ovoid with- out an ental process. Variations.—No variations of note, ex- cept the extent of reflexion of the spermi- ducal gland, were detected. Affinities. —Cambarincola bobbi clearly belongs to the group of species with a dif- ferentiated prostate with an ental bulb that Hoffman (1963:336-341) assigned to his ‘“‘Philadelphica section” and “group” of the genus. The “Fallax subgroup” in this scheme is distinguished by an equal (and odd) num- ber of teeth borne by each jaw. In this subgroup (consisting of C. fallax Hoffman, 1963, and C. holostomus Hoffman, 1963), Cambarincola bobbi most closely resembles C. holostomus. Cambarincola holostomus is described by Hoffman (1963:361) as characterized by an entire peristomium; that is, there is no di- vision of the peristomium into upper and lower lips, a 3/3 dental formula and a “‘slen- der elongate, fusiform” spermatheca. In ad- dition to differing from C. holostomus in these respects, C. bobbi also is distinguished by a larger and more reflexed spermiducal gland and a shorter prostate (Hoffman 1963: 359, fig. 64). Cambarincola bobbi resembles C. fallax in these features and the 5/5 dental formula, but lacks the characteristic elon- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gated peristomial tentacles of C. fallax and differs from both in its membranous penis. The structure of the penis of C. bobbi seems to reinforce the previously noted similari- ties between the bursal-penial complex of the genera Cambarincola and Sathodrilus Holt, 1968a (Holt 1982:254). However, in this case, it is most likely another example of convergence within the order, since the relatives of C. bobbi are presumably “ad- vanced”? members of the genus (Hoffman 1963:296-297). Host.—Cambarus bartonii bartonii. Distribution. —Known only from the type locality. Other similar streams of the Shen- andoah River system should harbor this species. Material examined. —The type series. Note. —The tributaries of the New River in western Virginia and eastern West Vir- ginia and the headwaters of the Holston, Roanoke, James, Shenandoah and Potomac rivers contain many similar species of the genus Cambarincola. Some specimens from this area are difficult to identify, but C. bobbi appears to be clearly distinct from any pre- viously described species, and is another ex- ample of the diversity and the incipient spe- ciation in these waters: waters undoubtedly linked by many stream captures in the re- cent past. Ellisodrilus carronamus, new species Fig. 2 Type specimens. —Holotype, USNM 119539 and two paratypes, USNM 119540- 119541, taken on Orconectes sp., from Carr Creek, Overton County, Tennessee, about 3.0 miles south of Livingston, Tennessee, on State Highway 42, by Perry C. and Virgie F. Holt, 26 Jul 1961. Diagnosis. —Small worms (holotype 1.95 mm in length); lips entire, no oral papillae; no dorsal ridges; upper jaw slightly longer than lower, dark brown in color, dental for- mula (?) 5/5; bursa slightly less than '2 body diameter in dorso-ventral length, antero- VOLUME 101, NUMBER 4 797 Fig. 2. Ellisodrilus carronamus, holotype. a, Lateral view of entire animal; b, Lateral view of male efferent apparatus; c, Optical section of bursa; d, Lateral view of jaws. posterior dimension (diameter) slightly greater than dorso-ventral one; penis re- duced to pore through ental portion of bur- sa, pseudo-penis formed by enlarged inner- directed atrial fold; ejaculatory duct short, thin; spermiducal gland oriented longitu- dinally in coelom, lying dorsad to gut, in length about twice its diameter; prostate arises entad to mid-point of spermiducal gland, less than 2 length, subequal in di- ameter to latter, non-differentiated, with small ental bulb; no spermatheca. Etymology. — From Greek, namos, creek, hence Carr Creek for the type locality and a family that includes friends of my youth for whom the creek was named. Description. —The holotype, which is comparable in size to the other specimens examined, has the following approximate dimensions: total length, 1.9 mm; greatest diameter, 0.4 mm; head length, 0.3 mm; head diameter, 0.2 mm; diameter segment I, 0.2 mm; diameter, sucker, 0.2 mm. The lips lack lobes; there are no oral pa- pillae. The one internal pharyngeal sulcus has no corresponding external one. The jaws are brown and prominent, the upper about 4, the length of the head, the lower somewhat less. No lateral teeth are apparent in the holotype at a magnification of 500 but some of the paratypes appear to have two lateral teeth at the side of the large median tooth: the dental formula would be, then, 5/5, a point requiring fur- ther confirmation. Dorsal ridges are absent from all seg- ments, but the inter-segmental furrow be- tween VI and VII is deeply incised and the clitellum of segment VII is prominent: this accounts for the greater diameter of segment 798 VII and gives to this segment a superficial appearance of a dorsal ridge, which, how- ever, entirely lacks supernumerary muscles. The spermiducal gland, though propor- tionately large, is unremarkable. There are no obvious deferent lobes and the organ ta- pers from its greatest diameter at the point of origin of the prostate to its ectal end. The prostate arises from the spermiducal gland slightly more than 4 the length of the latter from its ental end and is almost as great in diameter. It is non-differentiated with a small, but distinct, ental bulb. The bursa is large, filling the ventral half of its segment. Its ectal portion (atrium), which has an inner-directed circular fold (=atrial fold) whose lumen slants towards the postero-dorsal side of the bursa and opens into the bursal cavity opposite the opening of the ejaculatory duct into the atrium, is the most striking feature of the male reproductive system (Fig. 2c). In effect there is no structure which is homologous to the penis of other genera of the bran- chiobdellidans except the pore formed by the opening of the ejaculatory duct into the lumen of the bursa (atrium). The ejaculatory duct is thin, short and difficult to detect in available material. The atrial fold, a pseu- dopenis, would appear to be analogous to the penis of other branchiobdellidans. There is no spermatheca, hence some form of hypodermic impregnation must occur, though no packets of spermatozoa have been observed on the dorsal surface of any spec- imen of any member of the genus. Such packets of spermatozoa have been reported by Holt (1949:549) in Xironogiton instabilis (Moore, 1894). Variations. —In several of the specimens examined, the bursa lies along the dorso- ventral axis of segment VI, presumably compressed by a contraction of the body- wall. No other variations were noted. Affinities. —Ellisodrilus durbini (Ellis, 1919) and E. clitellatus Holt, 1960, have dorsal ridges on some segments: on segment VIII in E. clitellatus and segments II-V and VII, VIW in E. durbini. Ellisodrilus carron- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON amus lacks these ridges produced by super- numerary muscles (Holt 1960:171-172). Ellisodrilus carronamus further differs from its two congeners in its apparently larger size and somewhat larger and darker jaws with a dental formula of (?)5/5. The absence of dorsal ridges distinguishes E. carronamus. There appears to be a north-south gra- dient in the distributions of these species with E. carronamus the southern-most and E. durbini to the north, in part at least, in areas scoured by the Pleistocene glaciations (Holt 1960:171; 174). Branchiobdellidan associates. — Ellisodri- lus carronamus shares its type locality with four other branchiobdellidans: Cambarin- cola philadelphicus (Leidy, 1851), Xiron- odrilus formosus Ellis, 1918, Pterodrilus cedrus Holt, 1968b, and Oedipodrilus an- isognathus, newly described herein. Material examined. —The types; six spec- imens taken from Roaring River on Cam- barus tenebrosus Hay, 4.7 miles north of Rickman, Overton County, Tennessee, by Perry C. and Virgie F. Holt, 4 Jul 1958. Oedipodrilus anisognathus, new species Fig. 3 Type specimens. —Holotype USNM 119534 and one paratype, USNM 119535 taken from a small stream on Orconectes sp. in Montgomery Bell State Park, Dickson County, Tennessee, by Perry C. and Virgie F. Holt, 4 Jul 1958; three paratypes, USNM 119536-119537 taken on Orconectes sp. from Carr Creek, about 3.0 miles south of Livingston, Overton County, Tennessee, by Perry C. and Virgie F. Holt, 26 Jul 1961. Diagnosis. —Small worms (holotype 1.8 mm in length); lips entire; no oral papillae; no dorsal ridges; posterior segments mark- edly greater in diameter than anterior ones; jaws greatly dissimilar in size, upper one about 4 to '4 that of lower in length, dental formula 2/1; bursa relatively small, 2 body diameter in length; penis with (?) hooks; spermiducal gland lies longitudinally in coe- lom, with obscure deferent lobes; prostate VOLUME 101, NUMBER 4 799 Fig. 3. Oedipodrilus anisognathus, holotype. a, Lateral view of entire animal; b, Lateral view of male efferent apparatus; c, Optical section of bursa; d, Lateral view of jaws. with ectal end arising at about ental third of spermiducal gland, about '4 length of lat- ter, with ental bulb; spermatheca with ectal bursa, cylindrical bulb, narrow ental pro- cess. Etymology. —Greek: anisos, unequal; gnathos, jaw. Description. —The specimens of Oedipo- drilus anisognathus that constitute the type series have the following mean dimensions: total length, 1.4 mm; greatest diameter, 0.4 mm; head length, 0.4 mm; head diameter, 0.2 mm; diameter, segment I, 0.3 mm; di- ameter, sucker, 0.4 mm. The worms have a somewhat corpulent appearance with slender heads. There are no supernumerary muscles of the proso- mites of the body segments: the outline of the body is smooth. There is a slight con- striction delimiting the peristomium; an in- ternal pharyngeal sulcus lies at the greatest diameter of the head, often marked by a slight, shallow external one. The lips are entire, the dorsal somewhat longer than the ventral one. No oral papillae are apparent. The jaws are distinctive: unlike any oth- ers known for any member of the order. The upper jaw is small with two small, obscure teeth; the lower, three or four times the length of the upper, is massive, with flaring lateral flanges and a single, large, triangular tooth. The spermiducal gland is proportionately small, as is the entire male reproductive sys- tem, and lies along the longitudinal axis of the body. Deferent lobes are not prominent. The prostate consists of a rather thick lobe of the spermiducal gland, about ' the latter in length, with a clear ental bulb. It does not differ from the spermiducal gland in its histological appearance. The bursa is an elongated ellipsoid with a shallow constriction separating its atrial and penial sheath portions. There is a some- what irregular atrial fold. The penis is an elogated tube coiled inside the penial sheath, 800 attached only at the ectal end of the lumen of the latter and to its ental continuation as the ejaculatory duct. In its everted position the penis would be lined by a thin layer of tissue, possibly muscular. Penial hooks are few in number and difficult to detect with customary methods: there appear to be two or three pairs. The ejaculatory duct is a short, thick mus- cular tube, almost as great in diameter as in length. The ectal duct of the spermatheca is a short, thick, muscular spermathecal bursa. About 4 the total length of the spermatheca is composed of the spermatozoa storing bulb. There is a narrowed ental process with a thick muscular wall. Variations.—None were noted in the available material. Affinities. —Three other species of Oedi- podrilus have been described: O. oedipus Holt, 1967, the type species from Hum- phreys County, Tennessee; O. macbaini (Holt, 1955), from Tennessee northward to Illinois and Pennsylvania; O. cuetzalanae Holt, 1984, from the state of Puebla, Mex- ico. Oedipodrilus oedipus has dorsal ridges, jaws with a 5/4 dental formula, a much shorter penis with many penial hooks and lacks an ental process of the spermatheca, differing from O. anisognathus in all these features. Oedipodrilus macbaini principally differs from O. anisognathus in the 5/4 den- tal formula, the much longer penial sheath and remarkably longer penis with many prominent hooks and the very long ectal duct of the spermatheca and the absence of a spermathecal ental process. Oedipodrilus cuetzalanae has a 5/4 dental formula; a very small, often obscure, prostate; a long, coiled penis with many hooks; a slender sperma- theca with a long ectal duct, a thick-walled bulb and no ental process: O. anisognathus is different in all these features. Holt (1984: 40) said that “‘[t]he three known species of the genus form a coherent group and it is futile at this stage to speculate about which is more closely related to the other.” In gen- eral, this statement is still true, but perhaps PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the unusual (? gill-clipping) jaws of O. an- isognathus separates it more widely from the other species of the genus. Notice should be taken here of some similarities of these species with some members of the genus Sathodrilus, all of which likewise have an eversible penis. In most species of Sath- odrilus the penis is a straight tube attached by strands of tissue to the inner wall of the penial sheath and everts as a membranous cup-shaped structure. But the penis of S. villalobosi Holt, 1968a, is an elongated tube with shallow coils (Holt 1984:40, fig. 3a) that, however, lacks penial hooks. These penial hooks may be the only character that reliably separates the two genera. Habitat.— Oedipodrilus anisognathus has, though, not to as marked degree as some such species, the appearance of parasitic, gill-inhabiting worms: a thinner than usual body-wall, expanded posterior body seg- ments, elongated and tapering head and peristomium. The jaws, closer in shape than are those of any other species to those of Bdellodrilus illuminatus Moore, 1895, which is known to inhabit the gill chambers of its hosts and to be parasitic, seem well suited to piercing the thin cuticle of the host’s gills. More to the point, the gut of all the speci- mens examined of O. anisognathus is filled with a homogeneous coagulum, most likely blood, without any of the usual components of the slime found on the outer surfaces of the host. These worms most likely inhabit the gill chambers of the host. Hosts. —Orconectes juvenilis (Hagen). Distribution. —Oedipodrilus anisogna- thus is known only from the two localities cited: the type locality is in the Central (Nashville) Basin; the other in the eastern Highland Rim, the uplands surrounding the Nashville Basin. Oedipodrilus macbaini (Holt, 1955) Figs. 4, 5 Cambarincola macbaini (Holt 1955:27-—31). Oedipodrilus macbaini (Holt 1969:205; 1984:39). VOLUME 101, NUMBER 4 Type specimens. —Holotype, USNM 25952, six paratypes, PCH 134, taken from Charles Creek, eight miles west of Ashland on State Highway 5, Boyd County, Ken- tucky, on Orconectes sp. by Rodney MacBain, Jul 1948 (Holt 1955:29). Diagnosis (emended).—Small- to medi- um-sized worms (average length of five specimens of type series and five specimens from Powell County, Kentucky, 1.8 mm); lips entire; no oral papillae; no dorsal ridges; jaws small, dental formula 5/4; bursa ex- ceeds body diameter in length; penial sheath three to four times length of bursal atrium; penis long cuticular tube furnished with re- curved hooks, coiled within lumen of penial sheath; ejaculatory duct proportionately short, thick, muscular; spermiducal gland about 73 body diameter in length, its di- ameter about '2 its length; prostate short, lying along ental third of spermiducal gland; non-differentiated, with ental bulb; sper- matheca approximately equal to bursa and penial sheath in length, composed of long ectal duct, narrow median region, elongate bulb. Etymology.—For the collector, Rodney G. MacBain. Description. — Five specimens of the type series, collected in 70 percent ethanol have the following average dimensions: total length, 1.2 mm; greatest diameter, 0.3 mm; head length, 0.2 mm; head diameter, 0.1 mm; diameter, segment I, 0.1 mm; diam- eter, sucker, 0.2 mm (Holt 1955:27-28; modified to conform to current usage). In comparison, five specimens from Powell County, Kentucky, have the following av- erage dimensions: total length, 2.4 mm; greatest diameter, 0.3 mm; head length, 0.3 mm; head diameter, 0.2 mm; diameter, seg- ment I, 0.2 mm; diameter, sucker, 0.3 mm. The discrepancies in these measurements probably result from differences in the fluids used in collecting: material taken since 1958 by Holt has been fixed in a solution com- posed of five parts of formalin to 95 parts of 70 percent ethanol. The latter fluid is far superior to ethanol alone, in part because 801 of a lesser degree of contraction of the an- imals. The lips are entire; there are no oral pa- pillae. There is one internal pharyngeal sul- cus, but externally there is little indication (by an external sulcus) of its location. There are no supernumerary muscles of the dor- sum of any body segments: the body outline is smooth. The jaws present no distinctive features: they are small, delicate in appearance and light in color. The dental formula is 5/4. The gut is markedly narrowed in segments V and VI: most of the space in these seg- ments is occupied by the spermatheca and male reproductive organs. Diatoms and de- tritus constitute the food found in the gut. The anterior nephridiopore, undetectable in the entire specimens of the type series (Holt 1955:28) is, as determined by study of sectioned animals of the material from Powell County, Kentucky, a single minute opening on the dorsum of segment III. The spermiducal gland is unremarkable, about twice its diameter in length and usu- ally lying in a vertical position in the an- terior portion of segment VI. The prostate, composed of cells similar to those of the spermiducal gland, is about 3 the diameter and length of the latter and arises entad to the junction of the spermiducal gland and ejaculatory duct (Fig. 4b). It has a small ental bulb in the Powell County animals which is not apparent in the paratypes. This discrepancy may be attributed to the con- tracted state of the latter. The ejaculatory duct, as determined from sections (Fig. 5c) is a relatively short, thick and muscular tube distinctive only in the thickness of its inner layer of circular mus- cles. The bursa is a very long, cylindrical tube that is variously bent as it courses dorso- laterad over the gut to its union with the ejaculatory duct. The bursal atrium is com- posed of a moderately thick inner layer of circular muscles and an outer thinner one of longitudinal (in reference to the organ) muscles and encloses what is actually the 802 Cc PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Oedipodrilus macbaini (Holt, 1955), specimens from Powell County, Kentucky. a, Lateral view of entire animal with everted penis; b, Lateral view of reproductive systems (after Holt 1955); c, Section through ental end of spermathecal bulb; d, Cross section of ectal duct of spermatheca; e, Section through outlet pore of spermatheca. portion of the penis that is not enclosed in the penial sheath in its retracted state (Fig. 5a). Most of the bursa consists of the penial sheath whose wall is apparently composed of the same, but much thinner layers as is the bursal atrium. Its capacious lumen con- tains the loops of the cuticular penis (Figs. 5b, c). These dimensions of the bursa (atrium and penial sheath) accommodate the inor- dinately long cuticular penis. When com- pletely everted the penis may actually be longer than the animal itself (Fig. 4a shows the penis much foreshortened). Retracted, the penis appears to consist of a thin layer of tissue with a cuticular lining. These layers are continuous with those composing the penial sheath and ejaculatory duct (Fig. 5a, b, d). When everted, the ectal portion of the penis is provided with recurved hooks (Figs. 4a, 5d, e) that lie within the lumen of the uneverted penis (Fig. 5d, e) which are absent from its ectal- and ental-most portions (Fig. 5a, b). Such features of the penis, as might be expected, are matched by the structure which receives it in copulation: the spermatheca. There is a short ectal pore canal composed of the same layers as the body wall and the bursa (Fig. 4e) which passes shortly entad into the long and expanded ectal duct prop- er of the spermatheca (Fig. 4b, d, e). This duct courses dorsad, loops over the gut and is itself at least twice as long as the body diameter. The inner layer of the spermathe- cal duct is composed of cells with processes extending radially into the lumen of the duct (Fig. 4d). The spermatheca ends entally in a large, ventrally directed ovoid bulb. Affinities. —See the discussion of the af- finities of O. anisognathus above. Variations. —The loops and bendings of the penial sheath and spermatheca take var- ious positions within their segments. A careful comparison of paratypes with the other specimens herein assigned to the species lead to the conclusion that the dif- ferences in size and the apparent absence of VOLUME 101, NUMBER 4 803 Fig. 5. Oedipodrilus macbaini, specimens from Powell County, Kentucky. a, Optical section through bursal atrium and ecctal end of retracted penis; b, Optical section through ental end of penial sheath portion of bursa and penis; c, Optical cross section of ejaculatory duct; d, Optical cross section of penial sheath; e, Optical cross section of penis. an ental bulb of the prostate in these type specimens is accounted for by the differ- ences in fixation (see above). Hosts.—See ‘Material Examined” low. Distribution. —Oedipodrilus macbaini 1s widely dispersed throughout the Ohio River drainage in Tennessee, Kentucky, Ohio and Pennsylvania, and the Illinois River in II- linois. It is, therefore, a member of the di- verse branchiobdellidan fauna that is de- rived from the pre-Pleistocene inhabitants of the Cumberland River and ultimately the ancient Teays drainages that has followed the retreating ice northward (Hobbs et al. 1967:69 et seq.; Holt 1968b:5, 1969:199). Material examined.—(Unless otherwise noted all collections were taken by Perry C. and Virgie F. Holt). Illinois. Vermillion County, 9.2 miles north of Danville on U.S. Highway 136, on Orconectes propinquus (Girard), O. virilis (Hagen), O. immunis be- (Hagen), 25 Jul 1958. Indiana. Montgomery County, 6.4 miles north of junction of In- diana highways 47 and 234, 24 Jul 1958, on unknown host, PCH 802; Orange Coun- ty, 8.3 miles north of Crawford County line on Indiana Highway 145, 26 Jul 1958, on Cambarus laevis Faxon, O. virilis, PCH 812; Parke County, 5.6 miles west of Bellmore on U.S. Highway 36, 26 Jul 1958, on O. immunis, O. propinquus, PCH 807; Parke County, 4.2 miles north of Rockville on U.S. Highway 41, on unknown host, 26 Jul 1958, PCH 806; White County, 2.6 miles north of junction of U.S. highways 421 and 24 at Reynolds, 24 Jul 1958, on O. virilis, O. pro- pinquus, PCH 801. Kentucky. Boyd Coun- ty, 8 miles west of Ashland from Charles Creek on Kentucky Highway 5, Jul 1948, on Orconectes sp., R. G. MacBain, PCH 134. [Type locality]; Carter County, 1.9 miles northeast of Carter-Rowan County line on U.S. Highway 60, on unknown host, 804 23 Apr 1961, S. E. Neffand P. C. Holt, PCH 1360; Clarke County, 4.6 miles south of Winchester from Howard Creek on U.S. Highway 227, 30 Jul 1958, on O. rusticus (Girard), PCH 840; Fleming County, 1.5 miles northeast of Cowan on Kentucky Highway 32, on unknown host, 20 Apr 1961, S. E. Neff and P. C. Holt, PCH 1357; Har- rison County, 0.5 mile northwest of Renak- er on Kentucky Highway 36 from South Fork of Raven Creek, on O. rusticus, C. b. bartonii, 27 Jun 1964, J. F. Fitzpatrick, PCH 1788; Jackson County 1.8 miles south of junction of Kentucky Highway 30 and 1190, 29 Jul 1958, on O. sp., PCH 831; Johnson County, 6 miles northeast of Paintsville on U.S. Highway 460, on C. b. bartonii, 6 Aug 1961, P. C. Holt, PCH 1454 (USNM 37799); Madison County, 9.3 miles north of Rich- mond on U.S. Highway 227, on unknown host, 30 Jul 1958, PCH 841 (USNM 37175); Nicholas County, at Meyers, on unknown host, 20 Apr 1961, S. E. Neffand P. C. Holt, PCH 1358; (USNM 37565); Powell County, 1.4 miles east of Slade on Kentucky High- way 11-15, on unknown host, 29 Jul 1958, PCH 836 (redescription based on this col- lection); Powell County, 4 miles northeast of Nada on Kentucky Highway 77, on un- known host, 30 Jul 1958, PCH 845. Ohio. Jefferson County, 4.2 miles northeast of Bloomingdale, on U.S. Highway 22, on C. b. bartonii and O. obscurus (Hagen), 7 Aug 1960, PCH 1226; Stark County, 3.8 miles west of Alliance on U.S. Highway 62, 3 Aug 1960, on O. obscurus, PCH 1205; Wayne County, 3.9 miles west of Riceland on U.S. Highway 30, on O. propinquus, 3 Aug 1960, PCH 1204; Williams County, 0.8 mile west of junction of U.S. highways 6 and 127, on O. rusticus, 3 Aug 1960, PCH 1201. Penn- sylvania. Butler County, 4.6 miles northeast of crossing of Pennsylvania Highway 68 and Pennsylvania Turnpike near Zelienople, on C. b. bartonii and O. obscurus, 6 Aug 1960, PCH 1225; Butler County, Wolf Creek at crossing of Pennsylvania Highway 108, 4.9 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON miles northeast of Harlansburg, on O. ob- scurus, 6 Aug 1960, PCH 1224; Erie Coun- ty, French Creek at junction of U.S. high- ways 6 and 19, on O. obscurus, 5 Aug 1960, PCH 1216; Erie County, 2.6 miles west of Union City on U.S. Highway 6, on C. Db. bartonii and O. obscurus, 5 Aug 1960, PCH 1215. Tennessee. Putnam County on Ten- nessee Highway 56 at Baxter, on unknown host, 25 Jul 1961. Notes.—The streams from which collec- tions were taken by P. C. and V. F. Holt were all small- to medium-sized ones in which the hosts were taken by hand or with a dipnet. Most are in hilly, wooded country, but some (in Ohio) are in the plains on gla- cial silt. The gut contents of O. macbaini include diatoms and the usual detritus that adheres to the exoskeleton of the hosts. Sathodrilus rivigeae, new species Fig. 6 Type specimens. —Holotype USNM 119545 and two paratypes, USNM 119546, [PCH 1089] taken from clear cool pools in a medium-sized stream in Ouachita Na- tional Forest, 3.2 miles east of Joplin, Mont- gomery County, Arkansas, at crossing of U.S. Highway 270, on Orconectes palmeri longimanus (Faxon) 23 Jun 1960, by Perry C. and Virgie F. Holt. Diagnosis. —Medium-sized worms (ho- lotype 2.8 mm in length); lips entire, peri- stomium demarcated by distinct sulcus; no oral papillae; no dorsal ridges of body seg- ments; jaws small, delicate, in length 4, head length, dental formula (indeterminant in types) (?)1/4; bursa about '2 body diameter in length, constriction at mid-length marks junctions of bursal atrium and penial sheath; penis eversible, membranous, equal (when retracted) to penial sheath in length, atrial fold present; spermiducal gland large, about %, body diameter in length, with large prominent deferent lobes; prostate subequal in diameter, *%4 in length of spermiducal VOLUME 101, NUMBER 4 805 PY Fig.6. Sathodrilus rivigeae, holotype. a, Lateral view of entire animal; b, Lateral view of reproductive systems; c, Optical section of bursal complex and ejaculatory duct; d, Lateral view of jaws. gland, non-differentiated histologically; spermatheca with ectal duct, about '% total length, bulb elongate ovate. Etymology.—An anagram based on my co-collector’s first name. Description.—The paratypes are some- what smaller than the holotype which has the following dimensions: total length, 2.8 mm; greatest diameter, 0.5 mm; head length, 0.4 mm; head diameter, 0.3 mm; diameter, segment I, 0.4 mm; diameter, sucker, 0.4 mm. The lips lack lobes or tentacles, the peri- stomium is somewhat less in diameter than the greatest diameter of the head and a dis- tinct annular sulcus separates it from the remainder of the head. The prominent in- ternal pharyngeal sulcus is matched exter- nally by a broad shallow one. Oral papillae are absent. There are no dorsal supernu- merary segmental muscles: the body outline is smooth. The jaws are difficult to interpret in the three available specimens: they are small and delicate in appearance, dark in color in the holotype, lighter in the paratypes and in one paratype the lower jaw appears to bear four teeth and the upper only one median tooth. The spermiducal gland is of greatest di- ameter at the junction of the prominent de- ferent lobes and tapers gradually ectad to its union with the ejaculatory duct (the de- ferent lobes and ental portion of the gland are obscured in the illustration). The prostate is unusually large among the 806 members of the genus. It arises from the spermiducal gland about a third of the length of the latter from its ectal end and extends entally to the junction of the deferent lobes. It appears to be composed of the same gran- ular epithelium as the spermiducal gland: it is non-differentiated. There is no obvious ental bulb; but one paratype appears to have a short clear space between the investing peritoneum and its ental-most glandular cells. The bursa is elongate with a broad and deep constriction at its mid-length at the union of the penial sheath and bursal atrium. The atrium has thick muscular walls and the atrial fold is thin and irregular. The peni- al sheath is lined by an epithelium (? mus- cular) and encloses the penis in a commo- dious cavity. The latter is an almost straight cuticular tube with minute longitudinal folds and is likewise covered with a layer of pre- sumably muscular tissue. The diameter of the ejaculatory duct is more than half its total length and its lumen is greatly expanded. It has the structure of a bulb whose function is that of a pump. The spermatheca has a long ectal duct. Its bulb in the holotype is a thickened muscular elongate sac with a narrow lumen. Sper- matozoa may be absent. The spermathecal bulbs of the paratypes are of the more com- mon obovate type without ental processes. Variations. —Two were noted (an unusual number among branchiobdellidans): the dif- ferences in the spermathecal bulbs just not- ed and in the contents of the gut. In the holotype the middle sacculations of the gut contain dark brown particles of unidentifi- able detritus; the guts of the smaller para- types are filled with a homogeneous clear material similar in appearance to that found in gill-inhabiting species; but the jaws are ill-adapted for clipping the gill filaments or piercing the articular membranes of the host. Affinities. —Seven species of Sathodrilus have distinct prostates (as opposed to “‘prostatic protuberances” or “‘bulbs’’). In all of these species the prostate arises from PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the spermiducal gland entad to the junction of the latter with the ejaculatory duct. Of these, S. dorfus Holt 1977, S. lobatus Holt 1977, and S. inversus (Ellis, 1919) have eja- culatory ducts that are short and thickened (“bulb-like’’) and are the closest known rel- atives of S. rivigeae. Sathodrilus lobatus has an expanded peristomium furnished with 14 lobes, its spermiducal gland and prostate are proportionately lesser in diameter than those of S. rivigeae and the spermatheca has a prominent medial bulb and long ental pro- cess (Holt 1977:122-124, fig. 3). Peristo- mial lobes are absent in S. inversus; the sper- miducal gland is noticeably narrowed at its ectal end; the spermatheca has a median bulb and a long spermatozoa storing bulb without a muscular wall or ental process (Holt 1977:128-131, fig. 6, 1981:855). The spermiducal gland and prostate of S. dorfus are short and thick; the spermatheca is com- posed of a long ectal duct, a median bulb and an ental process (Holt 1977:120-121, fig. 2). All three of these species are inhab- itants of the Pacific versant of the north- western United States. Sathodrilus rivigeae differs from them most noticeably in the muscular wall of its spermathecal bulb and is, at least superficially, closest to S. inver- sus, from which it further differs in the short lobes of the peristomium and the frequent divergence of the prostate from the sper- miducal gland (Holt 1977, fig. 7) found in the latter. Branchiobdellidan associates. — Cambar- incola vitreus Ellis, 1918, a species of Cam- barincola that appears to be C. heterogna- thus Hoffman, 1963, and a widespread undescribed Ozarkian species of Xironodri- lus Ellis, 1918. Host.—Orconectes palmeri longimanus. Distribution. —Known only from the type locality. Material examined. —The types. Notes. —Branchiobdellidan associates of two species have been given herein because the data were readily available; it should always be borne in mind that rarely is only VOLUME 101, NUMBER 4 one species of branchiobdellidan found in any given locality. The resemblances of S. rivigeae to its con- geners of the Pacific northwest is another example of the homogeneity of the eastern and western branchiobdellidan faunas of North America, almost totally isolated now by the continental divide. The expanded, bulb-like ejaculatory ducts of the species of Sathodrilus discussed herein should be further investigated. Function is difficult to infer from form alone, but it is probable that, in species with long, eversible penes and long spermathecal ectal ducts, that these expanded ejaculatory ducts aid in pumping spermatozoa into the spermathe- cal bulb. Acknowledgments Drs. Brent D. Opell and Gerald H. Cross have read the manuscript; Dr. Horton H. Hobbs, Jr., in addition, identified the cray- fish hosts. Dr. Ernest R. Stout, Head, De- partment of Biology, Virginia Polytechnic Institute and State University and his sec- retaries, have been generous in their sup- port, financial and otherwise. I am grateful for their help. Literature Cited Ellis, Max M. 1918. Branchiobdellid worms (Anne- lida) from Michigan crayfishes.— Transactions of the American Microscopical Society 37(1): 49-51. . 1919. The branchiobdellid worms in the col- lections of the United States National Museum, with descriptions of new genera and species. — Proceedings of the United States National Mu- seum 55(2267):241-265, figs. 1-18, pls. 10-13. Hobbs, Horton H.., Jr., Perry C. Holt, & Margaret Wal- ton. 1967. The crayfishes and their epizo6tic ostracod and branchiobdellid associates of the Mountain Lake, Virginia, region. — Proceedings of the United States National Museum 123(3602):1-84, figs. 1-22. Hoffman, Richard L. 1963. A revision of the North American annelid worms of the genus Cam- barincola (Oligochaeta: Branchiobdellidae).— Proceedings of the United States National Mu- seum 114(3470):271-317, figs. 1-79. 807 Holt, Perry C. 1949. A comparative study of the reproductive systems of Xironogiton instabilius instabilius (Moore) and Cambarincola phila- delphica (Leidy) (Annelida, Oligochaeta, Bran- chiobdellidae).—Journal of Morphology 84(3): 535-572, pls. 1-4. 1955. A new branchiobdellid of the genus Cambarincola Ellis, 1912, (Oligochaeta, Bran- chiobdellidae) from Kentucky.—Journal of the Tennessee Academy of Science 30(1):27-—31, figs. 1-6. 1960. On a new genus of the family Bran- chiobdellidae (Oligochaeta).—The American Midland Naturalist 64(1):169-176, figs. 1-4. . 1967. Oedipodrilus oedipus, n. g., n. sp. (An- nelida, Clitellata, Branchiobdellida).—Trans- actions of the American Microscopical Society 86(1):58-60, figs. 1-4. . 1968a. New genera and species of branchiob- dellid worms (Annelida: Clitellata).— Proceed- ings of the Biological Society of Washington 81: 291-318, figs. 1-9. 1968b. The genus Prerodrilus (Annelida: Branchiobdellida).— Proceedings of the United States National Museum 125(3668):1—44, figs. 1-12. 1969. The relationships of the branchiob- dellid fauna of the southern Appalachians. Jn Perry C. Holt, ed., The distributional history of the biota of the Southern Appalachians, Part 1: Invertebrates.— Virginia Polytechnic Institute, Research Division Monograph 1:191-219, figs. 1-10. 1973. A summary of the branchiobdellid (Annelida: Clitellata) fauna of Mesoamerica. — Smithsonian Contributions to Zoology number 142:1-111, 1-40, figs. 1-19. 1977. An emendation of the genus Satho- drilus Holt, 1968 (Annelida: Branchiobdellida), with the description of four new species from the Pacific drainage of North America.—Pro- ceedings of the Biological Society of Washington 90(1):116-131, figs. 1-7. . 1981. New species of Sathodrilus Holt, 1968 (Clitellata: Branchiobdellida) from the Pacific drainage of the United States, with the synon- ymy of Sathodrilus virgiliae Holt, 1977.—Pro- ceedings of the Biological Society of Washington 94(3):848-862, figs. 1-3. 1982. A new species of the genus Cambar- incola (Clitellata: Branchiobdellida) from Illi- nois with remarks on the bursa of Cambarincola vitreus Ellis, 1919, and the status of Sathodrilus Holt, 1968.—Proceedings of the Biological So- ciety of Washington 95(2):251-255, fig. 1. 1984. On some branchiobdellids (Annelida: Clitellata) from Mexico with the descriptions of 808 new species of the genera Cambarincola and Oedipodrilus.—Proceedings of the Biological Society of Washington 97(1):35—42, figs. 1-3. . 1986. Newly established families of the order Branchiobdellida (Annelida: Clitellata) with a synopsis of the genera. — Proceedings of the Bi- ological Society of Washington 99(4):676-702, figs. 1-20. Leidy, Joseph. 1851. Contributions to helmintholo- gy.—Proceedings of the Academy of Natural Sciences of Philadelphia 5:205—209. Moore, J. Percy. 1894. On some leech-like parasites of American crayfishes. Proceedings of the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Academy of Natural Sciences of Philadelphia for 1893:419-428, pl. 12. 1895. The anatomy of Bdellodrilus illumi- natus, an American discodrilid.—Journal of Morphology 10(2):497—540, pls. 28.32. Department of Biology, Virginia Poly- technic Institute and State University, Blacksburg, Virginia 24061. Mailing ad- dress: 1308 Crestview Drive, Blacksburg, Virginia 24060. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 809-816 PYCNOGONIDA OF THE WESTERN PACIFIC ISLANDS V. A COLLECTION BY THE KAKUYO MARU FROM SAMOA Koichiro Nakamura and C. Allan Child Abstract.—A small shallow-water collection of Pycnogonida taken by the Japanese training vessel Kakuyo Maru in American Samoa and Western Samoa is described. The collection contains seven species, one of which is new: An- oplodactylus crassus. The species affiliations are discussed along with their limited known zoogeography. Little concerning zoogeographic affinities can be proposed due to lack of pycnogonid collections from most mid-Pacific localities. Pycnogonids of the Samoan Islands have remained unknown until these first records of specimens taken from this locality. There were only seven species collected in two well known shallow-water genera, with six species known and one new to science: Anoplodac- tylus crassus. The six known species have mostly Indo-West Pacific distributions based on what little is known, but one species, A. erectus, has been collected many times in shallow waters along the west coasts of the Americas at least as far south as Co- lombia. This species is also known from the Tuamotus and from Korea, but its apparent main distributional locality of the American coasts makes it the only apparent emigrant from the east in the Samoa collection. One species, A. perforatus, has been taken only in Japanese waters, while Ammothella stau- romata has been taken in the Marshall Is- lands and the northern and southern Phil- ippines, Anoplodactylus rimulus only from the western Indian Ocean, and the other two species, A. arescus and A. glandulifer, have broader known distributions in the Atlantic and Indo-Pacific. The distributional knowl- edge of these species is undoubtedly frag- mentary, as is that of most pycnogonids, and it will be extended for each species as more collecting is carried out in the vast areas still unsampled. None of the depths at which the six known species were taken extends the depth distribution for any of them. Family Ammotheidae Genus Ammothella Verrill, 1900 Ammothella stauromata Child Ammothella stauromata Child, 1982:271- MB ths NEMS Ss T/ Material examined.— American Samoa: Tutuila; Pago Pago Harbor, 3 m, 23 Oct 1985, 2 6 with eggs, 1 3. Distribution. —The type locality for this species 1s Enewetak Atoll, Marshall Islands, and it has also been taken in the Philippines in the south at Negros Island and in the north at Batan Island, Batanes Province. It is now known to inhabit American Samoa. All capture depths are shallow at 0-3 me- ters. Remarks.—This is an easily recognized species in a genus with many similar species. The dorsal trunk tubercles almost match the ocular tubercle and the abdomen size and length, and make this species very distinc- tive. There are several Ammothella species with median trunk tubercles, but none of the others have this tall shape nor does any other species have a tall tubercle on the ce- phalic segment. Other recognition charac- ters are the short first scape segment with small distal tubercles, anterolateral cephalic 810 segment tubercles, and dorsodistal tubercles on the lateral processes. None of these char- acters is unique but their combination in this species makes it unique. Family Phoxichilidiidae Genus Anoplodactylus Wilson, 1878 Anoplodactylus arescus du Bois-Reymond Marcus Anoplodactylus arescus du Bois-Reymond Marcus, 1959:105-107, pl. 21.—Stock, 1968:53 [text]; 1975:133, figs. 10-12.— Arnaud, 1973:954.—Child, 1988a:12. Material examined. —Western Samoa: Apia; Upolu Island, 10 m, 25 Oct 1985, 1 6,42. Distribution. —This species was first tak- en in the Red Sea in sand and has had sub- sequent captures in Madagascar and Tan- zania. It was recently found in the southern Philippines (Child 1988a), and is now also known from the mid-Pacific in Samoa, much farther east than previous records. This Sa- moan record marks the deepest capture for this species at 10 meters. The substrate at this locality is fine coral sand with small rubble bits. This species is one of the few pycnogonids known to prefer fine sand as a regular habitat. A few other Anoplodactylus species have been taken repeatedly from sand and most species of the genus Rhyn- chothorax are also known from this kind of substrate in at least one or more records. Remarks. —These specimens have a slightly shorter neck, a second smaller heel spine proximal to the main spine, and the proboscis is more cylindrical than the swol- len proboscis of the type. These specimens agree very well otherwise with the figures of the type. The integument is very papillose in all specimens examined. This is a very Fig. 1. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tiny species and is recognized by its distally extended tarsus, a round pad of integument at the heel base, a full propodal lamina, and a lack of any auxiliary claws, all characters shared by the 5 Samoan specimens. Anoplodactylus crassus, new species Fig. 1 Anoplodactylus viridintestinalis (Cole).— Kim, 1986:3—5, fig. 2.—Kim & Hong, 1986:44.—Hong & Kim, 1987:161. Material examined.— American Samoa: Tutuila; Pago Pago Harbor, 3 m, 23 Oct 1985, holotype 6 with eggs (USNM 234393), paratypes, 2 6 with eggs, 2 2, 1 juav (USNM 234394). Description. —Size moderately small, leg span 8.3 mm. Trunk compact, circular in outline, unsegmented, lateral processes con- tiguous proximally, slightly separated dis- tally, each armed with small conical dor- sodistal tubercle bearing seta. Neck short, flanked by large bulges denoting palp ru- diments. Ocular tubercle as tall as its di- ameter, capped by low anterodistal tubercle, eyes large, well pigmented. Proboscis short, with low midventral swelling. Abdomen broad in lateral view, only slightly longer than ocular tubercle, tapering distally, armed with 7-8 short distal setae. Chelifore scape slender, 5 times longer than its diameter, armed with 3 low dorsal tubercles each bearing seta. Chela palm nar- row, movable finger as long as palm, armed with several tiny serrate teeth and 3 ectal setae. Immovable finger shorter, more slen- der, armed with 2 tiny teeth. Fingers well curved, movable finger with greater curve, overlap at tips. Oviger short, setose, second to fifth seg- ments armed with lateral and distal setae, = Anoplodactylus crassus, holotype male: A, Trunk, dorsal view; B, Trunk, lateral view; C, Chela; D, Third leg with enlargement of cement gland tube; E, Leg terminal segments, enlarged; F, Oviger with several attached eggs; G, Oviger terminal segments, enlarged in lateral and ectal views. Paratype female: H, Trunk, dorsal view. VOLUME 101, NUMBER 4 811 a eed L OSs. re 812 sixth with dense field of short lateral setae on one side only. Second segment subequal to third which has proximal constriction, fourth slightly longer than fifth, sixth a short cone. Legs moderately short, robust, lightly se- tose. First coxa almost length of third, first armed with 2-3 short dorsodistal tubercles bearing apical setae. Femur the longest seg- ment, cement gland a single short truncate cone with broad tubular tip. First tibia long- er than second, both armed with low dorsal bulges bearing setae. Tarsus triangular with many short endal setae. Propodus with strong heel armed with 2 short stout spines and 3 setae. Sole with many short setae flanking long lamina on entire sole length. Claw robust, well curved, about 0.6 pro- podal length, auxiliaries entirely lacking. Female slightly larger than male, dimor- phic in trunk shape. Posterior pair of lateral processes well separated from third pair im- parting a more ovoid shape to trunk in dor- sal outline. Lateral process tubercles and those of first coxae reduced or lacking with corresponding reduction of coxal setae. Measurements (in mm).—Trunk length (chelifore insertion to tip 4th lateral pro- cesses), 0.86; trunk width (across 2nd lateral processes), 0.8; proboscis length, 0.38; ab- domen length, 0.29; third leg, coxa 1, 0.26; coxa 2, 0.37; coxa 3, 0.29; femur, 0.71; tibia 1, 0.6; tibia 2, 0.55; tarsus, 0.14; propodus, 0.51; claw, 0.34. Distribution.— Known from the type-lo- cality, Pago Pago, American Samoa, in 3 m, and from the Korean coast in littoral depths. Etymology. — The species name crassus is Latin (thick, fat, or stout) and refers to the thick or stout appearance of these speci- mens. Remarks. — This new species is very close to and forms a geminate pair with Anoplo- dactylus viridintestinalis (Cole, 1904). The two species are difficult to separate by su- perficial examination but close investiga- tion of both sexes in each species shows PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON many small differences between the eastern and the western Pacific species. Taken alone (as with Kim 1986), these differences could be attributable to variation in populations but examination of the cumulative effect of the many small differences suggests that iso- lation has produced speciation from what possibly was the original stock on the Amer- ican west coast. A number of specimens of A. viridintestinalis from California (Fig. 2A— F), and the female holotype appendages were examined for comparison with the Samoan specimens. The trunk and its additional ap- pendages in alcohol (not on slides made by Cole) were lost many years ago. The mainland American specimens show almost no variation among themselves and are consistently different from A. crassus in the following characters. The proboscis of A. crassus 1s shorter in length and thicker in diameter than the mainland species. The trunk shape 1s always broader in dorsal view in males than in A. viridintestinalis, and the chelifore scape has 3 low setose tubercles in almost all specimens rather than one or none, the chelae are longer and more slender or rectangular in shape, the abdomen is broader, carried at less of an erect angle, and has more distal setae, the lateral process tubercles each bear a seta at the tip rather than anterolateral to the tubercle itself, the 3 laterodistal tubercles of the first coxae are larger while the mainland species may have only one or none, the ocular tubercle is more rounded distally with a small cap tubercle while Cole’s species has a pointed conical ocular tubercle, and the ventrodistal sex pore tubercle on the second coxae is low and in- conspicuous while it is as long as half the segment diameter in A. viridintestinalis. The cement gland tube, although it has the same bottle shape and location on the femur, is shorter in length than that of Cole’s species. It is in the setation of the oviger terminal segment that the biggest difference is no- ticeable. In the new species, there is a field of many short setae, shorter than the max- VOLUME 101, NUMBER 4 imum segment diameter, which occurs only on one side of the conical segment. On the terminal oviger segment of Cole’s species, there are 6 to 8 setae, each longer than the maximum segment diameter, placed ran- domly on its endal and lateral surfaces. Sexual dimorphism of the lateral process placement on the trunk is evident in both species but the female proboscis of main- land specimens invariably has a rather large proximoventral bulge which is not found in A. crassus females. It would be rather difficult to separate these two species were specimens of both not at hand with which to compare the dif- ferences. The set of figures (Fig. 2A—F) of A. viridintestinalis from California is there- fore provided so that these differences can be more readily seen. Anoplodactylus erectus Cole Anoplodactylus erectus Cole, 1904:289-291, pl. XIV, fig. 12, pl. XXVI, figs. 1-9.— Child, 1970:288—289 [early literature]; 1979:52.—Kim & Hong, 1986:41, fig. 5.— Hong & Kim, 1987:161. Material examined.— American Samoa: Tutuila, Pago Pago Harbor, 3 m, 23 Oct 1985, 2 6 with eggs, 1 @. Distribution. —This species has long been known to inhabit shallow waters from Brit- ish Columbia, Canada, to Colombia, and it has also been taken in Hawaii, the Tuamotu Islands, and in Korea. This record extends its South Pacific distributional limits west- ward to Samoa, but adds nothing new to its shallow depth records. Remarks. — Males of this species are eas- ily recognized by the long subcutaneous ce- ment gland tube extending almost to the proximal end of the femur, dorsodistal lat- eral process turbercles of varying lengths, the long third oviger segment, and the two short spines on the well formed heel of the propodus. The chelae are short and the overlapping fingers have no teeth. Female 813 specimens, as with almost all females of this genus, need accompanying males which bear the critical characters essential for positive identification. Anoplodactylus glandulifer Stock Anoplodactylus glandulifer Stock, 1954:80- 84, fig. 36.—Child, 1982:273-274 [liter- ature]; 1988b:58—-59. Material examined.— American Samoa: Tutuila; Pago Pago Harbor, 2 m, 23 Oct 1985, 1 6 with eggs, 1 juv. Distribution. — This species has been tak- en in a number of Indian Ocean localities from Kenya and the Red Sea to Singapore, and it has been found at Enewetak Atoll, Marshall Islands. This is the first South Pa- cific record, but it probably will be found to inhabit many other western Pacific lo- calities. The Samoan capture depth con- tributes nothing new to the known depths of intertidal to 5 meters for the species. Remarks.—The 3 small cement gland cups on each femur serve to distinguish this species from most others known, although it is not the only species to possess multiple cement gland outlets. It has no other out- standing characters except for the closely spaced lateral processes and a long propodal lamina, making isolated females difficult to impossible to distinguish without accom- panying males. This species is little different from several other Pacific species except for the cement gland cups which may number from 2 to 4 per femur on the same specimen and for the very long propodal lamina which is another unusual character. The two char- acters serve to separate this species from any other small but similar species in the west- ern Pacific. Anoplodactylus perforatus Nakamura & Child Anoplodactylus perforatus Nakamura & Child, 1982:289-291, fig. 3; 1983:49. Fig. 2. Anoplodactylus viridintestinalis, California male: A, Trunk, dorsal view; B, Trunk, lateral view; C, Oviger terminal segments, enlarged; D, Third leg, cement gland tube enlarged. California female: E, Trunk, dorsal view; F, Proboscis, lateral view. Anoplodactylus rimulus, Samoan male: G, Trunk, lateral view; H, Oviger, enlarged; I, Femur, cement gland tube enlarged. VOLUME 101, NUMBER 4 Material examined.— American Samoa: Tutuila; Pago Pago Harbor, dredged in 15 m, 22 Oct 1985, 1 6. Western Samoa: Upolu Island; Apia, dredged in 10 m, 25 Oct 1985, 15 6 with eggs, 18 6, 44 9, 1 juv. Distribution. —This species is very com- monly found in large numbers in southern Sagami Bay, Japan (type locality), in depths of 7-15 and 113 meters. Its capture in 2 localities in the Samoa Islands greatly ex- tends its known distribution eastward and to the South Pacific, without expanding its known depth range. The single capture of 78 specimens in Apia emphasizes its ten- dency to be found in large groups, the same as in many of the Sagami Bay captures. Nothing can be said concerning this gath- ering of specimens except that sufficient suitable food must be available for such an aggregation to remain grouped in a small restricted area. Remarks.—This is another species in which males are made distinctive and easily identified by the cement gland openings which number from 17 to 25 pores on each femur, surpassing any other known species in number, and the very long oviger bearing only a tiny terminal segment. Similar species have a terminal segment half as long or longer in relation to the fifth segment, and always have far fewer cement gland open- ings. Other recognition characters are the very short propodal lamina, measuring less than 0.1 of the sole length, and the long chelae fingers without teeth. Anoplodactylus rimulus Child Fig. 2G-I Anoplodactylus rimulus Child, 1988b, fig. 3. Material examined.— American Samoa: Tutuila; Pago Pago Harbor, 3 m, 23 Oct 1985, 2 6 with eggs, 1 4, 2 2, 1 juv. Supplemental description.—(Male) Pro- boscis tapering only from 0.77 length dis- tally, with marked single proximoventral bulge lacking cleft. Femur less inflated, with cement gland from proximal rim to median dorsal point and cement gland tube, a long 815 straight tube half femoral diameter situated just proximal to median point. Oviger fairly short, second, third and fourth segments with few short randomly placed setae, fifth with many setae equal to segment diameter placed in rows, sixth a tiny cone with 6—7 setae circling cone, setae as long as those on fifth segment. Distribution.—The type locality for this species is Aldabra Atoll, Seychelles, Indian Ocean, in 3-6 meters. This is the second record for the species and it greatly extends the known distribution to the east in the mid-Pacific at Samoa. The depth of capture of the Samoan specimens is within that of the Aldabra records. Remarks.—The elaborate proximoven- tral tubercles of the females in this species are not carried over to the male to as great an extent. This is true for the so called “‘alar processes” of many females in this genus, but the male of this species, now known from the Samoan material, does have a sin- gle proximoventral bulge on the proboscis. The female proboscis bulge of the Samoan specimens is not as large or elaborate as those of the Aldabra Atoll specimens. It consists of a bulge with a barely perceptible longitudinal cleft but lacks the lateral cleft of the type specimens. All other female characters appear to fall within acceptable bounds of variation as the differences are only slight between the two sets of speci- mens. The male was previously unknown, but aside from the male characters of oviger and cement gland, it is very similar to the female including length of propodal lamina, heel and sole spination, leg segment lengths, chelae characters, and lateral process tu- bercles. These tubercles are slightly smaller in the female, as would be expected. The grossly clubbed chelifore scape of this species is rare in the genus and affords a good rec- ognition character along with the ventral proboscis bulges of this tiny species. Acknowledgments We wish to thank Professor Shigeo Abe, Captain of the fisheries training vessel Ka- 816 kuyo Maru for his many kindnesses while one of us (K.N.) was aboard, to Mr. Hiroshi Yoshimura, Mr. Hideaki Nishida, and the other members of the crew for their great help in collecting the specimens reported on here, and to the Department of Fisheries, University of Nagasaki, for permission to participate in the fisheries training cruise aboard the vessel. We also wish to acknowl- edge the constructive comments of Thomas E. Bowman and his excellent editing of the manuscript. The specimens are deposited in the col- lections of the National Museum of Natural History, Smithsonian Institution, under the catalog numbers of the old United States National Museum (USNM). Literature Cited Arnaud, F. 1973. Pycnogonides des récifs coralliens de Madagascar, 4. Colossendeidae, Phoxichili- diidae et Endeidae.— Tethys 4(4):953-960, figs. 1-8. Child, C. A. 1970. Pycnogonida of the Smithsonian- Bredin Pacific Expedition, 1957.— Proceedings of the Biological Society of Washington 83(27): 287-308, figs. 1-5. 1979. Shallow-water Pycnogonida of the Isthmus of Panama and the coasts of Middle America.—Smithsonian Contributions to Zo- ology 293:1-86, figs. 1-25. . 1982. Pycnogonida of the western Pacific Is- lands I. The Marshall Islands.— Proceedings of the Biological Society of Washington 95(2):270- 281, figs. 1-3. 1988a. Pycnogonida of the western Pacific Islands III. Recent Smithsonian-Philippine Ex- peditions.—Smithsonian Contributions to Zo- ology 468:1-32, figs. 1-12. 1988b. Pycnogonida from Aldabra Atoll.— Bulletin of the Biological Society of Washington 8:45-72, figs. 1-9. Cole, L. J. 1904. Pycnogonida of the West Coast of North America. — Harriman Alaska Expedition 10:247-298, pls. 11-26. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hong, J.-S., & I. H. Kim. 1987. Korean pycnogonids chiefly based on the collections of the Korea Ocean Research and Development Institute. — Korean Journal of Systematic Zoology 3(2):137— 164, 17 figs. Kim, I. H. 1986. Four pycnogonid species new to Korean fauna. — Proceedings of the Natural Sci- ence Research Institute, Kangreung National University 2(1):1-9, 4 figs. ——., & J.-S. Hong. 1986. Korean shallow-water pycnogonids based on the collections of the Ko- rea Ocean Research and Development Insti- tute.—Korean Journal of Systematic Zoology 2(2):35-52, 9 figs. Marcus, E. du B.-R. 1959. Ein neuer Pantopode aus Foraminiferensand.—Kieler Meeresforschun- gen 15(1):105-107, pl. 21. Nakamura, K., & C. A. Child. 1982. Three new species of Pycnogonida from Sagami Bay, Japan. —Pro- ceedings of the Biological Society of Washington 95(2):282-291, 4 figs. >, 6% . 1983. Shallow-water Pycnogonida from the Izu Peninsula, Japan.—Smithsonian Contributions to Zoology 386:1-71, 21 figs. Stock, J. H. 1954. Pycnogonida from the Indo-West Pacific, Australian and New Zealand waters. — Videnskabelige Meddelelser fra Dansk Natur- historisk Forening (Kobenhaven) 116:1-168, 81 figs. . 1968. Pycnogonida collected by the Galathea and Anton Bruun in the Indian and Pacific Oceans. — Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening (Kobenhaven) 131:7— 65, 22 figs. 1975. Biological results of the University of Miami Deep-Sea Expeditions, 108. Pycnogon- ida from the continental shelf, slope, and deep- sea of the tropical Atlantic and east Pacific.— Bulletin of Marine Science 24(4):957-1092, 59 figs. (KN) Ocean Research Institute, Univer- sity of Tokyo, 1-15-1, Minamidai, Nakano- ku, Tokyo 164, Japan; (CAC) Department of Invertebrate Zoology (Crustacea), Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 817-824 DARWINULOCOPINA (CRUSTACEA: PODOCOPA), A NEW SUBORDER PROPOSED FOR NONMARINE PALEOZOIC TO HOLOCENE OSTRACODA I. G. Sohn Abstract. —The new suborder Darwinulocopina is proposed in the order Po- docopida Sars, 1865, superorder Podocopa Kesling, 1981 to include living and fossil ostracode genera. The new category is coordinate with the Cypridocopina Kozur, 1972 and the Cytherocopina Grindel, 1967. Diagnostic characters of the Darwinulocopina are a posterior process instead of furcae and furcal at- tachments, and a rosette adductor muscle attachment scar comprised of large stigmata. The geologic range of the Darwinulocopina is Paleozoic to Holocene. The superfamily Darwinulacea Brady & Norman, 1889, contained, until after 1961, only one family, the Darwinulidae Brady & Norman, 1889, which in turn consisted of one genus: Darwinula, based on the type species Darwinula stevensoni, (Brady & Robertson 1870) (Swain 1961:Q253). Until 1972, the Darwinulacea Brady & Norman, 1889 have been classified in the suborder Podocopina Scott, 1961:Q86-—Q88, in the order Podocopida Sars, 1865. Grundel (1967:324) discussed the classification of the Podocopida in the Treatise (Moore 1961), in which he included 3 suborders: Meta- copina Sylvester-Bradley, 1961:358 (type genus Quasillites Coryell & Malkin, 1936, p. 18); Bairdiocopina new suborder (type genus Bairdia McCoy, 1844:164); and the Cytherocopina new suborder (type genus Cythere O. F. Miller, 1785, p. 49). The sub- order Bairdiocopina Grundel, 1967, includ- ed the Bairdiacea Sars, 1885. Cypridacea Baird, 1845, and the ?Darwinulacea. Kozur (1972:13-—15) referred the Darwin- ulacea to his emended suborder Cyprido- copina Jones, 1901 [Nom. transl. ex Cypri- dida Jones in Chapman, 1901:147, 154]. Kozur considered the suborder Metacopina Sylvester-Bradley, 1961, to be a junior syn- onym of the Cypridocopina (1972:13), to which he referred both the Darwinulacea and the Metacopina, and suggested that Darwinula Brady & Norman, 1889, devel- oped from Carbonita because ‘‘The stem group [of Darwinula] is probably the genus Carbonita, that has hitherto been referred to the Cypridacea. According to Pollard (1966) Carbonita claripunctata has some- times a rosette-like arrangement of the ad- ductor muscle scars (similar to Darwinula). Carbonita is also very similar in outline to Darwinula, also in that the right valve is larger than the left in this genus. Therefore Carbonita should be referred to the Dar- winulacea.” [my translation]. Kesling (1981) also referred the Darwinulacea to the Cy- pridocopina, but recognized the Metacopi- na as an order in the superorder Podocopa. Kesling’s (1981, p. 291, 300, 308) recom- mended classification of the Podocopida Sars, 1865, the superorder Podocopa is fol- lowed herein. The Darwinulacea should be excluded from the Cypridocopina based on both shell morphology and soft parts; consequently, the new suborder Darwinulocopina (type genus Darwinula Brady & Robertson, in Jones 1885:346), is proposed for the living and fossil taxa that have been referred to the Darwinulacea (Sohn 1987:151). 818 Superorder Podocopa Sars, 1865 Kesling (1981:291) elevated the orders in Moore (1961) to superorder categories, and included the orders Podocopida Sars, 1865, p. 10, and the Metacopida Sylvester-Brad- ley, 1961, p. Q358 in the Podocopa. Order Podocopida Sars, 1865 Kesling (1981:291) included in this order the suborders Bairdiocopina Grundel, 1967, Cypridocopina Jones (emend. Kozur, 1972) to include the superfamilies Cypridacea Baird, 1845, and Darwinulacea Brady & Norman, 1889, and the Cytherocopina Grindel, 1967. He described the Podocop- ida as follows: ““‘Duplicature present except perhaps in earliest forms, typically large an- teriorly and posteriorly and there set off from outer lamella by a vestibule. Contact groove not well developed if at all, at most a simple depression in inner lamella [infold].” (Kes- ling 1981:300). The new suborder Darwinulocopina is proposed for the Darwinulacea to remove them from the Cypridocopina. The new suborder is coordinate with both the Cy- pridocopina and the Cytherocopina. The distinguishing characters among the Dar- winulocopina, the Cypridocopina, and the Cytherocopina are listed in Table 1. Darwinulocopina, new suborder Diagnosis. —Small, smooth, elongate Po- docopida; without any nodes, ridges, sulci; with very narrow normal and radial pore canals, without calcified inner lamella on infold. Hinge simple, overlap variable; out- line of the adductor muscle-attachments scars (AMAS) pattern circular, consists of large individual scars (stigmata sensu Gramm 1982:201). Appendages of the type genus consist of pediform antennules and antennas; first thoracopod with respiratory plate; second and third thoracopods pedi- form: with posterior process; furcae lacking. Discussion.—Howe, Kesling, & Scott PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (1961: Q112-Q14) described “‘furca”’ as fol- lows: ‘‘Furcae (furcal rami of some authors) are appendage-like structures attached to the posterior end of the body .... In the Po- docopina the furcae are extremely variable .... In the Darwinulidae (fig. 8a) they are lacking or represented by a reflex process at the end of the thorax...” Rome (1969) described and illustrated the furcal attachments in different subfamilies in the Cypridae Baird, 1845. The Darwin- ulocopina differ from the Cypridocopina in having a unique AMAS, in absence of furcae and furcal attachments, in having a poste- rior process, and in lacking a wide calcified inner lamella on the infold. Living darwinulids differ from living Cy- pridocopina in having pediform antennules and antennas without swimming setae (Sohn 1987:154, text fig. 1, pl. 1, figs. 2, 4, 16), in absence of furcae and furcal attachments, and in presence of a posterior process (Sohn 1987:105, fig. 1). The darwinulids differ from the Cytherocopina in having smooth in- stead of ornamented carapaces, in simple instead of complex hingement, in round AMAS outline, in stigmata symmetrically arranged (Sohn 1987: figs. 12, 17-19) in- stead of in a single vertical row. Triebel (1941:219, fig. 48) illustrated with a photograph an AMAS of D. stevensoni with an accessory (frontal) scar in front and below the AMAS, and this scar was illus- trated in Moore (Swain 1961:Q253, fig. 183, 2c). Sharapova (1947:215, fig. 53), illus- trated with a drawing the frontal scars in Darwinula, and Starozhilova (in Lipatova & Starozhilova 1968:80, fig. 20a) published a drawing of two frontal scars of Darwinula. Frontal scars had not been recorded in other fossil and extant specimens of D. stevensoni until Swain (1986:pl. 4, fig. 14b) illustrated frontal scars on a Pliocene specimen from Idaho. However, I cannot explain why these scars have not been recorded in the nu- merous SEM micrographs of valves of liv- ing specimens of Darwinula (Keyser 1975; Sohn 1976, 1987). VOLUME 101, NUMBER 4 819 Table 1.—Comparison of selected diagnostic morphological features among the Darwinulocopina, the Cypridocopina, and the Cytherocopina. Darwinulocopina Cypridocopina Cytherocopina Shape Elongate-ovate variable variable Surface smooth or with ornamented ornamented ventroposterior or smooth or smooth spine Hinge simple simple complex Overlap narrow wide narrow or wide Vestibule absent or present present present Outline of AMAS rosette disorganised vertical row Number of spots 6-12 7 or less 4or5 Frontal & man- absent or present present present dibular scars Eye tubercle absent absent present or absent Appendages Antennules pediform natatory pediform (Sohn 1987:pl. 2, except candonids figs. 1-6 (Moore 1961: fig. 137, 1a) Antenna, endopodite pediform natatory pediform (Sohn 1987:pl. 2, except candonids with flagellum figs. 7-13) (Moore 1961: fig. 137, 2a) Antenna, exopodite vestigial vestigial flagellum with secreting duct (Moore, 1961: fig. 137, 3a) Posterior process Present absent absent Furcal rami absent present or reduced absent Reproduction asexual, ?sexual sexual, rarely asexual sexual Stratigraphic range.—Carboniferous to Holocene. Superfamily Darwinulacea Brady and Norman, 1889 Darwinulacea Swain, 1961:Q253. Diagnosis.—A superfamily in Darwinu- locopina with small (less than 1 mm), elon- gate, relatively thin shelled, smooth cara- paces; gently convex dorsal margins; simple hinges; narrow overlap, AMAS outlines round, with large stigmata. Family assigned.—Darwinulidae Brady & Norman, 1889. Discussion. —Only the Darwinulidae Bra- dy & Norman, 1889 and the Microdarwin- ulidae Kashevarova & Neustrueva, 1982 are based on living genera; the other families listed in Sohn (1987:151, table 1) are based on fossil taxa, and their referral to the Dar- winulacea is based on published data and is subject to change. Except for Whipplella Holland, 1934 (type species (original des- ignation) W. cuneiformis Holland, 1934:344, pl. 25, fig. 5. Permian), all the fossil genera in the list are based on European and Asiatic specimens that cannot be forwarded for study in the U.S. The only exception are two topotypes of Panxiania subelliptica Wang, 1978, the type species of the nomi- nate genus of the family Panxianidae Wang, 820 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1980, that Dr. Wang donated. The genus Vymella Kalis & Mischina in Mischina & Kalis 1975 (type species (original designa- tion) V. nazarovae Kalis & Mischina: 77, pl. 9, figs. 1, 2. Late Triassic) contains species with an infold, which bears a wide calcified lamella, that excludes that genus from the Darwinulacea. Stratigraphic range.—Carboniferous to Holocene. Family Darwinulidae Brady and Norman, 1889 Darwinulidae Brady & Norman, 1889:121. Darwinellidae Brady, Crosskey, & Robert- son, 1874:140. Microdarwinulidae Kashevarova & Neus- trueva, 1982:148. Suchonellininae Kukhtinov, 1985:68. Diagnosis. —Same as the superfamily. Genera assigned.—Darwinula Brady & Robertson in Jones, 1885; Microdarwinula Danielopol, 1968; Paradarwinula Kozur, 1970. Discussion. —Kukhtinov (1985:68) in- cluded in the Darwinulidae the genera Dar- winula (Darwinula), Gerdalia Belousova, 1961, and Paradarwinula Kozur, 1970, which he raised to generic rank, and he de- scribed the subfamily Suchonellininae in the family. Based on the illustrated darwinulid AMAS in the type species, D. (P.) spinosa (Kozur, 1970, fig. 1). Paradarwinula is con- firmed in the Darwinulidae. Based on the original description, Microdarwinula does not require a separate suprageneric catego- ry. Suchonellina is considered herein to be a synonym of Darwinula, and, consequent- ly, does not require a suprageneric category. Stratigraphic range.—Carboniferous to Holocene. Genus Darwinula Brady and Robertson in Jones, 1885 Darwinula Brady & Robertson in Jones, 1885: 346. Polycheles Brady & Robertson, 1870:25. Darwinella Brady & Robertson, 1872:50. Suchonellina Spizharsky, 1937:156. ?Cyprione Jones, 1885: 343. ?Gerdalia Belousova, 1961:140. Darwinula (Neudarwinula) Mischina, 1972: 49, 50. Type species (monotypy).—Polycheles stevensoni Brady & Robertson, 1870:25. Diagnosis.—Small, less than 1 mm in length, elongate, smooth darwinulids, with adont hinge, dorsal edge of one valve inserts into groove of other and ventral edge of groove of latter fits into groove of former valve, slight overlap of one valve over other along free margins. Apparently partheno- genetic; may or may not retain the eggs and brood the first two instars. Discussion. — Reversal of overlap in Dar- winula and the above synonyms has been recorded (Sohn & Chatterje 1979:584). Be- lousova described Gerdalia (type species (original designation) Gerdalia polenovae Belousova, 1961:141, pl. 1, figs. 1, 1a) from carapaces. Starozhilova (in Lipatova & Sta- rozhilova, 1968:91) included species having a reversed overlap. Sharapova (1947:215) published a drawing showing the hinge of Darwinula having a ridge in the right valve that fits into a groove in the left valve. This hingement differs from the hinge of Dar- winula illustrated by Sohn (1987: pl. 1) which has grooves in both valves. Shara- pova’s illustration of the hinge was pub- lished by Starozhilova (in Lipatova & Sta- rozhilova 1968:80, fig. 20; 91, fig. 26; 97, fig. 31) for Darwinula, Gerdalia, and Su- chonellina (type species (subsequent desig- nation) S. inornata (McCoy, 1844) sensu Spizharsky 1939:194, pl. 46, fig. 7). Mischina (1972) did not formally de- scribe the subgenus Neudarwinula; she based her subgenus on measurements of Darwin- ula parallela (Spizharsky) without docu- mentation. Darwinula parallela (Spizhar- sky) in Schneider 1948:24, pl. 1, figs. 5a, b, was a manuscript name ofa Permian species described as Suchonellina Spizharsky, 1937 that was validated by Schneider (1948:24, pl. 1, figs. 5a, b). Kashevarova and Neus- trueva (1982:143, 145) considered the sub- VOLUME 101, NUMBER 4 genus Darwinula (Neudarwinula) to be a ju- nior synonym of Suchonellina. Although a male copulatory appendage was illustrated by Brady and Robertson in the original description of the type species (1870:pl. 10, fig. 13), and the antennule and the tip of the second foot of a male D. im- provisa Turner, 1895 were illustrated (Tur- ner 1895:263, pl. 81, figs. 1, 2), there are no other published records of males in Dar- winula (Sohn, 1987:152). The presence of males in Darwinula and its suprageneric categories is considered dubious until such time that males are documented based on living material. Jaanusson (1985:79-81) differentiated between the brood care of eggs in the platy- copids and the podocopids. He named the platycopid egg care “Cytherella type,’ and the podocopid egg care, including Darwin- ula, *“‘Cyprideis type.’ In Cytherella abys- sorum Sars, 1865 the eggs are attached firm- ly to the inner epithelial layer inside the posterior part of the carapace where each egg leaves an impression in the inside of the valve; in the Cyprideis type the extruded eggs in the posterior part of the carapace are not attached and are freely movable. Ac- cording to Jaanusson there is no evidence that Cytherella does or does not retain the early instars within the carapace. Darwinula retains the first two instars within the car- apace where they can be observed with transmitted light or by video tape record- ings opening their valves and/or extruding their appendages. Ecology. —Mainly freshwater, rare in brackish water (salinity to 13%o according to Keyser 1975:266), terrestrial (damp moss), and phreatic. Geographic distribution. —Circumglobal. Geologic range.—Miussissippian to Ho- locene. Genus Microdarwinula Danielopol, 1968 Microdarwinula Danielopol, 1968:154. Type species (original designation). — (Darwinula zimmeri Menzel, 1916:487, figs. 821 16-21, and redescribed and reillustrated by Danielopol, 1968:figs. 1-28. Diagnosis. — Differs from Darwinula in having smooth tripartite hinge that consists of terminal elongate smooth bars and smooth central groove in right valve and opposing structures in larger left valve; in having internal tubercles at both ends of ventral margins of left valve; and in not brooding the young. Discussion. —Danielopol described, dis- cussed and illustrated in detail females of the type species, but did not mention males. Ecology. — Freshwater. Geographic distribution. — Africa, Mada- gascar, and Europe. Geologic range. —Oligocene to Holocene. Genus Paradarwinula Kozur, 1970 Darwinula (Paradarwinula) Kozur, 1970: 434. Type species (original designation). — Darwinula dreyerae Kozur, 1968:848, pl. 4, fig. 10. Diagnosis. — Differs from Darwinula and Microdarwinula in having ventroposterior spine on left valve. Discussion. — Based on the illustration of the AMAS by Kozur (1970:435, fig. 10) the genus belongs in the Darwinulidae. Because Kozur (1968; 1970) did not describe the inside of the valves, the inside morphology is unknown. Ecology. — Brackish water. Geographic distribution. — Europe. Geologic range. —Late Triassic. Acknowledgments I thank Dr. Wang Shang-qui, PRC, for two specimens of Panxiania; Dr. M. N. Gramm, USSR, and Dr. E. K. Kempf, West Germany, for reproduction of Russian pub- lications. This paper profited by discussions with Dr. L. S. Kornicher, and editorial sug- gestions by Dr. T. E. Bowman, both in the National Museum of Natural History. Pub- lication outside the U.S. Geological Survey 822 was approved by the Director on March 15, 1988. Literature Cited Belousova, Z. D. 1961. Ostrakody nizhnego Tria- sa.— Bulleten Moskovskogo Obshchestva Ispe- tateley Prorody, n. ser. v. 64, Otdel Geologi- cheskiy, Moscow Universitet 36(4):127—246, 2 pls. Brady, G. S., H. W. Crosskey, & D. Robertson. 1874. A monograph of the post-Tertiary Entomostra- ca of Scotland and Ireland.— Palaeontographi- cal Society of London. 231 pp., 16 pls. —, & A.L. Norman. 1889. A monograph of the marine and freshwater Ostracoda of the North Atlantic and of north-western Europe. —Scien- tific Transactions of the Royal Dublin Society. Ser. 2,4(2):63-270, 23 pls. —., & D. Robertson. 1870. The Ostracoda and Foraminifera of tidal rivers.—Annals and Mag- azine of Natural History. ser. 2, (4)6:1-33, 307— 309. —, & 1872. Contribution to the study of the Entomostraca. No. 6. On the distribution of the British Ostracoda.—Annals and Maga- zine of Natural History (4)9:48—70. Chapman, F. 1901. On some fossils of Wenlock age from Mulde, near Klinteberg, Gotland; with notes by T. Rupert Jones and F. A. Bather.— Annals and Magazine of Natural History. (7)7: 141-160, pl. 3. Coryell, H. N., & D. Malkin. 1936. Some Hamilton ostracodes from Arkona, Ontario.— American Museum Novitates no. 381, 20 pp., 1 pl. Danielopol, D. L. 1968. Microdarwinula n.g. et quel- — ques remarques sur la répartition de la Famille Darwinulidae Br. et Norm.— Annales de Lim- nologie 4(2):153-174, 45 figs. Gramm, M.N. 1982. The systematic position of the ostracod genus Healdianella Posner, 1951. Pp. 193-218, 5 pls., 9 figs. in R. H. Bate, E. Rob- inson, and L. M. Sheppard, eds., Fossil and Re- cent ostracods. Ellis Harwood Ltd., Chichester. Grindel, J. 1967. Zur Grossgliederung der Ordnung Podocopida G. W. Miiller, 1894 (Ostracoda). — Neues Jahrbuch fiir Geologie und Palaontolo- gie, Monatshefte, Jahrgang 1967 6:321-332. Holland, W. C. 1934. The ostracods of the Nineveh limestone of Pennsylvania and West Virginia. — Carnegie Museum Annals 22 (2—4):343-350, pl. DS, Howe, H. V., R. V. Kesling, & H. W. Scott. 1961. Morphology of living Ostracoda. Pp. Q3-Q17 in R. C. Moore, ed., Treatise on invertebrate paleontology, part Q, Arthropoda 3, Crustacea, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ostracoda. Geological Society of America and University of Kansas Press, New York and Law- rence, Kansas. Jaanusson, V. 1985. Functional morphology of the shell in platycope ostracodes—a study of ar- rested evolution.—Lethaia 18:73—84, 7 figs. Jones, T. R. 1885. On the Ostracoda of the Purbeck Formation; with notes on the Wealden species. — Quarterly Journal Geological Society of London 41:311-353. Kashevarova, N. P., & I. Yu. Neustrueva. 1982. The present state of the problem and classification principles of the ostracod superfamily Darwin- ulacea Brady et Norman, 1889. Pp. 121-154 in D. M. Rauzer, ed., Questions of micropaleon- tology. Systematics and morphology of micro- fossils. Academy of Sciences of the USSR, Order of the Red Banner of Labor Geological Institute, Micropaleontological Commission, number 25. ‘“‘Nauka,” (in Russian; English abstract). Kesling, R. V. 1981. Taxonomy of Ostracoda and the position of Microchelinella Geis.—Contri- butions from the Museum of Paleontology, Uni- versity of Michigan 25(15):209-315, 2 pls. Keyser, D. 1975. Ostracoden aus den Mangrovege- bieten von Siidwest-Florida.— Abhandlungen und Verhandlungen des Naturwissenschaftlich- en Vereins in Hamburg (NF) 18/19:255-—290, pls. 20-22, 13 figs. Kozur, H. 1968. Einige selten Ostracoden-Arten aus der germanischen Trias.—Monatshefte der Deutschen Akademie der Wissenschaften zu Berlin 10(11):848-872, 4 pls., 9 figs. 1970. Neue Ostracoden aus der german- ischen Mittel-und-Obertrias.—Geologie, Aka- demie-Verlag Berlin 19(4):434—-455, 4 pls. . 1972. Eine Bemerkungen zur Systematik der Ostracoden und Beschreibung neuer Platyco- pida aus der Trias Ungarns und der Slowekei. — Geologisch-Palaontologische Mitteilungen Innsbruck 2(10):1—27, 2 pls. Kukhtinov, D. A. 1985. Systema ostrakod nadse- meistva Darwinulacea.—Paleontologicheski Zhurnal 1985 (4):64—-69, 2 figs. (A systematic classification of the ostracode in the family Dar- winulacea. Paleontological Journal, Scripta Technica, Maryland 19(4):75-80.) Lipatova, V. V., & N. N. Starozhilova. 1968. Stra- tigrafiya i ostrakody triasovykh otlozhenty Sar- atovskogo Zavolzh’ya (Stratigraphy and ostra- codes of the Triassic deposits in the Saratov part of the Transvolga Region). Pp. 8-168, 20 pls., Izdatel’stvo Saratovs kogo Universiteta, Sara- tov, USSR. McCoy, F. 1844. A synopsis of the characters of the Carboniferous limestone fossils of Ireland. Uni- versity Press, Dublin, 190 pp., 20 pls. VOLUME 101, NUMBER 4 Menzel, R. 1916. Moosbewohnende Harpaticiden und Ostracoden aus Ost-Africa.— Archiv fur Hydro- biologie und Planktonkunde 1 1(3):485-489, 21 figs. Mischina, E. M. 1972. O sistematike iskopaemykh darvinulid (ostracody).—Paleontologicheskiy Sbornik, Izdatel’stovo L’vovskogo Universita 1(9):44—5 1, 6 figs. (in Russian: English abstract). —, & A. E. Kalis. 1975. Novyi rod ostrakod Severovostoka Russkoi Platformy. (A new os- tracode genus from the northeast of the Russian Platform.)— Paleontologicheskii Zhurnal 1975 (1):74-81, pls. 9, 10. (English Translation, Pa- leontological Journal 9(1):74—79.) Moore, R. C., ed. 1961. Treatise on invertebrate pa- leontology, part Q, Arthropoda 3, Crustacea, Ostracoda. Geological Society of America and University of Kansas Press, New York and Law- rence, Kansas, 442 pp., 334 figs. Muiler, O. F. 1785. Entomostraca seu insecta testacea que iun aquis Daniae et Norvegiae Reperit, de- scripsit et inconibus illustravit. Lipsae et Hav- inae, 135 pp., 21 pls. Rome, R. 1969. Morphologie de l’attache de la furca chez les Cyprididae et son utilisation en syste- matique. Pp. 168-193 in J. W. Neale, ed., The taxonomy, morphology and ecology of recent Ostracoda. Olivera and Boyd, Edinburgh. Sars, G. O. 1865. Oversigt af Norges marine ostra- coder, Saerskilt aftrykt af Videnskabe-Selska- bets Forhandlingen for 1865, 130 pp. (reprint, printed in 1966, Forhandlinger 1 Videnskabe- Selskabets 1 Kristiania.) Scott, H. V. 1961. Classification of the Ostracoda. Pp. Q74-Q92, figs. 32-34 in R. C. Moore, ed., Treatise on invertebrate paleontology, Part Q, Arthropoda 3, Crustacea, Ostracoda. Geological Society of America and University of Kansas Press, New York and Lawrence, Kansas. Schneider, G. F. 1948. Fauna ostrakod verkhneperm- skikh otlozheniy (Tatarskikiy 1 Kazanskiy ya- rusy) neftenosnykh rayonov SSSR. Pp. 21-48, 4 pls. in A. V. Fursenko, ed., Mikrofauna nef- tynosnykh mestorzhdeniy SSSR, Sbornik 1, Vtoroye Baku i Zapadnaya Sibir. Trudy Vse- soyuznogo Neftyanogo Nauchno-Issledova- tel’skogo Geologo-Razvedochnogo Instituta (VNIGRI), N. ser., 31. Sharapova, E. 1947. Otryad Ostracoda. Rakovicha- tye raki. Pp. 208-216, pls. 44, 45, in G. Krim- holx, ed., Atlas of the guide forms of the fossil faunas of the USSR. Volume 8. The Lower and Middle Jurassic. USSR Ministry of Geology, Ali- Union Geological Institute, Moscow-Lenin- grad. [In Russian]. Sohn, I. G. 1976. Antiquity of the adductor muscle attachment scar in Darwinula Brady and Rob- 823 ertson, 1885. Pp. 305-308, 3 pls. in G. Hart- mann ed., Proceedings of the 5th International Symposium on Evolution of post-Paleozoid Os- tracoda. Abhandlungen und Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg, (NF), 18/19 (Suppl.). 1985. Latest Mississippian (Namurian A) nonmarine ostracodes from West Virginia and Virginia.—Journal of Paleontology 59(2):446- 460, 6 figs. 1987. The ubiquitous ostracode Darwinula stevensoni (Brady and Robertson, 1870), rede- scription of the species and lectotype designa- tion.— Micropaleontology 33(2):150-163, 3 pls. , & S. Chatterje. 1979. Freshwater ostracodes from Late Triassic coprolite in Central India. — Journal of Paleontology 53(3):578—586, 2 pls. Spizharsky, T. N. 1937. Ostracoda from the Kolchu- gino series of the coal-bearing strata of the Kuz- netsk Basin.—Transactions Central Geological Prospecting Institute 97:139-172, 1 pl. (in Rus- sian; English summary). 1939. Order Ostracoda. Pp. 193-196 in B. Likharev, ed., Atlas rokovodyashchikh form is- kopaemykh faun SSSR. Volume 6, Permskaya Systema. Tsentral’nyi Nauchno-Issledova- tel’skiy Geologo-Razvedochnyi Institut (Ts- NIGRI). 193-196, pl. 46. (The atlas of the lead- ing forms of the fossil fauna USSR. Volume 6, Permian System. Central Geological and Pros- pecting Institute); (in Russian). Swain, F.M. 1961. Superfamily Darwinulacea Brady and Norman, 1889. Pp. Q253-Q254 in R. C. Moore, ed., Treatise on invertebrate paleontol- ogy, Part Q, Arthropoda 3, Crustacea, Ostra- coda. Geological Society of America and Kansas University Press, New York and Lawrence, Kansas. 1986. Freshwater Ostracoda from the Cache Valley Formation (Pliocene) of southeastern Idaho.—Revista Espanola de Micropaleonto- logia 18(3):363-385, 4 pls. Sylvester-Bradley, P.C. 1961. Suborder Metacopina. Pp. Q358-395, fig. 227 in R. C. Moore, ed., Treatise on invertebrate paleontology. Part Q, Arthropoda 3, Crustacea, Ostracoda. Geological Society of America and Kansas University Press, New York and Lawrence, Kansas. Triebel, E.M. 1941. Zur Morphologie und Oekologie der fossilen Ostracoden. Mit Beschreibung ein- iger neuer Gattungen und Arten.—Senckenber- giana 23(4/6):294-397, 15 pls. Turner, C. H. 1895. Fresh-water Ostracoda of the United States. Second Report of the State Ge- ologist.—Minnesota Geological and Natural History Survey, Zoological Series 2:277—337, pls. 67-81. 824 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Wang, Shang-qi. 1978. Late Permian and Early Trias- 1 geologov SSSR, Chlenostonogie, Trilobitoob- sic ostracodes of western Guizhou and north- raznye, Rakoobraznye. Gosudarstvennoe eastern Yunnan.—Acta Palaeontologica Sinica Nauchno-Tekhnicheskoe Izdatel’stvo Literatu- 17(3):299-308, 6 pls. ry po Geologiy 1 Okhrane Nedr, Moscow, vol- . 1980. Panxianidae, a new family of non-ma- ume 8. rine Ostracoda.—Acta Palaeontologica Sinica 19(4):302-310, 1 pl. U.S. Geological Survey, Room E-311, Zanina, I. E., N. F. Kashevarova, & E. N. Polenova. : 3 1960. Nadsemeistva Healdiacea. Pp. 337-3450, National Museum Natural History, Wash- figs. 855-871 in N. E. Chernysheva, ed., Osnovy ington, D.C. 20560. Paleontologiy; spravochnok diya paleontologiv PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 825-831 HYALOPONTIUS BOXSHALLI, NEW SPECIES (COPEPODA: SIPHONOSTOMATOIDA), FROM A DEEP-SEA HYDROTHERMAL VENT AT THE GALAPAGOS RIFT Arthur G. Humes Abstract.— Hyalopontius boxshalli, a new species of megapontiid copepod (Siphonostomatoida) from a depth of 2451 m at the Galapagos Rift, is char- acterized in the male by the exopod of leg 1 having the armature I-1; O-1; II,2,3, the third segment of the exopod of leg 3 with II,I,5, and the distal seta on the lateral margin of the free segment of leg 5 being one-half the length of the segment. Among many thousands of copepods re- covered by deep-sea submersibles from depths of 2000 m or more in the eastern Pacific (see Humes 1987, in press) two large male siphonostomatoids were found, both belonging to a new species of the genus Hy- alopontius Sars, 1909. Seven members of this genus have been described from plank- ton in the northeastern Atlantic (Sars 1909; Hulsemann 1965; Boxshall 1979). One species, Hyalopontius pleurospinosus (Heptner, 1968) is known from 3860-7100 m in the Kurile-Kamchatka Trench (Hept- ner 1968). Siphonostomatoida Thorell, 1859 Megapontiidae Heptner, 1968 Hyalopontius Sars, 1909 Hyalopontius boxshalli, new species Figs. 1—5 Type material.—2 6, in 2451 m at Ga- lapagos Rift, 00°48.0’N, 86°13.0'W, 7 Dec 1979, DSRV Alvin dive no. 990 (Hollis, Jones, and Tuttle). Holotype (USNM 234119) and 1 paratype (dissected) (USNM 235272) deposited in the National Museum of Natural History, Smithsonian Institu- tion, Washington, D.C. Male. — Body (Fig. 1a) elongate, 5.7 times longer than wide. Length (not including se- tae on caudal rami) 4.81 mm (4.75—4.86 mm) and greatest width 0.84 mm (0.84— 0.85 mm), based on 2 specimens in lactic acid. Epimera of segments bearing legs 1-4 pointed (Fig. 1b). Ratio of length to width of prosome 3.05:1. Ratio of length of pro- some to that of urosome 1.30:1. Segment bearing leg 5 (Fig. 1c) 330 x 363 um, a little wider than long. Genital segment 330 x 286 um. Four postgenital segments from anterior to posterior 495 x 215, 363 x 203, 209 x 198, and 308 x 258 um. Caudal ramus (Fig. 1d) 330 x 110 um, ratio 3:1, bearing 6 smooth setae, 2 lateral and subterminal (220 wm and 726 um), 3 inner and slightly dorsal (from outer to in- ner 1000, 902, and 803 um), and dorsal seta 638 um. Body surface mostly without ornamen- tation. Minute punctae over dorsal surface of rostrum (Fig. la). Anal segment with very small spinules over surface of anal oper- culum (Fig. lc) and ventral transverse row of minute spinules near insertions of both caudal rami (Fig. Id). Rostrum (Fig. le) projecting in lateral view with slightly pointed apex. First an- tenna (Fig. 2a) 1650 wm long not including setae. Lengths of its 11 segments (measured along their posterior nonsetiferous mar- oF 0.3 mm & 1.0 mm 0.5 mm Sy pr e = SS yet ale z AL Z. Zz Fig. 1. Hyalopontius boxshalli, male: a, Dorsal (scale A); b, Outline of epimera of segments bearing legs 1— 4, lateral (B); c, Urosome, dorsal (B); d, Caudal ramus, ventral (C); e, Rostrum, lateral (B). Fig. 2. Hyalopontius boxshalli, male: a, First antenna, anterodorsal (scale B); b, Second antenna, outer (D); c, Oral cone, lateral (D); d, Mandible, anterior (D); e, First maxilla, anterior (C); f, Second maxilla, anterior (C). al plate, anterior (D). cale D); b, Area between maxillipeds and first — 5 > = = i a i of legs, ventral (B); c, Leg 1 and intercoxal plate, anterior (D); d, Leg 2 and intercox Fig. 3. Hyalopontius boxshalli, male: a, Maxilliped, anterior pair VOLUME 101, NUMBER 4 oS <= 829 I { \ . \ ny \\ IN!) &) ~ Nai yy \ SN . WW ke Nas \ | WX ARKIN \ \ . \ YN \/ \ Wo} \ \ \ \ \ Fig. 4. Hyalopontius boxshalli, male: a, Leg 3 and intercoxal plate, anterior (scale D); b, Leg 4 and intercoxal plate, anterior (D). gins): 319 um (374 um along its anterior margin) 26, 26, 29, 35, 100, 55, 143, 155, 165, and 505 um, respectively. Setal for- MU ae Onl eo ae lie De 2s Dasand iS: Most setae smooth but | seta on segments 2, 6, 7, 8, and 11 with small setules. Second antenna (Fig. 2b) with small coxa and elongate basis bearing exopod 55 x 39 um carrying 3 long minutely feathered se- tae. Endopod 2-segmented. First segment with minute spinules along inner edge. Sec- ond segment greatly elongated, 495 um long, bearing proximal inner sparsely barbed seta located 67 um from base of segment. Be- yond this seta small spinules along edge of segment. Terminally very long outer seta 1,100 um barbed on proximal inner half and shorter inner seta 200 um with moderately long inner setules. Oral cone (Fig. 2c) 575 wm long. Man- 830 Fig. 5. lateral (B). dible (Fig. 2d) without palp, elongate, 573 um, and very slender with few minute ter- minal teeth. First maxilla (Fig. 2e) bilobed; elongate inner lobe with 3 smooth apical setae, outer lobe small with 1 barbed seta. Second maxilla (Fig. 2f) 2-segmented, first segment smooth, second segment slender and spinulose with blunt tip. Maxilliped (Fig. 3a) elongate, 3-segmented. First segment 385 um, with spinulose distal seta. Second seg- ment 990 wm, with smooth inner seta and ornamented with 2 rows of small spinules along posterior surface. Third segment 374 um, smooth, with 2 subterminal setae, lon- ger seta smooth and shorter seta spinulose, and | long terminal claw 850 um with inner row of small spinules proximally. Ventral surface of body between maxil- lipeds and first pair of legs (Fig. 3b) slightly protuberant. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hyalopontius boxshalli, male: a, Leg 5, ventral (scale B); b, Leg 6, ventral (B); c, Leg 5 and leg 6, Legs 1-4 (Figs. 3c, d, 4a, b) with 3-seg- mented rami. Spine and setal formula as follows (Roman numerals representing spines, Arabic numerals indicating setae): P, coxa O-1 basis 1-0 exp I-1; 0-1; I,2,3 enp 0-1; 0-2; 1,2,3 P, coxa O-1 basis 1-0 exp I-1; I-1; IJ,1,5 enp 0-1; 0-2; 1,2,3 P, coxa O-1 basis 1-0 exp I-1; I-1; IL1,5 enp 0-1; 0-2; 1,2,3 P, coxa O-1 basis 1-0 exp I-1; I-1; II,I,5 enp 0-1; 0-2; 1,2,2 Leg 5 (Fig. 5a, c) situated ventrally, with free segment 117 < 52 um, bearing 3 setae, proximal outer seta 385 um and minutely barbed, distal lateral seta 60 um and smooth, and apical seta 605 um and minutely barbed. Seta on body adjacent to free segment 460 um and minutely barbed. VOLUME 101, NUMBER 4 Leg 6 (Fig. 5b, c) forming posteroventral flap on genital segment bearing | long barbed seta 605 wm, | shorter smooth seta 88 um, and between them | very small seta 31 um. Color unknown. Female. —Unknown. Etymology.—The new species is named for Dr. Geoffrey A. Boxshall, who has con- tributed much to an understanding of the genus Hyalopontius. Remarks.—Hyalopontius boxshalli may be distinguished from its eight congeners (shown in Boxshall’s 1979 key to females) by a combination of characters. Unfortu- nately only the male of H. boxshalli is known. Congeners of the new species are known only from females, except for H. typ- icus Sars, 1909, where both sexes are known. Therefore, comparison of H. boxshalli with congeners must be made using characters not suspected of being sexually dimorphic. The formula for the armature of legs 1-5 seems to be constant in both sexes and thus is useful in making comparisons. Hyalopontius boxshalli possesses the fol- lowing distinctive characters that, used to- gether, are regarded as supporting its status as anew species: (1) the exopod of leg 1 with the armature I-1; 0-1; II,2,3, (2) the third segment of the exopod of leg 3 with II,I,5, and (3) the distal seta on the lateral margin of the free segment of leg 5 being about one- half the length of the free segment. Acknowledgments I thank Dr. Howard L. Sanders, Woods Hole Oceanographic Institution, for making 831 available to me the copepods collected by the manned submersible A/vin. The study of the specimens was supported by a grant (BSR-851461) from the National Science Foundation. Literature Cited Boxshall, G. A. 1979. The planktonic copepods of the northeastern Atlantic Ocean: Harpacticoida, Siphonostomatoida and Mormonilloida.— Bul- letin of the British Museum (Natural History) 35:201-264. Heptner, M. V. 1968. Description and functional morphology of Megapontius pleurospinosus sp. n. from the Pacific with some remarks on the status of the genus Megapontius within the sys- tem of families of Siphonostoma group (Copep- oda, Cyclopoida).— Zoologicheskii Zhurnal 47: 1628-1638. [In Russian. ] Hulsemann, K. 1965. A new genus and species of siphonostome cyclopoid copepod from deep North Atlantic waters.—Crustaceana 9:45-50. Humes, A. G. 1987. Copepoda from deep-sea hy- drothermal vents. Bulletin of Marine Science 41:645-788. [In press.]| New species of Stygiopontius (Copepoda: Siphonostomatoida) from a deep- sea hydrothermal vent at the East Pacific Rise. — Zoologica Scripta. Sars, G.O. 1909. Note préliminaire sur trois formes remarquables de Copépodes, provenant des campagnes de S.A.S. le Prince Albert de Mona- co. — Bulletin de l'Institut Océanographique 147: 1-8. Boston University Marine Program, Ma- rine Biological Laboratory, Woods Hole, Massachusetts 02543. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 832-837 DESCRIPTION OF MEMBRANOBALANUS ROBINAE, A NEW SPECIES OF SPONGE BARNACLE (CIRRIPEDIA, ARCHEOBALANIDAE) FROM BAJA CALIFORNIA, WITH A KEY TO THE GENUS Robert J. Van Syoc Abstract. —The California Academy of Sciences, Department of Invertebrate Zoology and Geology has recently undertaken a joint research program with the Centro de Investigaciones de Ciencia y de Educacion Superior de Ensenada (CICESE). The present paper documents a new species of the sponge barnacle Membranobalanus Pilsbry from Bahia de los Angeles collected during two expeditions in 1984. A key to the genus Membranobalanus is included. Systematic Account Subclass Cirripedia Burmeister, 1834 Order Thoracica Darwin, 1854 Suborder Balanomorpha Pilsbry, 1916 Superfamily Balanoidea (Leach) Newman & Ross, 1976 Family Archaeobalanidae Newman & Ross, 1976 Subfamily Archaeobalaninae Newman & Ross, 1976 Genus Membranobalanus Pilsbry, 1916 Membranobalanus robinae, new species Holotype. —Complete shell, opercular plates and body preserved in 75% EtOH, California Academy of Sciences (CAS), San Francisco, 061082. Dimensions of holotype. — Height 4.6 mm; carinorostral diameter 4.0 mm; lateral di- ameter 3.7 mm. Type locality. —Station BLA-11, 28°54'N, 113°30'W, Mexico, Baja California, Gulf of California, Bahia de los Angeles, 2 mile south of Casa Diaz, depth 10 feet, 6 Oct 1984. Material examined. —CAS paratypes 061083, 061084, 061086, one specimen each, and CAS 056184, about 100 speci- mens, Mexico, Baja California, Gulf of Cal- ifornia, Bahia de los Angeles, Punta Gringa, depth 10-15 feet. CAS paratype 061085, Mexico, Baja California, Gulf of California, Bahia de los Angeles, 2 mile south of Casa Diaz, depth 0-10 feet. Paratype distribution.—Paratypes have been deposited at the National Museum of Natural History (USNM 211487), Scripps Institution of Oceanography (SIO C9559), the Santa Barbara Museum of Natural His- tory (SBMNH 35076), the Los Angeles County Museum of Natural History (LACM 84-203.1), as well as the California Acad- emy of Sciences (as listed above). Diagnosis. —Shell thin, white, radii pres- ent, widening near orifice; orifice oval, sum- mits of wall plates nearly level, not deeply toothed; rostrum same length as other com- partmental plates or only slightly longer; base of wall plates rounded giving base a lobed appearance; opercular plates without chitinous lamellae; scutal adductor ridge in- cipient or lacking; tergal spur broad, ex- tending about % of basal margin, and short, only about % of basal margin width; tergal articular ridge gently sloped to sharp crest, bending 4 way between apex and basal mar- gin toward and descending to scutal margin then rising along margin to high point near basal margin; outer ramus of Cirrus IV with up to 6 recurved teeth per article; basal ar- ticles of Cirrus IV with several tooth-like spines at distal posterior margin. VOLUME 101, NUMBER 4 Fig. 1. 833 Membranobalanaus robinae: a, Shell, holotype CAS 061082; b, Inner view of scutum, paratype CAS 061083; c, Outer view of scutum, paratype CAS 061083; d, Inner view of tergum, paratype CAS 061083; e, Outer view of tergum, paratype CAS 061083. Scale: a, x 10; b-e, x50. Description. —Shell (Fig. 1a) cylindrical to high-conic, white, opening oval with summits of plates even and flattened by wear; compartmental plates solid but very thin and fragile, easily disarticulated or bro- ken; parietes with fine longitudinal striae perpendicular to growth lines; rostrum slightly longer than or equal to length of carina and laterals. Carinolaterals shorter and narrower than other plates; carina, ros- trum and laterals very similar in length and width, all 4 with curved triangular shape on upper half with narrower semioval lower half. Carina very deeply curved laterally in comparison to other plates. Radii very broad, wider than paries of carinolaterals; alae broad, extending about *% length of plates. Scutum (Fig. 1b—c) thick, convex, white; basal margin slightly shorter than or equal to length of tergal margin; tergal margin in- flexed; exterior with prominent growth ridges every second or third forming a tooth that extends onto occludent margin. Artic- ular ridge triangular and as high as long, extending from apex to lower third of scu- tum; articular furrow moderately shallow and narrow. Adductor ridge absent or in- conspicuous and rounded; triangular ad- ductor pit shallow on its broad, rounded open basal end to deep and narrow on its apical end; depressor muscle pit large and deep, oval or triangular extending up from basal margin over *% distance to base of ar- ticular ridge. Tergum (Fig. 1d-e) thin, white; about same width as scutum; basal margin slightly longer than scutal margin; growth ridges finer 834 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Membranobalanus robinae: a, Labrum, paratype CAS 061084; b, Palp, paratype CAS 061084; c, Mandible, paratype CAS 061084; d, First maxilla, paratype CAS 061084; e, Second maxilla, paratype CAS 061085; f, Cirrus I, paratype CAS 061085; g, Cirrus II, paratype CAS 061085; h, Cirrus IV, paratype CAS 061085. Scale: a—-e, g, x 100; f, x50; h, x75. and shallower than those of scutum, spur fasciole broad and delimited only by shifts in orientation of growth lines with no abrupt changes in elevation; tergal spur '/ to 73 width of basal margin and about 4 as long as basal width of spur, basally truncate at angle to basal margin; articular ridge slightly con- cave, distinct, reflexed 90 degrees toward scutal margin about '2 way from apex to basal margin, height decreases from apex as ridge approaches scutal margin, then in- creases as ridge bends back along lower ' of scutal margin; depressor crests may ap- pear as 2 or 3 distinct ridges or incipient ridges without discernable depth or height, or may be absent. Labrum (Fig. 2a) triangular, with deep notch at apex of crest; up to 3 teeth present on each side of notch. Palp (Fig. 2b) kidney-shaped, upper mar- gin concave, densely setose, setae slightly shorter than width of palp; tip of palp dense- ly setose, setae pectinate and longer than those of upper margin; lower margin devoid of setae over proximal * of palp. Mandible (Fig. 2c) with 3 large teeth and 2 smaller teeth; first tooth most robust, about same length as second tooth; second tooth not bifid, located near center of cutting edge; third tooth about *% length of first and sec- ond teeth; fourth and fifth about % length of first and second teeth; inferior angle armed VOLUME 101, NUMBER 4 with 2 small denticles or spines; inferior and superior margins setose. Maxilla I (Fig. 2d) with straight edge; up- per spine largest but equaled in length by sixth spine; second spine about '4 length of first spine; third, fourth, fifth and seventh spines 7% to 7% length of first spine; inferior angle with several short spines; superior margin setose; inferior margin lacking setae. Maxilla II (Fig. 2e) ovate, superior and posterior margins densely setose, hooked setae present on posterior margin. Cirrus I (Fig. 2f) with unequal rami; an- terior ramus about 3 times length of pos- terior ramus; articles of both rami slightly protuberant; both rami setose; hooked setae at apex of posterior rami; setae extremely pectinate near base, giving appearance of ferns or down feathers; short spines present near distal end of articles; Cirrus II (Fig. 2g) shorter than other cirri; inner ramus about 7 length of outer ramus; articles of both rami slightly protuberant; covered with pec- tinate setae; setae more fern-like near base; short spines present near distal end of ar- ticles; Cirrus III longer than Cirrus II; setae form and distribution similar to Cirri I and II; short spines present near distal end of articles; Cirrus IV (Fig. 2h) longer than Cir- rus III; outer ramus bearing large, recurved teeth on medial protuberant articles; rami of equal length and diameter; setae pecti- nate; spines in rows along distal margins of articles, 3-6 most posterior of these spines much larger and tooth-like than spines on other cirri; pedicel about '2 length of rami; Cirrus V longer than Cirrus IV; setae as in other cirri; small spines present along distal anterior margin as in Cirri I-III; Cirrus VI longer than Cirrus V; pectinate setae as in other cirri; small spines on anterior distal margin as in other cirri. Etymology.—The specific name robinae is in honor of Robin Ring. Discussion.—Membranobalanus robinae is unique among species of Membranobala- nus in possessing basally rounded com- partmental plates of approximately equal 835 length that give a lobed appearance to the whole barnacle when viewed from the side (Fig. la, Holotype). The new species bears recurved teeth on the outer ramus of Cirrus IV, indicating a relationship with Western Hemisphere Membranobalanus (Zullo & Standing 1983). Eastern Hemisphere mem- branobalanids have straight teeth or spines on the outer ramus of Cirrus IV. The other Western Hemisphere membra- nobalanids are M. declivis (Darwin, 1854) (western Atlantic south of Cape Fear, N.C.), M. costatus Zullo & Standing, 1983 (Cape Fear, N.C.), M. orcutti (Pilsbry, 1907) (southern California and Mexico), and /. nebrias (Zullo & Beach, 1973) (Galapagos Islands). Based on examination of approximately 50 specimens of M. robinae, the new species differs from M. orcutti, M. nebrias and M. declivis in the length of the rostrum. Both M. orcutti and M. declivis have rostra which are considerably longer than their other compartmental plates. The rostrum of M. nebrias while not as long as the rostra of M. orcutti or M. declivis, is decidedly longer than the other compartmental plates. There was very little variation in rostrum length relative to the length of other compartmen- tal plates in the specimens of M. robinae examined. M. robinae, like M. costatus, has a rostrum whose length is nearly equal to that of its other compartmental plates. However, M. robinae differs consistently from M. costatus in many respects: (1) the rami of Cirrus IV are equal rather than un- equal in length and diameter; (2) radii are present and well developed; (3) the parietes lack prominent costae; (4) the occludent margin of the scutum bears several teeth formed by extensions of prominent external growth ridges; (5) the bases of the parietes are rounded giving the base of the barnacle a lobed appearance when viewed from the side. Species in the genus Membranobalanus Pilsbry are obligate symbionts of sponges. Although most species have been found in 836 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Species of Membranobalanus and the species of sponge which they are known to inhabit. Authors of references used to compile this table are listed, complete citations may be found in Literature Cited. Barnacle species Sponge species Author M. declivis Spheciospongia vesparum Pearse 1932 Wells 1966 Zullo & Standing 1983 M. nebrias “Clionid”’ Zullo & Beach 1973 M. koreanus Cliona celata Kim & Kim 1983 M. cuneiformis Cliona sp. Hiro 1936 Utinomi 1968 M. longirostrum Spirastrella purpurea Rosell 1972 Utinomi 1968 M. longirostrum Suberites inconstans Utinomi 1968 M. costatus Anthosigmella varians Zullo & Standing 1983 M. orcutti Spheciospongia confoederata Jones 1978 Cliona celata californiana Jones 1978 “red clionid”’ “calcareous” M. brachialis M. basicupula M. robinae only one species of sponge (Table 1), ©. longirostrum in the western Pacific and Indo- Pacific inhabits at least two species of sponges (Utinomi 1968) as does M. orcutti (Jones 1978) in California and Baja Cali- fornia. The question of host specificity in Mem- branobalanus has been considered only with regard to M. orcutti (Jones 1978) and M. costatus (Zullo & Standing 1983). The other genus of sponge barnacle, Acas- ta Leach, has many species which inhabit more than one host sponge species. For ex- ample, Acasta cyathus inhabits Verongula ardis (de Laubenfels, 1950), Ircinia cam- pana (Lamarck, 1813), Erylus ministron- gulus Hectel, 1965, Ircinia felix (Duchasa- ing & Michelotti, 1864), Spinosella (=Callyspongia) vaginalis (Lamarck, 1814), and Spongia tubulifera Lamarck, 1814, and at least two other species of Demospongea in North Carolina (Wells 1966; Zullo & Standing 1983). To date, M. robinae has been found in only one species of sponge, Delaubenfelsia raromicrosclera Dickinson, 1945, at one lo- “probably Cliona sp.” Suberites inconstans Delaubenfelsia raromicrosclera Zullo & Beach 1973 Rosell 1973 Rosell 1972 Sukaimi 1966 Van Syoc herein cation, Bahia de los Angeles in the Gulf of California. If M. robinae lives in D. raro- microsclera throughout a greater part of the sponge’s range, we might expect to find it elsewhere in the Gulf of California (Dick- inson 1945, Hofknecht 1978). Key to the Genus Membranobalanaus la. Rostrum much longer than other compartmental plates .......... 2 b. Rostrum nearly as long, or as long as, other compartmental plates... 6 2a. Fourth cirrus with recurved teeth or spines on proximal articles... 3 b. Fourth cirrus with straight teeth or spines on proximal articles ..... 4 3a. Radii present .. M. declivis (Darwin) b. Radii lacking or very narrow M. orcutti (Pilsbry) 4a. Radii present RO Ueiaty Se M. koreanus Kim and Kim b. Radii lacking or very narrow ... 5 5a. Parietes with distinct horizontal growth lines .. M. brachialis (Rosell) VOLUME 101, NUMBER 4 b. Parietes without distinct horizon- talgonow thelimesieye i oe a Jes rele clleast las M. longirostrum (Hoek) 6a. Fourth cirrus with recurved teeth b. Fourth cirrus with straight teeth . eR eke ere M. cuneiformis (Hiro) 7a. Base of barnacle having a lobed appearance in side view due to rounded ends of compartmental plates. sess s. M. robinae, sp. nov. b. Base not lobed in appearance when viewed from side SasebanictesicOstatem ene. kines M. costatus Zullo and Standing b. Parietes not costate oe nee M. nebrias (Zullo and Beach) Acknowledgments Id like to thank Welton L. Lee for giving me the initial opportunity to study barna- cles. Special thanks to Victor A. Zullo for patiently training me in barnacle system- atics and being very generous with his en- couragement and continuing help and ad- vice with this manuscript and other cirripedian projects. William A. Newman and Terrence M. Gosliner read the manu- script and offered helpful criticisms. Sarah Ward Klontz identified the sponges inhab- ited by M. robinae. Lynette Cook provided the illustrations of M. robinae. Literature Cited Dickinson, M. G. 1945. Sponges of the Gulf of Cal- ifornia.—Allan Hancock Foundation Pacific Expeditions 11(1):1—55, 97 pl. Hiro, F. 1936. Report on the Cirripedia collected in the Malayan waters by the ship ““Zuihomaru.” — Japan Journal of Zoology 6(19):621-636. Hofknecht, G. 1978. Descriptions and key to the in- tertidal sponges of the Puerto Penasco area in the Gulf of California.—Journal of the Arizona and Nevada Academy of Sciences 13:51—56. 837 Jones, L. L. 1978. The life history patterns and host selection behavior of a sponge symbiont, Mem- branobalanus orcutti (Pilsbry) (Cirripedia). Ph.D. Dissertation, University of California, San Die- go, 112 pp. Kim, H. S., & I. H. Kim. 1983. A new species of Membranobalanus (Crustacea, Cirripedia) from the Korean waters.— Korea Journal of Zoology 26(1)1983:41-47. Pearse, A.S. 1932. VII. Inhabitants of certain sponges at Dry Tortugas. Papers from Tortugas Labo- ratory.—Carnegie Institution of Washington (Publication 435) 28:117—124. Rosell, N.C. 1972. Some barnacles (Cirripedia, Tho- racica) of Puerto Galera found in the vicinity of the U.P. Marine Biological Laboratory.—Nat- ural and Applied Science Bulletin 24(4):143- 285. . 1973. On two less well-known balanids (Cir- ripedia, Thoracica) from the Sulu Archipelago, Philippines. University of the Phillippines Nat- ural Science Research Center Technical Report 4:1-12. Sukaimi, A. 1966. A new species of Balanus (Crus- tacea: Cirripedia) from Singapore.— Bulletin of the National Museum of Singapore 33(9):65-— 68. Utinomi, H. 1968. Pelagic, shelf and shallow water Cirripedia from the Indo-Pacific.— Videnska- belige Meddeleser Dansk Naturhistorisk For- ening 131:161-186. Wells, H. W. 1966. Barnacles of the northeastern Gulf of Mexico. — Quarterly Journal of the Flor- ida Academy of Sciences 29:8 1-95. Zullo, V. A., & D. B. Beach. 1973. New species of Membranobalanus Hoek and Hexacreusia Zul- lo (Cirripedia, Balanidae) from the Galapagos Archipelago.—Los Angeles County Natural History Museum Contributions in Science 249: 1-16. , & J. D. Standing. 1983. Sponge-inhabiting barnacles (Cirripedia: Archaeobalanidae) of the Carolinian Province, Southeastern United States, with the description of a new species of Mem- branobalanus Pilsbry.— Proceedings of the Bi- ological Society of Washington 96(3):468—477. California Academy of Sciences, Depart- ment of Invertebrate Zoology and Geology, Golden Gate Park, San Francisco, Califor- nia 94118. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 838-842 ELASMOPUS BALKOMANUS, A NEW SPECIES FROM THE FLORIDA KEYS (CRUSTACEA, AMPHIPODA) James Darwin Thomas and J. L. Barnard Abstract. —Elasmopus balkomanus is described from Looe Key Reef in the Florida Keys. The species is very close to the eastern Pacific E. antennatus but in the male has equally extending rami on uropod 3, only 2 (versus 4—6) spines on each lobe of the telson in adults, a lateral ridge on the propodus of male gnathopod 2 and very heavily armed flagella of antenna 2 in the male. This species lives in a short-tufted algal turf community on coral rubble but apparently is rare because it has only been collected once in 10 years of sampling in the Florida Keys. Elasmopus balkomanus, new species Figs. 1-3 Etymology. —A.S. balko, ridge; L, manus, hand. Diagnosis of male. —Eyes ordinary, brownish-purple in alcohol. Flagellum of male antenna 2 densely setose. Mandibular palp article 3 deeply falcate. Palm of male gnathopod 2 with weak, sparsely spinose hump near dactylar hinge, no marginal teeth, inner face with longitudinal, cuspidate ridge, setae all posterior (“below’’) ridge and not organized into rows perpendicular to ridge; dactyl overriding palm onto face of prop- odus. Article 2 of pereopods 5-7 with only tiny setules and tiny serrations posteriorly; locking spines of pereopods 3-7 thin and almost straight, smooth; main subapical spine-seta on dactyl of medium thickness, smooth, with 2 accessory thin setules. Epi- mera |—2 with medium sharp posteroven- tral tooth, all ventral spines short, rarely paired. Uropod 3 with “long” rami (in ge- neric context), inner almost as long as outer. Telson with sharply and deeply incised api- ces each bearing pair of spines, one spine elongate, other spine short. Description of male.—Body generally as in other species of genus (for side views of body form in Elasmopus see Sars 1895, Bar- nard 1962, Bousfield 1973). Antenna 1 elongate, slender, article 1 with 3 ventral spines in tandem, accessory flagellum 3-ar- ticulate. Flagellum of antenna 2 densely armed with flags of setae. Upper lip rounded below, projecting slightly in front of epi- stome from side view. Incisors of right and left mandibles with 2 teeth, right lacinia mobilis bifid, proximal branch simple, dis- tal branch with 7 teeth, left lacinia mobilis with 4 teeth; right rakers 3, left 4; molars moderately triturative, each with plumose seta; palp article 1 weakly elongate, article 2 with 2-3 short medial setae, 2—3 apico- medial long setae, article 3 with 1-1 A setae, many D setae forming comb on falcate in- vagination, 3 E setae. Lower lip, maxilla 2 and maxilliped like figure 35 of Barnard (1979) with following minor exceptions. In- ner plate of maxilla 1 with 2 apical setae and 4 apicolateral setules, outer plate with 7 spines, palp article 2 with 2 sharp cusps, one apicolateral seta, 4 apicomedial mar- ginal elements (2 thick, 2 thin) and 4 facial setae. Both lobes of maxilla 2 equally thin, inner plate with only single apicomedial seta no longer than apical setae, outer plates asymmetrical, no cusps, 3 facial 5 apico- medial, 3 apicolateral setae. Inner plate of maxilliped with weak apicomedial cusp, 8 apical setae, 6 medial setae, outer plate with 10 medial blades, 5 apicolateral setae, palp article 3 with scaly apicolateral irregular VOLUME 101, NUMBER 4 839 Fig. 1. lobe, dactyl with thick nail and 4 accessory setules. Anterior margin of coxa | and posterior margin of coxa 4 weakly excavate, long setae on coxae 1-4 = 3-2-0-0. Gnathopod | or- dinary, see illustrations. Article 2 of gnatho- pod 2 with weak lateral ridge and hollow. Pereopods 3-4 slender, 4 smaller than 3, article 6 with 6—7 posterior sets of spine pairs including locking spines. Posterior margin of article 2 on pereopod 5 weakly excavate, posteroventral corner of Elasmopus balkomanus, male holotype ““m” 7.13 mm. Capital letters refer to parts; lower case letters to left of capital letters refer to specimens and to the right refer to adjectives as: C, coxa; G, gnathopod; H, head; M, mandible; O, outer pate or ramus; P, pereopod; R, uropod; T, telson; V, palp; W, pleon; X, maxilla; Z, gill; m, medial; r, right; s, setae removed; t, left. weak lobe sharp; pereopods 5-7 relatively slender in generic context, serrations on ar- ticle 2 tiny and numerous, dactyl relatively elongate. Broad gills present on coxae 2-6. Dorsolateral margin of peduncle on uro- pod 2 with only | spine. Female “‘f.’’-—Like male but gnathopod 2 of female form, see illustration; merus with sharp tooth. Narrow oostegites present on coxae 2—5. Inner ramus of uropod 3 short- ened. Illustrations. —Telson magnified more Fig. 2. Elasmopus balkomanus, unattributed figures = male holotype ““m’” 7.13 mm; f = female “f’ 6.15 mm. Letter codes, see Fig. 1. Fig. 3. Elasmopus balkomanus, unattributed figures = male holotype “m” 7.13 mm; f = female “‘f’ 6.15 mm. Letter codes, see Fig. 1. 842 than uropod 3. Pereopod 4 not illustrated, like pereopod 3 but significantly smaller. Holotype.—USNM No. 235007, male “mm” 7.13 mm. Type locality.—Florida: Florida Keys; Looe Key Reef, west end of rubble zone on backreef, formaldehyde wash of rubble in 2 m, 9 Oct 1983, coll. J. D. Thomas, associ- ated amphipod genera, Ceradocus, Maera, Spathiopus. Material.—Type locality, female “f’ 6.15 mm, and 4 other specimens. Relationship. — Differing from E. anten- natus as in Abstract. Differing from E. Jevis Smith (Bousfield 1973) in the equal rami of uropod 3, lack of medial hollow and con- figuration on propodus of male gnathopod 2; the heavily armed flagellum of antenna 2 in male and presence of lateral ridge on propodus of male gnathopod 2. Differing from E. ecuadorensis in the Ga- lapagos Islands (Barnard 1979) by the sub- equal rami of uropod 3, presence of tooth on epimeron 3, the excavate telsonic lobes with weak spination (versus truncate lobes bearing numerous spines), and the distinc- tive locking spine formation on pereopods 3-7. Differing from E. hawaiensis (as E. ec- uadorensis hawaiensis in Barnard 1970) in the equal rami of uropod 3, presence of tooth on epimeron 3 and the ordinary locking spines of pereopods 3-7. Distribution. —Florida: Florida Keys; Looe Key Reef, 2 m. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Acknowledgments We thank Dr. D. Challinor, Assistant Sec- retary for Science at Smithsonian Institu- tion, for providing funds for Barnard to join Thomas in Florida to undertake this study. Thomas was supported by NSF grant BSR- 8515186. Mrs. Linda B. Lutz of Vicksburg, Mississippi inked our drawings. We thank Janice Clark and Lori B. Jackintell for their assistance. Literature Cited Barnard, J. L. 1962. Benthic marine Amphipoda of southern California: 2. Families Tironidae to Gammaridae.— Pacific Naturalist 3:73-115. . 1970. Sublittoral Gammaridea (Amphipoda) of the Hawaiian Islands.— Smithsonian Contri- butions to Zoology 34:1-286. . 1979. Littoral gammaridean Amphipoda from the Gulf of California and the Galapagos Is- lands.—Smithsonian Contributions to Zoology 271:1-149. Bousfield, E. L. 1973. Shallow-water gammaridean Amphipoda of New England. Ithaca, New York, Cornell University Press. xii + 312 pp. Sars, G. O. 1895. Amphipoda: An account of the Crustacea of Norway with short descriptions and figures of all the species. Crustacea of Norway 1:1-711. Christiania: Cammermeyers. (JDT) Reef Foundation, P.O. Box 569, Big Pine Key, Florida 33043; (JLB) De- partment of Invertebrate Zoology, National Museum of Natural History, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 843-852 SYNALPHEUS DORAE, A NEW COMMENSAL ALPHEID SHRIMP FROM THE AUSTRALIAN NORTHWEST SHELF A. J. Bruce Abstract. — A new species of alpheid shrimp, Synalpheus dorae, is described and illustrated. It occurs in association with sponges of the genus Reniera on the Australian Northwest Shelf at depths of 37-82 m. The species is unusual in having only four carpal segments on the second pereiopod and all 162 specimens collected appeared to be male. A survey of the benthic fauna of the Aus- tralian Northwest Shelf carried out by the F.R.V. Soela of the Commonwealth Sci- entific and Industrial Research Organiza- tion’s Fisheries Division in 1983-1984 pro- vided abundant material of the family Alpheidae. The study of these shrimps was undertaken by Professor A. H. Banner and D. M. Banner and revealed numerous species new to the Australian fauna as well as many rare species and some undescribed species. On the death of Professor Banner the study was discontinued and the speci- mens returned to the Northern Territory Museum, Darwin. Amongst the material re- turned was one container with a note from Dora Banner enclosed, stating ““This is def- initely a new species. We had not had time to work on it.” This material is now de- scribed below. Synalpheus dorae, new species Figs. 1-6 Material examined. —18 6, operation 7/12, 19°51.9’S, 117°0.78’E, 57-58 m, trawl, F.R.V. Soela, Cr. 0283, 10 Apr 1983, coll. P. Blyth, NTM. Cr. 005060.—136 4, sta NWS-22, 19°05.0’S, 118°57.8’E, 82 m, trawl, F.R.V. Soela, Cr. 0283, 24 Apr 1983, coll. A. J. Bruce, NTM. Cr. 005059.—1 34, lo- cality D4, 19°29.6’S, 118°52.4’E, 37-38 m, trawl, F.R.V. Soela, Cr. 0583, 25 Oct 1983, coll. T. Ward, NTM. Cr. 005061.—7 2, lo- cality D7, 19°29.7'S, 118°51.4’E, 40-41 m, epibenthic sledge, F.R.V. Soela, Cr. 0583, 25 Oct 1983, coll. T. Ward, NTM. Cr. 005062. Description.—A small-sized, robust al- pheid shrimp of subcylindrical body form. Carapace smooth, glabrous, slightly com- pressed; rostrum small, blunt, short, de- pressed, slightly upturned, slightly exceed- ing orbital teeth, with distal dorsolateral setae, about 1.4 times longer than proximal width, reaching to about middle of proximal segment of antennular peduncle, without orbitorostral process, orbitorostral notch broadly rounded, orbital teeth triangular, slightly upturned distally, with simple setae laterally, anterolateral margin unarmed, very obliquely angular, posterior margin broadly rounded, cardiac notch distinct. Abdomen subcylindrical, glabrous, first 4 pleura small, rounded in female, similar in male, but first pleuron with acute postero- ventral tooth, fifth pleuron with acute ven- tral tooth; sixth segment about subequal to length of fifth, about 1.5 times longer than deep, posterodorsal margin unarmed, pos- teroventral angle well developed, acute, posterolateral angle acute. Telson about 1.6 times sixth segment length, triangular, dis- tally truncate, about 1.25 times longer than proximal width, lateral margins straight, convergent, posterior margin about 0.28 of anterior width, 2 pairs of large, erect dorsal spines at 0.35 and 0.82, posterior spines slightly longer than anterior, about 0.27 of 844 Fig. 1. telson length; 2 pairs of posterior marginal spines, submedian slightly larger than lat- eral, about 0.21 of telson length, short me- dian gap with 2 simple setae dorsally and 2 longer plumose setae ventrally; dorsal sur- face with sparse simple setae. Antennule well developed; proximal seg- ment of peduncle subcylindrical, with nor- mal statocyst with granular statolith, sty- locerite long, acute, reaching to about 0.8 of segment length, with short simple setae distolaterally; intermediate and distal seg- ments subequal, cylindrical, together sub- equal to proximal segment length; upper fla- gellum biramous, rami fused for 5 segments, robust; short ramus with 2 stout segments, about 8 groups of aesthetascs; lower ramus with about 12 slender segments, length sub- equal to fused portion plus short ramus; lower flagellum short, slender, about 0.75 times carapace length, exceeding long ramus of upper flagellum. Antenna with basicerite robust, with shorter acute dorsolateral and longer acute ventrolateral process, latter reaching or ex- ceeding distal end of proximal segment of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Synalpheus dorae holotype male, Northwest Shelf, Western Australia. Scale bar in millimeters. antennular peduncle, both with sparse sim- ple setae distolaterally; merocerite and is- chiocerite short; carpocerite subcylindrical, slightly flattened, about 4.5 times longer than wide, sparsely setose laterally; flagellum short, robust, about 1.3 times carapace length; scaphocerite with very robust lateral spine, slightly exceeding antennular pedun- cle, distinctly exceeding carpocerite, about 4.7 times longer than proximal width, la- mella greatly reduced, reaching to about 0.5 of lateral margin, spine with several short simple setae distolaterally. Eyes normal, completely covered dorsally by orbital hood, cornea well pigmented. Epistome normal, without special fea- tures. Mandible (left) with corpus moderately slender; palp 2-segmented, proximal seg- ment subcylindrical, proximally tapered, non-setose, distal segment oval, flattened, about 1.8 times longer than broad, distal margin with numerous short, plumose se- tae; molar process robust, obliquely trun- cate distally, margin densely fringed with short setae, with strong blunt tooth poste- VOLUME 101, NUMBER 4 | 845 Fig. 2. Synalpheus dorae paratype male: A, Anterior carapace and antennal peduncles, lateral; B, Same, dorsal; C, Antennule; D, Antenna; E, Scaphocerite; F, First abdominal segment, lateral; G, First pleopod; H, Second pleopod; I, Telson; J, Uropod. riorly; incisor process slightly expanded dis- tally, truncate with 7 small irregular acute teeth, larger teeth laterally. Maxillula with bilobed palp, upper lobe long, acute, lower lobe short with distally setulose spine; upper lacinia broad with about 20 short simple acute marginal teeth, with numerous short denticulate setae submarginally; lower la- cinia slender, curved, with several serrulate setae distally. Maxilla with short simple subcylindrical, distally tapering non-setose palp; basal endite broad, produced medi- ally, partly bilobed, with dense setal fringe medially; coxal endite simple, reduced, small acute lobe with several long setae; scaphog- nathite 4.0 times longer than broad, anterior lobe distally narrow, medial margin con- cave, 1.8 times longer than wide, posterior 846 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.1mm BCDFK B ; Fig. 3. NOW Synalpheus dorae paratype male: A, Mandible; B, Same, incisor process; C, same, molar process; D, Same, palp; E, Maxillula; F, Same, palp; G, Maxilla; H, First maxilliped; I, Second maxilliped; J, Third maxilliped; K, Same, Lateral plate. lobe about 0.35 of scaphognathite length. First maxilliped with 2-segmented palp, proximal segment about 2.2 times longer than broad, medial margin sparsely setose distally, distal segment about 0.8 of proxi- mal segment length, 3.0 times longer than wide, tapering slightly, with single short simple distal seta; basal endite normal me- dial margin densely setose; coxal endite simple, sparsely setose, 4 long setae disto- medially; exopod with well developed fla- gellum with numerous plumose setae dis- VOLUME 101, NUMBER 4 847 Fig. 4. Synalpheus dorae paratype male: A, Major first pereiopod; B, Same, chela, lateral; C, Same, fingers, medial; D, Same, fingers, lateral; E, Minor first pereiopod; F, Same, chela, lateral; G, Same, fingers; H, Same, ventral; I, Second pereiopod. tally, caridean lobe small, feebly setose; epipod large, transversely oval, simple. Sec- ond maxilliped of normal form; dactylar segment narrow, about 4.6 times longer than broad, densely setose medially, propodal segment broad, distally rounded, sparsely setose, basis broad, medially excavate and setose, exopod flagellum well developed with numerous plumose setae distally; coxa broad, medially produced with long setae distally, small rounded epipod without pro- dobranch laterally. Third maxilliped with endopod stout, exceeding carpocerite and antennular peduncle; isochiomeral segment 3.0 times longer than central depth, slightly tapered proximally, ventromedially exca- vate, margins feebly setose; carpal segment about 0.27 of ischiomeral segment length, 848 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. reiopod; D, Same, propod and dactyl; E, Fifth pereiopod; F, Same, propod and dactyl. as long as wide, subcylindrical, with 2 dor- solateral spines and several ventrolateral se- tae; distal segment about 0.3 times longer than proximal width, tapering distally, dis- tal width 0.4 of proximal, length 0.83 of ischiomerus, with 7 transverse rows of ser- rulate spines medially and 8 stout, blunt, finely denticulate, distally; basis about 0.3 of ischiomeral length, sparsely setose, exo- pod with well developed flagellum reaching to about 0.85 of ischiomeral length, with numerous plumose setae distally; coxa stout, ventrally medially rounded, with long low dorsolateral plate with small recurved distal tooth, without epipod or arthrobranch. Synalpheus dorae paratype male: A, Third pereiopod; B, Same, propod and dactyl; C, Fourth pe- Thoracic sternites narrow and unarmed, coxae of pereiopods in close apposition. First perelopods markedly unequal and dissimilar, apparently similar in males and females. Major chela enlarged, about 1.5 times carapace length, with palm subcylin- drical, centrally moderately swollen, slight- ly dorsoventrally flattened, smooth, gla- brous, about 1.8 times longer than greatest width, distodorsal angle produced, with slender acute tooth; dactylus strongly com- pressed, about 0.38 of palm length, semi- circular, with stout acute distal tooth, molar process small, feebly produced, equal to 0.3 of cutting edge length, medial and lateral VOLUME 101, NUMBER 4 849 Fig. 6. Synalpheus dorae paratype male: A, Distal spines of terminal segment of third maxilliped endopod; B, Second pereiopod, chela; C, Third pereipod, coxa; D, Same, dactyl; E, First pleopod, endopod; F, Second pleopod, endopod; G, Posterior telson spines; H, Distolateral spine of exopod of uropod. surfaces of dactyl with numerous submar- ginal groups of 2—3 simple setae; fixed finger short, curved, blunt distally, deeply exca- vate proximally for reception of dactylar molar process and cutting edge, with groups of simple setae; carpus very stout about 0.14 of palm length, expanded and excavate dis- tally, unarmed; merus robust, 2.0 times longer than greatest width, about 0.57 of palm length, unarmed; ischium, basis and coxa, all short, robust, without special fea- tures, basis without exopod, coxa as in ma- jor chela, without epipod and setobranch. Minor second pereiopod with chela about 0.4 of major chela length, about 0.5 of car- apace length; palm about 1.4 times longer than deep, smooth, slightly swollen cen- trally, compressed with scattered long sim- ple setae; dactylus about 0.75 of palm length, about 2.5 times longer than proximal depth, broadly subspatulate with thickened edge laterally, strong blunt hooked tooth distally with smaller tooth medially, with numerous groups of long simple setae dorsally; fixed finger deeply excavate, with strong blunt tooth distally with smaller tooth laterally, with long setae medially and laterally; car- pus about 0.6 of palm length, 0.9 times lon- 850 ger than wide, expanded distally deeply ex- cavate, unarmed; merus about 0.8 of chela length, 2.8 times longer than greatest width; ischium, basis and coxa without special fea- tures. Second pereipod moderately robust, ex- ceeding carpocerite by chela and distal car- pal segment, chela with palm compressed, about 1.2 times longer than deep, sparsely setose, fingers slender, tapering, with small hooked tips, cutting edges entire, with dense groups of coarsely serrulate setae proxi- mally, shorter simple setae distally, dactylus about 2.8 times longer than proximal depth, subequal to palm length; carpus with 4 stout segments, 4:1:1:2, proximal segment lon- gest; merus about 0.9 of carpus length, 4.0 times longer than wide; ischium 0.7 of me- rus length, 3.4 times longer than central width, tapering proximally; basis 0.6 of is- chium length, without exopod; coxa robust, without epipod or setobranch. Third pereiopod robust, exceeding car- pocerite by about half propod length; dactyl with unguis not clearly demarcated from corpus, compressed, feebly curved, about 2.6 times longer than proximal depth, with well developed accessory tooth at 0.6 of dac- tyl length, shorter and stouter than terminal tooth; propod about 3.7 times longer than deep, compressed, 4.0 times longer than dactyl, with 2 robust distoventral spines and 6 ventral spines, with numerous long simple setae dorsally and ventrally, carpus about 0.6 of propod length, 2.2 times longer than distal width, slightly tapered proximally, with well developed distodorsal lobe, dis- toventral margin with 2 spines, one robust, one slender; merus 2.0 times length of car- pus, 2.6 times longer than central width, unarmed; ischium about 0.33 of merus length as long as wide distally, tapered proximally, unarmed; basis 0.4 of merus length, unarmed; coxa robust with 2 sub- marginal ventral spines, without epipod or setobranch. Fourth pereiopod similar to third; propod 0.8 of third propod length, with single distoventral spine, 5 ventral PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON spines; carpus with single distoventral spine; coxa with single distoventral spine. Fifth pereiopod generally as third; propod 0.8 of third propod length, 4.2 times longer than central depth with 2 distoventral spines and 3 ventral spines and 5 transverse rows short of serrulate setae ventrolaterally; coxa with- out distoventral spine. Male first pleopod with basipodite 2.2 times longer than wide, exopod 1.15 times length of basipodite, 4.0 times longer than proximal width; endopod small, 0.25 of exopod length, 2.5 times longer than wide, tapering distally, with single short simple distal setae. Second pleopod with basipodite 2.5 times longer than wide; exopod about 3.6 times longer than proximal width, 1.1 times length of basipodite; endopod sub- equal to exopod length, without appendix masculina, appendix interna at 0.4 of en- dopod length, about 0.25 of endopod length, 5.5 times longer than central width with few (3-5) concinnull. Uropod with stout protopodite, with large acute dorsolateral tooth, with long simple setae medially, and smaller dorsal tooth; exopod broad, 1.4 times longer than wide, oval, lateral margin convex with acute dis- tolateral tooth with long mobile spine and small acute tooth medially, without distinct diaeresis, lateral margin with submarginal row of setae ventrally; endopod subequal to exopod length, 1.8 times longer than wide. Types.—An intact specimen from sta NWS-22 is selected as holotype and the oth- er specimens from that station are desig- nated as paratypes, catalog number NTM. Cr. 005059. Paratypes are deposited in the collections of the National Museum of Nat- ural History, Washington, D.C., catalog number USNM 234305, the Bishop Mu- seum, Honolulu, catalog number S10804 and the Rijksmuseum van Natuurlijke His- torie, Leiden, catalog number RMNH $7179, Measurements. —Holotype male, total length (approx.) 10.0 mm; carapace length, 4.0 mm, major chela, 5.5 mm; minor chela VOLUME 101, NUMBER 4 2.0 mm. Paratype male, total length (ap- prox.), 8.2 mm; carapace length, 3.5 mm; major chela, 4.15 mm; minor chela, 1.9 mm. Host.—The specimens from T/12 and NWS/22 were obtained from sponges. The 136 specimens from NWS/22 were all ob- tained from a single sponge host, identified as Reniera sp. (Haliclonidae). Parasites. —One specimen, sta NWS-22, was infected by an abdominal bopyrid par- asite, Eophryxus sp. (Isopoda: Hemiarthri- nae). Etymology.—The species is named in honor of the late Dora May Banner, in rec- ognition of her great contribution to alpheid taxonomy. Systematic position. —Morphological fea- tures of special significance in the assess- ment of the relationships of S. dorae are (1) the lack of an orbitorostral process, (11) the absence of a dense setal brush on the dactyl of the minor chela, (iii) absence of a strongly produced frontal region extending far be- yond eyes, (iv) lack of a reduced accessory spine on ambulatory dactyls, and absence of spines on merus of third pereiopod, (v) lack of a rostrum markedly exceeding or- bital teeth, and (vi) broad dentate fingers on minor chela. The first four items are char- acteristic of the ““Comatularum Group” and the others of the “Coutierei Group,” a species group largely characterized by its lack of consistent characters (Banner & Banner 1975). Synalpheus dorae appears most closely related to some of the species of the ‘Coutierei Group’ and can be immediately distinguished from most of them by the presence of only fgur segments on the car- pus of the second pereiopod, all other species except two having five segments. The two species with a 4-segmented carpus are S. quadriarticulatus Banner & Banner, 1975, and SS. redactocarpus Banner, 1953. The first species lacks an orbitorostral process but its presence or absence in S. redactodactylus is not recorded. Synalpheus dorae is most conveniently distinguished from Synalpheus quadriartic- 851 ulatus by the telson, which bears two pairs of very large dorsal spines and two pairs of large subequal posterior spines, with the in- terspace between the submedian spines ob- solete. In S. quadriarticulatus the dorsal spines are smaller, the posterior spines smaller, unequal, and the submedian spines well separated by a convex posterior mar- gin. In addition the fingers of the minor che- la are bidentate in S. dorae, simple in S. quadriarticulatus. Synalpheus dorae is less closely related to Synalpheus redactocarpus, which has a well developed lamella on the scaphocerite and a telson lacking dorsal spines, with a well developed convex posterior margin with slender unequal posterior spines. In |S. re- dactocarpus the orbital spines are longer, more acute and convergent, and the rostrum is longer and narrower. The palm of the major chela lacks a distodorsal spine and the fingers of the minor chela are less broad- ened and with simple finger tips. In its general morphology S. dorae also shows a close resemblance to Synalpheus pescadorensis Coutiére, 1905, from which it differs, in addition to the 4-segmented second pereiopod carpus, in having a much smaller lamella in the scaphocerite and lack- ing a very long posterolateral process on the protopodite of the uropod. The thoracic sternites in Synalpheus dor- ae are narrow and the coxae of the pereio- pods are medially flattened and apposed. The small spines on the coxa of the third and fourth pereiopods project posteriorly in this position, a feature that does not appear to have been noted in other species of Syn- alpheus. Remarks. —Of the 162 specimens avail- able, only a few are obvious juveniles on account of their small size. The rest are of adult size for a small Synalpheus species, yet no ovigerous females are present and all intact specimens are of the usual male form, with an angular posterior tooth on the first abdominal pleuron, and in the morphology of the first and second pleopods (Dardeau 852 1984). Similar populations have been re- ported in other sponge-inhabiting Synal- Dheus species: S. neptunus germanus (Ban- ner & Banner, 1975, 44 specimens), S. paradoxus (Banner & Banner, 1981, 241 4, 4 9), S. crosnieri (Banner & Banner, 1983, 144 3, 3 9). The condition was first noted in Atlantic species by Coutiére (1909) in S. pectiniger Coutiére, and Chace (1972) has also reported on a paucity of ovigerous fe- males in some West Indian populations of S. paraneptunus Coutiére, so the phenom- enon is clearly of widespread distribution. Banner & Banner (1981) have suggested that toxic or pheromone-like metabolites pro- duced by sponges may be the cause of ab- normal Synalpheus populations. Progress in the study of these peculiar populations is clearly handicapped by the lack of any de- tailed knowledge of the sponge hosts of many species of Synalpheus and of the degree of host specificity in most of the sponge-as- sociated species. Acknowledgments Iam particularly grateful to the late Dora May Banner for drawing my attention to these new shrimps, to J. N. A. Hooper for the identification of the sponge host, and to Dr. J. C. Markham for the identification of the bopyrid parasite. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Banner, A. H., & D. M. Banner. 1983. Annotated checklist of the alpheid shrimp from the West- ern Indian Ocean.— Travaux et Documents de PORSTOM 158:1-164, figs. 1-14. Banner, D. M., & A. H. Banner. 1975. The alpheid shrimp of Australia, II]. The genus Synal- pheus.—Records of the Australian Museum 29(12):267-389, figs. 1-29. ——.,, & 1981. Annotated checklist of the alpheid shrimp of the Red Sea and the Gulf of Aden.—Zoologisches Verhandelingen 190:1-99, figs. 1-12. Chace, F. A., Jr. 1972. The shrimps of the Smith- sonian-Bredin Caribbean Expeditions with a summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia).— Smithsonian Contributions to Zoology 98:i-x, 1-179, figs. 1-61. Coutiére, H. 1909. The American species of the snap- ping shrimp of the genus Synalpheus.—Pro- ceeding of the United States National Museum 659:1-93, figs. 1-54. Dardeau, M. R. 1984. Synalpheus shrimps (Crusta- cea: Decapoda: Alpheidae). I. The Gambarel- loides group, with description of a new species. — Memoirs of the Hourglass Cruises 7(2):1-125, figs. 1-54. Division of Natural Sciences, Northern Territory Museum, P.O. Box 4646, Darwin, Australia 5794. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 853-860 REDESCRIPTIONS OF TETRALIA CAVIMANA HELLER, 1861 AND TRAPEZIA CYMODOCE (HERBST, 1799) FIRST STAGE ZOEAS WITH IMPLICATIONS FOR CLASSIFICATION WITHIN THE SUPERFAMILY XANTHOIDEA (CRUSTACEA: BRACHYURA) Paul F. Clark and Bella S. Galil Abstract.—The first stage zoea of Tetralia glaberrima (Herbst, 1790) (now T. cavimana Heller, 1861) and Trapezia cymodoce (Herbst, 1799) of Gurney (1938) are redescribed and compared both with the original description and with the description of Tetralia glaberrima by Al-Kholy (1963). Differences between the two larvae are tabulated and larval characters that appear to support the separation of the families Platyxanthidae and Trapeziidae, as pro- posed in a classification of adult xanthoids by Guinot (1977 and 1978), are identified. Guinot (1978) proposed a new classifi- cation of adult Brachyura based primarily on the position of female and male genital openings. Eight families were recognized within the superfamily Xanthoidea, and she supported Ortmann (1897) in giving the rank of family to the Trapeziinae as defined by Miers (1886). But for a few minor changes, Guinot indicated that the family contained all the genera listed by Balss (1957) in his subfamily Trapeziinae. Guinot did not use any larval characters to corroborate her classification. Rice (1980) and Martin (1984) related larval groupings, based on chaetotaxy, to various classifications of adult xanthoids. Both found that many existing descriptions of xanthoid larvae were inadequate or un- reliable, and each attempted to use larval evidence to resolve incongruences between different schemes of adult classification. Rice found that his larval groupings did not cor- respond to the simple divisions of the Xan- thidae by Balss (1957), while Martin (1984) endorsed the scheme proposed by Glaessner (1969) because it was based on fossil and recent forms. But later, Martin et al. (1985) stated that evidence appeared partly to sup- port Balss, as their group I corresponded to the subfamily Xanthinae. Rice (1980) and Martin et al. (1985) agreed that the genus Homalaspis warranted separation from the remainder of the Xanthinae of Balss and thereby corroborated the Platyxanthidae of Guinot. Rice therefore tended to support the more complex divisions suggested by Guinot. The larval descriptions of Tetralia gla- berrima (Herbst, 1790) (now JT. cavimana Heller, 1861; see synonomy of Galil, 1988) and Trapezia cymodoce (Herbst, 1799) by Gurney (1938) and of Tetralia glaberrima by Al-Kholy (1963) are incomplete. There- fore, the aim of this paper is to redescribe these larvae and use this information to re- examine the classification of the xanthoids. Materials and Methods Material collected and hatched at the Bi- ological Station, Ghardaqa, Egypt, by Gur- ney (1938) was fixed originally in formalin and recently transferred to 70% ethanol. The female and the first zoea of 7Trapezia cy- 854 Fig. 1. pezia cymodoce. Scale bar = 0.1 mm. Carapace of a, Tetralia cavimana; b, Tra- modoce (Herbst) are stored in the British Museum (Natural History), registration number 1986:915, and those of Tetralia cavimana Heller, were registered as 1986: 53. Dissected appendages were mounted in polyvinyl lactophenol and examined using an Olympus BH-2 microscope with No- marski interference contrast. Drawings were made with the aid of a camera lucida. When comparing the first stage zoea of the two species, the setal arrangement on many ap- pendages was similar and it was only nec- essary to figure fully the chaetotaxy of one species, Tetralia cavimana, and illustrate the differences in Trapezia cymodoce. Tetralia cavimana Heller, 1861 Figs. la, 2a—d, 3a, b, 4c, d Tetralia glaberrima (Herbst, 1790).—Gur- ney, 1938:77, pl. HI, figs. 29-33.—Al- Kholy, 1963:138, pl. I, figs. 1-7.—Wil- liamson, 1970:37. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. na; c, Maxillule; d, Maxilla. Trapezia cymodoce: e, Se- tation of maxilla coxal, basial and endopod lobes. Scale bar = 0.1 mm. Tetralia cavimana: a, Antennule; b, Anten- non Tetralia glaberrima (Herbst, 1790).— Al-Kholy 1963:139-140, pl. I, figs. 8-21, pl. II, figs. 22-33. First zoea.—Carapace (Fig. la): Long dorsal and rostral spines spinulate; 2 pairs of lateral spines, dorsal pair j-shaped and spinulate on dorsal margin, ventral pair smaller than dorsal pair, unarmed; one pair of posterodorsal setae; ventral margin of carapace with minute denticles, marginal setae absent; one pair of posterodorsal car- apace setae; eyes sessile with small setule on each eye. Antennule (Fig. 2a): Endopod absent; exopod unsegmented with 4 terminal es- thetascs, 1 terminal seta and 1 minute ter- minal spine. Antenna (Fig. 2b): Spinous process dis- tally spinulate; endopod absent; exopod with unequal terminal setae. VOLUME 101, NUMBER 4 Fig. 3. Second maxilliped. Trapezia cymodoce: c, Second maxilliped. Scale bar = 0.1 mm. Tetralia cavimana: a, First maxilliped; b, Mandible: Endopod (palp) absent. Maxillule (Fig. 2c): Coxal endite with 7 setae; basial endite with 5 processes on inner margin plus 2 minute teeth, single seta ab- sent from outer margin; endopod 2-seg- mented, proximal segment with | seta, dis- tal segment with 1 subterminal and 4 terminal setae. Maxilla (Fig. 2d): Coxal endite bilobed with 4+3(+1 minute tooth) setae; basial endite bilobed with 4(+ 1 minute tooth)+4 setae; endopod bilobed with 2+ 3 setae; sca- phognathite (exopod) with 4 marginal setae plus 1 long stout posterior seta. First maxilliped (Fig. 3a): Basis with 10 setae arranged 2,2,3,3; endopod 5-seg- mented with 2,2,1,2,5 setae respectively; exopod 2-segmented, distal segment with 4 terminal natatory setae. Second maxilliped (Fig. 3b): Basis with 4 setae; endopod 3-segmented with 1,1,4 se- 855 | ed [ 22 mt Shi eee Xe 7 y ‘ i] N\N Hit hy N y WES avis Hats j\ | [4 1\\ b sire bye | hosel ji) Fig. 4. Trapezia cymodoce: a, Dorsolateral spines of somites 1-5; b, Abdomen from dorsal aspect. Te- tralia cavimana: c, Dorsolateral spines of somites 1— 5; d, Abdomen from dorsal aspect. Scale bar = 0.1 mm. tae respectively; exopod 2-segmented, distal segment with 4 terminal natatory setae. Third maxilliped: Undeveloped. Pereiopods: Undeveloped. Abdomen (Fig. 4c, d): 5 somites; somites 2 and 3 each bearing | pair of dorsolateral processes; somites 1-3 with rounded pos- terolateral margins; somites 4 and 5 with small posterolateral spines; somites 2—5 with pair of posterodorsal setae; pleopods absent. Telson (Fig. 4d): Each long fork with | prominent and | small lateral spine plus | prominent dorsal spine; posterior margin with 3 pairs of setae. Trapezia cymodoce (Herbst, 1799) Figs. 1b, 2e, 3c, 4a, b Trapezia cymodoce (Herbst, 1799).—Gur- ney, 1938:76, pl. II, figs. 23-28. First zoea. —Carapace (Fig. 1b): long dor- sal and shorter rostral spine lightly spinu- late; single pair of curved, lightly spinulate lateral spines; one pair of posterodorsal se- 856 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Differences in first stage zoea of Tetralia cavimana Heller as described by Gurney (1938) and as described in this study. Character Dorsolateral process on abdominal somite 3 Posteriolateral spines on abdominal somites Number of spines on telson fork Gurney (as T. glaberrima) This study (p. 77 & pl. 3, figs. 30, 31) (Figs. 3d & 4c, d) absent present *3-5 4&5 **D B (1 lateral, 1 dorsal) (2 lateral, 1 dorsal) * Gurney (p. 77), lists somites 3—5 with small lateral spines, but figures a first stage (pl. 3, fig. 30) with small lateral spines on somites 4 and 5 only. ** Gurney figures (pl. 3, figs. 29, 30 & 31) the telson without the minute third spine, but states (p. 77) that a specimen caught in the plankton had this spine. tae; ventral margin of carapace with minute denticles, marginal setae absent; one pair of posterodorsal carapace setae; eyes sessile with small setule on each eye. Antennule: Endopod absent; exopod un- segmented with 4 terminal esthetascs, | ter- minal seta and | minute terminal spine. Antenna: Spinous process distally spi- nulate; endopod absent; exopod with un- equal terminal setae. Mandible: Endopod (palp) absent. Maxillule: Coxal endite with 7 setae; ba- sial endite with 5 processes on inner margin plus 2 minute teeth, single seta absent from outer margin; endopod 2-segmented, prox- imal segment with | seta, distal segment with | subterminal and 4 terminal setae. Maxilla (Fig. 2e): Coxal endite bilobed with 5+3(+1 minute tooth) setae; basial endite bilobed with 4(+1 minute tooth)+4 setae; endopod bilobed with 3 +2 setae; sca- phognathite (exopod) with 4 marginal setae plus | long stout posterior seta. First maxilliped: Basis with 10 setae ar- ranged 2,2,3,3; endopod 5-segmented with 2,2,1,2,5 setae respectively; exopod 2-seg- mented, distal segment with 4 terminal na- tatory setae. Second maxilliped (Fig. 3c): Basis with 3 setae; endopod 3-segmented with 0,1,4 se- tae respectively; exopod 2-segmented, distal segment with 4 terminal natatory setae. Third maxilliped: Undeveloped. Pereiopods: Undeveloped. Abdomen (Fig. 4a, b): 5 somites; somites 2-5 each bearing a pair of dorsolateral pro- cesses; somites 1—3 with rounded postero- lateral margins; somites 3—5 with long pos- terolateral spines; somites 2-5 with pair of posterodorsal setae; pleopods absent. Telson: Each long fork with 1 prominent and 1 small lateral spine plus 1 prominent dorsal spine; posterior margin with 3 pairs of setae. Discussion The differences between the descriptions by Gurney (1938) and Al-Kholy (1963) of Tetralia cavimana Heller (as T. glaberrima Herbst) and the present study are listed in Tables 1 and 2. In his description of Tra- pezia cymodoce Gurney (1938) figured only abdominal somites 2 and 3 with dorsolat- eral processes, but on re-examination of this material they were found additionally on somites 4 and 5 (Fig. 4b). Although the first stage zoeas of Trapezia guttata Ruppell, 1830 and Trapezia maculata (MacLeay, 1838) have been described by Gurney (1938) and Al-Kholy (1963) respectively, these de- scriptions are inadequate for comparison. Differences in appendage chaetotaxy be- tween 7rapezia cymodoce (Herbst) and Te- tralia cavimana Heller first stage zoeas are listed in Table 3. Rice (1980), from larval descriptions, di- vided the Xanthidae into 4 groups. He com- VOLUME 101, NUMBER 4 857 Table 2.— Differences in first stage zoea of Tetralia cavimana Heller as described by Al-Kholy (1963) and as described in this study. Al-Kholy (as T. glaberrima) This study (p. 138 & pl. 1, figs. 2-7) (figs. 2-4) Character Terminal armature of antennule Terminal setae on exopod of antenna Setae on coxa of maxillule Terminal setae on endopod of maxillule Setae on bilobed coxa of maxilla Setae on bilobed basis of maxilla ~ Total no. of setae on scaphognathite Endopod setal formula of 1st maxilliped Basial setal formula of 1st maxilliped No. of endopod segments of 2nd maxilliped Setal formula of 2nd maxilliped endopod No. of basial setae of 2nd maxilliped Dorsolateral processes on 3rd abdominal somite Armature of telson fork 2 esthetascs 1 prominent lat- eral spine 4 esthetascs 1 seta & 1 minute spine 5 7 3 4 4+5 4+3 2+3 4+4 4 5 1,3,0,0,3 DDIEDES 5(2,3) 10(2,2,3,3) 2 3 2,3} 1,1,4 1 4 absent present 1 prominent & 1 small lateral spine + 1 prominent dorsal spine * Al-Kholy figures 3 terminal setae on antenna exopod (pl. 1, fig. 2), but scores an exopod with 4 terminal setae in his description on page 138. pared these groupings with several adult classifications, and concluded that they gave some support to the divisions proposed by Guinot (1978). Martin (1984) recognized six groups (I-VI) within the Xanthidae, based on zoeal characters, but gave his groupings no formal taxonomic status. He adopted the classification of xanthoids proposed by Glaessner (1969), because his larval groups did not correspond to the eight families of Guinot. Rice (1980) and Martin (1984) agree on the suite of characters that defines their xan- thoid group III. The genera that they as- signed to their respective groups III differs. Eriphia, Homalaspis, Ozius and Tetralia form the group III of Rice. The group III of Martin contains Baptozius, Carpilius, Epixanthus, Paramedaeus, Pilumnoides, Platyxanthus and Trapezia in addition to those of Rice. Both include ASM 26 in group III. However, the present redescription of Trapezia cymodoce (Herbst) and Tetralia cavimana Heller first stage zoeas indicates that the group III of both workers can be divided into two subgroups A and B, the characters of which are defined in Table 4. Group A includes the first stages zoeas of Tetralia cavimana Heller and Trapezia cy- modoce (Herbst) and corresponds to the Trapeziidae of Guinot (1978). Group B comprises her Platyxanthidae and contains the zoeas of Homalaspis plana (A. Milne Edwards) (Fagetti 1970), Platyxanthus Table 3.—Differences in chaetotaxy of appendages between first zoea stages of Trapezia cymodoce (Herbst) and Tetralia cavimana Heller. Trapezia_ Tetralia Character cymodoce cavimana No. of lateral carapace spines 1 2 Setation of maxilla endopod 3+2 2+3 Setation of maxilla coxa 5+3 3+4 Setal formula of endopod of 2nd 0,1,4 1,1,4 maxilliped Basial setae of 2nd maxilliped 3 4 Abdominal somites with dorso- 2-5 2&3 lateral processes Abdominal somites with dorso- 3-5 4&5 lateral spines (long) (short) 858 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 4.—Proposed characters that subdivide the xanthid group III of Martin (1984) and Rice (1980). Character Group A Group B Setation of distal endopod seg- ment of maxillule Setation of maxilla endopod only) Setation of basal endopod seg- 2 ment of Ist maxilliped Setation of distal endopod seg- 4 ment of 2nd maxilliped 1 subterminal + 4 terminal subterminal setae absent (2 or 3 terminal setae 2 subterminal + 4 terminal subterminal setae present (2 subterminal + 3 terminal setae) 73 ¥*6 * Lumare & Gozzo (1972) figure the zoeal stages of Eriphia verrucosa as variously having 2 or 3 setae on this segment. Examination of E. verrucosa \st stage zoeas from Ischia, Italy (plankton caught material by Galil) revealed 3 setae in this position. Hashmi (1970) described and figured the Ist zoeal stage of Eriphia laevimana smithii (MacLeay) with 2 setae, but re-examination of his material, BM(NH) registration number 1986:908, also revealed 3 setae. Wear (1968) illustrated the 1st maxilliped of Ozius truncatus H. Milne Edwards stage I zoea with only 1 seta on the basal endopod segment. Other zoeas in this group have 3 setae. ** The following Ist stage zoea all have 6 setae present on the distal segment of second maxilliped endopod; Baptozius vinosus (H. Milne Edwards), Eriphia laevimanus smithii MacLeay, Eriphia verrucosa (Forskal), Hom- alaspis plana (A. Milne Edwards), Monodeus couchii (Couch) & Platyxanthus patagonicus A. Milne Edwards. crenulatus (A. Milne Edwards) (Menu-Mar- que 1970) and P. patagonicus A. Milne Ed- wards (Iorio & Boschi 1986). Guinot (1978) also lists the genera Homalaspis and Pla- tyxanthus in her Platyxanthidae. Other lar- vae that fit in group B include the menippids Baptozius vinosus (H. Milne Edwards) (Saba et al. 1978a), Epixanthus dentatus (White) (Saba et al. 1978b), Eriphia laevimana smithii MacLeay (Hashmi 1970), E. spini- trons (Herbst) (Bourdillon-Casanova 1960; Hyman 1925; Paolucci 1910), E. verrucosa (Forskal) (Lumare and Gozzo 1972), Ozius rugulosus Stimpson (Kakati and Nayak 1977), O. truncatus H. Milne Edwards (Wear 1968) and the pilumnid Pilumnoides per- latus (Poeppig) (Fagetti & Campodonico 1973). None of these are listed in Guinot’s classification, but on the basis of larval char- acters these genera appear to have affinities with the Platyxanthidae. The xanthids Monodaeus couchi (Couch) (Ingle 1983) and Paramedaeus noelensis (Ward) (Suzuki 1979) are also grouped in the Platyxanthi- dae. In her classification of adult Xanthidae, Guinot (1978) placed Monodaeus and Par- amedaeus in the sub-family Euxanthinae Alcock, 1898. Apart from this anomaly, the larval evidence presented here appears to correspond to the classification of adult Pla- tyxanthidae and Trapeziidae as proposed by Guinot (1978). The status of the third stage zoea of “ASM 26 (Menippinae or Trapeziinae),”’ as de- scribed by Rice and Williamson (1977:52- 54, fig. 27) remains uncertain; it does not have a menippid type antenna, maxillule or maxilla endopod. Martin (1984) believed that ““ASM 26” was more likely to be in the Trapeziinae, but from evidence presented here it does not appear to share the char- acters defining group A (see Table 4). Carpilius was placed by Martin et al. (1985) in xanthid group III. Laughlin et al. (1983) described the zoeal stages of the coral crab Carpilius corallinus (Herbst) and noted a number of diagnostic characters: 5 zoeal stages, the separation of the 6th abdominal somite from the telson in stage II zoea, ap- pearance of pleopod buds in stage II, the large size of the zoea, the increased numbers of natatory setae in the maxillipeds in later stages (i.e., zoea IV with 15-16 and zoea V with 20-22 natatory setae), and the large size of the mandibles. In combination, these larval characters are interpreted by Laugh- VOLUME 101, NUMBER 4 lin et al. (1983) as evidence supporting the establishment of the family Carpiliidae by Guinot (1978). Careful re-examination of other xantho1- dean larval descriptions may further sub- stantiate the classification proposed by Guinot (1978). Literature Cited Al-Kholy, A. A. 1963. The zoeal stages of Tetralia glaberrima (Herbst), from the Red Sea.—Pub- lications of the Marine Biological Station, Ghar- daqa, Red Sea, Cairo No. 12:137-144. Balss, H. 1957. Decapoda VIII Systematik in H. G. Bronn, Klassen und Ordnungen des Tier- reichs Band 5, Abteilung 1, 7(12):1505-1672. Bourdillon-Casanova, L. 1960. Le méroplankton du Golfe de Marseille: les larves de Crustacés Déca- podes.—Recueil des Travaux de la Station d’Endoume, Marseille 30:1-—286. Fagetti, E. 1970. Desarrollo larval en el laboratorio de Homalaspis plana (Milne Edwards) (Crus- tacea, Brachyura, fam. Xanthidae).— Revista de Biologia Marina, Valparaiso 14:29-49. —., & I. Campodonico. 1973. Larval develop- ment of Pilumnoides perlatus (Brachyura: Xan- thidae) under laboratory conditions. Marine Bi- ology. First International Interdisciplinary Conference on Marine Biology, Washington 18: 129-139. Galil, B.S. 1988. Trapeziidae (Decapoda: Brachyura: Xanthoidea) of the Red Sea, Israel Journal of Zoology, Jerusalem 34(314):159-182. Glaessner, M. F. 1969. Decapoda. Pp. 399-533 in R. C. Moore, ed., Treatise on invertebrate pa- leontology Part R, Arthropod 2. Geological So- ciety of America and University of Kansas, Connecticut & New York. Guinot, D. 1977. Propositions pour une nouvelle classification des Crutacés Décapodes Brachy- oures.—Compte Rendu des Séances de la So- ciéte de Biologie, Paris sér. D, 285:1049-1052. . 1978. Principes d’une classification évolutive des Crustacés Décapodes Brachyoures.— Bulle- tin Biologique de la France et de la Belgique 112(3):211-292. Gurney, R. 1938. Notes on some decapod Crustacea from the Red Sea. VI-VIII.— Proceedings of the Zoological Society of London Ser. B. 108:73- 84. Hashmi, S.S. 1970. Study on the larvae of the family Xanthidae (Heteropanope, Eurycarcinus, and Eriphia) hatched in the laboratory. Brachyura: Decapoda.— Agriculture Pakistan, Karachi 21(4):457-478. Hyman, O. W. 1925. Studies on the crabs of the 859 family Xanthidae.— Proceedings of the United States National Museum 67:1-22. Ingle, R. W. 1983. A comparative study of the larval development of Monodaeus couchi (Couch), Xantho incisus Leach and Pilumnus hirtellus (Linnaeus) (Crustacea: Brachyura: Xanthi- dae).— Journal of Natural History, London 17: 951-978. Iorio, I. M., & E. E. Boschi. 1986. Studies on the larval stages of the crab Platyxanthus patagon- icus, from laboratory rearing and plankton sam- ples.—Journal of Agriculture in the Tropics 1(1): 7-24. Kakati, V. S., & V. N. Nayak. 1977. Larval devel- opment of the xanthid crab, Ozius rugulosus Stimpson (Decapod, Brachyura) under labora- tory conditions. — Indian Journal of Marine Sci- ences, New Delhi 6:26-30. Laughlin, R. A., P. J. Rodriguez, & J. A. Marval. 1983. Zoeal stages of the coral crab Carpilius coralli- nus (Herbst) (Decapoda, Xanthidae) reared in the laboratory. —Crustaceana 44(2):169-186. Lumare, F., & S. Gozzo. 1972. Sviluppo larvale de Crostaceo Xanthideae Eriphia verrucosa (For- skal, 1775) in condizioni di laboratorio. — Bol- lettino di Pesca, di Piscicottura edi Idrobiolo- gica, Roma 27(1):185—209. Martin, J. W. 1984. Notes and bibliography on the larvae of xanthid crabs, with a key to the known xanthid zoeas of the western Atlantic and Gulf of Mexico.—Bulletin of Marine Science 34(2): 220-239. , F. M. Truesdale, & D. L. Felder. 1985. Larval development of Panopeus bermudensis Bene- dict and Rathbun, 1891 (Brachyura, Xanthidae) with notes on zoeal characters in xanthid crabs. — Journal of Crustacean Biology 5(1):84—-105. Menu-Marque, S. A. 1970. Desarrollo larval de can- grejo Platyxanthus crenulatus (A. Milne Ed- wards, 1879) en el laboratorio (Decapoda, Brachyura, Xanthidae).— Physis, Buenos Aires 29(79):477-494. Miers, E. J. 1886. Report on the Brachyura collected by H.M.S. Challenger during the years 1873- 1876. Report on the Scientific Results of the voyage of H.M.S. Challenger during the years 1873-76, Zoology 17:xli + 362 pp. Ortmann, A. E. 1897. Die geographische Verbreitung de Decapoden-Familie Trapeziidae.—Zoolo- gische Jahrbucher. Abteilung fur Systematik 10: 201-216. Paolucci, C. 1910. La zoea di Eriphia spinifrons. — Rivista Mensile Pesca, Pavia 5:33-40. Rice, A. L. 1980. Crab zoeal morphology and its bearing on the classification of the Brachyura. — Transactions of the Zoological Society of Lon- don 35:271-424. , & Williamson, D. I. 1977. Planktonic stages 860 of Crustacea Malacostraca from Atlantic sea- mounts. — “Meteor” Forschungsergebnisse, Ge- briider Borntraeger, Berlin-Stuttgart Reihe D, Biologie, No. 26:28-64. Saba, M., M. Takeda, & Y. Nakasone. 1978a. Larval development of Baptozius vinosus (H. Milne Ed- wards). — Proceedings of the Japanese Society of Systematic Zoology, Tokyo 14:25-38. , & 1978b. Larval develop- ment of Epixanthus dentatus (White) (Brachy- ura, Xanthidae).— Bulletin of the National Sci- ence Museum (Zoology), Tokyo 4:151-161. Suzuki, H. 1979. Studies on the zoea of the two xan- thid crabs, Paramedaeus truncatus (de Haan) (Crustacea, Brachyura, Xanthidae).—Proceed- ings of the Japanese Society of Systematic Zo- ology 16:35—52. Wear, R. G. 1968. Life history studies on New Zea- land Brachyura 2. Family Xanthidae. Larvae of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Heterozius rotundifrons A. Milne Edwards, 1867, Ozius truncatus H. Milne Edwards, 1834, and Heteropanope (Pilumnopeus) serratifrons (Kin- ahan, 1856).—New Zealand Journal of Marine and Freshwater Research 2:698-707. Williamson, D.I. 1970. Onacollection of planktonic Decapoda and Stomatopoda (Crustacea) from the east coast of the Sinai Peninsula, northern Red Sea.—Bulletin of the Sea Fisheries Re- search Station, Israel No. 56:3-48. (PFC) Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD, England; (BSG) De- partment of Zoology, George S. Wise Fac- ulty of Life Sciences, Tel Aviv University, Israel. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 861-865 ALLOPOTAMON, A NEW GENUS FOR THE FRESHWATER CRAB POTAMON (POTAMONAUTES) TAMBELANENSIS RATHBUN, 1905 (CRUSTACEA: DECAPODA: POTAMIDAE) FROM THE TAMBELAN ISLANDS Peter K. L. Ng Abstract. —The types of the poorly known potamid crab, Potamon (Potamo- nautes) tambelanensis Rathbun, 1905, are reexamined, and its systematic po- sition clarified. The male first pleopod structure is unusually twisted, unlike any other potamid known from Southeast Asia, and warrants the establishment of a new genus, Allopotamon, for the species. Its affinities with allied Asiatic genera are also discussed. Potamon (Potamonautes) tambelanensis was described from the Tambelan Islands in the Straits of Kalimantan, between Su- matra and Borneo, by Rathbun in her im- portant revision of the world freshwater crabs in 1905. Although she provided a de- tailed description with figures, the drawings, especially of the male first pleopods, were too diagrammatic. In Bott’s (1970b) revi- sion of the Asian and Australian freshwater crabs, the species was mentioned only very briefly together with Jsolapotamon chaseni (Roux, 1934). He implied that Potamon tambelanensis should be placed in the genus Isolapotamon Bott, 1968, in the family Is- olapotamidae Bott, 1970a. A reexamination of the pair of types showed, however, that Rathbun’s species should be placed in a separate genus. In this paper I describe a new genus, A//opotamon, for Potamon tambelanensis. Detailed illus- trations of its gonopods are provided and its affinities with related taxa are discussed. Types are deposited in the National Mu- seum of Natural History (USNM), Smith- sonian Institution, Washington, D.C. The abbreviations G1 and G2 are used for the male first and second pleopods, respective- ly. Measurements are of the carapace breadth and length, respectively. Allopotamon, new genus Diagnosis. —Carapace quadrilateral, gas- tric and branchial regions distinctly inflated, dorsal surface strongly convex. Exopod of third maxilliped with long flagellum. G1 rel- atively stout, strongly twisted, terminal seg- ment distinctly demarcated from subter- minal, tip pectinated, slightly truncate, pointing towards sternum, dorsal fold di- lated. G2 with long flagellum, about half length of basal segment. Type species. —Potamon (Potamonautes) tambelanensis Rathbun, 1905. Etymology.—The name is derived from the Greek “‘allos’’ for ‘the other,’ and “‘Po- tamon,” the type genus of the family. The gender is neuter. Allopotamon tambelanense (Rathbun, 1905), new combination Figs. i Potamon (Potamonautes) tambelanensis Rathbun, 1905:182-183, pl. 15 fig. 4, fig. 46. Potamon (Potamon) tambelanensis Bott, 1970b:191. Diagnosis. — As for the genus. Description. —Only a few points need to 862 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Allopotamon tambelanense, holotype male. VOLUME 101, NUMBER 4 be added to supplement Rathbun’s (1905) excellent description of the species. The posterior margin of the epistome is pro- duced in three lobes; the median one 1s tri- angular, and delimited from the others by well developed sutures; the openings for the efferent branchial channels are concave. The G1 is relatively stout and appears twisted. The terminal segment is distinctly demar- cated from the subterminal; the dorsal lobe is dilated laterally, appearing swollen, ven- trolateral margin distal to the swollen area bears numerous short hairs, the distal part is slender, tapered, and the slightly truncate tip, pectinate. The subterminal segment ap- pears to be 2-segmented because of severe twisting, with the midlateral margin cleft, the proximal region of the distal part is strongly depressed. The flagellum of the G2 is long, about half the length of the basal segment, the latter is relatively straight, the tip of which is expanded into a cup-like structure, the proximal part of the basal seg- ment is broadly dilated. Material. —Holotype male (35.5 by 26.7 mm), paratype female (40.6 by 30.4 mm), USNM 23369, Big Tambelan Island (Pulau Tambelan Besar), South China Sea, Straits of Kalimantan, ca. 1°00’00’N, 107°33'26’E, leg. W. S. Abbott, 8-11 Aug 1899. Remarks.—Allopotamon tambelanense can be distinguished easily from all other potamids by its swollen carapace and char- acteristically twisted Gi. The G1 structure of A. tambelanense is like no other pres- ently known potamid from Southeast Asia, and warrants the establishment of a new genus for the species. The severity of the twisting, especially in the median region, resembles that in the genus Candidiopota- mon Bott, 1967, especially that of C. rath- bunae (De Man, 1914) from the Republic of China (Formosa, Taiwan) and C. oki- nawaense Minei, 1973, from Okinawa, Ja- pan. Direct comparisons of the Gls of C. rathbuni and A. tambelanense show how- ever, that the terminal segment of 4. tam- belanense is very different, being more con- ical, and the tip dilated. The carapace of 863 Fig. 2. Allopotamon tambelanense, holotype male. A-F, Left G1; G, Left G2; A, Dorsal view; B, Ventral view; C, Mesial (upper marginal) view; D, Mantel (low- er marginal) view; E, Terminal segment (dorsal view); F, Terminal segment (ventral view). Candidiopotamon is flatter, more nearly square, and the male abdomen is much more quadrate than are their respective structures in A. tambelanense. Bott (1970b), in transferring Potamon (Potamiscus) chaseni Roux, 1934, to the ge- nus [solapotamon and family Isolapotam- idae, commented that “... In diese Ver- wandtschaft gehort vermutlich auch Potamon (Potamon) tambelanensis Rath- bun 1904 ... von der Insel Tambelan zwischen Boreno und Singapore” (p. 191). Although he implied a relationship with the genus /solapotamon, he did not formally classify Rathbun’s species with any known genus. In the general structure of the cara- pace, shape of the male abdomen, and pres- ence of a distinct flagellum on the exopod 864 of the third maxillipeds Al/lopotamon re- sembles [solapotamon, but their Gls argue for the separation of the two taxa. The Gls of Isolapotamon, in sharp contrast to those of Allopotamon, are very long and slender, the terminal segment very elongate, being almost as long as the subterminal. Most of the Isolapotamon species also have flatter carapaces. Possessing a very different G1 and lacking a flagellum on the exopod of the third maxilliped, Potamon chaseni also can- not be retained in the genus Jsolapotamon as currently defined (Ng 1986a, 1987b), but should instead be transferred to the genus Stoliczia Bott, 1966, s.s. (Ng, 1988). Al- though Bott (1970b) established a new fam- ily for the genus Jsolapotamon and its kin, Ng & Yang (1985, 1986) and Ng (1986a, 1987b) regarded the differences between the Isolapotamidae and Potamidae s.s. as too minor to justify two separate families. They synonymized the two families, with the Po- tamidae having seniority, and this classifi- cation is adopted herein. Bott (1970b) gave the date of Rathbun’s species as 1904, but it should be 1905. Rath- bun’s classic monograph on the freshwater crabs was originally intended for publica- tion as one volume, but eventually came out in three consecutive volumes, each with the same title but different dates of publication (1904 to 1906). They should thus be treated as separate publications (Schmitt 1973). Po- tamon tambelanensis was described in the second volume (7), published in 1905. Although Bott (1970b) dismissed the val- ue of the exopod flagellum of the third max- illiped in the classification of freshwater crabs, there has been some revival in its use (Rodriguez 1982, Ng & Yang 1985, Ng 1985, 1987a). With regards to the Southeast Asian potamid freshwater crab fauna, only the genera [solapotamon, Ranguna Bott, 1968 s.s., Johora Bott, 1968, sensu Ng, 1987a, and Terrapotamon Ng, 1986b) (partim), have a distinct flagellum. The genus Ran- guna, as defined by Bott (1970b), contained taxa from peninsular Malaysia, but Ng PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (1987a, 1988) revised it, restricting Ran- guna to species found north of the Kra of Isthmus (ca. 8°N). The Gls of these genera however, are very different from those of Allopotamon. The G1 terminal segment of Allopotamon resembles that of many Ran- guna species, but in A/lopotamon, itis stout- er, and the subterminal segment is stout and twisted. The general shape of the G1 resem- bles that of Terrapotamon, but in this genus, neither the terminal nor the subterminal segments are as complex. The carapace of Allopotamon is quite close to that of. Ter- rapotamon, being inflated, probably asso- ciated with their semiterrestrial habits. The male abdomen of Terrapotamon however, is more distinctly triangular, and the fla- gellum on the exopod of the third maxil- lipeds is either short as in 7. aipooae Ng, 1986b, or vestigial as in T. abbotti (Rath- bun, 1898) (fide Ng 1986b, 1988). The Tambelan Islands are 140 km west of Borneo and 280 km east of the Riau Ar- chipelago, and in all likelihood, the pro- longed isolation of these islands accounts for the unusual G1 structure of A. tambe- lanense. The montane habitat of the crab (preferred by most potamids) would prob- ably also serve to isolate the taxon further (Ng 1985, 1987a). Acknowledgments The author is very grateful to Dr. Ray- mond B. Manning (USNM) for sending him the types of Allopotamon tambelanense and for his comments about nomenclature. Thanks are also due to Dr. Wang Chia Hsiang (Taiwan National Museum) for the gift of a pair of Candidiopotamon rathbun- ae. Literature Cited Bott, R. 1970a. Betrachtungen uber die entwicklungs- geschichte und verbreitung der siisswasser krab- ben nach sammlung des naturhistorischen mu- seums in Genf/Schweiz.— Revue Suisse de Zoologie 77,(2), (24):327-344. VOLUME 101, NUMBER 4 1970b. Die stisswasserkrabben von Europa, Asien, Australien und ihre stammesgeschichte. Eine revision der Potamoidea und Parathelphu- soidea. (Crustacea, Decapoda). — Abhandlungen der Senckenbergischen Naturforschenden Ge- sellschaft, Frankfurt 526:1—338, pls. 1-58. Ng, P. K. L. 1985. Freshwater decapod crustaceans from Pulau Tioman, West Malaysia.—Zoolo- gische Mededelingen, Leiden 59(14):149-162. 1986a. New freshwater crabs of the genus Isolapotamon Bott, 1968 from Kalimantan (De- capoda: Potamidae).—Treubia, Bogor 29(3): 215-223. . 1986b. Terrapotamon gen. nov., anew genus of freshwater crabs from Malaysia and Thai- land, with description of a new species, Terra- potamon aipooae gen. et sp. nov. (Crustacea: Decapoda: Brachyura: Potamidae).— Journal of Natural History, London 20:445-451. . 1987a. A revision of the Malayan freshwater crabs of the genus Johora Bott, 1966 stat. nov. (Decapoda: Brachyura: Potamidae).— Malayan Nature Journal, Kuala Lumpur 41:13—-44. 1987b. Freshwater crabs of the genus /sola- potamon Bott, 1968 from Sarawak, Borneo (Crustacea, Decapoda, Brachyura, Potami- dae). Sarawak Museum Journal, Kuching ns. 37(58):139-153, pls. 4-10. 865 . 1988. The freshwater crabs of peninsular Ma- laysia and Singapore. Department of Zoology, National University of Singapore, Shinglee Pub- lishers, Singapore. 156 pp., 63 figs., 4 pls. ——, & C. M. Yang. 1985. On three new species of freshwater crabs from Singapore and West Malaysia (Crustacea, Decapoda, Brachyura).— Malayan Nature Journal, Kuala Lumpur 39(1): 57-73. ——, & 1986. A new freshwater crab of the genus Jsolapotamon Bott, 1968 from Sara- wak, Borneo (Decapoda, Brachyura, Potami- dae). —Indo-Malayan Zoology, Rotterdam 3(1): 15-18. Rathbun, M.J. 1905. Les crabes d’eau douce. — Nou- velles Archives du Muséum d’Histoire Natu- relle, Paris (4)7:159-321, pls. 13-22. Rodriguez, G. 1982. Les crabes d’eau douce d’Amé- rique. Famille des Pseudothelphusidae. — Office de la Recherche Scientifique et Technique Outre- Mer, Faune Tropicale 22:1—223. Schmitt, W. L. 1973. Mary J. Rathbun 1860-1943.— Crustaceana 24:285-297, pl. 1. Department of Zoology, National Uni- versity of Singapore, Kent Ridge Campus, Singapore 0511, Republic of Singapore. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 866-871 THE STATUS OF CRYPTOCHIRUS HONGKONGENSIS SHEN, 1936 (BRACHYURA: CRYPTOCHIRIDAE) Roy K. Kropp Abstract. —Morphological differences in the extent of the inner orbital angle, the length of the dactylus of the claw, and the shape of the abdomen are presented which support the removal of Pseudocryptochirus viridis Hiro from the synonymy of Cryptochirus hongkongensis Shen. This evidence also indicates that C. hongkongensis should be transferred to Neotroglocarcinus and consid- ered a subjective senior synonym of N. monodi Fize and Seréne. In a short paper, Shen (1936) described two new species of the coral gall crab genus Cryptochirus. One of these, C. granulatus, was synonymized with C. crescentus Ed- mondson, 1925 [now Opecarcinus crescen- tus, see Kropp & Manning (1987)] by Uti- nomi (= Hiro, 1944), an action followed by all subsequent authors. The status of C. granulatus is under review and will be dis- cussed in a subsequent manuscript on the genus Opecarcinus in the Indo-Pacific. The second species described by Shen was C. hongkongensis. Utinomi (1944) trans- ferred it to his genus Pseudocryptochirus as a subjective senior synonym of P. viridis Hiro, 1938, but erred by continuing to use the latter as the valid name for the taxon. This mistake was followed by Fize & Seréne (1957), but corrected by Takeda & Tamura (1981). Both of the major works on Indo- West Pacific gall crabs published since Uti- nomi have upheld the synonymy (Fize & Seréne 1957, Takeda & Tamura 1981). Both noted apparent discrepancies between Shen’s account and the morphology of P. viridis, but neither disputed the synonymy of the two. Here, I discuss the discrepancies men- tioned by Fize and Seréne and Takeda and Tamura and present additional evidence supporting the dissolution of the synonymy between P. viridis and C. hongkongensis. Furthermore, I argue that C. hongkongensis is a subjective senior synonym of 7roglo- carcinus monodi Fize & Seréne (1955), the type species of Neotroglocarcinus Fize & Se- réne (1957). Types I have not been able to locate the holotype of Cryptochirus hongkongensis Shen, 1936. Contrary to the assertion of Fize & Seréne (1957:59), it is probably not in the British Museum (Natural History), London (BMNH). Dr. Raymond B. Manning (Smithsonian Institution, Washington, D.C.), in 1984 and 1987, examined the gall crab collection in the British Museum and did not see any types for the species al- though he did find the holotype of C. gran- ulatus. Shen (1936) did not report the dis- position of the holotype of C. hongkongensis. Although relatively crude, his figures and description do provide enough information by which comparisons to other species can be made. The types for Neotroglocarcinus monodi (Fize & Seréne, 1955) are also not available, however material identified by Seréne is available from the BMNH and the Muséum National d’Histoire Naturelle, Paris (MNHN). Additional material ex- amined was collected by the author in Mi- cronesia (HAP and PHAP denote my col- lection numbers) and is deposited in the National Museum of Natural History, VOLUME 101, NUMBER 4 Smithsonian Institution, Washington, D.C. (USNM). A series of morphological features that were easily interpreted was selected from the figures and description of C. hongkong- ensis published by Shen (1936). This series was compared among the species in ques- tion. As the description of Shen’s species is based on a male, only males were used in the comparisons. The results are presented below in the form of brief comparative di- agnoses, based on males only, for each species. I have reproduced Shen’s figures of C. hongkongensis and provided compara- tive figures of P. viridis and N. monodi. Abbreviations used in the text are: km, ki- lometers; m, meters; ov, ovigerous; P, per- eopod. Size ranges of material examined are given in millimeters as carapace length (cl) x width. At the first occurrence in the text of col- lection localities in the Caroline Islands, lo- cality names are given as the new ortho- graphic spelling (Motteler 1986) followed parenthetically by the former spelling. Pseudocryptochirus viridis Hiro, 1938 Figs. la—c, 2a—d Material. — Viet Nam: Nhatrang (12°14’N, 109°12’E), Rocher Noir, Rte. 1643, on Tur- binaria sp., 1 2 (ov), 1 6, BMNH. Guam: Toguan Bay (13°17'N, 144°39’E), reef front south of river channel, 6 m, 27 May 1984, HAP 229, on T. stellulata (Lamarck, 1816), 1 2 (ov), | 6; USNM; Cocos Lagoon (13°14'N, 144°39’E), southwest corner of la- goon just inside barrier reef, 1 m, 6 Mar, 3 Oct 1984, HAP 155, 272, on Turbinaria stellulata, 3 2 (2 ov), 1 6, USNM. Belau (Palau): Ngeruktabel (Urukthapel) Is. (07°15'N, 134°24’E), north shore, west end of rock islands, 2 m, 22 Jul 1984, PHAP 166, on 7. reniformis Bernard, 1896, 2 2 (ov), USNM. Same locality: 2 m, 23 Jul 1984, PHAP 193, 199, on T. cf. patula (Dana, 1846), 7. reniformis, 3 2 (2 ov), USNM. Pohnpei (Ponape): Main lagoon, 867 inside barrier reef about 1.6 km north of Main Passage (07°00'N, 158°13’E), 2 m, 14 Nov 1984, PHAP 244, on T. cf. mesenter- ina (Lamarck, 1816), 1 2 (ov), 2 6, USNM. Ant Atoll (06°47'N, 147°58’E), reef front off Imwinyap Is., 100 m west of pass, 8 m, 17 Nov 1984, PHAP 284, on 7. reniformis, | 2, 1 6, USNM. Size ranges. —Females, 1.8 xX 1.6 to 3.3 x 2.8; smallest ovigerous female, 1.8 x 1.6; Males EG) Xue) tore xX a1eS) Diagnosis.— Anterior third of carapace slightly depressed, not sharply set off from posterior carapace, latter lacking grooves or depressions; internal orbital angle greatly exceeding anterolateral angle of carapace. Basal segment of antennule with mesial margin straight, dorsal surface with longi- tudinal row of spines near mesial margin. Width of abdominal somite 6 about '4 that of somite 3. Dactylus of cheliped (P-1) lon- ger than dorsal margin of palm, latter with spines along entire length. Propodus of P-3 about 1.4 times longer than high, dorsal margin with tubercles. Gonopod tapering sharply, mesial and lateral margins with plumose setae originating just proximal to midlength. Neotroglocarcinus monodi (Fize & Seréne, 1955) Figs. 1g—i, 2i-l Material. —Viet Nam: Nhatrang, Rte. 1590, on T. peltata (Esper, 1797), 1 2 (ov), 1 6, MNHN. Bai Miew, 11 Apr 1956, Rte. 1637, on T. peltata, 1 6, BMNH. Rocher Noir, 8 May 1956, Rte. 1643, on T. peltata, 1 2 (ov), BMNH. Size ranges. —Females, 3.2 x 2.7 to 4.6 x 4.1: smallest ovigerous female, 3.2 x 2.7; males, 3.2 X 2.9 to 3.8 x 3.4. Diagnosis.— Anterior third of carapace markedly depressed, sharply set off from posterior carapace, latter with series of shal- low, longitudinal depressions; internal or- bital angle slightly exceeding anterolateral angle of carapace. Basal segment of anten- 868 nule with mesial margin convex, dorsal sur- face with scattered tubercles, lacking lon- gitudinal row of spines near mesial margin. Width of abdominal somite 6 about *% that of somite 3. Dactylus of P-1 shorter than dorsal margin of palm, latter with few tu- bercles proximally. Propodus of P-3 about 1.8 times longer than high, dorsal margin entire. Gonopod not tapering sharply, me- sial and lateral margins with plumose setae originating at about midlength. Cryptochirus hongkongensis Shen (1936) Figs. 1d—f, 2e—h From Shen (1936). Size. —Male, 2.3 x 2.0. Diagnosis.— Anterior third of carapace markedly depressed, sharply set off from posterior carapace, surface of latter uncer- tain; internal orbital angle slightly exceeding anterolateral angle of carapace. Basal seg- ment of antennule with mesial margin con- vex, dorsal surface with scattered tubercles, lacking longitudinal row of spines near me- sial margin. Width of abdominal somite 6 about *% that of somite 3. Dactylus of P-1 shorter than dorsal margin of palm, latter with few tubercles proximally. Propodus of P-3 about 1.7 times longer than high, dorsal margin entire. Gonopod tapering sharply, mesial and lateral margins with simple setae originating near or distal to midlength. Discussion From the above comparisons and the fig- ures provided, it is clear that C. hongkong- ensis 18 quite different from P. viridis, and further, that C. hongkongensis strongly re- sembles N. monodi. These relationships are most strongly supported by features such as the relative extent of the internal orbital an- gle compared to the anterolateral angle of the carapace, the relative demarcation be- tween the anterior and posterior parts of the carapace, the relative length of the dactylus of the P-1, and the shape of the abdomen PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON as indicated by the relative widths of somite 3 and 6. Some evidence is equivocal. Setation on the dorsal margins of the walking legs (Fig. 2) allies C. hongkongensis more closely to P. viridis than to T. monodi. However, I have noticed that setation can be variable within gall crab species. Also, setules on walking leg setae are often difficult to see and may have been missed by Shen. The gonopod figured by Shen is problematic. It tapers, as does the gonopod of P. viridis. Yet, the setation differs between the two, both in type (plumose in P. viridis, simple in C. hongkongensis) and position of origin (proximal to midlength in P. viridis, mid- length or just distal in C. hongkongensis). Shen may have erred in figuring the gono- pod. Shen’s figures and descriptions of the mouthparts and antenna of C. hongkongen- sis are too general to be of use in resolving the affinities of each taxon. The evidence presented by Utinomi (1944) for synonymizing P. viridis with C. hongkongensis 1s weak. He noted (p. 702) the antennule of “hongkongensis seems akin to viridis’ and (p. 703) the third maxilliped of hongkongensis “‘shows close similarity to that of viridis.”’ He further argued (p. 725) that the probable identity of the two was supported by “distributional evidences that both forms have been recorded together from neighboring seas.” Fize & Seréne (1957:142) noted in par- ticular the differences in the relative extent of the internal orbital angle and the relative length of the dactylus of the P-1 among the three taxa that I have mentioned. However, they did not ally C. hongkongensis with T. monodi, stating that Shen described the car- apace as having “‘petites épines,” a feature they attribute to P. viridis, not T. monodi. This is not true. Shen (p. 23) describes the carapace as “‘finely granulate,” not as having spinules. Takeda & Tamura (1981:16) noted the same two discrepancies mentioned above and the relative demarcation between the VOLUME 101, NUMBER 4 869 Fig. 1. Dorsal, lateral views of carapace, and abdomen: a—c, Pseudocryptochirus viridis, Pohnpei, USNM, cl = 1.9 mm; d-f, Cryptochirus hongkongensis, from Shen (1936), cl = 2.3 mm; g-i, Neotroglocarcinus monodi, Viet Nam, MNHN B-18762, cl = 3.8 mm. All males. Not to scale. anterior and posterior carapace regions. They declared that these differences are ‘“‘too small” to warrant separation of the two species; I disagree. Differences of the mag- nitude presented here are enough to sepa- rate species of gall crabs. The evidence presented here supports the restoration of Pseudocryptochirus viridis Hiro to valid status. This evidence further indicates that Cryptochirus hongkongensis Shen is a subjective senior syyonym of Troglocarcinus monodi Fize & Seréne, the 870 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Pereopods | and 3, antennule, and gonpod: a—d, Pseudocryptochirus viridis, USNM; e-h, Cryptochirus hongkongensis, from Shen (1936); i-1, Neotroglocarcinus monodi, MNHN B-18762. All males. Not to scale. type species of Neotroglocarcinus Fize & Se- réne. Shen’s species should now be known as Neotroglocarcinus hongkongensis (Shen). Acknowledgments I am grateful to the University of Guam Marine Laboratory and the Marine Mari- culture Demonstration Center for support- ing my field work in Micronesia and to the Division of Crustacea (USNM) for provid- ing support for my systematic studies. Spec- imens used in this study were kindly pro- vided by A. Crosnier (MNHN) and R. Ingle (BMNH). The corals I collected in Micro- nesia were identified by R. H. Randall. R. B. Manning and G. J. Vermeij reviewed the manuscript and provided encouragement. VOLUME 101, NUMBER 4 Financial support was provided in part by the University of Maryland’s Department of Zoology and Graduate School and a grant to G. J. Vermeij by the Biological Ocean- ography Section of the National Science Foundation. This is from a dissertation to be submitted to the Graduate School, Uni- versity of Maryland, in partial fulfillment of the requirements for the Ph.D. degree in Zoology and is Contribution No. 249 of the University of Guam Marine Laboratory. Literature Cited Fize, A., & R. Seréne. 1957. Les hapalocarcinidés du Viet-Nam.—Archives du Muséum National d’Histoire Naturelle, Paris (7)5:1—202, figs. 1- 43, pls. 1-18. Kropp, R. K., & R. B. Manning. 1987. The Atlantic gall crabs, family Cryptochiridae (Crustacea, Decapoda, Brachyura).—Smithsonian Contri- butions to Zoology 462:1-21, figs. 1-10. 871 Motteler, L. S. 1986. Pacific island names.—B. P. Bishop Museum Miscellaneous Publication 34: 1-91. Shen, C-J. 1936. Notes on the family Hapalocarcin- idae (coral-infesting crabs) with descriptions of two new species.— Hong Kong Naturalist, Sup- plement 5:21-26, pls. 1-2. Takeda, M., & Y. Tamura. 1981. Coral-inhabiting crabs of the family Hapalocarcinidae from Ja- pan. VIII. Genus Pseudocryptochirus and two new genera.—Bulletin of the Biogeographical Society of Japan 36:14—27, figs. 1-3, pls. 1-4. Utinomi, H. 1944. Studies on the animals inhabiting reef corals. III. A revision of the family Hapa- locarcinidae (Brachyura) with some remarks on their morphological peculiarities. — Palao Trop- ical Biological Station Studies 2(4):688-731, figs. 1-16, pls. 3-5. Department of Zoology, University of Maryland, College Park, Maryland 20742; (Present address: Battelle Ocean Sciences, 1431 Spinnaker Drive, Ventura, California 93001). PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 872-882 THE STATUS OF CRYPTOCHIRUS CORALLIODYTES HELLER AND LITHOSCAPTUS PARADOXUS MILNE EDWARDS (BRACHYURA: CRYPTOCHIRIDAE) Roy K. Kropp Abstract.—The types of Cryptochirus coralliodytes Heller and Lithoscaptus paradoxus Milne Edwards were examined and found to differ in sculpture of the carapace, the epistome, and relative lengths of the carpus and merus of the fifth pereopod among other features. Therefore, the latter is removed from the synonymy of the former. The type of Cryptochirus rugosus Edmondson was examined and found to be indistinguishable from C. coralliodytes; thus, C. rugosus is placed in synonymy with C. coralliodytes. The available data suggest that Cryptochirus bani Fize & Seréne is synonymous with L. paradoxus. Lec- totypes are designated for C. coralliodytes and L. paradoxus and are described and figured. Two years after the description of the first known coral gall crab, Hapalocarcinus mar- supialis Stimpson, 1859, the second record- ed species, Cryptochirus coralliodytes, was described by Heller (1861a:19). Milne Ed- wards (1862:F10) followed this with the de- scription of a third species, Lithoscaptus paradoxus. All three species were rather in- completely described, probably because they were reasonably different from most other brachyurans known at the time. Although this inadequacy has not been a problem for the first species it has resulted in some mis- understanding regarding the latter two species. The problem regarding the identities of C. coralliodytes and L. paradoxus can be traced to the failure of authors to examine type specimens or to carefully consider in- formation provided in the original descrip- tions that should have been useful in sep- arating the two species. Paulson (1875) was the first to place the two in synonymy, but did so by erroneously considering L. par- adoxus a senior synonym of C. corallio- dytes. He did not justify his action. Richters (1880) agreed, but did correct the order of synonymy. Rathbun (1897) also noted Paulson’s error. After Calman (1900) fol- lowed Paulson’s action, all authors up until the review of the family of Fize & Seréne (1957) attributed the synonymy of the two species to Paulson and/or Calman without question or examination of the types (e.g., Edmondson 1933, Shen 1936, Utinomi 1944). Fize & Seréne (1957) discussed Cryp- tochirus in detail and examined the syntypes of Lithoscaptus and Cryptochirus that are in the collection of the Muséum National d’ Histoire Naturelle, Paris. In spite of doing so, they upheld the synonymy of the two species. More recently, Takeda & Tamura (1980) reviewed Cryptochirus, but did not alter the status of the two species. I examined the syntypes of C. corallio- dytes and L. paradoxus and determined that they are not synonymous. Herein I desig- nate lectotypes for each species and con- clude that Heller’s species is a subjective senior synonym of Cryptochirus rugosus Ed- mondson, 1933. Because C. rugosus is the type species of the genus Favicola Fize & VOLUME 101, NUMBER 4 Seréne, 1957, the latter should now be con- sidered a subjective junior synonym of Cryptochirus Heller, 1861. Materials and Methods I examined the male and female syntypes of Cryptochirus coralliodytes Heller housed in the Muséum National d’Histoire Natu- relle, Paris (MNHN) and Naturhistorisches Museum, Vienna (NMW), respectively; the syntypes of Lithoscaptus paradoxus Milne Edwards in the MNHN; and the holotype of Cryptochirus rugosus Edmondson held in the B. P. Bishop Museum, Honolulu (BPBM). Additional material examined came from the BPBM and my own collec- tions (denoted as HAP and PHAP) made in Micronesia in 1984. The Micronesian ma- terial is deposited in the National Museum of Natural History, Smithsonian Institu- tion, Washington, D.C. (USNM). Place names for collection sites in the Caroline Islands are from Bryan (1971). At the first occurrence in the text of each locality, the new orthographic spelling (Motteler 1986) is given followed parenthetically by the for- mer spelling. Subsequently, only the new spelling is used. Some of the Micronesian material was used in the preparation of the figures and for study by Scanning Electron Microscopy (SEM). In preparation for SEM, specimens were dissected and cleaned by gentle me- chanical agitation, and brushing with a fine paint brush. Specimens were then trans- ferred to 100% ethanol via a graded series and air-dried overnight. Dried specimens were mounted on stubs and sputter-coated with gold-palladium and viewed with a Cambridge Stereoscan-100 microscope at an accelerating voltage of 10 kv. Drawings were made with a camera lu- cida mounted on a Wild M-5 microscope. The cheliped was drawn so that the outer surface of the manus is in the plane of the printed page. This distorts the relative pro- portions of the other segments, particularly the merus. Male pleopods were prepared for 873 illustration by lactic acid digestion and staining using methods described in Kropp & Manning (1987), except acid fuchsin was substituted for fast green. The carapace length and width of each specimen were measured to the nearest 0.1 mm with an ocular micrometer on a Wild M-5 microscope and are reported in mm as length 2 times that of spines; outer surface flat, with granules dorsodistally, ventrally; ven- tral margin relatively straight, with few tu- bercles, fringed with pappose setae; ven- 876 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Cryptochirus coralliodytes, 2? (Guam, from SEM micrographs): a, Epistome; b, MXP-3 (outer view); c, Antenna (ventral view); d, Endopod of MXP-1 (outer view). Scale: a = 0.3 mm; b = 0.5 mm; c, d = 0.2 mm. trodistal angle with prominent tubercles (tubercles worn on lectotype). Merus height <2 times that of carpus. Dorsal margins of carpus and propodus with robust spines, simple setae; outer surfaces with tubercles dorsally, ventrally; carpus subequal in length to propodus. Dactylus with proximal tooth dorsally; tip with subterminal pore. P-3, P-4 similar in form to P-2, stockier, P-4 less setose. P-5 elongate, smooth; carpus longer than other segments; propodus, dactylus di- rected anteriorly. Sternite of P-1 with few granules, that of P-4 with median suture. Female opening longitudinal, oval, with hood; PLP-2 bira- mous, PLP-3 uniramous. Variations. — May have spines instead of tubercles at inner orbital angles, anterolat- eral angles, and on anterior surface of car- apace. Spines vary as to number and sharp- ness. The width and depth of the grooves on the carapace varies considerably. The dactylus of P-2 may be missing. Smaller fe- males may have a deeper depression on the anterior carapace, may be less spiny, and have a more elongate projection of the an- tennule base than larger females. The car- apace length: width ratio ranged from 1.2 to 1.7 with most crabs within 1.4 to 1.6. Paralectotype male (Fig. 3).—Similar to, smaller than female. Carapace regions dis- tinctly marked as female, spines, rounded tubercles less pronounced. Projection of an- tennule base more elongate, pointed. P-1 robust, palm inflated, with tubercles; dac- tylus slightly longer than dorsal margin of palm. Abdominal somites 5-7 narrower than somites 3—4; telson broadly rounded. PLP-1 reaching middle of sternite of P-1; slightly curved, apex sharply pointed, di- rected slightly laterally; lateral margin with stout setae. Variations.—P-1 much less robust in proximal tooth on the dorsal surface of the VOLUME 101, NUMBER 4 smaller males, with the dactylus relatively longer than paralectotype. Regions of car- apace may have fewer tubercles. Lithoscaptus Milne Edwards, 1862 Lithoscaptus Milne Edwards, 1862:F10 [type species: Lithoscaptus paradoxus Milne Edwards, 1862:F10, by monotypy; gen- der masculine. ] Lithoscaptus paradoxus Milne Edwards Figs. 4-6 Lithoscaptus paradoxus Milne Edwards, 1862:F10 [TL: Reunion; lectotype, MNHN]. Cryptochirus coralliodytes var. rubrolineata Fize & Seréne, 1957:40, fig. 5D, pl. 14, figs. E-H [TL: Nhatrang, Vietnam; loca- tion of type unknown]. Cryptochirus coralliodytes var. cubrolinea- ta.—Fize and Seréne, 1957:201 [erro- neous spelling]. Cryptochirus coralliodytes var. fusca Fize and Seréne, 1957:40, fig. SB [TL: Nha- trang, Vietnam; location of type un- known]. Cryptochirus coralliodytes var. parvulus Fize and Seréne, 1957:40, fig. SC [TL: Nha- trang, Vietnam; location of type un- known]. Cryptochirus bani Fize and Seréne, 1957: 44, figs. SF, 6, pl. 1, fig. 7 [TL: Nhatrang, Viet-Nam; location of type unknown]. Type.—Two specimens are in the vial from the MNHN labelled Lithoscaptus par- adoxus “TYPE.” One is a nonovigerous fe- male that is herein designated the lectotype. The carapace of this female is somewhat misshapen, but still recognizable. Both P-2 are missing, but the remaining pereopods are present as are all mouthparts. The pleo- pods are present and are uniramous. The second specimen is Cryptochirus corallio- dytes Heller. Milne Edwards indicated which specimen on which he based his description by his reference to uniramous female pleo- 877 pods (1862:F12). For this reason the larger female is selected as the lectotype. Material examined. —Indian Ocean: Re- union Island [20°18’S, 57°29’E], lectotype, 126.4 x 5.3 (MNHN). Pacific Ocean: Car- oline Islands: Belau: Ngeruktabel Is., patch reef among rock islands on northeast shore, PHAP 045, 2 m, 2 Jul 1984, on Goniastrea pectinata (Ehrenberg, 1834), 2 2(1 ov), 1 6; Mariana Islands: Guam: Luminao Reef [13°28’N, 144°39’E], reef flat toward Ma- gundas, HAP 315, | m, 13 Oct 1984, on P. daedalea, 2 2 (ov), 1 6; Cook Islands: Rar- otonga [21°14'S, 159°46’W], Ararua, Wilder and Parks, Jun-Jul 1929, [no host], 13 2 (ov) (BPBM 83221). Description. — Female, based on lectotype in conjunction with a Guam female (Fig. 4, 5, USNM). Carapace about |.2 times longer than broad, widest just posterior to mid- length. Anterior carapace with broadly W-shaped depression having scattered spines; anterior gastric region slightly in- flated; median gastric with 2 depressions; mid to posterior carapace with many round- ed tubercles, regions of carapace not well defined; cardio-intestinal region rimmed anteriorly, laterally with depression. An- terolateral margin of carapace spinous. Anterolateral angle of carapace with sin- gle spine, apex exceeding inner orbital angle, latter swollen, with subterminal spine. Front concave, entire, width about '2 that at an- terolateral angle, latter 2/5 greatest carapace width. Orbit V-shaped. Epistome with subparallel longitudinal ridges laterally; median area swollen, lack- ing ridge; anterior margin entire, straight, with scant median indentation. Basal segment of antennular peduncle with suboval projection extending slightly be- yond eyestalk, rounded distally, no angled lateral lobe; dorsal surface flat, without tu- bercles; entire margin with subequal spines. Ventral surface of second antennal segment with few granules, distal margin with few raised granules. Eye directed anterolaterally, extending just 878 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Cryptochirus coralliodytes, 6, (a—d from paralectotype, e from Guam 6, USNM): a, Carapace (dorsal view); b, Abdomen; c, Left P-1; d, Right P-2; e, PLP-1 (outer view). Scale: a = 0.9 mm; b-d = 1 mm; e = 0.1 mm. beyond anterolateral angle; cornea subter- minal, occupying distal quarter of stalk dor- sally. Stalk mostly exposed, not broadening proximally; ventral surface smooth. MXP-3 with exopod, mesial margin of ischium slightly convex, outer surface with many distally-raised granules. Merus longer than broad, width less than half that of is- chium. Carpus shorter than length of prop- odus and dactylus combined. Endopod of MXP-1 subquadrate, mesial margin about 3/5 length of lateral margin; anterior margin with sharp median curve, lined with stout simple setae. Chelipeds (P-1) with many scattered sim- ple setae on upper margins. Dactylus longer than dorsal margin of palm; cutting edge with low tooth proximally. Dorsal margin of palm with few tubercles proximally, outer surface smooth. Manus much smaller than merus. Dorsal margin of merus of P-2 with spines distally, fringed with pappose setae of length >2 times that of spines; outer surface flat, with tubercles distally; ventral margin con- vex, with few tubercles, fringed with pap- pose setae. Merus height >2 times that of carpus. Dorsal margins of carpus, propodus VOLUME 101, NUMBER 4 879 Fig. 4. Lithoscaptus paradoxus, 2 (Guam, USNM): a, Carapace (dorsal view); b, Carapace (lateral view); c, Thoracic sternites; d—h, P-1 to P-5. Scale: a = 0.8 mm; b = 2.1 mm; c-h = 1 mm. with spines, simple setae; outer surfaces with few tubercles; carpus longer than propodus. Dactylus lacking proximal tooth dorsally, tip with subterminal pore. P-3, P-4 similar in form to P-2; outer surfaces of carpi, prop- odi with longitudinal row of rounded tu- bercles near upper margins, upper margins with simple, pappose setae. P-5 elongate, with tubercles dorsally on proximal 3 seg- ments; merus and carpus subequal in length, each longer than propodus; propodus, dac- tylus directed anteriorly. Sternite of P-1 smooth, that of P-4 with median suture. Female opening longitudi- nal, oval, with anterior hood; PLP-2, PLP-3 uniramous. 880 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Lithoscaptus paradoxus, ? (Guam, from SEM micrographs): a, Epistome; b, MXP-3 (outer view); c, Antenna (ventral view); d, Endopod of MXP-1 (outer view). Scale: a, c, d = 0.2 mm; b = 0.4 mm. Variations.—Relative sculpture of the carapace variable, particularly the median gastric area which may have depressions from two to four in number which may vary from obvious to faintly detectable. The ex- tent of the depression on the anterior car- apace varies from occupying the entire sur- face between the anterolateral margins to somewhat less. The size and number of spines and/or tubercles is highly variable. The carapace length: width ratio ranged from 1.2 to 1.5 with most crabs within 1.3 to 1.4. In some specimens the lateral pro- jection of the antennule base is more elon- gate than described. Also, the anterolateral angles of the carapace may extend only slightly beyond the inner orbital angles. Male. — Based on specimens from Micro- nesia. Similar to, smaller than female. Car- apace detail similar to females, spines pro- portionally smaller. Projection of antennule base elongate, with apical spine. P-1 robust, palm inflated, with few tubercles; dactylus longer than dorsal margin of palm. Abdom- inal somites 3-7 similar in width, telson broadly rounded. PLP-1 slightly curved, apex sharply pointed, directed slightly lat- erally; reaching posterior of sternite of P-1; lateral margin with stout setae. Variations. —Smaller males have a rela- tively smooth carapace, with the anterior depressions more distinct. The inner orbital angle may equal or exceed anterolateral an- gle of the carapace. Remarks.—The original description of Cryptochirus bani by Fize & Seréne (1957) did not provide information necessary to distinguish it from Lithoscaptus paradoxus. Seréne (1962), in a discussion of some ma- terial from Rarotonga sent to him by Ed- mondson, alluded that C. bani might be synonymous with C. coralliodytes. Seréne thought that this material was very similar to C. bani. I have examined material from the same collection (BPBM S3221) and found them to be L. paradoxus. Without examination of the type of C. bani, there is some uncertainty, but it is likely that C. bani VOLUME 101, NUMBER 4 is a subjective junior synonym of L. para- doxus. Fize & Seréne (1957) named three varieties of C. coralliodytes based primarily on color differences, giving no substantial morphological data by which to distinguish them. The varieties, rubolineata, fusca, and parvulus are therefore considered subjective junior synonyms of L. paradoxus. Discussion Paulson (1875) felt that many taxono- mists of the period were not making useful contributions to systematics as a science, and that their research “‘provides only a use- less ballast.”” He was particularly critical of Heller and A. Milne Edwards. Yet careful consideration of the original species de- scriptions by Heller and Milne Edwards and comparisons of those with material at hand might have prevented Paulson’s confusion of C. coralliodytes with L. paradoxus. Hel- ler’s original species account (1861a:19) was scanty, but the generic description included a characterization of the endopod of the first maxilliped that is useful. This was sup- ported by his later (1861b), more detailed, account which included an accurate figure of the appendage (1861b, pl. IV, fig. 39). The endopod of the first maxilliped is quite different in L. paradoxus (triangular in cor- alliodytes versus subquadrate in paradoxus, compare Fig. 2d and Sd herein). Heller’s figure of the female type (1861b: pl. IV, fig. 33) shows enough carapace detail, despite Edmondson’s (1933:4) complaint, to distin- guish the two species. Milne Edwards also gave information suf- ficient to separate the two species by refer- ring to the uniramous condition of the fe- male pleopods (1862:F12). Although female PLP-2 form is often not a reliable character (McCain & Coles 1979; Kropp & Manning 1987), in this case it is applicable because among the specimens of either species that I have examined, the PLP-2 is consistently uniramous in L. paradoxus and consistently biramous in C. coralliodytes. Fig. 6. Lithoscaptus paradoxus, 6 (Guam, USNM): a, Carapace (dorsal view; b, Abdomen; c-d, Right P-1, P-2; e, PLP-1 (outer view). Scale: a-d = 1 mm; e = 0.1 mm. Several features other than the above in- formation from the original literature allow separation of the two species. The most ob- vious difference is that the regions of the carapace of C. coralliodytes are well defined whereas those of L. paradoxus are not. Ad- ditionally, in the former, the epistome has a median ridge, and leg P-5 is smooth, with the carpus longer than the other segments whereas there is no median epistomal ridge and leg P-5 is tuberculate dorsally, with the carpus and merus subequal in length in L. paradoxus. Leg P-2 permits the two species to be distinguished because the merus is larger relative to the carpus in Lithoscaptus than in Cryptochirus. The changes made here have a bearing on the other species presently included in Cryp- tochirus and Favicola. | am reviewing these species and will clarify their status as a part of a revision of the genera of cryptochirids. 882 Acknowledgments I am grateful to the University of Guam Marine Laboratory and the Marine Mari- culture Demonstration Center for support- ing my field work in Micronesia, to the Di- vision of Crustacea (USNM) for providing support for my systematic studies, and to the SEM lab (USNM) personnel for assis- tance with the SEM work. Specimens used in this study were kindly provided by V. Stagl (NMW), A. Crosnier (MNHN), B. Burch (BPBM), and R. Ingle (BMNH). The corals I collected in Micronesia were iden- tified by R. H. Randall. R. B. Manning and G. J. Vermeij reviewed the manuscript and provided encouragement. Financial support was provided in part by the University of Maryland’s Department of Zoology and Graduate School and a grant to G. J. Ver- meij by the Biological Oceanography Sec- tion of the National Science Foundation. This is from a dissertation to be submitted to the Graduate School, University of Maryland, in partial fulfillment of the re- quirements for the Ph.D. degree in Zoology and is Contribution No. 250 of the Uni- versity of Guam Marine Laboratory. Literature Cited Bryan, E. H., Jr. 1971. Guide to place names in the Trust Territory of the Pacific Islands (the Mar- shall, Caroline and Mariana Is.). Pacific Science Information Center, Bernice P. Bishop Mu- seum, Honolulu. Calman, W. T. 1900. On a collection of Brachyura from Torres Straits.—Transactions of the Lin- nean Society of London, second series (Zoology) 8(1):1-50, pls. 1-3. Edmondson, C. H. 1933. Cryptochirus of the central Pacific.— Bernice P. Bishop Museum Occasion- al Papers 10(5):3—23, figs. 1-6, pls. 1-4. Fize, A., & R. Seréne. 1957. Les hapalocarcinidés du Viet-Nam.—Archives du Muséum National d’Histoire Naturelle, Paris (7), 5:1—202, figs. 1- 43, pls. 1-18 [also published as Mémoires de P’Institut Océanographique de Nhatrang, 10°, with the same pagination and date]. Heller, C. 1861a. Synopsis der im rothen Meere vor- kommenden Crustaceen.— Verhandlungen der k. k. Zoologisch-Botanisch Gesellschaft in Wien 11:1-32. 1861b. Beitrage zur Crustaceen-fauna des PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rothen Meeres.—Sitzungsberichte der Kaiser- lichen Akademie Wissenschaften, Mathema- tisch-Naturwissenschaftliche Classe, Wien 43: 297-394, pls. 1-4. Kropp, R. K., & R. B. Manning. 1987. The Atlantic gall crabs, Family Cryptochiridae (Crustacea, Decapoda, Brachyura).—Smithsonian Contri- butions to Zoology 462:1—21, figs. 1-10. McCain, J.C., & S. L. Coles. 1979. A new species of crab (Brachyura, Hapalocarcinidae) inhabiting pocilloporid corals in Hawaii.—Crustaceana 36(1):81-89, figs. 1-3, pl. 1. Milne Edwards, A. 1862. Faune carcinologique de Pile de la Réunion. Pp. F1-F16 in L. Maillard, Notes sur l’ile de la Réunion (Bourbon). Paris. Motteler, L. S. 1986. Pacific island names.—B. P. Bishop Museum Miscellaneous Publication 34: 1-91. Paulson, O. 1875. Podophthalmata and Endioph- thalmata (Cumacea). Studies of the Crustacea of the Red Sea with notes regarding other seas. Part I:xiv + 144 pp. [In Russian]. Kiev. Rathbun, M. J. 1897. A revision of the nomenclature of the Brachyura.— Proceedings of the Biologi- cal Society of Washington 9:153-167. Richters, F. 1880. Decapoda. Pp. 139-178 in K. Mobius, Beitrage zur Meeresfauna der Insel Mauritius und der Seychellen. Berlin. Seréne, R. 1962. Species of Cryptochirus of Ed- mondson 1933 (Hapalocarcinidae).— Pacific Science 16:30-41, figs. 1-5. . 1966. Note sur la taxonomie et la distribution géographique des Hapalocarcinidae (Decapoda- Brachyura).— Proceedings of the Symposium on Curstacea held at Ernakulam, 12-15 Jan 1965. Marine Biological Association of India. Part I: 395-398. Shen, C-J. 1936. Notes on the family Hapalocarcin- idae (coral-infesting crabs) with descriptions of two new species. — Hong Kong Naturalist, Sup- plement 5:21-26, pls. 1-2. Takeda, M., & Y. Tamura. 1980. Coral-inhabiting crabs of the family Hapalocarcinidae from Ja- pan. V. Genus Cryptochirus.—Researches on Crustacea 10:45—56, figs. 1-2, pls. 2-4. Utinomi, H. 1944. Studies on the animals inhabiting reef corals. III. A revision of the family Hapa- locarcinidae (Brachyura) with some remarks on their morphological peculiarities. — Palao Trop- ical Biological Station Studies 2(4):688—731, figs. 1-16, pls. 3-5. Department of Zoology, University of Maryland, College Park, Maryland 20742; (Present address: Battelle Ocean Sciences, 1431 Spinnaker Drive, Ventura, California 93001). PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 883-889 THE STATUS OF CALLIANASSA HARTMEYERI SCHMITT, 1935, WITH THE DESCRIPTION OF CORALLIANASSA XUTHA FROM THE WEST COAST OF AMERICA (CRUSTACEA, DECAPODA, THALASSINIDEA) Raymond B. Manning Abstract. — Callianassa hartmeyeri Schmitt, 1935, a species of uncertain iden- tity, is redescribed. It proves to be a member of Corallianassa Manning, 1987. Specimens from the Eastern Pacific previously identified with Callianassa hart- meyeri or Callianassa placida De Man are described as a new species, Coral- lianassa xutha. In 1924, Balss identified a callianassid from Kingston, Jamaica, with Callianassa grandimana (Gibbes, 1850) (as Glypturus grandimanus), a species then unknown since its original description. Balss’ poor figures (Fig. 1) distinctly showed a species with a three-spined front, a large terminal cornea, and a major cheliped with ventral spines on the ischium and merus. Schmitt (1935:2, 3, 4), in his review of the North American callianassids, recognized that Balss’s species differed from C. grandimanus in having ventral spines on the ischium and merus of the cheliped, and proposed the name Cal- lianassa hartmeyeri for Glypturus grandi- manus sensu Balss, 1924. Callianassa hart- meyer! has remained “incertae sedis’”’ since Schmitt’s account was published (see re- marks in Biffar 1971:640-641, 649). In 1938, Hult reported a specimen of Cal- lianassa hartmeyeri from the Galapagos Is- lands, and in 1939 Schmitt reported addi- tional material from Clipperton Island in the East Pacific. Chace (1962) studied ma- terial from Clipperton Island, identified it with Callianassa placida De Man, 1905, and synonymized Glypturus grandimanus sensu Balss, 1924, and Callianassa hartmeyeri Schmitt, 1935, with C. placida. No additional material from the Ameri- cas identified with either C. hartmeyeri or C. placida has been reported. In 1987 I showed that Callianassa gran- dimana Gibbes, 1850, was conspecific with the species then known as Callianassa bran- neri (Rathbun, 1900), and that Glypturus Stimpson, 1866, was distinct enough from Callianassa Leach, 1814, sensu stricto, to be recognized as a distinct genus. I also rec- ognized a new genus Corallianassa for Cal- lianassa longiventris A. Milne Edwards, 1870, from the Caribbean, and C. borra- dalei (De Man, 1928), from the Indo-West Pacific, and noted (p. 388) that ““The iden- tity of Callianassa hartmeyeri Schmitt is still uncertain.” Collection of a pair of chelipeds of a Cor- allianassa at Lake Worth Inlet, Florida, in March 1987, prompted me to try to deter- mine the identity of Callianassa hartmeyeri and to determine whether it was congeneric with or conspecific with Corallianassa lon- giventris (A. Milne Edwards, 1870). The chelipeds taken at Lake Worth Inlet proved to be those of C. /ongiventris. Through the kindness of H.-E. Gruner, Zoological Museum, Berlin, I was able to examine the type of Callianassa hartmeyeri. It proves to be a species of Corallianassa distinct from C. /ongiventris and C. borra- 884 dalei, the two species originally assigned to Corallianassa, from the Eastern Pacific population previously identified with P. hartmeyeri, and from C. placida, a species herein transferred to Corallianassa, as well. Roy Olerdéd, Swedish Museum of Natural History, Stockholm, allowed me to examine Hult’s specimen of Callianassa hartmeyeri from the Galapagos Islands; it proved to belong to the new species described below. All measurements are in millimeters (mm). Carapace length (cl) is the postorbital carapace length; total length (tl) is measured on the midline. Specimens have been de- posited in the collection of the National Museum of Natural History, Smithsonian Institution, Washington (USNM), the Swedish Museum of Natural History, Stockholm (SMNH), and the Zoological Museum, Berlin (ZMB). Corallianassa hartmeyeri (Schmitt, 1935) Figs. 1, 2 Glypturus grandimanus. —Balss, 1924:179, figs. 3, 4.—Schmitt, 1935:4 [footnote].— Biffar, 1971:640.—Manning, 1987:399. [Not Callianassa grandimana Gibbes, 1850.] Callianassa hartmeyeri Schmitt, 1935:3, 4; 1939:15.—Biffar, 1971:640, 641, 649, 651, 653.—Manning, 1987:388, 399. Material. —Jamaica: Kingston [Kingston Harbor = 17°57'N, 76°47'W]; Kiukenthal and Hartmeyer leg., 1907, 1 ovigerous fe- male (holotype, ZMB 20284). Description. —Carapace trispinous, with long rostral spine overreaching base of cor- nea. Anterolateral spines strong, separated from front by non-calcified membrane. Dorsal oval 0.8 carapace length. Eyes ex- tending to end of first segment of antennular peduncle, cornea large, terminal; eye with distinct ventromesial projection extending beyond cornea. Antennular and antennal peduncles incomplete; third maxilliped missing in holotype. Major cheliped large, distance from prox- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON imal end of merus to end of cheliped 1.6 times carapace length. Ischium lined ven- trally with tubercles and spines increasing in size distally. Merus longer than high, ven- tral margin convex, armed with small spines and tubercles decreasing in size distally. Carpus shorter than merus, about half palm length, much higher than long, cristate dor- sally and ventrally, inner side of ventral margin with few low serrations, appearing smooth in outer view, distal margin pro- duced into spine. Palm longer than high, longer than dactylus, with distinct proximal crest dorsally, cristate ventrally. Dactylus stout, hooked, shorter than palm, cutting edge with 2 low, obtuse teeth in proximal third. Fixed finger indistinctly toothed, with proximal notch in opposable margin. Fin- gers crossing when closed. Minor cheliped broken in type. Ischium spined ventrally, spines increasing in size distally. Merus longer than high, cristate dorsally and ventrally, with 4 low tubercles ventrally, only proximal (largest) visible in outer view. Carpus shorter than merus, slightly longer than high, cristate dorsally and ventrally, with ventrodistal spine. Second abdominal somite as long as sixth. Latter inflated, more than twice as long as telson. Telson trapezoidal, much shorter than uropods, lateral margins convergent posteriorly, posterior margin with rounded median projection. Size. —Ovigerous female holotype, bro- ken, cl 8 mm (tl, from Balss 1924, ca. 35 mm). Remarks. —Corallianassa hartmeyeri can be distinguished immediately from the only other Western Atlantic species of the genus, Corallianassa longiventris A. Milne Ed- wards, 1870, in that the carpus of the major cheliped is about half as long as the palm, whereas in C. /ongiventris it is longer than half the palm. This was one of the characters used by Schmitt (1935:4) to differentiate C. longiventris and C. hartmeyeri in his key. As both Chace (1962:619) and Biffar (1971: 649) pointed out, Schmitt inadvertently VOLUME 101, NUMBER 4 transposed the names in the couplet of his key differentiating C. hartmeyeri and C. lon- giventris. Corallianassa hartmeyeri differs from C. longiventris and resembles the species from the Eastern Pacific, named below, and dif- fers from C. placida (De Man), as well, in having a ventrodistal spine on the carpus of both first pereopods. The type of Corallianassa hartmeyeri could be compared with more than a dozen specimens of C. longiventris from Carib- bean localities in the collections of the Na- tional Museum of Natural History, as fol- lows: Bermuda (1 male, USNM 122449), Lake Worth, Florida (2 chelipeds, USNM 205698), Jamaica (1 male, | female, USNM 70799), Barbados (1 female, USNM 68939), Antigua (1 male, USNM 122448), and Car- rie Bow Cay, Belize (4 males, 1 ovigerous female, USNM 221700; 1 female and 2 che- lipeds, USNM 221701; 2 females (1 oviger- ous), USNM 205699; and 3 males, | female, USNM 205700). No specimen of the latter species was found to have the ventrodistal spine on the carpus of the chelipeds, and in all specimens of C. /ongiventris at all sizes, the carpus of the major cheliped is longer than half the dorsal length of the palm. The habitat of the type is unknown. The color pattern, distinctive in C. bor- radalei, C. longiventris, and C. xutha, new species, is unknown in C. hartmeyeri. Distribution. —Caribbean Sea, from Kingston, Jamaica. It is known only from the type locality. Corallianassa xutha, new species Fig. 3 Callianassa hartmeyeri.— Hult, 1938:7, figs. 1-4, pl. 1—Schmitt, 1939:15. Callianassa (Callichirus) placida. —Chace, 1962:617. [Part, not reference to Ed- mondson (1944). Not Callianassa placida De Man, 1905]. Callianassa placida. —Hernandez Aguilera et al., 1986:206. 885 DX b Fig. 1. Corallianassa hartmeyeri (Schmitt). Holo- type. a, Anterior margin of carapace; b, Major cheliped. (From Balss, 1924.) Material.— Mexico: Maria Madre Island, Baja California [21°35’N, 106°33’W], 4-10 fm (= 7-18 m), California Academy of Sci- ences, | female (USNM 142539). Clipperton Island [10°18’N, 109°13’W]: Shore collecting on rocks to south of landing place, Presidential Cruise sta 9, W. L. Schmitt leg., 21 Jul 1938, 1 male (holotype, USNM 77861); same data, | ovigerous fe- male (USNM 205906); Northeast side, low tide, reef flat, O-1 ft (= to 18 cm), collected with Endrin, Limbaugh, Chess, and Ham- bly leg., 13 Sep 1958, 1 male (USNM 110974); East end, coral reef, Reese, Bald- win, and Wintersteen leg., sta W58-289, 15 Aug 1958, 1 ovigerous female (USNM 110978); South shore, coral reef, Reese, Baldwin, and Limbaugh leg., sta W58-295, 1 ovigerous female (USNM 110979). Colombia: Port Utria [Ensenada Utria = 6°00'’N, 77°21'’W], mainland shore, first beach, intertidal, Velero III sta 418-35, 24 Jan 1935, 1 male (USNM 142538). Galapagos Islands: Academy Bay [0°45’S, 90°17'W], Indefatigable Island, lagoon, in sand at low tide, Rolf Blomberg leg., 9 Aug 1934, 1 male (SMNH 13883). All specimens other than the holotype are paratypes. Description. —Carapace trispinous, with long rostral spine overreaching base of cor- nea. Anterolateral spines strong, separated from front by non-calcified membrane. Dorsal oval 0.8 carapace length. Eyes ex- tending to end of first segment of antennular peduncle, cornea large, terminal; eye with distinct ventromesial projection extending beyond cornea. Antennal peduncle longer 886 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Corallianassa hartmeyeri (Schmitt). Holotype. a, Front; b, Front, in oblique lateral view; c, Major cheliped; d, Inner face of merus and carpus of major cheliped; e, Major chela, dorsal view; f, Minor cheliped (broken); g, Second abdominal somite; h, Sixth abdominal somite, telson, and uropods; i, Sixth abdominal somite and telson, lateral view. than antennular peduncle. Ischium and merus of third maxilliped not greatly ex- panded, ischium with spinous crest on inner surface, merus broader than long, dactylus simple, slender, slightly shorter and much narrower than propodus. Major cheliped very large, distance from proximal edge of merus to end of chela more than twice carapace length. Ischium lined ventrally with tubercles and spines, increas- ing in size distally. Merus slightly longer than high, ventral margin semicircular, dis- tinctly flared, with small spines and tuber- cles, decreasing in size distally. Carpus shorter than merus, more than half palm length, much higher than long, cristate dor- sally and ventrally, inner side of ventral margin with a few large tubercles, distal margin produced into spine. Palm slightly longer than high, about as long as dactylus, with distinct proximal crest dorsally, cris- tate ventrally. Dactylus stout, hooked, shorter than palm, cutting edge with 2 tri- angular teeth in proximal third, distal tooth more acute than proximal. Fixed finger in- distinctly toothed, with proximal notch on opposable margin. Fingers crossing when closed. Minor cheliped extending about to dac- tylus of major. Ischium spined ventrally, spines increasing in size distally. Merus longer than high, cristate dorsally and ven- trally, unarmed ventrally. Carpus shorter than merus, slightly longer than high, cris- tate dorsally and ventrally, with ventrodis- tal spine. Propodus longer than high, longer than carpus, cristate ventrally, cristate on proximal half dorsally. Movable finger lon- ger than palm, stout, with obtuse basal tooth. Fixed finger indistinctly toothed, fingers crossing when closed. Second abdominal somite as long as sixth. Latter inflated, twice as long as telson. Tel- son trapezoidal, much shorter than uro- VOLUME 101, NUMBER 4 Fig. 3. pods, lateral margins convergent posterior- ly, posterior margin with indistinct rounded median prominence. Size. — Males, cl 6.0 to 8.5 mm; non-ovig- erous females cl 3.8 and 4.1 mm; ovigerous females cl 6.1 to 9.8 mm. Largest male, tl 48 mm; largest female, ovigerous, tl 55 mm. Hult’s specimen, a male, is 37 mm long. Name.—The specific name is from the Greek, xouthos, yellowish brown, alluding to the color of the chelipeds reported in this species. Remarks. —This species is very similar to C. hartmeyeri in basic facies, agreeing with it and differing from all other species of Cor- allianassa in having a distinct distal spine on the ventral margin of the carpus of both first pereopods. The ventral border of the carpus is unarmed in C. borradalei, C. lon- giventris, and C. placida, the other species 887 Corallianassa xutha, new species (USNM 77861). a, Carapace, lateral view; b, Front, dorsal view; c, Third maxilliped; d, Major cheliped; e, Major chela, dorsal view; f, Merus and carpus of major cheliped, inner face; g, Minor cheliped; h, Abdomen; i, Sixth abdominal somite, telson, and uropods; j, Sixth abdominal somite, lateral view; k, Sixth abdominal somite and telson, lateral view; 1, Endopod of first pleopod. a—g, i, Female; h, j, k, Male. now assigned to Corallianassa. Coralli- anassa xutha differs from C. hartmeyeri in several features: the carpus of the major cheliped is more than half as long as the palm, and is distinctly serrate, almost spined, on its inner, ventral margin (these serrations are scarcely or not at all visible in external view); in C. hartmeyeri the carpus is no more than half the length of the palm, and the ventral serrations on the carpus are scarcely distinguishable. The movable finger of the major chela is armed with an obtuse and an acute tooth, separated by a gap; in C. hart- meyeri the two teeth are obtuse and are ad- jacent. The merus of the minor cheliped is twice as long as high and unarmed below; in C. hartmeyeri it is less than twice as long as high, armed below with a single denticle. Hult reported that the anterolateral spines of the carapace were not articulated in his 888 specimen; in all of the material reported here, including the specimen studied by Hult, these spines are distinctly separated from the anterior margin of the carapace by a non-calcified membrane. Little is known about the habitat of this species. Hult’s (1938) specimen from the Galapagos was taken in sand at low tide. The two specimens reported by Schmitt (1939) from Clipperton Island were taken while shore collecting. The material studied by Chace (1962) was taken on reef, coral cobble beach; coral reef; and on a reef flat, 0-1 foot at low tide. Hernandez Aguilera et al. (1986) reported it from under rocks of dead coral. Apparently most specimens tak- en so far were found in shallow water; the specimen from Maria Madre Island was taken in 7-18 meters. As is the case with C. Jongiventris, this appears to be a brightly colored species. Schmitt (1939:15) reported that “On the chelae of these specimens there were bright ochraceous to orange ochraceous (Ridgway) irregular color markings; both fingers to- ward their distal china-white tips were of this golden yellowish brown, the palm showed an irregular pattern of color and the upper surface of carpus and merus were like- wise suffused with it, the color being more concentrated and stronger on the merus and on the carpus.”’ Schmitt further commented that such a distinctive color pattern was un- usual in the callianassids, and Manning (1987:397) suggested that the bright color patterns might be characteristic of the genus Corallianassa. Distribution. —Eastern Pacific, from Ma- ria Madre Island, Mexico; Bahia Azufre [18°21'N, 114°44’W], Isla Clarion, Mexico (Hernandez Aguilera et al. 1986); Clipper- ton Island (Schmitt 1939, Chace 1962); Port Utria, Colombia; and Indefatigable Island, Galapagos Islands (Hult 1938). Discussion The genus Corallianassa Manning (1987: 393) was established for two species, the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON type species Corallianassa longiventris (A. Milne Edwards) from the Caribbean and C. borradalei (De Man) from the Indo-West Pacific. In the original account of Coralli- anassa, | noted that Callianassa placida De Man, 1905, differed from the two species placed in Corallianassa in that the second abdominal somite was not longer than the sixth. In the two species reported above, the second somite is subequal in length with the sixth, as is the case in C. placida (in contrast, in C. borradalei and C. longiventris the sec- ond somite is much longer than the sixth, being almost as long as the sixth and telson together). Inasmuch as C. hartmeyeri, C. xutha, and C. placida otherwise share sev- eral distinctive features with C. borradalei and C. /ongiventris, including the large, well- formed cornea, the anterior ventral projec- tion on the eye, and the anterolateral spines of the carapace which are separated from the carapace by a non-calcified membrane, all of these species are placed in Coralli- anassa. Other characters that may prove to be diagnostic for the genus are the low, round- ed median prominence on the posterior margin of the telson, and the inflated sixth abdominal somite. Members of the genus can be distin- guished by the following key: 1. Second abdominal somite longer than sixth, almost as long as sixth and telson combined ............ D, — Second abdominal somite subequal meleng che toysixt hese eee 3 2. Dorsal carina on palm of major che- liped extending over entire length of palm, ending in distinct flange ... Danii RAL Gieee SA C. borradalei (De Man) — Dorsal carina on palm short, not ex- tending to distal end, not ending in distinct flange ... C. longiventris (A. Milne Edwards) 3. Ventral margin of carpus of both chelipeds terminating in spine ... 4 — Ventral margin of carpus of both VOLUME 101, NUMBER 4 chelipeds unarmed distally ....... By rete ee eee ee ee C. placida (De Man) 4. Carpus of major cheliped more than half as long as palm, inner ventral margin distinctly serrate 4 ics REE A RE CR an be te Ee C. xutha, n. sp. — Carpus of major cheliped halfas long as palm, inner ventral margin with low, indistinct tubercles C. hartmeyeri (Schmitt) Acknowledgments I thank H.-E. Gruner, Zoological Mu- seum, Berlin, for the loan of the type of Callianassa hartmeyeri, Roy Olerod, Swed- ish Museum of Natural History, Stockholm, for the loan of the specimen of C. xutha from the Galapagos, and Richard Heard, Gulf Coast Research Laboratory, for read- ing a draft of the manuscript. William D. Lee, Smithsonian Marine Station at Link Port, Florida, collected the chelipeds of Cor- allianassa that led to this study. Lilly King Manning prepared the illustrations. This is contribution number 215 from the Smith- sonian Marine Station at Link Port. Literature Cited Balss, H. 1924. Westindische Decapoden.— Zoolo- gischer Anzeiger 61:177-182. 889 Biffar, T. A. 1971. The genus Callianassa (Crustacea, Decapoda, Thalassinidea) in south Florida, with keys to the western Atlantic species.— Bulletin of Marine Science 21(3):637-715. Chace, F. A., Jr. 1962. The non-brachyuran decapod crustaceans of Clipperton Island.— Proceedings of the United States National Museum 113(3466):605-635. Edmondson, C.H. 1944. Callianassidae of the central Pacific.—Occasional Papers of the Bernice P. Bishop Museum 18(2):35-61. Hernandez Aguilera, J. L., I. Lopez Salgado, & P. Sosa Hernandez. 1986. Crustaceos estomatopodos y decapodos de Isla Clarion. Fauna Carcinolo- gica Insular de Mexico, I. Investigaciones Oceanograficas/Biologia 3(1):183—250. Hult, J. 1938. Crustacea Decapoda from the Gala- pagos Islands collected by Mr. Rolf Blomberg. — Archiv for Zoologi 30A(5):1-18, pl. 1. Manning, R. B. 1987. Notes on western Atlantic Cal- lianassidae (Crustacea: Decapoda: Thalassinid- ea).— Proceedings of the Biological Society of Washington 100(2):386-401. Schmitt, W. L. 1935. Mud shrimps of the Atlantic coast of North America.—Smithsonian Miscel- laneous Collections 93(2):1—21, pls. 1-4. . 1939. Decapod and other Crustacea collected on the Presidential Cruise of 1938 (with intro- duction and station data).—Smithsonian Mis- cellaneous Collections 98(6):1—29, pls. 1-3. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 890-897 LABRAL MORPHOLOGY IN HEART URCHINS OF THE GENUS BRISSOPSTS (ECHINODERMATA: SPATANGOIDA), WITH AN ILLUSTRATED REVISED KEY TO WESTERN ATLANTIC SPECIES Richard L. Turner and Cathleen M. Norlund Abstract. —Existing keys to Brissopsis rely partly on the morphology of the labrum, an unpaired post-oral plate. As documented here, the unreliability of labral morphology has frequently resulted in misidentification of B. atlantica Mortensen, 1907 as B. elongata Mortensen, 1907. A revised key to western Atlantic species, using the number and shapes of plates below the periproct as substitutes for labral morphology, is provided. The current authority on systematics of the spatangoid genus Brissopsis in the At- lantic Ocean is Chesher’s (1968) revision based on analysis of 41 morphometric and meristic characters. His work includes a key that relies on only a few characters. The first couplet of this key separates B. elongata Mortensen, 1907 from other Brissopsis by the degree to which a post-oral plate, the labrum, extends posteriorly compared to adjoining plates. The basic format of Chesh- er’s key was incorporated by Serafy (1979) into his key to spatangoids of the Gulf of Mexico and vicinity. Labral extension is a convenient charac- ter to examine: a specimen need not be de- nuded of spines or dried, and damaged spec- imens often have the oral region intact. Brissopsis recently collected and examined by us included several specimens that were not B. elongata but had the labral condition of that species (Mortensen 1907). Subse- quent examination of hundreds of museum specimens revealed many that were mis- identified as B. elongata; nearly all of these had the labral condition of B. elongata. The purposes of this paper are to document the unreliability of labral extension as a taxo- nomic character in the genus and to provide for western Atlantic species a revised key that uses new characters of greater reliabil- ity. The Labrum The labrum is a tee-shaped, unpaired, in- terambulacral plate (plate 5.1, Lovén’s sys- tem) located just posterior to the peristome of spatangoids (Fig. 1). Its anterior margin forms the broad posterior lip of the peri- stome, giving the ventrum the appearance ofa carpenter’s plane. Projecting posteriorly from the lip, the narrowed stem of the la- brum abuts the first paired plates (sternal plates a.2 and b.2) of interambulacrum 5, which, along with the second pair (epister- nal plates 5.a.3 and 5.b.3), forms the broad plastron of the animal’s ventrum. The la- brum, plastron, and rest of interambula- crum 5 are bordered right and left by col- umns I.a and V.b, respectively, of the posterior paired ambulacra. The length of the labral stem varies in- terspecifically. In most Brissopsis it is short and does not reach the suture between plates 1 and 2 of the adjoining ambulacra (viz., I.a.1 and IJ.a.2 or V.b.1 and V.b.2) (Mor- tensen 1951, Chesher 1968, Serafy 1979). In B. elongata and B. obliqua Mortensen, 1948, the labrum typically extends beyond ambulacral plates I.a.1 and V.b.1 and abuts plates I.a.2 and V.b.2. [Mortensen’s (1907) statement that the labrum of B. pacifica (A. Agassiz, 1898) extends beyond the first am- bulacral plates is in error (Mortensen 1951).] VOLUME 101, NUMBER 4 Both Mortensen (1951:373, 377-378) and Chesher (1968:15) considered labral exten- sion to be a stable and useful character in the genus Brissopsis. Variability in labral extension has been known to occur in some species, but the specimens have been con- sidered abnormal. Mortensen (1907, 1951) found a few B. lyrifera (Forbes, 1841) in which the labrum extended beyond the first ambulacral plate on one (unilateral exten- sion) or both sides (bilateral extension) or at least reached the suture between the first and second plates (Fig. 2). Mortensen (1951) pointed out that the holotype of B. pacifica is “abnormal” in having an extended la- brum, and he included an appropriate cau- tionary note in his key (p. 379, footnote 3; see also p. 424 regarding a second speci- men). One specimen of B. columbaris A. Agassiz, 1898 was reported to have an ex- tended labrum (Mortensen 1951). Morten- sen (1951:416) examined few specimens of B. atlantica Mortensen, 1907 and did not report any cases of abnormal labra. Chesher (1968) reported 2 of 105 B. atlantica with labra extended unilaterally; he also de- scribed and illustrated a case of bilateral extension in an apparent hybrid from the Caribbean coast of Colombia, where B. at- lantica and B. elongata are sympatric. Variation in Brissopsis atlantica The problem of variability of labral ex- tension in Brissopsis atlantica first came to the attention of one of us (RLT) in 1983 while examining uncatalogued material at the Indian River Coastal Zone Museum (IRCZM), Harbor Branch Oceanographic Institution, Florida. The material, from the east coast of Florida, was tentatively la- belled B. elongata and consisted of 32 B. atlantica (IRCZM 72:346) with the labral conditions given in Table 1. Most recently, material collected specifically for this study in 1986 from the Gulf of Mexico included 24 B. atlantica, of which 19 had extended labra (6 lots; IRCZM 72:486-491). Thirty lots of B. elongata housed at the National 891 Fig. 1. Generalized ventral view of the test of a heart urchin, illustrating the alphanumeric designation of plates in the Lovénian system. Numbers of individ- ual plates are given for interambulacrum 4 and am- bulacrum V, and full designations are given for the sternal and episternal plates in column 5a. The sche- matic style follows David (1987). Abbreviations and conventions: ep, episternal plate; la, labrum; pa, pre- anal plate; ps, peristome; st, sternal! plate; light stipple, interambulacra; dense stipple, subanal fasciole; am- bulacra are not stippled. Museum of Natural History (USNM), Smithsonian Institution, have been exam- ined. The following misidentifications were found among them: (E10771), 2 B. atlantica with bilaterally extended labra; (E14283), 4 B. atlantica, 3 with bilaterally and 1 with unilaterally extended labra; (E14365), 1 B. atlantica with unilaterally extended labrum, redesignated E30961; 5 B. elongata remain in this lot; (E14373), 5 B. atlantica, 3 with bilaterally and 2 with unilaterally extended labra; (E14374), 2 B. atlantica with bilat- erally extended labra, redesignated E30962; 56 B. elongata remain in this lot; (E14377), 5 B. atlantica, 4 with bilaterally and 1 with unilaterally extended labra, redesignated E30960; 41 B. elongata remain in this lot; 892 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Examples of variation in extension and symmetry of the labrum (stippled). A, Labrum bilaterally extended between ambulacral plates I.a.2 and V.b.2. B, Labrum unilaterally extended to plate I.a.2. C, Labrum unilaterally extended to suture between I.a.1 and I.a.2. All B. atlantica, IRCZM 72:346. (E14389), 3 B. atlantica, 2 with bilaterally extended labra, 1 with unextended labrum; (E15552), 6 B. atlantica, 5 with bilaterally extended labra, 1 with unextended labrum; (E15560), 4 B. atlantica, 3 with bilaterally and 1 with unilaterally extended labra; (E15572), (E15573), and (E15574), each with 1 B. atlantica with a bilaterally ex- tended labrum; (E15587) and (E15588), each with 2 B. atlantica with bilaterally extended labra; (E15589), 5 B. atlantica, 2 with bi- laterally and 3 with unilaterally extended labra; (E15599), 1 B. atlantica with a bilat- erally extended labrum. The incidence of an extended labrum in some populations of B. atlantica is high, and misidentification of B. atlantica as B. elon- gata is frequent. It appears that the initial taxonomic assignment of USNM specimens was based only on the labral condition, for the course of the peripetalous fasciole over interambulacrum 3 and the ordinal number of ambulacral plates within the subanal fas- ciole in the specimens were typical of B. atlantica. Because existing keys separate B. elongata and B. obliqua from other Bris- sopsis spp. by labral morphology (Morten- sen 1951, Chesher 1968, Serafy 1979), the potential exists for continued misidentifi- cation. Pre-anal and Anal Plates Having eliminated labral extension as a useful character for Atlantic species of Bris- sopsis, we propose two replacement char- acters: the number of pre-anal plates and the shape of the first anal plates. The ar- rangement of the labrum, sternal plates, and episternal plates of interambulacrum 5 is described above. Following these plates in columns 5a and 5b is a variable number of pre-anal and anal plates (Fig. 3). Members of the pair(s) of pre-anal plates abut each other along their interradial margins, and none touches the periproct. Beyond the pre- anal plates, the two columns of interam- bulacral plates diverge around the peri- proct; where they border the periproct, members of these columns are called anal plates, and only members of the first and last pairs are partly in contact interradially. Each anal plate has by definition a margin along the periproct. In each column (a or b) of interambula- crum 5, Brissopsis alta Mortensen, 1907 has two pre-anal plates (5.4 and 5.5; Fig. 3C). Table 1.—Labral condition of 32 atypical Brissopsis atlantica in IRCZM 72:346. Symmetry of extension refers to extension on one (unilateral) or both sides (bilateral) of interambulacrum 5 (see Fig. 2). Degree of extension refers to the plate number in ambulacral col- umn I.a or V.b to which the stem of the labrum extends. Symmetry of extension Unilat- Degree of extension eral Bilateral Suture between ambulacrals 1 and 2 3 0 Beyond ambulacral plate 1 6 22 Mixed condition — 1 VOLUME 101, NUMBER 4 ie Fig. 3. Pre-anal and anal plates. A, B. atlantica, lectotype (USNM E10703, test length [TL] = 54 mm). B, B. elongata, (USNM 7117, TL = 30 mm). C, B. alta, lectotype (USNM E10704, TL = 67 mm). Out- 893 The first anal plate is, therefore, 5.6. All other western Atlantic Brissopsis spp. have one pre-anal plate (5.4), and plate 5.5 is the first anal plate. The shape of the first anal plate differs among species. Plate 5.5 of B. atlantica (Fig. 3A) is elongate and tapered, with a short periproctal margin and a much longer adradial margin, at which it abuts the adjoining ambulacral column. The in- terradial suture shared with the opposing member of the pair is long. The ratio of the adradial to periproctal margins is highly variable but always exceeds 2.4 (Fig. 4). In B. elongata (Fig. 3B), the periproctal margin is long, and the ratio is less than 2.4 (Fig. 4). The interradial margin shared with the opposing member of the pair is short. It is easy to distinguish B. atlantica and B. elon- gata by the shape of plate 5.5. The adradial- periproctal margin ratio for the lectotype of B. atlantica (USNM E10703) is 5.8. A pho- tograph of a type specimen of B. elongata (Mortensen 1907; pl. IV, fig. 18) gives a ratio of 1.4. Additionally, the ratio is 1.7 for a specimen of B. elongata (USNM 7117, ALBATROSS sta 2145) mentioned by Mor- tensen (1907:163, 426) to be conspecific with his type material, which we have not ex- amined. The first anal plate of B. mediter- ranea Mortensen, 1913 is of variable shape in the few nominal western Atlantic speci- mens available to us. The number and shapes of the pre-anal and first anal plates are partly correlated with overall body form in Atlantic Brissop- sis (Fig. 3). In contrast to the low profile of B. atlantica and B. elongata, the presence of a second pair of pre-anal plates gives the gas lined profiles give general shape of test, location of subanal fasciole, position of periproct (arrows), and angle of posterior margin of test for each species. Ab- breviations and conventions: aa, adapical suture; an, first anal plate; ao, adoral suture; ar, adradial suture; in, interradial suture; pa, pre-anal plate; pm, periproc- tal margin; light stipple, interambulacrum 5; dense stipple, subanal fasciole; periproct and ambulacra I and V are not stippled. 894 Ratio PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Test length [mm] Fig. 4. Ratio of lengths of the adradial suture and periproctal margin of the first anal plates (5.a,b.5) of 78 B. atlantica and 94 B. elongata. Conventions: open circle, B. atlantica; closed circle, B. atlantica, lectotype; open square, B. elongata; closed square, B. elongata (USNM 7117); broken line, B. elongata from Mortensen (1907, pl. IV, fig. 18, test length unknown); overlapping plots not indicated. posterior of B. alta a high profile. The in- clined posterior of B. atlantica results from the tapered adapical extension and length- ened interradial suture of the first anal plates. On the other hand, the vertical posterior of B. alta is produced by the squareness of the first anal plates; and that of B. elongata by the strap-like first anal plates with a short- ened interradial suture. Judging from illus- trations of other species of Brissopsis (Mor- tensen 1951), the number and shapes of these plates should be examined further for their taxonomic utility. Key to Western Atlantic Species of Brissopsis The key presented below is a revision of keys given in Chesher (1968) and Serafy VOLUME 101, NUMBER 4 895 1.a.6[7] F G Fig. 5. Diagrams of Brissopsis showing features used in key. A, Divergent posterior paired petals; adapical primary tubercle of each column in interambulacrum 5 is indicated. B, Confluent posterior paired petals; adapical primary tubercles as in A. C, Peripetalous fasciole without re-entrant angle on plate 3.a.4. D, E, Peripetalous fasciole with re-entrant angle on plate 3.a.4. F, Subanal fasciole crossing 4 plates of ambulacrum I. G, Subanal fasciole crossing 5 plates of ambulacrum I. H, Lip of labrum straight. I, Lip of labrum produced. Abbreviations and conventions: la, labrum; light stipple, interambulacra; dense stipple, fascioles; ambulacra, apical system, and peristome are not stippled. 896 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (1979). It resolves an apparently recurring problem of distinguishing B. atlantica and B. elongata by eliminating reference to la- bral morphology. Neither we nor Chesher had the opportunity to examine type ma- terial of B. mediterranea, for which pub- lished illustrations of interambulacrum 5 do not exist; our inclusion of this species in the key is tentative. Reference to globiferous pedicellariae is omitted because their scar- city on museum material [as noted also by Mortensen (1951:378)] makes their use im- practical. Brissopsis elongata with TL < 20 mm often do not have features described below and cannot be identified by using the key; e.g., the cardinal and ordinal numbers of plates crossed by the subanal fasciole in small specimens are not consistent, indi- cating a prolonged period of allometric growth in this species (McNamara 1987). Finally, we caution that most larger B. elon- gata(TL > 20 mm) we have examined from the western Atlantic have the peripetalous fasciole crossing ambulacrum III on plate 7 [contrary to Chesher (1968)], but smaller specimens (TL < 20 mm) often have it crossing III.6. Therefore, this character is omitted from the key. Characters used in the key but not illustrated in Figs. 1-4 are depicted in Fig. 5. Examination of speci- mens is aided by brushing spines from in- terambulacra 3 and 5 and from ambulacra I and V; sutures between plates can be made visible by streaking with a cotton swab moistened with xylene. 1. Posterior paired petals divergent, first primary tubercles of interam- bulacral columns 5.a, 5.b occurring 1-2 plates behind apical system (Fig. 5A); peripetalous fasciole without re-entrant angle in interambulacral column 3.a, crossing plates 3.a.4, 3.b.4 (Fig. 5C); 2 pairs of pre-anal plates (Fig. 3C); first anal plates are 5.a.6, 5.b.6; subanal fasciole cross- ing 4 ambulacral plates on each side (I.a.6-9, V.b.6-9) (Fig. 5F)....... A ee eee Cae B. alta Mortensen, 1907 Posterior paired petals confluent, first primary tubercles occurring 3— 7 plates behind apical system (Fig. 5B); peripetalous fasciole with re- entrant angle in interambulacral column 3.a, crossing at least plates 3.a.4, 3.a.5 (Fig. 5D, E); 1 pair of pre-anal plates (Fig. 3A, B); first anal plates are 5.a.5, 5.b.5; subanal fas- ciole crossing 4 or 5 ambulacral plates (Figs 5G) eee ae eee . Subanal fasciole crossing 4 ambu- lacral plates on each side (I.a.6—-9, V.b.6—9) (Fig. SF), and peripetalous fasciole in interambulacral column 3.b crossing only plate 5 (Fig. 5D) B. mediterranea Mortensen, 1913 Subanal fasciole usually crossing 5 ambulacral plates on each side (Fig. 5G); path of peripetalous fasciole in 3.b various: 20240 See eee . First ambulacral plates to enter sub- anal fasciole are I.a.7, V.b.7 (Fig. 5G); adradial suture of first anal plate less than 2.4 times the length of peri- proctal margin (Fig. 4), interradial suture short, adapical and adoral su- tures parallel, plate strap-like (Fig. 3B); lip of labrum straight (Fig. 5H); peripetalous fasciole crossing only plate 5 in interambulacral column 3:bi(Fig: SD) 0.2" 5. See eo eee Rtas B. elongata Mortensen, 1907 First ambulacral plates to enter sub- anal fasciole are IJ.a.6, V.b.6 (Fig. 5G); adradial suture of first anal plate more than 2.4 times the length of periproctal margin (Fig. 4), inter- radial suture long, adapical and adoral sutures distinctly not paral- lel, plate adapically elongate and ta- pered (Fig. 3A); lip of labrum pro- duced (Fig. 51); peripetalous fasciole crossing plates 3.b.4, 3.b.5 (Fig. SE) Tok Sit B. atlantica Mortensen, 1907 VOLUME 101, NUMBER 4 Acknowledgments The problem that formed the basis of this paper was recognized while working on echinoids collected during the 1983 cruise of the SEAMAP program of the Atlantic States Marine Fisheries Commission. For their invitation to participate and their lo- gistic support on the 1983-1985 cruises, we thank Elmer J. Gutherz and Gilmore Pel- legrin, National Marine Fisheries Service, Pascagoula, who served as field party chiefs, and William G. Lyons, Florida Department of Natural Resources, who served as chief scientist on the Florida leg (Florida East Coast Benthic Mapping Study). Gilmore Pellegrin provided additional specimens from an unrelated cruise. Loans of material and hospitality during visits were kindly provided by David L. Pawson, USNM, John E. Miller, IRCZM, and Robert M. Wool- lacott, Museum of Comparative Zoology, Harvard University. We thank D. Keith Serafy, Southampton College, for helpful discussions. John Miller offered construc- tive criticism of the manuscript. A Short- Term Visitor Award for travel funds from the Office of Fellowships and Grants, Smithsonian Institution, was provided through David Pawson for one of us (RLT) to examine USNM collections. During part of this study, RLT was a Harbor Branch 897 Institution Postdoctoral Fellow while on sabbatical leave from Florida Institute of Technology. HBOI contribution number 629. Literature Cited Chesher, R. H. 1968. The systematics of sympatric species in West Indian spatangoids: A revision of the genera Brissopsis, Plethotaenia, Paleo- pneustes, and Saviniaster. —Studies in Tropical Oceanography 7:1—168. David, B. 1987. Dynamics of plate growth in the deep-sea echinoid Pourtalesia miranda Agassiz: A new architectural interpretation. — Bulletin of Marine Science 40:29-47. McNamara, K.J. 1987. Plate translocation in spatan- goid echinoids: Its morphological, functional and phylogenetic significance. — Paleobiology 13: 312-325. Mortensen, T. 1907. Echinoidea (part 2).—The Dan- ish Ingolf-Expedition 4(2):1—200. Mortensen, T. 1951. A monograph of the Echinoidea, 5(2). Spatangoida, 2. Copenhagen, C. A. Reitzel, 593 pp. Serafy, D. K. 1979. Echinoids (Echinodermata: Ech- inoidea).—Memoirs of the Hourglass Cruises 5(3):1-120. Department of Biological Sciences, Flor- ida Institute of Technology, 150 West Uni- versity Boulevard, Melbourne, Florida 32901-6988; (CMN) Present address, De- partment of Zoology, University of Florida, Gainesville, Florida 32611. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 898-911 PATTERNS OF GEOGRAPHIC VARIATION IN THE ARABIAN WARBLER SYLVIA LEUCOMELAENA (AVES: SYLVUDAE) Steven M. Goodman Abstract. —The distribution and relationships of Sy/via leucomelaena, a species generally found in Acacia parkland, are reviewed. Analysis of seven morpho- logical and two plumage characters results in the recognition of three subspecies: S. 1. leucomelaena (Hemprich & Ehrenberg) from western Saudi Arabia, the Yemens, and the Dhofar region of Oman; S. /. somaliensis (Sclater & Mack- worth-Praed) from Somalia and Eritrea; and S. /. blanfordi Seebohm from ~ southeastern Egypt and eastern Sudan. Birds inhabiting the Arava of southern Israel are closest to nominate /eucomelaena but differ in several characters. A phytogeographical analysis is presented of the origin, dispersal, and modern distribution of this habitat, and how it relates to the present patterns of geo- graphic variation in S. /eucomelaena. Of the 15 species in the Palearctic genus Sylvia Scopoli, the Arabian or Blanford’s Warbler, S. /eucomelaena, is the only one breeding south of the Sahara (Watson et al. 1986). Its nesting range includes portions of southern Israel, the Arabian Peninsula, and the African Red Sea mountains and coastal plain, generally in relatively dense Acacia parkland. This habitat consists of widely dispersed clusters of trees, often in remote and relatively undisturbed country. Except for a recent study in southern Israel virtually nothing is known about the Arabian War- bler’s natural history and ecology. To date, no published study has examined patterns of geographic variation in this species. In the course of a 1985 biological survey of Gebel Elba in the Red Sea Mountains of southern Egypt, just north of the Sudanese border, a population of S. leucomelaena was discovered and a small series collected. Be- fore a subspecific designation could be placed on this material it was necessary to review patterns of geographic variation in this species. The purpose of this paper is to de- scribe these patterns and to present a phy- togeographical analysis that may account for them. Information is also given on distri- bution, sexual dimorphism, several aspects of natural history, and taxonomic conclu- sions. Materials and Methods Seven measurements were taken from museum specimens (all in mm): Wing— measured from the bend of flattened wing at carpal joint to longest primary tip. Tail- measured between the insertion of central tail rectrices to distal tip. Exposed culmen— measured from base of feathering on the forehead to maxilla tip. Bill from nostril— measured from anterior edge of nostril to maxilla tip. Bill width—measured across upper mandible at anterior edge of nostril. Gonys— measured from junction of rami to maxilla tip. Tarsus—measured from the junction of tibiotarsal-tarsometatarsal joint to distal edge of distal most undivided scute overlying the toes (little variation was found in the scale pattern of the feet). A rule was used to the nearest 1.0 mm for the wing and tail measurements and a dial calipers to the nearest 0.1 mm for the other measurements. VOLUME 101, NUMBER 4 Since virtually no information is available in the literature on the weight or food habits of S. Jeucomelaena this information, based on field and museum studies, is summa- rized herein. Weights of the Egyptian spec- imens were taken to the nearest 0.1 gm. All other weight data are from museum speci- mens and the unpublished field notes of Mr. G. Nikolaus or Mr. M. C. Jennings. All ref- erences to altitude are presumed to be m above sea-level. Four specimens were selected as stan- dards for comparison of crown-color. These are (darkest to lightest): BMNH 1925.11.20.26, male, Sogsode, Somalia; BMNH 1919.12.17.702, male, Erkowit, Su- dan; BMNH 1915.12.24.720, female, Er- kowit, Sudan; and BMNH 1919.12.17.686, female, Sinkat, Sudan. The contrast be- tween the crown and back did not enter into the selection of these standards. Four stan- dards were used to quantify the amount of white on the outer pair of rectrices (Fig. 1). Individual specimens were compared to these standards and scored accordingly. Statistical tests used MIDAS programs, written by the University of Michigan Sta- tistical Research Laboratory. Differences among means and variances were tested by Student’s f-test or analysis of variance (AN- OVA), as appropriate. Scheffe tests were also used in pairwise combinations to test dif- ferences in means between geographical re- gions. Probabilities of 0.05 or less are con- sidered statistically significant and sufficient to reject the null hypothesis that the means are equal. Review of Sylvia leucomelaena Taxonomy The original description of S. leucome- laena by Hemprich & Ehrenberg (1833) was based on material taken in Arabia. They placed the bird in genus Curruca. In a re- view of Hemprich and Ehrenberg speci- mens in the Berlin Museum, Dresser & Blanford (1874) described the type of S. /eu- comelaena, gave measurements and noted 899 TT Fig. 1. Four standards used to quantify the amount of white on the outer pair of rectrices. this species resemblance in details to Sylvia rueppelli. In 1878 Seebohm described Sy/- via blanfordi based on a single specimen tak- en at Rairo, Habab, Abyssinia (=Eritrea). Although he cited the paper of Dresser and Blanford, Seebohm believed that the Rairo specimen represented a new species, with some similarity to S. curruca, S. melano- cephala and S. rubescens (=S. mystacea). In a review of birds collected in the Ye- mens, Hartert (1917) named a subspecies, Parisoma blanfordi distincta. His placement of this form in Parisoma was unexplained but may have been due to the notion of his time that the Palearctic and the Afrotropical (=Ethiopian) biogeographic realms were completely distinct. It would thus have been difficult to explain a Sy/via breeding in sub- Saharan Africa. Sclater & Mackworth-Praed (1918) de- scribed the new form somaliensis from ma- terial collected at Mundara, Somalia, and retained this species in Parisoma. Their di- agnosis noted that somaliensis was distin- guishable from blanfordi and distincta by its smaller size and differences in back color- ation and tail pattern. Meinertzhagen (1949) reviewed this situation and considered Cur- ruca leucomelaena and Sylvia blanfordi conspecific. Further, he presented evidence that the species should be shifted from Par- isoma to Sylvia. Afik & Pinshow (1984) questioned the al- location of this species to Sy/via, primarily on several aspects of natural history, and suggested a reexamination of its generic po- sition. As pointed out by Dowsett-Lemaire 900 & Dowsett (1985), however, little compar- ative natural history information is cur- rently available on most Afrotropical Syl- viidae. Furthermore, the genus Sy/via shows a remarkable degree of plasticity in aspects of life history, and such characters probably provide little information on the natural- ness of the group. General Distribution of Sylvia leucomelaena by Country ISRAEL—In April 1972 this species was found nesting in the Arava (Rift Valley), 10 km south of the southern end of the Dead Sea (Zahavi and Dupai 1974) (Fig. 2). In more recent years it has been observed in relatively dense Acacia tortilis and A. rad- diana stands from the southern end of the Dead Sea south to within 40 km of Eilat (Afik & Pinshow 1984; H. Shirihai, pers. comm.). SAUDI ARABIA — The Arabian Warbler is a breeding resident of southwestern and western Saudi Arabia, often in areas with thick vegetation such as along the Tihama (Stagg 1985). Hutchinson (1975) reported several observations from central Saudi Arabia near Riyadh; these records were re- jected by Jennings (1981). Meinertzhagen (1954) designated the type locality of Cur- ruca leucomelaena Hemprich & Ehrenberg as Midian (perhaps Maidn Salah, 26°45'N, 37°55'E; Jennings 1981), a northern area of the country for which there are no modern records (M. C. Jennings, pers. comm.). THE YEMENS— During a trip to North Yemen from 7-29 Apr 1979 Cornwallis & Porter (1982) recorded this species on nine occasions in the Western Ramparts and Highland Plateau between 500 and 1700 m, invariably in Acacia groves. This species is a fairly common but local resident of the Tihama foothills, particularly in Acacia/ Commiphora bushland (Brooks et al. 1987). SULTANATE OF OMAN-—The Arabian Warbler is known from Dhofar, in the southwestern corner of the Sultanate, where PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON it tends to occur “on hillsides and in moun- tain wadis”’ (Gallagher & Woodcock 1980: 252). In this region it has been recorded in every month of the year and is considered a breeding resident (Gallagher 1986). The Oman distribution of the Arabian Warbler is exceptional. In some regions it inhabits areas of mixed woodland, without extensive tracts of Acacia spp. (M. D. Gallagher, pers. comm.). SOMALIA TO ERITREA —Archer and Godman (1961) noted that this species oc- curs in Somalia along the coastal plain.and plateau between sea-level and up to about 1800 m, and as far south as Galoli and Bu- rao. More recently it has been presumed to be a “fairly common” resident in the north- ern portion of Somalia above 8°30’N in aca- cias along the coastal plain and up to the plateau beyond the coastal mountains (Ash & Miskell 1983:59). This species has been noted throughout portions of Djibouti, par- ticularly in Acacia scrubland (G. Welch, pers. comm.). Smith (1957) noted that the Ara- bian Warbler was found in the Red Sea coastal hills of Eritrea, at least between Mas- sawa and Dankalia, generally in deep wadis below 320 m and with acacia cover. SUDAN—Cave & Macdonald (1955:255) listed this species as a “rather uncommon resident of the old Red Sea Province.” EGYPT — In early 1985 this species was dis- covered in the immediate vicinity of Gebel Elba (22°11'N, 36°21'E), Sudan Govern- ment Administration Area (Goodman & Meininger in press a). Although no nests were found, numerous singing males were observed in the dense Acacia groves and clear territorial interactions noted. Five adult specimens were collected, all of which had enlarged gonads. This species is presumed to be a local breeding resident in the Gebel Elba area. There are no specimen records from elsewhere in the country (Goodman and Meininger in press b); although it is plausible it occurs in extreme eastern Sinai just across the border from known Israeli populations. VOLUME 101, NUMBER 4 I eat sf 5 \ af \ Boa \ e-Jordan 5 if 2i0) 901 oh «> 25° o fo mo Riyadh 2 e Muscat Jebel Akbdar e Saudi Arabia 20° Erba Mountains/o\ . ! IN > ° og taniak 15° ! A srenesiove = Ethiopia Djibdutil) J 100 200 300 Kilometers La, Djibouti 45° 50° 55° 'Sultanate I j of Oman 0) 20° Fig. 2. Distributional map of Sylvia leucomelaena across its complete range. Open triangles represent spec- imens examined and open squares sight records from the literature or unpublished information. Notes on the Recently Discovered Egyptian Population The Egyptian population of S. /eucome- laena appears to be limited to the slopes and valleys of Gebel Elba in the extreme southeastern corner of the country. This mountain, which rises to 1428 m, is unique for Egypt in that a significant portion of the plants and animals are of Afrotropical rath- er than Palearctic origin. The Gebel Elba area has relatively heavy vegetation and contains dense Acacia parkland (Kassas & Zahran 1971). Acacia tortilis is common along the coastal plain, foothills and lower slopes of Gebel Elba, and 4. raddiana on the north slopes and in valleys bisecting the southern slope (Kassas & Zahran 1971). The Arabian Warbler was observed at various sites around the mountain but was more common on the northern side, where the Acacia parkland is most extensive. Gen- erally they were observed singly or in pairs. The first territorial aggression was noted on 13 Apr 1985, although songs were heard soon after our arrival in February. Inter- specific encounters were recorded on several occasions. On 13 April one adult S. /euco- melaena was noted flying in a mixed flock of three Cercomela melanura and six Sylvia melanocephala. Later that same day one adult S. leucomelaena was observed chasing 902 a male S. melanocephala. On 14 April four adult S. leucomelaena were noted foraging in a mixed flock with S. curruca and Cer- comela melanura. In the Gebel Elba area Cercomela is a local breeding resident and Sylvia curruca and melanocephala migrants and winter visitors. Results Sexual dimorphism in measurements. — No single geographically constant pattern of sexual dimorphism was found in this species (Table 1). Morphological differences were not detected between the sexes in the Arava or in the combined Somalian and Eritrean samples. Only a single female was available from the Gebel Elba, Egypt, population and no statistical comparison was made; how- ever, for all seven measurements the value of the female fell within the range of males from this locality. In both the Saudi Arabian and Yemen samples, males have statisti- cally longer wings and tails than females. Sudanese males have longer wings than fe- males. Sexual dimorphism in plumage color- ation within geographic regions.—In the Saudi Arabian, Somalian/Eritrean and Su- danese samples, adult males have darker crowns than adult females (Table 2); the contrast was most notable for Sudanese birds. These sexual differences are statisti- cally significant. No sexual variation was found in this character for the Yemen, Ar- ava or Egyptian populations, although in the latter two cases the sample size was lim- ited. Within the various geographical re- gions sexual difference was found in tail pat- tern only in the Saudi Arabian sample (Table 3), in which males tend to have more white on the outer rectrices than females (P = 0.018). Geographic variation in plumage color- ation.—No statistically significant differ- ence between regions was found in the head coloration of adults, whether partitioned by sex or lumped together. For the tail pattern PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the situation is more complex. Since no sex- ual variation was found in this character for any population inhabiting the western side of the Red Sea, the sexes were combined in this analysis. In this region there is clinal variation in the tail pattern. Adult birds from Egypt have less white on the outer rectrices (n = 5, mean = 2.0) than Sudanese (n = 21, mean = 2.9, P = 0.0001), and Sudanese birds have less white than Somalian/Eri- trean (n = 27, mean = 3.9, P = 0.0001). The pattern on the eastern side of the Red Sea is similar with an increasing amount of white on the tail from north to south, but the range of variation is not as great, per- haps in part due to small samples from some areas. Scores were only available from the Arava population for adult females. These birds have less white on the outer rectrices (n = 2, mean = 2.0) than Saudi Arabian females (n = 10, mean = 2.2) and Yemeni females (n = 7, mean = 2.4). None of these comparisons showed statistical significance. Males from Saudi Arabia had slightly more white (n = 15, mean = 2.7) than those from Yemen (n = 8, mean = 2.6). The single male from Oman has a tail score of 2. No differ- ence was found between the Arava and Egyptian populations. Geographic variation in morphology. — Since the wing and tail measurements of adult males and females are dimorphic in two populations they were partitioned with- in each sample (Table 4). No differences were found between birds from Saudi Ara- bia and Yemen, and these populations are combined in subsequent analyses. The wing lengths of both male and female populations living on the west side of the Red Sea are similar to one another; the only exception is that males from the Sudan have slightly longer wings than males from the Somalia/ Eritrea sample (F = 5.98, P = 0.017). There is considerable variation in tail length among the five geographic areas. Within either sex class no significant vari- ation was found between the Egyptian and Sudanese samples; however, when these VOLUME 101, NUMBER 4 samples were compared to birds from So- malia/Eritrea differences were found in both cases (Table 4). Males from Arava have longer tails than those from the Arabian Peninsula (F = 4.05, P = 0.048), while fe- males are similar to one another. Within each sex class there is a cline in tail length on both sides of the Red Sea, decreasing in size from north to south. Sylvia leucomelaena from Arava have substantially wider bills than the other four samples (Table 5). There appears to be clin- al variation in bill width on the western side of the Red Sea, decreasing in size from north to south. Analysis of the exposed culmen measure- ment showed a unique pattern of variation (Table 5). Significant differences were found in the Sudanese birds relative to those from the Arabian Peninsula (F = 21.60, P = 0.0000), Somalia/Eritrea (F = 33.54, P = 0.0000), and Egypt (F = 15.15, P= 0.0002). This variation shows no clear clinal pattern. The bill length from nostril also revealed an unusual pattern of variation (Table 5); the statistically significant differences were be- tween the Arabian Peninsula and Somalia/ Eritrea samples (F = 7.97, P = 0.005), the Arabian Peninsula and Sudan samples (F = 4.28, P = 0.04), and Somalia/Eritrea and Sudan samples (F = 19.28, P = 0.0000). The gonys length of Sudanese birds was the smallest of any of the samples and sta- tistically different from all others, except the Egyptian birds (Table 5). A comparison of F-statistics for tarsus length was not in- cluded in Table 5. The only statistically sig- nificant difference was between the Soma- lia/Eritrea and Egyptian samples (F = 5.50, P = 0.03), which most likely reflects two extremes of a cline. Weight. —Weights from three adults han- dled by Mr. Michael C. Jennings at Hejaz, Saudi Arabia, in mid-May 1976 are 14.8, 15.3, and 16.3 g. For 29 unsexed autumn birds weighed by Mr. Gerhardt Nikolaus during ringing operations near Erkowit, Sudan, the mean weight was 13.9 g and 903 range 12.5-16 g, excluding one bird which weighed 9 g. Four weights are available for wild birds from the Arava population, two adult males at 15 and 13.5 g, and two adult females at 16 (with “full grown egg”) and 15 g (UTM). Weights of Egyptian birds in- clude: five males—mean 12.3 g and range 11.2-13.2 g, and one female—12.5 g (UMMZ). Food. —Meinertzhagen (1954) consid- ered the Arabian Warbler an insectivore. Afik & Pinshow (1984) reported that it searched acacia bark for larvae, hawked flying insects, and dug for small insects on the ground under acacia. They further noted that this species frequently eats ripe fruits of three perennial shrubs: Nitraria retusa, Lycium shawii and Ochradenus baccutus. Of the five specimens taken at Gebel Elba, Egypt (UMMZ), three had empty stomachs and two contained unidentified fruits. Two birds collected at Sinkat, Sudan (BMNH), had seeds in their stomachs and no insects, while a third contained berries. One bird from Somalia (LSUM) had Coleoptera re- mains in the gizzard. Discussion The East African Rift Valley, the Red Sea, Gulf of Aqaba, and the Arava are portions of a single tectonic system, often referred to as the Afro-Arabian Rift Valley. The shear- ing of the Red Sea apparently commenced in the early Miocene (Ross & Schlee 1973), as the Arabian plate drifted away from the African plate. This rifting activity contin- ued with varying intensity through the up- per-most Miocene or lowest Pliocene, at which time the southern Red Sea opened into the Indian Ocean (Hotzl 1984). Pres- ently, the Red Sea is a little under 2000 km long. The southern strait at Djibouti, Bab el Mandab, is about 30 km wide. In the north the average width varies from 150 to 180 km and between 15° and 18°N the sea widens to 350 km. The Gulf of Aqaba is about 170 kms long and varies between 15 and 30 km in width. 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Both of these trees often occur $ mc aS 5 sympatrically and their local distribution is a 2 adjusted to the local micro-climate. A. tor- =| sete G S tilis is adapted to higher temperatures and 2 Noa B dryer conditions than raddiana; while rad- 3 foe) 5 . » ale aoAS = diana is more tolerant of a wider range of g al 9 temperature and moisture (Halevy and Or- . 13 5 shan 1972). The exception to this apparent 3 |? 2 Q 2 habitat requirement is in Oman, where the 3 2 al S bird occurs in mixed open woodland areas 8 2 of the Dhofar region. ) 5 ks Zohary (1962) hypothesized that before ol SI = 5 syle . . 2 ERs otis s the Miocene 4A. tortilis and raddiana mi- iS Z| 2 5 E Ala grated from Africa along the eastern coast 3 f= n = S| of the Red Sea across to and north along 906 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.— Percentages of adult Sylvia leucomelaena within a geographic region scored for different tail pat- tern standards.! Standard Region 1 2 3 4 Mean Arava Female (n = 2) 0.0 Saudi Arabia* 100.0 0.0 0.0 2.0 Male (n = 15) 0.0 26.7 73.3 0.0 2.7 Female (n= 10) 0.0 80.00 20.0 0.0 2.2 Yemen Male (n = 8) 0.0 37.5 62.5 0.0 2.6 Female (n = 7) 0.0 57.1 42.9 0.0 2.4 Somalia and Eritrea Male (n = 19) 0.0 0.0 5.9 94.1 3.9 Female (n = 8) 0.0 OW MS 875 39 Sudan Male (n = 14) 0.0 0.0 100.0 0.0 3.0 Female (n = 7) 00 43) 85.7 O0 2 Egypt Male (n = 4) 0.0 100.0 0.0 OM 20 Female (n = 1) 0.0 100.0 0.0 0.0 2.0 ' Within a geographical region sexual differences be- tween the means of the two sexes were tested with Student’s ¢-test, one asterisk P < 0.05. western Arabia, to the Gulf of Aqaba, into the Arava and other portions of the Levant. Given our present knowledge of plate tec- tonics of this area, these two land masses would have been at least partially connected at that time. Shmida & Or (1986) presented a strong case that the invasion of these plants into the Arava, although along the same route suggested by Zohary, has been since the end of the Pleistocene, long after the shearing took place. Their argument is pri- marily based on the low degree of local endemism in these Afrotropical elements, their wide distributional disjunctions, and adaptations to arid conditions and long-dis- tance dispersal. In either case, the modern range of Acacia spp. in the mountains and along the coastal plains bordering the Red Sea is evidence to support this dispersal route, whether it took place before or after rifting, and most importantly how this dis- tribution is relevant to that of Sylvia leu- comelaena. Several species of Acacia are common along the western Red Sea coastal plain from the East African Rift Valley (Beals 1970), north through Eritrea (Hemming 1961), to the Red Sea Mountains of Sudan (Kassas 1956, 1957). Just north of the Egyptian/Su- danese political border, near Gebel Elba, nine Acacia spp. are known to occur; these are dominated by fortilis and raddiana (Kassas & Zahran 1971, Tackholm & Bou- los 1972, Tackholm 1974). North of Gebel Elba, in the Egyptian Eastern Desert, there Table 4.—Comparison of F-statistics by geographic region for two sexually dimorphic characters in adult Sylvia leuacomelaena, wing length (upper right corner) and tail length (lower left corner).! Region 1 2 2 4.05* 0.42 3 ANS) Jaa HOSS oe 1337 One 22.41*** 4 NOAA Sle? Oia 4.44* 5.24* 5 13.88*** O25 0.39 0.09 WG BAA? 16.16*** 18.14*** 10.64** 14.78*** 7.46** 68.02*** 29825 IGT 102757* LWESSETe Sy 59S 0.03 0.62 0.79 23.00*** 1.56 4.69* 0.27 ORS ae 2.84 ' For each matrix the numbers on top within a geographic region comparison are males and below females. 1 = Arava, 2 = Arabian Peninsula (combined Saudi Arabia, Yemen, and Oman), 3 = Somalia and Eritrea; 4 = Sudan, 5 = Egypt. One asterisk P < 0.05, two asterisks P < 0.01, three asterisks P < 0.001. VOLUME 101, NUMBER 4 907 Table 5.— Matrices of F-statistics by geographic region for non-sexually dimorphic characters in adult Sylvia leucomelaena.' ; Bill width/exposed culmen Bill from nostril/gonys Geographic area 1 2 3 4 1 2 3 4 5 1 0.75 2.91 2.12 3.47 0.85 3.99* 1.60 2 SAO 4S ASS DNS 2.64 0.69 1.91 IQ) syil-*- 1.67 3 elt OO 33), 545** 0.52 OHO) Sih: NS NGA BTS 4 27.40*** 2.01 1.13 Salles PEAS GPRS OD see 0.12 5 13.64** 1.31 0.98 0.05 1.38 ' Character to left of slash is on bottom half of matrix, character to right of slash is on top half of matrix. 1 = Arava, 2 = Arabian Peninsula (combined Saudi Arabia, Yemen, and Oman), 3 = Somalia and Eritrea, 4 = Sudan, 5 = Egypt. One asterisk P < 0.05, two asterisks P < 0.01, three asterisks P < 0.001. are only small patches of Acacia raddiana and most of the other species fall out (Kas- sas & Imam 1959; Kassas & Zahran 1962, 1965; Boulos & Hobbs 1986). This is com- pared to the eastern side of the Red Sea, where relatively dense Acacia parkland oc- curs along the Arabian Peninsula coastal plain and above the Tihama in the Asir and Hejaz Mountains (M. C. Jennings, pers. comm.); forming a more or less continuous band from East Africa across to and along the eastern edge of the Arabian Peninsula (Vesey-Fitzgerald 1955, 1957). Based on the present distribution of Acacia along the western side of the Red Sea (including the Gulf of Suez), it appears that contiguous tracts have not existed in this region in the recent geological past; precluding the pos- sibility that these trees migrated through this pathway into the Levant. Clearly, this is based on the assumption that the modern distribution of Acacia in this area parallels that since the end of the Pleistocene, and that changes in climatic patterns and human land-use have not resulted in the extinction of areas of Acacia parkland. The main point of this phytogeographical analysis is that evidence exists that Acacia spp. spread from East Africa into the Levant via the Arabian Peninsula, and that a habitat corridor ex- isted along this pathway for dispersal from the northern Afro-Arabian Rift Valley into East Africa. Seventeen species of Sy/via are known to occur in Africa (Vaurie 1959, Etchécopar & Hue 1967, Moreau 1972, Watson et al. 1986). Of these, eleven have breeding pop- ulations on the continent. (This figure in- cludes S. sarda, but see Thomsen & Jacob- sen 1979.) For the most part the breeding ranges of the other ten species are restricted to the Mediterranean Sea coastal zone, in areas receiving about 100 mm of precipi- tation per year and basically with a Pale- arctic flora; a few species also occur south into coastal Mauritania. The exceptions are S. nana desertiand S. deserticola which pen- etrate into portions of the western Sahara; the latter species is endemic, as 1s the sub- species S. melanocephala norrisae known from the Faiyum of Egypt. Excluding /eu- comelaena, none of these Sy/via occur in areas with Afrotropical biota and the hab- itat of their African breeding range is ba- sically an extension of what they use in Eur- asia. Several workers have speculated that hor- tensis and leucomelaena form a superspe- cies, are part of the same subgenus, or each others closest relatives (Hall & Moreau 1970, Wolters 1980, Watson et al. 1986). Four subspecies of hortensis are currently recog- nized (Vaurie 1959, Watson et al. 1986), three of which are important for this review: nominate hortensis breeds in portions of central Europe and North Africa from Mo- rocco to Tripolitania (western Libya), and winters in portions of the Sahara and north- 908 ern sub-Saharan Africa; crassirostris breeds in eastern and southeastern Europe, the Middle East and Cyrenaica (eastern Libya), and winters in east Africa; and balchanica breeds in Transcaspia and Iran and winters in Arabia. Given the variety of ecotypes used by hortensis, ranging from temperate forest and high mountain country during the breeding season to desert scrub during the winter, and its close relationship with J/eu- comelaena, it seems plausible for the latter form to have adapted to one of these hab- itats, namely Acacia parkland. Throughout much of the Arabian and east African win- tering grounds hortensis occurs in scrub- land, often sympatrically with /eucome- laena; for example, in the Acacia plains of coastal Eritrea (Moreau 1972). Remembering that the genus Sy/via is of Palearctic origin, and that /euwcomelaena is the only Sy/via breeding in sub-Saharan Af- rica, it is presumed that /eucomelaena spe- ciated in the northern portion of the Afro- Arabian Rift Valley, then spread along the Acacia corridor of the western Arabian Pen- insula, and then crossed over the Red Sea (presumably at the narrows in the south) into east Africa. Once in Africa it spread north through the Red Sea Mountains and coastal plain until it reached the northern limit of dense Acacia groves. The modern sympatric occurrence of /eucomelaena and hortensis during the winter months in por- tions of Arabia and eastern Africa is pre- sumably secondary contact after the events of speciation and dispersal. The possibility that a population of the proto-modern hor- tensis/leucomelaena group wintering in east Africa became resident, speciated and spread northwards on both coasts of the Red Sea cannot be eliminated. However, this seems unlikely since no parallel exists for any other Sylvia species, the majority of which winter at least in part in Africa. Taxonomic Conclusions Sylvia 1. leucomelaena. —Meinertzhagen (1949) noted that the head and mantle colors PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of nominate /eucomelaena were paler than African birds. This conclusion is generally supported by the present study. Since sev- eral populations are sexually dimorphic in head coloration it is important to analyze this character only within a sex class. Some of Meinertzhagen’s other color comparisons were confounded by not taking this factor into account. Birds from the Arabian Pen- insula, including Yemen and Oman, tend to have less white on the outer pair of rec- trices than African birds, particularly those from Somalia/Eritrea. Specimens from the Arava differ from Arabian Peninsular birds in several ways: less white on the outer pair of rectrices (al- though the Arava sample consisted of only two birds); males have longer tails; and both sexes have substantially wider bills. These differences seem to warrant subspecific sep- aration of the Arava population. However, the naming ofa new form is suspended until further data are available.! Sylvia I. distincta. —No consistent differ- ence was found between birds from the Ye- mens and the balance of the Arabian Pen- insula. S. /. distincta is a synonym of S. /. leucomelaena. Sylvia I. somaliensis. —Populations from Somalia and Eritrea are easily recognizable from all others by the large amount of white on the outer pair of rectrices. No consistent difference was found in head coloration be- tween these populations and other African populations or those from the Arabian Pen- insula (cf. Meinertzhagen 1949). ' Shirihai, H. 1988. A new subspecies of Arabian Warbler Sylvia leucomelaena from Israel.— Bulletin of the British Ornithologists’ Club 108:64—-68, has re- cently proposed the name Sylvia leucomelaena negev- ensis for the Arava population. A copy of the current paper was sent to Mr. Shirihai on 26 June 1987 for his comments. In subsequent correspondence he made no mention that a manuscript describing this subspecies was in preparation and never provided the reciprocal courtesy of reviewing it. His paper was received by the Bulletin of the British Ornithologists’ Club on 10 Au- gust 1987. VOLUME 101, NUMBER 4 Sylvia |. blanfordi.— Morphologically the Gebel Elba, Egypt, and Sudanese popula- tions seem similar to one another, or dif- ferences are part of a north-south cline. The only exception is that Gebel Elba birds have distinctly longer exposed culmens than Su- danese birds; however, the degree of differ- ence may be partially an artifact of the small number of Egyptian birds available for comparison. Egyptian and Sudanese birds tend to have a shorter gonys. Both of these populations have significantly less white on the outer pair of rectrices than those So- malian/Eritrean. No important difference was found in head coloration among any of the African populations. Acknowledgments I am grateful to the following people for making material under their care available: D. M. Hamed, H. Hovel, H. Mendelssohn, J. V. Remsen, Jr., K.-L. Schuchmann, T. Shariv, E. Stickney, F. Vuilleumier, and M. Walters. A number of people kindly an- swered queries: D. Afik, J. Ash, M. D. Gal- lagher, M. C. Jennings (coordinator of the Atlas of Breeding Birds of Arabia), G. Ni- kolaus, U. Paz, B. Pinshow, R. F. Porter, H. Shirihai, and G. Welch. The work in Egypt was under the auspices of WWF/ IUCN Project no. 3612, and logistic support was provided by British Petroleum and the Egyptian Wildlife Service. The company and help of P. L. Meininger and R. W. Storer on the 1985 Gebel Elba expeditions is warmly acknowledged. The Frank M. Chapman Fund financed a portion of the museum work. The base map was kindly drafted by Mr. Edward Khounganian and Fig. 1 drawn by Ms. Margaret Van Bolt. For comments on an earlier draft of this paper Iam grateful to Dr. R. W. Storer and Messrs. M. D. Gallagher and M. C. Jennings. Literature Cited Afik (Aizik), D., & B. Pinshow. 1984. Notes on the breeding biology of the Arabian Warbler Sy/via 909 leucomelaena in the Arava (Rift Valley), Is- rael.—Ibis 126:82-89. Archer, G. F., & E.M. Godman. 1961. The birds of British Somaliland and the Gulf of Aden. Vol- ume 4. Oliver and Boyd, Edinburgh, 1043-1570 pp. Ash, J. S., & J. E. Miskell. 1983. Birds of Somalia. Scopus Special Supplement Number 1, 97 pp. Beals, E. W. 1970. Birds of a Euphorbia—Acacia woodland in Ethiopia: Habitat and seasonal changes.—Journal of Animal Ecology 39:277- 297. Boulos, L., & J. Hobbs. 1986. Three arboreal species new to the Eastern Desert of Egypt.—Candollea 41:183-191. Brooks, D. J., M. I. Evans, R. P. Martins, & R. F. Porter. 1987. The status of birds in North Ye- men and the records of the OSME expedition in autumn 1985.—Sandgrouse 9:4—66. Cave, F. O., & J. D. Macdonald. 1955. Birds of the Sudan. Oliver and Boyd, Edinburgh, 444 pp. Cornwallis, L., & R. F. Porter. 1982. Spring obser- vations on the birds of North Yemen.—Sand- grouse 4:1-36. Dowsett-Lemaire, F., & R. J. Dowsett. 1985. Breed- ing biology of the Arabian Warbler Sylvia leu- comelaena in Israel: Comments and suggestions for further research.—Ibis 127:567. Dresser, H. E., & W. T. Blanford. 1874. Notes on the specimens in the Berlin Museum collected by Hemprich and Ehrenberg.—Ibis 3(4):335- 343. Etchécopar, R. D., & F. Hiie. 1967. The birds ofnorth Africa from the Canary Islands to the Red Sea. Oliver and Boyd, Edinburgh, 612 pp. Gallagher, M. D. 1986. Additional notes on the birds of Oman, eastern Arabia 1980-86.—Sand- grouse 8:93-101. —., & M. W. Woodcock. 1980. The birds of Oman. Quartet Books, London, 310 pp. Goodman, S. M., & P. L. Meininger. (in pressa). The discovery of the Arabian Warbler Sylvia leu- comelaena in southeastern Egypt.—Courser 2. ——., & (eds.). (in press b.). The birds of Egypt. Oxford University Press, Oxford. Hall, B. P., & R. E. Moreau. 1970. An atlas of spe- ciation in African passerine birds. British Mu- seum (Natural History), London, 423 pp. Halevy, G., & G. Orshan. 1972. Ecological studies on Acacia species in the Negev and Sinai. I. Distribution of Acacia raddiana, A. tortilis and A. gerrardii ssp. negevensis as related to envi- ronmental factors. —Israel Journal of Botany 21: 197-208. Hartert, E. 1917. A few notes on the birds of Ye- men.—Novitates Zoologicae 24:454-462. 1961. The ecology of the coastal Hemming, C. F. 910 area of northern Eritrea.—Journal of Ecology 49:55-82. Hemprich, F. G., & C.G. Ehrenberg. 1833. Symbolae Physicae. Avium, Decas I. Berolini, ex Officina Academica. Hotzl, H. 1984. The Red Sea. Pp. 13-26 in A. R. Jado and J. G. Zotl, eds., Quaternary Period in Saudi Arabia, volume 2. Springer-Verlag, Wien. Hutchinson, M. 1975. Systematic list of birds ob- served at or near Riyadh. — Journal of the Saudi Arabian Natural History Society 14:12-30. Jennings, M. C. 1981. The birds of Saudi Arabia: A check-list. Private printing, Cambridge, 112 pp. Kassas, M. 1956. The mist oasis of Erkwit, Sudan. — Journal of Ecology 44:180-194. 1957. On the ecology of the Red Sea coastal land.—Journal of Ecology 45:187-203. —., & M. Imam. 1959. Habitat and plant com- munities in the Egyptian desert. IV. The gravel desert.— Journal of Ecology 47:289-310. —.,& M.A. Zahran. 1962. Studies on the ecology of the Red Sea coastal land. I. The district of Gebel Ataqa and el-Galala el-Bahariya.— Bul- letin de la Société de Géographie d’Egypte 35: 129-175. ——, & 1965. Studies on the ecology of the Red Sea coastal land. II. The district from el-Galala el-Qibliya to Hurghada.— Bulletin de la Société de Géographie d’Egypte 38:155-193. ———., & 1971. Plant life on the coastal mountains of the Red Sea coast, Egypt.—Jour- nal Indian Botanical Society 50A:571—589. Meinertzhagen, R. 1949. On the status of Parisoma leucomelaena (Hemprich and Ehrenberg). — Bulletin of the British Ornithologists’ Club 69: 109-110. . 1954. Birds of Arabia. Oliver and Boyd, Lon- don, 624 pp. Moreau, R. E. 1972. The Palaearctic-African bird migration systems. Academic Press, London, 384 pp. Ross, D. A., & J. Schlee. 1973. Shallow structure and geologic development of the southern Red Sea. — Bulletin of the Geological Society of America 84:3827-3848. Sclater, W. L., & C. Mackworth-Praed. 1918. A list of the birds of the Anglo-Egyptian Sudan. Part II. Alaudidae-Hirundinidae.—Ibis 10(6):602— Ale Seebohm, H. 1878. Ona new species of Sylvia from Abyssinia, and on some other Abyssinian Sy]l- vians. — Proceedings of the Zoological Society of London 1878:978—980. Shmida, A., & Y. Or. 1986. The Sudanian flora in Israel.—Society for the Protection of Nature in Israel, Rotem Bulletin Number 8 [in Hebrew], 150 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Smith, K. D. 1957. An annotated check list of the birds of Eritrea.—Ibis 99:307—337. Stagg, A. 1985. Birds of S.W. Arabia. 2nd. edit. Pri- vate printing, Riyadh, 59 pp. Tackholm, V. 1974. Students’ flora of Egypt. 2nd. edit. Cairo University, Beirut, 888 pp. , & L. Boulos. 1972. Supplementary notes to Students’ flora of Egypt.—Cairo University Herbarium Publication Number 5, 135 pp. Thomsen, P., & P. Jacobsen. 1979. The birds of Tu- nisia. Nature-Travels I/S, Copenhagen, 176 pp. Vaurie, C. 1959. The Birds of the Palearctic fauna. Passeriformes. H. F. and G. Witherby, London, 762 pp. Vesey-Fitzgerald, D. F. 1955. Vegetation of the Red Sea coast south of Jedda, Saudi Arabia. —Jour- nal of Ecology 43:477-489. 1957. The vegetation of the Red Sea coast north of Jedda, Saudi Arabia. — Journal of Ecol- ogy 45:547-562. Watson, G. E., M. A. Traylor, Jr., & E. Mayr. 1986. Family Sylviidae. Pp. 3-294 in E. Mayr and G. W. Cottrell, eds., Check-list of birds of the World, volume 11. Museum of Comparative Zoology, Cambridge, Massachusetts. Wolters, H. E. 1980. Die Vogelarten der Erde. Lie- ferung 5. Paul Parey, Hamburg, 321-400 pp. Zahavi, A., & R. Dupai. 1974. First breeding record of Blandford’s [sic] Warbler Sylvia leucome- laena. —Israel Journal of Zoology 23:55—56. Zohary, M. 1962. Plant life of Palestine. Ronald Press Co., New York, 262 pp. Museum of Zoology, The University of Michigan, Ann Arbor, Michigan 48109. Appendix | Material of Sylvia leuacomelaena Examined Museum codes: AMNH, American Mu- seum of Natural History, New York; BMNH, British Museum (Natural History), Tring; HH, Private collection of Haim Ho- vel, Haifa; LSUM, Louisiana State Univer- sity Museum, Baton Rouge; SGM, Sudan Government Museum, Khartoum; UMMZ, The University of Michigan Museum of Zo- ology, Ann Arbor; UTM, Zoological Mu- seum, University of Tel Aviv, Tel Aviv; YPM, Yale Peabody Museum, New Haven; ZFMK, Zoologisches Forschungsinstitut und Museum Koenig, Bonn. VOLUME 101, NUMBER 4 Israel.—Km 20 on Arava Road, UTM 8088, 8089; Yotvata, HH 82110, 82112, ZFMK 8416; Hatzeva, UTM 8153. Saudi Arabia.—Al Jurf, BMNH 1936.6.29.20-—23 (4); Ashaira (Asheira, Ushayeah), BMNH 1935.1.5.81—82 (2); Bir Askar, Taif Plateau, BMNH 1935.1.5.78- 79 (2); Bir Braman (Burayman), 15 km NE Jidda, BMNH 1940.4.1.24; Birka, BMNH 1950.58.2; Farha, BMNH 1935.1.5.80; Hadda, Wadi Fatima, BMNH 1936.6.29.18- 19 (2); Hamdha, upper Wadi Tathlith, BMNH 1937.4.17.275; Hejaz (al Hiyjaz), BMNH 1946.69.76-77 (2); Jidda (Jedda), 10 miles E., BMNH 1934.9.20.140- 142 (3), 1937.12.26.15; Khadra, Najran, BIVENS 937-4 lee SO Sq ela. 2a 7 Madriga (Madrakah), nr. Jidda, BMNH 1965.M.13775; Wadi Hafra, 80 km NE Jid- da, BMNH 1940.4.1.25; Wadi Harjab, Wadi Bisha, BMNH 1937.4.17.276, 1948.58.29: Wadi Hijla, BMNH 1946.69.78; Wadi Jaura, Tihama, Jizan, BMNH 1937.4.17.22;: Wadi Krarrar, near Taif, BMNH 1949.5.33. The Yemens.—A\| Kubar (el Kebir), Amiri Dist., South Yemen, BMNH 1903.8.12.25, AMNH 608810; Dhala (Dthala), Amiri Dist., South Yemen, BMNH 1903.8.12.26, 1965.M.13777; Gerba, Amiri Dist., South Yemen, BMNH 03.8.12.21-—24 (4), AMNH 608806 (type of distincta), 608807-809 (3); Habil, South Yemen, BMNH 1965.M.13776; Jajeilha, Yemen, AMNH 608811; Jebel Manif, north of Lahej, South Yemen, BMNH 1900.8.5.193; Lodar (Law- dar), South Yemen, BMNH 1965.M.13778: Ma’ir, Abian, South Yemen, BMNH 1900.8.5.194; Tullah, Hadhramaut, South Yamen, BMNH 1932.4.20.5; Wadi Awa, Hadhramaut, South Yemen, BMNH ili 1933.6.26.6; Wadi Khabb (Khubb), south- east Najran, Yemen, BMNH 1937.4.17.274: Wadi Thibi, South Yemen, BMNH NOS WEOe Role Oman. —Khadrafi, Jebel Qamr, Dhofar, BMNH 1977.1.10. Somalia.—Berbera Plain, BMNH 98.7.27.80; Bihendula (Bikendula), AMNH 608789-90 (2); Burao (Buraa), BMNH 1923.8.7.3038; Dubar, AMNH 608803-805 (3), BMNH 1905.11.27.164-165 (2); Eri- gavo, BMNH 1965.M.13773, LSUM un- cataloged (2); Galoli, BMNH 1923.8.7.3036: Gardo, BMNH 1945.10.11; Gidial (Gidil) Valley, Golis, AMNH 608800, BMNH 1923.8.7.3035; Las Khorai (Khoreh), AMNH 60879 1-792 (2); Mush Haled (Mush Aled, Musha Aled, Mash Caleed), Warsan- gli, 200 miles E. Berbera, AMNH 608793- 794 (2), YPM 32258; Mundara (Mandara), BMNH 98.6.13.76 (type of somaliensis); Sogsode (Sogsoda, Suksodi, Sugsade), BMINE 1923582730375 1925 s11e20.26. AMNH 608795-797 (3); Waghar (Wagr, Wogr), AMNH 608801-802 (2), BMNH 1905.11.27.168-169 (2). Eritrea. —near Massawa, BMNH 195222 551.9 Raino wes talbab aS MONI Er 69.10.16.85 (type of blanfordi.). Sudan. —Erba Mountains, BMNH 97.10.15.3; Erkowit, BMNH 1915.12.24.719- 720 (2), 1916.9.20.671, 1919.12.17.700-—703 (3), SGM 686, 2034, 2773, 3286; Port Su- dan, BMNH 1965.M.13774; Sinkat, BMNH 1919.12.17.683-699 (17). Egypt. —Wadi Aideib, Gebel Elba, UMMZ 224089-091 (3); Wadi Akwamtra, Gebel Elba, UMMZ 224092; Wadi Kansis- rob, Gebel Elba, UMMZ 224088. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 912-928 NEW SPECIES OF FOSSIL VAMPIRE BATS (MAMMALIA: CHIROPTERA: DESMODONTIDAE) FROM FLORIDA AND VENEZUELA Gary S. Morgan, Omar J. Linares, and Clayton E. Ray Abstract.—A new species of vampire bat is described from a partial skull and several postcranial elements from the early Pleistocene of Florida. Similar in size to the living Desmodus rotundus, it is considerably smaller than the only previously recognized species of North American fossil vampire, D. stocki. Characters that distinguish the new species include the large mastoid process, narrow occiput, reduced ventral flexion and posterior inflation of the braincase, and posteriorly directed foramen magnum. It occurs in fossil deposits ranging | in age from approximately 1.8 to 1.0 million years (early to middle Irvington- ian), and is more than one million years older than any previously known vampire bat. A second new species of giant vampire bat is described from two partial skeletons recovered from surficial deposits of uncertain Late Quaternary age in Cueva del Guacharo, Monagas, Venezuela. This new species is larger than any other described vampire and differs as well in its relatively slender and delicate skull, and mandible with straight ventral border and pockets behind incisors almost lacking. Skeletal fragments from caves in West Virginia and Yucatan represent vampires of similarly large size but undetermined affinities. Fossil remains, undoubtedly in part at least of Recent age, of the common vam- pire, Desmodus rotundus, were recognized by Lund from caves near Lagoa Santa, Mi- nas Gerais, Brazil, in the course of his ex- plorations of 1835-1844 (Ray et al. 1988). Remains thought to be Recent in age have been assigned to Desmodus rotundus from three caves, and to Diphylla ecaudata from one cave, in Venezuela (Linares 1968, 1970). The new species described here is the first evidence of an extinct vampire in South America. Fossil vampires were first report- ed from North America in a footnote to a paper on a new Species of Pleistocene ter- restrial gastropod from a cinnabar mine near Terlingua, Texas (Cockerell 1930). This large extinct vampire bat was later described as Desmodus stocki (Jones, 1958) from depos- its of late Pleistocene (Rancholabrean) age in San Josecito Cave, Nuevo Leon, northern Mexico. Gut (1959) described a second species, D. magnus, from the late Pleisto- cene Reddick 1 fauna, Marion County, Florida. Fossils of Desmodus from outside the Recent North American range of D. ro- tundus are known from late Pleistocene sites in northern Mexico, California, Arizona, New Mexico, Texas, Florida, West Virginia, and Cuba (Ray et al. 1988). Hutchison (1967) synonymized D. magnus with D. stocki on the basis of their similarity in size and other cranial features. Most recent workers have followed Hutchison in rec- ognizing D. stocki as the only extinct late Pleistocene species in North America. Ray et al. (1988) mapped and briefly summa- rized all known fossil occurrences of vam- pire bats. In Florida, Desmodus stocki has been re- corded from four Rancholabrean vertebrate faunas (Ray et al. 1988). Most of these fos- VOLUME 101, NUMBER 4 sils were referred originally to Desmodus magnus. Over the past 20 years numerous early Pleistocene (Irvingtonian) vertebrate fossil sites have been discovered in penin- sular Florida. Screenwashing for micro- vertebrates from these Irvingtonian locali- ties has revealed the presence of a new species of Desmodus in three faunas: the early Irvingtonian Inglis 1A Local Fauna, Citrus County, and the middle Irvingtonian Haile 16A and Haile 21A local faunas, Ala- chua County. These fossils represent the oldest known vampire bats. Meanwhile, in conjunction with field studies of bats and surveys of caves in Ven- ezuela, Linares in 1965 recovered two geo- logically young fragmentary skeletons of a very large new species of vampire bat. Ef- forts by Linares and Ray to determine the affinities of this form led to comprehensive study of the fossil record of vampires and to comparison of the hard parts of the three living species. Our mutually overlapping and converging interests in fossil vampires have resulted in presentation of a synopsis of our findings (Ray et al. 1988) and in a more detailed review we have in progress. Delay in publication of that review has caused pre- mature allusion in print to the new species from Florida and Venezuela. This, together with the intense current interest in vam- pires, has induced us to validate the names of the two new species without further de- lay. The fossils utilized for this paper are de- posited in the vertebrate paleontological collections of the Florida State Museum, University of Florida (UF) and the Florida Geological Survey (UF/FGS), also housed at the Florida State Museum; the Depart- ment of Paleobiology, U.S. National Mu- seum of Natural History, Smithsonian In- stitution (USNM); Department of Vertebrate Paleontology, Los Angeles County Museum of Natural History (LACM); Seccion de Pa- leobiologia, Museo de Ciencias Naturales, Universidad Simon Bolivar, Caracas, Ven- ezuela (MUSB). Recent comparative ma- 913 terial is from the Division of Mammals, Na- tional Museum of Natural History, Smithsonian Institution (USNM), the De- partment of Mammalogy, American Mu- seum of Natural History (AMNH), and the mammalogy collection of the Florida State Museum (UF). All measurements were tak- en with dial calipers and rounded to the nearest 0.1 mm. Descriptive morphological terms and cranial measurements follow DeBlase and Martin (1981). Systematic Paleontology Class Mammalia Linnaeus Order Chiroptera Blumenbach Family Desmodontidae Gill Genus Desmodus Maximilian Desmodus archaeodaptes, new species Fig. 1 D. praecursor. —Ray et al., 1988:20 (nomen nudum). Desmodus archaeodaptes. — Ray et al., 1988: 22 et sqq. (Nomen nudum). Holotype. —UF 94526, nearly complete braincase posterior to interorbital constric- tion, lacking only zygomatic arches, with associated left periotic. Collected by Law- rence H. Martin, Jr., in May 1983. Type locality. —Haile 21A Local Fauna of early Pleistocene (middle Irvingtonian) age, located 4.6 km northeast of Newberry, Ala- chua County, Florida. Referred material. —UF 94527, UF 94528, two proximal ends of left humeri from type locality; UF 40046, complete right humerus, from Inglis 1A Local Fauna of early Pleistocene (early Irvingtonian) age, located 3 km southwest of Inglis, Citrus County, Florida; UF 24206, distal end of left humerus and UF 40047, proximal end of left radius, from Haile 16A Local Fauna of early Pleistocene (middle Irvingtonian) age, located 5.8 km northeast of Newberry, Alachua County, Florida. Etymology. —From the Greek archaios, ancient, old, and daptes, eater of blood by 914 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Cranial and mandibular measurements of Desmodus archaeodaptes, D. rotundus, D. stocki, and D. draculae. Mean, standard deviation, observed range (in parentheses), and sample size are given for each mea- surement. Species and Total length Condylobasal Breadth Interorbital Zygomatic locality of skull length of rostrum constriction breadth Desmodus archaeodaptes (UF 94526, holotype) — — — 5.8 12.4 Haile 21A Florida Desmodus rotundus murinus 23.7 + 0.44 21.2 + 0.36 6.0 + 0.21 5.5 + 0.17 12.0 + 0.29 Mexico (22.7-24.3) (20.5-21.8) (5.5—6.5) (5.1-5.8) (11.5-12.7) 30 30 30 30 30 Desmodus r. rotundus 24.8 + 0.53 22.3 + 0.56 6.6 + 0.28 Sef ae O23 12.8 + 0.28 Chile and Paraguay 24.1-25.5 (21.5—23.3) (6.1—7.0) (5.4—6.0) (12.3-13.1) 10 10 10 10 10 Desmodus stocki D3) se O57 24.5 + 0.18 7.4 + 0.16 6.1 + 0.11 14.0+0 San Josecito Cave (26.5—28.2) (24.1-24.7) (7.1-7.6) (5.9-6.2) (14.0) Mexico 6 6 6 9 2 Desmodus stocki — — — 6.1 + 0.21 14.3 + 0.20 Reddick (5.8-6.3) (14.1-14.5) Florida 3 2 Desmodus draculae 31.2, — 29.7, — 9.2, — 5.7, 6.7 — (MUSB 152-85 PB, holo- type; USNM 23568, paratype) Cueva del Guacharo, Venezuela sucking. The name refers to the fact that this is the oldest known species of vampire bat. Diagnosis. — Differs from other known species of Desmodus in possessing broad plate-like mastoid process, narrower Occi- put, lateral connection of nuchal crest to paroccipital process; and from all except D. draculae in nearly vertical orientation of su- praoccipital, lack of inflation and ventral flexion of posterior portion of braincase, and posteriorly oriented foramen magnum with rounded dorsal margin. Desmodus archaeo- daptes differs from D. rotundus in presence of larger glenoid fossa, reduced postglenoid process, smaller occipital protuberance, and weakly inflated supraoccipital; from D. stocki in smaller size, longer and narrower braincase, weaker cranial crests, ventrally deflected paroccipital process, shallow basi- cranial pits separated by low indistinct ridge, and weakly inflated posteromedial process of basisphenoid; and from D. draculae in much smaller size. Measurements. —See Tables 1 and 2. Description and comparisons. —The braincase of the holotype of Desmodus ar- chaeodaptes from Haile 21A is nearly com- plete posterior to the interorbital constric- tion. The zygomatic arches are missing except for the base of the right squamosal process. This skull is within the size range of Recent D. rotundus and is considerably smaller than that of D. stocki and D. drac- ulae (see Table 1 and Fig. 1). Desmodus archaeodaptes more closely re- sembles D. rotundus than D. stocki in cer- tain cranial features: these include narrower more elongated braincase, reduced cranial crests, ventral deflection of the paroccipital VOLUME 101, NUMBER 4 915 Table 1.—Continued. Breadth of Length of Height Total length Length of mandibu- Height of Mastoid breadth braincase braincase of braincase of mandible lar tooth row coronoid 12.4 12.3 17.2 12.0 — = = 12.5 + 0.29 11.9 + 0.24 16.9 + 0.35 11.9 + 0.39 15.3 se 03333 4.7+ 0.14 6.1 + 0.20 (12.0-13.1) (11.5-12.5) (16.4-17.9) (11.1-12.6) (14.5-16.0) (4.4—5.0) (5.6-6.5) 30 30 30 30 30 30 3 13.0 + 0.22 12.6 + 0.32 17.4 + 0.33 12.3 + 0.36 15.8 + 0.40 4.8 + 0.09 6.3 + 0.26 (12.6-13.3) (12.1-13.0) (17.1-18.2) (11.9-12.6) (15.3-16.6) (4.7-4.9) (6.0—6.8) 10 10 10 10 10 10 10 14.1 + 0.33 13.8 + 0.20 19.2 + 0.32 3S) == OF17 — = = (13.5-14.5) (13.6—-14.2) (18.7-19.7) (13.2-13.8) 8 11 9 10 14.4 + 0.25 14.1 + 0.22 19.6 + 0.08 13.2 + 0.19 17.4 + 0.26 mas eae OP 0) 5 7/ ss WAI (14.1-14.7) (13.8-14.4) (19.5-19.7) (13.0-13.5) (17.0-17.6) (5.1-5.4) (6.2-6.9) 5 4 3 4 3 16 7 NS\2, 535) 14.8, 14.5 21.1, 22.4 14.8, 13.4 21.9, — 8.5, 8.3 9.4, — process, weakly inflated posteromedial pro- cess of the basisphenoid, and shallow basi- cranial pits. There are also many characters that distinguish the skulls of D. archaeo- daptes and D. rotundus. The occipital pro- tuberance only slightly overhangs the oc- cipital condyles in D. archaeodaptes, whereas in D. rotundus the braincase is ex- panded posteriorly and flexed ventrally so that the occipital protuberance occupies a more posteroventral position. The occipital protuberance is also larger in the living species and has a prominent posteriorly projecting process. In lateral view, the su- praoccipital of D. archaeodaptes is nearly vertical, but in D. rotundus, because of in- flation and ventral flexion of the braincase, it is tilted posteriorly. In the essentially un- modified braincase of D. archaeodaptes the foramen magnum faces posteriorly, where- as in D. rotundus it is oriented ventrally. The supraoccipital in D. archaeodaptes is essentially flat, but in D. rotundus the region along the midline between the foramen magnum and the occipital protuberance is noticeably inflated. The occiput of D. ar- chaeodaptes is narrower and higher than in the Recent species. The occipital protuber- ance is more dorsad, and the nuchal crests (forming the lateral borders of the occiput) make a more acute angle in D. archaeo- daptes than in D. rotundus because they connect laterally to the paroccipital pro- cesses rather than to the mastoid processes. Laterally, the nuchal crest turns abruptly ventrad in D. archaeodaptes and becomes confluent with the lateral edge of the par- occipital process; a weak secondary crest branches from the lateral edge of the nuchal crest and connects with the mastoid process. In D. rotundus the mastoidal connection is emphasized and the paroccipital connection is lost. The braincase of Desmodus stocki is larg- er overall and relatively shorter, broader, and more globose than that of D. archaeo- daptes and D. rotundus, but is intermediate between these smaller species in certain oth- 916 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.— Measurements of the humerus of Recent and fossil Desmodus. Mean, standard deviation, observed range (in parentheses), and sample size are given for all measurements, except those taken from Hutchison (1967). Species and locality Total length Desmodus archaeodaptes (UF 40046) Inglis 1A 39.7 Florida (UF 24206) Haile 16A — Florida (UF 94527) Haile 21A seh Florida Desmodus rotundus 37.0 + 2.33 (32.4—42.4) 19 Desmodus stocki San Josecito Cave 43.6 Mexico (39.3-47.5) (from Hutchison, 1967) 42 Reddick 41.8 + 1.34 Florida (39.4—44.3) 15 Desmodus draculae 51.0 (MUSB 152-85 PB, holotype) 1 Cueva del Guacharo Venezuela 1 Proximal width Distal width Medial width of shaft So 7/ 6.2 Dodd — 5.9 _ 5 - Dale 4.9 + 0.28 5.4 + 0.30 2.1 + 0.18 (4.4—5.6) (4.85.9) (1.7—2.4) 19 19 15 6.3 6.8 2.5 (5.8-6.8) (6.4-7.3) (2.0-2.9) 47 52 56 6.3 + 0.16 6.8 + 0.18 2.6 + 0.11 (6.0-6.7) (6.4-7.2) OQDT) 38 42 42 — 8.5 3.3 1 er characters. The posterior portion of the braincase of D. stocki is more inflated and the occipital protuberance and foramen magnum are located in a more ventral po- sition than in D. archaeodaptes. Compared with D. rotundus, however, the braincase of D. stocki is not expanded as far posteriorly, nor does it attain the same degree of ventral flexion. The dorsal margin of the foramen magnum is rounded in D. archaeodaptes in contrast to its hexagonal shape in D. stocki. The cranium of D. stocki resembles that of the living species in the more ventral po- sition of its occipital protuberance and the lateral connection of the nuchal crest to the mastoid process; these features give the oc- ciput the appearance of being broader, but shallower, dorsoventrally than in D. ar- chaeodaptes. The articular surface of the glenoid fossa is relatively large in Desmodus archaeo- daptes and D. stocki, and the postglenoid process is short and blunt. Desmodus ro- tundus has a smaller glenoid fossa and the postglenoid process is more elongated and triangular in shape. The smaller postglenoid = Fig. 1. Skulls of vampire bats in dorsal (A—C), ventral (D-—F), right lateral (G-I), and posterior (J—L) aspect. A, D, G, J, Desmodus rotundus, UF 20664, Recent, Paraguay; B, E, H, K, Desmodus stocki, UF/FGS 5646, Rancholabrean, Reddick 1, Florida; C, F, I, L, Desmodus archaeodaptes, UF 94526 (holotype), Irvingtonian, Haile 21A, Florida. Scale is one cm. + fo aa aa = S) Z S ea} = S) | ‘e) S 918 process of D. archaeodaptes and D. stocki, coupled with the larger articulating surface for the mandible, suggests that these extinct species may have had somewhat greater freedom of movement of the lower jaw than D. rotundus. The posteromedial process of the basisphenoid is weakly developed in the two smaller vampires, but is strongly in- flated in D. stocki. The basicranial pits lo- cated anterior and medial to the tympanic cavity are barely noticeable depressions in D. archaeodaptes, but are larger, deeper, and extend farther posteriorly in D. stocki. In the larger species these pits are separated by a thin, knife-like ridge of bone that extends posteriorly to a point about halfway be- tween the anterior and posterior edges of the tympanic cavity. This ridge is low and indistinct in D. archaeodaptes and disap- pears at the anterior edge of the tympanic cavity leaving the basioccipital almost per- fectly flat. The basicranial pits and ridge are intermediate in their development in D. ro- tundus. The paroccipital process is ventrally deflected in D. archaeodaptes and D. rotun- dus, but oriented posteriorly in D. stocki. The broad ventrally rounded mastoid pro- cess of D. archaeodaptes is larger than that of D. stocki or D. rotundus. The total length of the complete humerus from Inglis 1A referred to Desmodus ar- chaeodaptes is within the observed range of humeri of Recent D. rotundus, although it is longer than any humerus measured of Middle American specimens referred to D. rotundus murinus (Table 2). The proximal and distal widths of this specimen slightly exceed those measurements in the same sample of D. rotundus. The length of the Inglis humerus barely overlaps the lower end of the range of variation of D. stocki humeri from Reddick 1C, while the prox- imal and distal widths are less than those for any humerus of D. stocki measured (Ta- ble 2). Two proximal ends of humeri from Haile 21A anda single distal end from Haile 16A are slightly smaller than the humerus from Inglis 1A, but are well within the ob- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON served range of measurements for Recent D. rotundus. There are no apparent char- acters, other than size, that differentiate the humeri of D. rotundus, D. archaeodaptes, and D. stocki. Geologic age and localities. —Desmodus archaeodaptes is described on the basis of six specimens from three localities in north- ern peninsular Florida, all of which are early Pleistocene (Irvingtonian) in age. Biostrati- graphic correlations with the North Amer- ican Land Mammal biochronology estab- lish Inglis 1A as very early Irvingtonian in age (between about 1.8 and 1.5 Ma), while Haile 16A and Haile 21A are middle Ir- vingtonian (between about |.5 and 1.0 Ma). The oldest previously recorded vampire bats were samples of D. stocki from the late Pleistocene (early or middle Ranchola- brean) Arredondo 2A and Reddick 1A, 1B, and 1C faunas, also from northern Florida. With the exception of the probably Recent remains from San Miguel Island, all fossils of D. stocki from the western United States and northern Mexico are latest Pleistocene (late Rancholabrean) in age (Kurten & An- derson 1980, Ray et al. 1988), a time in- terval during which Desmodus appears to have been absent from Florida (Morgan 1985). Fossil sites representing former caves, fis- sures, and sinkholes are common in the karst terrain of northern peninsular Florida. Most of these sites are clastic infillings deposited in solution features developed in the soft marine limestones of the upper Eocene Oca- la Group. These fossiliferous terrestrial sed- iments are most often exposed by commer- cial limestone mining. Many Florida State Museum vertebrate fossil sites are identified by numbers and letters following the general locality name (e.g., Haile 21A). The name (e.g., Haile) refers to the geographic location of the site, the number identifies the partic- ular limestone quarry or specific area in which the site is (or was) located, and the letter refers to individual fossil deposits within one quarry or specific area. Even VOLUME 101, NUMBER 4 though two or more fossil sites may be found in the same limestone quarry, it cannot be assumed without strong evidence that they are necessarily close in age or formed through similar depositional processes. The Inglis 1A Local Fauna was collected from the north bank of the now-defunct Cross Florida Barge Canal in Citrus County, Florida, 3 km SW of Inglis (29°01’N, 82°42'W), SE 4, SE %4, Sec. 9, T17S, R16E, Yankeetown 7.5 minute quadrangle, U.S. Geological Survey (1955). The fossiliferous sediments, now completely excavated, con- sisted of alternating layers of sand and clay filling a large solution cavity in the Eocene Inglis Formation. The geology and stratig- raphy of Inglis 1A were discussed by Klein (1971). A diverse chiropteran fauna occurs at Inglis 1A supporting geological evidence that this deposit represents a former cave. Comprehensive studies have been con- ducted on the birds (Carr 1980) and snakes and lizards (Meylan 1982) of this fauna, and a list of the mammals was given by Webb & Wilkins (1984). Inglis 1A contains mam- mals found in both late Blancan and early Irvingtonian faunas in Florida, including Glyptotherium cf. G. arizonae (Gillette & Ray 1981), Megalonyx leptostomus (Mc- Donald 1977), Trigonictis cf. T. macrodon (Ray et al. 1981), Chasmaporthetes ossifra- gus (Berta 1981), and Capromeryx arizo- nensis (Klein 1971), but lacks forms re- stricted to the Blancan, including Borophagus, Nannippus phlegon, and Equus (Dolichohippus). An early Irvingtonian age for this fauna is further indicated by the presence of Geomys propinetis (Wilkins, 1984) and Sigmodon curtisi (Martin 1979). The Haile 16A Local Fauna was discov- ered in a limestone quarry 5.8 km NE of Newberry, Alachua County, Florida (29°41'N, 82°34’W), NE %4, NE 4, Sec. 25, T9S, R17E, Newberry 7.5 minute quadran- gle, U.S. Geological Survey (1968). The fos- sil vertebrates at this site occurred in a large fissure with massive dark silty clays. The depth, areal extent, and stratigraphy of this 919 deposit could not be determined accurately as it was destroyed by limestone quarrying operations before it could be excavated. Al- though all fossils from Haile 16A were ob- tained by screenwashing spoil piles, there is no evidence that the fauna is mixed. No list of the mammalian fauna from Haile 16A has been published; however, a few of the taxa have been studied, including Trigonic- tis (Ray et al. 1981), Geomys (Wilkins 1984), Erethizon (Frazier 1981), and Sigmodon (Martin 1979). According to Martin, Sig- modon libitinus, a species known only from Haile 16A, is intermediate between S. cur- tist from the early Irvingtonian Inglis 1A fauna and S. bakeri from the late Irving- tonian Coleman 2A fauna. A middle Ir- vingtonian age for Haile 16A is also indi- cated by the presence of Megalonyx wheatleyi (McDonald 1977), Tapirus haysii, and an early species of Pitymys, and by the stage of evolution of Holmesina. The Haile 21A Local Fauna 1s located 4.6 km northeast of Newberry, Alachua Coun- ty, Florida (29°41'N, 82°33’W), SE 4, NE 4 Sec. 26, T9S, RI17E, Newberry 7.5 mi- nute quadrangle, U.S. Geological Survey (1968). This site was discovered in 1983 by Eric Kendrew and was worked by the Flor- ida State Museum between October 1983 and March 1984. Fossils were initially re- covered from Haile 21A after the land sur- face had been graded with heavy equipment in preparation for mining. The fossiliferous sediments consist of clays, sands, and lime- stone breccias filling a former cave or sink- hole roughly circular in outline, some 10 m in diameter and 5 to 7 m in depth. There are no previous published references to the Haile 21A vertebrate fauna. The fauna is overwhelmingly dominated by a large species of Platygonus, similar to samples of Platygonus from other Irvingtonian sites in Florida. The occurrence of Smilodon graci- lis suggests a pre-late Irvingtonian age for Haile 21A (Berta 1987), and the presence of Tapirus haysii and the association of Canis edwardii and C. armbrusteri are in- 920 Natural Barriers is Present Limit of Living (Siphons) Desmodus rotundus Entraktes is IAA Elevation 1065 Meters O 500 Meters Approximate Scale ®A (Galeria Derecha N°3, D. draculae, Paratype) ®B (Galeria de los Italianos; D. draculae , Holotype) ®C (Arctothere Locality) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ss ff a B Fig. 2. Sketch map of Cueva del Guacharo, Monagas, Venezuela, the type locality of Desmodus draculae. In part after Anonymous, 1968, with modifications and additions by Linares. dicative of Florida middle Irvingtonian fau- nas. Desmodus draculae, new species Fig. 2-8 “‘Una forma de Desmodus posiblemente del Pleistoceno . . . mayor que todas las co- nocidas. . . una nueva especie.’’— Linares 1968:138-139. ““A new species of Desmodus from Vene- zuela.” —Martin 1972:326. “A new species of fossil vampire bat from Venezuela.’ — Woloszyn & Mayo 1974: 260. Desmodus sp.—Smith, 1976:66. “Fossil Desmodus larger than all previously known forms.” —Greenhall et al. 1983:1. Desmodus draculae. — Linares, 1987:1 1 (fig- ured on p. 10).—Ray et al., 1988:20 et sqq., figs. 2-5 (nomen nudum). Holotype. —-MUSB 152-85 PB, skull, mandible, skeletal parts. Skull lacks zygo- matic arches and all teeth save incisors; mandible essentially complete, retains right canine and all postcanine teeth and left P,; skeletal parts include left humerus, distal two-thirds (with ulnar sesamoid in place) and head of right humerus, distal half or more of left and right radii, with vestiges of coossified ulnae, and three vertebrae. All elements white in color, thoroughly leached and chalky in texture though retaining sharp surface detail, except for the radii, which are in part deeply pitted. Collected by Omar J. Linares, 10-18 Apr 1965. Type locality. —Cueva del Guacharo, District of Caripe, State of Monagas, Ven- ezuela (Fig. 2), 10°10’27’N, 62°33’07’W. This cave has been described, illustrated, and mapped by the Sociedad Venezolana de Espeleologia in its Boletin (Anonymous 1968, 1971). The cave is at least 9425 me- ters long, of which the first 1041 meters (known as the Tourist Zone) is the more readily accessible, and is terminated by nat- VOLUME 101, NUMBER 4 921 Fig. 3. Skulls of vampire bats in dorsal aspect: A, Diphylla ecaudata, USNM 269507: B, Diaemus youngi, AMNH 175654; C, Desmodus draculae, holotype, MUSB 152-85; D, Desmodus rotundus, USNM 114977. Scale is One cm. ural barriers in the form of siphons. The type specimen of D. draculae was collected at the surface on the floor of the Galeria de los Italianos, locality B on our sketch map (Fig. 2), at least 1973 meters from the pres- ent entrance of the cave, and some 1400 meters beyond the limit of living D. rotun- dus. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 922 "WD 9UO SI I[BIS “LLOVIT WNsn ‘snpunjod snpowusag “ ‘dd $8-7S1 ASAW “edAr0joy ‘anjnov4p snpowusad ‘O -vS9SL1 HNNVY 18unod snuaniq “ ‘L0S69T WNSN ‘Yepnvza vjcydiq “V :joodse [eso}¥] UI s}eq oIIduIeA Jo sTINAS “py “BI VOLUME 101, NUMBER 4 923 Fig. 5. Skulls of vampire bats in ventral aspect: A, Diphylla ecaudata, USNM 269507; B, Diaemus youngi, AMNH 175654. C, Desmodus draculae, holotype, MUSB 152-85 PB; D, Desmodus rotundus, USNM 114977. Scale is one cm. Paratype. —USNM 23568, incomplete skull, lacking most of facial region and all teeth; incomplete mandible lacking poste- rior ends of rami and all teeth except base of right canine; postcranial fragments, in- cluding fragment of right scapula, and frag- ments of left and right humeri, radii (with coossified portions of ulnae), and femora. 924 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Lene en jesse ia eee] Fig. 6. Skulls and jaws of vampire bats. Skulls in posterior aspect: A, Diphylla ecaudata, USNM 269507; B, Diaemus youngi, AMNH 175654. Skulls in anterior aspect: C, Diphylla ecaudata, USNM 269507; D, Diaemus youngi, AMNH 175654. Right mandibular rami in labial aspect; E, Diphylla ecaudata, USNM 269507; F, Diaemus youngi, AMNH 175654. Scale is one cm. Preservation similar to that of holotype, but not as good. Two proximal ends of right radii indicate still another individual. Col- lected by Omar J. Linares, 10-18 Apr 1965, at locality A, Cueva del Guacharo (Fig. 2). Etymology. —The specific epithet of this largest known chiropteran vampire com- memorates Count Dracula, the greatest hu- man vampire of folklore (Wolf 1975). Diagnosis. —A Desmodus larger than any other known form (greatest length of skull more than 31 mm vs. 27.4 maximum for D. stocki, length of humerus 51 mm vs. 47.5 maximum for D. stocki), but skull more VOLUME 101, NUMBER 4 925 Fig. 7. Skulls and jaws of vampire bats. Skulls in posterior aspect: A, Desmodus draculae, holotype, MUSB 152-85 PB; B, Desmodus rotundus, USNM 114977. Skulls in anterior aspect: C, Desmodus draculae, holotype, MUSB 152-85 PB; D, Desmodus rotundus, USNM 114977. Right mandibular rami in labial aspect: E, Desmodus draculae, holotype, MUSB 152-85 PB; F, Desmodus rotundus, USNM 114977. Scale is one cm. slender and delicate. Mandible with ventral Description and comparisons. —Except as border straight in lateral aspect and with otherwise indicated here, characters of skull pockets behind incisors virtually non- of Desmodus draculae essentially as in D. existent. rotundus. Skull long and narrow; facial re- Measurements. —See Tables 1 and 2. gion even more reduced; interorbital con- 926 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. Jaws and humeri of vampire bats. Mandibles in dorsal aspect: A, Diphylla ecaudata, USNM 269507; B, Diaemus youngi, AMNH 175654; C, Desmodus draculae, holotype, MUSB 152-85 PB; D, Desmodus rotundus, USNM 114977. Left humeri in cranial aspect: E, Desmodus draculae, holotype, MUSB 152-85 PB; F, Desmodus rotundus, USNM 114977. Left humeri in medial aspect; G, Desmodus draculae, holotype, MUSB 152-85 PB; H, Desmodus rotundus, USNM 114977. Scale is one cm, pertaining to A-D. Length of E and G is 51 mm, and F and H, 42.1 mm. striction even farther forward and relatively dian lambdoidal extension; nasals even greater; zygomatic ramus of maxilla direct- more reduced and depressed; maxillae ed more posteriad; occipital margin less scarcely expanded lateral to nasals. In lat- clearly lyriform, with only suggestion ofme- eral aspect, rostrum even more upturned VOLUME 101, NUMBER 4 and nares even more dorsally directed than in D. rotundus; dorsal profile of skull slopes forward from high braincase on a smooth line, uninterrupted by the slight nasal pro- jection seen in D. rotundus. In ventral as- pect, bullae are seen to be still more reduced than in D. rotundus, and are firmly coos- sified with skull anteriorly and posteriorly (as seen posteriorly in Diphylla ecaudata); palate longer and narrower, incisive foram- ina smaller than in D. rotundus. Mandible also as in D. rotundus except as indicated here. Pockets behind incisors for reception of tips of upper incisors hardly discernible. Tips of upper incisors appar- ently would have rested medial to inner lower incisors, not in part behind them. In lateral aspect, ventral border of mandible quite straight, not curved as in D. rotundus; posteriorly it curves abruptly dorsad toward angular process. Posterior part of mandible overall resembles that of Diaemus youngi rather than Desmodus rotundus. The postcranial parts preserved exhibit few distinctions from Desmodus rotundus and D. stocki other than larger size and con- comitant robustness. The entepicondylar process of the humerus is more slender in proximodistal expansion than is that of D. rotundus and D. stocki. Although all spec- imens of the radius and ulna are incomplete, deeply pitted, and poorly preserved, it does appear that the shaft of the ulna distal to its coossification with the radius is more re- duced than in D..rotundus and D. stocki. In none of the specimens can the ulnar shaft be traced with certainty far beyond the point of fusion; additional, better-preserved spec- imens will be needed to resolve the point. A few fragments each from cave deposits in Yucatan and West Virginia represent large vampires of uncertain affinities, but possi- bly related to D. draculae (Ray et al. 1988). Geologic age. —Quaternary. Unfortu- nately there is little basis on which to suggest an age for this material. The remoteness of the site within the cave, the occurrence of an arctothere in the same sector (Fig. 2, lo- 927 cality C), and the presumed extinct status of the species, all suggest an early, possibly late Pleistocene, age. The rotten condition of the bones and occurrence at the surface in a wet, active cave, in the same sector with similarly preserved remains of Desmodus rotundus and Diphylla ecaudata, are more consistent with a late, possibly Recent age. Only additional discoveries of fossils in a more revealing geologic context, or of living D. draculae, will resolve the problem. Acknowledgments Helpful comments on the manuscript were provided by A. M. Greenhall, C. O. Hand- ley, Jr., A. E. Pratt, and S. D. Webb. The holotype of Desmodus archaeodaptes was generously donated to the Florida State Mu- seum by L. H. Martin, Jr. The drawings for Figs. 2-6 were made by L. B. Isham. Final figures were prepared by M. Parrish. This is University of Florida Contribution to Pa- leobiology Number 311. Literature Cited Anonymous. 1968. Catastro espeleologico de Vene- zuela. Mo. 1—Cueva del Guacharo.—Boletin de la Sociedad Venezolana de Espeleologia 1 (2): 97-107. 1971. Catastro espeleologico de Venezuela. Mo. 1 —Cueva del Guacharo. 2da. Parte.— Bo- letin de la Sociedad Venezolana de Espeleologia 3(2):116-131. Berta, A. 1981. The Plio-Pleistocene hyaena Chas- maporthetes ossifragus from Florida.—Journal of Vertebrate Paleontology 1:341-356. 1987. The Sabrecat Smilodon gracilis from Florida and a discussion of its relationships (Mammalia, Felidae, Smilodontini).— Bulletin of the Florida State Museum, Biological Sci- ences 31(1):1-63. Carr, G. S. 1980. Early Pleistocene avifauna from Inglis 1A, Citrus County, Florida. Unpublished Ph.D. Dissertation, University of Florida, Gainesville, 146 pp. Cockerell, T. D. A. 1930. An apparently extinct Eu- glandina from Texas.— Proceedings of the Col- orado Museum of Natural History 9(5):52-53. DeBlase, A. F., & R. E. Martin. 1981. A manual of mammalogy, Second Edition. Wm. C. Brown Co. Dubuque, Iowa, 436 pp. Frazier, M. K. 1981. A revision of the fossil Erethi- 928 zontidae of North America.—Bulletin of the Florida State Museum, Biological Sciences 27(1): 1-76. Gillette, D. D., & C. E. Ray. 1981. Glyptodonts of North America.— Smithsonian Contributions to Paleobiology 40:1—255. Greenhall, A. M., G. Joermann, U. Schmidt [& M. R. Seidei]. 1983. Desmodus rotundus.—Mam- malian Species 202:1-6. Gut, H. J. 1959. A Pleistocene vampire bat from Florida.—Journal of Mammalogy 40:534—538. Hutchison, J. H. 1967. A Pleistocene vampire bat (Desmodus stocki) from Potter Creek Cave, Shasta County, California.—Paleobios 3:1-6. Jones, J. K., Jr. 1958. Pleistocene bats from San Jo- secito Cave, Nuevo Leon, Mexico. — University of Kansas Publications, Museum of Natural History 9(14):389-396. Klein, J.G. 1971. The ferungulates of the Inglis 1A local fauna, early Pleistocene of Florida. Un- published MS Thesis, University of Florida, Gainesville, 115 pp. Kurten, B., & E. Anderson. 1980. Pleistocene mam- mals of North America. Columbia University Press, New York, 442 pp. Linares, O. J. 1968. Quiropteros subfosiles encon- trados en las cuevas venezolanas. Parte I. De- posito de la Cueva de Quebrada Honda (De- signacion de Catastro Ar. 1).—Boletin de la Sociedad Venezolana de Espeleologia 1(2):119- 145. . 1970. Quiropteros subfosiles encontrados en las cuevas venezolanas. Parte III. Desmodus ro- tundus en la Cueva de la Brujula (Mi. 1) Mi- randa.—Boletin de la Sociedad Venezolana de Espeleologia 3:33-36. . 1987. Murciélagos de Venezuela. Cuadernos Lagoven, Caracas, Venezuela, 122 pp. Martin, R. A. 1972. Synopsis of late Pliocene and Pleistocene bats of North America and the An- tilles.—American Midland Naturalist 87:326- 335. 1979. Fossil history of the rodent genus Sig- modon.—Evolutionary Monographs 2:1-36. McDonald, H.G. 1977. Description of the osteology of the extinct gravigrade edentate Megalonyx, with observations on its ontogeny, phylogeny, and functional anatomy. Unpublished MS The- sis, University of Florida, Gainesville, 326 pp. Meylan, P. A. 1982. The squamate reptiles of the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Inglis 1A fauna (Irvingtonian), Citrus County, Florida. — Bulletin of the Florida State Museum, Biological Sciences 27(3):1-85. Morgan, G. S. 1985. Fossil bats (Mammalia: Chi- roptera) from the late Pleistocene and Holocene Vero fauna, Indian River County, Florida.— Brimleyana 11:97-117. Ray, C. E., E. Anderson, & S. D. Webb. 1981. The Blancan carnivore Trigonictis (Mammalia: Mustelidae) in the eastern United States.— Brimleyana 5:1-36. ——., O. J. Linares, & G. S. Morgan. 1988. Pa- leontology. Chapter 3, pp. 19-30 in A. M. Greenhall & U. Schmidt, eds., Natural history of vampire bats. CRC Press, Inc., Boca Raton, Florida. Smith, J.D. 1976. Chiropteran evolution, in Biology of bats of the New World family Phyllostomat- idae. Part 1. The Museum, Texas Tech Uni- versity, Special Publication 10:49-69. Webb, S. D., & K. T. Wilkins. 1984. Historical bio- geography of Florida Pleistocene mammals. Pp. 370-383 in H. H. Genoways & M. R. Dawson, eds., Contributions in Quaternary vertebrate pa- leontology: A volume in memorial to John E. Guilday. Carnegie Museum of Natural History, Special Publication 8. Wilkins, K. T. 1984. Evolutionary trends in Florida Pleistocene pocket gophers (genus Geomys), with description of a new species.—Journal of Ver- tebrate Paleontology 3:166-181. Wolf, L. 1975. The annotated Dracula. Clarkson N. Potter, New York, 362 pp. Woloszyn, B. W., & N. A. Mayo. 1974. Postglacial remains of a vampire bat (Chiroptera: Desmo- dus) from Cuba.— Acta Zoologica Cracoviensia 19:253-266. (GSM) Department of Natural Sciences, Florida State Museum, University of Flor- ida, Gainesville, Florida 32611; (OJL) De- partamento de Estudios Ambientales, Di- vision de Ciencias Biologicas, Universidad Simon Bolivar, Caracas, Venezuela; (CER) Department of Paleobiology, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 929-951 STUDIES IN THE LEPIDAPLOA COMPLEX (VERNONIEAE: ASTERACEAE) IV. THE NEW GENUS, LESSINGIANTHUS Harold Robinson Abstract. —The genus Lessingianthus is described as new for a series of 101 neotropical species typified by Vernonia buddleiifolia. The typical subgenus is characterized by pollen with type B areolation and has a few species with type D pollen. A second subgenus, Oligocephalus, is established for three species having type C pollen areolation. The genus contains some of the species pre- viously placed in the Vernonia series Buddletifoliae, Brevifoliae, Macrolepidae, Remotiflorae, and Flexuosae. The members of the new genus have consistently lophate pollen, as do members of the related genus Lepidaploa, but the new genus lacks the rhizomatous condition of the pollen crests, lacks the basal node of the style, lacks glands on the achenes, and often has pedunculate heads. Lessingianthus myrsinites, is described as new, using an apparently unpublished species name of Ekman. The present paper is the fourth in a series of seven devoted to the study of the neo- tropical Lepidaploa complex (Robinson 1987a, b, c, 1988). The element from the broad concept of Vernonia treated herein is the largest in the series other than Lepida- ploa itself. My interest in the group first developed during a study of style bases in the Vernonieae, which demonstrated that the members of the present group lacked a node. Interest was increased by the reali- zation that many species lacking nodes were of a group referred to in prior treatments of Vernonia as the section Lepidaploae Mac- rocephalae (Baker 1873). The fact that most species of the group have distinctive, large, lophate pollen (Jones 1979b), and the recent discovery of non-rhizomatous crests on the pollen grains has led to the conclusion that the group should be recognized as a new genus, herein named Lessingianthus. Prior taxonomic treatments of Vernonia sensu lato have provided very imperfect ap- proximations of the group treated here as Lessingianthus. Furthermore, the species were always included within the also im- perfectly defined Lepidaploa subgroup of Vernonia. Both the sections Lepidaploae Macrocephalae of Baker (1873) and the se- ries Buddleiifoliae of Jones (1982) were rec- ognized on the basis of the relatively large size of the heads in many of the species of the group. Although large heads remained a prime distinction of the group for Jones (1982), the 22 species included in his com- paratively refined treatment did not all have large heads. Subsequently, relationships have been traced to species included by Jones in his series Brevifoliae, Macrolepi- dae, Remotiflorae, and Flexuosae. In most of these taxonomic series, the members of Lessingianthus were intermixed with species placed in this series of studies in the related genus Lepidaploa. It is the accurate delim- itation of Lessingianthus from Lepidaploa that is the primary aim of the following dis- cussion. Four characters are most instruc- tive in this: the presence or absence of a basal stylar node, the pollen structure, the form of the inflorescence, and the form of the involucre. Each of these will be dis- cussed separately. 930 Basal Stylar Node The presence of basal stylar nodes has been noted in various Asteraceae since at least the work of Cassini (1818), and they have been found taxonomically useful in some groups such as the Eupatorieae (Rob- inson & King 1977, King & Robinson 1987). Nodes seem to have a function in species of the tribe Heliantheae where they expand belatedly and tear the corolla base loose from the achene. In other tribes such as the Eu- patorieae and Vernonieae, however, there is no function sufficiently important to pre- vent variability within the groups. In the Vernonieae, the stylar node ap- pears basic for the tribe, but seems to differ in commonness in the Eastern and Western Hemispheres. A highly developed node, such as that in the African-Indian Ocean genus Distephanus (Robinson & Kahn 1986), is otherwise found in comparatively few members of the tribe in the Eastern Hemisphere, where most Vernonieae lack nodes. In the Western Hemisphere, how- ever, a stylar node is much more widely distributed and its absences appear to be more significant. The distribution of the character has been surveyed and has yielded some interesting results. In the Western Hemisphere, the stylar node is lacking or poorly differentiated in a few small groups such as Lepidonia and Stramentopappus (Robinson & Funk 1987) in the Leiboldia group, and in Stenoceph- alum and Chrysolaena in the Lepidaploa group (Robinson 1987a, 1988), but it is well-developed in the genus Lepidaploa it- self. The node is present in most other ele- ments of Vernonia s.1., but is again lacking in a series of species traditionally placed in the genera Eremanthus and Lychnophora of the Lychnophorinae. In the latter group, the lack of a basal node correlates with the rath- er unique variation from the common cy- mose maturation pattern of the heads (Rob- inson 1980a). All these groups give the impression of many separate losses of a bas- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON al stylar node. Each loss seems correlated with other characters that favor recognition of the groups at generic or higher levels. The essential lack of a basal stylar node in Lessingianthus contrasts strongly with the rather marked structure seen in almost all members of the related Lepidaploa. In the species of Lessingianthus, there is no en- largement whatsoever, and there may or may not be a sclerified ring. The sclerified ring may appear enlarged when the softer upper tissue has collapsed. Some species included here, such as Vernonia regis and V. brevi- folia, seem to have a slight but distinct en- largement at the base of the style, but fully soaked material of styles shows the lower shaft equals the sclerified ring in width. The ring differs from that of Lepidaploa by its slight angle and the wide area of attachment that remains inside the ring. In Lepidaploa the sclerified surface cuts under the base of the style and leaves a very narrow area of attachment. Immaturity of style bases may lead to misinterpretation, but mistakes in obser- vation are usually caused by failure to ex- tract the complete style. The typical Lessin- gianthus style base closely resembles a broken Lepidaploa style base. It should be noted that Lessingianthus style bases may be distinguishable even from broken bases of Lepidaploa because they are often larger. Pollen Two aspects of the pollen of Lessingian- thus are significant for taxonomic discrim- ination. The pollen in both Lessingianthus and Lepidaploa is lophate with crests form- ing regular patterns. Regular patterns occur widely in both Eastern and Western Hemi- sphere Vernonieae, and they are regarded here as primitive in the tribe. The non-lo- phate types are regarded as reversions. The lophate types in both hemispheres show the perforated tectum restricted to the crests (Figs. 1-16) or completely lacking. The crests of the pollen of Eastern Hemisphere species VOLUME 101, NUMBER 4 and of genera such as Stokesia, Mattfeldan- thus, and Lessingianthus from the Western Hemisphere have stout, high baculae sep- arate from the point of attachment on the footlayer to the point above where they join the ridge of the crest (Figs. 3, 7, 9, 10, 15). This condition appears ancestral. The crests in the related Lepidaploa show a condition previously described as “‘rhizomatous’”’ (Robinson 1987a, b, c), where the baculae under the crests are joined into a transverse basal rhizome and where the attachment of the rhizome to the footlayer is compara- tively weak. The drawing by Stix (1960) of her general Vernonia type pollen seems to show this rhizomatous structure. If so, her inclusion of the Vernonia argyrophylla type pollen under this category was an error. The “rhizomatous” type of crest has a tendency to peel away from the pollen grain rather easily, a trait that can best be seen under SEM but which can also be seen under the light microscope. The Lepidaploa-type rhi- zomatous crest is reviewed more fully in the treatment of Lepidaploa. Only a few Bras- ilian species in Lepidaploa, mostly those having type D pollen, have an intermediate development of the rhizomatous character. Pollen with rhizomatous crests must be re- garded as a restricted type within the tribe, which probably evolved only once. The pre- cise form is known only in the Neotropical members of the tribe. There is reason to believe that all cther genera characterized by such pollen, e.g., Stenocephalum (Rob- inson 1987a), and Echinocoryne (Robinson 1987b), are related directly to Lepidaploa, and one other occurrence of such pollen in Eirmocephala may be derived from an in- tergeneric hybridization involving Lepida- ploa (Robinson 1987c). Because of the close relationship among the groups with such pollen, it is possible to conclude that the rather closely related Lessingianthus de- parts phyletically from a point slightly be- low the origin of the “‘rhizomatous’”’ apo- morphy. The pollen of Lessingianthus could be seen as being of a more ancestral type at 931 the same time the style base is of a more derived type. It is possible that the crest structure has undergone reversals in the evolution of the group, but everything ob- served could be explained without such re- versals. The precise form of the lophate pattern of the pollen also distinguishes almost all species of Lessingianthus from any Lepi- daploa. The Vernonia argyrophylla-type pollen described by Stix (1960), also called type B pollen by Jones (1979b), in its typical form appears to be restricted to Lessingian- thus and is completely absent from Lepi- daploa. Such pollen has colpar areolae ex- tending the whole length of the grain and meeting at the poles at the same time that the intercolpi have three areolae across at the equator (Figs. 1, 2, 5, 6, 8). The three areolae across the intercolpar equator may be part of a 1:2:3:2:1 pattern (Fig. 1) or a 1:2:1:2:1 pattern (Fig. 6). The type B grains tend to have diameters greater than the oth- er lophate types in the Lepidaploa relation- ship. They are mostly 50-80 um in diameter in fluid. Most Lepidaploa have pollen grains 40-55 um in diameter, rarely to 60 um. The larger grains of Lessingianthus tend to occur in the species with the most robust flowers and heads. All neotropical species seen with Vernonia argyrophylla or type B grains are recognized here as members of Lessingian- thus. A number of the species of Lessin- gianthus were included by Jones (1979b) under series Macrolepidae Benth. & Hook. with the pollen type listed as A, but the latter seems to have been a misprint. The Vernonia argyrophylla or type B pol- len has been adequately distinguished by both Stix (1960) and Jones (1979b) from other types of lophate grains found in Ver- nonia that have separate areolae at the pol- len poles or that have cross-walls across the colpar area above and below the pore. How- ever, distinction has not been ordinarily made between the larger type B pollen with three areolae across the intercolpar region and the generally smaller forms with only 932 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 14. Pollen of Lessingianthus argyrophyllus (Less.) H. Robins. 1. Intercolpar view showing 1:2:3:2:1 pattern. 2. Polar view showing the three colpi meeting at the pole. 3. Broken grain showing baculae. 4. Broken grain showing detail under perforated tectum. two areolae across the intercolpar region. The latter type has been recognized as the V. geminata-type by Robinson (1980b). Present observations show that the distinc- tion is more than technical. All species with V. geminata-type pollen prove to be mem- bers of the genus Lepidaploa or Echinoco- ryne. They are more closely related to species with type C and D pollens than to the Les- singianthus species with regular type B pol- len. The V. geminata type grains all have rhizomatous crests, which the typical type B grains do not have. Pollen with three ar- eolae across in many of its intercolpar re- gions has been seen in Lepidaploa in V. vio- liceps H. Robins. from Ecuador and in a specimen of an unnamed Vernonia received from Badillo in Venezuela, and in some in- tercolpar regions of such species as V. her- bacea, but the pollen involved is not true type B. The grains have rhizomatous crests and polar areolae that clearly indicate the identity of the species as Lepidaploa or Chrysolaena. The only pollen in a Lepidaploa that might qualify as a true type B is in V. psilostachya VOLUME 101, NUMBER 4 a 3. Aum 933 Figs. 5-8. Pollen of Lessingianthus. 5-7. L. tomentellus (Mart. ex DC.) H. Robins. 5. Polar view showing three colpi meeting at pole. 6. Intercolpar view showing 1:2:1:2:1 pattern. 7. Detail of pollen crest showing baculae. 8. L. cephalotes (DC.) H. Robins., polar view showing three colpi meeting at pole. DC. The latter has colpi reaching the poles and a variable number of 2 or 3 areolae across the intercolpus in | or 2 tiers. Careful examination shows that very few of the grains actually match true type B in all the intercolpi simultaneously. The habit of the plants is small compared to any Lessin- gianthus, and the involucre is a Lepidaploa 934 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SURAT ye ms & ee Eé Figs. 9-12. Pollen of Lessingianthus. 9. L. laurifolius (DC.) H. Robins., broken grain showing baculae. 10. L. niederleinii (Hieron.) H. Robins., broken grain showing baculae. 11, 12. L. brevifolius (Less.) H. Robins. 11. View with pole below middle showing three areolae above pores meeting at pole. 12. Colpar view showing crosswalls above and below pore. VOLUME 101, NUMBER 4 935 Tt 14.9um Figs. 13-16. Pollen of Lessingianthus. 13-15. L. simplex (Less.) H. Robins. 13. Polar view showing polar areola. 14. Intercolpar view showing 1:2:2:1 pattern. 15. Detail of pollen crest showing baculae. 16. L. virgulata (Mart. ex DC.) H. Robins., near-colpar view showing truncate areolate poles and 1:2:2:1 intercolpar pattern. 936 type. The approximation of the pollen to type B might incline one to place the species in Lessingianthus, but SEM study shows a clearly rhizomatous structure under the crests. There is one area of overlap in pollen areolation type. Vernonia simplex (Figs. 13- 15) and V. virgulata (Fig. 16) have a type C lophate pattern with polar areolae. The in- tercolpar region usually is only two areolae wide in a 1:2:2:1 pattern (Figs. 14, 16). In the pattern of the areolae the pollen of these species is like the pollen that occurs in such species as V. (Chrysolaena) flexuosa. The pollen in V. simplex and V. virgulata differs from that of Lepidaploa and is like that of Lessingianthus in that the baculae attach directly to the foot-layer without a “rhi- zome.” Previous treatments have associ- ated these Lessingianthus species with two specialized groups now placed in Chryso- laena (Robinson 1988) and Echinocoryne (Robinson 1987b). Vernonia simplex was included by Jones (1981) in his series Flex- uosae in his refined concept of the group. In this latter case, the pubescence of the plants and the lophate pattern of the pollen seemed to re-enforce the relationship be- tween species in the series. However, the pollen distinction seems to correlate rather well with the presence of pedunculate heads in the group. The series Flexuosae, in spite of initial appearances, actually consists of two totally separate elements, one in Les- singianthus and the other, Chrysolaena, having rhizomatous crests on the pollen and glanduliferous anther appendages. The lat- ter group is closer to Lepidaploa. Although its pollen has not been examined with the SEM, V. desertorum, which Jones (1981) also placed in series Flexuosae apparently is a Lessingianthus. Light microscope ob- servations of the crests indicate that they are not rhizomatous. The three pedunculate species with type C pollen, including the two from the series Flexuosae, have grains of ca. 50 wm diameter, in the size range common to both Lessingianthus and Lepidaploa. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Vernonia virgulata was placed by Jones (1979b) in his series Subulatae, which oth- erwise proves to belong to the genus Echi- nocoryne. The Jones (1979b) citation of type B pollen from V. virgulata seemed to cor- relate with the type cited for the remainder of the series, but none of grains involved were actually type B. As stated above, the pollen of V. virgulata is closest to type C; and as indicated by Robinson (1987b), the species of Echinocoryne have pollen of the V. geminata type with only two rows of intercolpar areoles and no polar areolae. Two additional pollen variations occur in Lessingianthus. The pollen of V. brevifolia (Figs. 11, 12) and V. dura have crests of exine aross the colpi above and below the pores. Such crosswalls were characteristic of the Vernonia arenaria-type of Stix (1960) and the type D of Jones (1979b). The V. brevifolia and V. dura pollen differs from type D and resembles other Lessingianthus pollen in the presence of non-rhizomatous crests with high baculae and by the presence of three areolae equatorially across the in- tercolpus. It should be recognized that these type D grains differ from the type B only in the presence of the crosswalls above and below the pores, and the genetic difference may not be very great. Although V. cephalotes normally shows type B pollen (Fig. 8), some specimens have grains with an irregular surface pattern best classified as type A. Type A pollen has been considered a reversion type in the tribe be- cause of the wider distribution of lophate types, but V. cephalotes is the only specific example of such a reversion thus far known in the Western Hemisphere. The grains measure ca. 45 um in fluid, larger than the common type A. The irregular pattern of colpi and spines suggests the influence of irregular meiosis. Another species in which only large type A pollen has been seen, V. glazioviana is included here in the belief it represents a similar reversion from a type A pollen. The size of pollen in Asteraceae often cor- VOLUME 101, NUMBER 4 relates with chromosome number, but the larger size of pollen in many species of Les- singianthus is not particularly correlated with known polyploidy, except to the extent that the Y = 17 basic to Neotropical Ver- nonieae is a polyploidy. Jones (1979a) pro- vided counts for four members of the genus. Two of the species, V. bardanioides, and V. glabrata include higher polyploids having N = 34, but each also has lower counts of N = 17. A similar mixture of lower counts and higher polyploidy is also seen in at least one member of Chrysolaena (as V. cognata in Jones 1979a). Inflorescence Form The general Lepidaploa and Lessingian- thus group is notable for a type of inflores- cence that can best be termed seriate-cy- mose. Lepidaploa itself exemplifies the structure where each head is technically ter- minal with a lateral branch from immedi- ately below the head bearing the rest of the series. The result looks like a series of lateral sessile heads except for the usual slight de- flection at each head. This form is seen in every species presently recognized as a true Lepidaploa, and it seems to derive from ancestors more like the Brasilian Vernonia diffusa Less. that have less regular cymes and are outside of the Lepidaploa complex. Such ancestral stocks seem to have pro- duced some other strongly seriate-cymose forms like those of the neotropical Cyrto- cymura and Eirmocephala (Robinson 1987c) and the Colombian Dipterocypsela Blake. The latter are considered to be ba- sically outside the Lepidaploa complex. All of the other marked examples of such cymes except V. peculiaris Verdcourt of Africa seem to be in Lepidaploa or its close relatives. These include Mattfeldanthus Robins. & King of Brasil that has lophate pollen. The cymes have been retained with smaller, less distinct heads in Stenocephalum (Robinson 1987a). In the closely related Echinoceph- alum the inflorescence is drastically altered 937 by the presence of peduncles, sometimes very long peduncles, under the heads. The latter condition seems derived within the complex. Lessingianthus has many species that show the same seriate-cymose condition noted for Lepidaploa, but includes varia- tions of this form that differ from anything in the latter genus. One variation is the de- velopment of pedunculate heads. This char- acter occurs in members of the genus, such as V. psilophylla and V. secunda that have a closer relationship to species with sessile heads than to each other. Many of the species such as V. sellowii, V. carvalhoi, V. santosii, and V. scaposa have scapose inflorescences from abbreviated vegetative plants, but the species do not form a related group within the genus. The pedunculate and scapose characters represent general trends in Les- singianthus that are lacking in Lepidaploa. The species group with type C pollen, which has been misplaced in the series Subulatae and Flexuosae, is characterized by pedun- culate heads. Vernonia virgulata is not dis- tinguishable from Echinocoryne on the ba- sis of that character, but differs in other features including its pollen. In both the pe- dunculate heads and the non-rhizomatous crests of the pollen, the latter species along with V. simplex and V. desertorum fall eas- ily within Lessingianthus. One other modification in a few species of Lessingianthus was discussed by Rob- inson (1980c) in connection with the de- scription of V. eitenii. Although the small group of species involved has inflorescences closely resembling the seriate-cymes of their relatives, the sequence of maturation of the heads is reversed, so that the uppermost head matures first and the progress is down- ward. This sequence indicates that devel- opmentally the inflorescence has converted to a cyme with the lower heads truly lateral. It is another variation that does not occur in Lepidaploa. The species with seriate cymes in Lessin- gianthus and Lepidaploa are often distin- 938 guishable by an additional feature. In those species having distinct foliose bracts in the cymes, the bracts in Lessingianthus never seem to have the abrupt reduction in size from leaves to bracts that is seen at the base of the cyme in many species of Lepidaploa. Involucral Bracts Lessingianthus is essentially the same as Lepidaploa in the ratio of involucral bracts to flowers in the head. The bracts are usually twice to three times as many as the flowers. Lessingianthus never has excessive num- bers of involucral bracts, as is seen in Echin- ocoryne. In other aspects of the involucre, however, there are characters that help to distinguish the two genera. One of these is the degree of differentiation between the outer and inner bracts. In Lessingianthus and Lepidaploa, the in- volucral bracts are graduated with the inner bracts progressively larger and longer than the outer ones. The bracts in Lessingianthus show only the differences to be expected in such graduated structures. In those where the bracts are narrow and pointed, as in V. bardanioides, or those where they are short, as is V. buddleiifolia and V. argyrophylla, the slender or obtuse tips occur throughout the involucre. In Lepidaploa the outer bracts are often very different in shape from the inner ones. The innermost bracts may often be rather abruptly wider and the outermost bracts are often very aristate and spreading. Such differences are not found in all Lepi- daploa, but they are found in almost no species of Lessingianthus. The condition is approached in Vernonia glabrata and V. neiderleinii which are otherwise like Lessin- gianthus in their rather large habit. A num- ber of species of Lessingianthus, such as V. arachniolepis and V. pseudosquarrosa, have outer bracts with spreading, rather foliose tips, a feature not noted in Lepidaploa. The tips of the involucral bracts in Les- singianthus are often very short in an almost reptilian scale pattern. Such a pattern is characteristic of the typical element of the genus characterized by larger heads, but it PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON also occurs in some smaller-headed species such as V. psilophylla. The narrowly point- ed bracts of V. bardanioides, V. eitenii, V. irwinii, and their relatives taper evenly to the tips and have a distinct longitudinal me- dian costa externally of a type not seen in Lepidaploa. They do not have the apical mucros or aristae seen in many Lepidaploa species. The latter short apical mucros oc- cur in Lessingianthus in only a few species such as V. elegans. The involucral bracts of most species of Lessingianthus are imbricate in many se- ries, but some species such as V. carvalhoi and V. brevifolia have involucres that ap- pear less imbricate because of the narrower bracts. The most significant examples of re- duced involucres in Lessingianthus are in the three species, V. virgulata, V. simplex, and V. desertorum that have been noted above as having type C pollen. These species are treated below as part of a distinct sub- genus within Lessingianthus. A number of other characters show vari- ations of interest in Lessingianthus al- though they are not as useful for delimiting the genus. In many species of Lessingian- thus the tendency for pedunculate heads seems correlated to some extent with the occurrence of distinct petioles. Neverthe- less, shortly petiolate and sessile leaves oc- cur in both genera. The corolla lobes of Les- singianthus show a broad fusion and enlargement of the vascular strands at the tip, a feature that seems rather characteristic of most of the Lepidaploa complex. The vascular strands of the corolla lobes of most other American Vernonieae are only nar- rowly fused or separated at the tips. The inner pappus in both genera is unfailingly capillary, but in Lessingianthus the bristles are comparatively short in a number of species such as V. eitenii and V. barda- nioides. The pappus in these species is shorter in relation to the involucre and co- rolla, and the corolla throat is mostly ex- posed. Short pappus bristles of this type do not occur in Lepidaploa. The achenes in the genus are normally densely setuliferous, but VOLUME 101, NUMBER 4 a few species such as V. pseudopiptocarpha and V. ixiamensis have glabrous achenes, whereas V. glabrata, V. saltensis, and V. westermanii have achenes with few setulae. The density of setulae on the achenes of most species allows little space for glands, and unlike many Lepidaploa species, no glands have been seen in the genus. A few species such as V. ammophila and V. pyc- nostachya have resin cells on the surface of the achene as single cells or small groups of two or three cells. The raphides in the achene wall are usually elongate, but are quadrate in some species such as V. regis, V. pseu- dopiptocarpha, and V. ammophila. In one species of Lessingianthus, V. san- tosii Robinson (1980b), the flowers in the heads have been noted as opening in two sets, the outer rows first and then the inner disk flowers later as a separate group. The initial phase offers the superficial appear- ance of a head with ray flowers. The full distribution of the character is not known, but it has been seen again in the compara- tively unrelated Vernonian genus Centra- therum, where the two stages occur on sep- arate days. The character is most obvious in the field, and studies in Brazil might show how common the character is in Lessin- gianthus. The character would obviously not be found in Vernonieae with smaller, fewer- flowered heads such as Stenocephalum. The two generic concepts Lessingianthus and Lepidaploa represent two elements with basically differing aspects and different ge- ography. The most obvious reason for rec- ognizing two genera is the form of the in- florescence, which in Lessingianthus frequently has pedicellate heads and in Lep- idaploa has all heads essentially sessile in the seriate cymes. It is the form of the pollen that seems to correlate best phyletically with the two groups, the one with many pedun- culate heads and mostly undifferentiated in- volucres, and the second with exclusively sessile heads and mostly differentiated in- volucres. The generic distinction is there- fore placed here technically at the point of the difference between non-rhizomatous 939 pollen crests with large discrete baculae ver- sus the mostly rhizomatous pollen. Every species of Lessingianthus can be assigned either by its pedunculate heads or by its type B or D pollen having no polar areolae in conjunction with 1:2:3:2:1 or 1:2:1:2:1 in- tercolpar patterns. This distinction creates problems in only a few species. One species falling into Lessingianthus on the basis of non-rhizomatous type D pollen having a 1:2:1:2:1 intercolpar pattern seems uncom- fortable in the genus on the basis of habit. Vernonia regis is most unusual in the genus by the rather prominent ring at the base of the style. Collapse of the softer shaft leaves a more marked enlargement than in any other Lessingianthus. Nevertheless, the structure is not truly broader than the shaft and it does not show the narrow attachment commonly seen in Lepidaploa. The species is further distinctive in its crowded narrow heads with distinct herbaceous outer bracts. Vernonia regis does not look like a Lepi- daploa but rather like a Stenocephalum. The differences from Stenocephalum are dis- cussed by Robinson (1987a). This particu- lar species may reflect some area of evolu- tionary complexity such as hybridization not yet resolved, but for the present the Type D non-rhizomatous pollen character is ap- plied rigorously to include it within Lessin- gianthus. In the present view, Lessingianthus may be circumscribed too broadly in some cases, but the genus is not too narrowly delimited. The new genus is named in honor of Carl F. Lessing 1809-1862 who was the author of many of the species of the genus. Lessingianthus, gen. nov. Plantae herbaceae perennes plerumque erectae ad 1.5—2.0 m altae in caulibus foltis pedunculis et bracteis involucri variabiliter pubescentes. Folia alterna breviter petiolata vel sessilia. Inflorescentiae simplices vel cy- mosae saepe scorpioideo-cymosae vel se- riate subcapitulate proliferatae vel rare stricte cymosae, pedunculis nullis vel var- iabiliter elongatis. Capitula late plerumque 940 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON campanulata; bracteae involucri 2-3-plo plus quam flores in capitulo subimbricatae plerumque multiseriatae plerumque late oblongae et obtusae interdum lineari-lan- ceolatae raro leniter imbricatae. Flores in capitulo 15—50; corollae plerumque lavan- dulae saepe in tubis et faucibus glabris in lobis distaliter plerumque minute glandu- liferae vel setuliferae apice in nervis con- junctae et incrassatae; appendices anthera- rum non glanduliferae; basi stylorum non noduliferi raro distincte annuliferi. Achaen- ia prismatica plerumque 5-costata plerum- que dense sericeo-setulifera non glanduli- fera raro cellulis resiniferis sparse obtecta; carpopodia cylindrica vel obturaculiformia et turbinata, cellulis plerumque oblongis in parietibus striate porulosis; setae pappi in- teriores capillares longitudinaliter ad corol- las subaequales vel multo breviores, serie- bus exterioribus longe squamiformibus. Grana pollinis (45—)50—75(—80) wm in dia- metro lophata in cristis non rhizomata, col- pis ad polos attingentibus, areolaris inter- colpis aequatorialiter triplicibus (typus B), raro grana ad polos areolata et in areolaris intercolpis aequatorialiter duplicibus (typus C). Type. — Vernonia argyrophylla Less. The genus is geographically mostly concentrated in Brasil with a comparatively few species occurring westward in Argentina, Paraguay, and Bolivia. The genus has two represen- tatives in Peru, only one, L. rubricaulis, as far north as Colombia, and only L. morilloi in southern Venezuela. The following 101 species are recognized in the genus. Lessingianthus adenophyllus (Mart. ex DC.) H. Robinson, comb. nov. Vernonia adenophylla Mart. ex DC., Prodr. 5:17. 1836. Brasil (Parana). Lessingianthus ammophilus (Gardn.) H. Robinson, comb. nov. Vernonia ammophila Gardn., Lond. J. Bot. 5:227. 1846. Cacalia ammophila (Gardn.) Kuntze, Rev. Gen. Pl. 2:969. 1891. Brasil (Goias, Minas Gerais). Lessingianthus arachniolepis (Ekman & Dusen) H. Robinson, comb. nov. Vernonia arachniolepis Ekman & Dusen in Malme, Kungl. Svenska Vetenskacade- miens Handlingar 12(2):17. 1933. Brasil (Parana, Rio Grande do Sul). Lessingianthus argenteus (Less.) H. Robinson, comb. nov. — Vernonia argentea Less., Linnaea 6:672. 1831. Cacalia argentea (Less.) Kuntze, Rev. Gen. Pl. 2:969. 1891. Brasil (Parana, Sao Paulo). Lessingianthus argyrophyllus (Less.) H. Robinson, comb. nov. (Figs. 1—4) Vernonia argyrophylla Less., Linnaea 6:627. 1831. Cacalia argyrophylla_ (Less.) Kuntze, Rev. Gen. Pl. 2:969. 1891. Brasil (D.F., Goias, Minas Gerais). Lessingianthus asteriflorus (Mart. ex DC.) H. Robinson, comb. nov. Vernonia asteriflora Mart. ex DC., Prodr. 5:29. 1836. Cacalia asteriflora (Mart. ex DC.) Kuntze, Rev. Gen. Pl. 2:969. 1891. Vernonia kuntzei Hieron., Engl. Bot. Jahrb. 22:678. 1897. Vernonia kuntzei (Hieron.) Kuntze, Rev. Gen. Pl. 3:138. 1898. Bolivia, Brasil (Parana, Sao Paulo, Santa Catarina). Lessingianthus bardanioides (Less.) H. Robinson, comb. nov. Vernonia bardanioides Less., Linnaea 6:669. 1831. Vernonia reticulata Gardn., Lond. J. Bot. 5:226. 1846. Vernonia lappoides Baker in Mart., Fl. Bras. 6(2):35. 1873. Vernonia cirsiiflora Mart. ex Baker in Mart., Fl. Bras. 6(2):36. 1873, nomen nu- dum Cacalia bardanioides (Less.) Kuntze, Rev. Gen. Pl. 2:969. 1891. Cacalia lap- VOLUME 101, NUMBER 4 poides (Baker) Kuntze, Rev. Gen. PI. 2: 970. 1891. Brasil (Bahia, D.F., Goias, Mato Grosso do Sul, Minas Gerais, Sao Paulo). Lessingianthus bishopii (H. Robinson) H. Robinson, comb. nov. Vernonia bishopii H. Robinson, Phytologia 49:261. 1981. Vernonia flavescens Glaz., Bull. Soc. Bot. France Mem. 3, 56:369. 1909, nomen nudum, non V. flavescens Less. Vernonia goiasensis S. B. Jones, Brittonia 34:107. 1982, nomen nudum Brasil (Goias, Mato Grosso do Sul). Lessingianthus brevifolius (Less.) H. Robinson, comb. nov. (Figs. 11, 12) Vernonia brevifolia Less., Linnaea 4:285. 1829; 6:659. 1831. Vernonia ericaefolia Hook. & Arnott, Comp. Bot. Mag. 1:236. 1835. Cacalia brevifolia (Less.) Kuntze, Rev. Gen. Pl. 2:969. 1891. Vernonia li- nosyrifolia Chod., Bull. Herb. Boiss., ser. 2, 2:300. 1902. Argentina, Brasil (Parana, Rio Grande do Sul, Sao Paulo, Santa Ca- tarina), Paraguay. Lessingianthus brevipetiolatus (Sch.Bip. ex Baker) H. Robinson, comb. nov. Vernonia brevipetiolata Sch.Bip. ex Baker in Mart., Fl. Bras. 6(2):85. 1873. Cacalia brevipetiolata (Sch. Bip. ex Baker) Kuntze, Rev. Gen. Pl. 2:969. 1891. Brasil (Minas Gerais). Lessingianthus buddleiifolius (Mart. ex DC.) H. Robinson, comb. nov. Vernonia buddleiifolia Mart. ex DC., Prodr. 5:45. 1836, as “Buddleiaefolia.”’ Vernon- la squamosa Gardner, Lond. J. Bot. 6: 419. 1847. Cacalia buddleiaefolia (Mart. ex DC) Kuntze, Rev. Gen. Pl. 2:969. 1891. Brasil (D.F., Goias, Mato Grosso, Minas Gerais, Sao Paulo). Lessingianthus bupleurifolius (DC.) H. Robinson, comb. nov. Vernonia laevigata var. bupleurifolia DC., Prodr. 5:56. 1836. Vernonia pupleurifolia (DC.) Sch.Bip. ex Malme, Arkiv. Bot. 24A(8):13. 1932. Brasil (Goias, Mato Grosso). Near L. obtusatus, see Malme (1932). Lessingianthus caiapoensis (H. Robinson) H. Robinson, comb. nov. Vernonia caiapoensis H. Robinson, Phy- tologia 45:171. 1980. Brasil (Goias). Lessingianthus carduoides (Baker) H. Robinson, comb. nov. Vernonia carduoides Baker in Mart., FI. Bras. 6(2):34. 1873. Cacalia carduoides (Baker) Kuntze, Rev. Gen. Pl. 2:969. 1891. Brasil (Minas Gerais). Lessingianthus carvalhoi (H. Robinson) H. Robinson, comb. nov. Vernonia carvalhoi H. Robinson, Phytolo- gia 53:394. 1983. Brasil (Bahia). Lessingianthus cataractarum (Hieron.) H. Robinson, comb. nov. Vernonia cataractarum Hieron., Bot. Jahrb. Syst. 22:681. 1897. Brasil (Santa Catari- na). Lessingianthus cephalotes (DC.) H. Robinson, comb. nov. (Fig. 8) Vernonia cephalotes DC., Prodr. 5:57. 1836. Brasil (Goias, Minas Gerais, Sao Paulo). DeCandolle (1836) in his original de- scription cited Chrysocoma oligophylla Vell. as a questionable synonym. Kuntze (1891) synonymized the two. The Vellozo descrip- tion and plate (1825, 1835) indicate a pilose plant with much more pointed, graduated involucral bracts. The Vellozo species 1s ev- idently not L. cephalotes but the habitally 942 very similar V. hypochlora Malme, which is a Chrysolaena. Lessingianthus chamaepeuces (Sch.Bip. ex Baker) H. Robinson, comb. nov. Vernonia chamaepeuces Sch.Bip. ex Baker in Mart., Fl. Bras. 6(2):31. 1873. Cacalia chamaepeuces (Sch.Bip. ex Baker) Kuntze, Rev. Gen. Pl. 2:969. 1891. Brasil (Goias, Mato Grosso). Lessingianthus compactiflorus (Mart. ex Baker) H. Robinson, comb. nov. Vernonia compactiflora Mart. ex Baker in Mart., Fl Bras. 6(2):44. 1873. Cacalia compactiflora (Mart. ex Baker) Kuntze, Rev. Gen. Pl. 2:969. 1891. Brasil (D.F., Goias, Minas Gerais). Lessingianthus cordiger (Mart. ex DC.) H. Robinson, comb. nov. Vernonia cordigera Mart. ex DC., Prodr. 5: 58. 1836. Cacalia cordigera (Mart. ex DC.) Kuntze, Rev. Gen. PI. 2:969. 1891. Brasil (Minas Gerais). Lessingianthus coriaceus (Less.) H. Robinson, comb. nov. Vernonia coriacea Less., Linnaea 6:661. 1831. Vernonia hecatantha DC., Prodr. 5:53. 1836 (as hexacantha, corr. 5:696. 1836). Cacalia coriacea (Less.) Kuntze, Rev. Gen. Pl. 2:969. 1891. Bolivia, Brasil (Bahia, D.F., Goias, Maranhao, Mato Grosso, Minas Gerais, Sao Paulo). Peru. Lessingianthus cristalinae (H. Robinson) H. Robinson, comb. nov. Vernonia cristalinae H. Robinson, Phyto- logia 45: 172. 1980. Brasil (Goias). Lessingianthus dorsiventralis (Chodat) H. Robinson, comb. nov. Vernonia dorsiventralis Chodat, Bull. Herb. Boiss. ser. 2, 1:410. 1901. Paraguay. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lessingianthus durus (Mart. ex DC.) H. Robinson, comb. nov. Vernonia dura Mart. ex DC., Prodr. 5:59. 1836. Cacalia dura (Mart. ex DC.) Kuntze, Rev. Gen. Pl. 2:970.1891. Ver- nonia macedoi Barroso, Arq. Jard. Bot. Rio de Janeiro 13:9. 1954. Brasil (D.F., Goias, Mato Grosso, Minas Gerais). Lessingianthus eitenii (H. Robinson) H. Robinson, comb. nov. Vernonia eitenii H. Robinson, Phytologia 46:109. 1980. Vernonia wasshausenii S. B. Jones, Brittonia 34:110. 1982. Brasil (D.F., Goias). The species concept includes one para- type of L. cristalinae (Irwin et al. 32817). Specimens with only a few apical heads can be distinguished from L. cristalinae most easily by the less appressed and more yel- lowish pubescence of the stems. Lessingianthus elegans (Gardn.) H. Robinson, comb. nov. Vernonia elegans Gardn., Lond. J. Bot. 6: 421. 1847. Cacalia elegans (Gardn.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (D.F., Goias, Mato Grosso, Minas Ger- ais, Sao Paulo). Lessingianthus erythrophilus (DC.) H. Robinson, comb. nov. Vernonia erythrophila DC., Prodr. 5:56. 1836. Cacalia erythrophila (DC.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (D.F., Minas Gerais). Lessingianthus exiguus (Cabrera) H. Robinson, comb. nov. Vernonia exigua Cabrera, Sellowia 13:166. 1961. Brasil (Parana, Santa Catarina, Sao Paulo). VOLUME 101, NUMBER 4 Lessingianthus farinosus (Baker) H. Robinson, comb. nov. Vernonia farinosa Baker. in Mart., Fl. Bras. 6(2):84. 1873. Cacalia farinosa (Baker) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (Bahia). Lessingianthus floccosus (Gardn.) H. Robinson, comb. nov. Vernonia floccosa Gardn., Lond. J. Bot. 5: 225. 1846. Cacalia floccosa (Gardn.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Ver- nonia robusta Glaz., Bull. Soc. Bot. France Mem. 3, 56:369. 1909, nomen nudum. Brasil (D.F., Goias, Minas Gerais). Lessingianthus fonsecae (H. Robinson) H. Robinson, comb. nov. Vernonia fonsecae H. Robinson, Phytologia 45:174. 1980. Brasil (Goias). Lessingianthus glabratus (Less.) H. Robinson, comb. nov. Vernonia glabrata Less., Linnaea 4:294. 1829; 6:661. 1831. Vernonia radula Mart. ex DC., Prodr. 5:52. 1836. Cacalia gla- brata (Less.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Cacalia radula (Mart. ex DC.) Kuntze, Rev. Gen. Pl. 2:971. 1891. Ver- nonia oxydonta Malme, Ark. Bot. 24A(6): 19. 1932. Brasil (D.F., Goias, Mato Gros- so, Minas Gerais, Parana, Rio Grande do Sul). Lessingianthus glaziovianus (Baker) H. Robinson, comb. nov. Vernonia glazioviana Baker in Mart., FI. Bras. 6(2):41. 1873. Cacalia glaziouana (Baker) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (Rio de Janeiro). Lessingianthus glomeratus (Baker ex Warming) H. Robinson, comb. nov. Vernonia glomerata Baker ex Warming, V1- 943 densk. Medd. Forh. Kjob. 1890:185. 1890. Brasil (D.F., Minas Gerais). Lessingianthus grandiflorus (Less.) H. Robinson, comb. nov. Vernonia grandiflora Less., Linnaea 6:660. 1831. Brasil (Minas Gerais, Parana, Sao Paulo), Paraguay. Kuntze (1891) equated this species with Chrysocoma pumilla of Vellozo (1825), but the plate of the latter (1835) is interpreted here as representing the species commonly known as Vernonia sessilifolia Less. Lessingianthus grearii (H. Robinson) H. Robinson, comb. nov. Vernonia grearii H. Robinson, Phytologia 45:175. 1980. Brasil (Goias). Lessingianthus heringeri (H. Robinson) H. Robinson, comb. nov. Vernonia heringeri H. Robinson, Phytolo- gia 53:395. 1983. Brasil (D.F., Goias). Lessingianthus hoveaefolius (Gardn.) H. Robinson, comb. nov. Vernonia hoveaefolia Gardn., Lond. J. Bot. 6:423. 1847. Cacalia hoveaefolia (Gardn.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Near L. obtusatus. Brasil (Goias, Minas Ger- ais). Lessingianthus hypochaeris (DC.) H. Robinson, comb. nov. Vernonia hypochaeris DC., Prodr. 5:45. 1836. Cacalia hypochaeris (DC.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (Par- ana). Lessingianthus irwinii (Barroso) H. Robinson, comb. nov. Vernonia irwinii Barroso, Loefgrenia 36:2. 1969. Brasil (D.F.). 944 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lessingianthus ixiamensis (Rusby) H. Robinson, comb. nov. Vernonia ixiamensis Rusby, Bull. New York Bot. Gard. 8:125. 1912. Bolivia. Lessingianthus lacunosus (Mart. ex DC.) H. Robinson, comb. nov. Vernonia lacunosa Mart. ex DC., Prodr. 5: 56. 1836. Cacalia lacunosa (Mart. ex DC.) Kuntze, Rev. Gen. PI. 2:970. 1891. Brasil (D.F., Goias, Minas Gerais). Lessingianthus laevigatus (Mart. ex DC.) H. Robinson, comb. nov. Vernonia laevigata Mart. ex DC., Prodr. 5: 56. 1836. Near L. obtusata; see Malme (1932). Brasil (D.F., Goias, Mato Grosso, Minas Gerais, Parana). Lessingianthus laurifolius (DC.) H. Robinson, comb. nov. (Fig. 9) Vernonia laurifolia DC., Prodr. 5:30. 1836. Cacalia laurifolia (DC.) Kuntze, Rev. Gen. PI. 2:970. 1891. Bolivia, Brasil, Peru. Lessingianthus ligulifolius (Mart. ex DC.) H. Robinson, comb. nov. Vernonia ligulifolia Mart. ex DC., Prodr. 46. 1836. as “‘ligulaefolia.”’ Cacalia lig- ulifolia (Mart. ex DC.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Vernonia cotaniensis Hi- eron., Bot. Jahrb, Syst. 40:352. 1908. Bra- sil (D.F., Goias, Sao Paulo). The species is close to L. coriaceus. Lessingianthus linearifolius (Less.) H. Robinson, comb. nov. Vernonia linearifolia Less., Linnaea 4:287. 1829. Cacalia linearifolia (Less.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (Minas Gerais). Lessingianthus linearis (Spreng.) H. Robinson, comb. nov. Vernonia linearis Spreng., Syst. Veg., ed. 16, 3:437. 1826. Cacalia linearis (Spreng.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (Bahia, D.F., Goias, Minas Gerais). Lessingianthus lorentzii (Hieron.) H. Robinson, comb. nov. Vernonia lorentzii Hieron., Engl. Bot. J ahrb. 22:674. 1898. Argentina, Paraguay. Lessingianthus macrocephalus (Less.) H. Robinson, comb. nov. Vernonia macrocephala Less., Linnaea 4: 298. 1829. Cacalia macrocephala (Less.) Kuntze, Rev. Gen. PI. 2:970. 1891. Brasil (Rio Grande do Sul, Sao Paulo), Uruguay. Lessingianthus macrophyllus (Less.) H. Robinson, comb. nov. Vernonia macrophylla Less., Linnaea 6:668. 1831. Cacalia macrophylla (Less.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (Bahia, Espirito Santo, Minas Gerais, Rio de Janeiro). Lessingianthus mansoanus (Baker) H. Robinson, comb. nov. Vernonia mansoana Baker in Mart., Fl. Bras. 6(2):84. 1873. Cacalia mansoana (Baker) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (Goias, Mato Grosso). Lessingianthus mollissimus (D. Don ex Hook. & Arn.) H. Robinson, comb. nov. Vernonia mollissima D. Don ex Hook. & Arn., Comp. Bot. Mag. 1:237. 1835. Ca- calia mollissima (D. Don ex Hook. & Arn.) Kuntze, Rev. Gen. Pl. 2:970. 1891. VOLUME 101, NUMBER 4 Argentina, Brasil (Mato Grosso, Parana, Rio Grande do Sul), Paraguay. Lessingianthus monocephalus (Gardn.) H. Robinson, comb. nov. Vernonia monocephala Gardn., Lond. J. Bot. 6:418. 1847. Cacalia monocephala (Gardn.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (Bahia, D.F., Goias). Lessingianthus morii (H. Robinson) H. Robinson, comb. nov. Vernonia morii H. Robinson, Phytologia 44: 290. 1979. Brasil (Bahia). Lessingianthus morilloi (Badillo) H. Robinson, comb. nov. Vernonia morilloi Badillo, Ernstia 1:2. 1981. Venezuela (Amazonas). Lessingianthus myrsinites H. Robinson, sp. nov. Vernonia myrsinites Ekman, in herb. Plantae fruticosae ad 0.5—1.0 m altae su- perne ramosae dense foliosae. Caules su- perne lanosi base xylopodiales. Folia ses- silia oblonga 2—7 cm longae et 1.0-3.5 cm latae base truncatae vel breviter cordatae margine integrae apice obtusae vel breviter acutae supra subglabrae et sublucidae sub- tus pilulosae et glandulo-punctatae utrinque in nervulis prominentes, nervis secundariis 3-8 erecto-patentibus. Inflorescentiae in ramis abrupte terminales fasciculatae ple- rumque ca. 3-capitatae; capitula late cylin- drica ca. 12 mm longa et 5 mm lata; squa- mae involucri subimbricatae appressae 36— 40 gradatim ca. 6-seriatae, bracteae exte- riores late ovatae 2-3 mm longae apice gla- brae breviter acutae et minute apiculatae extus plerumque dense pilosulae, bracteae interiores oblongae 4—7 mm longae apice roundatae vel saepe eroso-fimbriatae extus glabrae; corollae lavandulae vel purpureae ca. 11 mm longae plerumque glabrae, tubis 945 ca. 5 mm longis, faucibus ca. 1 mm longis, lobis ca. 5 mm longis et 0.6 mm latis apice minute scabridulis extus superne pauce glanduliferis; thecae antherarum ca. 3 mm longae, appendices antherarum ca. 0.7 mm longae et base 2.3 mm latae. Achaenia ca. 3 mm longa sericeo-setulifera; setae pappi interiores ca. 30 ca. 7 mm longae apice len- iter incrassatae, squamae exteriores lineares ca. 1.2 mm longae. Grana pollinis in dia- metro ca. 60 um distincte lophata (typus B). Type. — Brasil: Goias: Chapada dos Vead- eiros. 18-19 km N of Alto Paraiso. Wet campo at 4300 ft. elevation. Coarse suffru- tescent herb to | m tall. Florets lavender. Local in sandy areas within rocky outcrops. Jan 24 1980. R. M. King and F. Almeda 8281 (Holotype UB; Isotype US). Paratypes. —Brasil: Goias: Chapada dos Veadeiros, 14°S, 47°W, ca. 15 km W of Veadeiros. Elev. 1000 m. Much-branched subshrub ca. 60 cm tall. Heads lavender- purple. Frequent. Campo. Feb 8 1966. H. S. Irwin, J. W. Grear, Jr., R. Souza, R. Reis dos Santos 12342 (US); ca. 20 km W of Veadeiros. Elev. 1000 m. Erect subshrub ca. 1 m tall. Heads magenta. Creek margin, among rocks. Feb 9 1966. Irwin et al. 12408 (US); herb ca. | m tall. Heads magenta. Fre- quent. Rocky slopes and wet campo. Feb 11 1966. Irwin et al. 12569 (US); ca. 15 km W of Veadeiros. Shrub to ca. 50 cm tall. Heads cream. Rocky slope and creek mar- gin. Feb 14 1966. Irwin et al. 12839 (US); ca. 10 km S of Alto do Paraiso (formerly Veadeiros). Elev. 1000 m. Subshrub ca. 50 cm tall. Heads in fruit. Rocky slopes. Mar 23 1969. H. S. Irwin, R. Reis dos Santos, R. Souza, & S. F da Fonseca 24953 (US); ca. 19 km N of Alto do Paraiso, elev. ca. 1250 m. Subshrub ca. | m tall. Heads in fruit. Outcrops. Cerrado on steep rocky slopes, surrounded by campo. Mar 20 1971. H. S. Irwin, R. M. Harley, G. L. Smith 32812 (US); 18-19 km N of Alto Paraiso. Wet campo at 4300 ft. elev. Shrub 1 m tall. Pap- pus tawny at maturity. Jan 24 1980. R. M. King & F. Almeda 8282 (US); 20 and 30 946 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON km N of Alto Paraiso de Goias, along road to Monte Alegre de Goias. Elev. 4200 ft. Subshrub *% m tall, flowers lavender or pur- ple. Feb 7 1981. R. M. King & L. E. Bishop 8810 and 8818A (US); West of road to Monte Alegre de Goias, 12—20 km N of Alto Paraiso de Goias. Elev. 4000-4400 ft. Shrubs 3/4 m tall, flowers past anthesis and lav- ender. Feb 7 1981. King & Bishop 8827 and 8828 (US). The species is in herbaria under the name Vernonia myrsinites Ekman, and a type photograph has been seen ofa specimen that was once 1n the Berlin Herbarium annotated by Ekman in 1912. Unfortunately, no place of publication has been found, and the name does not appear in any of the indices. It is not among the Ekman names validated by Malme (1933). Part of the reason for the oversight is the restricted distribution of the species, with recollections occurring in quantity only comparatively recently. The species is therefore described here as new. Ekman is cited only in synonymy since he proposed the name in a different combi- nation. There is no evidence that the present species has been confused in the past with the vegetatively similar Lessingianthus cor- digera of Minas Gerais. The latter has a more branching inflorescence with densely hairy tips on the inner involucral bracts as the most obvious differences. Lessingianthus niederleinii (Hieron.) H. Robinson, comb. nov. (Fig. 10) Vernonia niederleinii Hieron., Bot. Jahrb. Syst. 22:681. 1897. Vernonia valenzuelae Chod., Bull. Herb. Boiss. ser. 2, 3:641. 1903. Argentina, Brasil (Parana), Para- guay. Lessingianthus obscurus (Less.) H. Robinson, comb. nov. Vernonia obscura Lessing, Linnaea 4:296. 1829; 6:663. 1831. Vernonia davalliifolia Gardn., Lond. J. Bot. 6:422. 1847, as ““davalliaefolia’’. Cacalia obscura (Less.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (Goias, Sao Paulo). Lessingianthus obtusatus (Less.) H. Robinson, comb. nov. Vernonia obtusata Lessing, Linnaea 6:662. 1831. Cacalia obtusata (Less.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Vernonia sub- acuminata Hieron., Bot. Jahrb. Syst. 22: 691. 1897. Bolivia, Brasil (Goias, Mato Grosso, Minas Gerais, Sao Paulo). — Lessingianthus octandrus (Sch.Bip. ex Baker) H. Robinson, comb. nov. Vernonia octandra Sch.Bip. ex Baker in Mart., Fl. Bras. 6(2):87. 1873. Cacalia oc- tandra (Sch.Bip. ex Baker) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (Goias?). Lessingianthus onoporoides (Baker) H. Robinson, comb. nov. Vernonia onoporoides Baker in Mart., FI. Bras. 6(2):36. 1873. Cacalia onoporoides (Baker) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil (D.F., Goias, Mato Grosso, Minas Gerais, Sao Paulo). Lessingianthus pentacontus (DC.) H. Robinson, comb. nov. Vernonia pentaconta DC., Prodr. 5:30. 1836, as pentacantha, corr. 5:696. 1836. Cacalia pentacantha (DC.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Brasil? Lessingianthus platyphyllus (Chod.) H. Robinson, comb. nov. Vernonia platyphylla Chod., Bull. Herb. Boiss., ser. 2, 2:299. 1902. Brasil (Mato Grosso), Paraguay. Lessingianthus polyphyllus (Sch. Bip. ex Baker) H. Robinson, comb. nov. VOLUME 101, NUMBER 4 Vernonia polyphylla Sch.Bip. ex Baker in Mart., Fl. Bras. 6(2):63. 1873. Brasil (Par- ana), Paraguay. Lessingianthus pseudopiptocarphus (H. Robinson) H. Robinson, comb. nov. Vernonia pseudopiptocarpha H. Robinson, Phytologia 45:180. 1980. Brasil (Goias, Mato Grosso). Lessingianthus plantaginodes (Kuntze) H. Robinson, comb. nov. Vernonia rubricaulis var. squarrosa Less., Linnaea 4:300. 1829. Vernonia squarrosa (Less.) Less., Linnaea 6:678. 1831, non V. squarrosa [Don] Less., Linnaea 6:627. 1831 which was given precedence by Kuntze (1891). Cacalia plantaginodes Kuntze, Rev. Gen. Pl. 2:969. 1891, non V. plantaginoides Hieron. Vernonia pseu- dosquarrosa Hieron., Bot. Jahrb. 22:685. 1897. Vernonia sancti-pauli Hieron., Bot. Jahrb. Syst. 22:687. 1897. Vernonia squarrulosa Mattfeld ex Malme, Ark. Bot. 24A(6):18. 1931. Argentina, Brasil (Par- ana, Rio Grande do Sul, Sao Paulo), Uru- guay. Lessingianthus psilophyllus (DC.) H. Robinson, comb. nov. Vernonia psilophylla DC., Prodr. 5:28. 1836. Vernonia graminifolia Gardn., Lond. J. Bot. 6:421. 1847. Cacalia graminifolia (Gardn.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Cacalia psilophylla (DC.) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (D.F., Goias, Minas Gerais, Para, Parana). Lessingianthus pulverulentus (Baker) H. Robinson, comb. nov. Vernonia pulverulenta Baker in Mart., FI. Bras. 6(2):42. 1873. Cacalia pulverulenta (Baker) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (D.F., Goias, Minas Gerais). 947 Lessingianthus pumilla (Vell.) H. Robinson, comb. nov. Chrysocoma pumilla Vell., Fl. Flum. 331. 1825; 8, pl. 32. 1835. Vernonia sessilifolia Less., Linnaea 6:659. 1831. Cacalia pum- ila (Vell.) Kuntze, Rev. Gen. Pl. 2:969. 1891. Cacalia sessilifolia (Less.) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (Par- ana). Lessingianthus pycnostachyus (DC.) H. Robinson, comb. nov. Vernonia pycnostachya DC., Prodr. 5:58. 1836. Cacalia pycnostachya (DC.) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (Minas Gerais). Lessingianthus regis (H. Robinson) H. Robinson, comb. nov. Vernonia regis H. Robinson, Phytologia 45: 181. 1980. Brasil (Bahia). Lessingianthus reitzianus (Cabrera) H. Robinson, comb. nov. Vernonia reitziana Cabrera, Sellowia 13: 160. 1961. Brasil (Parana, Santa Catari- na). Lessingianthus robustus (Rusby) H. Robinson, comb. nov. Vernonia robusta Rusby, Mem. Torrey Bot. Club 6:54. 1896, non V. robusta Glaz., nomen nudum. Bolivia. Lessingianthus roseus (Mart. ex DC.) H. Robinson, comb. nov. Vernonia rosea Mart. ex DC., Prodr. 5:59. 1836. Cacalia rosea (Mart. ex DC.) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (Minas Gerais). Lessingianthus rosmarinifolius (Less.) H. Robinson, comb. nov. Vernonia rosmarinifolia Less., Linnaea 4: 285. 1829. Cacalia rosmarinifolia (Less.) 948 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (Bahia, Minas Gerais). Lessingianthus rubricaulis (Humb. & Bonpl.) H. Robinson, comb. nov. Vernonia rubricaulis Humb. & Bonpl., PI. Aequin. 2:66. tab. 99. 1809. Vernonia in- termedia DC., Prodr. 5:27. 1836. Cacalia intermedia (DC.) Kuntze, Rev. Gen. PI. 2:970. 1891. Cacalia rubricaulis (Humb. & Bonpl.) Kuntze, Rev. Gen. Pl. 2:971. 1891. Argentina, Brasil (Mato Grosso, Minas Gerais, Parana, Santa Catarina), Colombia, Paraguay, Peru. Lessingianthus rugulosus (Sch.Bip. ex Baker) H. Robinson, comb. nov. Vernonia rugulosa Sch.Bip. ex Baker in Mart., Fl. Bras. 6(2):83. 1873. Cacalia ru- gulosa (Sch. Bip. ex Baker) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (Minas Ger- ais). Lessingianthus saltensis (Hieron.) H. Robinson, comb. nov. Vernonia saltensis Hieron., Bot. Jahrb. Syst. 22:691. 1897. Argentina, Bolivia. Lessingianthus santosii (H. Robinson) H. Robinson, comb. nov. Vernonia santosii H. Robinson, Phytologia 45:182. 1980. Brasil (Bahia). Lessingianthus scaposus (Barroso) H. Robinson, comb. nov. Vernonia scaposa Barroso, Loefgrenia 36: 2. 1969. Brasil (Minas Gerais). Lessingianthus secundus (Sch.Bip. ex Baker) H. Robinson, comb. nov. Vernonia secunda Sch.Bip. ex Baker in Mart., Fl. Bras. 6(2):93. 1973. Cacalia se- cunda (Sch.Bip. ex Baker) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (D.F., Goias). Lessingianthus sellowii (Less.) H. Robinson, comb. nov. Vernonia sellowii Lessing, Linnaea 4:301. 1829. Cacalia sellowii (Less.) Kuntze, Rev. Gen. Pl. 2:971. 1891. Vernonia has- sleriana Chod., Bull. Herb. Boiss., ser. 2, 2:302. 1902. Argentina, Brasil (Parana, Rio Grande do Sul, Santa Catarina), Uru- guay. Lessingianthus soderstroemii (H. Robinson) H. Robinson, comb. nov. Vernonia soderstroemii H. Robinson, Phy- tologia 45:183. 1980. Brasil (D.F., Goias). Lessingianthus souzae (H. Robinson) H. Robinson, comb. nov. Vernonia souzae H. Robinson, Phytologia 45:184. 1980. Brasil (Goias). Lessingianthus stoechas (Mart. ex Baker) H. Robinson, comb. nov. Vernonia stoechas Mart. ex Baker in Mart., Fl. Bras. 6(2):49. 1873. Cacalia stoechas (Mart. ex Baker) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (Goias, Minas Ger- ais). Lessingianthus subcarduoides (H. Robinson) H. Robinson, comb. nov. Vernonia subcarduoides H. Robinson, Phy- tologia 45:185. 1980. Brasil (Minas Ger- ais). Lessingianthus subobtusus (Malme) H. Robinson, comb. nov. Vernonia subobtusa Malme, Ark. Bot. 24A(8):13. 1932. Brasil (Mato Grosso). Lessingianthus syncephalus (Sch.Bip. ex Baker) H. Robinson, comb. nov. VOLUME 101, NUMBER 4 Vernonia syncephala Sch.Bip. ex Baker in Mart., Fl. Bras. 6(2):64. 1873. Cacalia syncephala (Sch.Bip. ex Baker) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (Goias?). Lessingianthus tomentellus (Mart. ex DC.) H. Robinson, comb. nov. (Figs. 5-7) Vernonia tomentella Mart. ex DC., Prodr. 5:59. 1836. Cacalia tomentella (Mart. ex DC.) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (Minas Gerais, Sao Paulo). Lessingianthus ulei (Hieron.) H. Robinson, comb. nov. Vernonia ulei Hieron., Bot. Jahrb. 22:686. 1897. Brasil (Minas Gerais). Lessingianthus varroniifolius (DC.) H. Robinson, comb. nov. Vernonia varroniifolia DC., Prodr. 5:56. 1836. Cacalia varroniifolia (DC.) Kuntze, Rev. Gen. Pl. 2:971. 1891. Bolivia, Bra- sil. Lessingianthus venosissimus (Sch.Bip. ex Baker) H. Robinson, comb. nov. Vernonia venosissima Sch.Bip. ex Baker in Mart., Fl. Bras. 6(2):30. 1873. Cacalia venosissima (Sch.Bip. ex Baker) Kuntze, Rev. Gen. Pl. 2:971. 1891. Vernonia ur- baniana Glaz., Bull. Soc. Bot. France Mem. 3, 57:369. 1909., nomen nudum. Brasil (D.F., Goias, Mato Grosso). Lessingianthus vepretorum (Mart. ex DC.) H. Robinson, comb. nov. Vernonia vepretorum Mart. ex DC., Prodr. 5:59. 1836. Cacalia vepretorum (Mart. ex DC.) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (Minas Gerais). Lessingianthus vestitus (Baker) H. Robinson, comb. nov. 949 Vernonia vestita Baker in Mart., Fl. Bras. 6(2):83. 1873. Cacalia vestita (Baker) Kuntze, Rev. Gen. Pl. 2:971. 1891. Brasil (Minas Gerais). Lessingianthus warmingianus (Baker) H. Robinson, comb. nov. Vernonia warmingiana Baker in Mart., FI. Bras. 6(2):68. 1873. Cacalia warmin- giana (Baker) Kuntze, Rev. Gen. PI. 2: 971. 1891. Brasil (Goias, Minas Gerais). Lessingianthus westermanii (Ekman & Dusen) H. Robinson, comb. nov. Vernonia westermanil Ekman & Dusen ex Malme, Kungl. Svenska Vetenskapsakad. Handl. 12(2):10. 1933. Brasil (Parana). Lessingianthus xanthophyllus (Mart. ex DC.) H. Robinson, comb. nov. Vernonia xanthophylla Mart. ex DC., Prodr. 5:58. 1836. Cacalia xanthophylla (Mart. exe DEG) ikuntzes RevaiGens Ria 2-97Ae 1891. Brasil (Bahia). Lessingianthus zuccarianus (Mart. ex DC.) H. Robinson, comb. nov. Vernonia zuccariana Mart. ex DC., Prodr. 5:55. 1836. Cacalia zuccariana (Mart. ex DC) Kuntze) Reve Genv Pl 2:97 189 1k Brasil (D.F., Goias, Mato Grosso, Minas Gerais). Lessingianthus subg. Oligocephalus H. Robinson, subg. nov. Vernonia sect. Lepidaploa subsect. Oligo- cephalae Baker in Mart., Fl. Bras. 6(2): 46. 1873, pro parte. Lessingianthis subg. Lessingianthis typi- cis similis sed squamae involucri subae- quales et granae pollines polariter areolatae et intercolpe aequatorialiter biareolatae; a 950 Lepidaplois in capitulis pedunculatis et in cristis pollinis non rhizomatibus differt. Type.—Vernonia simplex Less. (=Les- singianthus simplex (Less.) H. Robinson. Lessingianthus desertorum (Mart. ex DC.) H. Robinson, comb. nov. Vernonia desertorum Mart. ex DC., Prodr. 5:43. 1836. Vernonia campestris DC., Prodr. 5:43. 1836. Vernonia desertorum var. campestris (DC.) Baker in Mart., FI. Bras. 6(2):48. 1873. Vernonia desertorum var. /ongipes Baker in Mart., Fl. Bras. 6(2): 48. 1873. Cacalia desertorum (Mart. ex DC.) Kuntze, Rev. Gen. Pl. 2:290. 1981. Vernonia desertorum var. macrocephala Chod., Bull. Herb. Boissier, ser. 2, 2:300. 1902. Brasil (Bahia, D.F., Goias, Mato Grosso, Minas Gerais, Sao Paulo), Par- aguay. Lessingianthus simplex (Lessing) H. Robinson, comb. nov. (Figs. 13-15) Vernonia simplex Lessing, Linnaea 4:280. 1829. Vernonia simplex var. angustifolia Less., Linnaea 4:280. 1829. Vernonia simplex var. latifolia Less., Linnaea 4:280. 1829. Vernonia simplex var. regnellii Baker in Mart., Fl. Bras. 6(2):53. 1873. Vernonia erigerontis Mart. ex DC., Prod. 5:43. 1836, nomen nudum. Bolivia, Brasil (D.F., Goias, Mato Grosso, Minas Ger- ais, Santa Catarina, Sao Paulo). Lessingianthus virgulatus (Mart. ex DC.) H. Robinson, comb. nov. (Fig. 16) Vernonia virgulata Mart. ex DC., Prodr. 5: 42. 1836. Cacalia virgulata (Mart. ex DC.) Kuntze, Rev. Gen. PI. 2:971. 1891. Brasil (Goias, Minas Gerais). The species is placed in the subgenus be- cause of the polar areolae, but it is probably not closely related to the other two species. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Acknowledgments The pollen specimens were prepared by Mary Sangrey and Barbara Eastwood using facilities of the Botany Department Paly- nological Laboratory. The photographs were prepared by Suzanne Braden and Brian Kahn of the Smithsonian Museum of Nat- ural History SEM Laboratory using a Hi- tachi S-570 scanning electron microscope. Literature Cited Baker, J. G. 1873. Compositae I. Vernoniaceae. in C. F. P. Martius, Flora Brasiliensis 6(2):2-179. Cassini, H. 1818. Composées. Pp. 131-159 in G. Cuvier, ed., Dictionaire des Sciences Naturelles, 10. Paris. DeCandolle, A. P. 1836. Prodromus Systematis Na- turalis Regni Vegetabilis, vol. 5. Paris. Jones, S. B. 1979a. Chromosome numbers of Ver- nonieae (Compositae).— Bulletin of the Torrey Botanical Club 106:79-84. 1979b. Synopsis and pollen morphology of Vernonia (Compositae: Vernonieae) in the New World.—Rhodora 81:425-447. . 1981. Revision of Vernonia series Flexuosae (Compositae: Vernonieae).—Brittonia 33:214— 224. 1982. A revision of Vernonia series Bud- dleiifoliae (Compositae: Vernonieae). — Britton- ia 34:102-117. King, R. M., & H. Robinson. 1987. The Genera of the Eupatorieae (Asteraceae). Monographs in Systematic Botany from the Missouri Botanical Garden 22:i-x, 1-581. Kuntze, O. 1891. Revisio Generum Plantarum, vol 2. Leipzig. Malme, G. O. A.-N. 1932. Die Compositen der zweiten Regnellschen Reise. II. Matto Gros- so.—Arkiv for Botanik 24A(8):1—57. 1933. Compositae paranenses Dusenianae. Kungelige Svenska Vetenskakadamiens Han- dlingar 12(2):1-122, pl. I-VI. Robinson, H. 1980a. Notes on the Lychnophorine genera Chresta and Eremanthus (Vernonieae: Asteraeae).— Phytologia 45:89-100. 1980b. New species of Vernonieae (Astera- ceae) V. Additions to Vernonia from Brasil.— Phytologia 45:166-208. 1980c. New species of Vernonieae (Astera- ceae). VII. Five new species of Vernonia from Brasil.— Phytologia 46:107-119. 1987a. Studies of the Lepidaploa complex (Vernonieae: Asteraceae). I. The genus Steno- VOLUME 101, NUMBER 4 cephalum Sch.Bip.— Proceedings of the Biolog- ical Society of Washington 100:578-583. 1987b. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). II. A new genus, Echinocoryne.—Proceedings of the Biological Society of Washington 100:584—589. 1987c. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). III. Two new genera, Cyrtocymura and Eirmocephala. — Proceedings of the Biological Society of Washington 100: 844-855. 1988. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). V. A new genus, Chrysolaena.—Proceedings of the Biological Society of Washington 101:952-958. Robinson, H., & V. A. Funk. 1987. A phylogenetic analysis of Leiboldia, Lepidonia, and a new ge- nus Stramentopappus (Vernonieae: Astera- ceae).—Botanische Jahrbiicher fur Systematik 108:213-228. Robinson, H., & B. Kahn. 1986. Trinervate leaves, 951 yellow flowers, tailed anthers, and pollen vari- ation in Distephanus Cassini (Vernonieae: As- teraceae).— Proceedings of the Biological Soci- ety of Washington 99:493-501. Robinson, H., & R. M. King. 1977. Eupatorieae: Sys- tematic review. Pp. 437-485 in V. H. Heywood, J. B. Harborne, and B. L. Turner, eds., The Bi- ology and Chemistry of the Compositae, chapter 15); Stix, A. 1960. Pollenmorphologische Untersuchung- en an Compositae.—Grana Palynologica 2:41- 104. Vellozo, J. M. da Conceicao. nensis. 1835. Florae Fluminensis, atlas, vol. 8. 1825. Florae Flumi- Department of Botany, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 952-958 STUDIES IN THE LEPIDAPLOA COMPLEX (VERNONIEAE: ASTERACEAE) V. THE NEW GENUS CHRYSOLAENA Harold Robinson Abstract. —The genus Chrysolaena is described for seven neotropical species, most of which were previously placed in Vernonia series Flexuosae. The genus has the lophate and rhizomatous exine structure of Lepidaploa, but these xy- lopodial herbs differ in being mostly unbranched below the inflorescence, in having glanduliferous anther appendages, and in lacking basal nodes on the styles. The present paper is the fifth in a series of seven devoted to the study of the neo- tropical Lepidaploa complex (Robinson 1987a, b, c) The group treated in the present study includes some well-known species of central and southern Brazil and adjacent Argentina and Bolivia. General similarity of habit and pubescence has resulted in some accurate grouping of the species in previous studies. Jones (1981) made the observation that all the species associated with the group had his type C pollen. The discovery of ad- ditional anther appendage and style base characters for the group during the present study has resulted in the recognition of the new genus Chrysolaena. Previous treatments of species of Chry- solaena have recognized some of the rela- tionships, but these treatments have always shown some omissions and have usually in- cluded some species that are not Chryso- laena. Both Baker (1873) and Cabrera (1944) placed the species that they recognized with- in the overall Vernonia Sect. Lepidaploa. Jones (1981) placed the group under sect. Vernonia, of which he made sect. Lepida- ploaasynonym. In more detail, Baker (1873) placed the then known species now placed in Chrysolaena among others in his sub- sects. Macrocephalae and Scorpioideae. Two other species later considered to be related by Jones (1981), V. simplex and V. deser- torum, were placed by Baker among other species in his subsect. Oligocephalae. Ca- brera (1944) established subsect. Flexuosae for four species of Argentina, which all proved to be Chrysolaena; but like Baker, he kept V. verbascifolia separated in subsect. Macrocephalae. The recent study by Jones (1981) reduced the rank of the Flexuosae to a series. Jones included all the presently rec- ognized members of Chrysolaena in his se- ries, except the species commonly known as V. hypochlora (syn. Chysocoma oligo- phylla). He also included the two species of somewhat similar habit from Baker’s (1873) subsect. Oligocephalae. Jones’ (1981) de- scription of series F/exuosae emphasized the xylopodial habit, the usual lack of branching below the inflorescence, the yellowish to brownish trichomes, the few to numerous sessile heads, the lanceolate acuminate to aristate involucral bracts, the strigose to pi- lose achenes, and the type C lophate pollen. Only the two oligocephalous species violat- ed the cited characters by having pedun- culate heads. The circumscription of the Flexuosae of Jones (1981) was believed to be totally ac- curate by the present author at the time the present series of studies began because of the basic similarities of habit and pollen type. The concept did not come into ques- tion until the pollen of the Lepidaploa com- VOLUME 101, NUMBER 4 plex was subjected to detailed SEM study. The recognition of two groups based on pol- len differences was followed by the discov- ery of style base and glandular trichome characters that are of generic importance. The present redisposition of the series Flex- uosae of Jones is based on the characteris- tics discussed below under the headings pol- len, inflorescences, involucres, nonglandular trichomes, style bases, and glands. The lat- ter refer particularly to the glands of the anther appendage. The generic distinctions emphasized are those from the two larger members of the complex, Lepidaploa and Lessingianthus. Pollen Asin all members of the Lepidaploa com- plex, the pollen of Chrysolaena has the ex- ine organized into a lophate pattern. The actual pattern is one referred to by Stix (1960) in her major study of Asteraceous pollen as the Vernonia cognata-Type. The same type has more recently been desig- nated by Jones (1979a) as type C. The type is distinguished by having a single areolae at each pole, no cross-walls in the colpus immediately above or below the pores, and usually two areolae equatorially across the intercolpar region. There are sometimes three areolae in one or two of the intercolpar regions of a grain (Fig. 1). The type C pattern is found in all the species that have been placed in the series Flexuosae, and all those recognized here as Chrysolaena. Under detailed SEM examination, the pollen of series Flexuosae of Jones (1981) shows two types. The majority of the species shows the “‘rhizomatous”’ structure of the exine crests (Figs. 1—4) that is characteristic of the genus Lepidaploa and its close rela- tives Stenocephalum and Echinocoryne (Robinson 1987a, b). The remaining two species, examined by SEM and light mi- croscopy, V. simplex and V. desertorum, show no such rhizomatous structure. The latter species are the same ones that Baker 953 (1873) placed in his subsect. Oligocephalae, separate from his placement of those now included in Chrysolaena. These two species have pedunculate heads, which distinguish- es them from others included in the Flex- uosae by Jones. The characters by which the latter two species resemble Chrysolaena can be considered comparatively superfi- cial. The non-rhizomatous pollen crests, the pedunculate condition of the heads, and the lack of glands on the achenes and anther appendages are all characters that distin- guish the species from Chrysolaena and in- dicate their proper placement in Lessin- gianthus (Robinson 1988). Inflorescences The sessile condition of the heads in all the members of Chrysolaena that were rec- ognized by Jones in his series Flexuosae correlate with the rhizomatous pollen crests in indicating a close relationship to the ge- nus Lepidaploa. As in the latter genus, the inflorescences of Chrysolaena characteris- tically show a seriate-cymose condition where every head has a terminal position and the continuation of the apparent pri- mary branch is through a lateral branch. Each head except the terminal one appears sessile as a result of the lateral branch being immediately below the head. The only member of Chrysolaena in which the char- acter seems to fail is the species added to the group in this treatment, Chrysocoma oligophylla. Even in this species, displace- ment of the lateral branches from imme- diately below the heads seems rare. The pe- dunculate appearance seems to be mostly derived from the presence of many branches that usually have only one head. No other character indicates that the species should be excluded from the genus. Involucres The involucres of Chrysolaena have bracts of graduated lengths, but they show none of 954 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1?7.6um 4. 3um Figs. 1-4. Chrysolaena herbacea (Vell.) H. Robinson, pollen. 1. Polar view showing polar areola and variation between two and three intercolpar areolae at periphery. 2. Lateral view of crest showing weak basal attachment of rhizome. 3. Colpar view with colpus partially collapsed and appearing interrupted. 4. Crest stripped from surface showing underside with rhizome. the additional differentiation in shape and _ primitive condition, rather than a reduc- texture seen in many species of Lepidaploa. tion. The lack of differentiation is reminiscent of The ratio of involucral bracts to flowers Lessingianthus, and it may be a retained _ in the heads of Chrysolaena is 1:1-—2:1. The VOLUME 101, NUMBER 4 comparative number of bracts is generally less than in either Lepidaploa or Lessin- gianthus, both of which tend to have 2:1- 3:1 ratios. The shift in basic ratio is in the opposite direction from that seen in two other genera of the complex, Stenocephal- um and Echinocoryne, where the involucral bracts occur in comparatively higher num- bers. Nonglandular Trichomes The stems, leaves, and involucral bracts of Chrysolaena bear long sericeous or lanate hairs that contribute to the similarity in ap- pearance of the plants. The hairs are yel- lowish to brownish and vary from straight to slightly flexuous. The hairs on the leaves vary from very dense in V. herbacea to sparse in V. platensis. Pubescence density in the inflorescence on the branches and involu- cral bracts is comparatively uniform in all the species. The hairs as seen in the species V. herbacea are the basis of the generic name Chrysolaena. Style Bases The presence or absence of nodular style bases is discussed in greater detail in treat- ments of the genera Lessingianthus (Rob- inson 1988) and Lepidaploa (Robinson in prep). The presence of a node seems basic to many neotropical members of the tribe Vernonieae and is regarded as basic to the Lepidaploa complex. However, three dif- ferent genera of the complex appear to have lost the node. There is little or no node in Stenocephalum (Robinson 1987a), and Les- singianthus has essentially no node in its more than ninety species (Robinson 1988). Chrysolaena also lacks a distinct node. The only enlargements seen are the result of the basal row of sclerified cells holding their shape while the softer tissue above con- tracts. This condition contrasts strongly with the usually large and sometimes ornament- ed basal disk seen in Lepidaploa. Appar- ently, the three genera lacking basal stylar 955 nodes have separately lost this structure. No function is presently known for the structure in the tribe, and therefore no functional shift is known that could explain the three sep- arate losses. The essential coincidence of the character with generic concepts is conve- nient for the taxonomist, but it does raise unresolved questions about the precise na- ture of the phyletic gaps between the genera. Glands Structures sometimes referred to as short- stalked capitate glands are a type of a bi- seriate hair characteristic of the Asteraceae. They are notable in the family for the ses- quiterpene lactones that they produce. They can be found on stems, leaves, involucres, corollas, anthers, style branches and achenes, but their appearance on vegetative parts, involucres, or corollas does not necessarily correlate with occurrence on anthers or ach- enes. In fact, only the tribes Heliantheae and Vernonieae commonly have glands on the anther appendages. Chrysolaena is one ge- nus of the Vernonieae that seems to char- acteristically have glands on the anther ap- pendages, but it is the only such genus in the Lepidaploa complex. In Chrysolaena the glands occur over most of the length of the appendage. They have been seen in all spec- imens of all but one species. The anther appendages of V. herbacea usually lack glands, although a few have been seen in some specimens. The achenes of Chrysolaena also usually have glands among the setulae, at least near the base. Such glands do not occur in Les- singianthus, but they are found in various species of Lepidaploa. Glandular achenes and rhizomatous pollen crests both indicate a closer relationship of Chrysolaena to Lep- idaploa than to Lessingianthus. The various characters cited for Chryso- laena in some cases correlate with Lepi- daploa and in others with Lessingianthus. The rhizomatous pollen crests are consid- ered evidence that the relationship is closer 956 to Lepidaploa. The latter view is strength- ened by the basically sessile condition of the heads and by the common occurrence of glands on the achene. The lack of a basal stylar node and the habital resemblance to Lessingianthus subg. Oligocephalus seems to be the only basis for relating Chrysolaena to Lessingianthus, but these resemblances seem to be the result of independent reduc- tions. Chrysolaena violates the generic lim- its of the two larger genera in a number of minor characters, such as the involucre/ flower ratio and the pedunculate heads found rarely in one species. Still, one character, the glands on the anther appendages, is a unique departure from all other members of the Lepidaploa complex. Even though the glands are usually missing in one of the species, their presence is too consistent in the genus as a whole to be ignored. The combination of the glanduliferous append- ages and the other discrepancies from Lep- idaploa and Lessingianthus lead to the pres- ent recognition of Chrysolaena as a distinct genus. The generic name is derived from the words Chryso- meaning golden and /aena meaning cloak. Chrysolaena H. Robinson, gen. nov. Vernonia subsect. Flexuosae Cabrera, Dar- winiana 6:329. 1944. Type: Vernonia flexuosa Sims. Vernonia series Flexuosae (Cabrera) Jones, Rhodora 81:442. 1979. Type: Vernonia flexuosa Sims. Vernonia series Verbascifoliae Jones, Rho- dora 81:438. 1979 as to type but not as to intent. Type: Vernonia verbascifolia Less. Plantae herbaceae perennes ad 2.5-15.0 dm altae xylopodiales, caules folia et brac- teae involucri flave vel fulve sericeae vel lanatae. Caules erecti interdum abbreviati sub inflorescentia non ramosi. Folia alterna vel basilares saepe superne descrescentia sessilia ovata vel linearia vel obovata supra et subtus parce vel dense sericeae vel lan- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON atae, nervis secondariis ascendentiter pin- natis vel sublongitudinalibus. Inflorescen- tiae interdum scaposae distincte cymosae saepe seriate cymosae, ramis dense sericeis; capitula sessilia raro breviter pedunculata. Involucra leniter subimbricata subgraduata 1—2-plo quam floribus 3—5-seriata, bracteis lanceolatis acutis. Flores 10-65; corollae purpureae, lobis plerumque glanduliferis caetera glabris in nervis apice conjunctis et leniter incrassatis; appendices antherarum plerumque glanduliferae; basi stylorum non noduliferi. Achaenia dense setulifera et ple- rumque glandulifera plerumque 5-costata; carpopodia turbinata, cellulis oblongis in parietibus lateralibus porulosis; setae pappi interiores capillares elongatae apice non lat- eriores, Squamae exteriores breviores. Grana pollinis in diametro 40-50 um lophata ad polos uni-areolata intercolpe dupliciter ar- eolata (typus C), cristis rhizomataceis. Type. — Vernonia flexuosa Sims. Chromosome numbers of N = 17 and N = 34 have been reported in members of Chrysolaena (Jones 1979b). The genus is concentrated geographically in central and southern Brazil and northern Argentina, and extends westward into Bo- livia with one occurrence in Peru. The genus contains the following seven species of which six have been recently treated by Jones (1981) as members of the Vernonia series Flexuosae. Chrysolaena flexuosa (Sims) H. Robinson, comb. nov. Vernonia flexuosa Sims, Bot. Mag. 51, pl. 2477. 1824. Vernonia montevidensis Nees ex Otto & Dietr., Allg. Gartenzeitung 1: 229. 1833. Cacalia flexuosa (Sims) Kuntze, Rev. Gen. Pl. 2:970. 1891. Ar- gentina, Brazil (Rio Grande do Sul, Santa Catarina, Sao Paulo), Paraguay, Uruguay. Chrysolaena herbacea (Vell.) H. Robinson, comb. nov. Chrysocoma herbacea Vell., Fl. Flum. 330. 1825. Vernonia oboyvta Less., Linnaea 4: VOLUME 101, NUMBER 4 279. 1829. Vernonia densevillosa Mart. ex DC., Prodr. 5:43. 1836. Vernonia chry- sophylla Gardn., Lond. J. Bot. 6:417. 1847. Cacalia obovata (Less.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Vernonia pau- cifolia Rusby, Mem. Torrey Bot. Club 3: 50. 1893. Vernonia herbacea (Vell.) Rus- by, Mem. Torrey Bot. Club 4:209. 1895. Bolivia, Brazil (Amazonas, D.F., Goias, Mato Grosso, Minas Gerais, Parana, Rio de Janeiro, Sao Paulo), Peru (Junin). Chrysolaena lithospermifolia (Hieron.) H. Robinson, comb. nov. Vernonia lithospermifolia Hieron., Bot. Jahrb. Syst. 22:694. 1897. Brazil (Mato Grosso, Minas Gerais, Parana, Santa Ca- tarina, Sao Paulo), Paraguay. Chrysolaena oligophylla (Vell.) H. Robinson, comb. nov. Chrysocoma oligophylla Vell., Fl. Flum. 324. 1825, Atlas 8: pl. 2. 1835. Cacalia oli- gophylla (Vell.) Kuntze, Rev. Gen. PI. 2: 968. 1891. Vernonia hypochlora Malme, Kungl. Svensk Vetenskapsakad. Handl. 12(2):12. 1933. Brazil (Parana, Santa Ca- tarina, Sao Paulo). The identity of the Vellozo species is briefly discussed in the treatment of Lessingianthus (Robinson 1988). The species name has been in- correctly associated with the habitally very similar V. cephalotes DC. Chrysolaena platensis (Spreng.) H. Robinson, comb. nov. Conyza platensis Spreng., Syst. Veg. 3:509. 1826. Vernonia platensis (Spreng.) Less., Linnaea 4:312. 1829. Vernonia cognata Less., Linnaea 6:670. 1831. Vernonia se- necionea Mart. ex DC., Prodr. 5:54. 1836. Cacalia cognata (Less.) Kuntze, Rev. Gen. Pl. 2:969. 1891. Cacalia platensis (Spreng.) Kuntze, Rev. Gen. Pl. 2:970. 1891. Vernonia sceptrum Chod., Bull. Herb. Boissier, ser. 2, 2:303. 1902. Ar- gentina, Brazil (Minas Gerais, Parana, Rio 957 Grande do Sul, Santa Catarina, S40 Pau- lo), Paraguay, Uruguay. Chrysolaena propinqua (Hieron.) H. Robinson, comb. nov. Vernonia propinqua Hieron., Bot. Jahrb. Syst. 22:695. 1897. Vernonia lepidifera Chod., Bull. Herb. Boissier, ser. 2, 2:304. 1902. Argentina (Misiones), Brazil (Para- na, Rio Grande do Sul, Santa Catarina, Sao Paulo), Paraguay. Chrysolaena verbascifolia (Less.) H. Robinson, comb. nov. Vernonia verbascifolia Less., Linnaea 4: 310. 1829. Argentina (Corrientes, Misiones), Brazil (Parana), Paraguay. Acknowledgments The pollen specimens were prepared by Mary Sangrey using facilities of the Bot- any Department Palynological Labora- tory. The photographs were prepared by Suzanne Braden of the Smithsonian Mu- seum of Natural History SEM Laboratory using a Hitachi 570 scanning electron mi- croscope. Literature Cited Baker, J. G. 1873. Compositae I. Vernoniaceae. in C. F. P. Martius, Flora Brasiliensis 6(2):2-179. Cabrera, A. L. 1944. Vernonieas Argentinas (Com- positae).— Darwiniana 6:19-379. Jones, S. B. 1979a. Synopsis and pollen morphology of Vernonia (Compositae: Vernonieae) in the New World.—Rhodora 81:425-447. . 1979b. Chromosome numbers of Vernonieae (Compositae).— Bulletin of the Torrey Botanical Club 106:79-84. . 1981. Revision of Vernonia series Flexuosae (Compositae: Vernonieae). Brittonia 33:214— 224. Robinson, H. 1987a. Studies in the Lepidaploa com- plex (Vernonieae: Asteraceae). I. The genus Stenocephalum Sch. Bip.—Proceedings of the Biological Society of Washington 100:578—-583. 1987b. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). I]. A new genus, Echinocoryne.—Proceedings of the Biological Society of Washington 100:584-589. 958 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1987c. Studies in the Lepidaploa complex Stix, A. 1960. Pollenmorphologische Untersuchun- (Vernonieae: Asteraceae). III. Two new genera, gen an Compositen. Grana Palynologica 2:41- Cyrtocymura and Eirmocephala.— Proceedings 104. of the Biological Society of Washington 100: CASS Department of Botany, National Mu- —. 1988. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). IV. The new genus Lessingianthus. —Proceedings of the Biological Society of Washington 101:929-951. seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(4), 1988, pp. 959-965 STUDIES IN THE LEPIDAPLOA COMPLEX (VERNONIEAE: ASTERACEAE) VI. A NEW GENUS, 4 YNIA Harold Robinson Abstract. — Aynia pseudascaricida is described as a new genus and species on the basis of collections from Ayacucho, Peru. The plants superficially resemble Baccharoides of India and Africa, but belong to the Neotropical Lepidaploa complex. Numerous leaf-like basal involucral bracts distinguish the genus from others of the complex. The non-rhizomatous lophate pollen is unique among the Vernonieae in having three intercolpar areolae at the poles. Efforts to resolve the neotropical Lepi- daploa complex of the tribe Vernonieae (Robinson 1987a, b, c) have resulted in the discovery of a totally undescribed member of the complex from southern Peru. For nearly sixty years since the collection by Kil- lip and Smith the specimens have remained undetermined. The species has a superficial resemblance to Baccharoides anthelmintica of India that is sometimes cultivated as a medicinal plant, but the new genus is most easily distinguished from Baccharoides and from various potentially related neotropical genera by its more robust habit with larger heads (Fig. 1). The pollen is also unique in its details (Figs. 2-5). The non-type A pollen and the Lepidapioa-like ratio of involucral bracts clearly distinguish the genus from typical Vernonia, where it would be placed under older, artificial systems of classifica- tion of the tribe. The genus and species are as follows. Aynia pseudascaricida H. Robinson, gen. et sp. nov. (Figs. 1-5) Plantae alte herbaceae vel suffrutes- centes erectae ad 2.5 m altae laxae ramosae. Caules pallides teretes striati puberuli eva- nescentes, internodis ad 8 cm longis. Folia alterna, petiolis 1-2 cm longis anguste ala- tis; laminae late ellipticae ad 12—15 cm lon- gae et 4-8 cm latae base breviter acuminatae margine remote mucronato-denticulatae apice supra viridis erecte puberulae parce immerse glandulo-punctatae subtus ciner- eo-tomentosae obscure glandulo-punctatae, nervis secundariis patentiter pinnatis utrin- que 12-15. Inflorescentiae cymosae laxe la- teraliter ramosae; ramis pilosulis; capitula plerumque pedunculata raro sessilia, pe- dunculis plerumque 3-13 cm longis. Invol- ucrum hemisphaericum; bracteae involucri basilares foliiformes patentes elliptico-lan- ceolatae 2—7 cm longae et 0.5—2.0 cm latae base anguste cuneatae margine apice supra et subtus ut in foliis; bracteae ceterum ful- vae rubro-tinctae appressae chartaceae sub- imbricatae ca. 100 in seriebus 4—5 lineari- lanceolatae 10-25 mm longae et base 2-3 mm latae apice peranguste acutae extus ap- presse puberulae et leniter pallide tomen- tellae ad medio distincte longitudinaliter unicostatae. Flores ca. 50 in capitulo; co- rollae puniceae vel lavandulae plerumque glabrae, tubis cylindricis ca. 8 mm longis, faucibus leniter infundibularibus 3-4 mm longis, lobis angustis ca. 10 mm longis et 0.8 mm latis apice dense spiculiferis in ner- vis incrassatis sensim sclerificatis; filamenta angusta carnosa in partibus superioribus ca. 0.65 mm longa, cellulis oblongis vel laxe oblongis vix noduliferis vel incrassatis; the- 960 3 | lhulavilinali AYNIA PSEUDASGARTICIDA H. det H Robinson U S National Herbarium we Fig. 1. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Holotype Robinson, UNITED STATES NATIONAL MUSEUM EXPLORATION IN PERU tyl stance de i . , Dept. Ayacucho; Aina, between Huanta and Rio Apurimac; f ” alt. 750-1000 meters; open woods 4) R iE. P. Kietar \ i} Aynia pseudascaricida H. Robinson, Holotype, United States National Herbarium, Washington, D.C. VOLUME 101, NUMBER 4 961 20. Gum : Biase 7 eee eal oe Ui fii Figs. 2-5. Pollen of Aynia pseudascaricida. 2. Polar view showing three intercolpar polar areolae. 3. Colpar view showing incomplete muri above and below pore. 4. Intercolpar view showing two tiers of two areolae. 5. Detail of pollen crest showing underlying large baculae attached to footlayer. 962 cae antherarum ca. 5.5 mm longae, cellulis endothecialibus suboblongis leniter asym- metrice lineate ornatis; appendices anther- arum lanceolatae ca. 1.8 mm longae ad me- dio costatae margine leniter reflexae apice pungentes; basi stylorum distincte noduli- feri; rami stylorum ca. 5.5 mm longi et abaxialiter dense hirsuti non glanduliferi. Achaenia ca. 4.5 mm longa et 1.5 mm lata superne in zonis 1 mm longis laeva et scler- ificata inferne sensim 10-nervata a videtur glabra base et apice parce et minute glan- dulifera raro superne minute setulifera, ra- phidis plerumque quadratis; carpopodia ca. 0.5 mm alta et 1 mm lata ad foraminem valde incurvata, cellulis oblongis in parie- tibus incrassatis lateraliter porosis; setae pappi flavidae capillares rigidae ca. 75 in seriebus 2—3 longiores plerumque 13-17 mm longae apice sensim angustiores pungentes extus convexae extus et margine scabridae; setae exteriores breves paucae indistinctae ca. 0.5 mm longae squamiformes vel su- bulatae et base alatae. Grana pollinis in dia- metro ca. 60 um lophata, cristis non rhi- zomataceis, areolis intercolpi aequatoriale in seriebus duplicibus binis, areolis inter- colpi unicis superioribus et inferioribus ad polem attingentibus (Aynia-type, Figs. 2-5). Type. —Peru: Ayacucho: Aina (Ayna), be- tween Huanta and Rio Apurimac; alt. 750- 1000 meters; open woods. Herb to 6 ft; stem simple; corolla tubes deep pink; styles white to pink; stamens deep pink. May 7, 17 1929. E. P. Killip & A. C. Smith 22514 (Holotype US). Paratype: Peru; Ayacucho: Aina, be- tween Huanta and Rio Apurimac; alt. 750- 1000 meters. Thickets. Shrub 5-8 ft, with elongate branches; florets deep pink. May 7,17 1929.E. P. Killip & A. C. Smith 23101 (US). The generic name derives from the col- lection locality in northern Ayacucho in Peru. The species name derives from the resemblance to the well known Baccha- roides anthelmintica (L.) Moench of India that is sometimes introduced into the Neo- tropical Region. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The ease of distinction of Aynia as a genus leaves the question of its closest relationship unresolved. There is no problem once cer- tain assumptions are made. Some of the most important information that would de- termine the placement of the new genus in the paleotropical or neotropical members of the tribe is lacking in the present material. At present there is no information on the chemistry and chromosome number, char- acteristics that were found to differ in the new and old world groups (Jones 1977). Nevertheless, Aynia is presumed to be na- tive to the area where it was collected, and in the absence of any characters specifically relating the genus to paleotropical genera such as Baccharoides, it is supposed that relationship is to the neotropical members of the Lepidaploa complex. Examination of details shows ample differences in Baccha- roides such as the form of the inflorescence, the presence of appendages on the involu- cral bracts, the long, narrow, basal tube of the corolla rather abruptly expanding into a cylindrical throat, the lack of thickened veins and the presence of glands rather than numerous spicules at the tips of the corolla lobes, the smaller cells of the anther collar, the lack of broad appendages in the bases of the anther thecae, the transversely oblong and closely vertically striated endothecial cells, the smaller and blunter anther ap- pendages, the lack of a node at the base of the style, and the more elongate achenes densely covered with setulae and glands. For this reason the new genus is seen as unre- lated to Baccharoides and is regarded as a member of the Neotropical element of the tribe. The chromosome number is expected to be based on x= 17 when it is determined, rather than x= 9 or 10 as in paleotropical members of the tribe. The relationship of Aynia within the Neo- tropical Vernonieae can be understood best by a review of various individual features in detail as follows under the headings, pol- len, inflorescence, corolla lobes, anther, style base, pappus, and geography. VOLUME 101, NUMBER 4 Pollen The pollen of Aynia is lophate with non- rhizomatous crests attached to the footlayer by large baculae (Fig. 5). The general form occurs widely in the Vernonieae in both pa- leotropical genera such as Baccharoides and American genera such as Stokesia, Mattfel- danthus and Lessingianthus. Typical Bac- charoides differs by having more than one row of smaller baculae under the crests, but other species of that genus have a single row of larger baculae as in the American genera. Relationship to the paleotropical and American groups cannot be determined on the basis of the pollen alone. The pollen grains of Aynia are distinct from all the genera mentioned, both paleo- and neo-tropical, in the detail of the areo- lation. The unique form can be viewed as having intercolpar groups of six areolae (1: 2:2:1) that are unique in reaching the poles of the grain (Figs. 2—4). In other pollen types with similar intercolpar groupings, the groupings are always separated from each other at the poles by colpi or extra areolae. In all other lophate grains in the Vernonieae with three areolae grouped at the poles, the areolae involved are aligned with the pores. The phyletic value of a distinct pollen form must be judged by the ease with which it can be derived from other types in related genera. In this case, the pollen is not du- plicated in any other member of the Ver- nonieae. This is in spite of the fact that a 1:2:2:1 intercolpar pattern which does not reach the poles is one of the most common patterns in the tribe, and it is even seen in a crude form in most type A grains. Inflorescence The two specimens seen show an inflo- rescence with heads in one cymose series (Fig. 1). Most of the heads are terminal, with the remainder of the inflorescence arising laterally from the axil of a leaf-like bract. In only one of the heads is the lateral branch from immediately below the head with the 963 head appearing sessile. In two of the seven heads observed, the head rather than the branch is axillary, and it is less mature than the head to which it is lateral. The inflores- cence structure is simpler than any in Bac- charoides, where the branch origins often lack bracts. In the regularly bracteate con- dition, Aynia is more like the Neotropical Lepidaploa complex. Within the latter com- plex, the inflorescence superficially most re- sembles Lessingianthus by its usually long- pedunculate heads. The large, foliose, basal bracts of the head, by which Aynia superficially resembles typ- ical Baccharoides, are obviously not evi- dence of close relationship to the latter ge- nus. Still, the bracts readily distinguish Aynia from all members of the Lepidaploa com- plex to which Aynia is apparently related. The other members of the complex with foliose outer bracts have a gradual transi- tion, and their foliose bract tips are never very large. The bracts in Aynia are totally leaf-like and show almost no intergradation. The inner involucre of Aynia has bracts of a totally different texture from the basal series. The ratio of the inner bracts to the flowers is 2:1, a ratio characteristic of Lep- idaploa and Lessingianthus. Such a ratio is widely distributed in the Neotropical Ver- nonieae, but it is not found in typical Ver- nonia from eastern North America. Corolla Lobes On the basis of an incomplete survey, the veins in the corolla lobes of the Lepidaploa complex seem distinct among the Verno- nieae by the extent to which they fuse broad- ly and form an enlargement at the apex. The least amounts of fusion seen in the complex in the genus Chrysolaena are as strongly fused as those of any other genus examined in the tribe. In this respect Aynia is most clearly a member of the Lepidaploa com- plex. The veins at the tips of the corolla lobes expand into a large shield that is equalled in the complex only in the genus 964 Mattfeldanthus. The tracheids and fibers form a mass that makes the tip of the lobe very stiffin both Aynia and Mattfeldanthus. The type species of the latter genus, M/. mu- tisioides H. Robins. & R. M. King, further resembles Aynia by the densely spiculifer- ous outer surface of the lobe-tip. Mattfel- danthus nobilis (H. Robins.) H. Robins. dif- fers in having numerous small glands rather than spicules on the outer surface. The lobe character, more than any other, specifies the relationship of Aynia to the Lepidaploa complex. Anther The anther thecae of Aynia are like those of Mattfeldanthus and unlike those of Bac- charoides in both basal appendages and en- dothecial cells. The two Neotropical genera both have broadly truncate differentiated bases on the thecae. The marginal cells are obviously specialized and form slight lobes. Baccharoides, in contrast, has shortly point- ed anther bases with almost no differen- tiated marginal cells. The endothecial cells of the Neotropical genera are also alike in their curved, thickened bands that usually leave the median surfaces of the cells un- thickened. In Baccharoides the cells are shorter and have vertical bands usually rather evenly spaced and straight across the surface. The apical anther appendage of Aynia is distinct from those of all the other genera discussed by the lanceolate shape and the median costa. The appendage differs from Baccharoides additionally by the somewhat recurved margins and the laxer cells. The appendages of Baccharoides are much shorter, blunter, denser and flatter. The an- ther collars of Baccharoides are like the api- cal appendages in having smaller cells than in Aynia and Mattfeldanthus. Style Base The presence of a node at the base of the style is of some use in determining rela- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tionships in the Vernonieae. Aynia has a distinct node and in that way differs from the majority of paleotropical Vernonieae in- cluding Baccharoides. Those paleotropical Vernonieae having a stylar node such as Distephanus (Robinson & Kahn 1986) also tend to differ by having a much larger node. Nodes of the type seen in Aynia are most common in the neotropical Vernonieae and exactly the same form can be found in Matt- feldanthus. It is notable that distinct nodes are lacking in the other genus of the Lepi- daploa complex having non-rhizomatous pollen, Lessingianthus. Thus, the stylar node, like the corolla lobes, places Aynia closer to Mattfeldanthus, which differs in inflorescence form and corolla symmetry; and it tends to separate Aynia from Lessin- gianthus, which it resembles more in the latter two characters. Pappus The pappus of Aynia looks different from that of related genera in color and persis- tence, but the actual character differences are more subtle. The comparative persis- tence seems related to the presence of more rows of bristles. The color difference is, at least partly, due to the thicker bristles. The bristles are convex and scabrid on the outer surface. In contrast, the bristles of Mattfel- danthus are flattened and smooth on the outer surface, and the bristles of Baccha- roides are flattened and scabrid. A well- marked outer pappus series 1s not so ob- vious in Aynia as in most Vernonieae including Mattfeldanthus; however, some shorter outer papper segments are present. They are of various lengths and a few have broadened alariform bases with shortly awned tips. The form of the pappus is not unique within the Vernonieae, but it seems unique within the Lepidaploa complex. Geography The occurrence of Aynia in the Andes is a prime reason for relating the genus to oth- VOLUME 101, NUMBER 4 er neotropical Vernonieae rather than to pa- leotropical genera such as Baccharoides which has superficially similar basal invo- lucral bracts. All details of structure appear to reenforce the geographical evidence, and there is no reason to doubt that Aynia is related to the neotropical Vernonieae with which it occurs. Nevertheless, the geog- raphy does not correlate perfectly with the genera to which the new genus is evidently most closely related. Lepidaploa, which is common in the Andes, belongs to a more specialized element of the Lepidaploa com- plex that has rhizomatous crests on the pol- len. The two other genera of the complex that have non-rhizomatous pollen are con- centrated in Brazil. Both species of Matt- feldanthus are presently known only from Bahia in Brazil. Lessingianthus, which oc- curs in Peru, is represented there only by species extending their ranges from farther east. The geography seems to reenforce the distinct nature of Aynia in the Lepidaploa complex, with the new genus being the only element with non-rhizomatous pollen crests that is not centered in Brazil. Acknowledgments The pollen specimens were prepared by Barbara Eastwood using facilities of the Botany Department Palynological Labora- tory. The photographs were prepared by 965 Brian Kahn of the Smithsonian Museum of Natural History SEM Laboratory using a Hitachi 570 scanning electron microscope. The photograph of the holotype was made by Victor E. Krantz, Staff Photographer, Museum of Natural History. Literature Cited Jones, S. 1977. Vernonieae: Systematic review. Pp. 503-521 in V. H. Heywood, J. B. Harborne, & B. L. Turner, eds., The biology and chemistry of the Compositae, chapter 17, Academic Press, New York. Robinson, H. 1987a. Studies in the Lepidaploa com- plex (Vernonieae: Asteraceae). I. The genus Stenocephalum Sch. Bip.—Proceedings of the Biological Society of Washington 100:578-583. 1987b. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). IJ. A new genus, Echinocoryne.—Proceedings of the Biological Society of Washington 100:584—-589. 1987c. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). III. Two new genera, Cyrtocymura and Eirmocephala.— Proceedings of the Biological Society of Washington 100: 844-855. Robinson, H., & B. Kahn. 1986. Trinervate leaves, yellow flowers, tailed anthers, and pollen vari- ation in Distephanus Cassini (Vernonieae: As- teraceae). — Proceedings of the Biological Soci- ety of Washington 99:493-501. Department of Botany, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 101(4), 1988, p. 966 BIOLOGICAL SOCIETY OF WASHINGTON 115th Annual Meeting, 18 April 1988 The meeting was called to order by Austin Williams at 1:00 p.m. in the Waldo Schmitt Room, National Museum of Natural History. Austin announced the election results. The new elected officers are listed on the inside front cover of this issue. Don Wilson, Treasurer, reported that income from dues, subscriptions, page- charges, and sale of past issues of the Proceedings in 1987 was lower than for 1986. Total income for 1987 was $88,476.70. The estimated income for 1988 is $75,800.00, and the estimated expenditures are $68,550.00—leaving $7,250.00 available to subsidize page charges. Austin noted that the report had - been approved by the Council. Austin announced a change in the editorial staff of the Proceedings. Brian Kensley, Editor for the past seven years (Volume 94-100), understandably, had retired. He received our many thanks and a standing ovation from the members present. Kensley still served as editor for Bulletin No. 8 (Results of recent research on Aldabra Atol, Indian Ocean), which was published in 1988. The new editor, C. Brian Robbins, was introduced. Also, Stephen D. Cairns agreed to serve as an additional associate editor for Invertebrate Zoology papers. Brian Robbins, Editor, then reported on Volume 100 of the Proceedings. One hundred papers totaling 1050 pages, were published in 1987. It was noted that papers submitted by individuals unable to pay full page-charges were currently subject to a publications delay of one year. Unpaid papers now account for about 60 pages per issue. Don Wilson indicated that funds were available to increase this to 80 pages per issue; this will be implemented for Volume 101 (1988). Brian reported that a motion to limit the number of subsidized pages per author(s) per year to 12 printed pages (approximately 27 manuscript pages, including figures and tables), was approved by the Council. Also, Brian an- nounced that our manuscript review policy, including suggestions and guide- lines for reviewers and associate editor forms, was being revised and will affect all future manuscripts. The 100 Year Index is currently being prepared by Phyllis Spangler. A format has been agreed to by the editorial staff. Copy will be ready for comments, revisions, etc. by Fall of 1988. A motion was made and seconded that the meeting be adjourned; Kristian Fauchald, the new President, adjourned the meeting at 1:45 p.m. Respectfully submitted, C. Brian Robbins Editor ape 7 rogle ify) f ii! INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (botany, zoology, and paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with a summary in an alternate language when appropriate. Submission of manuscripts. —Submit manuscripts to the Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History NHB-108, Smithsonian Insti- tution, Washington, D.C. 20560. 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Costs. — Printed pages @ $60.00, figures @ $10.00, tabular material @ $3.00 per printed inch. One ms. page = approximately 0.4 printed page. Presentation. — Manuscripts should be typed double-spaced throughout (including tables, leg- ends, and footnotes) on one side of 8'2 x 11 inch sheets, with at least one inch of margin all around. Submit two facsimiles (including illustrations) with the original, and retain an author’s copy. Pages must be numbered consecutively. Underline singly scientific names of genera and lower categories; leave other indications to the editor. The sequence of material should be: Title, Author(s), Abstract, Text, Acknowledgments, Literature Cited, Author’s(s’) Address(es), Appendix, Figure Legends, Figures (each numbered and identified), Tables (double-spaced throughout, each table numbered with an Arabic numeral and with heading provided). Clarity of presentation, and requirements of taxonomic and nomenclatural procedures ne- cessitate reasonable consistency in the organization of papers. Telegraphic style is required for descriptions and diagnoses. Literature citations in the text should be in abbreviated style (author, date, page), except in botanical synonymies, with unabbreviated citations of journals and books in the Literature Cited sections. Direct quotations in the text must be accompanied by author, date, and pagination. The establishment of new taxa must conform with the requirements of the appropriate international codes of nomenclature. When appropriate, accounts of new taxa must cite a type specimen deposited in an institutional collection. Examples of journal and book citations: Eigenmann, C.H. 1915. The Cheirodontidae, a subfamily of minute characid fishes of South America.— Memoirs of the Carnegie Museum 7(1):1-99. Ridgely, R. S. 1976. A guide to the birds of Panama. Princeton, New Jersey, Princeton University Press, 354 pp. Olson, S. L. 1973. The fossil record of birds. Pp. 79-238 in D. Farner, J. King, and K. Parkes, eds., Avian biology, volume 8. Academic Press, New York. Figures and tables with their legends and headings should be self-explanatory, not requiring reference to the text. Indicate figure and table placement in pencil in the margin of the manu- script. Plan illustrations in proportions that will efficiently use space on the type bed of the Proceedings. Original illustrations should not exceed 15 x 24 inches. Figures requiring solid black backgrounds should be indicated as such when the manuscript is submitted, but should not be masked. CONTENTS Cryptotrochus, new genus and two new species of deep-water corals (Scleractinia: Turbinoliinae) Stephen D. Cairns Syringonomus dactylatus, a new species of bathyal marine nematode (Enoplida: Leptosoma- tidae) and a supplementary description of Syringonomus typicus Hope and Murphy, 1969 W. Duane Hope Notes on Antroselates Hubricht, 1963 and Antrobia Hubricht, 1971 (Gastropoda: Hydrobiidae) Robert Hershler and Leslie Hubricht A revision of the genus Themiste (Sipuncula) Edward B. Cutler and Norma J. Cutler A new species of polychaete, Scolelepis anakenae (Polychaeta: Spionidae) from Easter Island, South Pacific Ocean, with ecological comments N. Rozbaczylo and J. C. Castilla Description of Guaranidrilus oregonensis (Oligochaeta: Enchytraeidae) from North America, with additional comments on the genus Kathryn A. Coates and Robert J. Diaz Taxonomic revision of the Phallodrilus rectisetosus complex (Oligochaeta: Tubificidae) Christer Erséus Four new species of Cambarincolids (Clitellata: Branchiobdellida) from the southeastern United States with a redescription of Oedipodrilus macbaini (Holt, 1955) Perry C. Holt Pycnogonida of the western Pacific Islands V. A collection by the Kakuyo Maru from Samoa Koichiro Nakamura and C. Allan Child Darwinulocopina (Crustacea: Podocopa), a new suborder proposed for nonmarine Paleozoic to Holocene Ostracoda I. G. Sohn Hyalopontius boxshalli, new species (Copepoda: Siphonostomatoida), from a deep-sea hydro- thermal vent at the Galapagos Rift Arthur G. Humes Description of Membranobalanus robinae, a new species of sponge barnacle (Cirripedia, Ar- cheobalanidae) from Baja California, with a key to the genus Robert J. Van Syoc Elasmopus balkomanus, a new species from the Florida Keys (Crustacea, Amphipoda) James Darwin Thomas and J. L. Barnard Synalpheus dorae, a new commensal alpheid shrimp from the Australian Northwest Shelf A. J. Bruce Redescriptions of Tetralia cavimana Heller, 1861 and Trapezia cymodoce (Herbst, 1799) first stage zoeas with implications for classification within the superfamily Xanthoidea (Crustacea: Brachyura) Paul F. Clark and Bella S. Galil Allopotamon, a new genus for the freshwater crab Potamon (Potamonautes) tambelanensis Rathbun, 1905 (Crustacea: Decapoda: Potamidae) from the Tambelan Islands Peter K. L. Ng The status of Cryptochirus hongkongensis Shen, 1936 (Brachyura: Cryptochiridae) Roy K. Kropp The status of Cryptochirus coralliodytes Heller and Lithoscaptus paradoxus Milne Edwards (Brachyura: Cryptochiridae) Roy K. Kropp The status of Callianassa hartmeyeri Schmitt, 1935, with the description of Corallianassa xutha from the west coast of America (Crustacea, Decapoda, Thalassinidea) Raymond B. Manning Labral morphology in heart urchins of the genus Brissopsis (Echinodermata: Spatangoida), with an illustrated revised key to western Atlantic species Richard L. Turner and Cathleen M. Norlund Patterns of geographic variation in the Arabian Warbler Sylvia leucomelaena (Aves: Sylviidae) Steven M. Goodman New species of fossil vampire bats (Mammalia: Chiroptera: Desmodontidae) from Florida and Venezuela Gary S. Morgan, Omar J. Linares, and Clayton E. Ray Studies in the Lepidaploa complex (Vernonieae: Asteraceae) IV. The new genus, Lessingianthus Harold Robinson Studies in the Lepidaploa complex (Vernonieae: Asteraceae) V. The new genus Chrysolaena Harold Robinson Studies in the Lepidaploa complex (Vernonieae: Asteraceae) VI. A new genus, Aynia Harold Robinson Biological Society of Washington: 115th Annual Meeting Table of Contents, Volume 101 Index to new taxa, Volume 101 709 WY 730 741 767 773 784 794 809. 817 825 832 838 843 853 861 866 872 883 890 898 One 929 952 959 966 970 974 L prs i; ane = Dianne NG ice, > ~~ a SSS = > MITHSONIAN INSTITUTION | NOLLNIILSNI_NVINOSHLIWS (S31 uvu gl LIBRARIES SMITHSON# ~ fir z= { Ys oO o = Oo oe S » ise) ra w = w = ‘Yf, Pe) x Eo) 5 = E 4 2S = z ee a = Yj, = 5 ~ Es z en D z A es i= y z 1 VINOSHIIWS SaldvedIT_ LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHI] = Ye = 5 2 20h es | = Zz = = = = = aS Se S ss = ne =) i Zz = x S Zi rs 2 a > 2 2 S a = : MITHSONIAN INSTITUTION NOILNIILSNI_NVINOSHLINS S31YWvuaty LIBRARIES SMITHSON = Fa 5 ” y WD fas = Pq o we ar = Up ty Ly (ex wc S te c BS a) = aa) = aa) ey (2) a oO ca oO a. me =J PA, J > Th ANOSHLINS SAIYVYd!It LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHII Nk x fe) a ° — o = Sa Z = 5 E 2 SS = > fee > = = ‘ WQ. = 2 = 2 = 23 \ Se w he = Z wD = a Z ys », AITHSONIAN NOILALILSNI S3J!IYVYa!I1 LIBRARIES SMITHSON Zz ” z wo z ee wn j = = is = Sy NS = Kemp Zz = Zz = Z WY = oO ae oe) Be Oo We ASE 2) no n oO Y 2) RQ ~ se fe) 3e oO % ae \N oO = = = 2 = YS 2) on = oe > = > = < > Sinise Op) ; Pz 79) Zz 2p) Sr nike = “a JINOSHLINS S3IYVUEII INSTITUTION NOILNLILSNI NVINOSHIE ie 7p) 3 iG S ‘G S ox =e - s = Ss ~ o RE a = oe . a a a Ne < a = > lo > Ke 2 fs 2 = 22 - 4/ mn 5a Be cee m O° an = — wm = i VINOSHLINS (Sa iuYVvYydi _t! 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