PAD ne Ll pene Rey fentine ad She tintiasine es eet oe Gn Besse sa tieciasb PP NOTEN 0640010 ees -stiahp new a naman RAE GS eRte De Steak PLAT ANTI R eden mya Te URS APERE BEN TE Sa We say taptine, meni edintie fade aeons eduze over Wm vita SALVO Me Vom Rabe Peres Ara SIM sane Meteo Peter ara 1% SPA Treas DAg Mons rine Dathen rigamecdarh anita malts Dnt aA ca Cc ra * = Bathe Veal 6 Ped ode aye eT HOR nm tN Sore daa 4 b- ES ESE nS Oph EP ENT # rae EE OETA TI bP) staid tay Ses 9iths Halk Soe yes 81 Poa Bis AN rms ATR HT ‘ han Pas Ben erm Zar an nLP AW As Mheate fen denimath in Pata NEN TIN ENN Tania al Ay Syne UWA Nn Ne a P97 Hee EA My tne DAMN SIRNAS AUN Oe Burtt Ree M ee aie Av at= Asm thacynen aed el tm Sores aera aMatveten Betas pm TS EER em MPC Ae HN Vibe em 1 a aD at Bj Hai 295 NOTED Sash biny Ai Set fe Lent Tene NA AR an Mee ova CDE es NW RDS DN hs Fe REM entoe Fa MOOG os Est Ey SPR ADA Ms WT gore tr Lt IIS et SMA inatexn ang. PENI SAAS THR RENAN MAG As ana OO hile i aes or Cc Fone mer Sc a 5S a 2 TH 3 2 GQ 2 N's a z a z i a z a S a $ : a NI pene oe fyuVvug rr LIBRAR ce SMTBSONIAN INSTITUTION. NOLLALIZSN a saltuvyg Phe LIBRARIES SMITHSONIAN _— Mm a -_ — = = te ow NS = o S @ 2 Pa 2 © = o 7 WSs BK 5 “ =] a 5 a = a = Pe) = ra > i: > a > ~ > . > fe 2 a 2 = E E> 2» = 2 - 2 = Fs a x = Ps m 4 m 2 m 2 m a m 2 m | —— = = wn he 7) = 7) S SMITHSONIAN INSTITUTION |, NONLALILSNI_ NVINOSHLIWS $3 tuvuag rit BRARI ES SMITHSONIAN INSTITUTION. NOILNLILSNI NVINOSHLIWS = = ae = Z = me gas oer ee z < z . = z = Ye? eee SS = = \ = << = =o ; +t fyt , SS ; = is ~ z = z EGS 3 S ILA 2S 3 5 5 5 3 S = g = S Gam = \ 8 = = : 2 5 = > = > ; = > = Ss = - 2 7) es 7) z ” = a eee 7) 2 us s S BS ea I RSONIAN INSTITUTION, NOLLALILSHI NVINOSHLIWS. $2 fYvVYstit_ LIBRARIES SMITHSONIAN _ & es a ee i ul 2 & 8 a us & = o a! a. ¢ Md ae tc = ~ = o a — foe (= a 5: S c = a = > = OL = > E ae fe - a XS ra a re OL <7. a - 2 - uF Ww AN NS wo — a Za a — ~ z on = oO S\ = ie 2 m 2 mn 2 as = om == Ww = (27) . » = JI _NVINOSHLIWS S3 tyvUusit = LI Baan I ES, SMITHSONIAN INSTITUTION NOILNLILSNI_NVINOSHLINS $3 (Yuvyuyslt = LIBRARI ES SMITHSONIAN f < = eee = ae = < = ; sec 2 = z =z = z 4 = = Uy = = z : =| = - O ce S = Yd. z 5 Li =| La Ww > wo Fa : o ie ow = 4 o zt ow ist oc = os = 5 x ce 5 a 2 m. = m = as oO pa rs) = re) = Zz pa) = — = SMITHSONIAN INSTITUTION NOILONLILSNI NVINOSHLINS S31YVYSIT LIBRARIES SMITHSO! = z im z oe a y Pas) = = = ow = 2 LG oF Va 7 > 0] Ly ; ‘ PM > > | z > e ZL ” — n s = o” — NVINOSHLINS S3IYVHURIT LIBRARIES SMITHSONIAN NOILNLILSN!I NWINOSHI wn C4 sa 72) z wn Za = 4 = < = i 4 rs 4 Zz Se Ape Pare aM cS 2 = 2% * : a i = ee 5 “RAITLICONIAN {INCQTITIITION AIMGFINITICAlt NYVINOCUTIIAIC ecptuYvNuNaoIs4 LIRRARIES SMITHSOR aoe) -{ Ww i te ae re as PROCEEDINGS of the Biological Society of Washington VOLUME 99 1986 Vol. 99(1) published 14 May 1986 Vol. 99(2) published 4 June 1986 Vol. 99(3) published 17 October 1986 Vol. 99(4) published 12 December 1986 WASHINGTON PRINTED FOR THE SOCIETY EDITOR BRIAN KENSLEY ASSOCIATE EDITORS Classical Languages Invertebrates GEORGE C. STEYSKAL THOMAS E. BOwMAN Plants Vertebrates Davip B. LELLINGER RICHARD V. VARI Insects RoBERT D. GORDON All correspondence should be addressed to the Biological Society of Washington, Smithsonian Institution Washington, D.C. 20560 ALLEN PREss INC. LAWRENCE, KANSAS 66044 OFFICERS AND COUNCIL of the BIOLOGICAL SOCIETY OF WASHINGTON FOR 1985-1986 OFFICERS President AUSTIN B. WILLIAMS Vice President KRISTIAN FAUCHALD Secretary C. W. HART, JR. Treasurer DON C. WILSON COUNCIL Elected Members STEPHEN D. CAIRNS RICHARD P. VARI MASON E. HALE STANLEY H. WEITZMAN ROBERT P. HIGGINS DONALD R. WHITEHEAD TABLE OF CONTENTS Volume 99 Amin, Omar M. On the species and populations of the genus Acanthocephalus (Acan- thocephala: Echinorhynchidae) from North American freshwater fishes: a cladistic AMAL SUS ies ee a ea A See co em AC ee ek la er Pee Banks, Richard C. Subspecies of the Glaucous Gull, Larus hyperboreus (Aves: (@ hy ares hen Re eS) ee se Aare eee ean Nn Se eee Becker, Jonathan J. A new vulture (Vulturidae: Pliogyps) from the Late Miocene of Bertelsen, Rodney Duane. Speleobregma lanzaroteum, a new genus and species of Scal- ibregmatidae (Polychaeta) from a marine cave in the Canary Islands. Bishop, Gale A. Two new crabs, Parapaguristes tuberculatus and Palaeoxantho liber- tiensis, from the Prairie Bluff Formation (Middle Maastrichtian), Union County, INAS S1'S Sp) PTLD Neer seeRanRD OEET SBe ) eP k sO oere Oon eC Bowman, Thomas E. Tridentella recava, a new isopod from tilefish burrows in the New Works WB velott (VFleovel Uhtererne “rvs Keone re aS) eet Bowman, Thomas E. Paracymothoa tholoceps, a new freshwater parasitic isopod from southern Venezuela (Flabellifera: Cymothoidae) eee Bruce, A. J., and F. A. Chace, Jr. Paralebbeus zotheculatus, n. gen., n. sp., a new Byrnes, Thomas, and Roger Cressey. A redescription of Colobomatus mylionus Fukui from Australian Acanthopagrus (Sparidae) (Crustacea: Copepoda: Philichthyidae)....... Calder, Dale R. Symmetroscyphus, a new genus of thecate hydroid (Family Thyro- Soy Phi dae) sro ri Berm Uae eae ae a ee ee eee Chapman, Peter M., and Ralph O. Brinkhurst. Setal morphology of the oligochaetes Tubifex tubifex and Ilyodrilus frantzi capillatus as revealed by SEM... Clark, Janice, and J. L. Barnard. Tonocote, a new genus and species of Zobrachoidae from Argentina (Crustacea: Marine Amphipoda) eee eee eeeeeeeteeneeneeeeeeeeeeeee Coates, Kathryn A. Redescription of the oligochaete genus Propappus and diagnosis of a new family Propappidae (Annelida: Oligochaeta) eee eee eenenteeeeeeeeeeeeeeeeeeeee Cutler, Norma J., and Edward B. Cutler. A revision of the genus Nephasoma (Sipuncula: Gr oo lft oti Aa) ee A ees Ba a Se Delaney, Paul M. The synonymy of Cirolana tuberculata (Richardson, 1910) (Isopoda: lal tye late rea 3G inc earn Ge) fae ae le sae te eR oe ENE Dickerman, Robert W. Two hitherto unnamed populations of Aechmophorus (Aves: FO GO Gp bi Ae) ee Pee ee Erséus, Christer. Marine Tubificidae (Oligochaeta) at Hutchinson Island, Florida............. Ferrari, Frank D., and Ruth Bottger. Sexual dimorphism and a sex-limited polymor- phism in the copepod Paroithona pacifica Nishida, 1985 (Cyclopoida: Oithonidae) fromthe wRed Sea. ota eI, Se ee eR ee eC ed Frazier, J. G. Epizoic barnacles on pleurodiran turtles: is the relationship rare?.................. Froglia, Carlo, and Raymond B. Manning. Notes on the species of Allosquilla and Platysquilloidesn(@mstacca SLOMATOp OCA) meee eee aaa Frost, Darrel R. A new Colostethus (Anura: Dendrobatidae) from Ecuador........................... Gardner, Alfred L. The taxonomic status of Glossophaga morenoi Martinez and Villa, LOS 8i(Mammlalias Chiroptera e hyllostorm ce) esse eee eee George, Robert Y. Serolis agassizi, new species, from the deep sea off Cape Fear, North @aroliniai(Grustacea: 3180 pOG a) taser ee ee Ee Ea ee eee Gonzalez, Exequiel. A new record of Paracorophium hartmannorum Andres, 1975, from the Chilean coast, with a description of the adult (Amphipoda: Corophiidae)................. Graves, Gary R. Geographic variation in the white-mantled barbet (Capito hypoleucus) ofiColombia(Awves:Gapitomid ae) ta ee Graves, Gary R. Systematics of the gorgeted woodstars (Aves: Trochilidae: Acestrura). Graves, Gary R., and Storrs L. Olson. A new subspecies of Turdus swalesi (Aves: Passeriformes: Muscicapidae) from the Dominican Republic ..............-ccssssssssseeesseeeeseeeeeeeeeee 574-579 149-159 502-508 375-379 604-611 269-273 753-756 237-247 388-391 380-383 323-327 225-236 417-428 547-573 731-734 435-436 286-315 274-285 472-477 261-265 214-217 489-492 46-50 21-28 61-64 218-224 580-583 Harper, Donald E., Jr. Nephtys cryptomma, new species (Polychaeta: Nephtyidae) from theme rthern:Gilfio fa MGxi C0 tice ie ee ee ee ee a Heron, Gayle A., and David M. Damkaer. Two species of Urocopia, planktonic poe- cilostomatoid copepods of the family Urocopiidae Humes and Stock, 1972 _................ Hershler, Robert, and Glenn Longley. Hadoceras taylori, a new genus and species of phreatic Hydrobiidae (Gastropoda: Rissoacea) from south-central Texas... Heyer, W. Ronald, and Reginald B. Cocroft. Descriptions of two new species of Hylodes from the Atlantic forests of Brazil (Amphibia: Leptodactylidae) o..00000.ccccccccceeceeeeee Higgins, Robert P. Redescription of Echinoderes pilosus (Kinorhyncha: Cyclorhagida). Hobbs, Horton H., Jr., and Andrew G. Grubbs. Notes on the crayfish Procambarus (Ortmannicus) xilitlae (Decapoda: Cammbaridae) eee eeeeeeeeessecnceeeeceeeeeevesnenceeeeeeeseeseneeee Holt, Perry C. Newly established families of the order Branchiobdellida (Annelida: Clitellata) with a symopsis Of the Semera. ccc ceeccceeeceeceeeeeeeeeeeeeeeeetennnennnnnnnnnnnnnneneeeeesenneennnen Houbrick, Richard S. Discovery of a new living Cerithioclava species in the Caribbean (Mollusca: Prosobramchia: Cerith iid ae) cece ceeceeeeeceeceeeeeeeeeeeeeeeenvnnnnnnnnnnnnnnnnnnnneeneceeeceessesens Houbrick, Richard S. Transfer of Quadrasia from the Planaxidae to the Buccinidae (Mollusca: Gastropoda: Prosobrarch ia) cece eeeeeeceeeeeeceeeeeeeeeeevesevnnnnnnnnnnnnnnnnnnnnnenneesesesnenneesee Hutchings, P. A., and C. J. Glasby. Glossothelepus, a new genus of Thelepinae (Poly- chaeta: Terebellidae) from the Gulf of Califormia, Me@X1CO o..............0.cccccccscssssseeeeeeeeeeeeeeeeeeeeeeeeeees Jara, Carlos G. Aegla spectabilis, a new species of freshwater crab from the eastern slope of the Nahuelbuta Coastal Cordillera, Chatde 0.00... cccccccccccecccceccscccssseeseeeeceecseceeseeeeesecennesesseeeeseennueees Kornicker, Louis S. Redescription of Sheina orri Harding, 1966, a myodocopid ostra- code collected on fishes off Queensland, Australia ccc ceeeeeeceeeceeeceeeeeeeeeeeeeneeeneee Kritsky, Delane C., and Mary Beverley-Burton. The status of Pseudorhabdosynochus Yamaguti, 1958, and Cycloplectanum Oliver, 1968 (Monogenea: Diplectanidae)......... Kritsky, D. C., W. A. Boeger, and V. E. Thatcher. Neotropical Monogenea. 9. Status of Trinigyrus Hanek, Molnar, and Fernando, 1974 (Dactylogyridae) with descriptions of two new species from loricariid catfishes from the Brazilian Amazon... Kropp, Roy K. A new type designation for Petrolisthes tomentosus (Dana), and descrip- tion of Petrolisthes heterochrous, new species from the Mariana Islands (Anomura: Rorcellanid ac) meses 5.2 Be MARIS Ie eee RS tee ee ee ee Lew Ton, Helen M., and Gary C. B. Poore. Neastacilla falclandica (Ohlin), type species of the genus, and N. tattersalli, new species (Crustacea: Isopoda: Arcturidae)......... Manning, Raymond B. A small trap for collecting crustaceans in shallow water................ Manning, Raymond B., and Darryl L. Felder. The status of the callianassid genus Callichirus Stimpson, 1866 (Crustacea: Decapoda: Thalassinidea) ...... Manning, Raymond B., and Richard W. Heard. Additional records for Callianassa rathbunae Schmitt, 1935, from Florida and the Bahamas (Crustacea: Decapoda: GalliavasS1clte) eae a ees eee ee AN DPS OE HAE RE ES NU ee tank Manning, Raymond B., and L. B. Holthuis. Preliminary descriptions of four new species of dorippid crabs from the Indo-West Pacific region (Crustacea: Decapoda: BY GPEG) OI U E21) | es see easeere ee uLD een evs A GMted bel INE wT LCE E OME NANO MN Dye Gea ke REN aoirhdSIG UA BAER ASIA LETS TNI Manning, Raymond B., and L. B. Holthuis. Notes on Geryon from Bermuda, with the description of Geryon inghami, new species (Crustacea: Decapoda: Geryonidae).......... Maris, Robert C. Larvae of Xiphopenaeus kroyeri (Heller, 1862) Crustacea: Decapoda: Penaeidae) from offshore waters of Virginia, U.S.A. cece ccceeceecccccceeceecececeneeenennntttetttts McCranie, James R., and Larry David Wilson. A new species of red-eyed treefrog of the Hyla uranochroa group (Anura: Hylidae) from northern Honduras... Miller, John E., and Richard L. Turmer. Psolus pawsoni (Echinodermata: Holothuroi- dea), a new bathyal sea cucumber from the Florida cast COASt i cceececceceee ete Milligan, Michael R. Separation of Haber speciosus (Hrabé) (Oligochaeta: Tubificidae) from its congeners, with a description of a new form from North America.................... Modlin, Richard F.. Caecidotea dauphina, a new subterranean isopod from a barrier island in the northern Gulf of Mexico (Crustacea: Isopoda: Asellidae).. Ng, Peter K. L., and Gilberto Rodriguez. New records of Mimilambrus wileyi Williams, 1979 (Crustacea: Decapoda: Brachyura), with notes on the systematics of the Mim- ilambridae Williams, 1979, and Parthenopidae MacLeay, 1838, sensu Guinot, IO FR evens a eae. ANAT, AVG NLR RCE AUNT eee I. Weve UA aioe Lt Phe ee 1-7 140-148 121-136 100-109 399-405 735-738 676-702 257-260 359-362 84-87 34-41 639-646 17-20 392-398 452-463 191-195 266-268 437-443 347-349 363-365 366-373 602-603 51-55 478-485 406-416 316-322 88-99 Owens, Joan Murrell. Rhombopsammia, a new genus of the family Micrabaciidae (Coe- J@ritera tas SClGie titted): cence cee eee een ee ee eee ED Owens, Joan Murrell. On the elevation of the Stephanophyllia subgenus Letepsammia to generic rank (Coelenterata: Scleractinia: Micrabactidae) eee Pereira, Guido. Freshwater shrimps from Venezuela I: Seven new species of Palaemon- inael( Crustacea Decapodazslealac ma @rni ca c) kee eae anne Pettibone, Marian H. A new scale-worm commensal with deep-sea mussels in the seep- sites at the Florida Escarpment in the eastern Gulf of Mexico (Polychaeta: Polynoidae: Branchipoly moma) she hth ba eh a ce oe ee lee a ee ed aia et ae Petuch, Edward J. New South American gastropods in the genera Conus (Conidae) and Watirus (Hasciolaridae).2 22 sees = = ee 1 eee eee Seale nite el ame ene a Petuch, Edward J. The Austral-African conid subgenus Floraconus Iredale, 1930, taken OfsBenmiundan(Gastropodas orc ac) eee a ene ee ee Price, Roger D., and K. C. Emerson. New species of Cummingsia Ferris (Mallophaga: rinienoponidace) trom Renurangd \Viene Ziti e | ee een ne Randall, John E., and Ernest A. Lachner. The status of the Indo-West Pacific cardi- nalfishes Apogon aroubiensis and A. MigrOfASCLAtUs cece eeeeeeeeeeeeevevevvvvvvvnnnnnnnnnnennnnneneeeeee Rasmark, Berit, and Christer Erséus. A new species of Tubificoides Lastochkin (Oligo- chaeta: Tubificidae) from Bermuda and the Bahamas... ccc ceeeeeceesecneeeeeeeeeeenneeeeeee Reeder, Richard L., and Walter B. Miller. A new species of Helmonthoglypta (Gastrop- oda: Pulmonata: Helminthoglyptidae) from San Diego County, California... Reid, Janet W. A redescription of Microcyclops ceibaensis (Marsh, 1919) (Copepoda: Cyclopoida) from Marsh’s specimens in the National Museum of Natural History... Rios, Rubén. Caridean shrimps of the Gulf of California. V. New records of species belonging to the subfamily Pontoniinae (Crustacea: Decapoda: Palaemonidae) Robins, C. Richard. The status of the ophidiid fishes Ophidium brevibarbe Cuvier, Ophidium graellsi Poey, and Leptophidium proftundorvur Gl ccc eee Robinson, Harold, and Brian Kahn. Trinervate leaves, yellow flowers, tailed anthers, and pollen variation in Distephanus Cassini (Vernonieae: Asteraceae)... Riitzler, Klaus, and Shirley M. Stone. Discovery and significance of Albany Hancock’s microscope preparations of excavating sponges (Porifera: Hadromerida: Clionidae)... Savage, Jay M. Nomenclatural notes on the Anura (Amphibia)... eee Spangler, Paul J. Three new species of water scavenger beetles of the genus Chaetarthria from South America (Coleoptera: Hydrophilicae) 00 ceee eee eeeeneeeenteeneeee Stauffer, Jay R., Jr. and Kenneth R. McKaye. Description of a paedophagous deep-water cichlidi(@eleoster: Cichlidae) tromilakel MalawieyA tila Titgen, Richard H. Hawaiian Xanthidae (Decapoda: Brachyura) II. Description of Gar- thiella, new genus, with a redescription of G. aberrans (Rathbun, 1906)... Vari, Richard P. Serrabrycon magoi, a new genus and species of scale-eating characid (Bisces> Characifonmes)) fromthe wpper R10) Ne pro Vecchione, Michael, and Clyde F. E. Roper. Occurrence of larval J//ex illecebrosus and other young cephalopods in the slope water/Gulf Stream interface eee Wasshausen, Dieter C. The systematics of the genus Pachystachys (Acanthaceae)............. Weems, Robert E., and John F. Windolph, Jr. A new actinopterygian fish (Paleonisci- formes) from the Upper Mississippian Bluestone Formation of West Virginia................ Weitzman, Marilyn J., and Richard P. Vari. Astyanax scologaster, a new characid (Pisces: @stariophysi)mrompthe Rio wNEsros South yA tn eri Galea eee nnee nna Weitzman, Stanley H. A new species of Elachocharax (Teleostei: Characidae) from the IRGVORNESTOMESTONKOfMVENE Ze LAvAT Clim ES el 711 | eee aes ee a eae cae ne Weitzman, Stanley H., Naercio A. Menezes, and Heraldo A. Britski. Nematocharax venustus, a new genus and species of fish from the Rio Jequitinhonha, Minas Gerais, Brazalk(eleostei:Characidac)= ee ee ee Oe ee ee. ee ee Wicksten, Mary K., and Michel E. Hendrickx. Alpheopsis cortesiana, a new snapping Shinimpmnomluthes Grulite@ fe allt o rere a O Williams, Austin B., and Darryl L. Felder. Analysis of stone crabs: Menippe mercenaria (Say), restricted, and a previously unrecognized species described (Decapoda: Dan thidae) wo. ci.:8 er ees = A eS ee ae SE ee eee Williams, Ernest H., Jr., and Lucy Bunkley Williams. The first Ani/locra and Pleopo- 248-256 486-488 198-213 444-451 8-14 15-16 748-752 110-120 612-615 137-139 71-78 429-434 384-387 493-501 658-675 42-45 509-516 29-33 56-60 328-334 703-708 160-185 584-601 709-716 739-147 335-346 196-197 517-543 dias isopods (Crustacea: Cymothoidae) parasitic on Japanese fishes, with three new SC C1 CS ee ere NOIR RS LN I Ie ey 2 AN a A MN ei eh al Wilson, George D. F. Pseudojaniridae (Crustacea: Isopoda), a new family for Pseudo- janira stenetrioides Barnard, 1925, a species intermediate between the asellote su- perfamilies Stenetrioidea amd Jarirorde a nnn cece ceeeceeeeeecenneceeeneeeeetnenneceeneeeeeennnnunenennne Wolf, Paul S. A new genus and species of interstitial Sigalionidae and a report on the presence of venom glands in some scale-worm families (Annelida: Polychaeta)............ Wolf, Paul S. Three new species of Pilargidae (Annelida: Polychaeta) from the east coast of Florida, Puerto Rico, and the Gulf Of Me@xiCO o2...o..cccccccccececccseseeesceeecceeceeseecseeesesceseeeeeeeeeeeneesenee Wolf, Paul S. Four new genera of Dorvilleidae (Annelida: Polychaeta) from the Gulf of INAS xa pace ea eo cates Dee ee cern A NY No atc redeem aarh han Wolf, Paul S. Three new species of Dorvilleidae (Annelida: Polychaeta) from Puerto Rico and Florida and a new genus for dorvilleids from Scandinavia and North PENS Tal © ete a erect ca ee ene Jen Rn EN A ta Na aa tee eel an cae Yager, Jill, and Frederick R. Schram. Lasionectes entrichoma, new genus, new species, (Crustacea: Remipedia) from anchialine caves in the Turks and Caicos, British West Tin lat Speen cameo er elie reel ee a nae ee IN ee ae ad Zullo, Victor A., and William Miller, III. Barnacles (Cirripedia: Balanidae) from the Lower Pleistocene James City Formation, North Carolina coastal plain, with the description of a new species Of Balanus Da Costar iin.e....cccccccccccccccecceeeeecneeeceeeeeeeeeeteeneneeeeeeeeeetees 647-657 350-358 79-83 464-471 616-626 627-638 65-70 717-730 INDEX TO NEW TAXA VOLUME 99 (New taxa are indicated in italics; new combinations designated n.c.) COELENTERATA Hydrozoa SIV IVUIVICLY. OS GY DAUIS ecco ee cer a Be re De UR 381 Scleractinia ER LOVIAOPS CLYVUN AIT Cement ee ee er ere pk ree caren 249 TUUPD PLCLCL Ch tne i a Ee i el ON Se 252 ESC LLLN CS Lense ee ee KEP ag einai feline ere tee eee 1A A) eI 250 PLATYHELMINTHES Trematoda Trinigyrus acuminatus 393 tentaculoides 395 SIPUNCULA INIGpoI SCONCE ONION TUITON [OVS TV ONT VL FO CC pe 556 diaphanes corrugatum 558 pellucidum subhamatum n.c._ 564 WOCADISKsIWOGIANIS KAI 1 Coss ee eee a eee ee a 566 wOdjaniskiGelisae ini Cy Se tn een eae SE ee 566 ANNELIDA Polychaeta BrAanchipOlyMOess CCN Cris Sex Ose wees eee oe eee es EAS I ee ene ee 445 Di QPROTOSOMGA -cceecscnsceeecevecccveeeeeee 620 LST C1] Ca 620 Dorvillea (Dorvillea) clavata 631 LAR GTLCTLEIS Reet rei Ae Rie ME PRS EOE, Dh ER A Enna, Sr ee ee 628 (Schistomeringos) perkinsi 635 Eliberidens 622 i OF CCD Set ba eaniieuie) Sak tee Saves Rls WE ede eee eee 623 GIOSSOLNET C1) 11 Seinen ean Ais. PTS I DRL dee ee OUI GE EEE acted en een ie 84 IMOXIC ANU Seen 84 Litocorsa antennata 465 17 (2104) OXX01// 11/12 719 UEDE LACK CH Cl Cy merate tear NN Be Ie els ee OR See Db rac ee RO MM De era, 80 1 617 LCTUULAC TIE Ses seteles. Nt ctnns MeN a et See ENN Je es Ue ee ae ee ne 617 DPT OUST ser ee BJ Sa ls Beer ORT ON Br BIN Ree ee ce aE 636 RY 1420) 0) 541/10 376 lANZAVOLEUM oe 376 Synelmis acuminata 467 CANA died) Stcrecre aR RR Nas eR SU ty UR Bs ety. eRe cadre gon NC ibe NER A a Sn 469 WEST CIG GIG sie Be 1 an eS oh 8 LEE (eo Re en SE ee eee eee, 623 Oligochaeta TBE (TiS NALS UH ON cc ea rte Ce 302 HAS OS carck Se sts aa a Ee a ue ES 300 I YLCLGHO [DUOS UCLGULS area es ateee aed eto ae re Bee Noel INH eee eee Le cs ene dee 303 Coralliodrilus corpulentus 305 Tet terol Lin S 9/2 0:5 19 10a 01s ammeter Nera iin. ven nD tba Pr ee af ale ec gn ee 291 IDRIS ats se ee AS 2 JA RN ey 2 81 el eevee TE ET NEL 294 Olavius Jatus 307 PE irved evedreall Sea Oe hal Seared Sta a eee ese wesc aac ect ae 297 [EERSTE ae rE eh OST Ne 298 FIT) CDT) LCL G Cee a ee LD ed PNA ene ts 418 TEMS AON LSI OT Es ae PS a ODP ee orp ot 310 UG CTY LULA Cements Pedal aaah 612 Clitellata LBRONANID GSA NOLTT ee cD Ee eT EET RE IO OTE 688 BY CUTLG IODC LEG GC MMRME me ith oR cen aeo UA lh a eet Ue agentes AL el A go ee Fa 685 GOAT ATC ONCE Cl Cae ae nn sl eat og ude Tat Ae ees iy ee vee a AO 692 CON YELLOW UNIVE ee OEE ge ORD oe re Oe CN 692 NGL OTLOCLILUTCL Cr CaM eee es Peet a ee une lo la ese Al Nal ln pombe vga Ps 691 ARTHROPODA Insecta (Shreve Cea neE LMS ARC CCITT ate eS Rs See RS nt et A erp eee 511 PAVING meee 513 porknockeri 509 Cummiingsia (Acamthomemopon) SOPH ri... eee eeeeeeeee ene eneneeveennnnneeenneeevnnncnveevennnnessnnunnneseennnennsunneeeeeeee 750 (Gumimainesiayoankley ae. ee ee eee ee 748 Aegla spectabilis ...................... 34 Alpheopsis cortesiana 196 PATTI CHAR LCL CL ete aes aah Se Ze NB ees ho i ca a ae 651 FID TGLO TUL 1 me eR EATS BS SSP ETN elie IA PRO Eee PL a A ced ea oN de oe 647 Balanus neusensis 723 Caecidotea dauphina 316 Ginolamamtubye nc ell eat ety rs C5 aes eee 731 UVB) TRIN PSE FLO) AU Case sate ee TN ee aE US eC lel ea 363 TL LLCLLTD CS Pe Re cate gt hE sar toal | Cie Ete ee ed ga Ny AIST etal RRs 2 eee rae a 364 Garthiella 56 UDG HeIPeAT SBT essa acer ine ee cere eect ee ee ARAL eh a Oe Me ee ue alae ea 57 Geryon inghami 367 Y OKY (0) 161 ke 65 PLE ECILG 11] Cl came ssetaae ha ie rea nT a seen Noeae cera enn eRe eee ae Seat ee i ecee ene te ae 66 IMiaero brachii acaba Cris Cae tes aes ee ee a ee 202 CLC TGV UID Y TET] mn reeRE © cso Beek nce ae! Wana eA ele Se UR ase eae es 204 FED CELUI UU TY Waianae LER IL ed ele ee Si A PE oes ee eB 200 pumilum 208 ROS cect Beh ep et aa fa IR a ee RPS RPI Toa RPO ON a Ae 198 TO CLUS U1 (aaa ee IP REAR Var OM AN Tce Cd Nine Ooo enc een ee eee ea 206 NAY, (2500) 0) 0,0 (0 17/17 een 525 Neastacilla tattersalli 193 Nobilum arachnoides 364 Palaemonetes (Palaemonetes) mercedae 209 IPAIACOXN ATG rs ee ea es led Serie ate 607 SDC LUCTAG US eam te aaron bd URL alc ii ec ORS RE a eo tae ee 609 ParacymoOthoa th OLOCE pS sxc.1. 2 es Bee ee eee aN ee ere Ss ee Pe oe 753 PAtaAG OTP PC RCOL AY OIA ac a ea na ee 365 PGA CD CU Sf a a eee NE DON En eee ee 237 ZOE OCU AUUUS ioc ase ec age ee ne ee Es ee oa Se ee ae 238 PCT AD GSU EST CS fos ee Fo ea es RE ee 605 tuberculatus 606 Petrolisthes hererochrous 2 = ee Oe ee eee 458 PAT © pO GUS S U0 7 CHEE TAS eee Re eg Ree 656 IPSCUAO| QUT AAC ae a ce ee Freer BR el ee 351 Sr LS RASS S 0271 ses ec ee eo 46 hoi DOT magellani............ 228 Tridentella recava 269 LED OY OY of EY Ca SENAY Ve 0 ea re Se 146 MOLLUSCA Gastropoda STIL O CLA A aH CT cat ae Sea Ne ee 257 (@ leapt GLA ORT ee ee cease ee 359 (GOMUSHCOTI OC eS Sa ee a I 9 gibsonsmithorum 9 WES IUED OTE aire i Bi a arctan, ek OU 2k AR hh al Ne 16 MTD CTLGILCS LCA) ce a ch cB ee a 10 TLS caters ccna aPC ne ARE 8h re re ns tae es oe an Ate 10 LOST CS UR Na a Ren I eee ee ee 11 xanthocinctus 13 Hadoceras............. 122 taylori 125 Helminthoglypta montezuma 137 CAPITIS VOM IILEL Lire cee ee ee Ee ag Gol Reh tr hr ni nk Ee ope a een See EE 8 ECHINODERMATA Holothuroidea IES 1 NAS YE) CVS 111 eee eee ee nn eee ee ee 478 CHORDATA Pisces PAS EV AT AKASCOl OG SIE Toisas te ae ee 709 Diplotaxod onpereenwOOd ian ee 2 ee eee eee eee 29 Elachocharax mitopterus 740 Nematocharax 335 venustus 336 tS CET CLD Ta) CO To teat ae a ee ee) els 329 TYLEO Die Pee era RR eg at ar ree get ere VR eens, PONCE LEE 2 ee aS Se ee 329 TCT D ECT ICHEE/IIY Sets acter etaen rere adenine Nn at eee es a a 595 DLL CUS Y Sits saci a ere a ae ee Tee ee 595 Amphibia GOO StS tT S E7121 77 15 ee a re eee ee 0k esc 214 Ly ex SCLIV OVER Ca etx ce as ST og UR ete PY ra op 51 Hylodes charadranaetes 106 DNV LOM CS: rite Ee Nl See OR IE CY cE 1 yates Se 104 Aves AechintO phorus’ Clarks sie seo rats ee Se a eee eee ee 436 OCCIGEN ALIS SHS H ALES Ie A i eg at wee oe 436 Capito hypoleucus carrikeri : 62 hypoleucus extinctus......... 62 PALA OYERV DS, CLIO ke ea ent vee er DC ene ee Rte 503 sittindisas wale sind OGG Cram cornmeal inet UE Le Ale ee MeN te ss a 581 PLANTES Spermaphyta WISE MAMUSTAMCOLEMSIS i Cees ere ee ee ee ee aN ee OS 498 ATT ST UDNI EOL UTN RC Serene ese eed conten ees SE et ear en eet ots aa iY ea lel 499 TNS OC MACTOIMES RING Seer Niantic Sy an eos need eer ogee) Nee Be IAN We Lk oh 499 PENDS WN BONY OU see St ea ER oe Se 499 cloiselii n.c. 499 hia CALLS SEC wate im Neem ieee ie tee Ree ee Nea Meee tars UL ule 499 STATO PEA HIN Ste eee ere reer he eID Oe ed a ete 499 HE COOTEIGE SUED UBTNY Gemma e tic een aN Ee ener ate eC cera ee ety arpa er aces ta le Sei ae 499 (SEIT MSE NEST he hls eS rf oO 499 PL AMC UMICINCLUS RIC ee cet eae o the | MN, ee Cre vet a Ay arte athe ee 499 plutingsusynC ee Piso rela nena ete el erie) 499 NUS Tee US Uae spe sansa ly ae a ee de see cet ce ie oe A 499 ITTV ed Ha eal gs 6 © Bytes eee a ere eee rceeae es ialea. ian SOMES racemes eee eae Aner Sa RE Nunes NheeoBL 499 TNE UU RES OSG US) ane ee ee A el LE UI Os 499 MEN lll AC OTD ENV VE SWINE eres eet tec aN Ne ee een ae hae eels Lies Ua eras wine aca ed i alee I glen 499 NAN ATM VO) © TS TS Ie Cece ee esol ec a at ae ees tv SNe ln ee A ee RI fea AN OU a ae 500 PMB TT ATV OKC TNS1S 3 NE ese eee a ee eas a daar Nee) eaten eels 500 OUMTTAA ATU ATV AC fC) ULUL Siete Cae eae vier eo nd i lle le Leche ret ee a ps ahaa 500 OID LE UCU SINE CU ee aa pe ae cata Eee reas li et aL onal 500 MO lyealacto lawn Cet laws ie ck ees Cle nu) La oa eae aie eta nema 500 TOE NV TN TL OLE SHINN sesso oe era Cree alee Ek sla Sea Bee MOY ele UAE Saw A ni Mla a 500 streptocladus n.c. ........ 500 HUTS) NWSI USS i eat orp Cy eee Ae PER ae Pe CO Oo Py Neneh er Aare 500 SSVI 11 © ISIN Seer eS ee MAD NUNN NN eA oe a la ean ect 500 GvimmmanthemiumaCOlOPAvU I Ce 500 IZAChiy StAG MW SHI GALOS DICE ts meets chiA e Aas aks ee series Oar te od os alten eee ate ot aiee a Maen 168 BO SLC) Maem ae ORO et NOB copes cat ee eee Re EB RG eA Re em aS eee oe 168 EEG CATGYL CLL (eer ese ce Nea tees I ty ge I oa lt ae eae a ke 171 killipii 179 WOMEUDVACLCO LC ee me NE IANS eee ML A ONG Ae noe Oil ot eke clk A 163 DSS 0) Cl Cpememnee eset aa ey SR ce SONNE SNES Os Aa a shah eel ated dee NA a ate ae cn acca 180 173 178 166 SYMP SON a ae NN yn PN aR se DG Rn DO CSP 175 SS a te ae Pea. opheycl® rina tap ay gmat etal - oe Saas 7 : amen 2 i - a iP: oir. ae eo) A i ~ aed 7 a A : - : ithe " pi ee 7 tei. eos 7 . » — 7 (Rieatdh hg oD “eu <=> f - a : - free ee i Wy, 7 THIS ISSUE IS DEDICATED TO JOHN W. ALDRICH ON THE OCCASION OF HIS 80TH BIRTHDAY, AND IN RECOGNITION OF HIS LONG SERVICE Aine SOCIETY THE BIOLOGICAL SOCIETY OF WASHINGTON 1984-1985 Officers President: Donald R. Davis Secretary: Gordon L. Hendler Vice President: Austin B. Williams Treasurer: Leslie W. Knapp Elected Council J. Laurens Barnard Maureen E. Downey Frederick M. Bayer Louis S. Kornicker Isabel C. Canet Storrs L. Olson Custodian of Publications: David L. Pawson PROCEEDINGS Editor: Brian Kensley Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Richard P. Vari Insects: Robert D. Gordon 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: $18.00 within the U.S.A., $23.00 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 Instutution, 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. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 1-7 NEPHTYS CRYPTOMMA, NEW SPECIES (POLYCHAETA: NEPHTYIDAE) FROM THE NORTHERN GULF OF MEXICO Donald E. Harper, Jr. Abstract.—A new species of polychaetous annelid, Nephtys cryptomma, is described from the northern Gulf of Mexico. The species is most abundant on sandy bottoms and appears to reproduce during the spring. The species described herein as Nephtys cryptomma was first collected in 1978 dur- ing a benthic study conducted off Freeport, Texas. Specimens were subsequently col- lected off Louisiana in 1978-79 during the Central Gulf Platform Study (CGPS) con- ducted for the Bureau of Land Manage- ment, and off Cameron in 1983. The spec- imens were initially identified as Nephtys magellanica Augener (1912:208), based on the description by Hartman (1968:587), es- pecially because of obvious eyespots visible in young specimens. However, more de- tailed examination of the specimens and comparison with the redescription of the syntypes of N. magellanica from the Straits of Magellan by Perkins (1980:34) revealed several differences. The specimens were thus designated Nephtys “‘subdermal eyes” while the description was in progress. The type and additional specimens of Nephtys cryptomma have been deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). Other specimens were deposited in the Texas A&M University, Department of Oceanography reference collection, Col- lege Station (TAMU), and the Texas A&M Marine Laboratory reference collection, Galveston (AMML). Nephtys cryptomma, new species Figs. 1—4 Material examined. —TEXAS, Freeport, 19 km offshore, 28°44’N, 95°15’W, sandy to muddy sand bottoms, 21-m depth, 3 Apr 1980, Holotype (USNM 67778): 4 para- types (USNM 67779), 2 paratypes (AMML PLY-24); 24 Feb 1978, 1 adult (USNM 67780); 16 Dec 1979, 1 adult (TAMU 1-2289); 13 Feb 1980, 1 young (USNM 67781), 1 young (TAMU 1-2290); 10 Mar 1980, 7 young (AMML PLY-25); 30 Jun 1980, 1 young (USNM 67782); 24 Jul 1980, 3 young (AMML PLY-28); 25 Aug 1980, 3 young (USNM 67783); 17 Apr 1981, 1 adult (AMML PLY-27); Freeport, 9 km offshore, 28°41'N, 95°17'W, muddy bottom, 16-m depth, 28 Jan 1980, 1 young (USNM 67784). LOUISIANA, Grand Isle, 27 km off- shore, 29°02’N, 90°09'W, muddy sand bot- tom, 13-m depth; 27 May 1978, 1 young (AMML PLY-29); Grand Isle, 54 km SSW offshore, 28°39'N, 90°14’W, sandy to silty sand bottom, 36-m depth; 26 Aug 1978, 4 young (USNM 69962), 1 adult (TAMU 1-2291), 1 young (AMML PLY-26). VIRGINIA, York River, 3 to 10-m depth, Jan—Mar 1961, 4 adults (USNM 33327); Nov 1960, 1 adult (USNM 33326), 3-6 m; Mar 1961, 1 adult, 6 young (USNM 33328). FLORIDA, Seahorse Key, 6 Feb 1960, 3 adults (USNM 33330). Diagnosis.—Prostomium squarish ante- riorly, tapering posteriorly, with paired an- terolateral and ventrolateral antennae; mid- dorsal pigment spot. First or tentacular segment extended lateral to prostomium with weakly developed setigerous parapo- dia bearing small dorsal tentacular cirri and 2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1.0mm 1.0 mm b Fig. 1. 1.0mm 0.5mm d Nephtys cryptomma (holotype): a, Anterior end, dorsal view; b, Distal end of everted proboscis, dorsal view; c, Anterior end of juvenile showing location of eyespots and brain (dotted); d, right parapodium 3, anterior view (setae not shown). larger ventral tentacular cirri. Proboscis barrel-shaped, with 22 subterminal longi- tudinal rows of papillae in addition to long tapering middorsal and shorter midventral papillae. Parapodia biramous, with acicular lobes conical; notopodial presetal lamellae shorter than acicular lobes, with shallow ex- cavations; postsetal lamellae broadly oval, longer than acicular lobes. Branchiae sickle- shaped, beginning on setiger 3, terminating about 3 segments from pygidium. Dorsal cirri begin on setiger 2, small in anterior segments, becoming elongate, thin, strap- like, almost as long as branchiae by mid- body, continuing to posterior end. Preacicu- lar setae barred capillaries, postacicular se- tae longer flowing capillaries with mostly fine teeth. Pygidium with dorsal anus and single anal cirrus. Description. —Prostomium squarish an- teriorly, tapering posteriorly, with pair of antennae at anterior corners, pair of ven- trolateral antennae just anterior to neuro- podia of peristomium, pair of nuchal organs at posterolateral angles, and medial spot or streak of reddish-brown pigment (Fig. 1a). Peristomial or tentacular segment with bira- mous parapodia directed anteriorly; noto- podia with presetal lamellae smaller than acicular lobes, postsetal lamellae slightly larger, dorsal tentacular cirri small, noto- setae directed anterodorsally; neuropodia with weakly developed lamellae and larger tentacular cirri, neurosetae directed ante- VOLUME 99, NUMBER 1 0.5mm 0.5 mm 0.5mm d Fig. 2. Nephtys cryptomma: a, Right parapodium 10, anterior view; b, Right parapodium 38, anterior view; c, Right parapodium 50, anterior view; d, Right parapodium 70, anterior view. riorly (Fig. la). Mouth flanked by lateral lobes. Subterminal part of everted proboscis with long, tapering, middorsal papilla, shorter midventral median papilla and 22 longitudinal rows of conical papillae, usu- ally 4-6 per row, decreasing in size proxi- mally; on both dorsal and ventral surfaces, second row of papillae on either side of mid- line with only 2 (Sometimes 1) papillae; oc- casionally some ventral rows merge in V-shape; terminal part of proboscis with | 1 pairs of bifid papillae (Fig. 1b). Internal deep lateral grooves begin at proximal end and terminate about one-quarter of length be- fore distal end; single brown tooth in ter- minus of each groove. Pair of black eyespots on posterolateral surface of brain, visible through integument of juvenile and young 4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON specimens (Fig. 1c), but obscured by more opaque integument in adults. Notopodia of setiger 2 with small pre- and postsetal la- mellae and small dorsal cirri. Parapodia ful- ly developed from setiger 3 (Fig. 1d), reach- ing maximum size about setiger 10 (Fig. 2a). Acicular lobes conical or, on few anterior segments, slightly bilobed. Notopodial pre- setal lamellae shorter than acicular lobes, with shallow excavations in anterior seg- ments, becoming more pronounced toward mid-body; postsetal lamellae longer than acicular lobes, broadly oval, without exca- vations (Figs. 2a—d). Neuropodial presetal lamellae shorter than acicular lobes, oval; postsetal lamellae much larger than acicular lobes, oval. From mid-body posteriorly, la- mellae gradually reduced in size, and quite small in far posterior segments. Interramal branchiae present from setiger 3 (Fig. 1d), well-developed in mature specimens, large, recurved on all segments, with small conical projections just below insertion of dorsal cirri (Figs. 1d, 2a—d). Dorsal cirri attached to dorsal part of notopodia, small anterior- ly, attaining maximum size in anterior third of body and continuing posteriorly; when fully developed in mid-body and posterior segments, thin, lanceolate (Figs. 2a—d). Ven- tral cirri present from peristomial segment. Setae all capillaries of 2 basic types forming 2 fan-shaped rows in both rami. Preacicular setae shorter, straighter, appearing cross- barred internally (Fig. 3a). Postacicular se- tae longer, flowing, spinous; most neuro- setae finely spinous along entire blade, but 2 to 3 in middle of row with few large basal teeth in addition to fine spines (Fig. 3b); some upper neurosetae appearing smooth (Fig. 3c). Notosetae directed dorsolaterally, neurosetae directed ventrolaterally. Pygidi- um rounded, enclosed in posterior small segments, with dorsal anus and single ter- minal anal cirrus (Fig. 3d). Etymology. —The specific epithet is from the Greek “‘kryptos’”’ (hidden) and ‘“‘omma’”’ (eye) referring to the deep-set eyespots which are evident in juveniles, but hidden beneath integument in adults. Size range.—The holotype is a complete ovigerous female, 43 mm long, 3 mm wide (including parapodia), having 82 fully formed segments, followed by 3 incomplete ones, the last represented by setae only. The largest specimen, complete but broken, is 65 mm long, 4 mm wide, with 98 segments, including 2 incomplete ones (USNM 67780). Color. —Living worms are cream-col- ored, except for the nearly transparent pos- terior few segments and anal cirrus, reddish- brown prostomial spot and black eyes. Setae and tips of acicula dark golden. Blood red. Gut visible as dark area from about setiger 16 to end of body. Preserved worms are uniformly opaque white to pale yellow, ex- cept for the prostomial spot. Distribution.—The species is presently known from the northern and eastern Gulf of Mexico (Florida, Louisiana, and Texas) and Virginia. Remarks. —The dark prostomial spot could cause N. cryptomma to be mistaken for other species. Nephtys simoni Perkins (1980:37) has the spot and eyespots, but the paratypes (USNM 55684) have slightly bi- lobed acicular lobes rather than conical lobes, rounded notopodial presetal lamellae with the lamellae smaller than in N. cryp- tomma, and slightly bilobed postsetal lobes. Nephtys parva Clark and Jones (1955:143) has the pigment spot and eyes, and entire acicular lobes, but has reduced pre- and postsetal lamellae and branchiae beginning on setiger 4. Nephtys magellanica, the species with which Nephtys cryptomma was originally confused, lacks the prostomial pigment spot, has bilobed acicular lobes, shorter presetal lobes, and only 20 rows of papillae on the proboscis. Several lots of specimens collected from the Chesapeake Bay area by Wass (1965:16; USNM 33326-28, 38738) and Seahorse Key, Florida, by Taylor (1971:103-104; USNM 33330) were examined. These spec- VOLUME 99, NUMBER 1 imens, originally identified as N. magellan- ica were re-examined by Perkins (1980), de- termined to be similar to N. hombergii Savigny (1818:314) and labelled N. cf. hom- bergi. Perkins (1980:41—42) stated that these specimens differed from N. simoni in lack- ing dorsal cirri on segment 1, yet they all bear small papilla-like dorsal cirri on the tentacular segment, arising from the lateral surface of the parapodia. In some cases the cirri are very small and inconspicuous, causing the parapodium to appear bilobed. Furthermore, the dorsal presetal lamellae have shallow excavations as in N. cryptom- ma rather than the deeply cleft lamellae as in N. hombergii (Fauvel 1923:367-368, fig. 143c, d; Fauchald 1963:12, fig. 3E). These specimens have therefore been referred to N. cryptomma. Nephtys cryptomma is probably more abundant in the eastern Gulf of Mexico than the single record indicates. Re-examination of the specimens identified as N. simoni col- lected during the MAFLA study for the Bu- reau of Land Management (Taylor 1984:9) may show that some of the individuals are actually N. cryptomma. Ecological notes.—Nephtys cryptomma has been collected most frequently on sandy and muddy sand bottoms. Depths ranged from 16 m off the Texas coast to 36 m off Louisiana. The species was never a numer- ically dominant member of its assemblage. During a 6-year (1978-1983) benthic study off Freeport, Texas, in which 15 stations were sampled at each of two sites, the max- imum monthly abundance was 12 individ- uals at the deeper sandy bottom site. Data from both the Freeport and CGPS studies indicate that N. cryptomma is a spring breeder. The few ovigerous females ob- tained were collected in the spring. Off Free- port, only a few specimens were collected in 1978 and 1982. In 1979-1981, maxi- mum abundances occurred in early spring, followed by decreases through summer, while in 1983 the largest numbers were col- 0.1 mm 1.0 mm Fig. 3. Nephtys cryptomma: a, Barred preacicular capillary seta: inset showing details of cross-barring, highly magnified; b, Spinous postacicular neuroseta from middle of row; c, Smooth postacicular neuroseta from lower part of row; d, Posterior end and pygidium. 6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 14 We ay 1978-1979 ® 10 I aie a ea Sao 1980-1981 = Bilas deceh bes «ogee wacses 1982-1983 a > 6 Q Z 4 2 BA Ne at J oF M AM J J Aes ONE Ded F M AM JJ AS ON D Fig. 4. Seasonal distribution of Nephtys cryptomma off Freeport, Texas, 1978-1983 on sandy bottom study area in 21-m depth. lected in late spring (Fig. 4). The rapid de- cline in June 1979 coincided with the oc- currence of hypoxic, hydrogen sulfide-laden water (Harper et al. 1981) and was probably not indicative of the usual seasonal trend. During the 3-season, 36-station CGPS study off Louisiana in 1978-79, N. cryptomma abundances were: May—June— 84; August-— September—9; January—8, which agrees with the seasonal data collected off Free- port. Notes on living worms.—Living worms were collected on 3 April 1980 from the type-locality off Freeport, Texas, and ob- served in the laboratory in a finger bowl. The worms were generally passive, lying on their backs. When prodded they swam vig- orously by undulating the posterior three- quarters of the body. The proboscis was never everted unless the worm was im- mersed in 70% ethanol. Cilia on the branchiae and interramal body walls created water currents that flowed posteriorly in the interramal channel. The branchiae were kept away from the body wall in this current, and were frequently snapped backward, either singly or in uni- son, which dislodged small adhering parti- cles. The mid-dorsal longitudinal blood vessel was mostly obscured by musculature. It emerged from the musculature in the vicin- ity of setiger 25, and disappeared again at the base of the prostomium. Between setiger 25 and 15 the vessel was large, contractile and fixed in position. From setiger 15 to 9 the position of the vessel was not fixed, and it moved from side to side as the worm flexed. From setiger 9 to the base of the prostomium the vessel was fixed in the dor- sal midline. Paired contractile longitudinal ventral vessels, lying alongside the ventral nerve cord, were visible from about setiger 4 to the pygidium; they were largest ante- riorly. Blood flowed posteriorly in the right vessel and anteriorly in the left. Lateral ves- sels were small, forming networks of vessels in the body wall posterior to the parapodia, and in the branchiae (Fig. 2d). The gut was visible as a dark line from about setiger 16 to the posterior end of the body. Acknowledgments I thank the numerous graduate and un- dergraduate students who participated in the cruises off Freeport and sorted the samples, and particularly my associates Larry McKinney and Robert Salzer, who analyzed most of the benthic samples. This research VOLUME 99, NUMBER 1 was sponsored by the Department of Energy Contract No. DE-FC96-79P010114 admin- istered by the Texas A&M Research Foun- dation, directed by Roy W. Hann, Jr., and Robert E. Randall, both of the Environ- mental Engineering Division, Civil Engi- neering Department, Texas A&M Univer- sity. The Central Gulf Platform Study was supported by Bureau of Land Management Contract AA551CT817 to Southwest Re- search Institute, Houston, Texas, managed by C. A. Bedinger, Jr. Literature Cited Augener, H. 1912. Beitrag zur Kenntnis verschie- dener Anneliden und Bemerkungen tiber die Nephtys-Arten und deren epitoke Formen.— Archiv fur Naturgeschichte 87A(10):162-212. Clark, R. B.,and M. L. Jones. 1955. Two new Neph- tys (Annelida: Polychaeta) from San Francisco Bay.—Journal of the Washington Academy of Science 45:143-146. Fauchald, K. 1963. Nephtyidae (Polychaeta) from Norwegian waters.—Sarsia 13:1-32. Fauvel, P. 1923. Polychétes errantes.—Faune de France 5:1-488. Hartman, O. 1968. Atlas of the errantiate polychae- tous annelids from California. — Allan Hancock Foundation, University of Southern California 828 pp. Harper, D. E., Jr., L. D. McKinney, R. R. Salzer, and R. J. Case. 1981. The occurrence of hypoxic bottom waters off the upper Texas coast and its effect on the benthic biota.—Contributions in Marine Science 42:53-79. Perkins, T. 1980. Review of species previously re- ferred to Ceratonereis mirabilis and descrip- tions of new species of Ceratonereis, Nephtys and Goniada (Polychaeta). — Proceedings of the Biological Society of Washington 93:1-49. Savigny, J.C. 1818. Annélides. Jn Lamarck, J. B. de, Histoire Naturelle des Animaux sans Vertebrés 5:1-612. Paris. Taylor, J. H. 1971. Polychaetous annelids and ben- thic environments in Tampa Bay, Florida. Ph.D. Thesis, University of Florida 1,332 pp. 1984. Family Nephtyidae Grube, 1850. In Polychaetes of the northern Gulf of Mexico, J. M. Uebelacker and P. G. Johnson (eds). Vol. 5, Chapt. 35, pp. 1-20. Final report to Minerals Management Service, contract 14-12-001- 29091. Barry A. Vittor & Assoc., Inc., Mobile, Alabama. Wass, M. L. 1965. Check list of the marine inver- tebrates of Virginia.—Special Scientific Report Virginia Institute of Marine Science 24:1-55. Texas A&M Marine Laboratory, Bldg. 311, Ft. Crockett, Galveston, Texas 77550. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 8-14 NEW SOUTH AMERICAN GASTROPODS IN THE GENERA CONUS (CONIDAE) AND LATIRUS (FASCIOLARIIDAE) Edward J. Petuch Abstract. — Additions to the marine gastropod fauna of South America, in- cluding six new species of Conus (Conidae) and a new Latirus (Fasciolariidae), are described. Two of the new cones, Conus carioca n. sp. and C. riosi n. sp., range along the northern and eastern coasts of Brazil, while two others, C. tostesin. sp. and C. xanthocinctus n. sp., are restricted to the Patagonian region of southern Brazil and Uruguay. Two northern South American cones are also described: C. gibsonsmithorumn. sp. from the Paraguana Peninsula, Venezuela, and C. penchaszadehi n. sp. from the Goajira Peninsula of Colombia. The new fasciolariid, Latirus vermeiji n. sp., is endemic to Fernando de Noronha Island off northern Brazil. The northern and eastern coasts of South America, in particular Venezuela, Colom- bia, and Brazil, have recently been shown to harbor atypical Caribbean molluscan faunas with archaic appearances (Vermeij 1978:231-235; Petuch 1981). Through the works of Venezuelan authors, such as Gib- son-Smith, Penchaszadeh, Princz, and Flo- res, the gastropod fauna of that coastline is now becoming better known. The Brazilian coast, on the other hand, is still poorly known, with the compendium of Rios (1975) being the only comprehensive faunal guide to that area. Through the kindness of several South American malacologists, including Sr. Luiz Roberto Tostes, Rio de Janeiro, and Prof. E. C. Rios, Rio Grande do Sul, Brazil, and Dr. Pablo Penchaszadeh, Universidad Si- mon Bolivar, and Dr. and Mrs Gibson- Smith, Caracas, Venezuela, I was given study material of interesting new species from their respective countries. These unusual new gastropods include, amongst others, six new species of Conus. Dr. Geerat Vermeij, Uni- versity of Maryland, has also kindly donat- ed a number of specimens of a new Latirus from Fernando de Noronha Island off the northern Brazilian coast. These were col- lected during a research trip to the island in 1968. These important new additions to the fauna of South America are described here. Gastropoda Neogastropoda Fasciolariidae Latirus Montfort, 1810 Latirus vermeiji, new species Figs. 3, 4 Material examined.—Holotype: Length 25 mm, width 13 mm, on rocks at low tide line, south coast of Fernando de Noronha Island, Brazil, 1968, number 14243, type collection of the Museu Oceanografico Cen- tro de Ciencias do Mar, Funda¢ao Univer- sidade do Rio Grande, Brazil; Paratypes: length 27 mm, same locality and date as holotype, collection of the Division of Mol- lusks, National Museum of Natural Histo- ry, Smithsonian Institution, USNM 784685; three specimens, lengths 15—26 mm, collec- tion of Geerat J. Vermeij, Department of Zoology, University of Maryland. Description. —Shell fusiform in outline, spire protracted; siphonal canal short for VOLUME 99, NUMBER 1 genus; adults with 8 whorls; sculpture con- sisting of 8—10 large spiral cords with finer threads in between; whorls with 8—9 prom- inent axial ribs; shoulder sharply angled, producing faint knob at posterior end of each rib; columella with 3 large plications; outer lip with numerous lirae; lirae extending into aperture; siphon slightly umbilicate; shell uniformly deep orange colored; interior of aperture white; periostracum and opercu- lum dark brown. Etymology.—Named for Dr. Geerat J. Vermeij, of the University of Maryland, who collected the type material on Fernando de Noronha. Distribution.—Endemic to the island of Fernando de Noronha off northern Brazil. Discussion. —This new species somewhat resembles a stumpier version of the Abro- Ihos Islands endemic, Latirus ogum Pe- tuch, 1979, but differs in having a much shorter siphonal canal and more angled shoulder. In these last two characters, L. vermeiji resembles a small Leucozonia species. The new species is a true Latirus, however, in that it lacks the tooth on the outer lip and the color banding of that fas- ciolariid genus. Interestingly enough, Lati- rus vermeiji is morphologically closer to the Panamic L. socorroensis Hertlein and Strong, 1951, which is also endemic to off- shore island groups. Conacea Conidae Conus Linnaeus, 1758 Conus carioca, new species Figs. 1, 2 Material examined.—Holotype: Length 52 mm, width 24 mm, trawled by com- mercial fishermen from 100 m depth off Cabo Frio, Rio de Janeiro State, Brazil, 1975, collection of the Museu Oceanogra- fico de Fundacao Universidade de Rio Grande do Sul #20-915; Paratypes: length 45 mm, same locality and depth as holo- type, Museu Oceanografico #20-915; 2 specimens, lengths 53 mm and 53 mm, trawled by Brazilian fisheries research ves- sel, from 150 m depth off Recife, Pernam- buco, Brazil, 1968, Museu Oceanografico #14-242; length 57 mm, same depth and locality as holotype, collection of the Di- vision of Mollusks, National Museum of Natural History, Smithsonian Institution USNM 784686. Description. —Shell elongate, straight- sided, smooth and shiny, shoulder sharp- edged, carinated; spire flattened but slightly protracted in early whorls; color white with revolving bands of bright orange and or- ange-pink; orange bands overlaid with rows of pale brown dots and dashes; mid-body with white band; central white band bor- dered on both sides by bands of dark brown flammules; spire pale orange with crescent- shaped brown flammules; interior of aper- ture pale salmon; periostracum thin, smooth, translucent brown; operculum small, oval in shape. Etymology.—Named for the “‘cariocas,”’ the inhabitants of Rio de Janeiro. Discussion. —Conus carioca most closely resembles the deep water, slope species C. villepini Fischer and Bernardi, 1857. The new continental shelf species differs from its slope-inhabiting relative in being more elongate, in having a much flatter spire, and in being a much more colorful shell, with bands of bright orange and salmon-pink. Conus gibsonsmithorum, new species Figs. 5, 6 Material examined.—Holotype: Length 20 mm, width 18 mm, trawled by com- mercial shrimp boats from 35 m depth off north coast of the Paraguana Peninsula, Fal- con State, Venezuela, 1978, Museu Ocean- ografico #14244; Paratype: length 18 mm, same depth and locality as holotype, USNM 784687. Description. —Shell turnip-shaped, obese, thick and heavy; anterior one-third of shell greatly constricted; body whorl smooth and 10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON shiny, with anterior one-third having heavy grooving and thick spiral cords; aperture very narrow; spire elevated on early whorls, becoming planar on later whorls; shoulder sharp-edged, carinated; slight constriction just below shoulder carina; shell pale cream- yellow colored with pale tan band around middle; spire pale yellow, becoming tan on early whorls; periostracum thin, smooth, translucent yellow. Etymology.—Named for Dr. and Mrs. Jack Gibson-Smith (Jack and Winifred) of Caracas, Venezuela, in recognition of their contributions to Venezuelan malacology. Discussion. — This small new species is the only South American cone shell to have a squat, turnip-shaped body form. In this re- spect, C. gibsonsmithorum most closely re- sembles C. sennottorum Rehder and Ab- bott, 1951, from the Gulf of Mexico. The new Venezuelan species differs from its northern relative by lacking any spottings or color patterns, by having a more sharply carinated shoulder with sub-shoulder con- striction, and by being a smaller, stockier species. At present, C. gibsonsmithorum is known only from the Gulf of Venezuela region, to which it is most probably endemic. Conus penchaszadehi, new species Figs. 13, 14 Material examined.—Holotype: Length 18 mm, width 9 mm, trawled by commer- cial shrimpers from 35 m depth off Cabo La Vela, Goajira Peninsula, Colombia, 1974, Museu Oceanografico #14245. Description. —Shell elongate, thin, fragile; spire elevated; body whorl shiny, totally covered with numerous fine spiral threads; spiral threads becoming coarser at anterior end; shell pinkish-white with scattered or- ange flammules on body whorl; solid, dark orange band around anterior one-third of body whorl; anterior tip bright pinkish-or- ange; spire white with crescent-shaped or- ange flammules; interior of aperture pale salmon-pink; protoconch and early whorls bright orange; periostracum thick, brown, with rows of fine tufts. Etymology. —Named for Dr. Pablo Pen- chaszadeh, Department of Biology, Simon Bolivar University, Caracas, Venezuela. Discussion.—This distinctive little shell is unlike any other northern South Ameri- can cone. Conus penchaszadehi may be re- lated to C. atractus Tomlin, 1937, but dif- fers from that species by having the bright orange color band and flammules and by having finer spiral sculpture. The new Ven- ezuelan species, however, closely resembles pale color forms of the Panamic C. orion Broderip, 1833, and may be the Caribbean cognate species. Conus penchaszadehi is only known from off the Goajira Peninsula of Colombia, but it most probably ranges all along the Co- lombian coast and into the Gulf of Vene- zuela. Conus riosi, new species Fig. 7, 8 Material examined.—Holotype: Length 54 mm, width 32 mm, trawled from 50 m depth off Salvador, Bahia State, Brazil, by Brazilian fisheries research vessel R/V RIOBALDO, 1975, Museu Oceanografico #18757; Paratypes: length 37 mm, same lo- cality and depth as holotype, Museu Ocean- ografico #14242. Description. —Shell thick, heavy, wide- shouldered, tapering toward anterior end; spire completely flattened with only first few whorls being protracted; body whorl and spire shiny with waxy feel; shoulder sharp- edged with prominent carina; shells varying in color from yellow to orange with nu- merous crowded vertical flammules of dark brown or reddish-brown; mid-body with clear white band; white band bordered by, or sometimes covered by, band of dark brown checkers; second band of dark flam- VOLUME 99, NUMBER 1 mules around anterior end; anterior tip bright apricot-orange; spire orange with nu- merous crescent-shaped dark brown flam- mules; interior of aperture pale orange; peri- ostracum thin, translucent yellow-brown, with scattered tufts of large hairs. Etymology. —Named for Prof. E. C. Rios, of the Fundacao Universidade do Rio Grande, in recognition of his invaluable works on the Brazilian molluscan fauna. Discussion. —This new species is closely related to, and is often confused with, the common, widespread Conus daucus Hwass, 1792. The new Brazilian species differs from its Caribbean relative by being a larger, heavier shell, by having a different color pattern with a prominent white or brown central band, and by having an undulating suture on the spire whorls. Conus riosi also prefers deeper waters along the southern coast of Brazil, while C. daucus generally prefers shallow water reef areas around the Caribbean region. Of the specimens of C. riosi examined, none had the deep red-or- ange color or the rows of tiny spots seen on C. daucus. Conus daucus may range only into north- ern Brazil, since many of the records of that species from further south have turned out to be C. riosi. The ““C. daucus”’’ illustrated by Van Mol, Tursch, and Kempf (1967: plate 7, fig. 2) and that of Rios (1975: Fig. 542) both appear to be referable to the new species. The Patagonian Conus Species The last two new cones described here belong to a distinctive and close-knit species group that is restricted to the cold water Patagonian region, from Santa Catarina, Brazil, to south of Mar del Plata, Argentina. Included in this group are Conus clenchi Martins, 1943, C. tostesin. sp., and C. xan- thocinctus n. sp., from southernmost Brazil, and C. carcellesi Martins, 1945, C. iheringi Frenguelli, 1946, and C. platensis Fren- 11 guelli, 1946 from Uruguay and Argentina. All species in this group have highly pol- ished shells with rounded shoulders, rela- tively high to protracted spires, and large, mamillate protoconchs. Because of its sharp- angled shoulder and small protoconch, C. clerii Reeve, 1844, which is sympatric with C. clenchi, C. tostesi, and C. xanthocinctus along the southern Brazilian coast, does not appear to belong to this group. Instead, C. clerii appears to be related to southern Ca- ribbean species such as C. undatus Kiener, 1848, C. cingulatus Lamarck, 1810, and C. centurio Born, 1780. The six Patagonian cones represent rem- nants of a large cold water molluscan fauna that extended down a large part of the coast of Argentina during the Late Pliocene (Frenguelli 1932). This fauna was typified by conid species such as C. patagonicus Frenguelli, 1932, and a large number of in- teresting volutacean genera that includes Minicymbiola, Odontocymbiola, Zidona, Adelomelon, Pachycymbiola, Weaveria, Orbignytesta, Olivancillaria, and Olivina, and a remarkable trophonine (Muricidae) fauna. Descendants of these distinctive species radiations are still extant along the Patagonian coast, with the greatest number of species being found from southernmost Brazil to, and into, the Mar del Plata. In the Recent, the descendant Patagonian cones represent the southernmost limit of the family in the Americas. Conus tostesi, new species Figs. 9, 10 Material examined.— Holotype: Length 35 mm, width 18 mm, trawled by com- mercial fishermen from 100 m depth off Cabo Frio, Rio de Janeiro State, Brazil, 1977, Museu Oceanografico #14246; Para- types: length 29 mm, same locality and depth as holotype, Museu Oceanografico #14247; length 27 mm, USNM 784688. Description. —Shell elongate, thin fragile; 12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-16. Species of Latirus and Conus from South America. 1, 2, Conus carioca, dorsal and ventral aspects of holotype; 3, 4, Latirus vermeiji, dorsal and ventral aspects of holotype; 5, 6, Conus gibsonsmithorum, dorsal and ventral aspects of holotype; 7, 8, Conus riosi, dorsal and ventral aspects of holotype; 9, 10, Conus tostesi, dorsal and ventral aspects of holotype; 11, 12, Conus xanthocinctus, dorsal and ventral aspects of holotype; 13, 14, Conus penchaszadehi, dorsal and ventral aspects of holotype; 15, Conus clerii Reeve, 1844, specimen from 100 m off Cabo Frio, Rio de Janeiro State, Brazil; 16, Conus clenchi Martins, 1943, specimen from 100 m off Cabo Frio, Rio de Janeiro State, Brazil. VOLUME 99, NUMBER 1 spire protracted, slightly scalariform; body whorl shiny, with anterior one-third cov- ered with numerous fine spiral threads; shoulder produced but slightly rounded; color pale violet to darker violet with three wide bands of reddish-brown, one just be- low shoulder, one around mid-body, and One around anterior end; banded color pat- tern overlaid by 10-14 rows of brown dots; anterior tip darker violet on some speci- mens; spire white with numerous crescent- shaped flammules; protoconch large, mam- illate; periostracum thin, smooth translu- cent yellow. Etymology.—The new taxon honors Sr. Luiz Roberto Tostes of Rio de Janeiro, Bra- zil, who kindly donated large amounts of material for study. Discussion. — Conus tostesi is closest to the sympatric C. clerii Reeve, 1844, but differs in being a much smaller, more elongate shell, by having a higher, scalariform spire, by being of a violet color instead of white, and by having a much larger, mamillate proto- conch. A typical specimen of C. clerii, from the type locality of C. tostesi, is illustrated here for comparison (Fig. 15). This new Brazilian species actually shows a closer affinity to some of the rare Paoli- nian-Submagellanic species such as C. car- cellesi Martins, 1945 and C. platensis Fren- guelli, 1946 from the Mar del Plata (Frenguelli 1946). Conus tostesi differs from both of these species, however, by having a three-banded color pattern, finer body sculpture, and by lacking spiral grooves on the spire. Conus xanthocinctus, new species Figs. 11, 12 Material examined.—Holotype: Length 47 mm, width 22 mm, trawled by com- mercial fishermen from 100 m depth off Cabo Frio, Rio de Janeiro State, Brazil, Mu- seu Oceanografico #14248; Paratype: Length 45 mm, same locality and depth as holo- type, Museu Oceanografico #14249. 13 Description. —Shell elongate, slender, thin and fragile; spire protracted, stepped; body whorl and spire smooth, shiny; shoulder only slightly produced, rounded, anterior end of shell with few weak spiral striae; aperture narrow; shell color bright golden-yellow with three darker, orange-yellow bands, one just below shoulder, one around mid-body, and one around anterior end; mid-body band darkest, deep orange colored; bands over- laid with 12-14 spiral rows of brown dashes and scattered white flammules; spire gold- en-yellow with numerous crescent-shaped tan flammules; shoulder and suture of spire whorls ornamented with bands of alternat- ing dark tan and white flammules; interior of aperture pale golden colored; periostra- cum thin, smooth, translucent yellow; oper- culum small, oval. Etymology. —““Yellow-belted,”’ in refer- ence to the characteristic and distinctive bright orange-yellow band around the mid- body. Discussion. — This distinctive new species could only be confused with the sympatric C. clenchi Martins, 1943, and then only in general shell shape. The bright golden color bands and characteristic shoulder color- ation readily separates C. xanthocinctus from C. clenchi. A typical specimen of C. clenchi, from the type locality of C. xan- thocinctus, is illustrated here for compari- son (Fig. 16). Acknowledgments I thank Dr. M. G. Harasewych, Division of Mollusks, Smithsonian Institution, for taking the excellent photographs used in this paper. Literature Cited Frenguelli, Joaquin. 1932. Nuevos elementos faunis- ticos del Patagoniano de Comodoro Rivadavia (Chubut).— Boletin de Informacion de Petroleos (Buenos Aires) 9(98):785—790. 1946. Especies del genero Conus vivientes en el litoral Platense y fésiles en el Neozoico Superior Argentino-Uruguayo.— Notas del Mu- seo de la Plata 11 (Paleontology 33):231-—250. 14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Petuch, Edward J. 1981. A relict Neogene caenogas- Campagnes de la “Calypso” (Mollusques Proso- tropod fauna from northern South America. branches) 16:233-254. Malacologia 20(2):307-347. Vermeij, Geerat J. 1978. Biogeography and adapta- Rios, E.C. 1975. Brazilian marine mollusks iconog- tion. Harvard University Press, Cambridge, raphy. Fundacao Universidade do Rio Grande, Massachusetts. Pp. 1-332. Centro de Ciéncias do Mar, Rio Grande do Sul, Brazil. Pp. 1-331, 91 plates. Department of Geology, Florida Inter- Van Mol, J. -J., B. Tursch, and M. Kempf. 1967. Les Conidae du Brésil.— Résultats Scientifiques des national University, Miami, Florida 33199. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 15-16 THE AUSTRAL-AFRICAN CONID SUBGENUS FLORACONUS IREDALE, 1930, TAKEN OFF BERMUDA (GASTROPODA: CONIDAE) Edward J. Petuch Abstract. —Baited lobster traps have recently been used to sample the deep water molluscan fauna of the slope of Bermuda. Among the many unusual gastropods collected was the first known northern Atlantic member of the Austral-African conid subgenus Floraconus Iredale, 1930. The discovery of this species, which is geographically widely separated from related forms and is here named Conus (Floraconus) lightbourni new species, points to the relict nature of the subgenus. Mollusks from the deep waters around Bermuda have recently been taken by a nov- el new collecting technique. This method, undertaken by the well known Bermudan malacologists Messrs. John R. H. Light- bourn and Arthur T. Guest, involves drop- ping baited lobster traps into deep water and leaving them there for a number of days. At the end of that time, the traps are retrieved and the hermit crab-occupied shells are col- lected. Since dredging on the steep slope of the Bermuda seamount is difficult, this sim- ple method is proving itself to be the best way to explore the unknown slope fauna. To date, a large number of interesting new gastropod shells, all occupied by hermit crabs, have been collected from the lobster pots. These include unusual species such as the newly described Pterynotus lightbourni Harasewych and Jensen, a number of Pleu- rotomaria (Perotrochus) species, the “‘Ha- waiian endemic” Gyrineum loisae Lewis, the eastern and southern Atlantic Gyrineum olearium Linnaeus (Jensen, pers. comm.), and the first known northern Atlantic species of the Austral-African conid subgenus Fior- aconus Iredale, 1930. All of these elements give the Bermudan slope fauna a distinctly Indo-Pacific and Japonic appearance. The subgenus Floraconus was previously thought to be restricted to the southern and eastern shores of Australia and to the Cape Province region of South Africa. Because of this distributional pattern, this conid sub- genus may represent a Paleogene relict group, much like the sympatric Austral-Af- rican relict volutid genus Ath/eta (Petuch 1981:1127). The presence of Floraconus in the northern Atlantic indicates that the group was once very widespread but has survived into the Recent in only three widely sepa- rated areas. Figs. 1, 2. Dorsal and ventral aspects of the holo- type of Conus (Floraconus) lightbourni, from 497 m off Bermuda. 16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The new Floraconus is the only known truly endemic Bermudan cone shell. A large, common, shallow water species, named Co- nus bermudensis by Clench in 1942, was originally thought to be endemic to the is- land but has now been found to be conspe- cific with the widespread western Atlantic C. mindanus Hwass, 1792 (Walls 1979:726— 730). Along with C. mus Hwass, 1792, C. mindanus, and C. villepinii Fischer and Ber- nardi, 1857, the new Floraconus brings the total number of known Bermudan cone species to four. This unexpected mid-At- lantic member of an Austral-African group is described here. Gastropoda Neogastropoda Conacea Conidae Conus Linnaeus, 1758 Floraconus Iredale, 1930 Conus (Floraconus) lightbourni, new species Figs. 1, 2 Material examined.—Holotype: Length 35 mm, width 16 mm, occupied by hermit crab, taken in lobster pot from 180 fathoms (approx. 497 m), 12 miles (2.5 km) due south of Castle Island, Bermuda, July 1973, by J. R. H. Lightbourn; collection of the Department of Malacology, Delaware Mu- seum of Natural History, DMNH 134938; Paratypes: 3 specimens, lengths 26.0 mm— 47.7 mm, same depth and locality as ho- lotype, DMNH 134939; 5 specimens, lengths 22.4 mm—44 mm, collection of Mr. J. R. H. Lightbourn, Bermuda. Description. —Shell elongate, slender, ob- conical, thin, lightweight; spire elevated; shoulder and spire whorls rounded; body whorl and spire shiny, polished; anterior end of body whorl with numerous faint, raised spiral threads; aperture narrow, straight; base shell color bright orange, overlaid with two wide bands of deep salmon-pink, one at mid- body, one between mid-body and shoulder; some specimens with third salmon band around anterior end; salmon-pink bands or- namented with rows of large brown spots; spots often coalesce into large brown flam- mules, usually on either side of central band; some specimens with rows of small brown dots on orange bands; spire salmon-pink with large crescent-shaped brown flam- mules; protoconch mamillate; interior of aperture pale violet-purple; operculum and periostracum unknown. Etymology. —Named for Mr. John R. H. Lightbourn of Bermuda, in recognition of his many important discoveries of new deep water Bermudan mollusks. Remarks. —The narrow, highly polished body and rounded shoulder of C. Jight- bourni readily separates the new species from any other known western Atlantic cone shell. The new species is closest to the South Af- rican C. (Floraconus) pictus Reeve, 1843, in color pattern, but is more like the eastern Australian C. (Floraconus) wallangra (Gar- rard, 1961) or C. (Floraconus) angasi Tryon, 1883, in shape. Acknowledgments I thank Dr. M. G. Harasewych, of the Smithsonian Institution, Washington, D.C., for the excellent photographs. Literature Cited Petuch, Edward J. 1981. A volutid species radiation from northern Honduras, with notes on the Honduran Caloosahatchian secondary relict pocket.— Proceedings of the Biological Society of Washington 94(4):1110-1130. Walls, Jerry G. 1979. Cone shells: A synopsis of the living conidae. Tropical Fish Hobbyist Pub- lications, Inc., Neptune City, New Jersey. pp. 1-1011. Department of Geology, Florida Inter- national University, Miami, Florida 33199. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 17-20 THE STATUS OF PSEUDORHABDOSYNOCHUS YAMAGUTI, 1958, AND CYCLOPLECTANUM OLIVER, 1968 (MONOGENEA: DIPLECTANIDAE) Delane C. Kritsky and Mary Beverley-Burton Abstract. —Cycloplectanum Oliver, 1968, is considered a junior subjective synonym of Pseudorhabdosynochus Yamaguti, 1958. Diplectanum epinepheli Yamaguti, 1938, P. epinepheli Yamaguti, 1958, and C. hongkongensis Bev- erley-Burton and Suriano, 1981, are synonyms, resulting in the valid name of the species being P. epinepheli (Yamaguti, 1938) new combination. During our independent investigations on diplectanids from fishes in the Neotropics (DCK) and in the Pacific and Caribbean (MB-B), it became apparent that a problem exists concerning the status of the genera Pseudorhabdosynochus Yamaguti, 1958, and Cycloplectanum Oliver, 1968. In this paper, we present a historical review of these taxa and offer a solution to the systematic problem. Yamaguti (1958) established Pseudo- rhabdosynochus for P. epinepheli Yamaguti, 1958, from the gills of Epinephelus akaara (Temm. and Schleg.) collected from the In- land Sea of Japan. The genus was charac- terized, in part, by the presence of squa- modiscs reduced to membranous plaques with several curved, transverse ridges. Oli- ver (1968) proposed Cycloplectanum for di- plectanids in which the two interior rows of rods on the squamodiscs formed closed cir- cles. He designated Diplectanum america- num Price, 1937, as the type-species, of which D. epinepheli Yamaguti, 1938, D. serrani Yamaguti, 1953, D. amplidiscatum Bravo-Hollis, 1954, D. latesi Tripathi, 1957, D. melanesiensis Laird, 1958, and Pseu- dorhabdosynochus epinepheli Yamaguti, 1958, were considered junior synonyms. Beverley-Burton and Suriano (1981) emended Cycloplectanum on the basis of morphologic characteristics of the terminal genitalia (i.e., copulatory complex and va- gina). These authors did not accept Oliver’s (1968) synonymies of species, but consid- ered all six taxa listed to be distinct. Rec- ognizing that their arrangement would re- sult in homonymy between the then congeneric epinepheli (Yamaguti, 1938) and epinepheli (Yamaguti, 1958), Beverley-Bur- ton and Suriano (1981) proposed C. ya- magutii to replace the latter. It is evident that the proposals of Oliver (1968) concerning Pseudorhabdosynochus epinepheli and the establishment of Cyclo- plectanum are based on incorrect inter- pretations of the International Code of Zoological Nomenclature (ICZN). His de- termination that P. epinepheli was a junior synonym of Diplectanum americanum does not invalidate the status of the former as the name-bearing type of Pseudorhabdosy- nochus (Art. 61, ICZN). Thus, Cycloplec- tanum is a subjective junior synonym of Pseudorhabdosynochus since the taxon con- tains two type-species (P. epinepheli and D. americanum) with Pseudorhabdosynochus having priority. As long as D. americanum and P. epinepheli are congeneric, Cyclo- plectanum must be suppressed (Art. 23, ICZN). Based on the most recent revision of Cy- cloplectanum by Beverley-Burton and Sur- iano (1981), Pseudorhabdosynochus (= Cy- cloplectanum) currently contains the species listed in Table 1. From Yamaguti’s (1958) 18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Pseudorhabdosynochus species and their synonyms. Species Synonyms Pseudorhabdosynochus epinepheli (Yamaguti, 1938) (senior subjective synonym of the type species, P. epinepheli Yamaguti, 1958) P. americanum (Price, 1937) P. amplidiscatum (Bravo- Hollis, 1954) P. beverleyburtonae (Oli- ver, 1984) P. bocquetae (Oliver and Paperna, 1984) P. caballeroi (Oliver, 1984) P. cupatum (Young, 1969) P. lantauensis (Beverley- Burton and Suriano, 1981) P. latesi (Tripathi, 1955) P. melanesiensis (Laird, 1958) P. querni (Yamaguti, 1968) P. serrani (Yamaguti, 1953) P. summanae (Young, 1969) P. vagampullum (Young, 1969) Diplectanum epinepheli Yamaguti, 1938 Pseudorhabosynochus epinepheli Yamaguti, 1958 (type species) Cycloplectanum hongkongensis Beverley-Burton and Suriano, 1981 C. americanum (Price, 1937) Oliver, 1968 (partim) C. yamagutii Beverley-Burton and Suriano, 1981 americanum Price, 1937 . americanum (Price, 1937) Oliver, 1968 (partim) amplidiscatum Bravo-Hollis, 1954 . amplidiscatum (Bravo-Hollis, 1954) Beverley-Burton and Suriano, 1981 americanum (Price, 1937) Oliver, 1968 (partim) beverleyburtonae Oliver, 1984 americanum (Price, 1937) Oliver, 1968 (misidentification) americanum Price, 1937 of Euzet and Oliver (1965) (misidentification) AISA AAS Q's boquetae Oliver and Paperna, 1984 C. caballeroi Oliver, 1984 C. americanum (Price, 1937) Oliver, 1968 (partim) D. americanum Price, 1937 of Caballero and Bravo-Hollis (1961) (misidentifica- tion) D. cupatum Young, 1969 C. cupatum (Young, 1969) Beverley-Burton and Suriano, 1981 C. lantauensis Beverley-Burton and Suriano, 1981 . latesi Tripathi, 1955 latesi (Tripathi, 1955) Beverley-Burton and Suriano, 1981 . americanum (Price, 1937) Oliver, 1968 (partim) melanesiensis Laird, 1958 melanesiensis (Laird, 1958) Beverley-Burton and Suriano, 1981 americanum (Price, 1937) Oliver, 1968 (partim) querni Yamaguti, 1968 querni (Yamaguti, 1968) Beverley-Burton and Suriano, 1981 serrani Yamaguti, 1953 serrani (Yamaguti, 1953) Beverley-Burton and Suriano, 1981 summanae Young, 1969 summanae (Young, 1969) Beverley-Burton and Suriano, 1981 . vagampullum Young, 1969 . vagampullum (Young, 1969) Beverley-Burton and Suriano, 1981 ESE OR BONS (Oss (GiGy SEGiGes description of the internal anatomy and structure of the squamodisc of P. epinepheli, it might be argued that this species is not congeneric with others included in the table. Relevant to this is that Yamaguti (1958) indicated an intercecal ovary which does not loop the right intestinal crus. Pseudo- rhabdosynochus epinepheli was also consid- ered to have unarmed squamodiscs which distinguished it, at that time, from other known species of Diplectanidae. However, our study of the holotype and paratype of P. epinepheli Yamaguti, 1958 (Meguro Par- asitological Museum No. 22375) under No- VOLUME 99, NUMBER 1 - 30 2 i 6 3 a 8 Figs. 1-8. Pseudorhabdosynochus epinepheli (Yamaguti, 1938). 1, Diagram of median region of body (ventral); 2, Ventral anchor; 3, Dorsal anchor; 4, Vagina (ventral); 5, Cirrus; 6, Dorsal bar; 7, Hook; 8, Ventral bar. All figures are drawn to the same scale (30 micrometers) except Figure 1. marksi (direct interference contrast) illu- mination confirmed that the ovary does loop the right intestinal crus as it does in all other species of the complex (Fig. 1). While both the holotype and paratype lack scaled squa- modiscs, this feature also is not sufficient to exclude P. epinepheli from the complex since squamodisc scales are easily lost if fixation does not occur immediately after death of the diplectanid. Confusion concerning the valid name of the type-species of Pseudorhabdosynochus also exists. Originally indicated by mono- typy, the species, P. epinepheli Yamaguti, 1958, has undergone name changes (to C. americanum by Oliver, 1968, and to C. ya- magutii by Beverley-Burton and Suriano, 1981) as a result of the proposal and sub- sequent revision of Cycloplectanum. Now, our examination of holotypes and paratypes of Diplectanum epinepheli Yamaguti, 1938 (Meguro Parasitological Museum No. 22259), P. epinepheli Yamaguti, 1958 (Me- guro Parasitological Museum No. 22375) and C. hongkongensis Beverley-Burton and Suriano, 1981 (USNM Helm. Coll. Nos. 76720, 76726, 76727) has revealed that all of these forms are conspecific. The type- series of Diplectanum epinepheli includes specimens which have squamodiscs par- tially or completely lacking scales, and scler- ites of the haptor and terminal genitalia are indistinguishable from those of P. epineph- eli. Thus, since the three species listed above are herein considered conspecific, the senior available name (i.e., valid name, Art. 23a, ICZN) for this taxon is P. epinepheli (Ya- maguti, 1938). In his descriptions of D. epinepheli and P. epinepheli, Yamaguti (1938, 1958) did not provide detailed drawings of the scler- ites of the haptor and genitalia. Those pre- sented herein (Figs. 2-8) are based on the holotype and paratype of P. epinepheli Ya- 20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON maguti, 1958, the specimens on which Pseudorhabdosynochus was originally pro- posed. Discussion Blackwelder (1967:503-505) has shown that the 1961 ICZN was not clear regarding the definition of what the type of a genus is, 1.€., a species (a taxon) or a species name, although the preface (page v) to this edition of the Code clearly indicates the former. However, the Glossary of the 1985 edition of the ICZN expresses that a type-species is a nominal species, a nomenclatural concept having no defined taxonomic boundaries. Although many authors (taxonomists) have apparently believed that the type of a genus is a species rather than a species name, the definitions provided by the 1985 Code clearly indicate that the type of a generic taxon is a name. Thus, the type, designated by monotypy, of Pseudorhabdosynochus re- mains P. epinepheli Yamaguti, 1958, which is a junior subjective synonym of P. epi- nepheli (Yamaguti, 1938) n. comb. The nominal genus, Cycloplectanum Oliver, 1968, while a junior subjective syn- onym of Pseudorhabdosynochus, satisfies all criteria of the Code (Arts. 10—20) and is therefore an available name. If at some later revision of the species group, americanum and epinepheli are determined not to be congeneric, Cycloplectanum is available for the group containing americanum. Acknowledgments We wish to thank Dr. S. Kamegai, Di- rector of the Meguro Parasitological Mu- seum, Tokyo, Japan, for the loan of type- specimens of Pseudorhabdosynochus epi- nepheli Yamaguti, 1958, and Diplectanum epinepheli Yamaguti, 1938; Dr. J. R. Lich- tenfels, Animal Parasitology Institute, Beltsville, Maryland, for loan of the type- specimens of Cycloplectanum hongkongen- sis Beverley-Burton and Suriano, 1981; and Drs. R. C. Anderson, Idaho State Univer- sity, Pocatello, and R. L. Rausch, Univer- sity of Washington, Seattle, for critical and useful comments concerning our analysis. Financial support was provided by the Na- tional Sciences and Engineering Research Council of Canada (Grant No. 801-81). Literature Cited Beverley-Burton, M., and D. M. Suriano 1981. A revision of Cycloplectanum Oliver, 1968 (Monogenea: Diplectanidae) and descriptions of C. hongkongensis n. sp. and C. lantauensis n. sp. from Epinephelus spp. (Serranidae) in the South China Sea.—Canadian Journal of Zool- ogy 59:1276-1285. Blackwelder, R. E. 1967. Taxonomy. A text and ref- erence book. John Wiley and Sons, Inc., 698 pp. Oliver, G. 1968. Recherches sur les Diplectanidae (Monogenea) parasites de téléostéens du Golfe du Lion. I. Diplectaninae Monticelli, 1903.— Vie et Milieu, Serie A 19:95-138. Yamaguti, S. 1938. Studies on the helminth fauna of Japan. Part 24. Trematodes of fishes, V.—Jap- anese Journal of Zoology 8:15-74. 1958. Studies on the helminth fauna of Ja- pan. Part 53. Trematodes of fishes, XII. — Pub- lications of the Seto Marine Biological Labo- ratory 7:53-88. (DCK) Department of Allied Health Professions and Idaho Museum of Natural History, Idaho State University, Box 8002, Pocatello, Idaho 83209; (MB-B) Depart- ment of Zoology, University of Guelph, Guelph, Ontario NIG 2W1, Canada. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 21-28 A NEW RECORD OF PARACOROPHIUM HARTMANNORUM ANDRES, 1975, FROM THE CHILEAN COAST, WITH A DESCRIPTION OF THE ADULT (AMPHIPODA: COROPHIIDAE) Exequiel Gonzalez Abstract.—The adult male and female of Paracorophium hartmannorum Andres, 1975, are described, and P. chilensis Varela, 1983, is placed in syn- onymy. The main differences between adults and juveniles are longer dactyl teeth on the palm of the second gnathopod and the toothed posterior margin of the bases of pereopods 6 and 7 in the adults. The present material extends the geographical range of the species north of Valparaiso. Paracorophium hartmannorum was de- scribed by Andres (1975) from a juvenile male 2.8 mm in length collected by G. Hart- mann-Schroder from Estero Lenga (36°46’S, 73°10’W) near Concepcion, Chile. Andres recorded his new species from four localities between Constitucioén and Valdivia, in southern Chile. Collections made in 1976 from the mouth of Rio La Ligua (32°25’S, 71°26'W), north of Valparaiso (Fig. 1), ex- tend the known range to the north, and en- able me to describe the adults. The latter have been compared with Varela’s descrip- tion and figures of P. chilensis, from Val- divia, which is considered a synonym of P. hartmannorum. Paracorophium hartmannorum Andres Figs. 2-4 Paracorophium hartmannorum.— Andres, 1975:127-130, figs. I-V. Paracorophium chilensis. —Varela, 1983: 32-36, figs. 5-6. Description of male. —Length 3.8-—4.4 mm. Head with lateral lobes produced. An- tennae 1 and 2 as described by Andres (1975); segments 2 and 3 of antenna 2 al- most 3 times as wide as segment 4. Lower lip, mandible, maxilla 2, and maxilliped similar to Andres’ description. First max- illa, outer plate wide, distal margin with 8 to 9 bifid spines, second segment of palp with 5 simple apical spines. Second maxilla like that of Paracorophium excavatum Hur- ley, inner plate slightly shorter, inner mar- gin fringed with long fine setae along about */; of its length, setae also on distal margin; outer plate with setae only on distal margin. First gnathopod: coxal plate large, slightly constricted proximally, with 2 strong spines on posterior margin, 15 setae on ventral margin; basis proximally constricted, group of 3 long simple setae and 4th more distal seta on posterior margin, inner surface with 2 simple setae on middle region, distal end with 5 long simple setae on medial surface; ischium with same proportions as in P. ex- cavatum; merus with same proportions as ischium, 3 short plumose setae on posterior margin, inner surface of distal end toward posterior with 3 or 4 short plumose setae; carpus, anterior margin with 4 or 5 long simple setae in middle to distal region, an- teriodistal angle with 3 long plumose setae, posterior margin with 16 long slender plu- mose setae on outer surface and 10 long slender plumose setae on border, inner sur- face with oblique row of long and fine plu- 22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mose setae from posteroproximal to *3 an- terodistal end of segment; propod, anterior Margin with 2 long simple setae on middle region, anterodistal angle with 5 long simple setae, first shortest, inner surface with 2 or 3 setae, palm with 8 short spines on outer surface, and 5 or 6 on inner surface, margin of palm and distal part of posterior margin of propod with several fine teeth giving ap- pearance of little brush; dacty] slightly long- er than palm, with long spine on proximal anterior margin, posterior margin with in- conspicuous tooth at proximal end and strong spine about '4 from distal end, mar- gin finely serrate proximal to spine. Second gnathopod, coxal plate subrectangular with 7 or 8 short spines on ventral margin, | or 2 strong spines on posterior margin; basis constricted proximally, posteriorly convex as Hurley (1954) described for P. excava- tum, 2 or 3 long setae on posterior margin, 1 or 2 little spines on distal end of anterior margin, proximal end '2 width of distal end; ischium '3 basis length with 2 spines on dis- tal posterior margin; merus with distal end free, anterior margin divided, forming inner and outer borders, latter forms with pos- terior margin almost rectangle, merus length 3 times its width, distal end with 9 long strong plumose setae, most of them reach- ing distal end of propod, inner border of anterior margin and posterior margin form- ing lanceolate structure with 18 long plu- mose setae on inner border, posterior mar- gin with 3 or 4 simple setae on distal third; carpus, posterior end articulating with an- terior margin of merus, posterior margin of carpus with 13 or 14 long plumose setae, anterodistal angle with 4 or 5 long simple setae, inner surface with oblique row of long plumose setae directed from anterodistal to posteroproximal; propod quadrate, anterior margin with 6 or 7 plumose setae forming 2 or 3 groups, anterodistal angle with 3 or 4 simple setae, posterior margin produced distally into strong, narrow defining tooth, palm with strong median tooth '2 as long as defining tooth, deep excavation between teeth with 3 plumose setae, median tooth with 3 or 4 plumose setae on its base, de- fining tooth with 2 or 3 plumose setae on its tooth base, inner surface of palm with 2 or 3 short plumose setae on tooth base and 2 on deep excavation between teeth; dactyl strong, curved, reaching by half its length beyond defining tooth, anterior margin with simple spine near proximal end, posterior margin with 2 or 3 short simple spines on distal 4 and 3 or 4 on proximal end. Per- eopods: Third, Fourth and Fifth without dif- ferences from Andres’ (1975) description. Basis of Pereopod 5 with proximal posterior lobe. Basis of Pereopod 6 with proximal posterior lobe, distal lobe less evident but with conspicuous toothed margin, short plumose spines on margins of both lobes. Pereopod 7, basis with strongly toothed posterodistal lobe, short plumose spines on margin. Uropods: First similar to Andres’ description, peduncle produced into ventral process beneath rami as in P. excavatum and P. lucasi (Thompson), 4 spines on pe- duncle outer dorsal margin and 2 on inner dorsal margin; outer ramus with 2 lateral spines at about midlength and 3 or 4 dorsal spines at apex; inner ramus with one spine on lateral margin and 3 or 4 at apex. Second with single dorsal spine on peduncle dis- tolateral corner, outer ramus shorter than inner with single spine at midlength of lat- eral margin and 4 at apex; inner ramus with 2 spines near midlength on dorsal surface and 4 at apex. Third as in Andres’ descrip- tion, with a few more apical setae than shown by Varela (1983). Telson similar to Andres’ description, with slightly different setation as shown in Figure 4. Epimera: First with 3 or 4 plumose setae (Varela, 1983, shows 2 setae); second, with long plumose setae on ventral margin and distal part of posterior margin which is slightly toothed; third wide, without setae. Description of female.—Length 2.5—4.8 mm. Antenna 2 with second and third pe- VOLUME 99, NUMBER 1 23 LOPS aa — ame 5 —_ Fig. 1. Map of Central Chile showing the collection site. duncle segments not wide, just twice length Sixth with basis without teeth on posterior of fourth segment. Gnathopods: First much margin, posterior lobes not evident. Sev- as in male. Second with propod and carpus’ enth with basis without teeth on posterior slender, palm shorter than in male, without margin or if present, teeth very weak; lobes teeth, armed only with long plumose setae, not evident, general appearance as in Figure defining tooth inconspicuous. Pereopods: Vc of Andres (1975). 24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Paracorophium hartmannorum, male “‘u’ modified from Andres (1975); Male “‘f,” length 3.8 mm. A, Antenna; C, Head; G, Gnathopod; I, Inner plate or ramus; L, Lower lip; M, Mandible; P, Pereopod; R, Uropod; S, Maxilliped; T, Telson; W, Pleon; X, Maxilla. Lower case letters to left of capital letters refer to specimens cited in captions; lower case letters to the right are as follows: d, dorsal; r, right; 1, left. VOLUME 99, NUMBER 1 Fig. 3. Paracorophium hartmannorum, male “‘f,” length 3.8 mm. Symbols as in Fig. 2. 25 26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON uR 3td Fig. 4. Paracorophium hartmannorum, male “‘u” modified from Andres (1975); Male “‘f,” length 3.8 mm. Symbols as in Fig. 2. 1R ; General Remarks pendages are different as stated by Andres Paracorophium hartmannorum closely (1975). The most conspicuous characteris- resembles P. excavatum, but setation and _ ticsin the adult form are greater dactyl length proportional lengths of articles of the ap- and presence of teeth on palm of second VOLUME 99, NUMBER 1 gnathopod, and toothed posterior margins of basis in fourth and fifth pereopods. The geographical distributions given by Andres (1975) is from Constitucién (35°20’S, 72°25'W) to Valdivia (Niebla, Rio Valdivia, 39°51'S, 73°24'W); there were two inter- mediate localities, Rio Andalien (36°44’S, 73°01'W) in Concepci6n Bay and Estero Lenga (36°46'S, 73°01'W) in San Vicente Bay. Varela’s specimens of P. chilensis were from Playa Caleta, Mehuin (39°26’S, 73°13'W). The present material extends the northern geographical range to La Ligua (32°25’S, 71°26’W). All these localities have similar environmental characteristics, es- pecially fluctuating salinity, from 0.67% to 38.2%o (Andres 1975). In the mouth of Rio La Ligua the salinity varied from 8%o to 22%o depending on the effects of tides. The species is found mainly in fine sand (0.15- 0.20 mm in diameter). Key to the Species of Paracorophium _ (modified and expanded from Hurley (1954) and Andres (1975) 1. Outer plate of maxilliped with fringe of many fine setae along inner mar- gin; first, second and third epimera all with marginal fringes of long slender setae; pereopods 5-7 with long fine setae marginally on most segments; antenna 2 of male with 4th peduncle segment not produced forward markedly as lobe ........ 2S Oly Oe en eee P. lucasi Hurley — Outer plate of maxilliped with 8 or 9 slender spines along inner margin; third epimeral plate lacking margin- al fringe of setae; pereopods 5—7 with only a few marginal setae 2. Antenna 2 in male, 4th peduncle segment produced distoventrally into distinct lobe; median tooth on palm of gnathopod 2 almost as long as defining tooth; dactyl not reach- ing beyond defining tooth; posterior margin of basis of pereopods 6-7 enline eee P. excavatum (Thompson) Aull — Antenna 2 in male 4th peduncle seg- ment not produced distoventrally into lobe; median tooth on palm of gnathopod 2 half as long as defining tooth; dactyl overreaching defining tooth by about half its length; pos- terior margin of basis of pereopods G=/ toothedaena sn ane Paracorophium hartmannorum Andres Acknowledgments I wish to thank Dr. Juan Cancino, Mr. L. Nelson, Ms. I. Bahamondes, Ms. A. Mezey, and Mrs. S. Montalva who took the samples as part of a research project in a class on Marine Ecology (BIO-320, 1976) at the Pontificia Universidad Catolica de Chile, Laboratorio de Zoologia. I would also like to thank Dr. Juan Carlos Castilla of the Gru- po de Ecologia Marina, Pontificia Univer- sidad Catolica de Chile, who encouraged me to work on amphipods, for his constant ad- vice during my stay at the Laboratorio de Zoologia. I am also grateful to all who made this work possible, especially Mr. Leonel Jofré who drew the figures from my rough drafts. Drs. J. Laurens Barnard, Thomas E. Bowman, and Juan Carlos Castilla reviewed the manuscript and offered helpful com- ments. This work was finished while the writer was working at Facultad de Ciencias del Mar, Universidad del Norte, Coquim- bo, Chile. Literature Cited Andres, H. G. 1975. Zur Verbreitung eulitoraler Gammaridea (Amphipoda, Crustacea) an den von Kaltwasserstromen beeinflussten Kisten Sitidamerikas und Siidafrikas sowie Angaben tiber sublitorale Gammaridea vor der chilenischen Kiste. Dissertation zur Erlangung des Doktor- grades des Fachbereichs Biologie der Univer- sitat Hamburg, 139 pp. Barnard, J. L. 1969. The families and genera of ma- rine gammaridean Amphipoda. — Bulletin of the United States National Museum 271:1-535. Hurley, D. E. 1954. Studies on the New Zealand amphipodan fauna, No. 7: The family Coro- 28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Phiidae, including a new species of Paracoro- Facultad de Ciencias del Mar, Universi- phium.—Transactions of the Royal Society of : : jot Weslo dl GOAL. dad del Norte, Casilla 117, Coquimbo, Varela, C. 1983. Anfipodos de las playas de arena Chile. del sur de Chile (Bahia de Maiquillahue, Val- divia).—Studies on Neotropical Fauna and En- vironment 18:25—52. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 29-33 DESCRIPTION OF A PAEDOPHAGOUS DEEP-WATER CICHLID (TELEOSTEI: CICHLIDAE) FROM LAKE MALAWI, AFRICA Jay R. Stauffer, Jr. and Kenneth R. McKaye Abstract. —A deep-water paedophagous cichlid of the genus Diplotaxodon is described from Lake Malawi, Africa. Similarity of jaw morphology between this species and shallow-water paedophages which obtain eggs and larvae by ramming brooding females, suggest that this deep-water cichlid obtains food in the same manner. The exploitation of fish eggs, embryos, and larvae (i.e., paedophagy) was first re- ported for haplochromine fishes by Green- wood (1959, 1967, 1974). Recent studies of the shallow water fauna of Lake Malawi led to the discovery of a group of three fish species which specialized in ramming fe- males to take young from their mouths (McKaye and Kocher 1983). This ramming mode of feeding was suggested, contrary to hypotheses from earlier studies (Green- wood 1959, Fryer and Iles 1972, Wilhelm 1980), to be one of the general methods by which paedophagous cichlids gathered their prey in the lakes of Africa. The other meth- od is to steal eggs after they are laid, but before the female can gather them into her mouth, as done by Cyrtocara ovatus, Cyr- tocara insignis, and Cyrtocara labifer (McKaye 1984). Deep water trawls in Lake Malawi pro- duced four specimens of an undescribed species of Diplotaxodon, one of which had cichlid eggs in its stomach. The purpose of this paper is to describe this deep-water pae- dophage. External counts and measure- ments follow Barel et al. (1977). Diplotaxodon greenwoodi, new species Fig. | Holotype. —National Museum of Natural History (USNM) 270847, adult female (ripe) (Fig. 1), 198.7 mm standard length (SL), 8 km south of Mumbo Island, 34°45’E, 14°04’S, Cape Maclear, Lake Malawi, at 86 m. Collected by KRM and JRS, Field Col- lection Number JRS-84-29, 17 Apr 1984. Paratypes. -USNM 270848 (3 speci- mens, females, 167.2 (ripe) 111.9 (jmma- ture), 140.3 Gmmature) mm SL), same data as holotype. Description.—This description is based upon the holotype and 3 paratypes. Prin- cipal morphometric ratios and meristics are presented in Table 1. Body form.—The body is moderately compressed. Body depth ranges between 342-365 thousandths of standard length. Distances between snout and dorsal fin or- igin and snout and pelvic fin origin range between 382-404 and 452-472 thou- sandths of standard length, respectively. The holotype had 16 abdominal vertebrae and 18 caudal vertebrae. The three paratypes nal NO =P UY, WS se 17, acl US ae 1S alos dominal and caudal vertebrae (counts from radiographs). Head. — Head length ranges between 374- 381 thousandths of SL. The eye is large (horizontal eye diameter, 275-354 thou- sandths of HL). There are either two or three scale rows on the cheek. The jaw is prog- nathus and the gape inclination ranges be- tween 57-66 degrees. The mean length of the lower jaw is 487 thousandths of HL. 30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.— Principal morphometric and meristic characteristics of Diplotaxodon greenwoodi (n = 4 and includes holotype). Holotype Mean Range SD Standard length, mm 198.7 154.5 111.9-198.7 Siz! Head length, mm 75.1 58.3 42.1-75.1 14.2 Thousandths of HL Horizontal eye diameter DS 309 276-354 34 Vertical eye diameter 278 310 278-352 33 Snout length 317 299 242-342 43 Postorbital head length 406 389 380-406 12 Premaxillary pedicel 173 191 173-200 13 Lower jaw length 519 487 460-519 25 Interorbital width 189 176 166-193 18 Cheek depth DY) 185 159-201 28 Head depth 870 850 824-870 21 Thousandths of SL Head length 378 377 374-381 3 Snout to dorsal 383 396 383-404 10 Snout to pelvic 453 459 453-472 9 Body depth 355 354 342-365 9 Least caudal peduncle length 196 185 168-196 14 Least caudal peduncle depth NS 112 109-115 3 Pectoral fin length 329 320 306-332 13 Pelvic fin length 199 198 180-210 13 Dorsal fin base length 459 447 426-459 15 Gape inclination (degrees) 63 62.3 57-66 3.8 Lateral line scales 34 34 34 — Scale rows on cheek 2 25 2-3 0.58 Pectoral fin rays 13 2ES 12-13 0.58 Pelvic fin rays 5 5 5 — Anal fin spines 3 3 3 — Anal fin rays 10 10.3 10-11 0.5 Gill rakers on ceratobranchial 20 19.3 19-20 0.5 Gill rakers on epibranchial 6 5.3 5-6 0.5 There is a small ventral protrusion present at the symphysis of the dentaries. Teeth on the lower jaw of the holotype are in three rows, while those on the upper jaw are in four rows. Teeth in fourth row do not extend as far posteriorly as the oth- ers. All teeth are unicuspid. Those on the lower jaw are embedded in the soft tissue of the jaws. Gill rakers are single filaments and on the ceratobranchial range in number between 19-20. There are 6 on the epibranchial, with one in the angle between the ceratobranchial and epibranchial bones. Fins. —™Mean dorsal fin base length is 447 (426-459) thousandths of standard length. Pectoral fins are comprised of 12-13 seg- mented rays. Anal fins have three spines and 10-11 segmented rays. Lower pharyngeal bone. — The lower pha- ryngeal bone was dissected from the holo- type. It is triangular in outline with two tooth forms (Fig. 1). Both tooth forms are conical, with those on the posterior part of the bone straight, while those laterally are slightly re- curved. Squamation. —Scales are ctenoid, with 34 pored lateral line scales. Scales uniformly VOLUME 99, NUMBER 1 31 Fig. 1. Holotype (USNM 270847) of Diplotaxodon greenwoodi. cover the anterior third of the caudal fin, partially cover the middle third, and are essentially absent from the posterior third. Coloration. —Freshly collected speci- mens are dark dorsally, with a purple sheen. The lateral surfaces are silver, but fade to white ventrally. Diagnosis.—Trewavas (1935:68) distin- guished the genus Diplotaxodon as follows: “third, or third and fourth vertebrae with inferior apophyses which are short, and do not approach each other below dorsal aorta; vertebra 16 + 18; teeth conical, in two se- ries; premaxillaries not beak-like.”’ Diplo- taxodon greenwoodi differs from the other two described species in the genus, D. ar- genteus and D. eccelsi, by the increase in gape inclination, and vertebral number. There is also a fourth row of teeth on the upper jaw. An analysis of the phylogenetic relationships of D. greenwoodi must await revision of the genus, which probably con- tains some 20 undescribed species (Iles, pers. comm.). Discussion. —Most specimens trawled from below 70 meters had completely everted guts when they reached the surface. However, the gut of the holotype contained eggs, which were identical in shape and size with those contained in the ovaries of Leth- rinops gossei which were collected simul- taneously. It is conceivable that the eggs in the stomach of D. greenwoodi were a result of “trawl snatching.” However, the pres- ence of numerous ripe male and female L. gossei in the trawl sample indicates that the trawl was made over an L. gossei breeding arena where the presence of a paedophage would be expected. That the paedophage was relatively rare (< 0.1% of cichlids caught in trawl) is consistent with data collected on shallow-water paedophages (Fryer and Iles 1972, McKaye and Kocher 1983). The feeding behavior of D. greenwoodi 32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON has not yet been observed because it occurs at depths below our present diving capa- bilities. Nevertheless we hypothesize that D. greenwoodi is a paedophage which ob- tains eggs and larvae by ramming brooding females, in a manner convergent with that in Cyrtocara orthognathus and Cyrtocara liemi (McKaye and Kocher 1983). This be- havioral hypothesis is inferred from the morphology of D. greenwoodi. The head shape, particularly gape inclination (ranging between 57-66 degrees), closely resembles that of C. orthognathus and C. liemi, which are known ramming paedophages (McKaye and Kocher 1983). Examination of the gape inclination of the holotype (USNM 227497), and two paratypes (USNM 227501; 227503) of C. liemi showed that the gape inclination ranged between 41-45 degrees. Gape inclination of three speci- mens of C. orthognathus (British Museum Natural History [BMNH] 1973.3.26:88; 1956.6.4:4; 1969.3.11:17) ranged between 67-74 degrees. Both C. /iemi and C. or- thognathus obtain eggs and larvae by stalk- ing brooding females in the water column, and approaching these females from under- neath and behind before ramming them. Convergence in form among paedo- phages is further supported by D. green- woodi dentition. Teeth are restricted to the anterolateral parts of the upper and lower jaw bones and are buried in a thickened oral mucosa. A similar condition exists in both Lake Victoria haplochromine lineages of paedophagous fishes which are now placed in the genus Lipochromis and given subge- neric rank of Lipochromis and Clepto- chromis (Greenwood 1979). Etymology. —Named after P. H. Green- wood in recognition of his renowned studies of cichlid fishes, and for being the first to discover this unique feeding specialization in cichlids. Acknowledgments We especially thank the government of Malawi for providing the facilities to make this research possible, and the Cape Maclear fisheries technicians R. D. Makwinja, W. M. Menyani and O. K. Mhone, for help in making the collections. We benefitted from discussions with D. C. S. Lewis and D. Tweddle. L. Knapp of the Smithsonian In- stitution arranged for the shipment of spec- imens to the USNM. Financial support was from the National Science Foundation DEB- 79-12338, BSR-82-14603 (KRM) and In- ternational Programs (USAID), University of Maryland (JRS) and is Contribution No. 1624-AEL University of Maryland, Center for Environmental and Estuarine Studies. The original art work was completed by Ms. Michelle Katz. Literature Cited Barel, C. D. N., M. J. P. van Oijen, F. Witte, and E. L. M. Witte-Mass. 1977. An introduction to the taxonomy and morphology of the haplo- chromine Cichlidae from Lake Victoria. — Netherland Journal of Zoology 27(4):339-389. Fryer, G., and T. D. Iles. 1972. The cichlid fishes of the Great Lakes of Africa. Edinburgh: Oliver & Boyd. 641 pp. Greenwood, P. H. 1959. A revision of the Lake Vic- toria Haplochromis species (Pisces, Cichlidae), Part 3.—Bulletin British Museum of Natural History (Zoology) 5:179-218. 1967. A revision of the Lake Victoria Hap- lochromis species (Pisces, Cichlidae), Part 6.— Bulletin British Museum of Natural History (Zoology) 25:139-242. 1974. The cichlid fishes of Lake Victoria, East Africa: The biology and evolution of a species flock. — Bulletin British Museum of Nat- ural History (Zoology) Supplement 6:1-134. . 1979. Towards a phyletic classification of the ‘genus’ Haplochromis (Pisces, Cichlidae) and re- lated taxa. Part II; the species from Lakes Vic- toria, Nabugabo, Edward, George and Kivu. — Bulletin British Museum of Natural History (Zoology) 39:1-101. Mayr, E. 1963. Animal species and evolution. Har- vard University Press. Cambridge, Massachu- setts. 797 pp. McKaye, K. R. 1984. Behavioural aspects of cichlid reproductive strategies: Patterns of territoriality and brood defence in Central American sub- stratum spawners versus African mouth brood- ers. Pp. 425-273. In R. J. Wooton and G. W. Potts (eds.) Fish reproduction: Strategies and tactics. Academic Press, London. VOLUME 99, NUMBER 1 , and T. Kocher. 1983. Head ramming behav- ior by three paedophagous cichlids in Lake Ma- lawi, Africa.— Animal Behavior 31:206-210. , and C. Mackenzie. 1982. Cyrtocara liemi, a previously undescribed paedophagous cichlid fish (Teleostei: Cichlidae) from Lake Malawi, Africa.— Proceedings of the Biological Society of Washington 95:398—402. Trewavas, E. 1935. A synopsis of the cichlid fishes of Lake Nyasa.— Annuals and Magazine of Nat- ural History 10(7):133-152. Wilhelm, W. 1980. The disputed feeding behaviour 33 of a paedophagous Haplochromine cichlid (Pisces) observed and discussed. — Behaviour 17: 310-323. (JRS) School of Forest Resources, The Pennsylvania State University, University Park, Pennsylvania 16802. (KRM) Appa- lachian Environmental Laboratory, Uni- versity of Maryland, Frostburg, Maryland MNOS. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 34-41 AEGLA SPECTABILIS, A NEW SPECIES OF FRESHWATER CRAB FROM THE EASTERN SLOPE OF THE NAHUELBUTA COASTAL CORDILLERA, CHILE Carlos G. Jara Abstract. —A new species, Aegla spectabilis (Crustacea: Anomura: Aeglidae), is described from the Nahuelbuta region in central southern Chile. Morpho- logically it is related to A. rostrata and A. denticulata, both from Chile. Its most prominent feature is the robust spinulation on the margins of the carapace and dorsum of legs. It can be distinguished from all other species of Aegla by its spiniform palmar crest and the strong spine on the dorsum of carpus of the chelipeds. The Chilean aeglid fauna is currently rep- resented by 12 species of Aeg/a Leach (1821) (see Retamal 1981 and Jara 1982). Among them, three, namely A. denticulata Nicolet (1849), A. rostrata Jara (1977), and A. ba- hamondei Jara (1982) stand out from the remaining Chilean species mostly because of the prominent spinulation on the margins of the carapace (Jara 1982). All three occur in the region between Concepcion (36°50’S) and Valdivia (39°30’S) (see Jara 1977, 1980, 1982), but A. denticulata ranges farther to the south (Bahamonde and Lopez 1963). The Nahuelbuta Coastal Range, in which the type-locality of A. bahamondei is locat- ed (Jara 1982), extends into this part of the country. The same author mentions that specimens of Aeg/a, referable with doubt to A. bahamondei, were collected at a locality situated to the southeast of the Nahuelbuta Range. However, no previous record of Ae- gla from the eastern slope of Nahuelbuta proper exists. In this paper, a fourth strongly orna- mented form of Aeg/a, found in a river sys- tem that drains part of the eastern slope of Nahuelbuta, is described as a new species. Aegla spectabilis, new species Fig. 1 Holotype.—Instituto de Zoologia, Uni- versidad Austral de Chile, IZUA C-637, adult female, Chol Chol River under bridge on outskirts of Chol Chol, 29 km northwest of Temuco (38°36’S, 72°51'W) by road, Chile, 21 Dec 1982, coll. C. G. Jara. Paratype.—IZUA C-633, young female, Perquenco River at Galvarino town, 27 km north of Chol Chol (38°25'S, 72°47'W) by road, Chile, 22 Dec 1982, coll) C>G Warar Diagnosis. —Rostrum iong, styliform, acute; orbital spine well developed; first he- patic lobe spiniform; anterior and posterior branchial margins strongly denticulate; an- terolateral angle of second abdominal epi- meron upturned, spiniform; fourth thoracic sternum with median spiniform tubercle; palmar crest as single acute projecting spine; dorsum of carpus of chelae with long acute spine; dorsal margin of carpus of ambula- tory legs denticulate; merus of ambulatory legs with distodorsal and distoventral ends spiniform. Description of holotype.—Carapace ovoid, VOLUME 99, NUMBER 1 dorsoventrally depressed and laterally ex- panded in branchial regions. Middorsal line elevated but not forming longitudinal ca- rina; dorsum of carapace more ridge-roofed than convex, bearing sparse long fine setae arising in groups from small pits; setae dens- er, shorter, and stiffer on both sides of mid- dorsal line of abdominal terga. Precervical region of carapace much narrower than postcervical. Rostrum long, slender, tipped with acute conical scale; distal third slightly upturned and dorsally flattened. Rostral carina nar- row but well marked along proximal half of rostrum, bearing irregular row of small scales intermingled with setae; shallow grooves on both sides fading out beyond midlength of rostrum; height of rostral carina at level of corneae less than depth of ventral keel. Ros- tral tip surpassing eyestalks by 2.5 length of cornea. Rostral and orbital margins without scales. Orbits U-shaped, wide, moderately deep; orbital spine large, conical, acute, inclined toward anterolateral angle of carapace; ex- traorbital sinus wide, deep, subrectangular, its maximum aperture little less than half maximum orbital aperture. Anterolateral lobe of carapace not particularly flattened and produced as large conical spine reaching midlength of cornea; main part of spine di- rected laterally while acuminate tip inclined toward midline. External margin of antero- lateral lobe little scabrous but not scaly. Hepatic area expanded laterally, some- what flattened dorsally and limited inward- ly by shallow groove separated from more elevated gastric area; first hepatic lobe sep- arated from anterolateral lobe by wide sub- crescentic moderately deep notch, its free angle produced as strong, laterally diver- gent, acute spine. Second and third hepatic lobes relatively well defined and limited by small blunt-angled notches; dorsum of third moderately inflated. Epigastric eminences consisting of slender, arcuate, little protu- berant nodulose ridge behind which semi- 35 circular finely punctate depression extend- ing to base of protogastric eminences. Latter moderately prominent, forming small blunt conical protuberance tipped with small len- ticular scale separated from base of rostral carina by shallow concavity. Gastric area protuberant, its uppermost zone flat. Bran- chial areas expanded laterally, their free margins denticulate. Anterior branchial margin with 5 broad based, conical, some- what dorsally depressed denticles, their frontally recurved tips with acute scale, de- creasing 1n size posteriorly and separated by wide semicircular indentations. Posterior branchial margin with 6 (right) and 8 (left) small but clear cut, acute, conical denticles almost uniform in size, incurved frontally, and markedly upturned, especially the last. Dorsal surface of anterior branchial area slightly concave; posterior branchial area convex toward midline but concave behind free margin. Cardiac area subrectangular, slightly wider at frontal than at rear end. Areola limited laterally by subparallel grooves, subrectangular, about 1.5 times longer than broad, markedly protuberant, and regularly convex from side to side. Abdominal terga deeply sculptured, with narrow, shallow groove devoid of setae along midline. Anterolateral angle of second ab- dominal epimeron noticeably upturned and produced as broad based, triangular, later- ally compressed, sharp spine; tip rising above dorsal surface of epimeron reaching later- ally to level of tips of posterior branchial denticles; anteroventral margin of epimeron below anterolateral angle almost straight, ending in blunt ventral angle. Lateral sur- face of epimeron between anterolateral and ventral angle quite concave. Angles of third and fourth epimera also sharply acute. Tel- son plate dimerous, subpentagonal. Fourth thoracic sternum with projecting, subconi- cal, slightly compressed, corneous tipped, but almost blunt, ventromedial tubercle. Cheliped (only left one present) slender. Chela subrectangular in dorsal view, dor- 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 99, NUMBER 1 soventrally depressed, its dorsal surface with short stiff setae and tiny scales. Fingers slightly. curved; when closed leaving wide open gap between finely crenulate inner margins. Movable finger with small acute tubercle on its dorsal edge close to propo- dus. Anteromesial lobe of palm small, spi- niform, separated from palmar crest by nar- row deep notch. Palmar crest consisting of single, long, conical, sharply acuminate spine; its proximal base bearing 2 very small swellings tipped with setae. Carpal lobe pro- duced in slender, acute spine. Carpal crest forming 2 long, sharp, slightly recurved, and widely separated spines; proximal spine about two-thirds as long as distal one; row of 4 small blunt tubercles tipped with setae present above carpal crest and behind carpal lobe; another dorsal crest situated external to and clearly separated from former con- sisting of long sharp acuminate curved spine, as long as the second in inner crest, followed by blunt tubercle. Ventral face of carpus armed with long acute spine about size of second spine of inner carpal crest. Space between ventral spine and inner crest with several long thin setae. Merus with dorsal row of 4 spiniform tubercles, distalmost longest and second shortest; distodorsal vertex also spiniform; inner ventral margin smooth with single distal long curved spine; outer ventral margin with 3 spines of which proximalmost longest and second shortest. Ventromedial margin of ischium smooth, with only broad blunt tubercle proximally. Distodorsal and distal ventrolateral angles of merus of pereiopods projected in strong conical acute spine; merus of second left pereiopod also with minute acute tubercle on distal fifth of dorsal margin; distodorsal — Fig. 1. 3h7/ Table 1.—Somatometry of A. spectabilis, new species, type series. All measurements in mm. CL, carapace length, distance between rostral apex and posterior margin of cephalothorax; RL, rostral length, distance between rostral tip and midpoint of transverse line tangent to deepest points of orbital margins; PCL, pre- cervical length, distance between rostral tip and mid- point of cervical groove; FW, frontal width, distance between tips of anterolateral angles of carapace; PCW, maximum precervical width, distance across third he- patic lobes; CW, maximum carapace width; LCL, left cheliped length; RCL, right cheliped length; L2PL, length of second left pereiopod; L2DL, dactylar length of second left pereiopod; L4DL, dactylar length of fourth left pereiopod; TL, telson length. F: female. Collection IZUA-C 637 IZUA-C 633 Specimen Holotype Paratype Sex ; F F CL : 19.5 15.0 RL ; 6.1 4.1 PCL : 13.9 10.6 FW ; 6.1 4.9 PCW : 9.9 8.3 CW : 15.5 13.0 LCL 20.7 16.1 RCL — 15.4 L2PL 2 23.5 17.9 L2DL : 5.7 4.1 L4DL : 5.9 4.3 TL : 3.2 2.9 angle of carpus of pereiopods strongly spi- niform, preceded by 1 to 3 smaller but also strong spines in row decreasing in size proximally. Color (alcohol fixed specimens).— Dor- sum of carapace vinaceous (Ruber 3) as background color but with several changes of intensity in different regions. Margins of rostrum, orbits, and crest of rostral carina dark blood red (Ruber 8). Epigastric areas between proto and epigastric eminences Aegla spectabilis (all illustrations from holotype): a, Dorsal view; b, Left chela (dorsal); c, Carpus of left cheliped (dorsal), arrow points to large middorsal spine; d, Carpus of left cheliped showing large middorsal spine in profile; e, Dorsal view of rostral area; f, Carpus of left cheliped (ventral); g, Carpus of second left pereiopod; h, Lateral view of cephalothorax; i, Telson plate; k, Third and fourth thoracic sterna; 1, Lateral view of second abdominal epimeron; m, Ventral view of ischium of left cheliped. 38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON purplish black (Purpureus 9). Midline be- hind epigastric eminences and dorsum of posterior branchial areas sepia (Aurantia- cus 9). Distal half of marginal spines of branchial areas apricot (Aurantiacus 2) while corneous apex of all spines translucent saf- fron yellow (Aurantiacus 3). Dorsum of pe- reiopods orange (Aurantiacus 4) or honey yellow (Flavus 5). Ventral surface of body and appendages marmoreous white or slightly yellowish with some iridescences. (Color standards according to Paclt 1958). Variation and measurements.—The sin- gle female paratype differs from the holo- type in the following respects: rostrum shorter, surpassing eyestalks by 1.5 times length of corneae; dorsum of carapace more regularly convex; front comparatively nar- rower; branchial margins less upturned and expanded laterally but clearly denticulate; in dorsal view, anterior margin of antero- lateral angle of second abdominal epimeron closer to posterolateral angle of carapace; dorsal spiniform process of movable finger very small, lacking apical scale; ventrome- dial tubercle on fourth thoracic sternum blunt, without apical scale. Measurements of the holotype and para- type specimens are presented in Table 1. Measurements were made with calipers to the nearest 0.1 mm; the morphometric characters here considered are those defined by Jara and Lopez (1981). Distribution. — Known only from the type- locality and from Galvarino, 27 km north of the type-locality, in the Perquenco River which is a tributary of the Chol Chol River. Natural history. —The river section where the holotype of A. spectabilis was collected corresponds to an epipotamal facies; there the current velocity reached about 0.8 m sec’ '. The greenish turbid water flowed over a sand and gravel bottom limited on the sides by steep banks covered by bushes and bamboo-like vegetation. The water tem- perature on 21 December 1982 was 20°C, pH 7.9, and conductivity 53 uS cm™!; dis- solved oxygen reached 13.3 mg 1~!. The paratype was collected in a river section of hyporithral characteristics where current velocity reached an average of 0.6 m sec! flowing over boulders and stones without gravel or sand. Water was transparent, its temperature on 22 December 1982 was 17.5°C, ph 7.0, conductivity 95 uS cm", and dissolved oxygen 13.6 mg 1~!. At both localities A. spectabilis was collected to- gether with a second, largely more abun- dant, species of Aeg/a not yet identified but very similar to A. rostrata Jara. As sam- plings were done tracking counter-current for long stretches of the bottom with a sack- like kick net, the microhabitat conditions preferred by A. spectabilis are not known. Etymology. —From spectabilis (Latin), amazing. The specific name refers to the surprisingly heavy ornamentation of this species of Aeg/a. Comparison. —Aegla spectabilis resem- bles A. denticulata Nicolet, 1849, and A. rostrata Jara, 1977, both from Chile. With them it shares a rather short ovoid carapace greatly expanded at the branchial level, a long, tapered, acute rostrum, robust acute spines on the anterolateral angles of the car- apace, well developed orbital spines, an acute and somewhat exserted first hepatic angle, conspicuously denticulate branchial margins, and strongly armed chelipeds; moreover, the dorsodistal angle of the car- pus and merus of the ambulatory legs are acute or spiniform and the anterolateral an- gle of the second abdominal epimeron is spiniform and buttressed behind by a ridge which arises from the tergum. In 4. spec- tabilis, however, the dorsum of the carapace appears to be more depressed, the denticles on the branchial are better defined and more prominent, and the chelae are less massive than in the other two species. The areola of A. spectabilis is more protuberant than that of A. rostrata but less than that of A. den- ticulata though in A. spectabilis the areola has a faint keel along its midline. The an- terolateral angle of the second abdominal epimeron of A. spectabilis is more upturned VOLUME 99, NUMBER 1 than it is in A. rostrata and A. denticulata. With A. denticulata it shares the prominent conical tubercle on the fourth thoracic ster- num and the scarcely defined proto- and epigastric prominences, but A. spectabilis lacks the pronounced carina along the dor- sum of the carapace which distinguishes A. denticulata. On the other hand, A. specta- bilis resembles A. rostrata in possessing a long, narrow rostrum and wide U-shaped extraorbital sinuses but differs from it in several other respects, mainly in the form and size of the chelae and the distinctness of the branchial denticles. Neither A. denticulata nor A. rostrata has a row of spines along the dorsal margin of the carpus of the ambulatory legs as A. spec- tabilis has. Two spines on the distodorsal vertex of the carpus of the ambulatory legs are mentioned for A. /enitica Buckup and Rossi, 1977, from Brazil, and a row of spi- niform scabrosities along the carpal margin are drawn but not mentioned in the descrip- tion of A. parana Schmitt, 1942, also from Brazil. However, in no other species of Ae- gla are these spines as well developed as in A. spectabilis. A single-spined carpal crest of the che- lipeds, such as that found in A. spectabilis, is approached only by that drawn and de- scribed by Schmitt (1942) for the holotype of A. sanlorenzo Schmitt, 1942, from Ar- gentina; however, the character was absent in the specimens of A. sanlorenzo examined by Ringuelet (1949a). Finally, the long slen- der spine on the external half of the dorsum of the chelipedal carpus in A. spectabilis is a unique feature, not shared with any other aeglid. Whereas in A. parana Schmitt, 1942, a middorsal row of spiniform tubercles is located in a position similar to that occu- pied by the spine in A. spectabilis, no tu- bercle is markedly larger than the others. Remarks. —Aegla spectabilis is the most profusely ornamented form of Aeg/a yet de- scribed. Because of the shape of its rostrum, it could be included in the Pacific rostrum type group of Aegla proposed by Schmitt 39 (1942). However, because of its two most distinctive characters, namely the spiniform carpal crest and the spine on the dorsum of the chelipedal carpus, it clearly departs from the remaining species of Aeg/a. Due to these particular attributes it perhaps deserves to be allocated to a separate subgeneric or ge- neric taxon but such a decision seems pre- mature in view of the need for a revision of all the species assigned to the unique genus Aegla. The current diagnosis of the genus Aegla (see Leach 1821, Nicolet 1849, Gir- ard 1855, Hobbs, Hobbs et al., 1977) seems too broad, not adequately assessing the morphological variation displayed by ae- glids. At least several subgeneric categories should be recognized in order to arrange the species according to their presumed phy- logenetical relationships which, up to now, have been partially scrutinized only by Rin- guelet (1949b). The degree of spinulation varies greatly among aeglids and some ideas about their possible phylogeny have been advanced re- garding tendencies of morphological trans- formation among the species. Schmitt (1942: 442) supposed that “‘the least differentiated, least spiny or ornamented species stands nearest the ancestral Aegla.”’ If Schmitt’s opinion is correct then the spinulation of A. spectabilis would be apomorphic. At vari- ance with Schmitt’s opinion, Ringuelet (1949a, b) stated that a common trend among several Argentinian species is to- ward reduction of orbital spines and ex- traorbital sinuses. According to him (1949), the species of Aeg/a “‘nearest the ancestor” are those which possess wide concave ex- traorbital sinuses, large orbital spines, a wide front, long anterolateral spines, a prominent rostrum with a sharp carina not throughed on both sides, and a spiniform second ab- dominal epimeron. If Ringuelet’s opinion is correct then A. spectabilis morphology would be plesiomorphic in several respects. The tendencies of character transforma- tion observed among Chilean aeglids seem to indicate that an extreme reduction of or- 40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON namentation as well as an extreme devel- opment of spinulation may be considered as apomorphic conditions. In fact, the great- est reduction of spines is shown by A. ala- calufi Jara and Lopez, 1981, and, to a lesser degree, by A. papudo Schmitt, 1942. Both species share an undivided telson plate while the remaining aeglids have dimerous ones (Jara and Lépez 1981). The monomerous telson plate is undoubtedly apomorphic. Other Chilean aeglids which show reduced spinulation are: A. maulensis Bahamonde and Lépez, 1963, A. manni Jara, 1980, and A. concepcionensis Schmitt, 1940. With the exception of A. maulensis, all of these species live in small brooks and streams associated with the Chilean western continental slope, and possibly, they are phylogenetically re- lated. On the other hand, the trend toward pro- fuse spinulation seems to be correlated with living in lentic environments. Thus, A. ros- trata, from Lake Rinihue, most likely rep- resents a lacustrine lineage derived from less spiny riverine forms (see Jara 1977, 1982). The same trend is demonstrated by A. den- ticulata which is widespread in small, mod- erately fast running streams in the provinces of Valdivia, Osorno, and Llanquihue. Re- cently, a large population of this species was found in the sub-littoral of Lake Rupanco (Osorno); those specimens show a remark- ably more prominent and profuse spinula- tion than those collected from streams. Since Lake Rupanco, like Lake Rinihue, is a young lake (in geological time), it is unlikely that its population of A. denticulata is the stem population of the less spiny riverine form. Considered from this point of view, an ex- ceedingly prominent spinulation would be apomorphic. It is then possible that the spi- nulation of A. spectabilis was developed in a lentic environment. Such a possibility is strongly supported by the fact that during the Pliocene the region of Angol, about 70 km to the north of the type-locality of A. spectabilis, was occupied by lakes formed by accumulation of glacial waters along the eastern slope of the Nahuelbuta Coastal Range (Borgel 1983). If this is so, the low population density of A. spectabilis. in the Chol Chol river basin may be indicative of its relictual condition. Nahuelbuta has been advocated as a refugial area for the pregla- cial Tertiary freshwater fauna of central southern Chile (Arenas 1976). Jara (1982) hypothesized that A. bahamondei, from the western slope of Nahuelbbuta, might rep- resent the ancestral stock from which the lacustrine A. rostrata might have evolved. The finding of A. spectabilis on the eastern slope of Nahuelbuta reinforces the notion that this area harbours peculiar faunistic elements which render it important from a zoogeographical point of view. Acknowledgments The author thanks René Navarro for his collaboration during the field work in Na- huelbuta, Lic. José Arenas for many hours of enriching discussions about taxonomy and the zoogeography of Chilean aeglids, Mrs. Patricia Araya for correcting the first draft of this paper, Dr. Horton Hobbs Jr. for his most appreciated final revision, and Direccion de Investigaci6n y Desarrollo, Universidad Austral de Chile, for providing the funds for field trips through grant RS- 80-33. Literature Cited Arenas, J. 1976. La Cordillera de la Costa como re- fugio de la fauna dulcicola preglacial.—Archi- vos de Biologia y Medicina Experimentales 10: R40. Bahamonde, N., and M. T. Lopez. 1963. Decapodos de aguas continentales en Chile.—Investiga- ciones Zooldgicas Chilenas 10:123-149. Borgel, R. 1983. Geomorfologia. Instituto Geogra- fico Militar, Santiago, Chile. 182 pp. Buckup, L., and A. Rossi. 1977. O Genero Aegla no Rio Grande do Sul, Brasil (Crustacea, Deca- poda, Anomura, Aeglidae).— Revista Brasileira de Biologia 37(4):879-892. Girard, C. 1855. Description of certain Crustacea, brought home by the U.S. Naval Astronomical VOLUME 99, NUMBER 1 Expedition. — The U.S. Naval Astronomical Ex- pedition to the Southern Hemisphere, during the years 1849-50-51-52, 2:254—262. Wash- ington, D.C. Hobbs, H. H., Jr., H. H. Hobbs III, and M. D. Daniel. 1977. A review of the troglobitic decapod crus- taceans of the Americas.—Smithsonian Contri- butions to Zoology 244:1-183. Jara, C. 1977. Aegla rostrata, n. sp. (Decapoda, Ae- glidae), nuevo crustaceo dulceacuicola del Sur de Chile.—Studies on Neotropical Fauna and Environment 12:165-176. 1980. Taxonomia y distribucion del género Aegla Leach (Crustacea, Decapoda, Anomura) en el sistema hidrografico del rio Valdivia (Chile). Universidad Austral de Chile, Facultad de Ciencias, Thesis. 126 pp. . 1982. Aegla bahamondei, new species (Crus- tacea: Decapoda: Anomura) from the Coastal Mountain Range of Nahuelbuta, Chile.—Jour- nal of Crustacean Biology 2(2):232-238. , and M. T. Lopez. 1981. A new species of freshwater crab (Crustacea: Anomura: Aeglidae) from insular South Chile.— Proceedings of the Biological Society of Washington 94(1):88-93. Leach, W. E. 1821. Galatéadées, pages 49-56 in F. G. Levrault, ed., Dictionnaire des Sciences Na- turelles, 18(1820). Strasbourg. 41 Nicolet, H. 1849. In Gay, C., Historia Fisica y Po- litica de Chile. Zoologia 3:198-201. Paclt, J. 1958. Farbenbestimmung in der Biologie. Gustav Fischer Verlag, Jena. Retamal, M.A. 1981. Catalogo ilustrado de los Crus- taceos Decapodos de Chile.—Gayana (Zoolo- gia) 44:1-110. Ringuelet, R. 1949a. Los anomuros del género Aegla del noroeste de la Republica Argentina. —Re- vista del Museo de la Plata (Nueva Serie) 4 (Zoologia):1—45. 1949b. Consideraciones sobre las relaciones filogenéticas enntre las especies del género Aegla Leach (Decapodos Anomuros).— Notas del Mu- seo de la Plata 14(120):111-118. Schmitt, W.L. 1940. Two new species of Aeglea from Chile.—Revista Chilena de Historia Natural 44(1940):25-31, pl. 1. 1942. The species of Aegla, endemic South American freshwater crustaceans. — Proceed- ings of the United States National Museum 91: 431-520. Instituto de Zoologia, Universidad Austral de Chile, casilla 567, Valdivia, Chile. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 42-45 NOMENCLATURAL NOTES ON THE ANURA (AMPHIBIA) Jay M. Savage Abstract. — The status and availability of the family-group names Hylodidae, Allophrynidae, and Pelodryadidae and the generic name Crepidus are reviewed and clarified. Through the kindness of several col- leagues, a number of nomenclatural prob- lems or inconsistencies with the Interna- tional Code of Zoological Nomenclature (reference to pertinent articles of the Code are indicated as Art. in the rest of this paper) that have appeared or been perpetuated by my published works, have been recently called to my attention. Three of these relate to the use of family-group names in the clas- sification proposed for my biogeographical analysis of frog distribution (1973), the oth- er to the correct name of an endemic Central American bufonid. In the original manuscript (completed in 1971) of my 1973 paper, documentation for several novel features of classification was provided but later removed in the editing process. This circumstance makes ambig- uous certain family-group names used in that report and adopted by several subse- quent authors. The first of these is the name Hylodinae for a subfamily of the Leptodac- tylidae. The family-group name Hylodidae was originally proposed by Giinther (1859) for the genera Crossodactylus, Phyllobates, Hylodes, and Platymantis. The type-genus is Hylodes Fitzinger, 1826 (monotype: Hyla ranoides Spix, 1824, a synonym of Hyla nasus Lichtenstein, 1823). Several authors including Myers (1962) and Lynch (1971) argue that Fitzinger proposed exactly the same generic name again in 1843 for a sec- ond genus of frogs. It is clear from the 1843 work that this is not the case as the name is listed as Hylodes (Fitz.), which in this paper represents a genus previously named. As pointed out by Fitzinger (footnote p. 15), the parentheses around the author’s name indicate that the scientific name was pro- posed by that person, but is not used by Fitzinger strictly in its original sense (i.e., to have the same composition). Thus Lep- todactylus (Fitz.), Rana (Linné), and Hyla (Laurent) as well as others are so listed. Names newly proposed in the paper lack an indication of Fitzinger’s name, as for ex- ample Limnodynastes and Lithodytes. Fit- zinger (1843) designated Hylodes martini- censis Tschudi, 1838, as the type of the subgenus Hylodes, but because this form was not included in the original description of the genus-group name, it cannot be con- sidered the type-species (Art. 67g). In ad- dition, the type-species of Hylodes was es- tablished in 1826 as Hyla ranoides Spix (by monotypy) since the only other species in- cluded by Fitzinger in the genus was Hy- lodes gravenhorstii Fitzinger, a nomen nu- dum. Obviously, Fitzinger had no way to know that things would work out this way since there were no commonly accepted rules relating to nomenclature in his day. Con- sequently, contrary to the argument of Lynch (1971), the type of the family-group name Hylodidae (and Hylodinae) is Hylodes Fit- zinger, 1826. This name has priority over Elosiinae Miranda-Ribeiro (1926) under Art. 23 and the latter has no claim for ex- emption, under Art. 40b, as having won general acceptance prior to 1961. The second problem relates to my use of VOLUME 99, NUMBER 1 the family-group name Allophrynidae (monotypic for the genus A/lophryne Gaige, 1926). Mention of the family name by me did not constitute an appropriate proposal of a new name under Art. 13 of the Code and the name was not made available by my action. To remedy the situation, I here- with reintroduce the name Allophrynidae for the single genus and species A//ophryne ruthveni Gaige, 1926. This family is char- acterized by the features presented by Lynch and Freeman (1966), who give a description and definition of Allophryne which sepa- rates it from all currently recognized fami- lies of frogs. I have elsewhere (Savage 1973) suggested that the family is a derivative of a leptodactylid ancestor and cannot be placed in the Hylidae as proposed by Lynch and Freeman (1966) and Dowling and Duellman (1978). A third matter involves the use of the name Pelodryadidae Gunther, 1859, for the Australopapuan tree-frogs allied to the ge- nus Litoria and often placed in the family Hylidae. When I (1973) proposed that the Pelodryadidae be recognized, it was on the basis of the throat muscle characteristic used by Tyler (1971) to separate the Australo- papuan forms from the New World hylids. Since that time, although few have recog- nized the pelodryadids as a separate family, the family-group name Pelodryadinae has been used as a subfamilial appellation (Dowling and Duellman 1978; Tyler 1979). Unfortunately, the generic type of this tax- on, Pelodryas Ginther, 1859 (monotype: Rana caerulea White, 1790) is often re- garded as a subjective, junior synonym of Litoria Tschudi, 1838 (monotype: Litoria freycineti Tschudi, 1838). Some will argue that the proper name for this family-group should have been proposed as “‘Litoriidae”’ from the oldest generic name. However, Pe- lodryadidae is the oldest available name for this taxon, since the family-group name Pe- lobii Fitzinger, 1843 (type-genus Pelobius Fitzinger, 1843, with Litoria freycineti as monotype) is preoccupied in Coleoptera by 43 Pelobini Erichson, 1832 (generic type and senior homonym to Fitzinger’s genus: Pe- lobius Erichson, 1832) and no one has ever proposed “‘Litoriidae”’ as a name. In addi- tion, I was convined that the type-species, the well-known large, green tree-frog (Pelo- dryas caeruleus) was generically distinct from the type-species of Litoria (L. freyci- neti). Since that time a second species has been described (named Litoria splendida by Tyler, Davies, and Martin 1977) which is closely allied to caeruleus and should also be placed in Pelodryas according to the characteristics presented by Tyler and Da- vies (1978). The following features taken from the latter report diagnose the genus: large green frogs with broadly fringed and partially webbed fingers: cartilaginous in- tercalary elements in digits; hyoid plate with pedunculate alary processes; prominent parotoid or supracranial glands. For these several reasons, the appropriate family-group name for Australopapuan tree- frogs is Pelodryadidae. If subsequent work- ers decide to replace Pelodryas in the syn- onymy of Litoria, Art. 40a provides that the family-group name will remain Pelo- dryadidae. The final problem relates to the correct generic name for the lower Central Amer- ican toad originally described by Cope (1875) as Crepidius epioticus. In 1966, I pointed out that the name Crepidius Cope is preoccupied by Crepidius Candeze, 1859, in Coleoptera and used the replacement name Crepidophryne Cope, 1889, for epi- oticus, the sole species placed in the genus. In doing so I overlooked the use by Brocchi (1882) of the name Crepidus for this taxon. Brocchi lists the name as Crepidus Cope and cites Cope’s 1875 usage as Crepidus (sic), ‘““Cope, On the Batr. and Rept of Costa Rica .... The sole included species is also listed as Crepidus epioticus. Brocchi obviously had no intention of proposing a new name for Cope’s genus so that Crepidus must be either a correction of the original spelling, an un- justified emendation of the spelling of Cre- 44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pidius or an incorrect subsequent spelling. Under Art. 33c misspellings have no in- dependent status in nomenclature but a cor- rection of an original spelling (Art. 32d) or even an unjustified emendation (Art. 33b) must be regarded as an available name. If Crepidus is recognized as a correction of an original spelling or as an unjustified emen- dation it becomes available and has priority over Crepidophryne. In this case it is very difficult to determine which of the three possibilities apply. 1) Brocchi may have concluded that the cor- rect classical spelling for Crepidius was Cre- pidus (a correction of an incorrect original spelling); his citation of Cope as the author of Creidus supports this view. 2) on the oth- er hand, he may have misspelled Cope’s genus as Crepidus; 3) or then again, he may have preferred Crepidus over Crepidius, as an alternate acceptable classically correct spelling, to produce an unjustified emen- dation of the former name. It appears that alternate | is the most like- ly since Brocchi lists Cope as the author of Crepidus. There is no way to establish that alternate 2 is correct. Alternate 3 requires that Brocchi chose to modify Cope’s spell- ing for some other reason than to correct the original spelling. However, Art. 33b re- quires that Brocchi use of Crepidus instead of Crepidius is demonstrably intentional. Since this cannot be done, Crepidus must stand as an incorrect subsequent spelling (Art. 33c) and has no standing. The name Crepidophryne Cope, 1889 (monotype: Crepidius epioticus Cope, 1875) stands as the correct name for the taxon involved and replaces the preoccupied name Crepidius Cope, 1875, with the same monotype. Acknowledgments The following colleagues graciously called my attention to nomenclatural problems re- solved above: John S. Applegarth, Univer- sity of New Mexico; Darrel R. Frost, Uni- versity of Kansas; and Andrew Stimpson, British Museum of Natural History. They were most courteous in providing com- ments while permitting me to clarify these matters which my previous publications may have clouded; C. Richard Robins, Uni- versity of Miami, aided in the search of homonyms. My thanks to them all. Literature Cited Brocchi, M. 1882. Etude des Batraciens de 1’ Ame- rique Centrale.— Mission scientifique au Me- xique 3(2):57-96. Candeze, E. 1859. Monographie des Elaterides. Tome II.—Memoires de la Société Science Liége 14: 1-543. Cope, E. D. 1875. On the Batrachia and Reptilia of Costa Rica.—Journal of the Academy of Nat- ural Sciences of Philadelphia (letter-press) ser. 2,8:93-157. 1889. The Batrachia of North America.— Bulletin of the United States National Museum 34:1-515. Dowling, H. G., and W. E. Duellman. 1978. System- atic herpetology: A synopsis of families and higher categories. Herpetological Information Search Systems (HISS) Publications, New York. 118 pp. Erichson, W. F. 1832. Genera Dyticeorum. Disser- tatio inauguralis, Berlin, pp. 1-48. Fitzinger, L. J. F. J. 1843. Systema Reptilium. Vi- enna. 1826. Neue Classification der Reptilien. Vi- enna. Gaige,H.T. 1926. Anew frog from British Guiana. — Occasional Papers Museum of Zoology, Uni- versity of Michigan 176:1-3. Giinther, A. C. L. G. 1859. Catalogue of colubrine snakes in the collection of the British Mu- seum. Taylor and Francis, London. International Commission on Zoological Nomencla- ture. 1985. International Code of Zoological Nomenclature, 3rd. ed. London. Lichtenstein, H. 1823. Verzeichniss der Doubletten des Zoologischen Museums Konig]. Universitat zu Berlin. Berlin. 48 pp. Lynch, J. D. 1971. Evolutionary relationships, os- teology and zoogeography of leptodactylid frogs.— Miscellaneous Publications University of Kansas Museum of Natural History 53:1- 238. , and H. L. Freeman. 1966. Systematic status of a South American frog, A//lophryne ruthveni Gaige.— University of Kansas Publications Mu- seum Natural History 17(10):493-502. VOLUME 99, NUMBER 1 Miranda-Ribeiro, A. 1926. Notas para servirem ao estudio dos gymnobatrachios (Anura) Brasilei- ros.—Archivos Museo Nacional Rio de Janeiro 27:1-227. Myers, G. S. 1962. The American leptodactylid frog genera Eleutherodactylus, Hylodes (= Elosia), and Caudiverbera (= Calyptocephalus).—Co- peia 1962(1):195-202. Parker, H. W. 1940. The Australasian Frogs of the Family Leptodactylidae.—Novitates Zoolo- gicae, Tring Museum 42(1):1-106. Savage, J. M. 1973. The geographic distribution of frogs. In J. L. Vial, ed. Evolutionary biology of the anurans, 13. University Missouri Press pp. 351-445. Spix, J. B. 1824. Animalia nova sive species novae testudium et ranarum. Monaco. Tschudi, J. J. 1838. Classification der Batrachier. Neuchatel. Tyler, M. J. 1971. The phylogenetic significance of vocal sac structure in hylid frogs.— University 45 of Kansas Publications Museum of Natural His- tory 19(4):319-360. 1979. Herpetofaunal relations of South America with Australia.— University of Kansas Museum of Natural History Monographs 7:73- 106. , and M. Davies. 1978. Species-groups within the Australopapuan hylid frog genus Litoria Tschudi.— Australian Journal of Zoology, Sup- plement Series 63:1—47. ——,, M. Davies, and A. A. Martin. 1977. A new species of large, green frog from northern west- ern Australia.—Transactions of the Royal So- ciety of South Australia 101:133-138. White, J. 1790. Journal of a voyage to New South Wales, with sixty-five plates of non descript an- imals, birds, lizards, serpents, curious cones of trees and other natural productions. London. Department of Biology, University of Miami, Coral Gables, Florida 33124. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 46-50 SEROLIS AGASSIZI, NEW SPECIES, FROM THE DEEP SEA OFF CAPE FEAR, NORTH CAROLINA (CRUSTACEA: ISOPODA) Robert Y. George Abstract. —A new species of isopod Serolis agassizi is described. This blind deep-sea species was collected at 3840 meters over the continental rise off the coast of North Carolina. The abyssal species is the fourth member of the polytypic genus Serolis occurring north of the equator and its affinity with two shallow Northern Hemisphere species, S. carinata from the California coast, and S. mgrayi from the Georgia coast, is discussed. Its resemblance to the other abyssal blind species S. vemae is pointed out. The isopod species belonging to the fam- ily Serolidae are morphologicaly distinctive due to their flattened body form. Out of the approximately 50 known species of the ge- nus Serolis, only three have been found so far to occur north of the equator; all other species live in the Southern Hemisphere where they are particularly diverse and abundant in the cold waters of the Antarctic shelf and subantarctic islands (Nordenstam 1933; Sheppard 1933). Lockington (1877) discovered Serolis carinata, the first record of this genus in the Northern Hemisphere, in shallow waters off the coast of southern California. A second serolid species, S. mgrayl, was described by Menzies and Frankenberg (1966) from the continental shelf off Georgia. Hessler (1972) pointed out that these two shallow-water serolid species from the Northern Hemisphere, one from the Atlantic and the other from the Pacific coast, are closely related sibling species iso- lated geographically by the Central Ameri- can land bridge. The speciation of these two species took place as a consequence of their geographic separation. The third known Northern Hemisphere serolid species, S. vemae, was first reported from a depth of 5024 meters over the Ar- gentine Rise southeast of Rio Grande in the South Atlantic Ocean (Menzies 1962). Se- rolis vemae was later captured in good num- bers from a depth of 2862—4749 meters in the North Atlantic Ocean (Hessler 1967). The present paper describes the second abyssal species of Serolis from a depth of 3840 meters over the continental rise off North Carolina in the Northwest Atlantic Ocean. This new serolid species, S. agassiZzi, was collected during a cruise of R/V East- ward and is named in honor of Alexander Agassiz who made significant contributions to our understanding of the deep-sea fauna of the North Atlantic Ocean. Serolis Leach, 1818 Diagnosis. —Serolidae markedly flat- tened, not known to conglobate; coxal plates laterally expanded. Mandible with lacinia mobilis. Maxilliped with 3-articled palp. Pereopod 2 sexually dimorphic, in female ambulatory and in male subchelate. Pleo- pods 1-3 each with elongate peduncle and subelliptical rami. Uropoda biramous. Serolis agassizi, new species Figs. 1, 2 Diagnosis. — Serolis with coxal plates vis- ible dorsally in pereonal somites 2 to 4. Body configuration somewhat globular. Cephalon lacking eyes. Lateral margin of all pereonal VOLUME 99, NUMBER 1 47 Fig. 1. somites denticulate. Pereonal somite | fused with cephalon. Pleotelson globular; poste- rior margin broadly rounded, denticulate. Lateral margin of uropod biramous; exopod minute; endopod enlarged, with lateral mar- gin denticulate. Material examined.—Holotype female, length 3 mm, width 2.5 mm (USNM 138717). Type-locality. —Continental rise off North Serolis agassizi, holotype female, length 3.0 mm: A, Dorsal view; B, Uropod. Carolina coast. R/V Eastward Sta 6214, 1 female, 3840-3975 m, 33°01.5'—33°03.0'N, 75°06.2'-75°6.5'W, 5 Nov 1966; collected with Small Biological Trawl. Description.— Body shape globular with lateral margin denticulate and furnished with minute setae. Cephalon with large rostrum separated from lateral lobes by narrow fur- row on either side of rostrum. Eyes entirely absent, ocular ridge lacking. Cephalon al- 48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON most as long as pereonal somites 2—7 com- bined. First pereonal somite with lateral margin denticulate. Pereonal somites 2—3 showing distinct coxal plates. Somites 1 and 2 sub- equal in length at midline; fourth somite less than one-half as long at middorsal region than at lateral region. Fifth and sixth so- mites similar in shape, lateral parts pro- duced posteriorly to approximately half the length of pleotelson. Seventh pereonal so- mite uniformly narrow. Pleonite somewhat narrower than seventh pereonite. Pleotel- son broadly rounded; posterior margin studded with minute tubercles similar to those on lateral margin of pereonal somites. Dorsal surface of pleotelson smooth, com- pletely devoid of any ridges, tubercles or spines. First antenna composed of 9 articles. Bas- al 4 articles constitute peduncle; first article bulbous; second article longest article of pe- duncle; third article longer and broader than fourth. Flagellum consisting of 5 articles, all bearing sensory setae. Second antenna with well developed peduncle composed of 4 ar- ticles. Basal 2 articles small, subequal; third and fourth articles elongate. Flagellum with 7 articles; total length of flagellum less than length of fourth peduncular article. Outer margin of second antenna furnished with setae. Maxilliped with 3-jointed palp. Basal ar- ticle of palp about as long as terminal article; second article twice as wide as terminal ar- ticle. Endite furnished with 2 stout setae at distal end. Epipodite marked off from ba- sopodite by distinct suture. First pereopod with elongate basis longer than carpus, ischium and merus combined. Propodus enlarged, displaying row of spines along inner margin. Dactylus about as long as propodus; both combined into claw-like configuration. Seventh pereopod with sub- equal joints and reduced dactylus; merus bearing 3 leaf-like setae at outer distal edge. Third pleopod with triangular basopod- ite; endopodite larger than exopodite. Ex- opodite with 3 distal plumose setae. En- dopodite with long plumose setae along distal and outer margins. Uropoda biramous, exopod extremely small; endopod large and denticulate along margins. Affinity with Other Three North Atlantic Species of Serolis The new abyssal serolid species S. agas- sizi is markedly different from the two eye- bearing shallow water species, S. mgrayi from the Atlantic shelf (200—266 m) and S. carinata from the California coast (13-56 m) in the absence of eyes. As Hessler (1967) pointed out, these two shallow serolid species are quite similar in morphology but there are substantial differences to distin- guish them as distinct species which may have a common origin in the Northern Hemisphere. However, S. agassizi is closely related to S. vemae, the other blind serolid species known from the abyssal depths (2862-5024 m) in the North and South At- lantic Ocean. Both species are small (less than 5 mm). The presence of a minute ex- opod of the uropod and a flagellum shorter than the last peduncular article of the sec- ond antenna are features shared by these two species. There are obviously well-de- fined morphological differences between these two deep sea North Atlantic serolid species. Serolis agassizi has rounded an- terolateral angles of the cephalon but S. ve- mae has acephalon with acute anterolateral angles. The coxal plate is marked off in pe- reonal somite 5 in S. vemae but not in this new species. Serolis agassizi has a smooth dorsal surface but denticulate lateral mar- gin. Serolis vemae has a small tubercle on pereonal somites 5 and 6 and a smooth lat- eral margin. The pleotelson shape is very different between these two species, the apex broadly rounded in S. agassizi and some- what tapering in S. vemmae. The second ar- ticle of the maxilliped palp is elongate and the terminal third article small in S. vemae. The second article of the maxilliped palp is VOLUME 99, NUMBER 1 49 Fig. 2. Serolis agassizi, holotype female, length 3.0 mm: A, Maxilliped; B, Maxilliped, ventral view; C, first pereopod; D, Second pereopod; E-F, setae enlarged; G, Second pleopod. only slightly longer than the terminal third article in S. agassizi. These points of dif- ference clearly differentiate the two abyssal species of Serolis in the Northern Hemi- Besides these two species, there are only sphere. three other species of Serolis known to live Affinity with Abyssal Species of Serolis from the South Atlantic Ocean 50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON exclusively at depths greater than 2500 me- ters. These three abyssal species are known from the subantarctic regions in the Scotia Sea. Serolis maryannae Menzies, from the continental rise, south of Staten Island (55°31.2'S, 64°07.5'W) at 3839 meters, is about four times larger than S. agassizi. This species also has reduced eyes, middorsal tu- bercles on the pereonal somites and elon- gate lateral margins of the posterior pe- reonal somites. Serolis macdonellae Menzies, from the lower slope depths in a region west of South Sandwich Island (56°43'S, 27°41'W) at 2741 meters, is strik- ingly different from S. agassizi in general body dimension, in possessing tubercles on the dorsal surface of the pereonal somites and in having a middorsal carinae on the pleon. The uropods do not extend beyond the posterior margin of pleotelson and the lateral margins of the last two pereonal so- mites are elongate in S. macdonellae. The third abyssal species from the Southern Hemisphere, S. margaretae, is known from the continental rise in the northwestern part of the Scotia Sea, south of Staten Island (55°42.9’S, 62°21.6’W) at 3813 meters. This species resembles S. agassizi in general body shape shape and also in having a similar pleonal configuration. However, there are obvious differences between these two species in the cephalon, which has a mid- dorsal tubercle at the posterior end and con- spicuous eyelobes in S. margaretae. The epimera of the sixth pereonal somite extend slightly beyond the apex of the telson and this feature evidently demarcates this species from S. agassizi. The closest known relative of S. agassizi is undoubtedly S. vemae which inhabits the abyssal depths of both northern and southern hemispheres in the Atlantic Ocean. From a zoogeographic and evolutionary point of view, the distribution of the species of the genus Serolis is exceedingly interest- ing because of its dominant representation in the shelf-slope depths of the Antarctic and subantarctic zones. Their meager rep- resentation in the aybss (5 out of 50 known species) in depths greater than 2500 meters indicates that Serolis is generally a shelf- slope genus and speciation into the abyss probably took place after the Miocene gla- ciation when cooling of the deep sea came about. Literature Cited Hessler, R.R. 1967. A record of Serolidae (Isopoda) from the North Atlantic Ocean.—Crustaceana 12:159-162. 1972. The relationship between Serolis car- inata Lockington and Serolis mgrayi Menzies and Frankenberg (Isopoda, Flabellifera).— Crustaceana, suppl. 3:1-6. Lockington, W. N. 1877. Remarks on the Crustacea of the Pacific Coast, with descriptions of some new species.—Proceedings of the California Academy of Sciences 7(1):28-36. Menzies, R. J. 1962. The Isopods of abyssal depths in the Atlantic Ocean.— Vema Research Series 1:79-106. Menzies, R. J. and D. Frankenberg. 1966. Handbook on the Common Marine Isopod Crustacea of Georgia. University of Georgia Press, Athens, 93 pp. Nordenstam, A. 1933. Marine Isopoda of the families Serolidae, Idotheidae, Pseudidotheidae, Arctur- idae, Parasellidae and Stenetriidae mainly from the South Atlantic. — Further Zoological Results of the Swedish Antarctic Expedition 1901-1903 3:1-184. Sheppard, E. M. 1933. Isopod Crustacea Part I. The family Serolidae.—Discovery Reports 7:253- 362. Institute for Marine Biomedical Re- search, University of North Carolina at Wilmington, Wilmington, North Carolina 28403. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 51-55 A NEW SPECIES OF RED-EYED TREEFROG OF THE HYLA URANOCHROA GROUP (ANURA: HYLIDAE) FROM NORTHERN HONDURAS James R. McCranie and Larry David Wilson Abstract.—A new species of the Hyla uranochroa group is described from the Cordillera de Nombre de Dios, Departamento de Atlantida, Honduras. Relationships to other members of the group are discussed. Until recently, the Hyla uranochroa group was considered to be confined to lower Cen- tral America (Duellman 1970). However, in 1982 and 1983 we discovered specimens representing a new species of this group from western Honduras which we subsequently described as Hyla soralia Wilson and McCranie, 1985. In August 1982 and 1984 we collected additional material represent- ing another new species in this group from the Cordillera de Nombre de Dios in the Honduran department of Atlantida. The type-locality of this new species is alongside the Quebrada de Oro, a stream flowing out of the cordillera into the Rio Viejo, in turn a tributary of the Rio Cangrejal which flows into the Caribbean Sea at La Ceiba. Hyla salvavida, new species Fig. 1 Holotype.—University of Kansas Mu- seum of Natural History (KU) 200999, adult male, from Quebrada de Oro (15°38’N, 86°47'W), elevation 880 m, tributary of Rio Viejo, south slope of Cerro Bifalo, Cordi- llera de Nombre de Dios, Departamento de Atlantida, Honduras, collected 16 Aug 1984 by James R. McCranie, Kenneth L. Wil- liams, and Larry David Wilson. Original number LDW 6501. Paratopotypes.—KU 201000-008, 201010- 013, adult males, and KU 201009, adult female, 16 Aug 1982 and 16-18 Aug 1984, elevation and collectors as for holotype. Diagnosis.—A member of the Hyla ur- anochroa group distinguished from the oth- er members by the following combination of characters: dorsum uniform dark leaf green; venter pale yellow; pale lip stripe thin, diffuse, expanded below eye; lateral pale stripe poorly developed, consisting of bro- ken series of flecks; snout—vent length (SVL) 25.2-27.5 mm in males, 34.3 in single fe- male; tympanum diameter 38.7—43.8% of eye diameter in males; snout rounded in lateral profile; plantar surfaces of feet pig- mented; anterior arm of squamosal extend- ing one-third of distance to maxilla; quad- ratojugal present only as spur posteriorly. Description of holotype. — Adult male with SVL of 26.2 mm; tibia length 14.2 mm; tibia length/SVL 0.542; hand length 7.6 mm; hand length/SVL 0.290; foot length 11.0 mm; foot length/SVL 0.420; head length 10.0 mm; head length/SVL 0.382; head width 9.7 mm; head width/SVL 0.370; di- ameter of eye 3.1 mm; diameter of tym- panum 1.2 mm; tympanum/eye diameter 0.387. Snout in lateral profile rounded, in dorsal profile rounded; canthus rounded; lo- real region slightly concave; lips moderately thick and unflared; nostrils protuberant; in- ternarial distance 2.1 mm; internarial dis- tance/head width 0.216; top of head flat; interorbital distance 3.9 mm; interorbital distance/head width 0.402; width of eyelid 52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ‘ * PP hee ss Fig. 1. Female paratopotype (KU 201009) of Hyla salvavida. 2.2 mm; eyelid width/head width 0.227; snout length 2.7 mm; snout length/head length 0.270; snout length/eye diameter 0.871. Moderately heavy dermal fold ex- tending from posterior corner of eye above tympanum to point above base of forearm, obscuring upper edge of tympanum; tympa- num round, its diameter 1.25 times its dis- tance from eye. Forearm moderately robust, having well-developed dermal fold on wrist; raised dermal fold along outer edge of fore- arm; pollex slightly enlarged with poorly cornified patch of nuptial excrescences; sec- ond finger noticeably shorter than first; sub- articular tubercles round, those on third and fourth fingers bifid; discs on fingers mod- erate in size, that on third finger equal to diameter of tympanum; webbing vestigial between first and second fingers; webbing formula II 1'2-3 III 2'2—-2 IV (sensu Myers and Duellman 1982); heels broadly over- lapping when hindlimbs adpressed; tarsal fold moderately developed, extending length of tarsus; inner metatarsal tubercle ovoid, visible from above; subarticular tubercles rounded, conical; length of toes from short- est to longest 1-2-3-5-4; webbing formula I 2-2 II 1-22 III 14%2-3'2 TV 244-1 V; discs of toes distinctly smaller than those of fingers. Anal opening directed posteroventrally at upper level of thighs. Skin of dorsum smooth, that of throat and belly granular, that of ventral surfaces of thighs smooth. Tongue elongately ovoid, barely free be- hind; prevomerine teeth 4—5, situated on posteromedially-inclined ridges, narrowly separated and between moderately large ovoid choanae; vocal slits large, extending from posterolateral base of tongue to angle of jaws. Vocal sac single, median, and subgular. In life, the color pattern was as follows: dorsum dark leaf-green; exposed surfaces of limbs same; upper jaw with a thin, diffuse lip stripe that is expanded below eye; outer edge of forearm with series of interrupted pale dashes from elbow to wrist; outer edge of tarsus with thin, diffuse pale stripe; pale anal stripe poorly developed; tubercles be- low anal opening tipped with white; venter pale yellow; iris blood red. Variation in the paratopotypes. —The VOLUME 99, NUMBER 1 pertinent data on thirteen males (range fol- lowed by mean in parentheses) and one fe- male (separated from the former by a com- ma) are as follows (all measurements are in millimeters): SVL 25.2-27.5 (26.5), 34.3; tibia length 14.3-15.5 (14.8), 18.6; tibia length/SVL 0.532-0.587 (0.559), 0.542; foot length 10.2-11.5 (10.8), 13.8; foot length/ SVL 0.385-0.437 (0.408), 0.402; hand length 6.9-8.0 (7.6), 10.0; hand length/SVL 0.256—-0.317 (0.286), 0.292; head length 9.5— 10.3 (9.9), 12.3; head length/SVL 0.359- 0.398 (0.376), 0.359; head width 9.3-10.1 (9.8), 11.9; head width/SVL 0.358—0.388 (0.369), 0.347; diameter of eye 2.8—3.3 (3.1), 3.5; diameter of tympanum 1.2-1.4 (1.3), 1.3; tympanum/eye diameter 0.387-0.438 (0.409), 0.371; interorbital distance 3.6—4. 1 (3.9), 4.7; interorbital distance/head width 0.367-0.418 (0.399), 0.395; width of eyelid 2.1—2.5 (2.3), 3.0; eyelid width/head width 0.208—0.258 (0.235), 0.252; internarial dis- tance 2.0-2.3 (2.2), 2.9; internarial dis- tance/head width 0.198-0.237 (0.224), 0.244; snout length 2.7—3.0 (2.8), 3.3; snout length/head length 0.262—-0.303 (0.285), 0.268; snout length/eye diameter 0.87 1-1.00 (0.928), 0.943. Variation in color and pattern of the para- topotypes is minimal, with the exceptions of KU 201008 in which the labial stripe is well-developed, complete, and confluent with the relatively well-developed lateral stripe and in the presence of a large cream- colored spot just above the groin in the sin- gle female (KU 201009). Metamorphosing froglets.—On 16 Au- gust 1984 we collected two metamorphos- ing froglets (KU 201015-016) in stages 44 and 45 (Gosner 1960) respectively, at the type-locality. Mensural data on the speci- mens are as follows: SVL 15.2, 16.6; tail length 14.6, nub only in the latter; tibia length 7.8, 8.3; tibia length/SVL 0.513, 0.500; hand length 4.4, 4.6; hand length/ SVL 0.289, 0.277; foot length 5.6, 6.0; foot length/SVL 0.368, 0.361. Color notes on KU 201015 in life are as follows: dorsum metallic coppery green; lime 53 green on upper eyelid; venter yellow; limbs yellow with greenish-brown patina; enamel yellow spots at elbow, knee, and heel; un- dersurfaces of feet orangish-red; tail gray stippled with white; pale lip stripe present; iris red. Tadpole. —A single poorly-preserved tad- pole presumed to be of this species (KU 201014) was collected at 1070 m in the Que- brada de Oro. Allocation must be consid- ered tentative at best but the tadpole defi- nitely is of the type found in members of the Hyla uranochroa group (Duellman 1970; Wilson and McCranie 1985). Many features are obscured due to the desiccated condition of the specimen but the following features are determinable: mouth ventral, large, and funnel-shaped without lateral folds in the oral disc; oral disc entirely bordered by a row of minute papillae; large conical pa- pillae present within oral disc; beaks rela- tively small with long pointed serrations; denticle rows 7, second upper row narrowly interrupted medially, third lower row no- ticeably shorter than other rows. Osteology. —The following descriptive notes are based on a cleared and stained adult male specimen (KU 201017) of Hyla salvavida. The features of the skull are in complete agreement with those used by Duellman (1970) in diagnosing the uran- ochroa group (including a large frontopa- rietal fontanelle possessed by these frogs, Duellman’s 1970 illustration of the region as ossified in H. uranochroa notwithstand- ing). Furthermore, the skull of H. salvavida has the anterior arm of the squamosal ex- tending about one-third of the distance to the maxilla and the quadratojugal reduced to a spur posteriorly as is the case in rufi- oculis but not uranochroa (skull features are unknown for /ythrodes and soralia). Etymology.—The name salvavida is de- rived from the Spanish, meaning “‘lifesav- er,” in appreciation of the “lifesaving qual- ities” of our Honduran field companion, La Cerveza Salvavida. Natural history notes. —The vegetation at the type-locality may be characterized as of 54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the Subtropical Wet Forest formation of Holdridge (1967) and is described in more detail by McCranie et al. (in prep.). All the members of the hypodigm were collected at night on low vegetation one to two meters off the ground alongside a shal- low, broad slow-moving small stream and associated pools at a point just before it flows into the Quebrada de Oro. Extensive searches elsewhere in the area over several days in two different years produced no oth- er specimens. All the males were collected while calling. The call is a cricket-like chirp repeated three or four times. The single female collected on 16 August 1984 contains eggs about ready to be de- posited. The same evening two metamor- phosing froglets were found and a tadpole presumed to be of this species was collected on 4 June 1980. These data suggest an ex- tended breeding season. Relationships. —Hyla salvavida is a member of the uranochroa group as defined by Duellman (1970). He included two species, rufioculis and uranochroa, in the group. Myers and Duellman (1982) resur- rected Hyla lythrodes from the synonymy of H. rufioculis, and Wilson and McCranie (1985) described a fourth species, H. sor- alia, from Honduras. Hyla salvavida agrees in all features of the uwranochroa group as detailed by Duellman (1970), Duellman and Campbell (1982), and Wilson and Mc- Cranie (1985). Within this group of five species, Hyla salvavida most closely resembles H. rufi- oculis. They share a pale labial stripe ex- panded below the eye, pigmented plantar surfaces of the feet, similar tympanum/eye ratio, and the same features of osteology and larval denticle morphology. They differ in dorsal color (leaf green in salvavida, dull brown to olive green in rufioculis), nature of the lateral stripe (broken in salvavida, well-developed and complete in rufioculis), character of the labial stripe (broken and suffused with green in salvavida, well-de- veloped and complete in rufioculis), ventral color (pale yellow in salvavida, creamy white in rufioculis), and snout shape (rounded in lateral profile in sa/vavida, truncate in ru- fioculis). Wilson and McCranie (1985) discussed the relationships and biogeography of the uranochroa group, postulating a dispersal northward of a lower Central American stock across the Nicaraguan depression dur- ing a pluvial period which gave rise to Hyla soralia in western Honduras and the H. schmidtorum group in southern México. Hyla salvavida appears to fit in this scenario as an early offshoot of the northward-dis- persing stock that still shows major resem- blances to the less-derived forms in lower Central America. Acknowledgments Our thanks go to Kelly M. Hogan, Walter Holmes, and Kenneth L. Williams for their assistance in the field. Jay M. Savage and Brian I. Crother kindly loaned comparative material from the CRE collections housed at the University of Miami. Wilberto Agui- lar N., director of the Departamento de Re- cursos Naturales Renovables, as always, was instrumental in providing collecting per- mits. Our long-time friend, Jorge Porras Zu- niga, opened his home to us, obtained our rented field vehicle, and assisted in numer- ous other ways. To all we owe a debt of gratitude. Literature Cited Duellman, W. E. 1970. The hylid frogs of Middle America.— Monograph of the Museum of Nat- ural History of the University of Kansas 1:xi,1— WSS , and J. A. Campbell. 1982. A new frog of the genus Ptychohyla (Hylidae) from the Sierra de las Minas, Guatemala. — Herpetologica 38:374— 380. Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on iden- tification.— Herpetologica 16:183-190. Holdridge, L. R. 1967. Life zone ecology. Tropical Science Center, San José, Costa Rica. 206 pp. VOLUME 99, NUMBER 1 McCranie, J. R., L. D. Wilson, and K. L. Williams. [In prep]. A new genus and species of toad (An- ura: Bufonidae) from Honduras, with an ex- traordinary stream-adapted tadpole. Myers, C. W., and W. E. Duellman. 1982. A new species of Hyla from Cerro Colorado, and other treefrog records and geographical notes from western Panama.—American Museum Novi- tates 2752:1-32. Wilson, L. D., and J. R. McCranie. 1985. A new 55 species of red-eyed Hyla of the uranochroa group (Anura: Hylidae) from the Sierra de Omoa of Honduras. — Herpetologica 41:133-140. (JRM) 10770 S.W. 164th Street, Miami, Florida 33157; (LDW) Department of Bi- ology, Miami-Dade Community College, South Campus, Miami, Florida 33176. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 56-60 HAWAIIAN XANTHIDAE (DECAPODA: BRACHYURA) II. DESCRIPTION OF GARTHIELLA, NEW GENUS, WITH A REDESCRIPTION OF G. ABERRANS (RATHBUN, 1906) Richard H. Titgen Abstract. —The new genus Garthiella is erected to accommodate the xanthid species described as Chlorodopsis aberrans Rathbun, 1906. It is readily distin- guished from the genus Pilodius (=Chlorodopsis) by the sharp fingers of the chelipeds, the shape of the first pleopod of the male, and the wide orbital hiatus. Rathbun (1906) published the first com- prehensive study of Hawaiian Crustacea Decapoda. Her report was based primarily on the collections made in 1902 by the U.S.F.C. Steamer Albatross from waters surrounding the Hawaiian Islands, but in- cluded additional Hawaiian material from the U.S. National Museum of Natural His- tory, the Museum of Comparative Zoology, and the Philadelphia Academy of Natural Science. She also cited literature records of an additional 28 species. Her report remains the basis of decapod research in the Hawai- ian Islands. Of the 213 species and subspecies of Brachyura she reported, Rathbun (1906) placed 94 species in the family Xanthidae, 21 of which were described as new species. One new species, Chlorodopsis aberrans, was described and placed in the genus Chloro- dopsis A. Milne Edwards, 1873, because it ““*has much in common with C. woodmasoni Alcock.”” However, as noted by its specific epithet, Rathbun recognized that it was not a typical Pilodius Dana, 1851 (=Chloro- dopsis) by reason of pointed tips on the fin- gers of its chelipeds. All species of Pilodius are presently recognized as having chelipeds with the fingers hollowed at the tip. While studying the xanthid crabs in the Bishop Museum, the author noted that this species did not fit into the accepted concept for the genus Pilodius because of its aberrant chelipeds, nor could it be assigned to any other described xanthid genus. Accordingly, the new genus Garthiella is here erected to accommodate Chlorodopsis aberrans. Garthiella, new genus Diagnosis. —Carapace hexagonal to transversely subovate, about *% as long as wide, flattish from side to side, slightly con- vex longitudinally, front turned down and convex. Regions more or less separated an- teriorly, but vague posteriorly; surface var- iously sculptured with spines or tubercles. Front bilobed, approximately 3 carapace width; frontal-orbital border about *% car- apace width. Anterolateral border divided into 4 granulated lobes or teeth (excluding external orbital angle). Posterolateral bor- der slightly longer than anterolateral. Basal antennal article rather broad, inner angle touching ventral prolongation of front; out- er angle prolonged partway into wide orbital hiatus, not closing hiatus to antenna. Palatal ridges developed only posteriorly, not ex- tending forward to anterior boundary of buccal cavern. Chelipeds equal or subequal in both sexes, covered by tubercles; fingers pointed. Ambulatory legs anteriorly spi- nose, with scattered hairs. Male abdomen with 7 segments, 3—5 fused; gonopod curved and tapered, with several long setae near tip and conical spines on distal half. VOLUME 99, NUMBER 1 Type-species. —Chlorodopsis aberrans Rathbun, 1906. Etymology.—This genus is named for a student of the Xanthidae, Dr. John S. Garth of the Allan Hancock Foundation, Los An- geles, California; the gender is feminine. Remarks. —The new genus Garthiella is erected to accommodate the “aberrant” species Chlorodopsis aberrans Rathbun, 1906. Garthiella resembles Pilodius (=Chlorodopsis) in general appearance, but can be distinguished by the pointed fingers on the chelipeds and the tip of the first pleo- pod of the male. All species of Pilodius are presently recognized as having chelipeds with the fingers hollowed at the tip. The reduced beak at the tip of the gonopod of Garthiella aberrans is not like the larger, fuller beaks of species of Pilodius. In species of Pilodius the orbital hiatus varies from being open to give the antennal flagellum access to the eye, to being closed. In general, though, the orbital hiatus is not wide, whereas, in Garthiella aberrans the orbital hiatus is quite wide. Garniclla aberrans (Rathbun, 1906), new combination Figs. 1, 2 Chlorodopsis aberrans Rathbun, 1906:859, fig. 20.—Edmondson, 1925:43; 1946:295:; 1962:274, fig. 20a.—Seréne & Van Luom, 1958:90, 91; 1959:302, 328, text-figs. 2L, SL, pl. 2 fig. D, pl. 3 fig. K.—Forest & Guinot, 1961:89. Material examined. —3 males, 6 females, 2 juveniles, BPBM $1351, Johnston Island, coll. C. H. Edmondson, Jul 1923. Redescription.—Carapace hexagonal to transversely subovate, flattish from side to side, slightly convex longitudinally, front turned down and convex; length about % width (Fig. 1a). Regions and subregions sep- arated by wide smooth grooves lacking hair. Gastric and branchial regions defined and subdivided; 2M incompletely subdivided. Cardiac and intestinal regions blending into 57 branchial region with no clear divisions. Anterior of carapace sculptured with sharp tubercles decreasing in size posteriorly to granules; scattered short hairs originating at many tubercles and granules. Front bilobed and slightly greater than ' carapace width. Frontal lobes rounded, granular, and slightly produced adjacent to U-shaped median sinus; small lateral fron- tal lobes separated by shallow emargina- tions. Orbital margins spinulose with 1 me- dian dorsal and 1 ventrolateral notch; wide smooth groove just posterior to orbital mar- gins continuous behind front. Frontal-or- bital border about *4 carapace width. Anterolateral border of carapace granular and divided into 4 lobes or teeth (excluding external orbital angle). First lobe granular and blunt, originating behind and below outer orbital angle; posterior 3 lobes gran- ular and spinose. First 3 lobes obliquely di- rected anteriorly, increasing in size poste- riorly; fourth lobe slightly smaller than third, more laterally directed. Posterolateral bor- der slightly longer than anterolateral, and rather straight; posterior border slightly emarginate at midline. Pterygostomial region similar to dorsum, granular to spiny (Fig. 1b). Third maxilliped densely bristled medially, with scattered hairs over exterior surface (Fig. 2a). Palatal ridges developed only posteriorly, not ex- tending forward to anterior border of buccal cavern. Basal antennal article broad, inner angle touching ventral prolongation of front; outer angle prolonged partway into wide or- bital hiatus, not closing hiatus to antenna (Fig. 1b). Chelipeds slightly unequal, tubercular, with scattered short hairs. Tubercles on wrist and most of hand conical and sharp; lower surface of propodus with rounded tubercles. Two teeth on inner margin of carpus, in- dicated by slightly larger tubercles. Upper and lower borders of palm rounded; spi- nules on outer surface of palm tend to form lines, especially middle row. Fingers deeply grooved, acute (Fig. 2b); spinous tubercles 58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON oO Fig. 1. extending onto dorsal surface of dactylus about half its length, and onto outer face of pollex between grooves (more developed in females); teeth well developed; tip of dac- tylus curved, overlapping pollex internally when fingers closed, leaving no gap between teeth. Ambulatory legs lightly granular, upper margins spinulate (Fig. 2c). Upper margin of propodus, and especially carpus, bicari- AB VBA a2 Mb geo OE Cae ee et mle eae i > ar alee ae A aa A A Rn ned Bad aS mM A a, on aan BGA a BU Tie. Garthiella aberrans: a, Male, dorsal view; b, Front. nate. Legs with scattered long hairs, more dense on upper and lower margins. Abdomen of male with 7 segments, third to fifth fused, suture lines not recognizable (Fig. 2d). Abdomen of female oval, with 2 longitudinal grooves joining at seventh seg- ment (Fig. 2e); fringe of long hair around border, outer surface lightly covered by short hair. First pleopod of male curved, with sev- VOLUME 99, NUMBER 1 59 Fig. 2. Garthiella aberrans: a, Third maxilliped; b, Chela of male; c, Pereiopod; d, Abdomen of male; e, abdomen of female; f, g, First pleopod of male. eral long setae near tip; conical spines on distal half; beak reduced (Fig. 2f, g). Remarks. —Garthiella aberrans (Rath- bun, 1906) was originally described from a single male specimen (USNM 29434). It was dredged in the vicinity of Nihoa Island (Modu [Moku] Manu or Bird Island) at U.S.F.C. Steamer Albatross Station D.4146, at a depth of 42—48 meters, on a bottom of coarse coral, sand, and forams. The only other reported specimens were collected at Johnston Island (Edmondson 1925). The Bishop Museum catalog states that 15 spec- imens were collected at Johnston Island, as reported by Edmondson (1925). However, Edmondson (1962) stated that there were 60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 13 specimens in the Bishop Museum col- lections, of which only 11 can now be lo- cated. Unlike the Nihoa specimen from deeper water (42-48 meters), Edmondson (1962) reported that the Johnston Island speci- mens were collected in shoal water. This information is not on the specimen label, in the Bishop Museum catalogue, or re- ported previously by Edmondson (1925). Because Edmondson collected the Johnston Island specimens, it must be presumed that these depth records are from memory or, less probably, from private field notes. Distribution. —Near Nihoa Island, Northwestern Hawaiian Islands (Rathbun 1906); and Johnston Island (Edmondson 1925). Acknowledgments I would like to thank Drs. C. C. Chris- tensen, J. S. Garth, J. A. Titgen, and M. K. Wicksten for critically reading the manu- script and offering valuable comments. Dr. Garth was especially helpful by carefully ex- amining some of the specimens. Literature Cited Dana, J. D. 1851. Crustacea Grapsoidea (Cyclome- topa, Edwardsi1): Conspectus Crustaceorum quae in Orbis Terrarum circumnavigatione, Carolo Wilkes e Classe reipublicae Foederatae Duce. — Proceedings of the Academy of Natural Sciences of Philadelphia 5:247—254. Edmondson, C. H. 1925. Marine zoology of tropical Central Pacific. Crustacea.— Bernice P. Bishop Museum Bulletin 27:3-62, figs. 1-8. 1946. Reef and shore fauna of Hawaii.— Bernice P. Bishop Museum Special Publication 22:i-i1i, 1-381, figs. 1-223. . 1962. Xanthidae of Hawaii.— Occasional Pa- pers of Bernice P. Bishop Museum 22(13):215— 309, figs. 1-34. Forest, J.,and D. Guinot. 1961. Crustacés Décapodes Brachyoures de Tahiti et des Tuamotu. Jn Ex- pédition frang¢aise sur les récifs coralliens de la Nouvelle-Calédonie. Volume préliminaire.— Paris, Editions de la Fondation Singer-Polignac, pp. I-XI, 1-195, figs. 1-178, pls. 1-18. Milne Edwards, A. 1873. Recherches sur la faune carcinologique de la Nouvelle-Calédonie, Deux- iéme Partie.—Nouvelles Archives du Muséum d’Histoire Naturelle (Paris) 9:155—332, pls. 4— 18. Rathbun, M. J. 1906. The Brachyura and Macrura of the Hawaiian Islands. — Bulletin of the United States Fish Commission for 1903, part 3:827-— 930, text-figs. 1-79, pls. 1-24. Seréne, R., and N. Van Luom. 1958. Chlorodopsis (Brachyura) du Viet-Nam.—Annales de la Fa- culté des Sciences de Saigon 1958:87—147, text- figs. 1, 2, pls. 1-4. ——., and . 1959. Note additionnelle sur les espéces de Chlorodopsis (Brachyures).—An- nales de la Faculté des Sciences de Saigon 1959: 301-340, text-figs. 1-5, pls. 1-3. Division of Invertebrates, Bishop Mu- seum, P.O. Box 19000-A, Honolulu, Ha- wail 96817. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 61-64 GEOGRAPHIC VARIATION IN THE WHITE-MANTLED BARBET (CAPITO HYPOLEUCUS) OF COLOMBIA (AVES: CAPITONIDAE) Gary R. Graves Abstract. — Capito hypoleucus, a Colombian endemic currently considered monotypic, is divisible into three subspecies that differ in coloration. The nominate race is restricted to the foothills and lower slopes of the northern end of the central Andes from Valdivia, Antioquia, east to the eastern slope of the Serrania de San Lucas, Bolivar. Populations from the valley of the upper Rio Porce (C. h. carrikeri n. subsp.) and the middle Magdalena Valley (C. h. ex- tinctus n. subsp.) are described as new. Deforestation is responsible for the presumed extinction of C. h. extinctus. The White-mantled Barbet (Capito hy- poleucus), a poorly-known Colombian en- demic, is restricted to humid foothill forest (200-1500 m elevation) from the east bank of the lower Rio Cauca, east around the northern end of the central Andes, and south to the middle Magdalena Valley (Hilty and Brown, in press). The species was known from fewer than a dozen specimens until Melbourne A. Carriker, Jr. and assistants (1947-1952) collected a series of 22 skins from four localities (Fig. 1) spanning the geographic range of the species. Examina- tion of these specimens, now deposited in the National Museum of Natural History, Smithsonian Institution (USNM), indicates that C. hypoleucus, currently considered to be monotypic (Peters 1948), is divisible into three taxonomically distinct populations. Capito hypoleucus hypoleucus (Salvin) Capito hypoleucus Salvin, 1897:xvi. Val- divia (3800 ft.), Antioquia, Colombia. Characters. —The sides of the throat are lightly tinted with brown of the same color as that of the pectoral band. Dorsally, yel- low pigment is restricted to the mantle. The flanks and belly are lightly washed with pale yellow. Distribution. — Foothills and lower slopes at the northern end of the Central Andes from Valdivia, Antioquia, east to the east slope of the Serrania de San Lucas (Vola- dor), Bolivar. Specimens examined. — Antioquia: Puer- to Valdivia (1 2, AMNH); Valdivia (3 44, 1 2, USNM); La Frijolera (1 6, AMNH).— Bolivar: Volador (2 66, 3 92, USNM). Remarks. —The flanks of birds from Vo- lador tend to be more intensely yellow than those of specimens from the west slopes of the Central Andes in Antioquia, but there appear to be no other significant plumage differences between the two populations. Specimens taken in the Valdivia region more than 40 years apart do not differ noticeably in plumage color. There is little difference in size among any of the populations ex- amined. Carriker and an assistant collected along the road above Puerto Valdivia from 19 May to 19 June 1948. The four USNM spec- imens (401684—687) were collected on 25 May, in “badly cut over’’ forest on steep slopes below the road (Carriker’s field jour- nal deposited in USNM). One male had en- larged testes. Carriker collected at Volador from 5 May to 1 June 1947; the five C. hypoleucus 62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (USNM 392443-447) were taken on 19-31 May (6, 23 May, testes enlarged). Capito hypoleucus carrikeri, new subspecies Holotype. —USNM 425942, adult male from Botero, Antioquia, Colombia, eleva- tion 3600 ft (1098 m). Collected 31 Aug 1950 by M. A. Carriker, Jr.; original num- ber 18918. Characters.— Yellow pigments in carri- keri are more extensive throughout the non- black portions of plumage than in Aypoleu- cus. The yellow of the mantle extends up the nape to the rear of the crown, as opposed to being restricted to the mantle as in hy- poleucus. Ventrally, the yellow of flanks, belly, and breast below the pectoral band is brighter and much more extensively dis- tributed. The throat is white with a faint yellow tint, lacking the brown tint found in hypoleucus. The brown pectoral band is slightly lighter in shade than in hypoleucus. Measurements of holotype (mm).— Wing chord, 88.2; tail, 57.7; exposed culmen, 24.2; width lower mandible, 13.7. Distribution.—Known so far only from the type locality. Specimens examined.— Antioquia: Bo- tero (3 36, 6 92, USNM). Etymology.—Named for Melbourne A. Carriker, Jr., who was responsible for col- lecting and preparing the type series. Remarks. —This population is apparently restricted to the drainage of the northward flowing Rio Porce, a tributary of the Rio Nechi. Slopes above 1500 m elevation, de- forestation, and semiarid habitat around the periphery of the valley form barriers to the east, west and south. Contact with hypoleu- cus may have occurred in the past at the foot of the steep valley formed by the Rio Porce. Carriker worked at Botero, a small village on the west bank of the Rio Porce on the rail line to Medellin, from 17 August to 9 September 1950. Specimens of C. hypoleu- cus (USNM 425934—942) were collected on 30 August-8 September, from a “‘tall tree with small fruit’’ and “‘from large clumps of mistletoe in large tree,”’ in the patchwork of forest and pasture on the west bank of the river. Three females and one male had en- larged gonads; a single bird of each sex was in an immature plumage, which lacks the brown pectoral band. Capito hypoleucus extinctus, new subspecies Holotype.—USNM 436335, adult male from Hacienda Sofia, Rio Samana, Caldas, Colombia, elevation 3750 ft (1143 m). Col- lected 21 May 1951 by M. A. Carriker, Jr., original number 20235. Characters. — Feather tips on the mantle and nape are white with a faint buffy tint, not strongly yellow as in hypoleucus (man- tle) or carrikeri (mantle and nape). Yellow on the flanks and lower belly is much re- duced. The pectoral band tends to be darker brown than in hypoleucus and much more so than in carrikeri. The throat is similar to hypoleucus. Measurements of holotype (mm).— Wing chord, 89.0; tail, 58.0; exposed culmen, 23.5, width lower mandible, 14.0. Distribution. — Foothills on both sides of the valley of the Rio Magdalena in the vi- cinity of Honda. Specimens examined. — Tolima: within 20 miles (west) of Honda (2 46, 1 2, AMNH).— Cundinamarca: Carmen de Yacopi (1 8, AMNH).—Caldas: Hacienda Sofia (2 63, 2 22, USNM). Etymology.—Latin, extinctus, dead or destroyed, in reference to the probable ex- tinction of this taxon through deforestation of the middle Magdalena Valley. Remarks. —The series of specimens from the east bank of the Rio Magdalena are in- adequate to determine if populations east (Carmen de Yacopi) and west (Hacienda So- fia, west of Honda) of the Rio Magdalena are subspecifically different. VOLUME 99, NUMBER I 63 Fig. 1. Map showing the distribution of Capito hypoleucus in northwestern South America. Specimen lo- calities are identified with symbols (filled circles = C. h. hypoleucus; starred circle = C. h. carrikeri; empty circles = C. h. extinctus). Fine lines indicate provincial boundaries; cross-hatched areas indicate elevations above 1000 m. Carriker worked at Hacienda Sofia, along the road to Sons6n, from 10 May to 1 June 1951. Four specimens of C. hypoleucus (USNM 436334-337) were collected, two of which had enlarged gonads (6, 31 May; ? 29 May). Carriker’s only direct mention of C. hypoleucus from this locale was of an individual taken from a mixed-species flock. Discussion The middle and lower Magdalena and Cauca valleys have been heavily deforested since the 19th century and perhaps as far back as Pre-Columbian times (see Chap- man 1917, Hilty 1985). Hilty (1985) esti- mated that Capito hypoleucus, as well as a 64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON number of other species inhabiting humid forest on lower Andean slopes, have under- gone a historic range loss of more than 50% due to habitat destruction. Before the era of habitat destruction by humans, populations of C. hypoleucus were probably in genetic contact, and geographic variation in plum- age may have been more or less clinal in nature. The question of whether the rec- ognized subspecies are arbitrary subdivi- sions of an unperceived cline or discrete populations, in all probability can no longer be answered. There are no data available on the present status or distribution of any of the historically known populations, but the fragmented state of the remaining lower montane humid forest suggests that any re- maining populations are insular and genet- ically isolated. The complete deforestation of the floodplain and foothills of the middle Magdalena Valley during the past 30 years suggests that C. h. extinctus may already be extinct. Acknowledgments I thank the curators of the National Mu- seum of Natural History (USNM) and American Museum of Natural History (AMNH) for permission to examine speci- mens. William L. Brown, Steven L. Hilty, Storrs L. Olson, and J. V. Remsen, Jr. pro- vided information and commented on the manuscript. Olson supplied the base map. This work was supported by a Smithsonian Postdoctoral Fellowship. Literature Cited Chapman, F. M. 1917. The distribution of bird-life in Colombia.—Bulletin of the American Mu- seum of Natural History 36:1-—729. Hilty, S. L. 1985. Distributional changes in the Co- lombian avifauna: a preliminary blue list. Jn P. A. Buckley, M. S. Foster, E. S. Morton, R. S. Ridgely, and F. G. Buckley, eds., Neotropical Ornithology. — Ornithological Monographs No. 36. Pp. 992-1004. , and W. L. Brown. [In Press]. A Guide to the Birds of Colombia. Princeton, New Jersey, Princeton University Press. Peters, J. 1948. Check-list of birds of the World, vol. 6. Cambridge, Massachusetts, Museum of Comparative Zoology, 259 pp. Salvin, O. 1897. [Descriptions of five species of South American birds.]—Bulletin of the British Or- nithological Club 7:xv—xvii. Department of Vertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 65-70 LASIONECTES ENTRICHOMA, NEW GENUS, NEW SPECIES, (CRUSTACEA: REMIPEDIA) FROM ANCHIALINE CAVES IN THE TURKS AND CAICOS, BRITISH WEST INDIES Jill Yager and Frederick R. Schram Abstract. — Lasionectes entrichoma, a new genus and species of remipede from the Turks and Caicos, British West Indies, is described. It possesses characters which warrant its placement into a new genus of the crustacean class Remipedia. In 1979, Speleonectes lucayensis Yager, the first species representing the new class of Crustacea, Remipedia, was collected from Lucayan Cavern, an anchialine cave on Grand Bahama Island in the northern Ba- hamas. Subsequent exploration of other caves in the West Indies has resulted in the discovery of the new genus and species of remipede described here, as well as several other new species of remipedes to be de- scribed later. Although a separate country politically, the Turks and Caicos are geographically a southeast extension of the chain of islands making up the Bahamas. Their geologic makeup is also similar, in that the islands are the tops of exposed shallow water car- bonate banks separated by deep water chan- nels. Old Blue Hill Cave, the type-locality for these new remipedes, is located on the is- land of Providenciales. The submerged cave system is developed in an inland ridge and has two entrances along the collapsed mar- gin of a large sinkhole. The western opening is in a large, shallow, brackish pool which slopes abruptly down along the north mar- gin of the sinkhole into the dark passages of the cave system. The pool is entirely open to sunlight, and is rich in organic material which has turned the water brown and re- duced the level of visibility. The pool sup- ports a rich biota of algae, the cyclopoid copepod Apocyclops (Metacyclops) distans, the amphipod Spelaeonicippe provo Stock and Vermeulen, and a very dense popula- tion of the caridean shrimp Typhlatya gar- ciai Chace. The new remipede is found in the twilight zone which begins at about 5 m, and in the dark, deeper passages of the cave which have been explored to about 20 m. The eastern entrance into the cave sys- tem is at the bottom of a narrow, dimly illuminated fissure. In contrast to the west- ern entrance, the water in this pool is very clear and the numbers of Typhlatya are greatly reduced from thousands of individ- uals to less than a hundred. The new rem- ipede is relatively abundant, especially when compared to Speleonectes lucayensis. Hundreds of individuals have been seen while cave diving in both entrances of this cave system. Lasionectes, new genus Diagnosis. —Second maxilla and maxil- liped subchelate and distinctly more robust than first maxilla; terminal segments bear- ing trifid claw with comb-like row of spi- nules between large central spine and two flanking spines; both appendages with short setae along entire medial margin. Etymology. —From the Greek /asios meaning hairy, and nectes meaning swim- mer; a reference to the prevalence of fine 66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. hair-like setae on the second maxilla and maxilliped. Gender masculine. Type-species. —(By monotypy) Lasi- onectes entrichoma, new species. Lasionectes entrichoma, new species Figs. 1-3 Type-material.—Holotype. Adult, 31.5 mm, USNM 216978; in brackish water, Old Blue Hill Cave, Providenciales Island, Turks and Caicos, British West Indies, 6 Apr 1983.—Paratypes, 39 specimens, both juvenile and adult, from Old Blue Hill Cave and Airport Cave on Providenciales Island, and from Cottage Pond, Middle Caicos Island, Turks and Caicos, B.W.I., Coll. J. Yager and D. Williams. Retained for fur- ther studies in the collections of the San Diego Natural History Museum. Diagnosis. — With characters of the genus. Description. —Cephalon small, about 13% total body length (Fig. 1). Cephalic shield somewhat tapered anteriorly and folded over anterior margin of cephalon. Maximum of 32 postcephalic trunk segments; first trunk segment short, with greatly reduced pleura, partly covered by cephalic shield; pleura of trunk segments posterior to first well de- veloped, projecting laterally, reduced in size anterior to anal segment. Trunk sternites with distinct transverse bars, not developed Lasionectes entrichoma: Ventral view of animal showing swimming position. as plates; bar on segment 14 with large tri- angular process or flap on either end adja- cent to limb base and covering genital pore; transverse bars posterior to segment 14 and continuing to about segment 24; thereafter developed as small triangular processes, be- coming more prominent posteriorly in the series. Anal segment about as wide as long, anus terminal; caudal rami slightly less than length of anal segment, with 9 moderately long terminal setae and 2 distomedial setae (Fig. 2). Frontal filaments (Fig. 2A) small, rod-like, anteromedial to first antenna, with thumb- like process on posteromedial surface. An- tenna 1 (Fig. 2B) large, well developed, biramous. Basal segment of peduncle en- larged, bearing 3—4 rows of densely packed, long, lash-like esthetascs draped posteriorly over antenna 2 toward labrum. Dorsal ra- mus long, with 12 segments; ventral ramus about one-half to two-thirds length of dorsal ramus, 8—9 segments; segments of both rami slender with fine hair-like setae along ven- tral margins and in tufts distoventrally; dis- tal segment of both rami with 4-6 terminal setae. Antenna 2 (Fig. 2C) biramous, well de- veloped, moderate in size, not extending be- yond cephalic shield. Two-segmented pro- topod, medial margins of which bearing short simple setae. Three-segmented endo- VOLUME 99, NUMBER 1 67 AR Whe ree endites endopod Fig. 2. Left appendages and related structures of Lasionectes entrichoma: A, Frontal filament; B, Antenna 1; C, Antenna 2; D, Labrum; E, Mandible; F, Maxilla 1; G, Maxilla 2; H, Maxilliped; I, Tenth trunk limb; J, Terminal part of body with caudal rami. Scales = 0.2 mm. pod; first segment with about 13 plumose setae laterally, second segment with about 11 or 12, third segment with about 24 along entire margin and those most distal forming double row. Exopod a single, large, oval scale with about 35—40 long plumose setae along entire margin. Labrum (Fig. 2D) a large fleshy lobe, nar- row anteriorly, broad posterior section with transverse groove and fossa with densely packed short ribbon setae. Mandibles asym- metrical. Right mandible (Fig. 3B) with 3- cusped incisor process and 3-cusped lacinia mobilis. Left mandible (Figs. 2E, 3A) with 4-cusped incisor process and crescent- shaped lacinia mobilis. Molar processes densely spinose, semi-arcuate. Paragnaths round, flattened lobes, lateral and posterior to mouth, covered with fine ribbon setae. First maxilla (Fig. 2F) 6-segmented, uni- 68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON molar process incisor process Fig. 3. Appendages of Lasionectes entrichoma in greater detail. A, left mandible; B, right mandible; C, Maxilla 1, tip; D, Maxilla 2 anterior view; E, Maxilla 2 posterior view; F, Maxilla 1 endites of first segment. Scales = 100 wm. ramous, subchelate, robust. First or proxi- mal segment with 2 well developed endites (Fig. 3F); proximal endite terminates ven- tro-posteriorly in long stout spine, with 4— 6 stout spine-like setae dorsally; distal en- dite a broad, plate-like flap with 6 short, stout, spine-like setae along crest, flanked by many tiny setae, 8 short to long simple setae on anterodistal margin and 4 on pos- terodistal margin. Second segment with cone-like endite terminating in 2 robust spine-like sete and 1 or 2 small, simple se- tae. Third segment robust, with medial, subtriangular, double-crested endite bear- ing dense row of long, simple setae along each crest, and | short, stout terminal spine at apex of endite; anterior crest row with about 16 large setae, slightly longer than VOLUME 99, NUMBER 1 posterior row of about 24 short to moderate setae. Fourth segment long, robust, with about 8 simple setae on distomedial half of segment and 2 clusters of 8—10 simple setae on antero- and posterodistal margins. Fifth segment short, with 2 distal clusters of about 12 simple, moderately long setae on ante- rior and posterior medial margins and 2 dis- tal clusters of about 6 on anterior and pos- terior lateral margin. Sixth segment (fig. 3C) very short, terminating in single, long, tal- on-like claw with large terminal pore; tuft of about 10 long, fine, simple setae at medial base of claw. Principle flexion point of ap- pendage between segments 3 and 4, with segments 4-6 cradled in trough between double rows of setae on segment 3 when flexed. Secondary flexion occurring between segments 2 and 3, with 2 large setae of seg- ment 2 held in opposition to apex of sub- triangular endite of segment 3 during flex- ion. Second maxilla (Fig. 2G) 7-segmented, long, about twice the length of first maxilla, uniramous, subchelate; first segment with 3 digitiform endites increasing in size distally, 2 small, subconical lobes posteriorly asso- ciated with second and third endites each bearing 3-5 short to moderate setae; first endite small, apex with | short, terminal spine and about 4-6 tiny hair-like setae, row of 4-6 small to moderate, simple setae on lateral margin; second endite with 1 mod- erately long terminal spine, about 6—8 small, simple, setae apically, a row of about 3 mod- erate, simple setae on lateral margin; third endite with | large spine terminal seta, 8— 10 small, simple apical setae, a row of about 3-5 moderately long, simple, lateral setae. Segment 2 with large, medial, thumb-like lobe with clusters of moderate to long, sim- ple setae in 2 rows, the anterior row of setae longer. Segment 3 long, wide, with subtrian- gular endite bearing 2 rows of dense, short, simple setae along entire medial margin of segment. Segments 4, 5, and 6 with dense rows of subequal setae along entire medial margin of segments; segment 5 with 1-2 69 simple setae on distolateral margin; seg- ment 6 short, with 2 clusters of about 4 moderately long, simple setae on anterior and posterior distolateral margin and 2 clus- ters of about 8 long, fine setae on anterior and posterior distomedial margin. Segment 7 (Figs. 3D, 3E) very short, terminating in complex trifid claw with a central long spine and 2 shorter flanking spines, a comb-like row of several smaller spines between the central spine and posterior flanking spines. Thumb-like pad with fan-like setose margin opposed to claw. Maxilliped (Fig. 2H) similar to second maxilla but markedly longer and more ro- bust, with at least 8 segments. Segment 1 with several weakly developed median lobes, the most prominent of which with about 5 terminal, long, simple setae. Segment 2 with small, thumb-like lobe with two rows of clustered setae in V-shape, the anterior row with short to moderate simple setae, pos- terior with short, hair-like setae. Segment 3 long, wide, with subtriangular endite bear- ing 2 rows of setae along medial margin as in second maxilla. Segments 4, 5, 6 and 7 with dense rows of subequal setae as in sec- ond maxilla. Segment 6 with 1 short seta distolaterally. Segment 7 with several mod- erately long setae on anterior and posterior distolateral margin and 2 clusters of about 6 on anterior and posterior distomedial margin. Segment 8 with terminal trifid claw complex as in maxilla 2. Trunk appendages (Fig. 21) biramous, se- tose, laterally directed paddles. Protopod fleshy, exopod 3-segmented, endopod 4- segmented. Distal segments of both rami oval in shape. Etymology. —From the Greek entri- choma meaning eyelash, a reference to the long lash-like setae on the base of the first antenna; used as a noun in apposition. Remarks. — Additional specimens of La- sionectes, were collected from Airport Cave, another anchialine habitat on Providen- ciales. Associated fauna were a new family of caridean shrimp (C. W. Hart, Jr., pers. 70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON comm.), a new genus of leptostracan, Speo- nebalia cannoni Bowman, Yager, and Iliffe, 1985, and the amphipod Spelaeonicippe provo. Cottage Pond on Middle Caicos Is- land was also found to be inhabited by La- sionectes and another as yet undescribed species of remipede. Although blind cave fish are known from many West Indian caves, they are noticeably absent in the abovementioned caves. While Lasionectes bears some relation- ship to Speleonectes lucayensis Yager, 1981, the differences between the two taxa seem too great to maintain the two species within a single genus. The two species differ in the size ratio of the feeding appendages. The first maxilla of Speleonectes is the most ro- bust when compared to the second maxilla and maxilliped. Lasionectes has a small first maxilla when compared to the second max- illa and the very robust maxilliped. With additional remipede material now available from other West Indian caves and also the Canary Islands, it will be possible to assess the relationships within the group in greater detail. Acknowledgments The authors wish to acknowledge the help of the following people in collecting Turks and Caicos crustaceans: Howard Cos- grove, Mary Ellen Eckoff, Sam Harvey, Paul Hobbs, Tom Iliffe, Wayne Kafcsak, Dennis Williams, and the PRIDE Foundation on Pine Cay, Turks and Caicos. We would like to thank the following for specimen iden- tifications: Thomas E. Bowman, C. W. Hart, Jr., John R. Holsinger, Raymond B. Man- ning, Janet Reid. Liselotte Ohman (G@te- borg, Sweden) drew the swimming animal in Figure 1. Michael J. Emerson (San Diego Museum) prepared the illustrations for Fig- ures 2 and 3. Research was supported by NSF grant BSR 82-12335 to FRS. Literature Cited Bowman, T.E., J. Yager, and T. M. Iliffe. 1985. Speo- nebalia cannoni, gen. sp. nov. from the Caicos Islands, the first hypogean leptostracan (Neba- liacea: Nebaliidae). — Proceedings of the Biolog- ical Society of Washington 98(2):435-442. Yager, Jill. 1981. Remipedia, a new class of crustacea from a marine cave in the Bahamas.—Journal of Crustacean Biology 1:328-333. (JY) Department of Biological Sciences, Old Dominion University, Norfolk, Virgin- ia, 23508-8560; (FS) San Diego Natural History Museum, P.O. Box 1390, San Diego, California 92112. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 71-78 A REDESCRIPTION OF MICROCYCLOPS CEIBAENSIS (MARSH, 1919) (COPEPODA: CYCLOPOIDA) FROM MARSH’S SPECIMENS IN THE NATIONAL MUSEUM OF NATURAL HISTORY Janet W. Reid Abstract.—The neotropical freshwater cyclopoid copepod species Microcy- clops ceibaensis (Marsh, 1919) is redescribed from specimens from the type locality, deposited by Marsh in the National Museum of Natural History. Microcyclops diversus Kiefer, 1935, is a synonym. Locality records include Brazil, Costa Rica, Cuba, Haiti, Honduras, Uruguay, Venezuela, and possibly Mexico. Species of the freshwater cyclopoid co- pepod genus Microcyclops may be distin- guished by a complex of microcharacters. Because many characters now recognized to have taxonomic utility were not reported by earlier workers, and because some char- acters may be given different weight by con- temporary investigators, assignment of specimens to particular species is still a ten- uous business. Marsh (1919) described Cyclops ceibaeén- sis from a collection made at La Ceiba, Hon- duras, by F. J. Dyer. This species is now assigned to the genus Microcyclops Kiefer, 1929a. Some confusion has resulted from inadequacies in Marsh’s original descrip- tion: Kiefer (1929b) tentatively synony- mized C. ceibaénsis with Microcyclops var- icans (O. Sars, 1863), an opinion shared by Gurney (1933). On the basis of broader ex- perience with the range of variation within M. varicans, Kiefer later (1936) came to consider M. ceibaensis as a valid species and suggested that V/. diversus, described by him from Uruguay (1935) might be a synonym of ceibaensis. Lindberg placed M. ceibaensis as “‘species incertae sedis”’ in his list of Cy- clopoida from Central America and Mexico (1954b), though he later included it in a key to the genus (1957a). Smith and Fernando (1978) presented figures of a Cuban species which they attributed to M. ceibaensis, and commented that M. diversus is probably a synonym of the former species. On the other hand, Dussart (1984 and personal com- munication to the author) is of the opinion that M. diversus and M. ceibaensis are dis- tinct species. Marsh stated (1919) that the type speci- men of M. ceibaénsis had been “‘catalogued under No. 57392 in the collection of the United States National Museum.” This number is not traceable, and a careful search of the collections has failed to locate the type. However, several other specimens from the type locality, collected during the same period as the type, were found in the Marsh Collection of slides of Copepoda. The condition of the original slides permits a more extensive description than that fur- nished by Marsh. I here redescribe these specimens and compare them to closely re- lated forms. Marsh (1919) designated no paratypic specimens. Since, however, his comments and unpublished notes make it clear that his observations of the specimens described in the present article were used in part for the original description, they may be considered to be paratypes. Microcyclops ceibaensis (Marsh, 1919) Figs. 1-11 Cyclops ceibaénsis Marsh, 1919:546-547, figs. 1-5. (Honduras). Cyclops ceibaénsis. —Kiefer, 1929a:36-37, 72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 40; 1936:295—296.—Lindberg, 1954b: 486, 488. Cyclops varicans (part). —Gurney, 1933:255. Cyclops (Microcyclops) varicans (part).— Kiefer, 1929b:66; 1929a:36-37, 40. Microcyclops varicans (part).—Comita, 1950-37 2% Microcyclops diversus Kiefer, 1935:186— 187, figs. 10-13. (Uruguay).— Brehm, 1935:298-299, 305. (Uruguay).—Colla- do et al., 1984:94-97, figs. 26-28. (Costa Rica).— Dussart, 1984:41, 43, 57, fig. 18. (Venezuela).—Herbst, 1959:70.—Lind- berg, 1954a:214-216; 1954b:486-488; 1957a:168; 1957b:39.—Loffler, 1958:21; 1963:215; 1981:16.—Smith and Fernan- do, 1978:2020. Cyclops (Microcyclops) cf. diversus und cei- baénsis. —Kiefer, 1936:265, 267, 295— 296, figs. 68-70. (Haiti). Microcyclops ceibaensis.— Lindberg, 1954b: 488; 1957a:168.—Osorio Tafall, 1941: 343. (Mexico).—Smith and Fernando, 1978:2020, figs. 30-32. (Cuba); 1980:11, 18, figs. 7G—7I. (Cuba). Cyclops (Microcyclops) spec.— Herbst, 1959: 70-71, fig. 42. (Brazil). non Microcyclops diversus.— Harding, 1955: 243, figs. 68-69. (Peru). non Cyclops (Microcyclops) varicans. — Herbst, 1959:65-—66, figs. 29-31. (Brazil). Material examined. —Honduras, La Cei- ba, all dissected on slides, stained, and mounted in glycerine jelly: USNM 222298, 1 female (Marsh’s No. 4259, labelled “eggbearing’’), collected 19 Mar 1916; USNM 222299, 2 females (Marsh’s Nos. 4277 and 4280, labelled “‘eggbearing’’) and 1 female (Marsh’s No. 4283), collected 27 Sep 1916. All are labelled ““C. cubanensis”’ on the slides but “Cyclops ceibaénsis”’ in Marsh’s card file, and correspond to the de- scription of C. ceibaénsis. Marsh does not indicate whether collections were made from one or several ponds. Slides numbered 4281 and 4282, also in- dicated in the card file as being C. ceibaén- sis, are not present in the collection. Al- though No. 4277 was labelled “ant. abd. ft.”, only the antennule and antenna can now be located on the slide. Two copepo- dites (Marsh’s Nos. 4278 and 4284) were also listed in Marsh’s card file under C. cei- baénsis, but labelled only as ‘Cyclops’? on the slides. No. 4278 is partly dissected and the abdomen cannot be located on the slide. The caudal rami of specimen No. 4284, which is mounted whole, are very short. Since both are early copepodite stages (II or III), they cannot with certainty be assigned to M. ceibaensis. Slide No. 4280 contains the most com- plete specimen, which was used for the pres- ent diagnosis and figures. The slide con- tains, dissected: cephalothorax with antenna and some mouthparts attached; both anten- nules, separated; 1 maxilla; 1 maxilliped; legs 1, paired, having 3 external spines on exopod segment 2; 1 pair of legs having 4 external spines on exopod segment 2; | iso- lated leg with 3 external spines on exopod segment 2; | right and | left leg, attached one above the other, with 4 and 3 external spines on exopod segment 2 respectively; and abdomen with legs 5. Since 1 pair and 1 other leg (not leg 1) have 4 external spines on exopod segment 2, the spine formula of 3,3,4,3 given by Marsh for M. ceibaénsis is impossible. Marsh’s notes several times indicate con- fusion (which will be understood by anyone who has dissected a small copepod!) as to the dissections and therefore the armature of the swimming legs. The true spine for- mula is then 3,4,4,3, which is normal for the genus. The formula of 3,4,4,3 was also given by Smith and Fernando (1978) for Cuban specimens ascribed to M. ceibaensis. Assuming that Marsh dissected legs 3 and 4 together, a conclusion supported by the 2 strong terminal spines on endopod 2 of the supposed leg 4, the pair of legs with 4 ex- ternal spines on exopod 2 are therefore legs 2, and all 4 legs can be figured. Female.—Abdomen (Fig. 1) of 4 seg- ments, length 193 um. Anterior third of gen- VOLUME 99, NUMBER 1 73 b Figs. 1-6. Microcyclops ceibaensis, Marsh’s No. 4280, eggbearing female, La Ceiba, Honduras: 1, Abdomen and legs 5, ventral; 2, Antennule; 3, Antenna; 4, Maxillula; 5, Maxilla; 6, Maxilliped. Scale a, Fig. 1; scale b, Figs. 2-6; scales = 100 um. 74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 7-11. WTA WW yw Wy " Microcyclops ceibaensis; Figs. 7-10, Marsh’s No. 4280, eggbearing female, La Ceiba, Honduras: 7, Leg 1; 8, Leg 2; 9, Leg 3; 10, Leg 4. Fig. 11, Marsh’s No. 4259, female, La Ceiba, Honduras, Leg 5. Scale = 100 um. ital somite broadened, greatest width about equal to length; shape of seminal receptacle not distinguishable. Anal somite with spi- nules on posteroventral border. Caudal rami about 3.8 x longer than broad, bearing 4—5 spinules anterior to insertions of lateral and outer subterminal setae; lateral seta inserted at posterior fifth of ramus. Inner margins of rami smooth. Relative lengths of caudal se- tae as in Fig. 1 and Table 1. Antennule (Fig. 2) shorter than cephalo- thorax, of 12 articles; no hyaline lamella distinguishable on terminal articles. Anten- na, maxillula, maxilla and maxilliped as in VOLUME 99, NUMBER 1 Figs. 3-6 respectively. Mandible not ex- amined. Legs 1—4 (Figs. 7-10 respectively) with endopods and exopods of 2 articles; spine formula 3,4,4,3. Inner extension of basipod 2 of leg 1 with strong spine. Con- necting lamellae of legs 1-3 without con- spicuous ornamentation; lamella of leg 4 with 2 parallel rows of spinules on anterior surface. Exopods 2 of legs 2—4 each with group of 3-6 spinules on anterior surface. Endopod 2 of leg 4 about 2 longer than broad; inner terminal spine about 1.6 x longer than outer (Table 1). Free article of leg 5 (Figs. 1, 11) slightly tapering distally, about 3.5 x longer than broad, with minute spinule at midlength of inner margin and 1 long smooth terminal seta. 1 long seta in- serted anterolaterally to free article of leg 5. Leg 6 not visible. Comparative measurements of Marsh’s 3 most complete specimens appear in Table 1. The seminal receptacle and leg 6 are not visible on these specimens. Discussion A. The synonymy of Microcyclops cei- baensis (Marsh) and M. diversus (Kiefer): Kiefer (1935) cited as diagnostic features of M. diversus the length/width ratios of the caudal ramus and of the endopod 2 of leg 4; the proportions of the terminal spines of the latter article; leg 5 with its “‘kleines, aber gut ausgepragtes Dornchen’’; and the shape of the seminal receptacle. He also noted two horizontal rows of spinules on the connect- ing lamella of leg 4. As may be seen from Tables 1 and 2, the reported ranges of vari- ation in total length and proportions of cau- dal rami and 4th legs for most characters of M. diversus are similar to those of M. cei- baensis. The proportions of leg 5, with its tiny spinule on the inner ventral margin, the spinules on the anterior surface of leg 4 en- dopod 2, and the two parallel rows of spi- nules on the anterior surface of the basal lamella of leg 4 are also features shared by both species. Kiefer later (1936) com- mented on several similarities between M. 75 Table 1.—Measurements from Marsh’s specimens of Microcyclops ceibaensis, deposited in the National Museum of Natural History. Abbreviations: CR, cau- dal ramus; P4, leg 4; enp2, second (terminal) article of endopod. *Indicates specimen turned slightly laterally. Specimen no. 4280 4259 4283 Cephalothorax, length 330 wm — — Abdomen, length 193 296 296 Genital segment, length 90 90 94 Genital segment, width 90 90 100 CR, length 73 96* 76 CR, width 19 7 AO Antennule, length 220 215 — P4 enp?2, length 50 55 54 P4 enp2, width 24 24 23 P4 enp2, outer terminal spine 24 23 23 P4 enp2, inner terminal spine 40 40 36 CR, setae: Lateral 20 18 15 Dorsal 73 42 65 Inner terminal 55 48 50 Inner median terminal 275 300 280 Outer median terminal 173 180 165 Outer terminal 31 29 28 diversus and M. ceibaénsis, but since Marsh had provided neither exact measurements of ceibaénsis, the correct spinal formula nor a description of the connecting lamella of the 4th legs, Kiefer was unable to form a definite opinion as to the synonymy of these species. A useful diagnostic character in many cyclopoid copepod genera is the shape of the seminal receptacle. Unfortunately Marsh (1919) was unable to see this feature in his material, and it is not visible on any of his specimens. However, the forms of the sem- inal receptacle reported for different popu- lations of M. diversus vary so much (Figs. 12-16) that its value as a diagnostic char- acter in this case is dubious. Fig. 17 shows the shape of the receptacle of a specimen from Sao Paulo, Brazil, ascribable to M. ceibaensis. Dussart and co-workers’ reasons for considering M. diversus as distinct from M. ceibaensis appear to be the spine for- 76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Measurements of female specimensfrom different populations of Microcyclops ceibaensis. Ab- breviations: CR, caudal ramus; P4, leg 4; enp2, second (terminal) article of endopod; L:W, ratio of length to width. *Indicates measurement made from author’s figure. P4 enp2 Total length CR Terminal spines Length of enp2: Record (mm) L:W L:W inner : outer inner term. spine Honduras (1) _ 3.8-5.6:1 2.0-2.35:1 1.56-1.73:1 1.25-1.50:1 Uruguay (2) 0.96-1.0 3.76—4.27:1 2.09-2.43:1 1.26-1.41:1 1.85-1.97:1 Haiti (3) 0.54—0.60 4.0:1 2.38:1 1.61-1.80:1 1.19-1.35:1 Cuba (4) 0.60-0.65 about 5:1 about 3:1 1.6:1* 1.4:1* Venezuela (5) 0.88 4.5:1 DAM 1.4:1 1.9:1 Brazil (6) 0.92 5731 AASTUEN 1.44:1 1.44:1 Brazil (7) 0.75-0.76 3.62-3.72:1 2.21—2.31:1 1.52-1.58:1 1.43-1.75:1 Costa Rica (8) — 3.4:1* 2.0:1* 1.8:1* Neale (1) Marsh (1919), measurements presented in this article; (2) Kiefer (1935); (3) Kiefer (1936); (4) Fernando and Smith (1978); (5) Dussart (1984) and specimen from Camaguan, Venezuela; (6) Herbst (1959); (7) 3 specimens from Rio Una do Prelado, State of Sado Paulo, lent by C. E. F. da Rocha; (8) Collado et al. (1984). mulae, following Marsh’s erroneous de- scription; and the form of the seminal re- ceptacle in specimens from geographically distant populations ascribed either to M. diversus (Dussart 1984) or to M. ceibaensis (Collado et al., 1984), though they state that the species are “‘difficult to separate” (Collado et al., 1984). Although the shape of the receptacle is not distinguishable in the present specimens, it is my opinion that the reported variability of this character in these forms is such as to diminish its taxo- nomic utility. Microcyclops diversus should be considered a synonym of M. ceibaensis. Microcyclops ceibaensis as presently de- fined is at least a very variable species. Its records include freshwater ponds and lakes and one river in Brazil, Costa Rica, Cuba, Haiti, Honduras, Uruguay, and Venezuela. A record by Osorio Tafall (1941) from Lake Patzcuaro, Mexico, is dubious, since no fig- ures were furnished. Most records seem to be from the littoral zone. Loffler’s (1958) characterization of M. diversus as a pelagic species in cold polymictic lakes seems based on Harding’s record from the Peruvian pla- teau (Harding 1955); though this may not in fact be M. ceibaensis, as discussed below. B. Notes on some South American rec- ords of Microcyclops spp.: Harding (1955) reported that specimens ascribed to M. di- versus from Peru lacked a spine on the inner extension of basipod article 1 of leg 1. The neotropical species M. anceps (Richard, 1897), and its subspecies M. anceps paux- ensis Herbst, 1962 (=M. anceps minor Dus- sart, 1984); M. finitimus Dussart, 1984, known from Venezuela and French Guiana (Dussart 1983); and M. crassipes (O. Sars, 1927) from South Africa have also been re- ported as lacking this spine. However, Harding’s fig. 68 clearly shows two rows of spinules on the connecting lamella of the 4th leg, a diagnostic character shared by M. ceibaensis and M. finitimus. It is possible that Harding’s specimens from Lakes Umayo and Arapa refer to the latter species. Herbst (1959) described a Cyclops (Mi- crocyclops) varicans from the State of Sao Paulo, Brazil, with two rows of spinules on the leg 4 lamella and leg 4 endopod article 2 2.82—3.24x longer than wide. Microcy- clops varicans lacks the former character and the terminal article of leg 4 endopod is shorter (Dussart 1969; Gurney 1933). The placement of the inner spinule of leg 5 of Herbst’s specimen “‘neben der langen End- borste”’ is similar to varicans, but Herbst’s specimens are probably referable to M. fin- itimus or to M. ceibaensis. VOLUME 99, NUMBER 1 Vy Figs. 12-17. Seminal receptacles of Microcyclops: 12-13, M. diversus, Uruguay: 12, After Fig. 10 of Kiefer (1935); 13, After Fig. 13 of Kiefer (1935); 14, M. cf. diversus and ceibaensis, Haiti, after Fig. 68 of Kiefer (1936); 15, M. diversus, Camaguan, Venezuela, coll. 13 Oct. 1981, from specimen lent by Dr. B. H. Dussart; 16, M. ceibaensis, Costa Rica, after Fig. 28 of Collado et al. (1984); 17, M. ceibaensis, Rio Una do Prelado, State of Sao Paulo, Brazil, coll. 7 June 1984, from specimen lent by Dr. C. E. F. da Rocha. Scale = approximately 100 pm. Herbst (1959) also described a Cyclops (Microcyclops) spec. from Sao Paulo with caudal rami 5.19 longer than wide, and remarked on the similarity of this form to M. diversus. In view of the reported varia- tion of the length of the caudal ramus in M. ceibaensis, Herbst’s specimen should be re- ferred to this species. C. The Microcyclops varicans Group in South and Central America: Several species of Microcyclops with a spinule at or near the midpoint of the inner surface of leg 5 have been recorded from South and Central America and some Caribbean islands. These include M. varicans varicans (Sars, 1863), M. y. subaequalis (Kiefer, 1928), M. elon- gatus (Lowndes, 1934), as well as M. cei- baensis and M. finitimus. As discussed above, M. finitimus lacks a spine at the inner corner of basipod 2 of leg 1 and should therefore be considered part of the M. an- ceps group (Dussart 1983, 1984). Both sub- species of M. varicans have a relatively short caudal ramus (3.5—4.5 x longer than broad), and lack spinules on the connecting lamella of leg 4. Microcyclops elongatus is distin- guished by its very long caudal rami (5-6 x longer than broad); a row of spinules above the lateral seta of the caudal ramus, reaching to the midpoint of the dorsal and ventral surfaces of the ramus; and spinules on the anterior face of the terminal articles of the 78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON endopods of legs 2—4 borne on a “definite scale’’ (Lowndes 1934). Acknowledgments I am grateful to Dr. Bernard H. Dussart for lending a specimen, and for providing a prepublication draft of the article by Col- lado et al.; to Dr. Carlos Eduardo F. da Ro- cha for lending specimens; and to Dr. Thomas E. Bowman for helpful comments on the manuscript. Literature Cited Brehm, V. 1935. Uber die Siisswasserfauna von Uru- guay.— Archiv fiir Hydrobiologie 28:295-309. Collado, C., D. Defaye, B. H. Dussart, and C. H. Fer- nando. 1984. The freshwater Copepoda (Crus- tacea) of Costa Rica with notes on some species. —Hydrobiologia 119:89-99. Comita, G. W. 1951. Studies on Mexican cope- pods.—Transactions of the American Micro- scopical Society 70(4):367-379. Dussart, B. H. 1983. Copépodes d’eau douce de Guy- ane francaise.—Revue d’Hydrobiologie tropi- cale 16(4):321-325. 1984. Some Crustacea Copepoda from Ven- ezuela.— Hydrobiologia 113:25-67. Gurney, R. 1933. British Freshwater Copepoda. Vol. III. The Ray Society, London. xxix + 384 pp. Harding, J. P. 1955. The Percy Sladen Trust Expe- dition to Lake Titicaca in 1937. Report No. XV. Crustacea Copepoda. — Transactions of the Lin- nean Society of London, Series 3, 1(3):219-—247. Herbst, H. V. 1959. Brasilianische Sitisswassercyclo- poiden (Crustacea: Copepoda). — Gewasser und Abwasser 24:49-73. Kiefer, F. 1929a. Zur Kenntnis einiger Artengruppen der Stisswasser-Cyclopiden.—Zeitschrift fiir wissenschaftliche Zoologie 133:1-56. 1929b. Crustacea Copepoda. 2. Cyclopoida Gnathostoma. — Das Tierreich 53:xvi + 102 pp. . 1935. Neue Stisswassercyclopiden (Crustacea Copepoda) aus Uruguay.—Zoologischer An- zeiger 109:181-188. 1936. Freilebende Siiss- und Salzwasserco- pepoden von der Insel Haiti.—Archiv fiir Hy- drobiologie 30:263-317. Lindberg, K. 1954a. Cyclopides (Crustacés copé- podes) de l’Amérique du Sud.— Arkiv for Zoo- logi 7(11):193-222. . 1954b. Cyclopoides (crustacés copépodes) du Mexique.— Arkiv for Zoologi 7(23):459—489. 1957a. Cyclopides de la Céte d’Ivoire.— Bulletin de l'Institut francaise d’Afrique noire 19:137-179. 1957b. Cyclopides (crustacés copépodes) ré- coltés au Pérou par le Dr. Hernando de Macedo (Deuxiéme partie).—Folia Biologica Andina, Pars I1— Zoologica 1:39-52. Loffler, H. 1958. Die Klimatypen des holomiktischen Sees und ihre Bedeutung fiir zoogeographische Fragen.—Sitzungsberichte der Osterreichische Akademie von Wissenschaften, Mathematisch- naturwissenschaftliche Klasse, Abteilung !, 167: 1-33. . 1963. Zur Ostrakoden- und Copepodenfauna Ekuadors.— Archiv fiir Hydrobiologie 59:196— 234. 1981. Copepoda. Jn S. H. Hurlbert, G. Ro- driguez, and N. D. dos Santos, eds., Aquatic Biota of Tropical South America, Part 1. Ar- thropoda, pp. 14-19. San Diego State Uni- versity, San Diego, California. Lowndes, A. G. 1934. Copepoda from Brazil and Paraguay. — Journal of the Linnean Society (Zo- ology) 39:83-131. Marsh, C. D. 1919. Report on a collection of Co- pepoda made in Honduras by F. J. Dyer. — Pro- ceedings of the United States National Museum 55:545-548 + 1 plate. Osorio Tafall, B. F. 1941. Materiales para el estudio del microplancton del lago de Patzcuaro (Mex- ico). —Anales de la Escuela Nacional de Ciéncias Bioldgicas, Instituto Politécnico Nacional 2:33 1— 383. Smith, K. E., and C. H. Fernando. 1978. The fresh- water calanoid and cyclopoid copepod Crusta- cea of Cuba.— Canadian Journal of Zoology 56: 2015-2023. . 1980. Guia para los copépodos (Calanoida y Cyclopoida) de las aguas dulces de Cuba.—Aca- demia de Ciéncias de Cuba, Habana. 28 pp. NHB-163, Department of Invertebrate Zoology, National Museum of Natural His- tory, Smithsonian Institution, Washington, DNEL20560: PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 79-83 A NEW GENUS AND SPECIES OF INTERSTITIAL SIGALIONIDAE AND A REPORT ON THE PRESENCE OF VENOM GLANDS IN SOME SCALE-WORM FAMILIES (ANNELIDA: POLYCHAETA) Paul S. Wolf Abstract. —Metaxypsamma uebelackerae, a new interstitial polychaete genus and species of the family Sigalionidae, is described from the northern Gulf of Mexico. Metaxypsamma uebelackerae differs from other known sigalionids in lacking notopodia and elytra, but these represent neotenic features coincident with an adaptation to an interstitial habitat. The similarities in the structure of the piercing-type jaws and associated venom glands of the Sigalionidae, Polynoidae, Polyodontidae, Pholoididae, and Pisionidae are discussed, and these are compared with two other scale-worm families, Eulepethidae and Aphroditidae, both of which lack piercing-type jaws and venom glands. During 1975-1981, the U.S. Bureau of Land Management (now Minerals Manage- ment Service) funded several projects de- signed to characterize the fauna of the outer continental shelf along the northern Gulf of Mexico. In 1979, Barry A. Vittor & Asso- ciates, Inc., Mobile, Alabama, was funded by MMS to produce a taxonomic guide de- signed to standardize the identifications of polychaetes collected during these BLM projects (Uebelacker and Johnson 1984). Metaxypsamma uebelackerae, n. gen., n. sp., was discovered among the material exam- ined during the preparation of a chapter to be included in this publication. At that time, M. uebelackerae was considered a potential new family (Wolf 1984). The species is de- scribed herein as a neotenic, interstitial member of the family Sigalionidae. While examining M. uebelackerae, an in- ternal canal was found in each jaw. This raised the question of why an internal canal was present except perhaps to channel a fluid in or out through the large fang. Larger si- galionids were then examined, and a large gland (colored white in preserved material) was found associated with each jaw. This is probably a venom gland. This has led to the inclusion here of a report of the presence of venom glands in families heretofore not known to have such glands. Metaxypsamma, new genus Family B, Genus A, Wolf 1984:60-1. Type-species. —Metaxypsamma uebe- lackerae, new species. Gender. — Feminine. Diagnosis. —Body slender, with up to 24 segments. Prostomium with single median antenna arising frontally, lateral antennae lacking. Two pairs eyes present. Peristo- mium or tentacular segment achaetous, with tentaculophores directed anteriorly, each bearing dorsal and ventral tentacular cirrus. Pair of smooth digitiform palps emerging ventral to tentaculophores. Elytra lacking but low, fleshy mounds, each bearing 2—4 long, slender, knobbed papillae, present on segments 2, 4, 5, 7, continuing on alternate segments. Parapodia uniramous, with single internal aciculum. Setae as compound fal- cigers with short, serrate, unidentate blades. Proboscis eversible, muscular, with 2 pairs 80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of chitinous, piercing-type jaws and distal papillae; venom glands present, adhering to ventrolateral plates of each jaw. Etymology.—The generic name is de- rived from the Greek metaxy, between, and psammos, sand, referring to the interstitial habitat of the type-species. Remarks. —Metaxypsamma_ uebelacker- ae at first appears to differ considerably from other members of Sigalionidae since it lacks notopodia and, instead of elytra, it has paired mounds of papillae. Cazaux (1968:53 1-534) describes the larval nectochaete I and II stages of Pholoe synophthalmica Claparéde, 1868. He shows the nectochaete I stage as lacking notopodia and, instead of elytra, there are paired mounds of papillae. The nectochaete II develops notopodia but still retains the mounds of papillae, which are very similar to those of Metaxypsamma. It 1S apparent then, that Metaxypsamma ue- belackerae is a neotenic sigalionid. Among adult sigalionids, Metaxypsam- ma is most similar to Pholoe in lacking lat- eral antennae, in having an achaetous ten- tacular segment, in being of rather small size with few segments, in lacking branchiae, in having compound falcigers with short non- articulate blades, and in lacking compound spinigers. Metaxypsamma uebelackerae, new species Fig. 1 Family B, Genus A, Wolf 1984:60-3, figs. 60-1, 60-2a-e. Material examined. —FLORIDA: SO- FLA Sta 5A, 26°45.70'N, 84°00.13'W, coarse sand, 91 m, Aug 1981, 3 paratypes, including 1 female (USNM 86845).—SO- FLA Sta 5G, same location and date, 3 para- types (USNM 86844).—MAFLA Sta 2426G, 28°57'59.4"N, 85°23'00.2”W, fine sand, 82 m, Feb 1978, 1 specimen, | slide (USNM 89582).—Sta 2748, 27°37.2'N, 83°53.5’W, coarse sand, 50 m, Jul 1978, Holotype, female (USNM 86846).—Sta 2958J, 25°40'N, 83°50’W, medium fine sand, 120 m, Feb 1977, 1 specimen.—Sta 29581, same location, Aug 1977, 1 fe- male.—Sta 2959H, 25°40'N, 83°05'W, silty- very fine sand, 60 m, Aug 1977, paratype, ripe male (USNM 97801). Description. —Length to 2.5 mm, width to 0.5 mm, including parapodia, to 0.24 mm, excluding parapodia. Complete specimens with 21-24 segments. Segmentation distinct although intersegmental furrow present only on every other segment beginning at pos- terior margin of segment 9 and continuing to anterior margin of pygidium (Fig. la, c). Prostomium bilobed with median anten- na arising terminally between lobes (Fig. 1a); 2 pairs of eyes arranged trapezoidally; palps smooth, digitiform, emerging beneath pro- stomium; tentaculophores directed ante- riorly, without setae or acicula, with dorsal and ventral tentacular cirrus, similar to me- dian antenna in size and shape. Facial tu- bercle lacking. Dorsum without elytra but with paired fleshy mounds bearing long, filiform pa- pillae located above parapodia of segments 2, 4, 5, 7 and alternate segments thereafter (Fig. la, b). Anterior 5—6 mounds each with 2 papillae, following ones with 3—4 papillae. Beginning on segment 2, ventrum with 2 pairs of globular papillae per segment on either side of midline just posterior to para- podia; beginning on segment 9, alternate segments with additional midventral pa- pilla (Fig. 1c). Ventral (buccal) cirri of segment 2 twice as long as following ventral cirri. Parapodia uniramous, highly contractile, each sup- ported by single, pointed aciculum; para- podial stylodes and papillae lacking (Fig. 1a, b). Setae compound, unidentate falcigers only, about 6—7 per parapodium, 2 above aciculum in single row, 4—5 below aciculum in 2 rows. Falciger blades with minute teeth along concave margin; blades of anterior segments somewhat longer than those of fol- lowing segments and with shorter marginal teeth (Fig. 1d, e). Pygidium with 2 pairs of filiform anal cir- VOLUME 99, NUMBER 1 mAn \l \ ifs vtC IN Ss = pa \ é 3 VS: lp C= pap We ie =—— = Nie S— Sz. =~ + (aes aA we RS AG WO): “ — A Fig. 1. 0.05mm 81 Metaxypsamma uebelackerae: a, Anterior end, dorsal view; b, Parapodium from segment 9, posterior view; c, Segments 14-17, ventral view; d, Neuroseta from segment 2; e, Neuroseta from segment 9; f, Posterior end, dorsal view; g, Distal end of dissected proboscis, lateral view; h, Dorsal left jaw, inside lateral view. anC, anal cirrus; dtC, dorsal tentacular cirrus; mAn, median antenna; pa, palp; pap, papilla; vC, ventral tentacular cirrus (Figs. a, b, d—-f from Wolf, 1984:Figure 60-2a-e). ri located lateral to terminal anus; cirri about half as long as dorsal papillae (Fig. 1f). Proboscis eversible, muscular, with 2 pairs of chitinous, piercing-type jaws and distal circle of 9 pairs of papillae (Fig. 1g). Each jaw with a broad plate located ventrolateral to long stem, with internal canal extending to tip of large tooth (Fig. 1h). Venom gland not observed. Remarks.—Three mature females, in- cluding the holotype, were collected. The diameter of the largest egg was about 128 pum. About 30 eggs were present in the holo- type from segment 9 to the pygidium (Fig. 1c). One ripe male contained large amounts of sperm from setiger 10 to the end of the body. A peculiar phenomenon was noted for //. uebelackerae, namely the presence of inter- segmental furrows on alternate segments beginning on the posterior margin of seg- ment 9 (Fig. la, c). Coincident with this 82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON alternate arrangement of the furrows is the arrangement of the midventral papilla and the paired mounds of papillae dorsally. I then examined several undescribed species of Pholoe and found the intersegmental fur- rows on alternate segments, also, except that beginning on segment 23 the furrows are found on every segment. Segment 23 is also where elytra appear on every segment. I also examined larger sigalionids (Sthenelais sp. and Psammolyce ctenidophora) and found the intersegmental furrows on every seg- ment even in the region where elytra are present on alternate segments. This arrange- ment was also observed in Pholoides ber- mudensis (Pholoididae), two species of Pi- sione (Pisionidae), and several species of Polynoidae and Polyodontidae. In Pho- loides bermudensis, however, the furrows are faint along the entire body. The significance of the segmental arrange- ment of the furrows is not possible to assess here, since histological sectioning would be necessary to determine whether or not what appears externally to be reduced segmen- tation is actually the case internally. Such resources were not available to the author. The morphological adaptations of M. uebelackerae afford an interesting compar- ison with Pholoe swedmarki Laubier (1975: 671-678), an interstitial sigalionid from Bermuda. Pholoe swedmarki has notopo- dia, but they are reduced with few notose- tae, whereas M. uebelackerae has lost its notopodia completely. Pholoe swedmarki has retained its elytra; M/. uebelackerae has replaced them with paired mounds each bearing long, flexible papillae. Both species have somewhat reduced tentacular cirri. In P. swedmarki the dorsal tentacular cirri are longer than the median antenna and the ventral cirri are much smaller and of a dif- ferent shape than the dorsal ones. In M. uebelackerae, the dorsal and ventral tentac- ular cirri are similar to each other and to the median antenna in size and shape. Pho- loe swedmarki has up to 27 segments; M. uebelackerae has up to 24. Pholoe swed- marki measures up to 1.6 mm by 0.4 mm while M. uebelackerae measures up to 2.5 mm by 0.5 mm. Both species are considered to be inter- stitial worms, but the neotenic M. uebe- lackerae exhibits the more derived condi- tions, i.e., the loss of elytra, loss of notopodia, and reduction in the number of segments, even though it is a slightly larger species. It is body width, however, that de- termines the minimum diameter of the in- terstitial space an animal may inhabit (Westheide 1984:265). It is suggested here that M. uebelackerae has, through the loss of elytra, gained a functionally narrower body width than P. swedmarki. This could enable the former to invade smaller inter- stitial spaces than the latter. Etymology.—The species is named in honor of Ms. Joan Uebelacker who, as se- nior editor of ““Taxonomic Guide to the Polychaetes of the Northern Gulf of Mex- ico” (Uebelacker and Johnson 1984), edited every chapter and, therefore, is largely re- sponsible for the success of those volumes. Her editorial and scientific efforts are grate- fully acknowledged and deeply appreciated. Distribution. —Gulf of Mexico, off Flori- da, 50-120 meters. Observations on Scale-worm Jaws and Venom Glands The jaws of three other sigalionids, Sthe- nelais sp., Psammolyce ctenidophora, Pho- loe sp., and Ehlersileanira incisa were also examined. In all these specimens, the basic structure and appearance of the jaws were the same including the presence of an in- ternal canal and a large, white, spongy, glan- dular mass of tissue adhering to the dorsal side of the large, ventrolateral plate of each jaw. This glandular mass leads into the in- ternal canal within the fang of the jaw. This white mass is here interpreted to be a venom gland. A venom gland could not be found in Metaxypsamma uebelackerae, perhaps because of its minute size, but its presence is suggested by what appears to be a small VOLUME 99, NUMBER 1 duct and a volume of cellular material with- in the internal canal of each jaw. The jaws of other scale-worm families were also examined: Polyodontidae (Poly- odontes lupinus and Eupanthalis tubifex), Polynoidae (Lepidasthenia sp.), Pholoidi- dae (Pholoides bermudensis), Eulepethidae (Grubeulepis sp.), and Aphroditidae (Aphrodita sp.). The polynoid and pholoidid jaws are very similar in shape to that of the sigalionids and have venom glands associ- ated with their jaws. The polyodontid jaw differs in that its ventrolateral plate is fused to the concave margin of the main fang, and this plate is dentate along its outer edge. These jaws have a venom gland adhering to the dorsal side of the plate, but it appears smaller and less developed relative to the large size of the animal and its jaws, as compared to the large gland found in the Sigalionidae, Polynoi- dae, and Pholoididae. The Eulepethidae and Aphroditidae do not possess piercing-type jaws. Their jaws consist of broad, chitinous plates which do not have any associated venom glands. The jaws of the Pisionidae (Pisione sp.) were also examined and found to be re- markably similar to the sigalionid jaw. A venom gland, located as in the sigalionids, leads into an internal canal that extends to the tip of the fang. If the above described glands do indeed produce venom, the question remains as to how the venom is released. Glycerids, ac- cording to Gibbs and Bryan (1980:205), re- lease venom through a series of lateral pores located on the main fang of the jaw. No such pores were found associated with the jaws of the scale-worms examined. The internal canal does extend to the very tip of the main fang, but there did not appear to be an open pore at the tip. Acknowledgments The author wishes to thank the following colleagues for their critical reviews of the 83 manuscript: Mr. R. Michael Ewing of Old Dominion University; Dr. Marian H. Pet- tibone of the Museum of Natural History, Smithsonian Institution, Washington, D.C.; Dr. Mary E. Petersen of the Zoological Mu- seum, Copenhagen; Dr. Barry A. Vittor and Mr. Julian Stewart of Barry A. Vittor & As- sociates, Inc. Mr. Ben Jordan, also with Bar- ry A. Vittor & Associates, prepared many of the illustrations for publication and helped the author in his attempts to do the same for the remaining figures. Dr. Kristian Fau- chald, Smithsonian Institution, is thanked for the loan of material. Literature Cited Cazaux, C. 1968. Etude morphologique du dével- oppement larvaire d’Annélides polychétes. (Bassin d’Arcachon).—Archives de Zoologie Expérimentale et Générale 109(3):477-543. Claparéde, E. 1868. Les Annélides Chétopods du Golfe de Naples.— Mémoires de la Société de Physique et d’Histoire Naturelle de Genéve 19(2):313-584. Gibbs, P. E., and G. W. Bryan. 1980. Copper—the major metal component of glycerid polychaete jaws.—Journal of the Marine Biological Asso- ciation of the United Kingdom 60:205-214. Laubier, L. 1975. Adaptations morphologiques et biologiques chez un aphroditien interstitiel: Pholoe swedmarki sp. n.—Cahiers de Biologie Marine 16:671-683. Uebelacker, J. M., and P. G. Johnson [Eds.]. 1984. Taxonomic guide to the polychaetes of the northern Gulf of Mexico. Final Report to the Minerals Management Service, contract 14-12- 001-29091. Barry A. Vittor & Associates, Inc., Mobile, Alabama. 7 vols. Westheide, W. 1984. The concept of reproduction in polychaetes with small body size: adaptations in interstitial species.—Fortschritte der Zoolo- gie 29:265-287. Wolf, P.S. 1984. Family B. Jn Uebelacker and John- son, 1984 (see above). Taxonomic guide to the polychaetes of the northern Gulf of Mexico. Barry A. Vittor & Associates, Inc., Volume VII. Chapter 60. 3 pp. Barry A. Vittor & Associates, Inc., 8100 Cottage Hill Rd., Mobile, Alabama 36609. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 84-87 GLOSSOTHELEPUS, A NEW GENUS OF THELEPINAE (POLYCHAETA: TEREBELLIDAE) FROM THE GULF OF CALIFORNIA, MEXICO P. A. Hutchings and C. J. Glasby Abstract. —A new genus of Thelepinae, Glossothelepus, is described from deep water off Mexico, with the type species G. mexicanus, n. sp. The genus is closely related to Decathelepus Hutchings and Rhinothelepus Hutchings from Austra- lia, and Telothelepus Day from South Africa. While examining material in the collec- tions of the Allan Hancock Foundation, an undescribed genus was found. Dr. Kristian Fauchald had already separated this mate- rial, recognizing it as a new genus. We are thankful to Dr. Fauchald for allowing us to describe the new genus, which extends the distribution of those genera of Thelepinae with expanded tentacular lobes. Glossothelepus, new genus Diagnosis. —Expanded tentacular lobe with numerous buccal tentacles; dorsal buc- cal lobe T-shaped, and thickened. Numer- ous sessile simple branchial filaments on segments 2 and 3. Notopodia from segment 3, continuing for at least 23 segments. No- tosetae smooth-tipped, narrow-winged, broad-bladed capillaries. Neurosetae from setiger 7, arranged in single rows, avicular. Type species. —Glossothelepus mexican- Us. Etymology. —The generic name is a com- bination of the Greek ‘glossos,’ a tongue, and ‘thelepus’ and refers to the expanded glandular tentacular lobe. Glossothelepus mexicanus, new species Eig. 1 Holotype.—AHF Poly 1449, posteriorly incomplete, 10 mm length, 1.2 mm maxi- mum width, 23 setigers, Sta P51-59 (N15136-F3205): 25°31'5”N, 109°31'5’W; 9 fms. Paratypes: 1 spec. USNM 98572 pos- teriorly incomplete, 15 mm length, 2.5 mm maximum width, 20 setigers, Sta 496-36 (F3115): 23°21'55”N, 109°24'40”W: 80 fms, mud; | spec. AM W 199659 posteriorly in- complete, 20 mm length, 2 mm maximum width, 20 setigers, Sta 1732-49 (F3101): 23°24'45”N, 109°23'50”W to 23°24’30°N, 109°24'00”W; 50 fms; sand and mud. All material from Fraile Bay, Gulf of Califor- nia, Mexico. Description.—Body pale yellow, stout, with long golden setae. Prostomium with expanded tentacular lobe with convoluted margins, dorsally T-shaped, thickened, al- most papillate, from which some tentacles arising (Fig. la); ventrally forming elongate upper lip (Fig. 1b). Large numbers of grooved buccal tentacles of 2 types; few, thick, slightly bulbous ones resembling those found in Polycirrinae, and more numerous thin ones. Eye spots absent. Ventrum of peristomium thickened, slightly ridged, with maximum width in an- terior-posterior axis, forming discrete glan- dular pad. Segment 2 wedge-shaped later- ally, connected ventrally by very narrow strip, almost completely hidden by peri- stomium and thickened ridge of segment 3. Anterior lateral margins of segments 2 and 3 slightly thickened, glandular, but not forming discrete lateral folds. Ventrum of setigerous segments 1-8 slightly more glan- dular than subsequent segments, also slight- VOLUME 99, NUMBER 1 Fig. 1. 85 Glossothelepus mexicanus: a, Holotype, head end, dorsolateral view; b, Head end, ventral view; c, d, Paratype (AM) long and short notosetae respectively; e, f, Paratype (AM), uncini from setiger 8 and setiger 19, respectively. Scales in mm. ly granular in appearance. Thin medial ven- tral glandular stripe occurring along rest of body; stripe segmented, each segment hav- ing an additional intrasegmental fissure. Segmentation becoming more distinct pos- teriorly. Dorsum appearing slightly reticu- late, especially anteriorly. Branchiae consisting of numerous sessile filaments on segments 2 and 3. Each bran- chia consisting of laterally arranged group of sessile filaments with distinct medial gap. Segment 2 with approximately 20 filaments on each side arranged on arc-shaped thick- ened ridge, more laterally arranged than those on segment 3. Segment 3 with about 15 filaments on each side. Notosetae from segment 3, continuing for at least 23 segments. First pair of notosetae inserted at angle, in contrast to others. No- topodia elongate, flattened, rectangular po- 86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dia with setae arranged longitudinally. No- tosetae golden, long, arranged in 2 tiers, short broad-bladed, narrow-winged capillaries and longer wingless capillaries, both with fine tips (Fig. lc, d). Tips of setae under high magnification appearing slightly fuzzy. Neurosetae from setiger 7, uncini ar- ranged in single rows on long tori, on slightly raised glandular strip extending across body to medial ventral groove; uncini only oc- curring along part of ridge. Posteriorly un- cinial tori becoming slightly elevated. Un- cini minute, avicular with strongly crested head; dental formula MF: 6—10:00, with teeth arranged in 2 arcs above main fang; hooked dorsal button and rounded prow consistent throughout (Fig. le, f). Variation.—The two paratypes exhibit slight variations from the holotype. The paratype lodged in the Australian Museum is in poor condition but is lightly pigmented like the holotype, whereas the USNM para- type is very darkly pigmented, with setigers 1-3 having even darker pigmentation across the ventrum. This paratype exhibits marked lateral expansion from setigers 5—7, the body then resumes original width, which is far more marked than in the holotype. Discussion. — Within the subfamily The- lepinae to which this species clearly belongs, because of its sessile, simple branchial fil- aments, avicular uncini arranged in single rows throughout and the characteristically Thelepinae like uncini, the setiger on which the noto- and neurosetae begin is considered to be an important generic character (Hutchings and Glasby in press). No exist- ing described genus in the subfamily has notosetae beginning on segment 3 and neu- rosetae on setiger 7, and for this reason, this species is described in the new genus G/os- sothelepus. Within the Thelepinae, at least two distinct groups can be recognised, one group of genera with a compact prostomi- um, the other group with an expanded ten- tacular lobe. Glossothelepus clearly belongs to this latter group, which also includes Te- lothelepus Day, 1955, Rhinothelepus Hutchings, 1974, and Decathelepus Hutch- ings, 1977. Currently, Glossothelepus, Te- lothelepus, and Decathelepus are mono- specific genera, and Rhinothelepus is represented by two species. All these genera except Glossothelepus are currently only known from shallow intertidal areas. With the discovery of Glossothelepus from deep water off Mexico, the geographical distri- bution of these apparently closely related genera is expanded from South Africa (Telo- thelepus) and Australia (Rhinothelepus and Decathelepus) to Mexico. One may specu- late that the development of an expanded tentacular lobe is an advanced character in the Thelepinae and these four genera are closely related to each other, and that ad- ditional deep-water related genera may oc- cur in the South Pacific and Indian Ocean. An alternative hypothesis is that these gen- era with an expanded tentacular lobe have risen independently several times from the parent stock of Thelepinae. This seems un- likely, as Glossothelepus very closely resem- bles Rhinothelepus and Decathelepus in all major body characteristics except for the setiger on which the uncini begin. One may further speculate on the evolutionary sig- nificance of uncini beginning on setiger 6 in Rhinothelepus, setiger 10 in Decathelepus and on the first abdominal segment in Te- lothelepus, and on setiger 7 in Glossothele- pus. It would be very interesting to observe the feeding behavior in these genera, since presumably the uncini are used to retain the animal in its burrow. Etymology.—The name given by Dr. Fauchald has been retained and refers to the geographical location where the animal was collected. Habitat. —Deep water, in sand and mud- dy substrata. Distribution. —Gulf of California, Mexi- co. Acknowledgments We thank Dr. Kristian Fauchald for al- lowing us to describe the new genus and VOLUME 99, NUMBER 1 species which he had initially recognised, and Sue Williams for loaning us the mate- rial. The Australian Biological Resources study provided the salary of one of us (CJG). Literature Cited Day, J. H. 1955. The Polychaeta of South Africa. Part 3: Sedentary species from Cape shores and estuaries. — Journal of the Linnean Society (Zo- ology) 42:407—452. Hutchings, P. A. 1974. Polychaeta of Wallis Lake, New South Wales. — Proceedings of the Linnean Society of New South Wales, 98:175-195. 87 1977. Terebelliform Polychaeta of the fam- ilies Ampharetidae, Terebellidae and Tricho- branchidae from Australia, chiefly from More- ton Bay, Queensland.— Records of the Australian Museum 31:1-38. , and C. J. Glasby. [in press]. The Thelepinae (F. Terebellidae) from Australia together with a discussion of the generic and specific characters of the family.— Proceedings of The Biological Society of Washington. The Australian Museum, 6-8 College Street, P.O. Box A285, Sydney South, N.S.W. 2000, Australia. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 88-99 NEW RECORDS OF MIMILAMBRUS WILEYI WILLIAMS, 1979 (CRUSTACEA: DECAPODA: BRACHYURA), WITH NOTES ON THE SYSTEMATICS OF THE MIMILAMBRIDAE WILLIAMS, 1979, AND PARTHENOPIDAE MACLEAY, 1838, SENSU GUINOT, 1978 Peter K. L. Ng and Gilberto Rodriguez Abstract.—The family Mimilambridae was established by Williams (1979) to receive his new species Mimilambrus wileyi. The species is herein reported from several localities in Venezuela. The characters of Mimilambrus fit well within the definition of the subfamily Parthenopinae, family Parthenopidae MacLeay, 1838 (sensu Balss 1957) (equivalent to Guinot’s (1978) Partheno- poidea). There is agreement in many morphological details, such as in the ventral spines, fingers and dentition of the chelipeds, and the form of the second male pleopod. Some characters like the ischio-meral articulation of the che- lipeds and the articulation of segments 3 to 5 of the male abdomen, show a primitive condition. A comparison of the structures associated with respiration shows that the peculiar arrangement of afferent and efferent channels in Mim- ilambrus is merely a continuation of a trend already found in the more typical parthenopids. Placing this genus in a family of its own is thus unwarranted, and it should instead be included as a rather specialized genus with the rest of the parthenopids in the superfamily Parthenopoidea MacLeay, 1838, sensu Guinot, 1978. Mimilambrus wileyi Williams, 1979, was described from two males and one female collected at Tobago, West Indies, and was placed by Williams in a new monotypic family, Mimilambridae, close to the family Parthenopidae MacLeay, 1838, sensu Balss, 1957. Guinot (1977a, 1978a, b, 1979), in her revision of Brachyuran classification, el- evated the Parthenopidae to superfamily status. Bowman and Abele (1982) subse- quently elevated the Mimilambridae to its own superfamily, Mimilambroidea, prob- ably in accordance with the superfamilial method of classification proposed by Gui- not (1977a, b, 1978a). Williams (1979) suggested that his new family was intermediate between the tra- ditional “oxyrhynchan’”’ families Mayidae and Parthenopidae (sensu Balss 1957), and the traditional ‘“‘oxystomatan”’ families Leucosiidae and Calappidae. The current evidence, however, suggests that both the Oxystomata and Oxyrhyncha, are in fact heterogeneous groups, and not ‘good’ taxa (Guinot 1977a, 1978a, b, 1979; Stevci¢ and Gore 1980). Within this context, a reex- amination of the family Mimilambridae is of particular interest. During the study of samples taken on the Venezuelan coast, several specimens of Mimilambrus wileyi were collected. Oppor- tunity was taken to evaluate the characters used by Williams (1979) to differentiate his new family from the Parthenopidae sensu Balss, 1957. To this end, we have examined four American parthenopoids: Parthenope agona (Stimpson, 1871), Platylambrus ser- ratus (H. Milne Edwards, 1834), Leiolam- VOLUME 99, NUMBER 1 brus nitidus (Rathbun, 1901) and Hetero- crypta tomasii da Costa, 1959; and five Indo- Pacific ones: Daira perlata (Herbst, 1790), Daldorfia horrida (Linnaeus, 1758), Cryp- topodia fornicata (Fabricius, 1781), Rhin- olambrus longispinis (Miers, 1879), and Platylambrus echinatus (Herbst, 1790). We have also used morphological data available in the literature whenever appropriate. The systematics of the Parthenopoidea sensu Guinot, 1978a, b, are also briefly discussed in relation to several other characters. The material cited is deposited in the reference collection of the Instituto Venezolano de Investigaciones Cientificas (IVIC), Caracas, and the Zoological Reference Collection (ZRC) of the Department of Zoology, Na- tional University of Singapore. Mimilambrus wileyi Williams, 1979 Mimilambrus wileyi Williams, 1979:400, Figs. 1-5. Parthenope sp., Rodriguez, 1980:294, Fig. 70, Pl. 31A. Material. —Punta Arenas de Pariche, Golfo de Cariaco, 10°32'N, 64°12’W, 12 Jul 1959; Luis R. Salazar: 1 male, cl. 26.3 mm, cb. 28.6 mm, exploratory fishing with shore net, shallow water (IVIC).— Between Mar- garita Island and the Venezuelan mainland, 8 km west of Isla Coche, 10°46'N, 62°50’W: 1 male, cl. 14.8 mm, cb. 16.2 mm, 9 fath- oms (IVIC).—Paraguana Peninsula, north of Punto Fijo, 11°40’N, 70°10’W, 4 Mar 1982, G. Kremer & M. Bevilacqua: 1 male, cl. 29.7 mm, cb. 33.7 mm, 1 ovigerous, | spent female, cl. 25.6 and 30.5 mm, cb. 28.1 and 32.0 mm, respectively, between 10 and 17 fathoms (IVIC). (cb. measured at base of postero-lateral spines in all cases.) Discussion Our material agrees closely with Wil- liams’ (1979) original description. The eggs of our ovigerous female have a mean di- ameter of 0.32 mm (n= 15, largest 0.34, 89 smallest 0.31 mm). The present records ex- tend the range of the species to the north coast of South America and show that it is not an uncommon species. The superfamily Parthenopoidea (essen- tially the Parthenopidae of Balss (1957) with the exclusion of the Eumedoninae Miers, 1879) has been subjected to considerable scrutiny recently. Their origins are poorly understood, and they appeared rather sud- denly in the Middle Eocene (Glaessner 1969), some 45 million years ago. Most classifications have included them in the Oxyrhyncha, for they, like the majids and hymenosomatids, “have a general elongate- triangular, narrow fronted carapace and in- complete orbits which contrast strongly with the rest of the brachygnathous crabs” (Rice 1980:311). Acceptance of this systematic arrangement, however, is not unanimous. Flipse (1930) for instance, thought that the parthenopids were transitional between the Oxyrhyncha and the Cyclometopa. Guinot (1977a, b, 1978a, b), in her comprehensive review of Brachyuran systematics, disman- tled the Oxyrhyncha, and elevated the Par- thenopidae MacLeay, 1838, to their own superfamily, with the exclusion of the subfamily Eumendoninae Miers, 1879. Otherwise, they were essentially the Par- thenopinae of Alcock (1895), Rathbun (1925), and Balss (1957). She recognized four groups within the Parthenopoidea: 1, With only the thoracic sternal sutures 4/5 and 5/6 interrupted and having the III B type of endophragmal skeleton (e.g., Daldorfia Rathbun, 1904). 2, With all the thoracic sternal sutures interrupted and having the III C type of endophragmal skeleton (e.g., Parthenope Weber, 1795). 3, Containing the genera Daira de Haan, 1833, and Dairoides Stebbing, 1920 (=Asterolambrus Sakai, 1938). 4, Containing the ‘Parthénoxysto- mien’ group (Guinot 1967a, 1978b), with the genera Aethra Leach, 1816, Hepatus La- treille, 1802, Hepatella Smith, 1869, Osa- chila Stimpson, 1871, and Actaemorpha Miers, 1878 (sensu Guinot 1967a, 1978b). 90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A new genus, Sakaila Manning & Holthuis, 1981, was subsequently included in Gui- not’s ‘‘Parthénoxystomien”’ group. Al- though she did not designate these groups as separate families, it may prove worth- while to recognize each of these four groups as such, i.e., the families Daldorfidae, Par- thenopidae s. s., Dairidae, and Aethridae respectively. All further references to these families in this paper will be used as defined above. Guinot (1967b, 1977a, 1979) has also provided indications that the partheno- poids may have evolved from some xan- thoid-like stock. In fact, the genera Daira and Dairoides are almost perfect interme- diates for linking the xanthoids with the Daldorfidae and Parthenopidae s. s. The lar- val characters also suggest that the parthen- opoids evolved from some xanthoid stock via the more primitive portunoid families (sensu Guinot 1978a, b) (Rice 1980, 1983) or that both the parthenopoids and portu- noids evolved from the same xanthoid stock (Ng 1983). The subfamily Eumedoninae Miers, 1879, formerly placed in the family Parthenopi- dae sensu Balss, 1957, is almost certainly not closely affiliated with the true parthen- opoids, but is probably closer to the Xan- thoidea (Seréne, Tran, and Nguyen 1958), and may be related to (Seréne 1968) or part of the Pilumnidae Samuelle, 1819, sensu Guinot, 1978 (Ng 1983). Recent work on the larvae of the eumedonine Echinoechus pentagonus (A. Milne Edwards, 1878) lends support to Ng’s (1983) suggestion that the Eumedoninae be removed from the Par- thenopoidea and affiliated with the Pilum- nidae instead. The family Parthenopidae s. s. forms a sharply defined and presumably monophy- letic group, well characterized by the shape of the carapace and chelipeds. Mimilambrus wileyi shares with this family the following characters: heterotrematous gonopore ar- rangement (Guinot 1977a, b, 1978b, 1979; Saint Laurent 1980), carapace with frontal region narrow (the general outline is semi- circular in Mimilambrus and several other genera, but more usually triangularly elon- gate or pentagonal), orbits distinct and com- plete, cardiac and gastric regions so deeply marked off as to make the dorsal surface of the carapace trilobed, pseudorostrum (sen- su Stevcic and Gore 1980) simple (obscure- ly trilobed in Mimilambrus and some other species), second article of antennae small, short, not fused with the epistome or front, palp of the third maxilliped articulated at the antero-internal angle of the merus, che- lipeds vastly longer and more massive than the ambulatory legs, and the second male pleopods relatively short. Several characters that have been regard- ed as characteristic of the Mimilambridae are also not as definitive as they may ini- tially appear. Williams (1979) gives diag- nostic generic value to the articulation of segments 3 to 5 of the male abdomen. These segments do show a rather primitive con- dition in Mimilambrus since the articula- tion, although ankylosed, is still present as a thin line. In all parthenopoids, segments 3 to 5 are ankylosed (Flipse 1930; Ng 1983), but there is some variability as to whether the sutures are still distinct or not. In Par- thenope agona, the sutures are still visible near the midline of the abdomen, while in Platylambrus serratus, they are completely absent. In the supposedly primitive Daira and Dairoides, the sutures are still relatively distinct. Whether the segments 3 to 5 are ankylosed or free is itself an important clue to the cohesiveness of the taxa, and also provides valuable information about their affinities. The ankylosed segments in all the members of the Parthenopoidea as recog- nized by Guinot (1978a, b) (including Mim- ilambrus) not only lends support to the ex- clusion of the Eumedoninae, in which all the segments are free, but also suggests that the parthenopoids have close phylogenetic links with the Portunoidea (sensu Guinot 1978a, b), as well as the more primitive xanthoids, like the Carpiliidae Ortmann, VOLUME 99, NUMBER 1 1893. This relationship has in fact, already been suggested through the studies of their larvae (Rice 1980, 1983; Ng 1983). Wil- liams (1979) also regards the first male pleo- pod as characteristic, but although it is cer- tainly unusual, it could have been easily derived from ancestral parthenopid struc- tures. In fact, Cryptopodia concava Stimp- son, 1871 (Gore and Scotto 1979) and Pseu- dolambrus triangula (Stimpson, 1860) (Garth 1958) have very similar appendages, but their long subterminal spines are less well developed and the lateral hairs are still abundant. The most important character that dif- ferentiates Mimilambrus wileyi from most Parthenopidae is probably the position and structure of the afferent channels which are located inside the buccal cavity, and the ef- ferent channels, which are fused and dis- charge at the anteromedial part of the en- dostome. As a consequence of the reorganization of these respiratory chan- nels, the exognath of the third maxilliped is completely concealed behind the endog- nath, and the anterior part of the carapace is narrow and spoutlike, implanted at a sharper angle in relation to the upper surface of the carapace. In this respect it resembles the frontal areas of some Leucosiidae which have similarly structured respiratory chan- nels. The structure of the mouthparts is clearly an adaptation to a fossorial or semi- fossorial mode of life. Burrowing activities differ greatly in var- ious groups of crabs. The majids usually do not burrow into the substrate, whereas many catometopans tend to form permanent bur- rows in the sand. In contrast, many species of the traditional oxystomatan families Leucosiidae and Calappidae and many cy- clometopans which inhabit sandy or grav- elly substrata have burrowing habits. Their burrows, however, are never permanent channels in the sand but merely temporary excavations. Each of these different habits is associated with different morphological traits. In parthenopids, the crab is actually 91 embedded in the sand (Schafer 1950). The chelipeds are flexed to the underside of the anterolateral regions, with the fingers, which are flat and bend downwards and inwards, fitting closely at each side of the buccal cav- ity. The internal surfaces of the chelipeds are channeled and glossy. There is thus pro- duced on each side of the crab, between the chelipeds and carapace, a ventral channel which communicates with the afferent ap- erture located at the base of the chelipeds. Since the back of the crab is covered with sand, the teeth on the internal margins of the merus of the cheliped, and those of the anterior part of the carapace, act as a coarse sieve placed at the orifice of these accessory channels. In the most generalized condition, only these “‘exostegal channels” (Garstang 1897a, b) exist, and there are no accessory ridges or other structures on the pterygostomial regions to act as afferent channels. This con- dition is found in the genus Cryptopodia H. Milne Edwards, 1834 (A. Milne Edwards 1878, pl. 29, fig. la, 2a). In many parthen- opids, there are usually ridges that delimit the afferent channels. In Platylambrus ser- ratus for instance, there is a sinuous, cren- ulate ridge that runs transversely across the pterygostomial and distal subbranchial re- gions. In its distal part this ridge is provided with long hairs, which together with similar hairs on the anterior border of the coxa, basis and ischiomerus and third maxilliped, form a funnel-like entrance to the afferent aperture at the base of the cheliped. In Het- erocrypta, there is a submarginal ridge that, starting at the suborbital region, runs par- allel to the margin, and forms with it a chan- nel that is lined with hairs. This channel is joined by a second channel that runs per- pendicularly to the first, and ends at the base of the chelipeds. In Heterocrypta tommasii, the channels communicate with the upper surfaces of the carapace through an hepatic notch located at the junction of the two channels. In Solenolambrus Stimpson, 1871 (Rath- 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON bun 1925), Mesorhoea Stimpson, 1871 (A. Milne Edwards 1878, pl. 29, fig. 5a, 6a) and Aulacolambrus Paulson, 1875 (Flipse 1930), there is on each pterygostomial region, a sharp, elevated and crenulate crest that runs from the antero-external angle of the buccal area to the base of the chelipeds, separating the concave pterygostomial region from the subhepatic region, which is also concave and channellike. When retracted, the extremity of the hands of the chelipeds covers the pter- ygostomial regions, forming the afferent passages. In the very rare genus Tutankh- amen Rathbun, 1925, there are ridges sim- ilar to those of Solenolambrus, which ex- tend from the antero-external angle of the buccal area to the base of the chelipeds, but the structure is more complex because the channels are shorter and deeper, bordered by a laminar expansion of the hepatic and anterior branchial margin above, and by a parallel lamina having an emargination near the beginning of the branchial region below (A. Milne Edwards 1878). In Leiolambrus A. Milne Edwards, 1878, the exognath of the third maxilliped is part- ly concealed behind the endognath; the me- rus and ischium of the endognath are bor- dered by long, arched hairs which form with the hairs on the pterygostomial region, a ventral covering to the afferent channels. Mimilambrus has a similar arrangement, but the exognath is completely concealed be- neath the endognath, and the pterygosto- mial ridge is more prominent, particularly in its distal part. The efferent channels show a similar ten- dency towards a rearrangement within the buccal cavity. In Aethra Leach, 1816 (Gui- not 1967a) and some parthenopids, the ef- ferent channels are located on each side of the buccal frame, forming with the margin of the first maxilliped two separate sinuses. In Parthenope agona, Platylambrus serratus, Leiolambrus nitidus, and Heterocrypta tommasii, the anterior endostomial crests are sinuous, forming two adjacent openings that are separated by a medial endostomial septum, but in fact discharging near the midline of the concave epistome. A last step in what Guinot (1966:747) has called “la tendence des Parthenopidae vers le type oxystomien”’ is seen in Mesorhoea, with “the two efferent channels meeting at the middle of the endostome, which has there a trian- gular projection, and a deep notch in its vertical, laminiform wall’? (Rathbun 1925: 546). In Mimilambrus, a similar arrange- ment is found. The efferent respiratory channels empty through a common anterior median opening, into the concave epistome, which also has a ciliated notch. Williams (1979) has noted that the last pair of ambulatory legs of Mimilambrus is similar in shape to that of the calappid Acanthocarpus alexandri Stimpson, 1871, and its structure suggests that Mimilambrus is probably a more active burrower than most of the other parthenopids (with the possible exception of Mesorhoea). Its more fossorial habits could easily explain why its mouthparts have evolved convergently with the leucosiids and calappids, which spend their daylight hours almost completely bur- ied in sand or silt. Stevci¢ and Gore (1980), in summarizing the known characters of the Parthenopidae and Mimilambridae, also noted that where- as the Parthenopidae have antennules fold- ing slightly obliquely, Mimilambrus has them folding almost vertically. The folding of the antennule, however, appears to be a very variable character in the Parthenopi- dae, and appears to be dependent on the width of the pseudorostrum. In Parthenope agona, they are distinctly oblique, whereas in Leiolambrus nitidus, they are almost ver- tical. The presence of small antennae, with the peduncles in the orbital hiatus can also be found in many Parthenopidae. In our comparative studies of the Par- thenopoidea with the Mimilambridae, we also examined several other characters that are common to both. Of the genera dealt VOLUME 99, NUMBER 1 93 Fig. 1. Ventral view of Parthenopidae and Mimilambrus, showing locking mechanism of chelipeds: a, Mim- ilambrus wileyi; b, Platylambrus serratus; c, Platylambrus echinatus; d, Rhinolambrus longispinis; e, Leiolambrus nitidus; f, Cryptopodia fornicata; g, Daldorfia horrida. \1P, Left cheliped; 2P, Second pereopod. with, the only daldorfid is Daldorfia; Daira and Dairoides are dairids, Aethra the only aethrid, all the others being parthenopids. These characters will now be discussed sys- tematically. 1. Ventral spines.—These spines are lo- cated at each side of the sternum, near the bases of the chelipeds. When the chelipeds are moved backwards, the respective spine locks against the ischio-meral articulation and prevents any further displacement backwards of the appendage (Fig. 1). These spines have the same position in Mimilambrus (Fig. la) and Parthenope agona, P. hyponca (Stimpson, 1871), P. or- natus (Flipse, 1930, Fig. 24), Platylambrus 94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Inclination (in degrees) of the fixed finger and dactylus in some parthenopids and in Mimilambrus wileyi. Minimum and maximum values are given in parentheses. Fixed finger Dactylus n Left Right Left Right Platylambrus serratus 4 38 (30, 43) 38.5 (34, 43) 15 (10, 21) 7 s(2 0) Platylambrus echinatus 4 18 (16, 22) 18 (11, 26) 1.5 (0, 5) 3 (0, 6) Rhinolambrus longispinis 1 28 19 8 3 Leiolambrus nitidus 5 18 (17, 20) 20.6 (17, 25) 0 0 Cryptopodia fornicata 5 9.5 (6, 15) 9.5 (8, 13) 3 (2, 4) 1.5 (1, 2) Daldorfia horrida 2 12 (11, 13) NS) (I, 1) 8.5 (8, 9) 6 (4, 8) Mimilambrus wileyi 5 20.6 (14, 26) 14 (5, 20) 0.6 (—10, 10) 4 (-—9, 10) serratus (Fig. 1b), P. pourtalesii (Stimpson, 1871) (Rathbun 1925) and Heterocrypta tommasil. In Solenolambrus typicus Stimp- son, 1871, and S. portoricensis Rathbun, 1924 (both Rathbun 1925) the position of the ventral spine is occupied by a row of tubercles instead. In Leiolambrus hnitidus, the sternum is smooth, but the merus of the chelipeds has a triangular tooth on the an- teroproximal angle which accomplishes a similar function (Fig. le). Flipse (1930, fig. 40) shows a similar acute spine on the merus of Daldorfia semicircularis (Flipse 1930). Daldorfia horrida (Fig. 1g) however, lacks such an acute spine but has two blunt teeth in place of the ventral spine. The ventral spines of Rhinolambrus longispinis (Fig. 1d) and Platylambrus echinatus are both reduced to blunt tubercles. In P. echinatus (Fig. 1c), the antero-proximal angle has a blunt tooth which opposes the ventral tubercle. This tooth is absent in R. /ongispinis. The ventral spine of Cryptopodia fornicata (Fig. 1f) is quite well developed and opposes another sharp spine on the antero-distal angle of the coxa. The chelipeds are consequently much less mobile when compared to the other par- thenopids and daldorfids. The chelipeds of Daira are extremely mobile, with only a very small tubercle located far back behind the sternum, and no opposing spine or tubercle on any part of the cheliped. 2. Fingers of chelipeds.—In most par- thenopoids, the fingers of the chelipeds are bent downwards, and the dactylus bent in- wards in a very characteristic way. To eval- uate this character, we measured, in mate- rial available to us, (a) the angle formed by the fixed finger with the longitudinal axis of the cheliped (relative downward inclina- tion), and (b) the angle formed by the dac- tylus with the transverse axis of the cheliped (inward inclination) (Table 1). The fixed fin- ger is bent downwards in Platylambrus ser- ratus, P. echinatus, Rhinolambrus longispin- is, Leiolambrus hnitidus, Cryptopodia fornicata, Daldorfia horrida, and Mimilam- brus wileyi. The inclination of the dactylus in Platylambrus serratus is always inwards, but tends to be straight in Cryptopodia for- nicata and Platylambrus echinatus, and in Leiolambrus nitidus, it is in line with the palm of the cheliped. That on Mimilambrus wileyi shows considerable variation, with the inclination being either inwards or out- wards. 3. Dentition of the chelipeds. —Mimilam- brus wileyi shows dimorphic asymmetry in the dentition of the chelipeds (Fig. 21-1); the occlusive surface of the cheliped on one side (the crusher) has a double row of teeth that coalesce side-by-side to form molariform crushers (Fig. 2j, 1). The cheliped of the oth- er side (the cutter) has two rows of broad, thin shearing teeth over the occlusive sur- face of the dactylus, and a row of similar teeth on the occlusive surface of the fixed finger (Fig. 21, k). Additionally, the fixed finger has a lateral row, arched dorsally in the middle, bearing eight triangular teeth. Some variability is displayed in our mate- rial of Mimilambrus. In the male specimen VOLUME 99, NUMBER 1 95 Fig. 2. Dentition of chelipeds in American Parthenopidae and in Mimilambrus: a, b, c, d, Platylambrus serratus; e, f, g, h, Leiolambrus nitidus; i, j, k, 1, Mimilambrus wileyi. a, e, i, Dactylus of left cheliped; c, g, k, Fixed finger of left cheliped; b, f, j, Dactylus of right cheliped; d, h, 1, Fixed finger of right cheliped. from Golfo de Cariaco (cl. 26.3 mm) and Paraguana (cl. 29.7 mm), the molariform crushers have distinct and acute cusps, and the lateral row of teeth of the cutter cheliped are very prominent and convex, being sharp and distinct. In other specimens, the teeth of the crusher cheliped have coalesced to form single, oblong molars, the lateral row of the cutter cheliped is less convex and prominent, with smaller teeth. A similar dimorphic asymmetry is found in the members of the Parthenopidae. In Parthenope agona and Platylambrus serratus (Fig. 2a—d), there are two molar crushers on the fixed finger, but the cusps of the prox- imal one still have the cups discernible, and the cutter cheliped has a row of shearing teeth on both fingers and an additional oblique row of tubercles on the lateral sur- face of the fixed finger. Heterocrypta tom- masii has three molars with indiscernible cusps on the fixed finger of the crusher che- liped, with crenulations on the dactylus, and there is no lateral row of tubercles on the fixed finger of the cutter cheliped. The chelar dentition of Leiolambrus nitidus (Fig. 2e- h) shows a greater resemblance to that of Mimilambrus (Figs. 2e, 1). The crusher che- liped of Leiolambrus has four molars with- out distinct cusps, diminishing in size dis- tally on the fixed finger, while the dactylus is almost devoid of teeth except for a distal protuberance. The cutter cheliped has an indistinct double row of shearing teeth on the dactylus, some of which are fused to form molars. The lateral row of tubercles over the fixed finger is weak, with only mi- nute granules. The parthenopoids from the Indo-Pacific, Cryptopodia fornicata (Fig. 3m—p), Rhino- lambrus longispinis (Fig. 3i-l), Platylam- brus echinatus (Fig. 3e-h), and Daldorfia horrida (Fig. 3a—d) also show heterochely, with a distinct crusher and cutter cheliped. The crusher cheliped of Daldorfia is almost twice the size of the cutter, and the fixed finger has one very massive molar running through most of its length, without cusps, and slightly depressed medially (Fig. 3c). The dactylus bears a row of three strong, 96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. thenopoidea: a, b, c, d, Daldorfia horrida; e, f, g, h, Platylambrus echinatus; i, j, k, 1. Rhinolambrus longi- spinis; m, n, 0, p, Cryptopodia fornicata. a, e, 1, m, Dactylus of left cheliped; c, g, k, 0, Fixed finger of left cheliped; b, f, j, n, Dactylus of right cheliped; d, h, 1, p, Fixed finger of right cheliped. Dentition of chelipeds in Indo-Pacific Par- coniform molars (Fig. 3a). The crusher che- liped of Cryptopodia is slightly less well de- veloped than that of Daldorfia, but the mo- lars on both fingers appear confluent (Fig. 3n-—p). Platylambrus echinatus has two, al- most confluent molars on the fixed finger of the crusher cheliped which have low cusps (Fig. 3h) that occlude on a larger molar on the dactylus (Fig. 3f). The crushing appa- ratus of Rhinolambrus longispinis is quite close to Platylambrus serratus, with two large molars on the fixed finger (Fig. 31) and a row of smaller ones on the dactylus (Fig. 3j), but their cusps are rather lower. The crushing chelipeds are clearly asso- ciated with molluscivorous diets, and the form of the crushing and cutting chelipeds of Mimilambrus falls within the range of variation observed in the Parthenopidae s. s. The chelipeds of the Parthenopidae (in- cluding Mimilambrus) and the Daldorfidae however, show some minor, but distinct dif- ferences. In the daldorfids, heterochely is much more pronounced, and the crushing chelipeds have very much stronger and larg- er molariform teeth than the parthenopids. 4. Ischio-meral articulation of che- lipeds. —Guinot (1968) has stressed the im- portance of this articulation in the phylog- eny of the Brachyura. In Mimilambrus wileyi, the ischio-meral articulation is still mobile, and the two articles are joined to- gether by an arthrodial membrane. This character is commmonly found in the fam- ily Majidae (sensu Guinot 1978), and has been regarded by Guinot (1968) as primi- tive. In the Daldorfidae and Parthenopidae s. s., this character is rather variable, with a tendency towards ankylosis and total dis- appearance of the articulation and the ar- throdial membrane. In Platylambrus serra- tus, P. echinatus, Rhinolambrus longispinis, and Cryptopodia fornicata, no arthrodial membrane is present, and movement of the two articles is at best very slight. In the ge- nus Mesorhoea Stimpson, 1871, Daldorfia horrida, Leiolambrus nitidus, and Parthen- ope agona, there is a thin arthrodial mem- brane, but movement of the articles is still very restricted. In Daira, the arthrodial membrane is present, but it is thin, and the two main articles remain, for all purposes, immobile. 5. Second male pleopod (P2).—The P2 of all very primitive crabs (Dromiidae, Dy- nomenidae, and Homolidae) are long and whiplike, the cup is absent, and the basal portion of the flagellum cannot be differ- VOLUME 99, NUMBER 1 entiated. In some groups, e.g. in the super- family Xanthoidea (sensu Guinot 1978a, b), both long and short P2s can be found, and Ng (1983) has suggested that the long P2 represents the plesiomorphous condition. In the Parthenopoidea, the morphology of the P2 is slightly more homogeneous (Fig. 4). An examination of the material depos- ited in our collections, and the figures of these appendages that are available in the literature reveal that there are approxi- mately three main types within the super- family. In the following list, we give the ratio of the flagellum to the total length of the appendage (f/a). (A) Flagellum long and whiplike f/a > 0.4. Aethra scruposa (Linnaeus, 1764) (Guinot 1967a, fig. 40). Daldorfia horrida (Linnaeus, 1758). Daira perlata (Herbst, 1790). Dairoides margaritatus (Stebbing, 1920) (Guinot 1967b, fig. 14). (B) Flagellum short, in the form ofa spike, sometimes with basal cup, ratio f/a between 0.2 and 0.4. Parthenope macrochelos (Herbst, 1790) (Monod 1956, fig. 861). P. agona (Stimpson, 1871). Platylambrus pourtalesii (Stimpson, 1871) (Williams 1965, fig. 252C). P. fraterculus (Stimpson, 1871) (Williams 1965, fig. 252D). P. echinatus (Herbst, 1790). Rhinolambrus longispinis Miers, 1879. R. massena (Roux, 1830) (Monod 1956, fig. 856). Cryptopodia fornicata (Fabricius, 1781). C. concava Stimpson, 1871 (Gore and Scot- to 1979, fig. SI). Leiolambrus nitidus (Rathbun, 1901). Solenolambrus typicus Stimpson, 1871 (Gore and Scotto 1979, fig. 9C). (C) Flagellum very short, forming a cup- like structure, ratio f/a < 0.2. Platylambrus serratus (H. Milne Edwards, 1834). 97 imm Fig. 4. Second male pleopod: a, Platylambrus ser- ratus; b, Parthenope agona; c, Mimilambrus wileyi; d, Leiolambrus nitidus. Heterocrypta maltzami Miers, 1881 (Mon- od 1956, fig. 867). H. granulata (Gibbs, 1850) (Williams 1965, fig. 252E). Thyrolambrus erosus (Miers, 1879) (Guinot 1967a, fig. 37). All the genera with an f/a > 0.4 belong to the families Dairidae, Aethridae, and Daldorfidae, whereas genera with f/a < 0.4 constitute the Parthenopidae s. s. Mimilam- brus wileyi belongs to the second group of P2 type, since its f/a is 0.22 with the fla- gellum relatively short, spikelike, with a fairly well developed basal cup, i.e., the Par- thenopidae group. Also, as mentioned earlier, the genera Daira and Dairoides or their allies probably gave rise to the main line of the partheno- pids, and their male abdominal segments 3 to 5 are ankylosed, with the sutures still present, and their P2s are of the first group, i.e., with the flagellum long and whiplike. There is thus a very good chance that the evolutionary trend of the P2 observed in the Xanthoidea (Ng 1983) is also true for the Parthenopoidea. In summary, some of the characters of Mimilambrus, such as the articulation of the male abdominal segments 3 to 5, and the 98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ischio-meral articulation of the chelipeds appear to be rather primitive. The struc- tures of the afferent and efferent channels, on the other hand, are advanced and spe- cialized. The vertically folding antennules and form of the crusher cheliped of Mimi- lambrus finds an analogue in Leiolambrus, with their mouthparts and afferent channels also quite similar. The very unusual mouth- part arrangement of Mimilambrus with its medial efferent opening is not unique to it, but can also be found in Mesorhoea. The only unusual character of Mimilambrus seems to be the free chelar ischio-meral joints with an anthrodial membrane, which appears to be a plesiomorphous condition. In the more primitive parthenopoids like Daira, these joints are tightly appressed, but still distinct. The genera Mesorhoea and Leiolambrus also have the two joints dis- tinct, but basically immobile. The condition of the ischio-merus in Mimilambrus sug- gests that it may be a fairly primitive and specialized species. Presumably, the genus represents an early offshoot of the main par- thenopoid stock. The respiratory adapta- tions could be explained by convergent evo- lution, and are unlikely to have been derived from similarly advanced structures like those on Leiolambrus and Mesorhoea. Placing Mimilambrus in a family (and superfamily) of its own is, therefore, unwarranted since it shares far too many common characters with the Parthenopidae s. s., and should in- stead be regarded merely as a rather spe- cialized genus in the family. Acknowledgments The authors wish to thank Dr. Robert Gore for his kind information regarding the larvae of Echinoecus, Mrs. C. M. Yang of the ZRC for permission to examine the col- lections in her charge, and Dr. Leo W. H. Tan for his kind advice. This study has been partly supported by a 1984 grant to the first author from the Singapore Institute of Bi- ology. Literature Cited Alcock, A. 1895. Materials for a carcinological fauna of India N° 1. The Brachyura Oxyrhyncha.— Journal of the Asiatic Society of Bengal 64:157-— 291. Balss, H. 1957. Decapoda. Jn H. G. Bronns, Klassen und Ordnungen das Tierreichs. 5 Band, I Ab- teilung, 7 Buch, Decapoda (Zehnfusser), 12 Lie- ferung: 1505-1672. Bowman, T. E., and L. G. Abele. 1982. Classification of the recent Crustacea. Jn D. E. Bliss, ed., The Biology of the Crustacea. Volume 1. System- atics, the fossil record and biogeography. Ac- ademic Press, New York, pp. 1-27. Flipse, H. J. 1930. Die Decapoda Brachyura der Si- boga-Expedition. VI. Oxyrhyncha-Parthenopi- dae.—Siboga Expedition Monographie 39 C?: 1-96. Garstang, W. 1897a. Contributions to marine bio- nomics IJ. The function of antero-lateral den- ticulations of the carapace in sand burrowing crabs.—Journal of the Marine Biological As- sociation 4:396-401. 1897b. On some modifications of structure subservient to respiration in Decapod Crustacea which burrow in sand; with some remarks on the utility of specific characters in the genus Ca- lappa, and the description of a new species of Albunea. — Quarterly Journal of the Microscop- ical Society 40:211-—232, pls. 12-14. Garth, J. S. 1958. Brachyura of the Pacific Coast of America, Oxyrhyncha.— Allan Hancock Pacific Expeditions 21(1):i—xii, 1-499. Glaessner, M. F. 1969. Decapoda. Jn R. C. Moore, ed., Treatise on Invertebrate Paleontology, Pt. R, Arthropoda 4, vol. 2. University of Kansas and Geological Society of America Inc, pp. R399-533, R626-628. Gore, R. H., and Scotto, L. E. 1979. Crabs of the family Parthenopidae with notes on specimens from the Indian River region of Florida.— Memoirs of the Hourglass Cruises 3 (6):1—98. Guinot, D. 1966. Recherches préliminaires sur les groupements naturels chez les Crustacés Déca- podes Brachyoures. I. Les affinités des genres Aethra, Osachila, Hepatus, Hepatella et Actaeo- morpha. —Bulletin du Muséum National D’Histoire Naturelle, Paris (2) 38(5):744-762. 1967a. Recherches préliminaires sur les groupements naturels chez les Crustacaes Déca- podes Brachyoures. I. Les affinités des genres Aethra, Osachila, Hepatus, Hepatella et Actaeo- morpha (suite et fin). — Bulletin du Muséum Na- tional d’Histoire Naturelle, Paris (2) 38 (5):828- 845. 1967b. Recherches préliminaires sur les VOLUME 99, NUMBER 1 groupements naturels chez les Crustacés Déca- podes Brachyoures. III. A propos des affinités des genres Dairoides Stebbing et Daira de Haan.—Bulletin du Muséum National d’His- toire Naturelle, Paris (2) 39 (3):540-563. 1968. Recherches préliminaires sur les groupements naturels chez les Crustacés Déca- podes Brachyoures. V. Establissment d’un car- actére évolutif: l’articulation ischiomerale des chélipedes.— Bulletin du Muséum National d'Histoire Naturelle, Paris (2) 40 (1):149-166. 1977a. Données nouvelles sur la morpho- logie, la phylogenese et la taxonomie des Crus- tacés Décapodes Brachyoures. These de Doc- torat de’Etat és Sciences, soutenus le 21 Jun 1977 a Université Pierre-et-Marie-Curie. 2 vol. i folio, pp. I-XV, 1-486, X VI-XXIV, 56 feuilles n.n., 78 figs., 31 pls., 2 Fig. n.n., 14 Tabls. . 1977b. Propositions pour une nouvelle clas- sification des Crustacés Décapodes Brachy- oures.—Comptes Rendus hebdomadaires des Seances de l’Academie des Seances (D) 285: 1049-1052. 1978a. Principes d’une classification évolu- tive des Crustacés Décapodes Brachyoures. — Bulletin Biologique de la France et de la Bel- gique, nouvelle série 112(3):211-292. 1978b. Analyse morphologénétique d’une lignée de Crabes: la lignée “parthénoxysto- mienne” et position systématique de genre Dra- chiella Guinot (Crustacea, Decapoda, Brachy- ura). Jn volume jubilaire du Prof. Pierre Drach. — Archives de Zoologie Expérimentale et Génér- ale 119:7-20. 1979. Morphologie et phylogenése des Brachyoures.— Mémoires du Muséum National d’Histoire Naturelle, Paris, nouvelle série (A) 112:1-354, pls. 1-27. Manning, R. B., and L. B. Holthuis. 1981. West Af- rican brachyuran crabs (Crustacea: Decapo- da).—Smithsonian Contributions to Zoology 306:1-379. Milne Edwards, A. 1878. Etudes sur les Xiphosures et les Crustacés de la Région Mexicaine. Mission Scientifique au Mexique.—Recherches Zoolo- giques (5) 1:1-368. Monod, T. 1956. Hippidea et Brachyura ouest afri- caines.—Mémoires de l'Institut Frang¢ais de lAfrique Noire 45:1-674. Ng, P. K. L. 1983. Aspects of the systematics of the family Pilumnidae Samouelle, 1819 (Crustacea, Decapoda, Brachyura) and a study on evolu- 99 tionary trends within the superfamily Xanthoi- dea (sensu Guinot 1978). Unpublished Honours Thesis, National University of Singapore, pp. i— vili, 1-251, 12 pls. Rathbun, M. J. 1925. The Spider Crabs of Ameri- ca.— Bulletin of the U.S. National Museum 129: 1-613. 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. 1983. Zoeal evidence for Brachyuran phy- logeny. Jn F. R. Schram, ed., Crustacean Issues I. Crustacean Phylogeny. A. A. Balkema, Rot- terdam. Pp. 313-329. Rodriguez, G. 1980. Los Crustaceos Decapodos de Venezuela. Instituto Venezolano de Investi- gaciones Cientificas, Caracas, 428 pp. Saint Laurent, M. de. 1980. Sur la classification et la phylogenie des Crustacés Décapodes Brachy- oures. IJ. Heterotremata et Thoractremata Guinot, 1977.—Comptes Rendes des Séances de l’Academie des Sciences (D) 290:1317—1320. Schafer, W. 1954. Form und Funktion der Brachy- uren-Schere.—Abhandlungen der Senckenber- gischen Naturforschende Gesellschaft 489:1-65. Seréne, R. 1968. The Brachyura of the Indo-West Pacific region. Jn Prodromus for a Check List of the non-planctonic marine fauna of South East Asia. Special publication N° 1. Singapore National Academy of Science, Singapore. Pp. 33-112. , V. L. Tran, and V. L. Nguyen. 1958. Eu- medoinae du Vietnam (Crustacea) avec une bib- liographie de la sousfamille.—Treubia 24:135-— 242. Stevcic, Z., and R. H. Gore. 1981. Are the Oxyrhyn- cha a natural group?.— Thalassia Jugoslavia 17: 1-16. Williams, A. B. 1965. Marine Decapod Crustaceans of the Carolinas.— Fishery Bulletin 65:1—298. 1979. A new crab family from the shallow waters of the West Indies (Crustacea: Decapoda: Brachyura).— Proceedings of the Biological So- ciety of Washington 92:399-413. (PKLN) Department of Zoology, Nation- al University of Singapore, Singapore; (GR) Instituto Venezolano de Investigaciones Cientificas, Apartado 1827, Caracas 1010- A, Venezuela. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 100-109 DESCRIPTIONS OF TWO NEW SPECIES OF HYLODES FROM THE ATLANTIC FORESTS OF BRAZIL (AMPHIBIA: LEPTODACTYLIDAE) W. Ronald Heyer and Reginald B. Cocroft Abstract.— Analysis of available advertisement calls and morphology indi- cates that Hylodes babax and H. lateristrigatus occur at Santa Teresa, Espirito Santo, Brazil, and that several new species are present within the Hylodes lateristrigatus species cluster. We have adequate materials for description of two of these: H. charadranaetes from Teresopolis, Rio de Janeiro, and H. phyllodes from Boracéia, Sao Paulo. This study was begun to determine the nomenclatural status of the smaller species of Hylodes that occurs at Boracéia, Sao Pau- lo, Brazil. This interest was stimulated by a forthcoming summary of the frogs of Bor- acéia (Heyer, Rand, Cruz, Peixoto, Nelson, in prep.). In an effort to use the proper names for the Hylodes, it became apparent that we would have to evaluate data previously published on species from elsewhere in its range. W. C. A. Bokermann and Eugénio Izecksohn are involved in a long-term re- vision of the genus Hylodes (Bokermann, pers. comm.). At their suggestion (Boker- mann, pers. comm.), we report our data and conclusions to facilitate work on the Bora- céia fauna. Two names have been used in the liter- ature for the smaller Hylodes from Boracéia and/or Paranapiacaba: glabrus (as Elosia glabra, Bokermann 1967a) and /ateristri- gatus (as Elosia lateristrigata, Cochran 1955). Heyer (1982) demonstrated that H. lateristrigatus did not apply to the species from Paranapiacaba, and questioned whether glabrus was the correct name for the Paranapiacaba species. Izecksohn and Gouvéa (1983) conclude that Elosia glabra Miranda-Ribeiro, 1926, is a senior syn- onym of E£. pulchra Lutz, 1951. Through the courtesy of Professor Antenor Leitao de Carvalho, we were able to borrow and ex- amine the holotype of Elosia glabra, and we agree with their conclusion. The Boracéia/ Paranapiacaba species is clearly a /ateristri- gatus group member. Hylodes glabrus is not, since it lacks the diagnostic dorsolateral light stripe, so this name can not be applied to the Boracéia/Paranapiacaba populations. Based on proximity of localities in the same block of the Serra do Mar, Paranapiacaba and Boracéia would be expected to have the same species of Hylodes. The purposes of this report are: (1) to summarize data we have analyzed on the advertisement calls and external adult mor- phology of Hylodes; and (2) to describe new species for those populations lacking names and for which we have adequate materials for description. Variation in Calls and Morphology of Geographic Samples of the lateristrigatus Group Heyer (1982) proposed informal species clusters for Hylodes based on external mor- phology. We follow that system, recognizing that the groupings are ones of convenience and may not be monophyletic. With the ex- clusion of glabrus from this group, the fol- lowing names pertain to the /ateristrigatus group: Hylodes babax Heyer, 1982; Hylodes lateristrigatus (Baumann, 1912); Hylodes magalhaesi (Bokermann, 1964); Hylodes ornatus (Bokermann, 1967b); Hylodes ota- vioi Sazima and Bokermann, 1982; Hylodes VOLUME 99, NUMBER 1 101 Table 1.—Advertisement call characteristics of members of the Hylodes lateristrigatus group. 2 Dominant Frequency Call duration #Notes/ Note duration frequency modulation Population (s) call (s) #Notes/s (Hz). of notes H. babax 0.2-0.5 4-8 0.04—0.05 16-17 4290-5420 H. lateristrigatus 1.3-1.6 12-13 0.05—0.07 8-9 3700-4300 AP {I H. magalhaes? 1.0 26 0.02-0.03 20-25 1400-3000 +P Il H. otavior® 2.0 6-15 0.09-0.10 8 3800-5200 — H. regius 1.7 22-32 0.02-0.03 14-19 5200-6300 a5 || Boracéia =P || (A. phyllodes) 1.0—2.1 12-20 0.05—0.06 8-11 4100-5700 Paranapiacaba‘ a5 Il (H. phyllodes) 1.8—3.0 15-32 0.03-0.05 10-13 5400-60002 Santa Teresa A + (weak), | (A. lateristrigatus) 2.4 19 0.05-0.06 8 3100-3900 “Po | Santa Teresa B (H. babax) 0.6-0.8 10-12 0.03-0.05 14 3500-4900 +,] Teresopolis (H. charadranaetes) 1.1-1.3 2-4 0.06-0.17 2-4 4300-5500 arp {ll a Data from Bokermann 1964. > Data from Sazima and Bokermann 1982. ¢ Data from Bokermann 1964, 1967. 4 Published figures indicate dominant frequency range of 4200-5700 Hz. regius Gouvéa, 1979; Hylodes vanzolinii Heyer, 1982. Calls. — Advertisement calls are known for all named members of the Hylodes la- teristrigatus group except for ornatus and vanzolinii. It is likely that vanzolinii is voiceless (Heyer 1982). In addition to ad- vertisement calls from the type localities of the remaining named species, call data are available for the /ateristrigatus group from Boracéia, Sao Paulo; Paranapiacaba, Sao Paulo; two species from Santa Teresa, Es- pirito Santo; and a second species (in ad- dition to /ateristrigatus) from Teresopolis, Rio de Janeiro. Few recordings are available for most of these species; for the following comparisons, the number of calls analyzed ranges from three calls from one individual from the Santa Teresa B population to ten calls from two individuals of Hylodes char- adranaetes. Field observations (Heyer, pers. obs.), however, indicate that call character- istics in Hylodes are consistent within pop- ulations, so these samples are considered to be representative. None of the calls are identical when the major call features are compared (Table 1). In the two instances of sympatry (Santa Ter- esa and Teresopolis), the calls differ strik- ingly. At Teresopolis, /ateristrigatus and the second species (Hylodes charadranaetes n. sp., see below) differ by at least an order of magnitude in number of notes per call and number of notes per second. The two forms are also differentiated in terms of note du- ration, dominant frequency channel, and frequency modulation within notes (Table 1). The two forms from Santa Teresa differ strikingly in call duration, number of notes per call, and number of notes per second; the broadcast channel and note duration are also distinctive, though they show some overlap (Table 1). The distinctiveness of the calls of the two forms at Santa Teresa and TeresOpolis is consistent with species level differentiation. Most two-way comparisons of popula- tions for which call data are available dem- onstrate the same degree of differentiation observed in the sympatric pairs (Table 1). Certain two-way comparisons are not as distinctive, and each is discussed. 102 1) Hylodes babax-Santa Teresa B. None of the major call characteristics (Table 1) differ markedly between the two samples, although the calls are distinct in duration, number of notes per call, and number of notes per second. When tapes of these calls are played one after the other, the calls sound distinctive to the human ear, but it is not possible to tell whether the differences are due to recording (i.e., equipment, distance from specimen, etc.), individual, or species differences. 2) Hylodes lateristrigatus-H. otavioi. These calls differ somewhat in call duration, broadcast channel, and note duration. Comparison of audiospectrograms indi- cates more similarities than differences be- tween calls. The call data can not be un- ambiguously used to determine whether distinct species are involved. 3) Hylodes otavioi-Boracéia and Parana- piacaba. These calls differ slightly in terms of number of notes per call and distinctively in note duration. Comparison of audio- spectrograms indicates a different note structure for H. otavioi; the notes of the Bor- acéla and Paranapiacaba populations have a simpler and more vertical appearance on the audiospectrograms. Furthermore, there is no indication of pairing of notes in H. otavioi, which occurs in the other two pop- ulations. We believe these differences are indicative of species level differentiation. 4) Hylodes lateristrigatus-Santa Teresa A. The calls differ only in call duration and number of notes per call; this simply indi- cates a longer call. The broadcast channel, though somewhat distinctive, is overlap- ping. Comparison of audiospectrograms of the calls shows them to be very similar in other structural components. The differ- ences between calls are more of the level expected for geographic variation than for species level differentiation. 5) Boracéia-Paranapiacaba. The calls dif- fer somewhat in each of the major call char- acteristics (Table 1) but each characteristic shows some overlap. If the data from Bok- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ermann’s (1964, 1967a) figures are used rather than the values he presents in the text, the broadcast channels are identical. Comparison of audiospectrograms indi- cates that two features are shared by these two populations alone. First, there is no- ticeable frequency modulation within the call, which begins at a higher frequency and ends at a lower frequency. Other known /a- teristrigatus group members have either no noticeable frequency modulation of the call or an increase in frequency after the first one or two notes of the call. Second, the calls from the Boracéia population tend to have paired notes given at the end of the call. Bokermann’s (1964, 1967a) audio- spectrograms suggest the same pattern, though not as strongly as do the Boracéia audiospectrograms. We interpret the Bor- acéia and Paranapiacaba calls to represent the same species. Morphology. —Within the Jateristrigatus group, H. ornatus, H. regius, and H. van- zolinii are distinctive in size and/or pattern (see Heyer 1982). The remaining popula- tions are strikingly similar in morphology. The two forms occurring at Teresopolis dif- fer mostly in size, with subtle pattern dif- ferences (Heyer 1982). There is a much smaller size difference between the two forms from Santa Teresa (adult males 32.7—33.6 mm SVL versus 36.7—37.1 mm SVL) and we can find no other differences. In order to examine the morphological variation among populations of the /ateristrigatus group that are similar in appearance, a dis- criminant function analysis was performed. Although samples were limited, the results are instructive. Because only males were available for the Santa Teresa A and B forms, the analysis is limited to males. The vari- ables measured and used in the analysis were: SVL, head length, head width, vertical tympanum diameter, eye diameter, eye- nostril distance, internarial distance, width of third finger disk, width of fourth toe disk, hand length, femur length, tibia length, and foot length. Initially, three samples were VOLUME 99, NUMBER 1 103 7.07 O A 3.57 A ee @ . So © & w eo ee = O < 904 ee a ‘4 > a A & (S) FS ot Gd & A fo) A z A An A © A = 1/30) 5] I T T T -10 -5 (0) 5 CANONICAL VARIABLE | Fig. 1. Discriminant axis plot of males of four samples of the Hylodes lateristrigatus group. Triangles = TeresOpolis, solid triangles = Santa Teresa B, circles = Jateristrigatus, solid circles = Boracéia. used for the preformed groups: /ateristri- gatus (two males from Teresopolis and one from Correias), TeresOpolis, and Boracéia. These three groups were separated by the discriminant function analysis. The canon- ical variable coefficient results were used to plot four additional specimens on the plot of the first two canonical variables. Prior to the analysis, we were unable to differentiate morphologically the J/ateristrigatus males from the two Santa Teresa A males. The two Santa Teresa A males were adjacent to the Jateristrigatus males when plotted, and the two Santa Teresa B males were closest to the Santa Teresa A males. Next, four pre- formed groups were analyzed: /ateristriga- tus (five males from Correias, Teresopolis, and Santa Teresa), Boracéia (ten males), Teresopolis (ten males), and Santa Teresa B (two males). There is 100% discrimina- tion of cases among these groups. The plot of the first two canonical variables accounts for 95% of the total dispersion, and best separates the groups along the first canon- ical variable, which is size related (Fig. 1). In fact, the Boracéia, Santa Teresa B, and lateristrigatus groups are only differentiated on the first axis and the Santa Teresa B and lateristrigatus groups are very close to each other, even in size. Only the Teresdpolis group is distinctive along the second ca- nonical variable axis. The variable with the highest loading on this axis is the width of the third finger disk (20.4; the next highest value, —9.2, is for eye-nostril distance). Comparison of specimens indicates that the finger disk size does not allow visual dis- crimination of the Teresopolis specimens from the other samples. The morphological analyses indicate that: (1) the males of Santa Teresa A and Jateristrigatus form a single morphological group; (2) the Santa Teresa B group is very similar to the /ateristrigatus group; and (3) the four groups are quite sim- ilar, differing principally in size. There are too few specimens to perform a meaningful discriminant function analysis on external morphology between H. babax and Santa Teresa B males. Direct compar- isons indicate similarities in size, pattern, and color. There is overlap in male size (ba- bax 30.6-—32.6 mm SVL, Santa Teresa B 104 32.0-33.6 mm SVL). The belly patterns of boldly contrasting dark and light mottling are similar. The undersides of the femurs are red in life for both samples (Peter Wey- goldt, pers. comm. for Santa Teresa B spec- imens), a condition recorded only from these two samples and H. regius. We do not be- lieve the available data support recognition of Santa Teresa B as a species distinct from H. babax. We do point out, however, that differences in call parameters (noted above) and pattern (the lateral stripes are more dis- tinct in the Santa Teresa specimens) exist in the few individuals at hand, and encour- age additional call and morphological sam- ples to be gathered to understand the nature of the differences observed. Combining the available advertisement and morphological data with previously published analyses, we recognize the follow- ing species in the /ateristrigatus group: ba- bax (including Santa Teresa B), /ateristri- gatus (including Santa Teresa A), magalhaesi, ornatus, otavioi (more infor- mation is needed to clarify the specific sta- tus of otavioi relative to /ateristrigatus), re- gius, vanzolinii, a new species for the populations from Boracéia and Paranapia- caba, a new species for the second species from TeresOpolis, and a new species for an Itatiaia population which is morphologi- cally distinct, but for which only one adult and no call recordings are available. The two new species for which adequate samples are available are described in the following section. Hylodes phyllodes, new species Fig. 2 Holotype.—MZUSP 59934, male, from Brazil: Sao Paulo; Boracéia, 23°38’'S, 45°50'W. Collected by W. Ronald Heyer, 6 Dec 1976. Paratopotypes. -MZUSP_ 1700-1702, 1704-1706, 1708-1711, 1714, 1716-1721, 3308, 3527, 3529, 4040-4043, 4143, 23050- 23053, 23561, 23678-23697, 36874, 37573-37585, 37678-37687, 37701-37703, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 37712-37717, 38854, 56497; USNM 129156-129158, 243480-243506. Diagnosis. — Hylodes phyllodes has a light stripe from the eye to the groin, distinct at least posteriorly; H. vanzolinii lacks such a stripe. The dorsum of H. phyllodes lacks the distinct light spots found in H. regius. Hy- lodes phyllodes is larger (males 27.5—31.4 mm SVL, females 29.0-35.5 mm SVL) than ornatus (males and females 23.2—26.1 SVL) and smaller than /ateristrigatus (males 36.7— 39.2 mm SVL) and otavioi (males and females31.4—34.0 mm SVL). Hylodes phyl- lodes differs consistently from babax, char- adranaetes, and magalhaesi in advertise- ment call characteristics. In life, H. phyllodes lacks the brick red color on the under sur- face of the legs found in H. babax. The var- iegated belly of H. phyllodes never consists of a bold pattern of distinct light spots on a dark ground which is found in some in- dividuals of H. magalhaesi. Hylodes phyl- lodes is smaller than charadranaetes (males to 34.7 mm SVL, females to 37.7 mm SVL) and has a more slender body form. Male H. phyllodes differ from all other known male Hylodes in having nuptial thumb spines. Description of holotype.—Snout truncate from above, protruding in profile; canthus rostralis sharp; lores vertically concave in cross section; tympanum distinct, large, di- ameter about 73 diameter of eye; vomerine teeth in two small transverse patches be- tween and on line drawn across posterior edges of choanae, separated by less than length of one vomerine tooth patch; vocal slits present, near angle of jaw on each side; vocal sacs paired, lateral, inflated; finger lengths I = II < IV < III; middle of thumb with scattered small whitish spines in small ovate area on inner and dorsal surface; dor- sal texture finely etched, few scattered warts posteriorly, dorsal surfaces of legs with se- ries of longitudinal ridges; weak fold from eye to groin; throat and belly smooth, under surfaces of thighs smooth anteriorly, areo- late posteriorly; finger and toe tips with disks, disks about twice as broad as digit imme- VOLUME 99, NUMBER 1 105 Fig. 2. Holotype of Hylodes phyllodes (MZUSP 59934, a male): dorsal and ventral views. diately behind disk, finger and toe disks about equal size, upper surface of disks with pair of scutes; fingers fringed, most exten- sively on fingers III and IV; toes extensively fringed; subarticular tubercles moderate sized, rounded; inner oval metatarsal tu- bercle about twice as large as pungent, rounded outer metatarsal tubercle; exten- sive tarsal fold-flap extending 7% distance of tarsus, continuous distally with toe fringe on outer side of first toe; no metatarsal fold; outer tarsus and sole of foot smooth. SVL 29.2 mm, head length 10.2 mm, head width 8.9 mm, eye-nostril distance 2.1 mm, femur 13.1 mm, tibia 14.4 mm, foot 14.5 mm. Dorsum almost uniformly dark brown (fine bronze and gray mottle under micro- scope) with faint darker mid-dorsal pin stripe with small whitish dots spaced along stripe; upper limbs tan with narrow brown cross bands; flank a blending of dorsal and ventral patterns with distal 3 of eye-groin fold whitish and distinct; continuation of dorsal color as dark canthal stripe, rest of face lighter brown and white mottle, lightest under eye and continuing under tympanum to shoulder; mid-ventral brown stripe from almost tip of chin through chest to ante- riormost belly; rest of throat and belly mot- tled brown and white; under limbs mottled with brown and pigmentless areas; posterior surfaces of thighs indistinctly mottled dark- er and lighter browns. Variation. — Males range from 27.5-31.4 mm SVL, females 29.0—35.5 mm SVL. In preservative, the dorsum ranges from brown to brassy brown with various mark- ings, including an irregular mid-dorsal light stripe with almost regularly spaced small darker brown (than dorsum) spots, or a se- ries of short dark mid-dorsal dashes, or dor- sum scattered with small dark brown spots, or series of mid-dorsal large light blotches, or big darker brown blotches on a lighter brown background, or almost uniform with a series of faint light mid-dorsal dots; pair of dark round or U-shaped spots, one on each side of anus; upper limbs distinctly to indistinctly crossbarred brown on tan; flank with dark brown (almost black) band be- hind eye across tympanum just to or above arm, fading to mottled brown and white, ranging to flank almost uniformly dark. In life, iris copper to yellow, darker on sides forming black band with pupil; dor- 106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 8 eek. he pe oe Oe ee fous f eee ae. 6 t § $ : ; 3 9 A : ; * = Fd a» a ia om fa ia fo i fh a Re? a A 4 = j 3 Sige ff ff ff ff 7. eee KILOHERT Z SECONDS Fig. 3. Advertisement call of Hylodes phyllodes. From specimen MZUSP 59934, Brazil, Sao Paulo, Boracéia. Recorded 6 December 1976, air temperature 20.4°C. sum olive to dark and light brown; light stripes cream, copper, yellow, yellow-gold, or gold; flanks dark brown; upper limbs dark and light brown with or without red cast; groin yellow or not; throat and belly opal- escent, bright opalescent yellow, or yellow to golden with variable brown spotting, dis- tinct or not; under limbs colorless to dirty greenish yellow. Advertisement call.—Calls given sporad- ically, call duration 1.05—2.10 s; 12—20 notes per call given at rate of 8-11 per s; notes given at regular intervals at beginning of call, usually given in pairs at end of call; note duration 0.05—0.06 s; individual notes not pulsed, but weakly pulsatile; calls slight- ly frequency modulated, beginning higher, ending lower; calls not noticeably intensity modulated; fundamental frequency about 1500-2200 Hz; dominant frequency (=third harmonic) range at beginning of call 4300- 5700 Hz, at end of call 4100-5300 Hz, call with harmonic structure (Fig. 3). Etymology. —From the Greek phyllo (leaf) and oides (like) in allusion to the difficulty of visually distinguishing the frogs from leaves on or near the ground along streams during the day. Referred specimens. —Brazil: Sao Paulo; Caminho do Mar km 47 (very near Par- anapiacaba), MZUSP 10216, Ilha de Sao Sebastiao, MZUSP 9973-9974, 51669, Ilha dos Buzios, MZUSP 23952-23955, Sao Se- bastiao, MZUSP 58717. Hylodes charadranaetes, new species Fig. 4 Holotype.—MZUSP 60648, male, from Brazil: Rio de Janeiro; Alto do Soberbo, near TeresOpolis, 22°26’S, 42°59'W. Col- lected by Ronald I. Crombie, Maria Chris- tina Duchéne, and W. Ronald Heyer, 10 Dee 1977: Paratopotypes.—MZUSP 60649-60669, USNM 245894-245915, collected from Brazil: Rio de Janeiro; 2-5 km NE junction BR 116 and Teresdpolis bypass on various dates by various collectors. Diagnosis. —Hylodes charadranaetes has a light stripe from the eye to the groin, dis- tinct at least posteriorly; H. vanzolinii lacks such a stripe. The dorsum of H. charadran- aetes lacks the distinct light spots found in H. regius. Hylodes charadranaetes is larger (males 31.3-34.7 mm SVL, females 31.9- 37.7 mm SVL) than ornatus (males and fe- males 23.2—26.1 mm SVL) and smaller than lateristrigatus (males 36.7-39.2 mm SVL). Hylodes charadranaetes differs consistently from babax, magalhaesi, otavioi, and phyl- lodes in advertisement call characteristics. In life, H. charadranaetes is not brick red VOLUME 99, NUMBER 1 107 Fig. 4. Holotype of Hylodes charadranaetes (MZUSP 60648, a male): dorsal and ventral views. on the under surface of the legs as is H. babax. The variegated belly of H. charad- ranaetes never consists of a bold pattern of distinct light spots on a dark ground such as found in some individuals of H. magal- haesi. Hylodes charadranaetes is more ro- bust than H. otavioi, phyllodes, and babax and is somewhat larger than either phyllodes (males to 31.4 mm SVL, females to 35.5 mm SVL) or babax (males to 33.6 mm SVL). Male H. charadranaetes differ from male H. phyllodes in lacking nuptial thumb spines. Description of holotype. —Snout sub- ovoid from above, protruding in profile; canthus rostralis sharp; lores almost vertical in cross section; tympanum large, diameter greater than 2 diameter of eye; vomerine teeth in two small patches, separated by about length of one tooth patch, lying on line between posterior portions of choanae; vocal slits present, near angle of jaw; vocal sacs paired, lateral, inflated; first, second, and fourth fingers subequal, third longest; thumb lacking nuptial asperities; dorsal tex- ture finely etched, few scattered warts pos- teriorly; fold from eye over tympanum to groin, upper legs with series of longitudinal ridges; throat and belly smooth, under sur- faces of thighs smooth anteriorly, areolate posteriorly; finger and toe tips with disks, disks about '2 again as broad as digit im- mediately behind disk, finger and toe disks about same size, upper surface of disks with pair of scutes; fingers fringed, developed into extensive flaps on fingers III and IV; toes with extensive fringes produced into flaps; subarticular tubercles moderate, rounded; inner ovate metatarsal tubercle twice size of rounded outer metatarsal tubercle; exten- sive tarsal fold extending % distance of tar- sus, continuous distally with fringe on outer side of first toe; no metatarsal fold; outer tarsus and sole of foot smooth. SVL 34.4 mm, head length 12.2 mm, head width 10.4 mm, eye-nostril distance 2.6 mm, femur 16.2 mm, tibia 17.3 mm, foot 15.8 mm. Dorsum variegated tan and light brown with field of darker brown spots almost reg- ularly arranged in rows; upper limbs tan with narrow brown cross bands; dark brown spot on either side of anus; dark canthal 108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON KILOHERTZ SECONDS Fig.5. Advertisement call of Hylodes charadranaetes. From specimen USNM 245909, Brazil, Rio de Janeiro, near TeresOpolis. Recorded 8 December 1977, air temperature 21.2°C. stripe extending anteriorly past nostril to about '4 way to tip of snout, below canthal stripe light with scattering of brown mela- nophores, becoming lighter under eye and continuing as light stripe to under middle of tympanum; flank a blending of dorsal and ventral patterns with light stripe highlight- ing eye-groin fold; throat and belly varie- gated white and tan, tan mid-throat stripe; under arm with scattering of white pigment, under legs variegated tan and pigmentless areas; posterior thigh mottled tan and dark brown. Variation. —Males range from 31.3—-34.7 mm SVL, females 31.9-37.7 mm SVL. Variable number of dorsal spots, some- times definitely arranged in rows; few spec- imens with faint row of small light dots mid-dorsally; light lip stripe continuing to shoulder in some specimens; eye-groin fold more or less distinctly highlighted by white stripe, white always distinct in groin area; degree of ventral mottling ranging from mostly white with few tan markings to equal amounts of white and tan or brown ver- miculation. In life, iris bronze; lip stripe cream; eye- groin stripe tan anteriorly, cream posterior- ly, belly ranging from white, coppery white, coppery, to yellow, colors on belly same as those on front of thighs and ventral calves; posterior surface of thighs with faint red wash (belly colors noted for several indi- viduals; other color description based on USNM 245908). Advertisement call.—Calls given sporad- ically, call duration 1.1—1.3 s; 2-4 notes per call given at rate of 2—4 notes per s; note duration 0.06-0.17 s; notes not noticeably pulsed; each note frequency modulated, ris- ing then falling; calls not noticeably fre- quency or intensity modulated; fundamen- tal frequency range about 1000-1500 Hz, dominant frequency (third harmonic) range 4300-5500 Hz; calls with harmonic struc- ture (Fig. 5). Etymology.—From the Greek charadra (bed of mountain stream) and naetes (in- habitant), referring to the habitat of the species. Discussion Hylodes phyllodes is the only Hylodes with nuptial spines. As expected, these spines oc- cur only in male H. phyllodes. The nuptial spines in H. phyllodes likely serve a function different from the thumb spines occurring in the related genus Crossodactylus, where they occur in both sexes (a rare condition in frogs). Maxson and Heyer (1982) presented data on relationships among certain populations VOLUME 99, NUMBER 1 Table 2.—Cross reaction of certain Hylodes to al- bumin antisera against Hylodes pulcher and Megae- losia goeldi from Maxson and Heyer (1982). Values are in immunological distance units. Anti- Anti- Megae- Hylodes losia Species tested pulcher goeldi Hylodes charadranaetes 37 49 Hylodes lateristrigatus— Teresopolis 40 56 Hylodes lateristrigatus— Santa Teresa 42 51 Hylodes phyllodes 60 49 Hylodes sp.—Brejo de Lapa 108 86 Hylodes sp.—Eugenio Lefévre 39 71 of Hylodes, most without specific names, based on immunological microcomplement fixation analysis of albumin. The data for the /ateristrigatus species cluster are re- peated (Table 2), with the names recognized in this paper. The specimen from Brejo da Lapa represents the undescribed new species from Itatiaia. The specimen from Eugénio Lefévre was not recorded and its morphol- ogy 1s not distinctive; it may be from a pop- ulation of H. magalhaesi, but recordings will probably be required for certain identifi- cation. The molecular data are consistent with our recognition of /ateristrigatus from both Teresopolis and Santa Teresa and sup- port the distinctiveness of the other species recognized in this paper. Acknowledgments Dr. Peter Weygoldt (Albert-Ludwig-Uni- versitat, Freiburg) kindly provided data on the Santa Teresa specimens that he collect- ed, including call recordings and color notes. Ronald I. Crombie provided assistance in suggesting names for the new species. George R. Zug provided a thorough review of the manuscript. Research for this paper was supported in part by the Museu de Zoologia da Univer- sidade de Sao Paulo (MZUSP); the Schol- arly Studies Program, Smithsonian Insti- 109 tution; and the International Environmental Science Neotropical Lowland Research Program, Smithsonian Institution. Literature Cited Baumann, F. 1912. Brasilianische Batrachier des Ber- ner Naturhistorischen Museums.—Zoologisch Jahrbucher (Systematik) 33:87—172, pls. 4-8. Bokermann, W. C. A. 1964. Una nueva especie de Elosia de la Serra da Mantiqueira, Brasil. — Neo- tropica 10(33):102-107. 1967a. Notas sObre cantos nupciais de an- fibios brasileiros. I: O canto de “Elosia lateri- strigata’”’ e “Elosia glabra” (Anura).— Revista Brasileira de Biologia 27(3):229-231. 1967b. Una nueva especie de Elosia de Ita- tiaia, Brasil (Amphibia, Leptodactylidae).— Neotropica 13(42):135-137. Cochran, D.M. 1955. Frogs of Southeastern Brazil. — United States National Museum Bulletin No. 206:1—423 + 34 plates. Gouvéa, E. 1979. Uma nova espécie de elosiineo da Serra do Itatiaia (Amphibia, Anura, Leptodac- tylidae).— Revista Brasileira de Biologia 39(4): 855-859. Heyer, W.R. 1982. Twonew species of the frog genus Hylodes from Capara6, Minas Gerais, Brasil (Amphibia: Leptodactylidae).— Proceedings of the Biological Society of Washington 95(2):377— 385. Izecksohn, E., and E. Gouvéa. 1983. Sobre a iden- tidade de Elosia glabra Miranda-Ribeiro (Am- phibia, Anura, Leptodactylidae).— Arquivos da Universidade Federal Rural do Rio de Janeiro 1983:221-223. Lutz, B. 1951. Nota prévia sobre alguns anfibios an- uros do Alto Itatiaia.—O Hospital 39(5):705- 707. Maxson, L.R.,and W.R. Heyer. 1982. Leptodactylid frogs and the Brasilian Shield: an old and con- tinuing adaptive relationship. — Biotropica 14(1): 10-15. Miranda-Ribeiro, A. 1926. Notas para servirem ao estudo das Gymnobatrachios (Anuros) Brasi- leiros-Tomo primero.—Archivos do Museu Nacional do Rio de Janeiro 27:1—227, 22 pls. Sazima, I., and W. C. A. Bokermann. 1982. Anfibios da Serra do Cipo, Minas Gerais, Brasil. 5: Hy- lodes otavioi sp. n. (Anura, Leptodactylidae). — Revista Brasileira de Biologia 42(4):767-771. Department of Vertebrate Zoology, Am- phibians and Reptiles, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 110-120 THE STATUS OF THE INDO-WEST PACIFIC CARDINALFISHES APOGON AROUBIENSIS AND A. NIGROFASCIATUS John E. Randall and Ernest A. Lachner Abstract. — Apogon aroubiensis Hombron and Jacquinot (1853) from Malay- sia was named from a single 6 cm specimen which is not extant. It was described and illustrated as having four closely spaced black stripes on the side of the body and 12 pectoral fin rays. This nominal species cannot be equated to any apogonid fish known today. The species of Apogon most authors have identified as A. aroubiensis is A. nigrofasciatus Lachner which is known from the Red Sea and the western Pacific to French Polynesia. It is characterized as follows: five broad uniform dark brown stripes on body, the midlateral stripe ending in a dark spot at caudal base; no narrow short stripe extending posteriorly from upper margin of orbit; pectoral rays 14; gill rakers 20-24; least bony width of interorbital space 5.25—6.0 in head; last anal-fin ray not elongate, its length 2.55—4.0 in head. Apogon nigrofasciatus has been confused with A. angustatus (Smith and Radcliffe), A. cookii Macleay, A. fasciatus (Shaw), A. novemfasciatus Cuvier, and A. taeniophorus Regan. These five valid dark-striped species are differentiated from it in our discussion. Apogon aroubiensis was described by Hombron and Jacquinot (1853:31, pl. 1, fig. 1) from a single specimen nearly 6 cm in length collected at Aroub, Malaysia. The color was given as silvery green with four black stripes; the fins were green with a broad brown border; the pectoral fin ray count was 12. Their illustration, reproduced herein as Fig. 1, shows the four black stripes closely spaced on the side of the body with no stripe dorsally or ventrally; very broad dark mar- gins are evident on the second dorsal, anal, caudal, and pelvic fins. Jordan and Seale (1906:241, fig. 35) re- corded a dark-striped species of cardinalfish from Apia and Pago Pago in the Samoa Is- lands as Amia aroubiensis. However, it 1s clearly different from the A. aroubiensis of Hombron and Jacquinot. The illustration of the Samoan fish shows five black stripes— three broad ones on the side and a narrow one dorsally and ventrally; there are no dark borders on any fins. Radcliffe (1911:250, pl. 22, upper figure) recorded 125 specimens of the same car- dinalfish as that of Jordan and Seale as Amia aroubiensis from 50 localities in the Phil- ippines and Celebes collected by the U.S. Bureau of Fisheries steamer A/batross in 1908 and 1909. He placed Apogon fasciatus, as identified in Fische der Stuidsee by Gin- ther (1873:19), in part, in the synonymy of A. aroubiensis, specifically citing the broad- striped “‘variety”’ of figure A of Ginther’s plate 20. He pointed out that the true Apo- gon fasciatus is the fish described by White (1790:268, fig. 1) (actually Shaw in White) from Port Jackson (now Sydney), New South Wales. Since Shaw’s type has not been found, Lachner (1953:439, pl. 35 A) designated a neotype (USNM 59972, 80.5 mm SL) col- lected at ‘““Port Jackson” by D. G. Stead. Radcliffe (1911:249) had already provided a description of this fish and illustrated it on pl. 22 (lower figure, though obviously retouched). VOLUME 99, NUMBER 1 Fig. 1. Gunther (1873) included Apogon fascia- tus Quoy and Gaimard (1825:344) in his synonymy; however, this is a secondary homonym of Apogon fasciatus (Shaw). We have examined the holotype of Quoy and Gaimard’s A. fasciatus (MNHN 853, 54mm SL); it is a specimen of A. novemfasciatus Cuvier (in Cuvier and Valenciennes, 1828). McCulloch (1915:118) reported two spec- imens as Amia fasciata aroubiensis from Murray Island, Torres Strait and one from Suva, Fiji. He was followed by Marshall (1964:127). Fowler (1928:157) used the name Amia aroubiensis for what he believed to be the same species as Jordan and Seale (1906), listing their reference in his synonymy, and recording Bishop Museum specimens from Tubuai, Nuku Hiva, and Raiatea. Weber and de Beaufort (1929:302) erro- neously placed A. aroubiensis in the syn- onymy of A. novemfasciatus Cuvier. Holotype of Apogon aroubiensis, Aroub, Malaysia (after Hombron and Jacquinot 1853). Fowler and Bean (1930:53) reported on the same material from the Philippines and East Indies as Radcliffe; they listed the 104 lots by Albatross collection number, local- ity, date, and gave the lengths of the spec- imens. In his treatment of the fishes of the Phoe- nix and Samoan Islands, Schultz (1943:95) cited the Jordan and Seale reference and one lot of their material as Apogon aroubiensis; he recorded an additional specimen from Tutuila. Lachner (1953:466) described Apogon ni- grofasciatus from 132 specimens in 26 lots from the Marshall Islands, seven specimens from Guam, five from Western Samoa, and one from Tubuai, Austral Islands. He dis- tinguished his new species from what he regarded as its closest relative, A. aroubien- sis, principally by its narrower dark stripes. Randall (1955:71) recorded 17 specimens of A. nigrofasciatus from Onotoa, Gilbert 112 Islands (Kiribati). Noting the similarity to A. aroubiensis as defined by Lachner, he wrote, “It is possible that these two forms are merely subspecies of one wide-ranging species.” In 1978 the senior author noticed the dis- crepancy in color pattern of the illustration and description of A. aroubiensis by Hom- bron and Jacquinot and the fish the above authors have identified as this species. He wrote M. L. Bauchot of the Muséum Na- tional d’ Histoire Naturelle to request a loan of the holotype of A. aroubiensis if extant. She sent a specimen of what was listed as the type, but it proved to be A. kallopterus Bleeker, a very different species of cardi- nalfish. Clearly an error had been made and another fish substituted for the type. Much searching by Bauchot and associates failed to yield the true type of A. aroubiensis, and it must be presumed lost. There is little chance that A. aroubiensis Hombron and Jacquinot is the same as the species later authors identified with this name. Not only is the color pattern differ- ent, but the pectoral-fin ray count of 12 giv- en by Hombron and Jacquinot does not match the count of 14 that is consistently found in A. nigrofasciatus. These authors gave a count of 12 for the pectoral-fin rays of another species of Apogon they described as new, Apogon nigromaculatus. This nom- inal species is now regarded as a synonym of Sphaeramia orbicularis (Cuvier) which has 12 pectoral-fin rays. It might also be men- tioned that the misnamed A. aroubiensis is a clear-water fish usually found on well-de- veloped coral reefs, hence a Malaysian lo- cality for it would not be expected. It there- fore seems more likely that the true A. aroubiensis is a valid species that has not been correctly reported since the original description in 1853. Nevertheless, we rec- ommend that it be regarded as a doubtful species until material which conforms to the original description and figure is found. Recent collections of fishes in the western Pacific have failed to yield any broad-striped PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON specimens of the species that has been mis- identified as A. aroubiensis, but numerous lots of A. nigrofasciatus have been taken from the Philippines, New Guinea, and In- donesia, largely by Victor G. Springer and associates of the National Museum of Nat- ural History. It would seem that A. nigro- fasciatus has completely replaced the once abundant 4. “aroubiensis.’’ A close exam- ination of the color pattern of recent ma- terial of A. nigrofasciatus, however, has re- vealed that the pale interspaces have a slight dark pigmentation in a zone next to the ad- jacent dark stripe on each side. We now conclude that specimens initially preserved in alcohol, as was the A/batross material from the Philippines and East Indies, maintain the full dark pigmentation of the stripes in preservative, whereas specimens placed in formalin largely lose the dark edges of the stripes, thus giving a pattern of dark and light bands of about equal width. We present below a description of A. ni- grofasciatus, followed by discussion of re- lated or confused species and distribution. Key characters to many of the species of the dark striped “‘A. fasciatus” group are given in Radcliffe (1911) and Lachner (1953). Proportional measurements based on more than 30 specimens are rounded to the near- est 0.05. Specimens examined are deposited in the Australian Museum, Sydney (AMS); British Museum (Natural History), London [BM(NH)}]; Bernice P. Bishop Museum, Ho- nolulu (BPBM); Muséum National d’His- toire Naturelle, Paris (MNHN); J. L. B. Smith Institute of Ichthyology, Grahams- town (RUSI); and National Museum of Natural History, Washington, D.C. (USNM). The lengths given of specimens in Material Examined are standard lengths (SL) in millimeters. Apogon nigrofasciatus Lachner Figs. 25.3 Apogon nigrofasciatus Lachner, in Schultz and collaborators, 1953:440, 446, fig. 81, VOLUME 99, NUMBER 1 113 Fig. 2. Apogon nigrofasciatus, BPBM 8003, 51 mm SL, Enewetak, Marshall Islands. pl. 37, D, E (type locality, Bikini, Mar- shall Islands). Description. —Dorsal-fin rays VII-I,9; anal-fin rays II,8; pectoral-fin rays 14 (rare- ly 13); lateral line complete, the pored scales 24 or 25 (usually 25) (3 additional pored scales posterior to base of hypural plate); gill rakers 20 to 24 (x = 21.6); mouth ter- minal, oblique, the maxilla reaching to or slightly beyond a vertical through posterior edge of pupil; a broad band of villiform teeth in jaws, and a narrow band on vomer and palatines; anterior preopercular margin smooth; posterior preopercular margin ser- rate (the edge with a series of close-set, small spinules); margin of subopercle smooth; body depth 2.7-3.1 in SL; head length 2.4— 2.55 in SL; least bony width of interorbital space 5.25-6.0 in head; first dorsal spine very small, about 20 to 35% length of sec- ond dorsal spine, which in turn is about 35 to 50% length of third dorsal spine; third dorsal spine 1.85-2.1 in head; first dorsal soft ray longest, 1.35—1.6 in head; last anal- fin ray not prolonged, its length 2.55—4.0 in SL; caudal fin slightly forked, lobes round- ed; pectoral fins 1.45—1.7 in head; pelvic fins reaching beyond anus, sometimes to or slightly posterior to origin of anal fin, lon- gest ray 1.6-1.75 in head. Color in alcohol: pale brown with five uniform dark brown stripes, first narrow, middorsal on nape and along base of dorsal fins, extending broadly onto base of second dorsal fin except for last ray; second dark stripe beginning narrowly on dorsal part of snout, passing along upper edge of eye, enclosing anterior part of lateral line, and ending at upper base of caudal fin; third dark stripe from side of snout, through eye, along center of side of body to mid- caudal base where it is slightly enlarged and darker, thus appearing as an elliptical spot within stripe; fourth stripe from side of maxilla, along lower edge of eye, enclosing pectoral base, and passing to lower caudal fin base; fifth stripe narrow (often faint in preservative), passing from mandible 114 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nat 4 A Galt Fig. 3. through pelvic fin base, along lower abdo- men, and ending as a band basally in anal fin; second to fourth dark stripes as broad or broader than pale interspaces (distinctly broader if initial preservative was alcohol); ends of second and fourth stripes may con- verge slightly toward posterior end of mid- lateral stripe on caudal base, or even join with it (but not extending out onto fin); some dusky pigment anteriorly and basally in spi- nous dorsal fin; dark stripe at base of second dorsal and anal fins, that of dorsal not set off from base by unpigmented band, except narrow clear zone on last one or two mem- branes (dark basal band of second dorsal and anal fins more heavily pigmented on mature males than females); faint dusky midlateral streak on caudal fin. In life dark stripes are dark brown to black, suffused with red, lower stripe and anterior part of third and fourth stripes often mainly red; Underwater photo of Apogon nigrofasciatus taken in the Red Sea off Jeddah, Saudi Arabia. slight enlargement and intensification of black pigment to form spot at caudal fin base at end of midlateral body stripe; another darker area within fourth dark stripe ante- rior to pectoral fin base; pale interspaces white, sometimes pale yellowish or greenish yellow; edges of fin spines and rays light red, membranes clear (except where noted above as dusky or black). Remarks. —Apogon nigrofasciatus is mainly a coral reef inhabitant. It has been taken in the depth range of 1 to 45 m. It rarely occurs in less than 2 meters, however. Based on available material, this species exhibits a disjunct distribution. It is found throughout the islands of Oceania except the Hawaiian Islands, Easter Island, the Pit- cairn Group, and the Marquesas. (The Bish- op Museum specimen from Nuku Hiva re- ported by Fowler, 1928, as Amia aroubiensis has been lost. Extensive rotenone collec- VOLUME 99, NUMBER 1 115 Table 1.—Gill-raker counts of four dark-striped species of Apogon. 17 A. angustatus A. cookii 17 A. nigrofasciatus, Marshall Islands Other Pacific localities Red Sea and Gulf of Aden A. taeniophorus 21 tions by the senior author and associates in these islands failed to yield any A. nigro- fasciatus.) In the western Pacific, A. nigro- fasciatus ranges from Wakayama Prefec- ture, Honshu (Masuda et al. 1975) to the Capricorn Group of the southern Great Bar- rier Reef (Russell 1983). It probably occurs throughout Indonesia and New Guinea. It has been reported (as Apogon sp.) by Allen and Steene (1979) from Christmas Island in the eastern Indian Ocean. It is unknown from the rest of the Indian Ocean but is present in the Red Sea where it is among the more common apogonids on coral reefs. The largest specimen of A. nigrofasciatus examined, USNM 142290, from Bikini, measures 73 mm SL. Lachner (1953:470) noted that A. nigro- fasciatus from islands in Oceania attains a larger average size than individuals from the East Indies and Philippines. This is still ap- parent from the additional material we have examined. He also found that the gill-raker counts were higher in the Marshall Islands than in the Philippines. Our additional counts reveal Marshall Island material typ- ically higher than all other Pacific localities combined, though the two sets of counts are broadly overlapping (Table 1). We found no difference between the counts of specimens from western Pacific localities and those of islands of Oceania other than the Marshalls, and have combined these two sets of data. Specimens of A. nigrofasciatus from the Red Sea also show higher gill-raker counts than those from Pacific localities other than the Marshall Islands. 18 19 20 21 22 23 24 14 26 6 1 32 33 3 4 18 21 3 7 36 48 17 1 3 18 27 6 43 41 Apogon nigrofasciatus has been confused by some authors with A. novemfasciatus Cuvier. The latter, however, is easily dis- tinguished by two sectors of its midlateral stripe which are darker and slightly en- larged—one behind the gill opening and one in the middle of the body. Also the second and fourth stripes converge toward the mid- lateral stripe well out on the caudal fin (see Jordan and Seale 1906: fig.36). The three dark-striped, Indo-Pacific species most often confused with A. nigro- fasciatus are: A. cookii Macleay (1881), of which A. melanotaenia Regan (1905) and A. robustus (Smith and Radcliffe in Rad- cliffe, 1911) are junior synonyms; and 4A. angustatus (Smith and Radcliffe in Rad- cliffe, 1911); and A. taeniophorus Regan (1908), of which A. fasciata stevensi (McCulloch, 1915) and A. saipanensis (Fowler, 1945) are junior synonyms. Fowler was unable to locat the type spec- imens of his Lovamia saipanensis for Lach- ner, so the latter regarded it as a valid species (Lachner 1953:439). Fraser (in Bohlke 1984), however, placed it in the synonymy of A. novemfasciatus. Except for a diagonal dark stripe on the abdomen (which Fowler attributed to the blackish peritoneum show- ing through the body wall), the color pattern of Fowler’s figure of A. saipanensis resem- bles A. taeniophorus, not A. novemfasciatus. The second and fourth dark stripes do not converge on the caudal fin and the midlat- eral stripe is uniform in width and pigmen- tation. We made counts of 14 pectoral rays and 18 for the gill rakers of the holotype of 116 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Apogon cookii, BPBM 20834, 75 mm SL, Gulf of Aqaba, Red Sea. A. saipanensis. On the basis of the color pattern and these counts we place A. sai- panensis in the synonymy of A. taeniopho- rus. Apogon cookii (Fig. 4) may be distin- guished from A. nigrofasciatus by nearly al- ways having 15 pectoral rays (Fraser 1974, Table 1), by its lower gill-raker counts (Ta- ble 1 herein), its broader interorbital space (bony width 4.15—5.1 in head, compared to 5.25-6.0 for A. nigrofasciatus), and in color. Adults have a narrow dark stripe extending posteriorly from the upper edge of the orbit, often to below the second dorsal fin. Also the dark band in the second dorsal fin is separated from the base by a narrow whitish band. Grant (1982: pl. 130) illustrated A. cookii in color (as A. novemfasciatus). This species occurs in very shallow water, gen- erally less than 2 m. Although usually found on rocky substrata, as under ledges in tide- pools, it may also be seen in seagrass beds or around small coral heads. Apogon taeniophorus (Fig. 5) has the same pectoral-ray count of 14 as A. nigrofascia- tus; however, it is completely separable by its low gill-raker counts (Table 1). It also has a broad interorbital space, the least bony width 4.5—5.35 in head. It is very similar in color to A. cookii, differing in lacking a dis- tinct dark spot in the midlateral stripe at the caudal fin base. Its spot, if it can be distinguished at all, is elliptical, only slightly broader than the stripe, and at best slightly darker. By contrast, the caudal base spot of A. cookii is distinctly larger than the stripe width, round or nearly round, and darker than the stripe. The narrow dark stripe ex- tending posteriorly from the upper orbit is not as well developed as on A. cookii. Apo- gon taeniophorus is also a species of shallow water of variable habitat from surge chan- VOLUME 99, NUMBER 1 117 Fig. 5. Apogon taeniophorus, BPBM 16045, 58 mm SL, Guadalcanal, Solomon Islands. nels of exposed reefs to rock or debris in mud-bottom harbors. Apogon angustatus (Fig. 6) usually has 14 pectoral rays. Its gill-raker counts are lower than those of A. nigrofasciatus but do not provide complete separation (Table 1). The last anal rays of this species are elongate, thus the distal border of the fin is distinctly concave; the length of the last anal ray is contained 1.75-2.65 times in the head length, compared to 2.55-3.8 for A. nigro- fasciatus. The dark stripes of A. angustatus are narrower than the whitish interspaces; they are blackish at Indian Ocean localities and yellowish brown at islands of Oceania (on these Pacific fish the black spot at the caudal fin base is therefore more conspic- uous). Apogon angustatus has been collect- ed on coral reefs in the depth range of 6 to 40 meters. Mention should be made of Apogon fas- ciatus (Shaw) since some authors have placed A. aroubiensis in the synonymy of, or as a subspecies of this species. Apogon fasciatus is now regarded as a senior synonym of A. quadrifasciatus Cuvier. It has the short post- ocular dark stripe from the upper edge of the orbit asin A. cookii and A. taeniophorus, but lacks a well developed dark stripe below the midlateral stripe (though this was mis- takenly added to the illustration in Radcliffe 1911: pl. 22). The anus is black. The gill rakers range from 18-22; the neotype has Moe Material examined.—Apogon angusta- tus: In addition to the holotype of Amia angustata Smith and Radcliffe, we have ex- amined USNM material from Borneo and the Trobriand Islands, and two MNHN and 40 BPBM specimens, 21-78 mm, from the Pitcairn Group, Tuamotu Archipelago, So- ciety Islands, Line Islands, Palau Islands, New Caledonia, New Guinea, Seychelles, and Mauritius. Also we have examined ten RUSI specimens from Natal, 23-63 mm. Apogon cookii: We have examined the syntypes of Apogon cookii Macleay (AMS 1.16307-001, 6: 35.5—63) from the Endeav- 118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Apogon angustatus, BPBM 6946, 71 mm SL, Tahiti, Society Islands. or River, northern Queensland and the ho- lotype of Apogon robustus (Smith and Rad- cliffe) from Jolo, Philippines. Also 100 BPBM and USNM specimens from the fol- lowing major localities: New Caledonia, Great Barrier Reef, Taiwan, Western Aus- tralia, SE India, Seychelles, Natal, Tanza- nia, Kenya, and the Red Sea. Apogon nigrofasciatus: In addition to the type specimens as listed by Lachner (1953) and the USNM material given by A/batross field numbers in Fowler and Bean (1930), we have examined specimens from the fol- lowing localities: TUAMOTU ARCHI- PELAGO: Mangareva, BPBM 13558, 2: 49-— 56.8 mm. Rangiroa, BPBM 10258, 48 mm; BPBM 25224, 57.2 mm. SOCIETY IS- LANDS: Tahiti, BPBM 10283, 2: 26—54.2 mm. Moorea, MNHN 1984-118, 3: 32.2- 52.5 mm; BPBM 11969, 7: 40.5—53 mm. Raiatea, BPBM 1674, 48 mm; BPBM 1675, 46 mm. RAPA: BPBM 17268, 2: 55.7-69.6 mm. AUSTRAL ISLANDS: Tubuai, BPBM 794, 65 mm; BPBM 795, 62 mm. COOK ISLANDS: Aitutaki, BPBM 5612, 11: 33- 52.1 mm. FIJI: Viti Levu, USNM 176632, 2: 54-59 mm. VANUATU (NEW HEB- RIDES): Espiritu Santo, USNM 262508, 2: 37-47 mm. Efate, BPBM 5615, 55.5 mm. LOYALTY ISLANDS: Uvéa, BPBM 27071, 61.4 mm. NEW CALEDONIA: BPBM 11466, 46.2 mm. GREAT BAR- RIER REEF: Hook Island, BPBM 15556, 4: 33.4-51 mm. Yonge Reef, MNHN 1978- 588, 51 mm. SOLOMON ISLANDS: Guadalcanal, BPBM 5695, 41 mm; BPBM 15578, 2: 29-34.2 mm; BPBM 16157, 3: 32.5-41 mm; USNM 262511, 46 mm. NEW IRELAND: USNM uncat., 17: 32-48 mm. NEW GUINEA: Bagabag Island, USNM 262709, 31: 18-52 mm. Massas Island, BPBM 262612, 6: 19—48.5 mm. Louisade Archipelago, USNM 262710, 36: 30-59 mm. Madang, BPBM 15761, 44.2 mm. Ni- nigo Islands, BPBM 30345, 29: 31.2-46.8 mm; USNM 262426, 15: 16-49 mm. Her- mit Islands, USNM 261037, 32: 20—29.5 mm. INDONESIA: Ambon, BPBM 19312, 3: 35-53.7 mm. Banda Islands, USNM 262706, 13: 29-42 mm. Kai Islands, USNM VOLUME 99, NUMBER 1 262704, 12: 30-47 mm. Bali, BPBM 30181, 3: 42.5—49 mm. PHILIPPINES: Siquijor Is- land, BPBM 30210, 29: 31.2-46.8 mm; USNM 260967, 75: 15—49 mm. Apo Island, BPBM 30344, 24: 28.7-48 mm; USNM 262375, 3: 42-51.5 mm. Palawan, USNM 262369, 17: 38-46 mm. Negros, BPBM 28585, 2: 49-51.5 mm; BPBM 30343, 67: 29-48.9 mm; USNM 262366, 16: 13.5—48 mm. Balicasag Island, USNM 260963, 19: 32-51 mm. Cebu, USNM 262383, 15: 13.5— 47 mm. Mactan Island, USNM 262374, 10: 24-48 mm. TAIWAN: BPBM 23302, 33 mm; BPBM 23369, 36.5 mm. RYUKYU ISLANDS: Ishigaki, BPBM 8697, 5: 45-64.5 mm. PALAU ISLANDS (BELAU): BPBM 7423, 60.6 mm; BPBM 9808, 2: 44-47 mm. CAROLINE ISLANDS: Truk, BPBM 9062, 42.2 mm. MARSHALL ISLANDS: Enewe- tak, BPBM 8003, 5: 49-63 mm; BPBM 8220, 44.2 mm; BPBM 8254, 2: 56—-56.2 mm; BPBM 8256, 4: 44.5-59.3 mm; BPBM 8288, 54.5 mm; BPBM 29142, 15: 38.5-71 mm. GULF OF ADEN: MNHN 1977-639, 4: 48.5-52 mm; MNHN 1977-642, 57 mm; MNHN 1977-643, 54 mm. RED SEA: MNHN 1952-95, 52 mm. Gulf of Aqaba, BPBM 13397, 3: 35-52.2 mm; BPBM 18370, 3: 51.3-65 mm; MNHN 1977-815, 62 mm; MNHN 1977-816, 52.5 mm; USNM 212777, 12: 37.7-61 mm; USNM 212782, 21: 17.0-62 mm; USNM 213612: 35 mm. Apogon taeniophorus: We have examined the syntypes of A. taeniophorus Regan from the Maldives BM(NH) 1908.3.23.90-92, 3: 43-77 mm SL, and the holotype of A. sai- panensis (Fowler) from the Marianas (ANSP 71588, 22 mm SL). In addition, we have seen specimens listed by Lachner (1953) from the Marshall Islands and Marianas as Apogon robustus, and 128 BPBM speci- mens, 19—92.5 mm, from the following ma- jor localities: Pitcairn Group, Society Is- lands, Austral Islands, Rapa, Cook Islands, Line Islands, Gilbert Islands (Kiribati), Marshall Islands, Minami Tori Shima (Marcus Island), Taiwan, Indonesia, Fiji, 119 Solomon Islands, New Britain, SW Thai- land, SW India, Seychelles, Tanzania, Na- tal, Gulf of Aden, and the Red Sea. Acknowledgments We are grateful to Ofer Gon (RUSID and Douglass F. Hoese (AMS) for the loan of specimens and advice on species identifi- cation, and to John R. Paxton (AMS), A.C. Wheeler [BM(NH)], and Susan L. Jewett (USNM) for the loan of specimens. Thomas H. Fraser reviewed the manuscript and made valuable suggestions for its improvement. Literature Cited Allen, G. R., and R. C. Steene. 1979. The fishes of Christmas Island, Indian Ocean.— Australian National Parks and Wildlife Service, Canberra, Special Publication 2:1-81. Bohlke, E. B. 1984. Catalog of type specimens in the Ichthyological collection of the Academy of Natural Sciences of Philadelphia.—The Acad- emy of Natural Sciences of Philadelphia, Special Publication 14:vii + 246 pp. Cuvier, G., and A. Valenciennes. 1828. Histoire Na- turelle des Poissons. F. G. Levrault, Paris, xxi + 490 pp. Fowler, H. W. 1928. The fishes of Oceania.—Mem- oirs of the Bernice P. Bishop Museum 10:1ii1 + 340 pp. . 1945. Fishes from Saipan Island, Microne- sia.— Proceedings of the Academy of Natural Sciences of Philadelphia 97:59-74. , and B. A. Bean. 1930. Contributions to the biology of the Philippine Archipelago and ad- jacent regions.—Bulletin of the United States National Museum 100 (10):ix, 1-334. Fraser, T. H. 1974. Redescription of the cardinal fish Apogon endekataenia Bleeker (Apogonidae), with comments on previous usage of the name. — Proceedings of the Biological Society of Wash- ington 87(1):3-10. Grant, E.M. 1982. Guide to fishes. The Department of Harbours and Marine, Brisbane, Queensland, 896 pp. Giinther, A. C. L. G. 1873-1875. Andrew Garrett’s Fische der Stidsee.— Journal des Museum God- effroy 1:1-128. Hombron, J. B., and H. Jacquinot. 1853. Poissons. In Dumont d’Urville, J. Voyage au Pole Sud et dans l’Oceanie sur les corvettes “L’Astrolabe” et “La Zelee.”’ Gide et J. Baudry, Paris, volume 3, part 2:31—56. 120 Jordan, D. S., and A. Seale. 1906. The fishes of Sa- moa.— Bulletin of the Bureau of Fisheries (1905) 25:173-488. Lachner, E. A. 1953. Family Apogonidae: cardinal fishes. Jn Schultz and collaborators, Fishes of the Marshall and Marianas Islands. — Bulletin of the United States National Museum 202(1):xxxii, 1-685. Macleay, W. 1881. Descriptive catalogue of the fishes of Australia.— Proceedings of the Linnean So- ciety of New South Wales (1880) 5:302—444. Marshall, T.C. 1964. Fishes of the Great Barrier Reef and coastal waters of Queensland. Angus and Robertson, Sydney, 566 pp. Masuda, H., C. Araga, and T. Yoshino. 1975. Coastal fishes of Southern Japan. Tokai University Press, Tokyo, 382 pp. McCulloch, A. R. 1915. Biological results of the fish- ing experiments carried out by the F.I.S. ““En- deavour”’ 1909-14.— Department of Trade and Customs, Commonwealth of Australia, Report on the fishes 3(3):97-170. Quoy, J. R. C., and P. Gaimard. 1824-1826. Voyage autour du monde ... exécuté sur les corvettes de L.M. /’Uranie et la Physcienne, pendant les années 1817, 1818, 1819 et 1820; . . . Pillet Ainé, Paris, 712 pp. Radcliffe, L. 1911. Notes on some fishes of the genus Amia, family of Cheilodipteridae, with descrip- tions of four new species from the Philippine Islands. — Proceedings of the United States Na- tional Museum 41(1853):245-261. Randall, J. E. 1955. Fishes of the Gilbert Islands. — Atoll Research Bulletin 47:xi, 1-243. Regan, C. T. 1905. On fishes from the Persian Gulf, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the Sea of Oman, and Karachi, collected by Mr. F. W. Townsend.—The Journal of the Bombay Natural History Society 16:318-333. . 1907-1909. No. XIV. Report on the marine fishes collected by Mr. J. Stanley Gardiner in the Indian Ocean. — Transactions of the Linnean Society of London, 2nd Series, Zoology 12:217— 255. Russell, B. C. 1983. Annotated checklist of the coral reef fishes in the Capricorn-Bunker group Great Barrier Reef Australia. — Great Barrier Reef Ma- rine Park Authority, Queensland, Special Pub- lication Series 1:1-184. Schultz, L. P. 1943. Fishes of the Phoenix and Sa- moan Islands collected in 1939 during the ex- pedition of the U.S.S. “Bushnell.” — Bulletin of the United States National Museum 180:x, 1-— 316. Weber, M., and L. F. de Beaufort. 1929. The Fishes of the Indo-Australian Archipelago. 5. Anacan- thini, Allotriognathi. . . Percomorphi: Families: Kuhliidae . . . Centropomidae. E. J. Brill Ltd., Leiden, xiv + 458 pp. White, J. 1790. Journal of a voyage to New South Wales. Angus and Robertson, Sydney, xiii + 282 pp. (Reprint edition, 1962). (JER) Division of Ichthyology, Bernice Pauahi Bishop Museum, P.O. Box 19000- A, Honolulu, Hawaii 96817-0916; (EAL) Department of Vertebrate Zoology (Fishes), National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 121-136 HADOCERAS TAYLORI, A NEW GENUS AND SPECIES OF PHREATIC HYDROBIIDAE (GASTROPODA: RISSOACEA) FROM SOUTH-CENTRAL TEXAS Robert Hershler and Glenn Longley Abstract. — Hadoceras taylori, a new genus and species of Hydrobiidae, Lit- toridininae, is described from three localities in Real County, Texas. This phreatic species may well range throughout the southern portion of the Edwards Plateau and into northern Mexico. The detailed morphological description provided for this species includes aspects of the shell, operculum, pallial cavity, digestive system, and reproductive system of both sexes. Generic separation of Hadoceras from similar-shelled late Tertiary Orygoceras is warranted by both morphological and ecological considerations. A suite of unique characters, including the uncoiled, horn-like shell and concentric operculum with ventral process, separates Hadoceras from other known phreatic littoridinines of south- central Texas and northern Mexico. Taylor’s (1974) discovery of living phreatic snails from Roaring Springs, Texas referable to Orygoceras Brusina, 1882, a ge- nus previously known only as late Tertiary fossils from Idaho and southeastern Europe, ranks as a particularly exciting event in the recent history of freshwater malacology. The uncoiled, horn-like shell of Orygoceras is highly unusual among gastropods (Rex and Boss 1976) and the systematic placement of this genus has long been debated (Taylor 1974). Taylor (1974) concluded that living Orygoceras is a hydrobiid (Prosobranchia: Rissoacea), but did not describe this species. His morphological study was necessarily limited as he collected a single live speci- men, and various aspects of anatomy, in- cluding details of the male and female re- productive system critical to the systematic assessment of rissoacean snails, were not dealt with. As part of an ongoing survey and system- atic study of phreatic gastropods of south- central Texas (see Hershler and Longley, 1986) we collected more than 20 live Ory- goceras sp. (including a number of adults) from Roaring Springs and two other springs in Real County, Texas. We describe this snail as Hadoceras taylori, a new genus and species of Hydrobiidae; provide a detailed morphological description of this taxon, in- cluding aspects of the male and female re- productive systems; and discuss the system- atic relationships of Hadoceras, emphasizing comparisons with other members of the di- verse phreatic hydrobiid fauna of souih- central Texas and northern Mexico. Localities.—Hadoceras taylori was col- lected during May 22-24, 1985 from the following three springs (see Fig. 1): a) Roar- ing Springs (unnamed on USGS topograph- ic sheets), about 7.6 miles W of Camp Wood (water temperature 21.0°; conductivity 319.2 ymHOS; 5/23); b) Unnamed spring in South Prong Canyon, 6.6 miles W of Camp Wood (temperature 21.0°; conductivity 415.0 ymHOS; 5/23); and c) Unnamed spring at Jo Jan Van Camp, 9 miles N of Vance. All three springs are moderate-sized rheo- crenes, with numerous orifices. They occur in the Nueces River drainage in the region where the southern edge of the Edwards Pla- 122 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON EDWARDS (Balcones Fault Zone) AQUIFER REGION AUSTIN SUBREGION SAN ANTONIO SUBREGION 1 3 Steoravace ——_e——— Fig. 1. HTDROSEOLOSIC FEATURES ~ .\ i 3 4 \ ‘ ee ef + vas ‘ i A‘ ie . ’ ‘ d ; ca ' <<) : 7 / \W \ aa » Lge f i fj Le’ ; ~ Peery \ ps j | » / ¥ Fig. 8. Scanning electron micrographs of radulae of H. taylori. A—-C, Sections of the radular ribbon; D, Close- up of central teeth. Fig. 9. Scanning electron micrographs of radulae of H. taylori. A, Close-up of central teeth; B, Close-up of lateral, inner marginal and outer marginal teeth. Table 3. A phenogram, based on simple av- eraging of percent differences among generic pairs, is shown in Fig. 12. The most similar pair, linking at 33% dif- ference, are Paludiscala and Coahuilix, both endemic to the Cuatro Cienegas Basin, Mexico. The similarity between these gen- era extends to details of the anterior portion of the female reproductive system, with both taxa having a large-sized bursa (but no sem- inal receptacle) ventral to the pallial ovi- duct, and a sperm duct travelling from the right side of the bursa to enter the posterior portion of the albumen gland. While differ- ing in features involving shell form (Coa- huilix, planispiral; Paludiscala, elongate- conic), nature of the openings of the sper- mathecal duct and capsule gland (separate PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON vs. fused), and site of sperm storage (albu- men gland proper vs. albumen gland pouch), the two taxa are probably part of the same local radiation and perhaps should even be considered congeneric. Stygopyrgus and Balconorbis, both occurring in the Edwards (Balcones Fault Zone) Aquifer in south-cen- tral Texas, link at 48% difference and have rather dissimilar groundplans of the ante- rior portion of the female reproductive sys- tem. We have speculated (Hershler and Longley 1986) that Balconorbis may be- long to the Paludiscala-Coahuilix group, despite differences in characters such as pro- toconch microsculpture type, as its arrange- ment of the anterior portion of the female reproductive system is derivable (involving loss of the bursa) from that of the latter group. In Stygopyrgus the oviduct loops on the left side of the albumen gland, and enters the anterior end of this gland, with the sper- mathecal duct arising from this juncture and then receiving the duct of the seminal re- ceptacle well into the pallial cavity. This unique arrangement, coupled with the mammiform penial glands seen in this ge- nus (versus bulbous apocrine glands in the above three genera), suggests that Stygo- pyrgus belongs to a second radiation of phreatic littoridinines. Hadoceras differs from the other four gen- era by 57-68% of the characters considered and has a number of unique character-states, including the horn-like shell, concentric operculum with ventral process, intestinal loop on the right side of style sac, lack of penial lobes, and ventral displacement of the pallial gonoducts. The anterior portion of its female reproductive system does not closely resemble the arrangements seen in either of the other genera considered: note the unusual nature of the albumen gland, which is slender, non-glandular, and loops around the periphery of the bursa, and the elongate sperm duct, which joins the sper- mathecal duct well into the pallial cavity. Given the diversity of unique character- states seen in Hadoceras, we conclude that VOLUME 99, NUMBER 1 133 A Ast 0.50 mm Vd1 0.25 mm 0.125 mm Fig. 10. Stomach (A), penis (B), and prostate (C) of H. taylori. The right aspect is shown in A and B, and the ventro-right lateral aspect shown in C. The dark, curving line in C indicates the posterior end of the pallial cavity. Ast, anterior stomach chamber; In, intestine; Odg, opening of digestive gland; Oes, oesophagus; Pr, prostate; Pst, posterior stomach chamber; Sts, style sac; Vd, vas deferens; Vd1, posterior vas deferens; Vd2, anterior vas deferens. Sdu Oov 0.50 mm Fig. 11. Left lateral aspect of the anterior portion of the female reproductive system. The posterior portion of the intestine (In) has been removed. The thick curving line indicates the posterior end of the pallial cavity while the dashed line (to the left) indicates the mantle edge. Ag, albumen gland; Bu, bursa copulatrix; Cg, capsule gland; Cga, common opening of the capsule gland and spermathecal duct; In, intestine; Oov, opening of the oviduct into the albumen gland; Ov, oviduct; Osdu, opening of the sperm duct into the spermathecal duct; Sdu, sperm duct; Sd, spermathecal duct. 134 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.—Comparison of phreatic littoridinine gen- era involving 21 characters. B = Balconorbis, S = Sty- gopyrgus, P = Paludiscala, C = Coahuilix, and H = Hadoceras. Data for genera other than Hadoceras are from Hershler (1985) and Hershler and Longley (1986). Genus Character WS og) aa» en nisGa Shell 1. Maximum dimension >2.0 mm (0, 1) 0 O 1 Ora 2. Shell form: 0 1 1 Q » a) planispiral (0) b) elongate-conic (1) c) horn-like (2) 3. Protoconch micro- sculpture: One! 1 Leth a) spiral lines (0) b) punctate (1) 4. Teleoconch sculpture: OMON 21622) 2 a) spiral lines (0) b) collabral costae (1) c) absent (2) Operculum 5. Operculum concentric (0, 1) Yo @ @ @ i 6. Operculum with ven- tral process (0, 1) 0 0 @ @ I Nonreproductive anatomy 7. Ctenidium present (0, 1) oY @O it Ol © 8. Intestinal loop on style sac (0, 1) 0 OO @. @ 1 9. Intestinal loop in pal- lial cavity roof (0, 1) ill 0 Iu l Reproductive morphology 10. Pallial gonoducts dis- placed ventrally (0, 1) YO @ @ il 11. Number of penial lobes: I. 2 1 iL @) a) 0 (0) b) 1 (1) c) 2 (2) 12. Position of lobe(s): OL IO 20K = a) outer curvature of penis (0) b) inner curvature (1) 13. Penial gland type: oO tO MO = a) apocrine (0) b) mammiform (1) Table 3.—Continued. Genus Character B S P (C H 14. Anterior coil of oviduct: 0 1, 22a a) ventral to pallial oviduct (0) b) on left side of pallial oviduct (1) c) absent (2) 15. Oviduct opens into: 0 2 1 1 O a) posterior tip of al- bumen gland (0) b) posterior section of albumen gland (1) c) anterior end of al- bumen gland (2) 16. Albumen gland with posterior loop (0, 1) Lo © @ il 17. Bursa copulatrix present (0, 1) 0 0 1 I <4 18. Seminal receptacle present (0, 1) ot OF @ @ 19. Openings of sper- mathecal duct and capsule gland: 0 0 a) fused (0) b) separate (1) 20. Number of capsule gland tissue sections: Oo Oe tk O. a) 2 (0) b) 3 (1) 21. Capsule gland opening muscularized (0, 1) 0 1 0 1 O * Secondarily-derived seminal receptacle present. this genus represents yet another separate phreatic invasion within the Littoridininae. Epigean littoridinines having either a sim- ple penis, or a penis having mammiform or apocrine glands are known from Texas and northern Mexico (see Hershler 1985) and represent possible ancestors of the three phreatic radiations mentioned above. Fur- ther study of the diverse epigean littoridine fauna of the region will be necessary, how- ever, before the above phylogenetic specu- lations can be tested. VOLUME 99, NUMBER 1 Paludiscala Balconorbis Stygopyrgus Hadoceras 0.9 0.7 0.5 0.3 0.1 Fig. 12. Phenogram showing similarities among Hadoceras and other phreatic littoridinines from Texas and northeastern Mexico. Data used to generate the phenogram are given in Table 3. Acknowledgments We thank the owners of the Roaring Springs Ranch, Maly Ranch, and property at Jo Jan Van Camp for permission to sam- ple springs on their property. Mr. Ivan Woodard, of the Roaring Springs Ranch, was especially cooperative and even al- lowed the senior author to camp by the springs. Mr. Kevin Kuehn helped with the fieldwork. The senior author thanks W. Pratt for sharing both his ideas concerning the systematic position of Orygoceras and al- lowing us to cite his exciting discovery of Hadoceras sp. from a Nevada archaeolog- ical site. J. Koke and K. Auffenburg assisted with the scanning electron microscopy. The senior author acknowledges the receipt of a generous contract from Southwest Texas State University (Account # 29665), with- out which the project could not have been completed. Other logistic support to the se- nior author was provided by the Edwards Aquifer Research and Data Center (SWTSU) and the Florida State Museum. Literature Cited Ashworth, J. B. 1983 Ground-water availability of the Lower Cretaceous formations in the Hill 135 Country of south-central Texas.—Texas De- partment of Water Resources, Report 273:1- 172. Brune, G. 1975. Major and historical springs of Tex- as.— Texas Water Development Board, Report 189:1-94. Brusina, S. 1882. Orygoceras, eine neue Gastropo- dengattung der Melanopsidenmergel Dalma- tiens.— Beitrage Palaontologie Oesterreich-Un- gorns und des Orients 2:33-46. Dall, W. H. 1924. Discovery of a Balkan fresh-water fauna in the Idaho Formation of Snake River Valley, Idaho.— USGS Professional Paper 132G: 109-115. Davis, G. M. 1979. The origin and evolution of the gastropod family Pomatiopsidae, with emphasis on the Mekong River triculinae.—Academy of Natural Sciences of Philadelphia, Monograph 20:1-120. ——, M. Mazurkiewicz, and M. Mandracchia. 1982. Spurwinkia: Morphology, systematics, ecology of a new genus of North American marshland Hydrobiidae.— Proceedings of the Academy of Natural Sciences of Philadelphia 134:143-177. Davis, J. R. 1983. An additional record of living Orygoceras (Hydrobiidae) from Texas.—Nau- tilus 97:112-113. Fretter, V., and A. Graham. 1962. British Proso- branch Molluscs. Ray Society, London, xvi + 755 pp. Hershler, R. 1985. Systematic revision of the Hy- drobiidae (Gastropoda: Rissoacea) of the Cua- tro Cienegas Basin, Mexico.—Malacologia 26: 31-123. , and G. Longley. 1986. Phreatic hydrobiids (Gastropoda: Prosobranchia) from the Edwards (Balcones Fault Zone) Aquifer Region, south- central Texas.— Malacologia 27:127-172. Longley, G. 1981. The Edwards Aquifer: Earth’s most diverse groundwater ecosystem? — International Journal of Speleology 11:123-128. Papp, A. 1962. Die systematische Stellung von Or- ygoceras Brusina und aberrante Forme von Gy- raulus aus dem Steinheimer Becken.— Archiv far Molluskenkunde 91:203-—206. Rex, M. A., and K. J. Boss. 1976. Open coiling in recent gastropods. — Malacologia 15:289-297. Taylor, D. W. 1966. Summary of North American Blancan nonmarine mollusks.—Malacologia 4: 1-172. 1974. The tertiary gastropod Orygoceras found living. — Archiv fur Molluskenkunde 104: 93-96. Wenz, W. 1938-44. Gastropoda, Allgemeiner Teil und Prosobranchia. Jn O. H. Schindewolf, ed., 136 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Handbuch der Palaozoologie, 6. Borntraeger, Berlin, 1639 pp. Yen, T.-C. 1944. Notes on freshwater mollusks of Idaho Formation at Hammett, Idaho.—Journal of Paleontology 18:101-108. (RH) Department of Invertebrate Zool- ogy, Smithsonian Institution, Washington, D.C. 20560; (GL) Edwards Aquifer Re- search and Data Center, Southwest Texas State University, San Marcos, Texas 78666. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 137-139 A NEW SPECIES OF HELMINTHOGLYPTA (GASTROPODA: PULMONATA: HELMINTHOGLYPTIDAE) FROM SAN DIEGO COUNTY, CALIFORNIA Richard L. Reeder and Walter B. Miller Abstract. —Helminthoglypta montezuma is described from California. The species differs from its nearest consubgeners in having densely granulose shell sculpture. Relationships within the genus are discussed. Henry A. Pilsbry’s magnificent compila- tion of information on the western land snail genus Helminthoglypta Ancey, 1887 (Pils- bry 1939), has remained the most complete and authoritative reference to date for workers studying this genus. Anatomical data are lacking for many species and sub- species listed, however, and their systematic position may therefore be incorrect. During the period 1956-1964, the late W. O. Gregg and the second author conducted a series of field explorations in southern California for the express purpose of collecting from known populations of Helminthoglypta in the re- gion, and using the material for compara- tive morphological study. In the process, new populations were discovered and some were found to be undescribed species. One such new species from Montezuma Valley in San Diego County is described below. Helminthogyptidae Pilsbry, 1939 Helminthoglypta Pilsbry, 1939 Helminthoglypta montezuma, new species Figs. 1, 2 Description of holotype. —Shell small, de- pressed, helicoid, umbilicate, with umbili- cus contained about 8 times in diameter of shell. Color light brown with pale, chestnut band on rounded shoulder. Aperture nearly round, with peristome thickened, only slightly reflected, expanded slightly more at columellar insertion. Embryonic shell of 1' whorls, smooth. Post-embryonic whorls with radial growth wrinkles, covered with such dense numbers of minute papillae as to appear granulose. Papillae tend to be in descending spiral rows on second and third whorls, randomly distributed on later whorls and continuing strongly on to base of shell and into umbilicus. Diameter 17.4 mm, height 10.1 mm, diameter of umbilicus 2.2 mm, number of whorls 4%. Reproductive anatomy of holotype. —The genital system is typical of the genus, with an atrial sac having a dart sac at its proximal end, the latter structure being relatively small. There are two mucous glands with mucous bulbs, the ducts of which unite to form a single duct before entering the atrial sac. The spermatheca is spherical and the spermathecal duct bears a diverticulum of moderate length. The penis has a short, nar- row lower chamber and a longer, wider, double-walled upper chamber of uniform diameter. The lumen of the upper penis ex- pands broadly in its lower half. The penis forms a continuous tube with the epiphallus which is of moderate length and bears an epipallic caecum at its proximal end, also of moderate length. The penial retractor muscle attaches to the epiphallus. Measure- ments of distinctive organs are as follows: Penis 11.4 mm Epiphallus 12.3 mm Epiphallic caecum 9.1 mm Spermathecal duct 24.0 mm Spermathecal diverticulum 18.6 mm 138 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. views. Diameter 17.4 mm. Variation in paratypes.—A total of 19 adult and 15 juvenile shells was examined. The largest adult paratype is 19.6 mm in diameter and 11.0 mm in height, and the smallest 14.2 mm and 8.1 mm respectively. All of the shells exhibit the characteristic granulose condition and narrow umbilicus. Five reproductive anatomies were exam- ined; all exhibited the same cylindrical shape of uniform diameter, and the same upper and lower penis. Disposition of type material. —Holotype: Santa Barbara Museum of Natural History no. 33917. Paratypes: The Academy of Nat- ural Sciences of Philadelphia no. 359265; U.S.N.M. no. 842310; W. B. Miller collec- tion nos. 4302, 4306 and 7490; R. L. Reeder collection no. 685; H. L. Fairbanks collec- tion no. 459. Type locality. —San Diego County, Cali- fornia; in rocks among oaks along north side of Montezuma Valley Road from 1.0-2.0 road miles east of junction with San Felipe Shell of holotype of Helminthoglypta montezuma (SBMNH 33917). Dorsal, apertural, and umbilical Road; 33°13.0'N, 1040 m (3440 ft.). Discussion. —The anatomy of H. monte- zuma clearly indicates that the species be- longs in the nominate subgenus. Its nearest consubgeneric relatives are H. thermimon- tis Berry, 1953, H. waltoni Gregg and Mil- ler, 1976, and H. milleri Reeder, 1985. It differs from all three in that it has densely granulose shell sculpture while the others are only moderately papillose. It is also con- sistently smaller than the other species, with a maximum diameter ranging from 14.2 to 19.6 mm while the smallest of the others is 21.8 mm for H. thermimontis. Its umbili- cus, likewise, is narrower, ranging from 2.1 to 2.8 mm, while the others range from 3.2 to 4.3 mm. Its anatomy, while similar to that of the other three, does have certain distinctive features involving the shape of the penis. In H. waltoni the upper penis is decidedly club-shaped. In H. thermimontis, the upper penis is cylindrical for most of its 116°35.5'W; elevation VOLUME 99, NUMBER 1 10 mm Fig. 2. Portion of reproductive system of Hel- minthoglypta montezuma Reeder and Miller, showing diagnostic characters. Drawing made from projection of stained whole mount, #4302-DT; arrow shows junc- tion of upper penis and epiphallus. as—atrial sac; ds— dart sac; ec—epiphallic caecum; ep—epiphallus; go— genital orifice; lp>—lower part of penis; mb—mucous gland bulbs; mg— mucous gland membranes; ov—ovi- duct; pr—penial retractor muscle; pt—prostate; sd— spermathecal duct; so—spermatheca; sv —spermathe- cal diverticulum; up—upper part of penis; ut—uterus; va—vagina; vd—vas deferens. length and then tapers to the smaller initial diameter of the venturi-shaped lower penis; the venturi portion is extremely narrow. In H. montezuma, the upper penis is cylindri- 139 cal for all of its length, joining a lower penis of equal initial diameter. In H. milleri, the upper and lower penes have essentially the same characteristics as in H. montezuma except that the venturi portion is consis- tently wider. Helminthoglypta montezuma is currently known only from the type locality in Mon- tezuma Valley. Vegetation at this locality consists primarily of Quercus dumosa, Quercus agrifolia, Cercocarpus betuloides, Rhus ovata, Artemesia tridentata, Photinia arbutifolia, Adenostoma fasciculatum, Ber- beris pinnata, and Symphoricarpos sp. Etymology. —This species is named for Montezuma Valley, California, where it lives. Acknowledgments We are indebted to the late Wendell O. Gregg for providing some of the specimens and information used in preparation of this description, and to Susan J. McKee for pho- tographs and excellent assistance in the lab- oratory. Thanks also to The University of Tulsa for support of field work and to The University of Arizona for providing labo- ratory facilities. Literature Cited Pilsbry, Henry A. 1939. Land mollusca of North America (north of Mexico).— Academy of Nat- ural Sciences of Philadelphia Monographs (3)I(1): i-xvili + 1-573 + i-ix; figs. A, B, 1-377. (RLR) Faculty of Biological Science, Uni- versity of Tulsa, Tulsa, Oklahoma 74104; (WBM) Department of Ecology and Evo- lutionary Biology, The University of Ari- zona, Tucson, Arizona 85721. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 140-148 TWO SPECIES OF UROCOPIA, PLANKTONIC POECILOSTOMATOID COPEPODS OF THE FAMILY UROCOPIIDAE HUMES AND STOCK, 1972 Gayle A. Heron and David M. Damkaer Abstract. —Since the original description of the bathypelagic copepod Uro- copia singularis Sars, 1917, there have been only two other records of this species. New material from the eastern North Pacific Ocean has led to additional morphological details of the female and the first description of the male. A second poecilostome species, Sinoculosapphirina deeveyae Boxshall, 1981, is now considered to be in the genus Urocopia. The bathypelagic genus Urocopia was based by Sars (1917) on two female cope- pods collected in the North Atlantic Ocean during the 1913 cruise of the ARMAUER HANSEN. Sars (1917:3) placed this genus in the family Lichomolgidae because of the morphology of the oral appendages, but he stated that the copepods “differed essen- tially from other known Lichomolgidae.” Humes and Stock (1972) included the fam- ily Urocopiidae in the superfamily Lich- omolgoidea when they revised the family Lichomolgidae. In Humes and Stock’s key to the families (1972:122), the distinguish- ing characters of Urocopiidae are the 3-seg- mented exopods of legs 1 and 2 and endo- pod of leg 4, and leg 5 without a free segment. The family Lichomolgidae was separated on the basis of leg 4 endopod being 2-seg- mented, l-segmented, reduced to a small knob, or absent. There are characters in both families Uro- coplidae and Lichomolgidae that resemble those of some species of Sapphirina J. V. Thompson, 1829, but this genus may be separated by the 1-segmented leg 5. Addi- tional distinctive characters of Sapphirina species are two anterior cuticular lenses, a conspicuously depressed body with epi- meral plates expanded laterally, and the la- melliform caudal rami. An unpublished record of Urocopia sin- gularis from off Oregon and California in the eastern North Pacific Ocean (Olson 1949) was later published and corroborated with a record from the Bering Sea (Minoda 1971). The following new records from the eastern North Pacific further characterize the geographical range of U. singularis. Collections were made in 1964 and 1965 from the BROWN BEAR, former research vessel of the Department of Oceanography, University of Washington, with a specially- designed plankton net (mesh aperture 110 um) which accompanied a deep water-bot- tle cast (Heron and Damkaer 1978). The rare specimens of Urocopia singu- laris from the BROWN BEAR cruises and the loan of slides prepared by J. B. Olson have enabled us to supplement the pub- lished morphological details, as well as to describe the male. Figures were drawn with the aid of a Wild M2O' drawing tube. The slide of the male Urocopia singularis from which the male appendages were illustrated, was prepared by W. K. Peterson in 1965 with methyl blue stain and Turtox CMC mounting medium. The stain Solophenyl ' Reference to trade names does not imply endorse- ment by the National Marine Fisheries Service, NOAA. VOLUME 99, NUMBER 1 141 Table 1. Collection data. Length Date Depth (m) Location 2 (mm) 6 (mm) 6 V(mm) Previous records Sars VII 13 700-600 59°35.0'N, 20°40.0’E 2 (1.90) Olson 25V 39 800-0 44°46.0'N, 128°22.0'W 1 (1.97) 13 VI 39 800-0 35°42.0'N, 124°19.5'W 1 (1.28) Minoda 13 VI 61 883-743 56°23.0'N, 174°38.0’E 1 (2.10) New Records BROWN BEAR Cruise 344 4V 64 2600-0 45°29.3'N, 126°58.0’'W 1 (1.08) 28V 64 4310-0 47°49.2'N, 144°57.7'W 1 (1.05) 368 10 VIII 65 2700-0 45°22.0'N, 128°36.0’'W 1 (2.25) 1 (damaged) 13 VIII 65 2000-0 45°20.6'N, 134°56.5'W 1 (1.29) 14 VIII 65 4250-0 45°17.5'N, 139°09.6'W 1 (2.28) blue 2RL, dissolved in lactic acid, was used to study the other specimens (English and Heron 1976). The collection data for previous records and the new records of Urocopia singularis are shown in Table 1. Specimens collected from the BROWN BEAR described in this report, except the 10 VIII 65 male and female, have been de- posited in the Crustacea collection of the National Museum of Natural History (USNM), Smithsonian Institution, Wash- ington, D.C. The Olson male and female, mounted on slides, have also been depos- ited in the National Museum of Natural History. Poecilostomatoida Kabata, 1979 Urocopiidae Humes and Stock, 1972 Urocopia G. O. Sars, 1917 Sapphoncaea Minoda 1971:46. Sinoculosapphirina Boxshall 1981:307. Diagnosis. — Body cyclopiform, elongate. Urosome in female 5-segmented, in male 6-segmented. Caudal ramus elongate, la- mellar. Rostrum absent. First antenna 6-segmented; incomplete sutures between segments | and 2 or 2 and 3. Second antenna 4-segmented, sexually dimorphic; terminal segment of female with stout claw and 2 short apical setae; in male, length of ter- minal segment and width of claw reduced, outer apical seta very long. Key to the Species of Urocopia Females 1. Length 1.90-2.28 mm; caudal ra- mus length approximately equal to that of 3 preceding segments com- bined acess wero. U. singularis — Length 3.50 mm; caudal ramus length approximately equal to that of 3 preceding segments plus genital segment combined ....... U. deeveyae Urocopia singularis G. O. Sars, 1917 Figs. 1-4 Urocopia singularis G. O. Sars, 1917:3-11, figs. 1-15.—Lysholm and Nordgaard, 1921:29.—Humes and Stock, 1972:329, 330, fig. 183.—Gotto, 1979:6, 11, 13, fig. 26. Sapphoncaea moria Olson, [MS], 1949:112, pl. 27, figs. 3-11; pl. 28, figs. 1-12. — Min- oda, 1971:46, 47, pl. 4, figs. 1-12. Material examined. —2 °°, 3 66 (1 dam- aged), 1 6V; eastern Pacific (see Table). The 142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Urocopia singularis Sars, female: a, Habitus, dorsal (w); b, Segment of leg 5 and genital segment, lateral (x); c, First antenna, right ventral (y). Each scale bar equals 0.10 mm. VOLUME 99, NUMBER 1 male of this species is described for the first time. Female. —Lengths of two females 2.25 and 2.28 mm. Prosome relatively robust com- pared to slender urosome (Fig. la). Ratio of length of prosome to that of urosome 1.2: 1. Pediger 1 delimited from cephalosome dorsally by suture. Between segment of leg 5 and genital segment a pronounced ventral intersegmental sclerite (Fig. 1b). Genital segment shorter than length of caudal ramus. Genital areas located dor- solaterally anterior to middle of segment; lamellar fringe forming dorsal transverse line on one female. Each genital area with 2 se-. tae, posteriormost adjacent to sclerotized prong extending from operculum and over- lying 2 pores, anteriorly and posteriorly. Surfaces of genital segment and 3 postgen- ital segments with slightly crenulate ap- pearance. Caudal ramus elongate, dilated in area of lateral seta, terminating in inner tapered protuberance (that of smaller female twice as protuberant, shown by dashed line on Fig. 1a); minute spinules on inner and dor- sal surfaces. Setae broken or missing; based on hyaline circles of insertion (and those of male and stage V male specimens), a lateral, a dorsal, and three short terminal setae pres- ent. Innermost terminal seta absent. Rostrum absent. First antenna (Fig. Ic) 6-segmented, with partly coalesced suture between segments 2 and 3. Most setae bro- ken or missing; based on hyaline circles of insertion, armament formula may be: 2; 11; 7; 3, 1 esthete; 2, 1 esthete; 7, 1 esthete. Second antenna (Fig. 2a) 4-segmented; inner seta on first and second segments, 2 inner setae on short third segment, and stout claw plus 2 apical setae on fourth segment. Labrum (Fig. 2a) incised into 2 rounded, posteroventral lobes; inner third of each lobe delimited with thinner chitin; intricate sclerotization pattern extending from apex of incision. Mandible (Fig. 2b) with 2 outer setose 143 elements followed by denticulate ridge; ter- minating in spiniform lash; inner row of spinules on concave edge of blade. First maxilla (Fig. 2c) with row of short, minute spinules on outer anteroventral cor- ner; 4 setae on anterior half of segment; minute spinules on 2 longest setae, the ter- minal and antepenultimate. Single apo- deme, remarkably long, extending from base of segment. Second maxilla (Fig. 2d) 2-segmented; first segment with rugose ventral surface. Second segment bearing 2 elements on inner sur- face; shorter element barbed, distalmost ele- ment spinose and longer than terminal lash. Terminal lash, with vertical row of setules near base, carrying ventral row of setules adjacent to 3 or 4 triangular, inwardly-di- rected teeth. Maxilliped (Fig. 2e) 3-segmented. First segment with short, minute spinules on in- ner surface. Second segment with inner and distal patches of minute spinules plus 2 in- ner setae. Third segment bearing 2 inner setae near base and terminating in barbed claw. Legs 1—4 with trimerous rami. Spines with wide, serrate, hyaline flange. Minute spi- nules on posterolateral margins of coxae. First and second segments of leg 4 endopod with anterior digital projection overlapping base of wide, flat setae. Leg armament: see Table 2. Leg 5 (Fig. 1b) represented by 2 postero- lateral setae inserted on short pedicel. Leg 6 (Fig. 1b) probably represented by 2 setae and prong on posterodorsal margin of genital operculum. Male.—Lengths of two males 1.05 and 1.29 mm. Body compact, length of prosome approximately twice that of urosome (Fig. 3a, b). Pediger 1 delimited from cephalo- some by dorsal suture. Urosome (Fig. 3c) 6-segmented. Anal segment and caudal ra- mus with rows and patches of minute spi- nules on dorsal and ventral surfaces. Caudal ramus showing remarkable sexual dimor- 144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Urocopia singularis Sars, female: a, Second antenna and oral area, ventral (y); b, Mandible, right (y); c, First maxilla, right (y); d, Second maxilla, right (y); e, Maxilliped, right (y). phism; terminal protuberance of female de- veloped as articulated, blunt outgrowth, possibly the modification of innermost ter- minal seta. Rostral area, mandible, first maxilla, and second maxilla similar to those of female. Table 2. Leg armament. First antenna with line of coalescence be- tween segments 2 and 3 scarcely discernible; several setae and esthetes longer than those of female, posterior esthete reaching as far as coxa of leg 1. Second antenna (Fig. 3d) segments resem- Endopod 1 2 3 Leg Coxa Basis a Sigets ae Sian Sime anne Sans Ge ae 1 1 1 1 1 4 1 y; 1 1 1 2 3 II 3 1 1 1 D 2) II 4 1 i 1 1 - II Exopod 1 2 3 Se Se Si Se Si St Se I I 1 I 4 I Ill I I 1 I 5 I III I I 1 I 5 I Il - I 1 I 5 I II VOLUME 99, NUMBER 1 145 Fig. 3. Urocopia singularis Sars, male: a, Habitus, dorsal (w); b, Habitus, lateral (z); c, Urosome, dorsal (x); d, Second antenna, right (y); e, Maxilliped, right (y); f, Area between maxillipeds and first legs, ventral (y); g, Leg 1, anterior (y). 146 bling those of female in general form; ar- mament formula similar, but structure dif- fering on fourth segment, with reduced claw and increased length of 2 setae, 1 longer than second segment. Maxilliped (Fig. 3e) 4-segmented; second segment with two inner setae and patches of short spinules; third segment short and unarmed. Terminal claw bearing 2 proxi- mal setae and 2 inner rows of minute spi- nules. Distinct sclerotized band between bases of maxillipeds. Area between maxil- lipeds and first pair of legs (Fig. 3f) pro- truding ventrally as anterior and posterior lobes, separated by 2 constrictive sclero- tized bands. Legs 1—4 (Figs. 3g; 4a, b, c) segmented as in female, with same armament formula. Leg 5 (Fig. 3c) similar to that of female, except relatively longer setae. Leg 6 (Fig. 3b, c) probably represented by posterolateral flap on ventral surface of gen- ital segment, bearing 2 setae and sclerotized prong similar to those of female. Stage V male. —One specimen, 1.08 mm, resembles a mature male, including the ar- ticulated, blunt outgrowth on the caudal ra- mus and the rows of minute spinules on ventral surfaces of anal segment and caudal ramus (Fig. 4d). This stage lacks the third postgenital urosome segment of the adult male. Without dissection most oral appendages appear similar to those of the mature fe- male. Hyaline setae of leg 5 and the caudal ramus appear to be more transparent and fragile than those of the adults. Remarks. — Most setae of leg 5 and caudal ramus were broken or missing on all spec- imens. Setae which were not damaged ap- peared to be hyaline with sclerotized sup- port only in the proximal half, to that point where many of the setae were broken. Despite the omission of some swimming leg armament and slight differences in in- terpretations of details and illustrations of oral appendages, there seems little question that Olson’s, Minoda’s, and our specimens, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON all of similar size, are the same species as Urocopia singularis described by Sars C91): When Olson (1949:113) listed the ar- mament pattern of the swimming legs of Sapphoncaea moria, he inadvertently omit- ted a terminal spine on all exopods and en- dopods. All armament, including terminal spines, was on his illustrations, except 1 seta from the second segment of the female leg 3 endopod, the long terminal spine, 2 outer spines on the exopod, and | of the terminal spines on the endopod of female leg 4. Olson stated that this missing armament was pres- ent on his male specimen and he considered the armament formula to be similar to that of the female. Minoda (1971:47) identified a female specimen as Sapphoncaea moria, but his illustrations and formula omitted a seta on the second segment of leg 3 endopod and a spine from each exopod segment of leg 4. He also omitted a terminal spine of each swimming leg rami in the armament formula of swimming legs, although these spines were included on the illustrations. Urocopia deeveyae (Boxshall, 1981), new combination Sinoculosapphirina deeveyae Boxshall, 1981:307-311. figs. la—h, 2a—e (2 22, 3.50 mm). Material examined.—The paratype fe- male (USNM 173941) with legs dissected and mounted on a slide. Legs 1-4 have the same armament for- mula as U. singularis, but relative lengths of spines differ. The proximal spine of the third exopodal segment of legs 1-3 is rela- tively shorter for U. singularis as is also the outer terminal spine of leg 4 endopod. Remarks. —The swimming legs and leg 5 of Sinoculosapphirina deeveyae are similar to those of Urocopia singularis. The lack of cuticular lenses and the presence of unseg- mented leg 5 indicate that it is more closely related to Urocopiidae than to Sapphirini- dae. The 6-segmented first antenna of Uro- VOLUME 99, NUMBER 1 | i : i \ Fig. 4. Urocopia singularis Sars, male: a, Leg 2, anterior (y); b, Leg 3, anterior (y); c, Leg 4, anterior (y). Copepodid V, male: d, Urosome, ventral (x). Y % j| A| 148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON copia singularis has an incomplete suture between segments 2 and 3. Rather than 5-segmented, there is the possibility of an incomplete articulation on the long second segment of the first antenna of Sinoculo- sapphirina deeveyae. Although Boxshall de- scribed the second antenna as being 5-seg- mented, his illustration appears to have four segments with a strong terminal claw. The second antenna of the superfamily Lich- omolgoidea Humes and Stock (1972:122) was defined as being either 4-segmented or 3-segmented by a fusion of the last two seg- ments. Humes and Stock (1973:329) noted that Sars had mistakenly regarded the ter- minal claw as a fifth segment when he de- scribed the second antenna of Urocopia sin- gularis. Acknowledgments Weare indebted to Mr. Willis K. Peterson for devising the net and using it to collect deep-living copepods and for presenting the samples to us. We are grateful to Dr. Thom- as E. Bowman for the loan of the USNM slide of Urocopia deeveyae and for his con- structive criticisms of the manuscript. We also appreciate the loan from Dr. J. Bennet Olson of his slides of U. singularis. This is Contribution no. 1624 from the School of Oceanography, University of Washington. Literature Cited Boxshall, G. A. 1981. Sinoculosapphirina deeveyae, anew bathypelagic genus and species of the fam- ily Sapphirinidae (Copepoda: Poecilostomato- ida) from the Sargasso Sea off Bermuda. —Bul- letin of Marine Science 31(2):307-311. English, T. S., and G. A. Heron. 1976. A stain for morphological study of copepods.— Mono- graphs on Oceanographic Methodology, UNES- CO 4:288-289. Gotto, R. V. 1979. The association of copepods with marine invertebrates. — Advances in Marine Bi- ology 16:1-109. Heron, G. A.,and D. M. Damkaer. 1978. Seven Lub- bockia species (Copepoda: Cyclopoida) from the plankton of the Northeast Pacific, with a review of the genus.—Smithsonian Contributions to Zoology 267:1-36. Humes, A. G., and J. H. Stock. 1972. Preliminary notes on a revision of the Lichomolgidae, cy- clopoid copepods mainly associated with ma- rine invertebrates. — Bulletin van de Zoologisch Museum, Universiteit van Amsterdam 2(12): 121-133. ——., and 1973. A revision of the family Lichomolgidae Kossmann, 1877, cyclopoid co- pepods mainly associated with marine inver- tebrates.—Smithsonian Contributions to Zool- ogy 127:1-368. Kabata, Z. 1979. Parasitic Copepoda of British fish- es.—Ray Society Monographs 152:1—468. Lysholm, B., and O. Nordgaard. 1921. Copepoda collected on the cruise of the M/S ARMAUER HANSEN in the North Atlantic 1913.—Bergens Museums Aarbok 1918-19, Naturvidenskabe- lig Raekke 1(2):1—37. Minoda, T. 1971. Pelagic Copepoda in the Bering Sea and the northwestern North Pacific with special reference to their vertical distribution. — Memoirs of the Faculty of Fisheries, Hokkaido University 18(1/2):1-74. Olson, J. B. 1949. The pelagic cyclopoid copepods of the coastal waters of Oregon, California and Lower California. Thesis, University of Cali- fornia, Los Angeles. 208 pp. Sars, G. O. 1917. Urocopia singularis G. O. Sars, a peculiar semiparasitic copepod from great deeps of the North Atlantic Ocean.—Bergens Mu- seums Aarbok 1916-17, Naturvidenskabelig Raekke 1(4):1-11. Thompson, J. V. 1829. On the luminosity of the ocean, with descriptions of some remarkable species of luminous animals (Pyrosoma pig- maea and Sapphirina indicator) and particular- ly of the four new genera, Noctiluca, Cynthia, Lucifer and Podopsis, of the Schizopodae.— Zoological Researches and illustrations; or Nat- ural History of nondescript or imperfectly known Animals, in a Series of Memoirs 1(2):37-61. (GAH, DMD) School of Oceanography, University of Washington WB-10, Seattle, Washington 98195; and (DMD) Coastal Zone and Estuarine Studies Division, Northwest and Alaska Fisheries Center, Na- tional Marine Fisheries Service, NOAA, 2725 Montlake Boulevard East, Seattle, Washington 98112. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 149-159 SUBSPECIES OF THE GLAUCOUS GULL, LARUS HYPERBOREUS (AVES: CHARADRIIFORMES) Richard C. Banks Abstract. —Current writings treat the Holarctic Glaucous Gull, Larus hyper- boreus, either as a monotypic species or as having two or three subspecies; if divided, birds of Canada, Greenland, and Europe are considered to be of the nominate subspecies. This study shows that there are four subspecies, the birds of Canada and Greenland being separable from those of Europe; the name L. h. leuceretes Schleep, 1819, based on a Greenland bird, is available for the former. Alaskan birds (L. h. barrovianus) are relatively dark on the mantle, those of Canada and Greenland are pale, those of Europe and western Asia are dark, and those of Siberia (pallidissimus) are very pale. From the small Alaskan birds there is an increase in size to the east around the Holarctic to very large birds in Siberia. The Alaskan and Canadian populations intergrade in extreme northwestern Canada. Nonbreeding Glaucous Gulls along the Pacific coast of North America are of the Alaskan form, barrovianus; those east of the Rockies, previously referred to as barrovianus or hyperboreus, are all leuceretes or, in the western plains states, from the intergrade area. The nature and extent of geographic vari- ation in the Holarctic Glaucous Gull, Larus hyperboreus, and the nomenclatural recog- nition of this variation have been a matter of dissent since the species was first divided in the late 19th century. Authorities writing in the last two decades have considered the species to be either monotypic or composed of two or three subspecifically distinct pop- ulations. Authors who recognize subspecies have not agreed on the application of names or on the boundaries of the named popu- lations. A request for subspecific identification of a wintering specimen prompted a reexam- ination of the series of this species in the National Museum of Natural History (USNM), the American Museum of Natural History (AMNH), and the Academy of Nat- ural Sciences of Philadelphia (ANSP), and some individual specimens borrowed from other institutions (see Acknowledgments), as well as the taxonomic literature of the species. This has allowed the identification of some of the sources of earlier disagree- ment, and revealed that geographic varia- tion is more complex than has been rec- ognized. I believe that four populations are recognizable at the subspecific level. Taxonomic History The name Larus glaucus Brunnich, 1764, was used for the Glaucous Gull until the early part of the 20th century. The Ameri- can Ornithologists’ Union (A.O.U. 1908), citing a manuscript by C. W. Richmond (ap- parently never published), noted that Larus glaucus of Brunnich is preoccupied by Lar- us glaucus Pontippidan, 1763, a synonym of Larus canus Linnaeus, 1758, and that the next available name is Larus hyperboreus Gunnerus, 1767. The latter has been the accepted specific name ever since. Ridgway (1886) described Larus barro- vianus as an Alaskan species of gull that was smaller and darker than the related North Atlantic L. glaucus. The name barrovianus was applied to Bering Sea birds by Tacza- 150 nowski (1893), but was synonymized with glaucus by Saunders (1896). Dwight (1906) thought that the size difference between bar- rovianus and glaucus was insufficient for the recognition of the former, and although he did not comment on the color difference he also placed barrovianus in the synonymy of glaucus. After 1908, barrovianus was car- ried in the synonymy of hyperboreus, even by Ridgway (1919). Oberholser (1918), however, proposed recognition of barrovi- anus as a subspecies of hyperboreus, em- phasizing the color difference, and attrib- uted to it a breeding range in “Alaska and the territories of Yukon and western Mac- kenzie.”’ In a stinging rebuff, Dwight (1919) reemphasized the weakness of the charac- ters used by Ridgway and Oberholser; the latter repeated his argument (Oberholser 1919) only to have it rebutted again (Dwight 1925). Despite support for the recognition of barrovianus by Bishop (1927), that form was not recognized in compendia by the A.O.U. (1931) or Peters (1934). Portenko (1939), on the basis of a state- ment in a letter from Herbert Friedmann that “‘the characters given concerning the colouring of the mantle [of the type of bar- rovianus] ... do not hold for other speci- mens from the same region,” believed that the type of “barrowianus”’ was a hybrid be- tween L. hyperboreus and L. glaucescens and not “identical with the pale coloured Glau- cous Gulls from the Arctic shores of N.E. Asia and N.W. America.” Portenko (1939) proposed the name L. h. pallidissimus for the birds of eastern Asia, primarily on the basis of their paler mantle color relative to birds of Europe and western Asia but noting also their larger size. Portenko had a simi- larly pale bird from Ellesmere Island, and presumed that the range of pallidissimus ex- tended from Siberia eastward across arctic America at least to that island. Thus the Alaskan birds, first described as smaller and darker, were incorporated into a subspecies based on pallor and large size. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Witherby (in Witherby et al. 1941:112) noted that pallidissimus had been described, but stated that “specimens from Amur and Alaska do not appear to me to differ from European examples.” Bird and Bird (1941) examined the same specimens as Witherby had and suggested that Portenko had merely renamed barrovianus; they also were unable to find characters to divide the species in any part of its range, and considered hy- perboreus to be monotypic. Rand (1942) supported the earlier views of Oberholser and Bishop that barrovianus was subspecifically distinct from hyperbo- reus, without mentioning pallidissimus. Two subspecies, L. h. hyperboreus and L. h. bar- rovianus were recognized by the A.O.U. (1945). However, Hellmayr and Conover (1948:261) continued to follow Dwight in denying recognition of barrovianus. They used only the binomial Larus hyperboreus for the birds in North America, but in a footnote commented that “this form is re- placed by L. h. pallidissimus” on the arctic coast of Asia. They were equivocal on the validity of the latter, but stated that “birds from Greenland, Arctic America, and Alas- ka are not separable from those of northern Europe.” Dement’ev (in Dement’ev and Gladkov 1951) recognized two subspecies, hyperbo- reus from eastern Canada, Greenland, northern Europe, and extreme western Asia, and ‘“‘barrowianus”’ from eastern Asia, Alas- ka, and Canada east to Ellesmere Island. In this, he essentially followed Portenko’s di- vision of the species but considered palli- dissimus a synonym of barrovianus, reject- ing Portenko’s claim that the latter was based on a hybrid. He reinterpreted Friedmann’s statement to Portenko (see above), as had Bird and Bird (1941), as indicating that the type of barrovianus was merely atypical and that the gulls from arctic America are “in fact extremely light.’ He also noted that the east Siberian birds are larger than Atlantic ones, and assumed that Alaskan ones agreed VOLUME 99, NUMBER 1 with those in Siberia in this respect. De- ment’ev stated, however, that (p. 562) “‘the boundary lines of the range in America re- main completely obscure ...”’ and further that the identity of American birds with those of Siberia “‘is still unproved, although quite likely.” The A.O.U. (1957) continued to recog- nize barrovianus as an Alaskan form and hyperboreus as the single other subspecies, extending from western Canada around the Holarctic to and including Siberia. Walrus Island, in the Pribilofs, was mentioned in the breeding range of both forms. Gabriel- son and Lincoln (1959) followed the treat- ment by the A.O.U. but included Walrus Island and St. Matthew Island in the range only of hyperboreus. They indicated that the latter populations might belong to pallidis- simus, which they had not fully evaluated. Todd (1963:363) discussed the difference of opinion on the validity of barrovianus as distinct from hyperboreus. He noted an av- erage color difference, but also that individ- ual specimens from eastern and western Canada were “scarcely to be distinguished in general coloration.”’ He further remarked that “the western birds of this species run smaller, sex for sex, than those from the East.’ However, he agreed with Dwight (1906, 1925) that the degree of size differ- ence was insufficient for the recognition of subspecies. Ingolfsson (1970) similarly con- sidered L. hyperboreus as monotypic. Vaurie (1965:475) recognized three sub- species, with barrovianus restricted to “coasts and islands of Alaska ... to about Franklin Bay in northwestern Mackenzie.” He considered birds from central Macken- zie eastward through North America and Europe “to about the Taimyr Peninsula” in Siberia to be nominate hyperboreus. To pal- lidissimus he gave the range from the Tai- myr Peninsula to the tip of the Chuckchi Peninsula, including Anadyrland, Wrangel Island, and the Pribilofs. He considered bar- rovianus to be small and dark, with a slender 151 bill, and pallidissimus to be paler in adult and immature plumages than hyperboreus. Portenko (1973) followed in recognizing these three subspecies, as did Glutz von Blotzheim and Bauer (1982) and Cramp (1983). Several clues in this historical summary suggest that a pattern of variation in L. hy- perboreus has been overlooked. Bishop (1927) noted that individuals of hyperbo- reus from Siberia (actually Portenko’s later named pallidissimus) wander to Alaska, and suggested that such vagrants may have in- fluenced Dwight’s conclusions about the va- lidity of a small, dark Alaskan subspecies. Portenko (1939) recognized, on the basis of One specimen from Ellesmere Island, that eastern Canadian birds are paler than Eu- ropean birds, as are Siberian birds. Both Portenko and Dement’ev (1951) were mis- led by Friedmann’s letter to Portenko into thinking that Alaskan birds were also pale, despite Oberholser’s (1918, 1919) state- ments to the contrary. Dement’ev (1951) noted that Siberian birds were larger than European ones, and Todd (1963) com- mented on increasing size to the east across Canada. An additional clue to the reason for dis- agreement on the recognition of subspecies in North America is found in AMNH birds marked “‘Dwight ref. spec.,”’ individuals that apparently formed the basis for Dwight’s concept of various populations. A Green- land bird (AMNH 64142, unsexed, 2 Aug 1893, badly stained ventrally and soiled dorsally) and one from Sable Island, Nova Scotia (AMNH 358035, male, 22 Feb 1895) seem typical of their populations in size and color. However, the reference specimen from Pt. Barrow, Alaska (AMNH 358051, fe- male, 5 Sep 1897) is not an adult bird; it has a black band on the bill and some brown mottling in the wing and tail feathers. Both age and date indicate that it is not a breeding bird. Although it is the size of other Alaska specimens, it is very pale on the mantle, 152 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ad ali Co’ pbs istle Tare Eo ella Baas Ss 1 ) © @Mo E? pl vag Cc? ey Pe eS A. See eee /\6 sella metros [al Cd ¥ ee Evo cal [ae tS) 2 ° of@ QO © E? ol eae Cc? afte ed A? ered ene Ad Aina [E SUnaes oe Cd st ie el a Eo’ Babak fed ELD Ss 3 O° @Moo0o EQ en Cc? alien Ag? ee ae Fuca Sw ae Peo aS wl SS 1h a | ll ee ee 8 10 12 14 16 18 millimeters Fig. 1. Ad pe eee Ca pore) Cero Ect pam Ss 4 © 000 EQ? pale cg bey eae AQ Sellen Ad il ears (= oH maples Ed geil Ss 5 oo8 oO E? alk ce ecru A? 9 lee Ad te eer: Co a Ed reams Bb Ss 6 00000 Oo E? wilt Ee ail wer AQ ane Bere 2 a a a | 40 50 60 70 80 millimeters Range of measurements in populations of the Glaucous Gull: 1, Depth of upper mandible; 2, Depth of lower mandible at gonys; 3, Width of upper mandible; 4, Length of culmen; 5, Length of lower mandible; 6, Length of tarsus. Horizontal line represents the range, the longer vertical line the mean, and the shorter vertical lines one standard deviation to either side of the mean. Individual measurements are plotted for the Siberian birds, which are not separated by sex, because of the small sample size. even paler than the other two reference specimens. That reference specimen is probably why Dwight adamantly refused to recognize a dark-mantled Alaskan popula- tion. Methods of the Present Analysis Sexed birds in the adult, gray-mantled plumage, taken mainly in the breeding sea- son, were divided into subsets related to geographic origin— Alaska, eastern Canada and Greenland, Europe, and eastern Asia. The color of the mantle was judged in rel- ative terms as pale or dark and was related to the Munsell (1929-1942) color system. Six measurements were made on each bird: 1) depth of the upper mandible at the an- terior point of its lateral feathering; 2) depth of the lower mandible at the gonys; 3) length of the unfeathered culmen; 4) length of the lower mandible from the point of its lateral feathering; 5) width of the bill at the point of lateral feathering on the upper mandible; and 6) length of the tarsus. Several of these measurements are not standard, but are self- explanatory. The culmen length measure- ment (3) approaches “length of total cul- men’ more nearly than “length of exposed culmen” of Baldwin et al. (1931). These measurements were designed to show the “‘massiveness” of the bill. Wing and tail length were not measured, because exten- sive wear On many specimens renders them unreliable. Color comparisons were also made on the few downy young available, and on birds in the first-year brown streaked plumage. Col- or comparisons are not possible in the “‘sec- ond-year”’ fully white (“Autchinsii’’) plum- VOLUME 99, NUMBER 1 age. Measurements of birds in other than adult plumage were used for identification purposes after analysis had shown that the four samples represented separable popu- lations. Results Consideration of both color and size (Fig. 1) of birds from their breeding grounds in- dicates that four populations of Larus hy- perboreus are distinguishable. Birds from the mainland of Alaska and nearby islands have relatively dark mantles and are small, the latter especially noticeable in the bill. Those of the eastern Canadian archipelago and Greenland have considerably larger bills and are paler on the mantle. European birds have similarly large bills but are darker on the mantle. Birds from Siberia and some islands in the Bering Sea are very large, with mas- sive bills, and are very pale on the mantle. Thus, Siberian and eastern Canadian/ Greenland birds are similar in their pale color (Munsell N 8.5—9/0), as noted by Por- tenko (1939), but differ in size of bill. Alas- kan and European populations differ from the other two in being darker (Munsell N 7.5—8/0), and differ from one another in size of the bill. The relative color differences of the mantles of the adult birds are evident also in overall coloration of both downy young and first-year birds. In general, there is a trend of increasing size in all bill measurements and in tarsal length (Fig. 1) eastward from Alaska around the Holarctic to Siberia, with the result that the largest and smallest populations are ad- jacent (but separated by the Bering and Chuckchi Seas and Bering Straits). There are reversals or interruptions in the trend in several instances; European birds are smaller than expected if the trend were a smooth cline. This may be an artifact of the small samples of European and Siberian birds or it may indicate that European birds are in fact only slightly different in size from those of Canada and Greenland. The trend 153 of increasing size around the Holarctic can only provisionally be considered a true cline because too few populations, each from a large area, are considered. Systematic Treatment Larus hyperboreus barrovianus Ridgway, 1886 Larus barrovianus Ridgway, 1886:330. Holotype.—USNM 88913, adult male, Point Barrow, Alaska, 4 Aug 1882. This specimen has some feathers of the neck, back and rump edged with brown; a few are edged with black that looks rather like oiling. The tail is also discolored at the tip. These mark- ings may be what Friedmann was referring to in his letter to Portenko (see above). One other Point Barrow bird (USNM 93304) is similarly but less extensively marked. Breeding range.—Coasts and islands of Alaska from Bristol Bay north to Kotzebue Sound and Point Barrow, and eastward on the Alaskan north coast. I have not exam- ined specimens of barrovianus from east of Point Barrow. Discussion. —If the species’ range is con- tinuous across the northern coast of Cana- da, as indicated by Godfrey (1966) but not by Snyder (1957), barrovianus probably in- tergrades with the next form between the Mackenzie River Delta and Franklin Bay. A worn, pale bird (AMNH 119090) from the Mackenzie River Delta is intermediate in three of the six measurements. Manning et al. (1956) commented that birds from western Canada are smaller than those in the east and in Greenland, although speci- mens from the Mackenzie Delta show less approach to barrovianus in color than in size. Henri Ouellet (pers. comm., 1985) not- ed that specimens in the National Museum of Canada from Mackenzie Delta, Ander- son River, and Harrowhy Bay are closer to barrovianus in size and closer in average coloration of the mantle to /euceretes, and emphasized that Canadian birds are not 154 uniform in color; an occasional dark-man- tled bird may occur anywhere from Banks Island east to Baffin Island. A number of Alaskan specimens and non- breeding birds from the Pacific coast of North America are extensively dark on the primaries, perhaps a result of hybridization with the Glaucous-winged Gull, Larus glau- cescens, or the Herring Gull, L. argentatus (Ingolfsson 1970:357—358; Strang 1977). Some very large birds from Alaska seem to be nonbreeding vagrant pallidissimus from Siberia, as suggested by Bishop (1927). Larus hyperboreus leuceretes Schleep, 1819 Larus leuceretes Schleep, 1819:314. Holotype. —Schleep (1819) mentioned three young birds from unspecified locali- ties but based most of his description on an adult from Greenland in the collection of Herr Benicke of Schleswig. The range of the species was given as Greenland, Iceland, and Spitsbergen. Hellmayr and Conover (1948: 260) accepted the Greenland adult as the type, noting, however, that it is probably lost. Breeding range. —Northern Canada east- ward from Franklin Bay, Mackenzie, the Canadian archipelago, south in Hudson Bay to the Belcher Islands and on the Atlantic coast to northern Labrador (Hopedale), Greenland, probably Iceland. Discussion. —Ridgway (1919), Dwight (1925), and Hellmayr and Conover (1948) gave extensive synonymies for L. hyper- boreus, from which a name applicable to the Canada-Greenland population, here recog- nized as an entity for the first time, must be chosen. The earliest available and appro- priate name seems to be Larus leuceretes Schleep, 1819, based on specimens from Greenland, Iceland, and Spitzbergen, of which an adult bird from Greenland is con- sidered to be the type (Hellmayr and Con- over 1948:260). Meyer (footnote in Schleep 1819) indicated that /euceretes was a syn- onym of Larus giganteus Temm., but the latter name seems to be, at least in part, a PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON synonym of L. marinus Linnaeus (Saunders 1896; Ridgway 1919; Dwight 1925). Schleep (1819) contrasted /euceretes with glaucus by noting that the mantle of the former was “‘hellgraulichweiss” as opposed to “‘hell- blaugrau”’ in g/aucus. He also noted that the tarsus of /euceretes was slightly longer, and the bill was similar in form to that of glaucus but proportionately larger. In a table, Schleep further compared both /euceretes and glau- cus to L. marinus. It is clear that he distin- guished his Greenland bird from European specimens of glaucus (now hyperboreus). My sample from the Canada-Greenland population included birds from Coronation Gulf, Baffin Island, and various islands in the northern Canadian archipelago, as well as from Greenland. Most are definitely breeding birds, but a few September spec- imens (Cornwallis Island) may have moved some distance from their nesting area. Icelandic birds are provisionally placed with /euceretes, although the situation is far from clear. Ingolfsson (1970) has shown that Glaucous Gulls in Iceland now interbreed freely with Herring Gulls; recent specimens would be difficult to interpret on the sub- specific level. I have examined only three older specimens from Iceland. A male (AMNH 745238, June, year not specified) is pale on the mantle like birds from eastern Canada and Greenland. In most measure- ments this bird falls into the range of the latter population, but it is in the zone of overlap with European birds in three mea- surements; in depth of the upper mandible it matches only the small Alaskan birds. A female (AMNH 745239, 9 Sep 1898) is not fully adult but matches European birds best in mantle color. Most of its measurements are in the overlap zone. The third specimen (AMNH 745241, female, 30 Nov 1823) is the type of Larus minor Brehm = Larus medius Brehm, names now carried in the synonymy of L. hyperboreus. It is less than fully adult, having a slight dark ring on the bill and some brown in the crown. There is too much dark color on the primaries to be typical of hyperboreus, and the bird appears VOLUME 99, NUMBER 1 to be very small. In most measurements it fits with Canadian or Alaskan series, but the depth of the upper mandible is less than in any other specimen I measured. It is slightly paler on the mantle than European birds, but not quite as pale as Greenland speci- mens. I am not convinced that the bird is an example of hyperboreus, although it is too large to be an Iceland Gull, L. glau- coides. It may be a hybrid between these two species. Larus hyperboreus hyperboreus Gunnerus, 1767 Larus hyperboreus Gunnerus, in Leems 1767:226. Holotype.—Probably none extant, the name based on birds from northern Norway (Ridgway 1919; Hellmayr and Conover 1948; A.O.U. 1957). Breeding range. —Northern Europe from Jan Mayen and Spitzbergen east along the coast and islands of the U.S.S.R. to the Tai- myr Peninsula (Vaurie 1965). Discussion. —My breeding sample of this population is small, only six birds, but win- tering adults from Europe, presumed to rep- resent this subspecies on geographic grounds, are similar in all characters. It is possible that some birds from Greenland stray to Europe in winter, and that the presence of paler Greenland birds with the darker Eu- ropean ones has given the impression of a wider range of variability in the European population than is evident from the ex- amination of breeding birds alone. Larus hyperboreus pallidissimus Portenko, 1939 Larus hyperboreus pallidissimus Portenko, 1939:226. Holotype. —Male, settl. Naukan, Chukot- ski Peninsula (female paratype from settl. Uelen), in collection of L. Portenko, now presumably in the Leningrad Museum. Breeding range. —Arctic Siberia from about the Taimyr Peninsula eastward to the 155 tip of the Chuckchi Peninsula, Wrangel Is- land, St. Matthew Island, and Walrus Island in the Pribilofs (Vaurie 1965; Portenko 1973). Discussion. —An unsexed bird from Diomede Island, Jul 1881 (USNM 97255) is pale on the back like pallidissimus, but is in the size range of barrovianus. The Diomedes would be a reasonable place for intergradation of the two subspecies, if they do intergrade. I also consider a male from Cape Lisburne, Alaska, 1 Aug 1897 (USNM 745279) to be intermediate; it is as large as typical pallidissimus but the mantle color is dark like that of barrovianus. Nonbreeding Distribution in North America Glaucous Gulls are inclined to wander extensively in the nonbreeding season, and movement to the east or west into the breed- ing range of another subspecies has been responsible, I believe, for some of the mis- understanding of geographic variation in the species. One bird from Point Barrow, Alas- ka, taken 5 Oct 1897 (AMNH 358049) is typical of the Siberian breeding population, pallidissimus, in both color and size. This bird was formerly in Dwight’s collection and, if considered by him to be representative of the Barrow breeding population, may have been partly responsible for his (1906, 1919, 1925) rejection of Ridgway’s name for the smaller, darker Alaskan form. Oberholser (1918) placed this bird with barrovianus, presumably on geographic grounds. A bird from Unalaska Island in the Aleutians (USNM 230781, 9 Jun 1911) is typical of pallidissimus in size but is in the all white plumage. Several Alaska and Yukon birds appear to represent postbreeding or nonbreeding birds from the more eastern /euceretes. A bird from Tolugak Lake, Alaska (USNM 435222, 11 Jun 1949) in the all white plum- age has measurements more similar to Ca- nadian birds than to Alaskan ones. Juvenile birds from Bettles, Alaska (USNM 298495, 11 Oct 1924) and Old Crow, Yukon (USNM 156 469302, 28 Nov 1957) are both paler than typical juveniles from farther west in Alas- ka. As noted previously, Dwight’s reference bird from Point Barrow (AMNH 358051, 5 Sep 1897) is pale dorsally, and another immature bird from there (USNM 93301, 15 Sep 1882) is also pale. These may all represent wanders from a more eastern, per- haps intermediate, population. Most of the birds that move southward in the winter are young, in the all white plumage or some combination of that and a brown streaked plumage. For these birds color comparison is impossible and sub- specific identification must be from mea- surements alone. No one measurement will suffice to separate the forms, although there is little or no overlap in most mensural char- acters of Siberian and Alaskan birds. A se- ries or set of measurements, however, should indicate at least a strong probability of the breeding population from which a properly sexed specimen has been derived. Glaucous Gulls appear somewhat infre- quently in winter in Hawaii, more regularly along the Pacific coast of Canada and the United States to southern California and Baja California (Devillers et al. 1971), in- land south to Texas and the Gulf coast, and on the Atlantic coast to Florida and, rarely, Bermuda. Pacific coastal birds have gener- ally been referred to barrovianus when a subspecific determination has been made, and all west coast and Hawaiian birds that I have examined are indeed of that subspe- cies. A bird taken in Seattle, Washington (USNM 163899, 12 May 1896) was referred to barrovianus by Oberholser (1918) but to hyperboreus in distinction to barrovianus by Jewett et al. (1953). That identification to the larger form suggested that it might be a vagrant from the Siberian population, but measurements indicate that it is barrovi- anus. Devillers et al. (1971) mentioned an extremely large Glaucous Gull from the Sal- ton Sea, California. I have examined that male bird (SBCM 33216), which is white with extensive brown mottling, and find that PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON it, too, is best considered barrovianus al- though its measurements ovelap those of leuceretes. When subspecific identification has been reported, Atlantic coast specimens have been called hyperboreus. Those that I have examined should be referred to /euceretes, here separated from the European hyper- boreus. A Bermuda specimen (AMNH 783759, female, collected by D. Wingate, 18 Dec 1964) is a young bird, and its mea- surements are rather small, especially the depth of the lower mandible, but it best fits with /euceretes. Stevenson and Atherton (1984) have recently reviewed Florida rec- ords of L. hyperboreus, and noted that some specimens from that state are small. Bailey and Niedrach (1965) reported two specimens of Glaucous Gull from Barr, Col- orado, in the Denver Museum of Natural History, under the subspecific name bar- rovianus. Allan R. Phillips kindly provided measurements of these birds, taken to my specifications, and noted that both are in a brown mottled “‘first year” plumage. A male (DMNH 18800, | Apr 1938) is in the over- lap zone of Alaskan and Canadian birds in three measurements but resembles Alaskan birds in culmen length and width and depth of the upper mandible. A female (DMNH 18799, 28 Mar 1938) equals or exceeds Ca- nadian birds in all measurements except that of upper mandible width, leading one to wonder if it is missexed. If it were a male, two measurements would be those of Ca- nadian birds, two of Alaskan, and two in the overlap zone. Asked for an impression of bill massiveness relative to Barrow, Alas- ka, birds in the DMNH collection, Phillips (in litt.) reported that the Colorado birds’ bills were “heavier than most.’ With this information, and considering probably mi- nor differences in measuring techniques, I suggest that both birds are from a popula- tion in western Canada that is in the inter- grade zone. One specimen of Glaucous Gull has been reported from Kansas (Rintoul 1984) and VOLUME 99, NUMBER 1 two from Oklahoma (Anderson 1971; Ports 1976), all without subspecific identification. All three are in immature plumage with much brown mottling, and all are referred to /euceretes on the basis of measurements (KSTC B-1406, female, Cheyenne Bottoms, Barton Co., Kansas, 6 Mar 1967; UOMZ 7175, male, Salt Fork, Arkansas River, Al- falfa Co., Oklahoma, 5 Feb 1971; UOMZ 7913, male, Lake Hefner, Oklahoma Co., Oklahoma, 27 Dec 1974). Another bird (MCZ 33036, 17 Dec 1880) is claimed as a specimen record for both Oklahoma (Sutton 1967) and Texas (Ob- erholser 1974); it was taken on the Red Riv- er, which separates the states. Oberholser (1918, 1974) considered this bird to be bar- rovianus. The bird is labelled as a male, and if this sex determination is correct I agree with Oberholser’s identification. However, the depth of the upper mandible is less than in any male barrovianus that I measured and I suspect that the specimen is actually a female, in which case the measurements place in the low range of /euceretes. It may represent the population where the two sub- species intergrade. Ragsdale (1881) com- mented that the bird had been dead for six weeks before he obtained it; when and by whom the sex was determined is not indi- cated. A specimen from Gainesville, Cook Co., Texas (MCZ 32371; date unknown) was al- lotted to L. h. hyperboreus by Oberholser (1918, 1974). I refer this unsexed bird to leuceretes. A bird found dead at the Hag- erman National Wildlife Refuge, Grayson Co., Texas, by Karl Haller is also /euceretes. One other Texas specimen (WWF 1357, Mustang Island, Neuces Co., 10 Apr 1967) has also been listed as an example of L. h. barrovianus (Oberholser 1974). This bird was reexamined by R. G. McCaskie, who believed it to be an albinistic Herring Gull, L. argentatus. I have also studied this spec- imen and agree that it is a Herring Gull. Although its culmen length is within the range of L. h. barrovianus, all other mea- 157 surements are too small for any subspecies of hyperboreus. A similar all-white-plum- aged specimen from Louisiana (LSU 130496, female, 2 mi NNE Chalmette, St. Barnard Par., 28 Feb 1982) is also probably an albino, most likely of argentatus, rather than any form of hyperboreus. However, a partly gray-backed, nearly adult plumaged bird (LSU 103495, female) taken at the same time and place as the last is an example of L. h. leuceretes. It thus appears from the specimen record now available that Alaskan Glaucous Gulls (barrovianus) move to the south only along the Pacific coast, and not into the inland states. All specimens from east of the Rocky Mountains represent the Canadian popu- lation, /euceretes, although some seem to be from the western portion of the breeding range of that subspecies where intergrada- tion with barrovianus presumably occurs. Acknowledgments I thank authorities at the American Mu- seum of Natural History (AMNH) and the Academy of Natural Sciences of Philadel- phia (ANSP) for courtesies extended while I studied gulls in their collections. I also thank Raymond A. Paynter, Jr., Museum of Comparative Zoology (MCZ), Gene A. Blacklock, Welder Wildlife Foundation (WWF), Gary D. Schnell, University of Oklahoma Museum of Zoology (UOMZ), Eugene A. Cardiff, San Bernardino County Museum (SBCM), James Van Remsen and Stephen Cardiff, Louisiana State University Museum of Zoology (LSU), and Dwight L. Spencer, Emporia State University (KSTC) for the loan of specimens for examination. Allan R. Philips measured birds from the Denver Museum of Natural History (DMNH) and Lester L. Short examined cer- tain material in the AMNH for me. Roger B. Clapp provided references to the distri- bution and biology of this species, and George E. Watson gave help with literature written in German and Russian. Daniel D. Gibson, Charles O. Handley, Jr., David W. 158 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Johnston, and Watson made critical com- ments on an early draft of the paper. W. Earl Godfrey, Joe T. Marshall, and Henri Ouellet read a much revised version, and Ouellet provided important information about certain specimens in the National Museum of Canada. Claudia Angle pre- pared the figure. Literature Cited American Ornithologists’ Union. 1908. Fourteenth supplement to the American Ornithologists’ Union check-list of North American birds.— Auk 25:343-399. 1931. Check-list of North American birds. Fourth edition. Lancaster, Pennsylvania, 526 pp. . 1945. Twentieth supplement to the American Ornithologists’ Union check-list of North American birds.— Auk 62:436-449. 1957. Check-list of North American birds. Fifth edition. Baltimore, Maryland, 691 pp. Anderson, B. W. 1971. The Glaucous Gull in Okla- homa.—Bulletin of the Oklahoma Ornitholog- ical Society 4:31-32. Bailey, A. M., and R. J. Niedrach. 1965. Birds of Colorado. 2 volumes. Denver Museum of Nat- ural History, Denver, Colorado, 895 pp. Baldwin, S. P., H. C. Oberholser, and L. G. Worley. 1931. Measurements of birds. —Scientific Pub- lications, Cleveland Museum of Natural History 2, 165 pp. Bird, C. G., and E. G. Bird. 1941. The birds of north- east Greenland.—Ibis 1941:118-161. Bishop, L. B. 1927. The status of the Point Barrow gull.— Condor 29:204-205. Cramp, S. [Chief Ed.] 1983. Handbook of the birds of Europe, the Middle East, and North Africa. The birds of the western Palearctic. Vol. 3. Oxford University Press, Oxford, 913 pp. Dement’ev, G. P., and N. A. Gladkov. [Eds.] 1951. Birds of the Soviet Union. Vol. III. Moscow. [Translated 1969, Israel Program for Scientific Translations, Jerusalem, 756 pp.] Devillers, P., G. McCaskie, and J. R. Jehl., Jr. 1971. The distribution of certain large gulls (Larus) in southern California and Baja California. —Cal- ifornia Birds 2:11-26. Dwight, J. 1906. Status and plumages of the white- winged gulls of the genus Larus. —Auk 23:26- 43. . 1919. Reasons for discarding a proposed race of the Glaucous Gull (Larus hyperboreus).— Auk 32:242-248. 1925. The gulls (Laridae) of the world; their plumages, moults, variations, relationships and distribution.— Bulletin of the American Mu- seum of Natural History 52:63—401. Gabrielson, I. N., and F.C. Lincoln. 1959. The birds of Alaska. The Stackpole Co., Harrisburg, Pennsylvania, 922 pp. Glutz von Blotzhiem, U. N., and K. M. Bauer. 1982. Handbuch der Vogel Mitteleuropas. Vol. 8, pt. 1. Akademische Verlagsgesellschaft, Wiesba- den, 699 pp. Godfrey, W. E. 1966. The birds of Canada.—Na- tional Museum of Canada, Bulletin 203, 428 pp. Hellmayr, C. E., and B. Conover. 1948. Catalog of birds of the Americas and the adjacent islands. Part 1, no. 3.— Zoological Series, Field Museum of Natural History, Vol. XIII, Part 1, no. 3. 383 pp. Ingolfsson, A. 1970. Hybridization of Glaucous Gulls Larus hyperboreus and Herring Gulls L. argen- tatus in Iceland.—Ibis 112:340-—362. Jewett, S. G., W. P. Taylor, W. T. Shaw, and J. W. Aldrich. 1953. Birds of Washington State. University of Washington Press, Seattle, 767 pp. Leems, K. 1767. Beskrivelse over Finmarkens Lap- per, pt. I. Manning, T. H., E. O. Hohn, and A. H. Macpherson. 1956. The birds of Banks Island.— National Museum of Canada, Bulletin 143, 144 pp. Munsell Color Company. 1929-1942. Munsell book of color. Pocket edition. Munsell Color Com- pany, Baltimore, Maryland. Oberholser, H.C. 1918. The subspecies of Larus hy- perboreus Gunnerus.— Auk 35:467—474. 1919. The status of Larus hyperboreus bar- rovianus Ridgway.— Proceedings of the Biolog- ical Society of Washington 32:173-176. 1974. The bird life of Texas. 2 vols. Uni- versity of Texas Press, Austin, 1069 pp. Peters, J. L. 1934. Check-list of birds of the world. Vol. 2. Harvard University Press, Cambridge, 401 pp. Portenko, L. [A.] 1939. On some new forms of arctic gulls.—Ibis 1939:264—-269. 1973. [The birds of the Chukotsk Peninsula and Wrangel Island.] Vol. 2. Leningrad, 324 pp. [In Russian.] Ports, M. 1976. Third specimen of Glaucous Gull for Oklahoma.— Bulletin of the Oklahoma Or- nithological Society 9:6-7. Ragsdale, G.H. 1881. Larus glaucus in Texas. —Bul- letin of the Nuttall Ornithological Club 6:187. Rand, A. L. 1942. Larus kumilieni and its allies.— Canadian Field-Naturalist 56:123-126. Ridgway, R. 1886. On the Glaucous Gull of Bering’s Sea and contiguous waters.— Auk 3:330-331. . 1919. The birds of North and Middle Amer- VOLUME 99, NUMBER 1 ica. Part VIII.—United States National Mu- seum of Natural History, Bulletin 50(8):1-852. Rintoul, D. A. 1984. Glaucous Gulls in Riley Coun- ty.—Kansas Ornithological Society Bulletin 35: 22-23. Saunders, H. 1896. Gaviae in Saunders, H., and O. Salvin, Catalogue of the Gaviae and Tubinares in the collection of the British Museum. British Museum (Natural History), London, 475 pp. Schleep, B. C. 1819. Ueber eine bisher noch unbek- annte Mevenart.—Neue Annalen der Wetter- auischen Gesellschaft 1:314—320. Snyder, L. L. 1957. Arctic birds of Canada.—Uni- versity of Toronto Press, Toronto, 310 pp. Stevenson, H. M., and L. S. Atherton. 1984. Some comments about “white-winged” gulls in Flor- ida.— Florida Field Naturalist 12:99-103. Strang, C. A. 1977. Variation and distribution of Glaucous Gulls in western Alaska. — Condor 79: 170-175. Sutton, G. M. 1967. Oklahoma birds. University of Oklahoma Press, Norman, 674 pp. 159 Taczanowski, W. 1893. Faune ornithologique de la Sibérie orientale.—Memoires de 1l’Academie Imperial des Sciences de St. Petersbourg, no. 7. Todd, W.E.C. 1963. Birds of the Labrador Peninsula and adjacent areas. University of Toronto Press, Toronto, 819 pp. Vaurie, C. 1965. The birds of the Palearctic fauna. Non-Passeriformes. H. F. & G. Witherby, London, 763 pp. Witherby, H. F., F. C. R. Jordain, N. F. Ticehurst, and B. W. Tucker. 1941. The handbook of British birds. Vol. 5. H. F. & G. Witherby, London, 356 pp. Section of Biological Survey, Denver Wildlife Research Center, U.S. Fish and Wildlife Service, National Museum of Nat- ural History, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(1), 1986, pp. 160-185 THE SYSTEMATICS OF THE GENUS PACHYSTACHYS (ACANTHACEAE) Dieter C. Wasshausen Abstract.—In 1847, Nees von Esenbeck described the genus Pachystachys with a single species, P. riedeliana, from Brazil. Later that same year, he pub- lished six additional species including P. coccinea (Aublet) Nees, a species that has long been confused with P. riedeliana. Although the two species are su- perficially similar, they can readily be separated by the longer, linear calyx segments of P. coccinea. Recent field activities in South America have shown that in number of species and in morphological diversity, the genus reaches its zenith in Amazonian Peru. Nine new taxa are currently proposed, whereas four of Nees’ six species are reduced to synonymy. The genus is characterized by a prominent terminal spike of flowers with large, ringent, two-lipped corollas and two stamens that are about as long as the corolla, having equal, deeply sagittate anthers. Several species (P. coccinea, P. spicata) have been observed to be hummingbird pollinated, and the inflorescences of two additional species (P. ossolaea, P. puberula) are extensively covered with ants. However, floral morphology appears to preclude myrmecophily as a factor in the pollination system of the red-flowered rain-forested species of Pachystachys. All taxa have been studied in depth, including their pollen morphology, and analytical keys to the species have been prepared to facilitate the identification of the 12 species recognized in this treatment. The data from pollen morphology show only slight variation in the pollen of species of Pachystachys and support the reten- tion of the genus in Bremekamp’s (1938) tribe Odontonemeae subtribe Grap- tophyllinae. The genus Pachystachys Nees is a Neo- tropical genus of 12 species. A member of the subfamily Acanthoideae (=Imbricatae Lindau, 1893), these plants occur along edges of the lowland rainforest at relatively low elevations (below 1000 meters) from the West Indies southward to Peru and Bolivia and eastward to French Guiana and Ama- zonian Brazil. Members of Pachystachys are perhaps best known for their ornamental beauty, and many may be found in botanical gardens throughout the warmer regions of the world. Their prominent terminal spikes with bright red flowers are an impressive sight when the showy inflorescences reach full anthesis, and these shrubs or subshrubs are highly prized in the garden. Recent field activities in South America have shown that in number of species and in morphological diversity, the genus reach- es its zenith in Amazonian Peru. Since at- tempts to determine specifically the mate- rial from these collections proved unsatisfactory, I have decided to present a basic revision of the genus. History In 1847, Nees von Esenbeck described the genus Pachystachys with a single species, P. riedeliana, from Amazonian Brazil. Lat- er that same year in de Candolle’s Prodro- VOLUME 99, NUMBER 1 mus he published six additional species in- cluding P. coccinea, originally described as Justicia coccinea by Aublet in his “‘Histoire des plantes de la Guiane francoise” in 1775. A second epithet, Justicia spicata, pub- lished by Ruiz & Pavon (FI. Peruv. & Chil. Prodr. 1798) and based on one of their own collections from Peru, was cited by Nees in his second publication in synonymy under P. riedeliana. Unfortunately, this earlier ep- ithet (1798) has priority over riedeliana (1847), and therefore P. spicata becomes the correct name for this species. No further additions were made to the genus until 1947, when Carlos Rizzini de- scribed P. albiflora from Brazil. This species, collected in Acre, is synonymous with P. lutea from nearby Amazonian Peru. Morphology The plants are suffrutescent and climbing and are between | and 3 meters tall, with relatively large, entire or undulate, petiolate leaves. Their prominent terminal flowering spikes are often large and conspicuous, with variously shaped and colored bracts and with bright red, pinkish-red or white corollas. The large, showy corollas are ringent, slenderly obconic, curved, and two-lipped. The upper lip is usually erect or recurved, narrow and bilobed, whereas the lower lip is three-lobed, the lobes spreading, subequal, oblong or ovate. There are two fertile stamens; these are about as long as the upper corolla lip and are attached near the base of the corolla tube. Both anthers are deeply sagittate, with the sacs equal and muticous at the base. The pollen of Pachystachys is the typical tricol- porate, subprolate pollen characteristic of the tribe Odontonemeae. Here each colpus is flanked on either side by a pseudocolpus of approximately equal length. The colpi rarely fuse at one or both poles, whereas the pseudocolpi converge and are fused into three pairs just below the poles. The surface is commonly reticulate. While the pollen is 161 slightly variable, the variation does not ap- pear to be taxonomically significant. The red floral colors seem to suggest that the flowers are largely adapted to hum- mingbird-pollination, and indeed this is what I have personally observed in several of the species. Furthermore, nectar appears to be secreted by a cushion-like nectary which surrounds the base of the ovary. Another interesting phenomenon that I have observed is that the inflorescences of most specimens of Pachystachys, as well as cer- tain species of Aphelandra and Ruellia, are extensively covered with ants and homop- terous insects. It appears that the ants feed on the honeydew exudate produced by the armored scales and in return protect the plants from other indiscriminate feeders. Like most Acanthaceae, these taxa are of extreme local distribution in undisturbed forests and are rarely collected in secondary growth. There is no doubt in my mind that, as more field work is being undertaken in relatively uninhabited areas of Peru, addi- tional taxa will be discovered. Furthermore, the genus appears to be well suited for re- productive biology studies, especially pol- linator relationships and seed dispersal. Taxonomy Pachystachys Nees in Mart. Fl. Bras. 9:99. 1847. Type species: Pachystachys riede- liana Nees. Herbaceous or suffrutescent plants; leaves large, petioled; spikes terminal, dense; bracts conspicuous, herbaceous; bractlets small or none; flowers borne in terminal spike of ver- ticillasters consisting of 3 or 4 flowers each; calyx 5-parted, the segments relatively short; corolla ringent, slenderly obconic, curved, 2-lipped, upper lip narrow, 2-lobed at tip, lower lip 3-lobed, lobes subequal, oblong or ovate; stamens 2, about as long as corolla, attached near base of its tube; anthers deep- ly sagittate, basal lobes muticous, equal; staminodes, if present, rudimentary; cap- sule 4-seeded. 162 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Key to the Species of Pachystachys 1. Corolla white; bracts bright yellow or orange yellow ...................2.. P. lutea 1. Corolla red, crimson, pinkish-red or pink; bracts green, yellowish-green, green with purple at:tips.or brownishied, d4a0 9 eae A Aes ek. eee 2 2s BrACtS. 3=4..5) CHM IONG sce). wate daphan ss JEG OR eee Sk nest eee 3 3-_bracts elliptic-ovate,alis—ly7) mummwide ws see oe, eee P. longibracteata 3. Bracts oblanceolate, (6—8 mmb-wide .2i55205 2. bee). 252. eee 4 4. Leaf blades oblong to ovate, 8—8.5 cm wide; spikes 6—10 cm long; bracts 7-8 mm wide, terminated by mucro about 1 mm long, principal veins NHOtEpProniinernit +. 045s; Boeke. Des Thee ae ee P. schunkei 4. Leaf blades elliptic to oblong, 11—14.5 cm wide; spikes 19—20 cm long; bracts 6-7 mm wide, not terminated by mucro, three principal veins PLOMMIMNEHE ih. 48 Aa cea > Da ae ee, Bee oe ee P. fosterii 2.4Bracts less than:3- enmlongss. |. 4nc3.2 0 4b dna ee ee eee 5 5. Bracts brownish-red, lanceolate, 0.8 cm long, 1.5 mm wide ....P. badiospica 5. Bracts green, yellowish-green or green with purple at tips, lanceolate, ob- lanceolate, ovate or narrowly elliptic, 1.5—2.8 cm long, 6-18 mm wide .. 6 6-.Corolla.3d5=4em longa si. Ne eee 7 7. Bracts yellowish-green, membranous, sparingly glandular-pilose; co- rollaspini< upper lip erectae 22st oe See ee ee P. incarnata 7. Bracts dark-green, not membranous, densely puberulous; corolla CuIMSONAUppemlip recunved es ek ae aes ee P. puberula oF Corollas=7ssemdong ss et eS) Se ee ee eee 8 8. Calyx segments narrowly triangular, 3 mm long, 1 mm wide . P. spicata &. Calyx segments lanceolate, 5-10 mm long, 0.5-1.5 mm wide ..... 9 9. Calyx 9-12 mm long, segments 8—10 mm long; corolla pink, pink- ish=red: or,oranee-Tediyg! .. 0 oe ee ee oa oe 10 10. Bractlets minute, 2 mm long, 0.4 mm wide, glabrous; bracts Harrowly Ovate..2-2—2.5 cm long —~. 9)... ... 19. eee P. roseus 10. Bractlets 11 mm long, 1.25 mm wide, pilose; bracts broadly elliptical Cmiglone .f2 "05. a a os. e eee tee ae P. killipii 9. Calyx 5.5-7 mm long, segments 5 mm long; corolla bright red or CEUIMNSON ois 5 dics petra sixibong wn Meta rak eecee ous ocse eesti 4 a ee 11 11. Bracts green with purple at tip; corolla puberulous and glan- dular punctate, upper lip 2-lobed, lobes obtuse, 1.5 mm long, 2 am: Wide. s25..2.3 okt eee Seen tee ee P. ossolaea 11. Bracts green; corolla rather sparingly pubescent with minute spreading hairs (hirtellous), upper lip emarginate, lobes 1 mm long...0.5: mim, wide. .... eerie ee, seal, eee P. coccinea Pachystachys lutea Nees, DC. Prodr. 5-2. : len- 11:320. 1847 Erect, lax shrub 1.5—2.5 m tall; stem slen der, subterete, brownish, glabrous, the nodes Justicia lutea Ruiz & Pavonex Schult. Man- more or less tumid; leaf blades subsessile, tissa 1. 146. 1822, nom. nud. Pachysta- oblong-lanceolate, 8.5—18 cm long and 2-5 chys albiflora Rizzini, Bol. Mus. Nac. Rio cm wide, acuminate at apex, gradually nar- de Janeiro 8: 24, pl. 7. 1947. rowed from below middle to a rounded, VOLUME 99, NUMBER 1 clasping base, 1 cm wide in larger leaves, entire or undulate, membranous, glabrous except for puberulent midrib, cystoliths nu- merous and prominent both above and be- low; inflorescence consisting of terminal, solitary spike 6-10 cm long, rachis densely puberulous; bracts densely imbricate, bright yellow or orange-yellow, ovate, 1.5-—2 cm long, 1—1.5 cm wide (terminal bracts some- what smaller, basal ones larger), short-acu- minate at apex, subcordate at base, glan- dular-pilose, especially so along upper margins; bractlets lanceolate-spatulate to el- liptic, 9-12 mm long, 3.5—4 mm wide, acu- minate, glandular-pilose along margins; ca- lyx campanulate, 9-10 mm long, segments linear-lanceolate, 7-9 mm long and 1-1.1 mm wide near base, sparingly puberulous and ciliate; corolla white, 5.5 cm long, tube curved, 3 mm wide at base, narrowed to 1.75 mm at 5 mm above base, thence grad- ually enlarged to 8.5 mm at throat, pilose and glandular-punctate, upper lip erect, lan- ceolate, 1.8 cm long, 5—5.5 mm wide near base, gradually narrowed to | mm at tip, sparingly pilose, minutely 2-lobed, lobes obtuse, lower lip ovate, undulate, sparingly pilose, 3-lobed, lateral lobes 3.5 mm long, 2 mm wide, obtuse or rounded, middle lobe 3.5 mm long, 2.5 mm wide, obtuse or rounded; stamens attached near base of co- rolla tube, filaments 5 cm long, sparingly puberulous, anthers bright green, 5 mm long, deeply sagittate, basal lobes muticous; pollen grains 53 um long, 45 um wide, tricolporate, each colpus flanked by 2 pseudocolpi, tec- tum almost complete, psilate-punctate (Fig. 1); staminodes none; ovary glabrous; cap- sule clavate, 13 mm long, 5 mm broad, 2.75 mm thick, glabrous; retinacula 2 mm long, slightly curved, tip flattened, obtuse; seeds 2, brownish, cordate, flattened, about 4.5 mm long and 4 mm broad, glabrous. Material.—PERU: San Martin: Lamas, Matthews 1538 (syntype K); On trail from Lamas to San Antonio E of Rio Chupisena, Belshaw 3500 (NY, UC, US); Pongo de Cainarachi, Rio Cainarachi, 230 m, Klug 163 2655 (NY, US); Juanjui, 400 m, Klug 3838 (NY, US); Prov. Mariscal Caceres: Dtto. Tocache Nuevo: Quebrada de Huaquisha, 500-600 m, Plowman, Schunke & Rury 11389 (F, US); Quebrada de Canuto, ca. 500 m, Schunke-Vigo 12083 (NY, US); Que- brada de Saule Chico, Schunke-Vigo 4346 (F, NY, US); Between Uchiza and Puerto Huicte, 500-600 m, Ferreyra 4410 (US, USM); Fundo Melddia, road to Shunté, 800 m, Schunke-Vigo 7436 (US); Hudanuco: Chicolaya, Tafalla 345 (syntype G, MA); Prov. Tingo Maria: Vicinity of Tingo Ma- ria, 650-700 m, Ferreyra 6772 (US, USM): 625-1100 m, Allard 21622 (US); Junin: Near La Merced, Soukup 2531 (US); Cuzco: Prov. La Convencion: Hda. Luisiana, 620 m, Dudley 11525 (NA). BRAZIL: Acre: Varadouro S. Luiz, Kuhlmann 728 (R, ho- lotype of P. albiflora Rizz.). At edge of ravines in dense forests, some- times saxatile on very steep dry cliffs, usu- ally found on banks of streams at elevations between 230 and 1100 meters. This native of Peru is often found in cultivation in greenhouses. Pachystachys longibracteata Wasshausen, sp. nov. igsaleyZ Suffrutex; caules subquadrangulares, gla- bri; lamina foliorum elliptica, acuminata, basi angustata in petiolum amplectens, glabra; spicae solitariae, terminales, rhach- idi puberula; bracteae grandes, viridae, el- liptica-ovatae, 15—17 mm latae, apice acu- minatae vel obtusae; bracteolae lineari- lanceolatae; calycis segmenta anguste trian- gularia; corolla coccinea vel scarlatina, pu- berula, labio inferiore oblongo, lobo medio conduplicato apice curvato, lobis laterali- bus leviter rotundatis. Shrub 1-3 m high; stems subquadran- gular, glabrous; leaf blades elliptic, to 35 cm long and 13 mm wide, acuminate at apex, narrowed from about middle to rounded, clasping base 2—3 cm wide in larger leaves, 164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. SEM photomicrographs of Pachystachys pollen. A—-B, Pachystachys lutea (Mennega 7 1-136); C—D, Pachystachys longibracteata (Killip & Smith 28725). A, Polar orientation, < 1800; B, Portion of surface, including colpus at x 4600; C, Polar orientation, x 1700; D, Surface view of reticulate form of fractured grain, x 6600. entire or undulate, glabrous, cystoliths nu- merous, about 0.25 mm long; petioles to 7 cm long, glabrous; inflorescence to 23 cm long, internodes, bracts, calyx, and ovary densely but minutely puberulent; bracts green, elliptic-ovate, 3—4.5 cm long and 15-— 17 mm wide, acuminate to obtuse at apex, narrowed at base to short winged petiole; bractlets linear-lanceolate, to 17 mm long and 3 mm wide; calyx 2—3 mm long, seg- ments narrowly triangular; corolla scarlet, sparingly and minutely puberulent, 6 cm VOLUME 99, NUMBER 1 165 LOX SS OSS Fig. 2. A-E, Pachystachys longibracteata (Killip & Smith 27866). A, Flowering spike; B, Leaf blade; C, Bract and calyx; D, Corolla and exserted stamens and style; E, Corolla expanded, showing attachment of filaments. long, 3 mm broad at base, narrowed to 2 mm at 5 mm above base, and then gradually narrowed to 8 mm at throat, lobes minutely ciliate, those of lower lip oblong, about 13 mm long, 6 mm broad, two lateral rounded at top, middle conduplicate and curved at tip, upper lip 2 cm long, 4 mm at base, gradually narrowed to small, bilobed tip 166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON about 1 mm broad; filaments glabrous, an- thers about 4 mm long; pollen grains 58 um long, 40 wm wide, tricolporate, each me- socolpium flanked by two colpoid streaks, membrane densely granular, sexine su- prareticulate, reticulation heterobrochate (Fig. 1); style glabrous, persistent after fall of corolla; capsule wanting. Type.—PERU: Loreto: Puerto Arturo, lower Rio Huallaga below Yurimaguas, ca. 135 m, Killip & Smith 27866 (holotype US; isotype NY). Additional specimens.—PERU: Loreto: Santa Rosa, lower Rio Huallaga below Yu- rimaguas, ca. 135 m, Killip & Smith 28725 (NY, US); Yurimaguas, lower Rio Huallaga, ca. 135 m, Killip & Smith 28055 (US); Shi- tari, near Isla Santa Maria, Yurimaguas, 150-180 m, Ferreyra 10159 (US, USM). Occasional in dense forest at elevations of 135-180 meters. This species is well marked and distin- guished by its densely puberulous inflores- cence and large leaf blades with their bases clasping the petiole and the unusually broad or short lobes of the lower lip of the corolla. Pachystachys schunkei Wasshausen, sp. nov. Fig. 3 Suffrutex; caules subquadrangulares, bru- neoli, puberuli; lamina foliorum oblonga vel ovate, acuminata, basi angustata in petio- lum amplectens, glabra; spicae terminales, bifurcatae, aliquanto breves, rachidi dense puberula; bracteae viridae, oblanceolatae, acuminatae et minute mucronatae; bracteo- lae linear aristatae; calycis segmenta lan- ceolata, puberula, ciliolata; corolla cocci- nea, glabra, tubo leviter curvato, labio superiore recurvato bilobato, labio inferiore 3-lobato, lobis obovatis, lobo medio con- duplicato. Shrub 2-3 m tall; stem subquadrangular, brownish, puberulous; leaf blades oblong to ovate, 19-22 cm long and 8-8.5 cm wide, acuminate at apex, gradually narrowed from below middle to a rounded, clasping base 1.5 cm wide in larger leaves, entire or un- dulate, thin, glabrous, shining, cystoliths in- conspicuous; petioles 3—8.5 cm long, gla- brous; inflorescence consisting of terminal, bifuricate spike 6-10 cm long, rachis dense- ly puberulous; bracts green, oblanceolate, 3.5—4 cm long, 7-8 mm wide, acuminate, terminated by mucro about | mm long, gradually narrowed at base and attenuate, densely gladular pilose without, minute eglandular hairs intermixed with glandular ones, principal veins not prominent; bract- lets linear-aristate, about 2 cm long, 1 mm wide, pubescence similar to that of bracts; calyx campanulate, 4.5 mm long, segments lanceolate, 4 mm long and 1 mm wide near base, acuminate, densely puberulous and ciliate; corolla vivid red, 5—5.5 cm long, tube slightly curved, 4 mm wide at base, nar- rowed to 2 mm at 5 mm above base, thence gradually enlarged to 6.5 mm at throat, gla- brous, upper lip recurved, oblong to nar- rowly lanceolate, 1.4 cm long, 2.5 mm wide just below middle, gradually narrowed to 1.5 mm at tip, minutely 2-lobed, lobes ob- tuse, lower lip 3-lobed, lobes spreading, ob- ovate, rounded at tip, lateral pair 13 mm long, 6 mm wide, middle lobe 9 mm long and 4.5 mm wide, conduplicate; stamens attached near middle of corolla tube, fila- ments 3 cm long, sparingly and inconspic- uously puberulous, anthers 4 mm long, deeply sagittate, basal lobes muticous; staminodes none; ovary puberulous; cap- sule wanting. Type.-PERU: San Martin: Prov. Maris- cal Caceres: Dtto. Tocache Nuevo, Quebra- da de Santa Rosa de Cachiyacu, 500-700 m, Schunke-Vigo 7597 (Holotype US). Occasional in tall forest. Discussion. —Pachystachys schunkei is perhaps nearest in relationship to P. fosteri, another new species from Peru, but differs markedly in that the latter species has leaf blades elliptic to oblong, 11—-14.5 cm wide, spikes 19-20 cm long, and prominently veined bracts 6-7 mm wide, not terminated 167 VOLUME 99, NUMBER 1 UN RK SS ah % ae ey, | Zi) et hy Nee su Oe, SSS. SSE SSSR Z| Sasi I) cS) FF | ! f ii y a f ) SS if = a fH } | S ss | mS ’ SO 4 SSS Ss Fig. 3. A-D, Pachystachys schunkei (Schunke-Vigo 7597); E-I, Pachystachys badiospica (Schunke- Vigo 2714). A, Habit; B, Bract, bractlets and calyx; C, Corolla and exserted stamens and style; D, Corolla expanded, showing attachment of filaments; E, Habit; F, Bract and bractlets; G, Calyx; H, Corolla closed; I, Corolla expanded, showing attachment of filaments. 168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON by a mucro. In contrast, H. schunkei has leaf blades oblong to ovate, 8-8.5 cm wide, spikes 6—10 cm long, and bracts not prom- inently veined, 7-8 mm wide, terminated by a mucro about 1 mm long. Pachystachys fosteri Wasshausen, sp. nov. Fig. 4 Suffrutex; caules subquadrangulares, bru- neoli, glabri; lamina foliorum elliptica vel oblonga, acuminata, basi angustata in pe- tiolum amplectens, glabra; spicae termi- nales, bifurcatae, 19-20 cm longae, rachidi dense puberula; bracteae viridae, oblanceo- latae, acuminatae, glanduloso-puberulae, venis 3 principalibus prominentibus; brac- teolae lineares, puberulae; calycis segmenta lanceolata, puberula; corolla scarlatina, puberula, labio superiore erecto et recur- vato bilobato, labio inferiore trilobato, lobis obovatis, lobo medio conduplicato et re- tuso. Shrub 2-3 m tall; stem subquadrangular, brownish, glabrous; leaf blades elliptic to oblong, 33 cm long and 11-14.5 cm wide, acuminate at apex, gradually narrowed from below middle to a rounded, clasping base 2 cm wide in larger leaves, entire or undulate, glabrous, cystoliths numerous, especially below; petioles 3—9 cm long, glabrous; in- florescence terminal, consisting of bifuricate spike 19-20 cm long, rachis densely pu- berulous; bracts green, oblanceolate, 3.3 cm long, 6-7 mm wide, acuminate at apex, gradually narrowed at base and attenuate, densely glandular puberulous without, cil- late, three principal veins prominent; bract- lets linear, 2.3 cm long, 1 mm wide, densely glandular-puberulous and ciliate; calyx campanulate, 6 mm long, segments lanceo- late, 4.5 mm long and 1 mm wide near base, acuminate, densely puberulous and ciliate; corolla scarlet, 5 cm long, tube slightly curved, 3-3.5 mm wide at base, narrowed to 2 mm at 4 mm above base, thence grad- ually enlarged to 8 mm at throat, puberu- lous and occasionally interspersed with mi- nute glands, upper lip erect and recurved, oblong to narrowly lanceolate, 1.2 cm long, 2.5 mm wide just below middle, gradually narrowed to 1.5 mm at tip, minutely 2-lobed, lobes obtuse, lower lip 3-lobed, lobes spreading, obovate, lateral pair 12 mm long, 5 mm wide, rounded at tip, middle lobe 9 mm long and 6-6.5 mm wide, con- duplicate and retuse at tip; stamens attached near middle of corolla tube, filaments 3.5 cm long, sparingly puberulous, anthers 4.7 mm long, deeply sagittate, basal lobes mu- ticous; staminodes none; ovary densely to- mentose; capsule wanting. Type. —PERU: Huanuco: Pachitea; Puer- to Inca, 2-5 km E of town, 250-300 m, Foster 8686 (holotype US). Occasional, along forest trails. Discussion. —Named in honor of the col- lector, Robin B. Foster, in recognition of his valuable contributions to our knowledge of Peruvian botany. The major differences be- tween Pachystachys fosteri and P. schunkei, its nearest relative, are described under the discussion of the latter species. Pachystachys badiospica Wasshausen, sp. nov. Figs. 3, 5 Suffrutex; caules subquadrangulares, brunneoli, glabri; lamina foliorum elliptica vel ovata, acuminata, basi angustata in pet- iolum amplectens, glabra; spicae termi- nales, solitariae vel bifurcatae, rachidi dense puberula; bracteae badiae lanceolatae, mu- cronulatae, extus dense glanduloso-pilosae; bracteolae lanceolatae; calycis segmenta subaequalia, lanceolata, parce glanduloso- pilosa; corolla coccinea, parce pilosa, tubo leviter curvato, labio superiore erecto, bi- lobato, labio inferiore trilobato, lobis later- alibus oblongis, obtusis, lobo medio retuso; antherae rubrae. Shrub 1-2 m tall; stem subquadrangular, brownish, glabrous; leaf blades petiolate, el- liptic to ovate, 15—26 cm long and 5.5-11.5 cm wide, acuminate at apex, narrowed from VOLUME 99, NUMBER 1 169 MN Sos : i WA | i \ =z —— E_ —_——4 = ZF = _ HZ Fig. 4. A-E, Pachystachys fosteri (Foster 8686); F-I, Pachystachys rosea (Vargas C, 23159). A, Habit; B, Bract and bractlets; C, Calyx; D, Corolla and exserted stamens and style; E, Corolla expanded, showing attachment of filaments; F, Habit; G, Bractlets and calyx; H, Corolla and exserted stamens and style; I, Corolla expanded, showing attachment of filaments. 170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. SEM photomicrographs of Pachystachys pollen. A-B, Pachystachys badiospica (Schunke- Vigo 27 14); C-D, Pachystachys incarnata (Killip & Smith 28926). A, Equatoiral view, x 1700; B, Portion of surface, including pore at x3000; C, Equatorial view, x 2000; D, Portion of surface, including pore at 3700. below middle to rounded, clasping base 15 mm wide in larger leaves, moderately firm, entire or undulate, upper surface dark-green, nitid, lower surface paler green, glabrous, cystoliths numerous and conspicuous both above and below; petioles 2—6 cm long, can- aliculate, glabrous; inflorescence consisting of solitary or bifuricate, terminal spike 10- 16 cm long, rachis densely glandular pu- berulous; bracts brownish-red, lanceolate, 8 mm long, 1.5 mm wide, mucronulate at apex, densely glandular-pilose without, mi- VOLUME 99, NUMBER 1 nute eglandular hairs intermixed with glan- dular ones; bractlets lanceolate, 6.5 mm long, 1.25 mm wide, pubescence similar to that of bracts; calyx deeply 5-parted, 15 mm long, segments lanceolate, subequal, sparingly glandular-pilose, posterior segment 9 mm long, 1 mm wide, anterior pair 13 mm long, 1.5 mm wide, lateral pair 12 mm long, 1.25 mm wide; corolla red, 5 cm long, sparingly pilose, tube slightly curved, 5 mm wide at base, narrowed to 3 mm at 5 mm above base, thence gradually enlarged to 18 mm at throat, upper lip erect, oblong to narrowly lanceolate, 1.5 cm long, 4 mm wide, mi- nutely 2-lobed, lobes obtuse, lower lip ovate, 1 cm long, 3-lobed, lateral lobes oblong, 7 mm long, 3 mm wide, obtuse or rounded, middle lobe 8 mm long, 6 mm wide, retuse; stamens attached near base of corolla tube, filaments 2.7 cm long, sparingly puberulous, anthers red, 4.5 mm long, deeply sagittate, basal lobes muticous, pollen grains 50 wm long, 41 um wide, tricolporate, each me- socolpium with two colpoid streaks, tectum almost complete, psilate-punctate (Fig. 5); staminodes none; ovary glabrous; capsule clavate, long-stipitate, 2 cm long, puberu- lous, 2-seeded, solid basal portion about 1 cm long, seed-bearing cavity 8 mm broad, seeds flattened, orbicular, 7 mm long and broad, muricate, retinacula 3 mm long, slightly curved, tip flattened, obtuse. Type.—PERU: JUNIN: Satipo, Ridoutt s.n. (holotype US; isotype USM). Additional specimens. —PERU: Loreto: Prov. Coronel Portillo, Dtto. Iparia, Bosque Nacional de Iparia, along Rio Ucayali in the vicinity of Iparia, 300 m, Schunke V. 2714 (US). JUNIN: road between Santa Clara and Satipo, Ridoutt s.n. (US, USM). In openings and along the edge of dense, humid forests. Discussion.—This species is readily dis- tinguished by its narrow, brownish-red, lan- ceolate, glandular-pilose bracts, bractlets, and calyx segments. It is not very closely allied to any of the known species of Pach- ystachys. 171 Pachystachys incarnata Wasshausen, sp. nov. Figs. 5, 6 Frutex, caules erecti, teretes, brunneolli, glabri; lamina foliorum elliptica vel ovata, acuminata, basi angustata in petiolum am- plectens, membranacea, nitida, glabra; spi- cae terminales, rachidi dense puberula; bracteae dense imbricatae, stramineae, ova- tae, mucronulatae, parce glanduloso-pilo- sae; bracteolae lineari-lanceolatae, puberu- lae; calyx campanulatus, segmenta lanceolata, puberula; corolla rosea, puber- ula, tubo leviter curvato, labio superiore erecto, bilobato, labio inferiore trilobato, lobis patulis, lobis lateralibus oblongis vel anguste ovatis, rotundatis, lobo medio obovato, conduplicato et retuso. Erect shrub about 2 m tall; stem terete, brownish, glabrous; leaf blades elliptic to ovate, 20-25 cm long and 7.5—8.2 cm wide, acuminate at apex, gradually narrowed from below middle to a rounded, clasping base 1 cm wide in larger leaves, entire or undulate, membranous, nitid and dark-green above, much lighter-green below, glabrous, cysto- liths numerous and prominent both above and below; petioles 2-3 cm long, minutely puberulous; inflorescence consisting of ter- minal, solitary spike 13 cm long, rachis densely puberulous; bracts densely imbri- cate, yellowish-green, ovate, 2 cm long, 1.4 cm wide, mucronulate at apex, gradually narrowed and cuneate at base, sparingly glandular-pilose, especially near apex; bractlets linear-lanceolate, 11 mm long, 1.25 mm wide, puberulous and ciliate, bearing few glandular trichomes near apex; calyx campanulate, 12 mm long, segments lan- ceolate, 10 mm long and 1 mm wide near base, acuminate, puberulous and ciliate; co- rolla pink, 4 cm long, puberulous, tube slightly curved, 4 mm wide at base, nar- rowed to 1.8 mm at 5 mm above base, thence gradually enlarged to 6.5 mm at throat, up- per lip erect, oblong to narrowly lanceolate, 1.2 cm long, 3.5—-4 mm wide near base, 172 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON £11JTP laa SS SS y= INGEN | SSS ~=S SS I yf Yf, ¢ |! A= Wh Fig. 6. A-—D, Pachystachys incarnata (Killip & Smith 28926); E-G, Pachystachys killipii (Killip & Smith 2661). A, Habit; B, Bractlets and calyx; C, Corolla and exserted stamens; D, Corolla expanded, showing at- tachment of filaments; E, Habit; F, Bractlets and calyx; G, Corolla and exserted stamens and style. VOLUME 99, NUMBER 1 gradually narrowed to | mm at tip, minutely 2-lobed, lobes obtuse, lower lip 3-lobed, lobes spreading, lateral pair oblong to nar- rowly ovate, 15 mm long, 5 mm wide, rounded at tip, middle lobe obovate, 12 mm long and 4.5 mm wide, conduplicate and retuse, minutely lobulate on one side; sta- mens attached near base of corolla tube, filaments 4 cm long, minutely puberulous, anthers 5 mm long, deeply sagittate, basal lobes muticous; pollen grains 46 um long, 28 wm wide, tricolporate, each mesocol- pium with two colpoid streaks, tectum al- most complete, psilate-punctate (Fig. 5); staminodes none; ovary puberulous; cap- sule wanting. Type.— PERU: Loreto: Santa Rosa, lower Rio Huallaga below Yurimaguas, ca. 135 m, Killip & Smith 28926 (holotype US; iso- type NY). Occasional in dense forest. Discussion. —Pachystachys incarnata su- perficially resembles P. /utea Nees. How- ever, in P. /utea, the corolla is white, 5.5 cm long, the bracts bright yellow or orange-yel- low, and the leaf blades are oblong-lanceo- late, 8.5-18 cm long and 2-5 cm wide. In P. incarnata, the corolla is pink, only 4 cm long, the bracts yellowish-green, and the leaf blades are elliptic to ovate, 20-25 cm long and 7.5—8.2 cm wide. Pachystachys puberula Wasshausen, sp. nov. Figs. 7, 8 Suffrutex; caules subquadrangulares, brunnei, glabri; lamina foliorum oblonga vel ovata, breviter acuminata, basi angustata in petiolo amplectens, atrovirens et nitida, gla- bra; spicae terminales, solitariae, rachidi subtiliter puberula; bracteae laxe imbrica- tae, atrovirens, late ovatae, obtusae et api- culatae, basi angustatae ad petiolos brevi- alatae; calyx campanulatus, segmenta anguste triangularia, dense puberula; corol- la carminea, puberula, tubo leviter curvato, labio superiore erecto et recurvato, bilo- 173 bato, labio inferiore trilobato, lobis obo- vatis, patulis, lobo medio conduplicato ap- ice obtuso. Shrub 2-5 m tall; stem subquadrangular, dark-brown, glabrous; leaf blades very dark- green and glossy, oblong to ovate, 15—28 cm long and 7.5-13.5 cm wide, short-acumi- nate at apex, gradually narrowed from be- low middle to a rounded, clasping base 1 cm wide in larger leaves, entire or undulate, glabrous, cystoliths numerous, especially below; petioles 3-8 cm long; inflorescence consisting of solitary, terminal spike 7-15 cm or more long, rachis finely puberulous; bracts rather loosely imbricate, dark-green, broadly ovate, 1.5—2.5 cm long, 10-18 mm broad (terminal bracts somewhat smaller, basal ones larger), rounded and apiculate at apex, narrowed at base to short-winged pet- iole, densely puberulous; bractlets lanceo- late, about 4 mm long and 1.2 mm broad, densely puberulous; calyx campanulate, 3.5 mm long, segments narrowly triangular, about 2 mm long and | mm wide at base, acuminate, densely puberulous; corolla crimson, 3.5—4 cm long, tube slightly curved, 3.5—4 mm wide at base, narrowed to 2 mm at 7 mm above base, thence gradually en- larged to 7 mm at throat, puberulous, upper lip erect and recurved, lanceolate, 1.5 cm long, 3.5 mm wide near base, gradually nar- rowed to | mm at tip, minutely 2-lobed, lobes obtuse, lower lip 3-lobed, spreading, lobes obovate, rounded at tip, lateral ones 12 mm long, 6 mm broad, middle lobe 15 mm long and 5.5 mm broad, conduplicate, obtuse at tip; stamens attached near base of the corolla tube, filaments 3.5—4.3 cm long, glabrous, anthers 3.8-4 mm long, deeply sagittate, basal lobes muticous; pollen grains 52 um long, 35 wm wide, tricolporate, each mesocolpium flanked with two colpoid streaks, tectum almost complete, psilate- punctate (Fig. 8); staminodes none; ovary puberulous; capsule wanting. Type.—PERU: San Martin: Prov. Mar- tin: Puente Colombia, 24 km S of Tarapoto, 174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sgh ge , ti Hi) l] Gi) S \ LTA HH) My HY h. YS Fig. 7. A-D, Pachystachys puberula (Wasshausen & Encarnaci6n 1023). A, Habit; B, Calyx; C, Corolla and exserted stamens and style; D, Corolla showing attachment of filaments. 350-550 m, Wasshausen & Encarnaci6n 1023 (holotype US, isotypes USM, US-2). Additional specimens. —PERU: San Mar- tin: Prov. Lamas: trail San José de Sisa- Agua Blanca, 380-400 m, Ferreyra 7946 (US, USM); between Agua Blanca and Des- quito, 600-800 m, Ferreyra 7963 (US, USM); Prov. San Martin: Tarapoto, 830 m, VOLUME 99, NUMBER 1 Woytkowski 35110 (US); Juan-Guerra, Tarapoto, Ule 6489 (G, K); near Tarapoto, Spruce 3974 (K); Pucayacu, 11 km S of Ta- rapoto, 200-250 m, Ferreyra 7730 (US, USM); Granja El Porvenir, 25 km S of Ta- rapoto, 400-500 m Ferreyra 17855 (US, USM); Pucacaca, Plowman 6016A (GH, US); Dtto. Shapaja: Shapaja, 264 m, Schunke- Vigo 9815 (MO); 1-4 km S of Sha- paja, ca. 300 m, Belshaw 3133 (UC, US); Puente Colombia, 200-250 m, Ferreyra 17545 (US, USM); Prov. Huallaga: between Bellavista and Banos, 200—300 m, Ferreyra 4731 (US, USM); between Juanjui and Tin- go de Saposoa, 200-300 m, Ferreyra 4793 (US, USM); near Bellavista, 250-300 m, Ferreyra 10099 (US, USM); Prov. Mariscal Caceres: vicinity of Juanjui, 300-400 m, Ferreyra 4498 (US, USM); 4538a (US, USM). Distributed in open places and along mar- gins of trails in lowland rainforest. Discussion. —Pachystachys puberula is distinguished from P. spicata in that the co- rollas of the former species are 3.5-4 cm long, the upper corolla lip is recurved and 3.5 mm broad near base, the lower corolla lip lobes are obovate with the lateral ones 12 mm long and 6 mm broad and the bracts are broadly ovate, 10-18 mm broad, nar- rowed and apiculate at apex. In P. spicata the corollas are 5—7 cm long, the upper co- rolla lip is erect and 6.5 mm broad near base, the lower corolla lip lobes are linear with the lateral ones 17—24 mm long and 4 mm broad and the bracts are ovate-lanceo- late, 7-11 mm broad, acuminate at apex. Pachystachys spicata (R.&P.) Wasshausen, comb. nov. Justicia spicata Ruiz & Pavon, FI. Peruv. & Chil. Prodr. 1:8. pl. 9. 1798. Pachystachys riedeliana Nees in Mart. FI. Bras. 9:99. 1847. P. latior Nees in DC. Prodr. 11: 320. 1847. P. asperula Nees in DC. Prodr. 11:320. 1847. 175 Suffrutescent, erect, simple or sparingly branched plant 1-5 m tall; stem glabrous, terete or with upper portions subquadran- gular; leaves petioled, blades oblong, elliptic or broadly oblanceolate, 15—27 cm long and 7-11 cm wide, acute to acuminate, tip itself usually blunt, gradually to rather abruptly narrowed at base, glabrous, firm, margins entire, costa and lateral veins (9-12 pairs) rather prominent; petioles 2.5—7 cm long, glabrous; spikes solitary, terminal, 12—22 cm long, rachis finely puberulous; bracts im- bricate, green, ovate-lanceolate, 1.5—2.5 cm long, 7-11 mm broad, terminal bracts somewhat smaller, basal ones larger, acu- minate, narrowed at base, short-petiolate, both surfaces puberulous, lower surface more densely so, bearing additional glan- dular hairs; bractlets narrowly linear, about 3 mm long and 0.5 mm broad, sparingly puberulous; calyx campanulate, 4.5 mm long, segments narrowly triangular, about 3 mm long and | mm wide at base, acumi- nate, sparingly hirtellous; corolla scarlet, be- coming orange-red with age, 5.5—7 cm long, glabrous to sparingly hirtellous, tube curved, 4 mm wide at base, gradually enlarged to 8 mm at throat, upper lip erect, lanceolate, 6.5 mm broad near base, gradually nar- rowed to 1.5 mm at tip, 2-lobed, lobes ob- tuse, 1 mm long and wide, lower lip 3-lobed, lobes spreading, linear, rounded at tip, mid- dle lobe 17 mm long and 3 mm broad, lat- eral ones 1 7—24 mm long and 4 mm broad; stamens attached near base of corolla tube, filaments yellow, about 5 cm long, puber- ulous, anthers yellow, 7 mm long, deeply sagittate, basal lobes muticous; pollen grains 61 um long, 35 wm wide, tricolporate, each mesocolpium with two colpoid streaks, membrane densely granular, sexine supra- reticulate, reticulation heterobrochate (Fig. 8); staminodes rudimentary, puberulous; ovary glabrous; capsule clavate, 16 mm long, 5 mm broad, 4.5 mm thick, glabrous; reti- nacula 3 mm long, slightly curved, tips flat- tened, obtuse; seeds 4, brownish, cordate, 176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. SEM photomicrographs of Pachystachys pollen. A-B, Pachystachys puberula (Wasshausen & Encar- naci6n 1023); C-—D, Pachystachys spicata (Calder6n 2826). A, Equatorial view, x 1400; B, Portion of surface, including pore at x 4000; C, Polar orientation, x 1800; D, Portion of surface, including pore at 4400. flattened, 5.5 mm long, 4 mm wide, gla- brous, muricate. Material. —CUBA: Oriente: Monte Verde, Le6n 3832 (NY); Santiago, banks of Canas River, El Cobre, Clemente 5885 (NY, US); Sierra Maestra, Finca la Philanthropia, Ekman 15697 (NY, S, US); Sierra Maestra, Loma del Gato, banks of Canas River, Clement 562 (NY), La Perla to Santa Ana, 660 m, Shafer 8592 (NY, US). JAMAICA: Westmoreland: Bluefields Mountains, 650 m, Britton & Hollick 1975 (NY). Man- chester: Mandville, Smith s.n. (US). Clar- endon: 1 m NW of Rock River, 165-230 m, Proctor 32842 (NY). St. Catherine: Bog Walk, Crawford 814 (NY). St. Mary: Cas- telton district, 150 m, Harris 11933 (NY, US); Metcalf district, Constant Spring to Annatto Bay, Britton 825 (NY). St. Andrew: Manning Hill, 380 m, Powell 778 (NY, US); Stoney Hill, Smith s.n. (US). Portland: vi- cinity of Mill Bank, 200-300 m, Maxon & VOLUME 99, NUMBER 1 Killip 202 (US); vicinity of Windsor, 150- 175 m, Maxon & Killip 271 (NY, US); Blue Hole, Fosberg 59422 (US); near Long Bay, 15 m, Gentry 28287 (MO, NY). Without exact locality: Fredholm 3229 (US). LESS- ER ANTILLES: Guadeloupe: Sainte Rose, Quentin 1936 (P, US); Trois-Rivieres, Stehlé 1855 (US). Dominica: Carib Reserve, Hodge 3307 (US); Geneva, Nicolson 2172 (US); between Fond Baron Estate and Pichelin, Ernst 1591 (US); Lisdara, Hodge 2326 (GH, NY); road to Morne Plat Pays, Gillis 8118 (FTG, US); Portsmouth, Hodge 847 (NY, US); Salybia, Hodge 846 (GH); along Stew- art’s River near its mouth on Grand Bay, Eggers 641 (C), Wilbur et al. 8027 (US). Martinique: Balata to Trivoli, 250 m Stehlé 5742 (US); La Gracée, Rodriguez 3622 (US). St. Vincent: along Chateaubelair River, 300— 400 m, Morton 5104 (US); 160 m, Smith & Smith 981 (NY). Grenada: Mt. Pleasant, 500 m, Miller 160 (US). TRINIDAD: Caura River Valley, Britton & Hazen 1203 (NY, US); E of Sangre Grande near Cunapi River, Crosby 4 (DUKE, US); edge of Aripo Sa- vannah, Howard 10350 (NY, US); Arima Valley Road, North Range, 150-600 m, Cowan & Simmonds 1221 (US); 15.8 km above Arima, Harriman 17598 (OSH, US); Maracas waterfall, 160 m, Kallov B152 (NY). Without exact locality: Lockhart s.n. (K, holotype of P. asperula Nees); 1877-80, Fendler 534 (BM, NY); 1874, Kuntze 598 (NY, US); 1889, Broadway 3435 (US). GUYANA: Without exact locality: Parker s.n. (K, holotype of P. latior Nees). CO- LOMBIA: Meta: Rio Duida, Mt. Macarena, 330 m, Gilliard s.n. (NY); Llanos Orien- tales, Villavicencio to Guayuriba, 600 m, Garcia- Barriga 18944 (AMES). El Valle: Las Juntas, on the Rio Dagua, Lehmann 1887 (US); Cisneros, 300-500 m, Killip 35590 (US). ECUADOR: Esmeraldas: Rio Santia- go at Concepcion, 30 m, Holm-Nielsen et al. 25976 (AAU). Manabi: Balao, Eggers 14135 (US); Olmedo, 100 m, Haught 3484 (NY, US). Guayas: 3 km E of Olon, 75-190 m, Dodson & Thien 1660 (US); 2-4 km E NYY from Recinto Olon, Gentry 10045 (MO, US). Los Rios: near Quevedo, Canton Vinces, ca. 50 m, Mexia 6609 (US). El Oro: between Santa Rosa and La Chorita, 0-100 m, Hitchcock 21116 (NY, US). Pichincha: Santo Domingo, 570 m, Dodson & Thien 1626 (US); Rio Blanco below confluence with Rio Toachi, 300 m, Harling 4490 (S, US); Patricio Pilar, 45 km S of Santo Do- mingo, 400 m, Holm-Nielsen 16017 (AAU). Chimborazo: base of Volcan Chimborazo, 900 m, Spruce s.n. (K). Pastaza: Shiguaco- cha, ca. 5 km E of Puerto Sarayacu, Lugo 3852 (GB, US). Without exact locality: Pearce s.n. (K); Gilmartin 367 (US). PERU: Amazonas: Prov. de Bagua: Rio Maranon opposite Quebrada Mirana, 425-450 m, Wurdack 2010 (NY, US, USM). San Mar- tin: Prov. Moyobamba: Moyobamba, Ma- thews 1537 (K). Prov. Lamas: along Rio Mayo, Spruce 4872 (K); Lamas, 600-800 m, Ferreyra 17333 (US, USM); between Ta- balosos and Lamas, ca. 500 m, Belshaw 3414 (K, NY, UC, US). Prov. Huallaga: Saposoa, 200-300 m, Ferreyra 4614 (US, USM); 350 m, Sagastegui 6837 (US). Prov. Mariscal Caceres: Juanjui, Sandeman 35 (K); Dtto. Tocache Nuevo: Quebrada Cachiyacu de Huaquisha, ca. 500-600 m, Schunke- Vigo 12515 (US); Quebrada de Saule Chico, Schunke- Vigo 4349 (NY, US); Fundo “‘Cu- rare Land,” propriedad de José-Schunke V., 500-525 m, Schunke- Vigo 10951 (MO, US). Loreta: Prov. Maynas: San Antonio, on Rio Itaya, ca. 110 m, Killip & Smith 29426 (NY, US); Soledad, on Rio Itaya, ca. 110 m, Killip & Smith 29683 (NY, US); Caballococha, Williams 2367 (US); below mouth of Rio Ucayali, 130 m, Gentry et al. 29999 (MO, US); Rio Nanay, Torres 0224 (AMAZ, US). Prov. Alto Amazonas: Pumayacu, between Balsapuerto and Moyabamba, 600-1200 m, Klug 3219 (NY, US). Prov. Coronel Portil- lo: bank of Rio Neshuya, km 61 on Federico Basache road, Encarnacion 672 (US). Huanuco: Prov. Leoncio Prado: Dtto. Rupa Rupa: Tingo Maria, Plowman 7565 (US); Asplund 13218 (S); 600-700 m, Ferreyra 178 2278 (US, USM); 650 m, Wasshausen & Tovar 1260 (US, USM); E of Tingo Maria, 700-800 m, Schunke-Vigo 10518 (US). Junin: Prov. Tarma: La Merced, ca. 650 m, Macbride 5565 (F, NY, US); Rio Seco, 15 km from La Merced, Soukup 2494 (US); Rio Negro, 800 m, Woytkowski 5814 (MO, US); Rio Perené, Colonia Perené, 600 m, Killip & Smith 25228 (US); Rio Paucartam- bo Valley, near Perené Bridge, 700 m, Killip & Smith 25380 (US). Ayacucho: Prov. La Mar: Hda. Luisiana, 640 m, Wasshausen & Encarnacion 639 (US, USM). Cuzco: Prov. La Convencion: 3 km NE of Hda. Luisiana and Rio Apurimac, 660 m, Dudley 11450 (NA); 4 km E of San Francisco de Apuri- mac, 750 m, Wasshausen & Encarnacion 518 (US, USM). Prov. Paucartambo: Hda. Villa Carmen, 540 m, Vargas 14701 (CUZ, US); Dtto. Cosnipata: 9 km N of Pilcopata, 600 m, Wasshausen & Encarnaci6n 580 (US, USM); Valley de Cosnipata, between Mitiana and Keros, ca. 800 m, Scolnik 866 (NY). Prov. Quispicanchis: Marcapata Val- ley, 1200 m, Herrera 1170 (US). Madre de Dios: Rio Acre: Seringal Auristella, Ule 9808b (G, K). Without exact locality: Ruiz & Payon s.n. (MA, holotype of Justicia spi- cata Ruiz & Pavon; F, photo); 1878, Mar- tinet s.n. (P). BOLIVIA: Pando: W bank of Rio Madeira, opposite Abuna, Prance et al. 5695 (INPA, NY, US); 2 km above Ribe- rao, Prance 6499 (INPA, NY, US); Nicolas Suarez, ca. 30 km SW of Cobija on road to Naraueda, ca. 250 m, Sperling & King 6607 (NY, US). El Beni: junction of Rivers Beni and Madre de Dios, Rusby 1099 (NY, US). La Paz: Mapiri, Bang 1555 (NY, US); 750 m, Buchtien 1472 (US). Prov. S. Yungas: basin of Rio Bopi, San Bartolome, 750-900 m, Krukoff 10236 (NY, US); Bopi River Valley, 650 m, White 646 (NY). Cochabam- ba: Prov. Chapare, Todos Santos, 300 m, Steinbach 429 (NY, US). Without exact lo- cality: Pearce s.n. (K). BRAZIL: Amazonas: Borba, Riedel s.n. (LE, holotype of P. rie- deliana Nees; frag. GRZ); Parati, Traill s.n. (K); Rio Acre, Seringal Sao Francisco, Ule PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 9898 (K); 9809 (K); Boca do Acre, Rios Purus & Acre, track from Sao Paulo to Terra Firme, Rio Purus, opposite Boca do Acre, Prance et al. 2582 (INPA, NY); Manaca- puru, Spruce 1622 (K); Namorado Novo, watershed between Rio Curuqueté and Rio Madeira at Abuna, Prance et al. 14677 (INPA, NY, US). Rond6nia: Rio Jamari- Cachoeira de Sta. Cruz, Pires & Martin 9957 (UB, US); basin of Rio Madeira, 4 km N of Mutuparana, Prance et al. 5470 (INPA, NY, US); S bank of Rio Madeira, 2 km above Mutuparana, Prance et al. 5670 (INPA, NY, US); Sao Lourenco cassiterite mine, ca. 20 km NW of Rio Madeira, across from Mu- tuparana, Calderon et al. 2826 (US). Acre- Amazonas: municipio de Boca do Acre, vi- cinity of Rio Iaco, Cid & Nelson 2730 (INPA, NY). Acre: near mouth of Rio Ma- cauhan (tributary of Rio Yaco), Krukoff 5376 (NY); 5547 (NY). Para: 1908, Bakar s.n. (US); Taua, Spruce 266 (K); Pérto do Igarapé Mururetena, S. Miguel do Guama, Black 56-18888 (INPA, US). Edge of lowland rainforest, margins of thickets, along trails or river banks at ele- vations between 150 and 700 meters. West Indies southward to Bolivia and eastward to Amazonian Brazil (Amazonas, Rond6o- nia, Acre and Para). Discussion. —Leonard (1953:291) first noted that Pachystachys spicata (P. riede- liana) has often been confused with P. coc- cinea (Aublet) Nees of French Guiana and northern Amazonian Brazil. He further states: ““Although the two species are su- perficially similar, they can readily be sep- arated by the longer, linear calyx segments Of Pxcoceimeas: Only one chromosome count has been obtained for a species of Pachystachys, that by Takizawa (1957), for P. spicata, reported by him as Justicia coccinea, of n = 16. Pachystachys rosea Wasshausen, sp. nov Fig. 4 Frutex, caules subquadrangulares, brun- neoli, glabri vel parce puberuli; lamina fo- VOLUME 99, NUMBER 1 liorum elliptica vel anguste ovata, acumi- nata, basi angustata in petiolo amplectens, firma, glabra; spicae terminales, solitariae, rachidi puberula; bracteae imbricatae, vi- ridae, anguste ovatae, glanduloso-pilosae; bracteolae minutae, lineares, glabrae; calyx campanulatus, segmenta anguste lanceola- ta, minute puberula; corolla rosea, glabra vel parce puberula, glanduloso-punctata, tubo leviter curvato, labio superiore erecto, conduplicato, bilobato, labio inferiore tri- lobato, lobis patulis, obtusis, ciliatis, lobis lateralibus ellipticis vel oblongis, lobo me- dio elliptico. Shrub, 3 m tall; stem subquadrangular, brownish, glabrous or sparingly puberulous; leaf blades petiolate; elliptic to narrowly ovate, 15-22 cm long, 5—7.5 cm wide, acu- minate at apex, narrowed from below mid- dle to rounded, clasping base 5 mm wide, entire, undulate, moderately firm, glabrous, cystoliths conspicuous; petioles 2.5—6.5 cm long, glabrous or sparingly and inconspic- uously puberulous; inflorescence consisting of solitary, terminal spike 15-25 cm long, rachis puberulous; bracts imbricate, green, narrowly ovate, 2.2—2.5 cm long, 8-9 mm wide, acuminate and mucronulate at apex, gradually narrowed and attenuate at base, glandular-pilose and ciliate, eglandular hairs intermixed with glandular ones; bractlets minute, linear, 2 mm long, 0.4 mm wide, glabrous, ciliate; calyx campanulate, 9 mm long, segments narrowly lanceolate, 8 mm long, 1 mm wide near base, sparingly and inconspicuously puberulous and ciliate; co- rolla pink to orange-red, 6-6.5 cm long, gla- brous or sparingly and inconspicuously puberulous, glandular punctate, tube slight- ly curved, 4 mm wide at base, narrowed to 2mm at 7 mm above base, thence gradually enlarged to 9 mm at throat, upper lip erect, conduplicate, ovate, 2.5 cm long, 1.2 cm wide near base, gradually narrowed to 2.5 mm at tip, 2-lobed, lobes orbicular, 1 mm long and wide, obtuse, lower lip deeply 3-lobed, lobes spreading, obtuse and ciliate, lateral pair elliptic to oblong, 16 mm long, 179 6 mm wide, middle lobe elliptic, 18 mm long, 6 mm wide; stamens attached near base of corolla tube, filaments 5 cm long, sparingly puberulous, anthers 6 mm long, deeply sagittate, basal lobes muticous; staminodes rudimentary, puberulous; ovary glabrous; capsule wanting. Type. —PERU: Cuzco: Prov. Paucartam- bo, between Pilcopata and Carbon, 720 m, Vargas C. 23159 (holotype US). Paratype.— PERU: Cuzco: Prov. Paucar- tambo, Atalaya, near junction of Rio Car- bon with Rio Alto Madre de Dios, Foster 2398 (F, US). Hillside and riverbank, as well as margin of rainforest. Discussion. —Pachystachys rosea is per- haps nearest in relationship to the rather widespread P. spicata, but differs markedly in that the latter species has triangular calyx segments about 3 mm long, a scarlet corolla which may become orange-red with age, and lower corolla lip lobes which are all linear and 3—4 mm wide. In contrast, P. rosea has lanceolate calyx segments 8 mm long, a co- rolla that is pink to possibly orange-red with age, and lower corolla lip lobes of which the lateral pair is elliptic to oblong and the mid- dle lobe elliptic (all are 6 mm wide). Pachystachys killipii Wasshausen, sp. nov. Figs. 6, 10 Frutex; caules teres, brunneoli, glabri; lamina foliorum elliptica vel ovata, acu- minata, basi cuneata, modice firma, glabra; spicae terminales, solitariae, rachidi dense pilosa; bracteae laxe imbricatae, late ellip- ticae, acuminatae et mucronulatae, dense glanduloso-pilosae; bracteolae lineares, pi- losae; calyx campanulatus, segmenta lan- ceolata, glabra, ciliolata; corolla subrosea rubra, tubo leviter curvato, praecipue prope basin hirsuto, labio superiore erecto, oblon- go usque anguste lanceolato, minute bilo- bato, labio inferiore trilobato, lobis patulis, lobis lateralibus oblongis vel anguste ovatis, lobo medio ovato, conduplicato et obtuso. Erect shrub | m tall; stem terete, brown- 180 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ish, glabrous; leaf blades elliptic to ovate, 10.5-18 cm long and 3.5—6 cm wide, acu- minate at apex, narrowed and cuneate at base, moderately firm, entire or undulate, glabrous, cystoliths numerous both above and below; petioles 0.6-2.5 cm long, gla- brous; inflorescence consisting of terminal, solitary spike 15 cm long, rachis densely pilose; bracts rather loosely imbricate, green, broadly elliptic, 1.7 cm long, 6-7.5 mm broad, acuminate and mucronulate at apex, gradually narrowed at base and attenuate, rather densely glandular-pilose, glands pur- plish; bractlets linear, 11 mm long, 1.25 mm wide, pilose, bearing few glandular tri- chomes near apex; calyx campanulate, 12 mm long, segments lanceolate, 10 mm long and | mm wide near base, acuminate, bear- ing small mucro at apex, glabrous and cil- iate; corolla pinkish-red, 6.5—7 cm long, tube slightly curved, 4.5 mm wide at base, nar- rowed to 1.5 mm at 10 mm above base, thence gradually enlarged to 1 cm at throat, hirsute, especially near base, upper lip erect, oblong to narrowly lanceolate, 2.2 cm long, 7 mm wide near base, gradually narrowed to 1 mm at tip, minutely 2-lobed, lobes ob- tuse, lower lip 3-lobed, lobes spreading, lat- eral pair oblong to narrowly ovate, 17 mm long, 5 mm wide, rounded at tip, middle lobe ovate, 12 mm long and 5.5 mm wide, conduplicate and obtuse; stamens attached near base of corolla tube, filaments 5 cm long, glabrous, anthers 6 mm long, deeply sagittate, basal lobes muticous; pollen grains 46 um long, 45 um wide, tricolporate, each mesocolpium with two colpoid streaks, membrane densely granular, sexine supra- reticulate, reticulation heterobrochate (Fig. 10); staminodes none; ovary glabrous; cap- sule wanting. Type. —PERU: Junin: Puerto Bermudez, 355 m, Killip & Smith 26621 (holotype US; isotype NY). Occasional along clay river bank. Discussion. —Pachystachys killipii is characterized by a combination of a pink- ish-red corolla 6.5—7 cm long, rather small, loosely imbricate, glandular-pilose bracts 1.7 cm long, 6—7.5 mm broad, and a relatively large, campanulate calyx with the segments 12 mm long and bearing a small mucro at the tip. It is a pleasure to name this species for Ellsworth ““Buddy” Killip, who through his extensive field work, has added so much to our knowledge of South American floras. Pachystachys ossolae Wasshausen, sp. nov. Figs. 9, 10 Suffrutex, scandens; caules subquadran- gulares, brunneoli, glabri; lamina foliorum elliptica vel ovata, brevi-acuminata, basi angustata in petiolo amplectens, modice fir- ma, glabra; spicae terminales, solitariae, rachidi puberula; bracteae imbricatae, vi- ridae apice purpureae, oblanceolatae, mu- cronulatae, parce puberulae; bracteolae lanceolatae; calyx campanulatus, segmenta lanceolata, puberula, ciliolata; corolla coc- cinea, puberula et glanduloso-punctata, tubo leviter curvato, labio superiore erecto, ova- to, bilobato, lobis obtusis, labio inferiore profunde trilobato, lobis patulis, paribus la- teralibus oblongis vel anguste ovatis, ob- tusis, lobo medio anguste ovato, condupli- cato et obtuso. Climbing, suffrutescent shrub, 0.7—4 m tall; stem subquadrangular, brownish, gla- brous; leaf blades petiolate, elliptic to ovate, 17-30 cm long and 7-12 cm wide, short- acuminate at apex, narrowed from below middle to a rounded, clasping base 10 mm wide in larger leaves, entire or undulate, moderately firm, glabrous, cystoliths incon- spicuous; petioles 3-8 cm long, glabrous; inflorescence consisting of solitary, terminal spike 12-20 cm long, rachis puberulous; bracts imbricate, green with purple at tips, oblanceolate, 2—2.8 cm long, 0.6-1 cm broad, acute and mucronulate at apex, grad- ually narrowed and attenuate at base, spar- ingly puberulous and ciliate, cystoliths prominent without; bractlets lanceolate, 6 mm long, 1 mm wide, puberulous and cil- VOLUME 99, NUMBER 1 181 Fig. 9. A-D, Pachystachys ossolae (Wasshausen & Encarnacion 729). A, Habit; B, Bract, bractlets and calyx; C, Corolla and exserted stamens; D, Corolla expanded, showing attachment of filaments. 182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 10. SEM photomicrograph of Pachystachys pollen. A-B, Pachystachys killipii (Killip & Smith 26621); C-D, Pachystachys ossolae (Wasshausen & Encarnacion 741). A, Polar orientation, x 1000; B, Portion of surface, including pore at x 3000; C, Polar orientation, x 1700; D, Portion of surface, including colpus and pseudocolpi at x 3000. late; calyx campanulate, 7 mm long, seg- ments lanceolate, 5 mm long and 1.5 mm wide near base, puberulous and ciliate; co- rolla crimson, 7 cm long, puberulous and glandular punctate, tube slightly curved, 4 mm wide at base, narrowed to 2.5 mm at 8 mm above base, thence gradually enlarged to 1 cm at throat, upper lip erect, ovate, 2.2 cm long, 9 mm wide near base, gradually narrowed to 2.7 mm at tip, 2-lobed, lobes obtuse, 1.5 mm long, 1.2 mm wide, lower lip deeply 3-lobed, lobes spreading, lateral pair oblong to narrowly ovate, 2 cm long, 5 mm wide, obtuse, middle lobe narrowly VOLUME 99, NUMBER 1 ovate, 18 mm long and 4.5 mm wide, con- duplicate and obtuse; stamens attached near base of corolla tube, filaments 5.5 cm long, puberulous, anthers yellow, 5.5 mm long, deeply sagittate, basal lobes muticous; pol- len grains prolate, 53 wm long, 35 wm wide, tricolporate, each colpus flanked by two pseudocolpi, tectum almost complete, psi- late-punctate (Fig. 10); staminodes none; ovary glabrous; capsule wanting. Type. —PERU: Cuzco: Prov. de Quispi- canchis, 3 km E of Quincemil, 960 m, Was- shausen & Encarnacion 729 (holotype US; isotypes USM, US-3). Additional specimens. —PERU: Cuzco: 25 km SW of Quincemil, 1340 m, Wasshausen & Encarnacion 741 (US); 5 km E of Quin- cemil, 960 m, Wasshausen & Encarnacion 833 (US). Madre de Dios: 17 km N of Pil- copata, near Rio Carbon, 480 m, Gentry, Dillon, Aronson & Berry 23588 (MO). Par- que Nacional del Manu, Rio Manu, Cocha Cashu Station, 350 m Terborgh & Foster 6499 (F). Puno: Prov. de Carabaya, San Ga- ban, Lechler 2501 (K); vicinity of San Ga- ban, 900 m, Wasshausen & Salas 1246 (US, USM); below San Gaban on Rio San Ga- ban, 500-1000 m, Dillon, Aronson, Herra & Berry 1215 (MO); San Gaban to Ollachea, 1000-2000 m, Dillon, Aronson, Herra & Berry 1249 (MO); Ollachea-San Gaban road, Arica, Boeke & Boeke 3141 (NY). Occasional, along edge of lowland rain- forest. Discussion. —Pachystachys ossolae is nearest in relationship to the widespread species P. spicata, but differs markedly in that the latter species has a glabrous to spar- ingly hirtellous corolla 5.5—7 cm long, green, ovate-lanceolate, acuminate bracts, and narrowly triangular calyx segments about 3 mm long. In contrast, P. ossolae has a pu- berulous and glandular punctate corolla 7 cm long, green with distinctly purple, ob- lanceolate, acute and mucronulate bracts, and lanceolate calyx segments about 5 mm long. It is a great pleasure to name this new 183 species for my good friend and confidant, the late Charles A. Ossola, in recognition for his outstanding dedication and devotion to the advancement of systematic biology. Pachystachys coccinea (Aubl.) Nees in DC. Prodr. 11: 319. 1847. Justicia coccinea Aubl. Guyan. 1: 10, pl. 3. 1775. Jacobinia coccinea (Aubl.) Hiern in Warm. Symb. 23: 690. 1877. Shrub 1-2 m tall; branches puberulous when young, soon glabrous, constricted and easily broken just above nodes, bark thin and usually striate; leaves petiolate, blades elliptic to oblong-obovate, 18-25 cm long and 6—9 cm wide, acuminate (often abruptly SO) at apex, when not elliptic, narrowed from above middle to acute or obtuse base, entire or undulate, glabrous except costa and lat- eral veins, these puberulous but soon gla- brescent, cystoliths 0.25 mm long; petioles 2-6 cm long, glabrous; inflorescence con- sisting of terminal spike 9—22 cm long bear- ing conspicuous bracts and flowers, pedun- cles 1-3 cm long; bracts green, ovate to lanceolate, 1.5—2.5 cm long and 4-9 mm wide, lower bracts decussate, upper ones ternate and quaternate, narrower, all acute or subacuminate, apiculate, abruptly nar- rowed to short-winged base, more or less puberulent, usually glandular, margins and costa short-pubescent; bractlets linear-lan- ceolate, 5 mm long, 0.5 mm wide, puber- ulous; calyx campanulate, 5.5—6 mm long, slightly irregular, tube 1 mm long, segments lanceolate, equal, 5 mm long, 0.5 mm wide, sparingly and inconspicuously puberulent; corolla bright red, S—7 cm long, rather spar- ingly pubescent with minute spreading hairs (hirtellous), 4 mm wide at base, narrowed to 3 mm above base, 7-10 mm wide at throat, upper lip erect, 2—2.5 cm long, emar- ginate, lobes | mm long and 0.5 mm wide, lower lip oblong, obtuse, 1.7—2 cm long, 4— 5 mm wide; filaments 5.5 cm long, mature white-pubescent, densely so below, sparing- 184 Fig. 11. ly so above; staminodes when present mi- nute, vestigial; ovary and style glabrous. Material. —FRENCH GUIANA: Massif des Emerillons, centre N Sources de l’Ap- prouague, ca. 250 m, Cremers 6707 (CAY, US); Cayenne, Aublet s.n. (BM, holotype of Justicia coccinea Aubl.); Rothery 489 (K); Rorota, ile de Cayenne, Prévost 1259(CAY, US); vicinity of Cayenne, Montagne de Ma- hury, Sagot s.n. (K); 50-150 m, Maas, Men- nega & Koek-Noorman 2215 (NY, US); Matabon, Broadway 509 (NY, US); Mt. Bruyere, 4°5'N, 51°43’W, Irwin, Egler & Pires 47334 (NY, US); Village Kaw, Black & D’Aage 54-17539 (IAN, US). Without PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Isotype of Pachystachys nutans Nees (Ehrenbers 1060, BM). exact locality: Jul 1824, Poiteau s.n. (K, photo F, US); 1842, Melinon 263 (GZU, P, US); 1850, Leprieur s.n. (P, US); Perrottet 204 (GZU). BRAZIL: Territorio Amapa: Rio Oiapoque, Mt. Tipac, 3°36'N, 51°19'W, 0-100 m, Jrwin 48661 (NY, US); Porto Grande, Regiao do Vila Nova, Rosa 1051 (MG, NY, US). Para: Belém, Pires 14.871 (IAN, US). Common in shade of forest clearings and margins at elevations between 50 and 250 meters. Discussion. —Pachystachys coccinea is a very showy and beautiful species, with its large, conspicuously bracted terminal spike VOLUME 99, NUMBER 1 and bright red corollas. The species is near- est in relationship to P. spicata. The major difference between the two species is dis- cussed under the latter species. Excluded Species Pachystachys nutans Nees in DC., Prodr. 11: 320. 1847, = Jacobinia nutans (Nees) Hemsley, Biol. Centr. Amer., Bot. 2: 521. 1882. This species, based on Ehrenberg 1060 (holotype B, destroyed; isotype BM, sterile; fragment GZU), is known only from the type locality, Mexico, Barranca de Rio de Tolimea, near Ajuntas in Car- acol (Fig. 11). The description by Nees states that the plant is suffrutescent, pubescent; stem slen- der, very much branched; leaf blades ovate, 2.4-2.8 cm long, attenuate and obtuse at apex, short-acute at base, costa hirtellous; petioles 6—8 mm long; inflorescence termi- nal, sessile, short, dense, nodding, branched spike; bracts pale green, ovate, cuspidate- mucronulate at apex, membranous, ciliate; bractlets linear-subulate; calyx rather short; corolla short, apparently purple. The ultimate disposition of this taxon must await the recollecting of additional material. 185 Acknowledgments My sincerest thanks to Cathy Pasquale for the excellent line drawings and to Dr. Ste- phen F. Smith and the staff of the SEM lab- oratory for their high-quality pollen pho- tomicrographs. Literature Cited Bremekamp, C. E. B. 1938. Notes on the Acantha- ceae of Surinam.—Recuiel des Travaux Bota- niques Néerlandais 35:130-171. Leonard, E. C. 1953. The Acanthaceae of Colombia II.—Contributions from the United States Na- tional Herbarium 31:119-322. Lindau, G. 1893. Beitrage zur Systematik der Acan- thaceen. — Botanische Jahrbiicher 18:36-64, pls. I, Do Nees von Esenbeck, C. G. 1847. Acanthaceae. Jn K. F. P. von Martius, Flora Brasiliensis 9:6-163, pls. 1-31. 1847. Acanthaceae. In A. P. de Candolle, Prodromus Systematis Naturalis Regni Vege- tablis 11:46-519. Rizzini, C. T. 1947. Estudos Sébre As Acantha- ceae.— Boletim do Museu Nacional, Rio de Ja- neiro 8:1—38, pls. 1-11. Takizawa, Y. 1957. Die Struktur der Pachytanchro- mosomen einiger Acanthaceen, sowie eine Reihe neu bestimmter Chromosomenzahlen.—Cyto- logia 22:118-126. Department of Botany, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560. 186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE % BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD, LONDON, SW7 5BD 3 October 1985 The Commission hereby gives six months notice of the possible use of it plenary powers in the following cases, published in the Bulletin of Zoological Nomenclature, volume 42, part 3 on 30 September 1985, and would welcome comments and advice on them from interested zoologists. Correspondence should be addressed to the Executive Secretary at the above address, if possible within six months of the date of publication of this notice. Case No. 2464 Berytus Fabricius, 1803 (Insecta, Heteroptera, Berytidae): proposed desig- nations of a type species. 2490 Thylacites Germar, 1817; Brachyderes Schonherr, 1823; Cycloderes Sahl- berg, 1823; and Cycloderes Schonherr, 1823 (Insecta, Coleoptera): pro- posal to maintain current usage. 2467 Neodorippe Seréne & Romimohtarto, 1969 (Crustacea, Decapoda): proposed designation of type species. 2363 Sagartia luciae Verrill, 1898 (Coelenterata, Actiniaria): proposed conser- vation. P. K. TUBES Executive Secretary VOLUME 99, NUMBER 1 INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE % BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON, SW7 5BD 3 October 1985 The following Opinions have been published by the International Commission on Zoological Nomenclature in the Bulletin of Zoological Nomenclature, volume 42, part 3 on 30 September 1985: Opinion No. 1328 (p. 1329 (p. 1330 (p. 1331 (p. 1332 (p. 1333 (p. 1334 (p. 1335 (p. 1336 (p. 1337 (p. 1338 (p. 1339 (p. 1340 (p. 1341 (p. 1342 (p. 1343 (p. 1344 (p. 222) 226) 228) 230) 233) 236) 238) 241) 244) 249) 251) 255) 258) 261) 264) 266) 269) Belemnites mucronatus Schlotheim, 1813 (Coleoidea): conserved and neotype designated. Galago crassicaudatus E. Geoffroy, 1812 (Primates, Galigidae): neo- type designated. Prodorylaimus Andrassy, 1959 (Nematoda): type species desig- nated. SPHAERIIDAE Jeffreys, 1862 (1820) (Mollusca, Bivalvia) and MI- CROSPORIDAE Reichardt, 1976 (Insecta, Coleoptera): placed on the Official List. Calamoecia australica Sars, 1908 and Calamoecia australis (Searle 1911) (Crustacea, Copepoda): proposals to remove the con- fusion rejected. Ipnops murrayi Gunther, 1878 (Ostiechthyes): conserved. Harminius Fairmaire, 1851 (Coleoptera): type species designated. Nepa cinerea Linnaeus, 1758 (Insecta, Heteroptera): conserved. Five specific names proposed for Heterodera A. Schmidt, 1871 (Aschelminthes, Nematode) by B. A. Cooper, 1955 ruled to be available. Selkirkia columbia Conway Morris, 1977 designated as type species of Selkirkia Walcott, 1911 (Priapulida). Thrips rufus Haliday, 1836 (Insecta, Thysanoptera): conserved for the type species of Aptinothrips Haliday, 1836. Papilio fatima Fabricius, 1793 (Insecta, Lepidoptera): ruled to be exempt from the application of the Principle of Homonymy. Attus otiosus Hentz, 1846 (Arachnida, Araneae): conserved. Simulium amazonicum Goeldi, 1905 (Insecta, Diptera): neotype designated. Damalis planiceps Fabricius, 1805 designated as type species of Damalis Fabricius, 1805 (Insecta, Diptera). Kinosternon alamosae Berry & Legler, 1980 and Kinosternon oa- xacae Berry & Iverson, 1980 (Reptilia, Testudines): con- served. Mayorella Schaeffer, 1926 given nomenclatural precedence over Dactylamoeba Korotneff, 1880 (Rhizopoda, Amoebida). 187 188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1345 (p. 271) Laomedea flexuosa Alder, 1857, Sertularia volubilis Linnaeus, 1758 and Campanularia johnstoni Alder, 1856 designated as type species of Laomedea Lamouroux, 1812, Campanularia La- marck, 1816 and Clytia Lamouroux, 1812 (Coelenterata, Hy- droida) respectively. 1346 (p. 274) Cythereis distinguenda Neviana, 1928, Cythere crispata Brady, 1868 and Cythere pavonia Brady, 1866 (Crustacea, Ostracoda): type material conserved. 1347 (p. 277) Anthalia schoenherri Zetterstedt, 1838 designated as type species of Anthalia Zetterstedt, 1838 (Insecta, Diptera). 1348 (p. 279) Bos gaurus H. Smith, 1827 (Mammalia, Artiodactyla): conserved. 1349 (p. 281) Antilope depressicornis H. Smith, 1827 and Anoa quarlesi Ouwens, 1910 (Mammalia, Artiodactyla): conserved. 1350 (p. 283) Conus antiquus Lamarck, 1810 (Mollusca, Gastropoda): neotype suppressed. 1351 (p. 285) Galeopsomyia Girault, 1916 (Insecta, Hymenoptera): conserved. 1352 (p. 287) Eurhinus Schonherr, 1825 (Insecta, Coleoptera): ruled as a justified emendation of Eurhin Illiger, 1807. 1353 (p. 291) Myzus festucae Theobald, 1917 (Insecta, Hemiptera): conserved. The Commission regrets that it cannot supply separates of Opinions. P. K. TUBBS Executive Secretary VOLUME 99, NUMBER 1 189 INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE % BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON, SW7 5BD 11 December 1985 The Commission hereby gives six months notice of the possible use of its plenary powers in the following cases, published in the Bulletin of Zoological Nomenclature, volume 42, part 4 on 6 December 1985, and would welcome comments and advice on them from interested zoologists. Correspondence should be addressed to the Executive Secretary at the above address, if possible within six months of the date of publication of this notice. Case No 2434 Semionotus Agassiz, 1832 (Osteichthyes): proposed designation of Semio- notus bergeri Agassiz, 1833 as type species. 2470 Cephalopholis argus Schneider, 1801 and Cephalopholis sexmaculata (Rup- pell 1830) (Osteichthyes, Serranidae): proposed conservation by suppression of Bodianus guttatus Bloch, 1790, Anthius argus Bloch, 1792 and Serranus zanana Valenciennes, 1828. Cheirurus Beyrich, 1845 (Trilobita): proposed designation of Cheirurus in- signis Beyrich, 1845 as a type species. Eugynothrips Priesner, 1926 (Insecta, Thysanoptera): proposed designation of Cryptothrips conocephali Karny, 1913 as type species. HETEROGYNIDAE Rambur, 1866 (Insecta, Lepidoptera) and HETERO- GYNINAE Nagy, 1969 (Insecta, Hymenoptera): proposals to remove the homonymy. THAIDIDAE Jousseaume, 1888 (Mollusca, Gastropoda) and THAIDIDAE Lehtinen, 1967 (Arachnida, Araneae): proposals to remove the hom- onymy. Drasterius bimaculatus (Rossi, 1790) (Insecta, Coleoptera, Elateridae): pro- posed suppression of Elater bimaculata Fourcroy, 1785. Microchrysa Loew, 1855 (Insecta, Diptera): proposed conservation by the suppression of Chrysomyia Macquart, 1834. Musca trilineta Linnaeus, 1767 (Insecta, Diptera): proposed conservation by the suppression of Musca graeca Pontoppidan, 1763. The family names for the storm petrels and the dippers. P. K. TUBBS Executive Secretary 190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE % BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON, SW7 5BD 11 December 1985 The following Opinions have been published by the International Commission on Zoological Nomenclature in the Bulletin of Zoological Nomenclature, volume 42, part 4 on 6 December 1985. Opinion No 1354 (p. 330) Agrotis redimicula Morrison, 1874 (Lepidoptera): conserved from 1874. 1355 (p. 332) Lingula anatina Lamarck, 1801 is the type species of Lingula Bru- guiére, [1797] (Brachiopoda). 1356 (p. 335) Dactylopusia Norman, 1903 (Crustacea, Copepoda): type species designated. 1357 (p. 338) ANUROPODIDAE Bacescu, 1980 (Crustacea, Tanaidacea) and AN- UROPODIDAE Stebbing, 1893 (Crustacea, Isopoda): a ruling to remove the homonymy. 1358 (p. 341) Calaphis Walsh, 1862 and Callaphis Walker, 1870 (Insecta, He- miptera): a ruling to remove the confusion. 1359 (p. 344) UROPLAT~—as the stem of family-group names in Reptilia, Sauria and Insecta, Coleoptera: a ruling to remove the homonymy. 1360 (p. 347) Odceciacus vicarius Horvath, 1912 (Insecta, Hemiptera): conserved. 1361 (p. 349) Larentia capitata Herrich-Schaffer, 1839, given nomenclatural pre- cedence over Phalaena posticata Fabricius, 1794 (Insecta, Lep- idoptera). 1362 (p. 351) Phalaena coracina Esper, 1805, given nomenclatural precedence over Phalaena hirtata Fabricius, 1794 (Insecta, Lepidoptera). 1363 (p.353) Ancistroceroides Saussure, 1855 (Insecta, Hymenoptera): type species designated. 1364 (p. 355) Kassina Girard, 1853 (Amphibia, Anura): conserved. 1365 (p. 357) Allygus Fieber, 1872 (Insecta, Homoptera): type species designated. 1366 (p. 359) Mactra sachalinensis Schrenk, 1862 (Mollusca, Bivalvia): conserved. 1367 (p. 361) Alpheus lottini Guérin, 1829 (Crustacea, Decapoda): conserved. 1368 (p. 365) The generic names Pan and Panthera (Mammalia, Carnivora): avail- able as from Oken, 1816. The commission regrets that it cannot supply separates of Opinions. P. K. TUBBS Executive Secretary { fy . ( i at } i ray i ‘ i ——S 1 i) 5 i \ 7 | i | =, i x i | 4 8 2: ie { i 4 ‘ i} f ; ‘ hah i Z ah r Ay 1 i 4 : 7 . s} P « - i 2 ae. i | JOMENCLATUBBC. 2. SRE SCSI HATLBEAE. skid» = oe | SW? DOA 9 eo a o an |, DoNtewa 207 SD a ta > tee: eeiiemiry * wat iF TY 5 rr? iad 7) Wh vin) £9) Gea! 4 j - i f f r la 3 we) S 7 o hen 1 7 * ' ¢¢ I = nas Mh eno : ¥ i “ven SOK ie 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 will be published in English (except for Latin diagnosis/description of plant taxa which should not be duplicated by an English translation), with a summary in an alternate language when appropriate. Publication Charges.—Authors will be asked to assume publication costs of page-charges, tabular material, and figures, at the lowest possible rates. Submission of manuscripts.—Manuscripts should be sent to the Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History, 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. Manuscripts will be reviewed in order of receipt by a board of associate editors and appropriate referees. Presentation.—Clarity of presentation, and requirements of taxonomic and nomenclatural procedures necessitate reasonable consistency in the organization of papers. Telegraphic style is recommended for descriptions. Literature should be cited in abbreviated style (author, date, page), except in botanical synonymies, with full citations of journals and books (no abbrevia- tions) in the Literature Cited section. The establishment of new taxa must conform with the requirements of the appropriate in- ternational codes of nomenclature. Authors are expected to be familiar with these codes and to comply with them. New species-group accounts must designate a type-specimen deposited in an institutional collection. The sequence of material should be: Title, Author(s), Abstract, Text, Acknowledgments, Literature Cited, Author’s (s’) Address(es), Appendix, List of Figures (entire figure legends), Figures (each numbered and identified), Tables (each table numbered with an Arabic numeral and heading provided). Manuscripts should be typed, double-spaced throughout (including tables, legends, and foot- notes) on one side of 842 x 11 inch sheets, with at least one inch of margin all around. Manu- scripts in dot-matrix will not be accepted. Submit a facsimile with the original, and retain an author’s copy. Pages must be numbered on top. One manuscript page = approximately 0.5 printed page. Underline singly scientific names of genera and lower categories; leave other indications to the editor. Figures and tables with their legends and headings should be self-explanatory, not requiring reference to the text. Indicate their approximate placement by a pencil mark in the margin of the manuscript. Illustrations should be planned in proportions that will efficiently use space on the type bed of the Proceedings (12.5 x 20 cm) and should not exceed 15 x 24 inches. Figures requiring solid black backgrounds should be indicated as such, but not masked. Art work will be returned only on request. Proofs.—Galley proofs will be submitted to authors for correction and approval. Reprint orders will be taken with returned proof. Costs.—Page charges @ $60.00, figures @ $10.00, tabular material $3.00 per printed inch. All authors are expected to pay the charges for figures, tables, changes at proof stage, and reprints. Payment of full costs will probably facilitate speedy publication. CONTENTS Nephtys cryptomma, new species (Polychaeta: Nephtyidae) from the northern Gulf of Mexico Donald E. Harper, Jr. New South American gastropods in the genera Conus (Conidae) and Latirus (Fasciolariidae) Edward J. Petuch The Austral-African conid subgenus F/oraconus Iredale, 1930, taken off Bermuda (Gastropoda: Conidae) Edward J. Petuch The status of Pseudorhabdosynochus Yamaguti, 1958, and Cycloplectanum Oliver, 1968, (Monogenea: Diplectanidae) Delane C. Kritsky and Mary Beverley-Burton A new record of Paracorophium hartmannorum Andres, 1975, from the Chilean coast, with a description of the adult (Amphipoda: Corophiidae) Exequiel Gonzalez Description of a paedophagous deep-water cichlid (Teleostei: Cichlidae) from Lake Malawi, Africa Jay R. Stauffer, Jr. and Kenneth R. McKaye Aegla spectabilis, a new species of freshwater crab from the eastern slope of the Nahuelbuta Coastal Cordillera, Chile Carlos G. Jara Nomenclatural notes on the Anura (Amphibia) Jay M. Savage Serolis agassizi, new species, from the deep sea off Cape Fear, North Carolina (Crustacea: Isopoda) Robert Y. George A new species of red-eyed treefrog of the Hy/a uranochroa group (Anura: Hylidae) from northern Honduras James R. McCranie and Larry David Wilson Hawaiian Xanthidae (Decapoda: Brachyura) II. Description of Garthiella, new genus, with a redescription of G. aberrans (Rathbun, 1906) Richard H. Titgen Geographic variation in the white-mantled barbet (Capito hypoleucus) of Colombia (Aves: Capitonidae) Gary R. Graves Lasionectes entrichoma, new genus, new species, (Crustacea: Remipedia) from anchialine caves in the Turks and Caicos, British West Indies Jill Yager and Frederick R. Schram A redescription of Microcyclops ceibaensis (Marsh, 1919) (Copepoda: Cyclopoida) from Marsh’s specimens in the National Museum of Natural History Janet W. Reid A new genus and species of interstitial Sigalionidae and a report on the presence of venom glands in some scale-worm families (Annelida: Polychaeta) Paul S. Wolf Glossothelepus, a new genus of Thelepinae (Polychaeta: Terebellidae) from the Gulf of Cali- fornia, Mexico P. A. Hutchings and C. J. Glasby New records of Mimilambrus wileyi Williams, 1979 (Crustacea: Decapoda: Brachyura), with notes on the systematics of the Mimilambridae Williams, 1979, and Parthenopidae MacLeay, 1838, sensu Guinot, 1978 Peter K. L. Ng and Gilberto Rodriguez Descriptions of two new species of Hylodes from the Atlantic forests of Brazil (Amphibia: Leptodactylidae) W. Ronald Heyer and Reginald B. Cocroft The status of the Indo-West Pacific cardinalfishes Apogon aroubiensis and A. nigrofasciatus John E. Randall and Ernest A. Lachner Hadoceras taylori, a new genus and species of phreatic Hydrobiidae (Gastropoda: Rissoacea) from south-central Texas Robert Hershler and Glenn Longley A new species of Helminthoglypta (Gastropoda: Pulmonata: Helminthoglyptidae) from San Diego County, California Richard L. Reeder and Walter B. Miller Two species of Urocopia, planktonic poecilostomatoid copepods of the family Urocopiidae . Humes and Stock, 1972 Gayle A. Heron and David M. Damkaer Subspecies of the Glaucous Gull, Larus hyperboreus (Aves: Charadriiformes) Richard C. Banks The systematics of the genus Pachystachys (Acanthaceae) Dieter C. Wasshausen International Commission on Zoological Nomenclature. Opinions 100 121 137 140 149 160 186 ~ SE y AZ Se oC BRT a < 7 < ye <7 7 / YE THE BIOLOGICAL SOCIETY OF WASHINGTON 1985-1986 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 Kensley Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Richard P. Vari Insects: Robert D. Gordon 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: $18.00 within the U.S.A., $23.00 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 Instutution, 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. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 191-195 NEASTACILLA FALCLANDICA (OHLIN), TYPE SPECIES OF THE GENUS, AND N. TATTERSALLI, NEW SPECIES (CRUSTACEA: ISOPODA: ARCTURIDAE) Helen M. Lew Ton and Gary C. B. Poore Abstract.—Neastacilla falclandica (Ohlin) is redescribed and figured from topotypic material. It is shown to be different from material from New Zealand which Tattersall (1921) assigned to N. falclandica and on which he based the new genus Neastacilla. Tattersall’s specimen and more recent material are described as a new species, N. tattersalli. Tattersall (1921) erected the genus Neas- tacilla and recorded the type species, As- tacilla falclandica Ohlin, from New Zea- land. Comparison of Tattersall’s specimen with topotypes from the Falkland Islands has revealed that Tattersall misidentified his specimen. As there is no modern descrip- tion of Neastacilla falclandica a redescrip- tion was felt to be desirable and the New Zealand species should be described. It was not possible to locate the holotype of As- tacilla falclandica in Hamburg, Berlin, or Stockholm, and it is presumed lost. We are appealing to the International Commission for Zoological Nomenclature to rule that Astacilla falclandica Ohlin be type species of the genus Neastacilla to pre- serve Current usage. This contribution forms part of a larger work on Australian species of Neastacilla in which the genus is rediagnosed and con- trasted with Astacilla Cordiner. The format of the descriptions of the two species de- scribed here follows that of the larger work in preparation. Mouthparts of N. falclan- dica are figured but these contribute little to generic distinctions. Neastacilla falclandica (Ohlin) Figs. 1, 2 Astacilla falclandica Ohlin, 1907:266, pl. 20.—Schultz, 1981:91. Astacilla falclandicus.—Stebbing, 1914:353. Neastacilla falclandica. —Nordenstam, 1933:119-122, fig. 28.—Kussakin, 1967: 357.—Sivertsen and Holthuis, 1980:67, fig. 18c. Material examined. —Swedish Museum for Natural History, Stockholm, Isopoda 7305, female (6.2 mm), Falkland Islands, Port Louis, Swedish South Polar Expedition 1901-3, Station 43. USNM 222665, juve- nile (3.1 mm), Falkland Islands, off Light- house, 22 Apr 1927. USNM 222666, ju- venile (3.0 mm), Falkland Islands, Port Stanley, 14 Apr 1927, No. 107. Description. —Female: Anterolateral lobe of head rounded, rostral point absent. Eyes subtriangular, slightly prominent in dorsal view. Fusion of head and first pereonite in- dicated by obsolete groove, incomplete dor- sally and not extending to lateral margin; lateral margin not incised; combined length of head and pereonite | 1.7 times combined length of pereonites 2 and 3. Lateral margin of pereonite 1 not expanded ventrally. Pe- reonites 2 and 3 subequal, smooth; lateral margins rounded, visible in dorsal view. Pereonite 4 one-third total body length, slightly wider than preceding pereonites; an- terolateral margins produced, rounded; dor- solateral margin produced ventrally. Pereo- nites 5—7 smooth, shorter posteriorly. Pleon smooth, 1.5 times combined lengths of pe- 192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. U, Uropod. Scale = 1 mm. reonites 5—7; profile a smooth slope. Two pleonal somites indicated by lateral inden- tations; in dorsal view, second pleonal so- mite protrudes laterally. Pleotelson with poorly developed posterolateral expan- sions, apex rounded but with steeply sloping sides. Antenna 1 reaching one-third along ar- ticle 3 of antenna 2. Antenna 2 almost two- Neastacilla falclandica, female, 6.2 mm (NHRM Isopoda 7305): P1, P2, P5, Pereopods 1, 2 and 5; thirds body length, not excessively slender or stout; flagellum of 3 articles. Ratio of articles 3-5 1.0:2.1:1.1. Pereopod 1 dactyl with terminal setae, claw absent. Pereopods 2-4 of moderate build, densely setose; small claw-like dactyl present. Pereopod 5 with slight ventral expansion, barely more than pereopods 6 and 7; lacking marked ventral expansion. Pereopods 5—7 smooth, dactyl VOLUME 99, NUMBER 2 193 Fig. 2. Neastacilla falclandica, juvenile, 3.1 mm (USNM 222665): MD, Mandibles (left and right); MX1, MX2, Maxillae 1 and 2; MP, Maxilliped. with 2 well-developed claws, larger claw half length of dactyl. Uropod outer ramus, length 1.7 times base width, apex rounded; inner ramus, distal margin with single stout seta reaching beyond distal margin of outer ra- mus. Distribution. — Falkland Islands. Remarks. —Mouthparts of this species are figured because it is the nominal type species of the genus. Neastacilla tattersalli, new species Fig. 3 Neastacilla falclandica.—Tattersall, 1921: 244, pl. 10, fig. 1.—Hurley, 1961:264. Material examined.—Holotype, British Museum (Natural History) 1921:11:29:316, female (8.9 mm), New Zealand, North Cape (34°25'S, 173°02’E), British Antarctic (“Terra Nova’’) Expedition station 96, 129 m. Paratypes, Zoological Museum, Copen- hagen, female (7.8 mm), juvenile (3.2 mm), New Zealand, Hauraki Gulk, Kawau Island (36°25'S, 174°51'E), 10 fathoms, coll. Dr. Th. Mortensen, 29 Dec 1914. Description. —Female: Anterolateral lobe of head angular, with small spine-like pro- jection on ventral margin, rostral point small. Eyes large, ovoid, slightly prominent in dorsal view. Fusion of head and first pe- reonite indicated by poorly defined groove; lateral margin incised; combined length 1.7 times combined length of pereonites 2 and 3. Pereonite 1 smooth, lateral margin not expanded ventrally. Pereonites 2 and 3 sub- equal, smooth; lateral margins visible in dorsal view. Pereonite 4 less than one-third total body length, smooth; not markedly wider than preceding pereonites. Antero- 194 sc | V7 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Neastacilla tattersalli, holotype: A2, Antenna 2 flagellum; P4, Pereopod 4; U, Uropod. Scale = 1 mm. lateral margins produced, rounded. Dor- solateral margin angular, produced ven- trally in posterior one-third of pereonite. Pereonites 5—7 smooth, progressively short- er posteriorly. Pleon longer than pereonites 5-7 combined, smooth, profile a smooth slope; 2 pleonal somites indicated by ob- solete lateral grooves. Pleotelson tapering to truncate apex, posterolateral expansions ab- sent. Antenna 1 reaching distal margin of ar- ticle 2 of antenna 2. Antenna 2 little over half body length, not excessively slender or stout; flagellum of 3 articles. Ratio of arti- cles 3-5 1.0:2.0:1.5. Pereopod 1 dactyl with terminal setae, claw absent. Pereopods 2-4 of moderate build, densely setose; dactyl present. Oostegites on pereopods 1-4. Pe- reopod 5 coxa with slight ventral expansion, barely more than pereopods 6 and 7. Pe- reopods 5—7 smooth, dactyl with 2 well- developed claws, larger claw as long as dac- VOLUME 99, NUMBER 2 tyl. Uropod outer ramus length approxi- mately 1.5 times base width, apex angular; inner ramus less than half length of outer ramus, distal margin with 2 subequal setae; setae reach distal margin of outer ramus. Distribution. —Northern North Island of New Zealand. Remarks. —The holotype specimen is in poor condition with mouthparts and several limbs, including pereopod 1, missing. Ex- amination of the paratypes which bear pe- reopod | confirm the species’ generic place- ment. Although only three specimens were examined some variation was observed. The distinct spine on the anterolateral lobe of the head of the holotype is absent from the other specimens. This is one of few species of Neastacilla in which the females lack spines and tuber- cles. It most closely resembles Neastacilla falclandica. The most obvious differences separating the two are the size and shape of the eye, the number of setae on the inner ramus of the uropods, and the shape of the pleotelson. Acknowledgments We thank A. Andersson, Swedish Mu- seum for Natural History, Stockholm; B. Kensley, National Museum of Natural His- tory, Washington; R. Lincoln, British Mu- seum (Natural History), London; and T. Wolff, Zoological Museum, Copenhagen for the loan of material. 195 Literature Cited Hurley, D. E. 1961. A checklist and key to the Crus- tacea Isopoda of New Zealand and the Subant- arctic islands.—Transactions of the Royal So- ciety of New Zealand (Zoology) 1:259-292. Kussakin, O. G. 1967. Fauna of Isopoda and Tan- aidacea in the coastal zones of the Antarctic and Subantarctic water.— Biological Reports of the Soviet Antarctic Expedition (1955-58) 3:220- 380. [Translation by the Israel Program for Sci- entific Translations, Jerusalem, 1968, pp. 220- 389.] Nordenstam, A. 1933. Marine Isopoda of the families Serolidae, Idotheidae, Pseudidotheidae, Arctur- idae, Parasellidae and Stenetriidae mainly from the South Atlantic. — Further Zoological Results of the Swedish Antarctic Expedition, 1901-03 3(1):1-284. Ohlin, A. 1907. Isopoda from Tierra del Fuego and Patagonia. I. Valvifera.— Wissenschaftliche Er- gebnisse der Schwedischen Expedition nach den Magellanslandern 2(11):261-306. [Often erro- neously cited as 1901.] Schultz, G. A. 1981. Arcturidae from the Antarctic and Southern Seas (Isopoda, Valvifera), part 1.— Antarctic Research Series 32:63-94. Sivertsen, E., and L. B. Holthuis. 1980. The marine isopod Crustacea of the Tristan da Cunha Ar- chipelago.—Gunneria 35:1-128. Stebbing, T.R.R. 1914. Crustacea from the Falkland Islands collected by Mr. Rupert Vallentin, F.L.S. part II.— Proceedings of the Zoological Society of London 1914:341-378. Tattersall, W. M. 1921. Crustacea. Part VI. Tanaida- cea and Isopoda.—British Antarctic (“Terra Nova’) Expedition, 1910, Natural History Re- ports, Zoology 3:191-258. Department of Crustacea, Museum of Victoria, Russell Street, Melbourne, Vic- toria 3000, Australia. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 196-197 ALPHEOPSIS CORTESIANA, A NEW SNAPPING SHRIMP FROM THE GULF OF CALIFORNIA Mary K. Wicksten and Michel E. Hendrickx Abstract.—A new species of subtidal snapping shrimp has been dredged off Sinaloa, Mexico. The species, most closely resembling A/pheopsis harperi of the Gulf of Mexico, has a large chela with an arched dactyl. There are no grooves on the large chela. A key to the eastern Pacific species of Alpheopsis is provided. Species of Alpheopsis, like other members of the family Alpheidae, commonly are called snapping shrimps. In the eastern Pa- cific, species of Alpheopsis often are infaunal inhabitants of offshore bottoms. Most rec- ords come from material taken in box cores, grabs, and dredges. During benthic studies by the R.V. E/ Puma in the Gulf of California, a new species of snapping shrimp was discovered, and is described herein. The drawings are by Deb- bie Meier, Texas A&M University. Alpheopsis cortesiana, new species Fig. 1 Description. —Rostrum short, acute, car- inate, exceeding ocular spines, falling far short of distal end of first article of anten- nular peduncle. Orbital teeth acute. Eyes covered in dorsal view, covered or partly exposed in lateral view. Pterygostomial an- gle of carapace rounded. Antennular peduncle long, second article longest. Stylocerite long and acute, reaching beyond end of first antennular article. Scaphocerite reaching as far as or falling slightly short of distal end of antennular pe- duncle, its lateral spine overreaching blade. Carpocerite longer than scaphocerite and antennular peduncle. Basicerite with small lateral tooth. Large cheliped with chela having palm 1.5 as long as broad. Margins of chela without grooves or notches. Fixed finger with 7 large teeth and 5-7 denticles distal thereto. Dactyl with 5-6 weak teeth. Dactyl lamel- late, upper margin strongly convex. Carpus short, cup-shaped. Merus slender, 3 x as long as broad, with 2—3 spines on superior mar- gin. Ischium short. Small chela missing in both specimens. Second leg with ratio of carpal articles 10: 4:4:4:4. Third leg slender. Dactyl sickle-shaped, 7x as long as wide. Propodus 2 x length of dactyl, slender, without spines but bearing long setae. Carpus 0.8 x length of propodus, merus 1.5 x length of carpus. Ischium 0.5 x merus, without spines. Fourth and fifth legs similar to third. Telson more than 2 x long as wide, wider at proximal end than at distal; with 2 pair dorsolateral spines and pair of spines on distal margin. (Constriction at end of an- terior third, as shown in illustration, prob- ably due to damage to specimen.) Uropods longer than telson. Holotype. — Female, total length 19.7 mm. Off Rio Fuerte, Sinaloa, Mexico (25°48'N, 109°34’W), 90 m, mud, 12 May 1982, Van Veen grab, R.V. E/ Puma, CORTES cruise 1, Allan Hancock Foundation (University of Southern California) type number 82-2. Paratype. —Female, total length 9.0 mm. Off Mazatlan, Sinaloa, 20 m, compact mud, 13 March 1981, in oyster dredge, R.V. E/ Puma, Estacion Mazatlan catalog number EMU-2096. VOLUME 99, NUMBER 2 Fig. 1. Alpheopsis cortesiana, holotype female. a, Frontal region in dorsal view; b, Frontal region in side view; c, Major cheliped; d, Second pereopod; e, Third pereopod; f, Telson. Etymology.—The specific epithet refers to the Sea of Cortés, from which the species was collected. Discussion. — Alpheopsis cortesiana most closely resembles A. harperi Wicksten, 1984, from the Gulf of Mexico. Both are offshore infaunal species with large chelae lacking grooves but having dactyls with convex margins. Alpheopsis harperi, however, lacks teeth on the fingers of the chela. It has spi- nules on the propodus of the third leg. The second article of the carpus of the second leg is longer than in A. cortesiana, and the dactyl of the third leg is shorter. There are no published reports of other species of Alpheopsis in the Gulf of Cali- fornia or off the coast of Sinaloa. However, a single ovigerous female of a different species, resembling A. aequalis Coutiére, was 197 collected at Isla Candelero, Sonora (6 July 1983, 10 m, commensal with zooanthid, Alex Kerstitch, collector, Allan Hancock Foundation collections). This specimen agrees with the description of A. aequalis given by Banner and Banner (1973:342) ex- cept that the blade of the scaphocerite tapers to a point instead of having a broadly rounded margin. More specimens are need- ed before the species can be formally de- scribed. Key to the Species of Alpheopsis in the Eastern Pacific 1. Anterior region of carapace with rostrum and orbital teeth — Anterior region of carapace with rostrum only, no orbital teeth .... 3 2. Large chela with longitudinal groove. (Southern-central Califor- nia, U.S.A.) ..Alpheopsis equidactylus (Lockington) — Large chela without longitudinal groove. (Off Sinaloa, Mexico) .... ....Alpheopsis cortesiana, new species 3. Large chela with notch on dorsal surface. (Chile-Peru) See ae Alpheopsis chilensis Coutiére — Large chela without notch on dorsal surface. (Off Sonora, Mexico) .... ....Alpheopsis cf. A. aequalis Coutiére Literature Cited Banner, D. M., and A. H. Banner. 1973. The alpheid shrimp of Australia. Part I: The lower genera. — Records of the Australian Museum 28(15):291— 382. Wicksten, M. K. 1984. Alpheopsis harperi (Decapo- da: Alpheidae): A new species of snapping shrimp from Texas.— Northeast Gulf Science 7(1):97- 100. (MKW) Department of Biology, Texas A&M University, College Station, Texas 77842; (MEH) Estacion Mazatlan del In- stituto de Ciencias del Mar y Limnologia, Apartado Postal 811, Mazatlan, Sinaloa 82000, México. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 198-213 FRESHWATER SHRIMPS FROM VENEZUELA I: SEVEN NEW SPECIES OF PALAEMONINAE (CRUSTACEA: DECAPODA: PALAEMONIDAE) Guido Pereira S. Abstract. —Seven new species of palaemonid shrimps are described: Macro- brachium reyesi, Macrobrachium pectinatum, Macrobrachium atabapense, Macrobrachium rodriguezi, Macrobrachium pumilum, Macrobrachium diery- thrum, and Palaemonetes (Palaemonetes) mercedae. All the specimens were collected in Venezuela. Details of living color patterns, fecundity, and variation are provided. The new species are compared with previously described, and morphologically related, South American species. Resumen. —Se describen siete especies de camarones palaemonidos nuevos para la ciencia: Macrobrachium reyesi, Macrobrachium pectinatum, Macro- brachium atabapense, Macrobrachium rodriguezi, Macrobrachium pumilum, Macrobrachium dierythrum, and Palaemonetes (Palaemonetes) mercedae. To- dos los especimenes fueron colectados en Venezuela. Se dan detalles del patron de coloraci6on en vivo, de la fecundidad, y de la variacion morfoldgica. Se hacen comparaciones con especies previamente descritas y en especial con las especies suramericanas consideradas mas relacionadas morfologicamente. The decapod fauna of part of South America is relatively well known (Hulbert etal. 1981). The revision of Holthuis (1952) remains the most comprehensive taxonom- ic work treating the palaemonid shrimps. However, some geographical areas have not been extensively studied because of the in- accessibility of many parts of the continent, and the few available collections. Some of the poorer known regions are the Orinoco and Amazon basins, where recently several new species of palaemonids were found (Kensley and Walker 1982, Rodriguez 1982, Pereira 1985). While making a survey of the freshwater shrimps of Venezuela the author found seven undescribed species of palae- monid shrimps, six belonging to the genus Macrobrachium and one to the genus Pa- laemonetes. The purpose of the present pa- per is to present descriptions of these species. The following abbreviations are used: cl., for carapace length; tl., for total length; MBUCYV, Museum of Biology Universidad Central de Venezuela; and USNM, Nation- al Museum of Natural History, Smithson- ian Institution, U.S.A. Macrobrachium Bate, 1868 Macrobrachium reyesi, new species Figs: 1 .6€ Holotype. —Male MBUCYV (XI]I-1707), 44 mm tl.; 10.3 mm cl. Paratypes 18 males MBUCYV (XI-1706), 3 males and 3 females (USNM 228619). Collected 16 Jan 1982 by Guido Pereira and Matias Reyes. Type locality.—Quebrada (stream) Cor- ral de Piedra. El Limon, Maracay, Edo. Ara- gua, Venezuela, 10°15’N, 67°35’W. Etymology. —The species is dedicated to Matias Reyes who first collected specimens of this species in 1974, kindly gave me the sample, and helped me in the field. Additional material. —8 specimens, MBUCV (XI-1216), Bocono River, be- tween Barrancas and Portuguesa, Edo. Tru- jillo, Venezuela. 9°5’N, 70°10’W, Nov 1974; VOLUME 99, NUMBER 2 199 E Fig. 1. Macrobrachium reyesi, new species. A, Cephalic part of cephalothorax; B, Same in detail; C, Scaph- ocerite; D, Telson; E, Posterior part of telson; F, First pereiopod; G, Second pereiopod; H, Third pereiopod; I, Fifth pereiopod. All from paratypic male. Scale = 1 mm. coll. F. Mago.—5 specimens, MBUCV (XI- 1331), pond in the road between Guanare and Guanarito, Edo. Portuguesa, Venezue- la, 8°50'N, 69°10’W, May 1981; coll. D. Ta- phorn and J. Reid.—2 specimens, MBUCV (XI-1615), Stream Grande, Cojedes River system, Edo. Yaracuy, Venezuela, 10°5’N, 68°45'W, May 1978; coll. F. Mago. —3 spec- imens, MBUCYV (XI-1630), Stream Corral de Piedra, El Limon, Maracay, Edo. Aragua, Venezuela, 10°15’N, 67°35'’W, 1974; coll. M. Reyes.—4 specimens, MBUCV (XI- 1781), Stream Grande, Cojedes River sys- tem, Edo. Portuguesa, Venezuela, 9°50'W, 68°45’W, 1978; coll. F. Mago.—25 speci- mens, MBUCV (XI-1708), Stream Corral de Piedra, El Limon, Maracay, Edo. Aragua, Venezuela, 10°15’N, 67°35’W, Jan 1982; coll. G. Pereira and M. Reyes. Description of holotype. —Rostrum straight with tip slightly pointing upward, apex reaching slightly beyond distal end of scaphocerite; upper border bearing 9 (6 to 9 paratypes) regularly distributed teeth; 2 of which behind posterior margin of orbit; small subapical tooth frequently present; lower margin with 3 (3 to 4 paratypes) teeth. Carapace smooth. Scaphocerite 2.7 times longer than wide. Abdomen smooth, pos- teroventral angle of fifth pleuron acute. Sixth abdominal segment 1.3 times length of fifth, 0.4 times that of telson. Telson with 2 pairs 200 of dorsal spines, situated at 12 and % of length from base; posterior margin, tapering abruptly to median apex, bearing 2 pairs of lateral spines, inner of which overreaching median apex, and 8 plumose setae between inner spines. First pereiopods smooth and slender, with tip of dactyl overreaching sca- phocerite; palm cylindrical in cross section, thickest at midlength, and 0.9 times as long as dactyl; carpus 3.2 times length of palm, subequal to merus. Second pair of pereio- pods slender, smooth, equal in size, and overreaching distal border of scaphocerite by length of palm; fingers smooth, closing over entire length; proximal tooth on op- posable margin of each; palm cylindrical in cross section, 2.4 times longer than wide, 0.75 times length of dactyl; carpus 1.3 (1.2- 1.5 paratypes) times length of palm, sub- equal to merus; ischium smooth. Third pair of pereiopods reaching distal border of sca- phocerite; propodus with longitudinal row of 5 spines on inner margin, 2.2 times length of dactyl, 1.3 times that of carpus. Fifth pair of pereiopods reaching distal border of sca- phocerite; propodus, with longitudinal row of 4 (3-5 paratypes) spines on inner margin, 4.0 times length of dactyl, 2.0 times that of carpus. Size. —Largest male measures 38.6 mm tl. and 10.3 mm cl. Largest female 35 mm tl. and 9.0 mm cl. Fecundity. —One ovigerous female, 32.0 mm tl. and 8.5 mm cl. with 22 oval eggs, 2.2-1.5 mm diameter. Larval development. — Abbreviated (Per- eira, in prep). Color. —Translucent in life. Remarks. —This species is similar to M. jelskii (Miers) from which it can be distin- guished by the shape of rostrum: it is rela- tively shorter, and straight in M. reyesi, the apex overreaching the scaphocerite, while M. jelskii has the rostrum slightly arcuate over the eye, overreaching the scaphocerite by about % ofits length; a diagnostic feature of this species is the short length of the sec- ond pair of legs overreaching the scapho- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cerite by half the length of the palm, while in other American species, at least by the entire palm. Macrobrachium pectinatum, new species Figs, 2, 3468 Holotype. —Male MBUCV (XI-1759) 46.8 mm tl. and 9.3 mm cl. Paratypes, 2 males and 2 females MBUCV (XI-1760), 1 male and 2 females (USNM 228620) col- lected 17 April 1982 by Guido Pereira and Ramiro Ruyero. Type locality.—Atabapo River, Sta. Cruz, Territorio Federal Amazonas, Venezuela 3°20'N, 67°29'W. Etymology.—The specific name is de- rived from the Latin word pecten, a comb, to note the pectinate appearance of the cut- ting edges of fingers. Description of holotype. —Rostrum straight with anterior 3 curved upwards, and distal % overreaching distal border of scaphocerite; upper border bearing 8 teeth, 6 of them regularly distributed along pos- terior half, 2 basal ones located behind pos- terior margin of orbit; anterior half un- armed except for 2 small subapical teeth; lower margin with 7 (6-8 paratypes) teeth. Carapace smooth. Scaphocerite 2.3 times longer than wide. Abdomen smooth, pos- teroventral angle of fifth pleuron acute. Sixth abdominal segment 1.4 times length of fifth, 0.8 times length of telson. Telson with 2 pairs of dorsal spines, situated near 2 and ¥4 its length from base; posterior margin ta- pering abruptly to median apex, bearing 2 pairs of lateral spines, inner of which over- reaching median apex, and 8 plumose setae between inner spines. First pereiopods smooth and slender, reaching distal 4 of scaphocerite; palm cylindrical, thickest at midlength, 0.8 times length of dactyl; carpus 4.1 times length of palm, 1.2 times length of merus, 3.5 times length of dactyl; fingers gaping when closed; cutting edge bearing single series of strong hairs, giving pectinate appearance. Second pair of pereiopods smooth and subequal in size, overreaching VOLUME 99, NUMBER 2 > A | | > B tI C D E 201 Fig. 2. Macrobrachium pectinatum, new species. A, Cephalic part of cephalothorax; B, Same in detail: C, Scaphocerite; D, Telson; E, Posterior part of telson; F, First pereiopod; G, Second pereiopod; H, Third pereiopod; I, Fifth pereiopod; J, Detail of dactyl of second pereiopod. All from paratypic male. Scale = 1 mm. distal border of scaphocerite by entire palm; fingers gaping when closed, with cutting edges bearing single row of strong hairs giv- ing pectinate appearance; palm cylindrical in cross section, 1.6 times longer than wide, same length as dactyl; carpus 3.0 times length of palm, 1.6 times length of merus, 2.8 times length of dactyl. Third pair of pe- reiopods overreaching distal border of scaphocerite by 3 of propodus; propodus with row of 8 (7-9 paratypes) spines lon- gitudinally on inner margin, 2.1 times length of dactyl, 1.8 times length of carpus. Fifth pair of pereiopods overreaching distal bor- der of scaphocerite by entire propodus; pro- podus, with longitudinal row of 8 (7-9 para- types) spines on inner margin, 3.0 times length of dactyl, 1.8 times length of carpus. Size. — Largest male 33.9 mm tl. and 9.3 mm cl. Largest female 44.0 mm tl. and 10.7 mm cl. Fecundity. —One ovigerous female, 42.0 mm tl. and 9.9 mm cl. with 24 oval eggs, 2.3-1.5 mm diameter. Color.—Body ground color pink, with some darker zones on base of rostrum, on posterolateral and posterodorsal surfaces of cephalothorax. Abdomen striped with very dark, almost black, bands at junction of seg- ments. Telson with narrow longitudinal dorsal band; uropods with darker zone proximally. Remarks.—This species is similar to M. Jelskii (Miers) and M. reyesi, from which it can be distinguished by the gaping fingers of the chela, and the rostral shape, which is distally curved upwards in M. pectinatum, while straight in M. reyesi; the proximal half of the rostrum is curved over the eyes in M. Jelskii while straight in M. pectinatum, fi- nally there is one tooth behind the margin of the orbit in M. jelskii, while M. pectin- 202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Macrobrachium pectinatum, new species. Color pattern, lateral view of paratypic male. atum usually has two teeth behind the mar- gin of the orbit. Diagnostic features of this species are, additional to the gaping fingers, the walking legs being unusually long, and overreaching the distal border of the scaph- ocerite by the propodus. Macrobrachium atabapense, new species Figs. 4, 5, 6A Holotype. —Male MBUCV (XI-1781) 36.6 mm tl. and 11.0 mm cl. Paratypes: 6 males MBUCV (XI-1761), 4 males (USNM 228618) collected 17 April 1982 by Guido Pereira and Ramiro Ruyero at type locality. 2 males MBUCV (XI-1715), collected 16 Jan 1979 by Kate Clark, Temi River at Yav- ita, 31 Km NE of Maroa, Territorio Federal Amazonas, Venezuela 3°0'N, 67°0'W. Type locality.—Atabapo River, Sta. Cruz, Territorio Federal Amazonas, Venezuela 3°20'N, 67°29'W. Etymology.—The specific name ataba- pense is derived from the name of the Atabapo River. Description of holotype. —Rostrum straight, with apex reaching slightly beyond distal border of scaphocerite; upper border bearing 9 (9-11 paratypes) regularly distrib- uted teeth, 2 of which behind posterior mar- gin of orbit; lower margin with 3 (2-3 para- types) teeth. Carapace smooth. Scaphocerite 3 times longer than wide. Abdomen smooth, posteroventral angle of fifth pleuron acute. Sixth abdominal segment 1.4 times length of fifth, 0.75 times length of telson. Telson with 2 pairs of dorsal spines, situated near '’> and *% its length from base, posterior mar- gin tapering abruptly to median apex, bear- ing 2 pairs of lateral spines, inner of which overreaching median apex, and 9 (9-10 paratypes) plumose setae, between inner spines. First pereiopods smooth and slen- der, overreaching scaphocerite by distal part of merus; palm cylindrical in cross section, thickest at midlength, 1.7 times length of dactyl; carpus 3.4 times length of palm, 1.4 times length of merus. Second pair of pe- reiopods spiny, subequal in shape, about same length, but one more massive; Over- reaching distal border of scaphocerite by distal end of merus; fingers short and strong, gaping when closed; both fingers with con- spicuous teeth; dactyl with 2 teeth, larger VOLUME 99, NUMBER 2 Saal D E 203 Fig. 4. Macrobrachium atabapense, new species. A, Cephalic part of cephalothorax; B, Same in detail; C, Scaphocerite; D, Telson; E, Posterior part of telson; F, First pereiopod; G, Third pereiopod; H, Fifth pereiopod. All from paratypic male. Scale = 1 mm. situated at distal end of proximal third and smaller just proximal to it, remainder of cutting edge smooth. Fixed finger with strong tooth at end of proximal 4 and series of 3 (2-4 paratypes) smaller teeth more proxi- mal; no tubercles present on fingers; palm cylindrical in cross section, 3.0 times longer than wide, widest distally, with numerous longitudinal rows of short spines, those on lower surfaces larger than elsewhere, 1.9 times length of dactyl; carpus 0.9 times length of palm, 1.8 times length of merus; spines scarce distally on ventral margin, rest of merus and ischium smooth. Third pair of pereiopods overreaching distal border of scaphocerite by % of propodus; propodus with longitudinal row of 8 (8—9 paratypes) spines on inner margin, 3.0 times length of dactyl, 1.5 times length of carpus. Fifth pair of pereiopods reaching distal border of scaphocerite; propodus, with longitudinal row of 9 (8-10 paratypes) spines on inner margins; 3.8 times length of dactyl, 1.8 times length of carpus. Size. —Largest male 31.4 mm tl. and 12.0 mm cl. Largest female 26.0 mm tl. and 8.0 mm cl. Ovigerous females not available. Color.—Ground color violaceous with salmon pink longitudinal median stripe, ex- tending from rostrum to 6th abdominal seg- ment. Remarks.—This species is similar to M. quelchi (De Man). The main differences are: the second leg is relatively shorter in MM. atabapense, overreaching scaphocerite by distal end of carpus, while by % of the car- 204 Fig. 5. pereiopod. From a paratypic male. pus in M. quelchi. Also, the ratio carpus: palm is 0.9, and carpus: merus 1.8 in M. atabapense, while 0.75, and 1.2 respectively in M. quelchi; the second legs in M. ata- bapense have more numerous and more prominent spines than has M. quelchi. Fi- nally, they have very different color patterns (Holthuis 1952, Pereira 1985). Macrobrachium dierythrum, new species Figs: 7,78, 12 Holotype.—Male USNM 228323, 18.6 mm tl. and 4.2 mm cl. Paratypes: 3 males, 17 females USNM 228324, collected 15 Jan 1984, by Guido Pereira and Ernesto Panier. Type locality. — Aguaro River, Paso Gar- zerito, Edo. Guarico, Venezuela, 8°10'N, 66°25'W. Etymology.—The specific name is de- rived from the Greek word dierythros mean- ing variegated with red, to call attention to the red color of shrimp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Macrobrachium atabapense, new species. A, Second pereiopod; B, Distal part of chela of second Additional material. —2 males, and 470 females MBUCV (XI-2207); locality, date, and collectors same as above. Description of holotype. —Rostrum straight, with tip reaching distal border of scaphocerite; upper border bearing 9 (8-10 paratypes) regularly distributed teeth, 2 of which behind posterior margin of orbit, third just over or anterior to it; lower margin with 4 (3-5 paratypes) teeth. Carapace smooth. Scaphocerite 3.1 times longer than wide. Abdomen smooth, posteroventral angle of fifth pleuron acute. Sixth abdominal seg- ment 1.5 times length of fifth, 0.6 that of telson. Telson with 2 pairs of dorsal spines, situated at '2 and 4 its length from base; posterior margin acute, tapering abruptly to median apex, bearing 2 pairs of lateral spines, inner of which overreaching median apex, and 4 plumose setae between inner spines. First pereiopods slender, overreach- ing distal border of scaphocerite by dacty]; palm cylindrical in cross section, 1.5 times VOLUME 99, NUMBER 2 205 B C Fig. 6. Appendices masculinae. A, M. atabapense, new species; B, M. pectinatum, new species; C, M. reyesi, new species. From paratypes. Scale = 100 M. length of dactyl; carpus 2.2 times length of palm, 1.1 that of merus. Second pair of pe- reiopods similar in shape and size, over- reaching distal border of scaphocerite by 7 of carpus; fingers straight, both with con- spicuous teeth; dactyl with 2 teeth at 4 and % its length from base, fixed finger with 2 teeth just proximal to each tooth on dacty]; fingers without tubercles; palm cylindrical in cross section, 3.1 times longer than wide, with several longitudinal rows of spines, 1.5 times length of dactyl; carpus 0.8 times length of palm, 0.9 times length of merus, spinulation pattern as on palm; merus with three rows of ventral spines; ischium smooth, except for scattered ventral spines. Third pair of pereiopods not reaching distal border of scaphocerite; propodus with lon- gitudinal row of 7 spines on inner margin, 2.2 times length of dactyl, 1.9 times length of carpus. Fifth pair of pereiopods not reaching distal border of scaphocerite; prop- odus with longitudinal row of 7 spines on inner margin; 2.4 times length of dactyl, 2.2 times length of carpus. Size. — Largest male 18.4 mm tl. and 4.2 mm cl. Largest female 26.0 mm tl. and 6.4 mm cl. Fecundity.—No ovigerous female avail- able. Color.— Background color red, with nu- merous irregular pink spots over body. An- tennal and antennular flagella deep blue. Second leg overall red, some clear pink areas, fingers white. Pereiopods 3—5 pink, with 7 transverse red stripes, 1 basally on ischium, 2 on merus, one basally and other at mid- length, 2 on carpus, one basally and other at midlength, 2 or 1 on propodus, on second and third % length from base. Remarks. —Except for the report of Ro- driguez (1982), there are no records of Mac- 206 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. M. dierythrum, new species. A, Cephalic part of cephalothorax; B, Telson; C, Scaphocerite; D, First pereiopod; E, Third pereiopod; F, Fifth pereiopod; G, Second pereiopod; H, Second pereiopod detail. All from paratypic male. robrachium of such small size in America, probably because rivers in the Guayanas re- gion have not been sampled adequately. Comparisons are made in Table 1. Macrobrachium rodriguezi, new species Figs. 10, 12 Holotype.—Male USNM 228321, 16.2 mm tl. and 3.4 mm cl. Paratypes: 25 males USNM 228322, collected 13 Jan 1984 by Guido Pereira and Ernesto Panier. Type locality. —Caris River, El Tigre, Edo. Anzoategui, Venezuela, 8°45’N, 64°50'W. Etymology.—The species is named in honor of Dr. Gilberto Rodriguez, pioneer of carcinological studies in Venezuela. Additional material.—10 males, 18 fe- males MBUCV (XI-2200), locality, date, and collectors same as above. Description of holotype. —Rostrum arched, with apex reaching third antennular segment; upper border bearing 8 (8—9 para- types) regularly distributed teeth, one of which situated behind margin of orbit, sec- ond just over or behind it; lower margin with 3 teeth. Carapace smooth. Scaphoce- rite 2.5 times longer than wide. Abdomen smooth, posteroventral angle of fifth pleu- ron acute. Sixth abdominal segment 1.4 times length of fifth, 0.8 times length of tel- son. Telson with 2 pairs of dorsal spines, situated at '4 and *% of its length from base; posterior margin tapering abruptly to me- dian apex, bearing 2 pairs of lateral spines, inner of which overreaching median apex, with 7 (6—9 paratypes) plumose setae be- tween inner spines. First pereiopods slen- der, overreaching distal border of scapho- VOLUME 99, NUMBER 2 207 Fig. 8. MM. dierythrum color pattern, lateral view of paratypic male. cerite by dactyl; palm cylindrical in cross section, 1.6 times length of dactyl; carpus 1.7 times length of palm, same length as merus. Second pair of pereiopods smooth, equal in shape and length, overreaching an- terior border of scaphocerite by palm; fin- gers straight, without conspicuous teeth; palm cylindrical in cross section, 2.9 times longer than wide, 1.4 times length of dactyl; carpus 0.8 times length of palm, and 0.8 times length of merus. Third pair of pereio- pods reaching distal border of scaphocerite; propodus with longitudinal row of 7 spines on inner margin; 2.0 times length of dactyl, 1.7 times length of carpus. Fifth pair of pe- reiopods overreaching distal border of scaphocerite by 2 of dactyl; propodus with longitudinal row of 7 spines on inner mar- gin, 3.2 times length of dactyl, 2.2 times length of carpus. Size. —Largest male 16.2 mm tl. and 3.4 mm cl. Largest female 12.4 mm tl. and 3.0 mm cl. Fecundity. —Ovigerous females with 10-— 22 large and oval eggs, average largest di- ameter 1.3 mm. Color.— Background color either brown or blue, with longitudinal middorsal cream stripe, from tip of rostrum to sixth abdom- inal segment. Remarks. —Comparisons are made in Table 1. Fig. 9. MM. dierythrum color pattern, dorsal view of paratypic male. 208 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 10. MM. rodriguezi, new species. A, Cephalic part of cephalothorax; B, Telson; C, Scaphocerite; D, First pereiopod; E, Third pereiopod; F, Fifth pereiopod; G, Second pereiopod. All from paratypic male. Macrobrachium pumilum, new species Figs. 11, 12 Holotype.—Male USNM 228319, 10.5 mm tl. and 3.2 mm cl. Paratypes: 10 males, 16 females USNM 228320, collected 15 Jan 1984 by Guido Pereira and Ernesto Panier. Type locality. —Aguaro River, Cachimbo pass, Edo. Guarico, Venezuela, 8°10'N, 66°35'W. Etymology.—The specific name is de- rived from the Latin word pumilus, mean- ing little, referring to the small size of the shrimp. Additional material.—2 males, 250 fe- males MBUCV (XI-2204), locality, date, and collectors same as above. Description of holotype. —Rostrum strongly arched over eyes with apex reach- ing distal border of scaphocerite; upper bor- der bearing 8 (7-9 paratypes) regularly dis- tributed teeth, 2 of which situated behind posterior margin of orbit; lower margin with 1 (1-2 paratypes) tooth. Carapace smooth. Scaphocerite 3.1 times longer than wide. Abdomen smooth, posteroventral angle of fifth pleuron acute. Sixth abdominal seg- ment 1.8 times length of fifth, 0.8 times length of telson. Telson with 2 pairs of dor- sal spines, situated near '3 and % its length from base; posterior margin tapering abruptly to median apex, bearing 2 pairs of lateral spines, inner of which overreaching median apex, and 6 (6-8 paratypes) plu- mose setae, between inner spines. First pe- reiopods slender, overreaching distal border of scaphocerite by dactyl; palm cylindrical, 0.8 times length of dactyl; carpus 2.2 times length of palm, 0.9 times length of merus. VOLUME 99, NUMBER 2 209 Fig. 11. h 0.5 mm M. pumilum, new species. A, Cephalic part of cephalothorax; B, Telson; C, Scaphocerite; D, First pereiopod; E, Third pereiopod; F, Fifth pereiopod; G, Second pereiopod; H, Detail of second pereiopod. Second pair of pereiopods smooth, sub- equal in size, overreaching anterior border of scaphocerite by length of palm; fingers straight, without conspicuous teeth; palm cylindrical in cross section, 2.6 times longer than wide, 1.1 times length of dactyl; carpus 1.1 times length of palm, 0.8 length of me- rus. Third pair of pereiopods reaching '2 length of scaphocerite with dactyl; propodus with longitudinal row of 6 spines on inner margin, 1.9 times length of dactyl, 1.6 times length of carpus. Fifth pair of pereiopods reaching basal 4 of scaphocerite; propodus with longitudinal row of 6 spines on inner margin, 2.6 times length of dactyl, 2.0 times length of carpus. Size. —The largest male 10.5 mm tl. and 3.2 mm cl. Largest female 24.0 mm tl. and 5.2 mm cl. Fecundity. —One ovigerous female, 16.4 mm tl. and 4.7 mm cl. with 12 oval eggs, 1.6 mm largest diameter. Color.—Overall body lemon yellow in color. Remarks. —The last three species are of very small size as compared to other South American species in the genus. Table | gives a summary of their most distinguishing fea- tures. Palaemonetes Heller, 1869 Palaemonetes (Palaemonetes) mercedae, new species Fig. 13 Holotype. — Male MBUCYV (XI-1782) 3.2 mm cl. Paratypes 1 male and 1 female MBUCYV (XI-1782 B), collected 18 April 1982 by Guido Pereira and Ramiro Ruyero. Type locality.—Atabapo River at Cha- 210 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a == c 50 wm Fig. 12. Appendices masculinae. A, M. rodriguezi, new species; B, M. pumilum, new species; C, M. diery- thrum, new species. All from paratypic males. muchina, Territorio Federal Amazonas, Venezuela, 3°20’N, 67°29'W. Etymology. —The species is dedicated to my wife Mercedes. Additional material.—3 males, 17 fe- males, USNM 228318, Departamento Ature, Puerto Ayacucho, Territorio Federal Amazonas, Venezuela, 5°35'’N, 67°30'W, Dec 1984; coll. R. Vari.—4 males, 8 females USNM 228317, Departamento Rio Negro, between San Carlos and Solano, Territorio Federal Amazonas, Venezuela, 1°56’N, 67°2'W, Dec 1984; coll. J. Fernandez, and O. Castillo. Description of holotype.—Rostrum slen- der and straight, reaching distal end of scaphocerite; upper margin bearing 7 (6—7 paratypes) regularly distributed teeth, one of which placed behind orbit, and smaller than rest; lower margin with 2 teeth. Car- apace smooth. Branchiostegal spine sub- equal in length to antennal spine, not reaching anterior border of carapace; bran- chiostegal groove distinct, extending from midway between branchiostegal and anten- nal spines. Abdomen smooth, pleura of fifth segment antero- and posteroventrally rounded. Sixth segment 1.6 times as long as fifth, 0.7 length of telson. Telson bearing 2 pairs of dorsolateral spines, at midlength and *4 its length from base; posterior margin rounded, ending in median point and bear- ing 2 pair of spines, neither overreaching tip of telson; with 12 (10-14 paratypes) plu- mose setae between inner spines. Eyes nor- mal and well pigmented. Antennule with stylocerite sharp, reaching midlength of proximal segment of antennular peduncle; VOLUME 99, NUMBER 2 211 Table 1.—Distinguishing features between M. dierythrum, M. rodriguezi and M. pumilum; number of post- orbital teeth in parentheses; see additional color pattern in the descriptions. Character M. dierythrum M. rodriguezi M. pumilum Rostrum Straight, overreaching Arched over the eye, Arched over the eye, over- scaphocerite by apex reaching distal antennu- reaching scaphocerite by lar peduncle apex Rostral 8-9, (2) 8-9, (1-2) 7-9, (2) formula 3-5 3-5 1=2 Second leg Spiny, overreaching scaph- Smooth, overreaching Smooth, overreaching scaph- ocerite by 7 carpus. scaphocerite by palm. ocerite by palm. Palm 2.6 Palm 3.1 longer than Palm 2.9 longer than longer than wide, 1.1 wide, 1.5 length of dac- wide, 1.4 length of dac- length of dactyl tyl tyl Appendix With 7 spines on distal 14 With 21 spines over its en- With 12 spines on distal 2 masculina tire length outer margin of basal segment straight with strong anterolateral spines almost reaching midlength of second segment; second seg- ment about as broad as, but distinctly short- er than, third. Lateral antennal flagella with rami fused for 2 articles, free part of shorter ramus consisting of 7 articles. Scaphocerite 2.8 times as long as broad, with mesial mar- gin convex, lateral margin straight, latter ending in distal spine overreached by la- mella. Mouthparts typical of genus. First pereiopods reaching scaphocerite; palm cy- lindrical in cross section, widest at mid- length; fingers slightly longer than palm; carpus 2.9 times as long as dactyl and merus slightly shorter than carpus. Second pereiopods relatively strong, overreaching scaphocerite by length of chela, fixed finger and dactyl each with | tooth on proximal third of cutting edge, and 1 less prominent posteriorly, rest of margin entire, palm cy- lindrical in cross section, 3.6 times longer than wide, 1.7 times as long as dactyl, 0.8 times length of carpus, slightly shorter than merus and same length as ischium. Third pereiopod overreaching scaphocerite by tip of dactyl; propodus 2.5 times as long as dac- tyl, 2 times length of carpus, same length of merus, 2 times length of ischium. Fifth pe- reiopods overreaching scaphocerite by length of dactyl; propodus 3.4 times as long as dactyl, 2 times length of carpus, 1.2 times length of merus, and 2.5 times length of ischium. Pleopods and uropods of usual shape, lateral ramus of uropods with mov- able spine between fixed distolateral tooth and margin of blade. Appendix masculina with 8 spines on distal 2; 4 apical spines aligned in transverse row, and 4 subapical spines in a single longitudinal row (see Flemming 1969 for terminology). Size. —Male holotype tl. 13 mm; cl. 3.2 mm. Female tl. 15 mm; cl. 4.5 mm. Fecundity. —Ovigerous females with 15— 18 bright red spherical eggs, 1.0 mm in di- ameter. Color. —Translucent in life. Remarks. — This new species is similar to P. ivonicus Holthuis. I have compared my specimens with the holotype and paratypic females of the latter at the USNM. The main differences are: the chela of second leg in P. mercedae is relatively more robust, the ratio of dactyl : palm being 1:1.6 in P. mercedae and 1:1 in P. ivonicus. Another distinguish- ing feature is the shape of posterior margin of telson, which is rounded, with the median apex not overreached by the inner spines in the new species, while in P. ivonicus it is truncate and the apex is clearly overreached by the inner spines. There are 8—10 plumose setae on posterior margin of telson in the 212 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 13. Palaemonetes (Palaemonetes) mercedae, new species. A, 3-5 abdominal segments; B, Cephalic part of cephalothorax; C, Scaphocerite; D, Antennule; E, First pereiopod; F, Chela of second pereiopod; G, Second pereiopod; H, Third pereiopod; I, Fifth pereiopod; J, Telson and uropods; K, Appendices interna and masculina. All from a paratypic male. Scale = 1 mm. VOLUME 99, NUMBER 2 new species, while there are only two in P. ivonicus. The most striking characteristic of this species is the short carpus of the second legs, which is shorter than or equal to that of chela. This character sharply separates the species from the previously known freshwater South American Palaemonetes. Strenth (1976) used the morphology of the upper antennular flagellum to separate ma- rine and freshwater species occurring in North America. In this species the distal free portion of the shorter ramus in the lat- eral antennular flagellum is longer than the fused part, being similar in this feature to the group of South American freshwater species of Palaemonetes (P. argentinus No- bili, 1901; P. carteri Gordon, 1935; P. ivon- icus Holthuis, 1950). Acknowledgments The author is indebted to Drs. Horton H. Hobbs, Jr., Raymond B. Manning and Brian Kensley for their advice in styling the manu- script and their critical review. I am grateful to my friends Ramiro Ruyero, Matias Reyes, and Ernesto Panier for helping in the field collections. This work was partially fi- nanced by CONICIT grant S1-1259. Literature Cited Flemming, L. 1969. Use of male external genitalic details as taxonomic characters in some species 213 of Palaemonetes (Decapoda, Palaemonidae).— Proceedings of the Biological Society of Wash- ington 82:443-452. Holthuis, L. 1952. A general revision of the Palae- monidae (Crustacea: Decapoda: Natantia) of the Americas. II The Subfamily Palaemoninae.— Occasional Papers of the Allan Hancock Foun- dation 12:1-396. Hulbert, S., G. Rodriguez, and N. Dos Santos. 1981. Aquatic biota of tropical South America. Part 1 Arthropoda. San Diego State University, Xxli + 323 pp. Kensley, B., and I. Walker. 1982. Palaemonid shrimps from the Amazon basin, Brazil (Crustacea, De- capoda, Natantia).—Smithsonian Contribu- tions to Zoology 362:111, 1-28. Pereira, G. 1985. Freshwater shrimps from Vene- zuela III: Macrobrachium quelchi De Man and Euryrhynchus pemoni, n. sp. (Crustacea: De- capoda: Palaemonidae) from La Gran Sa- bana. — Proceedings of the Biological Society of Washington 3:615-621. Rodriguez, G. 1982. Fresh-water shrimps (Crustacea, Decapoda, Natantia) of the Orinoco basin and the Venezuelan Guayana.—Journal of Crusta- cean Biology 2:378-391. Strenth, N. 1976. A review of the systematics and zoogeography of the freshwater species of Pa- laemonetes Heller of North America (Crustacea: Decapoda). — Smithsonian Contributions to Zo- ology 228:11, 1-27. Universidad Central de Venezuela, Insti- tuto de Zoologia Tropical, Aptdo. 47058, Caracas 1041-A, Venezuela. Present ad- dress: Department of Zoology, University of Maryland, College Park, Maryland 20742, U.S.A. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 214-217 A NEW COLOSTETHUS (ANURA: DENDROBATIDAE) FROM ECUADOR Darrel R. Frost Abstract.—A new species of dendrobatid frog from southeastern Ecuador, Colostethus nexipus, is described. This species is distinguished from other members of Colostethus by the combination of extensive toe webbing and the presence of distinct dorsolateral and oblique lateral light stripes. An undescribed member of the dendro- batid genus Colostethus has come to my at- tention through the generosity of Alice G. C. Grandison of the British Museum (Nat- ural History). The specimens described are from a collection made by the British Los Tayos Archaeological Expedition and pre- sented to the British Museum. Assignment of this species to Colostethus is a matter of convenience because dendrobatids that lack the apomorphies of Atopophrynus, Den- drobates, or Phyllobates are currently re- ferred to the basal grade-genus Colostethus (see Lynch 1982, Lynch and Ruiz-Carranza 1982). Colostethus nexipus, new species Figs. 1, 2 Holotype. —-BMNH 1983.1061, adult male obtained at Los Tayos, Morona-San- tiago Province, Ecuador, 78°12’W, 3°10’S, 9 Jul 1976 by Philip Ashmole and J. K. Campbell. Paratypes.— All from the vicinity of the type locality: BMNH 1983.923 (small male), in a Shallow cave adjacent to the Rio Coan- gos (a tributary of the Rio Santiago), 1 Aug 1976 by Philip Ashmole; BMNH 1983.1060 (small male), same data as holotype; KU 194164 (small male) and BMNH 1983.924 (newly transformed) from bottom (50 m) of main cave shaft, on walls, 9 Jul 1976 by Philip Ashmole and J. K. Campbell. Diagnosis.—A dendrobatid frog not ex- hibiting bright coloration, loss of ears, or fusion of first and second toes; distinguished from all other members of Colostethus by the combination of extensive toe webbing and the presence of distinct dorsolateral and oblique lateral light stripes. Colostethus nexipus cannot be confused with any other known species. Description. —Head as wide as body, slightly wider than long; snout truncate and rounded in lateral view; nostrils small, slightly protuberant and directed anterolat- erally; canthus rostralis rounded but dis- cernible; loreal region concave; lips not flared; snout short; interorbital region flat, slightly narrower than width of upper eye- lid; no tubercles on head; weak supratym- panic fold; tympanum evident; postrictal tubercles not evident; choanae concealed by palatal shelf of maxillary arch when viewed from directly below; vomerine odonto- phores absent; numerous teeth on premax- illae and maxillae; tongue longer than wide, posterior notch very broad and shallow; posterior % not adherent to floor of mouth; vocal slits present; skin of dorsum finely shagreened, not easily abraded; folds on dorsum absent; skin of venter finely sha- greened; vent opening at upper level of thighs; no enlarged warts in vicinity of vent; ulnar fold and tubercles absent; palmar tu- bercle round, about 4 times larger than thenar tubercle; supernumerary palmar tu- bercles absent; subarticular tubercles indis- tinct, round, flat; lateral fringe on fingers present as weak ridge; third finger not swol- VOLUME 99, NUMBER 2 Fig. 1. len; tips of fingers expanded to form pads, approximately 1.5 times width of digit; pair of scutes atop tips of fingers and toes, not differing in coloration from digits; second finger shorter than first (anomalously so on right hand; in paratypes second longer than first); condition of thumbs of breeding males unknown; tubercles absent from knee, heel, or tarsus; tarsal fold weak; inner metatarsal tubercle nearly twice as long as wide, flat, at least twice as large as flat, round outer metatarsal tubercle; supernumerary plantar tubercles absent; subarticular tubercles round to ovoid, flat; toe tips expanded to form pads; toes with prominent lateral fringes confluent with toe webbing; toe web- bing formula (after Savage and Heyer 1967) I O-1 II O-1.7 II 1-2 IV 2-1 V; webbing 215 Paratype of Colostethus nexipus, BMNH 1983.923. Snout—vent length equals 21.3 mm. incised; when hind legs flexed and held at right angles to sagittal plane, heels touch. Coloration in preservative. —(Holotype is formalin-darkened so coloration has been determined, in part, from paratypes.) Dor- sum dark brown with broad dorsolateral stripe extending from eye to vent (stripe broken posteriorly); flanks dark brown with broken, white, oblique lateral stripe extend- ing from point above insertion of forelimb to groin; hind limbs pale brown with dark cross-bars; forelimbs tan with irregular dark blotches; venter pale brown with dark stip- pling, becoming darker in the gular region, not forming collar or discernible spots. Measurements of holotype in mm. — Snout—vent length 23.9; tibia 11.9; head width 8.2; head length 7.8; distance from 216 ES RUE PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Palmar views of left hand and left foot of holotype. Bar equals 5 mm. tip of snout to angle of jaw 8.2; upper eyelid width 2.6; interorbital distance 2.4; eye length 3.5; eye to nostril distance 2.0. Etymology.—The epithet, nexipus, is a Latin noun in apposition, derived from the Greek nexipous, meaning web-foot. Discussion. —The type locality is in rain forest. The types were collected in the vi- cinity of a 50 m deep limestone cave shaft that intermittently receives from the Rio Coangos a waterfall that carries with it tree trunks and other debris. Although two of the paratypes were collected from within this cave, it is doubtful that Colostethus nexipus 1s normally cave-dwelling. Non- cavernicolous species also found in the cave were the gymnophthalmine teiid lizards Alopoglossus buckleyi and Euspondylus guentheri. Colostethus nexipus is a denizen of stream banks and adjacent wet forest, like many of its congeners. The phylogenetic re- lationships of C. nexipus are unclear be- cause it lacks any of the striking apomor- phies (e.g., swollen third finger in males, dark collar, or chest spots) that characterize putative monophyletic groups within Co- lostethus. Acknowledgments I thank Alice G. C. Grandison (British Museum) for allowing me to describe this species and, with A. M. Hutson (British Mu- seum), for providing helpful information. David C. Cannatella, Rafael de Sa, William VOLUME 99, NUMBER 2 E. Duellman, Linda S. Ford, and John D. Lynch made useful comments. Literature Cited Lynch, J. D. 1982. Two new species of poison-dart frogs (Colostethus) from Colombia.— Herpeto- logica 38:366-374. Lynch, J. D., and P. M. Ruiz-Carranza. 1982. Anew genus and species of poison-dart frog (Am- phibia: Dendrobatidae) from the Andes of 217 northern Colombia.— Proceedings of the Bio- logical Society of Washington 95:557-562. Savage, J. M., and W. R. Heyer. 1967. Variation and distribution in the tree-frog genus Phyllomedusa in Costa Rica, Central America.—Beitrage zur Neotropischen Fauna 5:111-131. Museum of Natural History and Depart- ment of Systematics and Ecology, The Uni- versity of Kansas, Lawrence, Kansas 66045. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 218-224 SYSTEMATICS OF THE GORGETED WOODSTARS (AVES: TROCHILIDAE: ACESTRURA) Gary R. Graves Abstract. —Acestrura heliodor (sensu Peters 1945) forms a superspecies com- posed of two allospecies: A. astreans Bangs of the Sierra Nevada de Santa Marta; and A. heliodor (Bourcier) of the Andes of Venezuela, Colombia, and northeastern Ecuador. Acestrura h. meridae Zimmer and Phelps is synony- mized with A. h. heliodor. Acestrura h. cleavesi Moore of northeastern Ecuador is a valid subspecies. Wing and tail lengths are positively correlated with latitude in the heliodor superspecies. Gorget color in males is subject to postmortem change. The diminutive woodstars of the genus Acestrura of the Andean region in western South America are poorly known, under- represented in museum collections, and are frequently misidentified. Acestrura ber- lepschi, A. bombus, and A. mulsant are cur- rently considered to be monotypic. A fourth species, A. heliodor, exhibits considerable geographic variation that is recognized at the subspecific level (Peters 1945, Zimmer 1953): A. astreans of the Sierra Nevada de Santa Marta, Colombia; A. h. heliodor of the Eastern, Central and Western Cordil- leras of the Colombian Andes; A. h. meri- dae of the Venezuelan Andes; and A. h. cleavesi of northeastern Ecuador. The purpose of this paper is to re-evaluate the taxonomy of woodstars in the 4. helio- dor complex, incorporating data from pre- viously unreported series of specimens col- lected from 1946-1952 in Colombia by M. A. Carriker, Jr. and deposited in the Na- tional Museum of Natural History, Smith- sonian Institution (USNM). Differences among populations in plumage color in both sexes and tail configuration in males suggest that the Acestrura heliodor (Fig. 1) is ac- tually composed of two allospecies: A. as- treans (Santa Marta Woodstar) of the Sierra Nevada de Santa Marta, and A. heliodor (Gorgeted Woodstar) (A. h. heliodor and A. h. cleavesi) of the main Andes. Acestrura heliodor heliodor (Bourcier) Ornismya heliodor Bourcier, 1840:275. “Bogota.” Acestrura heliodor meridae Zimmer and Phelps, 1950:1. Paramo Conejos (4000 m), Mérida, Venezuela. Characters. — Male: Upperparts and flanks are green and the gorget purple. The formula of rectrix length in closed tail is: 3 > 4 > 5 > 2 > 1 (retrices numbered from the in- side outward). The width of rectrix 4 is greater than '2 the width of rectrix 3 and nearly intermediate in width between rec- trices 3 and 5 (Fig. 2). Females: The chin, throat, breast, and flanks are rich buffy cin- namon. In adult females, the tail is rufous with a broad black band across the center. A trace of green occurs just proximal to the band on the central rectrices. The lower rump and upper tail coverts are rufous; a few feathers have a green central spot. Distribution. —Scattered localities in the Andes of Venezuela, and the Eastern, Cen- tral, and Western Cordilleras of the Colom- bian Andes (Fig. 1). The reported occur- rence of A. heliodor at Cana, Cerro Pirre, Panama (Wetmore 1968, Ridgely 1976) is based on a misidentified specimen of Cal- liphlox mitchelli (Robbins et al. 1985). Specimens examined. —VENEZUELA: Pinos (USNM 1 32): ““Merida’” (USNM 1 4, 219 VOLUME 99, NUMBER 2 o eS ou 2) ag aS as} 38 ad 2s fe| 9 er BS =) Q | “el & Es oan—| o Oo Nn =| oO S| —_ iS) oO Q, Nn [=| o) ES) n so 3 aS} dey {2} — = 8 2 x2 3 S02 sod x Oo Ss ae ay (at; An Sc mo cape ovo’ ans ~ Os 6 Cr q@ Oss gE8 — e238 fe) fel Ss Pr nA, Bos (3) (She Xe : a» il 7 & ob 6 9 meso a fel . Square above 1000 feet elevation 220 heliodor (\s astreans e i Fig. 2. Tail patterns of Acestrura h. heliodor, A. h. cleavesi and A. astreans. Males on the left, females on the nght. Color of rectrices: stippling = green; un- marked = rufous or rich buff; shaded = black. 1.8 x natural size. 2 92; AMNH 19 44, 9 22).— Conejos (USNM 1 9; AMNH 1 6¢ type “meridae,” 1 2).— Tambor (USNM 1 4, 4 92; AMNH 9 24¢, 2 92).—Escorial (AMNH 4 _ 4é6).—Tierra (USNM 1 8). COLOMBIA: (unspecified lo- calities) “Bogota” (USNM 4 44, 2 92: AMNH 24 366 including type of A. h. heliodor, 9 22).—““Lower Magdalena’ (USNM 1 4, 1 ?).—““Colombia”’ (MLOC 3 646; USNM 4 28, 2 22; AMNH 2 8¢).—‘“‘Santiago”’ near Pasto? PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (FMNH 1 2).—‘“‘New Grenada” (USNM 1 6, 1 9; AMNH 2 24, 2 29). Norte de Santan- der: Buenos Aires (USNM 7 44, 3 22).— Ocafia (USNM 1 é4).—Ramirez (CM 3 44, 1 2; ANSP 1 2).—Cachiri (CM 1 2).—Las Ven- tanas (CM 2 929). Caldas: Laguneta (ANSP 1 6, 3 99). Huila: San Agustin (USNM 4 66, 2 92; AMNH 1 6; ANSP 3 64, 2 29).— Belén (USNM 1 3). Cauca: El Tambo (FMNH 1 6; WFVZ 1 6).—Tieras (FMNH 1 ).—Moscopan (USNM 1 9). Acestrura heliodor cleavesi (Moore) Chaetocercus cleavesi Moore, 1934:1. Cu- yuja, Ecuador. Characters. —Male: The formula for the length of rectrices in A. h. cleavesi is: 3 > 4> 2>5> 1, wstead of 34 2 > 1, asin A. h. heliodor and A. astreans. Rectrices 4 and 5 are similar in width, and much narrower than rectrix 3. Female: Sim- ilar to A. h. heliodor but more richly colored on chin and throat and with rump more extensively rufous. Distribution. —Known only from the Amazonian slope of the Andes in north- eastern Ecuador. Specimens examined. -ECUADOR: Baeza (MLOC 3 4¢, 6 92; USNM 1 4, 1 2; AMNH 1 4).—Cuyuja (MLOC 1 4, 1 2).— Pallatanga (MLOC 1 92).—Rio Hollin (MLOC | 2).—Rio Oyacachi Abajo (AMNH 1 3, 3 22).—Rio Tigre (MLOC 1 4, 3 22).— Tumbaco (MLOC | 9). Acestrura astreans Bangs Acestrura astreans Bangs, 1899:76. San Se- bastian (6600 feet), Sierra Nevada de San- ta Marta, Colombia. Characters. —Male: In adults, the back, rump, upper tail coverts, and flanks are me- tallic bluish-green, instead of green as in A. h. heliodor and A. h. cleavesi. The gorget is red or reddish-purple, depending on the an- gle of reflection, not purple as in A. h. he- liodor. The formula of rectrix length is the VOLUME 99, NUMBER 2 same as in heliodor. Rectrices 4 and 5 are very narrow, less than '2 the width of rectrix 3. Female: Adult females differ from those of A. heliodor in having the upper tail co- verts green instead of rufous. The central rectrices of A. astreans are green, instead of rufous with a black band like the other rec- trices, as in A. heliodor (Fig. 2). The venter of A. astreans is lighter, not as richly colored as in A. heliodor. Distribution. —Restricted to the Sierra Nevada de Santa Marta. Specimens have been taken on the western, eastern, and southern slopes at elevations between 2700 and 6600 feet (ca. 825-2010 m). Specimens examined. —Sierra Nevada de Santa Marta: San Sebastian (USNM 2 29). — Chinchicua (USNM 4 99).—Vista Nieve (USNM 1 4, 2 22).—Cincinati (ANSP 1 4, 1 ?).—El Mamon (AMNH 1 4).—No further locality data (USNM 2 6¢; AMNH 1 un- sexed). Variation in Male Gorget Color Gorget color in the Acestrura heliodor su- perspecies exhibits significant geographic variation. In contemporaneously collected specimens from the main Andes, gorget col- or varies clinally from pinkish-purple in northeastern Ecuador to purple in the East- ern Cordillera of Colombia and the Vene- zuelan Andes; specimens from the Western and Central Cordilleras are intermediate. The Santa Marta population is character- ized by a dark red gorget. Zimmer and Phelps (1950) separated the Venezuelan (A. h. meridae) populations from those of Colombia on the basis of ‘“‘a darker, more purplish, less reddish throat” in males. Unfortunately, assessment of this character is hampered by previously unrec- ognized postmortem change in gorget color. They compared their Venezuelan series, collected mostly between 1903 and 1921, with a large series of pre-1900 “Bogota” specimens, most of which were probably obtained from the Eastern Cordillera. These have consistently pinker, less purplish gor- 221 gets than more recently collected specimens from the Eastern Cordillera. For example, specimens from Buenos Aires, Norte de Santander, taken in 1946, have slightly more purplish, less pinkish gorgets than speci- mens collected in 1916 at Ramirez, Norte de Santander. In turn, the “1916” speci- mens have gorgets that are more purplish, less pinkish, than those of pre-1900 “‘Bo- gota’’ specimens. Specimens collected at ap- proximately the same time in the Eastern Cordillera and the Venezuelan Andes are indistinguishable in gorget coloration. Aces- trura h. meridae Zimmer and Phelps, 1950, should thus be considered a synonym of A. h. heliodor Boucier, 1840. In another ap- parent example of postmortem change from shorter to longer wave lengths, the gorgets of specimens (USNM 333521; MLOC 7015, 7016, 7023, 10367, 10377) included in the type series of A. h. cleavesi) are presently matched closely by Rose Color (capitalized names from Ridgway 1912), instead of Rhodamine Purple (Moore 1934). The magnitude of postmortem change observed in specimens collected from a sin- gle locality equals that of the contempora- neous geographic variation found among the Colombian populations of A. h. heliodor. The gorgets of birds from the Central (Be- lén, Tijeras, Laguneta) and Western Cor- dilleras (El Tambo) are pinker, less purplish than those of contemporaneously collected (1942-1957) specimens from the Eastern Cordillera, but match those of older speci- mens from the same region. These observations suggest that there are some discrete differences among heliodor populations. The lack of contemporaneous- ly collected series from key populations, however, prevents the subspecific partition of A. heliodor on the basis of gorget color alone. Size and Shape Variation Wing length is positively correlated with latitude in males (Fig. 3; n = 31, r? = 0.586, P < 0.0001) and females (n = 39, r? = 222 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 5s (0 y 0 £x15N LATITUDE Fig. 3. Relationship of wing length with latitude in the Acestrura heliodor superspecies. Squares = A. as- treans; triangles = A. h. heliodor; diamonds = A. h. cleavesi. 0.522, P < 0.0001). Tail length in males, an important taxonomic character (Moore 1934), is also positively correlated with lat- itude (Fig. 4; r> = 0.761, P < 0.0001) and wing length (r* = 0.658, P < 0.0001). These correlations conform to Bergmann’s Rule (cf. Handford 1983, Remsen 1984) and sug- gest that tail length is more closely linked with allometry and aerodynamic function, than to sexual display. Culmen length is un- correlated with either latitude or wing length (P > 0.05) in either sex. Tail and wing length in males of A. astreans and A. heliodor do not overlap. I performed a Principal Com- ponents Analysis of culmen, wing, and tail lengths of males. Not surprisingly, all three taxa (A. astreans, A. h. heliodor, A. h. cleavesi) had nonoverlapping distributions along the axis of the first principal com- ponent (which explained 60.8% of the vari- ance). These taxa are also readily identified by the proportional width of the outermost rec- trices (numbers 3-5) in males. The propor- tional width of rectrices is not clinal. Pop- ulations of A. h. heliodor from the Western, Central, and Eastern Cordilleras of Colom- bia and the Venezuelan Andes are not dis- tinguishable from one another on the basis 5 10° 15°N LATITUDE 16 18 20 22 24 26 Fig.4. Relationships of tail length with latitude and wing length in males of the Acestrura heliodor super- species. Squares = A. astreans; solid triangles = A. h. heliodor, empty triangles = pre-1900 ““Bogota”’ speci- men identified as A. h. heliodor on basis of plumage but lacking definite locality data; diamonds = A. h. cleavesi. of tail shape, but are easily distinguished from A. h. cleavesi and A. astreans. As ex- pected, ““Bogota” specimens have wing/tail ratios similar to specimens from known lo- calities in the Eastern Cordillera (Fig. 4). Taxonomic Conclusions Acestrura astreans differs from A. helio- dor in several important characteristics that are not the terminal states of observed clinal variation, and that support the recognition of A. astreans as a full species. The most important of these are: (1) the distinctive shape of the rectrices in the males; (2) the coloration of body plumage in both sexes; (3) pattern and color of the central rectrices of females; (4) the gorget color in males. Because of their allopatric distributions, the existence of reproductive isolating mecha- VOLUME 99, NUMBER 2 nisms between astreans and heliodor can only be surmised. However, the morpho- logical differences between these taxa are of the same scale as those observed among the other species of Acestrura and Chaetocercus Jourdanii. The high degree of endemism in the Santa Martas has long been recognized (Chapman 1917, Todd and Carriker 1922). At least 12 other avian taxa endemic to the Santa Marta massif, with affinities in the northern Andes, are recognized as specifi- cally distinct from their most closely related congeners (Pyrrhura viridicata, Campylop- terus phainopeplus, Coeligena phalerata, Ramphomicron dorsale, Synallaxis fusco- rufa, Cranioleuca hellmayri, Grallaria bangsi, Myiotheretes pernix, Myioborus fla- vivertex, Basileuterus basilicus, Anisogna- thus melanogenys, Atlapetes melanocepha- lus). The systematic status of Acestrura h. cleavesi is uncertain. Except for rectrix shape in males, plumage differences between A. h. cleavesi and A. h. heliodor appear to be pri- marily quantitative and clinal. The report of A. h. heliodor in Ecuador (Moore 1934) is based on three specimens with definite locality data: Two adult females (MLOC 3083, Pallatanga, 1°59’S; MLOC 3084, Tumbaco, 0°13’S); and an immature male (MLOC 3086, Pallatanga). Both localities are on the Pacific slope of the Andes. Be- cause the plumage of the immature male of A. h. cleavesi is unknown, the male speci- men cannot be identified to subspecies. The females resemble the more heavily pig- mented individuals of the nominate race from Colombia but also match the palest examples of A. h. cleavesi from Baeza, near the type locality. Adult male specimens will be needed to determine the racial affinity of the Pacific slope population in Ecuador. On geographical grounds alone, I tentatively consider these specimens as A. h. cleavesi. A female (FMNH 45429, received from the ““Museum Boucard’’) was collected by Delattre before 1850 at “Santiago, Colom- bia.” If correctly located (de Schauensee 223 1949, Paynter and Traylor 1981), Santiago (1°08’N) is ESE of Pasto, midway between populations of A. h. cleavesi at Cuyuja (0°24'N) on the Amazonian slope of the Ec- uadorian Andes and those of A. h. heliodor at San Agustin (1°53’N) in the Upper Mag- dalena Valley. Apparently, no specimens of Acestrura heliodor have been collected be- tween Cuyuja and San Agustin during the past century. The Santiago specimen is in- distinguishable from typical females of A. h. heliodor, and differs from all specimens of A. h. cleavesi in the intensity and distri- bution of rufous and cinnamon buff on the underparts and rump. Appropriate habitat is available in the distributional hiatus, and the two taxa al- most certainly intergrade or come into con- tact somewhere between Santiago and Cu- yuja. I recommend that cleavesi should best be regarded as a subspecies, rather than a full species, until specimens from the ap- propriate regions are obtained. Acknowledgments For the loan of specimens I thank the curators and staff of the American Museum of Natural History (AMNH), Academy of Natural Sciences, Philadelphia (ANSP), Carnegie Museum of Natural History (CM), Field Museum of Natural History (FMNH), Moore Laboratory of Zoology, Occidental College (MLOC), and the Western Foun- dation of Vertebrate Zoology (WFVZ). I thank Storrs L. Olson, Kenneth C. Parkes, J. V. Remsen, and Richard L. Zusi for many helpful comments. This work was support- ed by a Smithsonian Postdoctoral Fellow- ship. Literature Cited Bangs, O. 1899. Ona small collection of birds from San Sebastian, Colombia.— Proceedings of the New England Zoological Club 1:75-80. Bourcier, J. 1840. Oiseau-mouche nouveau. — Revue Zoologique, p. 275. Chapman, F. M. 1917. The distribution of bird-life 224 in Colombia.— Bulletin American Museum Natural History 36:1-729. de Schauensee, R. M. 1949. Birds of the Republic of Colombia. Part 2.—Caldasia 23:381-644. Handford, P. 1983. Continental patterns of morpho- logical variation in a South American spar- row.—Evolution 37:920—-930. Moore, R. T. 1934. A new species of hummingbird, genus Chaetocercus from eastern Ecuador.— Condor 36:1-6. Paynter, R. A., Jr., and M. A. Traylor, Jr. 1981. Or- nithological gazetteer of Colombia. Harvard University, Cambridge, Massachusetts, 311 pp. Peters, J. 1945. Check-list of birds of the world, vol. 5. Cambridge, Massachusetts: Museum of Com- parative Zoology. Remsen, J. V., Jr. 1984. Geographic variation, zoo- geography, and possible rapid evolution in some Cranioleuca spinetails (Furnariidae) of the An- des.— Wilson Bulletin 96:515—523. Ridgely, R.S. 1976. A guide to the birds of Panama. Princeton University Press, Princeton, New Jer- sey, 354 pp. Ridgway, R. 1912. Color standards and color no- menclature. Washington, D.C. [Published by the author. ] Robbins, M. B., T. A. Parker III, and S.E. Allen. 1985. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The avifauna of Cerro Pirre, Darien, eastern Panama. Jn P. A. Buckley, M. S. Foster, E. S. Morton, R. S. Ridgely, and F. C. Buckley, eds., Neotropical ornithology. Ornithological Mono- graphs No. 36, pp. 198-232. Todd, W. E. C., and M. A. Carriker, Jr. 1922. The birds of the Santa Marta region of Colombia: A study in altitudinal distribution.— Annals of the Carnegie Museum 14:1-611. Wetmore, A. 1968. The birds of the Republic of Pan- ama. Part 2—Columbidae (pigeons) to Picidae (woodpeckers).—Smithsonian Miscellaneous Collections 1:1-605. Zimmer, J. T. 1953. Studies of Peruvian birds. No. 63. The hummingbird genera Oreonympha, Schistes, Heliothryx, Loddigesia, Heliomaster, Rhodopis, Thaumastura, Calliphlox, Myrtis, Myrmia, and Acestrura.—American Museum Novitates 1604:1-26. , and W. H. Phelps. 1950. Three new Vene- zuelan birds.—American Museum Novitates 1455:1-7. Department of Vertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 225-236 TONOCOTE, A NEW GENUS AND SPECIES OF ZOBRACHOIDAE FROM ARGENTINA (CRUSTACEA: MARINE AMPHIPODA) Janice Clark and J. L. Barnard Abstract.—A new genus and new species of marine zobrachoid amphipod, Tonocote magellani, is described from the Magellan Strait. It is a fossorial amphipod digging in shallow sands like a sand-crab. It differs from Prantinus, an Australian genus, in the broad article 4 of antenna 2, the much reduced epimeron 2 and the absence of posterior spines on article 5 of pereopods 3-4. Like Prantinus it differs from other zobrachoids in the urothoe-like antenna 1 with article 3 elongate. The new genus Jonocote and its new species 7. magellani are described from Ar- gentina. Owing to the presence of small epi- meron 2 the inclusion of Tonocote in the Zobrachoidae requires emendation of the diagnosis in that family. Within the Zobrachoidae Tonocote re- sembles Prantinus, from Australia, in the urothoe-like antenna | with article 3 elon- gate. These two genera therefore comprise a subgroup differing from the other two gen- era of the family having a normal haustoriid antenna | with article 3 short. Owing to the early stages of investigation into these an- tipodeal groups, there may be future cause to divide these groups at subfamily level or higher. References used in the identification pro- cess are: Barnard and Drummond (1978, 1982), Barnard and Clark (1982a, b, 1984). Methods of morphological description fol- low Barnard and Drummond (1978). Master Legend Uppercase letters refer to parts; lower case letters to left of uppercase letters refer to specimens noted in legends; lower case let- ters to right of uppercase refer to adjectival modifications in list below: A, antenna; B, body; C, coxa; D, dactyl; E, eye (? or brain), subdivisions marked as E1, E2, E3, E4 (latter = ganglion); G, gnatho- pod; H, head; I, inner plate or ramus; J, pleopodal coupling hooks; L, labium; M, mandible; N, epimera; O, outer plate or ra- mus; P, pereopod; R, uropod; S, maxilliped; T, telson; U, labrum; W, pleon; X, maxilla; Y, pleopod; Z, gill; d, dorsal; r, right; s, setae removed; t, left. Zobrachoidae Barnard and Drummond, 1982 Diagnosis (emendations in italics). — Ros- trum well developed (for haustoriids), cheek poorly developed. Antenna | variable, ar- ticle 1 short (typical) or elongate (apo- morphic), articles 2-3 progressively shorter (typical) or elongate (apomorphic), flagella elongate (typical) or not (apomorphic), ar- ticles of peduncle weakly (typical) to strong- ly geniculate. Antenna 2 of haustorius form, article 4 expanded (plesiomorphic) or weak- ly so (apomorphic), article 5 shorter and narrower than article 4, these articles fur- nished with | or more longitudinal rows of facial armaments, ventral margin of article 4 with at least 3 kinds of setae: (1) elongate plumes, (2) shorter and stiffer glassy spines (often set in clusters) and, (3) bulbar-based penicillate setules; flagellum longer than ar- 226 ticle 4 of peduncle. Prebuccal complex mas- sive, upper lip usually dominant. Mandibles bearing elongate strongly toothed incisors, rakers almost simple and numerous (4 or more), molar large, strongly extended, weakly triturative but with several strong cusps, usually 1 of these forming accessory chopper; palp 3-articulate, article 3 with nu- merous outer setae, setae awned (apo- morphic) or not (typical and plesiomorph- ic). Lower lip with discrete inner lobes, mandibular extensions of outer lobes well developed. Maxilla 1 with uniarticulate palp, inner plate with more than 3 setae. Maxilla 2 ordinary, inner plate with oblique facial row of setae but poorly developed. Maxil- lipeds with unexpanded bases, normally enlarged plates, outer spinose; palp 4-ar- ticulate, article 2 expanded, article 4 cla- vate, at least 2+ setae apically. No baler lobes on maxillae or maxillipeds. Coxa 2 small to medium, larger than coxa 1 and forming stepped intergrade between coxa 1 and coxa 3, coxa 4 dominant, coxa 3 lacking deep posteroventral lobe. Coxal gills on segments 2—6 or 2-5. Brood plates slender. Gnathopods feeble, subchelate, grossly alike in proportions, wrists elongate, article 3 short. Article 5 of pereopods 3-4 broad, slightly expanded, not deeply lobate; dactyls of pereopods 3-5 well developed, those of pereopods 6-7 variable; pereopod 5 of haustorius form, articles 2, 4, 5 and 6 ex- panded, articles 5—6 with extensive facial rows of spines; pereopods 6-7 alike, article 4 broader than 6, articles 5-6 weakly ex- panded; no pereopod with underslung ar- ticulation. Pleopod 2 usually inferior in size, number of articles, or setation; peduncles of pleo- pods not longer than wide, inner rami in- ferior; coupling hooks paired on each pleo- pod, usually inner rami bearing one basal clothespin spine. Epimeron 1 moderately to strongly developed; epimeron 2 dominant in setation, often dominant in size. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Urosomites ordinary, though often fur- nished with lateral teeth. Rami of uropods 1—2 linguiform, setose (not spinose); uropod 3 of ordinary gammarid-phoxocephalid kind, outer ramus dominant, 2-articulate, peduncle short, flat, expanded; rami poorly setose apically. Telson variable in length, deeply cleft. Sexual dimorphism weak. Variables. — Right and left laciniae mo- biles not alike, right, if present, not distinct from raker row (Zobracho and Tonocote); palp article 3 outer setae awned (apomor- phic) or not (typical and plesiomorphic). Maxillipedal palp article 4 multisetose or with main nail and 2 setules (Prantinus). Article 5 of pereopods 3—4 with thick pos- terior spines or only distal spines present (Tonocote). Type genus.—Zobracho J. L. Barnard, 1961. Composition. —Bumeralius, Prantinus, Tonocote. Relationship. — Until our new genus came to light zobrachoids differed from urothoids in the absence of a ventral cephalic cheek, in the full development of the haustorius antenna 2, especially in the ventral arma- ment, and in the dominance of setation (or actual size) on epimeron 2. This epimeron in Tonocote is much smaller than epimeron 3 and barely has dominant setation (2 setae versus | on epimeron 3). Zobrachoids bear linguiform rami of uropods 1-2, in contrast to urothoids (but 1 genus of urothoid lacks rami). Prantinus is furnished with a uro- thoid antenna 1, and has epimeron 2 dom- inant, but antenna 2, though not fully ex- panded, lacks seriate ranks of spines, and bears the ventral spination diversity not typical of urothoids. There is not a great deal of difference between Urohaustoriidae and Zobrachoidae except that zobrachoids have gnathopod 1 subchelate, epimeron | is clearly defined, the mandibular molar is less strongly triturative, weaker and fur- nished more with side cusps, mandibular VOLUME 99, NUMBER 2 rakers are better developed, article 2 on the outer ramus of uropod 3 is usually better developed, coxae 1-2 are both smali and contrasted with a very large coxa 3, and the rostrum is larger. However, Prantinus of the Zobrachoidae intergrades some of these dif- ferences. In addition our new genus TJono- cote differs from urohaustoriids in the less expanded article 2 of the maxillipedal palp, the dactyl lacking inner setae. Key to the Genera of Zobrachoidae (Males) 1. Antenna 1 of urothoe form — Antenna 1 ofhaustorius form .... 3 2. Antenna 2 article 4 broad, epimeron 2 much smaller than 3, posterior spines on article 5 of pereopods 3- 4 absent (distals present) S36 6 Tonocote, new genus — Antenna 2 article 4 slender, epi- meron 2 as large as 3, posterior spines present on article 5 of pereo- WOOSISE4 Boies iat ge ds wae 3 Prantinus Barnard and Drummond, 1982 3. Telson elongate, rami of uropods 1- 2 with many medial setae, no ba- soventral setae Zobracho Barnard, 1961 — Telson short, rami of uropods 1-2 lacking medial setae, bearing baso- ventral setae Bumeralius Barnard and Drummond, 1982 ee ee ew we we ew ew ww Tonocote, new genus Diagnosis. — Rostrum short and broad but head extended strongly anteriad from an- tennal notch. Peduncle of antenna 1 some- what elongate, stout, articles 2 and 3 of pe- duncle progressively shortened, geniculate between articles 1 and 2, both flagella mod- erately long. Aesthetascs simple. Antenna 2 of haustorius form, article 4 expanded, ar- ticle 5 small, articles 4—5 with facial ar- maments, article 4 with long ventral setae, subventral clusters of simple setae and facial armament row. Mandibular incisors slight- 22a ly extended, of ordinary thickness, toothed; rakers 5 or more, serrate; molar small, thin, extended, with 1 main and 3-5 subapical cusps plus 3 marginal setae; setae of palp article 3 not awned, apically hooked. Man- dibular lobes of lower lip well developed. Inner plate of maxilla 1 of medium size, sparsely setose, outer plate with 8 spines, palp short. Inner plate of maxilla 2 with weakly submarginal row of setae. Inner plate of maxilliped ordinary; outer plate with spines; palp article 2 expanded, article 3 not extraordinarily elongate, slightly expanded apically, dactyl unguiform, elongate, bear- ing apical nail and subsidiary setae. Coxae 1—4 progressively larger, each slightly produced posteroventrally, coxae 1— 2 small, subequal in size, coxae 2-6 with simple gills; oostegites unknown. Gnathopods small, grossly alike, wrists elongate, hand somewhat smaller, mitelli- form, subchelate, but palm more transverse on gnathopod 2 than on gnathopod 1. Dac- tyls of pereopods 3-7 distinguishable, those of pereopods 3-5 large, those of pereopods 6—7 very small; dactyl of pereopod 5 blade- like, lacking spines. Article 2 of pereopods 5-7 expanded less strongly on pereopod 6 than on 5 and 7; pereopod 5 of haustorius form; distal articles of pereopods 6-7 not underslung, 6 moderately widened, 7 more expanded; pereopods 6—7 otherwise similar, dominating pereopod 5. Pleopod 2 slightly inferior, inner rami shorter than outer. Epimeron 2 dominantly setose, epimeron 3 dominant in size. Uro- somites weakly produced and weakly setose ventrally. Rami of uropods 1-3 styliform, each outer ramus bearing 2 apical plumose setae, each inner ramus bearing | apical plu- mose seta; peduncles weakly setose. Uropod 3 outer ramus biarticulate and dominant, inner ramus with one basomedial seta. Tel- son short, broader than long. Description. —Eyes weak, ocular ganglia visible. Dorsolateral surface of article 1 on antenna | furnished with small, poorly or- 228 ganized group of setae; article 2 moderately setose dorsolaterally; article 3 poorly setose. Article 3 of antenna 2 short, sparsely setose, flagellum much longer than article 4 of pe- duncle. No calceoli observed. No right lacinia mobilis; left slender, bi- fid. Lower lip lacking cones. Inner plate of maxilla 1 with sparse apical and medial se- tae; several spines on outer plate bifid. Inner plate of maxilliped with 3 stout apical spines (right side missing 2). Coxae 1-3 poorly setose, coxae 4 mod- erately setose. Gills forming stepped inter- grades with gill 2 dominant. Gnathopod 2 lacking surficial buttons. Pereopods 3—4 lacking extensive second- ary facial rows of spines and ventral spines on article 5. Uropods with sparse dorsal setae, medial margins of peduncles sparsely setose. Type species. — Tonocote magellani, new species. Etymology.—Named for a group of In- dians in South America; masculine. Composition. — Unique. Tonocote magellani, new species Figs. 1-6 Diagnosis. —With the characters of the genus. Description of male.— Holotype male “a,” 2.87 mm; head about 90 percent as long as wide, rostrum about 31 percent as long as remainder of head, eyes represented by granular tissue patches, no distinct omma- tidia, ocular ganglia visible. Facial formula of setae on article 1 of antenna 1, ventral = 4 penicillate, dorsal = 2 setae + 1 medium, 1 tiny penicillate, (noting that dorsal—ven- tral aspect reversed from normal, non-uro- thoid kind of antenna 1); article 2 with par- tial circle of 8 long plumose setae; primary flagellum with 4 articles, aesthetasc for- mula = 1-1-1-0; accessory flagellum of 4 ar- ticles. Article 3 of antenna 2 with 2 medium setae; facial formula of spines on article 4 = 2-2-1-2; article 5 = 1 seta dorsally and 2 setae, 1 tiny penicillate; flagellum of 4 articles. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Upper lip with granulations. Right and left mandibular incisors with 3 and 5 teeth; 6 right rakers (no lacinia mobiles), 5 left; each molar with large main cusp bearing 2 long thin basal accessory cusps and appos- ing minor cusps on each side more apical, plus longer thin seta from base opposite to most basal accessory cusp; article 3 of palp slightly longer than article 2, latter with 1 inner seta, spine formula on right and left article 3 = 4-1-2. Inner plate of maxilla 1 bearing 2 apical setae and pair of mediofa- cial setae; outer plate with 8 spines; palp with 3 apical setae. Inner plate of maxilliped with 3 stout spines, 2 medial and 2—4 apical setae; me- dial margin of outer plate with ragged mix- ture of spines and scattered small setae; apex with 1 seta; article 2 of palp with row of 5 mediofacial setae; article 3 with one subfa- cial seta and large serrate spine at base of dactyl. Coxa | subrectangular, convex anteriorly, bearing 1 seta on ventral margin and pos- teroventral long plume and setule; coxa 2 similar in shape to coxa | but anterior con- vexity greater, with 2 long plumes and 2 setules; coxa 3 similar to coxa 2 but more elongate, with 3 plumose posteroventral se- tae and | seta more anterior; coxa 4 adze- shaped with 7 long plumose setae along ventral and posterior margins, 1 short an- teroventral seta. Setal and spine formulas on pereopod 3 = 2,2, 2-0, 2+1+1; on pereopod 4 = 2,2, 2-0, 2+1+1; margins of articles 5—6 not serrate. Article 2 of pereopods 5—7 armed sparsely with long setae posteriorly; dactyl of per- eopod 5 with small anterior tooth. Peduncular spine formulas of pleopods 1-3 = 2 and 0,2 and 0,2 and 0; segmental formulas = 8-5, 7-4, 8-6; basal setal for- mulas = 7-0-1-1, 4-1-1-1, 6-1-1-2, one pe- duncular seta each on pleopods | and 3 (2 naked). Epimeron | rounded quadrate, with | tiny setule posteroventrally; epimeron 2 extend- ed posteroventrally, posterior margin ‘“‘crimped,”’ 1 marginal posteroventral seta VOLUME 99, NUMBER 2 Me) Fig. 1. Tonocote magellani, holotype male “a” 2.87 mm. 230 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Tonocote magellani, holotype male “a” 2.87 mm. VOLUME 99, NUMBER 2 231 \ i Ci eS ee Lk op SRS Q\ Se ph os ZL } aS “ ~ FEN EN = EY Na NFo~ Ditess Son LO BROS ONY She ESS. WS MARY ie. Be. SRN Wal See SSS Fig. 3. Tonocote magellani, holotype male “a” 2.87 mm. 232 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and | tiny facial seta near corner; epimeron 3 subsharply produced posteroventrally with 1 setule on posterior margin. Apicomedial corners of peduncles on uro- pods 1-3 with | plumose seta; apicolateral corner on peduncle of uropod 1 with 1 plu- mose seta; uropod 3 peduncle medial mar- gin with | small seta; inner rami of uropods 1-2 extending 50 percent along outer; setae of rami on uropods 1-3 = 1 plumose seta on medial margin of inner ramus, each out- er ramus with 2 apical setae. Telson about 1.5 times as wide as long, weakly alate laterally, cleft about 75 percent ofits length, each apex with one long plume, each side with 2 small penicillate setules. Glands discernible in coxae 1—3, pedun- cles of uropods 1-3, and telson. Male “‘b,” 2.78 mm: right mandible with 5 rakers, left with 4; epimeron 2 with an additional long facial seta at posteroventral corner; uropod 2 peduncle with 1 lateral seta; segmental formulas of pleopods 1-3 = 7-4, 7-4, 7-5, basal setal formulas = 8-1- 2-1, 5-1-1-1, 3-1-9-1. Relationship. —Tonocote resembles Prantinus in the following: shapes and se- tosity of coxae 1—4; similar molars; maxil- liped inner plate shape and spination, outer plate shape, dactyl and palp 3 shape; and pleopods. Tonocote differs from Prantinus in hav- ing: less reduced coxa 5; stouter antenna 1; larger article 4 of antenna 2; maxilla | inner plate more slender, outer plate with 8 spines (versus 11); maxilla 2 inner plate with many fewer medial setae; maxilliped outer plate spines fewer; gnathopods 1—2 lacking short penicillate spines on article 6, with posterior brushes on articles 2 and 3, more poorly developed on right; pereopods 3—4 lacking posterior spines on article 5, with sparser anterior setation on articles 4—5; pereopods 6-7 article 5 poorly spinose laterally; uro- pods 1-2 inner ramus short, peduncle poor- ly setose, outer ramus with 2 apical setae (versus 1); uropod 3 poorly setose; telson broader; epimeron 2 poorly developed in size and setation. Tonocote differs from Bumeralius in the following: urothoid form of antenna 1; no bifid setae on antenna 2 article 4; weak to absent right lacinia mobilis; non-triturative molar with marginal cusps; maxilla 1 outer plate with eight spines (versus 11), inner plate lacking basomedial setae; maxilla 2 inner plate with only one seta in facial row (versus many); maxilliped inner plate trun- cated, with thick blunt spines, few setae, outer plate poorly armed medially, arma- ments stout, dactyl with apical setae only; coxae 1-7 sparsely setose, coxa 2 lacking large posteroventral lobe, coxa 7 lacking an- gles (versus angular); pereopods 6—7 poorly spinose on faces of articles 5 and 6, pereo- pods 3-4 lacking marginal posterior spines on article 5, pereopod 3 article 5 poorly se- tose anteriorly; gnathopods 1-2 poorly se- tose, palm well serrate, dactyl heavily armed; inner rami of pleopods slightly shorter than in Bumeralius; uropods 1-2 outer rami lacking basomedial setae, peduncles very poorly setose, uropod 2 outer ramus lacking lateral setae, uropod 3 and telson poorly setose in adult; mandibular palp not as cla- vate and spinose. Tonocote differs from other zobrachoids in having epimeron 3 dominant in size (ver- sus epimeron 2) although the original dif- ference cited by Barnard and Drummond (1983) refers only to dominance in setation. Tonocote barely dominates in setation as epimeron 2 has only one to two long and one short setae compared to one short seta on epimeron 3. Other items of “‘variables”’ show Tonocote to be aberrant in the absence of right lacinia mobilis and lacking posterior spines on article 4 of pereopods 3-4 (but apical [?] spines remain). There is not a great deal of difference between Zobrachoidae and Urohaustoriidae except the latter have gnathopod | simple, loss of integrity in epi- meron 1, large differential in size of epi- meron 2 (tiny) and epimeron 3 (large and extended posteroventrally), small number of setae on palp article 1 of maxilla 1, re- duction in extension and presence of cusps of mandibular molar and usually a reduc- 233 VOLUME 99, NUMBER 2 oo y mod taht? aXe | 99 of 9, ; Caen aie 00D 6 9 = Eee By $e i Fig. 4. Tonocote magellani, holotype male ‘“‘a” 2.87 mm. 234 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Tonocote magellani, holotype male “a” 2.87 mm. VOLUME 99, NUMBER 2 235 Fig. 6. tion in article 2 of uropod 3 outer ramus and weaker rostrum. Prantinus of Zobra- choidae intergrades in the condition of epi- meron | and the outer ramus of uropod 3. Tonocote has antenna 2 article 4 expanded; pereopods 3-4 lacking posterior spines (only terminal spines on article 5); and epimeron 2 much smaller than epimeron 3. Illustrations.—Palp of mandible proba- Tonocote magellani, holotype male “‘a”” 2.87 mm. bly more clavate than shown owing to pres- ervational defects. Holotype. —USNM No. “a,” 2.87 mm (illustrated). Type locality. —Eastern Straits of Magel- lan, 52°29.9’S, 69°05.9’W, 11-12 m, 9 Apr 1976, coll. Dr. Victor A. Gallardo. Voucher material.—Type locality: male “b,”? 2.78 mm. 195148, male 236 Etymology. —Named for the type locali- ty. Distribution. —Straits of Magellan, 11- 2 To Acknowledgments We thank Dr. Victor A. Gallardo of Uni- versidad de Concepcion, Chile, for making this material available; Linda B. Lutz of Mobile, Alabama for inking our drawings; and Patricia B. Crowe for laboratory assis- tance. Literature Cited Barnard, J. L., and J. Clark. 1982a. Puelche oren- Sanzi, new genus, new species, a phoxocepha- lopsid amphipod from the shores of Argentina (Crustacea, Amphipoda, Phoxocephalopsi- dae). —Journal of Crustacean Biology 2:261-272. , and 1982b. Huarpe escofeti new ge- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nus, new species, a burrowing marine amphipod from Argentina (Crustacea, Amphipoda, Uro- haustoriidae).—Journal of Crustacean Biology 2:281-295. , and 1984. Redescription of Phoxo- cephalopsis zimmeri with a new species, and establishment of the family Phoxocephalopsi- dae (Crustacea, Amphipoda) from Magellanic South America. — Journal of Crustacean Biology 4:85-105. ,and M. M. Drummond. 1978. Gammaridean Amphipoda of Australia, Part III: The Phoxo- cephalidae.— Smithsonian Contributions to Zo- ology 245:1-551. , and 1982. Gammaridean Amphip- oda of Australia, Part V: Superfamily Hausto- rioidea.—Smithsonian Contributions to Zool- ogy 360:1-148. Department of Invertebrate Zoology, NHB-163, Smithsonian Institution, Wash- ington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 237-247 PARALEBBEUS ZOTHECULATUS, N. GEN., N. SP., A NEW HIPPOLYTID SHRIMP FROM THE AUSTRALIAN NORTHWEST SHELF A. J. Bruce and F. A. Chace, Jr. Abstract.—A new species of hippolytid shrimp found in association with hexactinellid sponges on the Australian Northwest Shelf at depths of 450-500 m is described, and a new genus designated for its accommodation. Paralebbeus zotheculatus, n. gen., n. sp., represents the first record of an association between a caridean shrimp and a hexactinellid sponge. Paralebbeus is most closely related to Lebbeus White, 1847, and is distinguished by the reduced, unarmed rostrum, and absence of lateral spines on the meri of the ambulatory pereiopods. A survey of the Australian Northwest Shelf area, off Port Hedland, Western Aus- tralia, carried out by the F.R.V. Soela of the Fisheries Laboratory, Commonwealth Sci- entific and Industrial Research Organiza- tions, has produced, from this little-studied region, numerous examples of crustacean taxa that are new to science or new to the Australian fauna. Among the caridean ma- terial collected were several examples of a commensal hippolytid shrimp. This could be referred to none of the described genera, and a new genus is proposed for its accom- modation. CL. refers to the postorbital carapace length. NIM refers to the Northern Terri- tory Museum, Darwin, Australia. Paralebbeus, new genus Diagnosis. —Carapace smooth; rostrum feebly developed, acute, edentate; supraor- bital, antennal, and pterygostomial spines present, branchiostegal and hepatic spines absent. Abdomen smooth, segments dor- sally rounded, sixth with posteroventral an- gle without moveable plate; first 3 pleura rounded, fourth convexly produced, fifth acutely produced; telson with 4-6 pairs of dorsal spines, about 6 pairs of posterior spines. Eyes pigmented. Antennule without statocyst, stylocerite large, peduncle with- out mobile distal plate; upper flagellum uni- ramous. Mandible with 2-segmented palp, molar and incisor processes present. First and second maxillipeds with exopods; epi- pod of second maxilliped with podobranch; third maxilliped without exopod, with epi- pod. All pereiopods without arthrobranchs; first and second pereiopods chelate, first pair with robust, similar, subequal chelae, sec- ond with chela small, carpus 7-segmented; ambulatory legs with dactyl biunguiculate, merus unarmed, first 3 pereiopods with epi- pod, all 5 pairs with mastigobranchs. Uro- pods normal. Type species. —Paralebbeus zotheculatus, Nn. sp. Etymology. —From para (Greek) beside, and Lebbeus, a hippolytid generic name cit- ed by White, 1847, from a manuscript by Leach. The etymology of the name Lebbeus is unknown. The gender is masculine. Systematic position. — Paralebbeus is most closely related to the genus Lebbeus White, 1847, and shares with it the following major characteristics: 1. Supraorbital and antennal spines present on carapace, branchiostegal spines ab- sent; pterygostomial spines generally present, minute. 238 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Paralebbeus zotheculatus in host sponge. 2. Abdominal segments smooth, dorsally rounded. 3. Posteroventral angle of sixth abdominal segment without moveable plate. 4. Mandible with incisor process and 2-seg- mented palp. 5. Third maxilliped without exopod, with epipod. 6. Arthrobranchs absent. One of the features that distinguish Para- lebbeus from Lebbeus is the markedly re- duced and toothless rostrum in Paraleb- beus; most species of Lebbeus show a well developed, strongly toothed rostrum, often continuous posteriorly with a very marked, strongly dentate postrostral carina. Also, Paralebbeus generally has only a minute pterygostomial spine, which is well devel- oped in all species of Lebbeus. Lebbeus has the meri of the ambulatory pereiopods strongly armed with spines laterally, which are completely absent in Paralebbeus. The commensal association of Paralebbeus with hexactinellid sponges is probably also a characteristic feature; some species of Leb- beus are known to be commensals but they are apparently associated with coelenter- ates, particularly actiniarians. Paralebbeus zotheculatus, n. sp. Figs. 1-6 Material examined. —(i) 1 2, F.R.V. Soe- la, cruise 0184, sta NSW/57, 17°30.1'S, 118°28.9’E, 505-506 m, 3 Feb 1984, coll. A. J. Bruce. Holotype, NIM Cr. 00574A (i1) 1 ovig. 2, collected with above. Paratype, NTM Cr. 00574B. (iii) 1 9, F.R.V. Soela cruise 0184, sta NWS/68, 16°14.0’S, 120°20.4’E, 456-452 m, 5 Feb 1984, coll. A. J. Bruce, NIM Cr. 00577 (Gr )eiee@): F.R.V. Soela cruise 0784, sta NWS/66, 16°45.3’S, 119°46.4’E, 502-504 m, 5 Feb 1984, coll. A. J. Bruce, NIM Cr. 00575. All specimens collected by prawn trawl. Description. —Adult female paratype. Body of stout, subcylindrical form, gener- ally glabrous (Fig. 2). Carapace with stout, acute, straight, lat- 239 VOLUME 99, NUMBER 2 ‘(€PLS000 “IO WLN) odAiered 9[BUay SNOIOBIAO ‘snynjn2aYj0Z SnaqqaviDg “T ‘BLy 240 erally compressed, toothless rostrum reach- ing slightly beyond eyes; smooth, with acute supraorbital and antennal spines, inferior orbital angle produced, subacute, antero- lateral angle rounded, not produced, with minute pterygostomial spine present on right, absent on left side of carapace (Fig. 3A). Abdomen with segments dorsally round- ed, not carinate or produced; pleura of first 3 segments broadly rounded, fourth con- vexly produced, fifth acutely produced. Sixth segment about 1.5 times length of fifth, 1.5 times longer than deep, posterolateral angle acutely produced, posteroventral angle bluntly produced. Telson (Fig. 3F) about 1.7 times length of sixth abdominal segment, about 2.7 times longer than anterior width, sides subparallel anteriorly, convergent pos- teriorly to broadly rounded posterior mar- gin, equal to about 0.5 of anterior width; with 5—6 lateral dorsal spines and 13 pos- terior spines, sublateral spines longest, about 1.7 times length of lateral spines and 1.3 times length of submedian spines (Fig. 6L). Eyes prominent, cornea hemispherical, well pigmented, without accessory pigment spot; stalk slightly flattened, central length subequal to corneal diameter (Fig. 3D). Antennular peduncle (Fig. 3B) with large, broad acute stylocerite reaching to base of intermediate segment, proximal segment about 2.2 times longer than central width, medial border setose, distolateral angle tri- dentate, with small distal ventromedial tooth; statocyst obsolete; intermediate seg- ment 1.5 times longer than distal width, ta- pered proximally, with acute distolateral tooth; distal segment shorter than width, half length of intermediate segment, with upper and lower distolateral teeth. Upper flagellum uniramous, short, proximal 18 segments stout with about 10 groups of es- thetascs, 8 distal segments slender; lower flagellum short and slender, with 28 seg- ments. Antenna (Fig. 3C) with basicerite bearing PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON small distolateral tooth, carpocerite about 3.0 times longer than wide, exceeding mid- dle of scaphocerite, flagellum slender, about 3.5 times postorbital carapace length; scaphocerite extending well beyond anten- nular peduncle, lamella about 2.7 times longer than broad, distal margin bluntly an- gular, exceeding tip of distolateral tooth, lat- eral margin straight. Mandible (Fig. 4A) with corpus stout, with feebly setose 2-segmented palp laterally; left molar process (Fig. 6A, B) robust, obliquely truncate distally, well developed marginal setose fringe dorsally and row of small teeth ventrally; incisor process (Fig. 6C) feeble, with 4 small, irregular acute teeth distally. Maxillula (Fig. 4B) with well developed, fee- bly bilobed palp (Fig. 6D), upper lobe with 2 slender simple setae, lower lobe with 1 stouter setulose seta; upper lacinia broad, with about 14 short stout simple spines dis- tally, upper and lower borders sparsely se- tose; lower lacinia slender, setose, with few slender spines distally (Fig. 6E). Maxilla (Fig. 4C) with well developed, distally narrowed palp, lateral border with short plumose se- tae, 3 longer simple setae distally; basal en- dite large, deeply bilobed, medially setose on both lobes, coxal endite small, simple, with sparser, longer setae medially, sca- phognathite about 3.5 times longer than central width, posterior lobe short and rounded, anterior lobe distally narrowed. First maxilliped (Fig. 4D) with 2-segmented palp, distal segment 2.0 times longer than wide and 2.0 times length of proximal, both sparsely setose medially; basal endite short and broad, medially setose, coxal endite fee- bly convex medially, sparsely setose; exo- pod with flagellum normally developed, nu- merous plumose setae distally and small caridean lobe proximally; epipod large, fee- bly bilobed, with triangular lobes. Second maxilliped endopod (Fig. 4E) with distal segment narrow, 3.0 times longer than wide, with numerous long finely setulose setae medially, propodal segment distomedially VOLUME 99, NUMBER 2 E F 241 Fig. 3. Paralebbeus zotheculatus: A, Anterior carapace and rostrum, lateral aspect; B, Antennule; C, Antenna; D, Eye; E, Uropod; F, Telson. A, D, Female holotype; B, C, E, F, Female paratype. angulate with medial margin setose; carpus and merus normal; ischiomerus medially excavate; coxa not medially produced, non- setose; with large oval epipod bearing small podobranch with 5 pairs of lamellae proxi- mally; exopod damaged on left, normal on right. Third maxilliped (Fig. 4F) elongate and slender, exceeding antennular peduncle by length of distal segment, reaching ante- riorly about as far as extremity of second pereiopod; terminal segment about 8.0 times longer than proximal width, subcylindrical and tapering distally, laterally setose, with numerous short robust cornified spines (Fig. 6F); penultimate segment short, twice as long as wide and 0.3 times length of antepenul- timate segment, medially setose; ischio- merus and basis completely fused, com- bined segment about 5.0 times longer than wide, slightly narrowed proximally; coxa not 242 medially produced, without oval lateral plate, small hooked epipod present. Branchiae as follows: Maxilli- peds Pereiopods | es es a ee a alas ah Plemmmbmimnedns = = = fe == 4b se GE Arthrobranchs — — — — — — — — Podobranchs st AS ee Miastigobranchsy) =) =" ate at tee a Epipods +++ 4+4+4+-- Exopods un ees) a ee Sas Thoracic sternites narrow and armed with slender acute submedian spines on sixth to eighth thoracic segments. First pereiopods (Fig. 5A) robust, short, subequal and similar, exceeding carpocerite by chela and carpus. Chela (Fig. 5B) robust with palm subcylindrical, smooth, about 2.4 times longer than wide, slightly tapered dis- tally, with grooming setae proximally, dac- tyl (Fig. 6G) strongly curved, acute, 3.2 times longer than deep, about 0.5 of palm length, with 2 strongly cornified teeth distally, me- dial and lateral cutting edge entire; fixed fin- ger (Fig. 6H) similar, with 1 cornified distal tooth; carpus short and stout, about as wide as long, subcylindrical, distal end feebly ex- cavate, unarmed but provided with groom- ing setae; merus about 2.4 times length of carpus and 2.8 times longer than cen- tral width, subcylindrical and unarmed, obliquely articulated with ischium; ischium about 2.0 times longer than wide and twice length of basis, with short longitudinal row of 5 spines proximally; coxa robust, without processes but with hooked epipod. Second pereiopods (Fig. 5C) slender, sub- equal and similar, reaching to tip of third maxilliped. Chela (Fig. SE) small with palm subcylindrical, about 2.2 times longer than deep, dactyl 0.65 of palm length, acute, fee- bly curved, with unarmed cutting edge, 2 strongly compressed cornified teeth distally, fixed finger similar, with single distal tooth only; carpus (Fig. 5D) slender, 3.5 times length of chela, 7-segmented, with segments PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON in following ratios from proximal end 2.0: 1.0:3.3:1.8:1.0:1.0:2.0, merus undivided, about 0.6 of carpus length, 7.0 times longer than wide, uniform and unarmed; ischium subequal to merus, about 7.6 times longer than deep, ventral border sparsely setose, with 3 conspicuous spines proximally; basis short, without special features; coxa robust, with hooked epipod and small setose pro- cess ventrally. Ambulatory pereiopods moderately slen- der. Third pereiopod (Fig. 5F) extending to about 0.6 of terminal segment of third max- illiped and exceeding carpocerite by propod and dactyl; dactyl (Fig. 61) with corpus com- pressed, about 1.8 times longer than deep, about 0.15 of propod length, unguis distinct from corpus, about 3.0 times longer than wide, ventral border of corpus with 6 spines, distal spine shorter but more robust than unguis, other spines decreasing in size proximally; propod (Fig. 5G) about 6.5 times length of dactyl, 8.0 times longer than wide, with 2 rows of about 12 spines ventrally and tufts of setae dorsally; carpus barely 0.5 of propod length, about 6.3 times longer than wide, unarmed; merus about 1.2 times pro- pod length, 6.3 times longer than deep, un- segmented, without lateral spines or disto- ventral tooth; ischium about 0.45 of merus length; basis and coxa without special fea- tures, latter with hooked epipod (Fig. 6J). Fourth pereiopod similar to third. Fifth pe- reiopod (Fig. 5H) similar, more slender than third, dactyl (Fig. 6K) with 7 ventral spines, propod 1.1 times length of third and 10.5 times longer than deep, with fewer ventral spines, transverse rows of short, serrate se- tae distolaterally. Pleopods (Fig. 5I, J) normally developed, biramous, with protopodite broadly ex- panded laterally on second to fourth ab- dominal segments, endopods of second to fifth segments with appendix interna. Uropods (Fig. 3E) with protopodite acutely angulate posterolaterally; exopod 2.5 times longer than broad, lateral margin fee- bly convex, with small acute distal tooth VOLUME 99, NUMBER 2 243 1.0 mm SS WW TN\ N' Fig. 4. Paralebbeus zotheculatus, ovigerous female paratype: A, Left mandible; B, Maxillula; C, Maxilla; D, First maxilliped; E, Second maxilliped; F, Third maxilliped. accompanied medially by larger mobile Adult female holotype.—Generally very spine; endopod subequal to exopod, about similar to described paratype, smaller and 3.1 times longer than broad. less robust. Rostrum slightly longer rela- Ova few and large, length about 1.5mm. ___ tively and more acute, distally slightly up- 244 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Paralebbeus zotheculatus: A, First pereiopod; B, Same, chela; C, Second pereiopod; D, Same, carpus and chela; E, Same, chela; F, Third pereiopod; G, Same, propod and dactyl; H, Fifth pereiopod; I, First pleopod; J, Second pereiopod. A-H, Female paratype; I, J, Female holotype. VOLUME 99, NUMBER 2 eee 245 0.5 mm CDJL Fig. 6. Paralebbeus zotheculatus, female paratype: A, Molar process of mandible, anterior aspect; B, Same, posterior aspect; C, Incisor process of mandible; D, Palp of maxillula; E, Lower lacinia of maxillula, distal spine; F, Third maxilliped, distal end of terminal segment; G, First pereiopod, dactyl; H, Same, fixed finger; I, Third pereiopod, dactyl and distal end of propod; J, Same, epipod; K, Fifth pereiopod, dactyl and distal end of propod; L, Telson, posterior margin. turned. Small pterygostomial spine present on each side of carapace. Additional paratypes.—In close agree- ment with holotype. Specimen from NWS/ 68 lacking first and fifth pereiopods and one of fourth pereiopods. Pterygostomial spines absent from both sides. Telson with 4 dorsal spines on one side, 5 on other. Specimen from NWS/66 complete and also with 4 dorsal spines on one side of telson and 5 on other. Pterygostomial spines present on both sides of carapace. Coloration.—Body mainly transparent, abdomen whitish speckled with small red chromatophores increasing in density pos- teriorly, caudal fan reddish; antennae whit- ish; third maxilliped deep red; fingers of first pereiopod reddish, third to fifth pereiopods red, especially distally. Cornea black. Intra- thoracic organs orange. Ova turquoise. Host. — All specimens collected from small chambers in hexactinellid sponges, proba- bly of the genus Euplectella. Habitat.—Bottom temperatures 7.88°— 8.70°C. Parasites. —One specimen with pair of 246 bopyrid isopods in right branchial chamber. These have been identified as Bopyroides lamellatus (Kr@yer), a species not previ- ously recorded from Southern Hemisphere. Etymology.—The specific epithet is de- rived from ‘“zothecula’’ (Latin), a small chamber. Discussion The specimens of Paralebbeus zothecu- latus described above were all obtained from inside trawl-caught hexactinellid sponges, in small closed chambers only found on breaking up the sponges, leaving their as- sociation with the sponge beyond any doubt (Fig. 1). Most of the sponges collected were damaged and incomplete, which probably accounts for only single specimens of shrimps being found; it seems probable that they were originally in heterosexual pairs. The discovery of Paralebbeus zothecu- latus is of particular interest as it represents the first established example of an associ- ation between a caridean shrimp and a hex- actinellid sponge. This niche has been large- ly taken over by stenopodid shrimps, such as Spongicola, Spongiocaris, and Spongi- coloides (Saint Laurent and Cleva 1981). The genus Lebbeus White contains about 25 species (Wicksten and Méndez 1982, Butler 1980) which occur predominantly in northern seas, although one species, L. in- dicus, has been described from Indonesian waters (Holthuis 1947). Most of these have been trawl-caught specimens and no details of their associations, if any, have been pre- served. One of the boreal species found in shallow waters has been observed to live in association with sea anemones (Butler 1980, L. grandimanus [Brazhnikov] on Cribri- nopsis and Tealia spp.) but none have been reported in association with hexactinellid or any other sponges. The only hippolytid shrimp so far recorded as an associate of Porifera is Gelastocaris paronae Nobili, which lives on the external surface of shal- low-water Indo-West Pacific sponges. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Three specimens of a hippolytid shrimp that probably belongs to this genus and pos- sibly to the same species were collected dur- ing the Albatross Philippine Expedition, 1907-1910, but, as each specimen differs in some respects from the above description, they are not considered to be part of the type series of Paralebbeus zotheculatus. A female (CL 9.7 mm) from Verde Island Pas- sage, Philippines, agrees most closely with the Australian series but it has a more slen- der rostrum, the anterior margin of the car- apace not deeply recessed below the anten- nal spine, the scaphocerite with the lateral margin slightly concave rather than faintly convex at midlength, and the chela of the first pereiopod with the distal corneous teeth on the dactyl subequal rather than stag- gered. A male (CL 6.9 mm) from west of Halmahera, Indonesia, has a small supraor- bital spine on the left side and none on the right, no pterygostomial denticle on either side, the telson with only two and three left and right lateral dorsal spines, and the cor- neous subdistal spines on the third maxil- liped more numerous and extending some- what farther proximally on the terminal segment. A female (CL 11.7 mm) from southern Celebes, Indonesia, resembles the male in the spination on the third maxil- liped but it has no supraorbital spine on either side, a vestige of a pterygostomian denticle on the left side of the carapace but none on the right, the telson with two lateral dorsal spines on the left side and one on the right, and the right scaphocerite shorter than the left. These specimens will be discussed in greater detail in the report now in prep- aration by the junior author on the alpheoid families of the A/batross Philippine Expe- dition. Acknowledgments We are most grateful to Dr. T. Davis and Dr. T. Ward of the Fisheries Laboratory, C.S.I.R.O., Cronulla, for the opportunities and facilities provided aboard the F.R.V. VOLUME 99, NUMBER 2 Soela, and to Dr. J. C. Markham, for the bopyrid identification. Literature Cited Butler, T. H. 1980. Shrimps of the Pacific coast of Canada.—Canadian Bulletin of Fisheries and Aquatic Sciences 202, 280 pp. Holthuis, L. B. 1947. The Hippolytidae and Rhyn- chocinetidae collected by the Siboga and Snel- lius expeditions with remarks on other species. Part IX in The Decapoda of the Siboga Expe- dition. —Siboga-Expeditie 39a®, 100 pp. Saint Laurent, M. de, and R. Cleva. 1981. Crustacés Décapodes: Stenopodidea. Jn Résultats des 247 campagnes MUSORSTOM. I. Philippines (18- 28 Mars 1976) 1(7).— Collection Mémoires OR- STOM 91:151-188. Wicksten, M. K., and M. Méndez G. 1982. New records and new species of the genus Lebbeus (Caridea: Hippolytidae) in the eastern Pacific Ocean.—Bulletin of the Southern California Academy of Sciences 81:106-120. (AJB) Division of Natural Sciences, Northern Territory Museum, P.O. Box 4646, Darwin, Australia, 5794; (FAC) De- partment of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 248-256 RHOMBOPSAMMIA, A NEW GENUS OF THE FAMILY MICRABACIIDAE (COELENTERATA: SCLERACTINIA) Joan Murrell Owens Abstract. —Rhombopsammia, a new genus of deep-water micrabaciid corals, and two new species, R. squiresi and R. niphada, are described and figured. Rhombopsammia bears a marked superficial resemblance to Letepsammia, but examination of the corallum structure, microstructure, and microarchitecture of the new genus through thin-section and light microscope study reveals that it differs from Letepsammia in having essentially imperforate septa, prominent vepreculae which alternate in position on either side of the septum, ridge-like tracings of trabeculae on septal flanks and dentate columella. Its known geo- graphic distribution (China and Philippine seas) is more restricted than that of Letepsammia, but its bathymetric range (68—930 m) is only slightly less. Al- lusions by Squires (1967 and ca. 1967) to an undescribed species of micrabaciid coral from the Pliocene of Italy suggest that this species may belong to the new genus. If true, Rhombopsammia will conform with the evolutionary trend of the family towards increasing depth of occurrence with progressive geologic age. The Micrabaciidae is a small family of solitary, ahermatypic corals known from the shallow waters of Cretaceous continental shelves and inland seas to the deep waters of today’s Indo-Pacific. When Vaughan (1905) defined the family, he included in it Micrabacia Milne-Edwards and Haime, Di- afungia Duncan, Microsmilia Koby, Po- doseris Duncan, and Antilloseris Vaughan, largely on the basis of their solid septa and perforate walls. Later, Wells (1933) emend- ed the family to emphasize the structural significance of an alternation of septa and costae. He thus excluded from it all but /7- crabacia and Diafungia, and included in it Stephanophyllia Michelin and Leptopenus Moseley. In revising the Scleractinia, Vaughan and Wells (1943) determined that Micrabacia and Diafungia were synony- mous, and defined the Micrabaciidae as consisting of three genera: Micrabacia, Stephanophyllia, and Leptopenus. Most current workers accept this definition of the family. For many years most of the species of these genera were classified primarily on the basis of external characters observable in whole-mount study; consequently, many species were grouped together more or less according to superficial resemblances and broadly defined generic characteristics that sometimes masked structural and micro- structural differences that are significant enough to warrant generic separation. This was particularly true of many of the species comprising Stephanophyllia. On re-exami- nation of this genus, Yabe and Eguchi (1932) detected subtle but fundamental differences in morphology and component elements among the species which led them to erect four subgenera of Stephanophyllia, one of which was Letepsammia, a group of micra- baciid corals distinguishable by their highly perforated septa. Wells (1956) considered VOLUME 99, NUMBER 2 Letepsammia to be synonymous with Stephanophyllia, but more recent workers, particularly Squires (1967 and ca. 1967) and Owens (1984), have deemed septal perfo- rations to be a valid generic characteristic and have therefore referred to Letepsammia as a discrete genus of micrabaciid corals. In examining the Albatross collection of deep-sea corals, Squires (ca. 1967) separat- ed from those sorted by National Museum workers as Stephanophyllia a group which differed from that genus in having essen- tially imperforate septa. Although he ob- viously felt this group deserved further study, he left it undescribed. This group was later included in a study of the structure, microstructure, and microarchitecture of the Micrabaciidae (Owens 1984). As a result of this latter study of the family both in whole mount and in thin section, a new genus, Rhombopsammia, is designated in this pa- per. Rhombopsammia consists at present of two species, R. squiresiand R. niphada, both Recent. Squires (1967 and ca. 1967) alluded to a fossil coral of the Pliocene of Italy that apparently belongs to this new taxon; how- ever, his fossil species could not be found or verified for this study. This paper, there- fore, describes only the two Recent species, but acknowledges the possible existence of a Pliocene member. In light of the evolu- tionary trend of the family (Steinmann 1908, Squires 1967, Owens 1984) towards in- creasing depth of occurrence with progres- sively younger geologic age, a Pliocene member of this genus with a depth range similar to the Recent species is highly prob- able. Order Scleractinia Bourne, 1900 Suborder Fungiida Duncan, 1881 Superfamily Fungioidea Vaughan and Wells, 1943 Family Micrabaciidae Vaughan, 1905 Rhombopsammia, new genus Diagnosis. —Corallum large, loosely built, 249 strongly convex orally and nearly flat to pa- tellate aborally, with narrow to wide mar- ginal shelf formed by non-elevated exten- sions of septa alternating with costae. Septa essentially imperforate, coarsely dentate, rising steeply proximally and distally, form- ing a crown above the basal wall. Tracings of trabeculae visible on septal flanks. Deltas broad, porous. Vepreculae numerous, alter- nating in position on either side of septa. Synapticulae scarce, mainly near base of septa. Calicular depression deep, elongate. Costae thin, finely serrate or complex with recumbent teeth, connected by closely spaced concentric rows of synapticulae. In- tercostal loculi broader than costae. Wall perforate, moderately to strongly laterally flattened. Columella porous, deeply set, elongate, and dentate on exposed upper sur- face. Corallum completely invested in soft tissue of the polyp. Occurrence. —(?)Pliocene, Italy; Recent, China and Philippine seas; 68-930 m. Type species.—Rhombopsammia squire- Si, Mew species. Etymology. —The generic name refers to the convexo-patellate shape of R. squiresi, and is derived from the Greek rhombos = spinning top + psammos = sand. Gender: feminine. Discussion. —Specimens of both Rhom- bopsammia and Letepsammia have gener- ally been sorted by museum workers as Stephanophyllia. The three do bear a resem- blance, though the former two are more alike than either is like the latter. The resem- blances are greatly diminished, however, when specimens of Rhombopsammia are closely examined by use of a light micro- scope and thin sections, or when septal per- forations are taken into account. Rhombo- psammia differs from Stephanophyliia in its imperforate septa, convexo-patellate coral- lum, marginal shelf, spongy columella, and large, loosely built corallum; it differs from Letepsammia in its imperforate septa, prominent vepreculae which alternate in 250 position on either side of the septum, ridge- like tracings of trabeculae on septal flanks, and dentate upper margin of the columella. In thin section, the trabecular structure in Rhombopsammia is seen to be more widely spaced than in Stephanophyllia, \ess sin- uous than in Letepsammia. Only two species of Rhombopsammia are described below. A third species, a fossil form from the Pliocene of Italy, was re- ported by Squires (1967 and ca. 1967), but was not described and could not be located for this study. Steinmann (1908), Squires (1967), and Owens (1984) have noted an evolutionary trend of the Micrabaciidae towards increas- ing depth of occurrence with progressively younger geologic age. Rhombopsammia, with its bathymetric range of 68—930 m and its probable age of Pliocene to Recent, fits well into this trend. However, its virtually imperforate septa present a departure from another apparent trend of the family to- wards increasing septal perforations with depth (Steinmann 1908, Owens 1984). Stephanophyllia is moderately perforated, whereas Letepsammia is highly perforated. Inasmuch as Rhombopsammia falls be- tween Stephanophyllia and Letepsammia in both age and depth of occurrence, one would anticipate its also falling between the two in septal perforations. In many respects, Rhombopsammia ap- pears to be a transitional genus between Stephanophyllia and Letepsammia, for its species have characters found in each genus, and though its bathymetric range is close to that of Letepsammia, its maximum depth of occurrence nonetheless falls more than 50 m shorter. However, the evolutionary placement of Rhombopsammia between two genera with well-developed septal perfora- tions is troublesome. The question arises as to whether Rhombopsammia is indeed part of a linear evolutionary progression toward adaptation to life at deeper and deeper depths and, possibly, also a transitional ge- nus between Stephanophyllia and Lete- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON psammia, or a divergent member of the family following a different but parallel evolutionary path. More study along the lines of alternative solutions to the problem of adapting to life in deep waters is needed before that question can be satisfactorily an- swered. Rhombopsammia squiresi, new species Figs. 1; 2A Description. —Corallum large, loosely built, and strongly convexo-patellate, with wide marginal shelf. Calicular depression deep and long, but narrow. Wall finely per- forate. Deltas broad, porous, with proximal edges of secondaries distinct. Diameter of specimens 30.3-—32.6 mm; height 13.3—13.7 mm; average H:D ratio 43:100. Ninety-six septa. Costae thin, finely granulated axially, coarsening distally; costae and synapticulae adorned with irregular, recumbent teeth, beginning about midway between axis and periphery. Intercostal loculi as broad or broader than costae axially, but obscured distally. Costae begin as 6 (first cycle) at apex of wide cone forming basal wall and bifurcate immediately (second cycle); all new costae immediately bifurcate again (third cycle); outer costae bifurcate before inner at about one-tenth and one-seventh, respec- tively, distance from center to periphery (fourth cycle); final bifurcations (fifth cycle) begin at one-third distance to periphery with first and fourth costae of each quartuplet in each system bifurcating first. Septa imperforate and thin, with inter- spaces as wide or wider than septa. Vepre- culae radially aligned and alternate in po- sition on either side of each septum, giving septal margins a fluted appearance. Den- tation slightly lobulate, becoming slightly lacerate at inner margins of all but prima- ries. All but highest cycle septa tall and ap- proximately same height; inner margins of primaries more rounded than those of the deltas. Primaries inserted straight and free VOLUME 99, NUMBER 2 251 Fig. 1. Rhombopsammia squiresi: A, Oral view of holotype (USNM 72797), Albatross 5423: 9°38'30’N, 121°11’E, 930 m; B, Side view of holotype; C, Aboral view of holotype; D, Thin section of septum of a paratype (USNM 72798), under reflected light, showing thin, wavy trabeculae with growth laminae. Holes in septum were caused by mechanical grinding to make thin section. Albatross 5427: 9°11'30’N, 118°37'08’E, 68 m. Scale bars = 2 mm. 252 and extend to columella. Secondaries straight and free until joined with others in deltas. Tertiaries divide just beyond deltas, and these septa successively divide sym- metrically, with final division occurring at inner edge of shelf. Slightly porous sheets mark points of division. Synapticulae scarce, mainly at base of septa, particularly distally. Columella porous, spongy, terminating at upper surface in mass of lacerate-like teeth. Trabeculae in slightly diffuse fan system, visible along septal flanks as faint tracings. Growth laminae prominent, expressed as undulating ridges along the sides of the sep- ta. Types. — Holotype: USNM 72797; Alba- tross Sta 5423 (9°38'30’N, 121°11’E; 930 m). Paratypes: USNM 72799, 72800, 72798: Albatross Sta 5513 (8°16'45”N, 124°02'48’E; 924 m), 5425 (9°37'45’N, 121°11’E; 906 m), and 5427 (9°11'30’N, 118°37'08”E; 68 m), respectively. Occurrence. —Recent, China Sea, Phil- ippine Sea; 68—930 m. Remarks.—The pronounced convexo- patellate corallum, high crown of septa, wide marginal shelf, and recumbent costal and basal synapticular teeth make R. squiresi a very distinctive species, quite unlike any other micrabaciid coral save R. niphada, with which it bears a slight structural but stronger microstructural resemblance. That it is a micrabaciid coral is obvious from its alternation of septa and costae. Its pattern of septal insertions (Fig. 2A) is very similar to that seen in other micrabaciids which possess true deltas, and typically has the quinaries proximal to the primaries merging with the quaternaries closer to the center than the inner quinaries. One or two random pores were observed in a few septa in each specimen, but as these pores lacked any regularity in frequency, po- sition, or even cycle of septa in which they occurred, the septa are considered essen- tially imperforate. In thin section, the arcuate manner in which the ends of the sclerodermites of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON neighboring trabeculae merge proved to contribute to the fluted appearance of the septa. The arcs described by the scleroder- mites persist not only in the plane of the septum, but also swing laterally to form cusps along the flanks. This arcuate pattern between trabeculae is found also in R. ni- phada but to a lesser degree, and explains why the septa in the latter species are less wavy. Etymology. —The species name is in hon- or of Donald F. Squires, whose recognition that this group of corals differed from the other species of Stephanophyllia inspired this study. Rhombopsammia niphada, new species Figs. 2B, 3 Description. —Corallum large, delicate, loosely built, with narrow to wide promi- nent shelf; strongly convex orally and slight- ly patellate to nearly flat basally, but with pronounced basal apex. Calicular depres- sion deep, long, and narrow. Deltas broad, porous, slightly spinose on inner margin. Diameter of specimens 21.5-36.0 mm; height 6.7—12.0 mm; H:D ratio 31:100. 144 septa. Costae smooth, very thin, finely serrate on outer surface. Intercostal loculi broader than costae, but interrupted by closely spaced synapticulae connecting costae to bases of septa. Basal synapticulae smooth, lacking serration. Costae begin as 6 (first cycle) at apex of protuberant basal tip and bifurcate immediately (second cycle); each costa immediately bifurcates again (third cycle); outer pairs bifurcate fractionally be- fore inner ones at about one-tenth distance from center to periphery (fourth cycle); bi- furcations of fifth cycle begin about one- sixth distance from center with outermost costae of each system dividing first, fol- lowed closely by next outer and then in- nermost, other inner costae do not divide in this cycle; sixth cycle, also incomplete, begins again with outermost and next outer VOLUME 99, NUMBER 2 253 Fig. 2. Diagrammatic representations of one system: A, Rhombopsammia squiresi, B, Rhombopsammia niphada. Heavy lines represent septa; thin lines, costae; dotted lines, non-elevated extensions of septa. dividing; followed symmetrically by 3 ad- jacent inner costae, those flanking each side not dividing. Final bifurcations occurring about one-third the distance to periphery. Septa imperforate and thin, with inter- spaces wider than septa. Vepreculae radially aligned and alternate in position on either side of a septum, but produce only slight fluting effect. Dentation lobulate-like on up- per and outer margins of septa, less so on inner margins of primaries. Synapticulae broad relative to width of septa and irreg- ularly spaced deep in calice. Primaries straight, free, extending from shelf to col- umella. Secondaries straight and free to del- tas. Tertiaries divide just beyond deltas, and quaternaries thus formed divide halfway between deltas and outer edge of calice, with outer quaternaries dividing slightly before inner ones. Quinaries proximal to the pri- maries and secondaries do not bifurcate un- til three-fourth distance from delta to shelf whereas of the two inner quinaries, the one closer to secondary does not bifurcate at all. At inner edge of shelf, beginning with no bifurcation of septa proximal to primaries, every other septum bifurcates for a final time, giving each system total of 24 septa. Coarsely porous sheets mark points of di- vision. Columella porous, spongy, elongate, but broader than in R. squiresi; ornamented at center with cluster of small, slightly spinose teeth. Trabeculae, often ramified near distal Margins, in moderately well-spaced fan sys- tem, and clearly visible as tracings along septal flanks. Ends of sclerodermites either joining together in arcuate pattern between trabeculae or diverging normally from axis to form vepreculae. Types. —Holotype: USNM 72802; Alba- tross Sta 4911 (31°38'30’N, 129°19’E; 716 m). Paratypes: USNM 72801, 72804, 72803, 72805; Albatross Sta 4911 (same as holo- type), 5283 (13°48'30’N, 120°28’40’E; 512 m), 5348 (10°57'45’N, 118°38'15”E; 686 m), and ND (no data), respectively. Occurrence. —Recent, China Sea and Philippine Sea; 512-716 m. Remarks. —Rhombopsammia niphada bears a close resemblance to species of Le- tepsammia, in that it shares with them a delicate, lacy corallum with a generally nar- row marginal shelf, narrow columella, and highly perforated wall. However, close ex- 254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Rhombopsammia niphada: A, Oral view of holotype (USNM 72802), Albatross 4911: 31°38'30’N, 129°19’E, 716 m; B, Aboral view of holotype; C, Side view of holotype; D, Thin section of septum of a paratype (USNM 72804), under reflected light, showing thin, wavy trabeculae with ends of sclerodermites merging in an arcuate pattern in the interarea. Note ramified trabecula at distal margin of septum in upper left. A/batross 5283: 13°48'30"N, 120°28'40’E, 512 m. Scale bars = 2 mm. VOLUME 99, NUMBER 2 amination of the structure, microstructure, and microarchitecture of R. niphada dis- tinctly separates it from Letepsammia and reveals its closer relationship with R. squire- Si. Both R. niphada and R. squiresi have es- sentially imperforate septa, numerous well- developed vepreculae, and strong fan-like trabeculae that impart ridge-like tracings on the sides of the septa and project along the outer margins as lobulate-like teeth. Also, in both the vepreculae alternate in position on either flank, giving the septa a fluted ap- pearance. Both have a very protuberant bas- al apex, and their costal-intercostal spacings are similar. As in all micrabaciid corals, R. niphada has an alternation of septa and costae. Its pattern of septal insertions (Fig. 3) is similar to that of other micrabaciids with true del- tas, and is virtually identical with that seen in R. squiresi for an equal number of cycles and septa. Rhombopsammia niphada differs most conspicuously from R. squiresi in its total number of septa (144) and its very slightly patellate to nearly flat base. It also lacks the recumbent basal teeth found in R. squiresi. Although the marginal shelf in R. niphada varies from narrow to wide, the shelf, even when as wide as in R. squiresi, is not as prominent due to the less steeply sloping distal edges of the septa. Etymology.—The species name is de- rived from the Greek niphados = snowflake, and refers to the intricate design of the oral surface of the corallum. Gender: feminine. Discussion.—Rhombopsammia niphada is distinguished by its 144 septa, which poses the most perplexing problem in its generic identity with R. squiresi, which has 96. In all other micrabaciid corals studied or de- scribed in the literature, the number of sep- ta, except in an occasional deviant individ- ual, has been so consistent within a genus as to suggest that it is a valid generic char- acteristic. But in Rhombopsammia a sufh- cient number of specimens in each species USS) was examined to indicate that the difference in number of septa between the two species is not due to random deviation. Within each species the number of septa is consistent. The structural and microstructural similar- ities between R. niphada and R. squiresi ar- gue convincingly for their generic identity in spite of the difference in septal number. Perhaps too much emphasis should not be placed on that character as a generic char- acteristic. Interestingly, 144 septa are also found in the lectotype of Letepsammia formosissi- ma, though all other specimens of that species studied had 120. Even if the lecto- type of L. formosissima should prove to be a deviant individual, the occurrence of this unusual number of septa in both it and all specimens of R. niphada may be more than coincidental. When the similarities of R. ni- phada with R. squiresi and with species of Letepsammia are thoroughly considered, the strong suggestion arises that R. niphada may be an intermediate between Rhombopsam- mia and Letepsammia. Acknowledgments I wish to thank Stephen D. Cairns of the Smithsonian Institution for making avail- able to me the specimens used in this study, and for his thoughtful criticism of this manuscript. A special debt of gratitude is owed to Donald A. Dean of the Smithson- ian Institution for his assistance in thin-sec- tioning these specimens, and Thayer McKell for typing this manuscript. I am also espe- cially indebted to my husband, Frank A. Owens, for photographing the specimens used in this paper. Finally, special acknowl- edgment is due to Donald F. Squires whose perceptive recognition of disparities within the micrabaciids collected by the Albatross Expedition inspired my study of this group of corals. Literature Cited Owens, J. M. 1984. Microstructural changes in the scleractinian families Micrabaciidae and Fun- 256 giidae and their taxonomic and ecologic impli- cations. Doctoral dissertation, The George Washington University, Washington, 242 pp., 14 pls., 15 text figs. Squires, D. F. 1967. The evolution of the deep-sea coral family Micrabaciidae. — Proceedings of the International Conference on Tropical Ocean- ography 5:502-510. . [ca. 1967]. Tertiary and extant scleractinian corals of the family Micrabaciidae. Unpub- lished partial manuscript on deposit at the Smithsonian Institution. Steinmann, G. 1908. Die geologischen Grundlagen der Abstammungslehre. Wilhelm Engelmann, Leipzig. 284 pp. Vaughan, T. W. 1905. A critical review of the liter- ature of the simple genera of the Madreporaria Fungida, with a tentative classification. — Pro- ceedings of the U.S. National Museum 28:37 1- 424. , and J. W. Wells. 1943. Revision of the sub- orders, families, and genera of the Scleracti- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nia.—Geological Society of America, Special Papers, number 44, 363 pp., 40 pls., 32 text figs. Wells, J. W. 1933. Corals of the Cretaceous of the Atlantic and Gulf coastal plains and western interior of the United States.—Bulletins of American Paleontology 18 (67), 206 pp., 16 pls., 4 text figs. . 1956. Scleractinia. Jn R. C. Moore, ed., Trea- tise on invertebrate paleontology. Part F. Coel- enterata: pp. 328-444, figs. 223-339. Geological Society of America, Lawrence: University of Kansas Press. Yabe, H., and M. Eguchi. 1932. Some Recent and fossil corals of the genus Stephanophyllia H. Michelin from Japan.—Tohoku Imperial Uni- versity Scientific Reports, series 2, Geology 15: 55-63, 2 pls., 3 text figs. Department of Geology and Geography, Howard University, Washington, D.C. 20059. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 257-260 DISCOVERY OF A NEW LIVING CERITHIOCLAVA SPECIES IN THE CARIBBEAN (MOLLUSCA: PROSOBRANCHIA: CERITHIIDAE) Richard S. Houbrick . Abstract. — Cerithioclava garciai occurs on the shallow-water banks of the Caribbean off Honduras and is the only living member of the Pliocene Ca- loosahatchian genus Cerithioclava Olsson and Harbison, 1953. The group was previously thought to be extinct and is here accorded full generic status. The species is the largest of all Caribbean cerithiids and closely resembles members of the Indo-Pacific genera Rhinoclavis and Pseudovertagus. Specimens ofa very large, distinctive cer- ithiid have recently been collected by fish- ermen in the western Caribbean on the shal- low-water, continental shelf off Honduras. These shells, sent to me for identification by Dr. E. Garcia, proved to represent an undescribed species in the genus Cerithio- clava Olsson and Harbison, 1953, a taxon previously thought to be extinct and limited to the Tertiary Caloosahatchian Province of Florida. The genus Cerithioclava includes two described and one undescribed fossil species. Although only three specimens of the new species were available for study, the dis- covery of a living Cerithioclava was deemed sufficient justification for immediate de- scription. A description of the new species and a discussion of the genus Cerithioclava follows. Genus Cerithioclava Olsson and Harbison, 1953 Type species, by original designation: Cer- ithium caloosaensis Dall, 1892. Remarks. — This taxon was regarded as a subgenus of Cerithium Bruguiére by Olsson and Harbison (1953) although they noted a close affinity to Rhinoclavis Swainson and indicated that it might be considered a di- vision of that genus rather than of Ceri- thium. In a previous monograph (Houbrick 1978:119), I considered Cerithioclava to be a subgenus of Pseudovertagus Vignal but pointed out that Cerithioclava lacks the dis- tinctive cancellate sculpture of the early whorls so distinctive of Pseudovertagus species and, in addition, does not have the central columellar fold characteristic of members of the genus Rhinoclavis. Ceri- thioclava species differ considerably from Caribbean Cerithium species by being much larger and in having a long narrow anal ca- nal that is an adpressed extension of the outer lip. In retrospect, I think Cerithiocla- va, comprising four species, differs suff- ciently from other cerithiid genera to be giv- en full generic status. It represents an independent cerithiid radiation, having its roots in the Caloosahatchian Province (Pe- tuch 1981) of the northwestern Caribbean and Florida, and has as its sole survivor the new species described herein. There are two described fossil species: Cerithioclava ca- loosaensis (Dall, 1887) and C. dalli (Olsson and Petit, 1964). Another fossil species from the Burmont Formation of the Miami area, is undescribed. Cerithioclava garciai, new species Fig. 1, Table 1 Description. —Shell large, up to 86.5 mm in length, robust, turreted, comprising 15- 258 18 slightly convex whorls sculptured with 0-9 axial plications per whorl and with nu- merous fine spiral and sinuous axial striae. Axial plicae strongly developed on earlier whorls, weaker on last 3. Each whorl having broad subsutural constriction that divides axial plicae into 2 rows, producing an upper subsutural spiral cord with smooth nodular elements, lower remainder of whorl having axial plicae and giving shell an overall wrin- kled appearance. Axial plicae absent on body whorl and sometimes very weak on last 3— 4 whorls. Fine axial and spiral striae coarser at base of each whorl and on entire body whorl, producing a microscopic beaded tex- ture. Large, prominent varix opposite outer lip of body whorl. Suture impressed, slightly wavy. Aperture (excluding long extension of anal canal) round-ovate, nearly one-fourth shell length. Anterior siphonal canal long, reflected upwards toward the left. Colu- mella concave, smooth, thick and with strong lip. Outer lip smooth, thick, and greatly extended and adpressed to body whorl, where it joins upper columellar lip to form long anal canal, latter almost closed. Tip of anal canal slightly detached from body whorl. Distal end of anal canal lying adja- cent to suture of body whorl. Protoconch unknown. Shell color white to light brown, with light tan spiral threads and spiral band of darker brown subsutural nodules. Aper- ture white. Operculum and radula un- known. Type locality.—Off Great Corn Island, Nicaragua; 24 meters. Holotype (Fig. 1, A).—USNM 849023; length 77.4 mm, width 24 mm. Paratypes (Fig. 1, B—C).—Two speci- mens, USNM 849024; lengths 84.7 mm and 66 mm. Etymology. —Named for Dr. Emilio Gar- cia of Lafayette, Louisiana, who sent the specimens to me. Remarks. —This large, distinctive species is unlike any other cerithiid in the Carib- bean or Florida. At first glance it resembles the large Rhinoclavis and Pseudovertagus PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON species found in the Indo-Pacific, but al- though it shares with them a long reflected anterior canal, it is very different in other characters. Aside from size, the chief dis- tinguishing character of Clavocerithium garciai is the long extension of the outer lip, adpressed onto the body whorl and forming a long, narrow anal canal. The three spec- imens upon which this description is based were obviously collected alive as may be seen from shell condition and traces of the animal’s mantle, dried in the aperture of the shells. Unfortunately, there were no re- mains of the radula or operculum. Although only three specimens are available for com- parison, the range of sculptural variation is considerable, one specimen lacking or hav- ing weaker axial plications on the last four whorls. Shell color is likewise variable, one specimen being highly pigmented, the oth- ers being whiter and having only weak color patterns. Clavocerithium garciai does not resemble either of the two described fossil species from Florida, but an undescribed fossil species from the Miami area, shown to me by Dr. E. Petuch, is undoubtedly closely related, ancestral, or even conspecific. However, Cerithioclava garciai does not appear to be a secondarily derived species, sensu Petuch (1981:1125), since it shows only slight di- vergence from the Florida fossil species. The Honduran shelf region is poorly known, but recently many unusual species have been collected there by shrimp and lobster boats based at Roatan Island, Hon- duras. Large carnivorous prosobranchs, such as volutes, miters, and fasciolariids have turned up in this locality and several new species have been described. Petuch (1981: 1110-1111) noted that this region con- tained many extant elements of the ““Neo- gene Caloosahatchian Molluscan Prov- ince,” and that a number of relict genera and species complexes have undergone sec- ondary speciation. The discovery of a living Cerithioclava species in this locality sup- ports Petuch’s (1981:1125, 1982:292) sug- VOLUME 99, NUMBER 2 Fig. 1. gestion that the region is a Caloosahatchian relict pocket. A map showing the extent of this relicit pocket region was presented by Petuch (1981:1122, fig. 21). Due to the limited collecting data, not much may be said about the ecology of this species. Live-collected specimens show cal- careous and fine algal growth and other foul- 259 A-C, Cerithioclava garciai, 24 meters off Great Corn Is., Nicaragua: A, Holotype (USNM 849023, length 77.4 mm); B, Large, darkly pigmented specimen with weak axial sculpture, paratype (USNM 849024, length 66 mm); C, Paratype (USNM 849024, length 84.7 mm); D, Cerithioclava caloosaensis (Dall, 1887), type species of the genus, from the Caloosahatchee of Florida, Tertiary (USNM 22383, length 60.3 mm). ing organisms on the entire dorsal surface of the shell while the ventral portion is smooth and clean, indicating that this species is a partial burrower on sandy or rubble bottoms. The long anterior and anal siphons allow the animal to maintain a clear flow of water through the mantle while the shell is partially buried. Breakage scars and the bro- 260 Table 1.—Shell measurements of Cerithioclava gar- ciai. (Total length and number of whorls estimated in shells with damaged tips. All measurements in mm.) Statistic (n = 3) x SD Var. Range Length 79.33 7.01 32.7 72.5-86.5 Width 24.33 1.53 1.56 23-26 Length body whorl 31.93 2.18 3.18 30-34.3 Aperture length 20.30 2.87 5.49 18.4-23.6 Aperture width 13.70 1.30 1S 9 = 1522 No. axial plicae 6.33 4.62 14.22 0-9 No. whorls WG.333) S93! 1.55 15-18 ken shell tips show previous unsuccessful crab attacks on all three specimens. It is remarkable that a species as large and distinctive as Cerithioclava garciai has re- mained unknown for so long in the Carib- bean. One would expect that a large ceri- thiid, in a family characterized by herbivorous species that usually occur in abundance, would be common in its habitat and taken more frequently than the rarer carnivorous volutes. I suspect that this species is common, but as cerithiids are not regarded as collector’s items they are prob- ably thrown back by fishermen. This may explain why only a few specimens exist, to date. It is possible that other Caloosahatch- ian cerithiid taxa, such as Ochetoclava Woodring, previously regarded as extinct in the Western Atlantic (Houbrick 1978:89), may be found here, as well as other unusual molluscan elements that are of little interest to shell collectors. The region needs a thor- ough scientific survey. Acknowledgments I thank Dr. Emilio Garcia for first calling my attention to this unusual species and for PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sending me the specimens described herein. Dr. David Reid, Smithsonian Postdoctoral Fellow, critically read the original draft of the manuscript and offered helpful sugges- tions. Photography was done by Mr. Victor Krantz, Smithsonian Photographic Ser- vices, National Museum of Natural Histo- ry. Literature Cited Dall, William H. 1892-1897. Contributions to the Tertiary fauna of Florida with especial reference to the Miocene-Silex beds of Tampa and the Pliocene beds of the Caloosahatchie River. Part II. Streptodont and other Gastropods, conclud- ed.— Transactions of the Wagner Free Institute of Science of Philadelphia 3(1—6):1—1654, pls. 13-22, 1 map. Houbrick, Richard S. 1978. The family Cerithiidae in the Indo-Pacific. Part 1: The genera Rhino- clavis, Pseudovertagus, and Clavocerithium. — Monographs of Marine Mollusca 1:1—130, 98 pls. Olsson, Axal A., and A. Harbison. 1953. Pliocene Mollusca of southern Florida.—Academy of Natural Sciences of Philadelphia, Monograph 8: 1-457, 65 pls. , and Richard E. Petit. 1964. Some Neogene mollusca from Florida and the Carolinas. — Bul- letins of American Paleontology 47(217):509- 574, pls. 77-83. Petuch, Edward. 1981. A volutid species radiation from northern Honduras with notes on the Hon- duran Caloosahatchian secondary relict pock- et.—Proceedings of the Biological Society of Washington 94:1110-1130. 1982. Geographical heterochrony: Contem- poraneous coexistence of Neogene and Recent molluscan faunas in the Americas.—Palaeo- geography, Palaeoclimatology, Palaeoecology 37: 277-312. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 261-265 NOTES ON THE SPECIES OF ALLOSQUILLA AND PLATYSQUILLOIDES (CRUSTACEA: STOMATOPODA) Carlo Froglia and Raymond B. Manning Abstract.—A second specimen of an Allosquilla from the Adriatic Sea is reported, and A. adriatica Stevcic, 1979, is shown to be asynonym of A. africana (Manning, 1970). Allosquilla is compared with the related Platysquilla, Mex- isquilla, and Platysquilloides, and the second species of Allosquilla, A. lillyae Manning, 1977, is transferred to Platysquilloides. The genus Allosquilla was erected by Manning (1977:64) for two species from off West Africa, A. africana (Manning, 1970) and A. Jillyae Manning, 1977. Allosquilla africana was known from the unique ho- lotype, a female 39 mm long, taken at a depth of 174-148 m off the Niger delta, and A. lillyae was known from fragments of two specimens taken in depths of 260—225 and 345 m in the Azores. Two years later a third species, A. adriatica Stevcic (1979:642), based on our then unpublished account (Manning and Froglia 1979:178), was de- scribed from a female, 67 mm long, taken in the Adriatic Sea in 130-150 m. Manning (1977:65) pointed out that the species then assigned to Allosquilla differed from other genera of lysiosquilloids in lack- ing the mandibular palp, having four rather than five epipods, and in the number of papillae on the antennal protopod. A/llo- squilla africana was characterized as having two papillae on the antennal protopod, one small one mesially and one larger one ven- trally, and A. /illyae was characterized as having one mesial and two ventral papillae. Allosquilla adriatica also was characterized as agreeing with A. /illyae and differing from A. africana in having one mesial and two ventral papillae (Manning and Froglia 1979: 179). Subsequently, Holthuis (1984:132) re- ported two specimens of A. /illyae from the Azores in 52 m; one of these, a female, 42 mm long, was the first intact specimen of the species to be studied. The discovery and description by Hol- thuis of a complete specimen of A. Jillyae and the capture of a second specimen of a species of Allosquilla in the central Adriatic Sea by one of us (C.F.) prompted us to reex- amine the species of Al/osquilla, for the sec- ond specimen differed from the holotype of A. adriatica in several features that initially suggested it might belong to a different species. The Status of Allosquilla adriatica The second specimen of A. adriatica from the Adriatic (Fig. 1), an adult female 70 mm long with ripe ovaries, was taken at a depth of 216-222 m during trawling investiga- tions in the western Pomo Pit (net shot 42°50’/N, 14°33’E, hauled 42°48’N, 14°47’), a few miles south of the type locality, on 7 March 1981. It is preserved in the reference collection of the Istituto di Ricerche sulla Pesca Marittima in Ancona. We are able to add some color notes made in the field from the living specimen, and some data on mor- phology to the original description. In life, the background color is olive brown, with the deep pink ovaries clearly visible through the background color. The rostral plate, ocular peduncles, and antennal peduncles are light brown, and the cornea is green. The raptorial claw is covered with 262 Fig. 1. view. pink chromatophores, with a band of orange chromatophores proximally and brown chromatophores distally on the merus, and an orange spot on the propodus at the ar- ticulation with the dactylus. The thoraco- pods, pleopods, and uropods are clear. The edge of the telson is marked by a brownish white line. Other measurements of this specimen, in mm, are as follows: carapace length 11.7; rostral plate length 3.3, width 3.2; cornea width 3.2; antennal scale length 3.9; telson length 6.1, width 10.4. There are only nine teeth on the dactylus of the claw (Fig. 1a); there are ten teeth on the claw of the ho- lotype. There are five movable teeth on the uropod, the proximalmost very small, tri- angular, the remainder larger, spatulate; the holotype had three or four movable teeth on the uropod. The rounded lobe on the inner distal margin of the uropodal endopod is ornamented with 10-11 setae. The main morphological difference be- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON <4 @) ¢C A fete" Allosquilla africana, 2, TL 70 mm: a, Claw; b, Sixth abdominal somite and telson; c, Telson, ventral tween this specimen and the holotype of A. adriatica is in the posterior armament of the telson. The second marginal projection denticle is enlarged, so that the telson has three rather than two pairs of major, fixed marginal teeth lateral to the movable sub- medians, each separated by an intervening denticle. Also, the median dorsal projection on the telson (Fig. 1b) is asymmetrical; it probably has been damaged. This specimen, like the holotype of A. adriatica (Fig. 2a, b), has one mesial and two ventral papillae on the antennal pro- topod (Fig. 2c, d). Reexamination of the holotype of A. africana reveals that it, too, has the same number of papillae (Fig. 2e, f). That was one of the more important char- acters used by Manning and Froglia (1979: 180-181) to distinguish these two species. Manning and Froglia also used the fol- lowing characters to separate the two species: 1. The submedian denticles are in two convex rows in A. africana, one transverse VOLUME 99, NUMBER 2 263 Fig. 2. Antennal protopod in dorsal (a, c, e, g) and lateral (b, d, f) views to show antennal papillae in: a, b, Alllosquilla adriatica, 2? holotype, TL 67 mm; c¢, d, A. africana, Pomo Pit, ¢, TL 70 mm; e, f, A. africana, 2 holotype, TL 39 mm; g, Platysquilloides enodis, ? holotype, TL 57.5 mm (note dorsal spine on antennal protopod). row in A. adriatica. In the second Adriatic specimen, these denticles are arranged in two convex rows (Fig. 1c), as in the holotype of A. africana. 2. Both species were interpreted as having four intermediate marginal denticles on each side of the midline of the telson, with the first and third bluntly triangular and larger than the spiniform second and fourth den- ticles in A. adriatica and the first and third blunter but not much larger in A. africana. The second Adriatic specimen indicates that the telson has four distinct marginal teeth separated by intervening denticles, with the mesial one or two intermediate teeth small- er than the lateral two; the smallest of these marginal teeth is a low lobe flanking the movable submedian tooth. In this feature the holotypes of both A. africana and A. adriatica differ from the second Adriatic specimen. 3. The anterior prominences of the telson are less prominent in A. adriatica than in A. africana. This may be a function of size, for the holotype of the latter species is much smaller than either of the two Adriatic spec- imens. 4. Differences were noted in the armature of the basal segments of the walking legs in the two species. A reexamination of this fea- ture in the types of the two species reveals that it is the same in all three specimens. We conclude that A. adriatica is a syn- onym of A. africana, and that the holotype of A. africana is much younger than either of the two specimens collected in the Ad- riatic. Allosquilla can be characterized as having 1 mesial and 2 ventral papillae on the an- tennal protopod and 4 pairs of fixed mar- ginal teeth on the telson, the inner one or two of which can be much smaller than the lateral two; the marginal teeth are separated by smaller intervening denticles. In addi- tion, Allosquilla lacks a distinct dorsal spine on the antennal protopod (Fig. 2a—f) and has 264 —— PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Eyes of: a, Allosquilla adriatica, 2 holotype, TL 67 mm; b, A. africana, 2 holotype, TL 39 mm; c, Platysquilla enodis, 2 holotype, TL 57.5 mm. the cornea distinctly bilobed (Fig. 3a, b). In other features Allosquilla closely resembles Platysquilloides. The Status of Allosquilla lillyae The account of a complete specimen of A. lillyae by Holthuis (1984:132) shows that it shares several features with Platysquil- loides enodis (see Manning and Camp 1981: 593, fig. 2), as follows: the cornea is ex- panded laterally (Fig. 3c) but not distinctly bilobed as in A. africana (Fig. 3a, b); the rostral plate is subrectangular rather than distinctly cordiform; there is a distinct dor- sal spine on the antennal protopod (Fig. 2g); the marginal armature of the telson com- prises, on each side of the midline, four fixed teeth separated by intervening denticles; and the inner two pairs of teeth are small and distinctly spatulate, whereas the outer two pairs are produced into large, slender, sharp teeth. In these features, A. /illyae closely resem- bles the west Atlantic Platysquilloides en- odis (Manning, 1962) and differs from A/- losquilla africana. We conclude that A. lillyae was incorrectly placed in Allosquilla, and we transfer it here to Platysquilloides, which until now was considered to be monotypic. Genera Related to Allosquilla and Platysquilloides Allosquilla and Platysquilloides share many characters with two other genera of the family Nannosquillidae, Platysquilla Manning, 1967, and Mexisquilla Manning and Camp, 1981 (see Manning and Camp 1981 for accounts of these other genera). They differ from Platysquilla in having only one mesial papilla on the antennal proto- pod, a much broader cornea, only four rath- er than five epipods, and a much smaller median projection on the telson; they also differ in lacking a pair of spines ventrally on the sixth abdominal somite. The eyes of Platysquilla resemble those of Mexisquilla and Platysquilloides in being somewhat broadened, not distinctly bilobed as in A/- losquilla. Allosquilla and Platysquilloides differ from Mexisquilla Manning and Camp, 1981, in several features, including the an- teriorly spined rostral plate, having four rather than three epipods, and in overall size; species of Mexisquilla are not known to exceed 20 mm in length, whereas species of Allosquilla are as large as 70 mm and species of Platysquilloides as large as 67 mm are known (Howells, Karp, and Langton 1980). Platysquilla differs from all of these gen- VOLUME 99, NUMBER 2 era in having four papillae on the antennal protopod and slender, non-spatulate mov- able spines on the outer margin of the uro- podal exopod, and Platysquilloides differs from all of these genera in having a distinct dorsal spine on the antennal protopod. Acknowledgments We thank Lilly K. Manning for preparing the illustrations. Literature Cited Holthuis, L. B. 1984. On some Crustacea Stomatop- oda collected by the CANCAP expeditions in the waters of N.W. Africa and the Azores.— Zoologische Mededelingen 58(8):131-139. Howells, R. G., C. Karp, and R. W. Langton. 1980. Occurrence ofa rare mantis shrimp, Platysquilla enodis (Manning, 1962), in the Middle Atlantic Bight region (Stomatopoda).—Crustaceana 38(1):101-104. 265 Manning, R. B. 1977. A monograph of the West Af- rican stomatopod Crustacea. — Atlantide Report 12:25-181. ,andD.K.Camp. 1981. Areview of the Platy- squilla complex (Crustacea, Stomatopoda, Nan- nosquillidae), with the designation of two new genera.— Proceedings of the Biological Society of Washington 94(2):591-597. , and C. Froglia. 1979. Description of a new Allosquilla with notes on other Adriatic stoma- topod Crustacea. — Quaderni del Laboratorio di Tecnologia della Pesca 2(4):177-190. [pub- lished November 1979] Stevcic, Z. 1979. Vabici (Stomatopoda) Jadrana.— Pomorski Zbornik 17:639-651. [published Sep- tember 1979] (CF) Istituto di Ricerche sulla Pesca Mar- ittima, Molo Mandracchio, 60100 Ancona, Italy; (RBM) Department of Invertebrate Zoology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 266-268 A SMALL TRAP FOR COLLECTING CRUSTACEANS IN SHALLOW WATER Raymond B. Manning Abstract. — Results from using a small, inexpensive trap, made from plastic water-sampling bottles, to collect crustaceans in shallow water, are reported. The traps can be used to collect both decapods and microcrustaceans. In 1982, M. Turkay published a summary of results on the use of a small trap for col- lecting shrimps. In the Aegean Sea, the trap, developed by H. Thiel, University of Ham- burg, and Turkay, yielded representatives of eight species of caridean shrimps, of which one, Lysmata seticaudata (Risso), was col- lected only with the trap. Turkay’s paper was originally presented at the COLLOQUIUM CRUSTACEA DE- CAPODA MEDITERRANEA II, held in Ancona, Italy, in 1979, and I had an op- portunity to hear the original presentation. While listening to Turkay’s talk I realized that he had come up with an excellent sup- plement to any field kit: a small, inexpen- sive, portable trap for decapods. The Trap The Thiel-Turkay traps (Fig. 1) can be made from a 1-liter narrow-mouthed poly- ethylene bottle, with both the top and bot- tom cut off. The top is inverted and fastened into the end of the bottle to form a funnel- shaped opening. Completed traps measured 12.5 cm long, 9 cm in diameter, with an opening of 2.5 cm. The bottom is replaced by any mesh desired, held in place with a hose clamp. A weight may be attached at the mouth end to hold the trap on the bot- tom. The narrow mouth allows only small animals to enter the trap, which can be bait- ed and emptied by removing the hose clamp. As many traps as desired can be run in a string. Results My first opportunity to use the trap came while I was on Carrie Bow Cay, Belize, in 1982. Three traps, baited with leg segments of spiny lobsters, were set in about four feet of water in coral and shell rubble off the dock on the western side of the island. Over a 24-hour period, no decapods were taken in the traps, and after about 36 hours still no decapods were taken. The traps were then essentially abandoned, forgotten for another two days. When the traps were finally raised, they were allowed to drain upon the dock, where a variety of small crustaceans appeared in the drain water. Included were more than 50 specimens of a red nebaliacean; one cyclopoid, one calanoid, and eight harpac- ticoid copepods; three gammaridean am- phipods; two cirolanid isopods; eight my- odocopid and three podocopid ostracodes; and one small spider crab. This catch dem- onstrated that the traps could be used to collect crustaceans other than decapods. In October 1982, these traps were used in several of the marine caves in Bermuda (see Sket and Iliffe 1980), where Thomas M. Iliffe, C. W. Hart, Jr., and I were studying the cave shrimps (Hart and Manning 1981). These little traps, usually baited with fish, yielded up to four specimens of the Cuban cave shrimp, Barbouria cubensis (Von Mar- tens), when left overnight. In addition, in Wonderland Cave, formerly open as a com- mercial cave for tourists, a trap set and left VOLUME 99, NUMBER 2 =| H a a tf] a a ol Fi a a ty i _| | | Qe Su e B ze Ss CI Coa Fig. 1. Diagrammatic cross-section of the Thiel- Tirkay bottle trap. for a week yielded a specimen of Barbouria as well as an undescribed ostracode related to the Thaumatocyprididae. In October 1983 the traps were again used on Bermuda in several different caves in the Walsingham system. Traps left in the water for 24 to 48 hours yielded a fish, Bathygo- bius, several crabs of the genus Panopeus, at that time not known to occur in the cave systems, and up to 10 specimens of Bar- bouria. These traps all had been baited with pieces of fresh fish. Three traps baited with crushed speci- mens of Panopeus were left in an open pool in an area near the caves for five days, and two baited with fish heads were left in a small sink hole in the Walsingham cave sys- tem for four days. Both yielded the same kind of organisms. Those in the open pool yielded eight specimens of the calanoid co- pepod, Exumella, and four harpacticoid co- pepods. Those in the sink hole yielded 23 specimens of Exumella and five harpacti- coids. In September 1984 these traps, baited with fish, were left overnight in mangrove roots on the east side of Little Jim Island, in the Indian River just north of the Fort Pierce Inlet, St. Lucie County, Florida (=RBM field station 84-10, 20 Sep 1984). There several specimens of Periclimenes americanus (Kingsley) and Leander tenuicornis Say were taken. Periclimenes americanus was one of 267 the most abundant decapods taken in grass beds in the Indian River in an earlier study (Gore, Gallaher, Scotto, and Wilson 1981), but L. tenuicornis apparently was not found in the grass beds in that survey. Remarks The Thiel-Tiirkay trap is an effective piece of gear for collecting a wide variety of crus- taceans in the field. With a fine mesh cover over the opening at the bottom of the bottle, small crustaceans are collected readily. The traps should be left in the water for rela- tively long periods of time, even for several days, when used to collect smaller crusta- ceans. When larger crustaceans are sought, the traps can be left out for much shorter periods, even overnight. Some experimen- tal work on effectiveness of different types of baits in different habitats for varying pe- riods of time would be most interesting. Acknowledgments The samples of microcrustaceans from Carrie Bow were sorted and identified to group by Anne C. Cohen; Douglas C. Barr sorted and identified the copepods from Bermuda. Thomas M. Iliffe spent many hours introducing C. W. Hart, Jr. and me to the caves in Bermuda. Elizabeth Allen, Lilly King Manning, William D. Lee, and Marilyn Schotte provided assistance in the field and in the laboratory. This paper is contribution no. 1052 from the Bermuda Biological Station and contribution 152 from the Smithsonian Marine Station at Fort Pierce. Field work on Bermuda was supported by the National Geographic Society under grant 2485-82 and by a grant from the Scholarly Studies Program of the Smithson- ian Institution; that program also supported work on Carrie Bow Cay, Belize. Literature Cited Gore, R. H., E. E. Gallaher, L. E. Scotto, and K. A. Wilson. 1981. Community composition, 268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON structure, biomass and species-areal relation- ships of seagrass and drift algae-associated mac- rocrustaceans. Studies on decapod Crustacea from the Indian River region of Florida, XI.— Estuarine, Coastal, and Shelf Science 12:485- 508. Hart, C. W., Jr., and R. B. Manning. 1981. The cav- ernicolous caridean shrimps of Bermuda (AI- pheidae, Hippolytidae, and Atyidae).— Journal of Crustacean Biology 1(3):441—456. Sket, B., and T. M. Iliffe. 1980. Cave fauna of Ber- muda.— International Revue der Gesamten Hy- drobiologie 85(6):87 1-882. Tirkay, M. 1982. Results of collecting decapods with small traps in the Aegean Sea.—Quaderni del Laboratorio di Tecnologia della Pesca, Ancona 3(2-5):339-345. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 269-273 TRIDENTELLA RECAVA, A NEW ISOPOD FROM TILEFISH BURROWS IN THE NEW YORK BIGHT (FLABELLIFERA: TRIDENTELLIDAE) Thomas E. Bowman Abstract. — Tridentella recava, n. sp., described from tilefish burrows at depths of 180-304 m on the New York Bight, is characterized by its large size (up to 25.8 mm), its excavated telsonic apex, and the spines on the margins of the uropods. It lives in secondary burrows within the tilefish burrows. The tilefish, Lopholatilus chamaeleonti- ceps Goode and Bean, a large fish reaching 60 pounds is the object of a major fishery along the east coast of the United States and the Gulf of Mexico. It occurs on the outer continental shelf at depths between about 100 m to more than 300 m, and is restricted to a rather narrow band of bottom water that remains between 9-14°C year round. It constructs burrows whose function appears to be to provide refuge from predators. Sev- eral species of crustaceans and fishes are associated with the tilefish burrows (Able et al. 1982, Grimes et al., in press). The isopod described below is a previously undescribed associate in tilefish burrows. Family Tridentellidae Bruce, 1984 Genus Tridentella Richardson, 1905 Tridentella recava, new species Figs. 1-3 Material.— All collected with the R/V Johnson Sea Link IT.—Dive 901, sample 3, 40°00.91'N, 70°05.79'W, 242-304 m, 29 Jul 1984: juv. 69.9 mm; juv. 2 10.1 mm, USNM 227074.— Dive 906, sample 10, 40°01.05'N, 70°20.98'W, 196-230 m, 31 Jul 1984: 2 9 without oostegites, 20.0 and 24.6 mm, USNM 227073.—Dive 909, sample 6, 40°02.96’N, 70°20.84’W, 180-195 m, 2 Aug 1984: 2 2 without oostegites, 24.1 and 25.0 mm; ¢ 20.5 mm; ¢ in 2 fragments; 4 mancas, 5.9, 6.0, 6.3, and 6.3 mm, USNM 227072.—Dive 909, sample 8, 40°02.69’N, 70°21.35'W, 180-195 m, 2 Aug 1985: 9 without oostegites, 25.2 mm; juv. 2 10.2 mm; juv. 2, damaged; ¢ without head, USNM 227071; 2 with empty marsupium, 25.8 mm (holotype, USNM 227070). All specimens listed, except the holotype, are paratypes. Description. —Length up to about 26 mm, about 2.4 x as long as wide. Head with small pointed rostrum meeting frontal lamina and slightly separating bases of antenna 1. Eyes large, with well developed facets. Pereonite 1 with complete transverse furrow; pereo- nites 2-3 with lateral furrows. Coxae 2-3 rounded posteriorly, not produced poste- riorly; coxae 4—7 progressively more point- ed and produced posteriorly, coxa 7 reach- ing pleonite 3. Pleonites 1-3 progressively wider and more produced posteriorly; pleonite 4 about as wide as pleonite 3, pro- duced posterolaterally to midlength of pe- duncle of uropod. Pleonite 5 covered lat- erally by pleonite 4. Pleotelson nearly 1.5 x as wide as long, subtriangular; lateral mar- gins slightly convex, armed with short setae; apex with U-shaped excavation, base of ex- cavation crenulate, with short setae between the crenulations. Antenna 1 peduncle seg- ment 3 about 1.7 x length of segments | and 2 combined; flagellum about 21-merous, segments distal to segment 4 each bearing 2-3 esthetes. Antenna 2 reaching pereonite 3 or 4; peduncle segment 5 about '4 longer 270 Fig. 1. Clay from margin of tilefish burrow, col- lected from submersible by grab sampler on manipu- lator arm and broken to show Tridentella recava in its burrow. than segment 4; flagellum with about 34 segments. Frontal lamina pentagonal, moderately elongate. Clypeus narrow, extending later- ally well beyond labrum; latter with convex free margin. Mandible with narrow 2-pronged incisor; molar triangular, fleshy, with hirsute margin; segment 2 of palp with several rows of long setae on distal %; seg- ment 3 with closely spaced row of shorter setae and a few longer apical setae. Maxilla 1 exopod with 5 long terminal spines and 5 short subterminal spines; endopod less than half length of exopod, with 2 short subter- minal setae. Maxilla 2 apex with about 5 scalelike spines with 3 or 4 teeth; proximal to these a patch of smaller bicuspid spinules. Maxilliped with narrow endite bearing a few terminal setae, reaching distal margin of palp segment 3. Pereopods 1-3 similar, prehensile; pe- reopod 1 somewhat stouter than pereopods 2-3. Carpus with strong spine on postero- distal corner; posterior margin of merus with row of shorter spines. Pereopods 4—7 slen- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON der; distal corners of segments, except basis, with groups of long spines. Pleopod 1 exopod with proximal spine on lateral margin. Pleopods 1-5 with 8, 7, 7, 6, O retinacula. Exopods of pleopods 3-— 5 divided by transverse suture. Appendix masculina of pleopod 2 widening distally into lanceolate tip reaching slightly beyond distal margin of endopod. Uropods reaching beyond telson, endo- pod slightly longer than exopod, both rami rather narrow, pointed, margins armed with a few spines and close-set rows of setae. Etymology. —From the Latin “recavus,” hollowed or arched inward, referring to the apex of the telson. Comparisons. — Tridentella recava is eas- ily distinguished from the 10 known species of the genus (listed in Delaney and Brusca 1985) by its smooth pleotelson with an ex- cavated apex. Seven of the 10 species have highly ornamented pleotelsons, and the three species with smooth telsons have smoothly rounded entire pleotelsons. Marginal spines on the uropods are found in T. /aeviceph- alax Menzies (see Carvacho 1977), T. quin- icornis Delaney and Brusca, 1985, and T. recava, but not in other species of Triden- tella. Tridentella recava is by far the largest known species, with a length of up to 25.8 mm. The other species range in length from 9 mm (T. virginiana [Richardson, 1905]; T. japonica Thielemann, 1910) to 20 mm in T. ornamenta (Menzies and George, 1972). Remarks. —Thanks to the insight of Bruce (1984), Tridentella has been retrieved from the family Corallanidae, in which it had re- mained since 1905, and deservedly given its own family. Bruce’s discussion of the features of the new family is clear and con- cise, except for the molar of the mandible. He describes the molar as “‘present,”’ but in table 1 he states ““molar vestigial.” Else- where he says that the mandible is “essen- tially similar’”’ to that of the Aegidae. The mandible of the Aegidae lacks a molar. Ac- VOLUME 99, NUMBER 2 271 Fig. 2. Tridentella recava, 25 mm 2, dive 909, sample 6: A, Habitus, dorsal; B, Head, ventral; C, Coxae 2— 7, lateral; D, Uropod, ventral; E, Antenna 1; F, Antenna 2, proximal segments; G, Right mandible; H, Molar of right mandible; I, Maxilla 1; J, Maxilla 2; K, Maxilliped; L-P, Pereopods 1-5. 272 N\\ \ i Hl i» UK WY Fig. 3. Pereopod 6; B, Pereopod 7; C—G, Pleopods 1-5. tually, as pointed out by Delaney and Brus- ca (1985), the molar in the Tridentellidae is essentially like that of the Cirolanidae, in which it is conical, fleshy, spinose on one margin, and sometimes setose. It cannot cut or grind, and presumably its function is to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Tridentella recava, A-C, E-G, 25 mm 2, dive 9, sample 6; D, 20.5 mm 4, dive 9, sample 6; A, push food into the mouth. Its structure is clearly specialized, i.e. apomorphic, a fact overlooked by systematists who state that the Cirolanidae have generalized mouth- parts. Habitat.—Tridentella recava was found VOLUME 99, NUMBER 2 in secondary burrows within the tilefish bur- rows. Clumps of clay collected showed iso- pod burrows of two kinds: larger burrows about 5.0 mm in diameter with a distinct whorled pattern of narrow marks resulting from excavation; smaller burrows about 2.0 mm in diameter without distinct excavation marks on the walls. These burrows are oc- cupied by large and small specimens of 7. recava, respectively. A third kind of bur- row, 8—10 mm in diameter with excavation marks larger in scale and more widely spaced, may be produced by a goneplacid crab that is associated with tilefish burrows. Acknowledgments The investigation on tilefish in the Mid- Atlantic Bight was carried out cooperatively by Kenneth W. Able, Rutgers University, New Brunswick, New Jersey; Churchill B. Grimes, then of Rutgers University, now with the National Marine Fisheries Service, Southeast Fisheries Center, Panama City, Florida; Robert S. Jones, then of Harbor Branch Foundation, Ft. Pierce, Florida, now with the University of Texas, Marine Sci- ence Institute, Port Aransas, Texas; and Da- vid C. Twitchell, U.S. Geological Survey. Crustaceans associated with tilefish burrows were sent to Austin B. Williams, National Marine Fisheries Service, Systematics Lab- oratory, by Dr. Able, and Dr. Williams kindly gave me the isopods from this col- lection for study. Dr. Able loaned me the color transparency from which Fig. | was prepared. Literature Cited Able, Kenneth W., Churchill B. Grimes, Richard A. Cooper, and Joseph R. Uzmann. 1982. Bur- 273 row construction and behavior of tilefish, Lo- Dholatilus chamaeleonticeps, in Hudson Sub- marine Canyon.—Environmental Biology of Fishes 7(1):199-205. Bruce, Niel L. 1984. A new family for the isopod crustacean genus Tridentella Richardson, 1905, with description of a new species from Fiji.— Zoological Journal of the Linnean Society 80: 447-455. Carvacho, Alberto. 1977. Isopodes intertidaux de cétes du centre et du Nord du Chili. I. Familles des Cirolanidae, Excorallanidae et Corallani- dae.—Crustaceana 32(1):27—44. Delaney, Paul M., and Richard C. Brusca. 1985. Two new species of Tridentella Richardson, 1905 (Isopoda: Flabellifera: Tridentellidae) from Cal- ifornia, with a rediagnosis and comments on the family, and a key to the genera of Tridentellidae and Corallanidae.—Journal of Crustacean Bi- ology 5(4):728-742. Grimes, Churchill B., Kenneth W. Able, and Robert S. Jones. [In press]. Tilefish, Lopholatilus cha- maeleonticeps, habitat, behavior and commu- nity structure in Mid-Atlantic and southern New England waters.—Environmental Biology of Fishes. Menzies, Robert J., and Robert Y. George. 1972. Iso- pod Crustacea of the Peru-Chile Trench. —An- ton Bruun Report Number 9:9.1—9.124. (Texas A&M Press.) Richardson, Harriet. 1905. A monograph on the iso- pods of North America. — Bulletin of the United States National Museum 54:I-LIII, 1-727. Thielemann, Martin. 1910. Beitrage zur Naturge- schichte Ostasiens. Herausgegeben von Dr. F. Doflein. Band II, No. 9. Beitrage zur Kenntnis der Isopoden fauna Ostasiens.— Abhandlungen der Mathematisch-Naturwissenschaftlichen Klasse der K. Bayer. Akademie der Wissen- schaften, suppl. vol. 2, Abhandling 3:1—109, pls. 1-2. Department of Invertebrate Zoology, Na- tional Museum of Natural History, NHB- 163, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 274-285 SEXUAL DIMORPHISM AND A SEX-LIMITED POLYMORPHISM IN THE COPEPOD PAROITHONA PACIFICA NISHIDA, 1985 (CYCLOPOIDA: OITHONIDAE) FROM THE RED SEA Frank D. Ferrari and Ruth Bottger Abstract.—Paroithona pacifica Nishida, 1985, from the Red Sea is the first species of the genus known to exhibit polymorphism. Among females the number of external spines on swimming legs 3 and 4 exopods is variable; a pair of legs on one animal also may express variation in external spine number. Males do not exhibit similar variation within or among animals. Sexual di- morphism in the genus Paroithona is manifested in 18 differences in number, shape, and position of various exoskeletal elements. Differences in the number of segments and/or setae on swimming legs 2—4 are unique to this group relative to three other oithonid lineages. Farran (1908) established the genus Pa- roithona from an unspecified number of fe- male specimens of Paroithona parvula found in two samples (SR 193—54°50’N, 10°30'W; SR 197—54°57'N, 10°51'W) collected be- low 1000 m off the northwest coast of Ire- land with an open, conical net (mesh size about 20 microns). Farran’s initial descrip- tion noted that Paroithona differed from Oithona in having three well-developed lobes on the first maxilla (Mx1), swimming legs with 3-segmented exopods but 2-seg- mented endopods, and leg 5 with only one seta. In describing four females of another species, P. pulla, taken in shallow, near- shore waters around Christmas Island in the Indian Ocean, Farran (1913) mentioned 2-segmented endopods on the swimming legs and the form (unspecified) of the man- dibular palp as differentiating Paroithona from Oithona. Paroithona pulla differed from P. parvula by the reduced number of exopodal segments (2) on swimming leg 1 and the reduced number of outer spines on the exopods of swimming legs 3 and 4. Lat- er, Sars (1918) redescribed females of P. parvula from the west coast of Norway, and emphasized the 2-segmented endopods of the swimming legs and the simple structure of the mandibular palp. Kiefer (1929) for- malized these differences for Paroithona— reduced number of endopodal segments of swimming legs 1—4 and the single spine on the tip of mandible basipod 2—in his widely accepted definitions of the oithonid genera. Greze (1963), Shmeleva (1964), and Ra- zouls (1968) recently reported P. parvula from the Ionian Sea, Adriatic Sea, and Gulf of Lyon respectively in the Mediterranean Sea, and Nishida et al. (1977) collected P. pulla from Tokyo Bay, Sagami Bay, and the adjacent Kuroshio, in the northwest Pacific Ocean. Two new species, P. flemingeri from the North Atlantic Ocean off Venezuela and P. pacifica from the Pacific Ocean were de- scribed respectively by Ferrari and Bowman (1980) and by Nishida (1985). Descriptions of P. pulla females are still incomplete and one of us (FDF) recently determined that P. flemingeri has a one-segmented exopod with four setae on maxilla 1 and a swimming leg 1 exopod with five inner margin setae, not four as in the description of Ferrari and Bowman (1980). The genus Paroithona was not mentioned in Halim’s (1969) review of VOLUME 99, NUMBER 2 Red Sea copepods. However Delalo (1966) reported P. pulla from the northern part of the Red Sea in the winter of 1961/1962. In describing P. parvula, Farran (1908: 90) noted that the numbers of setae ob- served “must be regarded as somewhat doubtful” due to the “minute size and ex- treme transparency” of the animals. In the intervening years, observations on oithonid copepods of the genus Paroithona have been hampered not only by small size of the an- imals but also by relatively small numbers of animals collected from one or a few lo- calities. Morphological observations of pre- served animals often are difficult because their exoskeleton does not stain easily. In this paper we report 48 females and 11 males of P. pacifica from the Red Sea and expand the description to include a female sex-limited polymorphism in external ex- opodal spines. We also compile differences between female and male skeletal mor- phology, and compare this degree of sexual dimorphism to three other oithonid lin- eages. Material and Methods Zooplankton samples were collected aboard the R/V Valdivia between 9 October and 9 November 1980 in the northern and central Red Sea, and in February 1981 in the central Red Sea. Stations are located over the Kebrit Deep and Atlantis II Deep, or a northern reference station located north of Kebrit Deep and sampled only in October 1980 (Table 1). Consecutive depth strata (usually 50 m intervals between 0-450 m) were sampled with vertical tows using a multiple opening-closing net (Weikert and John 1981) with 0.1 mm mesh size and an opening area of 0.25 m*. Plankton samples were fixed in 4% formaldehyde-seawater so- lution buffered with hexamethylenetetra- mine. Specimens were sorted and preserved in 5% propylene glycol, 0.5% propylene phenoxetol and 94.5% filtered seawater (Steedman 1976). 275 Specimens of Paroithona pacifica were separated from samples during a compre- hensive study of metazoan composition, abundance, and vertical distribution, whose results are presented by Bottger (1985). Af- ter examination of Paroithona pacifica specimens from the central Red Sea (Atlan- tis II Deep) in October, single samples from each region and season were checked for geographical and seasonal distribution of the species. Specimens were cleared for morphologi- cal examination in lactic acid. After initial observations, including measurements of body length, each was stained by adding a solution of chlorazol black E dissolved in 70% ethanol/30% water. After staining, specimens were transferred to lactic acid for dissection and final observations. The following morphological abbrevia- tions are used in the descriptive text and illustration legends: Body Segments Pr—prosome Cph—cephalosome Pg—pediger Ur—urosome CR—caudal ramus Appendage Elements Bspd— basipodal segment Re—exopodal seg- ment Ri—endopodal seg- ment Li—inner lobe Le—outer lobe ( )—adjacent seg- ments fused ) (—adjacent seg- ments incom- pletely fused Appendages AlI—antenna 1 AlI—antenna 2 Mn—mandible MxI—maxilla 1 MxII—maxilla 2 Mxp—maxilliped P—swimming leg Appendage Armament Se—external spine or seta Si—internal spine or seta St—terminal spine on P Fig. 1. Paroithona pacifica female: A, Lateral; B, Ur, lateral; C, Url—2, dorsal; D, Head, dorsal; male: E, Lateral: F, Ur, lateral; G, Ur1-3, dorsal; H, Head dorsal; female: J, Genital segment with spermatophore, lateral. VOLUME 99, NUMBER 2 Table 1.—Occurrence of Paroithona pacifica in the Red Sea in autumn and winter 1980/1981. Station Region Position Date Depth (m) Specimens Va 29-03 Northern Reference 34°56.69’E, 9 Oct 80 50-100 12 females Station 26°45.36'N 5 males Va 29-23 Kebrit Deep 36°16.63’E, 11 Oct 80 50-100 10 females 24°43.32'N 2 males Va 29-125 Atlantis II Deep 38°04.20’E, 27 Oct 80 60-80 16 females 21°23.28'N 4 males Va 29-676 Atlantis II Deep 38°04.66’E, 23 Feb 81 50-100 10 females 21°22.56'N Results Paroithona pacifica Nishida, 1985 Figs. 1-6 Paroithona pacifica Nishida, 1985:109, figs. 64, 65. Description.—Female: length 0.43-0.47 mm (33 specimens); Pr/Ur-1.3. Rostrum absent (Fig. 1 A); forehead rounded dorsally; internally with paired clusters of polygonal cells (Fig. 1D). CR length 1.7 x width, armed (Fig. 2B) with 1 lateral seta, ventral surface hairs, | dorsal and 4 terminal setae; 2 setae, apparently dorsal and 2nd terminal, elon- gate (but see description of male). Al (Fig. 2A) apparently 9 free segments, armature undetermined. A2 (Fig. 6A) 2 segments, 2nd with incomplete suture; respectively with 2 lateral and 5 lateral + 7 terminal setae. Mn gnathobase simple; Baspd2 (Fig. 5A) with 1 large terminal spine and 2 subterminal setae; Re 4 segments, 4—5 fused, with 5 se- tae, 2 terminal; Ri a non-articulated seg- ment with 4 setae, 1 larger with setules. Mx1 (Fig. 5B) Lil with 4 thick + 3 thin spines; Bspd2 with 1 large terminal spine and 1 subterminal seta; Re 1 segment with 4 setae; Ri 1 unarmed segment. Mx2 segmentation unclear, 3 inner and | terminal lobes armed as in Fig. 6B. Mxp 4 segments armed as in Fig. 6C. P1—4 (Figs. 3A, B, 4A, B); each Bspd2 with 1 small, outer seta. Bspd1P1 with 1 inner seta; Bspd2P1 with 1 inner spine + hairs—these elements absent on P2— 4. Re 2-3P1 fused; Re 2—3P2 fused on an- terior surface; RiP1-—4 2 segments. ReSe 1-(3), 1-)3(, 1-1-2 or 1-0-2 or 1-0-1, 1-0-2 or 1-1-1 or 1-0-1 or 0-0-1 (not all combi- nations illustrated); Si 1-(5), 1-)6(, 1-1-5, 0-0-5; RiSe 0-1, 0-1, O-1, 0-1; Si 1-6, 0-3, 0-3, 1-3; all setae simple except Ri 1P4 thicker. P5 (Fig. 1B) 2 simple, unarticulated, lateral cylindrical segments, with 1 seta each, dorsal with setules. P6 (Fig. 1B, C) 2 small spines, dorsal-most thicker. Male: Length 0.37-0.40 mm (10 speci- mens); Pr/Ur-1.7. Rostrum absent (Fig. 1E); forehead rounded dorsally; small clusters of polygonal cells internal to upper lip (Fig. 1H). CR length 1.7 x width, armed (Fig. 2D) with | lateral, 1 dorsal, and 4 terminal setae; middle 2 terminal setae elongate; ventral hairs absent. Al (Fig. 2C) a complicated digeniculate appendage. A2 (Fig. 6D) sim- ilar to female except distinctly 3-segmented. Mn gnathobase simple; Baspd2 (Fig. 5C) with 1 small thick spine and 1 thin seta terminally; Re segments incompletely fused, with 5 setae; Ri an articulated segment with 4 setae, largest with setules. Mx1 (Fig. 5D) Lil with 4 thick + 3 thin spines; Bspd2 1 thick spine and 1 seta terminally; Re | seg- ment with 4 setae; Ri 1 unarmed segment, fused to Bspd2. Mx2 segmentation unclear, 4 inner and 1 terminal lobes, armed as in Fig. 6E. Mxp apparently with 4 segments armed as in Fig. 6F. P1—4 (Figs. 3C, D, 4C, D); each Bspd2 with 1 small outer seta. Bspd1P1 with 1 inner seta; Bspd2P1 with 1 inner spine. Re 2 and 3P1 fused; ReP2-—4 3 segments; RiP1 and 4 2 segments; RiP2-3 278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Paroithona pacifica female: A, Al; B, CR; male: C, Al; D, CR. 3 segments. ReSe 1-(3), 1-1-2, 1-1-2, 1-1-2; Si 0-(5), 0-1-5, 0-1-5, 0-1-5; RiSe 0-1, 0-0-1, 0-0-1, 0-1; Si 1-6, 0-1-5, 1-2-5, 1-6; all setae simple except Ri 1P4 thicker. P5 (Fig. 1F) 2 unarticulated knobs, dorsal-most smaller, with 1 seta each. P6 (Fig. 1F, G) 2 unarti- culated knobs, dorsal-most smaller, with 1 seta each. Remarks.—Males of P. pacifica differ from Paroithona pulla, the only other Pa- roithona males described (Nishida et al. 1977) in swimming leg 2 endopod with 0-1-6 setae (1-2-6 for P. pulla, as illustrated) and a 2-segmented endopod (vs. 3-segmented for P. pulla by Nishida et al.) on swimming leg 4. Males available for this study exhib- ited variation in lateral curvature of the gen- ital segment in dorsal view. This variation seems dependent on the degree of devel- opment of the spermatophores. Specimens VOLUME 99, NUMBER 2 with large, round spermatophores exhibited broadly curved, rounded genital segments; those with small, poorly-developed sper- matophores had cylindrically-shaped geni- tal segments. Discussion Zooplankton samples taken over the At- lantis II Deep in late February 1981 con- tained only females of P. pacifica. Both sexes were found at two northern stations and over the Atlantis II Deep in October-November 1980; females bearing spermatophores were confined to the Northern Reference Station. Spermatophores on females (Fig. 1J) are sub- oval or kidney-shaped structures attached laterally by a fertilization tubule with a disc- like plug. Spermatophores occurred in pairs (three females) or multiple pairs (one female with four). Two females carried single sper- matophores with remnants of a second plug; the spermatophore associated with this plug is considered to have been lost during sam- pling. Paroithona pacifica occurred mainly at 50-100 m at all stations with an abundance of 60 adults/m? estimated from 60-80 m at Atlantis II Deep in October. The species did not show a clear relation to the thermohal- ocline which was situated between 40-80 m in October-November and 80-130 m in February. This subsurface distribution in the Red Sea is somewhat shallower and more compact than that reported by Nishida and Marumo (1982) for Paroithona spp. in the Indian Ocean and South China Sea. Females of Paroithona available for this study exhibit variability in the number of external spines on swimming legs 3 and 4 exopods. Detection of such variability is complicated by the small size of these spines, especially on leg 4 where the proximal spine on exopod 3 may be found closely appressed to the exopod (Fig. 4B2). We carefully stained and examined all swimming legs of 28 females; 22 females with symmetrical 279 exopods on legs 3 and 4 had 3 armament types: leg 3 1-0-2, leg 4 1-0-2; 14 specimens leg 3 1-0-2, leg 4 1-0-1; 7 specimens leg 3 1-1-2, leg 4 1-1-1; 1 specimen tp. D Six females had asymmetrical armament on 1 pair of legs 3 and 4 (left/right): leg 3 1-1-2/1-0-2, leg 4 1-0-2; 1 specimen leg 3 1-1-2/1-0-2, leg 4 broken off; 1 spec- imen leg 3 1-0-2/1-0-1, leg 4 1-0-2; 1 specimen leg 3 1-0-2, leg 4 1-0-1/1-0-2; 2 specimens leg 3 1-0-2, leg 4 1-0-2/1-0-1; 1 specimen In some cases the variation may be artifi- cial, resulting from difficulties in determin- ing the presence of the thin subterminal |st spine on exopod 3 of leg 4 (Figs. 4B1, 2). Presence of a thick, terminal spine on ex- opod 2 of leg 3 (two asymmetrical speci- mens) or leg 4 (one specimen) provides conclusive evidence of a sex-limited poly- morphism occurring in Paroithona pacifica. Discovery of this sex-limited polymor- phism in exopod armature has temporarily compromised use of these traditional struc- tures in differentiating species of Paroi- thona. Whether this variation is wide- spread in the genus, confined to this species, or is a local phenomenon— perhaps a prod- uct of secondary contact between Atlantic and Indo-Pacific species in the vicinity of the Suez Canal—remains to be determined. Some morphs of P. pacifica from the Red Sea have the same spine count as Atlantic species P. parvula and P. flemingeri (Table 2). However, there is no overlap with the other Pacific species, P. pulla. In Paroithona, a number of differences between females and males reflects the de- gree of sexual dimorphism. They comprise a) changes in numbers of elements, b) rel- ative changes in shape or size of elements, c) changes in position of elements as fol- lows: changes in number include: 280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Paroithona pacifica female: A, P1; B, P2; male: C, P1; D, P2. 1) antenna 1: male more segments and se- tae than female. 2) urosome: female 1 less segment than male. 3) swimming leg 2 endopod: female | less segment. 4) swimming leg 2 endopod: female 3 fewer setae. VOLUME 99, NUMBER 2 281 Fig. 4. Paroithona pacifica female: A, P3; B, P4; male: C, P3; D, P4. 5) swimming leg 3 endopod: female 1 less 7) swimming leg 4 exopod: female 1 less segment. seta. 6) swimming leg 3 endopod: female 5 fewer 8) swimming leg 4 endopod: female 3 fewer setae. setae. 282 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Paroithona pacifica female: A, Mn; B, Mx1; male: C, Mn; D, Mx1. 9) swimming leg 2 exopod: segments 2 and 3 partly fused in male, completely fused in female. Changes in shape or size include: 1) antenna 1: many segments and setae. 2) mandible exopod and basipod 2 arma- ture: male smaller than female. 3) maxilla 1: male smaller. 4) maxilla 2: male smaller. 5) maxilliped: male smaller. 6) leg 5: male smaller. 7) leg 6 + setae: female smaller than male. 8) swimming leg 1: external spines of ex- opod, long and smooth in female, short- er and dentate in male. Changes in position include: 1) leg 6: mid dorsolateral on fused urosome 2-3 in female, posterior and ventrolat- eral in male urosome 2. Comparisons of degree of sexual dimor- phism in Paroithona with other oithonid lineages is hampered by lack of descriptions of many males. At present we hypothesize four lineages: 1) “‘paroithona,” all species of Paroithona; 2) “‘limnoithona,” all species of Limnoithona; 3) “‘dioithona”’ all species of the subgenus Oithona (Dioithona) plus O. bjornbergae; 4) “‘oithona”’ all other species of subgenus Oithona (Oithona). We rely on descriptions of both sexes of Limnoithona in Ferrari and Orsi (1984), of Oithona (Dioithona) in Nishida (1985), of Oithona (Oithona) in Ferrari and Bowman (1980), Ferrari and Orsi (1984) and Uchima (1979) for coastal zone animals, and Nishida (1985) for several oceanic species. In all lineages many changes including an- tenna 1, leg 6 and fusion of the urosome segments appear similar. Relative to “lim- noithona” the following changes are unique VOLUME 99, NUMBER 2 Fig. 6. Paroithona pacifica female: A, A2; B, Mx2; C, Mxp; male: D, A2; E, Mx2; F, Mxp. to “‘paroithona’’—in number of segments in swimming leg 2 exopod and swimming legs 2 and 3 endopods, in number of setae on swimming legs 2, 3, and 4 endopods and swimming leg 4 exopod, in size of male ex- opod and basipod armature, and in shape of external spines of swimming leg 1 exo- pod. A sexual dimorphism expressed in the caudal rami (segment and armature size slightly smaller in males) is unique to Lim- noithona. Unique differences in degree of sexual di- morphism of “‘dioithona”’ relative to “‘pa- roithona” include changes in shape of an- tenna 2 segment 3 (longer in males), mandibular basipod (segmented in males), shape of setae on swimming leg 4 endopod (modified in females). Unique differences in ‘“‘paroithona”’ relative to “‘dioithona” in- clude changes in shape of external spines on swimming leg 1 exopod, and changes in number of segments and setae on swimming legs 2-4 endopods, and segments or setae on swimming legs 2 and 4 exopods. The “oithona”’ lineage shows a wide range in degree of sexual dimorphism among var- ious species occupying a wide number of aquatic niches. All members of this lineage 284 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Number of spines previously reported on exopodal segments of Paroithona swimming legs 1-4. Name Leg 1 Leg 2 Leg 3 Leg 4 Source P. parvula 1-1-2 1-1-2 1-1-2 1-1-1 Farran 1908, Sars 1918 P. pacifica 1-)3( 1-1-2 1-0-2 1-0-2 Nishida 1985 P. flemingeri 1-(3) 1-1-2 1-0-2 1-0-1 Ferrari and Bowman 1980 P. pulla 1-(3) 1-1-2 1-0-1 0-0-1 Farran 1913, Nishida et al. 1977 ()—adjacent segments fused. ) (—adjacent segments incompletely fused. exhibit differences relative to “‘paroithona”’ in shape of antenna 2 segment 3 (longer in males), ornamentation on coupler of swim- ming leg 4 (present in females, absent in males), shape of setae on swimming leg 4 endopod (modified in females), changes in number of integumental organs on cepha- losome (more numerous and forming an un- usual pattern in males). Unique differences in degree of sexual dimorphism of “‘paro- ithona”’ relative to “‘oithona’’ include changes in shape of external spines on swimming leg 1 exopod, and changes in number of segments and setae on swimming legs 2-4 endopods and segments or setae on swimming legs 2 and 4 exopods. Each set of differences can be considered a character unique to one lineage and exclusive of the other three. Differences in degree of sexual dimorphism clearly separate the “‘paro- ithona” lineage from the remaining lineages in the family. Acknowledgments This study was supported in part by a Deutsche Forschungsgemeinschaft research grant to R.B. Cruise 29 of R/V Valdivia was funded by the Red Sea Joint Commission, Jeddah, and the Bundesministerium fur Forschung und Technologie, Federal Re- public of Germany. We wish to thank R. Cressey for use of his Nomarsky interfer- ence microscope, D. Schnack and K. Hiilse- mann for helpful comments, and S. Nishida for a careful review of the manuscript. Literature Cited Bottger, R. 1985. Untersuchungen zur Verteilung der kleinen Metazoa im Plankton des Roten Meeres, unter besonderer Beriicksichtigung cyclopoider und harpacticoider Copepoden. Dissertation, Fachbereich Biologie, Universitat Hamburg, 248 pp. Delalo, E. 1966. Distribution of the zooplankton bio- mass in the Red Sea and the Gulf of Aden, win- ter 1961/62.—Okeanologicheskiye Issledova- niya 15:131—139. [in Russian, English abstract] Farran, G. 1908. Second report on the Copepoda of the Irish Atlantic Slope.—Scientific Investiga- tions Fisheries Branch, Ireland 1906 2:3-104. 1913. Plankton from Christmas Island, In- dian Ocean.—II. On Copepoda of the genera Oithona and Paroithona.—Proceedings of the Zoological Society of London, 1913, pp. 181- 193, pls. XX VII-XXXI. Ferrari, F., and T. E. Bowman. 1980. Pelagic cope- pods of the family Oithonidae (Cyclopoida) from the east coasts of Central and South America. — Smithsonian Contributions to Zoology 312:1- Di , and J. Orsi. 1984. Oithona davisae, new species, and Limnoithona sinensis (Burckhardt, 1912) (Copepoda: Oithonidae) from the Sacra- mento-San Joaquin Estuary, California. —Jour- nal of Crustacean Biology 4:106-126. Greze, V. 1963. Zooplankton of the Ionian Sea.— Okeanologicheskiye Issledovaniya 9:42—59. [in Russian, English abstract] Halim, Y. 1969. Plankton of the Red Sea.—Ocean- ography and Marine Biology, Annual Review 7: 231-275. Kiefer, F. 1929. Cyclopoida Gnathostoma. Das Tier- reich 2. Walter de Gruyter und Co., Berlin und Leipzig, xii + 104 pp. Nishida, S. 1985. Taxonomy and distribution of the family Oithonidae (Copepoda, Cyclopoida) in the Pacific and Indian oceans.— Bulletin of the Ocean Research Institute, University of Tokyo 20:1-167. VOLUME 99, NUMBER 2 ,and R. Marumo. 1982. Vertical distribution of cyclopoid copepods of the family Oithonidae in the western Pacific and eastern Indian oceans. — Bulletin of the Plankton Society of Ja- pan 29:99-118. , O. Tanaka, and M. Omori. 1977. Cyclopoid copepods of the family Oithonidae in Suruga Bay and adjacent waters. — Bulletin of the Plank- ton Society of Japan 24:119-158. Razouls, C. 1968. Présence du genre Paroithona Far- ran (Copepoda, Cyclopoida) dans le Golfe du Lion.— Vie et Milieu ser. A, 19:153-158. Sars, G. 1918. Copepoda, Cyclopoida. An account of the Crustacea of Norway, vol. VI. Bergen Museum, Bergen, Norway, xiii + 295 pp. Shmeleva, A. 1964. New Copepoda species in the Adriatic Sea and the characteristic features of their distribution. Okeanologiya 6:1066—1072. [in Russian] Steedman, H. 1976. Examination, sorting and ob- servations fluids. Pp. 182-183 in H. Steedman, Ed., Zooplankton fixation and preservation, 285 monographs on oceanographic methodology 4, 350 pp. UNESCO Press, Paris. Uchima, M. 1979. Morphological observation of de- velopmental stages in Oithona brevicornis (Co- pepoda, Cyclopoida).— Bulletin of the Plankton Society of Japan 26:59-76. Weikert, H., and H.-C. John. 1981. Experiences with a modified Bé multiple opening-closing plank- ton net.—Journal of Plankton Research 3:167-— 176. (FDF) Smithsonian Oceanographic Sort- ing Center, Smithsonian Institution, Wash- ington, D.C. 20560, U.S.A.; (RB), Institut fiir Hydrobiologie und Fischereiwissen- schaft, Hydrobiologische Abteilung, Uni- versitat Hamburg, Zeiseweg 9, D 2000 Hamburg 50, Federal Republic of Ger- many. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 286-315 MARINE TUBIFICIDAE (OLIGOCHAETA) AT HUTCHINSON ISLAND, FLORIDA Christer Erséus Abstract. —Twenty-five species of marine Tubificidae are recorded from off- shore, largely coarse, shelly sediments at Hutchinson Island on Florida’s lower central east coast. Nine species are described as new: Heterodrilus hispidus, H. perkinsi (subfamily Rhyacodrilinae), Phallodrilus hirsutus, Bathydrilus ingens, B. formosus, B. macroprostatus, Coralliodrilus corpulentus, Olavius latus (Phal- lodrilinae), and Tubificoides annulus (Tubificinae). Redescriptions of Hetero- arilus bulbiporus Erséus, 1981, and Phallodrilus acochlearis (Erséus and Loden, 1981), n. comb., are provided. Heterodrilus bulbiporus, H. occidentalis Erséus, 1981, H. pentcheffi Erséus, 1981, and Adelodrilus magnithecatus Erséus, 1979, are recorded from Florida for the first time. The Hutchinson Island tubificid fauna comprises a mixture of warm-temperate (American east coast) and trop- ical (Caribbean) elements. In 1943, Gates described an intertidal earthworm, Pontodrilus gracilis [now re- garded as a synonym for P. /itoralis (Grube, 1855), family Acanthodrilidae; Easton 1984] from the banks of a brackish river at Fort Myers, Florida. However, it was not until the middle of the 1960’s that a marine fauna of aquatic oligochaetes was discovered in this part of the southeastern U.S.A. The first, very scanty, records of Tubificidae from marine waters in Florida were given by Brinkhurst (1965, 1966), while Kennedy (1966) used material from off Panama City, Florida, when describing the marine enchy- traeid, Grania americana. During a collecting trip to Miami in 1977, the author had the opportunity to gather a rich material of marine oligochaetes in the area just south of the city (largely Biscayne Bay). The Tubificidae dominated this ma- terial, and most of the tubificid species have now been accounted for in various taxo- nomic publications (Erséus 1979a, b, c, 1980a, 1981a, 1982a, b, c, 1983a, 1984b). Additional collections of Tubificidae from other parts of the state have been treated by Erséus (1981b, 1984a), Erséus and Baker (1982), Erséus and Loden (1981), and Lod- en (1980). The species thus recorded from Florida to date number 34 (species not as- terisked in Table 1). One enchytraeid species, Grania mono- spermathecus Erséus and Lasserre, 1976, was recently reported from the Miami area by Coates and Erséus (1985), and additional enchytraeid material will be treated else- where by the same authors (in preparation). In 1981, Erséus and Loden reported on a small collection of Tubificidae (four species of the subfamily Phallodrilinae) from sub- tidal sites off Hutchinson Island on Flori- da’s lower central east coast. The material was obtained in 1976-1977 during a sam- pling program funded by Florida Power and Light Company (FP&L). The program was a continuation of an earlier, three-year (1971-1974) environmental baseline study conducted in connection with the location of the FP&L St. Lucie Unit No. 1 nuclear power plant on Hutchinson Island. The study was designed to generate a marine ecology data base for the future assessment of possible changes in the environment of the area influenced by the heated effluent VOLUME 99, NUMBER 2 Table 1.—Checklist of marine Tubificidae reported from Florida, with asterisks denoting new records. “Florida only” means species hitherto only reported from Florida. E. = Erséus. Flor- ida Species only Reference Adelodrilus magnithecatus E., present paper 1979* Aktedrilus floridensis E.,1980 x! E. 1980a Bathydrilus adriaticus (Hrabé, E. 1979a, 1985 1971) B. formosus, n. sp.* present paper B. ingens, n. sp.* present paper B. macroprostatus, n. sp.* x present paper Bermudrilus peniatus E., E. 1984a 1979 Coralliodrilus corpulentus,n. x present paper sp.* Heterodrilus bulbiporus E., present paper 1981* Hi. hispidus, n. sp.* X present paper H. minisetosus E., 1981 E. 1981a; pres- ent paper H. occidentalis E., 1981* present paper H. pentcheffi E., 1981* present paper H. perkinsi, n. sp.* X present paper Inanidrilus bulbosus E.,1979 x E.1979c I. ernesti E., 1984 x E. 1984b; pres- ent paper I. extremus (E., 1979) x E. 1979b, 1984b I. mexicanus (E. and Baker, x EE. and Baker 1982) 1982 I. triangulatus E., 1984 x E. 1984b I. vacivus E., 1984 x EE. 1984b, pres- i ent paper Kaketio ineri Righi and Kan- E. 1980b ner, 1979 Limnodriloides baculatus E., E. 1982c 1982 L. barnardi Cook, 1974 E. 1982c L. hastatus E., 1982 x E. 1982c L. monothecus Cook, 1974 E. 1982c; pres- ent paper L. rubicundus E., 1982 E. 1982c L. vespertinus E., 1982 E. 1982c; pres- ent paper Marcusaedrilus hummelincki E. 1983a Righi and Kannner, 1979 M. luteolus E., 1983 Monopylephorus rubroniveus Levinsen, 1884 (=M. he- lobius Loden, 1980) E. 1983a; pres- ent paper Loden 1980; Baker and Brinkhurst 1981 287 Table 1.—Continued. Flor: ida Species only Reference Olavius caudatus (E., 1979) x E. 1979b, 1984b O. latus, n. sp.* x present paper O. macer E., 1984 x E. 1984b O. tenuissimus (E., 1979) E. 1979b, 1984b O. sp. A* x? present paper O. sp. B* X present paper Parakaketio longiprostatusE., x E. 1982b 1982 Phallodrilus acochlearis (E. E. and Loden and Loden, 1981), n. 1981; present comb. paper P. biprostatus (Baker and E., E. and Loden 1979) 1981; present paper P. hirsutus, n. sp.* x present paper P. rectisetosus E., 1979 P. sabulosus E., 1979 E. 1979b, 1985 E. 1979b; pres- ent paper; E. and Loden 1981 Smithsonidrilus marinus ma- x3 Brinkhurst rinus Brinkhurst, 1966 1966; E. 1982a “2 Spiridion insigne Knollner, ? Brinkhurst 1935” (=indeterminable) 1965; Brink- hurst and Baker 1979 Tectidrilus bori (Righi and E. 1982c Kanner, 1979) T. squalidus Erséus, 1982 x E. 1982c Thalassodrilides gurwitschi E. 1981c (Hrabé, 1971) Tubificoides annulus, n. sp.* x T. “gabriellae’ (not gabriellae Marcus, 1950) present paper Brinkhurst 1965; Brink- hurst and Baker 1979 present paper; Brinkhurst (in prepara- tion) ~ T. sp.* ? 1 Known also from Barbados (Erséus unpublished). 2 A very similar form, probably the same species, known from Belize (Erséus unpublished). 3 Subspecies westoni Erséus, 1982, known from New Jersey, Delaware, Maryland, Virginia, North Carolina, Bermuda, and Bahamas (Erséus 1982a). 288 discharge of the plant. An introduction to this extensive study was provided by Gal- lagher and Hollinger (1977), and several re- ports have been given in Florida Marine Research Publications (1977-1979), a series published by the Florida Department of Natural Resources Bureau of Marine Re- search (FDNR). During the 1971-1974 sampling pro- gram, about 2400 specimens of marine Oli- gochaeta were collected at five offshore sta- tions within a few kilometers from the FP&L plant site. Through the kind efforts of Mr. Thomas H. Perkins (FDNR), this very large collection was recently made available to the author. However, as a large majority of the specimens proved to be sexually im- mature and thus very difficult to identify to species in most cases, only those worms with well-developed genital organs, about 400 individuals, were more carefully scruti- nized. Almost 90% of the mature oligo- chaetes examined are members of the Tubi- ficidae, and they are the subject of the present account, which is devoted primarily to the taxonomy of the species. Twenty-two named and three unnamed tubificid species are rec- ognized. Of these, nine are described as new, and four others are reported from Florida for the first time. Heterodrilus bulbiporus Erséus, 1981, H. minisetosus Erséus, 1981, Bathydrilus ingens, n. sp., and B. formosus, N. Sp., are simultaneously reported from the Gulf of Mexico; B. formosus is reported also from Belize in Central America. The enchytraeids in the Hutchinson Is- land material will be treated separately. Stations Sampled The five stations off Hutchinson Island were described by Gallagher and Hollinger (1977), and their sediments more carefully examined by Gallagher (1977). A brief sum- mary is provided here. Sta 1. 27°22.2'N, 80°14.1'W, at seaward margin of beach terrace, 0.5 km offshore. Depth 6-9 m (mean sampling depth 7.5 m). Gray, hardpacked, fine to very fine sand. Note: all oligochaetes here reported from PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sta 1 were from the May 1972 collection, which was taken slightly east (seaward) of the typical Sta 1, in a sediment resembling that of Sta 5 (Gallagher 1977:18). Sta 2. 27°21.6'N, 80°13.1'W, 1.5 km ENE of Sta 1 in the offshore “trough,” approxi- mately midway between the beach terrace and offshore shoal. Depth 9-12 m (mean 11 m). Clean shell hash; very coarse to coarse sand with varying amount of granule-sized shell fraction. Sta 3. 27°21.9'N, 80°12.4'W, Pierce Shoal, 3 km from Sta 1, in line with Sta 1 and Sta 2. Depth 5—8 m (mean 7 m). Clean, medium sands with very small amount of granule- sized shell. Sta 4. 27°20.8'N, 80°12.9'W, 1.6 km SSE of Sta 2, 0.6 km W of southernmost tip of Pierce Shoal. Depth 9-12 m (mean 10 m). Shell hash, very similar to sediment of Sta 2, but with lesser amount of large shell. Sta 5. 27°22.6'N, 80°149I"Wo) 228k NNW of Sta 2, 2.1 km E of the beach in similar depth as Sta 4 (9-12 m). Very coarse shell sands with granule-sized shell making up 28-45% of sample, and with some mud. Material and Methods Five replicate Shipek grab samples (each 0.04 m?’) were taken at each station bi- monthly between September 1971 and July 1973. Samples were preserved in 10% buff- ered formalin-seawater. For further infor- mation on the sampling and sorting pro- cedures, see Gallagher and Hollinger (1977). Sexually mature tubificids, which oc- curred in about one-third of the 300 sam- ples, were sorted and separated from the juveniles by the author. A few specimens of Heterodrilus perkinsi, n. sp., and Phallo- drilus biprostatus were longitudinally sec- tioned and stained with Azan. The remain- ing worms were all stained in paracarmine and mounted whole in Canada balsam. Material from other areas included in this study comprises (1) a few worms from off- shore localities in the NE Gulf of Mexico, originating from a Bureau of Land Man- agement baseline study by personnel at VOLUME 99, NUMBER 2 Dauphin Island Sea Lab, Alabama (cour- tesy Dr. M. Susan Ivester); (2) a single spec- imen of Bathydrilus ingens, n. sp., collected by Mr. Michael R. Milligan (Mote Marine Lab, Sarasota, Florida) on the west coast of Florida; (3) a number of specimens of B. formosus, n. sp., collected by the late Dr. H. Randall Baker (University of Victoria, B.C., Canada) in the vicinity of Carrie Bow Cay on the barrier reef of Belize in the west- ern Caribbean Sea (courtesy Dr. Ralph O. Brinkhurst). These worms were all studied as whole mounts. Holotypes of all the new species are de- posited at the U.S. National Museum of Natural History (USNM), Washington, D.C.; paratypes (when available) and rep- resentative specimens of most of the pre- viously known species are lodged in the Florida Department of Natural Resources Invertebrate Collection (museum prefix FSBC I; = old abbreviation for Florida State Board of Conservation), Bureau of Marine Research, St. Petersburg, Florida. In descriptions, the following abbrevia- tions are used: sec = sectioned; spm/spms = specimen/specimens; whm = whole- mounted. Length and width measurements refer to fixed, whole-mounted, and slightly compressed specimens. Roman numerals denote segment number. Abbreviations used in figures are as fol- lows: a, atrium; bs, blind sac; cs, copulatory sac; ep, epidermal pad; fp, flap-like papilla; g, glandular body of unknown function; gs, glands associated with spermathecal seta; m, muscles; mp, male pore; p, penis; pa, papilla; pp, pseudopenis; pr, prostate gland; pr 1, anterior prostate gland; pr 2, posterior prostate gland; ps, penial seta; s, sperma- theca; se, seta; ss, spermathecal seta; sz, spermatozeugma; vd, vas deferens. Descriptions Subfamily Rhyacodrilinae Heterodrilus Pierantoni, 1902 Heterodrilus minisetosus Erséus, 1981 Clitellio arenicolus (part).—Lasserre and Erséus, 1976:45 1-452. 289 Heterodrilus minisetosus Erséus, 198 1a: 113-114, fig. 2. New material examined. —FSBC I 31852- 31853, 2 whm spms from Sta 4 (10 May 1972).—Author’s collection: 2 whm spms from Sta 2, 14 from Sta 4; 1 whm spm from NE part of Gulf of Mexico, off Hernando Co., Florida, 28°36’N, 83°30’W, about 25 m, sediment unknown (27 Aug 1977; cour- tesy M. S. Ivester). Remarks.—Heterodrilus minisetosus, which was described on the basis of exten- sive material from Bermuda (type locality), Bahamas, Florida (Miami area), North Car- olina, and Virginia (Erséus 1981la), was named for the minute penial setae (one at each side of worm) present in most indi- viduals. Specimens from Hutchinson Island lack penial setae (as did most worms from North Carolina and Bahamas in the original material) but otherwise fit the description completely. Distribution and habitat. —Both coasts of Florida (new record from Gulf coast), North Carolina, Virginia, Bermuda, and Bahamas. Subtidal sands, 0.5-33 mm depth. Heterodrilus pentcheffi Erséus, 1981 Fig. 1 Clitellio arenicolus (part).—Giere, 1979: 304. Heterodrilus pentcheffi Erséus, 1981a:121- 123, fig. 12.—Erséus, 1984c:196. Heterodrilus sp.—Erséus, 1981a:123-124, fig. 14. New material examined. —FSBC I 31854, 3 whm spms from Sta 2 (9 May 1972).— Author’s collection: 7 whm spms from Sta 2, 2 from Sta 3, 1 from Sta 4, and 1 from Sta 5. Remarks. — Heterodrilus pentcheffi was originally described on the basis of material from Bermuda, New Jersey, and North Car- olina in the Northwest Atlantic, and the Ga- lapagos Islands in the East Pacific (Erseus 1981la; see also Erséus 1984c). The species is characterized by (1) trifid setae in seg- ments II-IX (sometimes —VIII, —X, —XI, or —XII), 2 per bundle, (2) thereafter bifid se- 50 um Fig. 1. Heterodrilus pentcheffi, postclitellar setae: A, From segment immediately posterior to clitellum; B-E, From various posterior segments; F, From pos- terior end of worm. tae, 1 per bundle, with more or less devel- oped lower tooth (cf. Fig. 1), and (3) straight, single-pointed, ectally wide, erect penial se- tae, 2(3) per bundle. Vasa deferentia are coiled in spirals, and atria are long and cy- lindrical, generally M-shaped, terminating in simple, somewhat bulbous pseudopenes. As illustrated in Fig. 1, bifid (postclitellar) setae of the new material are highly vari- able. In segments immediately posterior to the clitellar region, the lower tooth of the setae is conspicuous, almost as large as the upper one (Fig. 1A), but the lower tooth generally becomes reduced on more poste- rior setae (Figs. 1B—D), so much so that se- tae may appear single-pointed, at least from certain angles (Fig. 1E). At the posterior end of the worm, setae become smaller and their bifid nature generally becomes more pro- nounced again (Fig. 1F). In fact, the new material from Hutchin- son Island has bifid setae very similar to those of the form described as *“‘Heterodrilus sp.”’ from Miami (Florida) by Erséus (198la). This species was tentatively re- garded as separate from H. pentcheffi, large- ly also because of its small size. As the new material contains both large (with posterior setae up to 145 wm long) and small speci- mens (posterior setae only about 45 um PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON long), it is now concluded that “‘Heterodrilus sp.” is identical to H. pentcheffi, and that the Florida populations differ slightly from the more northern forms in terms of the appearance of the postclitellar setae. Ranges of body length and segment num- ber of the new material lie within those not- ed previously, if descriptions of H. pentch- effiand H. sp. are united (4.1—11.3 mm, 43- 88 segments). In one of the new individuals, the penial setae are only about 65 um long, which extends the length range of these setae for the species to 65-155 yum. The small “epidermal papilla,’ which perhaps should be termed an “epidermal pad” and which is located mid-ventrally be- tween the spermathecal pores in “Hetero- drilus sp.” (cf. Erséus 1981a:124), is present also in many of the original specimens of H. pentcheffi (Erséus, unpublished) as well as in most specimens from Hutchinson Is- land. Distribution and habitat.—East coast of Florida (new record), North Carolina, New Jersey, Bermuda, and Galapagos Islands. Largely coarse sands, 0.5—39 m depth. Heterodrilus occidentalis Erséus, 1981 Fig.2 Heterodrilus occidentalis Erséus, 1981a:121, fig. 11.—Davis, 1985:170. New material examined. —FSBC I 31855- 31856, 2 whm spms from Sta 2 (3 Jan and 9 May 1972, respectively).—Author’s col- lection: 3 whm spms from Sta 2, 6 from Sta 4. Remarks.—This species was originally described from shallow waters in North Carolina and off New Jersey (Erséus 198 1a), and was recently reported from Georges Bank off Massachusetts by Davis (1985). It is closely related to H. pentcheffi (genitalia and penial setae are virtually identical in the two species), but differs from that species in the appearance of the somatic setae. In the original material, trifid setae were found only in about the first five setigerous VOLUME 99, NUMBER 2 segments, followed by a few segments with bifid setae (cf. Erséus 1981a:figs. 11A—C); thereafter (segment X and backwards, ex- cept a few segments in posterior end), setae are sharply single-pointed (Erséus 198 la:fig. 11D). New specimens from Florida deviate slightly from this pattern. The lower tooth of the trifid setae gradually diminishes pos- teriorly, but this tooth can generally still be discerned in segments VIII and IX (Fig. 2A). Similarly, “‘single-pointed”’ setae, com- mencing in segment X, initially are actually somewhat bifid; a faint lower tooth can be seen from certain angles (Fig. 2B). Further back, however, most setae are truly single- pointed (Fig. 2C). Distribution and habitat.—East coast of Florida (new record), North Carolina, New Jersey, and Massachusetts. Coarse sand and gravel, 4.5-138 m depth. Heterodrilus hispidus, new species Fig. 3 Holotype. —-USNM 98130, whm spm from Sta 2 (15 Sep 1971). Paratypes. —FSBC I 31857, 2 whm spms from Sta 4 (14 Mar 1972). Other material examined. — Author’s col- lection: 5 whm spms from type locality (Sta Dys Description. —Length (2 complete spms) 13.6-14.8 mm, 82-86 segments; third spm only 4.5 mm long, about 38 segments, but its posterior end regenerating. Width at XI, 0.29-0.41 mm. Clitellum extending over %X—XII. Setae 2 per bundle in II-IX, 1 per “bundle”’ thereafter. Anterior setae (from II to IX) bifid, with teeth of approximately same length and both oriented along long axis of seta (Fig. 3A). Bifids 60-100 um long, 2.5—5 wm thick. Posterior setae (from X to end of worm) stout, sharply single-pointed, and with curved inner end; dorsal ones (Fig. 3C) much larger (95-135 um long, 7-9.5 um thick) than ventral ones (Fig. 3B; 60-95 wm long, 3.5-6 wm thick). Penial setae (Figs. 3D, E, ps) 2 per bundle (bundle ‘“‘doubled”’ 50 um Fig. 2. Heterodrilus occidentalis, setae: A, From segment VIII; B, From segment XII; C, From posterior segment. at one side in one aberrant spm), slender and slightly curved, single-pointed, with in- ner ends generally somewhat tilted over to posterior. Penials 125-135 um long, 4.5—5 um thick. Male pores paired, slightly ventral to lines of ventral somatic setae, posteriorly in XI. Spermathecal pores paired, slightly ventral to lines of ventral setae, anteriorly in X. Pharyngeal glands very poorly developed (absent?). Granulated coelomocytes numer- ous. Male genitalia (Fig. 3E) paired. Vas def- erens 9-12 wm wide, very long and tightly coiled in spiral. Atrium somewhat C-shaped, about 300 um long, 25—35 wm wide entally, narrower ectally, with thin outer lining and densely granulated inner epithelium; lobes of prostate glands attached to ventral sur- face of atrium. Ectal part of atrium non- granulated, tapering, opening to exterior through simple pore (possibly a very simple pseudopenis). Spermathecae (Fig. 3E, s) variable in shape and extension, with slen- der ducts and large ampullae, latter with large mass of sperm in irregular lumen and with large granules of secretion in wall. Remarks. —Most species of Heterodrilus have trifid setae in at least some anterior segments; only H. subtilis (Pierantoni, 1917) 292 A B f D 50 um | (A-D) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Heterodrilus hispidus, n. sp.: A, Anterior bifid seta; B, Posterior ventral seta; C, Posterior dorsal seta; D, Penial setae; E, Lateral view of spermatheca and male genitalia in segments X—XI. and this new species lack such setae alto- gether. Heterodrilus hispidus is distin- guished from the other Mediterranean and poorly known form by the consistently sin- gle-pointed tips of its setae from segment X to the posterior end of the worm (penial setae absent in H. subtilis). The great difference in size between the dorsal and ventral postclitellar setae is a striking feature of H. hispidus; a similar case has not been noted for any other species of the genus, and it may thus prove to be a reliable specific character. In most species of the genus, penial setae are either erect or with inner ends obliquely pointing toward the anterior, but in H. his- pidus and the aberrant H. perkinsi, n. sp., described below, inner ends of penial setae tend to be directed toward the posterior. Etymology. —The specific name hispidus is Latin for “shaggy, rough” and alludes to the appearance of the worm with regard to the very large setae. Distribution and habitat.—Known only from off Hutchinson Island, east coast of Florida. Shell hash, 10-12 m depth. Heterodrilus bulbiporus Erséus, 1981 Fig. 4 Heterodrilus bulbiporus Erséus, 1981a:123, fig. 13. New material examined. —FSBC I 31858, 2 whm spms from Sta 2 (1 Mar 1972).— Author’s collection: 2 whm spms from Sta 1 (but see note under “Material and meth- ods” above), 18 from Sta 2, 3 from Sta 4, and 21 from Sta 5; 1 whm spm from NE part of the Gulf of Mexico, off Clearwater, Florida, 27°55'N, 83°28’W, 20 m, sediment unknown (26 Aug 1977; courtesy M. S. Ivester). Redescription (based on the new materi- al).— Length (5 worms measured) 6.5—11.3 mm, 48-65 segments (original material: 5— 12 mm, 38-58 segments). Width at XI, 0.31- 0.56 mm. Clitellum extending over 2X—- XII. Setae 2 per bundle in II-IX, 1 per “bun- dle’’ thereafter. Anterior setae (in IJ-IX) tri- fid (Fig. 4A), with upper and middle teeth pointed and basally narrow, lower tooth ba- sally much wider. Trifids 75-125 um long, 4.5-—7 um thick (original material: 40-115 VOLUME 99, NUMBER 2 | | 50 um 6 Somer era 293 100 pum Fig. 4. Heterodrilus bulbiporus: A, Anterior trifid seta; B, Posterior seta; C, Penial setae; D, Spermatheca in segment X; E, Lateral view of male genitalia in segment XI. um, 3-5 wm). Posterior setae (from X and backwards) bifid (Fig. 4B), with upper tooth slightly thinner than lower, both teeth ba- sally wide. Bifids 65-140 um long, 4.5—9 um thick (original material: 50-115 wm, 4—7 pm). Penial setae (Figs. 4C, E, ps) 2 per bundle, erect, single-pointed, 80—95 um long (original material: 95-125 wm), about 7 wm thick at middle, 12—13 wm wide at ental end. Male pores paired in line with ventral setae, posteriorly in XI. Spermathecal pores paired, slightly ventral to lateral lines, an- teriorly in X. Pharyngeal glands in IV—V, but poorly developed. Granulated coelomocytes nu- merous. Male genitalia (Fig. 4E) paired. Vas deferens 16-21 um wide (original material: 7-13 wm), long and coiled. Atrium cylin- drical, slender, about 0.8 mm long (original material: 0.4—1.0 mm), 25—40 wm wide, with 2—4 wm thick outer muscular lining (original material: up to 5 um thick), and granulated inner epithelium along most of its length; narrow ectal portion of atrium not granu- lated, but containing numerous nuclei. Atrium ectally terminating in bulbous cop- ulatory apparatus, consisting of inner wid- ened part (most ectal part of atrium) and outer, narrow, tapering penis, latter en- closed in copulatory sac (whole complex probably somewhat eversible; cf. Erséus 198la:fig. 13C). Small lobes of prostate glands present along granulated part of atrium. Spermathecae (Fig. 4D) consisting of short but broad ducts and large, more or less oval ampullae, latter with thick walls containing large granules of secretion and irregular lumen containing large bundles of sperm. Remarks. —Heterodrilus bulbiporus was originally described from Bermuda, New Jersey, and North Carolina. Some dimen- sional differences between the old and the new material are indicated in the redescrip- 294 tion above. In addition, the new material deviates slightly from the original descrip- tion with regard to the morphology of the male terminalia; the copulatory organ of H. bulbiporus was previously perceived as a pseudopenis (Erséus 198 1a), but in the new material it rather appears more like a proper pendant penis and should be described as such. This penis discriminates H. bulbipo- rus from all the other species of the genus. Distribution and habitat. — Both coasts of Florida (new records), North Carolina, New Jersey, and Bermuda. Subtidal sands, 6.5-— 130 m depth. Heterodrilus perkinsi, new species Fig. 5 Holotype. —USNM 98131, whm spm from Sta 4 (4 Jan 1972). Paratypes. —FSBC I 31860-31861, 1 sec (10 May 1972) and 2 whm spms (4 Jan 1972) from Sta 4 (type locality). Other material examined. — Author’s col- lection: 2 sec and 16 whm spms from Sta 2; and 1 sec and 24 whm spms from Sta 4 (type locality). Description. —Length (5 complete spms measured) (5.7)8.9-11.2 mm, (42)58-72 segments; smallest spm still growing pos- teriorly. Width at XI 0.41-0.50 mm. Cli- tellum extending over '2X—XII. Somatic se- tae 2 per bundle in IIJ-IX (—VIII in single spm with genitalia shifted one segment for- ward), 1 per ““‘bundle’’ thereafter. Anterior setae (from II to IX) trifid (Fig. SA), with upper tooth smaller than other two, middle tooth longer than upper and lower teeth. Trifids 65—110 um long, 4—7 um thick. Pos- terior setae (from X to end of worm) bifid (Fig. 5B), with lower tooth slightly larger than upper, both teeth with broad bases. Bifids 60-120 um long, 4.5-—9.5 wm thick. Penial setae (Figs. SC, D, ps) small and straight, (1)2 per bundle, often retracted into copulatory sac and with inner ends generally directed obliquely toward posterior; ectal ends single-pointed and hooked. Penials 60— PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 85 um long, 4.5—5.5 um thick. Male pores paired, located close together posteriorly in mid-ventral bursa in XI (X in aberrant spm); bursa laterally delimited by two flap-like pa- pillae (Fig. 5D, fp). Spermathecal pores paired in line with ventral setae anteriorly in X (IX in aberrant spm). Pharyngeal glands in IV—V(VI). Granu- lated coelomocytes present. Male genitalia (Fig. 5D) paired. Vas deferens remarkably short and wide, with very thin and densely ciliated wall; vas deferens about 150-175 um long, 34—45 wm wide, abruptly narrow- ing at entrance into atrium. Atrium long and slender, basically cylindrical but with some- what varying width, coiled and extending through 2-3 segments. Atrium about 1.4— 3.2 mm long (exact length difficult to estab- lish), maximally 50-70 wm wide (many parts narrower), with 9-15 wm thick outer layer of muscles, and poorly granulated inner ep- ithelium; cilia not observed. Lobes of pros- tate glands scattered along most of atrium. Atrium tapering ectally, terminating in somewhat pointed, conical penis. Penis 45— 80 um long, basally 23-37 um wide, with thin cuticular lining, located in hollow inner part of deep copulatory sac. Spermathecae (Fig. 5D, s) with elongate, thick-walled ducts, and ampullae of variable shape, latter con- taining at least one large bundle of sperm in postcopulatory spms; ampullae often dis- located into IX. Remarks.—The most striking feature of Heterodrilus perkinsi is its extremely long and muscular atria, which discriminate the species from all its congeners. Noteworthy, and somewhat paradoxical, is that the vasa deferentia are among the shortest in the ge- nus, measuring only about one-tenth or less of the atrial length. Etymology. —This species is named for Mr. Thomas H. Perkins, who placed the Hutchinson Island Oligochaeta at my dis- posal and provided most valuable infor- mation, moral support, and assistance dur- ing the preparation of this paper. VOLUME 99, NUMBER 2 a 50 ym 295 Fig. 5. Heterodrilus perkinsi, n. sp.: A, Anterior trifid seta; B, Posterior seta (from segment immediately posterior to clitellum); C, Penial setae; D, Somewhat lateral view of spermatheca and male genitalia in segments X—XIII (note that flap-like papillae and penial setae of both sides of worm are shown; the spermatheca and the vas deferens depicted are not from the same side as the atrium in the specimen used). Distribution and habitat.—Known only from off Hutchinson Island, east coast of Florida. Shell hash, 10-12 m depth. Subfamily Phallodrilinae Phallodrilus Pierantoni, 1902 Phallodrilus sabulosus Erséus, 1979 Fig. 6 Phallodrilus sabulosus Erséus, 1979b:188- 189, figs. 1-2.—Erséus and Loden, 1981: 820-821, fig. 1A.—Erséus, 1984d:813. New material examined. —FSBC I 31862, 3 whm spms from Sta 4 (14 Mar 1972).— Author’s collection: 2 whm spms from Sta 2, 9 from Sta 4, and 1 from Sta 5. Remarks.—Phallodrilus sabulosus was first described from a coral reef off Miami (Erséus 1979b), but as the two type speci- mens were both precopulatory, the mature spermathecae of the species were not de- scribed until new material was recovered from off Hutchinson Island by Erséus and Loden (1981). Recently P. sabulosus was found off New Jersey, further north along the North American east coast (Erséus 1984d). The new material from Hutchinson Is- land is highly variable in terms of overall size: length 2.8-9.0 mm, segments 22-63 (previously studied individuals 5.5—6.1 mm long, 44-61 segments). According to the original description, bi- fid setae either have an upper tooth that is slightly longer than the lower one (anterior bundles) or the two teeth are equally long (posterior bundles). In the present material, however, many posteriormost setae have a lower tooth which is clearly longer than the upper one (Fig. 6A); in fact, the lower tooth appears prolonged as compared to that of anterior setae. The most striking feature of this species is the possession of both penial and sper- mathecal setae. Penial setae (illustrated by Erséus 1979b:figs. 1-2) are stout, somewhat 296 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 25 um Fig. 6. Phallodrilus sabulosus: A, Seta from one of posteriormost segments; B, Spermathecal seta with associated glands. spoon-shaped, always one per “bundle,” and located near the atrial openings in segment X. These setae are 60-80 um long and up to 7 um wide in the present material. Spermathecal setae (Fig. 6B, ss) are more slender than penial setae, ectally deeply forked (now established; cf. statement by Erséus 1979b:189), one or two per bundle and located either posterior to spermathecal pores in segment X (in 13 of the 16 present worms), or anterior to these pores, poste- riorly in segment IX (in the remaining three worms). The middle part of each seta is en- closed in a glandular body, which bears a few lobes of external glands (Fig. 6B, gs), which are somewhat reminiscent of prostate glands; these external glands were not noted in the original description. Spermathecal se- tae are 60-100 um long. Male efferent ducts of the new material conform to the original account, although the length range of the atrium is now ex- tended to 60-140 um. Spermathecae are variable in size and shape but basically con- form to the description provided by Erséus and Loden (1981:fig. 1); they are very slen- der, consisting of (1) a long duct, which is ectally distended and often contains sperm, and (2) an oval, thin-walled ampulla, gen- erally with very dense mass of sperm. Distribution and habitat.—East coast of the U.S.A., from Florida to New Jersey. Largely coarse sands, 3—15.5 m depth. Phallodrilus biprostatus (Baker and Erséus, 1979) Peosidrilus biprostatus Baker and Erséus, 1979:506-508, figs. 1-2.—Erséus and Loden, 1981:819-820. Phallodrilus biprostatus. —Erséus, 819-820.— Davis, 1985:table 1. New material examined. —FSBC I 31863- 31864, 3 whm spms from Sta 2.—Author’s collection: 1 sec and 12 whm spms from Sta 2, 2 whm spms from Sta 3, 2 sec and 19 whm spms from Sta 4, and 1 whm spm from Sta 5. Remarks. —This species, which occurred at several stations in the present material, 1984d: VOLUME 99, NUMBER 2 was originally described as a member of a monotypic genus Peosidrilus Baker and Er- seus, 1979, from New Jersey and North Carolina. It was reported from Hutchinson Island by Erséus and Loden (1981), and lat- er also from Georges Bank off Massachu- setts (Erséeus 1984d; Davis 1985). The species was recently transferred to Phallo- drilus (Erséus 1984d); its voluminous penes are now regarded as an elaboration of, and thus homologous to, the smaller copulatory structures (pseudopenes and true penes) possessed by several other Phallodrilus forms. One of the new specimens consists of 70 segments (previously studied worms: 44—62 segments). Distribution and habitat.—East coast of U.S.A., from Florida to Massachusetts. Largely coarse sands, 5.5—73 m depth. Phallodrilus acochlearis (Erséus and Loden, 1981), new combination Fig. 7 Adelodrilus acochlearis Erséus and Loden, 1981:821-823, figs. 1B-C, 2.—Erséus, 1983b:77-78. New material examined. —FSBC I 31865, 2 whm spms from Sta 4 (4 Jan 1972).— Author’s collection: 7 whm spms from Sta 4. Redescription (modified after Erseus and Loden 1981).—Length 2.5—3.9 mm, 27-39 segments. Width at XI, 0.15—0.25 mm. Cli- tellum extending over '12X—XII. Somatic se- tae bifid, 3-5 per bundle anteriorly, (2)3- 4(5) per bundle in postclitellar segments. Anterior dorsal setae and all ventral setae with upper tooth shorter and much thinner than lower. Several setae in dorsal bundles of most posterior segments modified with lower tooth extremely long, curved down- wards and parallel to setal shaft (Erséus and Loden’s fig. 1C). Bifids 42-62 wm long, 1.5- 2.5 um thick. Penial setae straight, 30-50 um long, 0.5—1 wm thick at middle, each with ectal “‘club’”’ bearing apical hook (Fig. 297 7A). Penial setae about 8—14 per bundle (ex- act number difficult to establish as setae sit- uated very near each other within bundle), with ectal ends protruding into small cop- ulatory sac (sac everted in Fig. 7B) imme- diately posterior to atrial opening. Male pores paired in line with ventral somatic setae posteriorly in XI. Spermathecal pores paired in lateral lines in anteriormost part of X. Pharyngeal glands inconspicuous, but ex- tending as far back as VII or VIII. Male genitalia (Fig. 7B) paired. Vas deferens very wide and heavily muscular, longer than atrium, often convoluted (as shown in Er- seus and Loden’s fig. 2); cilia observed along inner epithelium of vas deferens in well- preserved spms, but sperm not stored in lumen (cf. Remarks). Vas deferens entering atrium sub-apically together with anterior prostate gland. Atrium 58-82 um long, 35— 43 um wide, divided into two histologically different portions; ental portion with gran- ulated and ciliated inner epithelium, ectal portion neither granulated nor ciliated but eversible to form bulbous pseudopenis (everted in Fig. 7B; cf. also Erséus and Lod- en’s fig. 2); muscular lining of atrium thin. Prostate glands moderately large, both sub- apical on atrium, with attachments more or less opposite to each other. Spermathecae (Fig. 7B, s) consisting of short ducts and slender ampullae, latter 90-180 um long, maximally 33—44 um wide and variable in shape, with sperm in random masses. Remarks.—This species was originally described on the basis of specimens from a site very near that of the present material. One additional individual was subsequently reported from shallow inshore water of North Carolina (Erséus 1983b). The new material has necessitated a re- description of the species, as the true ap- pearance of its vasa deferentia was not pre- viously recognized; the vasa deferentia are ciliated, but they do not store sperm as ini- tially conceived, and thus the species does not qualify for inclusion in the genus Ade- 298 A B Ss 100 um Fig. 7. genitalia in segments X—XI. lodrilus, where it was originally placed. In- stead it should be assigned to Phallodrilus as defined by Erséus (19844). The revised description of P. acochlearis shows that the species is closely related to another North American east coast form, P. boeschi Erséus, 1984, which has posterior dorsal setae (with prolonged lower teeth; cf. Erséus 1984d:fig. 2B), penial setae, atria, and spermathecae that are very similar to those of P. acochlearis. However, the latter species is easily distinguished from P. boeschi by its conspicuously muscular vasa deferentia, its more elaborate pseudopenes and even longer lower teeth of the posterior dorsal setae (cf. Erséus and Loden 1981:fig. 1C). Distribution and habitat.—East coast of Florida and North Carolina. Very coarse sands, 5.5—11 m depth. Phallodrilus hirsutus, new species Fig. 8 Holotype. —USNM 98132, whm spm from Sta 2 (5 Mar 1973). Paratype. —FSBC I 31866, 1 whm spm from Sta 5 (5 Jan 1973). Description. —Length of holotype (para- type not complete) 16.1 mm, 86 segments. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON at ss PISS WVSSZs Wy ©, Lk, Phallodrilus acochlearis, n. comb.: A, Tip of penial seta; B, Lateral view of spermatheca and male Width at XI, 0.44 mm. Clitellum extending over '2X—XII. Somatic setae 2 per bundle in IJ-IX(X), 1 per “‘bundle”’ thereafter. An- terior setae (Fig. 8A) bifid, with upper tooth reduced; these setae 70-140 um long, 3.5— 10 wm thick. Setae from about X and back- wards straight and single-pointed (occasion- ally bifid, similar to anterior setae); these setae dorsally (Fig. 8A) 140-165 um long, 12-13 wm thick, ventrally (Fig. 8C) 85-95 um long, about 7 wm thick. Ventral setae of XI modified into penial setae (Figs. 8D, F, ps), 4-6 per bundle, more or less straight, ectally single-pointed and hooked; in ho- lotype (Fig. 8D), tips of penial setae oblique- ly pointing toward posterior; in paratype (Fig. 8F), penial setae and male genitalia somewhat distorted and dislocated. Male pores paired in line with ventral setae in XI. Spermathecal pores paired in lateral lines, anteriorly in X. Pharyngeal glands in IV—VI. Male geni- talia paired. Vas deferens 16-19 um wide, longer than atrium, entering apical end of latter. Atrium (Fig. 8F, a) somewhat spin- dle-shaped and curved, about 105 um long, about 40 um wide, with very thin outer lin- ing, and thick, granulated outer epithelium. Prostate glands (Fig. 8F, pr 1-2) very large VOLUME 99, NUMBER 2 299 Fig. 8. Phallodrilus hirsutus, n. sp.: A, Anterior setae; B, Posterior dorsal seta; C, Posterior ventral seta; D, Penial setae; E, Spermatheca; F, Male genitalia. and lobed, anterior one subapical on atrium, posterior one attaching to posterior face of atrium, at some distance from atrium open- ing. Copulatory sac apparently present, but details not clear in available spms. Sper- mathecae (Fig. 8E) elongate, consisting of indistinct ducts and long, thin-walled am- pullae, latter somewhat constricted at mid- dle, and containing large bundle of sperm. Remarks.—This very large species of Phallodrilus is distinguished from all other members of the genus by its very large and modified setae. In particular, the postclitel- lar dorsal (single-pointed) setae (Fig. 8B) are very conspicuous and make the species su- perficially very similar to Heterodrilus his- pidus described above, and also the NW European species Bathydrilus rarisetis (Er- séus, 1975) (cf. Erséus 1975; 1979a:fig. 7). The latter form also inhabits very coarse sand and gravel, and it appears likely that the convergent evolution of very stiff and large setae in different groups of marine Tubificidae (including many other species of Heterodrilus) is explained by their func- tional advantage in such coarse substrates. Etymology.—The specific name hirsutus is Latin for “shaggy, bristly, rough” and al- ludes to the conspicuous setation. Distribution and habitat.—Known only from off Hutchinson Island, east coast of Florida. Coarse, clean or somewhat muddy, sands, 10-11 m depth. Adelodrilus Cook, 1969 Adelodrilus magnithecatus Erséus, 1979 Adelodrilus magnithecatus Erséus, 1979d: 419-421, fig. 1. New material examined. —FSBC I 31867, 1 whm spm from Sta 2 (3 Nov 1971).— Author’s collection: 1 whm spm from Sta 2. Remarks. —Adelodrilus magnithecatus, 300 originally described from Bermuda and in- shore waters of North Carolina, is distin- guished from its congeners by its large sper- mathecae. Its penial setae are of two types as is typical for the genus; within each bun- dle are one “spoon-shaped” giant seta and a number of smaller, straight setae, the latter clubbed and each with an apical hook. The new specimens from Florida are larg- er (6.2—6.5 mm, 55-57 segments) than the original material (2.9-5.1 mm, 26-47 seg- ments) but otherwise fit the description well. Distribution and habitat.—East coast of Florida (new record), North Carolina, and Bermuda. Coarse sands, 4.5—17 m depth. Bathydrilus Cook, 1970 Bathydrilus ingens, new species Fig. 9B—C, E-H Bathydrilus sp.—Erséus, 1979a:146. Holotype. —USNM 98133, whm spm from Sta 2 (2 Nov 1972). Paratype. —FSBC I 31868, 1 whm spm from type locality (2 Nov 1972). Other material examined. — Author’s col- lection: 1 whm spm from Sta 4; 1 whm spm from NE Gulf of Mexico, 1 mile off Withlacoochee River mouth, Florida, 28°58'48"N, 82°48'27” W, 3—4 m, shell hash (Nov 1984; courtesy M. R. Milligan); 2 whm spms from between Carrot Island and Mid- dle Marsh, near Beaufort, North Carolina, 34°42'06’N, 76°37'13”W, 5 m, shells and shell gravel with some coarse sand (19 Oct 1977; cf. Erséus 1979a). Description. —Length (2 complete spms) 25.6-—27.0 mm, from about 127 (posterior end not fully differentiated) to 139 seg- ments. Width at XI, 0.26-0.42 mm. Epi- dermal glands (Fig. 9B) as continuous dorsal band along most of body, starting few seg- ments anterior to clitellum. Clitellum ex- tending over 'sX—XII. Somatic setae of 2 types. Anterior and some posterior ones bi- fid (Fig. 9E), with upper tooth smaller than lower; many posterior ones (particularly those in dorsal bundles; Fig. 9F), however, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON single-pointed and somewhat enlarged. So- matic setae 35-85 um long, 2.5—6 wm thick at node, 2—3(4) per bundle anteriorly, 2 per bundle in postclitellar segments. Ventral se- tae of XI (XII in aberrant spm) modified into penial setae (Figs. 9C, H, ps), 3 per bundle; penial setae straight with somewhat curved, single-pointed tips, latter directed towards mid-ventral line, exiting at sum- mits of 2 large papillae; tips of bundles often crossing each other (as in Fig. 9C). Penial setae 150-200 um long (only about 80 um long, but probably not fully grown, in spm f.om Gulf of Mexico), 6—9 um thick. Male pores paired in line with ventral somatic setae in posterior part of XI (XII in aberrant spm). Spermathecal pores paired in lateral lines in anteriormost part of X (XI in aber- rant spm). Pharyngeal glands extending into VII(VIII-IX). Male genitalia (Fig. 9H) paired. Vas deferens about 7 um wide (full length not seen in available material), en- tering anterior face of ectal-to-middle part of atrium together with anterior prostate gland. Atrium with ental end obliquely di- rected towards posterior, often extending into next segment. Atrium elongate, spindle- shaped, about 220-270 um long (difficult to measure), 30—90 um wide, with about 2 um thick outer lining of muscles, and granulated and ciliated inner epithelium. Atrium open- ing at outer end of deep blind sac, atrial opening hidden behind lateral fold of body wall; whole complex forming elaborate pseudopenis. Posterior prostate gland large, attached to apex of atrium. Spermathecae (Fig. 9G) consisting of short, triangular ducts and large elongate, somewhat bilobed am- pullae; former opening into middle of latter. Sperm as rhomboid or elongate spermato- zeugmata, as bundles, or as loose masses, in ampullae of postcopulatory spms. Remarks.—One specimen from Hutch- inson Island has its sexual organs shifted backwards to occupy segments XI—XII in- stead of X—XI. Bathydrilus ingens is a very large species, VOLUME 99, NUMBER 2 301 mp Fig. 9. Bathydrilus longus (A, D) and B. ingens, n. sp. (B, C, E-H): A and B, Patterns of dorsal epidermal glands (shown for a few successive segments in postclitellar part of body); C, Ventral view of penial setae and male pores in segment XI (ingens); D, Posterior dorsal seta (/Jongus); E, Anterior seta (ingens); F, Posterior dorsal seta (ingens); G, Spermatheca (ingens); H, Lateral view of male genitalia in segment XI (ingens). a feature shared with the very similar B. longus Erséus, 1979, reported from deeper (70-165 m) stations off New Jersey (Erséus 1979a) and Massachusetts (Davis 1985). Both species have two or three very con- spicuous, straight penial setae per bundle. The new, southern form is distinguished from B. longus by four important charac- ters: (1) the dorsal epidermal glands form a continuous dorsal band along body except 302 for anteriormost segments (Fig. 9B; in B. longus these glands in rhomboid, segmen- tally arranged patches as shown in Fig. 9A); (2) the aberrant appearance (Fig. 9F) of many posterior setae (in B. Jongus posterior setae always have a small upper tooth; Fig. 9D); (3) the thin atrial musculature (only about 2 wm thick; about 5—7 um thick in B. Jon- gus); and (4) the possession of a pair of deep invaginations lateral to ectal parts of atria, and at the outer ends of which atria open (such blind sacs absent in B. /ongus). Etymology. —The species name ingens is Latin for ““huge, enormous.” Distribution and habitat.—North Caro- lina and both coasts of Florida. Coarse sands, 3-11 m depth. Bathydrilus formosus, new species Fig. 10 Holotype. -USNM 98134, whm spm from Carrie Bow Cay, Barrier reef of Belize, edge of Thalassia bed near north end of island, subtidal shallow water, mixed coral sand with some organic debris (6 Apr 1982; coll. H. R. Baker). Paratypes.—USNM 98135, 1 whm spm from type locality. USNM 98136, 1 whm spm from near reef crest at Carrie Bow Cay, Belize, about 0.5 m, coral sand and Hali- media debris in small pocket of bedrock coral (12 Apr 1982; H. R. Baker).—FSBC I 31869, 1 whm spm from Hutchinson Is- land; Sta 5 (5 Jan 1973). Other material examined. — Author’s col- lection: 4 whm spms from type locality; 2 from Tobacco Reef, Barrier reef of Belize, back reef of South Water Cay (about 0.5 km N of the cay), 1 m, small patch of coral sand (9 Apr 1982; H. R. Baker).—1 from Carrie Bow Cay, Belize, Thalassia bed very near Research Station, barely subtidal, coral sand with some organic debris (12 Apr 1982; H. R. Baker).—1 whm spm from NE Gulf of Mexico, off Clearwater, Florida, 27°55'N, 83°28'W, 20 m, sediment unknown (26 Aug 1977; courtesy M. S. Ivester). Description.—Length (only 3 complete spms) 15.9-18.0 mm, 84-108 segments. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Width at XI, 0.27-0.40 mm. Epidermal glands as continuous dorsal band in many posterior segments. Clitellum extending over ’3X—XIIT. Somatic setae (Fig. 10A) bifid, sig- moid, with upper tooth thinner and shorter than lower; 45-55 um long, 3—3.5 wm thick at node, 3 per bundle in anteriormost seg- ments, 2 per bundle from few segments an- terior to clitellum and along rest of worm. Ventral setae of XI modified into penial se- tae (Figs. 10D, C, E, ps), 2(3) per bundle; penial setae strongly curved with tips di- rected towards and located near mid-ven- tral line, exiting at summit of oval epider- mal pad mid-ventrally in middle of XI (cf. Fig. 10C, ep). Penial setae 100-155 um long, 4.5-7 um thick at swelling (modified node) near ectal end, with bifid tips; upper tooth shorter and much thinner than lower, but not always visible (hidden behind lower tooth; cf. 1 seta in Fig. 10D). Male pores paired in line with ventral somatic setae in posterior part of XI, on 2 bulbous protu- berances (= pseudopenes; cf. below and Fig. 10C, pp). Spermathecal pores paired in lat- eral lines in anteriormost X. Pharyngeal glands extending into VIII. Male genitalia (Fig. 10E) paired. Vas def- erens 9-12 wm wide, longer than atrium, entering middle of anterior face of latter to- gether with stalk of anterior prostate gland. Atrium with ental end obliquely directed towards posterior, sometimes extending into XII (indicated in Fig. 10C). Atrium elon- gate, pear-shaped or spindle-shaped, 150- 235 um long, 50-80 wm wide, with 1-2.5 um thick outer lining of muscles and ciliated inner epithelium, latter densely granulated except for most ectal part. Atrium termi- nating ectally into bulbous and muscular pseudopenis, latter more or less protruded in all spms available. Prostate glands mod- erately developed, anterior one attached to middle of atrium, ectal one to apex of atrium. Spermathecae (Fig. 10B, s) consisting of short, triangular ducts and large, extremely thin-walled ampullae; latter containing roundish, rhomboid or elongate spermato- zeugmata in postcopulatory spms. VOLUME 99, NUMBER 2 20 pm 50 um 303 Fig. 10. Bathydrilus formosus, n. sp.: A, Somatic seta; B, Spermathecae in segment X; C, Ventral view of penial setae and male pores in segment XI; D, Penial setae; E, Lateral view of male genitalia in segment XI. Remarks. —This species is distinguished from the closely related B. longus and B. ingens, n. sp., by its externally more con- spicuous and bulbous pseudopenes and by its curved penial setae. In addition, B. for- mosus is the only species within the genus known to possess bifid penial setae; in all congeners, penial setae (when present) are described as single-pointed. [Note, how- ever, that Baker (1983) reported that in a single specimen of the NE Pacific B. litoreus Baker, 1983, penial setae in one bundle were bifid.] Etymology. —The species name formosus is Latin for “finely formed, beautiful’’; here primarily referring to the penial setae, which in the author’s opinion are esthetically ap- pealing. Distribution and habitat. —Belize and both coasts of Florida. Subtidal sands, 0.5— 11 m depth. Bathydrilus macroprostatus, new species Fig. 11 Holotype. —USNM 98137, whm spm from Sta 2 (15 Sep 1971). Paratype. —FSBC I 31870, 1 whm spm from type locality (3 Jan 1972). Description. —Length more than 5.7 mm, more than 25 segments (no complete spm 304 Fig. 11. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bathydrilus macroprostatus, n. sp.: A, Somatic setae; B, Penial setae; C, Somewhat lateral view of atrium (of one side only), penial setae and spermathecae in segments IX—X]I (note that this location is an anomaly, characterizing the holotype only; see text). available). Width at male pores, 0.37—0.47 mm. Epidermal glands not observed. Cli- tellum extending over '2X—XII in paratype, not developed in holotype. Somatic setae (Fig. 11A) bifid, slender in most anterior segments, only 50-55 um long, about 2.5 um thick at node, thereafter stouter and larger, 70-85 wm long, 4—6 um thick at node; teeth variable, but upper tooth always smaller than lower. Bifids 2—3(4) per bundle anteriorly, 2 per bundle in postclitellar seg- ments. Ventral setae of XI (X in holotype) modified into penial setae (Fig. 11B), 2 per bundle, with tips directed towards and lo- cated near mid-ventral line (cf. Fig. 11C). Penial setae straight to somewhat curved, 70— 90 um long, 4—5 wm thick at swelling (mod- ified node) near ectal end, with flat, hooked tips. Spermathecal and male pores paired in line with ventral somatic setae, former in most anterior part of X (IX in holotype), latter in posterior part of XI (X in holotype). Pharyngeal glands extending into VIII. Male genitalia (Fig. 11C) paired. Vas def- erens thin, 8—9 wm wide, longer than atrium, but whole length not visible in available ma- terial. Vas deferens entering anterior face of ental part of atrium together with anterior prostate gland (indicated by ciliation inside atrium), but exact position of entrance not ascertained. Atrium erect or somewhat tilt- ed over to posterior, totally (including peni- al organ) 230-245 wm long, separated into two main parts by constriction: ental part 115-140 um wide with broad, somewhat rounded, truncate apex, ectal part 70-85 um VOLUME 99, NUMBER 2 wide, somewhat spindle-shaped and ter- minating in elongate (permanently pen- dant?) penis protruding from male pore. Pe- nis lacking cuticular sheath. Lumen of vas deferens continuous with ciliated narrow canal running beneath apex of ental part of atrium, perpendicular to main lumen of atrium. Inner epithelium of ental part of atrium with (paratype) or without (holo- type; Fig. 11C) granulation (=different de- grees of maturation), that of ectal part of atrium granulated in both spms. Muscular lining of atrium 2.5—4 um thick. Prostate glands attached by thick stalks to anterior and posterior faces of ental part of atrium, almost opposite to each other, anterior prostate somewhat more ectal than poste- rior one. Microtubules of prostatic cells con- spicuous at entrances into atrium. Prostate glands large and lobed, posterior gland ex- tending into XII (XI in holotype). Sper- mathecae (Fig. 11C, s) not fully developed in available material (spms pre-copulatory), but nevertheless very large, consisting of short, muscular ducts and elongate, folded ampullae, latter with glandular, thick inner epithelium. Remarks. — The holotype has its genitalia in segments IX—X and not in X—XI as is normal for a tubificid; this is regarded as an anomaly of this particular specimen. The atrial morphology (bipartite, erect atrium terminating in an elongate penis) and the very large prostate glands and sperma- thecae distinguish B. macroprostatus from other species of the genus. Etymology. —This species has very large prostate glands; hence the name macro- prostatus. Distribution and habitat.—Known only from off Hutchinson Island, east coast of Florida. Shell hash, 10-11 m depth. Coralliodrilus Erséus, 1979 Coralliodrilus corpulentus, new species Fig. 12 Holotype. —USNM 98138, whm spm from Sta 4 (4 Jan 1972). 305 Paratypes. —FSBC I 31871-31872, 2 whm spms, from Sta 2 (4 Jan 1973) and 4 (5 Jan 1973), respectively. Description. —Length 5.4—5.9 mm, 56-63 segments. Width at XI, 0.47—0.53 mm. Pro- stomium rounded triangular, not very large. Fixed spms very stout; segments short. Sec- ondary annulation present, several annuli per segment. Clitellum over /2X—XII (in one paratype '2I1X—X]I). Somatic setae bifid, of general phallodriline type in most bundles, with upper tooth slightly smaller than lower and with subdental ligament (Fig. 124A). Posterior dorsal setae (Fig. 12B) modified, with minute upper tooth (sometimes not visible; absent?) and prolonged, sharply pointed lower tooth, and without or with very inconspicuous subdental ligament. Anterior setae 55-70 um long, 1.5-3 wm thick, (2)3 per bundle. Posterior setae (from few segments posterior to clitellum and backwards) up to 80 um long, 3—5 um thick. Ventral setae of XI (X in one paratype) modified into penial bundles, each contain- ing about 12 straight, ectally strongly hooked single-pointed setae (Fig. 12C), 80-95 um long, about 2.5 wm thick (shown at one side only in Fig. 12D). Male pores paired in line with ventral setae, posteriorly in XI (X in one paratype). Spermathecal pores paired in lateral lines, in most anterior part of X (IX in One paratype). Pharyngeal glands poorly developed in (I1DIV—VI. Male genitalia (Fig. 12D) paired. Vas deferens not observed in complete length, but appearing short; ectally about 20 wm wide with strong circular muscles. Atrium spindle-shaped, somewhat sigmoid, 210-290 um long, 55-80 um wide at mid- dle, with 3.5—12 um thick outer layer of muscles, in which fibers arranged in some- what spiral pattern; inner epithelium of atrium ciliated and for most parts granu- lated. Atrium tapering ectally into short, narrow duct surrounded by very strong muscles; duct terminating in simple, minute male pore (only that of 1 side shown in Fig. 12D). Spermathecae (Fig. 12D, s) consisting 306 A B C 25 um PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON l 100 um | Fig. 12. Coralliodrilus corpulentus, n. sp.: A, Somatic seta; B, Posterior dorsal seta; C, Penial seta; D, Ventral view of spermathecae and male genitalia in segments X—XI (note that musculature is shown at one of the two male openings, penial setae at the other). of ducts about 80 um long, entally about 25 um wide, ectally strongly muscular and nar- rower, and oval, thin-walled ampullae, 130- 140 um long, about 75 wm wide; latter con- taining large bundle of sperm. Remarks. —In one paratype, the clitellum and the genitalia are shifted forward to oc- cupy segments IX—X, not X—XI which is the normal position for the Tubificidae. Coralliodrilus corpulentus is distin- guished from all its shallow-water congeners by its very thick atrial muscles, and is unique within the genus in having modified bifid setae in the posterior dorsal bundles. The atrium of the new species is somewhat sim- ilar to that of the South Atlantic deep-sea species C. /ongiductus Erséus, 1983 (Erséus 1983c), but the latter lacks the voluminous muscular mass around the ectal part of the atrial duct which characterizes C. corpulen- tus, and has only five to seven penial setae per bundle. It should be noted that the gutless species described as Coralliodrilus avisceralis by Erséus (1981b), which also has heavily mus- cular atria, is now regarded as a member of the genus Olavius Erséus, 1984 (subgenus Coralliodriloides Erséus, 1984; see Erséus 1984b). Etymology. —The species name corpulen- tus, which is Latin for “‘stout, corpulent,” refers to the shape of the fixed specimens. VOLUME 99, NUMBER 2 Distribution and habitat.—Known only from off Hutchinson Island, east coast of Florida. Shell hash, 10-11 m depth. Inanidrilus Erséus, 1979 Inanidrilus ernesti Erséus, 1984 Phallodrilus sp.—Erséus and Loden, 1981: 821. Inanidrilus ernesti Erséus, 1984b:251, fig. 14. New material examined. —FSBC I 31873, 2 whm spms from Sta 2 (9 May 1973).— Author’s collection: 5 whm spms from Sta 2, 5 from Sta 4. Remarks. —The gutless species [nanidri- lus ernesti was described (Erséus 1984b) on the basis of four specimens found in another collection of oligochaetes from off Hutch- inson Island, provided by Applied Biology, Inc., first to M. S. Loden (Louisiana State University) (cf. Erséus and Loden 1981), subsequently to the author. The new ma- terial conforms to the original description. Distribution and habitat.—Known only from off Hutchinson Island, east coast of Florida. Coarse sediments, 10—12 m depth. Inanidrilus vacivus Erséus, 1984 Inanidrilus vacivus Erséus, 1984b:249-250, fig. 12. Holotype. -USNM 96540, whm spm from Sta 2 (3 Jan 1972). Paratype.—FSBC I 31263, 1 whm spm from Sta 4 (6 Jul 1972). Remarks.—Inanidrilus vacivus is de- scribed elsewhere (Erséus 1984b) on the ba- sis of two specimens in the present material. Distribution and habitat.—Known only from off Hutchinson Island, east coast of Florida. Shell hash, about 10 m depth. Olavius Erséus, 1984 Olavius latus, new species Fig. 13 Holotype. -USNM 98139, whm spm from Sta 2 (7 Sep 1972). 307 Description.—Length 9.5 mm, 80 seg- ments. Width at XI, 0.40 mm. Body flat with elongate prostomium and with pygid- ium possessing long filiform caudal process (Fig. 13A). Secondary annulation indistinct (this may be a preservation artifact of the only spm at hand). Epidermal glands scat- tered over most of body surface (the fact that they are visible may be due to the pres- ervation method used and should not be regarded as a specific character). Clitellum well developed over /2X—XII. Somatic setae (Fig. 13B) bifid, but variable, with upper tooth thinner and shorter than lower, and with conspicuous subdental ligament; upper tooth not always visible (oblique view?). Bifids 3—4 per bundle anteriorly, 2—3 per bundle in postclitellar segments. Penial se- tae (Figs. 13C, D, ps) 5—6 per bundle, more or less parallel and somewhat spread out within bundle, about 35 wm long, entally about 2 um thick, ectally much thinner, sin- gle-pointed and curved. Male pores paired in line with ventral setae posteriorly in XI. Spermathecal pores paired between lateral lines and lines of dorsal setae, anteriorly in X. Alimentary canal absent. Male genitalia (Fig. 13D) paired. Vas deferens longer than atrium, up to 10 wm wide, but whole length not visible in available spm. Atrium some- what comma-shaped, 50-60 um long, 25 um wide, with thin outer lining and thick inner epithelium, but details of lumen not clear. Atrium tapering ectally into short duct opening at inner end of very complex, fold- ed and deep penial sac; one “‘fold’’ of sac appearing as somewhat granulated and pen- dant(?) papilla. Prostate glands two per atrium, large and lobed; posterior one at- tached by long stalk to ectal part of atrium. Pair of glandular bodies of unknown func- tion located anterior to male pores near body wall. Spermathecae (Fig. 13D, s) consisting of inconspicuous ducts, about 20 um long, about 15 wm wide, and bipartite ampullae, 135-165 wm long; parts of each ampulla separated from each other by consiriction, ectal part 30-35 wm wide and devoid of 308 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (ee ee ee Fig. 13. Olavius latus, n. sp.: A, Pygidium with caudal process; B, Somatic setae; C, Penial seta; D, Lateral view of spermatheca and male genitalia in segments X—XI. sperm, ental part 35-45 um wide and con- taining broad bundle of sperm. Remarks.—This species appears closely related to O. planus (Erséus, 1979) from Bermuda, another large species of Olavius characterized by a flat body shape and pos- session of a long caudal process (Erséus 1979b). However, the new species has a more complex penial sac than has O. plan- us, and its penial setae are single-pointed, not bifid as in the other species. Etymology.—The species name /atus is Latin for “broad, wide,”’ and refers here to the flattened shape of the worm. Distribution and habitat.—Known only from off Hutchinson Island, east coast of Florida. Shell hash, 11 m depth. Olavius sp. A Material.—Author’s collection: 1 whm spm from Sta 2. Remarks.—This single specimen is very similar to O. /atus described above, but lacks penial setae and is probably a separate species. However, it is not considered ap- propriate to describe it as such until addi- tional material becomes available. Olavius sp. B Material.—Author’s collection: 2 whm spms from Sta 2. Remarks. —This large, most probably new, gutless species is briefly characterized as follows: body wide and flat; pygidium rounded, without filiform appendage; peni- al setae 6-10 per bundle. Unfortunately, male genital organs are not clearly visible in the two specimens at hand. Subfamily Limnodriloidinae Limnodriloides Pierantoni, 1903 Limnodriloides vespertinus Erséus, 1982 Limnodriloides vespertinus Erséus, 1982c: 215-216, fig. 2. New material examined. —FSBC I 31874, 1 whm spm from Sta 4 (10 May 1972). Remarks.—This species was described from near Miami, Florida, and from Andros Island, Bahamas, by Erséus (1982c). The VOLUME 99, NUMBER 2 single specimen from Hutchinson Island has poorly developed spermathecae and its pos- terior end is not fully grown (specimen only 3.5 mm long, consisting of about 36 seg- ments), but otherwise it fits the original de- scription. Distribution and habitat.—East coast of Florida and Bahamas. Subtidal, generally somewhat muddy sands, down to 11 m depth. Limnodriloides monothecus Cook, 1974 Limnodriloides monothecus Cook, 1974: 131-132, fig. 2.—Brinkhurst and Baker, 1979:1664.—Erséus, 1982c:250-—253, figs. 28-29. Bohadschia monotheca Hrabé, 1975:112- 114, figs. 1-3. New material examined. —FSBC I 331875, 1 whm spm from Sta 2 (2 Nov 1972).—Author’s collection: 2 whm spms from Sta 2. Remarks. — This species is widely distrib- uted in North America (including the Pa- cific coast) and the Caribbean area (cf. Er- seus 1982c:fig. 14B for map), and it has been reported also from the Mediterranean Sea (as Bohadschia monotheca Hrabé, 1975, which is regarded as both a synonym and a homonym for L. monothecus). The new material largely conforms to the previous descriptions, but a slight deviation from the latter regarding some setal characteristics (cf. Erséus 1982c:table 1) should be noted. In the present specimens, preclitellar setae are up to four (occasionally five) per bundle, which is more than the two to three pre- viously stated, and ventral setae are present in X in two of the three worms (these setae absent in both X and XI of previously stud- ied material). In one specimen, the sper- matozeugmata are short and stout as noted for material from Barbados by Erséus (1982c), but in the other two individuals they are of the “normal,” very slender type. Distribution and habitat. — Pacific coasts of British Columbia, California, and Mex- ico; Atlantic coast of Florida through New 309 Fig. 14. Marcusaedrilus luteolus, spermatheca. Jersey; Gulf coast of Florida; Bermuda; Bar- bados; Yugoslavia. Euryoecious, occurring also in brackish water; in various kinds of sand and silt, generally with rich organic material, down to 370 m depth. Marcusaedrilus Righi and Kanner, 1979 Marcusaedrilus luteolus Erséus, 1983 Fig. 14 Marcusaedrilus luteolus Erséus, 1983a:27—- 29, fig. 3. New material examined. —FSBC I 31876- 31877, 2 whm spms, from Sta 2 (2 Nov 1972) and Sta 5 (10 May 1972), respective- ly.—Author’s collection: 4 whm spms from Sta 2, 2 from Sta 4, and 45 from Sta 5. Remarks. —Marcusaedrilus luteolus was described on the basis of material from Bar- bados and from near Miami, Florida (Er- seus 1983a). New specimens are consis- tently larger (12.6-15.7 mm long, 77-88 segments) than the original material (6.1- 6.9 mm, 53-68 segments), and spermathe- cae of all individuals from Florida (see Fig. 14) are larger than those of the original ma- terial from Barbados (Erséus 1983a:29, fig. 3D). 310 Distribution and habitat.— Barbados and east coast of Florida. Subtidal muds and sands, down to 21 m depth. Subfamily Tubificinae Tubificoides Lastockin, 1937 Tubificoides annulus, new species Fig. 15 Holotype. —USNM 98140, whm spm from Sta 4 (14 Mar 1972). Paratypes. —FSBC1 31878-31879, 2 whm spms from Sta 4 (type locality; 6 Mar 1973) and Sta 5 (1 Mar 1972), respectively. Description.—Length (only 1 complete spm) 7.8 mm, about 47 segments (still grow- ing posteriorly). Width at XI, 0.20—0.22 mm. Prostomium rounded, well demarcated from peristomium, appearing somewhat retrac- tile within latter. Body wall smooth, without adhering foreign particles, but cuticle thick and finely ridged. Clitellum poorly devel- oped over XI—'2XII. Anterior dorsal bun- dles with 1-2 single-pointed setae (possibly bifid with teeth very close together) and 1-— 2 hair setae; single-pointed setae (Fig. 15A) 35-55 um long, hair setae 85—150 um long. Anterior ventral bundles with 40-50 um long bifid setae (Fig. 15B), with upper tooth thin- ner and slightly shorter than lower. Dorsal and ventral bundles in postclitellar seg- ments each represented by | bifid seta (Fig. 15C), similar to anterior bifids but generally more curved entally. Ventral setae absent in XI. Male pores paired in line with ventral setae, posteriorly in XI. Spermathecal pores paired immediately anterior to ventral setae in X. Pharyngeal glands in IV—V. Part of esoph- agus enlarged and somewhat glandular in IX. Male genitalia (Figs. 15D-E) paired. Vas deferens 11-18 wm wide, thin-walled and densely ciliated, at least 5 times as long as atrium, entering latter sub-apically. Atrium about 200 um long, cylindrical and curved, entally (at caecum) about 30 um wide, at middle about 20 um wide. Atrium with wide, distinct outer layer of muscles, 1—2 wm thick, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON arranged in circles around long axis; mus- cles more developed in ental than in ectal part. Inner epithelium of atrium histologi- cally bipartite; inner part (caecum and area with entrances of vas deferens and prostate gland) with glandular, discrete bodies, lu- men wide and containing few cilia near en- trance of vas deferens; outer part evenly granulated, lumen not observed. Prostate gland attached by short stalk to atrium, clearly more ectal than, but on opposite side to, that of entrance of vas deferens. Penial sheath short (only about 15 um long), some- what ring-shaped, but with inner end some- what wider (about 20 um) than outer. Sper- mathecae (Fig. 15D, s) with very slender ducts and oblong ampullae; sperm trap in- distinct but appearing to be present. Sper- matozeugmata slender with rounded tips, but not very long (as compared to most con- geners). Remarks.—The characteristic shape of penial sheaths distinguishes 7. annulus from the other species of the genus. Sheaths are reminiscent of those of 7. brevicoleus Ba- ker, 1983, a littoral form from the Pacific coast of Canada, which also possesses hair setae and very long vasa deferentia. Tubi- ficoides brevicoleus, however, bears hair se- tae in postclitellar segments as well as an- teriorly (hairs restricted to anterior segments in 7. annulus) and papillae along most of the body (naked in 7. annulus). Etymology.—The name annulus, which is Latin for “ring,” alludes to the shape of the penial sheaths in this species. Distribution and habitat.—Known only from off Hutchinson Island, east coast of Florida. Coarse, clean or somewhat muddy, sands, 10—11.5 m depth. Tubificoides sp. Material examined. —Author’s collec- tion: 4 whm spms from Sta | (but see note under ‘material and methods’ above), 23 from Sta 2, 6 from Sta 4, and 23 from Sta 5. Remarks. —This species, which occurred VOLUME 99, NUMBER 2 311 Fig. 15. at four of the stations, belongs to a complex of species within Tubificoides lacking hair setae. The complex is currently under re- vision by Dr. R. O. Brinkhurst in Canada, and, therefore, the species will not be further treated here. It appears closely related to T. wasselli Brinkhurst and Baker, 1979. Discussion Summarized data on the local distribu- tion of the Hutchinson Island Oligochaeta show that all but one of the 25 species oc- curred at either Sta 2 or Sta 4, or, for most of the species, at both stations (Table 2). Specimens from these two stations clearly dominate the material, although the sam- pling effort was the same for all stations. The overwhelming similarity between Sta 2 and Sta 4, in terms of distance from shore, depth, sediment composition (very coarse, shell hash), and oligochaete fauna, appears to suggest that they can be “united”’ for a total characterization of the particular trough in which they are located. Thus, it can be concluded that the benthic community of the coarse, shelly bottom of this trough in- cludes an association of at least 24 intersti- tial, tubificid species. This is a remarkably high figure, as it outnumbers all other sim- Tubificoides annulus, n. sp.: A, Anterior, single-pointed, dorsal seta; B, Anterior ventral seta; C, Posterior seta; D, Lateral view of spermatheca and male genitalia in segments X—XI; E, Penis sheaths. ilar oligochaete associations that have been more carefully examined to date; for in- stance, only eight coarse-sand tubificids were found in Cape Cod Bay, Massachusetts (Cook 1971), and the corresponding species were only four in some Norwegian fjords studied by Erséus (1976). In a recent paper on the Oligochaeta of Georges Bank off Massachusetts, Davis (1985) noted that when sampling in some medium-to-coarse- sand habitats, as many as 20 species could occur in the same sample, but this figure also includes some species belonging to families other than the Tubificidae. High diversity of interstitial tubificids will also be reported from medium to coarse sands off Virginia (approximately midway between Massachusetts and Florida) (Diaz et al., in press), but nowhere reaching the level re- corded at Hutchinson Island. By adding nine species new to science and a further four (named) species new to the state’s fauna, the Hutchinson Island mate- rial has considerably increased our knowl- edge about the marine Tubificidae of Flor- ida. The present checklist now comprises 50 species (Table 1). It should be noted that several additional records of estuarine and offshore species along the Gulf of Mexico 312 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Numbers of sexually mature specimens of Tubificidae collected (1971-1973) at Stations 1-5 off Hutchinson Island. Numbers are pooled for all replicates over the whole sampling period for each station. Species are ranked after abundance. Total Sta Sta Sta Sta Sta Rank Species n 1 2 3 4 5 1 Tubificoides sp. 56 4 23 6 23 2 Marcusaedrilus luteolus 53 6 2 45 3 Heterodrilus perkinsi 47 18 29 4 Heterodrilus bulbiporus 46 D, 20 3 21 5 Phallodrilus biprostatus 40 16 D Di 1 6 Heterodrilus minisetosus 18 D 16 7 Phallodrilus sabulosus 15 yD 12 1 8 Heterodrilus pentcheffi 14 10 2 1 1 9 Inanidrilus ernesti 12 7 5 10 Heterodrilus occidentalis 11 5 6 11 Phallodrilus acochlearis 9 9 12 Heterodrilus hispidus 8 6 2 13 Bathydrilus ingens 3 2 1 14 Coralliodrilus corpulentus 3 1 2 15 Limnodriloides monothecus 3 3 16 Tubificoides annulus 3 2 1 17 Adelodrilus magnithecatus 2 2 18 Bathydrilus macroprostatus 2 2 19 Inanidrilus vacivus D, 1 1 20 Phallodrilus hirsutus 2 1 1 21 Olavius sp. B 2 2 22 Bathydrilus formosus 1 23 Olavius latus 1 24 Limnodriloides vespertinus 1 25 Olavius sp. A 1 All samples: 355 * For habitat purposes, this should be regarded as Sta 5 (cf. ““Material and methods” section). coast will soon be available (M. R. Milligan, pers. comm.). The east coast of Florida has been rec- ognized as a transitional zoogeographic zone for shallow-water marine invertebrates (see review by Briggs 1974). The area at Hutch- inson Island is characterized by an overlap of warm-temperate and tropical faunal ele- ments (e.g., Work 1969; Camp et al. 1977); in the terminology of Valentine (1973) and Dobzhansky et al. (1977), the area is at the border between the Carolinian and Carib- bean provinces. The present material and previous records of Florida east coast Tubi- ficidae (cf. Table 1) seem to support this view, although the distribution of marine oligochaetes is still not very well known, as indicated by the fact that about 40% of the species known from Florida are known from nowhere else. Heterodrilus bulbiporus, H. occidentalis, H. pentcheffi, L. barnardi, L. rubicundus, O. tenuissimus, P. prostatus, and P. sabulosus appear to be warm-temperate species, pres- ent as they are along a great part of the U.S. east coast, several even as far north as off Massachusetts. Adelodrilus magnithecatus, B. ingens, and P. acochlearis are possibly more restricted to the southeastern states (the first species known also from Bermu- da). Caribbean forms include B. formosus, M. luteolus, M. hummelincki, K. ineri, and T. bori. None of the latter has been taken north of Florida on the U.S. east coast, but the last three are present at Bermuda, which can be regarded as a Caribbean “‘satellite.” VOLUME 99, NUMBER 2 Some species occurring along the Florida east coast are so widely distributed in the world that they may be regarded as circum- tropical (B. adriaticus, P. rectisetosus, L. monothecus, and T. gurwitschi) or even cos- mopolitan (MVM. rubroniveus; cf. Baker and Brinkhurst 1981). The only more general discussion pub- lished on marine tubificid zoogeography and diversity is a tentative review by Baker (1984). Acknowledgments I am indebted to Mr. Thomas H. Perkins for placing the Hutchinson Island Oligo- chaeta at my disposal, and for his support for this study; to Drs. M. Susan Ivester and Ralph O. Brinkhurst, and Mr. Michael R. Milligan, for supplementary material; to Ms. Barbro Lofnertz for skillful technical assis- tance; and the Swedish Natural Science Re- search Council for financial support. Particular thanks are due to the Florida Department of Natural Resources Bureau of Marine Research and the Florida Power and Light Company for defraying the pub- lication costs, and to Mr. T. H. Perkins and Mr. W. G. Lyons for their many suggestions for improvements of the manuscript. This paper is a contribution of the Florida Department of Natural Resources Bureau of Marine Research, 100 8th Ave., St. Pe- tersburg, Florida 33701-5095, from which reprints are available. Literature Cited Baker, H.R. 1983. New species of Bathydrilus Cook (Oligochaeta; Tubificidae) from British Colum- bia.—Canadian Journal of Zoology 61:2162- 2167. . 1984. Diversity and zoogeography of marine Tubificidae (Annelida, Oligochaeta) with notes on variation in widespread species.—Hydro- biologia 115:191-196. , and R. O. Brinkhurst. 1981. A revision of the genus Monopylephorus and redefinition of the subfamilies Ryacodrilinae and Branchiuri- nae (Tubificidae: Oligochatea).— Canadian Journal of Zoology 59:939-965. 313 , and C. Erséus. 1979. Peosidrilus biprostatus N.g., N. sp., a marine tubificid (Oligochaeta) from the eastern United States.— Proceedings of the Biological Society of Washington 92:505-509. Briggs, J.C. 1974. Marine Zoogeography. McGraw- Hill Book Company, New York. 475 pp. Brinkhurst, R.O. 1965. Studies on the North Amer- ican aquatic Oligochaeta II. Tubificidae. — Pro- ceedings of the Academy of Natural Sciences of Philadelphia 117:117-172. 1966. A contribution to the systematics of the marine Tubificidae (Annelida, Oligochae- ta).— Biological Bulletin 130:297-303. , and H. R. Baker. 1979. A review of the ma- rine Tubificidae (Oligochaeta) of North Amer- ica.—Canadian Journal of Zoology 57:1553- 1569. Camp, D. K., N. H. Whiting, and R. E. Martin. 1977. Nearshore marine ecology at Hutchinson Is- land, Florida: 1971-1974. V. Arthropods.— Florida Marine Research Publications 25:1-63. Coates, K.,and C. Erséus. 1985. Marine enchytraeids (Oligochaeta) of the coastal northwest Atlantic (northern and mid U.S.A.).— Zoologica Scripta 14:103-116. Cook, D. C. 1971. The Tubificidae (Annelida, Oli- gochaeta) of Cape Cod Bay. II: Ecology and sys- tematics, with the description of Phallodrilus parviatriatus nov. sp.— Biological Bulletin 141: 203-221. 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. Davis, D. 1985. The Oligochaeta of Georges Bank (NW Atlantic), with descriptions of four new species. — Proceedings of the Biological Society of Washington 98:158-176. Diaz, R. J., C. Erséus, and D. F. Boesch. In press. Distribution and ecology of Middle Atlantic Bight Oligochaeta.— Hydrobiologia. Dobzhansky, T., F. J. Ayala, G. L. Stebbins, and J. W. Valentine. 1977. Evolution. W. H. Freeman and Company, San Francisco, 572 pp. Easton, E.G. 1984. Earthworms (Oligochaeta) from islands of the southwestern Pacific, and a note on two species from Papua, New Guinea. — New Zealand Journal of Zoology 11:111-128. Erséus, C. 1975. Peloscolex amplivasatus sp. n. and Macroseta rarisetis gen. et sp. n. (Oligochaeta, Tubificidae) from the west coast of Norway. — Sarsia 58:1-8. 1976. Marine subtidal Tubificidae and En- chytraeidae (Oligochaeta) of the Bergen area, western Norway.—Sarsia 62:25—48. 314 1979a. Taxonomic revision of the marine genera Bathydrilus Cook and Macroseta Erséus (Oligochaeta, Tubificidae), with descriptions of six new species and subspecies.— Zoologica Scripta 8:139-151. 1979b. Taxonomic revision of the marine genus Phallodrilus Pierantoni (Oligochaeta, Tubificidae), with descriptions of thirteen new species. —Zoologica Scripta 8:187—208. 1979c. Inanidrilus bulbosus gen. et sp. n., a marine tubificid (Oligochaeta) from Florida, USA.—Zoologica Scripta 8:209-210. . 1979d. Bermudrilus peniatus n.g., n. sp. (Oli- gochaeta, Tubificidae) and two new species of Adelodrilus from the Northwest Atlantic.— Transactions of the American Microscopical Society 98:418—427. 1980a. Taxonomic studies on the marine genera Aktedrilus Knollner and Bacescuella Hrabé (Oligochaeta, Tubificidae), with descrip- tions of seven new species.— Zoologica Scripta 9:97-111. 1980b. Two new records of the Caribbean marine tubificid Kaketio ineri Righi and Kanner (Oligochaeta).— Proceedings of the Biological Society of Washington 93:1220-1222. 198la. Taxonomic revision of the marine genus Heterodrilus Pierantoni (Oligochaeta, Tubificidae).— Zoologica Scripta 10:111-132. 1981b. 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. . 1981c. Taxonomy of the marine genus Thal- assodrilides (Oligochaeta: Tubificidae).— Transactions of the American Microscopical Society 100:333-344. . 1982a. Revision of the marine genus Smith- sonidrilus Brinkhurst (Oligochaeta, Tubifici- dae).—Sarsia 67:47—54. . 1982b. Parakaketio longiprostatus gen. et sp. n., a marine tubificid (Oligochaeta) from Flor- ida, U.S.A.— Zoologica Scripta 11:195-197. 1982c. Taxonomic revision of the marine genus Limnodriloides (Oligochaeta: Tubifici- dae).— Verhandlungen des naturwissenschaft- lichen Vereins in Hamburg (Neue Folge) 25: 207-277. . 1983a. Taxonomic studies of the marine ge- nus Marcusaedrilus Righi & Kanner (Oligo- chaeta, Tubificidae), with descriptions of seven new species from the Caribbean area and Aus- tralia.— Zoologica Scripta 12:25-36. 1983b. New records of Adelodrilus (Oligo- chaeta, Tubificidae), with descriptions of two new species from the Northwest Atlantic. — Hy- drobiologia 106:73-83. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1983c. A new bathyal species of Corallio- drilus (Oligochaeta: Tubificidae) from the Southeast Atlantic. — Proceedings of the Biolog- ical Society of Washington 96:272-275. 1984a. A record of Bermudrilus peniatus (Oligochaeta: Tubificidae) from the Gulf of Mexico.— Gulf Research Reports 7:381. . 1984b. Taxonomy and phylogeny of the gut- less Phallodrilinae (Oligochaeta, Tubificidae), with descriptions of one new genus and twenty- two new species. — Zoologica Scripta 13:239-272. 1984c. Interstitial Fauna of Galapagos. XXXIII. Tubificidae (Annelida, Oligochae- ta).—Microfauna Marina 1:191-198. 1984d. 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. 1985. Annelida of Saudi Arabia. Marine Tubificidae (Oligochaeta) of the Arabian Gulf Coast of Saudi Arabia.— Fauna of Saudi Arabia 6:130-154. , and H. R. Baker. 1982. New species of the gutless marine genus /nanidrilus (Oligochaeta, Tubificidae) from the Gulf of Mexico and Bar- bados. — Canadian Journal of Zoology 60:3063- 3067. , and M. S. Loden. 1981. Phallodrilinae (Oli- gochaeta: Tubificidae) from the east coast of Florida, with description of a new species of Adelodrilus. —Proceedings of the Biological So- ciety of Washington 94:819-825. Gallagher, R. M. 1977. Nearshore marine ecology at Hutchinson Island, Florida: 1971-1974. II. Sed- iments. — Florida Marine Research Publications 23:6-24. ,and M. L. Hollinger. 1977. Nearshore marine ecology at Hutchinson Island, Florida: 1971- 1974. I. Introduction and rationale.— Florida Marine Research Publications 23:1—5. Gates, G. E. 1943. On some American and oriental earthworms. Part II. Family Megascolecidae. — Ohio Journal of Science 43:99-1 16. Giere,O. 1979. Studies on marine Oligochaeta from Bermuda, with emphasis on new Phallodrilus species (Tubificidae). — Cahiers de Biologie Ma- rine 20:301-314. Hrabé, S. 1975. Second contribution to the knowl- edge of marine Tubificidae (Oligochaeta) from the Adriatic Sea.— Vestnik Ceskoslovenské Spolecnosti Zoologiské 39:111-119. Kennedy, C. R. 1966. A taxonomic revision of the genus Grania (Oligochaeta: Enchytraeidae).— Journal of Zoology 148:399-407. Lasserre, P., and C. Erséus. 1976. Oligochétes marins des Bermudes. Nouvelles espéces et remarques sur la distribution géographique de quelques VOLUME 99, NUMBER 2 Tubificidae et Enchytraeidae. — Cahiers de Biol- ogie Marine 17:447—462. Loden, M. S. 1980. A new euryhaline species of Monopylephorus (Oligochaeta: Tubificidae) from the southeastern United States.—Bulletin of Marine Science 30:600-603. Pierantoni, U. 1917. Sull’ Heterodrilus arenicolus Pierant. e su di una nuova species del genere Clitellio. —Bollettino della Societa di Naturalisti 1 Napoli 29:82-91. Valentine, J. W. 1973. Evolutionary paleoecology of 315 the marine biosphere. Prentice-Hall, Englewood Cliffs, New Jersey. Work, R. C. 1969. Systematics, ecology and distri- bution of the mollusks of Los Roques, Vene- zuela.— Bulletin of Marine Science 19:614—711. 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. 99(2), 1986, pp. 316-322 CAECIDOTEA DAUPHINA, A NEW SUBTERRANEAN ISOPOD FROM A BARRIER ISLAND IN THE NORTHERN GULF OF MEXICO (CRUSTACEA: ISOPODA: ASELLIDAE) Richard F. Modlin Abstract.—Caecidotea dauphina, a phreatobitic asellid of the Hobbsi group, is described and identified. Its type habitat is located on a sandy barrier island in the Gulf of Mexico off the coast of Alabama. Records of subterranean isopods of Ala- bama are limited to four. Caecidotea ala- bamensis was reported originally from a well located in the Piedmont of east-central Al- abama (Stafford 1911), while C. bicrenata (Steeves 1963) and C. meisterae (Lewis and Bowman 1981; Modlin, unpublished rec- ord) were collected in caves in the northern Appalachian region. Steeves (1964) syn- onymized C. bicrenata with C. alabamen- sis, but Lewis and Bowman (1981) rein- stated its species status. The species list of Fleming (1972) includes C. richardsonae in addition to C. alabamensis from locations in Alabama. Morphologically, all four species align with Steeves’ Stygius group (Steeves 1963, 1966). Herein is the descrip- tion of a new species of the Hobbsi group collected on Dauphin Island, Mobile Coun- ty, Alabama, a sandy barrier island in the southernmost part of the state. Caecidotea dauphina, new species Figs. 1-4 Material examined.—Alabama, Mobile County, Dauphin Island, Audubon Sanc- tuary, 9 Aug 1984, R. F. Modlin, 1 male (7.5 mm) holotype, partly dissected on 5 slides and in alcohol (USNM 227076); 8 females (3.3—4.5 mm) and 1 juvenile (2.4 mm) paratypes, in alcohol (USNM 227077). 27 Aug 1984, R. F. Modlin, 1 female (4.2 mm), dissected on 7 slides, in author’s col- lection (082784F22); same locality. Type- specimens are deposited in the U.S. Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. (USNM). Diagnosis. — Blind, unpigmented. Anten- na | esthete formula 0-1-0. Male pereopod 1 palm defined by robust proximal spine, small triangular mesial process, and trun- cate distal process. Male pleopod 1 longer than pleopod 2, protopod with 6 retinacula. Endopodial tip of male pleopod 2 with short straight cannula, triangular mesial process with tip curved medially, and truncate lat- eral process. Description. —Lateral sides of body slightly concave, narrowest at pereonites 3 and 4. Average width of 7.5 mm male is 1.6 mm. Head about 0.6 x wider than long; an- terior margin concave. Telson about 0.8 x longer than wide; lateral margins parallel, posterior margin with distinct caudomedial lobe. Antenna | reaching to about midlength of last segment of antenna 2 peduncle. Fla- gellum composed of 10 segments, esthete formula 0-1-0. Single esthete located on seg- ment 9. Antenna 2 flagellum about 2.5 x length of peduncle, reaching to about 6th pereonite. Mandible with 4-cuspate incisors and la- cinia, spine-row with 8 and 10 spines re- spectively on right and left. Medial spine row on segment 3 of mandibular palp in- VOLUME 99, NUMBER 2 317 \S \ \ Pal Peg Fig. 1. Caecidotea dauphina, A-E, L = 7.5 mm male holotype; F—K = 4.2 mm female: A, Habitus, dorsal; B, Antenna 1; C, Antenna 2, peduncle; D, Mandibular palp; E, Left mandible; F, Right mandible; G, Labrum; H, Left labium; I, Maxilla 1; J, Maxilla 1 outer lobe; K, Maxilla 2; L, Maxilliped. 318 creasing slightly in length distally, 2 distal- most spines about 2.5 x longer than others; distal row of spines on segment 2 strongly increasing in length proximally, minute se- tae arranged in linear clumps located lateral to spine row. Maxilla 1 inner lobe with 5 apical plumose setae; outer lobe with 13 apical robust spines (not all spines shown in Fig. 1], J) and 2 subapical setae, one sim- ple and one plumose. Maxilliped with 5 ret- inacula in male and 3—4 in females. Male pereopod 1 propodus about 0.9 x as wide as long; palm defined by robust proximal articulated spine, triangular me- sial process separated from lower truncate distal process by shallow cleft, distal process about twice as wide as mesial process; car- pus with 3 robust articulated spines and 3 slender simple setae on anterodistal margin. Female pereopod 1 propodus about 1.6 x as wide as long; palm defined by robust proximal articulated spine, without mesial and distal processes; carpus with 1 robust and 2 smaller articulated spines and | long simple seta on anterodistal margin. Sexual dimorphism in pereopods 2-7 exhibited only by fewer spines on female pereopods. Male pleopod 1 longer than pleopod 2, protopod with 6 retinacula; endopod about 0.4 x as long as wide, medial and distal mar- gins convex, lateral margin slightly concave in distal half, distal margin with 4 long plu- mose setae and 5 shorter simple setae, lat- eral margin with 8-10 very small simple setae in distal half and about 5 simple setae of moderate length in proximal half, about 5 small setae subterminal to distal margin. Male pleopod 2 protopod with 3 simple setae on medial margin; exopod basal seg- ment triangular with 3 short simple setae on lateral edge, distal segment oval, about 0.8 x as long as wide, with 14 plumose mar- ginal setae; endopod with rounded lateral basal apophysis, distal end of lateral margin weakly striated, tip with 4 processes: 1) can- nula straight, subapical, slightly directed medially, partly obscured in anterior view by mesial process, but completely visible in posterior view; 2) mesial process with apex PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rising above other processes, apex twisted strongly in medial direction, proximal *4 of medial edge with many striae, lateral edge almost straight forming medial edge of en- dopodial groove; 3) lateral process rectan- gular, height about 0.7 x distance from base of endopodial groove to tip of the mesial process, apex truncate, lateral edge drop- ping straight to small shelf; 4) caudal pro- cess low, rounded, traversing entire tip of endopod. Female pleopod 2 with 7 plumose marginal setae. Male pleopod 3 exopod with about 20 long plumose setae along and around distal margin, short plumose setae interspersed with simple setae along distal 7/, of lateral margin, short simple setae line lateral margin proximal to suture, many short simple setae on anterior surface. Pleo- pod 4 exopod with about 5 long plumose setae near lateral edge of distal margin, many minute setae interspersed with long simple setae on proximal 4 of lateral margin, false suture pattern A with transverse suture ter- minating on lateral margin just below mid- length. Pleopod 5 with 2 false sutures and 5 simple setae on lateral margin proximally. Uropods spatulate, about 0.6 x length of tel- son, endopods 1.7 x longer than exopods. Etymology. —This species’ name refers to Dauphin Island, Alabama on which the specimens were collected. Habitat.—The type habitat of C. dau- phina is located in the Audubon Sanctuary on Dauphin Island, Mobile County, Ala- bama (30°15’N, 88°05'W) (Fig. 5). Speci- mens were collected in a depression located about 1—2 m from the outfall of a tile drain pipe that crosses under a service road. This depression is on the periphery of a black gum, Nyssa sylvatica, swamp and it is pe- riodically dry. The swamp is a remnant of the once expansive Alligator Swamp system that covered much of the eastern part of Dauphin Island. Consequently, the sanc- tuary and contiguous areas contain several artesian wells that become active during the rainy season (McNeely 1974). Specimens were collected in the company of the epi- gean isopod C. obtusus several days after a VOLUME 99, NUMBER 2 Fig. 2. Caecidotea dauphina, Pereopod 1: A, 7.5 enlarged; B, detail of palmar margin of propodus and major rain storm. Caecidotea dauphina was probably flushed from the tiles by the in- creased water flow. Some physical and chemical data were collected to characterize the habitat; water temperature ranged from 25—26°C, pH 5.3- 5.6 and conductivity 236-360 wMhos at Us Os Interestingly, the type habitat is about 500 m from the saline water of the Gulf of Mex- ico and on an island whose last surface land connection to the mainland occurred during the Pleistocene (Price 1954). Relationships. —Caecidotea dauphina shows affinity to the Hobbsi group because it has all the morphological characteristics that define this group (Steeves 1964; Lewis 1982). An evolutionary link may exist be- tween C. dauphina and C. tridentata, C. 319 SGALIE. tin tia ©.10 AG O05 8B mm male holotype, structure of multiflagellated spine anterior margin of carpus; C, 4.2 mm female. spatulata, C. teresae, and C. parvus. All these species have a mesial process on the tip end of the endopod of male pleopod 2 that curves, to varying degrees, medially. Setal structure and arrangement on the dis- tal margin of the exopod of male pleopod 1, endopod of male pleopod 2 and margins of the female pleopod 2 suggest that C. dau- phina has more in common with C. teresae than with the others. However, the geo- graphic ranges of C. teresae, C. tridentata and C. spatulata are in close proximity and may overlap, but they are greatly separated from that of C. dauphina. The former three are known from the midwestern United States (Lewis and Bowman 1981; Lewis 1982), while C. dauphina is located in the coastal region of Alabama, slightly west of the Hobbsi geographic range originally pro- 320 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Caecidotea dauphina, A-C, E-G = 7.5 mm male holotype; D, 4.2 mm female: A-C, Pereopods 2- 4; D, Pereopod 4; E-G, Pereopods 5-7. VOLUME 99, NUMBER 2 321 SCALE, mm 0.50 | 0,25 A,B,F-H O16 J 0.10 E 0.025 C, D Fig. 4. Caecidotea dauphina, A-D, F-J = 7.5 mm male holotype; E = 4.2 mm female: A, Pleopod 1; B, Pleopod 2; C and D, Pleopod 2 endopodial tip, ventral and dorsal views, can = cannula, c.p. = caudal process, l.p. = lateral process, m.p. = mesial process; E, Pleopod 2; F, Pleopod 3; G. Pleopod 4; H, Pleopod 5; I, Telson- uropod complex; J. Uropod. posed by Steeves (1964). The type habitat of C. dauphina appears contiguous with that of C. parvus which occurs in north central Florida (Steeves 1964). However, C. parvus does not belong to the Hobbsi group (Lewis 1982). Its only similarity to C. dauphina and the other three species above is the structure of the mesial process on the en- dopodial tip of the male pleopod 2. Addi- tionally, C. dauphina along with C. teresae, C. tridentata, and C. spatulata are phreato- bitic and occupy drain tile habitats (Lewis and Bowman 1981; Lewis 1982), while C. parvus is a troglobite inhabiting caves (Steeves 1964). Recently C..parvus was placed into the new genus Remasellus be- 322 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON DAUPHIN ISLAND ALABAMA \S MOBILE X BAY ¢\ MISSISSIPPI SOUND a OG Wik MEX1CO Fig. 5. Dauphin Island, Alabama, showing the Audubon Sanctuary where specimens of Caecidotea dauphina were collected. Inset indicates the position of Dauphin Island along the northern coast of the Gulf of Mexico. cause it was found to be a swimming isopod (Bowman and Sket 1985). Acknowledgments I wish to thank Dr. Thomas E. Bowman, USNM, for critically reviewing this manu- script and for providing helpful comments, and the Marine Environmental Sciences Consortium, Dauphin Island Sea Lab, Dau- phin Island, Alabama, for providing labo- ratory space and other facilities during this project. This is M.E.S.C. Contribution #088. Literature Cited Bowman, Thomas E., and Boris Sket. 1985. Rema- sellus, anew genus for the troglobitic swimming Florida asellid isopod, Asellus parvus Steeves. — Proceedings of the Biological Society of Wash- ington 98(3):554—-560. Fleming, Laurence E. 1972. The evolution of the east- ern North American isopods of the genus Asel- lus (Crustacea: Asellidae) Part I.—International Journal of Speleology 4:221-256. Lewis, Julian J. 1982. A diagnosis of the Hobbsi group, with descriptions of Caecidotea teresae, n. sp., and C. macropropoda Chase and Blair (Crustacea: Isopoda: Asellidae).— Proceedings of the Biological Society of Washington 95(2): 338-346. , and Thomas E. Bowman. 1981. The subter- ranean asellids (Caecidotea) of Illinois (Crus- tacea: Isopoda: Asellidae).—Smithsonian Con- tributions to Zoology 355:1-66. McNeely, S. B. 1974. The development of Dauphin Island, Alabama.—The Mobile Area Chamber of Commerce, 93 pp. Price, W. Armstrong. 1954. Shoreline and coasts of the Gulf of Mexico. Jn Galtsoff, P. S. (ed.), Ge- ology of the Gulf of Mexico, its origin, waters and marine life.—U.S. Fish and Wildlife Ser- vice, Fishery Bulletin 55(89):39-85. Stafford, B. E. 1911. A new subterranean freshwater isopod. — Pomona Journal of Entomology 3:572- 575. Steeves, Harrison R., III. 1963. The troglobitic asel- lids of the United States: The Stygius group. — American Midland Naturalist 69(2):470-481. 1964. The troglobitic asellids of the United States: The Hobbsi group.— American Midland Naturalist 71(2):445-451. . 1966. Evolutionary aspects of the troglobitic asellids of the United States: The Hobbsi, Sty- gius, and Cannulus groups.—American Mid- land Naturalist 75(2):392-403. Department of Biological Sciences, Uni- versity of Alabama in Huntsville, Hunts- ville, Alabama 35899. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 323-327 SETAL MORPHOLOGY OF THE OLIGOCHAETES TUBIFEX TUBIFEX AND ILYODRILUS FRANTZI (CAPILLATUS) AS REVEALED BY SEM Peter M. Chapman and Ralph O. Brinkhurst Abstract. —Scanning electron microscope observations of Tubifex tubifex re- veal details of the serrations on hairs, pectinations on dorsal setae, and more extensive pectinations on the ventral setae than previously described. Lack of serrations on I/yodrilus frantzi (capillatus) hairs is documented as are details of dorsal setal pectinations. Three new features of this species are shown: the concave nature of the upper and lower setal teeth, pectinate ventral setae, and an apparent ability to retract setal bundles. Scanning Electron Microscopy (SEM) is an extremely valuable technique for estab- lishing and documenting, at a greater level of resolution than is possible with light mi- croscopy, the presence of fine differences in organism morphology. Previous uses of SEM with oligochaetes include documen- tation of external sense organs (Chapman 1979; Smith 1983) and of fine setal differ- ences in two species of Naididae (Smith 1985). In previous studies (Chapman and Brinkhurst, in press; Brinkhurst and Chap- man, in prep.), we used the SEM technique to document setal variations inducible in naidid and tubificid oligochaetes under varying environmental conditions. As a re- sult of these previous studies, we obtained various new descriptive observations on the structure of the setae of Tubifex tubifex and Ilyodrilus frantzi (capillatus) which are pro- vided herein, related to taxonomic descrip- tions of these species. Materials and Methods Tubifex tubifex specimens were collected from the Fraser River, B.C.; I. frantzi (cap- illatus) specimens were collected from the Fraser River and from the Columbia River, Oregon. Worms were fixed in cold 3% buff- ered glutaraldehyde, washed in 0.1 M phos- phate buffer and post-fixed in 1% buffered osmium tetroxide for 2 h at room temper- ature. Specimens were then washed in 0.1 M phosphate buffer and dehydrated through an ascending ethanol series before being critical-point dried with CO,, mounted on aluminum stubs, and sputter coated with gold. A Cambridge S100 Stereoscan Scan- ning Electron Microscope was used for viewing. Results Details of serrations on 7. tubifex hairs, pectinations of dorsal setae, and pectina- tions of ventral setae, are provided in Fig. 1. Hairs in the same setal bundle can vary from virtually non-serrate to heavily ser- rate. Similarly, the degree of ventral setal pectination is highly variable, even within the same bundle and can approach the dor- sal setal condition. Details of hairs, dorsal and ventral setae of I. frantzi (capillatus) are provided in Fig. 2. Hairs are shown to be essentially non- serrate. Dorsal and ventral setae have con- cave outer teeth, may have varying degrees of pectination, and can apparently be re- tracted. Discussion Tubifex tubifex.—T. tubifex are de- scribed as having serrate or non-serrate hairs PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 99, NUMBER 2 325 Fig. 2. Details of setal morphology of I/yodrilus frantzi (capillatus): a—b, Variable amounts of dorsal pectin- ation, hairs show only traces of serrations; c, Detail of dorsal pectinate seta showing concave lower tooth (upper tooth is similar); d, Retracted, non-pectinate dorsal setae; e, Non-pectinate ventral setae; f, Pectinate ventral seta. Scale bars 10 um for b and c, 20 um for a and d to f. ee Fig. 1. Details of setal morphology of Tubifex tubifex: a—b, Variable amounts of serrations on dorsal hairs, note serrations begin around top of pectinate setae; c, Detail of dorsal pectinate seta; d—i, Ventral pectinations observed in various specimens. Scale bars 5 wm for c, 10 wm for a and d, 20 um for b and e toi. 326 (Brinkhurst and Jamieson 1979; Brinkhurst 1982; Brinkhurst, 1986). The nature of these serrations, as documented in Fig. la— b, is variable even within a single dorsal bundle, which may simply reflect differen- tial exposure to abrasion or other environ- mental factors causing shredding. SEM pho- tographs indicate that the observed serrations are due to shredding; serrations generally begin immediately distal to the top of the dorsal setae, suggesting that the setae provide some measure of protection against abrasion. The hypothesis of serra- tions being formed through shredding was first advanced by Smith (1985), for Dero spp. Since serrations appear to be a variable characteristic, probably influenced by en- vironmental conditions, their use in oligo- chaete taxonomy should be discontinued. The nature of dorsal setal pectination is illustrated (Fig. la—c). Pectinations have ap- proximately equal lateral teeth shorter in length than the upper and lower setal teeth. The degree and variability of pectination possible on ventral setae is large (Fig. 1d— h). The degree of pectination can vary within a single bundle (Fig. 1d), and shows a range including single (Fig. 1e—f), bifid (Fig. lg—h) and trifid pectinations (Fig. 1i). The degree of ventral pectination may approx- imate that seen dorsally (compare Fig. lc and 1i) and is shown to be more extensive than that described by Brinkhurst and Jamieson (1971:455), “rarely with a single intermediate tooth,” Brinkhurst (1982:40), “sometimes a few short intermediate teeth on ventral setae,” or Brinkhurst (1986:149), ““sometimes with a small intermediate tooth.” Ilyodrilus frantzi (capillatus).—Descrip- tions of I. frantzi (capillatus) have not in- cluded lateral serrations on hairs. This lack of serrations is confirmed by SEM (Fig. 2a— b). Brinkhurst (1978) provided the first re- port of dorsal pectinate setae; Brinkhurst (1986:173) states that there are a “few pec- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tinate setae” dorsally. Dorsal setae may have none, one (Fig. 2a), or two pectinations (Fig. 2b). SEM reveals that the inner surfaces of the setae of I. frantzi capillatus have gutter- shaped (concave) upper and lower teeth (Fig. 2c, e-f). Apparently setal bundles can be retracted (Fig. 2d). These features have not previously been described for this or for any other aquatic oligochaete. The presence of ventral pectinations was suspected by Bginkhurst (1978:2173) who noted that the ventral setae in some speci- mens “seem to be slightly ornamented, but this needs confirmation by scanning elec- tron microscopy.” Such confirmation is now provided. Ventral setae may be apectinate (Fig. 2e), or may have a single intermediate pectination (Fig. 2f). Acknowledgments Technical assistance was provided by S. Cross (E.V.S. Consultants), D. Little (Uni- versity of Victoria), and M. Johns (Institute of Ocean Sciences). The paper was word- processed by M. Mees. Literature Cited Brinkhurst, R. O. 1978. Freshwater Oligochaeta in Canada.— Canadian Journal of Zoology 56:2166- 2175. . 1982. British and other marine and estuarine oligochaetes. Cambridge University Press, N.Y. 127 pp. 1986. A guide to the freshwater aquatic microdile oligochaetes of North America. —Ca- nadian Special Publication of Fisheries and Aquatic Sciences 84, 259 pp. , and P. M. Chapman. [In Preparation.] Vari- ations in somatic characteristics of aquatic oli- gochaetes. , and B. G. M. Jamieson. [In press]. Aquatic Oligochaeta of the World. University of Toron- to Press. 866 pp. Chapman, P. M. 1979. The prostomial pit in Both- rioneurum vejdoyvskyanum Stole (Oligochae- ta)—a note on detail revealed by SEM.—Pro- VOLUME 99, NUMBER 2 ceedings of the Biological Society of Washington 93:812-813. , and R. O. Brinkhurst. [In press.] Hair today, gone tomorrow (induced setal changes in tubif- icid oligochaetes). — Hydrobiologia. Smith, M. E. 1983. External sense organs of Tubifex tubifex and Limnodrilus hoffmeisteri (Tubifici- dae).— Freshwater Invertebrate Biology 2:154— 158. . 1985. Setal morphology and its intraspecific variation in Dero digitata and Dero nivea (Oli- 327 gochaeta: Naididae).—Transactions of the American Microscopical Society 104:45—-51. (PMC) E.V.S. Consultants Ltd., 195 Pem- berton Avenue, North Vancouver, B.C. V7P 2R4, Canada; (ROB) Ocean Ecology Sec- tion, Institute of Ocean Sciences, P.O. Box 6000, 9860 West Saanich Road, Sidney, B.C. V8L 2Y2, Canada. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 328-334 SERRABRYCON MAGOTI, A NEW GENUS AND SPECIES OF SCALE-EATING CHARACID (PISCES: CHARACIFORMES) FROM THE UPPER RIO NEGRO Richard P. Vari Abstract.—Serrabrycon magoi, a previously undescribed genus and species of scale-eating tetragonopterin characid, is described from the black waters of the Rio Negro system in the region of the Rio Casiquiare, Venezuela. The genus and species are unique in the combination of the presence of a series of out- wardly oriented teeth in the upper and lower jaws, the relatively low number of scales in a longitudinal series to the hypural joint, and in the limitation of pores for the laterosensory canal system of the body to the anteriormost scales of the lateral line. Scale-eating (lepidophagy) has been de- scribed as a significant food habit in a va- riety of both freshwater and marine fishes inhabiting diverse regions of the world (see Sazima 1983 fora review). Within the fresh- waters of the Neotropical realm, this spe- cialized food habit has been described for a number of species belonging to the order Characiformes. Neotropical characiforms of the family Characidae for which lepi- dophagy has been previously described in- clude Cataprion mento Miller and Trosch- el (Kner 1860:34; Gosline 1951:54, Géry 1964:460) and Serrasalmus elongatus Kner (Goulding 1980:162) of the subfamily Ser- rasalminae, Exodon paradoxus Miiller and Troschel, Roeboexodon guanensis (Puyo), and the various species of the genus Roe- boides (Breder 1927:127; Géry 1964:459- 460) all presently assigned to the subfamily Characinae, and Probolodus heterostomus Eigenmann (Roberts 1970:384; Sazima 1977:510) and Bryconexodon juruenae Géry (1980:1) of the subfamily Tetragonopteri- nae. These taxa, with the exception of Ser- rasalmus elongatus, are characterized by specialized dentition, typically consisting of everted teeth, often mammilliform in over- all shape, which form very irregular series along the outer margins of the jaws. The outwardly directed teeth function in dis- lodging scales from the host species, with the removed scales then being ingested di- rectly if taken into the mouth, or gathered from the water column or substrate if knocked free (Sazima and Machado 1982). Although these dental modifications are characteristic for, and very similar in, the majority of lepidophagous Neotropical characiforms, such distinctive teeth have been implicitly hypothesized to have arisen independently in the diverse lineages of scale-eaters which are presently assigned to four different subfamilies of the Characidae (but see also comments under “Relation- ships’’). Recent collecting activities in the upper portions of the Rio Negro have revealed a previously undescribed genus and species of lepidophagous tetragonopterine characid described herein. This new form shares many of the dental modifications noted above as “‘typical’’ for lepidophagous Neo- tropical characiforms, but is quite distinc- tive in other attributes, most notably in its incompletely pored lateral line. Materials and methods.—A\ll measure- ments are given as proportion of standard length (SL) except for subunits of the head which are presented as proportions of head VOLUME 99, NUMBER 2 length. Vertebral counts were taken from radiographs, and cleared and counter- stained specimens. This number includes the four vertebrae incorporated into the Weberian apparatus and considers the fused PU,+U, as a single element. In the counts of median and pelvic fins, lower-case Ro- man numerals indicate unbranched fin rays, and Arabic numbers indicate branched fin rays. In the meristic values presented, the range for each measurement for the para- types and holotype is presented first, with the value for the holotype indicated in brackets. The following abbreviations are used for institutions: BMNH, British Museum (Nat- ural History), London; MBUCV, Museo de Biologia, Instituto de Zoologia Tropical, Universidad Central de Venezuela, Cara- cas; MZUSP, Museu de Zoologia da Uni- versidade de Sao Paulo, Sao Paulo; and USNM, National Museum of Natural His- tory, Smithsonian Institution , Washington, IDC Serrabrycon, new genus Diagnosis. —Tetragonopterine characid with mammilliform teeth in both jaws. Teeth of outermost tooth row of premaxilla directed anteriorly or anteroventrally rather than ventrally, forming irregular series. Sec- ond and fifth dentary teeth distinctly rotated anterodorsally. Lateral line pores present only on anterior seven or eight scales of lat- erosensory canal series. Scales in longitu- dinal series from supracleithrum to hypural joint 29 to 31. The combination of an incompletely pored lateral line, the relatively low number of scales in a longitudinal series, and the noted modifications of the dentition in the upper and lower jaws distinguishes Serra- brycon in the Characiformes. Type species. —Serrabrycon magoi, n. sp. Etymology. —Serrabrycon from the Lat- in, serra for saw, and Brycon a genus of Neotropical characiforms, in reference to the 329 Table 1.—Morphometrics of Serrabrycon magoi, new species. Standard length is in millimeters; measure- ments | to 11 are proportions of standard length; 12 to 16 are proportions of head length. Paratypes (37) Holo- type Range Average Standard length 27.5 21.3-31.8 1. Greatest body depth 0.29 0.28-0.30 0.288 2. Snout to dorsal-fin origin 0.53 0.53-0.57 0.540 3. Snout to anal-fin origin 0.68 0.68-0.71 0.689 4. Snout to pectoral-fin origin 0.30 0.29-0.33 0.319 5. Snout to pelvic-fin origin 0.52 0.51-0.53 0.520 6. Origin of dorsal fin to hypural joint 0.48 0.45-0.50 0.480 7. Pectoral-fin length 0.20 0.19-0.22 0.207 8. Pelvic-fin length 0.17 0.17-0.20 0.185 9. Length of base of anal fin 0.23 0.21-0.24 0.223 10. Least depth of caudal peduncle 0.12 0O.11-0.12 0.115 11. Head length 0.31 0.30-0.33 0.318 12. Snout length 0.24 0.23-0.27 0.248 13. Orbital diameter 0.39 0.36-0.40 0.375 14. Postorbital length 0.41 0.39-0.43 0.401 15. Length of upper jaw 0.46 0.45-0.48 0.466 16. Interorbital width 0.32 0.29-0.33 0.305 saw-like appearence of the outwardly point- ing teeth in the upper and lower jaws. Serrabrycon magoi, new species Figs. 1, 2, 3, Table 1 Holotype. —MBUCV 14270, 27.5 mm standard length (SL). Venezuela, Territorio Federal Amazonas, Departamento Rio Ne- gro, lower portion of Cano Manu, which drains into the Rio Casiquiare about 250 m upstream of Solano (approx. 02°00'N, 66°57'W); collected by R. P. Vari, C. J. Ferraris, Jr., O. Castillo, and J. Fernandez; 7 Dec 1984. Paratypes. —Taken with the holotype; 27 specimens: USNM 270260, 11 specimens, 21.5—27.5 mm SL (3 specimens cleared and 330 Fig. 1. counterstained for cartilage and bone); MBUCYV 14271, 10 specimens, 21.3-—25.4 mm SL; MZUSP 28749, 3 specimens, 23.5— 24.6 mm SL; and BMNH 1985.4.9:4—-6, 3 specimens, 21.3—23.5 mm SL. Venezuela, Territorio Federal Amazonas, Departa- mento Rio Negro, drying lagoon northeast of airport at San Carlos de Rio Negro (ap- prox. 01°55’N, 67°02’W); collected by A. Machado-Allison, R. P. Vari, C. J. Ferraris, Jr., J. Fernandez and O. Castillo; 4 Dec 1984: 10 specimens: USNM 270259, 5 specimens, 24.9-31.8 mm SL; and MBUCV 14272, 5 specimens, 24.7—26.5 mm SL. Diagnosis.— As for the genus. Description. —Table 1 gives morphomet- rics of holotype and paratypes. Body rela- tively slender, moderately compressed lat- erally. Greatest body depth at origin of rayed dorsal fin. Dorsal profile of body gently curved from tip of snout to interorbital re- gion, nearly straight from that point to in- sertion of rayed dorsal fin. Body profile at base of rayed dorsal fin straight, postero- ventrally sloped. Dorsal profile of body slightly convex from rear of insertion of last dorsal fin ray to caudal peduncle. Ventral profile of head distinctly convex from tip of fleshy lower jaw to below opercle; nearly straight from that point to anus other than for slight convexity at insertion of pelvic fins. Ventral surface of body transversely flattened anterior to insertion of pelvic fins. Base of anal fin straight or very slightly con- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Serrabrycon magoi, new species, holotype, MBUCV 14270, 27.5 mm SL. vex. Ventral profile of caudal peduncle slightly convex. Head obtusely pointed in profile; mouth terminal, lower jaw longer than upper. Up- per jaw extending posteriorly to vertical through anterior third of pupil. Mammil- liform teeth of outer row of premaxilla ex- tending through fleshy covering ofjaw. Nos- trils approximate; anterior round, posterior crescent-shaped. Eye relatively large. Fron- to-parietal fontanel extensive, completely separating parietals, frontals in contact only at epiphyseal bar. Fontanel wider poste- riorly, extending onto dorsomedial portion of supraoccipital. No supraorbital present. Four branchiostegal rays, three attached to anterior ceratohyal, one to posterior cer- atohyal. Anterior cartilage of basihyal sub- divided longitudinally into lateral halves. Pharyngeal teeth all unicuspidate; broad tri- angular patch of teeth on fifth ceratobran- chial; smaller groupings of teeth on fourth and fifth upper pharyngeal tooth plates, and on tooth plate associated with third infra- pharyngobranchial. Gill rakers elongate. Gill rakers on first gill arch 6 or 7+1+10 or 11. Outer row of premaxillary teeth consist- ing of four mammilliform teeth, reoriented anteriorly or slightly anterodorsally (Fig. 2). Tips of teeth in very irregular line. Second tooth from midline distinctly oriented more anterodorsally than others. Teeth of inner row on premaxilla tricuspidate, somewhat anteroposteriorly flattened, not mammilli- VOLUME 99, NUMBER 2 Fig. 2. Serrabrycon magoi: A, Left premaxilla, anterior view; B, Left premaxilla, lateral view; C, Left maxilla, lateral view, first and fourth teeth from bottom in process of replacement. form; tips oriented ventrally or slightly an- teroventrally. Five teeth in inner row. Max- illa with single series of 15 or 16 very slightly mammilliform teeth along entire anterior edge. Dorsalmost tooth in series somewhat mammilliform and laterally directed. Re- maining teeth having typical tetragonopter- in orientation, with tips approximately in plane of main body of maxilla; some very slightly mammilliform. Lower lip very fleshy, forming distinct pad. Teeth on dentary becoming progres- sively smaller posteriorly (Fig. 3). First four or five dentary teeth tricuspidate; lateral cusps relatively small. Remaining dentary teeth slightly mammilliform or conical. Sec- ond tricuspidate tooth distinctly rotated lat- erally relative to primary axis of first and third teeth. Fourth dentary tooth more lat- erally oriented than third, fifth more than fourth; sixth through eighth teeth progres- sively less laterally aligned. Remaining den- tary teeth vertically oriented. Scales cycloid. Scales in longitudinal se- ries between supracleithrum and hypural joint 29 to 32 [32]. Two or three series of scales extend beyond hypural joint onto base of caudal fin. Pores communicating with lat- erosensory canal system of lateral line de- veloped only on anteriormost seven or eight scales of longitudinal series. Scales in trans- verse series from origin of rayed dorsal fin to ventral midline 12 or 13 [13]. Rayed dorsal fin obtusely pointed, first and second branched rays longest, last un- branched ray slightly shorter. Dorsal-fin rays 332 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Serrabrycon magoi, left dentary, anterior portion: A, Lateral view; B, Dorsal view. 11,8 or 11,9 [11,8]. Adipose dorsal fin rela- tively well developed, length about one-half diameter of orbit; unscaled. Anal-fin rays ii1,15 or iv,15 or 16 [iv,15]; first two un- branched rays very short; last unbranched ray and first and second branched rays long- est, subequal. Margin of anal fin falcate; posteriormost ten branched rays about one- third to one-half length of longest rays. Pec- toral fin pointed, reaching to or slightly be- yond origin of pelvic fin. Pectoral fin rays 12 to 14 [13]. Pelvic fin reaching to origin of anal fin. Pelvic fin rays 1,6,1 [1,6,1]. Vertebrae: 31 (6), 32 (18), 33 (1). Coloration in alcohol.—Overall ground coloration of specimens fixed in formalin and preserved in alcohol light tan. Head with dense field of small chromatophores on upper lip, snout, and dorsal portion of head. Line of dark pigmentation along mar- gin of lower lip. Lateral surface of head with scattered chromatophores, largest on oper- cle Body with diffuse field of chromato- phores on lateral surface. Field more dense anteriorly, particularly in region of pored portion of lateral line. Chromatophore field extending to caudal peduncle, with discrete straight dorsal margin; field gradually thin- ning out ventrally in region between pec- toral and anal fin origins; reaching to ventral margin of body along base of anal fin and ventral surface of caudal peduncle. Scales dorsal of lateral chromatophore field with margins distinctly outlined by relatively wide series of small chromatophores. Over- all body pigmentation darkest along dorsal midline. Caudal fin with well developed field of dark pigmentation at base of caudal fin rays. Pigmentation patch extending anteriorly on middle rays to slightly posterior of hypural joint; smaller chromatophores extending posteriorly from main portion of pigmen- tation patch to tips of those rays. Series of very small chromatophores outlining other caudal-fin rays. Dorsal-fin rays outlined by series of small chromatophores; fin dusky distally. Distal portion of anteriormost branched anal fin-rays dusky. Rays of pec- toral and pelvic fins outlined by small chro- matophores. Adipose dorsal fin covered with scattered small chromatophores. Relationships. —Weitzman and Fink (1983) have recently summarized the diffi- culties and uncertainties associated with the present classificatory scheme applied to New World characiforms. Serrabrycon exempli- fies many of the problems inherent in the present taxonomy of the group. Under the traditional classification the genus could be assigned to the Hemigrammus-Pristella grouping of tetragonopterines that is char- acterized, in part, by an incompletely pored VOLUME 99, NUMBER 2 lateral line. However, no member of that assemblage has the distinctive dental mod- ifications characteristic of Serrabrycon. As noted by Fink and Weitzman (1974) and again by Weitzman and Fink (1983) an in- completely pored lateral line is a question- ably valid character for delimiting hypoth- esized monophyletic lineages within characids. The relationships of Serrabrycon could also be hypothesized to lie with gen- era such as Exodon, Roeboexodon, and Roeboides that authors have placed in the characid subfamily Characinae. An alter- native alignment of Serrabryon could be with Probolodus and Bryconexodon which have been assigned to the Tetragonopteri- nae. Although all of the genera just listed have completely pored laterosensory canal systems on the body, they are also charac- terized by mammilliform dentition similar to that of Serrabrycon. Furthermore the de- fining characters of the two subfamilies have not been based on derived characters sup- porting a hypothesis of their monophyly, and the present partitioning of the genera between the Characinae and Tetragonop- terinae is open to reinterpretation (Menezes in Sazima 1983:88). Thus Serrabrycon to- gether with Exodon, Roeboexodon, Roe- boides, Probolodus and Bryconexodon may constitute a monophyletic group delimited, at least in part, by the presence of everted mammilliform dentition. On the other hand the assemblage may be polyphyletic with the mammilliform dentition homopla- siously present in two or more lineages. Further research is necessary to resolve the above questions and to arrive at a hy- pothesis of the relationships between the contained species. Pending such a study and in light of the various uncertainties noted above, in particular the lack of a readily apparent sister group to the new species, a new genus of the Tetragonopterinae is pro- posed to contain the undescribed form. Etymology.—-magoi, in honor of Dr. Francisco Mago-Leccia of the Instituto de Zoologia Tropical of the Universidad Cen- 333 tral de Venezuela who has contributed sig- nificantly to our knowledge of the Vene- Zuelan fish fauna. Food habits.—Serrabrycon magoi has a very muscular stomach that is large relative to the remainder of the intestinal tract. Stomach content analysis of three individ- uals showed that the stomachs were com- pletely and exclusively filled with series of scales of different sizes. Ecology.—The holotype and 27 para- types were collected in the lower portion of Cano Manu, a very slow-flowing black water tributary of the Rio Casiquiare. The stream was well-shaded by rain-forest canopy, and had few small emergent plants and no float- ing vegetation. The remaining ten paratypes were collected in the shallow black waters of a small sluggish stream running through the center of a drying lagoon. That locality was characterized by large amounts of sub- merged logs and detritus. The site had nu- merous large emergent plants, but was dis- tant from the rain-forest canopy and poorly shaded. Resumen.—Serrabrycon magoi, un gé- nero y una especie no descritos anterior- mente que comprende pequenos peces comedores de escamas pertenecientes a la subfamilia Tetragonopterinae, Characidae; son descritos a partir de muestras proven- lentes de aguas negras del sistema del Rio Negro de la region del Rio Casiquiare, Ven- ezuela. El género y especie son unicos en la com- binacion de los siguientes caracteres: la pre- sencia de una serie de dientes en las man- dibulas superior e inferior orientados exteriormente; el numero relativamente bajo de escamas en la serie longitudinal hasta la union hipural y en la limitacion de poros del sistema del canal laterosensorial a las escamas anteriores de la linea lateral. Acknowledgments The specimens that served as the basis for this study were collected by a joint MBUCV-USNM expedition. I thank Dr. 334 Antonio Machado-Allison (MBUCV) for his efforts in organizing the expedition and as- suring its success. He along with Mr. Carl J. Ferraris Jr., Lic. Justa M. Fernandez, and Lic. Otto Castillo enthusiastically partici- pated in all aspects of the collecting efforts. Mr. Andrew G. Gerberich, Mr. Kurt A. Bru- welheide, and Ms. Ann M. Williams pro- vided technical assistance at USNM. The Spanish translation of the ““Resumen’”’ was provided by Sr. Hernan Ortega. Figure 1 was prepared by Mr. Theophilus Britt Gris- wald. The collecting expedition was made possible by a grant from the Scholarly Stud- ies Program of the Smithsonian Institution. Research associated with this study was supported in part by the I.E.S.P. Neotrop- ical Lowland Research Program of the Smithsonian Institution. This paper was improved by the comments and suggestions of Dr. Stanley H. Weitzman, Dr. Wayne C. Starnes, and Mr. Carl J. Ferraris, Jr. Literature Cited Breder, C. M. 1927. The fishes of the Rio Chucu- naque drainage, eastern Panama.—Bulletin of the American Museum of Natural History 57(3): 91-176. Fink, W. L.,andS.H. Weitzman. 1974. Theso-called cheirodontin fishes of Central America, with de- scriptions of two new species (Pisces: Charac- idae).—Smithsonian Contributions to Zoology 172:1-46. Géry,J. 1964. Poissons characoides nouveaux ou non signalés de Vilha do Bananal.—Vie et Milieu, supplement 17 (Volume Jubilaire dédié a Georges Petit):447—471. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1980. Un nouveaux poisson characoide oc- cupant la niche des mangeurs d’écailles dans le haut rio Tapajoz, Brézil.—Revue frangaise de Aquariologie 7:18. Gosline, W. A. 1951. Notes on the characid fishes of the subfamily Serrasalminae.— Proceedings of the California Academy of Sciences 27(2):17— 61. Goulding, M. 1980. The fishes and the forest. Uni- versity of California Press, Berkeley, 280 pp. Kner, R. 1860. Zur Familie der Characinen, II.— Denkschriften der Akademie der Wissenschaft- en, Wien 18:9-62. Roberts, T. 1970. Scale-eating American characoid fishes, with special reference to Probolodus het- erostomus. —Proceedings of the California Academy of Sciences 38(20):383-390. Sazima, I. 1977. Possible case of aggressive mimicry in a Neotropical scale-eating fish. — Nature 270: 510-512. 1983. Scale-eating in characoids and other fishes. —Environmental Biology of Fishes 9(2): 87-101. , and F. A. Machado. 1982. Habitos e com- portamento de Roeboides prognathus, um peixe lepidofago (Osteichthys, Characoidei).—Bole- tim do Zoologia, Universidade de Sao Paulo 7: 36-56. Weitzman, S. H., and W. L. Fink. 1983. Relation- ships of the neon tetras, a group of South Amer- ican freshwater fishes (Teleostei, Characidae), with comments on the phylogeny of New World Characiforms.—Bulletin of the Museum of Comparative Zoology 150(6):339-395. Department of Vertebrate Zoology (Fish- es), National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 335-346 NEMATOCHARAX VENUSTUS, A NEW GENUS AND SPECIES OF FISH FROM THE RIO JEQUITINHONHA, MINAS GERAIS, BRAZIL (TELEOSTEI: CHARACIDAE) Stanley H. Weitzman, Naércio A. Menezes, and Heraldo A. Britski Abstract. —Nematocharax venustus, a new genus and species, is described from the lower portion of the Rio Jequitinhonha in Minas Gerais, Brazil. The adult male of this species is distinguished from all known characins by the combination of elongate, filamentous dorsal, anal, and pelvic fins, and a nearly full series of maxillary teeth. The possible relationships of the new genus are briefly discussed but no corroborated hypothesis of relationship is advanced. The status of the knowledge of the fish fauna of the Atlantic forest of eastern Brazil is briefly discussed. The description given below of a new species of characid from the Rio Jequitinho- nha is a small contribution towards increas- ing the data base for studies of the biogeog- raphy of the small and modest-sized littoral streams of eastern and southeastern Brazil. Although the characid fish described herein is probably quite common in at least the middle and lower course of the Rio Jequi- tinhonha it apparently has been collected only twice, first in 1966 from the Rio Je- quitinhonha at Itaobim, Minas Gerais and again in 1985 further east in the same river near Salto da Divisa. It was very common at both collecting sites. That a relatively common, distinctive, easy-to-capture fish of eastern Brazil is now being made known to science is a fact representative of our poor knowledge of the fish fauna of the region. Much future descriptive, phylogenetic, and faunal research must be published before adequate biogeographical studies of this re- gion’s fishes can be accomplished. Methods and materials. —Counts and measurements are those described by Fink and Weitzman (1974:1-2). All measure- ments other than standard length (SL) are expressed as a percentage of SL except sub- units of the head which are expressed as a percentage of head length. Ranges and means of the meristics and morphometrics are giv- en in the tables, with information for the holotype given separately. Ranges and means for counts in the tables are repre- sented by one set of figures for both sexes combined except in cases of obvious sexual dimorphism where the information is given by sex. Logarithmic and square root trans- formed data were used in covariance anal- yses of sexual differences in measurements and counts respectively. Specimens examined for this study are deposited in the Museu de Zoologia da Uni- versidade de Sao Paulo (MZUSP), the Na- tional Museum of Natural History, Smith- sonian Institution (USNM), Museum of Zoology, University of Michigan (UMMZ), the Academy of Natural Sciences of Phila- delphia (ANSP), the British Museum (Nat- ural History) (BMNH), Field Museum of Natural History (FMNH), and the Museu Nacional de Rio de Janeiro (MNRJ). Nematocharax, new genus Type species.—Nematocharax venustus, new species. Diagnosis.—A “‘tetragonopterine”’ char- acid distinguished from all other genera of American and African characids by having a combination of elongate branched dorsal-, 336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Morphometrics of Nematocharax venustus, Itaobim, Rio Jequintinhonha, holotype MZUSP 5131, paratypes from same locality, MZUSP 28480-28509, 28661-28689, and USNM 232955 and 232956. Standard length expressed in mm. Following 19 measurements are percentages of standard length and last 4 measurements are percentages of head length. Entries including both sexes and juveniles (n = 52 to 75) are for specimens 17.5 to 32.0 mm SL. Entries listed by sex include males (m) between 30.0 and 52.0 mm SL and females (f) between 33.0 and 40.5 mm SL. Holotype n Low High iG Standard length 50.5 75 N17 52.0 — Depth at dorsal-fin origin m 40.6 Di 37.5 43.0 41.0 Depth at dorsal-fin origin f — 30 36.0 40.4 38.6 Snout to dorsal-fin origin S357 US 50.8 55.8 53.4 Snout to pectoral-fin origin 26.5 75 24.7 31.5 27.4 Snout to plevic-fin origin 44.2 75 41.3 52.4 45.3 Snout to anal-fin origin 60.4 27 56.9 6222 59.6 Caudal peduncle depth 13.1 75 10.8 13.5 12a Caudal peduncle length ES WS 9.9 13.5 11.2 Pectoral-fin length 23.8 74 14.7 26.2 232 Pelvic-fin length m SES 23 40.2 64.2 54.8 Pelvic-fin length f — 30 20.0 30.9 23.2 Dorsal-fin base length 14.9 75 13.0 17.0 15.0 Longest dorsal-fin ray length m 73.3 24 59.9 78.8 68.2 Longest dorsal-fin ray length f —_ 30 32.9 36.7 34.7 Anal-fin base length m 333}.3" 27 30.0 34.9 33.1 Anal-fin base length f — 30 28.1 3255 30.6 Anal-fin lobe length m 50.3 24 41.0 58.5 50.2 Anal-fin lobe length f — 29 22.4 34.5 26.3 Bony head length Ast iS 26.4 SKS 28.4 Horizontal eye diameter 35.9 75 34.8 43.0 38.4 Snout length 27.9 WS 22.9 33.1 27.6 Least width interorbital 30.9 74 27.6 36.4 30.8 Upper jaw length 47.9 74 36.4 51.4 44.1 anal-, and pelvic-fin rays, two rows of pre- maxillary teeth in adults, and an almost complete row of teeth along the free ventral maxillary border. The possible relationships of this genus to other characids are obscure and we rec- ognize the genus Nematocharax because we are unable to hypothesize a particular rel- ative of the single known species, N. ven- ustus (see ““Relationships.”’) Etymology. —Nemato from the Greek for thread and charax from the Greek characo meaning a pointed stake, here used in ref- erence to the characid genus Charax, hence a characid fish. Nematocharax thus means a thread-bearing characid fish in reference to the long threadlike extensions of the dor- sal, anal, and pelvic fins. Nematocharax venustus, new species Figs. 1-4, Tables 1, 2 Holotype. —MZUSP 5131, male, SL 50.5 mm, Brazil; Minas Gerais, Municipio Me- dina, Rio Jequitinhonha at Itaobim, about 16°40’'S, 41°23’W, 25 Jun 1966, by Heraldo Britski and Naércio Menezes. Paratypes.—1, MZUSP 28480, female (photographed), SL 35.4 mm.—29, MZUSP 28481-28509, SL 17.7-52.0 mm.—239, MZUSP 28661-28689, SL 33.0-51.5 mm.—80, MZUSP 28690 (data not tak- en).—6, USNM 232955, SL 34.0-51.7 mm.—7, USNM 232956, SL 34.7-52.0 mm.—4, UMMZ 213448, SL 34.7-51.9 mm.—2, ANSP 157373, SL 34.3-48.0 mm.—2, BMNH 1985.11.10.1—2, SL 37.5- VOLUME 99, NUMBER 2 337 Table 2.—Meristics of Nematocharax venustus, Itaobim, Rio Jequitinhonha, holotype MZUSP 5131, paratypes from same locality, MZUSP 28480-28509, 28661-18689, and USNM 232955 and 232956. Holotype n Low High x Branched dorsal-fin rays 9 73 9 10 9.0 Branched pectoral-fin rays 12 70 11 14 11.8 Branched pelvic-fin rays 6 73 7 7 7.0 Branched anal-fin rays 23 73 Dy) 26 B31 Branched caudal-fin rays 17 73 17 17 17.0 Horizontal scale rows, between dorsal- and anal-fin origins 13 73 11 13 12.7 Predorsal scales 11 68 10 12 10.8 Perforated lateral-line scales 10 66 8 12 9.7 Lateral series scales 34 66 32 35 33.9 Horizontal scale rows around caudal peduncle 14 52 13 14 14.0 Dorsal limb gill rakers 6 73 5 7 6.2 Ventral limb gill rakers 9 73 8 10 9.4 Outer row premaxillary teeth 4 73 yD, 4 3.1 Inner row premaxillary teeth 5 73 4 6 5.1 Maxillary teeth m 12 31 7 12 10.8 Maxillary teeth f — 27 9 12 10.0 Dentary teeth 11 73 9 13 11.0 Vertebrae — 36 33 34 33.7 50.3 mm.—2, FMNH 96633, SL 42.8-49.4 mm.—4, MNRJ 11.285, SL 33.6-—49.6 mm. All preceding paratypes with same collec- tion data as holotype.— 15, USNM 270816, 29.5—41.7 mm, Brazil, Minas Gerais, Muni- cipio Jequitinhonha, Rio Jequitinhonha, near km 205, road between Salto da Divisa and Jacinto, about 16°02’S, 40°00’W, 22 Mar 1985 by Naércio Menezes, Ricardo Castro, Marilyn Weitzman, and Stanley H. Weitz- man.—18, MZUSP 28811, SL 23.0-41.5 mm, with same data as USNM 270816. Diagnosis. —Since Nematocharax is monotypic, the species and the generic di- agnoses are the same. See also discussion below under “Relationships.” Description. —See Tables | and 2 for mor- phometric values and counts. Body mod- erately deep, sides compressed, greatest depth at dorsal-fin origin. Predorsal body profile gently convex in females and juve- niles, more steeply arched in males (com- pare Figs. 1 and 2). Body profile somewhat elevated at dorsal-fin origin, nearly straight along dorsal-fin base and continuing almost in straight line in females and juveniles to origin of adipose fin. In mature males, pro- file between posterior dorsal-fin base and adipose-fin origin slightly convex. Dorsal- fin origin about equidistant between caudal- fin base and snout tip. Dorsal-fin origin closer to posterior border of eye than to cau- dal-fin base. Ventral body profile gently convex to anal-fin origin in both sexes. Body profile along base of anal fin slightly convex to nearly straight in males, almost straight in females. Both dorsal and ventral profiles of body of distal to anal-fin border concave in both sexes. Head and snout relatively deep. Jaws about equal, mouth terminal or lower jaw slightly included, ventral to upper jaw. Gape nearly horizontal. Maxilla extending pos- teriorly approximately to point on line drawn vertically at anterior one-third to one- half of pupil of eye. Dorsal fin with last ray not split to its base. First to fourth branched dorsal-fin rays filamentous and much more elongate in males than females (see Table 1), rays ex- 338 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Gerais, Municipio Medina, Rio Jequitinhonha at Itaobim. tending posteriorly beyond caudal-fin base. In females these rays occasionally reaching as far posteriorly as anterior origin of adi- pose fin. Branched rays two and three usu- ally shorter than branched rays one and four; may occasionally equal rays two and three in length. Dorsal fin with no well-developed hooks but with small bony spinules scat- tered distally on branched rays two to six. Any given ray with more spinules on pos- terior borders of posterior branches than on anterior branch. (For example, a cleared and stained male, USNM 232955, 47.6 mm SL with 2 spinules on anterior branch, 10 on posterior branch of ray two; rays three and four damaged; ray five with 2 spinules on anterior branch, 3 on posterior branch; ray six with 2 on anterior branch and 4 on pos- terior branch; ray seven with | on anterior branch and 6 on posterior branch.) Adipose fin present, situated at position bisected by or slightly anterior to imaginary line drawn vertically from posterior anal- fin base termination. Posterior anal-fin ray split to its base. Anal fin of both sexes with broad anterior lobe (Figs. 1-3). Anal-fin rays of anterior lobe Nematocharax venustus, new species, holotype, MZUSP 5131, male, 50.5 mm SL; Brazil, Minas much longer in males than in females, reaching posteriorly beyond caudal-fin base. Males with fourth unbranched anal-fin ray and first to about sixth branched anal-fin rays forming major part of anal-fin lobe. Fifth branched anal-fin ray usually longest. See Table 1 and section on sexual dimor- phism for comparison of anal-fin lobe lengths of males and females. Males with hooks present on third, fourth and/or fifth unbranched and on first to tenth or eleventh branched rays; see Fig. 3 of anterior un- branched and first six branched rays. Hooks small, 1 to 2 per ray segment. Hooks nu- merous (more than 10) on unbranched ray four or five and also on branched rays one to six or seven. Rays eight to eleven with 1 to about 3 hooks per ray, with 1 or 2 hooks on hook-bearing segments. Remaining rays may occasionally bear single spinule. Hooks occur near or on those divided, Y-shaped, segments forming initial branching point for each ray. Tiny spinules continuing on branched portions of rays five to six, espe- cially one to four, most common on pos- terior branch of each ray. Distal tips of pectoral-fin rays of males VOLUME 99, NUMBER 2 339 Fig. 2. Nematocharax venustus, new species, paratype, MZUSP 28480, female, 35.4 mm SL; Brazil, Minas Gerais, Municipio Medina, Rio Jequitinhonha at Itaobim. and females extending posteriorly beyond pelvic-fin origin. Pelvic fin sexually dimor- phic, with first unbranched ray greatly elon- gate in males. See discussion on sexual di- morphism below. Pelvic fin with about | to 5 spinules present distally on most branches of rays two to five. Principal caudal-fin rays 10/9 in all counted specimens (73). Hooks and spinules absent on caudal-fin rays. Scales cycloid, lateral line complete. Pre- dorsal scale series complete. See Table 2 for scale counts. Premaxillary teeth in two distinct rows (Fig. 4). Two to four, usually three, teeth in outer row (see Table 2); first to fourth teeth of outer row cusped, with lateral most tooth having one to two cusps, and medial tooth up to four cusps. Four to six, usually five, teeth in inner row; inner row teeth with five or six, usually six, cusps. Outer row teeth circular in cross section, not compressed. Inner-row teeth compressed, especially dis- tally along cusps. Maxillary teeth five to twelve, x = 9.5 in 73 specimens 17.7 to 52.0 mm SL. Adult specimens over 33.0 mm SL, with seven to twelve teeth, x = 10.5. Larger specimens usually having more teeth. See also discussion on sexual dimorphism. Anterodorsal four to five maxillary teeth with three or four cusps, remaining pos- teroventral teeth large, strong, and unicus- pid. Dentary teeth nine to thirteen, in single row. Anterior six to eight teeth large, with three to five cusps; posterior five to seven teeth usually unicuspid and noticeably smaller than anterior teeth. See Figure 4 of jaws and teeth. Color in alcohol. —See Figs. 1 and 2 for preserved color pattern of males and fe- males respectively. Entire body pale yellow- ish brown (muscle tissue color) in specimens with guanine destroyed by formalin. Body silvery in specimens with guanine pre- served. Series of small chevron-shaped marks occurring along horizontal junction of dorsal and ventral myomeres. These marks especially distinct in region posterior to middle of body cavity and anterior to dark shield-shaped caudal spot. Caudal spot extending posteriorly and terminating at distal tips of three or four middle caudal- fin rays. Caudal spot tapering anteriorly to narrow Stripe and then terminating anterior to imaginary vertical line through posterior 340 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Nematocharax venustus, USNM 232955, male, 47.6 mm SL, cleared and stained. Anal fin, lateral view, right side. end of anal-fin insertion. Dorsal body sur- face dusky in area of predorsal scales, along base of dorsal fin, and along median part of back to base of dorsal caudal-fin lobe. Head dusky brown dorsally, pale ventrally, no dark markings. Dark crescent-shape mark on opercle in Figs. 1 and 2 formed by shadow within gill chamber, not from pigment. Bel- ly pale, without dark marks. Dorsal fin dusky in both sexes, with fil- amentous rays of males especially dark. Caudal fin dusky with pale brown chro- matophores on fin rays. Anal fin dusky in both sexes with filamentous rays of males especially dark. Pelvic and pectoral fins slightly dusky, brown chromatophores few, scattered over rays. Color in life.—Specimens taken from near Salto da Divisa in silty waters with very little visibility had body silvery with dark chevron-shaped marks pale, partly ob- scured by guanine pigment on body. Body with pale pink and greenish reflections on silvery sides. Back pale brown, top of head dark brown. Dorsal fin with same dark pat- tern as preserved specimens but with pale rosy color in fin rays. Adipose fin pale rosy. Dorsal caudal-fin lobe pale yellow distally and pale rosy proximally. Ventral caudal- fin lobe with similar color but somewhat yellowish throughout. Anterior lobe of anal fin in both sexes pale rosy, posterior part of anal fin pale yellow. Pectoral fin hyaline, pelvic fin pale rosy. Belly, sides, and side of head bright silvery flushed with small amount of rose color. Iris of eye silvery with rosy dark spot dorsally. In clear waters or in dark tea-colored waters which may have once been common in the tributaries of the lower regions of the Rio Jequitinhonha, the colors of this fish may have been darker and/or more intense. Sexual dimorphism.—Nematocharax venustus is obviously sexually dimorphic in three characters. Table 1 shows gaps in ranges of ratios between males and females for the length of the longest dorsal-fin ray (measured between the origin of the fin and the distal tip of the longest ray), anal-fin lobe length, and pelvic-fin length. Measurements were taken from adult specimens between 33.3 to 52.0 mm SL for males and 33.3 to 40.5 mm SL for females. Inspection of upper jaw length relative to head length in adult specimens shows a greater jaw length in males than in females. VOLUME 99, NUMBER 2 The percent of upper jaw length relative to head length in 30 males (standard length 33.3 to 52.0 mm) from Itaobim ranged from 45.5 to 51.4, x = 47.7. Twenty-seven fe- males from the same locality (with standard lengths of 33.3 to 40.5 mm) ranged from 39.0 to 45.9, X = 42.5. Analysis of covari- ance by sex using logarithmic transforma- tions of original data for upper jaw and head lengths of the same population samples showed no significant difference in regres- sion coefficients (slopes), t = 0.696, P > 0.05, determined in a two-tailed test but did show significant differences in the Y-inter- cepts (elevations), t = 2.600, P < 0.001, also determined in a two-tailed test. The upper jaw length differences in adult males and females may be due to a growth rate difference during sexual maturation since specimens of shorter standard length were not separable into two groups on the basis of upper jaw length. Inspection of the population sample of Nematocharax venustus from Itaobim showed that ten juvenile specimens between 17.7 and 32.8 mm SL had 5 to 7 maxillary teeth, and a mean of 5.8. Twenty-seven adult males had 7 to 12, and a mean of 10.8 max- illary teeth. Thirty-one adult females be- tween 33.3 and 40.5 mm SL had 9 to 12, and a mean of 10.0 maxillary teeth. Co- variance analysis of the same two adult samples by sex using logarithmic transfor- mations of upper jaw length measurements and square root transformations of maxil- lary tooth numbers showed a difference in regression coefficients (slope), t = 2.117, P < 0.05 and in the Y-intercept (elevations), t = 2.003, P < 0.05, both determined in a two-tailed test. In Nematocharax venustus there does seem to be some sexual dimor- phism in maxillary tooth numbers. Sexual dimorphism in maxillary (or oth- er) tooth number is apparently rare in te- tragonopterine characids, especially in those species with one to three or so teeth in each maxilla. When the count is higher, maxil- lary tooth count differences might be ex- 341 Fig. 4. Nematocharax venustus, USNM 232955, male, 47.6 mm SL, cleared and stained. Lateral view, right side of premaxillary, maxillary, and dentary bones. pected to display sexual dimorphism, es- pecially if the maxilla is longer in males than in females. At some stage in the develop- ment of all tetragonopterines with maxillary teeth there are fewer teeth (or none) in the young than in older specimens. In species with many maxillary teeth and long max- illae the increase in tooth numbers and in maxillary length may continue evenly over a relatively long interval of growth and then differentiate during sexual maturation. Body depth in Nematocharax venustus increases at a greater rate relative to body length during growth. Males and females are evidently sexually dimorphic in the ratio of body depth to standard length. In the pop- ulation from Itaobim, ten juveniles 17.7 to 32.0 mm SL had a body depth ratio of 33.0 to 37.3 percent of the standard length with a mean of 34.8 percent. In thirty-one fe- males of 33.3 to 40.5 mm SL the rate ws 36.0 to 40.4, with a mean of 38.6 percent while in twenty-seven males of 33.3 to 52.0 342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mm SL the rate was 37.5 to 43.0, with a mean of 41.0 percent. Covariance analysis using logarithmic transformations of origi- nal measurements of these same males and females showed no differences in regression coefficients (slopes), t = 0.000, P > 0.05 and Y-intercepts (elevations), t = 0.256, P > 0.05. It would seem that differences are size related and that females appear to be less deep only because they do not reach as large a size as the males in the two population samples at hand (Itaobim and Salto da Div- isa). Relationships. —The phylogenetic rela- tionships of Nematocharax venustus re- main obscure. A combination of a com- pressed, relatively deep body form, long dorsal fin and a nearly fully toothed maxilla is also found in members of the Rhoadsi- inae from western Ecuador, Colombia, Pan- ama (an undescribed species), Costa Rica, and Lake Nicaragua in Nicaragua; see Ei- genmann and Myers (1929:457), Dahl (1960:472), and Fink and Weitzman (1974: 26). These fishes, geographically distant from Nematocharax, lack the derived, elongate anal- and pelvic-fin rays of Nematocharax. Also, Nematocharax with a vertebral count of 33-34 differs from the higher vertebral count, 36-38, of species of the Rhoadsiinae [based on specimens of Rhoadsia altipinna Fowler (FMNH 83870), Parastremma sad- ina Eigenmann (FMNH 56023), Carlana eigenmanni (Meek) (USNM 272646) and specimens of an undescribed species of Car- lana collected recently in Panama (USNM uncatalogued)]. Furthermore, Nematochar- ax venustus does not have the “‘cheirodon- tine” teeth (flattened and with a single series of several more or less subequal cusps) found in young specimens and as at least partly present in adults of the Rhoadsiinae, Eigen- mann and Myers (1929:457), Fink and Weitzman (1974:25). Small specimens, down to 17.7 mm SL, of Nematocharax venustus have teeth similar to those of adults, relatively elongate, rather flattened and with cusps of different sizes on each tooth. The geographical distance between Nemato- charax and members of the Rhoadsiinae make it unlikely that Nematocharax ven- ustus is most closely related to members of Rhoadsiinae, but the possibility cannot be ignored. If Nematocharax is most closely related to the Rhoadsiinae it is less derived in hav- ing fewer conical teeth in the maxilla than members of the Rhoadsiinae, having no “‘cheirodontine”’-like teeth at any life his- tory stage, and possibly in vertebral num- bers as described above. Nematocharax does have its own derived features relative to the Rhoadsiinae. These are the elongate anal and pelvic fins. The color pattern of Nematocharax ap- pears less derived, having a caudal spot and chevron-shaped marks along the midsides similar to those of many tetragonopterine characids. It lacks the bright red color of some (all?) of the Rhoadsiinae and the large midside blotch (posterior humeral spot?) in life of some of the Rhoadsiinae. At this time, little phylogenetic information can be gath- ered from the color patterns of characid fish- es since so little of that system has been cladistically analyzed. Only in those cases where a species or a group of species have a unique (not found in any other taxa) color pattern as in Paracheirodon (see Weitzman and Fink 1983:354) can color pattern cur- rently be used in a cladistic analysis of char- acids. Another fish, Hyphessobrycon elachys M. Weitzman (1985) from Paraguay, has elon- gate dorsal- and pelvic-fin rays. This fish differs from Nematocharax venustus in hav- ing a different color pattern, derived ante- rior jaw teeth in which the central cusp ex- tends in an opposite plane from the lateral cusps, and a small adult standard length of nearly 20.0 mm. The only evidence sug- gesting a possible relationship with N. ven- ustus are the elongate dorsal and pelvic fins, a character we suspect is convergent. Again, VOLUME 99, NUMBER 2 phylogenetic studies are necessary to ex- amine possible relationships between the two species. Other “‘tetragonopterine characids”’ have long fins. The glandulocaudine characid Ge- phyrocharax martae Dahl (1942:4) from northern Colombia has an elongate anterior anal-fin lobe that becomes filamentous. Other species of glandulocaudines such as Corynopoma riisei and Pseudocorynopoma doriae tend to have fin rays produced or to have entire fins elongate. There is no evi- dence that Nematocharax venustus is a glandulocaudine characid since it lacks the glandular tail structure of these fishes. Some species of “‘rosy” tetras in the genera Mega- lamphodus Eigenmann (1915:50), and Hy- phessobrycon, Weitzman (1977:335) have relatively elongate dorsal and anal fins which are not, however, filamentous as in Ne- matocharax venustus. Sometimes speci- mens of Hyphessobrycon bifasciatus also have a somewhat elongate dorsal fin and somewhat filamentous anterior pelvic-fin rays, but these are never as long as in Ne- matocharax venustus. It is our opinion that elongate fins in characids probably evolved independently several times and that for such fins to be utilized as synapomorphies relating species and genera, the morpholog- ical similarity of the fins should involve ho- mologous rays. Such characters would be best used when they are congruent with oth- er synapomorphies indicating the mono- phyletic relationships of particular taxa. Two genera, Hollandichthys Eigenmann (1910) and Rachoviscus Myers (1926) have species which, like Nematocharax venustus, bear well-toothed maxillae. Since these species are part of the Atlantic forest fish fauna, relatively close geographically to the known localities for NV. venustus, they should be considered as possible candidates for re- lationships with Nematocharax. Weitzman and Cruz (1981:999) briefly discussed some possible relationships of Rachoviscus and concluded that the species 343 were “‘tetragonopterine”’ rather than “‘chei- rodontine” characids as suggested by Myers (1926:1). Weitzman and Cruz (1981:1003) were unable to corroborate any phyloge- netic hypotheses of the relationships of Ra- choviscus to specific “‘tetragonopterine”’ genera. In addition to a rather heavily toothed maxilla, the two species of Ra- choviscus have a deep red adipose fin. The adipose fin of Nematocharax venustus is a pale rosy color but has none of the deep red pigment found in the species of Rachovis- cus. Considering the differences in these pigments we cannot consider the adipose fin color of Rachoviscus and Nematocharax a synapomorphy based on similar color pat- tern, although the rosy color could be con- sidered a transition state between no red color and deep red. The details of the jaw morphology of Ne- matocharax are different from those of Ra- choviscus. Nematocharax venustus has the mouth horizontal, not angled. Both species of Rachoviscus have somewhat upturned mouths with the gape angled posteroven- trally (Weitzman and Cruz 1981:1005, 1009). The condition in Rachoviscus is probably more derived and may represent a synapomorphy for the species of that ge- nus, although it must be kept in mind that an angled jaw is not uncommon in other ‘‘tetragonopterine” characids. The maxilla of Nematocharax venustus is well toothed (x = 10 teeth) (Fig. 3 and Table 2), while Rachoviscus graciliceps Weitzman and Cruz has 3 to 8 teeth and Rachoviscus crassiceps Myers has 8 to 14 teeth. Evolution ofa well- toothed maxilla may have taken place with- in Rachoviscus crassiceps and Nematochar- ax independently. We have at present no way of evaluating the cladistic significance of maxillary teeth in these genera since the character appears in several other “‘tetra- gonopterine”’ genera and we have found no other synapomorphies possibly indicative of a close relationship between Nemato- charax and Rachoviscus. 344 The monotypic Hollandichthys multifas- ciatus Eigenmann and Norris (1900:358) might be related to Nematocharax venustus, both having well-toothed maxillae. Because of its nearly fully toothed maxillae Hollan- dichthys multifasciatus was considered a species of Pseudochalceus by Schultz (1966: 26) and Géry (1972:30). The relationships of Hollandichthys and putatively related characids assigned to Pseudochalceus were reviewed by Schultz (1966:26), and Géry (1972a:24-25; 1972b:933-942). We here make no comment on these reviews pending a cladistic treatment of these and other, pos- sibly related fishes. Hollandichthys multi- fasciatus, apparently confined to Atlantic coastal streams between and including the states of Sao Paulo and Santa Catarina, has a maxilla with up to 17 teeth. There are several differences between Hollandichthys and the other species assigned to Pseudo- chalceus, Géry (1972a:37; 1972b:942; 1977: 414). Considering these differences and the geographical isolation of Hollandichthys (southeastern Brazil) from other species as- signed to Pseudochalceus (western Ecuador and Colombia) we propose to retain the name Hollandichthys until a cladistic phy- logenetic study demonstrates monophyly of these geographically disparate genera. At present we see no more reason to relate Ne- matocharax venustus to Hollandichthys multifasciatus or to any species of Pseudo- chalceus than to relate Nematocharax to the species of Rachoviscus. The only possible synapomorphy found by us relating these genera and species is a high maxillary tooth count. For some taxa, a high maxillary tooth count is undoubtedly a synapomorphy in- dicating monophyly of a group formed by these taxa; for example, the species of Pseu- dochalceus from Ecuador recorded by Géry (1972). But high maxillary tooth counts have appeared a number of times in characids, for example in Rachoviscus Weitzman and Cruz (1981:1000) and xenurobryconin characids subgroup C, Weitzman and Fink (1985:55). Because of the possibility that PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Nematocharax may have evolved its max- illary teeth independently and especially be- cause we have been unable to find any other synapomorphies common to Nematochar- ax and other characid genera with high maxillary tooth counts we are unable to ac- cept a hypothesis that Nematocharax forms a sister group with any of them. Without extensive and detailed morpho- logical and phylogenetic research, we have no fully tested hypotheses concerning the relationships of Nematocharax venustus. It appears to be a “tetragonopterine” characid but that group itself is not currently defined by synapomorphies (see Weitzman and W. Fink (1983:342) and Weitzman and S. Fink (1986) for our opinions about the mono- phyly of the Tetragonopterinae). Etymology. —The name venustus is from the Latin Venus, meaning like Venus, hence beautiful, elegant, or graceful. The name is given in reference to the graceful flowing shape of the dorsal, anal, and pelvic fins of this fish. Remarks.—The yet-to-be-accomplished studies of fish biogeography and fish species composition of the streams in areas for- merly occupied by the Atlantic forests of eastern and southeastern Brazil have now been made more difficult by inadequate sampling in the past and by much habitat alteration within the last eighty years and especially within the last thirty years. It is no surprise that new species of fishes have been collected and described from many of the Atlantic forest areas in recent years. For example, see Britski and Ortega (1983), Buckup (1981), Garavello (1977), Nijssen and Isbriicker (1976, 1980), and Weitzman and Cruz (1981). The Rio Jequitinhonha, a relatively large coastal river, has been very little explored ichthyologically, and unfortunately for sys- tematic and biogeographic ichthyological research, in part has been long subject to habitat alteration by man’s agricultural ac- tivities. Arising in the cerrado area of the Serra do Espinhaco of Minas Gerais, the VOLUME 99, NUMBER 2 Rio Jequitinhonha flows north then east through Minas Gerais and Bahia to the At- lantic Ocean. In parts of Minas Gerais and in Bahia this river originally traversed the Atlantic forest. Only about two percent of the forest area remains, the rest is now es- sentially replaced by agriculturally devel- oped land. A combination of cattle grazing, farming, and pulpwood plantations of pine and eucalyptus trees constitute most of the land use along the coastal plain. The ecology of the middle and lower Rio Jequitinhonha drainage is greatly altered from its preagri- cultural condition and this has undoubtedly greatly affected the composition of the fish fauna. At the present time this river bears a heavy silt load, at least during the rainy season. Furthermore, in 1985 two of us (Menezes and Weitzman) observed an in- troduced species of the African cichlid ge- nus Tilapia to be common in the vicinity of Salto da Divisa and we suspect this fish is Common in most parts of the river. 77- lapia, just as other factors influencing the altered ecology, undoubtably has affected the relative composition of the fish fauna in the Rio Jequitinhonha. We will never know the preagricultural composition of the fish fauna of this river. Some species, less adapt- able to the altered ecology of the river, may have become rare or extinct. Other species, not impacted by Tilapia, adaptable to a heavy silt load, increased sunlight and prob- ably to higher water temperatures, and to a non-forest water chemistry, especially in the small tributaries in the lower courses of the river, may now be far more common than formerly. Eventually some presumptive measure of the original fish fauna of middle and lower parts of the Rio Jequitinhonha and its tributaries may be discerned by sam- pling the shorter coastal rivers in adjacent areas where some forest remains, such as those flowing within and from the Parque Nacional de Monte Pasqual just to the south. However, the shorter coastal rivers now have and probably formerly had a poorer fish fau- na than the Rio Jequitinhonha because they 345 lacked access by stream capture to the richer ichthyofauna of more interior water courses such as the Rio Sao Francisco drainage fur- ther to the west. Furthermore, the larger Rio Jequitinhonha may have had a longer geo- logical history than the shorter coastal rivers, allowing the evolution of more endemic species. Thus the fish faunas of the short adjacent coastal rivers that drain only the Atlantic forest are unlikely to represent fully the original fauna of the Rio Jequitinhonha. Acknowledgments Ricardo M. C. Castro, and Marilyn Weitzman provided field assistance in col- lecting the new species. Financial support and/or transportation for field work was provided by Paulo Vanzolini, Director of MZUSP and Gustavo W. Nunan, MNRJ. Marilyn Weitzman and Lynn Norrod pro- vided technical assistance. Sara Fink pre- pared Figs. 3 and 4. The manuscript ben- efitted from comments and suggestions from Richard Vari, Wayne C. Starnes, and Mar- ilyn Weitzman. We thank all these individ- uals for their interest and contributions. The research and part of the field support was in part financed by the Smithsonian Insti- tution Lowland Tropical Forest Ecosystem Research Program under the direction of W. Ronald Heyer. Literature Cited Britski, H. A., and H. Ortega. 1983. Trichogenes lon- gipinnis, novo género e espécie de Trichomyc- terinae do sudeste do Brasil (Pisces, Siluri- formes).— Revista Brasileira de Zoologia, Sao Paulo 1(3):211-216. Buckup, P. A. 1981. Microlepidogaster taimensis sp. n., novo Hypoptopomatinae do Estacao Eco- logica do Taim, Rio Grande do Sul, Brasil (Os- tariophysi, Loricariidae).—Iheringia, Porto Ale- gre 60:19-31. Dahl, G. 1960. New fresh-water fishes from western Colombia.—Caldasia, Bogota 8(39):45 1-484. Eigenmann, C. H. 1915. The Cheirodontinae, a subfamily of minute characid fishes of South America.—Memoirs of the Carnegie Museum 7(1):1-99. . 1921. The American Characidae, 3.—Mem- 346 oirs of the Museum of Comparative Zoology, Harvard College 43(3):209-310, plates 30-32, 40-55, 61-62, 64, 66, 69, 85, 87, 89, 92. , and G. S. Myers. 1929. The American Characidae, 5.—Memoirs of the Museum of Comparative Zoology, Harvard College 43(5): 429-515, supplement pages 516-558, plates 57, 63, 70-74, 81-83, 94. Fink, W.L., and S.H. Weitzman. 1974. Theso-called cheirodontin fishes of Central America with de- scriptions of two new species (Pisces: Charac- idae).—Smithsonian Contributions to Zoology 172:1-46. Garavello, J.C. 1977. Systematics and geographical distribution of the genus Parotocinclus Eigen- mann and Eigenmann, 1889 (Ostariophysi, Lor- icariidae).—Arquivos de Zoologia, Museu de Zoologia da Universidade de Sao Paulo 28(4): 1-37. Géry, J. 1972a. Contribution a l’etude des poissons characoides de l’Equateur.—Acta Humboldt- iana, Wiesbaden, series Geologica, Palaeonto- logica et Biologica 2:1—110, 8 plates. 1972b. Remarques sur quelques poissons characoides de la Colombie et de l’Equateur avec la description d’une nouvelle espéce de Pseu- dochalceus.—Revue Suisse de Zoologie 79(2): 931-945. . 1977. Characoids of the World. Neptune City, New Jersey, TFH Publications. 672 pp. Myers, G. S. 1926. Eine neue Characiniden Gattung der Unterfamilie Cheirodontinae aus Rio de Ja- neiro, Brasilien.—Blatter fur Aquarien- und Terrarienkunde 37 Jahrgang 24:1-2. Nijssen, H., and I. J. H. Isbriicker. 1976. The South American plated catfish genus Aspidoras R. von Ihering, 1907, with descriptions of nine new species from Brazil (Pisces, Siluriformes, Cal- lichthyidae). — Bijdragen tot de Dierkunde 46(1): 107-131. ——, and 1980. Aspidoras virgulatus, n. sp., a plated catfish from Espirito Santo, Brazil (Pisces, Siluriformes, Callichthyidae).— Bulletin PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON van de Zoologisch Museum, Universitet van Amsterdam 7(13):133-139. Schultz, L. P. 1966. Pseudochalceus kyburzi, a new characid fish from Colombia.—Ichthyologica, the Aquarium Journal 37(1):25-30. Weitzman, M. 1985. Hyphessobrycon elachys, a new miniature characid from eastern Paraguay (Pisces: Characiformes).—Proceedings of the Biological Society of Washington 98(4):799-808. Weitzman, S. H. 1977. Hyphessobrycon socolofi, a new species of characoid fish (Teleostei: Charac- idae) from the Rio Negro of Brazil.—Proceed- ings of the Biological Society of Washington 90(2):326-347. , and C. A. Goncalves da Cruz. 1981. The South American fish genus Rachoviscus, with a description of a new species (Teleostei: Charac- idae).— Proceedings of the Biological Society of Washington 93(4):997-1015. , and S. V. Fink. 1985. Xenurobryconin phy- logeny and putative pheromone pumps in glan- dulocaudine fishes (Teleostei: Characidae).— Smithsonian Contributions to Zoology 421:1- lie , and W. L. Fink. 1983. Relationships of the neon tetras, a group of South American fresh- water fishes (Teleostei, Characidae), with com- ments on the phylogeny of New World chara- ciforms.—Bulletin of the Museum of Comparative Zoology, Harvard University 150(6):339-395. (SHW) Department of Vertebrate Zool- ogy (Fishes), National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C. 20560; (NAM and HAB) Museu de Zoologia, Universidade de Sao Paulo, Caixa Postal 7172, 01051, Sao Paulo, Brazil and (HAB) Bolsista, Conselho Nacional de Desenvolvimento Cientifico e Technologi- co—CNPg. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 347-349 ADDITIONAL RECORDS FOR CALLIANASSA RATHBUNAE SCHMITT, 1935, FROM FLORIDA AND THE BAHAMAS (CRUSTACEA: DECAPODA: CALLIANASSIDAE) Raymond B. Manning and Richard W. Heard Abstract. —Callianassa rathbunae Schmitt is far more abundant in shallow water habitats than suggested by records in the literature. More than 40 spec- imens are recorded from localities in the Bahamas and the Indian River, Flor- ida. The mud shrimp Callianassa rathbunae was described by W. L. Schmitt (1935:15) from two male specimens, 59 mm and 61 mm long, that had washed ashore at Blue- fields, Jamaica [18°10'N, 78°03’W], in 1899. Biffar (1971:699) redescribed the species from two other specimens, one male, 95 mm long, and one female, 82 mm long, dredged in an undisclosed depth off the east coast of Key Biscayne, Miami, Florida, in 1969. Suchanek (1985) reported the results of investigations of the biology of C. rath- bunae and three other species of Callianas- sa at three localities on St. Croix, U.S. Vir- gin Islands: Tague Bay, 17°46'N, 64°36'W; Great Pond Bay, 17°43’N, 64°39'W, in depths to 5 m; and Salt River Canyon, 17°47'N, 64°45'W, in depths to 40 m. So far as we are aware, there are no other rec- ords in the literature for this species. Independent collections by each of us has uncovered much additional material of this species. On 2 October 1980, one of us (RWH) found this species on a silt and cor- alline sand bottom in 10 to 15 feet of water north of the seaplane ramp on the west side of the channel at Bimini, Bahamas [25°44’N, 79°15'W]. A total of 19 specimens was taken with a suction pump: 10 6, carapace lengths (cl) 11.0-17.8 mm (total lengths (tl) 37-68 mm); 5 non-ovigerous 8, cl 11.5 to 15.4 mm (tl 42-57 mm); and 4 ovigerous 2, cl 11.7 to 19.0 mm (tl 42-57 mm) [1 ¢, 1 ovigerous 2, GCRL 180:1107; 2 6, 2 2 (1 ovigerous), USNM]. One specimen of Callianassa branneri (Rathbun), one of the commonest shallow water species in the tropical western Atlantic, and two specimens of an unde- scribed species, closely related to Calli- anassa quadracuta Biffar, were taken at the same time. Twenty-four other specimens have been collected by one of us (RBM) at two local- ities in the southern Indian River, Florida, one just inside the St. Lucie Inlet, Martin County, the other just inside the Fort Pierce Inlet, St. Lucie County. All of these col- lecting sites were on sand flats exposed at low tide; that on the south side of the Fort Pierce Inlet was on a sand bar with scattered seagrass, most abundant on the edges of the bar where the bottom became muddier, whereas the site on the north side of the Fort Pierce Inlet and the St. Lucie site lacked vegetation. The following specimens were taken in the Indian River (unless otherwise indicated, all specimens are in the collection of the Smithsonian Institution, USNM): Florida, Martin County, flat just inside St. Lucie Inlet, 27°10.3'N, 80°10.4'W, clean sand: Sta RBM FP-82-8, R. B. Manning, M.E. Rice, J. Piraino, H. Reichardt, 16 Jul 1982: 1 6, cl 16.3 mm (tl ca. 64 mm).—Sta RBM FP-83-2, R. B. Manning, W. D. Lee, H. Schiff, 11 Feb 1983: 1 4, cl 10.1 mm (tl 33 mm), 1 2, cl 8.9 mm (tl 30 mm). 348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Callianassa rathbunae Schmitt. Male, cl 16.3 mm, Sta RBM FP-82-8: a, Front; b, Inner face of third maxilliped; c, Chela; d, Sixth abdominal somite, telson, and uropod (stippled areas distally on uropods mark regions of dense setation). St. Lucie County, south side of Fort Pierce Inlet, 27°27.7'N, 80°18.7'W, sand flat with sparse seagrass: Sta RBM FP-83-5, R. B. and L. K. Manning, 12 Jul 1983: 1 2, cl 14.5 mm (tl 51 mm).—Sta RBM FP-84-8, R. B. Manning, D. L. Felder, 14 Jul 1984: 1 3, cl 15.8 mm (tl 65 mm), 8 9, cl 16.1—20.5 mm (tl 69-81 mm).—Sta RBM FP-85-1, R. B. Manning, D. L. Felder, 17 Jul 1985: 6 2 (2 ovigerous), non-ovigerous cl 17.2—21.1 mm (tl 62-76 mm), ovigerous specimens dam- aged, cl ca. 17 mm (tl ca. 62 mm) [1 spec- imen to Indian River Coastal Zone Mu- seum, Fort Pierce; 1 specimen to Gulf Coast Research Laboratory (GCRL)].—Sta RBM FP-85-4, R. B. Manning, D. L. Felder, 23 Jul 1985: 1 6, cl 19.1 mm (tl 81 mm), 1 ovigerous 2°, damaged (female taken with 2 specimens of Pinnixa retinens Rathbun).— Sta RBM FP-85-8, R. B. Manning, M. L. Reaka, W. D. Lee, B. Tunberg, 15 Aug 1985: 2 4, cl 18.8-19.0 mm (tl 76-80 mm). St. Lucie County, north side of Fort Pierce Inlet, south side of Coon Island, 27°28.2'N, 80°18.3’W, muddy, hard packed sand near shore: Sta RBM FP-85-2, R. B. Manning, VOLUME 99, NUMBER 2 D. L. Felder, W. D. Lee, 18 Jul 1985: 1 9, cl 19.0 mm (tl 74 mm). The specimens from both the Indian Riv- er (Fig. 1) and Bimini were pink overall in color, with deeper pink to red on the claws. Unlike many callianassids, which appear to be listless when taken from their burrows, the smaller specimens taken at the St. Lucie Inlet site were very active and tried to es- cape capture. All of the specimens reported here were taken with a suction or yabby pump. Those from Bimini were taken with an open ended pump, as described by Manning (1975), which can be used by divers as well as from shore, whereas those from the Indian River were collected with a closed-end Australian commercial yabby pump (Hailstone & Ste- phenson 1961), which can only be used with the lower part of the tube under water. Use of either kind of suction pump greatly sim- plifies the collection of callianassids and other burrowing crustaceans. These samples demonstrate that C. rath- bunae is far more abundant and widespread than suggested by records in the literature, and that it occurs in intertidal habitats as well as in shallow, sublittoral habitats. Cu- riously, whereas Biffar’s collections from shallow water dredging operations off Miami (Biffar 1971:704) yielded about 40 speci- mens of C. branneri and two specimens of C. rathbunae, our collections from Bimini yielded 19 specimens of the latter and only one C. branneri. Acknowledgments We thank Lilly King Manning for pre- paring the figure. This paper is contribution 349 number 153 from the Smithsonian Marine Station at Fort Pierce. The junior author’s research was spon- sored in part by NOAA Office of Sea Grant, U.S. Department of Commerce under Grant #NA81AA-D-00050, the Mississippi-Ala- bama Sea Grant Consortium and the Gulf Coast Research Laboratory. The U.S. Gov- ernment is authorized to produce and dis- tribute reprints for governmental purposes notwithstanding any copyright notation that may appear hereon. Literature Cited 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. Hailstone, T. S., and W. Stephenson. 1961. The bi- ology of Callianassa (Trypaea) australiensis Dana, 1852 (Crustacea, Thalassinidea).— Uni- versity of Queensland Papers 1(12):259-285. Manning, R. B. 1975. Two methods for collecting decapods in shallow water. —Crustaceana 29(3): 317-319. Schmitt, W. L. 1935. Mud shrimps of the Atlantic coast of North America.—Smithsonian Miscel- laneous Collections 93(2):1—21. Suchanek, T. H. 1983. Control of seagrass commu- nities and sediment distribution by Callianassa (Crustacea, Thalassinidea) bioturbation. —Jour- nal of Marine Research 41:281-298. (RBM) Department of Invertebrate Zo- ology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560; (RWH) Gulf Coast Research Laboratory, East Beach Road, Ocean Springs, Mississippi 39564. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 350-358 PSEUDOJANIRIDAE (CRUSTACEA: ISOPODA), A NEW FAMILY FOR PSEUDOJANIRA STENETRIOIDES BARNARD, 1925, A SPECIES INTERMEDIATE BETWEEN THE ASELLOTE SUPERFAMILIES STENETRIOIDEA AND JANIROIDEA George D. F. Wilson Abstract.—The South African marine isopod Pseudojanira stenetrioides is redescribed because characters of its male copulatory structures may be inter- mediate between the asellote superfamilies Stenetrioidea and Janiroidea. Spe- cifically, the endopod of pleopod II has a stylet similar to that of the Janiroidea, but it lacks a sperm tube; pleopod I is similar to the stenetrioid condition but has distal stylet guides as in the Janiroidea. The female copulatory structure, the cuticular organ, is not on the anterodorsal surface as in most janiroideans, and has special auxiliary structures not described from any other asellotan. The species and genus is assigned to a new family, Pseudojaniridae, but its super- family designation is left undecided. Pseudojanira stenetrioides Barnard (1925) is a small isopod from South Africa, re- cently redescribed by Kensley (1977) and classified by that author as a member of the family Janiridae, superfamily Janiroidea. This species showed a stenetrioid habitus and male pleopods I-II that display simi- larities with both the Stenetrioidea and the Janiroidea (Kensley 1977: his figures 9-10). An examination of two specimens of this species from the South African Museum confirmed these observations and, in ad- dition, revealed a new type of female genital organ. The discovery of unusual female and male copulatory organs in Pseudojanira ste- netrioides Barnard (1925) made it necessary to prepare a redescription of this unique species. Because of the possibly transitional characters found in P. stenetrioides, the new family Pseudojaniridae is erected for the ge- nus Pseudojanira. The superfamilial clas- sification, however, is left undecided; prop- er determination of the superfamilies must be based on a complete morphological sur- vey of all the families of the “‘lower’’ Asel- lota. This paper adds to and corrects infor- mation in Kensley (1977) for the purpose of providing the data on P. stenetrioides for such a morphological survey. A survey of asellotan female copulatory organs and oth- er characters (manuscripts in preparation) will extend the results reported here. Materials and Methods M. G. van der Merwe, Marine Biology Technical Officer of the South African Mu- seum (SAM) kindly loaned the holotype fe- male and male specimen of Pseudojanira stenetrioides (catalogue numbers SAM A6295 and SAM A15345, respectively). The illustrations in this paper were inked from pencil drawings made using a Wild M20 microscope fitted with a camera lucida drawing tube. To study the reproductive or- gans of the female specimen, it was stained with methylene blue dissolved in lactic acid. Previous discussion of the evolution of the Asellota has typically relied on simple out- line drawings of limbs for comparison. The fine details of asellotan construction, how- ever, are often phylogenetically important VOLUME 99, NUMBER 2 (Wagele 1983). For example, an outline of the endopod of male pleopod II would not show the difference between the stylet of Pseudojanira and that of the Janiroidea. Therefore, this paper will provide more pic- torial information than has been typically offered in the past. In the illustrations of body parts, anterior is toward the top of the page, mouthparts and pleopods are shown in ventral view unless noted otherwise. Order Isopoda Latreille, 1817 Suborder Asellota Latreille, 1803 Superfamily [ncertae Sedis Pseudojaniridae, new family Figs. 1-3 Type genus.—Pseudojanira Barnard, 1925, by original designation. Previous assignments of type. —Jaeridae: Barnard, 1925:406. Janiridae: Wolff, 1962: 252; Kensley, 1977:251. Ianiridae: Kensley, NOTED Se Diagnosis. —Asellota with broad pe- reonal tergites extending laterally and ven- trally, hiding coxae from dorsal view. Ceph- alon with dorsal eyes, broad lateral lappets, and large frontal rostrum. Pleotelson with only 1 free pleonite visible dorsally, 2 ven- trally. Pereopod I robust, with enlarged se- tose propodus; prehension between dacty- lus and propodus; carpus _ short, quadrangular, setose, not participating in grasping. Male first pleopods with basal seg- ments fused, distal rami separate; distolat- eral corners with dorsal grooves; distal mar- gins truncate, with simple setae. Male pleopod II basal segment enlarged, with en- dopod and exopod projecting medially; pos- terior end of basal segment enlarged, thick- ened, with transverse distomedial groove supplied with fine setae; endopod distal seg- ment stylet-shaped, with open ventral groove and distolateral barbs; endopod proximal segment with thickened cuticular ridge; exopod comprising only single short, robust segment, with thickened dorsal hook on setose anterodistal corner. Male pleo- 351 pods I and II together not opercular. Female second pleopods (not seen by me) fused into single opercular segment lacking setae on margins. Pleopod III exopod broad, round- ed, with fringe of simple setae; endopod with 3 large plumose setae; in male, exopod oper- cular. Uropods short, biramous, setose, barely extending beyond posterior margin of pleotelson. Pseudojanira stenetrioides Barnard Figs. 1-3 Pseudojanira stenetrioides Barnard, 1925: 406-407.—Kensley, 1977:251-253. Holotype.— Adult female, 2 poorly pre- served fragments (cephalon and pereon), pleotelson missing, original reported length 3 mm, width 1.3 mm, SAM 6295. Type locality: “Zululand coast, in a coral (H. W. Bell-Marley, 1920)...” (verbatim from original description, Barnard 1925). Additional material. —Partially dissected adult male, with removed limbs on a slide, length (including rostrum) 2.8 mm, width at sixth pereonite 1.4 mm, SAM A15345. Locality: “°. . . 24°53’S, 34°56’E, 55 metres, from fine gray sand” (verbatim from Kens- ley 1977). Description (in addition to Kensley 1977).—Body characters (Fig. 1A, B): Lat- eral margins of pereonites oval. Body sur- faces covered with fine setae. Body dorso- ventrally thin but highly vaulted: tergites extending beyond main part of body and angling sharply downward. Pereonite 1 sex- ually dimorphic, longer and more robust in males than in females. Female cuticular organ (Fig. 1B, 3): De- scribed below in the section on the female copulatory organ. Cephalon (Fig. 1A, D): Rostrum: ante- riorly rounded; thin, broad, nearly as long as short antennulae; projecting anteriorly from frons, below anterior margin of ce- phalic dorsum. Lateral margins broad, flat- tened, with small anterior spine. Eyes pro- jecting dorsolaterally from domed central 352 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Pseudojanira stenetrioides. A, C-H, Male, 2.8 mm; B, Holotype female, reported intact length 3 mm; A, Dorsal view, setae on right side omitted; B, Dorsal view of female pereonal fragment, co— position of cuticular organ seen through dorsal cuticle, sp—spermatheca seen through dorsal cuticle; specimen was stained in lactic acid and methylene blue to make this possible; C, Pereopod I of male, distal segments only, with enlargement of opposing setation on propodus and dactylus (carpus and propodus have many long tubular setae; setal insertions are indicated by 'u’ or circular marks, and a few are drawn in to show an approximate length of the ones omitted. Some of the setae in the enlargement are illustrated in the same manner.); D, Ventral view of left side of cephalon VOLUME 99, NUMBER 2 portion of cephalon, positioned roughly halfway between midline and lateral mar- gins. Pleotelson (Fig. 1A, E): Broader than long. Pleopodal cavity small, width half width of pleotelson, cavity separate from anus. Lat- eral margins entire, smoothly curving. Antennula (Fig. 1H): Very short, length approximately length of antennal segments 1-4, basal segment largest. Broom setae on segments 2 and 3; aesthetascs on distal 3 segments. Antenna (Fig. 1D): Basal segment 3 with large, basally articulated scale (or squama) extending beyond segment 4. Right mandible (Fig. 1G): Spine row with 10 members. Articular condyle on dorsal surface distinctly shorter than length of ro- bust molar process. Molar process with ap- proximately 9 setae on posterior part of den- ticulate circumgnathal surface. First pereopod (Fig. 1C): Claw of dactylus opposing large spine-like serrate seta on propodus. Row of small tapering setulate setae leaning toward more posterior large spine-like seta. Opposing margin of dacty- lus armed with row of short multiply- toothed setae. Carpus and propodus with several dense groups of long, thin setae. Dactylar claws of walking legs (Fig. 1F): Distal tips of walking legs with 2 robust claws of similar size, and more proximal small claw-like accessory seta. Male Pleopod I (Fig. 2A, B): Length 0.42 pleotelson length, distal segments covering rami of pleopod II. Basal segments quad- rate, fused medially. Distal rami separate, distally truncate with fringe of simple setae posteriorly and laterally. Dorsal side of dis- —_— 353 tolateral corners with stylet grooves (sg in Fig. 2B). Male pleopod II (Fig. 2C, D): Length of basal segment subequal to pleopod I, with endopod and exopod inserting in center of medial margin. Basal segment distally broad, curving laterally to subacute angle, with se- tose groove in medial part of margin. Lat- eral margin of basal segment with row of simple setae. Endopod proximal segment robust, with pronounced ridge on ventro- medial edge; distal segment styliform, with distally tapering groove on ventral surface and 4 small denticles on distolateral margin. Exopod robust, powerfully muscled, with rounded hook and fine setae on anterodistal edge. Pleopod III (Fig. 2E): Exopod broad, fringed with simple setae, covering pleo- pods IV and V; endopod somewhat less broad, dorsal to exopod. Pleopod IV (Fig. 2F): Endopod broader than exopod. Exopod with 2 free, laterally rounded segments, and 7 plumose setae on distal tip. Pleopod V (Fig. 2G): Endopod longer and broader than endopod of pleopod IV. Basal segment and endopod fused, exopod absent. Discussion Characters important to the classification of the Asellota. —Because the current clas- sification of the superfamilies of the Asel- lota is based on the pleopods (Wolff 1962), the unique combination of the male pleo- pod characters (Fig. 2A—D) in Pseudojanira stenetrioides make it difficult to place in the accepted superfamilies. I will not discuss all the superfamilies here because the Pseu- (right side had been dissected); I—antennula, II—antenna, r—rostrum, m—maxilliped, mnd—mandible; note how the rostrum is nearly as long as the antennula, the tip of which is protruding past the basal articles of the antenna; E, Ventral view of male pereonite 7 and pleotelson, with pleopod I shown at the same scale; I—pleopod I, II—pleopod II, I1I—pleopod III, p—penile papillae, pl.1—presumed pleonite 1, pl.2—presumed pleonite 2, a—anus, u—uropod; F, Dactylus of pereopod mounted on slide, possibly pereopod VII as in Kensley (1977); note presence of 2 subequal claws and a more proximal accessory seta on dactylus; G, Right mandible, dorsal view, palp omitted; H, Right antennula, ventral view. 354 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON al NLR | My) \ Fig. 2. Pseudojanira stenetrioides. Dissected parts on slide from male paralectotype. A, Pleopod I, ventral view; B, Pleopod I, dorsal (interior) view of distolateral corner; sg—stylet groove; C, Pleopod II, ventral view, exopodal musculature shown through cuticle; D, Pleopod II rami, d—denticles, en—endopod, ex—exopod, h— position of dorsally directed hook on exopod, r—ridge on proximal segment of endopod, s—stylet (distal segment of endopod), spg—sperm groove (note the ridge on proximal segment of the endopod; if homologous to the janiroidean condition, this ridge may allow the well-muscled exopod to hook onto the endopod during copu- lation); E-G, Pleopods III—V respectively, plumose seta on pleopod IV enlarged. VOLUME 99, NUMBER 2 dojaniridae, the Stenetrioidea, and the Jan- iroidea will be shown in another paper to form a monophyletic group in exclusion to all the other asellote taxa. The primary syn- apomorphies of this group are a short, sin- gle-articled exopod of male pleopod II and a cephalon with a frontal projection. The following discussion is nonphylogenetic and is only meant to highlight the intermediate nature of Pseudojanira. The unique female copulatory organs, also of taxonomic sig- nificance, are described below. The first male pleopod of Pseudojanira has a mixture of janiroidean and stenetrioid characters. As in Stenetrium, the basal seg- ment is large, quadrate, and medially fused. The two sides of the distal segment are free from each other. In contrast, all janiro- ideans have highly reduced basal segments of the male pleopods I and their distal rami are medially fused forming a central sperm tube. However, the distal tips of the first pleopod in Pseudojanira are setose, and their distolateral corners have deep, laterally- curving grooves on the dorsal surfaces, ho- mologous to the same structures in the Jan- iroidea that function as guides for the stylet of the second pleopod. This determination of homology is made on the basis of the position and presumed functional relation- ships of the stylets in both taxa. The male second pleopod is interesting not only in its similarity to the janiroidean condition, but also for specializations that are seen only in this species. Derived char- acters shared with the Janiroidea are the pointed endopodal stylet, the ridge on the proximal segment of the endopod, and the club-like and hooked form of the exopod with its enlarged musculature and distal group of fine setae. The stenetrioid endopod is club-shaped and lacks the proximal cou- pling groove, and the exopod, although short, is not stout and hooked. Unlike any janiroidean, however, the stylet of Pseu- dojanira has only a ventral groove and ter- minates with tiny barbs. The distal end of the basal segment is also unusual and does 355 not occur in either the Stenetrioidea or the Janiroidea: it narrows just posterior to the exopod, and then becomes broad more dis- tally. The distal end is curved, grooved, and covered with tiny, fine setae. The distal part of the stylet rests in the groove of the basal segment’s tip. It may function as an auxil- lary stylet guide, or perhaps as the top part of an enclosed sperm channel. The diagnosis (above) of the Pseudojan- iridae states that one free pleonite is visible dorsally (Fig. 1A), and two ventrally (Fig. 1E). This observation is made with some misgivings because the only whole speci- men had been damaged by Kensley’s dis- section in the region of the ventral pleonites. If more specimens come to light, the pleo- nites should be re-examined. If this obser- vation is correct, Pseudojanira possesses another intermediate character between the Janiroidea (1 free pleonite) and the Stene- trioidea (2 free pleonites, 1 reduced). I agree, however, with Fresi et al. (1980) that the number and fusion of free pleonites is less important than other characters in the clas- sification, simply because the reduction of characters may recur independently, with- out apparent differences in unrelated taxa. The chaetotaxy and form of the first per- eopod require special mention: in many re- spects, both features are similar to those of Stenetrium. The typical janiroidean first pereopod has fewer setae, a larger carpus, and a smaller propodus. Wagele (1983) makes a strong case for the similarity of the chaetotaxy of the Stenasellidae, Atlantasel- lidae, and Microcerberidae of the superfam- ily Aselloidea (see his figure 1, p. 253), as- serting that the microcerberids belong in the Aselloidea based on these similarities. Al- though I don’t doubt his overall conclusions on the placement of the Microcerberidae, these setal similarities may be primitive characters at the level of the Asellota, be- cause many of the same types of setae are also seen in Pseudojanira, Stenetrium, and Gnathostenetroides, all ‘out groups” to the Aselloidea. 356 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Female reproductive system of Pseudojanira, preparatory female. A, Semi—diagrammatic dorsal view of the reproductive organs, showing what they would look like if the dorsal surface of the pereon were removed (anterior is to the right, cephalon and pleotelson broken off); B, Ventral view of pereonite 5, left side, showing oopore region and spermatheca through the ventral surface; C, Enlargement of oopore region showing structures beneath the cuticle, ov—ovary, o0o—oopore, co—cuticular organ, sp—spermatheca, sr—stylet receptacle, b— basis of pereopod V truncated (shown only partially). The accessory seta on the dactyli of per- eopods II—VII is close in position to the third accessory claw found in the janiro- idean family Janiridae and also in the Pro- tojaniridae, and is nearly identical in posi- tion to an accessory seta on the dactyl of the Stenasellidae (see Magniez 1974:33). This seta is presumed to be homologous to the VOLUME 99, NUMBER 2 third “‘claw”’ of the Janiridae, and could well be common to all of the Asellota. The female copulatory organ. — Because only the preparatory female holotype of Pseudojanira stenetrioides was available, no specimens were macerated in potassium hy- droxide (a useful technique for studying cu- ticular structures). The female did stain well in lactic acid and methylene blue, allowing inspection of the cuticular organ (see Veuille 1978 for a description of the homologous structure in Jaera) close to the ventral sur- face. The cuticular organ opens on the an- terior edge of the attachment of the oviduct to the ventral cuticle, and is adjacent to a cuticular fold that continues posteriorly into a blind tube just below the ventral surface. This closed tube opens anteriorly to a broad groove in the anteroventral edge of the fifth pereonite that curves dorsally. The opening of the cuticular organ is surrounded by a bulbous, thickened funnel that appears to open almost directly into a large spermathe- cal sac. The cuticular organ is also posi- tioned anterior to the oopore and is almost separate from it. The spermatheca pro- trudes posteriorly into the sixth pereonite and was observed to contain translucent, heavily staining material similar to sperm masses seen in other species of Asellota. A pocket-like structure protrudes internally and dorsally from the external position of the oopore. The systematic position of Pseudojan- ira.—Much of the unusual nature of P. ste- netrioides derives from its possession of unique characters or characters seemingly intermediate between two superfamilies of the Asellota. This taxon is assigned to the Pseudojaniridae n. fam. because it cannot be effectively placed in any of the existing asellote families without diluting the poten- tial recipient family’s or superfamily’s con- cept. Moreover, Pseudojanira has repro- ductive specializations that are unique to the Asellota, thereby warranting the new family for the genus. Its higher classifica- tion, however, can be stated no more ac- curately than “Isopoda Asellota superfam- 357 ily incertae sedis.’ Conceivably a new superfamily could be created as well, but this would result in continued ‘superfamily inflation’ with the perceived differences be- tween the superfamilies becoming smaller and more difficult to reconcile. Note, for example, Schultz’s (1978) ill-fated (Sket 1979; Wilson 1980) attempt to create a new asellotan superfamily based on the presence of a free coxa on the first pereopod. The male pleopods I and II define the superfamily Janiroidea extremely well: all the diverse members of that taxon have male copulatory structures identical in their gen- eral form. The homologous structures in all the remaining families of the Asellota, on the other hand, show radical variations, with differences in the endopod of male pleopod II often characterizing species and genera, rather than superfamilies (e.g. species of the asellid genus Caecidotea, Lewis and Bow- man 1981; genera of the Stenasellidae, Mag- niez 1974, his figure 11). This variation her- alds a need for a re-evaluation of the current classification. Describing the second pleo- pod’s endopod of the janiroidean male as ““stylet-like” provides insufficient data be- cause the detailed structure of the stylet is different between the Janiroidea, the Pseu- dojaniridae, and the Protojaniridae (Fresi et al. 1980). More morphological data must be collected on all the major groups of the Asellota to allow effective comparisons. A study of the female copulatory organs may be of considerable value, but other features require attention. Pseudojanira stene- trioides will provide a useful datum in this evolutionary discourse. Acknowledgments This work was removed from a larger manuscript on the evolution of copulatory structures in the Asellota. The earlier manu- script was reviewed at various stages by Richard C. Brusca, Los Angeles County Museum of Natural History, and by Robert R. Hessler, William A. Newman, Richard H. Rosenblatt, William R. Riedel, and Dan L. Lindsley, all of the University of Cali- 358 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON fornia, San Diego. The final version was re- viewed by Thomas E. Bowman, U.S. Na- tional Museum of Natural History, and Torben Wolff, Zoologisk Museum, Copen- hagen. I thank these gentlemen for their ad- vice and editorial help. This work was par- tially supported by a grant from the National Science Foundation, Systematic Biology and by acontract from the National Oceanic and Atmospheric Administration, administered by G. L. Taghon, Oregon State University. Literature Cited Barnard, K. H. 1925. Contributions to the Crusta- cean fauna of South Africa. 9. Further additions to the list of Isopoda.— Annals of the South Af- rican Museum 20:381-412. Fresi, E., E. Idato, and M. B. Scipione. 1980. The Gnathostenetroidea and the evolution of prim- itive asellote isopods.—Monitore Zoologico Italiano (New Series) 14:119-136. Kensley, B. 1977. New records of marine Crustacea Isopoda from South Africa. — Annals of the South African Museum 72(13):239-265. Lewis, J. J., and T. E. Bowman. 1981. The subter- ranean asellids (Caecidotea) of Illinois (Crus- tacea: Isopoda: Asellidae).—Smithsonian Con- tributions to Zoology 335:1-66. Magniez, G. 1974. Données faunistiques et écolo- giques sur les Stenasellidae.—International Journal of Speleology 6:1—180. Schultz, G. A. 1978. Protallocoxoidea new superfam- ily (Isopoda Asellota) with a description of Pro- tallocoxa weddellensis new genus, new species from the Antarctic Ocean. —Crustaceana 34(3): 245-250. Sket, B. 1979. Atlantasellus cavernicolus, n. gen., n. sp. (Isopoda, Asellota, Atlantasellidae n. fam.) from Bermuda.—Bioloski Vestnik (Ljubljana) 27(2):175-183. Veuille, M. 1978. Biologie de la reproduction chez Jaera (Isopode Asellote) II. Evolution des or- ganes reproducteurs femelles.— Cahiers de Biol- ogie Marine 19:385-395. Wagele, J. 1983. On the origin of the Microcerberidae (Crustacea: Isopoda).— Zeitschrift fuer Zoolo- gische Systematik und Evolutionsforschung 21(4):249-262. Wilson, G. 1980. Superfamilies of the Asellota (Is- opoda) and the systematic position of Stene- trium weddellense (Schultz). — Crustaceana 38(2): 219-221. Wolff, T. 1962. The systematics and biology of bathy- al and abyssal Isopoda Asellota.—Galathea Re- port 6:1—320. A-002, Scripps Institution of Oceanog- raphy, La Jolla, California 92093. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 359-362 TRANSFER OF QUADRASIA FROM THE PLANAXIDAE TO THE BUCCINIDAE (MOLLUSCA: GASTROPODA: PROSOBRANCHIA) Richard S. Houbrick Abstract. — Quadrasia hidalgoi Crosse, previously assigned to the Planaxidae, is herein transferred to the genus Clea H. and A. Adams, family Buccinidae, on the basis of anatomical characters. The animal has a long muscular inhalant siphon, closed pallial gonoducts, a pleurembolic proboscis, and a rachiglossate radula. The poorly known genus Quadrasia Crosse, 1886, has been cited by Crosse (1886), Cossmann (1906:197), Thiele (1929: 204), and Wenz (1940:722) as the only freshwater representative of the family Planaxidae Gray, 1850. Very few references to this monotypic genus occur in the liter- ature, and despite a thorough search of ma- jor museum collections throughout the world only three specimens were found, one of which was the holotype. A single, alco- hol-preserved specimen, from which soft parts were extracted, was studied. Anatom- ical examination showed that Quadrasia hi- dalgoi Crosse, 1886, is not a planaxid as originally suggested, but should be allocated to the genus Clea H. and A. Adams of the family Buccinidae. A redescription of this genus and species follows with a discussion of the evidence placing them in the Buccin- idae. Genus Clea H. and A. Adams, 1855 Clea H. and A. Adams, 1855:119. Type species: Clea nigricans A. Adams, 1855, by monotypy and original designation; 1858:623; Thiele, 1931:316—-317. Quadrasia Crosse, 1886:159-161. Type species: Quadrasia hidalgoi Crosse, 1886, by monotypy; Cossmann, 1906:197; Thiele, 1929:204; Wenz, 1940:722, fig. 2090. Diagnosis. —Shell solid, ovate, moderate- ly elongate with 5-6 convex, inflated, trans- versely striated whorls of overall smooth aspect. Protoconch large, smooth, usually missing. Aperture large, about one-half the shell length, fusiform. Outer lip smooth, convex. Columella concave, abruptly trun- cate with basal fold. Anterior canal short. Shell cream-colored but with horny, dark brown periostracum. Operculum corneous, ovate, with subterminal nucleus. Radula ra- chiglossate, buccinid. Remarks. —This little-known taxon has been found in the Philippines, Borneo, and in parts of Indonesia. It is noteworthy in being a freshwater representative of the largely marine Buccinidae. Clea hidalgoi (Crosse), new combination Figs. 1, 2 Quadrasia hidalgoi Crosse, 1886:161-163, pl. 8, fig. 7; (Holotype: MNHNP).—Coss- mann, 1906:197.—Thiele, 1929:204.— Wenz, 1940:722, fig. 2090. Description. —Shell moderate in size (ho- lotype 13.4 mm in length), solid, moder- ately elongate with overall smooth aspect, comprising about 6 inflated whorls. Shell sculptured with minute axial riblets and spi- ral threads producing wrinkled look under 360 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. A-—F. Apertural, side, and dorsal views of holotype of Quadrasia hidalgoi Crosse, 1886 (MNHNP, no number, 13.5 mm length). Bottom row shows holotype whitened with ammonium oxide to enhance sculptural details. VOLUME 99, NUMBER 2 361 Fig. 2. Radula of Clea hidalgoi from Palawan, Philippines (bar = 110 um). magnification. Protoconch large, 1.5 whorls, smooth. Large body whorl with elongate, fusiform aperture spanning about one-half of shell length. Anal canal elongate. Ante- rior siphonal canal short, but well-devel- oped. Outer lip smooth, slightly sinuous. Columella concave with prominent basal tooth and slight columellar callus. Suture moderately impressed, slightly overhung by each consecutive whorl. Shell flesh-colored, early whorls dark purple-brown. Periostra- cum well-developed, wrinkled, light brown, giving shell overall tan color. Operculum corneous, elongate-ovate, paucispiral with subterminal nucleus. Radula (Fig. 2) ra- chiglossate, long and narrow. Rachidian tooth strongly arched with cutting edge of 7-8 tiny, sharp denticles and expanded lat- eral extensions. Lateral teeth large, angular in outline, and tricuspid with 2 sharp, curved inner basal cusps and large sickle-shaped distal cusp. On basal radular membrane, laterals slightly separated from rachidian teeth. Animal with long muscular inhalant siphon, closed pallial gonoducts, and pleu- rembolic proboscis. Remarks.—Quadrasia hidalgoi was de- scribed from freshwater streams on the is- land of Balabac, Philippine Islands. The original description (Crosse 1886:160) stat- ed that according to the observations of Quadras, it lives on sandy bottoms in quiet streams. Both the shell and radula of Quadrasia are similar to those of some buccinids which have been described from freshwater hab- itats in this region. Among these freshwater buccinid genera the genus Clea H. and A. Adams best matches the description of Quadrasia. Crosse (1886:161—162) referred Quadrasia hidalgoi to the Planaxidae, but this allocation was based on shell characters only. He pointed out that other marine prosobranch families also had freshwater representatives, specifically referring to Canidia H. Adams, 1862 (not Thompson, 362 1857), which is a synonym of Anentome Cossmann, 1901, a ribbed, buccinid genus closely related to Clea. Crosse (1886) was apparently unaware of the description of Clea by H. and A. Adams in 1855. Had he been familiar with the type species of this genus, Clea nigricans, he would have noted the close resemblance of it to Quadrasia hi- dalgoi. Thiele (1931:317) considered Can- idia to be a subgenus of Clea. H. and A. Adams subsequently (1858:623) referred Clea to the subfamily Melanopsidae (su- perfamily Cerithiacea). The radula of Quadrasia is very similar to that of Clea nigricans as depicted by Thiele (1931:317) and also to Macron (Thiele 1931:316). The tricuspid lateral teeth of Quadrasia clearly distinguish it from the similar radulae of nassariids, which have bicuspid laterals (Cernohorsky 1984:12-—13). As may be seen in the above discussion, Quadrasia hidalgoi was previously allocat- ed to the Planaxidae and Melanopsidae, both in the superfamily Cerithiacea. Examina- tion of the anatomy of Quadrasia revealed a long, tubular inhalant siphon, closed pal- lial gonoducts, a pleurembolic proboscis, and a rachiglossate radula. These anatom- ical and radular features exclude Quadrasia from the Cerithiacea and point to the Buc- cinidae as the proper taxonomic assign- ment. Within this family, the genus Clea appears best to fit the shell and radular char- acters seen in Quadrasia, which is here con- sidered to be conspecific with Clea. Quad- rasia hidalgoi is herein transferred from the Planaxidae and assigned to the genus Clea in the family Buccinidae. Acknowledgments I thank Dr. Philippe Bouchet of the Mu- seum National d’Histoire Naturelle, Paris, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON for loans of the type specimen and the pre- served material. Mr. Victor Krantz of the Smithsonian’s Photographic Services kind- ly supplied the photographs. Drs. David Reid and M. G. Harasewych critically re- viewed the manuscript. Literature Cited Adams, H. 1862. Descriptions of some new genera and species of shells from the collection of Hugh Cuming, Esq.—Proceedings of the Zoological Society of London 1862:383-385. , and A. Adams in A. Adams. 1855. Descrip- tions of two new genera and several new species of mollusca, from the collections of Hugh Cum- ing, Esq. — Proceedings of the Zoological Society of London 23:119-120. 1853-1858. The genera of recent mollusca, vol. 1, xl + 484 pp.; vol. 2, 66 pp; vol. 3, 138 pls. London. Cernohorsky, W. O. 1984. Systematics of the family Nassariidae (Mollusca, Gastropoda).— Bulletin of the Auckland Institute and Museum No. 14: 356 pp., 51 pls. Cossmann, M. 1901; 1906. Essais de Paléoconcho- logie Comparée, Vol. 5:293 pp., 10 pls.; Vol. 7: 261 pp., 14 pls. Paris. Crosse, H. 1886. Description du nouveau genre Quadrasia.—Journal de Conchyliologie 34:159- 163, pl. 8. Gray, M. E. 1850. Explanation of plates and a list of genera. Figures of molluscous animals vol. 4, pp. i-iv, 1-219. London, Longmans, Brown, Green & Longmans. Thiele, J. 1929-1930. Handbuch der Systematischen Weichtierkunde Vol. 1, part 1. 376 pp. Jena. Wenz, W. 1940. Gastropoda. Teil 1, Allgemeiner Teil/ Prosobranchia, in Schindewolf, O. H., Hand- buch der Paldozoologie, 6:vi + 1639 pp. Berlin. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution NHB-163, Washington, DC] 20560: PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 363-365 PRELIMINARY DESCRIPTIONS OF FOUR NEW SPECIES OF DORIPPID CRABS FROM THE INDO-WEST PACIFIC REGION (CRUSTACEA: DECAPODA: BRACHYURA) Raymond B. Manning and L. B. Holthuis Abstract.—The following species are diagnosed: Dorippe irrorata, from the Andaman sea; Dorippoides nudipes, from the western Indian Ocean; Nobilum arachnoides, from the Inland Sea of Japan; and Paradorippe cathayana, from China. Since our review of the west African brachyuran crabs was published (Manning and Holthuis 1981), we have been working on a revision of the crabs of the subfamily Dorippinae, family Dorippidae, from the Indo-West Pacific region. The two Atlantic representatives of this subfamily were re- vised by us in 1981. Publication of our re- vision of the Dorippinae has been delayed much longer than anticipated, and several colleagues have inquired about the status of our new species, some of which have been labelled as new in museum collections (Brit- ish Museum (Natural History), London; Muséum National d’Histoire Naturelle, Paris; Zoological Museum, Copenhagen; and our institutions as well) for several years. We take this opportunity to make available the names of these new taxa. The diagnoses given below will differen- tiate the new species from others in their respective genera, defined in Manning and Holthuis (1981:30, 31). The following ab- breviations are used: BM(NH), British Mu- seum, Natural History, London; RMNH, Rijksmuseum van Natuurlijke Historie, Leiden; USNM, National Museum of Nat- ural History, Smithsonian Institution, Washington; cb and cl, carapace breadth and carapace length, respectively, in mm. Dorippe irrorata, new species Fig. la, b Holotype. — Andaman Sea, south of Mer- gui Archipelago; 09°54'N, 97°42’E; 73 m; International Indian Ocean Expedition, An- ton Bruun (Cruise I, Sta AB-21; 24 Mar 1963: 1 male, cl 21.5, cb 22.0 (USNM 172495). Diagnosis. —Carapace broader than long, with distinct tubercles dorsally and with dis- tinct lateral branchial tooth. Anterolateral margin of carapace, between bases of ex- orbital teeth and cervical groove, smooth. Inner dorsal margin of exorbital tooth smooth. Lower orbital margin with teeth and denticles mesially. Carpus of cheliped with granules, palm of chela with granules over most of surface. Teeth on second and third somites of male abdomen very low, distinctly granular. Remarks. —This species differs from the three species of Dorippe now recognized as follows: It can be distinguished from Do- rippe frascone (Herbst, 1785) in having the carpus of the chelipeds granular and in hav- ing granules on the teeth of the male ab- domen. It differs from D. sinica Chen, 1980, in having tubercles or denticles on the an- terolateral margin of the carapace, behind 364 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. a, b, Dorippe irrorata, male holotype, Andaman Sea: a, Cheliped; b, Abdomen; c, Dorippoides nudipes, female paratype, Harkiko Bay, Ethiopia, Red Sea, propodus and dactylus of third pereopod; d, Nobilum arach- noides, male holotype, Inland Sea of Japan, third pereopod; e, Paradorippe cathayana, male holotype, Jimei, Fujian Province, China, cheliped. the anterolateral spine, and differs from D. tenuipes Chen, 1980 [=D. miersi Seréne, 1981] in having shorter walking legs, with the merus of the second pereopods only five rather than six times as long as high in males. Etymology.—The specific name is from the Latin, “irroratus,’’ covered with gran- ules. Dorippoides nudipes, new species Fig. lc Holotype. —Massawa, Ethiopia, Red Sea; Israel South Red Sea Expedition no. E62/ 4115; trawled: 1 male, cl 17 mm, cb 19 mm (RMNH no. D.35530). Diagnosis. —Carapace broader than long, lacking both dorsal tubercles and distinct lateral branchial tooth. Surface distinctly granular laterally on branchial region. Ex- orbital teeth overreaching frontal teeth. Dactyli of second and third pereopods broad throughout their lengths, broadest in distal fourth. Remarks. — Differs from the only other species in the genus, Dorippoides facchino (Herbst, 1785), in having the carapace more granular and in having the dactylus of the second and third pereopods broadest in dis- tal fourth rather than at midlength. Etymology. —The name is from the Lat- in, ““nudus,”’ naked, and “‘pes,”’ foot. Nobilum arachnoides, new species Fig. ld Holotype. —Japan, Inland Sea, near Kobe; 34°38'N, 135°01’E; dredged in 8—50 fms (15- VOLUME 99, NUMBER 2 92 m); sand; Challenger Sta 233A; 19 May 1875: 1 male, cl 15.8, cb 16.7 (BM(NH) 84.44). Diagnosis. —Carapace wider than long, surface lacking erect tubercles, grooves well defined. Exorbital teeth falling short of front. Posterior margin of orbit lacking erect spine on outer side of orbital fissure. Gastric re- gion lacking erect prominences. Merus of second and third pereopods about 7 times longer than high. Remarks. —This species agrees with No- bilum japonicum (von Siebold, 1824) and differs from N. histrio (Nobili, 1903) in lacking a tooth on the exorbital margin and five prominences on the gastric region. It differs from both of these species in length and slenderness of the second and third per- eopods. Etymology. —The name is from the Greek, “‘arachnes,”’ spider, and “‘-oides,” like. Paradorippe cathayana, new species Fig. le Holotype.—China, Jimei, Fujian Prov- ince; S. F. Light, coll.; 24 Jun 1923: 1 male, cl 16.8, cb 18.2 (USNM 57762). Diagnosis.—Carapace wider than long, surface lacking erect tubercles or promi- nences, appearing smooth, grooves well de- fined. Orbital fissure narrow, closed. Carpus 365 of chelipid smooth. Propodus of third leg less than 3 times longer than high. Remarks. —This species differs from Paradorippe australiensis (Miers, 1884) in smoothness of carapace and in having the carpus of the cheliped smooth, not granular, and it can be distinguished from P. polita (Alcock and Anderson, 1894) in having the orbital fissure closed and the propodus of the third pereopod less than three times longer than high. Etymology. —The name is derived from Cathay, the name used for China in the Middle Ages. Acknowledgments We thank Lilly King Manning for pre- paring the figure. Literature Cited Manning, R. B., and L. B. Holthuis. 1981. West Af- rican Brachyuran crabs.—Smithsonian Contri- butions to Zoology 306:i—x1i, 1-379. (RBM) Department of Invertebrate Zo- ology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560; (LBH) Rijksmuseum van Na- tuurlijke Historie, Post Box 9517, 2300 RA Leiden, The Netherlands. PROC. BIOL. SOC. WASH. 99(2), 1986, pp. 366-373 NOTES ON GERYON FROM BERMUDA, WITH THE DESCRIPTION OF GERYON INGHAMI, NEW SPECIES (CRUSTACEA: DECAPODA: GERYONIDAE) Raymond B. Manning and L. B. Holthuis Abstract.—Two species of the deep water crab genus Geryon, taken in com- mercial traps, G. fenneri Manning and Holthuis and G. inghami, n. sp., are reported from Bermuda. Geryon incertus Miers, 1886, originally described from Bermuda, is shown to be a junior synonym of the portunid Bathynectes longi- spina Stimpson, 1871. Commercial trapping operations in depths in excess of 400 fathoms (732 m) around the Bermuda Islands [32°20’N, 64°45’W] by John P. Ingham in 1984 have revealed the occurrence of two distinct species of Ger- yon. One of these, G. fenneri Manning and Holthuis, 1984, was originally described from localities around Florida. The second species proves to be undescribed and is named below. Luckhurst (1985) has pro- vided a preliminary account of the fishery. There are two previous records of a species of Geryon from Bermuda. The first is that by Miers (1886:224), who described Geryon incertus from an immature specimen taken off Bermuda. As both Chace (1940:39) and Manning and Holthuis (1984:666) pointed out, the status of Miers’ species has re- mained uncertain since its description. We believe that G. incertus is the juvenile of a portunid described by Stimpson in 1871, Bathynectes longispina. Although adults of B. longispina have not yet been recorded from Bermuda (Markham and McDermott 1981), we provide records of adults taken in traps off Bermuda. The second record was given by Mark- ham and McDermott (1981), who included G. quinquedens Smith in a checklist of Ber- muda decapods. We tentatively identify their material with G. fenneri, as we have found neither records in the literature nor material of G. quinquedens from Bermuda. Species of Geryon from Bermuda Geryon fenneri Manning and Holthuis, 1984 Figs. la, 2a ?Geryon quinquedens. —Markham and McDermott, 1981:1274 [not G. quinque- dens Smith, 1879]. Geryon fenneri Manning and Holthuis, 1984:666, figs. 1, 2a, b, 3a—c, 4a, b.— Soto, 1985:482, 483, 486, 487. Material. —Off Bermuda [32°20’N, 64°45'W], 430-450 fms (787-824 m), traps, Oct 1984, John P. Ingham, Eugene Lambe, R. B. Manning, D. L. and J. Felder, and B. Luckhurst, collectors, from vessel Jrilogy: 2 males, 8 females (3 ovigerous), 3 dry car- apaces (USNM 205334, 205335; 2 females to Rijksmuseum van Natuurlijke Historie, Leiden; 2 females to British Museum (Nat- ural History), London).— Off south shore of Bermuda, 500 fms (915 m), 5 Nov 1984, John P. Ingham, collector: 1 male (USNM).—Same, ca. 550 fms (1007 m), 27 Dec 1984: 1 male, 1 female (USNM).—Off Bermuda, 1985: 2 females (1 with sacculin- id) (USNM).—Same, 1985: 8 males, 10 fe- males (2 ovigerous) (USNM). Measurements. —Carapace lengths (cl) of examined males 57 to 160 mm, of non- ovigerous females 68.5 to 116 mm, of ovig- erous females 91 to 118 mm, of unsexed VOLUME 99, NUMBER 2 Fig. 1. measurement of height and width); b, G. affinis, male, cl 134 mm, off Scotland (USNM 210904). dry carapaces 97 to 98 mm. Carapace widths (cb) of examined males 81 to 190 mm, of non-ovigerous females 89 to 146 mm, of ovigerous females 113 to 146 mm, of un- sexed dry carapaces 121 to 124 mm. Luckhurst (1985) reported that 73 spec- imens examined by him in October and No- vember 1984 had carapace widths ranging from 108 to 177 mm, and that all of the largest crabs he examined were males. Remarks.—These specimens agree well with the original account. The carapace length ranges from 0.76-0.84 times the width. The distance between the first and third anterolateral teeth of the carapace is 0.93 to 1.10 times the distance between the third and fifth anterolateral teeth. The orbit is narrower than the front in most speci- mens. On the fifth leg, the merus is about half as long as the carapace and its length is 4.1 to 5.1 times its height; and the pro- podus is 3.5 to 4.3 times longer than high. The third maxilliped of the largest male (cl 160 mm) differs from that of G. affinis A. Milne Edwards and Bouvier (male, cl 134 mm), from off Scotland, as shown in Fig. 1. In G. affinis the merus of the third maxilliped is slightly longer than high, Merus of third maxilliped of: a, G. fenneri, male, cl 160 mm, Bermuda (arrows indicate points of whereas in G. fenneri it is slightly wider than high. The outer angle differs in the two species. As we noted above, we tentatively assign Markham and McDermott’s (1981) record for G. quinquedens to this species, the more abundant of the two species of Geryon now known with certainty from Bermuda. Soto (1985) reported this species from the Straits of Florida, on mud bottoms in depths of 322 to 470 m. He characterized it as one of the species taken only on the continental side of the Straits. Geryon inghami, new species Figs. 2b, c, 3, 4, 5a Material. —Off Bermuda [32°20’N, 64°45’W], 1400 fathoms (2562 m), traps, summer 1984, John P. Ingham, collector: 2 males, 1 female (USNM: largest male is ho- lotype, USNM 205336, other specimens are paratypes, USNM 205333); 2 males (Riks- museum van Natuurlijke Historie, Leiden, paratypes); 1 male (British Museum (Nat- ural History), paratype); 5 males (Bermuda Division of Fisheries, paratypes).— Ber- muda, off south shore, 500-550 fms (915- 368 A Fig. 2. Dorsal view of: a, G. fenneri, ovigerous fe- male, cl 91 mm, Bermuda; b, c, G. inghami, male holotype, cl 64 mm, Bermuda (carapace enlarged in c to show detail). 1007 m), 25 Feb 1985, J. P. Ingham, col- lector: | male (paratype, USNM 228196).— Off Bermuda, 1985: 1 male (USNM 228197, paratype). Diagnosis. —A small Geryon, clto 82 mm in adults. Carapace broader than long, length 0.68 to 0.77 (mean 0.72) times width. Me- dian pair of frontal teeth large, overreaching PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON larger lateral frontal teeth, separated by me- dian V-shaped sinus, latter usually posterior to level of acute, pointed lateral frontal teeth. Distance between submedian frontal teeth variable, usually more than half distance from each to lateral frontal tooth, but much less in largest male. Anterolateral teeth 5, first, third, and fifth strongest, second and fourth smaller but well developed, all sharp. Distance between first and third teeth sub- equal to distance from third to fifth (less in some specimens, more in others). Subor- bital spine large, extending to level of apex of lateral frontal teeth. Carapace with curved ridge extending mesially from lateral tooth. Posterolateral margin of carapace irregular- ly granular. Cheliped rather rough, surface distinctly granular, with sharp subdistal and smaller distal spine dorsally on merus; car- pus with distinct outer spine, anterior mar- gin with small, sharp granules and large in- ner spine; propodus with distinct dorsal spine. Meri of second to fifth pereopods with distinct distal dorsal spine. Fifth leg: merus slender, length 6.1 to 6.9 times height, with erect distal dorsal spine; carpus with distinct line of granules dorsally; propodus slender, length 5.5 to 6.0 times height, with sharp dorsal granules proximally; dactylus 0.74 to 0.87 times as long as propodus, compressed laterally, height at midlength greater than width, longitudinally channelled dorsally. Telson triangular, 1.3 to 1.6 times broader than long. Gonopod as figured. Size.—Carapace lengths of males 40 to 82 mm, of single female examined 44 mm; carapace widths of males 55 to 110 mm, of female 64.5 mm. Color. —In life this species is dark red. Remarks. —Geryon inghami shares many features with G. gordonae Ingle (1985:90) from the northeastern Atlantic, but a com- parison of specimens from the two areas reveals that G. inghami differs as follows: overall the body is much more coarsely granulated, especially on the chelae and the anteroventral surfaces of the carapace; the suborbital teeth are stronger, and there is a VOLUME 99, NUMBER 2 369 Fig. 3. distinct line of tubercles on the suborbital margin; the frontal teeth are larger; the fourth anterolateral tooth is much larger; the sec- ond and fifth anterolateral teeth are strong- er; the longitudinal carina on the outer sur- face of the palm of the cheliped is more distinct, and, in the female, the inner surface of the palm is much more granular. Ingle (1985:95) noted that in G. gordonae the an- terolateral spines of the carapace are smaller and blunter with increasing size; this trend is not at all apparent in the available ma- terial of G. inghami. All of the anterolateral spines of the carapace are present and well developed, even in the largest male exam- ined. In G. gordonae the fourth lateral tooth is very small even in small specimens; it may be essentially obsolete in adults. The holotype of G. inghami, the illus- trated specimen, has the fourth anterolat- eral tooth of the left side damaged; in other specimens it is as large as shown for the right fourth tooth in Fig. 3. This species, apparently less common off Bermuda than G. fenneri, was found in Geryon inghami, male holotype, cl 64 mm, dorsal view. depths between 430 and 1900 fms (787- 3477 m) (Luckhurst 1985). Etymology.—This species is named for Mr. John P. (Sean) Ingham of Bermuda, whose interest in deep-water trapping in Bermuda led to its discovery. The Status of Geryon incertus Miers (Fig. 6) Miers (1886:224, pl. 16, fig. 3) described Geryon incertus from off the Bermudas (32°21'30"N, 64°35'55”W) in 435 fms (796 m). Miers’s account was based on an im- mature specimen with a carapace length and width of 5 mm. Miers noted (1886:224) that “It [G. incertus] may belong to a genus of the Portunidae, near to Bathynectes, where I originally placed it.’’ The status of Miers’s species has remained uncertain since it was described. Chace (1940:39) remarked that “it obviously does not belong in this genus.” Geryon incertus, although characterized by Miers as having four anterolateral teeth, actually has but three primary teeth on each side, as shown in Miers’s original figure (pl. 370 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON NWA 4 oy Sex Zs C Fig. 4. Geryon inghami, male holotype, cl 64 mm: a, Suborbital margin; b, Cheliped; c, Chela; d, Fifth pereopod; e, Dactylus of fifth pereopod, dorsal view; f, Telson; g, Gonopod; h, Tip of gonopod. 16, fig. 3), parts of which are reproduced here in Fig. 6. The posteriormost two teeth have a small denticle between them. This denticle is shown on both sides in Miers’s figure, but was described as occurring on one side and being obsolete on the other. In his original account, Miers remarked (1886:225): “From all the described species of this genus, Geryon incertus is distin- guished by the form of the front, and from the typical species of Bathynectes, Stimp- son, not only by this character, but also by the structure of the basal antennal joint, which is free and not united with the lateral subfrontal process.” The basal joint of the antenna also is free in specimens of B. /ongi- spina Stimpson from off New Jersey and in our material of Geryon (Fig. 5) as well. R. W. Ingle of the British Museum (Nat- ural History) (in litt.) examined the type of Geryon incertus at our request, and pointed out similarities between it and juveniles of B. longipes (Risso, 1816), reported in Ingle and Rice (1984). A comparison of the figure of the carapace of the type of G. incertus in the collections of the British Museum (Nat- ural History) (Fig. 6a) with that ofa juvenile of B. longispina (Fig. 7) from off New Jersey reveals that they share many similarities, even to the pattern of granulation on the carapace. At the size figured by Miers, the lateral spine of the carapace has not yet be- VOLUME 99, NUMBER 2 371 Fig. 5. Ventral view of orbit and antenna in: a, G. inghami, male holotype, cl 64 mm; b, Bathynectes longispina, male, cl 45 mm, off New Jersey (USNM 185422). gun to enlarge in relation to the other spines. We consider Geryon incertus Miers, 1886, to be a subjective junior synonym of Bath- ynectes longispina Stimpson, 1871. Until recently, Bathynectes longispina was identified with the East Atlantic species then known as B. superbus (Costa, 1853). Man- ning and Holthuis (1981:80) pointed out that the East Atlantic B. superbus was a junior synonym of B. maravigna (Prestandrea, 1839) and that material from the West At- lantic until then identified with B. superbus actually should be known as B. longispina Stimpson, 1871. Bathynectes longispina has not been re- corded from Bermuda as an adult. Williams and Wigley (1977:11) noted that it was known from Martha’s Vineyard to the Yu- catan Channel. Two specimens were sent to one of us (RBM) for identification by W. Sterrer, then Director of the Bermuda Bi- ological Station. They were taken a few miles south of Bermuda in May 1981, in about 500 meters. Both specimens are males, with carapace lengths of 32.7 and 41.5 mm and carapace widths of 59.6 and 72.9 mm, re- spectively. A second lot of this species from Bermuda was found in the Smithsonian col- lections (USNM 169237). It comprises two females with carapace lengths of 28.5 and 37.2 mm and carapace widths of 52.0 and 68.2 mm, respectively. These latter speci- mens were collected in 400 fathoms (732 meters) by R. T. Abbott on 26 September 1974. Bathynectes longispina recently has been recorded from localities off North Carolina (Williams, McCloskey, and Gray 1968), in the Chesapeake Bight (Lewis and Haefner 1977), the Middle-Atlantic Bight (600 spec- Fig. 6. 1886: pl. 16, fig. 3, 3a). Holotype of Geryon incertus Miers, cl ca. 5 mm: a, Dorsal view; b, Suborbital margin. (From Miers 372 Fig. 7. mm, off New Jersey (USNM 185623). Carapace of juvenile B. /ongispina, cl 5.0 imens at 33 stations in depths between 90 and 751 meters; Wenner and Boesch 1979), the middle Atlantic shelf (Bowen et al. 1979), and the Straits of Florida in 174 to 403 m (Soto 1985). It apparently is fairly common in moderate depths off the eastern United States (see Rathbun 1930:28, for earlier rec- ords), and its occurrence in Bermuda is not too surprising. Acknowledgments Our studies of members of the crab genus Geryon have been supported by the Food and Agriculture Organization of the United Nations (FAO); that support is gratefully acknowledged. Our studies of Atlantic species of Geryon were initiated with the support of the Smithsonian Marine Station at Fort Pierce; this is contribution number 160 from that station and contribution number 1070 from the Bermuda Biological Station. We thank Roy Kropp for taking the photographs, Lilly King Manning for pre- paring the figures, and Austin B. Williams, Systematics Laboratory, National Marine Fisheries Service, for reviewing the manu- script. R. W. Ingle, British Museum (Nat- ural History), examined the type of G. in- certus and pointed out the similarity between it and juveniles of Bathynectes. This work PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON would not have been possible without the support and cooperation of John P. Ingham and Eugene Lambe of Bermuda, and Brian Luckhurst, Bermuda Division of Fisheries. Literature Cited Bowen, M. A., P. O. Smyth, D. F. Boesch, and J. van Montfrans. 1979. Comparative biogeography of benthic macrocrustaceans of the Middle At- lantic continental shelf. — Bulletin of the Biolog- ical Society of Washington 3:214—255. Chace, F.A., Jr. 1940. The brachyuran crabs. Reports on the scientific results of the Atlantis expedi- tions to the West Indies, under the joint auspices of the University of Havana and Harvard Uni- versity. —Torreia 4:1—67. Ingle, R. W. 1985. Geryon gordonae sp. nov. (De- capoda Brachyura, Geryonidae) from the north- eastern Atlantic Ocean. — Crustaceana 48(1):88— 98. ,and A. L. Rice. 1984. The juvenile stages of eight swimming crab species (Crustacea: Brachyura: Portunidae); a comparative study. — Bulletin of the British Museum (Natural His- tory), Zoology 46(4):345-354. Lewis, E., and P. A. Haefner, Jr. 1977. Intraspecific setal variation in Bathynectes superbus (Costa) (Decapoda, Portunidae).—Crustaceana 33(2): 163-166. Luckhurst, B. 1985. Discovery of deep-water crabs (Geryon spp.) at Bermuda—a new potential fish- ery resource.— Proceedings of the 37th Annual Gulf and Caribbean Fisheries Institute, 1984 [in press]. Manning, Raymond B., and L. B. Holthuis. 1981. West African Brachyuran Crabs.—Smithsonian Contributions to Zoology 306:1-379. ——, and . 1984. Geryon fenneri, anew deep- water crab from Florida (Crustacea: Decapoda: Geryonidae).—Proceedings of the Biological Society of Washington 97(3):666-673. Markham, John C., and John J. McDermott. 1981. A tabulation of the Crustacea Decapoda of Ber- muda.— Proceedings of the Biological Society of Washington 93(4):1266—-1276. 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:i—xli, 1-362. Rathbun, Mary J. 1930. The cancroid crabs of Amer- ica of the families Euryalidae, Portunidae, Ate- lecyclidae, Cancridae and Xanthidae.— United States National Museum Bulletin 138:i—xvi, 1- 609. Soto, Luis A. 1985. Distribution patterns of deep- VOLUME 99, NUMBER 2 water brachyuran crabs in the Straits of Flori- da.—Journal of Crustacean Biology 5(3):480- 499. Wenner, E. L., and D. F. Boesch. 1979. Distribution patterns of epibenthic decapod Crustacea along the shelf-slope coenocline, Middle Atlantic Bight, U.S.A.—Bulletin of the Biological Society of Washington 3:106-133. Williams, A. B., L. R. McCloskey, and I. E. Gray. 1968. New records of brachyuran decapod Crustaceans from the continental shelf off North Carolina, U.S.A.—Crustaceana 15(1):41-66. Williams, A. B.,and R. L. Wigley. 1977. Distribution of decapod Crustacea off northeastern United S75 States based on specimens at the Northeast Fisheries Center, Woods Hole, Massachu- setts.—NOAA Technical Report NMFS Cir- cular 407:i-111, 1-44. (RBM) Department of Invertebrate Zo- ology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560; (LBH) Rijksmuseum van Na- tuurlijke Historie, Post Box 9517, 2300 RA Leiden, The Netherlands. sma apaiile kane ee dro a Pe ee taba shale very a nadlaanan sie «noe wink? soingAait oe 1 PRA foqak lmpinriga’? AML, ~ wise adeags 7 WTP ns a m a : 4 7 t rr : “} ah stitch al Wot afl { > pigitt ainabé sthbi satanse 34 4 « ide _ ot ‘a. sipdiiabe omen ye uleot? sciaaimminaine vie vl Bil imu 0 most Tnpnite) oral i can Hisrysihee ry aoe ghey ie rir Ppgies yl Ajay a Soe ve. nay nie els (TOD sey OAT hoger nny! ia? Yo Ae BO ee © 170 eA Acryl oreseiti ee Aj WAT ES Mater ras Me ri 4 % Tbe oa ae Uh hl ok wTlavial (91 I Se ni AIRS. J ‘A, vy Thee OR" Pr Cre ae lely 0 eng eds 1) OA Bad eh veal uy st ~ ae S, a ‘ / (é nee Mm: ¥ 2 U Lj 14 id ’ la ae Madly: ol: : & F bpd PO Meee Om i i idiay Ft Neer, * Foal 4 Nesdlncad Wadia Th rind % € r Land , ‘ L 4 {> i0hep LEN fat 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 will be published in English (except for Latin diagnosis/description of plant taxa which should not be duplicated by an English translation), with a summary in an alternate language when appropriate. Publication Charges.—Authors will be asked to assume publication costs of page-charges, tabular material, and figures, at the lowest possible rates. Submission of manuscripts.—Manuscripts should be sent to the Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History, 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. Manuscripts will be reviewed in order of receipt by a board of associate editors and appropriate referees. Presentation.—Clarity of presentation, and requirements of taxonomic and nomenclatural procedures necessitate reasonable consistency in the organization of papers. Telegraphic style is recommended for descriptions. Literature should be cited in abbreviated style (author, date, page), except in botanical synonymies, with full citations of journals and books (no abbrevia- tions) in the Literature Cited section. The establishment of new taxa must conform with the requirements of the appropriate in- ternational codes of nomenclature. Authors are expected to be familiar with these codes and to comply with them. New species-group accounts must designate a type-specimen deposited in an institutional collection. The sequence of material should be: Title, Author(s), Abstract, Text, Acknowledgments, Literature Cited, Author’s (s’) Address(es), Appendix, List of Figures (entire figure legends), Figures (each numbered and identified), Tables (each table numbered with an Arabic numeral and heading provided). Manuscripts should be typed, double-spaced throughout (including tables, legends, and foot- notes) on one side of 82 x 11 inch sheets, with at least one inch of margin all around. Manu- scripts in dot-matrix will not be accepted. Submit a facsimile with the original, and retain an author’s copy. Pages must be numbered on top. One manuscript page = approximately 0.5 printed page. Underline singly scientific names of genera and lower categories; leave other indications to the editor. Figures and tables with their legends and headings should be self-explanatory, not requiring reference to the text. Indicate their approximate placement by a pencil mark in the margin of the manuscript. Illustrations should be planned in proportions that will efficiently use space on the type bed of the Proceedings (12.5 x 20 cm) and should not exceed 15 x 24 inches. Figures requiring solid black backgrounds should be indicated as such, but not masked. Art work will be returned only on request. Proofs.—Galley proofs will be submitted to authors for correction and approval. Reprint orders will be taken with returned proof. Costs.—Page charges @ $60.00, figures @ $10.00, tabular material $3.00 per printed inch. All authors are expected to pay the charges for figures, tables, changes at proof stage, and reprints. Payment of full costs will probably facilitate speedy publication. CONTENTS Neastacilla falclandica (Ohlin), type species of the genus, and N. tattersalli, new species (Crus- tacea: Isopoda: Arcturidae) Helen M. Lew Ton and Gary C. B. Poore Alpheopsis cortesiana, a new snapping shrimp from the gulf of California Mary K. Wicksten and Michel E. Hendrickx Freshwater shrimps from Venezuela I: Seven new species of Palaemoninae (Crustacea: De- capoda: Palaemonidae) Guido Pereira S. A new Colostethus (Anura: Dendrobatidae) from Ecuador Darrel R. Frost Systematics of the gorgeted woodstars (Aves: Trochilidae: Acestrura) Gary R. Graves Tonocote, a new genus and species of Zobrachoidae from Argentina (Crustacea: Marine Am- phipoda) Janice Clark and J. L. Barnard Paralebbeus zotheculatus, n. gen., n. sp., anew hippolytid shrimp from the Australian northwest shelf A. J. Bruce and F. A. Chace, Jr. Rhombopsammia, a new genus of the family Micrabaciidae (Coelenterata: Scleractinia) Joan Murrell Owens Discovery of a new living Cerithioclava species in the Caribbean (Mollusca: Prosobranchia: Cerithiidae) Richard S. Houbrick ‘Notes on the species of Ad/osquilla and Platysquilloides (Crustacea: Stomatopoda) Carlo Froglia and Raymond B. Manning A small trap for collecting crustaceans in shallow water Raymond B. Manning Tridentella recava, a new isopod from tilefish burrows in the New York Bight (Flabellifera: Tridentellidae) Thomas E. Bowman Sexual dimorphism and a sex-limited polymorphism in the copepod Paroithona pacifica Ni- shida, 1985 (Cyclopoida: Oithonidae) from the Red Sea Frank D. Ferrari and Ruth Bottger Marine Tubificidae (Oligochaeta) at Hutchinson Island, Florida Christer Erséus Caecidotea dauphina, a new subterranean isopod from a barrier island in the northern Gulf of Mexico (Crustacea: Isopoda: Asellidae) Richard F. Modlin Setal morphology of the oligochaetes Tubifex tubifex and Ilyodrilus frantzi (capillatus) as revealed by SEM Peter M. Chapman and Ralph O. Brinkhurst Serrabrycon magoi, a new genus and species of scale-eating characid (Pisces: Characiformes) from the upper Rio Negro Richard P. Vari Nematocharax venustus, a new genus and species of fish from the Rio Jequitinhonha, bes Gerais, Brazil (Teleostei: Characidae) Stanley H. Weitzman, Naércio A. Menezes, and Heraldo A. Britski Additional records for Callianassa rathbunae Schmitt, 1935, from Florida and the Bahamas (Crustacea: Decapoda: Callianassidae) Raymond B. Manning and Richard W. Heard Pseudojaniridae (Crustacea: Isopoda), a new family for Pseudojanira stenetrioides Barnard, 1925, a species intermediate between the asellote superfamilies Stenetrioidea and Janiroidea George D. F. Wilson Transfer of Quadrasia from the Planaxidae to the Buccinidae (Mollusca: Gastropoda: Proso- branchia) Richard S. Houbrick Preliminary descriptions of four new species of dorippid crabs from the Indo-West Pacific region (Crustacea: Decapoda: Brachyura) Raymond B. Manning and L. B. Holthuis Notes on Geryon from Bermuda, with the description of Geryon inghami, new species (Crus- tacea: Decapoda: Geryonidae) Raymond B. Manning and L. B. Holthuis 3395 347 350 359 363 366 RG ty us? eed : “ 4 X : a 4 - j ' : oy t 3 ; A SZ) ey i, q | Lo THE BIOLOGICAL SOCIETY OF WASHINGTON 1985-1986 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 Kensley Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Richard P. Vari Insects: Robert D. Gordon 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: $18.00 within the U.S.A., $23.00 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 Instutution, 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. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 375-379 SPELEOBREGMA LANZAROTEUM, A NEW GENUS AND SPECIES OF SCALIBREGMATIDAE (POLYCHAETA) FROM A MARINE CAVE IN THE CANARY ISLANDS Rodney Duane Bertelsen Abstract. —Speleobregma lanzaroteum, a new genus and species of polychaete in the family Scalibregmatidae, is described from a marine lava tube cave, Jameos del Agua, on Lanzarote Island, Canary Islands. It differs from all scal- ibregmatid genera by possessing hirsute hooded geniculate setae. Its placement within the family is discussed. During a recent cave—diving expedition on Lanzarote in the Canary Islands (Fig. 1), a new genus and species of scalibregmatid was collected by Dennis Williams. The ex- pedition was part of an ongoing study of insular aquatic cave fauna (Iliffe et al. 1984). Although only one specimen was found, AFRICA Fig. 1. LANZAROTE it is in excellent condition due to careful collection and gentle handling. The speci- men contains unique and important sys- tematic characters that warrant a descrip- tion. Due to the difficulties of cave diving and potential scarcity of material, addition- al specimens may not be available in the pyameos "del Agua Map of Lanzarote, Canary Islands. The “‘x”’ indicates location of Jameos del Agua, the collection site of Speleobregma lanzaroteum. The inset shows the relationship of the Canaries to the coast of Africa and Europe. Lanzarote is the darkened island. 376 foreseeable future. The holotype is depos- ited in the National Museum of Natural History, Smithsonian Institution (USNM). Speleobregma, new genus Etymology. —‘“‘Speleo”’ referring to cave, ‘bregma’ from the stem of the family. Gen- der: Neuter. Type species. —Speleobregma lanzaro- teum, new species Diagnosis. — Body elongate, slender, and cylindrical. Prostomium blunt, entire, with lateral horns. Acicular setae and furcate se- tae lacking; branchiae absent; capillary setae in both rami with hirsute hooded geniculate setae in neuropodia. Speleobregma lanzaroteum, new species Figs. 2, 3 Material examined.—Jameos del Agua lava tube cave, Lanzarote, Canary Islands, 4 Mar 1984, 29°12'N, 13°38’W, 20 m, SCU- BA, coll. D. Williams, holotype, USNM 98275. Diagnosis. —Neuropodial hirsute hooded geniculate setae present on all setigers. Para- podia with neuropodial postsetal lamellae and interramal papillae. Pygidium lacking anal cirri; with 2 ventral spherical papillated lobes. Description. —Holotype 8 mm in length, 0.8 mm in width, 23 setigers. Body color- less, elongate, cylindrical and slender; se- tigers 7-11 slightly inflated (reduced aren- icoliform). Dorsal surface of anterior segments weakly biannulate and areolated. Prostomium (Fig. 2a) triangular, ante- riorly blunt with a pair of tapered lateral horns, eyes absent. Nuchal organs not evi- dent. Proboscis ventral, forming soft, smooth, eversible pouch. Buccal segment apodous and achaetous. Each parapodium with single spherical interramal organ and cylindrical neuropo- dial postsetal lamella (Fig. 2b). Lamellae conspicuous on all but last 2 setigers. Bran- chiae and cirri absent. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON All notosetae simple capillaries inserted in single row; up to 20 per fascicle. Neu- rosetae inserted in 2 rows. Anterior row shorter than posterior row, consisting of 5 to 9 hirsute, distally abruptly tapered, hood- ed geniculate setae (Fig. 3b). Posterior row all capillaries. Pygidium with 2 heavily papillated ven- tral spherical lobes. Lobes with anteroven- tral groove extending halfway across ventral surface (Fig. 3a). Anal cirri lacking. Distribution. —Known only from type lo- cality, Jameos del Agua Cave, Lanzarote, Canary Islands. Etymology. —The specific epiphet is tak- en from the type locality, Lanzarote Island. Discussion Lanzarote and its large marine lava tube cave, Jameos del Agua, are noted for their unusual fauna (Dinkins 1969; Iliffe et al. 1984). This species, typical of many cave organisms, is eyeless and lacks pigment. The thickened areolated epidermis, common among scalibregmatids, is reduced in this species. Staining with fast-blue aided in re- vealing this character. Following the organization of the family Scalibregmatidae proposed by Blake (1981), Speleobregma belongs to body type I (ar- enicoliform with T-shaped prostomium) and parapodial type D (dorsal and ventral cirri absent; prolonged postsetal lamellae pres- ent). This classification includes the genera Scalibregmides (Hartmann-Schroder 1965) and an undescribed genus (for Asclerochei- lus californicus Hartman, 1963). Speleo- bregma differs from them in having long postsetal lamellae only on the neuropodia and in lacking furcate setae. The only other species lacking furcate se- tae is Kebuita minuta Hartman, 1967. Spe- leobregma differs in body form in that Ke- buita minuta is maggotlike. Speleobregma differs from all scalibreg- matid genera by possessing hirsute hooded geniculate setae. VOLUME 99, NUMBER 3 25mm Fig. 2. Speleobregma lanzaroteum: A, Anterior end, dorsal view; B, Setigers 3-5, lateral view. 377 378 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON JS. : : 4) eee ca ee as the he ots, . ee rt Pay, Be, toes: SERRE A OY Lis 4 uke uy by Fig. 3. Speleobregma lanzaroteum: A, Pygidium, ventral view; B, Neuropodial hirsute hooded geniculate seta. VOLUME 99, NUMBER 3 Acknowledgments I wish to thank Jill Yager and Dennis Williams for permission to examine their collection of marine cave polychaetes and for helpful discussions about their research. Drs. John R. Holsinger and Marian Petti- bone gave valuable support and advice. Dr. Kristian Fauchald examined the specimen. The manuscript benefitted from reviews by Dr. Fauchald and an anonymous reviewer. The collection of material studied is sup- ported by a grant from the National Science Foundation to Dr. Thomas M. Iliffe (BSR- 8215672). Literature Cited Blake, J. A. 1981. The Scalibregmatidae (Annelida: Polychaeta) from South America and Antarctica collected chiefly during the cruises of the R/V Anton Bruun, R/V Hero and USNS Eltanin. — Proceedings of the Biological Society of Wash- ington 94(4):1131-1162. B79 Dinkins, S. 1969. Lanzarote, the strangest Canary. — National Geographic Magazine 135(1):117-139. Hartman, O. 1963. Submarine canyons of southern California. Part 3. Systematics: Polychaetes. — Allan Hancock Pacific Expeditions 27(3):1-93. . 1967. Polychaetous annelids collected by the USNS Eltanin and Staten Island cruises, chiefly from Antarctic seas.—Allan Hancock Mono- graphs in Marine Biology 2:1-387. Hartmann-Schroder, G. 1965. Die Polychaeten des Sublitorals. Jn Hartmann-Schroder, G., and G. Hartmann. Zur Kenntnis des Sublitorals der chilenischen Kiiste unter besonderer Beriick- sichtigung der Polychaeten und Ostracoden. — Mitteilungen aus dem Hamburgischen Zoolo- gischen Museum und Institut, Supplement 62: 59-305. lliffe, T. M., H. Wilkens, J. Parzefall, and D. Williams. 1984. Marine lava cave fauna: Composition, biogeography, and origins.—Science 225:309- 311. Department of Biological Sciences, Old Dominion University, Norfolk, Virginia 23508, U.S.A. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 380-383 SYMMETROSCYPHUS, A NEW GENUS OF THECATE HYDROID (FAMILY THYROSCYPHIDAE) FROM BERMUDA Dale R. Calder Abstract. — A new genus, Symmetroscyphus, is proposed for the single species Thyroscyphus intermedius Congdon, 1907, a shallow water hydroid of the family Thyroscyphidae Stechow, 1920, from Bermuda. Unlike the closely allied genus Thyroscyphus Allman, 1877, with its erect hydroid colonies and bilat- erally symmetrical hydrothecae, Symmetroscyphus typically has stolonial col- onies and radially symmetrical hydrothecae. Symmetroscyphus intermedius is superficially similar to but morphologically distinct from Calamphora parvula Allman, 1888, Sertularella solitaria Nutting, 1904, and Sertularella campan- ulata Warren, 1908. Thyroscyphus intermedius Congdon, 1907, is a small thecate hydroid presently known only from Bermuda, where it is com- mon to abundant in shallow water on turtle grass (Thalassia). Colonies are brownish in color when alive due to the presence of algal symbionts in the coenosarc. Hydroids of this species superficially resemble those of Cal- amphora parvula Allman, 1888, Sertular- ella solitaria Nutting, 1904, and Sertular- ella campanulata Warren, 1908, in their colony form, which is typically stolonial, and in having hydrothecae with annulated walls and four marginal teeth. Because of the similarity of these species, 7. interme- dius has been included as a synonym of C. parvula (as Sertularella parvula) by Ver- voort (1968) and as a questionable synonym of S. campanulata by Stechow (1919). Spec- imens of 7. intermedius from Bermuda dif- fer from type material of both Calamphora parvula and Sertularella campanulata, as well as Sertularella solitaria, in having ra- dially rather than bilaterally symmetrical hydrothecae, a centrally located rather than an excentric hydropore, an annular fold about the hydranth base instead of an ab- cauline diverticulum, perisarc of uniform thickness around the perimeter of the hy- drothecal wall instead of thicker perisarc on the side opposite the excentric hydropore, and clusters of large nematocysts in an in- trathecal sheath of ectoderm. Unlike C. par- vula, the hydrothecal orifice of JT. inter- medius is perpendicular to the axis of the hydrotheca, and intrathecal teeth are lack- ing. Based on such major differences, T. in- termedius cannot be retained in the syn- onymy of either C. parvula ors. (a ] \ Figs. 1-3. Symmetroscyphus intermedius: 1, Hy- drotheca with renovated margin, ROMIZ B313; 2, Hy- drotheca, ROMIZ B314; 3, Erect colony, ROMIZ B314. Scale bars equal 500 um. VOLUME 99, NUMBER 3 381 Fig. 4. Large nematocysts from ectodermal sheath lining hydrothecal cavity of Symmetroscyphus intermedius, ROMIZ B314. Scale bar equals 20 um. campanulata, nor can it be assigned to Cal- amphora Allman, 1888, Sertularella Gray, 1848, Thyroscyphus Allman, 1877, or any other described genus. A new genus, Sym- metroscyphus, is established for the species in this report. Institutional abbreviations used in this paper are: BMNH—British Museum (Nat- ural History); NM— Natal Museum; ROM- IZ—Royal Ontario Museum, Department of Invertebrate Zoology. Materials and Methods Type materials of Calamphora parvula (BMNH 1888.11.13.65A) from Bass Strait, Australia, and Sertularella campanulata (NM 751) from Natal, were obtained on loan and compared with specimens of Thy- roscyphus intermedius (ROMIZ B186, ROMIZ B205, ROMIZ B313, ROMIZ B314) from Bermuda. Nematocysts of for- malin-preserved materials of 7. interme- dius (ROMIZ B314) were examined in un- stained tissue squashed under a coverslip. All observations were made using bright- field microscopy. Symmetroscyphus, new genus Figs. 1-4 Diagnosis. —Hydroid colonies occasion- ally with erect hydrocauli, but typically sto- lonial. Hydrothecae pedicellate in both sto- lonial and erect colonies, radially symmetrical, barrel-shaped, margin with 4 teeth, submarginal teeth absent. Operculum pyramid-shaped, composed of 4 triangular valves. Diaphragm present, with centrally located hydropore. Hydranth with annular fold basally, abcauline diverticulum absent, with large nematocysts in body wall. Inner 382 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON surface of hydrothecal wall lined to some extent with ectoderm, ectodermal layer with aggregation of large nematocysts beneath each marginal tooth. Gonophores unknown. Type species.—Thyroscyphus interme- dius Congdon, 1907, designated herein. Etymology.—The name is derived from the Latin (originally Greek) words sym- metria (symmetry) and scyphus (goblet) in allusion to the radial symmetry of the hy- drotheca; the gender is masculine. Remarks.—In being stolonial, Symme- troscyphus superficially resembles the gen- era Calamphora and Sertularella of the family Sertulariidae Fleming, 1828, but dif- fers in having hydrothecae that are radially instead of bilaterally symmetrical, a hydro- pore that is centrally located instead of ex- centric, an annular fold about the hydranth base rather than an abcauline diverticulum, and large nematocysts of an undetermined category (Fig. 4) in the ectoderm. Symme- troscyphus is most closely allied to Thyro- scyphus Allman, 1877, of the family Thy- roscyphidae Stechow, 1920, from which it differs in having a stolonial instead of an erect colony form, and in having symmet- rical hydrothecal walls instead of a more protuberant adcauline than abcauline wall. The gonophores of Symmetroscyphus are unknown. Gonothecae were present in hy- droids identified by Leloup (1935) as Thy- roscyphus intermedius forme peculiaris. However, Leloup’s specimens are regarded here as belonging to a species other than S. intermedius, and in fact correspond with di- agnoses of the genus Sertu/arella rather than Symmetroscyphus. Symmetroscyphus is a monotypic genus, referable to the family Thyroscyphidae. The only species assigned here to the genus is its type species, Thyroscyphus intermedius. Acknowledgments I am indebted to Dr. P. F. S. Cornelius and Mr. S. J. Moore of the British Museum (Natural History) for loan of the type ma- terial of Calamphora parvula, and to Dr. Londt of the Natal Museum for loan of the type material of Sertularella campanulata. This study was supported by funds from the Royal Ontario Museum, the Canadian As- sociates of the Bermuda Biological station, and the Exxon Corporation. The paper is contribution number 1103 from the Ber- muda Biological Station for Research, Inc. Literature Cited Allman, G. J. 1877. Report on the Hydroida collected during the exploration of the Gulf Stream by L. F. de Pourtalés, assistant United States Coast Survey.— Memoirs of the Museum of Compar- ative Zoology at Harvard College 5(2):1-66. 1888. Report on the Hydroida dredged by H.M.S. Challenger during the years 1873-86. Part II. The Tubularinae, Corymorphinae, Campanularinae, Sertularinae, and Thalamo- phora.— Report on the Scientific Results of the Voyage of H.M.S. Challenger during the Years 1873-76, Zoology 23:1—90. Congdon, E. D. 1907. The hydroids of Bermuda.— Proceedings of the American Academy of Arts and Sciences 42:463—485. Fleming, J. 1828. A history of British animals, ex- hibiting the descriptive characters and system- atical arrangement of the genera and species of quadrupeds, birds, reptiles, fishes, Mollusca, and Radiata of the United Kingdom. Bell and Brad- fute, Edinburgh, 565 pp. Gray, J. E. 1848. List of the specimens of British animals in the collection of the British Museum. Part I. Centroniae or radiated animals. London (British Museum), 173 pp. Leloup, E. 1935. Hydraires calyptoblastiques des Indes Occidentales.— Mémoires du Musée Royal d’Histoire Naturelle de Belgique 2,2:1-73. Nutting, C.C. 1904. American hydroids. Part II. The Sertularidae.—Smithsonian Institution, United States National Museum Special Bulletin 4(2): 1-325. Stechow, E. 1919. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer Ge- biete, nebst Angaben tiber einige Kirchenpa- uer’sche Typen von Plumulariden.—Zoolo- gische Jahrbiicher, Abteilung fiir Systematik, Geographie und Biologie der Tiere 42:1-172. . 1920. Neue Ergebnisse auf dem Gebiete der Hydroidenforschung.—Sitzungsberichte der Gesellschaft fiir Morphologie und Physiologie in Miinchen 31:9-45. VOLUME 99, NUMBER 3 Vervoort, W. 1968. Report on a collection of Hy- droida from the Caribbean region, including an annotated checklist of Caribbean hydroids.— Zoologische Verhandelingen 92:1-124. Warren, E. 1908. Onacollection of hydroids, mostly from the Natal coast.—Annals of the Natal Government Museum 1:269-355. 383 Department of Invertebrate Zoology, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6, Canada, and Department of Zoology, University of To- ronto, Toronto, Ontario MSS 1A1, Canada. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 384-387 THE STATUS OF THE OPHIDIID FISHES OPHIDIUM BREVIBARBE CUVIER, OPHIDIUM GRAELLSI POEY, AND LEPTOPHIDIUM PROFUNDORUM GILL C. Richard Robins Abstract.—Ophidium brevibarbe Cuvier is shown to be the oldest valid name for a common and wide-ranging species of Lepophidium, of which Ophidium graellsi Poey is a junior synonym and Lepophidium profundorum auctorum is a common misattribution. Leptophidium profundorum Gill, type species both of Leptophidium (preoccupied) and of Lepophidium, its replacement name, is a senior synonym of L. cervinum (Goode and Bean), a common species in shelf waters along the eastern seaboard of the United States and southeastern Canada. Several nomenclatural problems have de- layed a revision of the ophidiid genus Le- pophidium, now nearing completion. The answers to these problems are important because they involve: a) the most common shallow-water species which ranges from the southeastern United States and the North- ern Gulf of Mexico to the southern coast of Brazil, b) the earliest name in the genus, and c) the type species of the genus. The solution of these problems is the object of this paper. Because the names of two common species are changed as a result, this information is being made available separately so that they will be available to regional works and checklists now in preparation. Counts and measurements used in this paper (Table 1) are as defined by Robins (1962:487—488). Ophidium brevibarbe Cuvier, 1829 Cuvier (1829:359) simply stated ““Nous en connaissons une troisiéme espéce du Brésil (Oph. Brevibarbe N.) brune, a bar- billons plus courts; ...’’ This species can- not be identified with any particular species on the basis of this statement alone. How- ever, type material of the species exists. We have examined MNHM 5772, the type ma- terial of Ophidium brevibarbe. There are two specimens. One, the Lepophidium illustrat- ed by Kaup (1856a:95; 1856b:154, pl. 16, fig. 4) clearly shows the rostral spine and head squamation which uniquely define the genus Lepophidium. The other is a speci- men of Ophidion holbrooki (Putnam). Le- pophidium brevibarbe and Ophidion hol- brooki are the most common and widespread species of their respective genera, occurring together from the southeastern and Gulf coasts of the United States to the southern coast of Brazil. Both are species of the inner shelf. They are common “‘jar mates” in mu- seums because early collectors commonly preserved one specimen of each type of or- ganism which they obtained. It also was common practice to describe one specimen as a new species and return it to its jar which contained other species. Nothing in Kaup’s description suggests a second specimen, and, since Ophidion holbrooki has very long pel- vic rays, it could well be that Cuvier paid no attention to the second, smaller speci- men. Kaup (1856a:95) recorded the length of the specimen as 200 mm. I measure it at 202 mm, but it is now quite soft. There is little question but that this is the specimen described by Kaup. If both specimens in MNHM 5772 are regarded as syntypes, we may regard Kaup (1856a, b) as having re- VOLUME 99, NUMBER 3 stricted the name brevibarbe to the larger specimen, which becomes the lectotype, the specimen of Ophidion holbrooki becoming a paralectotype of L. brevibarbe. Alterna- tively, one can regard the largest specimen as the holotype, and the smaller specimen as lacking type status as it did not enter into the description. I have followed the first course here. Counts and measurements of both specimens are given in Table 1. That brevibarbe was a Lepophidium was correctly noted long ago by Gill (1863:210). Ophidium graellsi Poey, 1861 Poey (1861:425-—426) described Ophi- dium graellsi in some detail, based on a specimen 230 mm long (=total length). He noted that the head was scaled except for the very tip of the snout. He also said that it lacked pyloric caeca. His later accounts (1868:402; 1876:137) added nothing, the latter being a mere listing of the name. Jor- dan and Evermann (1898:2488) correctly emended the spelling to graellsi and rede- scribed the species based on a small speci- men from Havana, sent to them by Poey. They noted six developed gill rakers on the lower limb of the first gill arch (O. holbrooki has only four), a sharp spine at the snout tip, and a lanceolate swimbladder. Poey also sent a drawing of his type and a description, a short translation of the original, which Jordan and Evermann included in their ac- count. Apparently two specimens of cusk- eels were sent to the Museum of Compar- ative Zoology as Ophidium graellsi and I consider them to be Poey’s original mate- rial. They are now catalogued as MCZ 12440 and 12441 and bore Poey’s original num- bers 729 and 720 respectively. MCZ 12440 is a specimen of Lepophidium brevibarbe (Cuvier) 216 mm standard length. It is gut- ted as expected if Poey has removed the gut for description. The specimen agrees with all of the salient features noted by Poey (1861) and Poey (in Jordan and Evermann 1898), except that one cannot attest to the 385 pyloric caeca. All species of Lepophidium always have at least a few small pyloric cae- ca. These could have been overlooked or the statement could have been based on MCZ 12441, a specimen of Ophidion hol- brooki (Putnam). It also is gutted but all species of this genus lack pyloric caeca. Al- though it bears a label “‘Poey’s orig.,” it can- not be the type as it is too small and lacks scales on the head and a forward-projecting rostral spine. MCZ 12440, the specimen of L. brevibarbe (Cuvier), is the holotype as Poey mentioned only one specimen. Poey commonly described species from one spec- imen stored with other, not necessarily re- lated, species. There is no reason, except for the statement concerning the absence of py- loric caeca, to regard the specimen of Ophi- dion holbrooki (=MCZ 12441) as a paralec- totype. Ophidium graellsi Poey is therefore a junior synonym of Lepophidium brevi- barbe (Cuvier). Counts and measurements of both specimens are given in Table 1. Leptophidium profundorum Gill, 1863 Gill described this species from a single specimen collected by Commodore Rogers off the east coast of Florida in 60 fms (277 m). It is the type species, by monotypy, of Leptophidium Gill, 1863 (preoccupied) and also of the replacement genus Lepophidium (Gill, 1895). This name, which represents a valid species, has commonly been misap- plied to the common shallow-water species whose correct name is L. brevibarbe. Counts and measurements of the holotype (USNM 6247), are given in Table 1. Its gill chamber and gill bars are blackish, the peritoneum is silvery, and the esophagus is black. All these structures are without pigment in L. brevibarbe. The specimen is badly faded so that no external pigment pattern is evident. The counts, proportions, and the internal pigment noted above are identical with those of Lepophidium cervinum (Goode and Bean, 1885), a common shelf species character- ized by a row of pale spots on a tan back- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 386 "(4x91 99s) adAjojOg[RIeg 41 ‘adAjojoy 10 adAjo}097T | 6=L506 8=9+7 9L=09/91 ¢L=09/S1 CC-CT PC-ET Lil Lil Lel cel € — m= oT 97 Sc Be 8E cs €¢ VC €¢ [tet 3 Oe 0's 0 gs Uv $8 6L 98 c8 v8 £8 £6 86 86 C6 cl IT 9€ ce £7 CC LI 91 BIZ OT7 6=L+7 LL=8S/91 €C-EC vit vel LI CEC 6=S+7 ¢L=8S/L1 CC-CC 9 9EI £ LI LC 9€ vs CT ce 67 8'P OL Ol Ol Ol 68 cl ve 16 LI 891 9=7+7 89=€S/SI Oc-I¢ vil Ovl vi Lé OV cS Ic ORS el 68 val! le LT 81 O9T OI=L+t CC-CC 601 Tel 81 61 9IC 9=7+C 99=05/91 CC-CC 901 vel 0 el VC vi Iv 9C Ic €cl 6=9+E€ TL=9S/S1 61-61 LO! 9TI 91 (YG cS 9¢ Ic vs OL SIOCIT[LO (jepnes + [epnevooid) seIqoLIO A sAvl UY-[210}990g sAvl ug-[euy sAvl uy-[es10q 20989 OLIO[AG x[e1gio19jur Auog xTOJOUILIP WQIO ,2][IXeU JO pus 0} dj ynous ,U41SUI] [211G101]SOg ,41dUI] NOUS (19UUT) IOTLIOISOg (7 (19]NO) JOLINUY (| yisug] Avi SIAlog yi3us] Uy-jepneD yisus] UY-[e10}09g ynd1990 1yV (¢€ uIsLIO Ug-jeuR ly (Z uIdLIO UY-[eSIOP 1V (1 Apog jo yidaq dul] [e191e] JO pus 0} dy jnous ynd1990 0} dj jnous 0uejsIp (UY) [euUvoIg 90UvjSIP [eSIOPIIg yisus] peop (Ww) 413U9] prepurisS —— ae - c I 4igsz7plzt WNSN 4p9L87 WNSN 47L¢9 WNSN Ippcl ZOW L0bpzI ZOW 447 c7LS WHNW 41 TLLS WHNW mn “y3u9] peay (,) 10 prepurys yuddI0d ul possoidxa oie suonsodolg (8SZPLTZ ‘P9L8Z WNSM) Winuidsas uinipiydojdaT pue (7PL9 WNSM) Wnsopunfoad uunipiydojdaT “(IppZI ‘ObPZI ZOW) ssyjan43 wnipiydo (ZLLE NHNW) 2940qIA04q wnipiydgC Jo susutioads juRAa|aI Joyo pue sodA} ay) Joy BIepP SLNEWOYdIOW —"] I4eL VOLUME 99, NUMBER 3 ground, which ranges from Florida to east- ern Canada. Although the probable identity of profundorum has been known to me for years, new species in the genus were being encountered regularly, and I wished to be certain that there was not a very similar, but unpatterned species, in the region. That no other species shares this combination of characters is now clear. Lepophidium cer- vinum (Goode and Bean) is, therefore, a ju- nior synonym of Lepophidium profundo- rum Gill. Goode and Bean (1885:422) designated one specimen, USNM 28764, 262 mm total length, as the type. The jar bearing this number contained three spec- imens, the smaller two of which are recat- aloged as paratypes, USNM 274258. Counts and measurements of these three specimens are given in Table 1. Lepophidium cervinum is not of sufficient importance under guide- lines laid down by the International Com- mission of Zoological Nomenclature, to merit a request to the Commission for con- servation of the name. The use of Lepophidium brevibarbe by Robins (1958: fig. 1d) for a species with short pelvic fins and a dark gill chamber was in error. That species lacks a name and is being described elsewhere. Acknowledgments Many persons have supplied information and loaned material relevant to this study, or made facilities available to me at their institutions. In particular, I am indebted to Marie-Louise Bauchot (Muséum National d’Histoire Naturelle, Paris; the late Henry B. Bigelow, Myvanwy Dick, and Karsten Hartel (Museum of Comparative Zoology, Harvard University); Giles W. Mead and Daniel M. Cohen (then of the Systematics Laboratory, National Marine Fisheries Ser- vice); and Leonard P. Schultz, Ernest A. Lachner, Robert H. Kanazawa, and Susan L. Jewett (National Museum of Natural History). Aspects of this study were dis- cussed with the late Luis Howell-Rivero and, most thoroughly, with Robert N. Lea, Cal- ifornia Department of Fish and Game, to 387 whom I am especially indebted. Catherine H. Robins commented on the manuscript. Some of this work was done with grant sup- port through NSF-G-7116, G105. The Maytag Chair of Ichthyology has supported me throughout. Literature Cited Cuvier, G. L. C. F. D. 1829. Le Régne Animal. 2nd ed., Paris, vol. 2, i-xv, 1-406. Gill, T. 1863. Description of a new generic type of ophidioids.— Proceedings of the Academy of Natural Sciences of Philadelphia 1863:209-211. . 1895. The genus Leptophidium.— American Naturalist 29:167-168. Goode, G. B., and T. H. Bean. 1885. Description of Leptophidium cervinum and L. marmoratum, new fishes from deep water off the Atlantic and Gulf coasts.— Proceedings of the United States National Museum 8(533):422-424. Howell Rivero, L. 1938. List of the fishes, types of Poey, in the Museum of Comparative Zoolo- gy.—Bulletin of the Museum of Comparative Zoology 83(3):169—227. Jordan, D. S., and B. W. Evermann. 1898. The fishes of North and Middle America. — Bulletin of the United States National Museum 47(3):i—xxiv + 2183a—3136. Kaup, J.J. 1856a. Einige iiber die Unterfamilie Ophi- diinae.— Archiv fiir Naturgeschichte 22(1):93- 100. 1856b. Catalogue of apodal fish, in the col- lection of the British Museum. British Museum, London, vii + 163 pp. Poey, F. 1858-1861. Memorias sobre la historia nat- ural de la Isla de Cuba. Habana, vol. 2:1—442. [Cited pages were published in 1861]. 1866-1868. Repertorio fisico-natural de la Isla de Cuba. Habana, vol. 2:1—484. [Cited pages were published in 1868]. . 1876. Enumeratio Piscitum Cubensium. Part 2.—Anales de la Sociedad Espanola de Historia Natural 9:131-218 (89-176). Robins, C. R. 1958. Studies on Fishes of the family Ophidiidae. I. A new species of Lepophidium from the Caribbean Sea.—Bulletin of Marine Science of the Gulf and Caribbean 81(4):360- 368. . 1962. Studies on Fishes of the family Ophi- diidae. VII. The Pacific species of Lepophi- dium. —Copeia 1962(3):487—498. Maytag Professor of Ichthyology, Rosen- stiel School of Marine and Atmospheric Sci- ences, University of Miami, 4600 Ricken- backer Causeway, Miami, Florida 33149. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 388-391 A REDESCRIPTION OF COLOBOMATUS MYLIONUS FUKUI FROM AUSTRALIAN ACANTHOPAGRUS (SPARIDAE) (CRUSTACEA: COPEPODA: PHILICHTHYIDAE) Thomas Byrnes and Roger Cressey Abstract.—Colobomatus mylionus Fukui has been recollected from Austra- lian Acanthopagrus. The original description was based on a single female from a Japanese Acanthopagrus macrocephalus. The species is unique because of the presence of rings of fine processes on the surface of the lateral processes of the female. The male is described for the first time. In 1965 Fukui described a new species of Colobomatus from the sparid fish Acantho- pagrus macrocephalus from Japan. This de- scription was based on a single damaged female. Recently, the first author, as part of a survey of the parasitic copepods of Aus- tralian bream, collected several specimens of the same species from three species of Acanthopagrus. This paper is based on that additional material including the first de- scription of the male. Colobomatus mylionus Fukui, 1965 Figs. 1-15 Material examined.—51 females and 2 males collected from the cephalic canal sys- tem adjacent to the nasal cavity of Acan- thopagrus australis (Gunther), A. berda (Forsskal), and A. /atus (Houttuyn) from off Eden, Newcastle, Brisbane, Yeppoon (all east coast Australia), and Point Sampson (west coast Australia). Description.—Female: Body form as in Figs. 1 and 2. Length of body, including processes, 4.75 mm (3.37—5.71 mm). Great- est width 0.94 mm (0.61-1.1 mm) mea- sured near base of posterior thoracic pro- cess. Measurements based on 10 specimens from Point Sampson. Color in life yellowish with or without black gut contents visible in vivo. Cephalon bearing 2 dorsolateral process- es and shorter ventromedial process. First antenna (Fig. 3) indistinctly segmented, bearing naked, blunt setae as figured. Mouth ventral, with mandibles, first maxillae, and second maxillae within. Mandible armed with single blade. First maxilla with robust base bearing row of spinules and 3 spines at base of distal process (process with row of denticles along inner edge). Second max- illa with globose basal segment and short naked terminal process. Maxilliped absent (see Fig. 4). Single anteriorly directed pro- cess arising from center of posterior margin of mouth cone. Leg 1 (Fig. 5) 2-segmented with single naked seta on last segment. Legs 4—5 absent. Genital segment bearing 2 lateral, pos- teriorly directed processes. Small genital process bearing | short and 1 long setae (Fig. 6) on dorsolateral posterior margin of gen- ital segment near origin of lateral processes. Abdomen indistinctly 3-segmented. Each caudal process bearing small seta-like pro- cess (Fig. 7), possibly representing caudal ramus. Egg sacs typical of genus and loosely at- tached (often dropping off when specimens handled). Male: Body form as in Fig. 8. Total length 0.95 mm, greatest width 0.14 mm, based on single specimen. Cephalothorax indis- tinct. VOLUME 99, NUMBER 3 389 Figs. 1-7. Colobomatus mylionus Fukui, female: 1, Dorsal; 2, Ventral; 3, First antenna; 4, Oral area; 5, Leg 1; 6, Leg 2; 7, Leg 3. First and second antennae separated from oral area (Fig. 9). First antenna 5-segment- ed, armed with setae as in Fig. 10. Second antenna (Fig. 11) 4-segmented, last 2 seg- ments bearing spines and setae as in figure. Remaining oral appendages as in Fig. 12. Leg | (Fig. 13) coxopod with inner naked seta; basipod with outer naked seta; exopod first segment with outer stout spine; second segment with 4 outer stout spines and 3 inner setae (all spines barbed along both margins); endopod first segment with inner 390 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 8-15. Colobomatus mylionus Fukui, male: 8, Dorsal; 9, Cephalon and leg bearing thoracic segments, ventral: 10, First antenna; 11, Second antenna; 12, Oral area; 13, Leg 1; 14; Leg 2; 15, Leg 3. VOLUME 99, NUMBER 3 seta; second segment with 2 outer barbed spines and 4 inner setae (all setae naked). Leg 2 (Fig. 14) as in leg 1 except exopod with 1 less spine on last segment and en- dopod last segment with 2 less setae. Leg 3 (Fig. 15) represented by 3 naked setae (Fig. 2), Acknowledgments We wish to thank Mr. K. Izawa of Mie University for opinions relative to the taxo- nomic status of the new material from Aus- tralia and for suggesting the collaborative arrangement with the second author that 391 resulted in the above redescription of C. mylionus. Literature Cited Fukui, T. 1965. Onsome parasitic copepods of Japan [in Japanese].— Researches on Crustacea No. 2. The Carcinological Society of Japan. 60-65. (TB) 69 Louis Drive, Farmingdale, New York 11735; (RC) Department of Inverte- brate Zoology, National Museum of Natu- ral History, Smithsonian Institution, Wash- ington D.C. 20560. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 392-398 NEOTROPICAL MONOGENEA. 9. STATUS OF TRINIGYRUS HANEK, MOLNAR, AND FERNANDO, 1974 (DACTYLOGYRIDAE) WITH DESCRIPTIONS OF TWO NEW SPECIES FROM LORICARIID CATFISHES FROM THE BRAZILIAN AMAZON D. C. Kritsky, W. A. Boeger, and V. E. Thatcher Abstract. —The generic diagnosis of Trinigyrus Hanek, Molnar, and Fernan- do, 1974, is emended to incorporate the following characters: presence of con- fluent intestinal caeca, overlapping gonads (testis dorsal or dorsoposterior to ovary), a ventral anchor/bar complex, and a haptor with 4 pairs of appendages on which hook pairs 2, 3, 4, 6, and 7 are located. The genus is transferred from the Dactylogyrinae to the Ancyrocephalinae based on the presence of the ventral anchor/bar complex, the absence of hook pair 4A, and its relationship to Hamatopeduncularia Yamaguti, 1953. Two new species of Trinigyrus are de- scribed from the gills of Loricariidae: T. acuminatus from Acanthicus hystrix Spix, and 7. tentaculoides from Hypoptopoma thoracathum Gunther. Hanek, Molnar, and Fernando (1974) proposed TJrinigyrus for their new species, T. hypostomatis, infesting the gills of Hy- postomus robinii (Valenciennes), Loricari- idae, in Trinidad. The genus was originally placed in the Dactylogyrinae of the Dactyl- ogyridae presumably because of the pres- ence of a single anchor/bar complex in the haptor. Gussev (1978) postulated that members of this subfamily would be rare or absent in the native Neotropical fauna since their primary hosts (cyprinid fishes) do not occur naturally in the region. Gussev (1978), who felt the types of 7. hypostomatis were probably damaged specimens in which the second anchor/bar complex had been torn away, suggested that Trinigyrus belongs to the Ancylodiscoidinae Gussev, 1961, based on the morphology of the anchors and bar and on the reported host. However, he did not formally make the transfer. In the pres- ent study, Trinigyrus is emended and reas- signed at the subfamily level, and two new species of the genus are described. Hosts were collected from the environs of Manaus, Amazonas, Brazil, during Sep- tember and November 1984. Methods of host and parasite collection, preparation of helminths for study, measurement, and numbering of haptoral hook pairs are those used by Kritsky, Thatcher, and Boeger (1986). Measurements are in micrometers; averages are followed by ranges in paren- theses. Cirrus length was approximated us- ing a Minerva curvimeter on camera lucida drawings. Type specimens are deposited in the collections of the Instituto Nacional de Pesquisas da Amazonia (INPA), the U.S. National Museum Helminthological Col- lection (USNM), and the University of Ne- braska State Museum (HWM.L) as indicated below. Trinigyrus Hanek, Molnar, and Fernando, 1974 Emended diagnosis. —Dactylogyridae, Ancyrocephalinae. Body divisible into ce- phalic region, trunk and haptor (peduncle absent). Tegument thin, smooth. Cephalic lobes, head organs, cephalic glands present. Eyes absent. Mouth subterminal, midven- VOLUME 99, NUMBER 3 tral; pharynx muscular, glandular; esopha- gus short or absent; intestinal caeca 2, con- fluent posterior to gonads, lacking diverticula. Gonads intercaecal, overlap- ping; testis dorsal or dorsoposterior to ovary. Vas deferens looping left intestinal caecum; seminal vesicle a dilation of vas deferens; prostatic reservoirs 2; copulatory complex comprising accessory piece and tubular cir- rus. Oviduct short; uterus delicate or well developed; vagina dextral; seminal recep- tacle lying diagonally to right of midline. Genital pore midventral. Vitellaria consist- ing of 2 bilateral bands in trunk and anterior portion of haptor. Haptor with variable number of glandular reservoirs, ventral an- chor/bar complex, 7 pairs of similar hooks. Haptor exhibiting 4 pairs of hook-bearing appendages: 2 bilateral pairs bearing hooks pairs 2, 7; single posteroventral pair branched, bearing hook pairs 3, 4; pair of posterodorsal appendages bearing hook pair 6; hook pairs 1, 5 sessile. Parasites of gills of Loricariidae, Siluriformes. Type species, host, and locality. —Trini- gyrus hypostomatis Hanek, Molnar, and Fernando, 1974, from Hypostomus robinii (Valenciennes), Talparo River near Talpa- ro, Trinidad. Other species.—Trinigyrus acuminatus, n. sp. from Acanthicus hystrix Spix, Rio Ne- gro near Manaus, Amazonas, Brazil; T. ten- taculoides, n. sp. from Hypoptopoma thor- acathum Gunther, Rio Solim6es near Marchantaria Island, Manaus, Amazonas, Brazil. Remarks. —In adults of Trinigyrus species described below, we observed that the testis is seldom developed, although the ovary ex- hibits large oocytes and the seminal vesicle and receptacle are filled with spermatozoa. These observations suggest that some form of protandry exists in Trinigyrus, though we could not determine the developmental se- quence from available specimens. The oc- currence of spermatozoa in the seminal ves- icle of specimens apparently lacking a testis implies that this gonad develops and sub- 393 sequently regresses before the ovary ma- tures. Bychowsky (1957) reports that the testes form prior to the ovary in Microcotyle spinicirrus, but that both gonads apparently remain in the fully developed adult. Pro- tandry has also been reported in hexaboth- riids by Brinkmann (1952) and Mayes, Brooks, and Thorson (1981). Trinigyrus acuminatus, new species Figs. 1-7 Host and locality.—Acanthicus hystrix Spix, Loricariidae; Rio Negro near Manaus, Amazonas, Brazil, 1 Nov 1984. Type specimens.—Holotype, INPA PA 284-1; paratypes, INPA PA 284-2 + PA 284-4, USNM 79139, HWML 23306. Description (based on 32 specimens).— Body subconical, 320 (190-407) long; great- est trunk width immediately anterior to haptor. Cephalic region elongate, with 2 ter- minal lobes and subterminal dilation; head organs poorly developed, present in ce- phalic lobes and expanded cephalic area; cephalic glands indistinct, lying postero- lateral to pharynx. Pharynx spherical, 26 (16-33) in diameter. Haptor 164 (110-196) wide, 93 (63-129) long, an expanded por- tion of body; haptoral appendages short; glandular reservoirs variable, conspicuous. Anchor 41 (33-45) long, with short shaft, elongate point, sharply recurved tip, base lacking roots; base 9 (7-11) wide; anchor filament double. Bar 66 (60-79) long, with ventral longitudinal groove, tapered ends. Hooks similar, delicate; each 13 (11-16) long, with erect thumb, enlarged proximal shank; FH loop '2 shank length. Cirrus 98 (94-101) long, an elongate slender tube with sinistral loop, base reduced; accessory piece 24 (19-36) long, comprising a proximal rod, distal dumbbell-shaped appendage. Gonads subovate. Testis 25 (21-30) wide, 46 (39- 53) long; seminal vesicle elongate, fusiform; prostatic reservoirs ovate, with thick walls. Ovary 52-53 wide, 26-27 long; oviduct, ootype, uterus not observed; numerous con- spicuous glands surrounding presumed 394 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-7. Trinigyrus acuminatus: 1, Composite drawing of whole mount (ventral); 2, 3, Copulatory com- plexes; 4, Hook; 5, Vagina and distal portion of seminal receptacle; 6, Bar; 7, Anchor. All figures are reproduced to the same scale (30 micrometers) except Fig. 1 (100 micrometers). VOLUME 99, NUMBER 3 ootype; vagina with exterior flower-like ap- pendage, short internal sclerotized tube di- rected posteriorly; seminal receptable ovate, ventral in trunk, extending posteriorly to midline. Vitellaria coextensive with intes- tinal caeca, vitelline commissure indistinct. Remarks. —Trinigyrus acuminatus dif- fers notably from 7. hypostomatis Hanek, Molnar, and Fernando, 1974, and 7. ten- taculoides by possessing an elongate cirrus and a complex accessory piece. In 7. hy- postomatis and T. tentaculoides, the cirrus comprises a short curved tube associated with a simple rod-shaped accessory piece. Based on the morphology of the haptoral armament, 7. acuminatus is most closely related to 7. hypostomatis in that the an- chor/bar complexes are similar. Etymology.—The specific name is from Latin (acumin/o = pointed) and refers to the elongate anchor point with recurved tip. Trinigyrus tentaculoides, new species Figs. 8-14 Host and locality.—Hypoptopoma thor- acathum Gunther, Loricariidae; Rio Soli- moes near Marchantaria Island, Manaus, Amazonas, Brazil, 13 and 26 Sep 1984. Type specimens.—Holotype, INPA PA 285-1; paratypes, INPA PA 285-2, PA 285-3, USNM 79138, HWML 23305. Description (based on 19 specimens).— Body robust, stout, 222 (165-307) long; greatest trunk width immediately anterior to haptor. Cephalic margin rounded or with 2 terminal, 2 bilateral cephalic lobes; head organs well developed in lobes and adjacent cephalic area; cephalic glands comprising small spherical cells posterolateral to phar- ynx. Pharynx subspherical, 21 (16-30) in diameter. Haptor 110 (81-145) wide, 59 (42-73) long; posteroventral appendages conspicuously bifurcated; glandular reser- voirs poorly developed. Anchor 47 (43-50) long, easily distorted by coverslip pressure (Fig. 14), roots absent, base tear-drop shaped, shaft and point a continuous smooth 395 curve, tip of point recurved; base 10 (8-11) wide; anchor filament delicate. Bar 75 (60— 93) long, with tapered ends, ventral longi- tudinal groove, flat posteromedial projec- tion. Hooks 10 (7-12) long, delicate; each with erect thumb, proximal dilation of shank; FH loop '2 shank length. Cirrus 29- 30 long, a curved shaft arising from simple base; accessory piece 28 (22-33) long, rod- shaped, recurved distally. Gonads ovate. Testis 33 (30-36) wide, 41 (40-43) long; seminal vesicle fusiform, lying medially an- terior to gonads; prostatic reservoirs subov- ate, with thick wall. Ovary 35 (27—45) wide, 49 (36-57) long; oviduct, ootype not ob- served; well-developed glands surrounding presumed ootype; uterus with thick mus- cular wall distally; vagina comprising a small funnel and irregularly sclerotized tube reaching to ovate seminal receptable; vitel- laria coextensive with intestinal caeca, com- missure indistinct. Egg distorted, with ex- ceptionally elongate proximal filament. Remarks. —Based on comparative mor- phology of the haptoral sclerites and cop- ulatory complex, 7. tentaculoides is most similar to the type species, 7. hypostomatis Hanek, Molnar, and Fernando, 1974. These species are easily differentiated by 7. ten- taculoides possessing conspicuous haptoral appendages, which are short in 7. hyposto- matis. Trinigyrus tentaculoides also shares fea- tures of the haptoral armament with several species of the marine genus Hamatopedun- cularia Yamaguti, 1953. A flat postero- medial projection of the bar similar to that of 7. tentaculoides occurs in H. arii Ya- maguti, 1953 of Bychowsky and Nagibina (1969), and H. thalassini Bychowsky and Nagibina, 1969. Glandular reservoirs in the haptor have been described in most Ham- atopeduncularia species. These features and the fact that all species of Hamatopedun- cularia and Trinigyrus occur on siluriform fishes suggest a close relationship of the two genera (see discussion). 396 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON or 12 13 14 Figs. 8-14. Trinigyrus tentaculoides: 8, Ventral view of whole mount (composite); 9, Vagina; 10, Copulatory complex; 11, Bar; 12, Hook; 13, Anchor; 14, Typically distorted anchor as a result of coverslip pressure. All figures are drawn to the 30-micrometer scale except Fig. 8 (100 micrometers). VOLUME 99, NUMBER 3 Etymology.—The specific name is from Neolatin (tentacul/o = tentacle + oides = like) and refers to the well-developed hap- toral appendages. Discussion Trinigyrus was briefly diagnosed by Ha- nek, Molnar, and Fernando (1974) based on observations of unstained specimens of the type species, 7. hypostomatis. Our discov- ery of two new species of the genus from Loricariidae in the Brazilian Amazon, some specimens of which were prepared for study of the internal anatomy, provided new in- formation which necessitated an emended generic diagnosis. Findings that supplement the original diagnosis include: (1) the pres- ence of a bifurcated gut with intestinal caeca confluent posterior to the gonads (not ob- served by Hanek et al. 1974); (2) overlap- ping gonads with the testis lying dorsal or dorsoposterior to the ovary (described as tandem by the original authors); (3) an an- chor/bar complex that is ventral in the hap- tor with the bar lying on the superficial (ven- tral) surface of the anchor bases (anchors and bar are said to be dorsal by Hanek et al. 1974) and (4) a haptor with short to well- developed appendages on which hook pairs 2, 3, 4, 6 and 7 are located (haptoral ap- pendages were not reported in the type species). We examined a paratype of T. hy- postomatis (USNM 73183) and have veri- fied the presence of short haptoral append- ages and a ventral anchor/bar complex in this species. The cleared, unstained and highly contracted paratype precluded de- termination of the position of the gonads and the pattern of the haptoral appendages. Although reduced hooks (pair 4A of Miz- elle and Price 1963) are absent, Hanek, Mol- nar, and Fernando (1974) considered Tri- nigyrus a member of the Dactylogyrinae apparently because of the presence of a sin- gle anchor/bar complex. Our finding that the complex is ventral in the haptor, along with the fact that 4A hooks are absent, sug- 397 gests that the genus belongs in the Ancy- rocephalinae as a group expressing derived characters which include the loss of the dor- sal anchor/bar complex and eyes, and the development of haptoral appendages. Thus, Trinigyrus is the only genus of Ancyroce- phalinae characterized by species possess- ing a single anchor/bar complex. The transfer of Trinigyrus from the Dac- tylogyrinae to the Ancyrocephalinae is fur- ther supported by the apparent relationship of Trinigyrus species with those of the an- cyrocephaline genus Hamatopeduncularia Yamaguti, 1953. As far as we are aware, the characteristic flat posteromedial projection of the bar in Trinigyrus tentaculoides is also found only in species of Hamatopeduncu- laria (H. arii of Bychowsky and Nagibina 1969, and H. thalassini Bychowsky and Na- gibina, 1969). Although not unique to the genus, nearly all Hamatopeduncularia species have been described with conspic- uous glandular reservoirs (usually four) in the haptor. These structures seem similar to those we report from 7. acuminatus and T. tentaculoides, in the former of which they are well developed. The relationship is fur- ther substantiated by the fact that species of both genera possess haptoral appendages and parasitize fishes of the Order Siluri- formes. ‘“‘Fahrenholz Rule” states that the natural classification of some parasite groups usu- ally corresponds directly with the natural relationships of their hosts (Eichler 1948). The Ariidae include primarily marine cat- fishes believed derived from freshwater si- luriform ancestors (Darlington 1957); and the Hamatopeduncularia are known only from marine ariid hosts. Our conclusion that the species of 7rinigyrus and Hamatope- duncularia are related suggests that the Ha- matopeduncularia and species of related marine genera (Chauhanellus Bychowsky and Nagibina, 1969; and Hargitrema Tri- pathi, 1959) represent derived taxa whose common ancestors secondarily invaded the 398 marine environment. This invasion likely occurred concomitantly with the same event for their hosts. The Hamatopeduncularia apparently retain more of the primitive characters of their ancestor than do the 777- nigyrus species. For example, the loss of the dorsal anchor/bar complex and eyes and the development of confluent intestinal caeca in Trinigyrus likely occurred after isolation of the two ancestral populations. Diver- gence in the marine population appears less dramatic, although it has also progressed to the generic level as expressed in the related genera Hamatopeduncularia, Chauhanel- lus, and Hargitrema. Acknowledgments The authors are grateful to Dr. J. Ralph Lichtenfels, USNM, for providing the para- type of Trinigyrus hypostomatis; and Lucia Py-Daniel, INPA, for identifying the fish hosts. The Max Planck Institute, Plon, Ger- many, kindly provided financial and tech- nical support for the collection of fish hosts, and the Conselho Nacional de Desenvol- vimento Cientifico e Tecnologico provided a study grant (20.0115/84) to WAB. Literature Cited Brinkmann, A., Jr. 1952. Fish trematodes from Nor- wegian waters.— Universitetet 1 Bergen Arbok, Naturvitenskapelig Rekke, 134 pp. Bychowsky, B. E. 1957. Monogenetic trematodes, their systematics and phylogeny.— Akad. Nauk SSSR, 1-509. [Translation from Russian, AIBS, Washington, D.C.] , and L. F. Nagibina. 1969. Ancyrocephalinae (Dactylogyridae, Monogenoidea) from fishes of the family Ariidae.— Parazitologiia 3:357-368. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Darlington, P. J. 1957. Zoogeography: the geograph- ical distribution of animals. John Wiley & Sons, Inc., 675 pp. Eichler, W. 1948. Some rules in ectoparasites. —An- nals and Magazine of Natural History (12) 1: 588-598. Gussev, A. V. 1978. Monogenoidea of freshwater fishes. Principles of systematics, analysis of world fauna and its evolution.—Parazitologicheskii Sbornik 28:96-199. Hanek, G., K. Molnar, and C. H. Fernando. 1974. Three new genera of Dactylogyridae (Monoge- nea) from freshwater fishes of Trinidad. —Jour- nal of Parasitology 60:91 1-913. Kritsky, D. C., V. E. Thatcher, and W. A. Boeger. 1986. Neotropical Monogenea. 8. Revision of Urocleidoides (Dactylogyridae, Ancyrocephali- nae). — Proceedings of the Helminthological So- ciety of Washington 53:1-—37. Mayes, M. A., D. R. Brooks, and T. B. Thorson. 1981. Potamotrygonocotyle tsalickisi, new genus and species (Monogenea: Monocotylidae) and Par- aheteronchocotyle amazonensis, new genus and species (Monogenea, Hexabothriidae) from Po- tamotrygon circularis Garman (Chondrich- thyes: Potamotrygonidae) in northwestern Bra- zil.—Proceedings of the Biological Society of Washington 94:1205-1210. Mizelle, J. D., and C. E. Price. 1963. Additional hap- toral hooks in the genus Dactylogyrus. —Journal of Parasitology 49:1028-1029. (DCK) Department of Allied Health Professions and Idaho Museum of Natural History, Idaho State University, Box 8002, Pocatello, Idaho 83209; (WAB) Depart- ment of Biological Sciences, Idaho State University, Box 8007, Pocatello, Idaho, and Instituto Nacional de Pesquisas da Ama- zonia, Manaus, Brazil; (VET) Instituto Na- cional de Pesquisas da Amazonia, Manaus, Brazil. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 399-405 REDESCRIPTION OF ECHINODERES PILOSUS (KINORHY NCHA: CYCLORHAGIDA) Robert P. Higgins Abstract.—Echinoderes pilosus Lang is redescribed from a paratypic male and female collected from kelp washed onto a beach at South Georgia Island in the southwestern Atlantic. The presence of lateroventral adhesive tubes on the fourth segment, patterns of cuticular hairs, scars, and sensory spots, and the morphological differences between the sexes are noted. The species is com- pared with other species having the same spine formula. Although the phylum Kinorhyncha gen- erally is associated with marine benthic sed- iment, several species have been found in association with either other invertebrates or plants (mostly algae). Echinoderes du- jardinii Claparéde, 1863 was discovered in washings of estuarine algae and oysters by the French zoologist Felix Dujardin (1851). In his “Monographie der Echinodera,”’ Ze- linka (1928) again reported E. dujardinii and added E. ferrugineus Zelinka, 1928, several additional juvenile cyclorhagids, and a homolorhagid, Pycnophyes rugosus Zelin- ka, 1928 to the list of algal inhabitants. Karl Lang (1936) found P. kielensis Ze- linka, 1928 and P. flaveolatus Zelinka, 1928 in “red algae associated with stone and mud’; 13 years later, he described E. pi- losus Lang, 1949 “‘from washings of old kelp cast up on shore. S. Georgia. Grytviken.” Pallares (1966), apparently studying the plankton associates with red algae along the southeastern coast of Argentina, also found Lang’s species. The only other published record of kinorhynchs in association with algae is that of Moore (1973) who found Campyloderes macquariae in samples of kelp holdfast fauna from several localities in the British Isles. The precise relationship between these kinorhynchs and their algal habitat remains a matter of conjecture. Since all of the kino- rhynchs reported from algae also are known from sediments, it is most probable that their presence, in most instances, has been a matter of their being washed into inter- tidal algae or transferred to the algae as it is washed along the sediment. At the time of its description, E. pilosus needed only to be compared with three oth- er species having the same or at least similar spine formula: E. dujardinii, E. ferrugineus, and E. worthingi Southern, 1914. Since its description, ten additional species with this same spine formula have been described. In the genus Echinoderes, the presence or ab- sence of one spine in particular, the lateral spine or adhesive tube on the fourth seg- ment (L-4), is often difficult to determine. In some species such as one most recently described, E. nybakkeni Higgins, 1986, it is a robust spine equal to the other lateral spines in its cuticularization as well as di- mensions. In others, such as E. brevicau- datus Higgins, 1966, it may be smaller and less cuticularized. In E. bookhouti Higgins, 1961 and a few others, only a round cutic- ular scar, possibly a pore, may replace the L-4 spine or adhesive tube. Some species appear to have neither the L-4 spine or any vestige of it. In some cases such as E. fer- rugineus, the presence of this spine is men- tioned in the description (Zelinka 1928) but not shown in the illustration. In other cases such as E. pilosus, the L-4 spine was neither mentioned in the description nor indicated 400 in the illustration (Lang 1949). The appar- ent absence of the L-4 spine and the exact nature of several other taxonomic charac- ters prompted the re-examination of this species which is the subject of this report. Suborder Cyclorhagae Zelinka, 1896 Family Echinoderidae Butschli, 1876 Echinoderes Claparéde, 1863 Echinoderes pilosus Lang, 1949 Figs. 1-10 Echinoderes pilosus Lang, 1949:17, fig. 4a, b (type locality: Grytviken, South Geor- gia Island).—Karling, 1954:189.—Hig- gins, 1960:88; 1964:491; 1966:518.— Pallares, 1966:103, pl. 1: figs. A—D; pl. 2 (Puerto Deseado to Sorrel, Argentina). — Kozloff, 1972:121.—Schmidt, 1974: 189.—Higgins, 1977a:12; 1977b:353; 1983:9; 1986:267. Diagnosis. —Echinoderes with middorsal spines on segments 6-10, lateral spines on segments 4, 7-12; middorsal spines 36-74 um long, becoming longer in posterior pro- gression; lateral spines 36—48 um long; lat- eral terminal spines 175-184 um long, 43- 46% of trunk length, with serrulate lateral margins; trunk length 400-408 um (400- 460 um, Lang’s data); lateral terminal ac- cessory spine (of female) 55 um long, 29% of lateral terminal spine; pectinate fringe very fine; perforation sites abundant, pat- tern distinctive but not well defined, cutic- ular hairs appear more abundant than per- foration sites suggest; trichoscalid plates large ventrally, very small dorsally. Redescription. —Paratypic adult male (Figs. 1, 2, 5, 6, 8-10) trunk length 408 um; MSW-8 (maximum sternal width, at seg- ment 8), 84 wm, 20.5% of trunk length; SW (standard width, =sternal width at segment 12), 78 wm, 19.1% of trunk length; cuticle with many hairs, perforation sites not uni- formly distinct; posterior margins of trunk segments finely striated with very fine pec- tinate fringe. Segment 2 with 16 placids, about 24 um PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON long, widest posteriorly, narrowest ante- riorly; midventral placid wider (22 um) than others (14 um); trichoscalid plates of head region (segment 1) overlapping alternate placids, ventral trichoscalid plates larger (12 wm X 15 yum) than dorsal trichoscalid plates (8 wm X 5 wm), modified midventral scalid prominent, 12 um long, pointed at posterior end, overlapping midventral placid. Segment 3, 44 um long (at lateral optical view); sternal width (measured along ante- rior pachycycli) 76 wm; two middorsal cu- ticular scars at midline, subdorsal cuticular scar on either side; single cuticular scar near anterolateral margins on ventral surface. Segment 4, 42 um long; pachycyclus moderately developed, no evidence of cu- ticular thickening along midventral line; middorsal and more lateral (than on seg- ment 3) subdorsal cuticular scars; small an- gular muscle scar lateral to each dorsolateral cuticular scar; single cuticular scar on either side of ventral midline, near anterior mar- gin; prominent lateral spines (adhesive tubes), 38 um long, midway in segment, in line with ventrolateral articulation zones of remaining segments. Segment 5, 44 wm long; sternal width 80 um; pachycyclus well developed, midven- tral articulation of sternal plates clearly vis- ible, well cuticularized; middorsal cuticular scar only, small angular muscle scar more lateral, in line with previous scar; single cu- ticular scar on either side of ventral midline, near anterior margin, additional cuticular scar near each lateral margin of ventral plates. Segment 6, 48 um long; sternal width 82 uum; pachycyclus similar to preceding one; middorsal spine, 42 wm long, sensory spots laterally adjacent and slightly posterior to middorsal spine; dorsolateral cuticular scars and small angular muscle scar similar to those of segment 4; cuticular scar near mid- dle of ventral plate, in line with those of previous segments. Segment 7, 48 um long; sternal width 83 uum; pachycyclus similar to preceding one; VOLUME 99, NUMBER 3 401 hie HATH PETG a } tC oe he aR aa io i ina x pee jiuitt | mi es a cana ; 100m F Nitsieainsepe co sacuensee i fe Nien D eee se cre nereeegen Aah Te WW HH t ii BEL Oa Ben aoe I o% ou a? mals . - eee @ Ar ee hi TULLE nami f it Huth Tuy PUTA sith +f ae in yh GAATTATNLAR IS fiat eee re Ss I See - Serer ese " TD) FTONTLLAMUAVCU HALA O.°. aman } hn aera hat HI city mya ied rH ri Hiei Nh Ha A il i Ny tr ees HAT TEA VFRraeSRR H en 23 Din i Neate ili Paratenict aoe ail ny oe nt FINA Uy Seneass LO AN, ar i Ve Figs. 1-4. Echinoderes pilosus: 1, Adult male, neck and trunk segments, ventral view; 2, Same, dorsal view; 3, Adult female, segments 12, 13, ventral view; 4, Same, dorsal view. 402 middorsal spine, 42 um long, with adjacent sensory spots, cuticular scars and muscle scars as in segment 6; lateral spine 36 wm long (probably adhesive tube) on tergal plate adjacent to junction with each ventral plate; cuticular scar near middle of ventral plate as in previous segment, slight evidence of small angular muscle scar slightly more lat- eral, near origin of arthrocorium where marginal striation begins. Segment 8, 50 um long; maximum sternal width (MSW) 84 um; middorsal spine, 46 um long; lateral spine 36 um long; otherwise similar to segment 7. Segment 9, 50 um long; sternal width 84 um; middorsal spine 50 um long; lateral spine 40 um long; otherwise similar to seg- ment 8. Segment 10, 60 um long; sternal width 83 um; middorsal spine 74 um long; lateral spine 46 um long; no evidence of small, angular muscle scar on each ventral plate; otherwise similar to segment 9. Segment 11, 60 um long; sternal width 80 um; without middorsal spine; lateral spine 42 um long; subdorsal cuticular scars closer to midline, small, angular muscle scars less distinct, otherwise dorsal and ventral mor- phology similar to segment 10. Segment 12, 61 wm long; SW-12 78 um; lateral spine 40 um long, more flexible in appearance than preceding lateral spines, more dorsally elevated on tergal plate; 2 middorsal cuticular scars, no subdorsal cu- ticular scars or evidence of small, angular muscle scars, perforation sites weak, sparse- ly distributed near posterior half of segment both dorsally and ventrally; ventral cutic- ular scars similar to those on preceding seg- ments. Segment 13, 35 um long; sternal width 50 um; lateral terminal spines 175 um long; serrulate on lateral margins beginning about one-fourth the distance from base; ventral plates with slightly pointed margins, some- what conforming to margin of tergal plate, with hair-like projection; pachycycli widely divergent at anterior margin; tergal plate PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with single middorsal cuticular scar, plate bifurcated into two pointed tergal exten- sions; 3 pairs of penile spines (P-1, 45 wm long, dorsally displaced; P-2, 28 um long, with blunted tip, ventrolateral; P-3, 50 um long, adjacent to P-2). Mean length of middorsal spines (D 6- 10) 50.8 wm, 12.5% of trunk length; mean length of lateral spines (L 4, 7-12) 39.7 um, 9.7% of trunk length. Paratypic female (Figs. 3, 4, 7) TL about 400 um (broken specimen); MSW-8 86 um, 21.5% of trunk length; SW 80 um, 20.0% of trunk length; middorsal spine lengths: D-6 36 um, D-7 42 wm, D-8 44 um, D-9 52 um, D-10 74 wm; mean length of middorsal spines (D 6-10) 49.6 um, 12.2% of trunk length; lateral spine lengths: L-4 34 um, L-7 38 um, L-8 34 wm, L-9 38 um, L-10 48 wm, L-11 42 um, L-12 40 wm; mean length of lateral spines (L 4, 7-12) 39.1 wm, 9.7% of trunk length; lateral terminal spines 184 wm long, 46.0% of trunk length; lateral terminal accessory spines (female only) 54 um long, 13.5% of trunk length, 29.3% of lateral ter- minal spine length. With exception of minor morphological differences in the structure of the last two segments, the female (Figs. 3, 4, 7) closely resembles the male. Material examined.—1 paratypic male and | paratypic female (broken specimen), permanently mounted (by Higgins) in Cobb aluminum slide frames with modified Hoy- er’s medium (Higgins 1983), material from the Swedish South Pole Expedition (1901- 1903), 23 May 1902, Grytviken, South Georgia Island, “Sample 11.’ Repository: Swedish Natural History Museum (Section of Invertebrate Zoology), Stockholm. Remarks. —Forty-two species of Echi- noderes are based on adult specimens and identifiable on the basis of their description. Three others are so poorly described that I consider them species inquirenda; 27 others are based on juvenile states which are not likely to be reconciled with any adult. With- in this genus, the most common middorsal VOLUME 99, NUMBER 3 Figs. 5-10. Echinoderes pilosus: 5, Adult male, segments 12, 13, ventral view; 6, Same, dorsal view; 7, Adult female, ventral view; 8, Adult male, segments 10 (partial), 11, dorsal view; 9, Same, segments 2—4, ventral view; 10, Same, dorsal view. (Interference contrast photographs.) spine formula is D 6-10; it is shared by 23 species, 12 of which have lateral spines on segments 4, 7-12. Two of these have an additional lateral accessory spine (LA-10) dorsally adjacent to the L-10 spine and two others differ only by the absence of the L-4 spine or adhesive tube, a character that can be difficult to see. The current re-exami- nation of EF. pilosus, in large part, was to determine whether or not this spine was ab- 404 sent as indicated in the original description, or present as has now been established. Of the 12 species having the same spine formula, FE. dujardinii, from northern Eu- ropean coasts, and its sibling species, E. ger- ardi Higgins, 1978, from the Mediterra- nean, have a lateral accessory spine on segment 10 leaving only nine remaining species to compare with E. pilosus. The L-4 spine or adhesive tube is not present in either E. truncatus Higgins, 1983 or E. bookhouti, although the latter species has either a pore or a cuticular scar in the L-4 position. Several distinctive as well as unique char- acters in the recently described E. nybak- keni separate it from E. pilosus; the former species is very small, 185 um long, has a midventral cuticularization on segment 4, extremely prominent pectinate fringe, and very prominent spines, including a D-7 spine which is twice the length of those on adja- cent segments. Similarly, E. krishnaswamyi Higgins, 1985 is equally distinctive in hav- ing very long, flexible spines and a unique perforation site pattern of one or two trans- verse rows. This species is the only other with the same spine formula that has ser- rulate lateral margins on the lateral terminal spines as in E. pilosus. Echinoderes ferrugineus is not as well de- scribed as the remaining species closely re- lated to E. pilosus. Lang (1949) separated E. pilosus from E. ferrugineus on the basis of differences “‘in the size of the 3rd and the shape of the last zonite.” Echinoderes fer- rugineus females, at least, have a prominent seta adjacent to the L-12 spine, and the lat- eral terminal accessory spine is only 13.5— 16.4% of the lateral terminal spine length as contrasted with 29.3% in E. pilosus. Al- though the significance of total length is questionable because of annual variation (Higgins and Fleeger 1980), E. ferrugineus, from European coasts, is much smaller (TL 210-220 um) than E. pilosus (TL 400-408 um; or by Lang’s measurements, 400—460 um). In addition, Lang’s species appears to have relatively shorter lateral terminal PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON spines (42.9—46.0% of the trunk length) in contrast with the longer (up to 71% of the trunk length) in E. ferrugineus. Lang also compared his species with E. worthingi (redescribed by Higgins 1985). The latter northern European species also is smaller (242-265 um) than E. pilosus, has relatively longer lateral terminal spines (60.3—72.7% of the trunk length), distinctly longer terminal tergal extensions, and a D-10 spine twice the length of the D-9 spine. Distinctive brace-shaped muscle scars on certain ventral plates of E. koz/offi Higgins, 1977, E. ehlersi Zelinka, 1913, E. imper- foratus Higgins, 1983, E. pacificus Schmidt, 1974, and E. sublicarum Higgins, 1977, separate them from E. pilosus. Echinoderes pilosus has been reported only once since its original description. Pal- lares (1966) found numerous specimens of this species, including juvenile stages as small as 149 wm long, mostly in various collections of red algae made between 1949- 1965 along the southeastern coast of Ar- gentina. Pallares reported the presence of red eyespots in the animals but, like Lang, did not observe the L-4 spines. As in the case of Lang, Pallares’ measurements of body length are not defined. Using the stan- dard trunk length measurement (from the anterior margin of the first trunk segment to the posteriormost margin of the last seg- ment) and the measurement scales or other information provided by each of these au- thors I would suggest that their reported length measurements are excessive. The lengths and relative proportions of various body regions and spines that can be ob- tained through Pallares’ illustrations agree remarkably well with those I have derived from the two paratypes used in the rede- scription. Similar data derived from Lang’s illustrations tend to be much less compat- ible. In addition to confirming the presence of the L-4 spine in E. pilosus, Lang’s (1949) original description requires two additional emendations. His notation on the presence VOLUME 99, NUMBER 3 of ‘“‘a ring of extremely fine ‘hair’ quite close to the anterior border of the zonite,”’ refers to the very fine pectinate fringe along the posterior margin of each segment which overlaps the anterior part of the posteriorly adjacent segment. Lang also was in error in his interpretation of placid morphology. Correctly noting the presence of 16 placids comprising segment 2, he stated: “Neither of the two ventro-median placids is percep- tibly wider than the others.” There is, in E. pilosus, as in all known species of this genus, a single midventral placid which is wider than the laterally adjacent placids. In this case, the midventral placid of E. pilosus is nearly twice as wide as the others, mor- phology which also escaped notation by Pal- lares (1966). Acknowledgments I thank Dr. R. Olerod, Section of Inver- tebrate Zoology, Swedish Museum of Nat- ural History for his generous assistance in obtaining access to the type material in this study. Appreciation is also extended to Dr. Raymond B. Manning for his comments on this manuscript. Literature Cited Dujardin, F. 1851. Sur un petit animal marin, /’Ech- inodere, formant un type intermédiaire entre les Crustacés et les Vers.—Annales des Sciences Naturelles, Zoologie, series 3, 15:158-160. Higgins, R. P. 1960. A new species of Echinoderes (Kinorhyncha) from Puget Sound.—Transac- tions of the American Microscopical Society 79: 85-91. . 1964. Three new kinorhynchs from the North Carolina Coast.— Bulletin of Marine Science of the Gulf and Caribbean 14:479-493. 1966. Echinoderes arlis, a new kinorhynch from the Arctic Ocean. — Pacific Science 20:518- 520. 1977a. Redescription of Echinoderes dujar- dinii (Kinorhyncha) with description of closely related species. —Smithsonian Contributions to Zoology 248:1-26. 405 . 1977b. Two new species of Echinoderes from South Carolina. — Transactions of the American Microscopical Society 96:340-354. . 1983. The Atlantic barrier reef ecosystem at Carrie Bow Cay, Belize, II. Kinorhyncha.— Smithsonian Contributions to Marine Sciences 18:1-131. . 1985. The genus Echinoderes (Kinorhyncha: Cyclorhagida) from the English Channel.— Journal of the Marine Biological Society of the United Kingdom 65:785-800. . 1986. A new Echinoderes (Kinorhyncha: Cy- clorhagida) from a coarse sand California beach.— Transactions of the American Micro- scopical Society [in press]. , and J. W. Fleeger. 1980. Seasonal changes in the population structure of Echinoderes coulli (Kinorhyncha).— Estuarine and Coastal Marine Science 10:495-—505. Karling, T. G. 1954. Echinoderes levanderi n. sp. (Kinorhyncha) aus der Ostsee. — Arkiv for Zool- ogi, series 2, 7:189-192. Kozloff, E. N. 1972. Some aspects of development in Echinoderes (Kinorhyncha).—Transactions of the American Microscopical Society 91:119- 130. Lang, K. 1936. Undersékningar 6ver Oresund. XXI. Einige Kleintiere aus dem Oresund.— Kungliga Fysiografiska Saliskapets Handlingar, N. F. 46: 1-8. 1949. Echinoderida, pp. 1-22. In N. H. Odhner, ed. Further Results of the Swedish Ant- arctic Expedition, 1901-1903. Moore, P.G. 1973. Campyloderes macquariae John- ston, 1938 (Kinorhyncha: Cyclorhagida from the Northern Hemisphere.—Journal of Natural History 7:341-354. Pallares, R. 1966. Nota sobre Echinoderes pilosus Lang, 1949 (Aschelminthes, Kinorhyncha).— Physis 26:101—106. Schmidt, P. 1974. Interstitielle Fauna von Galapagos X. Kinorhyncha.—Microfauna des Meeresbo- dens 43:1-15. Zelinka, C. 1928. Monographie der Echinodera. Wil- helm Engelmann, Leipzig, iv + 396 pp. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C., 20560. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 406-416 SEPARATION OF HABER SPECIOSUS (HRABE) (OLIGOCHAETA: TUBIFICIDAE) FROM ITS CONGENERS, WITH A DESCRIPTION OF A NEW FORM FROM NORTH AMERICA Michael R. Milligan Abstract. —The diagnosis of Haber Holmquist is expanded to include species of previously questionable position. The members of this genus are unique among the Tubificinae in possessing similar penial and spermathecal setae and have characteristic genital morphology. Haber now includes eight species: H. speciosus (Hrabé), H. dojranensis (Hrabé), H. monfalconensis (Hrabé), H. amu- rensis (Sokolskaja and Hrabé), H. pyrenaicus (Juget and Giani), H. turquini (Juget and Lafont), H. hubsugulensis (Semernoi and Akinshina), and H. svi- renkoi (Lastockin). Haber simsi (Brinkhurst) is confirmed to be a synonym of H. speciosus. Based on setal morphology, H. speciosus is separated into four forms: speciosus, zavreli, simsi and fluminialis. This complex is compared to all congeners. Until Brinkhurst (1981) recorded the first appearance of Haber in the state of New York, this genus was only known from lo- calities in Europe. Subsequent material from Maryland, South Carolina, and Florida in- dicates that this genus, represented by a sin- gle species, is well established in extreme upper estuarine locations along the eastern seaboard and Gulf of Mexico in North America. Eight species have been incorpo- rated into the genus Haber, some based on incomplete descriptions. The species are: H. speciosus (Hrabé, 1931), H. svirenkoi (Las- tockin, 1939), H. dojranensis (Hrabé, 1958), H. monfalconensis (Hrabé, 1966), H. amu- rensis (Sokolskaja and Hrabé, 1969), H. pyrenaicus (Juget and Giani, 1974), H. tur- quini (Juget and Lafont, 1979), and H. hub- sugulensis (Semernoi and Akinshina, 1980). Holmquist (1978) established the genus (Haber) to accommodate those Tubificinae previously identified as either Peloscolex or Tubifex sharing the unique combination of similarly shaped penial and spermathecal setae, and an “apparent” penis sheath. Early descriptions of many of these species de- scribed a “‘cuticular penis sheath.” Holm- quist (1978) has shown, through the analysis of serial sections, the structure actually to be a thickened basement membrane, not of cuticular origin, surrounded by epithelial cells, which acts as an attachment for the musculature of the penial apparatus. Two forms of H. speciosus (zavreli and simsi) were once regarded as distinct species, but Hrabé (1981) regarded zavreli as a subspe- cies of speciosus; he placed both in the genus Tubifex. Brinkhurst and Jamieson (1971) had synonymized both plus the subspecies monfalconensis with H. speciosus. The most recent account by Brinkhurst and Wetzel (1984) suggests that some species in the ge- nus (including zavreli and simsi) would not survive a revision, but their status was left in doubt. Holmquist (1979) elevated mon- falconensis to specific rank, along with spe- ciosus and zavreli, but she indicated a pos- sible relationship between simsi and speciosus based on the single specimen of simsi available at that time. She listed other VOLUME 99, NUMBER 3 possible Haber species as dojranensis Hrabé, amurensis Hrabé and svirenkoi Finogeno- va. The North American forms are herein de- scribed and compared with their congeners. Materials and Methods Samples were collected from the Head Springs region of the Crystal River as part of a study of estuaries on the west coast of Florida sponsored by the Southwest Florida Water Management District. Samples were taken using a 12.5 x 12.5 x 20 cm diver operated core. A 15% solution of magne- sium sulfate was added to narcotize the an- imals. The samples were subsequently sieved through a 0.5 mm mesh screen, fixed in for- malin with Rose Bengal stain, then trans- ferred to 70% isopropyl alcohol. The pre- served worms were sorted from the residue and mounted on microscope slides in Am- man’s lactophenol. Additional specimens were stained in Grenacher’s alcohol borax carmine, cleared in terpineol, and either dis- sected or mounted whole in Canada balsam. Measurements referred to in the description were taken from fixed, mounted worms. Representative material has been deposited in the United States National Museum of Natural History, Washington, D.C. (USNM). Specimens of Haber from South Carolina collected by Dr. Dale Calder were supplied to me courtesy of Dr. R. O. Brinkhurst. These worms were stained with Rose Bengal and mounted whole in Canada balsam. Ma- terial from Maryland was received courtesy of Mr. Michael T. Barbour (EA Engineering, Science and Technology, Inc.). Specimens from New York identified as Haber cf. spe- ciosus by Dr. Brinkhurst were received courtesy of the USNM. The Maryland and New York material was mounted whole and cleared in CMC-10. Specimens of Haber from England were mounted in polyvinyl lactophenol and provided by Dr. Michael 407 Ladle (Freshwater Biological Association, England). Systematics Haber Holmquist, 1978 Definition (modified).—Limnetic tubifi- cids. Dorsal bundles include smooth or his- pid hair setae, and bifid or pectinate crotch- ets. Somatic ventral setae all bifid crotchets. Body wall usually smooth, without adherent foreign particles. Male and spermathecal pores paired. Coelomocytes absent or in- conspicuous. Modified spermathecal and penial setae present: thin and hollow-tipped, inserted in glandular sacs. Vas deferens long, bipartite in some species, entering ental por- tion of atrium apically. Glandular prostate present, attached to atrium medially in most species. Ectal region of atrium often mod- ified into narrow ejaculatory duct. Penial pouch present. Cuticular penis sheath ab- sent. Thick basal membrane resembling a penis sheath lining internal canal of penis. Male pore and penial setal sac with common opening. Spermathecae bipartite: ectal nar- row canal; entally, elongate ampullae. Sper- matozeugmata vermiform. Type species.—Tubifex speciosus Hrabé, 1931. Remarks. —The presence of modified penial setae has been reported in only two tubificine species, Tubifex nerthus Michael- sen, and 7. thompsoni Southern, other than those now considered in the genus Haber. Brinkhurst and Baker (1979:1554) deter- mined the penial setae reported in a single specimen of 7. nerthus from Europe prob- ably to be “no more than broken somatic setae.”” Tubifex thompsoni, synonymized with 7. costatus by Brinkhurst (1963), has been reported as having unmodified penial setae according to Holmquist (1979). How- ever, the morphology of the male efferent duct and the presence of palmate setae pre- cludes any relationship to the Haber group, and the penial setae may simply be bifids 408 retained. Tubifex costatus was described as lacking penial setae (Brinkhurst and Baker 1979). General Remarks. —Eight species are in- cluded in this genus as suggested by Holmquist (1978) and Brinkhurst (1981, 1984) based on the presence of penial and spermathecal setae inserted in a glandular sac. These species are separated primarily by the morphology of the male genitalia. Secondary characters are setal morphology, position of the spermathecal pore, and modifications of the body wall. Distribution. —North America, Europe. Haber speciosus (Hrabé, 1931) Tubifex (Tubifex) speciosus Hrabé, 1931: 24-27; 1964:108. Peloscolex zavreli Hrabé, 1942:23-26.— Brinkhurst, 1963:41, (equals Peloscolex speciosus (Hrabé), Brinkhurst, 1971:514). Peloscolex speciosus (Hrabé), Brinkhurst, 1962:304-305; 1963:43; 1966:735 (par- tim).—Brinkhurst and Jamieson, 1971: 514-515. Peloscolex simsi Brinkhurst, 1966:735—-736 (equals Peloscolex speciosus (Hrabé), Brinkhurst, 1971:514). Tubifex speciosus speciosus (Hrabé), Hrabé, 1966:68-70. ?Peloscolex zavreli Hrabé, Kasprzak, 1973: 421-422. Haber speciosus (Hrabé), Holmquist, 1978: 188. 191, 193) 195. 196, 200,201. 203: 204, 206; 1979:51, 52, 58.—Brinkhurst, 1984:52. Haber simsi (Brinkhurst), Bird and Ladle, 1981:493-498. Haber cf. speciosus (Hrabé), Brinkhurst, 1981:1062-1064. Tubifex speciosus zavreli Hrabé, Hrabé, 1981:87. Type material. —1593-10-P-II Hrabé Oligochaeta collection, from Ochrida Lake, Yugoslavia. Material examined. —NEW YORK: One PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON wholemounted specimen from Susquehan- na River, 4.6 kilometers upstream of rail- road bridge, above Gaudey Generating Sta- tion, Binghamton, New York, shallow riffle, large cobbles, USNM 065223; collector, Kurt Stimpson, 2 Aug 1976. MARYLAND: 7 wholemounted specimens from Piscata- way Creek, tributary of Potomac River south of Washington, D.C., 1 m, tidal freshwater marsh, sandy silt; collector, Michael T. Bar- bour, May-June 1983. SOUTH CAROLI- NA: 9 wholemounted specimens; 7 from upper estuarine region of Black River, 5 m, sand, 5 Apr 1977, 2 from upper estuarine region of Pee Dee River, 3 m, sand, collec- tor, Dr. Dale Calder; 5 Jan 1977. FLORI- DA: 30 wholemounted specimens, 3 dis- sected specimens, Head Springs region of Crystal River off Banana Island, Crystal River, 1 m, medium clean sand; collector, Michael R. Milligan, Feb—Nov 1984. EN- GLAND: 2 wholemounted specimens from Bere Stream, tributary of River Piddle, Dor- set, stream bed with corase flint gravel and sand; collector, Michael Ladle, 27 Oct 1980. Diagnosis (combined from literature and current studies).—Length (fixed, whole- mounted, complete specimens) 10-15 mm, width at clitellum 0.15—0.4 mm (Table 1). Body wall generally naked. Clitellum cov- ering X—XII. Preclitellar dorsal bundles with 1—3(4) smooth or hispid setae 150-290 um, and 1—4(5) pectinate setae 25—65 wm with maximally 8 intermediate teeth, nodulus distal. Postclitellar dorsal bundles with 1(2) hair setae 95-442 um, and 1(2,3) bifid or pectinate setae 35—52 um (Table 2). Pecti- nate setae with maximally 3 intermediate teeth. All somatic ventral setae bifid. Pre- clitellar ventral setae with upper tooth as long as or longer, and thinner than, lower tooth, 3-8 per bundle 35-65 um (Table 2). Postclitellar ventral setae generally with up- per tooth as long as, and thinner than, lower tooth, 1-3 per bundle 43-62 um (Table 2). Spermathecal and penial setae thin, hollow- tipped, embedded in glandular sac, gener- VOLUME 99, NUMBER 3 Dorsal setae Ventral setae ett i anterlor posterlor anterlor posterior Teen a ae S ya A =e vdw ley ee pr om S\ 1 oe Wh Cae )y @ IN J Z Fig. 1. 409 spermathecal seta y= / = ee a aa — ~ 4 Ys i} ; (Ales Male efferent duct and characteristic setae of Haber speciosus, simsi form, from Florida. Abbreviations: a, atrium; bm, basement membrane; gs, glandular sac; pr, prostate gland; ps, penial setae; sf, sperm funnel; vdn, vas deferens, narrow part; vdw, vas deferens, wide part. ally of equal length 42-70 um (Fig. 1). Sper- mathecal pore dorsal to ventral seta. Male funnel small. Vas deferens, long, bipartite. Thinner ental half 16-25 wm wide, ciliated, distinctly separated from wider ectal half 38-45 wm wide. Ectal region partly or wholely ciliated, enters atrium of similar width. Glandular prostate attached medi- ally to relatively elongate, spindle-shaped atrium (Figs. 1—2). Ejaculatory duct present, terminating in bulb-like penial apparatus. Basal membrane lining the penial canal forming a tube longer than wide. Canal of spermatheca c. 20 um wide and c. 100 um long; ampulla c. 100 wm wide, length vari- able. Remarks.—This species can be divided into four distinct “‘forms” based on setal morphology: speciosus, zavreli, simsi, and fluminialis form. Differences between the male efferent ducts of the different forms are insufficient to warrant separation as dis- tinct species. The differences are summa- rized in Tables 1 and 2. The term “‘subspe- cies” usually refers to a geographic race differing slightly from another group of the same species (Steen 1971). Form is more appropriately applied to the variants of H. speciosus since the variations observed may be attributable to changes in the environ- ment, primarily conductivity (Brinkhurst, pers. comm.), not necessarily due to geo- graphic isolation.! Two of these forms have only been col- lected from Europe: the speciosus form from 1 According to Article 45g(1) of the 1985 Interna- tional Code of Zoological Nomenclature adopted by the XX General Assembly of the International Union of Biological Sciences, a new name proposed as a form after 1960 is infrasubspecific and not officially recog- nized. 410 Dorsal setae PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ventral setae hee eR anterior posterior anterlor posterior Fig. 2. Male efferent duct and characteristic setae of Haber speciosus, fluminialis form, from South Carolina. Abbreviations as in Fig. 1. Ochrida Lake, Yugoslavia, and the zavreli form from Poland and Slavakia. In the orig- inal description of Haber speciosus (Hrabé, 1931) Hrabé referred to glandular hypo- dermal swellings on the body wall. This character was absent from the types Holmquist (1979) examined. However Holmquist (1979) does report the body of H. speciosus and H. zavrelito be finely ringed post-clitellarly. Hrabé (1981) regarded the latter to be a subspecies of H. speciosus. The simsi form has been found on both sides of the Atlantic: two streams in Dorset, England, and the Head Springs region in Crystal River, Florida. Brinkhurst (1966) described simsi as a separate species of Pe- loscolex, synonymized it with speciosus in 1971, and subsequently transferred it to the genus Haber (Brinkhurst 1981) as one of a string of possible species. Bird and Ladle (1981) redescribed H. simsi. None of the previous references described the male ef- ferent ducts. Bird and Ladle (1981) cleared their material in polyvinyl lactophenol. Consequently, a description of the internal male genitalia is not possible. However, from examination of their specimens, all observable characters agree with the speci- mens from Florida, which have been deter- mined to be a separate form of H. speciosus (Fig. 1). A few specimens from Florida and England have been found with hispid hair setae. The fluminialis form of Haber speciosus was previously described as Haber cf. spe- ciosus (Brinkhurst 1981). Examination of subsequent material from Maryland and South Carolina has determined it to be another form of H. speciosus (Fig. 2) unique to the east coast of the United States. The term fluminialis refers to the streams from which it has been collected. VOLUME 99, NUMBER 3 Table 1.—Body size and setal length of the forms of Haber speciosus. simsi simsi fluminialis fluminialis fluminialis speciosus zavreli (England) (Florida) (New York) (Maryland) (South Carolina) Number of segments 70 50-68 42 94-107 19* 74-80 70 Length (mm) 10 12 11 12 * c. 15 NA Width (mm) 0.16 0.3 0.4 0.24—-0.64 0.15 0.30-0.47 0.22-0.29 Spermathecal setae (um) NA 62-67 >35 (broken) 46-54 60 50-72 47-67 Penial setae NA 62-67 62 47-62 63 54-70 46-64 Dorsal crotchets (anterior) (um) NA NA 65 46-59 54 55-70 44-65 Dorsal crotchets (posterior) (um) NA NA 38 40-60 43 37-52 40-45 Hair setae (anterior) (um) 290 448 300 157-250 200 205-245 137-232 Hair setae (posterior) (um) 290 NA 362 287-442 142 147-215 95-170 Ventral setae (anterior) NA 90 55 40-53 54 55-63 45-51 Ventral setae (posterior) NA c. 90 49 40-50 43 44-62 43-48 * = incomplete. NA = not available. The fluminialis form is most similar to the speciosus form, but differs in number and morphology of anterior pectinate and ventral bifid setae (Table 2). It differs from the zavreli form by having an equal or slightly subequal lower tooth on all ventral setae, fewer intermediate pectinate teeth, and bifid posterior dorsal crotchets. It may be separated from the simsi form by possessing bifid crotchets instead of pectinate setae and shorter hair setae in posterior dorsal bun- dles. Discussion Separation of species within the genus Haber is difficult because of the extensive intraspecific morphological variation of se- tae. Brinkhurst and Chapman (pers. comm.) have demonstrated that a change in the con- ductivity of the water can alter the degree of pectination of dorsal crotchets in two un- related tubificid species, can change the length of hair setae and cause them to have a hispid appearance, and can even remove hair setae altogether. Additional experi- ments have indicated that setal variations have been associated with salinity and ionic concentrations (Giere and Pfannkuche 1982). The relative length of the teeth on the crotchets may also be subject to varia- tion due to wear (Brinkhurst 1971). Con- sequently, the basic criteria for distinguish- ing Haber speciosus from its congeners should be based primarily on the configu- ration of the male efferent ducts and sec- ondarily on the setal shape and distribution, the latter characters being most useful in the separation of “forms” or distinct popula- tions exhibiting the same genital morphol- ogy. The two largest species of Haber are no- table exceptions, because their setae are so distinct from their congeners. Haber sviren- koi and H. dojranensis are the only mem- bers of this genus lacking pectinate setae, and having the upper tooth of dorsal and ventral bifid setae much shorter than the lower. Haber svirenkoi, the only species (in mature specimens) lacking preclitellar hair setae, has a greater number of anterior dor- sal crotchets, and is the largest (Tables 3 and 4). Although these characters distinctly separate H. svirenkoi from the other seven congeners, comparison of the vas deferens will further distinguish this species from H. 412 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.— Distinguishing characters of the species of Haber. dorsal hair setae Haber per bundle dorsal crotchet setae per bundle Holmquist, 1978 svirenkoi Lastokin, 1937 7 tat ia 4 lee dojranensis Hrabe, 1958 — monfalconensis Hrabe, 1966 >| _=S amurensis Sokolskaja and Hrabe, 1969 Ppyrenaicus Juget and Giani, 1974 turquini Juget and Lafont, 1979 hubsuglensis Semernoi and Akinshina , 1980 (Si *Peloscolex sp.” 1-2 Giani and Martinez— Ansemil, 1981 see NA = NOT AVAILABLE speciosus. The ental portion of the vas def- erens in H. svirenkoi is narrower (c. 12 wm vs. c. 20 um), and the ectal region is much greater (c. 40 wm vs. >65 um) and lacks ciliation. Whereas Finogenova (1972) de- scribes a chitinous tubular structure at the male sexual orifice, further investigation may reveal it to actually be synonymous with the basal membrane of H. speciosus, as indicated by Holmquist (1979). ante- Iposte-| anterior posterior rior rior morphology of male efferent ducts ventral bifid setae per bundle anterior NA The shape of the male efferent duct can also be used to differentiate H. speciosus from H. dojranensis. The vas deferens of the latter is of uniform width, not distinctly set off from the very narrow atrium (Table 4). Haber speciosus can be distinguished most reliably from the remaining taxa by com- paring the position of the spermathecal pore in relation to the ventral seta, and the shape VOLUME 99, NUMBER 3 413 Table 3.—Size, position of spermathecal pore and type-locality of the species of Haber. Segment Length Width at Species number (mm) clitellum (mm) speciosus 70-170 10-12 0.16-0.47 svirenkoi 180 22-38 NA dojranensis 140-150 20-25 NA monfalconensis NA NA 0.28 amurensis NA NA 0.44 pDyrenaicus 50-70 9-15 0.4—0.5 turquini 13-27 1-3 0.14-0.32 hubsugulensis 38 10 0.36 NA = not available. of the male efferent duct. The spermathecal pore is located in the line of the ventral setae in all species except H. speciosus, where it lies in the lateral line dorsal to the ventral seta (Table 3). Haber monfalconensis was originally de- scribed as a subspecies of H. speciosus (Hrabé 1966), but Brinkhurst and Jamieson (1971) synonymized it with H. speciosus along with Peloscolex simsi and P. zavreli. Holmquist (1979) elevated it to species sta- tus because the position of the spermathecal pore is in line with the ventral setae. The narrow region of the vas deferens is also shorter as is the ejaculatory duct which en- ters the atrium more gradually (Table 4). A further separation is based on the shape of the pectinate setae. In H. monfalconensis the lateral teeth are acute and very long, the upper tooth longer than the lower (Hrabé 1966) (Table 4). In H. speciosus the lateral teeth are obtuse and of equal length (Table 3). Somatic setae of H. speciosus and H. pyr- enaicus are very similar. Conversely, the genital setae are distinctive. Penial setae are rarely present in H. pyrenaicus, and the spermathecal setae are more than twice as long (120-130 um) as those observed in H. speciosus. The male genitalia of H. pyren- aicus are also considerably different: the vas deferens is of uniform width, the prostate is attached posteriorly on the atrium, and the ejaculatory duct is absent (Table 4). As in H. svirenkoi, the basal membrane of the Spermathecal pore in lateral line in line of ventral setae in line of ventral setae in line of ventral setae in line of ventral setae in line of ventral setae in line of ventral setae in line of ventral setae Type-locality Lake Ochrid, Yugoslavia Dnepr River, U.S.S.R. Lake Dojran, Yugoslavia Timova River, Europe Amur River, U.S.S.R. Mountain Lakes, High Pyrenees Puits de Rappe, France Lake Khubsugul, Mongolia penis may have been misinterpreted as a cuticular sheath. The only way to separate reliably H. spe- ciosus from H. hubsugulensis is to compare the morphology of the male genitalia. The vas deferens of the latter species is similar in shape to that of H. speciosus, but is ap- proximately one-half the width. The atrium is sacciform, leading directly into a cone- shaped penis and the ejaculatory duct is ab- sent (Table 4). Semernoi and Akinshina (1980) describe the penis of H. hubsugu- lensis as being covered by a thickened cu- ticle, but without a separate penial case. Descriptions of H. turquini and H. amu- rensis were derived from poorly preserved material and are here only tentatively con- sidered as distinct species. Sokolskaja (1961) briefly described H. amurensis as Tubifex sp. No. | from the examination of a single incomplete specimen in poor condition. The subsequent conflicting description by Hrabé (1969) as Tubifex amurensis was based on a single series of damaged longitudinal sec- tions. Although the description is adequate enough to commit it to the genus Haber, I agree with Brinkhurst (1971) that a specific designation should be reserved until addi- tional specimens of better quality are ex- amined. At present, the location of the sper- mathecal pore in line with the ventral setae of H. amurensis is the only definitive char- acter to separate it from H. speciosus. The description of H. turquini (Juget and Lafont, 1974) omits the morphology of the 414 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 4.— Distinguishing characters of the forms of Haber speciosus. morphology of male efferent ducts Ga) NA Haber dorsal crotchet dorsal hair ventreyplt¢ . setae setae per bundle ae ee setae per bundle per bundle orm lke rior |rior 1-2 Iv 2 3-4 1-2 | speciosus 3-4 lia 2 simsi (England) NA zavreli [7 simsi (Florida) fluminialis (Maryland) 4 (5) 1 (2) 2-4] 1 fluminialis (South Carolina) \ 17 1-2 1 “possible congener” (New York) \ 1 2-3 fluminialis (New York) NA = NOT AVAILABLE genitalia (other than the presence of mod- ified spermathecal and penial setae) because of poorly preserved material, but somatic characters are distinctive enough to sepa- rate it from H. speciosus. Haber turquini is the smallest species in this group, and the only one to have a papillate cuticle remi- niscent of Peloscolex. The shape of the “gaine cuticulisée”’ covering the penis is of the form characteristic of the basal mem- brane in other species of Haber, and may iiaeal ea : 4-8 2 Py | SZ bia | 4-5 2 ol oth ae 3-6 2-3 NA prove to be the same structure upon further examination of additional material. Two specimens of indeterminable status have been described in the literature and tentatively referred to this genus (Brink- hurst 1981; Brinkhurst and Wetzel 1984). Giani and Martinez-Ansemil (1981) col- lected a mature specimen, identifying it as ‘‘Peloscolex sp.’ They compared it to H. speciosus and H. zavreli. Penial setae are absent, but the penis is enclosed by a “cu- VOLUME 99, NUMBER 3 ticular sheath” similar to H. zavreli. The shape of the spermatheca and spermato- phore also conform to the genus Haber. Un- til conspecifics can be more thoroughly ex- amined, this species is best left undesignated. Brinkhurst (1981) mentioned a “‘possible congener” from the Hudson River, New York, of a species he referred to as Haber cf. speciosus described from the Susquehan- na River, New York. The specimen was de- posited in the United States National Mu- seum of Natural History (USNM 65224) and identified as Peloscolex speciosus. The specimen was mounted in a clearing me- dium. Consequently, the genital morphol- ogy cannot be determined. However, the presence of spermathecal and penial setae suggests its placement in the genus Haber. The morphology of the somatic setae is dif- ferent from any form thus far encountered in North America (Table 2). Examination of additional specimens will probably es- tablish it as either a new species of Haber or as an additional “‘form”’ within the com- plex of Haber speciosus, most similar to the zavreli form. As Bird and Ladle (1981) indicated, im- mature specimens of H. simsi, confirmed herein as a synonym of H. speciosus, may be confused with Tubifex ignotus. They pre- sented a comparison between the setae of the two species, and used this character as a basis for separation. Unfortunately, due to the plasticity of the setal morphology, this distinction is unreliable. Care must be ex- ercised when referring to the taxonomic key prepared by Stimpson and Klemm (1982) on the Tubificidae of North America. An immature specimen of H. speciosus will be misidentified as 7. ignotus. Careful exam- ination of mature specimens is essential for accurate identification. Summary The genus Haber is newly reported from eastern North America. Specimens were ex- amined from New York, Maryland, South Carolina, and Florida, and have been de- termined to be the same species, Haber spe- 415 ciosus, based on similar morphology of the male efferent duct. The eight species of Ha- ber are separated, primarily by differences within the male genitalia, and secondarily by setal morphology and modifications of the body wall. Four distinct forms of H. speciosus can be distinguished on the basis of their setal morphology. Two of these forms are represented in the North Amer- ican fauna, the fluminialis form and simsi form. The former has been found only on the east coast of North America. The latter has been collected from the Gulf of Mexico, Florida, and from England. The material from England was originally described as a distinct species, P. simsi. The remaining two forms, zavreli and speciosus, have only been reported from Europe. Acknowledgments I am very indebted to Dr. R. O. Brink- hurst (Institute of Ocean Sciences, British Columbia) for his valuable guidance and critical evaluation of the manuscript; to Mr. Michael T. Barbour (EA Engineering, Sci- ence and Technology, Inc.) and Dr. Michael Ladle (Freshwater Biological Association, England) for providing me with oligochaete material; to Dr. Christer Ers¢us (Swedish Museum of Natural History) for reviewing the manuscript; to Mote Marine Laboratory for the time and resources to complete the study; and to the Southwest Florida Water management District for financial support. Literature Cited Bird, G. J., and M. Ladle. 1981. Aspects of the mor- phology and ecology of two British tubificids (Oligochaeta).—Journal of the Zoological So- ciety of London 194:493-504. Brinkhurst, R. O. 1963. Taxonomical studies on the Tubificidae (Annelida, Oligochaeta).—Interna- tionale Revue der Gesamten Hydrobiologie, Systematische Beihefte 2:1-89. 1962. A re-description of Peloscolex varie- gatus Leidy (Oligochaeta, Tubificidae), with a consideration of the diagnosis of the genus Pe- loscolex.—Internationale Revue der Gesamten Hydrobiologie 47(2):301—306. . 1966. Taxonomical studies on the Tubifici- dae (Annelida, Oligochaeta) supplement.—In- 416 ternationale Revue der Gesamten Hydrobiolo- gie 51(5):727-742. 1981. A contribution to the taxonomy of the Tubificinae (Oligochaeta: Tubificidae).—Pro- ceedings of the Biological Society of Washington 94:1048-1067. , and H. R. Baker. 1979. A review of the ma- rine Tubificidae (Oligochaeta) of North Amer- ica.—Canadian Journal of Zoology 57(8):1553- 1569. , and B. G. M. Jamieson. 1971. Aquatic Oli- gochaeta of the world. University of Toronto Press, Toronto, Canada, 860 pp. ,and M. J. Wetzel. 1984. Aquatic Oligochaeta of the world: Supplement. A catalogue of new freshwater species, descriptions, and revi- sions.—Canada Technical Report of Hydrog- raphy and Ocean Sciences Number 44, 101 pp. Finogenova, N. P. 1972. [Newspecies of Oligochaeta from Dnieper and Bug Firth and Black Sea and revision of some species.] [in Russian].— Trudy Zoologichesko Instituta Akademii Nauk SSSR 52:94-116. Giani, N., and E. Martinez-Ansemil. 1981. Contri- bution a la connaissance des oligochétes aqua- tiques du Bassin de L’Argens (Var, France).— Annales de Limnologie 17(2):121-141. Giere, O., and O. Pfannkuche. 1982. Biology and ecology of marine Oligochaeta, a review.— Oceanography and Marine Biology Annual Re- view 20:173-308. Holmquist, C. 1978. Revision of the genus Peloscolex (Oligochaeta, Tubificidae). 1. Morphological and anatomical scrutiny; with discussion on the ge- neric level.— Zoologica Scripta 7:187—208. 1979. Revision of the genus Peloscolex (Oli- gochaeta, Tubificidae). 2. Scrutiny of the species. — Zoologica Scripta 8:37-60. Hrabé, S. 1931. Die Oligochaeten aus den Seen Och- rida und Prespa.—Zoologische Jahrbiicher Ab- teilung fiir Systematik Oekologie und Geogra- phie der Tiere 61:1-62. 1942. Poznamky o zvirene ze studni a pra- menu na Slovensku.—Sbornik Prirodovedeck- eho klubu v Brne 24:23-30. . 1958. Die Oligochaeten aus den Seen Dojran und Skadar.—Spisy Vydavane Prirodovedeck- ou Fakulton Masarykovy University 397:337- 354. . 1962. Oligochaeta limicola from Onega Lake collected by B. M. Alexandrov.—Spisy Priro- dovedecke Fakulty University J. E. Purkyne v Brne 435:277-333. 1964. On Peloscolex svirenkoi (Jarosenko) and some other species of the genus Peloscol- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ex.—Spisy Prirodovedecke Fakulty University J. E. Purkyne v Brne 450:101-112. 1966. New or insufficiently known species of the family Tubificidae.—Spisy Prirodovedecke Fakulty University J. E. Purkyne v Brne 470: 57-77. 1969. On certain points in the structure of Tubifex minor Sokolskaja, and Tubifex amu- rensis n. sp. Sokolskaja and Hrabé.—Spisy Pri- rodovedecke Fakulty University J. E. Purkyne v Brne 506:259-263. . 1981. The freshwater Oligochaeta (Annelida) of Czechoslavakia.—Acta Universitatis Caro- linae-Biologicica 1979(-2):1—167. [in Czech with English summary] Juget, J., and N. Giani. 1974. Repartition des oli- gochétes lacustres du massif de Neovielle (Hautes-Pyrenees) avec la description de Pelo- scolex pyrenaicus, n. sp.—Annales de Limnolo- gie 10(1):33-53. , and M. Lafont. 1979. Déscription de Pelo- scolex turquini, n. sp. et redéscription de Pelo- scolex moszynskii, Kasprazak, 1971 (Tubifici- dae, Oligochaeta), avec quelques remarques sur la répartition du genre Pe/oscolex dans les eaux douces Fran¢aises.— Bulletin Mensuel de la So- ciété Linnéene de Lyon 48(2):75-80; 113-118. Kasprzak, K. 1973. [Notes on Oligochaeta fauna of Poland, I.].—Fragmenta Faunistica 18:405—434. [in Polish with Russian and English summary] Lastockin, D. A. 1937. New species of Oligochaeta Limicola in the European part of the USSR.— Doklady Akademii Nauk SSSR 17:233-235. Semernoi, V. P., and T. V. Akinshina. 1980. [Oli- gochaeta worms of Lake Khubsugal and some other water bodies of Mongolia].— Proceedings of Soviet-Mongolian Multidisciplinary Khub- sugal Expedition. Irkutsk State University Press, Irkutsk, USSR 1980:117-134. Sokolskaja, N. L. 1961. Materialy po faune pres- novodnych malostetinkovych cervej bassejna Amura.—Sborniktrudov Zoologicheskogo Muzeya Moskovskogo Universiteta 8:79-101. Steen, E. B. 1971. Dictionary of biology. Harper and Row, Publishers, New York, New York. 630 pp. Stimpson, K. S., and D. J. Klemm. 1982. A guide to the freshwater Tubificidae (Annelida: Clitellata: Oligochaeta) of North America. —Environmen- tal Monitoring and Support Laboratory. EPA- 600/3-82-033, 61 pp. Mote Marine Laboratory, 1600 City Is- land Park, Sarasota, Florida 33577. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 417-428 REDESCRIPTION OF THE OLIGOCHAETE GENUS PROPAPPUS, AND DIAGNOSIS OF THE NEW FAMILY PROPAPPIDAE (ANNELIDA: OLIGOCHAETA) Kathryn A. Coates Abstract. Examination of type material of Propappus glandulosus Michael- sen, 1905, and P. volki Michaelsen, 1916, as well as other material of the latter, led to a reconsideration of the classification of Propappus Michaelsen, 1905, within the Enchytraeidae. Species included in Propappus, P. volki, P. glandu- losus, and P. arhyncotus Sokolskaja, 1972, possess character-states that are unique. They have sigmoid, nodulate, bifid setae; large epidermal glands pos- terior to each setal bundle; spermathecal pores at septum 3/4; and glandular parts of the vasa deferentia located posterior to 11/12. Another significant, unique character state is the location of the single pair of ovaries in XIII, except in P. arhyncotus, rather than in the male pore segment, XII. This character, in addition to the others, warrants the removal of Propappus from the Enchy- traeidae. The new, monotypic family Propappidae is erected for species of Propappus. The genus Propappus is redefined. Propappus glandulosus and P. volki are redescribed, and a description of P. arhyncotus is given. Propappus Michaelsen, 1905, has been accepted as being nearest the stem form of the Enchytraeidae (Michaelsen 1916a, 1923; Stephenson 1930; Cernosvitov 1937a; Timm 1981). These authors thought that Propappus exhibited a high number of an- cestral states for the Enchytraeidae, remi- niscent of a freshwater origin of the family. All species of the genus, P. glandulosus Mi- chaelsen, 1905, P. volki Michaelsen, 1916, and P. arhyncotus Sokolskaja, 1972, are known from temperate, fresh waters. The presence of both P. glandulosus and P. volki in Lake Baikal, an ancient lake, enhanced the view of the genus as, itself, relatively ancient. Propappus volki, the species most studied, is regarded as characteristic of pure waters with sandy bottoms, and strong or moderate currents (Bird 1982). Michaelsen (1905) established the genus Propappus and described its type species (by monotypy), P. glandulosus. In his diagnosis of this genus he noted that one characteristic was unique for the Enchytraeidae: the setae on all the segments were distinctly double- pointed (now see also Barbidrilus Loden and Locy, 1980). His species description includ- ed other character states anomalous for En- chytraeidae: 1) each setal bundle was as- sociated with a large, glandular organ; 2) setae occurred ventrally in the male pore segment; 3) the sperm funnel was not glan- dular, but the ental part of the vas deferens, posterior to 11/12, was thickened and glan- dular; and 4) the ovary was located on the posterior of 12/13, in the segment posterior to the male pore segment. According to the latter character state the species of Propap- pus were plesiopore, like all other enchy- traeids, but had a sterile segment between the two fertile segments, as in many earth- worms (Lumbricida Brinkhurst, 1982) and some Haplotaxidae and Lumbriculidae. Michaelsen (1916a, b) described a second species, Propappus volki. It differed from P. glandulosus in that it was slightly larger and possessed a well-developed, finger-like, prostomial proboscis. Michaelsen (1916a) 418 noted that the ectal pore of the spermatheca was anterior to the setae of IV, as (““wie auch bei...”’) in P. glandulosus. This was, in fact, an amendment to the original description of P. glandulosus in which the spermathecal pore had been described in the usual en- chytraeid position, at 4/5. In his early papers on Propappus, Michaelsen (1905, 1916a) described the ovaries in XIII, with the egg cell string extending from 12/13, and clearly recognized (Michaelson 1905) that this was not the usual enchytraeid condi- tion. He did not include the location of the ovaries in the diagnosis of Propappus. Later, Michaelsen (1923) described the ovaries in P. volki as lying on the posteroventral part of 11/12. He reported that he had never clearly seen the female funnel or female pore but thought that both were in 12/13. Ma- terial from the Volga River was used for these (Michaelsen 1923) amendments to the description of P. volki. Michaelsen (1916a) suggested that, be- cause of the double-pointed setae, the lack of integrity of the pharyngeal glands, the loose structure of the nephridial postseptale, and the structure of the sperm funnel (=the preseptal part of the male duct), Propappus was very likely near the stem form of the Enchytraeidae. Cernosvitov (1937a) stated categorically that Propappus was the most primitive genus of the Enchytraeidae. This, he thought, was most clearly shown by the setal form and the structure of the sperm funnels. He also suggested that a number of character states pointed to a close affinity between Propappus and a subgroup of Mes- enchytraeus Eisen, 1878, including nephrid- ial form, structure of the pharyngeal glands, development of the sperm funnel, and structure of the longitudinal muscles. Niel- sen and Christensen (1959) suggested that Propappus was rather doubtfully placed in the Enchytraeidae at all. At no time was the diagnosis of that family amended to include the gonadal arrangement originally de- scribed in Propappus, nor were derived character states unifying Propappus with PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON other Enchytraeidae recognized. Instead, many of the derived character states pos- sessed by Propappus are unique and those states that are shared with Enchytraeidae can only indicate some more remote com- mon ancestor. Materials and Methods Cotype material was borrowed from the British Museum (Natural History). All ma- terial was examined under a compound mi- croscope. Drawings were made with the aid of a drawing tube (camera lucida). Photo- graphs were made using a Zeiss-Universal photo-microscope. The means of the lengths of the probosces of Propappus volki in dif- ferent populations were compared using the two-sample ¢ test for two-tailed hypotheses, all means were tested for significant differ- ence from 0 and the samples were compared by a non-parametric Mann-Whitney U test (Zar 1974). Propappidae, new family Type genus. —Propappus Michaelsen, 1905. Diagnosis. — As for type genus. Distribution. — Palearctic. Discussion. —Species of the genus Pro- pappus do not exhibit apomorphies shared with other Enchytraeidae. A group Enchy- traeidae, including Propappus, cannot be di- agnosed. The few, obviously derived char- acter states of Propappus do not allow an immediate, intuitive choice for the precise position of Propappus within the existing classifications of the Oligochaeta. A new family of Oligochaeta, the Propappidae with type genus Propappus, is established here for the single genus and its included species P. glandulosus, P. volki, and P. arhyncotus. The family Propappidae is phylogenet- ically near both the Enchytraeidae and some species of Haplotaxidae (Coates MS). Those species of Haplotaxidae from which the lin- eage of the family Propappidae is not yet clearly resolved include Metataxis ameri- canus (Cernosvitov 1939) and M. brink- VOLUME 99, NUMBER 3 hursti (Cook 1975) in the recently recog- nized genus Metataxis Righi, 1985. Along with at least some Enchytraeidae, species of Propappidae are in a monophyletic lineage including Metataxis falcifer (OQmodeo 1958). The initial implications of this are that Metataxis is paraphyletic. Characteristic of all these nominal taxa is the presence of a pair of testes in at least segment XI, and, with the exception of Enchytraeidae, the oc- currence of paired ovaries only in segment XIII. In order to conserve a consistent tax- onomy ofthe sistergroup Enchytraeidae, and to avoid attenuating the diagnosis of that family, the new family Propappidae is erect- ed for the genus Propappus. The outgroup criterion for determining relative polarity of character states should be applied and phy- logenetic analyses performed to resolve these genealogical problems. The primary distinguishing characteris- tics of this monotypic family are discussed following the diagnosis of Propappus. Propappus Michaelsen, 1905 Propappus Michaelsen, 1905:24—25; 19 16a: 51-55; 1923:37—42.—Stephenson, 1930: 765—766.—Cernosvitov, 1937a:264—267, 269-270, 277, 279.—Nielsen and Chris- tensen, 1959:29-30.—Cekanovskaja, 1962:370-377. Palpenchytraeus Michaelsen, 1916a:52; nomen nudum. 1916b:1; Type species. —Propappus glandulosus Michaelsen, 1905, by monotypy. Diagnosis. —Setae in 4 bundles per seg- ment, including genital segments; bundles ventrolateral and dorsolateral. Three setae, rarely 2, per bundle; setae sigmoid, bifid, nodulate. Setal gland present immediately posterior to each setal bundle (Figs. la, g; 2a & 2b). Head and other dorsal pores ab- sent. Prostomium with or without probos- cis. Clitellum single-layered, usually ex- tending over XII—XIV. Pores of male and female reproductive systems paired. Sper- mathecal pores anterolateral in IV. Male 419 pores (Fig. la, m) anterior to ventral setal bundles and setal glands of XII. Female pores ventral, in or just posterior to 13/14. Anus opening dorsally. Cavity of prosto- mium almost filled by elongate, glandular, epidermal cells (Fig. la, pg). Brain deeply cleft posteriorly, divided into almost sepa- rate, lateral lobes. Dorsal pharyngeal pad (Fig. la, p) slightly thickened, protrusile. Bodies of glandular cells of pharyngeal pad not organized into compact pharyngeal glands. Gut without appendages or diver- ticula; expansion at intestinal origin may be abrupt. Holonephridial, preseptal parts of nephridia small (Fig. 1b), including funnel only; postseptal parts lobed, with little in- terstitial tissue. Spermathecae paired (Fig. la), originating in IV, not communicating with gut; ectal duct thick-walled, aglandu- lar; ampulla abruptly expanded, thin-walled, extending posteriad through a few seg- ments. One pair of testes (Fig. la, t) ventro- lateral on posterior of 10/11. Seminal ves- icle (Fig. la, sv) unpaired, extending as far anteriad as VI. Sperm funnels (Fig. 1a, sf) simple, on 11/12; vasa deferentia confined to XII, anteriorly thickened (?glandular). Atria, prostates and other copulatory glands lacking. One pair of ovaries (Fig. 1a, 0) ven- trolateral on the posterior of 12/13 (see P. arhyncotus). Female ducts simple, on 13/14 (as above). Distribution. — Palearctic. Central and eastern USSR; England; France; Austria; West Germany; Finland; and Norway. Lakes or running waters with moderate to strong currents; in sand or gravel. Discussion.—The Enchytraeidae can be characterized by their possession of a num- ber of shared character states: discrete, well- developed pharyngeal glands; testes only in XI, male pores and ovaries in XII; presep- tal, glandular sperm funnels in XI; dorsal setal bundles, when present, (usually) in a mid-lateral position; spermathecal pore/ pores at or just posterior to 4/5; and, fre- quently, modified glandular structures at the penial pore. 420 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Propappus volki: a, Drawing of anterior of whole-mount, to segment XIV; b, Nephridium from post- clitellar segment; specimens from southern England. Abbreviations—a, spermathecal ampulla; d, spermathecal duct; g, setal gland; m, male pore; 0, ovary; p, pharynx; pg, prostomial glandular epidermis; s, seta; sf, sperm funnel; sv, seminal vesicle; t, testis; v, dorsal blood vessel. VOLUME 99, NUMBER 3 The species of Propappus share a set of character states distinct from those of the Enchytraeidae: setae with bifid ectal ends, usually three per bundle; setal glands in as- sociation with every setal bundle; sper- mathecal pores at or near 3/4; thickened vasa deferentia in XII; and ovaries only in XIII with female funnels on 13/14. The species of Achaeta Vejdovsky, 1877, a genus of the Enchytraeidae, lacking setae entirely, may possess sac-like organs in po- sitions similar to those occupied by either the setal follicles of setate genera or by the setal glands of species of Propappus. The homologies of the organs in Achaeta have been debated by several authors (Michael- sen 1923, Stephenson 1930, Nielsen and Christensen 1959). In megadrile and some haplotaxid oligochaetes various glandular appendages of setal follicles, which may not all be homologous structures (Stephenson 1930), have also been termed setal glands. The organs of Achaeta were first de- scribed (Vejdovsky 1879) as homologues of the setae, possibly as the empty follicles. Michaelsen (1923) did not think that it was possible to distinguish the setal follicles of Achaeta from the setal glands of Propappus; the organs in Propappus were just composed of fewer, larger cells. Thus the glands were not a unique character of Propappus. Both Stephenson (1930) and Cekanovskaja (1962) also saw the structures of Achaeta as setal glands but Nielsen and Christensen (1959) referred to these as setal follicles or setal sacs for which the true homology (“‘nature”’) was doubtful. In sectioned and whole-mounted speci- mens of Achaeta that I have examined using a compound light microscope, cellular structure inside the cuticular sac of the setal gland could not be discerned. The contents of the cuticular sacs were granular and not densely packed. The internal cellular struc- ture of the organs of Propappus is easily discernible at similar magnifications, al- though it does not stain with borax carmine in whole mounts. The cells of the gland do 421 stain with haematoxylin and are quite ob- vious in sectioned material. A cuticle-like outer layer can be distinguished in both whole-mounts and in sections. There ap- pear to be two types of secretory cells pres- ent in each gland, distinguished by the stained color and by the coarseness of the granular inclusions. Such a glandular-type complement is common in invertebrate ad- hesive-gland systems. Cekanovskaja (1962) observed that specimens of P. volki adhere to stone and sand grains by means of some secretion from the setal glands of the pos- terior segments. With the present information it is not possible to establish the probable homolo- gies of the setal glands of either Propappus or Achaeta. Their distribution in Propappus specimens, co-occurring with all setal bun- dles, is unique among the microdrile oli- gochaetes. Other characteristics of Propappus are not regarded as derived. These are the presence of unmodified, ventral setae in the male pore segment; dorsolateral location of the dorsal setal bundles; unmodified gut; diffusion of the pharyngeal gland cell bodies; and the lack of interstitial tissue associated with the postseptal part of the nephridia. These char- acteristics are considered “primitive” for the class Oligochaeta (Jamieson 1978; Brink- hurst 1982, 1984). Propappus glandulosus Michaelsen, 1905 Figs. 2a, 2c & 2d Propappus glandulosus Michaelsen, 1905: 25-28, figs. 4-5.—Cernosvitov, 1937b: 273-274, figs. 3—4.—Cekanovskaja, 1962: 371-272, fig. 187. ?Propappus volki.—Sokolskaja, 1968:21. ?Propappus arhyncotus Sokolskaja, 1972: 83-85, fig. 5; 1973:64—-66, fig. 8. Material examined.—Syntypes of Pro- pappus glandulosus, BMNH 1949.3.1.226, mature whole-mount; BMNH 1949.3.1.227, a mature, sagittally sectioned specimen; and BMNH 1949.3.1.228, a mature, transverse- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON v Fig. 2. a, Propappus glandulosus, setal gland from sectioned cotype specimen BMNH 1949.3.1.227; b, Propappus volki, setal gland and seta from sectioned cotype specimen BMNH 1949.3.1.230; c, Propappus glandulosus, sagittal section of prostomium and pharyngeal pad of cotype BMNH 1949.3.1.227; d, Propappus glandulosus, prostomium of whole-mounted cotype specimen BMNH 1949.3.1.226. Scale bar = 50 wm. ly sectioned specimen. Collected in 1901 or 1902. Type locality. —Lake Baikal, middle and southwest side, in sand or gravel at 2-8 me- ters depth. Description of material examined. —Four to 5 mm long, with 28-32 segments; body diameter at IV—V (specimens slightly com- pressed) 175-224 (175, 219, 224) um, at IX about 300 (301, 307) um. Prostomium short and rounded, without proboscis (Figs. 2c & 2d). Spermathecal pores at 3/4. Setae 55- 73 (59-dorsal, 55, 73-ventral) um up to 90 um long (according to Michaelsen 1905); nodulus in distal one-third of seta; upper tooth much shorter and thinner than lower; ventral and dorsal setae alike in size. Female pore at 13/14. Setal glands (Fig. 2a) 80, 106 um long, 22-32 (22, 26, 32, 32) wm in di- ameter, flask-shaped with narrow ectal neck; entire gland appearing filled with secretions. Pharyngeal gland cells ventral in V—VIII. First pair of nephridia at 7/8 or 8/9, ne- phridia absent in clitellar and most of more posterior segments. Dorsal vessel origin at abrupt transition from esophagus to intes- VOLUME 99, NUMBER 3 tine, in VIII or IX. Thick-walled ental parts of vasa deferentia about 120 um long, 14, 24 wm in diameter; ectal part 2—3 times as long as ental, only 6-12 (6, 8, 7.5, 12) um in diameter. Diameter of vas deferens at male pore may be slightly greater, about 13 pm, canal sometimes with short, abrupt widening just ental to pore. Spermathecal duct extending to one-half IV, ampulla ex- tending into VI or more posteriad; duct nar- rowing gradually from pore to ampulla, from about 22 wm to 8-10 um, then widening slightly at ampulla (according to Cernos- vitov 1937b). Remarks. —The cotype specimens of Pro- pappus glandulosus examined here came from the collections of L. Cernosvitov held at the BMNH. This material was originally held in Hamburg and is from the type series (Cernosvitov 1937b). Discussion. —Although Michaelsen (1923), in redescribing Propappus_ volki, amended the position of the ovaries from 12/13 to 11/12 he did not include this char- acteristic in a formal generic diagnosis. In fact, P. glandulosus was never redescribed with the “new” ovarial position. Michaelsen (1905) reported that the ori- gin of the dorsal blood vessel and the abrupt widening of the esophagus to the intestine was in VIII. This was the condition seen here in P. glandulosus and also in the new material of P. volki from Lake Baikal. Fur- ther comments on the determination of the intestinal origin are made following the de- scription of P. arhyncotus. Propappus arhyncotus, also aproboscid- ial, may be a synonym of P. glandulosus, as indicated. Three characteristics possibly distinguish the species. In P. arhyncotus the origin of the intestine is reported to be post- clitellar, the diameters of the vasa deferentia are greater ectally than entally, and the ova- ries are located in the anterior of XII (So- kolskaja 1972, and pers. comm.). These characteristics are discussed following the description of P. arhyncotus. Distribution. —Lake Baikal, shallow (Mi- 423 chaelsen 1905) to abyssal (820 m) (Semer- noj 1983); Yenisey River from the mouth of the Angara River to Igarka (Cekanov- skaja 1962). As Propappus arhyncotus: Kamchatka Peninsula, Zhupanova River basin, Karymai Reach inflow to Bol’shaya River, and Kamchatka River basin (Sokol- skaja 1972). On sand and gravel bottoms. Propappus volki Michaelsen, 1916 Figs. 1, 2b & 3 Palpenchytraeus volki.—Michaelsen, 191 6b: (in Michaelsen, 1916a) 1. [momen nu- dum]. Propappus volki. Michaelsen, 1916a:51—55; 1923, 37-42; 1926:4.—Cernosvitov, 1937b:273, figs. 1-2.—Chekanovskaja, 1962:372-373, fig. 188.—Wachs, 1967: 322, fig. 9.—Bird, 1982:67-75. Non So- kolskaja, 1968:21. Material examined.—Cotypes from the Cernosvitov collection, BMNH 1949.3. 1.230, mature, sagittally sectioned speci- men, and BMNH 1926.927.1-5, 3 speci- mens in fluid, collected by W. Michaelsen, in the Lower Elbe River, near Hamburg; Nov 1915 to Jan 1916. Nineteen mature specimens collected from gravel at East Bur- ton on the River Frome, Dorset, England, during period Jan 1979 to Dec 1980. All stained in alcoholic borax carmine. Fifteen specimens subsequently mounted whole in Canada balsam; 3 specimens sectioned transversely and 1 specimen sectioned sag- ittally, at 7 wm intervals. Twenty mature and 15 immature specimens collected from River Tarn, tributary of Garonne River, Massif Central near Albi, France; 17 Oct 1979. Four specimens attributed to Pro- pappus volki from Lake Baikal. Two with developing gonads, one also with develop- ing spermathecae and vasa deferentia. All mounted whole, as above. Type locality.—Elbe River, near Ham- burg, at 3-5 m depth. Description of material examined. — Four to 7 mm long, with 29, 33-43 segments; 424 Fig. 3. and anterior of XIII with part of an ovarial strand; drawn from a sagittally-sectioned specimen from southern England. Abbreviations—m, male pore; 0, Ovary; s, seta; vd, vas deferens. Propappus volki, segment XII with male pore body diameter at IV—V about 150-300 um, at XI about 300-400 um; body of mature specimens usually constricted through XII and XIII. Prostomium with annulate, non- retractile proboscis (Fig. la), up to 97 wm long; proboscis with distinct lumen. Setae (Fig. la, s; 3, s) 40-80 um long, ventrals (preclitellar: X = 69.6 +/— 6.7 um [n = 17]; postclitellar: x = 60 +/— 12.4 wm [n = 5]) on average just longer than dorsals (precli- tellar: x = 64.0 +/— 14.0 um [n= 14]; postclitellar: t = 56.0 +/— 8.0 um [n = 8)); preclitellar setae slightly longer than post- clitellar setae; upper tooth much thinner and shorter than lower. Epidermal papillae in transverse ring at level of setae; other epi- dermal glands in 12-15 transverse rows per segment. Spermathecal pores just posterior to 3/4. Clitellar gland cells small, scattered over XII through 2 XIV. Male pores each on small, conical papilla (Fig. 6, m), body wall usually depressed around papillae. Brain with median, anterodorsal projection. Anterior dorsal blood vessel dividing in front of brain, vessel originating in VII (Fig. 1a, v) or in clitellar segments. Abrupt expansion of gut in VII or XIV to XV. Setal glands (Figs. la, s; 2b) flask-shaped, with narrow ectal neck; glands in anterior segments with maximum diameter about 17-25 um (x = PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 20.1 +/— 3.1 um [n = 10]), about 40-70 wm (x = 57.4 +/— 8.8 um [n = 9]) long. Ectal end filled by large cells with dense, finely or coarsely granular contents. Gland enclosed by cuticle-like layer. Nephridia (Fig. 1b) from 5/6. Seminal vesicle (Fig. la, sv) ex- tending as far anteriad as VI or VII and posteriad into XII. Testes (Fig. la, t) large, compact. Vas deferens sometimes with bit of thicker-walled ental part appearing to protrude into XI; ental part 20-52 um (Xx = 40 +/— 10 um [n = 15]) in diameter, 154, 171, 186 wm long; ectal part of vas about 7-16 wm (X = 12 +/ — 3.4 um [n= 17) in diameter, widening to 14-23 wm (x = 18.6 +/— 4.2 wm [n = 5]) at pore (Fig. 3, m), 2—4 times as long as ental part. Segment XII criss-crossed by muscular strands (Fig. 3). As many as 3 maturing eggs in XIII, extending into XII and/or XIV. Spermathe- cal ectal duct (Fig. la, d) bulbous just ental to pore, 21-29 wm (X = 26 +/— 3.2 um[n = 12]) in diameter at pore to 11-17 wm (x = 13 +/— 1.7 wm [n = 12]) between expan- sion and ampulla. Ampulla (Fig. la, a) ex- tending from 4/5 as far as XII, walls very thin. Remarks.—As for Propappus glandulo- sus, the specimens of P. volki held at BMNH are cotypes. The comprehensive description of P. volki by Michaelsen (1923) included specimens from the Volga River but none of these could be obtained for this study. The Volga material and additional cotypes are held at the Zoologisches Museum Uni- versitat Hamburg. Discussion. —The ovarial segment in the material examined was always XIII, and there was no evidence for lability of this character in Propappus volki (Michaelsen 1916a, 1923). The origin of the dorsal blood vessel and gut transition in Propappus volki are con- sistently (Michaelsen 191l6a, 1923) de- scribed as in postclitellar segments, XIV or XV. In the new material from Lake Baikal, none of which was fully mature, an abrupt expansion of the gut (?transition from VOLUME 99, NUMBER 3 esophagus to intestine) was located in VII or VIII, as for P. glandulosus, and the dorsal blood vessel origin was at the expansion or just anterior to it. The esophageal wall im- mediately posterior to the pharyngeal pad was highly folded in all specimens of P. volki, perhaps facilitating the eversion of the pad. In many specimens, the ental part of the setal glands appeared empty. Here the cu- ticular layer was collapsed forming a “‘tail”’ (Fig. 2b) on the gland. The glands appeared to be larger and full in more mature speci- mens. The length of the proboscis in specimens of Propappus volki obtained from England, France and Lake Baikal varied from 29 um to 97 um. Statistical comparisons of the pro- boscideal length for each population by the t test showed that the mean probosces lengths of the French and English popula- tions were not significantly different at the 95% level (mean lengths were 56.2 +/— 11.2 wm [n = 26] and 64.6 +/— 15.9 um [n = 15], respectively). The mean length of pro- bosces (38.5 +/— 5.4 um [n= 4]) for Baikal specimens was significantly different from the mean lengths of both French and En- glish populations. The Mann-Whitney U test gave the same results. All mean lengths of probosces were significantly different (P< 0.001) from a mean of 0 when compared by a ¢ test. The presence of a prostomial proboscis may be the only character state that can be used easily to distinguish P. volki from P. glandulosus. These species differ little in size or in the size and shape of var- ious taxonomic characters including setae, spermathecae and male ducts. Both Propappus volki and P. glandulosus are recorded from Lake Baikal. However, P. glandulosus is known from only a few additional locations near Lake Baikal, whereas P. volki is recorded from numer- ous, widespread locations in the USSR and Europe. Distribution. —Widely distributed in western Europe: Germany (Michaelsen 191l6a, Stephenson 1930, Wachs 1967); 425 Austrian Tyrol (Bird 1982); France (Juget 1980, 1984); and northwest Spain (Marti- nez-Ansemil 1982). Britain, southern En- gland (Ladle and Bird 1980, Bird 1982). Central and eastern Europe: rivers of the Baltic Sea Basin (Parele 1983); Latvia (Cekanovskaja 1962); Estonia (Bird 1982); Ukraine and Moldavia (Grigelis 1980); Po- land: Tatra and Carpathian Mountains (Bird 1982), Welna and other Polish rivers (Kas- przak 1980); Czechoslovakia (Bird 1982); and Romania (Bird 1982). Central USSR: large rivers, especially Volga and tributar- ies, and reservoirs (Michaelsen 1923, 1926; Cekanovskaja 1962; Grigelis 1980; Bird 1982); Lakes Baikal, Chud and Pskov (Cekanovskaja 1962). In springs and phreatic waters (moving ground waters); small stony streams, on sand beds in large rivers, in the sandy substrates of lakes and reservoirs (Bird 1982). Propappus arhyncotus Sokolskaja, 1972 Propappus arhyncotus Sokolskaja, 1972:83- 85, fig. 5; 1973:64-66, fig. 8. Propappus volki. —Sokolskaja, 1968:21. Type material. —No. 1001 in the Zoolog- ical Museum of Moscow University (not seen). Collected 11 Aug 1964 by V. Ya. La- vanidov. Type locality.—Kamchatka, Lake Aza- bach’e at 7 cm depth; gravel substrates. Description from the literature. —Six to 12 mm long, maximum diameter 300—400 um; with 29-40 segments. Prostomium large, without proboscis. Setae commonly 3-4, rarely 5—6 per bundle in preclitellar seg- ments, usually 3 posteriorly. Dorsal setae 55-71 um long, ventral setae 59-76 um long; length of setae increasing slightly from an- terior to posterior segments; upper tooth much shorter and thinner than lower; nod- ulus distal. Spermathecal pores approxi- mately lateral at or in 3/4. Small epidermal tubercles irregularly distributed on prosto- mium and peristomium, in more regular rings in following segments. Male pores on 426 cone-shaped papillae. Setal glands flask- shaped, about 55 um by 27 wm at segments VII—X. Most anterior nephridia in V (? at 5/6). Esophagus widening to intestine at XV. Vasa deferentia about 310 um long, wid- ening from about 17 wm in diameter entally to 29 um near pore, distal end further wid- ened to 34—38 wm but with narrow channel. Ovaries paired in anterior of XII. Sper- mathecal duct about 160 um long, extend- ing through IV, ampulla may extend into VI. Remarks.—It has not been possible to borrow or otherwise obtain specimens of Propappus arhyncotus, and some questions concerning its structure could not be re- solved. Discussion. —Sokolskaja (1972) recorded that bundles of three through six setae were present; however, she consistently depicted (1972, fig. SA, obviously schematic) only three setae per bundle. Other species of Pro- pappus have only two or three setae per bun- dle. As in some enchytraeids, replacement setae may be present at the same time as the seta or the bundle being replaced. This was observed here in P. volki. The entire bundle complement can be replaced simul- taneously, leading, in Propappus species, to bundles appearing to have four or six and possibly five setae. The condition of the male duct described by Sokolskaja (1973) for Propappus arhyn- cotus is quite different from that found in either P. volki or P. glandulosus. In the for- mer the vas deferens widens from the ental to the ectal end whereas in the latter two species the ental part (about one-quarter of the total length of the vas deferens) is no- tably wider and the walls more “‘glandular”’ than in the ectal part. In P. arhyncotus there is a considerable widening of the vas def- erens at the male pore, to about 34-38 um. In P. volki and P. glandulosus the terminal part of the vas deferens is either not mea- surably wider or may be a few micrometers wider than the ectal part, with a maximum diameter of 23 um in P. volki. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON It was originally stated (Sokolskaja 1972), and has been confirmed (Sokolskaja, pers. comm.) that the ovaries are in segment XII in Propappus arhyncotus. It would seem that little doubt could remain about this char- acter state. However, the same character state has erroneously persisted in the liter- ature regarding both P. glandulosus and P. volki for more than 60 years. Other shared characters point to a very close relationship of all three species and similarity of the go- nadal condition seems probable. Sokolskaja (1972) pointed out four cate- gories of differences of Propappus arhyn- cotus from P. volki: 1) the shape of the pro- stomium and absence of proboscis; 2) the thickness and sculpturing of the cuticle; 3) the location of the spermathecal pores in 3/4; and 4) the structure of the spermatheca, with a short ampulla extending only into V. Michaelsen (1923) mentioned that the cu- ticle in Propappus was very thick and that there were rings of prominent epidermal cells around the segments. The epidermal pa- pillae are similarly described by Sokolskaja (1972). The location of the spermathecal pore in all the specimens of Propappus ex- amined was in or very near the furrow marking 3/4. Sokolskaja (1973) modified the description of the spermatheca of P. arhyn- cotus to indicate that the ampulla was more sac-like and extensive in mature specimens. Apparently only the proboscidial (and ovar- ial) character state separates P. arhyncotus from P. volki. Propappus arhyncotus was distinguished (Sokolskaja 1972) from P. glandulosus by the postclitellar location of the esophageal- intestinal transition in the former. This is a character that may be difficult to determine without examination of the cellular struc- ture of the gut. In much of the fixed material examined here the location of the gut tran- sition was, in fact, difficult or impossible to observe. Present estimations of the location of the gut transition are not consistently re- liable. This species remains a systematic prob- VOLUME 99, NUMBER 3 lem. If the ovaries are in XIII, then Pro- pappus arhyncotus probably should be syn- onymized with P. glandulosus. However, the condition of the male ducts also requires examination. If the ovaries are in XII, P. arhyncotus may differ from the other species of Propappus at the generic or even familial level. Only re-examination of specimens of P. arhyncotus can resolve its specific, ge- neric and familial relationships and conse- quent classification. Distribution. —Kamchatka Peninsula: Zhupanova River basin, inflow of Bol'shaya River, Kamchatka River basin, including Lakes Azabach’'e and Klyuchevskoe. Acknowledgments Drs. G. Bird, of the Freshwater Labora- tory East Stoke, Wareham, England, N. Gi- ani, and R. O. Brinkhurst generously loaned new material of Propappus volki from En- gland, France, and Lake Baikal, respective- ly. Mr. G. Easton and Mr. R. W. Sims made available the cotype material of P. glan- dulosus and P. volki held at the BMNH. Dr. N. P. Finogenova transmitted information between N. Sokolskaja, the author of P. ar- hyncotus, and myself. Facilities for this study were provided by the Ocean Ecology Division, Institute of Ocean Sciences, Fisheries and Oceans, Sid- ney, British Columbia, and, during the com- pletion of the work, both facilities and sup- port were provided by the Department of Invertebrate Zoology of the Royal Ontario Museum, Toronto. Drs. C. Erséus, G. Allen and D. R. Calder read the manuscript and made numerous helpful and provocative suggestions. B. A. Ruddell is gratefully ac- knowledged for preparing the successive versions of the manuscript. Literature Cited Bird, G. J. 1982. Distribution, life cycle and popu- lation dynamics of the aquatic enchytraeid Pro- Ppappus volki (Oligochaeta) in an English chalk- stream.— Holarctic Ecology 5:67—75. Brinkhurst, R.O. 1982. Evolution in the Annelida. — Canadian Journal of Zoology 60:1043-1059. 427 1984. The position of the Haplotaxidae in the evolution of oligochaete annelids.— Hydro- biologia 115:25-36. Cernosvitov, L. 1937a. System der Enchytraeiden. — Bulletin de l’Association pour les Russe Re- cherches Scientifiques 4 Prague 5:262—295. . 1937b. Zur Kenntnis der Enchytraeidae. IV. Revision einiger Michaelsenscher Typen.— Zoologischer Anzeiger 118:273-281. . 1939. The Percy Sladen Trust Expedition to Lake Titicaca in 1937. VI. Oligochaeta. — Trans- actions of the Linnaean Society, London 1 (se- ries 3):80—116. Cekanovskaja, O. V. 1962. Aquatic Oligochaeta of the USSR.—Akademiya Nauk SSSR, Zoologi- cheskii Institut. Opredeliteli po Faune SSSR 78: 1-411. [Translated from Russian for the U.S. Department of the Interior. New Delhi: Amer- ind Publishing Co. 1981]. Cook, D. G. 1975. Cave-dwelling aquatic Oligo- chaeta (Annelida) from the eastern United States.— Transactions of the American Micro- scopical Society 94:24—37. Grigelis, A. 1980. Ecological studies of aquatic oli- gochaetes in the USSR, pp. 225-240. In R. O. Brinkhurst and D. G. Cook, eds., Aquatic Oli- gochaete Biology. New York: Plenum Press, ix + 529 pp. Jamieson, B.G. M. 1978. Phylogenetic and phenetic systematics of the opisthoporous oligochaetes (Annelida: Oligochaeta). — Evolutionary Theory 3:195-235. Juget, J. 1980. Aquatic Oligochaeta of the Rh6ne- Alpes area: current research priorities, pp. 241— 252. In R. O. Brinkhurst and D. G. Cook, eds., Aquatic Oligochaete Biology. New York: Ple- num Press, ix + 529 pp. . 1984. Oligochaeta of the epigean and under- ground fauna of the alluvial plain of the French upper Rh6éne (biotypological trail).—Hydro- biologia 115:175-182. Kasprzak, K. 1980. Oligochaeta community struc- ture and function in agricultural landscapes, pp. 411-432. In R. O. Brinkhurst and D. G. Cook, eds., Aquatic Oligochaete Biology. New York: Plenum Press, ix + 529 pp. Ladle, M., and G. J. Bird. 1980. Aquatic Oligochaeta of southern England, pp. 165-174. In R. O. Brinkhurst and D. G. Cook, eds., Aquatic Oli- gochaete Biology. New York: Plenum Press, ix + 529 pp. Loden, M.,andS. Locy. 1980. Barbidrilus paucisetus, new genus, new species (Oligochaeta: Enchy- traeidae) from eastern North America.—Pro- ceedings of the Biological Society of Washington 93:1173-1176. Martinez-Ansemil, E. 1982. Les oligochétes aqua- 428 tiques de la Peninsule Ibérique (2e note) avec la description de Lumbricillus brunoi n.sp. (En- chytraeidae).— Bulletin de la Société d’ Histoire Naturelle de Toulouse 118:145-151. Michaelsen, W. 1905. Die Oligochaeten des Baikal- Sees. — Wissenschaftliche Ergebnisse einer Zoo- logischen Expedition nach dem Baikal-See 1900- 1902. Kiev and Berlin, R. Friedlander und Sohn. 69 pp. 1916a. Ein eignetiimlicher neuer Enchy- traeide der Gattung Propappus aus der Nie- derelbe.— Verhandlungen der Naturwissen- schaftlicher Verein, Hamburg n.s.3, 23:51-55. 1916b. Palpenchytraeus volki n.gen. n.sp.— Hamburger Nachrichten 53 (30 January 1916): 1. [in Michaelsen, 191 6a]. 1923. Die Oligochaeten der Wolga.— Arbei- ten der Biologischen Wolga-Station, Saratov 7: 30-43. 1926. Oligochaeten aus dem Gebiete der Wolga und der Kama.—Arbeiten der Biolo- gischen Wolga-Station, Saratov 9:1-11. Nielsen, C. O., and B. Christensen. 1959. Studies on Enchytraeidae. 7. Critical revision and taxon- omy of European species.—Natura Jutlandica 8-9:1—160. Omodeo, P. 1958. La reserve naturelle intégrale du Mont Nimba. IV. 1. Oligochétes.— Mémoires de l’Institut Fondamental d’Afrique Noire 53: 9-17. Parele, E.A. 1983. Oligochaete fauna of the big rivers in the Baltic Sea basin, pp. 76-81. Jn B. E. Ku- rashvili, ed., Proceedings of the 4th All-Union Symposium, Tbilisi, 5—7 October, 1983. Aquat- ic Oligochaeta. [Translated from Russian. Ca- nadian Translation of Fisheries and Aquatic Sci- ences #5091, 1984. 143 pp.] Righi, G. 1985. Dois novas Oligochaeta da Ama- zonia Venezuelana.—Papeis Avulsos de Zoo- logia, Sao Paulo 36:23-30. Semernoj, V. P. 1983. Species composition and dis- tribution of oligochaetes in the abyssal zone of Lake Baikal, pp. 100-104. Jn B. E. Kurashvili, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ed., Proceedings of the 4th All-Union Sympo- sium, Tbilisi, 5—7 October, 1983. Aquatic Oli- gochaeta. [Translated from Russian. Canadian Translation of Fisheries and Aquatic Sciences #5091, 1984. 143 pp.]. Sokolskaja, N. L. 1968. The aquatic oligochaete fau- na of the Soviet Far East.—Nauchnye doklady vysshei shkoly. Biolohiyi Nauki 6:21. [in So- kolskaja, 1972]. 1972. New data on the fauna of aquatic Oli- gochaeta from Kamchatka.—Sbornik Trudov Gosudarstvennogo Zoologicheskogo Muzeya 12: 74-90. [Translated from Russian. Translation bureau, Multilingual Services Division, De- partment of the Secretary of State of Canada]. 1973. New Tubificidae species from Kam- chatka and new findings of oligochaetes in the basins of the Peninsula.—Byulletyen Moskov- skogo Obshchestva Ispytatelei Prirody. Otdel Biologicheskii 78:54—67. [Translated from Rus- sian. Translation bureau, Multilingual Services Division, Department of the Secretary of State of Canada Fisheries and Marine Services Trans- lation series #3144. 30 pp.]. Stephenson, J. 1930. The Oligochaeta. Oxford: Clar- endon Press. xiv + 978 pp. Timm, T. 1981. On the origin and evolution of aquat- ic Oligochaeta.— Eesti nsv teaduste Akadeemia Toimetised, 30. Kdide Bioloogia 3:174-181. Vejdovsky, F. 1879. Beitrage zur Vergleichenden. I. Monographie der Enchytraeiden. Prague. 62 pp. Wachs, B. 1967. Die Oligochaeten-Fauna der Fliess- gewasser unter besonderer Beriicksichtigung der Beziehungen zwischen der Tubificiden-Besied- lung und dem Substrat.—Archiv fir Hydro- biologie 63:310-386. Zar, J. H. 1974. Biostatistical analysis. New Jersey: Prentice Hall. xiv + 620 pp. Department of Invertebrate Zoology, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, Canada M5S 2C6. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 429-434 CARIDEAN SHRIMPS OF THE GULF OF CALIFORNIA. V. NEW RECORDS OF SPECIES BELONGING TO THE SUBFAMILY PONTONITNAE (CRUSTACEA: DECAPODA: PALAEMONIDAE) Rubén Rios Abstract. —Extensions in the known distribution of three species of ponto- niine shrimps are presented. The first known male of Neopontonides dentiger has been collected in Bahia Kino, Sonora, some 800 km northwestward from its previously recorded range. Two other species of Panamic pontoniine shrimps are reported for the first time from the Gulf of California: Periclimenaeus hancocki and P. spinosus were found in two different sponges in Bahia Con- cepcion, B.C.S. The male holotype is the only previous record of the latter species. Diagnoses and illustrations of all three species are provided. Resumen. —Se incrementa el rango de distribuciOn conocida de tres cama- rones pontoninos. Las especies panamicas Periclimenaeus hancocki y P. spi- nosus han sido colectadas por vez primera en el Golfo de California: en Bahia Concepcion dentro de dos esponjas diferentes. De P. spinosus solo se conocia el holotipo macho. En Bahia Kino, Sonora, fué hallado el primer macho cono- cido de Neopontonides dentiger; ésto significa un incremento de unos 800 km hacia el NW, en el area de su distribucion conocida. Se incluyen diagnosis e ilustraciones de las tres especies mencionadas. Because of their small size and typical associations, pontoniine shrimps are very likely to go unnoticed. Detailed examina- tion of sponges collected in Bahia Concep- cion (25°40'N, 11°50'W) has provided sev- eral specimens of two _ species of Periclimenaeus. Another specimen avail- - able through the kindness of Prof. Marco Escalante, from the Escuela de Ciencias del Mar (University of Sinaloa at Mazatlan) has proved to be the first male specimen known of Neopontonides dentiger. All specimens are deposited in the reference collection of the Centro de Investigacion Cientifica y Ed- ucacion Superior de Ensenada (CICESE). Neopontonides dentiger Holthuis, 1951 Fig. 1 Neopontonides dentiger Holthuis, 1951:193, pl. 61; 1952:18.—Hendrickx, Wicksten, and van der Heiden, 1983:70.—Wick- sten, 1983:20. Material examined.—One male (cara- pace length 1.6 mm), Bahia Kino, Sonora (28°45' N, 111°55’W), 4 m, macroalgae, coll. M. Escalante, 1983. Previous records.—Cabo San Francisco (Ecuador); Mazatlan and Punta Piaxtla in Sinaloa (Mexico). Diagnosis. —Carapace smooth, only an- tennal spines present. Rostrum laterally ex- panded in proximal portion and laterally compressed in distal part; 9 to 11 dorsal teeth, ventral margin unarmed. Basal seg- ment of antennular peduncle armed with spine on anterolateral border. Stylocerite sharply pointed, reaching middle of basal segment. Scaphocerite well developed, with lateral spine overreached by lamella. Sec- ond and third maxillipeds without exopods, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 430 ‘gurljnoseu xipuodde puv eusojur xtpusdde jo [Iejop ‘oureg ‘py :podosjd puosas “7 ‘podogyd ising “y ‘ejoyo ‘oureg ‘f ‘podiosod ysis ‘[ :podi[ixeur puossg “YH ‘podiyjixew piyy “D ‘TAjoep ‘aures ‘4 ‘podotssed pry. y, “gq Spodyyrxews IsILj °C ‘MIA [BSIOP “UOTSAI IOLIOJUY “D -MIA [PI9}e| ‘uowulopqe pue aoedeies ‘g ‘MoIA [esUdA ‘spodoresod puooses pur isIY JO oseg “VW :o[eU ‘ad7zjUap sapiuojuodoan *{ “3ty H VOLUME 99, NUMBER 3 widely separated at base. Ventral two-point- ed process between coxae of first and second pereiopods. Third pereiopod with dactylus simple, without basal protuberance. Second pleopod with appendix masculina reaching beyond middle of appendix interna and armed with 3 apical setae. Remarks. —Holthuis (1951) described N. dentiger from a single incomplete ovigerous female. Apparently, the specimens exam- ined by Wicksten (1983) are those reported in Hendrickx, Wicksten and van der Heiden (1983): an ovigerous female and an unde- termined specimen. This is the first report of a male of the species, unfortunately lack- ing both second pereiopods. The most con- spicuous differences with the holotype de- scription are the number of rostral teeth and the ventral mesial spiniform bifurcate pro- cess between pereiopods I and II. Neopon- tonides dentiger was previously collected in mud and rocks (Holthuis 1951) and on sand- rock and muddy bottoms with gorgonians on stones (Hendrickx, Wicksten and van der Heiden 1983, Wicksten 1983). It is now re- corded along with macroalgae. Proper host identity remains to be established. Periclimenaeus hancocki Holthuis, 1951 Fig. 2 Periclimenaeus hancocki Holthuis, 1951:97, pl. 29.—Abele, 1975:70, Fig. 28; 1976: 270—Wicksten, 1983:8. Material examined.—One male (c.l. 2.9 mm), one female (c.l. 3.5 mm), one oviger- ous female (c.1. 3.5 mm); Bahia Concepcion, Baja California Sur, 2.3 m, sponge; coll. H. Licon, 10 May 1981. Previous records.—Bahia Pinas, Pacific coast of Panama; Malpelo Island. Diagnosis. —Carapace smooth, only an- tennal spines present. Rostrum laterally compressed, longer than eye, reaching be- yond first antennular peduncle; 4 or 5 dorsal teeth, ventral margin entire. Scaphocerite shorter than antennular peduncle; antero- lateral spine overreached by lamella. First 431 pereiopods with fingers a little shorter than palm. Second pereiopods unequal, larger with fingers almost half as long as palm; smaller with palm almost 3 times as long as fingers. Larger second pereiopod with fin- gers obliquely inserted on palm. Posterior pereiopods with dactylus strongly biunguic- ulate. Second pelopod of male with appen- dix masculina not reaching middle of ap- pendix interna, armed with 2 apical and 2 subapical setae. Remarks.—The holotype of this species came from soft mud (Holthuis 1951). Abele (1976) found it in what he called ‘‘Pocillo- pora damicornis habitat’’. Specimens from Bahia Concepcion were taken from a non- calcareous sponge with acicular spicules. A male and a female were living together, which suggests that P. hancocki occurs in pairs as in other sponge associate species. Wicksten (1983) included P. hancocki in a list of four species that may occur in the Gulf of California because of its presence on the southwestern Mexican coast. Unfor- tunately I did not find any precise records from that region. Abele (1975) refers to the holotype as being a male, while Holthuis (1951) in the original description mentions a female as the only specimen examined. Dr. Raymond B. Manning of the Smithson- ian Institution kindly examined that spec- imen and found that Prof. Holthuis was cor- rect. Periclimenaeus spinosus Holthuis, 1951 Fig. 3 Periclimenaeus spinosus Holthuis 1951:113, pl. 35. Material examined.—One male (c.l. 1.8 mm), one ovigerous female (c.l. 2.3 mm), Bahia Concepcion, B.C.S.; sponge, coll H. Licon, 10 May 1981. Previous records.—South Viradores Is- lands off Pacific coast of Costa Rica. Diagnosis.—Carapace smooth with su- praorbital and antennal spines. Rostrum laterally compressed and directed slightly Fig. 2. Periclimenaeus hancocki A-J, ovigerous female: A, Carapace, lateral view; B, Anterior region, dorsal view; C, Chela of first pereiopod; D, Left second pereiopod, outer view; E, Same, inner aspect; F, Right second pereiopod, outer view; G, Dactyl of third pereiopod; H, Telson; I, Mandible; J, Second pleopod with egg attached. K-L, male: K, Second pleopod; L, Same, detail of appendix masculina. VOLUME 99, NUMBER 3 433 Fig. 3. Periclimenaeus spinosus A—M, ovigerous female: A, Carapace, lateral view; B, Anterior region, dorsal view; C, Fourth pereiopod; D, Same, finger; E, Telson; F, Right second pereiopod, outer aspect; G, Same, fingers; H, Same, dorsal view; I, Left second pereiopod, outer aspect; J, Same, dorsal view; K, First pereiopod; L, Same, chela; M, Second pleopod with egg attached. N—O, male: N, First pleopod; O, Second pleopod. 434 downwards, armed with 5 or 6 dorsal and one subterminal ventral teeth. Basal seg- ment of antennular peduncle with a spine on anterolateral margin. Stylocerite sharp- pointed, reaching middle of basal segment. Scaphocerite well developed, with lateral spine overreaching lamella. First pereio- pods with slender chela and small round tooth on proximal margin of fixed finger. Second pereiopods different in size but sub- equal in shape. Posterior legs with dactylus biunguiculated and accessory spinules on ventral margin. Accessory spinules also present on posterior margin of propodus and merus of third leg and propodus of fourth and fifth legs. Second pleopod of male with appendix masculina reaching beyond mid- dle of appendix interna and furnished with single long apical seta. Remarks. —Periclimenaeus spinosus was known only from the male holotype from Costa Rica. Specimens examined showed the following small differences with the de- scription of the holotype: third maxilliped does not reach middle of scaphocerite; sec- ond pereiopods with small spinules on pos- terior margin of merus and ischium; carpus of larger second pereiopod is about one fourth as long as chela. Periclimenaeus spi- nosus has been collected with corals (Hol- thuis 1951). Both specimens from Bahia Concepcion were drawn from conspicuous channels of a small sponge with a loosely reticular mesenchima. Acknowledgments Prof. Alberto Carvacho critically re- viewed the manuscript. Mme. Clara Yanez made the drawings. Dr. Raymond B. Man- ning from the Smithsonian Institution, checked the sexual identity of the holotype PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of Periclimenaeus hancocki and kindly re- viewed the final manuscript. Marco Esca- lante provided the specimen from Bahia Kino. Literature Cited Abele, L.G. 1975. The macruran decapod Crustacea of Malpelo Island. — Smithsonian Contributions to Zoology 176:69-85. . 1976. Comparative species composition and relative abundance of decapod crustaceans in marine habitats of Panama.— Marine Biology 38:263-278. Hendrickx, M. E., M. K. Wicksten, and A. M. van der Heiden. 1983. Studies of the coastal marine fauna of southern Sinaloa, Mexico. IV. Report on the caridean crustaceans. — Proceedings of the Biological Society of Washington 96:67-78. Holthuis, L. B. 1951. A general revision of the Pa- laemonidae (Crustacea Decapoda Natantia) of the Americas. I. The subfamilies Euryrhynchin- ae and Pontoniinae.— Occasional Papers of the Allan Hancock Foundation 1 1:1—332. . 1952. The Palaemonidae collected by the Si- boga and Snellius expeditions, with remarks on other species. II. Subfamily Pontoniinae.—Si- boga Expeditie 39a!°:1—253. 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. Division de Oceanologia, Centro de In- vestigacion Cientifica y de Educacion Su- perior de Ensenada (CICESE), Apartado Postal 2732, Ensenada, B. C., México. Note added in proof: Another record of Neopontonides dentiger has re- cently appeared: Heard, R. W. 1986. Pontoniine shrimps (Decapoda: Caridea: Palaemonidae) of the northwest Atlantic. I. The genus Neopontonides Hol- thuis, 1951, with the description of N. chacei, new species, and the erection of Pseuwdopontonides, new ge- nus, to receive N. principis Criales, 1980.—Journal of Crustacean Biology 6:471-484. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 435-436 TWO HITHERTO UNNAMED POPULATIONS OF AECHMOPHORUS (AVES: PODICIPITIDAE) Robert W. Dickerman Abstract.—The Mexican nesting populations of Aechmophorus occidentalis are named A. o. ephemeralis, and the larger birds of the northern populations of Aechmophorus clarkii in the United States and Canada are named A. c. transitionalis. The Western Grebe (Aechmophorus oc- cidentalis) was named as Podiceps occiden- talis by Lawrence (1858:892, 894), from “Pacific coast from Washington Territory to California” (=Fort Steilacoom, Wash- ington, fide American Ornithologists’ Union Checklist, 6th Edition [1983]). On the fol- lowing page Lawrence described Podiceps clarkii. These were dark-backed, dark-lored and pale-backed, white-lored forms, re- spectively. Two of the three specimens of clarkii listed by Lawrence were wintering birds from California of the large, pale morph that occurs throughout the range of the species in the United States and Canada. Deignan (1961:7-8) pointed out that the third specimen, from Laguna Santa Maria, Chihuahua, should be considered the type of P. clarkii, and Dickerman (1963) redun- dantly so restricted the type locality. The critical differences between the two morphs are the yellow-orange bill and white face of the pale morph, and darker bill and black face of the dark morph (Storer 1965); but all color characters are highly variable (Sto- rer and Nuechterlein 1985). Podiceps clarkii was considered a syn- onym of 4. occidentalis until 1963, when Dickerman (1963) demonstrated the lack in overlap in wing chord measurements (and very little overlap in bill length) between northern and Mexican populations, and rec- ognized clarkii as a small subspecies with pale and dark morphs as in nominate oc- cidentalis. Mayr and Short (1970:29) over- looked or ignored the 1963 paper, stating “no races are recognized.” Later, Dickerman (1973) further charac- terized the small Mexican subspecies. He also pointed out that there was a degree of geographical segregation of the two morphs, in that more specimens from the central pla- teau and northern Mexico were of the pale type, while more from the western and southern portions of the plateau were of the dark morph. However, the trenchant char- acteristic of the Mexican versus northern populations is the small size of Mexican birds. In the ensuing years, studies of the breed- ing behavior of northern populations (see review in Storer and Nuechterlein 1985) in- dicated than an apparently significant de- gree of assortative mating occurs between the two morphs. Based on this, the Amer- ican Ornithologists’ Union Committee on Classification and Nomenclature (AOU 1985) concluded that the two color morphs should be recognized by the specific names Aechmophorus occidentalis and A. clarkii. This decision was made in the absence of studies of the critical Mexican populations, and is contrary to recommendations pub- lished earlier in the same volume of the Auk (Storer and Nuechterlein 1985). Further, starch gel electrophoresis revealed no sig- nificant difference between the two color morphs in preliminary studies (Ratti 1979). In view of the AOU Committee’s deci- sion, and in view of the great size disparity 436 between birds of the northern and southern populations, it now becomes necessary to name the large pale northern population and the small dark southern population. These may now be known as: Aechmophorus occidentalis ephemeralis, new subspecies Holotype.—Adult male, American Mu- seum of Natural History 708,559, collected at Laguan de San Pedro Lagunillas, Nayarit, Mexico, 25 May 1962 by Robert W. Dick- erman; field number RWD 10650. Addi- tional data: with a large young; weight 1050 gms.; pectoral muscles 17 and 16.3 gms.; moderately to rather fat; extent 851 mm.; wing chord 179 mm., culmen (from anterior edge of nostril) 56.7 mm., tarsus 70.4 mm. Diagnosis.—Small, inseparable in size from the co-existing population of A. clarkii, but dark-lored as in nominate occidentalis. Distribution. —Specimens recorded to date from Nayarit, western Jalisco, Puebla and Guererro (Dickerman 1973). Aechmophorus clarkii transitionalis, new subspecies Holotype.—Adult male, National Mu- seum of Natural History 158016, collected at Silver Lake, Lake County, Oregon, 22 Jun 1897 by Vernon Bailey; original number 20. Additional data: wing cord 194 mm., cul- men (from anterior edge of nostril) 63.4 mm., tarsus 77.5 mm. Diagnosis. —Large, similar in size to nominate A. occidentalis, but coloration pale as in A. clarkii. Distribution. —Sympatric with A. o. oc- cidentalis in western North America, north of the Mexican border, becoming rarer northwards. Discussion. —For detailed size compari- son of nominate 4. o. occidentalis and A. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON clarkii transitionalis see Storer and Nuech- terlein (1985). Acknowledgments I thank M. Ralph Browning for his selec- tion and measurement of the type of A. clarkii transitionalis, and Kenneth C. Parkes and Robert W. Storer for their comments on the manuscript. Literature Cited American Ornithologists’ Union. 1983. American Ornithologists’ Union Checklist of North Amer- ican Birds, 6th Edition. xxix + 877 pp., Amer- ican Ornithologists’ Union, Washington, D.C. 1985. Thirty-fifth Supplement to the Amer- ican Ornithologists’ Union Checklist of North American Birds.—Auk 102:680-686. Deignan, H. G. 1961. Type specimens in the United States National Museum.—Bulletin of the United States National Museum 221:1-718. Dickerman, R. W. 1963. The Grebe Aechmophorus occidentalis clarkii as a nesting bird of the Mex- ican Plateau.—Condor 65:66-67. 1973. Further notes on the Western Grebe in Mexico.—Condor 75:131-132. Lawrence, G. H. 1858. Jn Baird, Cassin, and Law- rence, Report of Explorations and Surveys, Railroad Mississippi Pacific 9:894—-895. Mayr, E., and L. Short. 1970. Species taxa of North American Birds.—Publication of the Nuttall Ornithological Club 9:1—127. Ratti, J. T. 1979. Reproductive separation and iso- lating mechanisms between sympatric dark- and light-phased Western Grebes. — Auk 96:573-586. Storer, R. 1965. The color phases of the Western Grebe.— Living Bird 6:59-63. ——, and G. L. Nuechterlein. 1985. An analysis of plumage and morphological characters of the two forms of the Western Grebe (Aechmopho- rus).—Auk 102:102-119. Cornell University Medical College, 1300 York Avenue, New York, New York, 10021, and American Museum of Natural History, 79th Street at Central Park West, New York, New York, 10024. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 437-443 THE STATUS OF THE CALLIANASSID GENUS CALLICHIRUS STIMPSON, 1866 (CRUSTACEA: DECAPODA: THALASSINIDEA) Raymond B. Manning and Darryl L. Felder Abstract.—The callianassid genus Callichirus Stimpson is redefined and re- stricted to its type species, Callianassa major Say, 1818, and three other species. The callianassid genus Callichirus was es- tablished by Stimpson (1866:47) with Cal- lianassa major Say, 1818, from the south- eastern United States, as its type species. Since then, Callichirus has been treated in various ways, including: (1) as a subgenus of Callianassa Leach, 1814 (e.g., by Bor- radaile 1903; De Man 1928a, b; Schmitt 1935; Rodrigues 1971); (2) as a distinct ge- nus, comprising numerous species formerly assigned to Callianassa (e.g., by de Saint Laurent 1974, Le Loeuff and Intés 1974, De Saint Laurent and Le Loeuff 1979); or (3), as a synonym of Callianassa (e.g., by Biffar 1971, Poore and Griffin 1979). We disagree with all of these interpretations for reasons discussed below, and we give here charac- ters that distinguish Callichirus from all other genera of the Callianassidae. Part of the problem has been that most authors, at least up to de Man (1928a:30), had seen no material of Callianassa major, and that until very recently the only avail- able illustrations of the species were pho- tographs (Hay and Shore 1918:pl. 29, fig. 10; De Man 1928a:fig. 14c, d). De Man had examined and figured two chelipeds (1928a: fig. 14-14b), but had studied no complete specimens. The species was not well figured until specimens from Brazil were studied by S. A. Rodrigues (1971:192, figs. 1-20). In his original account of Callichirus, Stimpson (1866:47) noted that the cheliped was elongate (“carpus and hand of the great- er cheliped very long’’), that the uropodal endopod was narrow (‘‘very narrow, almost styliform”’), and that the telson was short and broad, with a posterior emargination. We suggest that these features and the dis- tinctive ornamentation of the third to fifth abdominal somites (Figs. 1f, 2e, 3e) are di- agnostic for the genus Callichirus, and will serve to distinguish its members from all other callianassids. De Saint Laurent (1974:513) divided the family Callianassidae into two subfamilies, the Callianideinae and the Callianassinae. She assigned eight genera, including Calli- chirus Stimpson, 1866, to the Callianassi- nae. Refinement of diagnoses for most of these genera was attempted later by Le Loeuff and Intés (1974) and by de Saint Lau- rent and Le Loeuff (1979). However, not all of the genera recognized by de Saint Laurent have been accepted by other workers. Poore (1975:205) comment- ed on the limitations of de Saint Laurent’s genera, and Poore and Griffin (1979:245) noted, ““We have had some difficulty in plac- ing some of the Australian callianassid species in the new genera erected by Saint Laurent (1973) . . . and therefore prefer not to follow her arrangement. The exception is her new genus Gourretia which forms a rel- atively homogeneous group of species clear- ly distinct from the remainder of the fam- ily.” Rabalais, Holt, and Flint (1981:98) abandoned use of de Saint Laurent’s genera because “‘the distinctions proposed. . . were not clear cut for Western Hemisphere cal- lianassids.”” Williams (1984:180) com- mented, ““According to de Saint Laurent, the 438 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Callichirus major (Say), male, tl 77 mm, Indian River, Florida (USNM 228086): a, Front; b, Third maxilliped; c, Major cheliped; d, Minor cheliped; e, Sixth abdominal somite, telson, and uropod (stippled areas on uropod show areas of dense setation). Male, tl 56 mm, Indian River, Florida (USNM 228087): f, Third to fifth abdominal somites (top is anterior). VOLUME 99, NUMBER 3 known species from the western Atlantic formerly placed in Callianassa belong in Callichirus. The differences between Calli- anassa and Callichirus depend upon mouthparts and seem somewhat overlap- ping. For the time being, therefore, I am retaining the well-known name Callianassa for these species, awaiting further research which may place de Saint Laurent’s system on a firmer basis.” We redefine Callichirus, as follows: Genus Callichirus Stimpson, 1866 Callichirus Stimpson, 1866:47. Definition.—Ocular peduncles elongate, either in precorneal region or as postcorneal spine. Major chela of adult male greatly en- larged, with 3 distal segments equal to 2 or more times middorsal carapace length. Third to fifth abdominal somites with dis- tinctive ornamentation, visible dorsally, each somite with midlateral patch of setae. Telson much shorter than uropods, much wider than long, with medial emargination. Uropodal endopods strap-shaped, very slender, 2.5 to 4 times longer than wide, with setae most concentrated in apical patch. Type species.—Callianassa major Say, 1818, by original designation and mono- typy. Included species. —Four, as follows: 1. Callichirus major (Say, 1818:238) (Fig. 1), from the southeastern United States, Gulf of Mexico, and Brazil. 2. Callichirus islagrande (Schmitt, 1935: 5) (Fig. 2), from the northern and western Gulf of Mexico. 3. Callichirus seilacheri (Bott, 1955:47) (Fig. 3), from El Salvador. 4. Callichirus adamas (Kensley, 1974: 266), from West Africa and South Africa. Remarks.—We believe that these four species comprise a distinctive group of species within the Callianassidae recogniz- able as a distinct genus, Callichirus. The 439 ornamentation of the third to fifth abdom- inal somites alone will distinguish these species from all other species assigned to Callianassa or other genera of the Calli- anassidae. Preliminary anatomical studies of the third to fifth abdominal somites suggest that the dorsal sculpture and setation may com- prise the exoskeletal components of an ab- dominal glandular apparatus, perhaps re- lated to cementing of burrow walls. If so, this apparatus would represent a unique an- atomical specialization among the thalas- sinideans. A previous study (Thompson 1972) has demonstrated that cement pro- duction is centered in a hindgut gland in Upogebia pugettensis (Dana) and in glands at the bases of mouthparts in Callianassa californiensis Dana. Additional histological studies are planned to resolve the function of these abdominal glands in Callichirus. Our corrected definition of Callichirus also is corroborated in part by recent ecological and physiological studies. Osmoregulatory capacities and habitat preferences in C. ma- jor and C. islagrande are very similar; nei- ther species can osmoregulate and both can tolerate limited reductions in salinity (Feld- er 1978). This is in striking contrast to the case in Callianassa jamaicensis Schmitt and Callianassa kraussi Stebbing, both of which are euryhaline, hyperosmotically regulate at and tolerate low salinities, withstand abrupt changes of salinity, and show marked reg- ulation of volume (Forbes 1974, Felder 1978). But the four above species were treat- ed as congeneric in the broadly defined Cal- lichirus of de Saint Laurent and Le Loeuff (1979). In fact, the tendency for osmocon- formation and limited tolerance of reduced salinity, characteristic of Callichirus major and Callichirus islagrande, bears the great- est similarity to those features in Calli- anassa californiensis, Callianassa filholi A. Milne Edwards, and Callianassa affinis Holmes (see Gross 1957, Thompson and Pritchard 1969), Felder 1978), none of which 440 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Callichirus islagrande (Schmitt), male holotype, TL 87 mm, Grand Isle, Louisiana (USNM 69369): a, Front; b, Third maxilliped; c, Major cheliped; d, Minor cheliped; e, Third to fifth abdominal somites (top is anterior); f, Sixth abdominal somite, telson, and uropod (stippled areas on uropod show areas of dense setation). VOLUME 99, NUMBER 3 441 Fig. 3. Callichirus seilacheri (Bott), male holotype, tl 85 mm, Los Blancos, El Salvador (Forschungsinstitut Senckenberg no. 2184): a, Front; b, Third maxilliped; c, Right first cheliped; d, Left first cheliped; e, Third to fifth abdominal segments (top is anterior); f, Sixth abdominal segment, telson, and uropod (stippled areas on uropod show areas of dense setation). Note that both first chelipeds are of minor form. 442 is assignable to Callichirus. We believe that this evidence from the physiology of these organisms suggests an artificial and heter- ogeneous grouping within Callichirus sensu de Saint Laurent and Le Loeuff (1979). We provide here original sketches (Figs. 1-3) of three of the species that we assign to Callichirus. We have not seen material of Callichirus adamas, but the accounts and figures of this species provided by Kensley (1974:266, figs. 1, 2), who first pointed out the resemblance between C. adamas, C. major, and C. islagrande, and by de Saint Laurent and Le Loeuff (1979:67, figs. 14f, l6a, 17a, 19f, 20e-g, 23f-i), clearly identify C. adamas as a member of Callichirus. The characteristic ornamentation of the abdo- men in C. adamas is clearly shown by de Saint Laurent and Le Loeuff (1979) in their figure 17a. Of the species assigned to or described in Callichirus by Le Loeuff and Intés (1974) and de Saint Laurent and Le Loeuff (1979), only those four listed above can be assigned to Callichirus sensu stricto. We consider all of the other species assigned to Callichirus by those authors to be members of Calli- anassa. Acknowledgments We thank Austin B. Williams, System- atics Laboratory, National Marine Fisheries Service, for reviewing this paper. M. Tur- kay, Forschungsinstitut Senckenberg, Frankfurt-am-Main, West Germany, loaned us the holotype of Callianassa seilacheri. The figures were prepared by Lilly King Manning. This paper is contribution num- ber 163 from the Smithsonian Marine Sta- tion, Fort Pierce, Florida, and contribution number 2 from the Center for Crustacean Research, University of Southwestern Lou- isiana. Literature Cited Biffar, Thomas A. 1971. The genus Callianassa (Crustacea, Decapoda, Thalassinidea) in south PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Florida, with keys to the western Atlantic species. — Bulletin of Marine Science 21(3):637— TS: Borradaile, L. A. 1903. On the classification of the Thalassinidea.—Annals and Magazine of Nat- ural History (7)12:534—-551. Bott, Richard. 1955. Litorale Dekapoden, ausser Uca. Dekapoden (Crustacea) aus El Salvador, 2.— Senckenbergiana biologica 36(1/2):45-70. Felder, D. L. 1978. Osmotic and ionic regulation in several western Atlantic Callianassidae (Crus- tacea, Decapoda, Thalassinidea).— Biological Bulletin 154:409-—429. Forbes, A. T. 1974. Osmotic and ionic regulation in Callianassa kraussi Stebbing (Crustacea: De- capoda: Thalassinidea).—Journal of Experi- mental Marine Biology and Ecology 16:301-311. Gross, W. 1957. An analysis of response to osmotic stress in selected Crustacea.— Biological Bulle- tin 112:43-62. Hay, W. P., and C. A. Shore. 1918. The decapod crustaceans of Beaufort, N. C., and the sur- rounding region. — Bulletin of the United States Bureau of Fisheries, 35(for 1915 and 1916):369— 475, pls. 25-39. Kensley, Brian. 1974. The genus Callianassa (Crus- tacea, Decapoda, Thalassinidea) from the west coast of South Africa with a key to the South African species.—Annals of the South African Museum 62(8):265-278. Le Loeuff, P., and A. Intés. 1974. Les Thalassinidea (Crustacea, Decapoda) du Golfe de Guinée. Sys- tématique-Ecologie.—Cahiers O.R.S.T.O.M., série Océanographie 12(1):17-69. ; De Man, J. G. 1928a. A contribution to the knowl- edge of twenty-two species and three varieties of the genus Callianassa Leach.—Capita Zoo- logica 2(6):1—56, plates 1-12. . 1928b. The Thalassinidae and Callianassidae collected by the Siboga-Expedition with some remarks on the Laomediidae. The Decapoda of the Siboga-Expedition, Part 7.—Siboga Expe- dition, Monographie 39a6:1—187, pls. 1—20. Poore, Gary C. B. 1975. Systematics and distribution of Callianassa (Crustacea, Decapoda, Macrura) from Port Phillip Bay, Australia, with descrip- tions of two new species. — Pacific Science 29(2): 197-209. , and D. J. G. Griffin. 1979. The Thalassinidea (Crustacea: Decapoda) of Australia.— Records of the Australian Museum 32(6):217-321. Rabalais, Nancy N., Scott A. Holt, and R. Warren Flint. 1981. Mud shrimps (Crustacea, Deca- poda, Thalassinidea) of the northwestern Gulf of Mexico.—Bulletin of Marine Science 31(1): 96-115. ; Rodrigues, Sergio de Almeida. 1971. Mud shrimps VOLUME 99, NUMBER 3 of the genus Callianassa Leach from the Bra- zilian coast.— Arquivos de Zoologia, Sao Paulo 20(3):191—223. De Saint Laurent, Michéle. 1974. Sur la systématique et la phylogénie des Thalassinidea: Définition des familles des Callianassidae et des Upoge- biidae et diagnose de cinq genres nouveaux (Crustacea Decapoda).—Compte rendus Aca- demie des Sciences, Paris (D)277[1973]:513- 516. , and P. Le Loeuff. 1979. Upogebiidae et Cal- lianassidae. Crustacés Décapodes Thalassinid- ea, 1. Campagnes de la Calypso au large des cotes Atlantiques africaines (1956 et 1959)(suite), 22.—Résultats scientifiques des campagnes de la Calypso 11:29-101. Say, Thomas. 1818. An account of the Crustacea of the United States [Part 5].—Journal of the Acad- emy of Natural Sciences of Philadelphia 1(part 2, number 1):235-253. Schmitt, Waldo L. 1935. Mud shrimps of the Atlantic coast of North America.— Smithsonian Miscel- laneous Collections 93(2):1—21, pls. 1-4. Stimpson, William. 1866. Descriptions of new genera and species of macrurous Crustacea from the coasts of North America.— Proceedings of the Chicago Academy of Sciences 1:46-48. 443 Thompson, L. C., and A. W. Pritchard. 1969. Os- moregulatory capacities of Callianassa and Upogebia (Crustacea: Thalassinidea).— Biolog- ical Bulletin 136:114—129. Thompson, R. K. 1972. Functional morphology of the hind-gut gland of Upogebia pugettensis (Crustacea, Thalassinidea), and its role in bur- row construction. Unpublished Ph.D. Thesis, Department of Zoology, University of Califor- nia, Berkeley. Williams, Austin B. 1984. Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida: xviii + 550 pages. Smithson- ian Institution Press, Washington, D.C. (RBM) Department of Invertebrate Zo- ology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560; (DLF) Department of Biology and Center for Crustacean Research, Uni- versity of Southwestern Louisiana, Lafay- ette, Louisiana 70504. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 444-451 A NEW SCALE-WORM COMMENSAL WITH DEEP-SEA MUSSELS IN THE SEEP-SITES AT THE FLORIDA ESCARPMENT IN THE EASTERN GULF OF MEXICO (POLYCHAETA: POLYNOIDAE: BRANCHIPOLY NOINAE) Marian H. Pettibone Abstract.—Some polynoid polychaetes found living commensally in mantle cavities of deep-sea mussels at the seep-sites near the Florida Escarpment, are described as a new species of Branchipolynoe, B. seepensis. They show some differences from the previously described B. symmytilida, commensal with deep-sea mussels in the Galapagos hydrothermal vents. Among the abundant biological com- munities recently discovered in the abyssal eastern Gulf of Mexico near hypersaline seep-sites were large dense mats of mussels (Florida Escarpment Cruise Participants 1984, Paull et al. 1984, Turner and Lutz 1984, Denoux et al. 1985, Hecker 1985). They were collected in March 1984 during four dives of the submersible A/vin in 3266- 3270 meters (26°02'N, 84°55'’W). The mus- sels were found in dark iron-sulfide sedi- ments at near-ambient temperatures in an area where dense hypersaline fluids seep out onto the sea floor. The communities were similar in many respects to the prolific com- munities found around the Galapagos hy- drothermal vents and elsewhere in the Pa- cific. Some polynoid polychaetes were found living commensally in the mantle cavities of these mussels. Six specimens were given to me for study by Dr. Barbara Hecker: five adults removed from the mussels by Dr. Ruth D. Turner, and a juvenile collected from washings of a clump of mussels. It was of particular interest to compare these polynoids, commensal with the un- named, non-hydrothermal, large, Gulf of Mexico mussels (to be described and named by Ruth Turner and Barry Wilson), with the polynoids commensal with the large mus- sels in the Galapagos hydrothermal vents, recently described under the name Bathy- modiolus thermophilus by Kenk and Wilson (1985). The Galapagos polynoids were re- ferred earlier to a new genus and species, Branchipolynoe symmytilida, in a new subfamily of the Polynoidae, Branchipoly- noinae (Pettibone 1984). The Gulf of Mex- ico commensals are referred herein to a new species of Branchipolynoe, B. seepensis. The types are deposited in the National Museum of Natural History, Smithsonian Institution (USNM). Branchipolynoinae Pettibone, 1984 Branchipolynoe Pettibone, 1984 Type species.—Branchipolynoe symmy- tilida Pettibone, 1984. The generic diagnosis is emended to in- clude Branchipolynoe seepensis, new species, commensal with deep-sea mussels in the Gulf of Mexico. Diagnosis, emended.—Branchiae small to well developed, arborescent, on all para- podia from segment 2 or 3. Prostomium bilobed, with anterior lobes subtriangular or rounded, with or without minute frontal filaments; median antenna with ceratophore indistinct, style short; palps short to long. First or tentacular segment visible dorsally as short ring or fused to prostomium. Par- apodia subbiramous or biramous; notopo- VOLUME 99, NUMBER 3 dia small, nodular or conical with projecting acicular process; notosetae few (0-3) or moderate in number (5-20). Neuropodia with or without truncate lobes enclosing conical acicular lobes; neurosetae very nu- merous, upper ones much stouter or only slightly stouter than lower ones. Commen- sal with deep-sea mussels at hydrothermal vents or hypersaline seep-sites. Key to the Species of Branchipolynoe 1. Elytra small, leaving greater part of dorsum uncovered (Pettibone 1984, figs. 1, 2A). Arborescent branchiae beginning on segment 2 (Pettibone 1984, figs. 2A, 3A). Anterior lobes of prostomium with minute frontal filaments (Pettibone 1984, fig. 2A). First or tentacular segment visible dorsally as small ring (Pettibone 1984, fig. 2A). Parapodia subbira- mous, with notopodia small and 0 to few notosetae (Pettibone 1984, fig. 3F). Neuropodia with truncate lobes enclosing conical acicular lobes (Pet- tibone 1984, fig. 3F). Upper neuro- setae much stouter than lower neu- rosetae (Pettibone 1984, fig. 3I, K). Ventral papillae on segments 11 and 12 short (Pettibone 1984, fig. 4C). Commensal with deep-sea mussels in Galapagos hydrothermal vents . wel ee B. symmytilida Pettibone, 1984 — Elytra large, leaving middle third of body uncovered (Fig. 1A). Arbores- cent branchiae beginning on segment 3 (Fig. 2B). Anterior lobes of prosto- mium without frontal filaments (Fig. 1A). Tentacular segment fused to prostomium, not visible dorsally as separate ring (Fig. 1A). Parapodia biramous, with notopodia large, con- ical, with moderate number of no- tosetae (Fig. 2C, D). Neuropodia without truncate lobes enclosing conical acicular lobes (Fig. 2D). Up- per neurosetae only slightly stouter than lower neurosetae (Fig. 2H, I). Ventral papillae on segments 11 and 12 long (Fig. 1D). Commensal with deep-sea mussels in hypersaline seeps in Gulf of Mexico ie oC Ce cry Branchipolynoe seepensis, new species Figs. 1-4 Material. —Gulf of Mexico, from dives of Alvin in 1984, associated with mussels near hypersaline seeps at base of Florida Escarp- ment, in dark iron-sulfide sediment: Dive 1343, 9 Mar, 26°02'24’N, 84°54'48’W, 3266 m, from washings of clump of mussels, ju- venile paratype (USNM 98782).—Dive 1344, 12 Mar, 26°02’N, 84°54’ 48” W, 3270 m, from mantle cavities of mussels, holo- type (USNM 98779), 2 paratypes (USNM 98780).—Dive 1346, same data, 2 para- types (USNM 98791). Measurements. —Holotype (USNM 98779) 31 mm long, 13 mm wide, including setae, with 21 segments. Two adult para- types (USNM 98780) 27-28 mm long, 11- 13 mm wide, with 21 segments. Two small- er adult paratypes (USNM 98781) 12-16 mm long, 5—7 mm wide, with 21 segments. Juvenile paratype (USNM 98782) 8 mm long, 3.5 mm wide, with 18 segments, last 2 very small. Description. — Body short, spindle-shaped, slightly tapered anteriorly and posteriorly, flattened ventrally and strongly arched dor- sally. Fully-developed worms having 21 segments, first achaetous, with 10 pairs of elytra attached on large, low elytrophores on segments 2, 4,5, 7,9, 11, 13, 15, 17 and 19. Elytra (Figs. 1A, E-G, 3A, B, 4C, D) moderately large, oval, covering lateral thirds of body on larger adults and nearly covering body on juvenile. Elytra oval, opaque, smooth, soft, and attached near middle on first pair and eccentrically at- tached near their anterior borders on fol- lowing elytra, sometimes showing branched “veins.” Dorsal cirri on non-elytra-bearing 446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON G Fig. 1. Branchipolynoe seepensis, holotype (USNM 98779): A, Dorsal view of anterior end, right first elytron removed; B, Ventral view of anterior end; C, Dorsal view of posterior end showing segments 19 to 21 and pygidium, tenth elytra of segment 19 removed; D, Ventral view of right side of segments 10 to 13 showing elongate ventral papillae on segments 11 and 12; E, Right first elytron from segment 2, with “veins”’ indicated; F, Right fifth elytron from segment 9; G, Right tenth elytron from segment 19. Scales = 1.0 mm for A-C; 2.0 mm for D-G. segments having short cylindrical cirro- phores and short styles with slender tips, extending to about tips of neurosetae (Figs. 2B, E, 3D, 4E). Dorsal tubercles on cirri- gerous segments indistinct. Branchiae well developed, arborescent, with rather short terminal filaments (Fig. 2B—F, J). Branchiae emerging from body in two short trunks, lower smaller group attached dorsal to bases of notopodia or cirrophores of dorsal cirri and upper larger one attached to lateral bas- es of elytrophores or dorsal tubercles. — Fig. 2. Branchipolynoe seepensis, holotype (USNM 98779): A, Right elytragerous parapodium from segment 2, anterior view, acicula dotted; B, Right cirrigerous parapodium from segment 3, posterior view, cirrophore of dorsal cirrus hidden from view; C, Right cirrigerous parapodium from segment 8, posterior view; D, Right elytragerous parapodium from segment 9, anterior view, acicula dotted, branchiae cut off; E, Lower branchia from same; F, Upper branchia from same; G, Three neurosetae from same, with detail of tips; H, Upper neuroseta from same, with detail of tips; I, Lower neuroseta from same, with detail of tips; J, Right elytragerous parapodium from segment 15, anterior view, acicula. Scales = 1.0 mm for A-F; 0.1 mm for G-I. 447 VOLUME 99, NUMBER 3 448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ee : SI =} aH fe H Fig. 3. Branchipolynoe seepensis, smaller adult paratype of 12 x 5 mm (USNM 98781): A, First elytron, with “veins” indicated; B, Middle elytron; C, Ventral view of right side of segments 11 and 12, showing elongate ventral papillae; D, Right middle cirrigerous parapodium, posterior view; E, Right middle elytragerous para- podium, anterior view, acicula dotted; F, Three notosetae from same, with detail of tips; G, Upper neuroseta from same, with detail of tips; H, Two lower neurosetae from same, with detail of tips. Scales = 0.5 mm for A, B; 1.0 mm for C; 0.5 mm for D, E; 0.1 for F—H. Branches of branchiae somewhat flattened and spread anteriorly and posteriorly be- tween parapodia. Branchiae beginning on segment 3 as 2 small groups (Fig. 2B), be- coming larger with more numerous branch- es in middle region (Fig. 2C), and smaller posteriorly (Figs. 1C, 2J). Branchiae smaller on smaller adults (Fig. 3D, E), very small and incompletely developed on juvenile (Fig. 4E, F). VOLUME 99, NUMBER 3 449 Fig. 4. Branchipolynoe seepensis, juvenile paratype, 8 < 3.5 mm, (USNM 98782): A, Dorsal view of anterior end, median antenna and palps missing; B, Ventral view of anterior end; C, Right first elytron; D, Right middle elytron; E, Right cirrigerous parapodium, posterior view; F, Right elytragerous parapodium, anterior view, acicula dotted; G, Notoseta from same, with detail of tip; H, Upper neuroseta from same, with detail of tip; I, Two lower neurosetae from same, with detail of tips. Scales = 1.0 mm for A, B; 0.5 mm for C—F; 0.1 mm for G-I. Prostomium (Figs. 1A, B, 4A, B) oval, bilobed, with rounded anterior lobes lack- ing frontal filaments. Median antenna, lack- ing distinct ceratophore, inserted in anterior notch; short, conical, with slender tip; ven- trolateral palps rather short, thick, smooth, and tapered, extending beyond prostomi- um; lateral antennae and eyes lacking. First or tentacular segment fused to prostomium and not distinct dorsally; tentaculophores, lateral to prostomium, achaetous, short and bulbous, each with pair of short cirriform tentacular cirri with slender tips, about as long as palps (Figs. 1A, B, 4A, B). Second or buccal segment bearing first pair of large elytrophores and elytra, biramous parapo- dia with smaller conical notopodia with up to 9 notosetae, and larger rounded neuro- podia with numerous slender neurosetae and lower buccal cirri attached basally, similar to tentacular cirri (Figs. 1A, B, 2A, 4A, B). Mouth ventral with lateral and posterior lips, enclosed in tentacular and buccal segments (Figs. 1B, 4B). Thick muscular pharynx ex- tended on one of paratypes (USNM 98780), with opening pulled tightly together; when cut open, 5 pairs of small, delicate, oval papillae around opening and 2 pairs of small jaws, lacking denticulated bases (similar to that figured for B. symmytilida, Pettibone 1984, fig. 2F, G), revealed. Biramous parapodia having short conical 450 notopodia and long conical neuropodia (Figs. 2B—D, J, 3D, E, 4E, F). Notosetae moderate in number (up to 20 on adults, up to 5 on juvenile), short to long, extending nearly to tips of neuropodia; stouter than neurosetae, acicular, smooth or slightly roughened distally, with tips rounded (Figs. 2G, 3F, 4G). Presetal acicular lobes of neu- ropodia slightly longer than postsetal lobes. Neurosetae very numerous, long, slender, separated by slight gap into upper supra- acicular and lower subacicular groups. Up- per neurosetae slightly stouter than lower ones, minutely serrated, with blunt tips (Figs. 2H, 3G, 4H), some with terminal bulb (Fig. 2H, upper right). Lower neurosetae more slender and having hooked tips with some longer denticles (Figs. 21, 3H, 41). Ventral cirri, attached to middle of neuropodia, short, tapered, and not reaching tips of neu- ropodia (Fig. 2C, D). Pygidium rather large and bulbous, with terminal anus and pair of stout anal cirri basally fused; enclosed in parapodia of last two posterior segments (20, 21) and their long dorsal cirri (Fig. 1C). Distinct segmen- tal or nephridial papillae not present, except for 2 pairs of long ventral papillae on seg- ments 11 and 12, projecting posteriorly (Fig. 1D). Papillae shorter on smaller adult para- type (Fig. 3C), absent on juvenile. Etymology. —Neo-Latin adjective see- pensis, formed from seep, in reference to the distribution of the species in the hypersaline seeps in the Gulf of Mexico. Remarks. — According to the preliminary report by Denoux et al. (1985), the che- mosynthetic vent-type taxa found in hy- persaline, cold water, hydrocarbon seeps in the Gulf of Mexico are associated with an abundance of non-chemosynthetic species and this enhanced community diversity suggests the possibility that the oil seep communities may be the progenitors of those at the deep-water vent sites and, thus, more primitive. There are some indications that the Gulf of Mexico commensal, B. seepen- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sis, might very well be more primitive than the Galapagos commensal, B. symmytilida. In B. seepensis, the parapodia are biramous, with large notopodia and a moderate num- ber of notosetae; the neuropodia have the form of conical acicular lobes, following the usual pattern in the Polynoidae. In B. sym- mytilida, the parapodia are subbiramous, with the notopodia small, nodular, and the notosetae few to none; the neuropodia have an extra truncate lobe enclosing the acicular lobe, thus, the development of a new struc- ture. There is greater development of the arborescent branchiae in B. symmytilida than in B. seepensis. Acknowledgments I wish to thank the Florida Escarpment Cruise Participants and especially Drs. Bar- bara Hecker and Ruth Turner for the ma- terial on which this study is based. The manuscript benefited from the reviews of Drs. Meredith L. Jones, Nancy J. Maciolek, and James A. Blake. Literature Cited Denoux, G. J., M. C. Kennicutt, IJ, R. R. Bidigare, J. M. Brooks, R. R. Fay, M. L. Jones, and R. D. Turner. 1985. Description of a hydrocarbon- seep community on the Louisiana slope.— American Zoologist 25(4):10A, Abstract 35. Florida Escarpment Cruise Participants. 1984. The seeps find at the Florida Escarpment. — Oceanus 27(3):32-33. Hecker, B. 1985. Fauna from a cold sulfur-seep in the Gulf of Mexico: Comparison with hydro- thermal vent communities and evolutionary implications. — Bulletin of the Biological Society of Washington 6:465-473. Kenk, V. C., and B. R. Wilson. 1985. A new mussel (Bivalvia, Mytilidae) from hydrothermal vents in the Galapagos nft zone.— Malacologia 26(1— 2):253-271. Paull, C. K., B. Hecker, R. Commeau, R. P. Freeman- Lynde, C. Neumann, W. P. Corso, S. Golubic, J. E. Hook, E. Sikes, and J. Curray. 1984. Bi- ological communities at the Florida Escarpment resemble hydrothermal vent taxa.—Science 226: 965-967. VOLUME 99, NUMBER 3 Pettibone, M. H. 1984. A new scale-worm commen- sal with deep-sea mussels on the Galapagos hy- drothermal vent (Polychaeta: Polynoidae).— Proceedings of the Biological Society of Wash- ington 97(1):226—239. Turner, R. D., and R. A. Lutz. 1984. Growth and distribution of mollusks at deep-sea vents and seeps. — Oceanus 27(3):54-62. 451 Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 452-463 A NEOTYPE DESIGNATION FOR PETROLISTHES TOMENTOSUS (DANA), AND DESCRIPTION OF PETROLISTHES HETEROCHROUS, NEW SPECIES, FROM THE MARIANA ISLANDS (ANOMURA: PORCELLANIDAE) Roy K. Kropp Abstract. — Petrolisthes tomentosus (Dana) is determined to be a senior syn- onym of P. penicillatus (Heller) and a neotype of the former is designated. Petrolisthes pubescens Stimpson, a species previously confused with P. tomen- tosus, is redescribed and compared to the latter. The two can be distinguished primarily by the presence of elevated, setae-topped bosses on the carapace of P. tomentosus as compared to the flat carapace of P. pubescens. A new species, P. heterochrous, is also described. For over 100 years carcinologists have confused several Indo-Pacific species of Pe- trolisthes characterized by having a very to- mentose carapace and a pronounced fringe of plumose setae on the posterior margin of the manus. Lewinsohn (1969, 1979) has un- raveled part of the problem. He (1969) rec- ognized P. pubescens Stimpson as distinct from a group that includes P. tomentosus (Dana 1852), P. penicillatus (Heller 1862), and P. villosus (Richters 1880). Moreover, he (1979) summarized the confusion over possible synonymy ofthe latter three species. Briefly, Ortmann (1897) considered P. vil- losus to be a junior synonym of P. penicil- latus. However, since then, the matter of synonymy between P. tomentosus and P. penicillatus has been unresolved. Lewin- sohn (1979) examined a syntype of P. pen- icillatus and described it thoroughly. He (as well as Haig 1983) speculated that P. pen- icillatus might be a junior synonym of P. tomentosus, whose type has been lost. How- ever, Lewinsohn declined to select a neo- type for the latter because the type locality for P. tomentosus is the Tuamotu Archi- pelago, and he did not have specimens from French Polynesia. Here I select and describe a neotype for P. tomentosus based on a specimen from Tahiti and compare it to P. penicillatus and P. pubescens. Also, I describe a new species of Petrolisthes from a collection of porcel- lanids from the Mariana Islands. Materials and Methods Most of the material on which this report is based was collected by the author in 1979-— 1981 and 1984. Additional material was ob- tained from the collection of the National Museum of Natural History (USNM), Smithsonian Institution, Washington, D.C. Some specimens have been deposited in the Allan Hancock Foundation, Los Angeles, California, and the B. P. Bishop Museum, Honolulu, Hawaii. Specimens were measured to the nearest 0.1 mm using an ocular micrometer on a Wild M-5 microscope. Measurements are given in the text as carapace length (along the midline) < carapace width (at the great- est dimension) for the type specimens and the largest male, the largest female, and the smallest ovigerous female. Other abbrevi- ations used in the text are: OV = ovigerous females; juv. = juveniles; m = meters. Setal terminology is based on Kunze and Anderson (1979). Setal measurements were VOLUME 99, NUMBER 3 made, but were extremely variable within and among individuals so are not included. The figures were prepared with the aid of a drawing tube on a Wild M-5 or M-8 mi- croscope. Systematic Account Petrolisthes tomentosus (Dana) Fig. 1 Porcellana tomentosa Dana, 1852:420; 1855: pl. 26, fig. 10. Petrolisthes tomentosus. —Stimpson, 1858: 227 (list); 1907:182 (list).—Kropp et al., 1981:40 (list). Porcellana penicillata Heller, 1862:523; 1865:79. Porcellana villosa Richters, 1880:160, pl. 17, figs. 11-12. Petrolisthes villosus.—Ortmann, 1892:259 (key), 264; 1894:27 (key).— Ward, 1942: 3. Petrolisthes penicillatus. —Ortmann, 1897: 287 (key), 288.—Lenz, 1910:566.—Mi- yake, 1942:347, text-figs. 11-12; 1943:55, 83, text-fig. 18.—Haig, 1964:368; 1966: 42; 1983:283.—Lewinsohn, 1979:45.— Yang, 1983:2, pl. 2. Petrolisthes villosus?— Miers, 1884:559. Not Petrolisthes tomentosus. —Ortmann, 1892:259 (key), 264; 1894:27 (key); 1897: 287 (key), 288.—Grant and McCulloch, 1906:39.—Mliyake, 1943:55 (key), 85, text-figs. 19-21; 1956:310.—Haig, 1964: 364; 1966:47 (key).—Nakasone and Mi- yake, 1971:4. = Petrolisthes pubescens Stimpson. Type data.—Much of the type material collected by Dana during the United States Exploring Expedition (1838-1842) was studied by Stimpson in Chicago. This ma- terial, along with Stimpson’s North Pacific Exploring Expedition (1853-1856) was de- stroyed in the 1871 Chicago fire (Deiss and Manning 1981). Although some of Dana’s types have since been rediscovered (Deiss and Manning 1981), I have been unable to locate the type of Porcellana tomentosa. I 453 have searched the USNM, Yale Peabody Museum, and the Philadelphia Academy of Sciences without success. Dr. R. B. Man- ning has compiled an unpublished list of Dana material at the Museum of Compar- ative Zoology (Harvard) and the major Eu- ropean museums. A type of P. tomentosa is not among them. It is very probable that the type for this species has been destroyed. I located a disarticulated, but complete, specimen of Dana’s species in the USNM. It is from Tahiti, close to the original type locality (Tuamotu) and is herein designated as the neotype. Neotype.—TAHITI: Arue; under dead coral; 1963; Coll. H. A. Rehder; 2 (OV); 5.3 X 4.8 mm; USNM 190776. Other material examined. —OKINAWA: W of Onna, S of Manzamo Precipice; reef flat; 20 May 1984; Coll. R. B. Manning et al.; 1 2 (OV). MARIANA ISLANDS: PA- GAN: Puntan Lagona; on dead coral; 1 m; 13 Mar 1981; 1 2. Bandeera Peninsula; in- tertidal; 11 Mar 1981; 1 °. Palapala Bay; subtidal under rock; 1.5 m; 14 Mar 1981; 1 6. Liyan; on base of Pocillopora setchelli; 1.5 m; 9 Mar 1981; 1 2(OV). Apaan Santati; reef flat under rock; <1 m; 10 Mar 1981; 1 6, 1 29. SAIPAN: Tanapag Barrier Reef; on dead coral; 6 m; 20 Nov 1980; 1 2. Agingan Pt.; reef margin on coralline algae; <1 m; 19 Nov 1980; 3 6 (1 with bopyrid isopod). GUAM: Tanguisson Pt.; reef front on dead branching coral; 11 m; 25 May 1984; 1 é. Piti Bay; outer reef flat S of Camel Rock; intertidal and subtidal on dead coral; 0—1.5 m; 13 Jun 1980; 5 6, 2 2 (OV). Luminao; reef flat on dead coral or under rocks; 1 m; 28 Mar 1980, 5 Apr 1980; 4 6, 6 2 (2 OV). Neye Is.; on Pocillopora sp.; 29 Jan 1981; Coll. R. H. Randall; 1 2. Toguan Bay; at reef margin on reef rock; <1 m; 14 Feb 1984; Coll. J. H. Dominguez; 1 6. Pago Bay; reef flat on coralline algae or intertidal under rock; 0-1 m; 18 May 1981, 16 Feb 1984; 2 6. FIJI: Nananu-i-Ra; intertidal pool; 3 Jun 1980; 1 2. Measurements. —Largest male—5.4 xX 454 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mente) i) AGN Ney Vaca ACG N WO Fig. 1. Petrolisthes tomentosus (Dana), neotype 2: a, Carapace and chelipeds; b, c, d, Right walking legs 1, 2, 3; e, Right antennule base; f, Right eye (dorsal view); g, Right eye (lateral view); h, Carapace (lateral view). Scale: 3 mm (a); 2 mm (b-e); 1 mm (f, g); 4 mm (h). [Surface setae on carapace and right cheliped not shown.] VOLUME 99, NUMBER 3 5.2 mm; largest female—5.9 ) o..c:.25 03 eee 11 11. Posterior end tail-like; anal aper- ture surrounded by many cycles of plates; sole comparatively small, rectangular; midventral ambula- crum complete [Norway to New England, 0-400 m] ...P. phantapus (Strussenfeldt, 1765) — Tail absent; few periproctal cycles of plates; sole large, oval; midven- tral ambulacrum incomplete [Nor- way to New England, 0-1800+ m] .. P. fabricii (Diiben and Koren, 1846) Acknowledgments We thank the following colleagues for the invitation to participate in the SEAMAP program and for logistic support: Gilmore Pellegrin, National Marine Fisheries Ser- vice, Pascagoula, who served as field party chief; William G. Lyons, Florida Depart- VOLUME 99, NUMBER 3 ment of Natural Resources, who served as chief scientist on the Florida leg (Florida East Coast Benthic Mapping Study). We thank Haydee Hernandez, Florida Institute of Technology, and Stephen E. Stancyk, University of South Carolina, for translat- ing parts of Tommasi (1971). Patricia Lin- ley, Harbor Branch Foundation, Inc. (HBF), assisted with SEM photography. The ho- lotype was illustrated by Charissa Lounibus. Loan of type material from the MCZ was made possible by Robert M. Woollacott. This paper is HBF Contribution no. 530. Literature Cited Deichmann, E. 1930. The holothurians of the western part of the Atlantic Ocean. — Bulletin of the Mu- seum of Comparative Zoology 71(3):41—226, 24 pls. . 1941. The holothurians collected by the Ve- lero III during the years 1932 to 1938. Part I, Dendrochirota.— Allan Hancock Pacific Expe- ditions 8(3):61-194, pls. 10-30. Diiben, M. W., and J. Koren. 1846. Ofversigt af Skandinaviens Echinodermer.— Kungliga Svenska Vetenskaps-Akademiens Handlingar, for Ar 1844:229-328, pls. 6-11. Mortensen, T. 1927. Handbook of the echinoderms of the British Isles. 471 pp., 269 figs. Oxford University Press. Ostergren, H. 1904. Eine neue Psolus-Art, Ps. val- vatus. —Zoologische Anzeiger 27:659-662. Pawson, D. L. 1967. The psolid holothurian genus Lissothuria. —Proceedings of the United States National Museum 122(3592):1-17, 5 figs. Note added in proof: 485 1968. A new psolid sea cucumber from the Virgin Islands.— Proceedings of the Biological Society of Washington 81:347—350, 1 fig. , and J. F. Valentine. 1981. Psolidium pros- tratum, new species, from off the east coast of the U.S.A. (Echinodermata: Holothuroidea).— Proceedings of the Biological Society of Wash- ington 94(2):450—454, 2 figs. Pourtalés, L. F. de. 1868. Contribution to the fauna of the Gulf Stream at great depths. — Bulletin of the Museum of Comparative Zoology 1(7):121- 142. Strussenfeldt, A.M. von. 1765. Beskrifnig pa et Syo- krak, Hafs-Spoke kalladt.—Kungliga Svenska Vetenskaps-Akademiens Handlingar 26:256- 266, pl. 10. Théel, H. 1886. Report on the Holothurioidea. Re- ports on the results of dredging, under the su- pervision of Alexander Agassiz, in the Gulf of Mexico (1877-78), in the Caribbean Sea (1879- 80), and along the eastern coast of the United States during the summer of 1880, by the U.S. Coast Survey Steamer “Blake.” — Bulletin of the Museum of Comparative Zoology 13(1):1—21, 1 pl. Tommasi, L. R. 1971. Equinodermes do Brasil. I. Sobre algumas espécies novas e outras pouco conhecidas, para o Brasil. — Boletim do Instituto Oceanografico de Sao Paulo 20:1-21, 23 figs. (JEM) Indian River Coastal Zone Mu- seum, Harbor Branch Foundation, Inc., RR 1, Box 196, Ft. Pierce, Florida 33450. (RLT) Department of Biological Sciences, Florida Institute of Technology, 150 W. University Blvd., Melbourne, Florida 32901. We thank Patricia A. Lindsay and John B. Wise, South Carolina Department of Wildlife and Marine Re- sources, for bringing to our attention 4 more specimens of Psolus pawsoni: IRCZM 71:341, 7-11 mm TL, R/ V LADY LISA, Golden Crab Project, Year II, 19 May 1986, vicinity of 31°21’N, 79°34’W (east of Brunswick, Georgia), ~450 m, rocket grab, clinging to rock. A specimen of P. operculatus (IRCZM 71:342) from the same rock represents a considerable range extention for this species. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 486-488 ON THE ELEVATION OF THE STEPHANOPHYLLIA SUBGENUS LETEPSAMMIA TO GENERIC RANK (COELENTERATA: SCLERACTINIA: MICRABACIIDAE) Joan Murrell Owens Abstract.—The differentiating macrostructural and microstructural charac- teristics of the subgenus Letepsammia Yabe and Eguchi are described, and the subgenus is officially elevated to the taxonomic rank of genus. Letepsammia differs most significantly from Stephanophyllia and other genera of the Micra- baciidae in its numerous coarse perforations in all septa, and its slender, wavy, and irregularly spaced trabeculae. Fossil specimens of Letepsammia are found in the Plio-Pleistocene of Japan, while living specimens have been retrieved from depths of less than 100 meters to nearly 1000 meters over a wide area of the Pacific and Indian Oceans. Letepsammia was described as a subge- nus by Yabe and Eguchi in 1932, primarily on the basis of the well-developed, spongy columella in S. formosissima Moseley, the type species. Although they acknowledged differences in septal perforations between Letepsammia and the other subgenera of Stephanophyllia, they evidently did not consider these septal differences to be of ge- neric significance. Similarly, Vaughan and Wells (1943) and Wells (1956) considered Letepsammia to be synonymous with Ste- phanophyllia. More recent authors, most notably Squires (1965, 1967), Keller (1977), Cairns (1982), and Owens (1984a, b, 1986), have tacitly accepted the septal and colu- mellar differences as worthy of generic sep- aration, and have informally recognized the subgenus as Letepsammia. A review of the literature, however, reveals that Letepsam- mia has never been officially elevated to generic status. This paper, therefore, is in- tended to do so. Order SCLERACTINIA Bourne, 1900 Suborder FUNGIIDA Duncan, 1881 Superfamily FUNGIOIDEA Vaughan and Wells, 1943 Family MICRABACIIDAE Vaughan, 1905 Genus LETEPSAMMIA Yabe and Eguchi, 1952 Stephanophyllia (Letepsammia) Yabe and Eguchi, 1932:58.—Wells, 1933:51.— Yabe and Eguchi, 1934:280—281, figs. 1-3. Letepsammia Vaughan and Wells, 1943: 145.—Wells, 1956:F390.—Squires, 1965: 878; 1967:505-508.—Keller, 1977:41- 43.—Cairns, 1982:9.—Owens, 1984a: 519-521, fig. 4; 1984b:88, fig. 4; 1986: 248-255. Diagnosis. —Corallum large, loosely built, with highly perforated septa and wall, well- developed deltas, and prominent marginal shelf. Wall horizontal or nearly so, very thin, with septa alternating in position with cos- tae. Septa slender, with varied, complex dentation; interspaces wider than septa. Ve- preculae inconspicuous or absent. Costae slender, serrated on outer edge, and sepa- rated by broad intercostal loculi interrupted by closely spaced, concentric rows of syn- apticulae. Columella broad, spongy, and elongate. Trabeculae thin, wavy, poorly or- ganized, ramified in some species along un- perforated growing edge; 10-20 in number. VOLUME 99, NUMBER 3 Low height : diameter ratio. Generally 120 septa. Occurrence. —Plio-Pleistocene, Ryukyu Islands; Recent, Japan Sea, Celebes Sea, Banda Sea, Coral Sea, Philippine Islands, Hawaiian Islands, Australia, New Zealand, East African Coast. 50-960 meters. Type species.—Stephanophyllia formo- sissima Moseley, 1881, by monotypy, by Yabe and Eguchi, 1932. Deposited in the British Museum (Natural History). Taxa included. — Letepsammia formosis- sima Moseley, 1881, and Letepsammia su- perstes Ortmann, 1888. Discussion. —When Yabe and Eguchi (1932:58) established Letepsammia, they based it solely on their type of the subgenus, S. (L.) formosissima, and described the sub- genus as follows: “Skeletal elements very delicate. Col- umellar fossa distinct, deep; columella spongy, well developed. Septa usually and horizontal wall always coarsely perforat- ed, being a network of slender trabeculae. Pores of the horizontal wall circular, coarser, and are easily distinguished from the smaller, elongate or oval perforations in the other subgenera .. .” Later, Yabe and Eguchi (1934:280-28 1) included in this subgenus a fossil form from Japan, S. (L.) japonica nov. Although this form was subsequently determined to be a fossil representative of the living species S. superstes, it nonetheless proved the validity of perforated septa and spongy columella as identifying characters of their subgenus. Re- cently I acquired from the National Mu- seum of Natural History a new, undescribed species that also has these characters. No- where else among the genera and subgenera of the Micrabaciidae are septa of both lower and higher cycles so highly and conspicu- ously perforate. On this basis, if on no other, recent authors of studies on the Micraba- ciidae have informally recognized Lete- psammia as a genus. 487 The diagnosis given earlier in this paper is intended to expand and emend Yabe and Eguchi’s original description of their sub- genus so as to incorporate other common characters of the species of that group that seem to be of generic significance. Letepsammia, at present, contains two described species: L. formosissima Moseley, 1881. Recent, Celebes, Banda, and Coral Seas, Philip- pine and Hawaiian Islands. L. superstes Ortmann, 1888. Plio-Pleisto- cene, Ryukyu Islands; Recent, Japan Sea. Squires (1967:505) alluded to six species of letepsammid corals found in the Pacific and Indian Oceans. Undoubtedly, this number included the two described species, and probably also the new, undescribed species from the National Museum of Nat- ural History; the remaining species to which he refers have not, to date, been either de- scribed or verified. Although Letepsammia was originally as- signed as a subgenus of Stephanophyliia, it bears only a superficial resemblance to that genus. The corallum in Letepsammia is rel- atively large and loosely built, with all septa highly perforated and adorned with prom- inent, complex dentation. In Stephano- pDhyllia, the corallum is small-to-medium and generally quite compact; perforations are present, but they are restricted to the lower portions of higher cycle septa and are often sparsely and irregularly distributed. Septal edges are noticeably dentate, but the dentation is of the simple acute or lacerate type. The basal wall in Letepsammia is thin and generally projects as a prominent shelf, whereas it forms a rather tight rim in Ste- phanophyllia with little or no marginal shelf. In each case, the columella is elongate, but it is very porous in Letepsammia, lamellar in Stephanophyliia. Septal microstructure also differs in the two genera. Trabeculae in Stephanophyllia are regularly spaced in a well-organized fan 488 system, with ends of the sclerodermites pro- jecting laterally to form vepreculae along the flanks of each septum. In Letepsammia, the trabeculae are thin, wavy, irregularly spaced in a very loose fan, and are reduced in number. The sclerodermites rarely pro- ject beyond the plane of the septum, but when they do, they form only rudimentary vepreculae. Superficially, Letepsammia does resem- ble Stephanophyllia in that both possess true deltas, dentate septa, and septal perfora- tions to some degree. However, the differ- ences between them far outweigh their sim- ilarities. These differences, moreover, are fundamental enough to warrant official el- evation of Letepsammia to the rank of ge- nus. Acknowledgments I wish to thank Richard Duffield, Howard University, for suggesting to me the need for this paper; Stephen D. Cairns, Smith- sonian Institution, for carefully reading my manuscript and providing helpful com- ments; and my husband, Frank, for editing and proofreading this manuscript. Literature Cited Cairns, S.D. 1982. Antarctic and Subantarctic Scler- actinia.— Antarctic Research Series 5, 34:1-74, 18 pls. Keller, N. B. 1977. New species of genus Leptopenus and some peculiarity of the deep-sea aherma- typic corals.— Trudy Instituta Okeanologii 108: 37-43, 3 figs., 1 pl. Owens, J. M. 1984a. Microstructural changes in the Micrabaciidae and their ecologic and taxonomic PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON implications.— Palaeontographica Americana 54:519-522, 5 figs. 1984b. Evolutionary trends in the Micra- baciidae: An argument in favor of preadapta- tion.—Geologos 2(1):87—93, 5 figs. 1986. Rhombopsammia, a new genus of the family Micrabaciidae (Coelenterata: Scleracti- nia).— Proceedings of the Biological Society of Washington 99(2):248-256, 3 figs. Squires, D. F. 1965. A new record for Leptopenus, a rare deep-water coral.— Nature 207:878-879, 1 fig. 1967. The evolution of the deep-sea coral family Micrabaciidae.—Studies in Tropical Oceanography, 5 (Proceedings of the Interna- tional Conference on Tropical Oceanography): 502-510. Vaughan, T. W., and J. W. Wells. 1943. Revision of the suborders, families, and genera of the Scler- actinia.— Geological Society of America Special Papers Number 44, 363 pp., 40 pls., 32 text figs. Wells, J. W. 1933. Corals of the Cretaceous of the Atlantic and Gulf coastal plains and western interior of the United States.—Bulletins of American Paleontology 18(67):1—206, 16 pls., 4 text figs. . 1956. Scleractinia. In R. C. Moore, ed., Trea- tise on invertebrate paleontology. Part F. Coe- lenterata, pp. 328-444, figs. 223-339. Geolog- ical Society of America, Lawrence, University of Kansas Press. Yabe, H., and M. Eguchi. 1932. Some Recent and fossil corals of the genus Stephanophyllia H. Michelin from Japan.—Tohoku Imperial Uni- versity Scientific Reports, series 2, Geology 15: 55-64, 2 pls., 3 text figs. 1934. Probable generic identity of Stephan- ophyllia Michelin and Micrabacia M. Edwards and J. Haime.—Imperial Academy of Japan, Proceedings 10(5):278-281, 5 figs. Department of Geology and Geography, Howard University, Washington, D.C. 20059. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 489-492 THE TAXONOMIC STATUS OF GLOSSOPHAGA MORENOI MARTINEZ AND VILLA, 1938 (MAMMALIA: CHIROPTERA: PHYLLOSTOMIDAE) Alfred L. Gardner Abstract.—A review of the nomenclatural history of Mexican Glossophaga, the identity of the neotype of G. morenoi, and the content of the species described as G. mexicana leads to the conclusion that G. morenoi is the correct name for the species currently known as G. mexicana. The taxonomy and nomenclature of Mex- ican species of the genus Glossophaga have a complex history. Confusion has resulted from the misallocation of Monophyllus leachii Gray, 1844; the apparent loss of the type and paratypes of Glossophaga morenoi Martinez and Villa, 1938; and the recently confirmed presence of four species of Glos- sophaga in Mexico (Webster and Jones 1980) instead of only one, the ubiquitous G. soricina (Pallas, 1766). Based on Miller’s (1913) revision of Glos- sophaga, G. soricina leachii (Gray, 1844) was the name applied to all North American mainland populations and G. soricina mu- tica Merriam, 1898, to the Tres Marias Is- lands populations. In 1938, Martinez and Villa described G. morenoi based on ma- terial from Xiutepec, Morelos. Davis (1944) described G. soricina alticola from 13 km northeast of Tlaxcala, Tlaxcala, but did not mention G. morenoi. Villa (1953) considered G. soricina alti- cola to be a synonym of G. soricina mor- enol. In contrast, Davis and Russell (1954) continued to recognize G. soricina alticola, treating G. morenoi as a synonym of G. sor- icina leachii, the name by which all other Mexican mainland populations of Glos- sophaga were known. Hall and Kelson (1959:114) followed Davis and Russell (1954) but commented on their doubt con- cerning “the correct taxonomic arrange- ment of the central Mexican material.” Gardner (1962) described G. commissar- isi (type locality 10 km SE of Tonala, Chia- pas) based on material from Nayarit, Co- lima, and Chiapas, Mexico. This prompted Villa (1964) to review the names that had been applied to Mexican Glossophaga. He concluded that G. soricina alticola and G. commissarisi were junior synonyms of G. morenoi, which he again recognized as a species distinct from G. soricina. In addi- tion, Villa (1964:387) designated a neotype for G. morenoi because the original material had disappeared. Alvarez (1966) reported on the discovery of specimens of small mammals in the Mu- seo Nacional de Historia Natural that in- cluded some long missing types of bats and four specimens of Glossophaga, which he identified as G. soricina. Although admit- ting that there was no proof, Alvarez be- lieved the four Glossophaga were used by Martinez and Villa (1938) when they de- scribed G. morenoi. Alvarez stated that Vil- la’s (1964) designation of a neotype of G. morenoi was invalid, because Villa’s action did not conform with Article 75 of the In- ternational Code of Zoological Nomencla- ture (Code). Alvarez (1966) did not consider Villa’s work a revision of a genus because the description of the neotype did not per- mit its recognition and its characterization was not consistent with the original descrip- tion. Thus Alvarez concluded that G. mor- enoi was best considered a “‘nomen dubius.” 490 Baker (1967) did not use the name G. morenoi. Instead he used the names G. al- ticola and G. commissarisi, in addition to G. soricina, based on identifications by J. Knox Jones, Jr., for Mexican species. In their review of North American G/os- sophaga, Webster and Jones (1980) clarified relationships among the Middle American and Mexican species. They recognized four sympatric species in Mexico: G. soricina, G. leachii, G. commissarisi, and G. mexicana. The latter they described as new. They as- signed G. soricina alticola and G. morenoi to G. leachii, citing Villa’s (1953, 1964, 1967) argument that G. morenoi was an ear- lier name for G. soricina alticola. Therefore, this made G. morenoi a junior synonym of G. leachii. Webster and Jones (1980:5) also described G. soricina handleyi to provide a name for the subspecies that had been known for so many years as G. soricina leachii. Glossophaga soricina leachii (with G. mutica and G. morenoi as synonyms), G. alticola, and G. commissarisi were the three taxa recognized by Hall (1981) as occurring in Mexico. In 1982, Webster and Jones named and described the northwestern Mexican population of G. commissarisi as G. c. hespera. Urbano and Sanchez (1983) agreed with Webster and Jones’ (1980) decision to place G. morenoi in the synonymy of G. leachii. They incorrectly identified the neotype of G. morenoi as an example of G. soricina and reiterated Alvarez’s (1966) opinion that Vil- la (1964) had not conformed to the require- ments of Article 75 (sections a and c) of the Code when designating a neotype for G. morenol. Webster and Jones (1984a) proposed the name G. mexicana brevirostris for the northern population of G. mexicana. They also commented (p. 4), “It is of passing note that the specimen (UNAM 7383) designat- ed as the ‘neotype’ of Glossophaga morenoi by Villa-R. (1964) actually represents G. m. brevirostris.. . .”’ Inthe mammalian species account on G. /eachii, Webster and Jones PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (1984b) provided a key to the species, re- marked on the taxonomic history of G. leachii, and included G. morenoi in the syn- onymy of G. leachii. The latest published report concerning the systematics and natural history of Mexican Glossophaga is the mammalian species ac- count on G. mexicana by Webster and Jones (1985). This account contained no mention of G. morenoi. The argument advanced by Alvarez (1966) and Urbano and Sanchez (1983) that Villa’s (1964) designation of the neotype of G. morenoi was invalid because he did not conform with the requirements of Article 75 of the Code (Stoll et al. 1964) is not supported by the evidence. Villa’s review of the taxonomy of the Mexican species of Glossophaga known to him, although not a revision of a genus, is clearly “revisory”’ in the sense of Article 75(a) of the Code. Fur- thermore, Villa’s designation conforms with the requirements of Article 75(c) of the Code. In other words, although Villa’s (1964) taxonomic conclusions were wrong, his ac- tions were nomenclaturally correct. There- fore, the name G. morenoi has standing based on the neotype. The conspecificity of the neotype of G. morenoi and specimens identified by W. David Webster as G. mexicana has been independently confirmed by Oscar Sanchez and Rodrigo Medellin (pers. comm.). In the original description, G. morenoi was char- acterized as similar to G. soricina in dental formula and morphology of the upper and lower incisors. However, differences includ- ed larger size, absence of a tail, and unde- fined differences in the ear and nose leaf. The upper incisors of G. mexicana are more similar to those of G. soricina in size and procumbency than are those of either G. leachii or G. commissarisi, but the lower incisors of G. mexicana are distinctly more similar to those of G. leachii and G. com- missarisi. In Mexico, Glossophaga leachii (with G. alticola a synonym) averages larger than G. soricina in external measurements VOLUME 99, NUMBER 3 and in breadth of braincase whereas G. commissarisi averages smaller than G. sor- icina in all dimensions; however, G. mex- icana brevirostris is approximately the same size or slightly larger than G. soricina and G. mexicana mexicana is even larger (Web- ster 1983). The purported absence of a tail cannot be evaluated at this time and may represent an oversight on the part of Mar- tinez and Villa (1938). A factor that undoubtedly influenced Opinions concerning the taxonomy of Mex- ican Glossophaga was the number of species believed to be present in the country. The prevailing opinion up to the time Gardner (1962) described G. commissarisi was that there was one species (G. soricina) with one, two, or possibly three subspecies. When de- scribing G. commissarisi, Gardner (1962:4) recognized the presence of two additional species besides G. soricina. One represents G. leachii as understood today; the other was later described as G. mexicana (see Webster and Jones 1980:6). Nevertheless, Villa (1964, 1967) apparently believed that only two (G. soricina and G. morenoi) were present. It remained for Webster and Jones (1980) to sort out the taxa and to confirm the presence of four species in Mexico. Clearly, the neotype of G. morenoi rep- resents the same species described by Web- ster and Jones (1980) as G. mexicana. Vil- la’s (1964) designation of the neotype of G. morenoi conforms to the requisites of Ar- ticle 75 of the International Code of Zoo- logical Nomenclature. Therefore, G. mexi- cana is a junior synonym of G. morenoi, whose synonymy is described as follows: Glossophaga morenoi Martinez and Villa, 1938 Moreno’s long-tongued bat Glossophaga morenoi Martinez and Villa, 1938:347; type locality ““Xiutepec,’’ Mo- relos, Mexico. Type and paratypes lost (Villa 1953, 1964, 1967). Neotype des- ignated by Villa (1964:387); type locality 491 ““Cueva del Idolo, 1 Km. S. tequesqui- tengo 970 m., Estado de Morelos,”’ Mex- ico. Glossophaga mexicana Webster and Jones, 1980; type locality ““Rio GuamoOl, 34 mi. S (by Hwy. 190) La Ventosa Jct., Oaxaca, Mexico.” Glossophaga mexicana brevirostris Webster and Jones, 1984a:2; type locality ““6 mi. W Yautepec, 4500 ft., Morelos, México.” The northern subspecies distinguished by Webster and Jones (1984a) as G. mexicana brevirostris should be known as G. morenoi morenoi Martinez and Villa, 1938, and the southern populations as G. morenoi mexi- cana Webster and Jones, 1980. Literature Cited Alvarez, T. 1966. Redescubrimiento de algunos tipos de murcielagos Mexicanos que se suponian per- didos.— Revista de la Sociedad Mexicana de Historia Natural 27:191-198. Baker, R. J. 1967. Karyotypes of bats of the family Phyllostomidae and their taxonomic implica- tions.—The Southwestern Naturalist 12:407— 428. Davis, W. B. 1944. Notes on Mexican mammals. — Journal of Mammalogy 25:370—403. , and R. J. Russell. 1954. Aves y mamiferos del Estado de Morelos. — Revista de la Sociedad Mexicana de Historia Natural 14(1—-4):77-148. Gardner, A. L. 1962. A new bat of the genus Glos- sophaga from Mexico.—Contributions in Sci- ence, Los Angeles County Museum 54:1-7. Gray, J. E. 1844. Mammalia. Pp. 7-36, pls. 1-18, in R. B. Hinds, ed., The zoology of the voyage of H.M.S. Sulfur. Vol. 1. Hall, E.R. 1981. The mammals of North America. Second edition. John Wiley and Sons, New York, 1:xviii + 1-600 + 90. , and K. R. Kelson. 1959. The mammals of North America. The Ronald Press Company, New York, 1:xxx + 1-546 + 79. Martinez, L., and B. Villa R. 1938. Contribuciones al conocimiento de los murcielagos de Mexi- co.—Anales del Instituto de Biologia, México 9: 339-360. Merriam, C. H. 1898. Mammals of Tres Marias Is- lands, off western Mexico.— Proceedings of the Biological Society of Washington 12:13-19. Miller, G. S., Jr. 1913. Revision of the bats of the genus Glossophaga. — Proceedings of the United States National Museum 46:413-429. 492 Pallas, P. S. 1766. Miscellanea Zoologica. Hagae Comitum, xii + 224 pp. Stoll, N. R., R. P. Dollfus, J. Forest, N. D. Riley, C. W. Sabrosky, C. W. Wright, and R. V. Melville. 1964. International code of zoological nomen- clature. The International Commission on Zoo- logical Nomenclature, xx + 176 pp. Urbano, V. G., and O. Sanchez-H. 1983. Type spec- imens of mammals in the collection of the In- stitute of Biology, National University of Mex- ico.—Occasional Papers, The Museum, Texas Tech University 87:1—7. Villa R., B. 1953. Mamiferos silvestres del Valle de Mexico.—Anales del Instituto de Biologia, Mexico 23:269-492. 1964. Reflexiones acerca de la posiciOn tax- onomica de los murcielagos siricoteros de Mex- ico, genero Glossophaga.—Anales del Instituto de Biologia, México 34:381-391. 1967. Los murcielagos de Mexico. Instituto de Biologia, Universidad Nacional Aut6noma de México, xvi + 491 pp. Webster, W. D. 1983. Systematics and evolution of bats of the genus Glossophaga. Unpublished Ph.D. dissert., Texas Tech University, Lub- bock, ix + 332 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON , and J. K. Jones, Jr. 1980. Taxonomic and nomenclatorial notes on bats of the genus Glos- sophaga in North America, with description of a new species.—Occasional Papers, The Mu- seum, Texas Tech University 71:1—12. , and . 1982. Anew subspecies of Glos- sophaga commissarisi (Chiroptera: Phyllostom- idae) from western Mexico.—Occasional Pa- pers, The Museum, Texas Tech University 76: 1-6. , and . 1984a. Anewsubspecies of Glos- sophaga mexicana (Chiroptera: Phyllostomi- dae) from southern Mexico.—Occasional Pa- pers, The Museum, Texas Tech University 91: 1-5. , and 1984b. Glossophaga leachii.— Mammalian Species 226:1-3. , and . 1985. Glossophaga mexicana. — Mammalian Species 245:1-3. Biological Survey Section, Fish and Wild- life Service, National Museum of Natural History, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 493-501 TRINERVATE LEAVES, YELLOW FLOWERS, TAILED ANTHERS, AND POLLEN VARIATION IN DISTEPHANUS CASSINI (VERNONIEAE: ASTERACEAE) Harold Robinson and Brian Kahn Abstract. —The genus Distephanus Cassini is resurrected for a series of mostly western Indian Ocean and eastern African species of Vernonieae including the only members of the tribe having trinervate leaves and yellow flowers. The genus is characterized by distinct basal stylar nodes, sclerified basal appendages on the anther thecae, and a unique form of pollen. Twenty-six species are recognized in the genus, 24 being newly transferred, and additional possible members are listed. The broad concept of Vernonia suffers from most of the ills of a core genus, being excessively paraphyletic, distracting from more appropriate phyletic comparisons be- tween its parts and other genera of the tribe, and being defined primarily by what it is not rather than by what it is. Such concepts ultimately encourage shoddy taxonomy, and Vernonia is no exception, having come to include elements that do not even fit the broadest definition of the genus when ex- amined critically. It is by virtue of such lapses of careful study that the Paleotropical genus Diste- phanus has fallen into the synonymy of Ver- nonia. Distephanus is a genus with predom- inantly yellow flowers in a tribe that otherwise lacks them, and the genus in- cludes the only members of the tribe with trinervate leaves. Furthermore, the anther thecae of the genus are almost as promi- nently appendaged at their bases as those of the American genus Piptocarpha which is distinguished by that character. Distephanus was first described by Cas- sini (1817) on the basis of a single species, Conyza populifolia Lam., from Mauritius. Cassini (1819) distinguished the genus by its pappus, described as 10 short outer squa- mellate scales alternating with 10 longer in- ner linear-squamellate scales. His descrip- tion mentioned the yellow-flowered heads, and a microfiche of the specimen in the Jus- sieu herbarium shows that the poplar-like leaves are trinervate. The claimed pappus distinction seems to have been ignored in subsequent literature as Distephanus has fallen into the synonymy of Vernonia, and the pappus was not mentioned by Humbert (1960) who related the yellow-flowered Ver- noniae of Madagascar to the genus. The ge- nus name has been transferred to sectional status by Bentham and Hooker (1873) for the single species, and to subsectional status by Jones (1981) for mostly unrelated species of Malayasia. With these latter exceptions, the name Distephanus has been ignored in recent literature. It is the African members of the same group that were later segregated from Ver- nonia as the genus Gongrothamnus Steetz, a genus that was transferred to the Seneci- oneae by Bentham and Hooker (1873) on the basis of its yellow flowers and trinervate leaves. Gongrothamnus was reduced to syn- onymy under Vernonia by Hoffmann (1890- 1894) at the same time that he named a new genus Newtonia in the same Senecioneae on the basis of still another species of the group from Angola. The name Newtonia was a 494 later homonym and a new name, Antunesia was provided by Hoffmann in 1893. Hoff- mann (1902) ultimately corrected his error, recognizing Gongrothamnus at generic level and placing Antunesia in its synonymy. Brown (1909) briefly summarized some of the work in Gongrothamnus, but reduced it again to synonymy under Vernonia, being unaware of the importance of the trinervate leaves, and knowing no other character ex- cept the yellow flowers. Almost all recent students of the Verno- nieae have treated the yellow-flowered species as part of Vernonia. Wild (1978) placed the yellow-flowered species of the Zambezica area in Vernonia along with some trinervate species with reddish to purplish flowers. Humbert (1960) placed the yellow- flowered species of Madagascar in two groups with the note that the name Diste- phanus would apply to his Group V. Jones (1981) recognized the yellow-flowered species as a subsection of Vernonia in his classification of the Paleotropical members of the genus, but as mentioned above, treat- ed Distephanus as a distinct indirectly re- lated subsection. The retention of the name Gongrotham- nus at the generic level by Robinson et al. (1980) was based on the difference in flower color along with the realization that sub- division of Vernonia seemed inevitable. A firm basis for the distinction of the yellow- flowered species has arisen from the present study initiated as a review of stylar nodes in the tribe. The nodes as well as the leaf trinervation, flower color, anther tails, en- dothecial cells, and pollen variation all in- dicate a distinctive element of small shrub- by or scandent Vernonieae distributed primarily in the western Indian Ocean. The characters involved warrant the following individual analyses. Trinervate leaves. —Early observations of Gongrothamnus by the senior author, and studies by most other authors (Humbert 1960, Wild 1978) were without recognition of the importance of the trinervate character PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON in the tribe. Only Bentham and Hooker (1873) seem to have appreciated how un- usual the character is in the tribe when giv- ing their reasons for transferring Gongro- thamnus to the Senecioneae. Trinervation versus pinnate venation is certainly highly variable in most other tribes in the family, commonly varying within genera. It was only with more detailed studies of Neotropical Vernonieae that the senior author noted the fundamentally pinnate nature of leaf ve- nation in the Vernonieae (Robinson et al. 1980) and commented upon the lack of tri- nervation in the tribe in comparisons with the Liabeae (Robinson 1983, Robinson et al. 1985). The species of Distephanus can now be seen as an exception in the Verno- nieae, an exception in which most of the species are trinervate or have leaves reduced to an ill-defined venation pattern. Triner- vation has been seen in no other Vernonieae and is regarded as uniquely derived in this Indian Ocean element of the tribe. Yellow flowers. —Closely correlating with the trinervate leaves in the Vernonieae are the yellow flowers. The latter character is sufficiently exceptional in the tribe to have caused both Bentham and Hooker (1873) and Hoffmann (1890— 1894) to place mem- bers of the genus in the tribe Senecioneae. A few species on the African mainland were noted by Wild (1978) as having trinervate leaves but not having yellow flowers. These are scandent like the yellow-flowered species of Distephanus on the mainland and have stylar nodes, basal appendages on the anther thecae, and mostly unstriated endothecial sclerified shields as in that genus. These red- dish species appear to be members of Dis- tephanus, but occur outside of the apparent center of distribution of the genus, where they seem to be a derived element. Since hybridization is common in the Asteraceae, such convergences in the Vernonieae may well represent borrowing of traits from the associated reddish-flowered Vernonieae on the continent. In any case, the character fail- ure is regarded as a de-differentiation be- VOLUME 99, NUMBER 3 tween two well-defined elements of the tribe and not a relict of any ancestral type. Basal appendages of the anther.—The Vernonieae, like almost all Cichorioideae, have spurred bases on the anthers, but in addition to the fertile extensions, most the- cae have at least a fringe of sterile tissue at the lower end. In typical Vernonia and most species that have been placed in the genus, the sterile tissue is usually unsclerified and easily overlooked, but some species in ba- sically unappendaged groups, such as V. megaphylla Hieron. have a small sclerified basal appendage. In Neotropical Verno- nieae, prominent narrow sclerified basal ap- pendages are the character by which Pip- tocarpha has traditionally been distinguished from Vernonia. In Paleotropical members of the tribe such as Distephanus, basal an- ther appendages have not been noted, prob- ably because they usually are broad and do not have the form of a narrow tail. One African segregate of Vernonia, Bacchar- loides (Vernonia subgenus Stengelia), has no sclerified appendage, but all of Diste- phanus, typical Gymnanthemum, and some species of Humbert’s (1960) Group IV from Madagascar show distinct basal append- ages. A narrow basal appendage in the form of a broad tail does occur in Distephanus glandulicincta. The character technically places these elements outside of the tradi- tional definition of Vernonia, even though the less obvious appendages of these Afri- can Vernonieae have traditionally been ig- nored. The appendages in Gymnanthemum and Group IV may indicate relationship of these groups to Distephanus, but other char- acters such as the trinervate leaves and flow- er color are different. Of these, only some species of Group IV have a distinct stylar node, but they differ by having deciduous inner bracts in the involucre. Endothecial cells. —A\l members of Dis- tephanus examined have median endothe- cial cells with a single unlined sclerified shield on the outer surface. These superfi- cially appear like a series of non-contiguous 495 cells in the endothecium. This contrasts with the more annulated appearance of other Vernonieae that have been seen. A tendency toward an intermediate condition occurs in D. angulifolius, a species of continental Af- rica noted above for possible introgression with other Vernonieae in flower color. Gym- nanthemum and Humbert’s Group IV both can have sclerified shields approaching those of Distephanus in form but with multifid ends. Those of typical Gymnanthemum dif- fer further in their frequently oblique or nearly transverse direction. Apical anther appendages. —Glands oc- cur on the apical anther appendages of typ- ical Vernonia and on many other Neotrop- ical species placed in the genus. Such glands are lacking in some of the American species which seem to have generally lost the ability to produce glands anywhere on the plant. The African groups seen in the present study show no glands on the appendages in any species, even when glands occur on other parts. Stylar node.—An expanded node with sclerified cells at the base of the style occurs commonly in the Vernonieae, and the char- acter may be basic to the tribe. The char- acter is most common, however, in Neo- tropical members of the tribe; it has a comparatively restricted distribution among Paleotropical species. A few African entities such as V. bainesii Oliv. & Hiern, have slight nodal development, but distinct nodes as large as any in the tribe seem almost totally restricted to and characteristic of Diste- phanus among the Paleotropical Verno- nieae. The mature node of Distephanus is puck-like, narrowing abruptly above into the stalk partly as a result of shrinkage of un- sclerified stalk tissue. A few species from Madagascar placed by Humbert (1960) in his Group IV, such as V. andrangovalensis Humb. and V. appendiculata Less., have nodes as large, but the latter group differs by being larger shrubs and trees, having red- dish flowers normal for the tribe, and hav- ing deciduous inner bracts of the involucre. 496 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-6. Equatorial and polar views showing sublophate intercolpar surface; 3-6, D. divaricatus (Steetz) R. & K., Colpar view; 4, Polar view, showing lack of regular sublophate intercolpar surface; 5, End of colpus; 6, Area of broken tectum showing basal columellae. Pollen.—The so-called Lychnophora-type pollen (Stix 1960) or Type A pollen (Keeley and Jones 1979), the most widely distrib- uted pollen type in the Vernonieae, is found in most species of Distephanus and in Gym- nanthemum and Humbert’s Group IV. The grains seen in Distephanus and Group IV are ca. 38—40 wm in diameter while those of typical Gymnanthemum ca. 45—47 wm in diameter. In Distephanus the Type A pollen occurs in all non-Madagascar species (Figs. 3-4) and six species seen from Madagascar, D. antandroy, D. glandulicinctus, D. ma- lacophytus, D. nummulariaefolius, D. ro- chonioides and D. subluteus. The Type A pollen grains of Distephanus seem unusual in the irregularity of the surface pattern, an irregularity not seen in Gymnanthemum (Figs. 1-2) and believed here to be related to the irregularity seen in the surface pattern Pollen of Gymnanthemum and Distephanus, lines = 5 wm. 1-2, G. coloratum (Willd.) R. & K.., of the non-Type A grains found in other species of Distephanus. Another aspect of these Type A grains that might be unusual shows in broken areas of D. divaricatus pol- len, where separate smaller basal columellae occur in the spaces between the primarily basal columellae under the spines (Fig. 6). Seven other species of Distephanus from Madagascar, D. cloiselii, D. eriophyllus, D. mahafaly, D. mangokensis, D. ochroleucus, D. swinglei, and D. trinervis, have a dis- tinctive type of pollen not seen elsewhere in the tribe (Figs. 7-12). It is lophate with areolae distinct but lacking any definite or- der. The ridges are not pitted or perforated on the edges, but a weakly perforated tectum lines the lower sides and bottoms of the areolae (Fig. 10). The direct attachment of the crests to the foot layer seems intermit- tent within the area covered by perforated VOLUME 99, NUMBER 3 497 Figs. 7-12. Pollen of Distephanus ochroleucus (Baker) R. & K., lines = 5 um. 7, Colpar view; 8, Polar view; 9, Oblique view of intercolpar area showing irregular lophate pattern; 10, Areolae showing non-perforate edges of tectum ridges and perforate basal parts; 11-12, Broken sections of ridges showing parts with and without direct median attachment of the foot layer. tectum. The pores are located in long colpi into which walls partially intrude in an al- ternating pattern (Fig. 7). The alternating pattern of the intruding walls seems char- acteristic and is apparently directly linked to the characteristic irregularity of the tec- tum areolation. The ridges of these species of Distephanus (Figs. 11-12) are reminis- cent of those in the pollen of Cyanthillium Blume, Phyllocephalum Blume, Stokesia L’Her., or the Elephantopinae, but the latter have more regular reticulations, have more truncated colpi or complete cross-walls above and below the pores which interrupt any colpi, and have the ridges raised on a series of small columellae or a fenestrated curtain above the surface of the foot layer. None have the partial perforated tectum seen in Distephanus. The presence of variation of pollen type within the well defined related group Dis- tephanus is not the first example of such variation in the Vernonieae. As in other ex- amples, the variation involves the Type A pollen and is not a variation between two of the lophate types. This type of anomalous occurrence along with the general pattern of distribution of Type A in the tribe suggests that the Type A is often the product of re- version and not necessarily the primitive form. In this particular case, one can theo- rize that it is only necessary developmental- ly to limit the type of structure seen on the ridge margins to the isolated spine tips and allow the perforated tectum to become the continuous structure of the surface instead. In any case, the presence of Type A pollen is not regarded here as evidence of either primitiveness or direct relationship be- tween the species in which it occurs. Con- 498 sidering the specialized lophate pollen of Distephanus alone seems to present a more realistic picture of relationship, with the type being restricted to the distinctive genus cen- tered geographically in Madagascar. Distephanus is a genus differing in two obvious characters from all other Verno- nieae. All the species have either the yellow flowers or the trinervate leaves, neither fea- ture occurring elsewhere in the tribe. The naturalness of the group is supported by other less obvious but nearly as unique fea- tures such as the puck-like stylar node, and the simple broad sclerified shields of the endothecial cells. The basal appendages of the anther thecae place Distephanus with Gymnanthemum and parts of Humbert’s Group IV technically outside of the tradi- tional definition of Vernonia to whose North American type they have only an extremely paraphyletic relationship. Species of Diste- phanus showing flower color or endothecial cells approaching the type seen in “‘Verno- nia” are seen as recent introgressions. The genus Distephanus is resurrected and de- fined as follows. Distephanus Cassini Distephanus Cassini, Bull. Soc. Philom. 1817:151. 1817. Type Conyza populifolia Lam. Vernonia sect. Distephanus (Cassini) Ben- tham & Hooker f., Gen. Pl. 2:228. 1873. Vernonia subsect. Distephanus (Cassini) Jones, Rhodora 83:68. 1981. Gongrothamnus Steetz ex Peters, Reise Mossamb. Bot. 336. 1862. Type Gongro- thamnus divaricatus Steetz. Vernonia subsect. Gongrothamnus (Steetz) Jones, Rhodora 83:65. 1981. Newtonia O. Hoffmann, Engler & Prantl., Naturl. Pflanzenfam. 4(4):285. 1892. Type Newtonia angolensis O. Hoffmann. Not Newtonia Baill. 1888. Antunesia O. Hoffmann, nom. nov., Bolet. Soc. Brot. 10:178. 1893. Type Newtonia angolensis O. Hoffmann. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Small shrubs or vines. Leaves alternate, blades usually trinervate, often with trun- cate or subcordate bases, less often narrow with cuneate bases and irregularly pinnate venation. Inflorescences terminal on branches, of corymbose cymes, with minute bracts, peduncles short but distinct. Heads campanulate; involucral bracts mostly 21— 24, ca. 75 in D. forrestii, persistent, multi- seriate, graduated, unappendaged apically; receptacle epaleaceous. Flowers mostly 10— 16 in a head, ca. 75 in D. forrestii. Corollas usually yellow, purplish in a few continental African species; thecae of anthers with dis- tinct broad sclerified basal appendages; en- dothecial cells with simple, broad, non-con- tiguous, sclerified shields; apical appendages of anthers without glands; style base with large abruptly broadened node. Achenes cy- lindrical to prismatic, sometimes subtri- quetrous or quadrangular, with 5—12 ribs, usually 10, setulae or glands present or ab- sent; carpopodium turbinate; pappus of 10 or more shorter outer squamellae alternat- ing with 10 or more longer capillary or lin- ear inner bristles or squamellae. Pollen in many species Type A with continuous in- tercolpar perforated tectum and subreticu- lately arranged spines, in some Madagascar species irregularly lophate with perforated tectum restricted to lower sides and bases of crests, with distinct colpi intruded upon by short alternating spurs of reticulate tec- tum. The species recognized in the genus in this study are as follows: Distephanus angolensis (O. Hoffmann) H. Robinson & B. Kahn, comb. nov. Newtonia angolensis O. Hoffm., Natirl. Pflanzenfam. 4(5):285. 1892. Antunesia angolensis (O. Hoffm.) O. Hoffm., Bolet. Soc. Brot. 10:178. 1893. Gongrothamnus angolensis (O. Hoffm.) Hiern, Cat. Welw. Afr. Pl. 1:592. 1898. Vernonia angolensis (O. Hoff.) N. E. Brown, Kew Bull. 1909: 116. 1909. SW Africa. VOLUME 99, NUMBER 3 Distephanus anisochaetoides (Sond) H. Robinson & B. Kahn, comb. nov. Vernonia anisochaetoides Sond, Linnaea 23: 61. 1850. SE Africa. Distephanus angulifolius (DC.) H. Robinson & B. Kahn, comb. nov. Vernonia angulifolia DC., Prodr. 5:29. 1836. SE Africa. Distephanus antandroy (H. Humbert) H. Robinson & B. Kahn, comb. nov. Vernonia antandroy H. Humb., Not. Syst. Paris 8(1):7. 1939. Madagascar. Distephanus cloiselii (Moore) H. Robinson & B. Kahn, comb. nov. Vernonia cloiselii Sp. Moore, Journ. Bot. 44:145. 1906. Madagascar. Distephanus divaricatus (Steetz) H. Robinson & B. Kahn, comb. nov. Gongrothamnus divaricatus Steetz in Pe- ters, Reise Mossamb. Bot. 2:342. 1864. Not V. divaricatus Swartz, 1806. Gongro- thamnus aurantiacus O. Hoffm., Bot. Jahrb. 30:433. 1901. Vernonia aurantia- ca (O. Hoffm.) N. E. Brown, Kew Bull. 1909:116. 1909. Vernonia vitellina N. E. Brown, Kew Bull. 1909:117. 1909. East Africa. Distephanus eriophyllus (Drake) H. Robinson & B. Kahn, comb. nov. Vernonia eriophylla Drake, Bull. Soc. Bot. Fr. 46:230. 1889. Madagascar. Distephanus forrestii (Anthony) H. Robinson & B. Kahn, comb. nov. Vernonia forrestii Anthony, Notes Bot. Gard. Edinb. 18:35. 1933. Yunnan. Distephanus garnieriana (Klatt) H. Robinson & B. Kahn, comb. nov. 499 Vernonia garnieriana Klatt, Linnaea 37:508. 1872. Vernonia parviflora (error for par- vifolia) Klatt, Ann. Sc. Nat. 5° sér. Bot. 362. 1873. Vernonia lyallii Baker, J. Linn. Soc. 20:174. 1883. Vernonia moqui- nioides Baker, J. Linn. Soc. 20:177. 1883. Vernonia alboviridis Baker, J. Linn. Soc. 25:325. 1890. Madagascar. Distephanus glandulicinctus (H. Humb.) H. Robinson & B. Kahn, comb. nov. Vernonia glandulicincta H. Humb., Not. Syst. Paris 8(4):306. 1948. Madagascar. Distephanus glutinosus (DC.) H. Robinson & B. Kahn, comb. nov. Vernonia glutinosa DC., Prodr. 5:18. 1836. Vernonia scariosa Baker, Journ. Bot. 20: 169. 1882, hom. illeg. Vernonia lepido- phylla Drake, Bull. Soc. Bot. Fr. 46:229. 1889. Madagascar. Distephanus lastellei (Drake) H. Robinson & B. Kahn, comb. nov. Vernonia lastellei Drake, Bull. Soc. Bot. Fr. 46:232. 1899. Vernonia goudotii Drake, Bull. Soc. Bot. Fr. 46:239. 1899. Mada- gascar. Distephanus mahafaly (H. Humb.) H. Robinson & B. Kahn, comb. nov. Vernonia mahafaly H. Humb., Not. Syst. Paris 8(1):10. 1939. Madagascar. Distephanus majungensis (H. Humb.) H. Robinson & B. Kahn, comb. nov. Vernonia majungensis H. Humb., Not. Syst. Paris 13:308. 1948. Madagascar. Lecto- type designated here, Humbert & Perrier BUNTY (OP). Distephanus malacophytus (Baker) H. Robinson & B. Kahn, comb. nov. Vernonia malacophyta Baker, Journ. Linn. Soc. 25:323. 1890. Vernonia rampans 500 Baker, Journ. Linn. Soc. 25:323. 1890. Vernonia grandidieri Drake, Bull. Soc. Bot. Fr. 46:240. 1899. Madagascar. Distephanus manambolensis (H. Humb.) H. Robinson & B. Kahn, comb. nov. Vernonia manambolensis H. Humb., Not. Syst. Paris 13:306. 1948. Madagascar. Distephanus mangokensis (H. Humb.) H. Robinson & B. Kahn, comb. nov. Vernonia mangokensis H. Humb., Bull. Soc. Bot. Fr. 87:347. 1940. Madagascar. Distephanus nummulariaefolius (Klatt) H. Robinson & B. Khan, comb. nov. Decaneurum (Gymnanthemum) nummu- lariaefolium Klatt, Ann. Sc. Nat. 5° sér. Bot. 18:363. 1873. Gongrothamnus mul- tiflorus Klatt, Flora 68:205. 1885. Ver- nonia leucolepis Baker, Journ. Bot. 25: 322. 1890. Madagascar. Distephanus ochroleucus (Baker) H. Robinson & B. Kahn, comb. nov. Vernonia ochroleuca Baker, Journ. Linn. Soc. 20:179. 1885. Vernonia trichantha Baker, Journ. Linn. Soc. 21:416. 1884. Madagascar. Distephanus polygalaefolia (Less.) H. Robinson & B. Kahn, comb. nov. Vernonia polygalaefolia Less, Linnaea 6: 628. 1831. Vernonia arbutifolia Baker, J. Bot. 20:169. 1882. Vernonia perrieri Drake, Bull. Soc. Bot. Fr. 46:229. 1899. Madagascar. Distephanus populifolius (Lamarck) Cassini Conyza populifolia Lamarck, Encyc. 2:87. 1786. Distephanus populifolius (La- marck) Cassini, Bull. Soc. Philom. 1817: 151. 1817. Vernonia populifolia (La- marck) Spreng., Syst. 3:434. 1826. Mau- ritius. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Distephanus rochonioides (H. Humb.) H. Robinson & B. Kahn, comb. nov. Vernonia rochonioides H. Humb., Bull. Soc. Bot. Fr. 87:346. 1940. Madagascar. Distephanus streptocladus (Baker) H. Robinson & B. Kahn, comb. nov. Vernonia streptoclada Baker, Journ. Linn. Soc. 21:416. 1885. Madagascar. Distephanus subluteus (S. Elliot) H. Robinson & B. Kahn, comb. nov. Vernonia sublutea S. Elliot, Journ. Linn. Soc. 29:26. 1891. Madagascar. Distephanus swinglei (H. Humb.) H. Robinson & B. Kahn, comb. nov. Vernonia swinglei H. Humb., Not. Syst. Paris 8(1):8. 1939. Madagascar. Distephanus trinervis Boj. ex DC. Distephanus trinervis Boj. ex DC., Prodr. 5: 75. 1836. Distephanus capitatus Boj. ex DC., Prodr. 5:74. 1836. Vernonia capi- tata (Boj. ex DC.) Drake in Grandidier, Hist. Madag. Pl. VI, Atlas, pl. 464. 1897. Vernonia trinervis (Boj. ex DC.) Drake, Bull. Soc. Bot. Fr. 46:228. 1899. Vernonia rusillonii Hochr., Ann. Cons. Genéve, | 1— 12:117. 1908. Madagascar. Additional species having the characters of the genus according to Humbert (1960) but not seen in this study are as follows: Vernonia ambongensis H. Humb., V. bara H. Humb., V. bakeri Vatke, V. capuronii H. Humb., V. Grevei Drake, V. ibityensis H. Humb., V. madagascariensis Less., V. pois- sonii H. Humb., V. polytricholepis Baker, V. quartziticola H. Humb., V. rhodopappa Baker, V. spiciforma Klatt. In the course of the present study it has seemed best to recognize Gymnanthemum at the generic level also, since it also falls outside of the definition of traditional Ver- nonia, and the status is inevitable in any VOLUME 99, NUMBER 3 final revision of the tribe. This would not be so important at the present time except for the fact that the oldest name for the type species G. cupulare Cass. has never been transferred to the genus. There seems to be no reason for the precedence usually given to the Persson name Baccharis senegalensis since the Willdenow name antidates it by four years, unencumbered by inadequate description or homonymy. Gymnanthemum coloratum (Willd.) H. Robinson & B. Kahn, comb. nov. Eupatorium coloratum Willd., Sp. Pl. 3: 1768. 1803. Acknowledgments The photographs of the pollen were pre- pared by the Smithsonian Institution Mu- seum of Natural History SEM laboratory supervised by Walter Brown. An Hitachi 570 scanning electron microscope was used. The microscope was operated by Heidi Wolf. Dr. Joan Nowicke is thanked for use of the facilities of the Palynological Laboratory in the Smithsonian Department of Botany. Literature Cited Bentham, G., and Hooker, J. D. 1873. —Genera Plantarum 2(1):1—554. Brown, N. E. 1909. List of plants collected in Nga- miland and northern part of the Kalahari Des- ert, chiefly in the neighbourhood of Kwebe and along the Botletle and Lake rivers.—Kew Bul- letin of Miscellaneous Information 1909:89-146. Cassini, H. 1816-1817. Apercu des genres nouveux dans la famille des Synanthérées.— Bulletin de la Société Philomatique 1816:198-—200; 1817: 10-13, 31-34, 66-70, 115-118, 137-140, 151- 154. 1819. Distéphane. Jn G. Cuvier, ed., Dic- tionnaire des Sciences Naturelles 13:361. 501 Hoffmann, O. 1890-1894. Compositae. In A. Engler and K. Prantl, ed., Die Natiirlichen Pflanzen- familien 4(5):87—387. Leipzig. 1893. Compostas de Africa portugueza.— Boletim da Sociedade Broteriana 10:170-185. “1892.” 1902. Compositae. Jn Engler, A., Berichte uber die botanischen Ergebnisse der Nyassa-See und Kinga-Gebirgs-Expedition der Hermann- und Elise- geb. Heckmann-Wentzel-Stifting. IV. Die von W. Goetze am Rukwu-See und Nyassa- See sowie in den zwischen beiden Seen gelege- nen Gebirgslandern insbesondere dem Kinga- Gebirge gesammelten Pflanzen, nebst eigigen Nachtragen (durch bezeichnet) zu Bericht III.— Botanische Jahrbiicher 30:421-445. Humbert, H. 1960. 189° Famille Composées (Com- positae) Tome 1.—Flore de Madagascar et des Comores (Plantes vasculaires) 1:1-—338. Jones, S. 1981. Synoptic classification and pollen morphology of Vernonia (Compositae: Verno- nieae) in the Old World.—Rhodora 83:59-75. Keeley, S., and Jones, S. B. 1979. Distribution of pollen types in Vernonia (Compositae).—Sys- tematic Botany 4:195-202. Robinson, H. 1983. A generic review of the tribe Liabeae (Asteraceae).—Smithsonian Contribu- tions to Botany 54:1-69. , Bohlmann, F., and King, R. M. 1980. Che- mosystematic notes on the Asteraceae. III. Nat- ural subdivisions of the Vernonieae.—Phyto- logia 46:421-436. , Powell, A. M., King, R. M., and Weedin, J. F. 1985. Chromosome numbers in Compositae, XV: Liabeae.— Annals of the Missouri Botani- cal Garden 72:469-479. Stix, E. 1960. Pollenmorphologische untersuchungen an Compositen.—Grana Palynologica 2(2):41- 104, pl. 1-21. Wild, H. 1978. The Compositae of the Flora Zam- besiaca Area 8. Vernonieae (Vernonia). —Kirkia 2:31-127. Department of Botany, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 502-508 A NEW VULTURE (VULTURIDAE: PLIOGYPS) FROM THE LATE MIOCENE OF FLORIDA Jonathan J. Becker Abstract.—A new vulture, Pliogyps charon n. sp., from the late Miocene (latest Clarendonian) Love Bone Bed locality, Alachua Co., Florida, is described from a complete tarsometatarsus and referred distal ends of the tibiotarsus and humerus. Pliogyps charon is distinguished from Pliogyps fisheri Tordoff (1959) by smaller size, and by proportions and qualitative characters of the tarso- metatarsus. Pre-Quaternary vultures are poorly known. Recently, Olson (1985) reviewed the fossil history of this family. The earliest rec- ord of the family Vulturidae is from the late Eocene and early Oligocene deposits of the Phosphorites du Quercy, France, and in the early Oligocene of Mongolia. In South America, the oldest vulture is Dryornis pampeanus Moreno and Mercerat, from the Monte Hermoso Formation in Argentina, which is probably close to the living genus Vultur (Tonni 1980). The oldest records of vultures in North America are Sarcoram- phus kernensis from the late Miocene (mid- Hemphillian) of Kern River, California, and a specimen under study from the mid-Bar- stovian Sharkstooth Hill local fauna (Em- slie, pers. comm.). While there are several described genera and species of vultures from the Pliocene and Pleistocene (Brod- korb 1964, Olson 1985), the true number of valid taxa among them remains to be determined. This paper describes a new species of New World vulture (Family Vulturidae), dis- cusses functional aspects of the genus Plio- gyps, and comments on the generic status of living and fossil members of this group. Material and Methods Recent specimens examined are in the Florida State Museum, the collection of Pierce Brodkorb, and the National Museum of Natural History. Recent skeletons of Vul- turidae examined: Gymnogyps californi- anus, 7; Vultur gryphus, 9; Sarcoramphus papa, 9; Coragyps atratus, 16; Cathartes aura, 4. Fossil specimens are in the verte- brate paleontology collections of the Florida State Museum (UF) and the Museum of Pa- leontology, University of Michigan. Ana- tomical terminology follows Baumel et al. (1979). Measurements, defined in Table 1 and Figs. 2 and 3, were taken with dial cal- ipers accurate to 0.05 mm, and rounded to the nearest 0.1 mm. Systematics Order Accipitriformes (Vieillott, 1816) Family Vulturidae (Illiger, 1811) Pliogyps Tordoff, 1959 Emended generic diagnosis. —Tarso- metatarsus of Pliogyps differing from that of other living and fossil genera of vultures in having a proportionately large trochlea for digit III, proximal articular surface wide and deep in comparison to length of bone, a generally columnar form, with symmet- rical lateral and medial flaring, both proxi- mally and distally; shaft wide in comparison to length of bone; hypotarsus merging dis- tally with shaft by means of broad, rounded ridge (as in Vultur, Breagyps, Gymnogyps, and Geranogyps; more narrow in Coragyps, Cathartes, and Sarcoramphus). This last VOLUME 99, NUMBER 3 character may be strictly size dependent and if so, not of value as a generic character. Remarks. —Two characters (shaft less deeply and extensively excavated anterior- ly; groove of trochlea for digit III ending anteroproximally in a shallow, but distinct pit) used by Tordoff (1959) to define this genus are variable within a given species, and therefore should not be used as generic characters. Pliogyps charon, new species Fig. 1 Holotype. —UF 25952, complete right tarsometatarsus, missing a small portion of hypotarsus. Paratype. —UF 25886, distal end left tib- iotarsus. Referred material. —UF 25719, fragment of shaft of left humerus, tentatively referred. Type locality. —Love Bone Bed local fau- na, along State Road 241, near Archer, Ala- chua County, Florida (NW 4, SW %4, NW %, Sec. 9, T. 11 S., R. 18 E., Archer Quad- rangle, U.S. Geologic Survey 7.5 minute se- ries topographical map, 1969). Fossil ver- tebrates occurring in the Alachua Formation (Williams et al. 1977) are considered latest Clarendonian in age (Webb et al. 1981), about 9 million years before present. The known avifauna of the Love Bone Bed is primarily aquatic, with a few terrestrial species being present (Becker 198Sa, b). Diagnosis. —Tarsometatarsus distin- guished from that of Pliogyps fisheri Tordoft (1959) in smaller size, in having a narrow ridge extending from hypotarsus farther down shaft (caudal view), in having sulcus extensorius more excavated and extending farther down shaft (cranial view), and in having shaft and trochlea III proportion- ately less deep. Etymology. —The specific name charon, Greek, masculine, is a noun in apposition. In Greek mythology Charon, portrayed as a robust old man, ferries the souls of the dead across the River Styx (considered by 503 Table 1.—Measurements of the tibiotarsi and tar- sometatarsi of the vultures Coragyps atratus atratus (n = 16, 8 males, 8 females), Pliogyps fisheri, and Plio- gyps charon, new species. Data are mean + standard deviation and range. Measurements of tibiotarsus are W-DIST-CR, Transverse width of distal end, mea- sured across cranial portion of condyles; D-MCON, Greatest depth of medial condyle. Measurements of tarsometatarsus: LENGTH, Greatest length from in- tercondylar eminence (Eminentia intercondylaris) through trochlea for digit III (Trochlea metatarsi III). W-PROX, Greatest transverse width proximal artic- ular surface, measured across dorsal surface. D-PROX, Depth of proximal end, measured from dorsal edge of the proximal articular surface through the lateral hy- potarsal crest (Crista lateralis hypotarsi). W-DIST, Greatest transverse width of distal end. W-TRIII, Greatest transverse width of trochlea III. D-TRIII, Greatest depth of trochlea III. Measurements of Cor- agyps are included to show amount of variation present in a living population of vultures. P. fisheri P. charon Measurements C. a. atratus Tibiotarsus W-DIST-CR 12.79 + 0.41 — 19.5 12.1-13.6 D-MCON 13.67 + 0.41 — [18.1] 13.2-14.6 Tarsometatarsus LENGTH 84.43 + 1.54 94.0 86.6 80.4-87.1 W-PROX 15.11 = 0253" 21-9 21.1 14.1-16.2 D-PROX 11.71 + 0.42 — — 11.1-12.4 W-DIST 16.59 + 0.57 33.0 — 15.6-17.6 W-TRIII 6.43 + 0.21 9.6 9.2 6.0-6.7 D-TRIII 9.99 + 0.32 15.2 13.5 9.4-10.6 some authorities to be the River Acheron) into the lower world. This name also reflects the proximity of a modern River Styx, lo- cated 18 miles due east of the Love Bone Bed locality on the northern edge of Orange Lake, Alachua County, Florida. Description. —In cranial view, the proxi- mal vascular foramina are large and are ap- proximately equal in size. The papilla for PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Photographs of holotype and paratype of Pliogyps charon, new species. A-C, UF 25886, distal end of left tibiotarsus, paratype. D-G, UF 25952, right tarsometatarsus, holotype. A, D, Cranial (dorsal) view; B, E, Caudal (plantar) view; C, G, Distal end view; F, Proximal end view. VOLUME 99, NUMBER 3 W-SHAFT 80 90 100 505 110 120 130 140 LENGTH Fig. 2. Bivariate plot of greatest length from intercondylar eminence (Eminentia intercondylaris) through trochlea for digit III (Trochlea metatarsi III) (LENGTH) versus transverse width of shaft (W-SHAFT) of the tarsometatarsi of the following species of vultures: open square— Pliogyps charon, new species, open triangle— Pliogyps fisheri, solid triangles—Sarcoramphus papa, open circles— Vultur gryphus, solid circles—Gymnogyps californianus. the attachment of M. tibialis cranialis is rounded and is in two parts. The sulcus ex- tensoris, with a sharp lateral border, extends down the shaft to the distal foramen. A dis- tinct intermuscular line extends obliquely through this sulcus (separating attachments for the extensor digitorum brevis pars hal- lucis and extensor digitorum brevis pars ad- ductor-extensor digiti IV; Jollie 1977:43). In caudal view, the tarsometatarsus has a long ridge extending down the shaft from the hypotarsus. This ridge terminates in an intermuscular line that extends to the level of the articular facet of metatarsal I. The distal end of tibiotarsus has a broad extensor sulcus. There is a slight projection of bone on the lateral surface of the distal end (approximately 4 cm from distal end) for attachment of the fibula. The intercon- dylar sulcus is broad, and the external con- dyle merges evenly into it. In distal end view, the intercondylar sulcus is asymmetrical, with the lateral border sloping gradually, and the medial border sloping abruptly, up from the base of the intercondylar sulcus (sym- metrical or U-shaped in Coragyps, Sarco- ramphus, Breagyps, Gymnogyps; asymmet- rical in Cathartes). Measurements are given in Table 1. Humerus fragment tentatively referred. Discussion. — There is a strong correlation between the cross-sectional area of the tib- iotarsus of a given avian species and its live weight (Campbell and Tonni 1983, Prange et al. 1979). The following empirically de- rived regression equation, log Y = 2.54 log X — 0.19906 where Y is the live body weight (gms) and X is the least shaft circumference of the tib- iotarsus (mm), has a correlation coefficient of 0.986, showing that the predictions of the 506 26 24 22 20 18 FLEXOR 16 14 12 10 70 80 90 100 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 120 110 130 140 LENGTH Fig. 3. Bivariate plot of greatest length (LENGTH) versus length from intercondylar eminence to middle of tubercle for tibialis anterior (Tuberositas m. tibialis cranialis) (FLEXOR) of tarsometatarsi of the following species of vulture: open square—Pliogyps charon, new species, open triangle— Pliogyps fisheri, solid triangles— Sarcoramphus papa, open circles— Vultur gryphus, solid circles—Gymnogyps californianus. live weight should be very accurate. The least shaft circumference of the paratype of Pliogyps charon (UF 25886) measures 32 mm, yielding a predicted weight of 5.2 kg. Sarcoramphus papa, which has a similar tarsometatarsal length, weighs between 3.0 and 3.75 kg (5 individuals, Brown and Amadon 1968). This supports Tordoffs (1959:341ff) contention that Pliogyps is rel- atively a heavy-bodied, short-legged vul- ture. Figures 2 and 3 show the length of the tarsometatarsus plotted against the width of the shaft and the flexor length. Species of Pliogyps have an average flexor length, but a very broad tarsometatarsus in comparison to its length. When considered in conjunc- tion with the well-developed muscle attach- ments discussed above, a powerful pelvic limb is suggested. It is possible that P. char- On was more rapacious than other living or fossil vultures, or that this powerful pelvic limb is merely a reflection of a heavy body. As additional fossil material of this species becomes available, these tentative sugges- tions should be examined further. The intergeneric relationships of living and fossil vultures are difficult to determine, owing to a paucity of pre-Pleistocene fossil specimens and the conservative nature of the tarsometatarsus (the holotypical ele- ment in most fossil species). Pliogyps shares some tarsometatarsal characters with Sar- coramphus (anterior fossa continuing down shaft to the distal foramen, a similar size of the distal foramen and a similar shape of the hypotarsal ridge), but differs from Sar- coramphus in the degree of elevation of trochlea III (proximal border merging smoothly with shaft [plantar surface] in all modern skeletons of Sarcoramphus exam- ined), and the amount of excavation of the VOLUME 99, NUMBER 3 lateral parahypotarsal sulcus. Pliogyps also shares the following characters with Vultur and Gymnogyps: the lateral side of the area proximal to trochlea IV is inclined and the excavation of the anterior fossa extends to the distal foramen (although to a lesser de- gree than in Sarcoramphus). Mayr and Short (1970) proposed that Pliogyps, Vultur, and Gymnogyps should be viewed as congeneric. The proportions of the tarsometatarsus of both species of Plio- gyps are quite distinct from all species of Gymnogyps and Vultur. Considering the conservative nature of the tarsometatarsus in this family, I would maintain Pliogyps as separate from all other large vultures. It does appear possible though, that the large condors (species of Vultur and Gym- nogyps and probably Geranogyps and Brea- gyps) are all part of one radiation that differ primarily in the degree of cranial special- ization (Vultur being primitive; Gymnogyps and Breagyps more specialized, Jollie 1977: 110; Geranogyps unknown). Fisher (1944, 1946) discusses a number of characters of the skull and locomotor apparatus that unites the condors as a group separate from the more primitive genera Cathartes and Coragyps. Acknowledgments This paper is a portion of a dissertation completed at the Department of Zoology, University of Florida. I thank P. Brodkorb, R. A. Kiltie, G. S. Morgan, S. D. Webb, E. S. Wing, and R. G. Wolff for their com- ments on the dissertation; S. L. Olson and R. L. Zusi for their comments on this paper. The following individuals made fossil and/or Recent specimens available for study: G. Barrowclough, F. Vuilleumier, Ameri- can Museum of Natural History; P. Brod- korb, University of Florida; J. Hardy, B. J. MacFadden, G. S. Morgan, S. D. Webb, T. Webber, Florida State Museum; P. Ginger- ich, R. Payne, University of Michigan; H. James, S. L. Olson, D. Steadman, National 507 Museum Natural History, Smithsonian In- stitution. Financial support was received from the Frank M. Chapman Memorial Fund, Amer- ican Museum of Natural History, and from Sigma Xi Grants-In-Aid of research. Literature Cited Baumel, J. J.. A. S. King, A. M. Lucas, J. E. Breazile, and H. E. Evans. 1979. Nomina Anatomica Avium. Academic Press, London. 637 pp. Becker, J. J. 1985a. Pandion lovensis, a new species of osprey from the late Miocene of Florida.— Proceedings of the Biological Society of Wash- ington 98:314—320. . 1985b. The fossil birds from the late Miocene and early Pliocene of Florida. Ph.D. disserta- tion. University of Florida, Gainesville. 245 pp. Brodkorb, P. 1964. Catalogue of fossil birds. Part 2. (Anseriformes through Galliformes).— Bulletin of the Florida State Museum, Biological Sci- ences 8:195-335. Brown, L. H., and D. Amadon. 1968. Eagles, hawks, and falcons of the world. Country Life, London. 945 pp. Campbell, K. E., Jr., and E. P. Tonni. 1983. Size and locomotion in teratorns (Aves: Teratornithi- dae).— Auk 100:390-403. Fisher, H.I. 1944. The skulls of cathartid vultures. — Condor 46:272-296. 1946. Adaptations and comparative anato- my of the locomotor apparatus of New World vultures.—American Midland Naturalist 35: 545-727. Jollie, M. J. 1977. A contribution to the morphology and phylogeny of the Falconiformes. Part 4.— Evolutionary Theory 3:1-142. Mayr, E., and L. L. Short. 1970. Species taxa of North American birds. A contribution to comparative systematics.— Publication of the Nuttall Orni- thological Club 9:1-127. Olson, S. L. 1985. The fossil record of birds. Jn D.S. Farner, J. R. King, and K. C. Parkes, eds., Avian biology. Vol. 8, pp. 79-252. Academic Press, New York. Prange, H. D., J. F. Anderson, and H. Rahn. 1979. Scaling of skeletal mass to body mass in birds and mammals. — American Naturalist 113:103- 122. Tonni, E. P. 1980. The present state of knowledge of the Cenozoic birds of Argentina.—Contribu- tions in Science, Natural History Museum of Los Angeles County 330:105—114. Tordoff, H. B. 1959. A condor from the upper Plio- cene of Kansas.—Condor 61:338—343. 508 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Webb, S. D., B. J. MacFadden, and J. A. Baskin. 1981. Resources, Report of Investigations No. 85:1- Geology and paleontology of the Love Bone Bed 98. from the late Miocene of Florida.—American Journal of Science 281:513-544. Divisi . ivision of Birds, Department of Verte- Williams, K., D. Nicol, and A. Randazzo. 1977. The P Sle geology of the western part of Alachua County, brate Zoology, National Museum of Natu- ural Resources, Florida Department of Natural ington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 509-516 THREE NEW SPECIES OF WATER SCAVENGER BEETLES OF THE GENUS CHAETARTHRIA FROM SOUTH AMERICA (COLEOPTERA: HYDROPHILIDAE) Paul J. Spangler Abstract. —Three new species of water beetles, Chaetarthria porknockeri from Guyana, and Chaetarthria ayacuchana and Chaetarthria gavilana from Ven- ezuela are described. Distinguishing characters for the three species are illus- trated with pen and ink line drawings and their habitats are described. A key is provided to separate the new species from two previously described species. The species of Chaetarthria from the Western Hemisphere were revised by Miller in 1974; at that time 32 species and sub- species were known. Since Miller’s revision was published three new species from Ec- uador were described by Spangler (1977). During the past several years three addi- tional new species have been collected from South America. One species was collected in December 1983 during an Earthwatch expedition to the Takutu Mountains of Guyana. A second species was collected at Puerto Ayacucho, Venezuela, in January 1985 while I participated in an expedition to the tepui Cerro de la Neblina, in southern Venezuela and the third species was col- lected near Puerto Ayacucho, Venezuela, in February 1986. With the description of these new taxa, 38 species are now known from the Western Hemisphere for the genus Chaetarthria. The three new species de- scribed below key to couplet 41 in Miller’s key where C. granulata Miller and C. bra- silia Miller are separated. The five species now keying to couplet 41 may be identified by using the key following the description of C. gavilana, new species. Chaetarthria porknockeri, new species Figs. 1, 2 Holotype male.—Body form: Very con- vex, hemispherical. Length, 1.34 mm; greatest width, 1.1 mm, at about midlength. Color.—Black dorsally except reddish brown around margins of pronotum and lat- eral margins of elytra. Ventral surface red- dish brown with antennae and palpi light yellowish brown. Head.—Punctures indistinct, extremely fine and sparse; punctures between eyes sep- arated by 6 to 8 times their diameter; punc- tures slightly more dense along anterior margin of clypeus. Clypeus with lateral mar- gins moderately arcuate and rimmed; an- terior margin shallowly, broadly emargin- ate. Labrum more distinctly punctate than head, strongly rounded anteriorly. Eyes al- most round viewed dorsally but oblong viewed ventrally; shallowly emarginate where clypeus extends into anterior margin of eye. Ventral surface of head microalu- taceous behind eyes. Mentum smooth and shiny, strongly rounded apically. Submen- tum between maxillae microalutaceous. Antenna, 8 segmented; basal segment long, sinuous; second segment almost globular; third, fourth, and fifth segments slender, platelike; sixth, seventh, and eighth seg- ments increasingly larger; sixth segment with few setae apically; seventh and eighth seg- ments with long, rather dense setae; eighth segment ending in narrow apical projection. Maxillary palpus, 4 segmented; basal seg- ment tiny; second (pseudobasal) segment al- most as long as apical segment; third seg- ment one-third as long as second segment; 510 Figs. 1 and 2. Chaetarthria porknockeri, n. sp.; male genitalia. 1, Dorsal view. 2, Lateral view. apical segment longest; pseudobasal seg- ment when lying along stipes not attaining base of stipes. Labial palpus, 3 segmented; basal segment very small; second segment longest and bearing 2 long, golden setae near apex; apical segment about one-fourth shorter than second segment and bearing 2 long, golden, hairlike setae dorsally just be- fore apex. Thorax. —Pronotum strongly convex; sides, posterolateral angles, and anterolat- eral angles moderately rounded; lateral margins finely rimmed; anterior margin ar- cuate medially and slightly angulate behind eyes; disc essentially impunctate, punctures indistinct, very fine and sparse. Scutellum an equilateral triangle. Elytron 1.0 mm long; strongly convex; widest near midlength; dis- tinctly rimmed along base laterally; lateral margins finely rimmed, sinuate in lateral view; punctures on disc fine, indistinct, and sparse like those on pronotum, separated by 4 to 8 times their diameter; sutural stria extending and widening from slightly before midlength to apex; sides strongly declivous, without obvious epipleura. Metathoracic wings present. Prosternum, mesosternum, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and metasternum glabrous and shiny except middle of metasternum with fine, sparse, seta-bearing punctures. Prosternum narrow in front of procoxae. Mesosternum with a small, transverse, carinate process between and slightly in front of mesocoxae. Meta- sternum with slightly raised discal area; sides shallowly depressed. Profemur and meso- femur with dense, hydrofuge pubescence on basal two-thirds; apices glabrous. Metafe- mur finely, sparsely punctate; without dense pubescence except marginally along ante- rior third. Protarsus, mesotarsus, and meta- tarsus with basal segment shortest; second segment of mesotarsus and metatarsus slightly longer than third and fourth seg- ments combined; fifth segment slightly shorter than second segment. Metatrochan- ter moderately elongate and pubescent. Abdomen. — First and second abdominal sterna with a common, deep concavity. First sternum shiny; with numerous, very long, golden, hairlike setae along anterior margin and extending posteriorly as far as third sternum and holding a hyaline mass in con- cavity. Midline feebly raised longitudinally on first and second sterna. Sterna 3, 4, and 5 covered with short, dense, golden setae; fifth sternum rounded, not emarginate ap- icomedially. Male genitalia.—As illustrated (Figs. 1, 2). Female. —Unknown. Type data. —Holotype male: GUYANA: Mazaruni-Potaro District: Takutu Moun- tains, 6°15’N, 59°05'W, 18 Dec 1983, P. J. Spangler, M. Levine, E. Vystricil; deposited in the National Museum of Natural History, Smithsonian Institution. Paratype: Same data as holotype, 1 male. Etymology.—On weekends, when guy- anese gold miners would arrive in town late in the evening after stores were closed, the hungry miners would knock on the doors and demand to buy pork and other food. Sometimes, with their new-found wealth, they would buy the entire contents of the store. Consequently, those miners earned VOLUME 99, NUMBER 3 the name “‘pork-knockers.” This new species is named for the friendly gold miners who were panning gold in the lower reaches of the stream from which we “‘mined”’ this and other new species of aquatic beetles. Habitat.—The specimens were obtained by placing leaves in berlese funnels and re- covering the specimens from the berleseate. The wet, rotting leaves were collected from the margin of a small brook heavily shaded by the rainforest. Chaetarthria ayacuchana, new species Figs. 3-5 Holotype male.—Body form: Very con- vex, hemispherical. Length, 1.24 mm; greatest width, 1.0 mm, at about midlength. Color.—Black dorsally except reddish brown around margins of pronotum and lat- eral margins of elytra. Ventral surface of head dark reddish brown except antennae and palpi light yellowish brown; remainder of venter light reddish brown. Head.—With moderately coarse, sparse punctures; punctures between eyes separat- ed by 2 to 4 times their diameter; punctures slightly more dense along inner margin of each eye. Clypeus with lateral margins mod- erately arcuate and rimmed; anterior mar- gin shallowly, broadly emarginate. Labrum more distinctly punctate than head and al- most truncate anteriorly. Eyes almost round viewed dorsally but oblong viewed laterally; shallowly emarginate where clypeus extends into anterior margin of eye. Ventral surface of head microalutaceous behind eyes. Men- tum smooth and shiny, strongly rounded apically. Submentum finely, sparsely punc- tate between maxillae. Antenna, 8 seg- mented; basal segment long, sinuous; sec- ond segment almost globular; third, fourth, and fifth segments slender, platelike; sixth, seventh, and eighth segments increasingly larger; sixth segment with few setae apically; seventh and eighth segments with long, rather dense setae; eighth segment ending 511 Figs. 3 and 4. Chaetarthria ayacuchana, n. sp.; male genitalia. 3, Dorsal view. 4, Lateral view. in slender apical projection about one-sixth as long as swollen basal part. Maxillary pal- pus, 4 segmented; basal segment tiny; sec- ond (pseudobasal) segment swollen and al- most as long as apical segment; third segment one-third as long as second segment; apical segment longest; pseudobasal segment when lying along stipes not attaining base of stipes. Labial palpus, 3 segmented; basal segment very small; second segment longest and bearing numerous, long, golden setae me- dially and 1 near apex laterally; apical seg- ment about one-fourth shorter than second segment and bearing numerous, long, gold- en setae medially and 3 long, golden, hair- like setae dorsally just before apex. Thorax. —Pronotum strongly convex; sides, posterolateral angles, and anterolat- eral angles moderately rounded; lateral margins finely rimmed; anterior margin ar- cuate medially and slightly angulate behind eyes; punctures on disc moderately coarse, sparse, separated by 4 to 8 times their di- ameter. Scutellum an equilateral triangle. Elytron 0.97 mm long; strongly convex; widest near midlength; distinctly rimmed along base laterally; lateral margins finely PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Amazonas, Venezuela. rimmed, slightly sinuate in lateral view; seta- bearing punctures on disc coarse, distinct, and rather dense and much more coarse and dense than those on pronotum, separated by 3 to 4 times their diameter; sutural stria deep, extending and widening from slightly before midlength to apex; sides strongly de- clivous, without obvious epipleura. Meta- thoracic wings present. Prosternum, meso- sternum, and metasternum glabrous and shiny. Prosternum narrow in front of pro- coxae. Mesosternum with small, transverse, carinate process between and slightly in front of mesocoxae. Metasternum with slightly raised discal area; sides shallowly de- pressed. Procoxa with 5 very stout spines apicoventrally. Profemur with dense, hy- drofuge pubescence on basal two-thirds; apices glabrous. Mesofemur with fine, sparse, seta-bearing punctures; punctures denser along anterior third. Metafemur with fine, sparse, seta-bearing punctures margin- ally along anterior third. Protarsus, meso- tarsus, and metatarsus with basal segment shortest; second segment of mesotarsus and metatarsus slightly longer than third and fourth segments combined; fifth segment j Chaetarthria ayacuchana, n. sp.; biotope. ““Tobogan” area, 40 km south of Puerto Ayacucho, T.F. slightly shorter than second segment. Meta- trochanter moderately elongate and sparse- ly pubescent. Abdomen. — First and second abdominal sterna with a common, deep concavity. First sternum shiny; with numerous, very long, golden, hairlike setae along anterior margin and extending posteriorly as far as third sternum and holding a hyaline mass in con- cavity. Midline feebly raised longitudinally on first and second sterna. Sterna 3, 4, and 5 covered with short, sparse, golden setae; fifth sternum rounded, not emarginate ap- icomedially. Male genitalia.—As illustrated (Figs. 3, 4). Female. —Similar to male except length of female is greater. Type data. —Holotype: VENEZUELA: Territorio Federal Amazonas: Puerto Aya- cucho (40 km S) at ““Tobogan,”’ 22 Jan 1985, P. J. and P. M. Spangler, R. A. Faitoute, W. E. Steiner; deposited in the National Mu- seum of Natural History, Smithsonian In- stitution. Allotype: Same data as holotype. Paratypes: Same data as holotype, 3 males, 4 females; same locality, 25 Feb 1986, P. J. VOLUME 99, NUMBER 3 Spangler, 1 male; same locality, 27 Feb 1986, P. J. Spangler, 1 male. Paratypes deposited in the Instituto de Zoologia Agricola, Fa- cultad de Agronomia, Maracay, Venezuela, and the British Museum (Natural History), London. Etymology. —The trivial epithet ayacu- chana (adjective) is named for the town near which this new species was collected. Habitat. —Enroute to Cerro de la Nebli- na, the participants in the expedition had to layover in Puerto Ayacucho for two days awaiting air transportation. During that time we visited an interesting area about 40 km south of Puerto Ayacucho known locally as the ““Tobogan’’; so named because water running Over an extensive area of bedrock had polished the rock and formed a slide- like channel on one side of the outcropping. The water dropped into a pool at the base of the slide area and drained away in a shal- low stream with a sandy substratum. Col- lections of aquatic insects were made from the stream and its sandy margins. The type specimens of C. ayacuchana were collected from the sandy margins in the sunny part of the stream before it entered the shade of the forest (Fig. 5). Colorimetric water chem- istry tests provided the following data: Oxy- gen, 15 ppm; pH, 6; hardness, 0. The water temperature was 28°C and the air temper- ature was 37°C at the time the data were collected. Chaetarthria gavilana, new species Figs. 6-8 Holotype male.—Body form: Very con- vex, hemispherical. Length, 1.54 mm; greatest width, 1.04 mm, at about mid- length. Color.—Black dorsally except reddish brown around margins of pronotum. Ven- tral surface of head, thorax, abdomen and appendages reddish brown. Head.—With moderately coarse, sparse punctures; punctures between eyes separat- ed by 4 to 6 times their diameter; slightly 513 Figs. 6 and 7. Chaetarthria gavilana, n. sp.; male genitalia. 6, Dorsal view. 7, Lateral view. alutaceous along inner margin of each eye. Clypeus with lateral margins moderately ar- cuate and rimmed; anterior margin shallow- ly, broadly emarginate. Labrum more dis- tinctly punctate than head; almost truncate; with short, dense fringe of setae antero- medially. Eyes almost round viewed dor- sally but oblong viewed laterally; shallowly emarginate where clypeus extends into an- terior margin of eye. Ventral surface of head microalutaceous behind eyes. Mentum and submentum finely, sparsely punctate be- tween maxillae. Antenna, 8 segmented; bas- al segment long, sinuous; second segment almost globular; third, fourth, and fifth seg- ments slender, platelike; sixth, seventh, and eighth segments increasingly larger; sixth segment with few setae apicolaterally; sev- enth and eighth segments with long, rather dense setae; eighth segment ending in nar- row elongate apical projection longer than swollen basal part. Maxillary palpus, 4 seg- mented; basal segment tiny; second (pseu- dobasal) segment swollen apically and al- most as long as apical segment; third segment slightly more than half as long as second segment; apical segment about a sixth long- 514 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. Ayacucho, T.F. Amazonas, Venezuela. er than second segment; pseudobasal seg- ment when lying along stipes not attaining base of stipes. Labial palpus, 3 segmented; basal segment very small; second segment longest; both basal and second segments bearing numerous, long, golden setae es- pecially on medial and lateral surfaces; api- cal segment about one-fourth shorter than second segment and bearing several, long, golden setae subapically and 3 long, golden, hairlike setae dorsally on apex. Thorax. —Pronotum strongly convex; sides, posterolateral angles, and anterolat- eral angles moderately rounded; lateral margins finely rimmed; anterior margin ar- cuate medially and angulate behind eyes; punctures on disc moderately coarse, sparse, separated by 4 to 8 times their diameter. Scutellum an equilateral triangle; very finely and sparsely punctate. Elytron 1.15 mm long; strongly convex; widest near mid- length; distinctly rimmed along base later- ally; lateral margins finely rimmed, slightly sinuate in lateral view; seta-bearing punc- tures on disc coarse, distinct, and rather Chaetarthria gavilana, n. sp.; biotope. Cano near the village of Gavilan, 35 km southeast of Puerto dense and much more coarse and dense than those on pronotum, separated by 3 to 4 times their diameter; punctures on sides of elytra very coarse; sutural stria deep, extending and widening from slightly before mid- length to apex; sides strongly declivous, without obvious epipleura. Metathoracic wings present. Prosternum, mesosternum, and metasternum mostly glabrous and shiny; metasternal disc with sparse, fine punctures. Prosternum narrow in front of procoxae. Mesosternum with small, transverse, cari- nate process between and slightly in front of mesocoxae. Metasternum with slightly raised discal area; sides shallowly de- pressed. Procoxa with 3 very stout spines apicoventrally. Profemur and mesofemur with dense, hydrofuge pubescence on basal two-thirds; apices glabrous. Metafemur with fine, sparse, seta-bearing punctures margin- ally along anterior third. Protarsus, meso- tarsus, and metatarsus with basal segment shortest; second segment of mesotarsus and metatarsus slightly longer than third and fourth segments combined; fifth segment VOLUME 99, NUMBER 3 slightly shorter than second segment. Meta- trochanter moderately elongate and sparse- ly pubescent. Abdomen. — First and second abdominal sterna with a common, deep concavity. First sternum shiny; with numerous, very long, golden, hairlike setae along anterior margin and extending posteriorly as far as third sternum and holding a hyaline mass in con- cavity. Midline feebly raised longitudinally on first and second sterna. Sterna 3, 4, and 5 covered with short, sparse, golden setae; fifth sternum rounded, not emarginate ap- icomedially. Male genitalia.—As illustrated (Figs. 6, 7). Female. —Unknown. Type data. —Holotype: VENEZUELA: Territorio Federal Amazonas: Puerto Aya- cucho (35 km SE), Gavilan area, 20 Feb 1986, P. J. Spangler and W. Sanchez; de- posited in the National Museum of Natural History, Smithsonian Institution. Etymology. —The trivial epithet gavilana (adjective) is named for the Indian village and river of the same name near which this new species was collected. Habitat. —The specimen described above was collected from the sandy margins of an unnamed cano 2 km west of the village of Gavilan (35 km southeast of Puerto Aya- cucho). The cano (Fig. 8) was about 0.5 m deep and 1 m wide. Colorimetric water chemistry tests provided the following data: Oxygen, 12 ppm; pH, 6; hardness, 0. The water temperature was 27°C and the air temperature was 37°C at the time the data were collected. The following key should serve to distin- guish the five species which key to couplet 41 in Miller’s (1974) key. 1. Head, pronotum, and elytra shiny; punctures indistinct, extremely fine and sparse. Male genitalia as illus- trated (Figs. 1, 2). Guyana SD eh a porknockeri, new species — Head, pronotum, and elytra dis- tinctly punctate 2. Elytra with moderately coarse, moderately sparse, seta-bearing punctures on discal area; lateral punctures very coarse and dense, es- pecially at midlength; setae ar- ranged in widely separated rows. Male genitalia as illustrated (Figs. 6, 7). Venezuela .. gavilana, new species — Elytra with fine or moderately coarse and moderately sparse, seta-bearing punctures on discal area; lateral punctures moderately coarse at midlength; setae in narrowly sepa- rated rows or scattered over surface 3. Elytra with rows of moderately coarse, dense, seta-bearing punc- tures Over entire surface; setae very dense; rows of punctures becoming substriate toward apex. Brazil .... PBN RN ME granulata Miller — Elytra with fine or moderately coarse punctures in indistinct rows or scat- tered over surface; without indica- tions of any striae except sutural stria 4. Elytra with discal area finely, sparse- ly punctate; moderately coarsely punctate laterally and toward apex. Brazile. © aia lee tects brasilia Miller — Elytra with discal area moderately coarsely, sparsely punctate; punc- tures becoming coarse laterally and toward apex. Male genitalia as il- lustrated (Figs. 3, 4). Venezuela .. ayacuchana, new species Acknowledgments The Earthwatch expedition to Guyana was organized by Margaret Collins and sup- ported by the Center for Field Research, Boston, Massachusetts. The following par- ticipants also provided generous support and field assistance: John Byrd, Robin Faitoute, Martin Hegyi, James Hill, Fred Holtzclaw, William Johnson, Michael Levine, Molly Levine, Philip Perkins, Warren Steiner, 516 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Matthew Stevens, and Edouard Vystricil. I am deeply grateful and thank all of the con- tributors for their help in making that field- work possible. The expedition to Cerro de la Neblina was organized and directed by the Foundation for the Development of Physics, Mathe- matics, and Natural Sciences of Venezuela, with the patronage of the following Vene- zuelan institutions—the Ministry of Edu- cation, the Ministry of the Environment, the Venezuelan Air Force, the National Council of Scientific and Technological Re- search, and National Institute of Parks. The expedition was coordinated by Charles Brewer Carias and was conducted in col- laboration with the National Science Foun- dation of the United States, the American Museum of Natural History, the Field Mu- seum of Natural History, the Missouri Bo- tanical Garden, the New York Botanical Garden, and the Smithsonian Institution; biologists from several universities and oth- er institutions also participated. I thank all of the above organizations and their ad- ministrators for their extensive contribu- tions to this biotic survey. I also thank the following for their assis- tance: Robin A. Faitoute, Phyllis M. Span- gler, and Warren E. Steiner for collecting and preparing specimens during the expe- dition to Cerro de la Neblina; Young T. Sohn, biological illustrator, for the pen and ink drawings; the administrators of the Smithsonian Institution’s Scholarly Re- search Fund and the Research Opportu- nities Fund for supporting the fieldwork in Venezuela; and Phyllis Spangler for typing the manuscript into the word processor. Literature Cited Miller, David C. 1974. Revision of the New World Chaetarthria (Coleoptera: Hydrophilidae). — Entomologica Americana 49(1):1-123. Spangler, Paul J. 1977. Three new Ecuadorian species of the aquatic beetle genus Chaetarthria (Co- leoptera: Hydrophilidae).— Proceedings of the Biological Society of Washington 90(3):566—578. PROC. BIOL. SOC. WASH. 99(3), 1986, pp. 517-543 ANALYSIS OF STONE CRABS: MENIPPE MERCENARIA (SAY), RESTRICTED, AND A PREVIOUSLY UNRECOGNIZED SPECIES DESCRIBED (DECAPODA: XANTHIDAE) Austin B. Williams and Darryl L. Felder! Abstract.—The stone crab, Menippe mercenaria (Say, 1818) sensu lato, in the Caribbean and Carolinian Provinces of the western North Atlantic, is divisible into two morphologically distinct populations with almost separate, narrowly overlapping geographic ranges. These populations are here recognized as distinct species. The species differ in color, carapace morphometry, and stridulatory patches on chelae of the chelipeds. Five measured factors subjected to discriminant function analysis give good separation of the species at P < 0.05 for all specimens measured, and at P < 0.01 for specimens more than 15 mm in carapace length. Menippe mercenaria, restricted, ranges from Cape Lookout, North Carolina, through peninsular Florida, the Bahamas and Greater Antilles, to Yucatan peninsula, Mexico, and Belize. Menippe adina, new species, ranges from northwestern Florida around the Gulf of Mexico to Tamaulipas State, Mexico. The two species hybridize in the Apalachee Bay region of north- western Florida. Recent field studies in the Gulf of Mexico indicate that stone crabs assignable to Men- ippe mercenaria (Say, 1818), sensu lato, from the northwestern Gulf differ in color and morphometry from those found in penin- sular Florida, and furthermore, that indi- viduals exhibiting characters intermediate between these groups can be observed in the Apalachee Bay region of northwestern Flor- ida. Questions arise from these recent sur- veys concerning variation over the range as a whole, i.e., whether the preliminary ob- servations in the Gulf of Mexico were com- prehensive enough to have validity, and whether still other populations within the range may exhibit character states with suf- ficient cohesion to be recognized as taxo- nomically distinct. The chelipeds (claws) of this large crab are prized as a delicacy throughout a geo- ! Order of authorship was decided by flip of a coin. graphic range that extends through the Ca- ribbean and Carolinian Provinces of the western North Atlantic Ocean from Cape Lookout, North Carolina, southward around peninsular Florida, through the Bahamas and Greater Antilles, to the Yucatan pen- insula and Belize, and around the northern and western margin of the Gulf of Mexico to Tamaulipas State, Mexico, from intertid- al burrows, crevices and pools, to a depth of 51 m (Williams 1984, in part). Through most of that range, the crabs are incidentally taken for human consumption as a byprod- uct of other fisheries, but along western and especially southwestern Florida, they are the basis for a seasonal commercial fishery (Costello et al. 1979). Since Florida record- keeping began in 1962-63, the fishery grew to a peak annual landing of 2.6 million lb of claws during the 1981-82 season, and thereafter declined somewhat (preliminary estimate by National Marine Fisheries Ser- vice, Southeast Fisheries Center, 1.7 million 518 Fig. 1. Diagram of Menippe mercenaria (from Rathbun, 1884) showing carapace dimensions mea- sured in mm: cl = carapace length in midline; wbls = width to base of lateral spine, wls = width between tips of lateral spines (wls — whbls = length of al5); al3 = width of anterolateral tooth 3, al4 = width of antero- lateral tooth 4. lb for 1984-85). There is a sizeable fishery in Cuba as well where the crab is known as cangrejo moro (dappled crab). Interrelationships of populations of these crabs seem important from standpoints of both biology and fisheries, and it is therefore the purpose of this paper to analyze the pop- ulation structure from a systematic point of view. We have attempted to accomplish this by studying color and morphometry over selected parts of the range. Evaluation of these analyses demonstrates that two pop- ulations of the crabs can be distinguished. We interpret these populations to be two closely related species which hybridize in the panhandle region of northwestern Flor- ida, and we give diagnostic characters for each in comparative tables, graphs, illustra- tions, and discussion. Materials and Methods General observations of color over the entire geographic range of the species were Fig. 3. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Diagram showing inner surface of major chela bearing stridulatory patch, ¢ from USNM 7509, Key West, Florida. accomplished by DLF, recorded photo- graphically, and comparatively analyzed. Color pattern on the dactyl of the major chela was studied on specimens in the field, in museum collections, and on photographs by both of us. Measurements (Fig. 1) to the nearest 0.1 mm were recorded for carapace length in the midline, carapace width to the base of the notch between the 4th and 5th anterolateral teeth, carapace width includ- ing the 5th anterolateral tooth, and widths of the 3rd and 4th anterolateral teeth. Mea- surements of specimens from the northern Gulf of Mexico taken by DLF and of spec- imens in the crustacean collection of the National Museum of Natural History (USNM) taken by ABW were combined with a comparable data set from Charleston, South Carolina, plus a few measurements from specimens in other museums, photo- graphs, and illustrations in literature, for combined statistical analysis. Density of parallel striae/mm in the stridulation patch on the inner surface of the major chela, and the maximum height of that chela, were re- corded for representative populations by DLF (Fig. 2). Statistical tests were performed on the Honeywell computer of the Smithsonian In- = Menippe adina. Louisiana.— a, 6 cw 61.7 mm, Bayou Fourchon, Fourchon Parish; b, ? unmeasured, Cameron, Cameron Parish M. adina Fig. 6. Comparisons of morphological variables in Menippe populations assigned to M. adina (crosses), M. mercenaria (circles), and the hybrid zone (asterisks). A, Width anterolateral tooth 3 (AL3) as a function of ao Atypical coloration of some specimens from the northeastern Gulf (Fig. 3c—f) first led us to suspect hybridization between populations in northwestern Florida. As early as 1979, we recognized the Apalach- icola to Apalachee Bay region as the general area of transition or break between mor- phological extremes of “M. mercenaria,” and noted that these two extremes appeared to differ somewhat in life history and hab- itat. However, not until adequate samples were measured and photographed did we conclude that introgressive hybridization between species was occurring there; si- multaneously, an independent study of iso- zyme variations in populations of Menippe resulted in similar conclusions regarding populations in northwestern Florida (Bert, in press). Also supportive of this conclusion was evidence that stone crab fishermen in northwestern Florida have long recognized that a hybrid is involved in the fishery there. When asked to comment on coloration of local stone crabs in his catch (but otherwise unprompted), Mr. Clint Wood of Taylor Co., Fla., volunteered the following: “There is the ‘dominicker’ . . . spotted and with marks on the legs, there is the ‘stone crab’ . .. more solid brown . . . they catch more of them further to the west, and there is the ‘cross’ . a mix of the other two.” He further noted that the ““dominicker”’ was the only kind found south of Cedar Keys. Small juvenile to large adult specimens were included in our two-group discrimi- —_— width anterolateral tooth 4 (AL4); MZ. mercenaria, Y = 0.46776 + 1.18854 X; M. adina, Y = 0.66283 + 0.93972 X; M. hybrid, Y = 0.36809 + 1.05838 X; B, Width AL4 as a function of carapace length (CL); M. mercenaria, Y = —0.67632 + 0.18536 X; M. adina, Y = —0.73003 + 0.22309 X; M. hybrid, Y = —0.39580 + 0.20494 X; C, Length AL5 (equal to WLS — WBLS in Fig. 1) as a function of CL; M. mer- cenaria, Y = —0.25118 + 0.04756 X; M. adina, Y = —0.45164 + 0.08246 X; M. hybrid, Y = —0.11821 + 0.05568 X. All measurements in mm and taken as indicated in Fig. 1. 532 160 140 A 120 100 e 80 : L Ss 60 + 40+ oO 8 20 + * o 10 30 i 50 ; 70 90 110 Oo 20 40 60 80 100 CL 40; oOo : B 35+ ie) ce) 30 + one T) *o +; s 20t «¢ ° T : R ° 15+ 10+ ait Ait 10) Fig. 7. Comparisons of morphological variables in Menippe populations assigned to M. adina (crosses), M. mercenaria (circles), and the hybrid zone complex (asterisks). All measurements in mm. A, Width of cara- pace excluding anterolateral tooth 5 (WBLS) as a func- tion of carapace length (CL), taken as indicated in Fig. 1; M. mercenaria, Y = —1.63382 + 1.42285 X; M. adina, Y = 0.04080 + 1.38674 X; M. hybrid, Y = —0.47386 + 1.40583 X; B, Number of striae per mm (STR) in stridulatory patch on major chela asa function of maximum chela height (CHH), as shown in Fig. 2; M. mercenaria, In Y = 4.07954 — 0.92235 In X; M. adina, In Y = 4.60272 — 0.94048 In X; M. hybrid, In Y = 4.07787 — 0.81343 In X. nant analysis of M. mercenaria and M. ad- ina, and subsequent assignment of the sam- ple from the hypothesized hybrid zone (Panama City to Steinhatchee, Fla.) (Fig. 8B). The analysis included five variables, all PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of which provided for an increase in group separation. The analysis correctly classified 97% of individuals assigned a priori to Group | (M. mercenaria) or Group 2 (M. adina), thus demonstrating morphological distinction of these species over a broad range of size classes. The resultant canonical correlation of 0.883 indicates that about 78% of the separation between Groups 1 and 2 is thus explained. Exclusion of individuals less than 15 mm carapace length from the analysis improved correct classification to over 99% on the basis of the same five vari- ables (data not figured). Superposition of a sample from the hy- pothesized hybrid zone (Fig. 8B, upper axis) on discriminant scores from populations as- signable to M. mercenaria and M. adina (lower axis) defines a broadly intermediate but morphologically heterogeneous group which appears to include M. mercenaria, M. adina, hybrids of the two, and possible backcrosses. However, when discriminant scores for this group are independently sep- arated into subgroups based upon color, they resolve into means a, b, and c as indicated, where color morph “‘a”’ matches color of M. adina, “‘c’’ matches color of M. mercenaria and “‘b”’ is intermediate. The significance of differences among these three morphs is made clear by statistical comparisons among sample means of their discriminant scores: a, xX—.79063 = SD .86266\ vsbieslSs25—= SD 1.2334, t= —5.071, 53 DE Bwsiers 1.76489 + SD .92149, t = —4.165, 40 DF; IP (Me The sample comprising subgroup “a” (Fig. 8B) of the hybrid zone population in our analysis is slightly larger than those com- prising subgroups “‘b” and “‘c,”’ and likely includes a few individuals from just west of the area in which introgression of the two species occurs. This consists of 11 speci- mens from the vicinity of Panama City, Fla. In retrospect, all the latter are assignable to M. adina, and we have to date obtained no hybrid specimens from west of Cape San Blas. Thus, while we include specimens from ss (2) = I5 ag Li aa) = =) Zz -40 -2.0 O 20 40 DISCRIMINANT SCORES = (@ = O Lu aa) = =) = -4.0 -2.0 e) 2.0 4.0 6.0 DISCRIMINANT SCORES Fig. 8. A, Distribution of discriminant scores based upon R1, R2, R3 and R4 for specimens =15 mm carapace length: lower axis—a priori geographic Groups 1 and 2 (arrows mark centroids), assignable to M. mercenaria (cross-hatched bars) and M. adina (open bars), respectively; upper axis—ungrouped sample from Charleston, S.C. (arrow » ———_, and C. E. Kalman. 1974. Claw ex- traction during molting ofa stone crab, Menippe mercenaria (Decapoda, Brachyura, Xanthi- dae).— Florida Department of Natural Re- sources, Florida Marine Research Publications 4:5 pp. ; , and 1975. An analysis of stone crab (Menippe mercenaria) landings on Florida’s west coast, with a brief synopsis of the fishery.—Florida Department of Natural Re- sources, Florida Marine Research Publications 13:37 pp. 541 Say, T. 1818. An account of the Crustacea of the United States. —Journal of the Academy of Nat- ural Sciences of Philadelphia 1, Part 2:235-253, 313-319, 374-401, 423-444, 445-458. Schlieder, R. A. 1980. Effects of desiccation and au- tospasy on egg hatching success in stone crab, Menippe mercenaria.—Fishery Bulletin 77(3): 695-700. Schroeder, W. C. 1924. Fisheries of Key West and the clam industry of southern Florida. — United States Department of Commerce Bureau of Fisheries, Report of the United States Com- missioner of Fisheries for the Fiscal Year 1923, with Appendixes. Appendix XII:1—74. Scotto, L.E. 1979. Larval development of the Cuban stone crab, Menippe nodifrons (Brachyura, Xan- thidae), under laboratory conditions with notes on the status of the family Menippidae. — Fish- ery Bulletin 77(2):359-386. Siebenaler, J. B. 1952. Studies of “trash” caught by shrimp trawlers in Florida.— Proceedings of the Gulf and Caribbean Fisheries Institute, Fourth Annual Session, Miami Beach, November, 1951: 94-99. Simmons, E. G. 1957. An ecological survey of the upper Laguna Madre of Texas. — Publications of the Institute of Marine Science, University of Texas 4(2):156—200. Simonson, J.L. 1985. Reversal of handedness, growth, and claw stridulatory patterns in the stone crab Menippe mercenaria (Say) (Crustacea: Xanthi- dae). — Journal of Crustacean Biology 5(2):281- 293. , and P. Steele. 1980. Menippe mercenaria (stone crab) claw regeneration with reversal. — Florida Scientist 43(supplement):6. [abstract] , and 1981. Cheliped asymmetry in the stone crab Menippe mercenaria with notes on claw reversal and regeneration. — Northeast Gulf Science 5(1):21-30. Sinclair, M.E. 1977. Agonistic behaviour of the stone crab, Menippe mercenaria (Say).— Animal Be- haviour 25(1):193-207. Springer, S., and H. R. Bullis, Jr. 1956. Collections by the Oregon in the Gulf of Mexico. List of crustaceans, mollusks, and fishes identified from collections made by the exploratory fishing ves- sel Oregon in the Gulf of Mexico and adjacent seas 1950 through 1955.—United States Fish and Wildlife Service, Special Scientific Report, Fisheries No. 196:134 pp. Stephenson, T. A., and A. Stephenson. 1952. Life between tidemarks in North America. II. North- ern Florida and the Carolinas.— The Journal of Ecology 40(1):1—49, pls. 1-6. Stewart, V. N. 1972. Observations on the potential use of thermal effluent in mariculture.—Pro- 542 ceedings of the Third Annual Workshop, World Mariculture Society, St. Petersburg, Florida, pp. 173-178. Stimpson, W. 1859. Notes on North American Crus- tacea, No. 1.—Annals of the Lyceum of Natural History of New York 7[for 1862](2):49-93[1- 47], pl. 1. 1871. Notes on North American Crustacea in the Museum of the Smithsonian Institution. No. IIJ.—Annals of the Lyceum of Natural His- tory of New York 10(6):92-136 [119-163]. Suarez Alvarez, G., and R. Xiqués Diaz. 1969. Phys- iological aspects of some marine invertebrates of commercial interest in Cuba.—FAO Fisher- ies Report 71.1:155. , and 1971. Aspectos fisioecologicos de algunos invertebrados marinos de interes commercial para Cuba.—FAO Fisheries Re- ports 71.2:279-281. Sulkin, S. D., and W. F. Van Heukelem. 1980. Eco- logical and evolutionary significance of nutri- tional flexibility in planktotrophic larvae of the deep sea red crab Geryon quinquedens and the stone crab Menippe mercenaria. — Marine Ecol- ogy, Progress Series 2(2):91—96. Sullivan, J. R. 1979. The stone crab Menippe mer- cenaria in the southwest Florida USA fishery. — Florida Department of Natural Resources, Flor- ida Marine Research Publications 36:37 pp. Sushchenya, L. M., and R. Claro. 1966. Quantitative regularities of feeding and their connection with the balance of energy of the commercial crab Menippe mercenaria (Say). In Issled. Tsen- tral’no-Amerik. Morei [Soviet-Cuban Marine Exped. 1964-65].—Akademiya Nauk Ukrain SSR, Inst. Biol. Yuzhykh Morei A. O. Koval- evsk, Kiev, 1:217—230 [in Russian with English and Spanish summaries]. , and 1973. Quantitative regularities of feeding and their connection with the balance of energy of the commercial crab Menippe mer- cenaria (Say). Investigations of the Central American seas. [Published for the Smithsonian Institution and the National Science Founda- tion, Washington, D.C. by the Indian National Documentation Centre, New Delhi, pp. 311- 335.] [Translated from Russian by Fisheries Re- search Board of Canada, ““Naukova Dumka,” Kiev, 1966.] , and R. Claro Madruga. 1967. Datos cuan- titativos de la alimentacion del cangrejo co- mercial Menippe mercenaria (Say) y su relacion con el balance energético del mismo. — Estudios Instituto de Oceanologia, La Habana 2(1):75— 97 [in Spanish with English summary]. Tabb, D. C., and R. B. Manning. 1961. A checklist of the flora and fauna of northern Florida Bay and adjacent brackish waters of the Florida PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mainland collected during the period July, 1957 through September, 1960.— Bulletin of Marine Science of the Gulf and Caribbean 11(4):552- 649. , and 1962. Part II. Aspects of the biology of northern Florida Bay. Jn D. C. Tabb, D. L. Dubrow, and R. B. Manning, The ecology of northern Florida Bay and adjacent estuaries, pp. 39-79.—State of Florida Board of Conser- vation, Technical Series 39:1-81. Telford, M. 1982. Chapter 10. Shrimps, lobsters, and crabs. Jn E. H. Kaplan, ed., A field guide to coral reefs of the Caribbean and Florida, pp. 150-168. The Peterson Field Guide Series, Houghton Mifflin Co., Boston, xv + 189 pp. Tucker, R. K. 1978. Free amino-acids in developing larvae of the stone crab Menippe mercenaria. — Comparative Biochemistry and Physiology 60A(2):169-172. Vernberg, F. J. 1956. Study of the oxygen consump- tion of excised tissues of certain marine decapod Crustacea in relation to habitat.— Physiological Zoology 29(3):227—234. Walker, J. F. 1953. A check list of marine inverte- brates collected at the Gulf Coast Research Lab- oratory from 1947 through 1950.—Journal of the Mississippi Academy of Science 4:192—207. Warner, G. F. 1977. The biology of crabs. Elek Sci- ence, London, Van Nostrand Reinhold Com- pany, New York, xv + 202 pp. Wass, M. L. 1955. The decapod crustaceans of Al- ligator Harbor and adjacent inshore areas of northwestern Florida.—The Quarterly Journal of the Florida Academy of Sciences 18(3):129- 176. Weiss, H. B., and G. M. Ziegler. 1931. Thomas Say, early American naturalist. Charles C. Thomas, Publisher, Springfield, Illinois, and Baltimore, Maryland, xiv + 260 pp. Wenner, E. L., and A. D. Stokes. 1983. Preliminary observations on the distribution and abundance of the stone crab, Menippe mercenaria, in South Carolina waters.—South Carolina Marine Re- sources Center, Technical Report 55:28 pp. Whetstone, J. M., and A. G. Eversole. 1978. Pre- dation on hard clams, Mercenaria mercenaria, by mud crabs Panopeus herbstii.— Proceedings of the National Shellfish Association 68:42-48. Whitten, H. L., H. F. Rosene, and J. W. Hedgpeth. 1950. The invertebrate fauna of Texas coast jetties; a preliminary survey.— Publications of the Institute of Marine Science, University of Texas 1(2):53-87, 1 pl. Williams, A. B. 1965. Marine decapod crustaceans of the Carolinas.—Fishery Bulletin 65(1):xi + 298 pp. 1984. Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine VOLUME 99, NUMBER 3 to Florida. Washington, D.C., Smithsonian In- stitution Press, xviii + 550 pp. , and T. W. Duke. 1979. Crabs (Arthropoda: Crustacea: Decapoda: Brachyura). Jn C. W. Hart and S. M. Fuller, eds., Pollution ecology of es- tuarine invertebrates. Chapter 6, pp. 171-233. Academic Press, New York, 406 pp. Yang, W. T. 1971. Preliminary report on the culture of the stone crab.—Proceedings of the Second Annual Workshop, World Mariculture Society. Louisiana State University, Division of Con- tinuing Education, Baton Rouge, pp. 53-54. 1972. Notes on the successful reproduction of stone crabs, Menippe mercenaria (Say) reared from eggs.—Proceedings of the Third Annual Workshop, World Mariculture Society, St. Pe- tersburg, Florida, January 26—28, 1972, pp. 183— 184. —, andG. E. Krantz. 1976. “Intensive” culture of the stone crab, Menippe mercenaria. —Uni- versity of Miami Sea Grant Program, 15 pp. (mimeo). Young, A. M. 1978. Superorder Eucarida, order De- capoda. Jn R. G. Zingmark, ed., An annotated 543 checklist of the biota of the coastal zone of South Carolina, pp. 171-185. University of South Car- olina Press, Columbia, S.C., xii + 364 pp. Zeiller, W. 1974. Tropical marine invertebrates of southern Florida and the Bahama Islands. Wiley Interscience, New York, xii + 132 pp. Zuboy, J. R., and J. E. Snell. 1980. Assessment of the Florida stone crab fishery. —NOAA Tech- nical Memorandum, NMFS-SEFC-21:32 pp. , and . 1982. Assessment of the Florida stone crab fishery, 1980-81 season.—NOAA Technical Memorandum, NMFS-SEFC-79:21 pp. (ABW) National Marine Fisheries Ser- vice, Systematics Laboratory, National Mu- seum of Natural History, Washington, D.C. 20560; (DLF) Department of Biology and Center for Crustacean Research, University of Southwestern Louisiana, Lafayette, Lou- isiana 70504. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE % BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD, LONDON, SW7 5BD 14 April 1986 The Commission hereby gives six months notice of the possible use of its plenary powers in the following cases, published in the Bulletin of Zoological Nomenclature, volume 43, part 1 on 9 April 1986, and would welcome comments and advice on them from interested zoologists. Correspondence should be addressed to the Executive Secretary at the above address, if possible within six months of the date of publication of this notice. Case No. 2472 Dasyurus hallucatus Gould, 1842 (Mammalia, Marsupialia): proposed conservation by the suppression of Mustela quoll Zimmermann, 1783. 2485 Cholus Germar, 1824 (Insecta, Coleoptera): proposed conservation by the suppression of Archarias Dejean, 1821. 2486 Dryophthorus Germar, 1824 (Insecta, Coleoptera): proposed conservation by the suppression of Bulbifer Dejean, 1821. 2487 Lachnopus Schoenherr, 1840 (Insecta, Coleoptera): proposed conservation by the suppression of Menoetius Dejean, 1821 and Ptilopus Schoenherr, 1823. 2488 Nemocestes Van Dyke, 1936 (Insecta, Coleoptera): proposed conservation and desig- nation of type species. 2489 Zygops Schoenherr, 1825 (Insecta, Coleoptera): proposed conservation by the suppres- sion of Eccoptus Dejean, 1821. 2505 Tylocidaris Pomel, 1883 (Echinoidea, Cidaroidea): proposed designation of Cidaris clavigera Mantell, 1822 as type species. 2479 Ammonites perarmatus J. Sowerby, 1822 (Cephalopoda, Ammonoidea): proposed ex- emption from the Principle of Homonymy. Clausilia Draparnaud, 1805 (Mollusca, Gastropoda): proposed correction of Opinion 119. De la Cépéde, 1788-1789, “Histoire naturelle des Serpens” and later editions: proposed rejection as a non-binominal work. ATYIDAE De Haan, [1849] (Crustacea, Decapoda) and ATYIDAE Thiele, 1926 (Mol- lusca, Gastropoda): proposals to remove the homonymy. Carcharias Rafinesque, 1810 (Chondrichthyes, Lamniformes): proposed conservation by the use of the relative precedence procedure. Pyralis nigricana Fabricius, 1794 (Insecta, Lepidoptera): proposed conservation by the suppression of Phalaena rusticella Clerck, 1759. Apanteles ornigis Weed, 1887 (Insecta, Hymenoptera): proposed conservation by the suppression of Microgaster robiniae Fitch, 1859. Strongylaspis Spaeth, 1936 (Insecta, Coleoptera) non Strongylaspis Thomson, 1860: proposed designation of Cassida atripes LeConte, 1859 as type species. Nomadacris Uvarov, 1923 (Insecta, Orthoptera): proposed conservation by setting aside the first-reviser action of Jago. Type species of the genus Ca/ymene Brongniart (Trilobita) in Brongniart & Desmarest, 1822 and proposed suppression of the name tuberculatus Briinnich, 1781. Tubulanus Renier, [1804] and 7. polymorphus Renier, [1804] (Polychaeta): proposed reinstatement under the plenary powers. P. K. TUBBS Executive Secretary INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE % BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD, LONDON, SW7 5BD 14 April 1986 The following Opinions have been published by the International Commission on Zoological Nomenclature in the Bulletin of Zoological Nomenclature, volume 43, part 1 on 9 April 1986. Opinion No. 1369 (p. 15) Astacilla Cordiner, 1793 (Crustacea, Isopoda): conserved. 1370 (p. 17) Neadmete okutanii Petit, 1974 designated as type species of Neadmete Habe, 1961 (Mollusca, Gastropoda). 1371 (p. 19) Pachycephalosaurus Brown & Schlaikjer, 1943 and Troodon wyo- mingensis Gilmore, 1931 (Reptilia, Dinosauria): conserved. 1372 (p. 21) Donax hanleyanus Philippi, 1847 (Mollusca, Bivalvia): conserved. 1373 (p. 23) Panesthia saussurii Wood-Mason, 1876 designated as type species of ; Caeparia Stal, 1877 (Insecta, Dictyoptera). 1374 (p. 25) Boga Fitzinger, 1826 (Reptilia, Serpentes): conserved. 1375 (p. 27) Glossodoris Ehrenberg, 1831, Hypselodoris Stimpson, 1855 and Chro- modoris Alder & Hancock, 1855 (Mollusca, Gastropoda): con- served. 1376 (p. 30) Cuspidaria (Rhinoclama) adamsi Morgan & Heppell, 1981 designated as type species of Rhinoclama Dall & Smith, 1866 (Mollusca, Bivalvia). 1377 (p. 33) Chelydra osceola Stejneger, 1918 given nomenclatural precedence over Chelydra laticarinata Hay, 1916 and Chelydra sculpta Hay, 1916 (Reptilia, Testudines). 1378 (p. 35) Phalaena bellatrix Stoll, 1780 designated as type species of Crinodes Herrich-Schaffer, 1855 (Insecta, Lepidoptera). 1379 (p. 37) Gonodontis rectisectaria Herrich-Schaffer, [1855] designated as type species of Pero Herrich-Schaffer, 1855 (Insecta, Lepidoptera). 1380 (p. 39) Euphaedra Hubner, [1819] (Insecta, Lepidoptera): conserved. 1381 (p. 42) Ourocnemis Baker, 1887 (Insecta, Lepidoptera): conserved. 1382 (p. 45) Zeugophora Kunze, 1818 (Insecta, Coleoptera): conserved. The Commission regrets that it cannot supply separates of Opinions. P. K. TUBBS Executive Secretary i, - fn a 7 5 7 _ . a my See | ; a a) : : i ; a iAaIOOICO.S 4 ye PARAL EY os os Lia! fe t ‘“ > MUPTR ROAR LATURE Oo pete EL OAT TR LATA, MC ee i Owe * OA Se | aut (eae =e wea i i nae +t i 7 1 var Lars » ai rf Se sare @ ‘ in ; * na ‘ > oa | / _ i may 1px) . / ae 4k95 6 \4i oh) aE Uy. ape le atanink, ‘IRE f s LhAY oe a, Lae | i “ee. ae } a ay mipelsrumees : sl ; : ah reer v nw : | , You y 4 j | ¥ i f 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 will be published in English (except for Latin diagnosis/description of plant taxa which should not be duplicated by an English translation), with a summary in an alternate language when appropmiate. Publication Charges.—Authors will be asked to assume publication costs of page-charges, tabular material, and figures, at the lowest possible rates. Submission of manuscripts.—Manuscripts should be sent to the Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History, 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. Manuscripts will be reviewed in order of receipt by a board of associate editors and appropriate referees. Presentation.—Clarity of presentation, and requirements of taxonomic and nomenclatural procedures necessitate reasonable consistency in the organization of papers. Telegraphic style is recommended for descriptions. Literature should be cited in abbreviated style (author, date, page), except in botanical synonymies, with full citations of journals and books (no abbrevia- tions) in the Literature Cited section. The establishment of new taxa must conform with the requirements of the appropriate in- ternational codes of nomenclature. Authors are expected to be familiar with these codes and to comply with them. New species-group accounts must designate a type-specimen deposited in an institutional collection. The sequence of material should be: Title, Author(s), Abstract, Text, Acknowledgments, Literature Cited, Author’s (s’) Address(es), Appendix, List of Figures (entire figure legends), Figures (each numbered and identified), Tables (each table numbered with an Arabic numeral and heading provided). Manuscripts should be typed, double-spaced throughout (including tables, legends, and foot- notes) on one side of 842 x 11 inch sheets, with at least one inch of margin all around. Manu- scripts in dot-matrix will not be accepted. Submit a facsimile with the original, and retain an author’s copy. Pages must be numbered on top. One manuscript page = approximately 0.5 printed page. Underline singly scientific names of genera and lower categories; leave other indications to the editor. Figures and tables with their legends and headings should be self-explanatory, not requiring reference to the text. Indicate their approximate placement by a pencil mark in the margin of the manuscript. Illustrations should be planned in proportions that will efficiently use space on the type bed of the Proceedings (12.5 x 20 cm) and should not exceed 15 x 24 inches. Figures requiring solid black backgrounds should be indicated as such, but not masked. Art work wil! be returned only on request. Proofs.—Galley proofs will be submitted to authors for correction and approval. Reprint orders will be taken with returned proof. Costs.—Page charges @ $60.00, figures @ $10.00, tabular material $3.00 per printed inch. All authors are expected to pay the charges for figures, tables, changes at proof stage, and reprints. Payment of full costs will probably facilitate speedy publication. CONTENTS Speleobregma lanzaroteum, a new genus and species of Scalibregmatidae (Polychaeta) from a marine cave in the Canary Islands Rodney Duane Bertelsen Symmetroscyphus, a new genus of thecate hydroid (Family Thyroscyphidae) from Bermuda Dale R. Calder The status of the ophidiid fishes Ophidium brevibarbe Cuvier, Ophidium graellsi Poey, and Leptophidium profundorum Gill C. Richard Robins A redescription of Colobomatus mylionus Fukui from Australian Acanthopagrus (Sparidae) (Crustacea: Copepoda: Philichthyidae) Thomas Byrnes and Roger Cressey Neotropical Monogenea. 9. Status of Trinigyrus Hanek, Molnar, and Fernando, 1974 (Dac- tylogyridae) with descriptions of two new species from loricariid catfishes from the Brazilian Amazon D. C. Kritsky, W. A. Boeger, and V. E. Thatcher Redescription of Echinoderes pilosus (Kinorhyncha: Cyclorhagida) Robert P. Higgins Separation of Haber speciosus (Hrabé) (Oligochaeta: Tubificidae) from its congeners, with a description of a new form from North America Michael R. Milligan Redescription of the oligochaete genus Propappus, and diagnosis of the new family Propappidae (Annelida: Oligochaeta) Kathryn A. Coates Caridean shrimps of the Gulf of California. V. New records of species belonging to the subfamily Pontoniinae (Crustacea: Decapoda: Palaemonidae) Rubén Rios Two hitherto unnamed populations of Aechmophorus (Aves: Podicipitidae) Robert W. Dickerman The status of the callianassid genus Callichirus Stimpson, 1866 (Crustacea: Decapoda: Thalassi- nidea) Raymond B. Manning and Darryl L. Felder A new scale-worm commensal with deep-sea mussels in the seep-sites at the Florida Escarpment in the eastern Gulf of Mexico (Polychaeta: Polynoidae: Branchipolynoinae) Marian H. Pettibone A neotype designation for Petrolisthes tomentosus (Dana), and description of Petrolisthes heterochrous, new species, from the Mariana Islands (Anomura: Porcellanidae) Roy K. Kropp Three new species of Pilargidae (Annelida: Polychaeta) from the East coast of Florida, Puerto Rico, and the Gulf of Mexico Paul S. Wolf Epizoic barnacles on pleurodiran turtles: is the relationship rare? J. G. Frazier Psolus pawsoni (Echinodermata: Holothuroidea), a new bathyal sea cucumber from the Florida east coast John E. Miller and Richard L. Turner On the elevation of the Stephanophyllia subgenus Letepsammia to generic rank (Coelenterata: Scleractinia: Micrabaciidae) Joan Murrell Owens The taxonomic status of Glossophaga morenoi Martinez and Villa, 1938 (Mammalia: Chirop- tera: Phyllostomidae) Alfred L. Gardner Trinervate leaves, yellow flowers, tailed anthers, and pollen variation in Distephanus Cassini (Vernonieae: Asteraceae) Harold Robinson and Brian Kahn A new vulture (Vulturidae: Pliogyps) from the Late Miocene of Florida Jonathan J. Becker Three new species of water scavenger beetles of the genus Chaetarthria from South America (Coleoptera: Hydrophilidae) Paul J. Spangler Analysis of stone crabs: Menippe mercenaria (Say), restricted, and a previously unrecognized species described (Decapoda: Xanthidae) Austin B. Williams and Darryl L. Felder International Commission on Zoological Nomenclature: Notices and Opinions 374 380 384 388 392 399 406 417 429 435 437 444 452 . 464 472 478 486 489 493 502 509 517 544 a — — — THE BIOLOGICAL SOCIETY OF WASHINGTON 1985-1986 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 Kensley Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Richard P. Vari Insects: Robert D. Gordon 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. 99(4), 1986, pp. 547-573 A REVISION OF THE GENUS NEPHASOMA (SIPUNCULA: GOLFINGIIDAE) Norma J. Cutler and Edward B. Cutler Abstract. —The species in the sipunculan genus Nephasoma (formerly a sub- genus of Golfingia) are reviewed and evaluated in light of a critical examination of 13 historically used morphological characters. Whenever possible, type ma- terial was examined. The monograph of Stephen and Edmonds (1972) was used as a starting point. Of the 46 putative species listed in that work or described since then, 23 remain as valid species, three are reduced to subspecific rank, five are considered either species inquirendum or incertae sedis, and the remainder are considered junior synonyms. One new subspecies is described (N. diaphanes corrugatum). The known distribution of each species is sum- marized. A key to all species is provided. This paper is part of a continuing series begun in Cutler and Murina (1977) where all of the taxa then considered subgenera of Golfingia were reviewed in a preliminary fashion. The sipunculan genus Golfingia was created by Lankester in 1885 to commem- orate a pleasant outing on the greens. In 1950 W. K. Fisher created five subgenera including Phascoloides. One of the species in the group was G. (P). glacialis which Cut- ler and Murina (1977) subsequently showed to be conspecific with Nephasoma marinki Pergament. This species name is a junior synonym but, unfortunately, the genus- group name is senior to Phascoloides and therefore must be retained as the name of this taxon. The retention of Nephasoma as a separate monotypic genus by Stephens and Edmonds (1972) is unfortunate and poten- tially confusing. When Cutler and Gibbs (1985) elevated Nephasoma to generic rank it had a different meaning, i.e., a polytypic taxon with the species formerly included within Golfingia (Phascoloides). This con- fusion is further compounded since Gibbs (1982) determined that N. glacialis is a ju- nior synonym of N. /illjeborgi. This review is one of a series critically examining the morphological characters used by taxonomists to characterize species within a genus and evaluating their mean- ingfulness within populations made up of varying size and age worms. We have ex- amined the type material wherever possible and reevaluated the original descriptions based on our current understanding of a species as a group of reproductively isolated individuals within which there is variation. This effort has already been concluded for four of the six taxa considered Golfingia subgenera in Cutler and Murina (1977): Mitosiphon (Cutler 1979), Golfingiella and Siphonoides (Cutler et al. 1983), and Thysa- nocardia (Gibbs et al. 1983). The remaining group (Golfingia sensu stricto) is currently being revised and will be presented in the near future. The genus Nephasoma is more difficult than many because of the small size of adult worms (many with diameters less than 1 mm) and the paucity of obvious characters. Many of the ‘useful’ features are located on the distal end of the introvert which is only rarely extended in preserved material (hook size/shape/arrangement and tentacle num- ber/form). This genus is also very common (hundreds of individuals per sample) in deep-water collections made in recent years. 548 The need for a useful and biologically valid classification is especially pressing. This preliminary effort is directed towards this need. In the following text we use these abbre- viations for the indicated institutions: Zoo- logical Museum, University of Bergen (ZMUB); British Museum of Natural His- tory, London (BMNH); National Museum of Natural History, Washington (USNM); Zoological Institute, Academy of Sciences, Leningrad (ZIAS); Museum fur Natur- kunde, Berlin (MNHU),; Irish National Mu- seum, Dublin (INMD); Institut voor Tax- onomische Zodlogie, Amsterdam (ZMUA). Remarks on the ‘minutum’ Section Since the early part of this century biol- ogists have been confounded by one subset within this group of worms, 1.e., those with reduced tentacular lobes, small, scattered, transparent, possibly deciduous hooks, nor- mal body proportions, a nondistinct inter- nal anatomy, and no unique external fea- ture. This paper retains several names which have, at various times, been included in this complex. Now that N. minutum has been more narrowly defined (Gibbs 1975, 1977a) and N. abyssorum can be distinguished, we are left with N. diaphanes (with a new sub- species) and N. /Jilljeborgi. While these may be conspecific, only subsequent work using different methods of analysis will tell with any certainty. When one has only a few an- imals, some large and smooth and others small and papillated, it is easy to separate them into two sets. However, when one is working with hundreds of specimens the distinctiveness blurs and it becomes very difficult to draw a clear line. In terms of almost all the ‘standard’ characters, these taxa are identical, but after long deliberation we have decided to retain the two names (adding one subspecies) with much ambiv- alence. The written descriptons of these taxa cannot adequately communicate what one sees and is only an approximation. Anyone conducting zoogeographical PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON analyses of marine animals would be wise to exclude these species (as well as all their junior synonyms, especially N. minutum) from their works. There has been so much confusion over the years that it would be necessary to reexamine each collection in order to revalidate them within the current usage. Nephasoma Pergament, 1946 Type species. —Nephasoma marinki Per- gament, 1946 [=Onchnesoma glaciale Dan- ielssen & Koren: Cutler & Murina, 1977; =Phascolosoma lilljeborgii Danielssen & Koren: Gibbs, 1977] by monotypy. Diagnosis.—Species generally small- to medium-sized (trunk less than 5 cm in length). Introvert about equal to, or shorter than, trunk. Hooks present, usually scat- tered (arranged in rings in WN. rimicola (Gibbs), in spirals in N. abyssorum (Koren and Danielssen)). Body wall with continu- ous muscle layers. Oral disk carrying ten- tacles arranged around the mouth but ten- tacles may be reduced in both size and number and restricted to dorsal region. Two introvert retractor muscles often partially fused. Contractile vessel without villi. Spin- dle muscle not attached posteriorly. Two nephridia. This genus now contains all those species previously assigned to the Golfingia sub- genus Phascoloides Fisher, 1950, since Ne- phasoma Pergament has been shown to have priority over Phascoloides (Cutler and Mu- rina 1977). The endings of some species names have been modified in accordance with the ICZN as Nephasoma is a neuter name. Morphological Characters of Nephasoma In their description of species, biologists have used a variety of morphological char- acters to differentiate these entities. These descriptions were legitimate attempts but not always based on a good understanding of possible variation within a population (species). In this section we evaluate these VOLUME 99, NUMBER 4 Fig. 1. N. multiareneusa; D, N. confusum; E, N. constricticervix; F, N. abyssorum. All drawn to same scale. characters in light of our experience with large sample sizes. 1. Introvert hooks.—The presence or ab- sence of hooks on the distal portion of the introvert throughout the life of a worm has been presumed to be a species-specific char- acter by many, but not all authors (e.g., Ge- rould 1913; Selenka 1885; Southern 1913: 22). Our experience confirms the sugges- tions that while this may be true in some cases there are many species which have hooks as young individuals but lose them with age. For example, NV. abyssorum seems to retain its hooks throughout its life and N. eremita is said to never have hooks. Our concern, which can only be answered by breeding and rearing worms, is: If one had young N. eremita in hand which did have hooks, one would probably put another name on it, i.e., thet of a hooked species. Contrariwise, if there are large N. abysso- rum without hooks, one would be inclined 549 60 ym Introvert hooks of Nephasoma species; A, N. laetmophilum (after Fisher 1952); B, N. minutum; C, to identify these as some hookless species. An additional problem arises if one has an incomplete/damaged worm or one which is regenerating an introvert missing as the re- sult of predation. There are some useful attributes when hooks can be seen. A few species have very characteristic shapes or sizes (see N. con- fusum, N. constricticervix, or N. multiara- neusa) or unique arrangements (N. abys- sorum, N. rimicola). However, most species have small, bluntly triangular, transparent, scattered hooks about 20—40 um tall. There are six species with hooks 50-150 um tall (N. abyssorum, N. confusum, N. cutleri, N. laetmophilum, N. rimicola, N. schuttei), and only two with hooks frequently more than 150 um tall (NV. vitjazi and N. constricticer- VIX). While we are uncertain and skeptical, our operational assumption is that three sets of species exist within this genus: Those which 550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. never have hooks, those which have hooks throughout their lives, and those with de- ciduous hooks. To use the absence of hooks in a few individuals, by itself, to establish a new species would be a mistake. 2. Tentacles. —Most members of this ge- nus have one of two general types of ten- tacular arrays. About half the species have a ‘normal’ crown of flattened, digitate ten- tacles (more than 10, number increasing with age, Fig. 2A), while the other half have few (eight or less) short, lobate tentacles. Often these latter species exhibit only two tenta- cles plus four to six small lobes (Fig. 2B). There are two with unique arrangements (see Tentacles of Nephasoma species; A, N. rimicola; B, N. minutum (A and B from Gibbs 1977a); C, N. novaezealandiae (after Benham 1904); D, N. rutilofuscum (from Cutler and Cutler 1979a). N. novaezealandiae and N. rutilofuscum, Fig. 2C, D). In summary, this character can be helpful if it can be seen. A major problem is that most preserved specimens do not have their introverts extended and it is very dificult accurately to interpret the mor- phology of these structures in dissected ma- terial, especially since most have introverts less than 0.5 mm in diameter. 3. Caudal appendage. — This is an easily seen character. It is best exhibited in N. fla- griferum as a thin, rat-like tail. In N. bul- bosum it is less clearly set off and more like a narrowed tapered portion of the trunk, not an appendage. The posterior end of the trunk VOLUME 99, NUMBER 4 in the other species may form a conical point (like a blunt pencil) but not a tail. 4. Trunk length to width ratio.—These species fall into two general groups; elon- gate, slender worms (trunk length more than 10 times the width) and the shorter, stouter worms (length less than 6-8 times the width). Due to the ontogenetic or elastic properties of these worms there may be wide variation in this ratio. One must also be sure the trunk one is measuring has not been damaged/ truncated. Except in a very general way, this measurement is of minimal value to the tax- onomist and can be misleading if inter- preted too narrowly. 5. Introvert length.—This body region ranges from ¥, (in N. constricticervix) to twice the trunk length in a few species. Most species have introverts shorter than the trunk (0.4—0.8 x). When the introvert is re- tracted or only partially extended, this mea- surement will be shorter than it would be if the introvert was completely extended due to its extreme elasticity. This character can have limited usefulness if accurately deter- mined. Some confusion has resulted from how this measurement is made. Our prac- tice is to use the nephridiopores to mark the anterior end of the trunk so that anything anterior to this point is considered introvert (in taxa where the anus is anterior to the nephridiopores the anus is used instead). Earlier authors have been less explicit and therefore one cannot be certain what they considered to be the introvert. Another often overlooked issue is an ontogenetic one. It is clear that the trunk grows faster than the introvert, so that as a worm ages, the intro- vert appears to get shorter, i.e., represent a smaller fraction of the total length (see Ap- pendix). 6. Anus/nephridiopores relationship. — The middorsal anus usually is located at the anterior end of the trunk as are the pair of ventrolateral nephridiopores. There may be a small but measureable difference in the relative position of these openings. The ne- phridiopores are at essentially the level of the anus in most species in this genus. In 551 six species these pores are anterior to the anus, commonly by 3-10% of the trunk length (see N. flagriferum in Appendix). Seven species are reported to have their ne- phridiopores posterior to the anus. In small worms where this distance is less than 1 mm the decision is subjective as to whether or not they are at the same level. This can be a useful character for some species. 7. Papillae distribution, size, and shape. — As with other genera these secretory organs are more concentrated and larger at the two ends of the trunk, sparser and smaller in the center. There are two basic morphs in these taxa; most species have low, inconspicuous ‘skin bodies.’ There are only a few species with well developed, obvious, mammiform papillae. The shape of these structures, when present, is quite variable and should not be considered taxonomically useful. It seems likely that a genetic potential for producing papillae exists and the degree to which this is expressed is determined by the nature of the microhabitat an individual occupies, 1.e., analogous to human calluses. 8. Shields. —This term has caused much confusion within this phylum. In the As- pidosiphonidae there is a hardened epider- mal structure (calcium or scleroprotein?) at the anterior end of the trunk (sometimes posterior also). This same term has been used to describe a very different situation in this genus, i.e., an aggregation of close- packed papillae around the distal ends of the trunk giving it a dark, rugose appear- ance. This condition is very different from a shield as used in other genera and the use of this term is misleading and should not be used in this taxon. These epidermal pa- pillae may be gathered into longitudinal ridges and darkly pigmented (e.g., NV. vitjazi or N. wodjanizkii). In addition, under cer- tain conditions, the anterior end of the trunk is pulled in to give a flattened appearance accentuating this pseudoshield morph. 9. Spindle muscle. —In most sipunculans there is a thin thread-like muscle running through the gut coil and connecting to it at intervals. In some genera it extends through 552 and out of the posterior end anchoring the coil to the posterior end of the trunk. In many, it extends out of the anterior end of the coil, along the rectum, and attaches to the body wall just anterior to the anus. How- ever, there are several variations on this theme. In Nephasoma the muscle does not extend beyond the posterior end of the coil and in most species it does not extend be- yond the anterior end either. In a few species one can see the muscle coming out the an- terior end but it quickly terminates on the wall of the rectum. In a few (e.g., NV. flagrife- rum) this muscle is well developed, some- times branching, and extends from the rec- tum on to the body wall near or anterior to the anus. In a few species (e.g., N. abysso- rum, N. lilljeborgi) one can rarely find any trace of the spindle muscle. One does find variation within a population (see N. con- strictum in Appendix) and in small worms itis very difficult to find, even when present. Therefore, the taxonomic value of this char- acter is limited as most species have a sim- ilar condition, it is not well developed in small individuals, and it is not consistently present within a population so that its ap- parent absence in an individual can be mis- leading. 10. Retractor muscles’ point of origin. — The muscles that insert behind the ‘head’ and function to retract the introvert have their origins on the inner surface of the trunk wall. The position of this attachment along the anterior/posterior axis has been as- sumed to be of value. The most common condition is for these muscles to originate between 30-70% of the distance towards the posterior end of the trunk. Within a species this relative position may vary as the worm increases in size (appearing to move anteri- ad with an increase in trunk length, see Ap- pendix). There are four species having these origins in the 15-30% range (N. constricti- cervix, N. novaezealandiae, N. tasman- iense, N. vitjazi) and three with their origins between 75-90% of the distance to the pos- terior end of the trunk (N. filiforme, N. ru- tilofuscum, N. wodjanizkii). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 11. Intestinal coiling.— Most species have a tightly wound double helix but some of the elongate, slender species have this helix stretched out or loosely wound with space between the individual coils easily seen. This coiling seems to be a constant difference. What is not constant is the number of coils. This is clearly correlated to the size of the animal and not a species-specific character (see Appendix; also Ditadi and Migotto 1981). 12. Intestinal fixing muscles. —The num- ber of fine, thread-like muscles attaching the gut coil to the body wall varies from 0-7 according to published accounts. This has been alleged to be a species-specific char- acter but our experience suggests that A: these are very fragile structures and can be easily broken, and B: these are easily over- looked and even if present not mentioned by certain authors, and C: The number of muscles within one population does vary (see Appendix). In a population of 52 N. confusum examined by Ditadi and Migotto (1981) most had five of these muscles but the range was one to seven. Therefore, we place very little weight on this feature. 13. Rectal caecum. —This is not a useful character for two reasons. Given the small size of many of these worms and the diffi- culty of dissection, even if it was present it could be easily overlooked. Many authors simply do not mention its presence or ab- sence. Also, in populations where this is present some individuals may lack it— which does not exclude the individual from this species; there is variation. However, there are species which consistently lack one (N. constrictum) while others seem to consis- tently have one (N. flagriferum) so in a few particular cases it may be helpful as a di- agnostic character. Key to Nephasoma Species 1. Posterior end of trunk with caudal appendage (tail)? 2 eee DD — Posterior end of trunk rounded or blunthy;pomted sae eee 3 VOLUME 99, NUMBER 4 Ds . Trunk rusty-red Posterior end of trunk with large papillae, and tail thin, whiplike .. RP ran ce tttd Sedan cok N. flagriferum Posterior end of trunk without large papillae, and tail formed from nar- rowed portion of trunk .. N. bulbosum Ae he N. rutilofuscum Trunk white, yellow, or brown; not NO CME Arcee Oh he AION RIS TS Af 4 . Trunk with obvious, pigmented, MISC EPA ILAC ae eee tat: elon 5 Trunk may have papillae (skin bodies) but these are unpigmented and barely raised above the surface 7 . Tentacles present, papillae uni- formly distributed all over trunk, trunk length rarely exceeds width by more than 8 times (stout cyl- inder or flask shaped), shallow to bathyal depths Tentacles reduced to lobes, pa- pillae rare in midtrunk, trunk length commonly exceeds width by more than 8 times (slender cylin- der), deep, cold water .. N. diaphanes . Trunk flask shaped, anus on nar- rowed anterior region which usu- ally also exhibits an indented con- striction, nephridia posterior to the anus N. constrictum Trunk sausage shaped, nephridia not posterior to anus .. N. pellucidum . Elongate, very slender transparent trunk, sometimes threadlike (trunk width usually less than 4, the length), gut with separated coils and no spindle muscle Cylindrical translucent or opaque trunk (trunk width rarely less than %o the length), gut coils close to- gether, spindle muscle usually PRESS Ure we ata ss ieee es pth 13 . About 30 dark longitudinal epi- dermal ridges at anterior end of trunk sometimes giving the impression of a hardened pseu- doshield Anterior end of trunk without dark epidermal ridges ite 12. NS 14. We 553 Retractor muscles originate in an- terior '4 of trunk, introvert less than ¥; trunk length, distal hooks over 150 um N. vitjazi Retractor muscles originate in pos- terior third of trunk, introvert '2— 7 times the trunk length, distal hooks less than 25 wm APES tn oii evans N. wodjanizkii Anterior end of trunk in shape of short cone with epidermal ridge around base of cone (Fig. 3C) ... SETA TG Reels. N. tasmaniense Anterior end of trunk not cone shaped but often swollen around Me PMiMGdOPOLES eee eee Il Introvert longer than trunk (Fig. 3A), retractor muscles originate from 35-50% of distance to pos- terior end, hooks less than 30 wm eT: AER RE, N. capilleforme Introvert shorter than trunk, re- tractor muscles originate from 15— 30% of distance to posterior end, distal hooks more than 100 um 12 Introvert less than 25% trunk length, distal hooks exceed 200 um SAE OMEN cu Ree Re N. constricticervix Introvert 50-75% trunk length (Fig. 3B), distal hooks 150 um or less... i OAL AEA RINE DRA ea ie MR ER N. cutleri Hooks in distinct rows around in- trovert N. rimicola Hooks, if present, notin rings .. 14 Hooks with unusual series of ra- diating filaments from base (Fig. NCA BS. PRESEN. N. multiaraneusa Hooks, if present, without basal fil- INET ect NMA Re fore A WS) . Retractor muscles originate in pos- terior % of trunk ....... N. filiforme Retractor muscles originate in mld @ilee/2ro fatnunikennne eee eee 16 . Tentacular crown reduced to short lobes, dorsal pair largest ........ V7/ Normal array of digitiform tenta- CIESIPRESEN Gs Say ke oO ae ee 19 Hermaphroditic species from shal- low, Northeastern Atlantic Ocean, trunk length usually 4-5 x diam- eter N. minutum — Dijoecious species, trunk length 6- 10x diameter 18. Larger, opaque, from bathyal depths) 2s 03. 80Gals Ja N. lilljeborgi — Smaller, transparent/translucent, bathyal and abyssal depths i ret: HOS a N. diaphanes 19. Medium sized (more than 50 um) dark hooks present — Hooks apparently absent . Hooks in spiral arrangement, unique shape (Fig. 1F) N. abyssorum 1 HOOKSiscattered: Ae ae 21 21. Hooks robust and blunt (Fig. 1D) aes Siete are Re See N. confusum — Hooks tall, spinelike with soft cor- tical layer (Fig. 1A) N. laetmophilum 22. Less than 50 digitiform tentacles, introvert 1—2 times trunk length. . BS See PRO esas Be N. eremita — More than 50 threadlike tentacles, introvert less than '2 trunk length RAEN naees Oak N. novaezealandiae Taxonomic Section This section has those species we consider to be valid arranged alphabetically. Follow- ing this is a section including the species placed in incertae sedis, species inquiren- dum or moved to other genera (see Table i): Nephasoma abyssorum (Koren and Danielssen, 1875) Phascolosoma abyssorum Koren and Dan- ielssen, 1875:129-130.—1877:131- 132.—Selenka et al., 1883:30-31.— Fischer, 1895:14; 1925:16; 1929:471- 472.—Théel, 1905:78.—Gadd, 1911:88- 89.—Southern, 1913:27—28.—J. Fischer, 1914:99-100.— Wesenberg-Lund, 1933: 9-11. Phascolosoma abyssorum var. punctatum Herubel, 1925:261-262. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Golfingia abyssorum. —Wesenberg-Lund, 1955:201.—Murina, 1964a:220-221; 1964b:56-57; 1968:195.—Stephen and Edmonds, 1972:134-135.—Cutler and Cutler, 1980b:452.—Gibbs, 1982:121.— Cutler et al., 1984:268—269.—Saiz-Sali- nas, 1984:182-183. Not Physcosoma (=Phascolosoma) abysso- rum Southern 1913:12. Phascolosoma incompositum Sluiter, 1912: 16-17, pl. 1, figs. 9-10. Golfingia incomposita.—Stephen and Ed- monds, 1972:145-146. Nephasoma incompositum.—Gibbs, 1986: 339-340. Material examined. —N. abyssorum: ZMUB, type; other recently collected North Atlantic specimens. G. iIncomposita: ZMUA, type. G. benhami: BMNH, type. In general shape and appearance (smooth, white, sometimes iridescent trunk) this species resembles WN. /illjeborgi and N. mi- nutum. It is distinguished by having devel- oped tentacles and unique, spirally ar- ranged, medium-sized, dark hooks (50—150 uum). While this species is alleged to lack a spindle muscle we did observe a weakly de- veloped muscle within the gut coil in two out of eleven specimens closely examined. Herubel (1925) described a variety (P. a. punctatum) which Stephen and Edmonds (1972) elevated to subspecific rank. Since the reference material cannot be located and the putative difference (small elliptical pa- pillae scattered thinly over the trunk) is so slight, we have reduced this taxon to the status of a junior synonym of the nominate form. The single specimen of N. incompositum has the iridescent skin, dark recurved hooks and bathymetric range of N. abyssorum. Therefore, we concur with Gibbs (1986) in reducing it to the status ofa junior synonym. Distribution. —Common in northeast At- lantic and Arctic oceans, rare in northwest Atlantic and Pacific oceans, and Mediter- ranean Sea at bathyal and abyssal depths. VOLUME 99, NUMBER 4 Table 1.—Species considered and proposed taxonomic changes. Present name Nephasoma abyssorum (Koren and Danielssen, 1875) Nephasoma benhami (Stephen, 1948) Nephasoma bulbosum (Southern, 1913) Nephasoma capilleforme (Murina, 1973) Nephasoma chuni (Fischer, 1916) Nephasoma cinctum (Gerould, 1913) Nephasoma cinereum (Gerould, 1913) Nephasoma confusum (Sluiter, 1902) Nephasoma constrictum (Southern, 1913) Nephasoma constricticervix (Cutler, 1969) Nephasoma cutleri (Murina, 1975) Nephasoma delagei (Herubel, 1903) Nephasoma depressum (Sluiter, 1902) Nephasoma diaphanes (Gerould, 1913) Nephasoma elachea (Fisher, 1952) Nephasoma elisae (Murina, 1977) Nephasoma eremita (Sars, 1851) Nephasoma filiforme (Sluiter, 1902) Nephasoma fimbriatum (Sluiter, 1902) ‘Nephasoma flagriferum (Selenka, 1885) Nephasoma glacialis (Koren and Danielssen, 1880) Nephasoma improvisa (Théel, 1905) Nephasoma incompositum (Sluiter, 1912) Golfingia intermedia (Southern, 1913) Nephasoma laetmophilum (Fisher, 1952) Nephasoma lilljeborgi (Danielssen and Koren, 1880) Golfingia macra (Sluiter, 1891) Nephasoma marinki Pergament, 1940 Nephasoma minutum (Keferstein, 1863) Nephasoma mucidum (Sluiter, 1902) Nephasoma multiaraneusa (Murina, 1967) Nephasoma nicolasi (Thompson, 1980) Nephasoma novaezealandiae (Benham, 1904) Golfingia pavlenkoi (Ostroumov, 1909) Nephasoma pellucidum (Keferstein, 1865) Nephasoma prioki (Sluiter, 1881) Nephasoma rimicola (Gibbs, 1973) Nephasoma rutilofuscum (Fischer, 1916) Nephasoma schuttei (Augener, 1903) sensu Cutler and Murina Nephasoma sluiteri (tenBroeke, 1925) Nephasoma subhamatum (Sluiter, 1902) Nephasoma tasmaniense (Murina, 1964) Nephasoma verrilli (Gerould, 1908) Nephasoma vitjazi (Murina, 1964) Nephasoma vitreum (Roule, 1898) Nephasoma wodjanizkii (Murina, 1973) 555 Proposed name No change N. abyssorum benhami No change No change incertae sedis N. diaphanes diaphanes N. pellucidum pellucidum No change No change No change No change incertae sedis species inquirendum N. diaphanes diaphanes N. eremita N. wodjanizkii elisae No change No change species inquirendum No change N. lilljeborgi N. minutum N. abyssorum Phascolion strombus No change No change Aspidosiphon macer N. lilljeborgi No change N. filiforme No change N. wodjanizkii wodjanizkii No change Thysanocardia nigra N. pellucidum pellucidum Thysanocardia nigra No change No change No change N. diaphanes corrugatum N. pellucidum pellucidum N. pellucidum subhamatum No change N. pellucidum pellucidum No change incertae sedis N. wodjanizkii wodjanizkii 556 Nephasoma abyssorum benhami (Stephen, 1948), new status Phascolosoma benhami Stephen, 1948:218- 219, pl. 1, figs. 2-4. Golfingia benhami. —Stephen and Ed- monds, 1972:135-136. The two bathyal Antarctic specimens of G. benhami are said to be similar to N. /ill- Jeborgi differing only in papillae and hook size. However, our examination showed a much greater similarity to N. abyssorum with its larger, dark hooks and ‘normal’ ten- tacles. Stephen measured the body regions in the holotype from a point about 3 mm anterior to the nephridiopores. If one uses these pores to mark the anterior end of the trunk then the retractors are in the middle third, not posterior. The two specimens are 13 mm and 22 mm long. The presence of a spindle muscle within the gut coil (not ex- tending out onto the rectum) and its loca- tion (no records of N. abyssorum in the southern latitudes) rule against combining these two taxa. Therefore, we have retained the name but reduced it to the rank of a geographically separated subspecies whose major morphological distinction is a strong- ly developed spindle muscle. Distribution. —Off Kemp Land, Antarc- tica 600 m. Nephasoma bulbosum (Southern, 1913) Phascolosoma bulbosum Southern, 1913: 23-24, pl. 5, fig. 6. Golfingia bulbosa. —Stephen and Edmonds, 1972:136.—Cutler, 1973:152-153. Nephasoma bulbosum. —Gibbs, 1986:337. Material examined.—INMD, Type ma- terial. This taxon is similar to N. flagriferum in having a tail and papillae on the posterior end of the trunk but in both cases they are very much smaller in this species. The tail is not really a caudal appendage but more a narrowing of the trunk into a point. The nerve cord extends into the distal tip of this PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tail unlike the condition in N. flagriferum. Aside from the shape of the body this species is like N. eremita in many ways, including the apparent absence of hooks, and may be closely related. Distribution. —Off Ireland and northeast United States at bathyal depths. Nephasoma capilleforme (Murina, 1973) Golfingia capilleformis Murina, 1973a:943-— 944.—Frank, 1983:14—-15. Golfingia glacialis. —Murina, 1964b:57-59; 1974a:234.—Cutler and Cutler, 1980b: 453-454. Material examined. —ZIAS, Type mate- rial and other specimens identified by Mu- rina; recently collected Atlantic Ocean worms. This is one of the group of thin deep-water species with loosely wound gut coils. When present and extended the long introvert (1— 2 times the trunk) and small hooks (20-25 um) are diagnostic. In general this has the most elongate thread-like body of the genus and it is often irregularly contracted to give a beaded appearance. The anterior end of the trunk often is swollen and golden col- ored in the vicinity of the anus and ne- phridiopores. Distribution. — Pacific and Atlantic oceans from 920-4900 m. Nephasoma confusum (Sluiter, 1902) Phascolosoma confusum Sluiter, 1902:38- SOL sal 3h, lie, 5 Golfingia confusa. —Murina, 1957:993-994; 1972:298-299; 1974a:234; 1978:123.— Stephen and Edmonds, 1972:138-139.— Cutler and Cutler, 1980c:199-200.—Di- tadi and Migotto, 1981:125-134.—Cut- ler et al., 1984:269. Golfingia confusa zarenkovi Murina, 1974b: 1716-1717, fig. 3. Material examined. —ZMUA, Type specimen; 2 specimens from Japan; ZIAS, Murina’s subspecies holotype. VOLUME 99, NUMBER 4 This species has been redescribed in de- tail by Ditadi and Migotto (1981) on the basis of 106 specimens ranging in length from 2.5—25 mm. The scattered hooks are medium sized (up to 90 wm) and have a distinctive rim of reinforcing chitin. The skin lacks papillae but does have round skin bod- ies. In their paper Ditadi and Migotto (1981: 131-132) compare this species to N. cine- reum but it is our belief, based on an ex- amination of the type, that their interpre- tation of the papillae/skin bodies and hooks was in error. Nephasoma cinereum does not have this type of hook and does have dis- tinct papillae. Therefore, following the lead of Cutler and Murina (1977) we consider it to be a junior synonym of N. pellucidum. Murina’s subspecies was described from two specimens of 4 mm and 5 mm. The supposed differences from the nominate form are due to its small size and these in- dividuals do not warrant subspecific rank. Distribution. — Around the Antarctic, off Brazil, southern Australia, Indonesia, and the northwest Pacific Ocean from 44600 m. Nephasoma constrictum (Southern, 1913) Phascolosoma constrictum Southern, 1913: 25-27, pl. 6, fig. 7. Golfingia constricta. —Stephen and Ed- monds, 1972:139. Nephasoma constrictum. —Gibbs, 1986: 337. Material examined. —INMD, Type spec- imens; several recently collected specimens from near the type locality. This species shares many attributes with N. pellucidum but differs most notably by the flask-shaped trunk, short stubby tenta- cles and the constriction at the introvert/ trunk junction. The Appendix contains morphological information on 17 worms ranging in trunk length from 7-30 mm. The known distribution is distinct and limited. Distribution. — Off Ireland at 1100-1300 m plus unpublished records in northeastern Atlantic (35-51°N) at 1900-4000 m. 557 Nephasoma constricticervix (Cutler, 1969) Golfingia constricticervix Cutler, 1969:215- ANT? MIPS VNSI.—Cuilere eyarel (Counalere, 1980b:452; 1980c:200. Material examined. —Type material plus several recently collected specimens. This is one of two species in this genus which has hooks exceeding 200 um. It is one of several elongate, slender deep-water members of this genus and if the introvert is withdrawn or broken off it can be difficult to identify with certainty (see discussion of N. cutleri). The range in hook size is from 40-250 um and only the more distal hooks exceed 200 um. Therefore, if only the small- er hooks are seen it is possible to mistake this for another species e.g., NV. cutleri. Distribution.—North Atlantic at 1500- 5500 m. Unpublished records extend this to 22°S in the eastern Atlantic. Nephasoma cutleri (Murina, 1975) Golfingia cutleri Murina, 1975:1087—1088; 1978:123. Material examined. —ZIAS, Type mate- rial. On paper this species has many similar- ities to N. constricticervix. However, the hooks are considerably smaller (less than 150 um) and paler, the introvert is longer (50-75% of the trunk length, not 20-25%) and the worm is much less robust. This species has only been recorded from the Pa- cific Ocean while the other appears to be an Atlantic Ocean species. Distribution. —Pacific Ocean from 40°S— 11°N at abyssal depths. Nephasoma diaphanes diaphanes (Gerould, 1913) Phascolosoma diaphanes Gerould, 1913: 395. Golfingia diaphanes.—Cutler and Cutler, 1980b:452-453; 1980c:201-—202.— Frank, 1983:15-16.—Cutler et al., 1984:269- 270. 558 Phascolosoma improvisum. —Gerould, 1913:395-396.— Wesenberg-Lund, 1930: 32-34; 1939:22-23. Golfingia improvisa. —Wesenberg-Lund, 1955:11; 1963:110—111.—Murina, 1958: 1625-1628; 1968b:197; 1971b:42; 1972: 300; 1973b:69; 1978:123.—Stephen and Edmonds, 1972:145.—Amor, 1975:113- 115.—Edmonds, 1976:222-—224. Phascolosoma minutum. —Théel, 1911: 31.—Sluiter, 1912:10-—11.—Fischer, 1922a:34; 1922b:237; 1929:464-467 (partim).— Wesenberg-Lund, 1930:30; 1932:9-10; 1937a:9-10; 1937b:12-13. Golfingia minuta. —Wesenberg-Lund, 1955: 11; 1963:110-111.—Murina, 1957:994— 995: 1958:1628—-1634; 1968b:197; 1971b: 42: 1972:300; 1973b:69; 1974a:235; 1976:65; 1978:124.—Stephen and Ed- monds, 1972:149-150.—Cutler, 1973: 155-159; 1977a:143.— Cutler and Cutler, 1979a:957-958.—Saiz-Salinas, 1984:183. Golfingia sectile Murina, 1974a:228-—230. Phascolosoma anceps. —Sluiter, 1912:10.— Wesenberg-Lund, 1925:90. Phascolosoma cinctum Gerould, 1913:398—- 400, pl. 59, fig. 6, text—fig. 8. Golfingia cincta.—Stephen and Edmonds, 1972:137-138. Phascolosoma sabellariae.—Gerould, 1913: 392-395. Material examined. —USNM #8584, Gerould’s other material, G. cincta type; hundreds of specimens from the East Coast of North America; ZIAS, type specimen of N. sectile. As noted in the introduction to this sec- tion this taxon is very similar to N. minu- tum and N. lilljeborgi but is generally small (less than 10 mm), with a transparent/trans- lucent body wall, cosmopolitan in deep water, and dioecious. Gerould (1913) first suggested using the name diaphanes for the western North Atlantic animals that matched the description of P. minutum but were not hermaphroditic (he used the name Phascolosoma sabellariae for that collec- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tion). However, it was not until Cutler and Cutler (1980c:201) that this idea was dis- cussed, adopted, and the name G. diaphanes was formally used. We recently examined Gerould’s specimens of P. sabellariae which now serve as the type series for this species and selected one sample for a lectotype. The single N. sectile from 6200 m in the South Atlantic is not as slender as most members of this species (1.6 X 6.7 mm) but is within the range of possible variation and does not exhibit any unique set of features. Thus, we consider it to be a junior synonym of this species. In Cutler (1973) G. cincta was placed in synonymy with G. minuta but since the recent redefinition of the latter, G. cincta must now be considered a junior syn- onym of this species. Distribution. —Cosmopolitan, cold water, most from bathyal and abyssal depths. Nephasoma diaphanes corrugatum, new subspecies Golfingia schuttei.—Murina, 1964a:238- 242; 1967c¢:54; 1971a:81; 1971b:43; 1973b:69-70; 1974a:235; 1978:124.— Cutler and Cutler, 1980b:453; 1980c: 204.—Frank, 1983:17-18. Description. — These pear- to cylindrical- shaped animals have trunks up to 10 mm long (occasionally up to 30 mm). The skin is tan to greyish-brown, translucent to opaque, with irregular, longitudinal epider- mal ridges (wrinkles or folds) commonly on the base of the introvert and the anterior part of the trunk (occasionally extending to the posterior end). Often there are papillae on the posterior end which may be darker than the surrounding skin. The introvert is about equal in length to the trunk but may vary from 50—150% of the trunk length, and bears small (usually 20-30 um tall), scat- tered, pale, triangular hooks. The tentacular crown is reduced to a few (6-8) short lobes plus two longer dorsal tentacles similar to that in N. minutum. Internally the ventral pair of introvert re- VOLUME 99, NUMBER 4 tractor muscles originate near the posterior end (75-85%) in small worms (less than 4 mm) but in the middle (50-70%) in larger ones. The pair of short free nephridia are at the level of, or just posterior to, the anus. The gut forms a double helix with a weakly developed spindle muscle within the coil, not extending onto the rectum. No fixing muscles or caeca were seen. Differential diagnosis. —The main differ- ence from the nominate form is the nature of the epidermis, this one having rough, opaque skin with wavy ridges, N. diaphanes diaphanes having smoother, translucent/ transparent skin, although some do have raised, pigmented papillae, especially on the posterior third. The subspecific name re- flects its ridged epidermal appearance. Type material.—Deposited at the Na- tional Museum of Natural History (USNM) Washington, D.C., holotype USNM 98775, 10 paratypes USNM 98776. Remarks.—This subspecies is not well defined and future systematists using other characters may reach different conclusions. The use of the name N. schuttei in the sev- eral works by Cutler and Murina listed in the synonymy is the result of looking at the original description and interpreting it too broadly. The deep waters of the world’s oceans are densely populated with small members of this genus which have been in- tensively collected over the past two de- cades. A dependable and universally ap- plied classification of this fauna is desirable, but not yet in our hands. Distribution. — Atlantic and Pacific oceans and the Mediterranean and Red seas from 80-5900 m. Nephasoma eremita (Sars, 1851) Sipunculus eremita Sars, 1851:197. Sipunculus (Phascolosomum) borealis Qua- trefages, 865:620. Phascolosoma boreale Keferstein, 1865: 437-—438.—Baird, 1868:84.—Diesing, 1851:555; 1859:760.—Verrill, 1874: 387.—Koren and Danielssen, 1877:134. 559 Phascolosoma digitatum Théel, 1875:11; 1905:72. Phascolosoma eremita. —Selenka et al., 1883:35-36.—Fischer, 1895:17; 1914:10; 1922b:237; 1925:17; 1929:463-464.— Théel, 1905:72-74.—Gadd, 1911:88.— Sluiter, 1912:8.—Gerould, 1913:385- 387.—J. Fischer, 1914:98-99.—Cham- berlain, 1920:4.— Wesenberg-Lund, 1930: 28; 1932:67; 1937b:11-12.—Leroy, 1936: 425. Golfingia eremita. —Stephen and Edmonds, 1972:141-142.—Cutler, 1973:150- 152.—Cutler and Cutler, 1980a:1-2; 1980c:202—204.— Frank, 1983:16-17. Phascolosoma eremita australis Benham, 1922:17-18.—Fischer, 1929:483. Golfingia eremita var. australe. —Wesen- berg-Lund, 1963:111.—Stephen and Ed- monds, 1972:14.—Murina, 1972:299- 300; 1974a:234.— Not Wesenberg-Lund, 1959:182-183. Golfingia eremita californica Fisher, 1952: 396-397.—Stephen and Edmonds, 1972: 142. Phascolosoma eremita scabra Gerould, 1913:387-388. Golfingia eremita scabra. —Stephen and Edmonds, 1972:142. Golfingia elachea Fisher, 1952:399—400, pl. 25, figs. 1-3. Material examined. —USNM, types of G. eremita californica, G. eremita scabra, G. elachea; MNHU, P. boreale type; recently collected North Atlantic specimens. This species appears to be one of the few members of this genus to lack hooks. How- ever, as pointed out earlier, if only the young worms do have hooks (as has been shown for other members of this phylum) they probably would be identified as some other species. A study of the early life stages of this species would be most valuable. Mature worms have a stocky nonpapillated trunk, often with transverse grooves in the thick body wall, and the tentacles are well devel- oped. 560 Fisher’s single example of N. elachea from Baja California presumably differs from this species because it has seven fixing muscles. The present condition of the type precludes any confirmation of this. Fisher did not pro- vide a differential diagnosis but did put this item in italics thereby stressing its impor- tance to him. The number of fixing muscles is quite variable throughout this phylum and especially within this genus. We see no real differences here and therefore consider G. elachea to be a junior synonym. The three subspecies listed in Stephen and Edmonds (1972) are not considered by us to merit such rank and are hereby reduced to junior synonyms. We reaffirm Cutler’s (1973) observations about Gerould’s dark variety scabra being an environmentally (not genetically) determined attribute. Benham’s (1922) variety australis was based on a sin- gle worm and the primary distinction was the distribution and relative size of the pa- pillae, not a solid basis. Fischer’s (1929) re- cord was simply a repeat of Benham’s, not new material. Fisher’s (1952) variety cali- fornica has a ‘strong’ spindle muscle and no functional contractile vessel. Our compar- ison of this material showed these structures to be near one end of their particular con- tinuum but not outside the range found in Atlantic populations. Distribution. —Common in the Arctic and North Atlantic, present in South Atlantic and Antarctic, rare in the eastern Pacific Oceans from 20-2000 m. Nephasoma filiforme (Sluiter, 1902) Phascolosoma filiforme Sluiter, 1902:37-38, pl. 4, figs. 1-4. Golfingia filiformis. —Stephen and Ed- monds, 1972:143. Phascolosoma mucidum Sluiter, 1902:40. Golfingia mucida. —Stephen and Edmonds, L9F2=2TS 0: Material examined. —ZMUA, Type ma- terial of N. filiforme and N. mucidum. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The three N. filiforme specimens vary more than the description suggests, espe- cially in the form of their papillae on the two ends of the trunk. Only one of these has long hairlike papillae on the front end and the mushroomlike form of the posterior pa- pillae is an oversimplification. One addi- tional problem concerns the location of the anus and nephridiopores. Thompson (1980) said that these openings were on the intro- vert and Sluiter reported the introvert length to be half that of the trunk. This confusion results from the fact that the circular mus- cles in the anterior part of the trunk and the proximal part of the introvert have con- tracted in an atypical fashion. This situation has been the cause for confusion in N. ma- rinki also (see Cutler and Murina 1977). Our operational definition of the trunk/introvert junction being that region just anterior to the nephridiopores resolves this problem so that the trunk is 40 mm long (not 60) with an introvert 1.2 times longer in this worm. In the other two type specimens (trunk lengths 15 mm and 45 mm) the introvert is about 0.8 and 0.3 times the trunk length, hardly a constant species-specific character. This is not a solidly based species and needs some clarification and a larger data base but we propose no changes at this time. There are also some problems with N. mucidum. Our measurements of the two re- maining specimens showed their length (6 mm and 7 mm) exceeds their width by only three times, not 10. It is clear that these specimens are damaged and incomplete so these measurements are of limited value. The long papillae Sluiter mentioned are very few in number. The grey felt-like covering does not seem to be a part of the animal but rather some adhering substrate and very easily rubbed off. These animals came from the same location and we see no significant differences. Since N. mucidum appears later in the publication it becomes the junior syn- onym. Distribution. — Off Indonesia (6°S, 134°E) at 1788 m. VOLUME 99, NUMBER 4 Nephasoma flagriferum (Selenka, 1885) Phascolosoma flagriferum Selenka, 1885: 13-16, pl. 3, fig. 17.—Sluiter, 1900:12.— Fischer, 1914:10.—Gerould, 1913:391- 392. Golfingia flagrifera.—Murina, 1968b:196; 1978:123.—Stephen and Edmonds, 1972: 144.—Cutler, 1973:153-155; 1977a:142— 143.—Cutler and Cutler, 1980b:453. Material examined. —-BMNH, Type specimen; many recently collected speci- mens. The large vesicular, bulbous papillae on the posterior end of the trunk and the dis- tinct caudal appendage make this species easy to identify and there is no confusion about its validity. The Appendix contains morphological information on 27 worms ranging in trunk length from 3.5-120 mm. Distribution.—An abyssal species (few bathyal records) common in the North At- lantic, present in the South Atlantic and Pa- cific. Nephasoma laetmophilum (Fisher, 1952) Golfingia laetmophila Fisher, 1952:397- 399, pl. 25, figs. 4-6.—Stephen and Ed- monds, 1972:148. Material examined. —USNM, Type spec- imen. This single bathyal California specimen has many characters in common with the Atlantic Ocean N. abyssorum. The larger number of tentacles (40) and the strong wing muscle may be important or may be arti- facts of its larger size. The one striking dif- ference is the nature of the hooks; they have a spine-like form with a cortical layer which can be easily rubbed off. Whether this single specimen is an anomaly or part of a bio- logical population is an open question. It is puzzling why no additional specimens from this heavily sampled region have been re- covered in the intervening decades. Despite our reservations we propose no change in the status of this species at this time. 561 Distribution. —Off southern California, 1900 m. Nephasoma lilljeborgi (Danielssen and Koren, 1880) Phascolosoma lilljeborgii Danielssen and Koren, 1880:463-464; 1881:63-64.—Se- lenka et al., 1883:40—41.— Fischer, 1895: 14; 1929:471-472.—Sluiter, 1912:9.— Théel, 1905:79-80. Golfingia lilljeborgi. —Wesenberg-Lund, 1954:9-10.—Stephen and Edmonds, 1972:148.—Gibbs, 1982:121-122.— Frank, 1983:17. Onchnesoma glaciale Danielssen and Ko- ren, 1880:464; 1881:64. Phascolosoma glaciale. —Roule, 1896: 474.—Théel, 1905:80—81.—Fischer, 1929:472.— Wesenberg-Lund, 1930:30; 1932:8—9. Golfingia glacialis.—Stephen and Ed- monds, 1972:144.—Gibbs, 1982:119- 120.—Not Murina, 1964b: 57-59; 1974a: 234.—Cutler and Cutler, 1980b:453-—454. Nephasoma marinki Pergament, 1940:55-— 61.—Stephen and Edmonds, 1972:214— Daley Material examined. —ZMUB, Type spec- imens of N. /illjeborgi and N. glaciale; other recently collected specimens from near the type-locality. This taxon has recently been revised by Gibbs (1982). On paper there is very little to differentiate this taxon from JN. diaph- anes. The type material is larger (10-30 mm), more opaque, does not live in foraminiferan tests as many N. diaphanes do, and was col- lected from shallower water. The posterior end of the trunk in many worms comes to a blunt point or has a posterior nipple on a rounded base. This name has only been used a few times and only for specimens collected in the northeast Atlantic Ocean. Nephaso- ma marinki was synonymized in Cutler and Murina (1977). Since the recent reevalua- tion of this species it has been determined 562 that Cutler and Murina had used this name for some N. capilleforme. Distribution. —Far northeast Atlantic Ocean from bathyal depths. Nephasoma minutum (Keferstein, 1863) ?Sipunculus johnstoni Forbes, 1841:254. Phascolosoma johnstoni. —Southern, 1913: 28.—Lindroth, 1941:449-450. Phascolosoma minutum Keferstein, 1863: 40, pl. 3, figs. 7-10; 1865:438.—Cuenot, 1922:9-10.—Fischer, 1925:19-—20; 1929: 464-—467(partim).— Wesenberg-Lund, 1939:20-22.—Stephen, 1934:167-168. Petalosoma minutum. —Selenka et al., 1883: 129.—Southern, 1908:83-86.— Paul, 1909:1-50. Golfingia minuta. —Akesson, 1958:33- 46.—Stephen and Edmonds, 1972:149- 150.—Gibbs, 1973:73-86; 1975:69-82; 1977a:16-17. Phascolosoma anceps Théel, 1905:84—-86.— Wesenberg-Lund, 1925:90. Phascolosoma improvisum Théel, 1905:82-— 83, pl. 5, figs. 51-58, pl. 12, figs. 177-178, pl. 14, figs. 202—203.— Wesenberg-Lund, 1939:22-23. Golfingia improvisa. —Stephen and Ed- monds, 1972:145. Phascolosoma sabellariae Théel, 1905:81. Material examined. —Several recently collected specimens identified by P. Gibbs collected near the type locality. This name has been used in a variety of ways over the past century. Gibbs (1977a: 79-80) has been the most recent to examine this situation and he redefined the taxon in a narrow way including only those from shallow water in the northeastern Atlantic Ocean which are hermaphroditic (see also Paul 1910 and Akesson 1958). It is often very difficult to differentiate between sev- eral populations as discussed in the intro- duction and it is likely that very often this name has been used as a taxonomic waste- basket. The present definition still includes N. improvisa, 1.e., those with larger darker PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON papillae. Some of the uses of this latter name may actually refer to N. diaphanes but as noted above, without reexamining the ac- tual specimens one cannot be certain. We here divide the records based on location assuming that this is correct most of the time. Distribution. —Northeast Atlantic Ocean from shallow water. Nephasoma multiaraneusa (Murina, 1967) Golfingia multiaraneusa Murina, 1967b: 1332-1333, fig. 2.—Stephen and Ed- monds, 1972:151. Material examined. —ZIAS, Type mate- rial. The hooks are the unique attribute of this species. They are 15-30 um tall with a series of radiating filaments from the base giving each hook a spider-like appearance. The single 2 mm specimen is not a solid basis for a species but we propose no change at this time. If future collections in this region do not uncover additional specimens the status of this worm should be reconsidered. Distribution. —Cuba at 4 m. Nephasoma novaezealandiae (Benham, 1904) Phascolosoma novae-zealandiae Benham, 1904:301-303, pl. 15, figs. 1-2, pl. 16, fig. 8; 1909:82. Golfingia novae-zealandiae. —Edmonds, 1960:162-163. Golfingia novaezealandiae.—Stephen and Edmonds, 1972:151. Material examined. —None. The type of this species was taken from the stomach of a dogfish and five years later a second worm was found in ooze. Edmonds added two more specimens, all four coming from near New Zealand. The trunk length ranges from 25-235 mm with introverts much shorter than the trunks. As noted by Edmonds this species is morphologically VOLUME 99, NUMBER 4 similar to NV. eremita but because of its geo- graphic separation, the large number of thin, thread-like tentacles, and large size, it has been considered a separate taxon. Subspe- cific rank may be more appropriate but we propose no change at this time. Distribution. —Off New Zealand and Chatham Is. from 65-70 m. Nephasoma pellucidum pellucidum (Keferstein, 1865) Phascolosoma pellucidum Keferstein, 1865: 433, pl. 32, figs. 26-27.—Baird, 1868: 86.—Selenka et al., 1883:32—34.—Shi- pley, 1899:155.—Sluiter, 1902:34.—Au- gener, 1903:299-300.— Lanchester, 1905: 28.—Southern, 1913:6.—Fischer, 1914:8; 1919:281; 1922a:17; 1923:23.—ten- Broeke, 1925:83.—Leroy, 1936:425. Golfingia pellucida.—Murina, 1968a:421- 422; 1972:302-303.—Stephen and Ed- monds, 1972:152-153.—Cutler, 1973: 159-162; 1977a:143; 1977b:152.—Cut- ler and Cutler, 1979b:105; 1980a:2.— Cutler and Murina, 1977:177.—Thomp- son, 1980:258.—Cutler et al., 1984:270. Sipunculus (Phascolosomum) pellucidus Quatrefages, 1865:620. Phascolosoma riisei Keferstein, 1865:437.— Baird, 1868:96. Phascolosoma cinereum Gerould, 1913: 396-398, figs. 6-7. Golfingia cinerea. —Stephen and Edmonds, 1972:138. Phascolosoma sluiteri tenBroeke, 1925:84- 86. Golfingia sluiteri.—Stephen and Edmonds, 1972:156-157.—Cutler and Murina, IDVVENV I, Meee. Phascolosoma verrillii Gerould, 1908:488-— 489; 1913:388-391. Golfingia verrillii.—Murina, 1964a:243- 246.—Stephen and Edmonds, 1972:158. Golfingia coriacea. —Fisher, 1950:551; 1952:396.—Not Murina, 1972:298. Not Phascolosoma coriaceum Keferstein, 1865:432. 563 Golfingia eremita var. australe.—Wesen- berg-Lund, 1959:181-182. Material examined. -MNHU, Type specimens; numerous recently collected specimens; USNM, G. cinerea type, Fish- er’s specimens of G. coriacea; ZMUA, G. sluiteri; ZLAS, Murina’s G. coriacea; UZMK, Wesenberg-Lund’s G. eremita var. australe. This species with large uniformly distrib- uted papillae is well founded and widely distributed. In Cutler and Murina (1977) G. sluiteri was reduced to a junior synonym since it turned out to be merely N. pelluci- dum with its esophagus everted through the mouth. Golfingia verrillii was reduced in status in Cutler (1973). Golfingia cinerea was alleged to be dis- tinct because of its stouter trunk (length 3 vs. 5—9 times the diameter), a greater variety of papillae shapes, and the relationship be- tween the nephridiopores and anus. Our analysis of some large collections showed there to be sufficient variation within one population to include this species. When Gerould erected this species he did not com- pare it to N. pellucidum nor did he record any from the U.S. east coast. Ditadi and Migotto (1981) suggested that G. cinerea should be considered a junior synonym of N. confusum. As discussed above we disa- gree and place it here. In Cutler (1973) G. coriacea was synon- ymized with this species based on an ex- amination of Fisher’s material. While we still believe Fisher’s worms to be this species, Keferstein’s species name has been trans- ferred to Themiste with some uncertainty due to the loss of the type material (Gibbs et al., 1983). Murina’s specimens of this species are without doubt a Themiste. Wesenberg-Lund’s G. eremita var. aus- trale bears distinct hooks and in other ways clearly resembles this taxon. Murina’s (1972) record from near Prince Edward Island (Subantarctic waters) is based on three worms all less than 6 mm long. We have been unable to confirm this record and 564 question its validity. Edmonds (1982) ques- tions the validity of earlier N. pellucidum records from Australian waters and suggests that they are probably N. schuttei. In Thompson (1980) G. pellucidum is listed in a brief checklist but no morphological or location data are given. This is the only sug- gestion that it occurs in the eastern Pacific and one hopes for a fuller statement in the near future. Distribution. —Shallow-water species (few bathyal records) from the Western Atlantic and Caribbean down to Brazil, in the South Pacific (Indonesia/Australia), southern Ja- pan, and one record from Cape Town. Nephasoma pellucidum subhamatum (Sluiter, 1902), new status Phascolosoma subhamatum Sluiter, 1902: 35-36, pl. 3, figs. 10-12. Golfingia subhamata.—Stephen and Ed- monds, 1972:157.—Cutler et al., 1984: 270-271. Material examined. —ZUMA, Type specimens. This taxon compares with the nominate form in most ways but is a bathyal western Pacific population (not shallow warm water). The only morphological feature which dif- fers is that the hooks in this form are larger (100 vs. 60 wm) and thinner. The type ma- terial has not been preserved well and the internal organs are macerated. One of the worms from Sluiter’s Sta 126 is a Thysano- cardia. Distribution. —Indonesia and central Ja- pan at 440-2050 m. Nephasoma rimicola (Gibbs, 1973) Golfingia rimicola Gibbs, 1973:74-80; 1977a:18.—Saiz-Salinas, 1980:54—56. Material examined. —Several specimens from type locality identified by P. Gibbs. This is the only species in the genus to have its hooks arranged in distinct rings, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON which serves to differentiate it from species such as the very similar N. minutum. It also has its anus posterior to the nephridiopores; N. minutum has the reverse relationship. Distribution. —Southwest England and northern Spain in intertidal waters. Nephasoma rutilofuscum (Fischer, 1916) Aspidosiphon rutilofuscus Fischer, 1916:17. Phascolosoma aspidosiphonoides Fischer, 1922c:11-12, pl. 2, fig. 8. Golfingia rutilofusca Fisher, 1952:395.— Stephen and Edmonds, 1972:153-154.— Cutler, 1977a:143-—144.— Cutler and Cut- ler, 1979a:958—961. Material examined. -MNHU, Type specimen; many recently collected speci- mens. This western Indian Ocean species is the most distinctive member of this genus and may well merit separate generic rank if one chose to weight the unique tentacular crown more heavily; we do not at this time. The rusty red color makes this form easy to iden- tify. The posterior end of the trunk some- times appears shield-like but this is not permanent as it varies with body wall contraction. It may be misleading as it was for Fischer who originally placed it in the genus Aspidosiphon. See Cutler and Cutler (1979a:958—961) for a redescription and il- lustrations. Distribution. —Western Indian Ocean from 1—1562 m. Nephasoma schuttei (Augener, 1903) Phascolosoma schuttei Augener, 1903:335-— 337, figs. 17-18. Golfingia schuttei. —Stephen and Edmonds, 1972:156.—Edmonds, 1980:25-—27.—Not as per Cutler, Cutler and Cutler, or Mu- rina. Material examined. -MNHU, Type specimen. This name has been widely misapplied in recent decades by Murina and Cutler. They VOLUME 99, NUMBER 4 have used the name for a small deep-water species very Common in two major oceans (see N. diaphanes). This error became ob- vious when we examined the type. Ed- monds’ (1980) material and excellent de- scription are correctly based on Augener’s species. The large dark papillae and coarse skin are distinctive. Edmonds gives a de- tailed description and discusses the simi- larities to N. pellucidum. This is a distinct and valid species but the unfortunate con- fusion may continue to be a problem for some time until all workers understand and accept this narrower, original definition. Distribution. —South and West Australia from intertidal waters. Nephasoma tasmaniense (Murina, 1964) Golfingia tasmaniensis Murina, 1964a:242-— 243, fig. 13a—b.—Stephen and Edmonds, IODT2RNS I Material examined.—ZIAS, Two type specimens and one additional specimen identified by Murina. Both type specimens are incomplete and damaged worms lacking introverts. What were described as tentacles are actually the broken introvert retractor muscles. The third worm (unpublished record from 9°S, 71°E at 2218 m) is intact with an extended in- trovert. Small pale hooks and a few reduced tentacles are present on a swollen, bulb-like terminal part of the short introvert (less than half the trunk length). The anterior 4 mm of the trunk has the conical shape with a collar at the base of the cone, as figured by Murina (total trunk length is 18 mm). The trunk/introvert junction is constricted into a narrow neck. This species has an uncertain foundation being based on two incomplete worms and the flawed description. The ab- sence of any differential diagnosis is another problem. In general size, shape, looseness of gut coil, constricted neck, and short introvert with bulbous tip, it resembles N. constric- ticervix. The hooks are much smaller and 565 Fig. 3. Introvert and anterior end of three elongate, slender, deep-water Nephasoma species; A, N. capil- leforme; B, N. cutleri; C, N. tasmaniense. Drawn as if trunk lengths were equal to show comparative lengths of introverts. the collar at the base of the anterior cone may be diagnostic. In several ways this species is similar to other long, slender, deep- water members of this genus. A more ade- quate description and diagnosis is needed. In anticipation of this by Murina, we pro- pose no change in the status of this species. Distribution. —Tasman Sea at 1330 m. Nephasoma vitjazi (Murina, 1964) Golfingia vitjazi Murina, 1964a:246-248, fig. 1l6a—b.—Stephen and Edmonds, 1972:158. Material examined. —ZIAS, Type speci- men. 566 This single specimen is incomplete with a torn posterior end, and measures 15 xX 0.08 mm. The anterior end has 30-35 par- allel, longitudinal ridges forming what has been called a “‘shield.”’ The 4 mm introvert bears large hooks (210-280 um) which is unusual in this genus. While there is only this single specimen known, we propose no change in the status of this species. Distribution. —Northwest Pacific Ocean at 4150 m. Nephasoma wodjanizkii wodjanizkii (Murina, 1973), new status Golfingia wodjanizkii Murina, 1973a:944— 945; 1973b:70.—Frank, 1983:18-19. Golfingia nicolasi Thompson, 1980:951- 956. Material examined. —ZIAS, Type speci- men and one other N. wodjanizkii; several specimens of N. nicolasi from type locality identified by B. Thompson. This slender, bathyal species, with its two subspecies, has nephridia posterior to the anus, retractors originating in the posterior Y and an introvert of varying lengths. In very young animals it is shorter than the trunk, it grows to two to three times the trunk in mature worms but in some popu- lations, when completely extended, it may reach six to seven times the trunk length. Small hooks may be present and the ten- tacles are few and reduced. The two N. wodjanizkii have 4 mm and 6 mm trunks with longitudinal lines along most of the trunk and part of the introvert. These worms were said to have indistinct shields at both ends of the trunk but we would not use this term. The introverts are about three times the trunk length but in- completely extended so nothing can be said about tentacles or hooks. The California population of Nephasoma nicolasi is based on several hundred speci- mens with 7-36 mm trunks. The introverts range from 0.5 (in the smallest) to seven times the trunk length, longer than most PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON species in this genus. No hooks have been seen. The introvert and the anterior end of the trunk have a series of longitudinally ar- ranged fine brown lines (ridges in the epi- dermis). A comparison of these two forms showed the differences to be artifacts of small sam- ple size on the one hand and a failure to make a comparison on the other hand. When Thompson was analyzing his population he probably did not consider Murina’s species since one had hooks and the other did not. There was also an apparent difference in trunk size and introvert length. Murina’s use of the term shield could also have been misleading. It is our conclusion that these two taxa are conspecific and, despite the fact that Thompson’s data base is superior his species must be submerged as a junior syn- onym. Distribution. —Sea of Okhotsk, Peru-Chile Trench, and southern California from 1000-— 2400 m. Nephasoma wodjanizkii elisae (Murina, 1977), new status Golfingia elisae Murina, 1977:133-134. Nephasoma elisae. —Gibbs, 1986:338—339. Material examined. —ZIAS, Type speci- men; UZMK, cotypes; 2 specimens from the eastern Atlantic. There are two problems with the descrip- tion of this taxon: rather than the 11-13 longitudinal grooves forming the anterior shield there are 25—30; secondly, hooks have only been observed once (Murina) and ap- pear to be deciduous, a feature shared with many species in this genus. A comparison of these two taxa, as corrected, shows them to be remarkably similar except that the introvert length in N. w. elisae is less than twice the trunk length. Our decision to retain this name at the subspecific rank was largely based on the fact that this represents the Atlantic Ocean population while the nominate form is a Pacific Ocean popula- tion. VOLUME 99, NUMBER 4 Distribution. —Gulf of Guinea at 1520 m and from 43-58°N from 1600-2300 m in the northeast Atlantic Ocean. Species Names Transferred to Other Genera or Considered species inquirendum or incertae sedis Nephasoma chuni (Fischer, 1916) Phascolosoma chuni Fischer, 1916:15; 1922c:9. Golfingia chuni.—Stephen and Edmonds, 1972:136-137.—Murina, 1973b:68. Material examined. -MNHU, Type specimen. One of the two bottles in Berlin labelled P. chuni contained a poorly preserved nematode. The second bottle contained a worm with completely disintegrated inter- nal organs except for the one pair of retrac- tor muscles. Therefore, it is impossible to verify anything about its internal anatomy. Externally it resembles the N. pellucidum complex but there are too many unanswer- able questions (see Stephen and Edmonds 1972:137) to do anything other than place this name on the list of incertae sedis. Nephasoma delagei (Herubel, 1903) Phascolosoma delagei Herubel, 1903:100; 1907:115-117. Golfingia delagei.—Stephen and Edmonds, 1972:139-140. Material examined. —None. In 1922 Cuenot treated this as a junior synonym of Golfingia elongata. Gibbs (1973) agreed but since the type has been lost he considered it to be indeterminable. We concur and place this name on the list of incertae sedis. Nephasoma depressum (Sluiter, 1902) Phascolosoma depressum Sluiter, 1902:39- 40. Golfingia depressa.—Murina, 1964a:227-— 228.—Stephen and Edmonds, 1972:140. 567 Material examined. —ZMUA, Type specimen; ZIAS, Murina’s (1964) specimen. The type specimen has a 4 mm trunk, is poorly preserved, and the introvert is miss- ing. Murina’s worm is 2.5 mm long and she noted that her comparison was hampered by the ‘poor intactness’ of her worm and Sluiter’s incomplete description. We place this name on the list of species inquirendum pending future clarification because of the poor condition of these two very small spec- imens and the puzzling nature of the de- scriptions. Nephasoma fimbriatum (Sluiter, 1902) Phascolosoma fimbriatum Sluiter, 1902:34— 35. Golfingia fimbriata. —Stephen and Ed- monds, 1972:143.—Murina, 1976:64. Material examined. —ZMUA, Type specimen; ZIAS, two of Murina’s worms. This bathyal species is not solidly found- ed and the descriptions contain some dis- crepancies. The anterior part of the trunk is narrowed but by definition, the position of the nephridiopores coincide with the intro- vert—trunk junction. The nephridiopores open 2 mm posterior to the anus and not on the introvert. The introvert is retracted to an unusual degree so that the anterior 5 mm of the trunk is also retracted and while the introvert is shorter than the trunk (7 and 25 mm) this is not the normal condition. The papillae on the introvert are not un- usually long. The retractor muscles origi- nate very near the posterior end of the trunk (85-95%) which is not common in this ge- nus. Of greatest significance is the presence of contractile vessel villi. These structures are present but not on the free portion of the esophagus, only in the beginnings of the gut coil. We suspect that this is an artifact of the extreme state of contraction and in the extended state this portion of the gut would be uncoiled. This removes the ambiguity of earlier statements (Cutler and Murina 1977). 568 Although it is not extended the tentacular crown appears to be short festoons. These two features point towards Thysanocardia but the short introvert and anus—nephridio- pore relationship do not fit that pattern. While we are inclined to place this in the synonymy of 7. nigra we are hereby placing this name on the list of species inquirendum pending future clarification based on addi- tional specimens. Golfingia intermedia (Southern, 1913) Phascolosoma intermedium Southern, 1913: 3-5, pl. 1, figs. 1-8.—Stephen, 1948:219. Golfingia intermedia.—Stephen and Ed- monds, 1972:147.—Gibbs, 1977b:109- 112. Material examined. —None. When Gibbs (1977b) examined the type material he concluded that these specimens are merely immature Phascolion strombus. We concur with that conclusion. Golfingia macra (Sluiter, 1891) Phascolosoma macer Sluiter, 1891:114—115, pl. 2, figs. 13-14; 1902:34. Golfingia macra.—Stephen and Edmonds, 1972:149.—Cutler and Murina, 1977: 183. This species was discussed in Cutler and Murina (1977) after examining the type ma- terial. It clearly belongs in the genus Aspi- dosiphon and is an emended spelling of the original P. macer. Golfingia pavlenkoi (Ostroumovy, 1909) Phascolosoma pavlenkoi Ostroumovy, 1909: 323: Golfingia pavlenkoi. —Stephen and Ed- monds, 1972:152.—Cutler and Murina, 1977:175.—Gibbs et al., 1983:301. Material examined. —None. Cutler and Murina (1977) discussed this species and concluded that it was a junior PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON synonym of Golfingia (Thysanocardia) catharinae. However, when Gibbs et al. (1983) reviewed Thysanocardia and elevat- ed it to generic rank, this species name went into the synonymy of Thysanocardia nigra. Nephasoma prioki (Sluiter, 1881) Phascolosoma prioki Sluiter, 1881:152-153, pl. 1, figs. 5 and 9; 1891:115; 1902:34.— Selenka et al., 1883:37.—Selenka, 1885: 12-13. Golfingia prioki.—Stephen and Edmonds, 1972:153.—Cutler, 1977b:153. Material examined. —ZMUA, Type specimen. This taxon has a very uncertain founda- tion. The first four references are all repe- titions of the single holotype. Selenka’s 1885 record was based on one additional worm. The three worms in Cutler (1977b:153)“.. . seem to fit this poorly known species.”” What has been overlooked is the presence of small but real contractile vessel villi and the na- ture of the tentacles which are withdrawn. At present, after our work on the genus Thy- sanocardia, it is clear that these four worms fit the criteria for that genus and are closest to 7. nigra. The age and preservation his- tory of the material makes positive identi- fication difficult. The depth of Sluiter’s worm (962 m) is also greater than other 7. nigra. Nevertheless, NV. prioki now becomes a ju- nior synonym of 7. nigra. Nephasoma vitreum (Roule, 1898) Phascolosoma vitreum Roule, 1898:386; 1906:86-90. Golfingia vitrea.—Stephen and Edmonds, 1972:158-159. Material examined. —None. The type and only representative of this species cannot be located and in the inter- vening 88 years no additional specimens fit- ting this rather peculiar description have been collected from these bathyal eastern VOLUME 99, NUMBER 4 Atlantic waters. We therefore place this name on the list of incertae sedis. Acknowledgments We are indebted to P. Gibbs (Plymouth, England) and G. Murina (Sevastopol, USSR) for their constructive criticisms, loan of specimens, and interest. G. C. Steyskal (USNM) provided guidance in emending the species names. Financial support was pro- vided by U.S. National Science Foundation Grant #BSR 83-14301 and Hamilton Col- lege. The cooperation of the following per- sons and institutions in the loan of reference material and/or providing access to their collections was essential to the completion of this project and is appreciated: E. Wil- lassen (ZMUB); R. Sims (BMNH); M. Jones (USNM); A. V. Ivanov (ZIAS); G. Hartwich (MNHU); B. Thompson, Long Beach, CA; Irish National Museum, Dublin; Institut voor Taxonomische Zoologie, Amsterdam. Literature Cited Akesson, B. 1958. A study of the nervous system of the sipunculoideae, with some remarks on the development of the two species Phascolion strombi Montagu and Golfingia minuta Kefer- stein.— Undersokninger 6ver Oresone 38:1-249. Amor, A. 1975. Notas sobre Sipuncula de la Argen- tina, Brasil y Peru.—Physis 34(88):113-120. Andreae, J. 1881. Zur Anatomie des Sipunculus nu- dus L.—Zoologischer Anzeiger 4:477—48 1. Augener, H. 1903. Beitraége zur Kenntnis der Ge- phyreen nach Untersuchung dere im Gottinger zoologischen Museum befindlichen Sipunculi- den und Echiuriden.—Archiv fiir Naturge- schichte 69:297-371. Baird, W. 1868. Monograph on the species of worms belonging to the subclass Gephyreae. — Proceed- ings of the Zoological Society of London, 1868: 77-114. Benham, W. B. 1904. Further notes on the sipun- culids of New Zealand.—Transactions of the Royal Society of New Zealand 37:301-308. 1909. Annelida and Sipunculoidea.—Rec- ords of the Canterbury Museum 1:71-82. 1922. Gephyrea Inermia.— Australian Ant- arctic Exposition 1911-14, Scientific Reports Series C, Zoology and Botany VI (5):5—21. Chamberlain, R. V. 1920. Gephyrea.— Report of the Canadian Arctic Expedition 1913-18 9:3D-7D. 569 Cuenot, L. 1922. Sipunculiens, Echiuriens, Priapu- liens. —Faune de France, Paris 4:1-30. Cutler, E. B. 1969. New species of Sipuncula from the western North Atlantic. — Proceedings of the Biological Society of Washington 82:209-218. . 1973. Sipuncula of the western North Atlan- tic.— Bulletin of the American Museum of Nat- ural History 152(3):105-204. . 1977a. The bathyal and abyssal Sipuncula. — Galathea Report 14:135-156. 1977b. Shallow water sipunculans collected by the Galathea Expedition (Sipuncula).— Steenstrupia 4:151-155. . 1979. A reconsideration of the Golfingia sub- genera Fisherana Stephen, Mitosiphon Fisher, and Apionsoma Sluiter (Sipuncula).—Zoologi- cal Journal of the Linnean Society 65:367—384. ——,, and N. J. Cutler. 1979a. Madagascar and Indian Ocean Sipuncula.— Bulletin du Muséum National d’Histoire Naturelle, Paris 4e(1):941- 990. ,and . 1979b. Campagnes de la Calypso au large des cotes Atlantiques Africaines (1956 et 1959) et Sud-Américaines (1960-1961). 23. Sipuncula.—Campagnes de la Calypso XI:103- 109. , and . 1980a. Sipuncula from Southern Brazil.— Boletim do Institut Oceanografico, Sao Paulo 29(1):1-8. , and 1980b. Deep water Sipuncula from the Gulf of Gascogne. — Journal of the Ma- rine Biological Association of the United King- dom 60:449-459. , and 1980c. Sipuncula collected by the R/V “Vema.” —Journal of Zoology, London 190:193—209. ———_,, ———., and T. Nishikawa. 1984. The Sipun- cula of Japan: Their systematics and distribu- tion.— Publications of the Seto Marine Biolog- ical Laboratory 29(4/6):249-322. , and P. E. Gibbs. 1985. A phylogenetic anal- ysis of the higher taxa in the phylum Sipuncu- la.—Systematic Zoology 34(2):162-173. , and V. V. Murina. 1977. On the sipunculan genus Golfingia Lankester, 1885.—Zoological Journal of the Linnean Society 60:173-189. Danielssen, D. C., and J. Koren. 1880. New northern Gephyrea.— Annals and Magazine of Natural History (5)6:462-465. , and 1881. Fra den norske Norhavs- expedition (1876-1878), Gephyreer. — Nytt Ma- gasin for Naturvidenskapene 26:44-66. Diesing, K. M. 1851. Systema Helminthum. Brau- miller, Vindobonae. pp. 1-558. . 1859. Revision der Rhyngodeen.—Sitzungs- berichte der mathematisch-naturwissenschaft- liche Klasse, Akademie der Wissenschaften in Wien 37:719-782. 570 Ditadi, A. S. F., and A. E. Migotto. 1981. On the sipunculan Golfingia confusa (Sluiter, 1902).— Papeis Avulsos de Zoologie, Sao Paulo 34(10): 125-134. Edmonds, S. J. 1960. Sipunculids from New Zealand and the Chatham Islands.—New Zealand De- partment of Scientific and Industrial Research Bulletin 139(4):159-167. 1976. Three sipunculan species (two new) from New Zealand.—New Zealand Journal of Marine and Freshwater Research 10(1):21 7-224. 1980. A revision of the systematics of Aus- tralian sipunculans (Sipuncula).— Records of the South Australian Museum (Adelaide) 18(1):1- 74. Fischer, J. 1914. Die Sipunculiden der Nord und Ostsee, unter Beriichsichtigung von Formen des nordatlantischen Gebietes.— Wissenschaftlich- en Meeresuntersuchungen Abteilung Kiel 16:85— 125. Fischer, W. 1895. Die Gephyreen des Naturhisto- rischen Museums zu Hamburg.—Abhandlun- gen aus dem Gebiet der Naturwissenschaften, Hamburg 13:1-24. 1914. Weitere Mitteilungen tiber die Gephy- reen des Naturhistorischen (Zoologischen) Mu- seums zu Hamburg.—Jahrbuch der Hamburg- ischen Wissenschaftlichen Anstalten 31:1—28. 1916. Die Gephyreensausbeute der deutsch- en Tiefsee Expedition (1898-1899), vorlaufige Mitteilung.— Zoologischer Anzeiger 48:14—20. 1919. Gephyreen der Siid-westkiiste Aus- traliens.— Zoologischer Anzeiger 50:277-285. 1922a. Gephyreen des Reichsmuseums zu Stockholm.— Arkiv fiir Zoologie 14(19):1-39. 1922b. Gephyreen der Arktischen Meere. — Wissenschaftlichen Meeresuntersuchungen Ab- teilung Helgoland 13:229-246. . 1922c. Gephyreen der deutschen Tiefsee-Ex- pedition, auf dem Dampfer “Valdivia” 1898- 1899.—Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition, Valdivia 22(1): 1-26. 1923. Gephyreen des Golfes von Siam.— Videnskabelige Meddelelser fra Dansk Natur- historisk Forening 76:21-27. 1925. Echuiriden, Sipunculiden, und Pria- puliden. Jn G. Grimpe and E. Wagler, eds. Die Tierwelt der Nord-und Ostsee, Akademische Verlagsgesellschaft, Leipzig, pp. 1-55. 1929. Die Sipunculiden, Priapuliden und Echiuriden der Arktis. Jn F. Romer and F. Schaudinn, eds. Fauna Arctica, pp. 451-490. Fisher, W. K. 1950. The sipunculid genus Phasco- losoma.— Annals and Magazine of Natural His- tory (12)3:547-552. 1952. The sipunculid worms of California PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and Baja California. — Proceedings of the United States National Museum 102:371-450. Forbes, E. 1841. A history of British starfishes and other animals of the class Echinodermata. Lon- don, J. van Voorst, 267 pp. Frank, P. G. 1983. A checklist and bibliography of the Sipuncula from Canadian and adjacent waters. —Syllogeus 46:1-47. Gadd, G. G. 1911. Verzeichnis der Gephyreen des Kolagolfes und zwei neue Species der Phasco- losoma.—Trudy Leningrad Obschchestva Es- testvois Pytatelei 42:79-93. Gerould, J. H. 1908. A comparison of the cephalic organs of certain sipunculids. — Science 27:488- 489. . 1913. The sipunculids of the eastern coast of North America.—Proceedings of the United States National Museum 44:373-437. Gibbs, P.E. 1973. On the genus Golfingia (Sipuncula) in the Plymouth area with a description of a new species.—Journal of the Marine Biological As- sociation of the United Kingdom 53:73-86. . 1975. Gametogenesis and spawning in a her- maphroditic population of Golfingia minuta (Si- puncula).— Journal of the Marine Biological As- sociation of the United Kingdom 55:69-82. . 1977a. British sipunculans.—Synopses of the British Fauna 12:1-35. . 1977b. On the status of Golfingia intermedia (Sipuncula).—Journal of the Marine Biological Association of the United Kingdom 57:109-112. . 1982. The synonymy of the Golfingia species assigned to the Abyssorum section (Sipuncu- la).—Sarsia 67:119-122. 1986. The taxonomy of some little-known Sipuncula from the north-east Atlantic region including new records.—Journal of the Marine Biological Association of the United Kingdom 66:335-341. , E. B. Cutler, and N. J. Cutler. 1983. A review of the genus Thysanocardia Fisher (Sipuncu- la).— Zoologica Scripta 12(4):295-304. Herubel, M. A. 1903. Sur la distribution et les affini- tés réciproques des Sipunculides.—Bulletin de la Société Zoologique de France 28:99-111. 1907. Recherches sur les Sipunculides.— Mémoires de la Société Zoologique de France 20:107-418. . 1925. Quelques echiurides et sipunculides de cétes du Maroc.—Bulletin du Société des Sci- ences Naturelle et Physiques du Maroc 5:260- 263. Keferstein, W. 1863. Beitrage zur Kenntnis der Gat- tung Phascolosoma F. S. Leuck. Untersuchun- gen liber niedere Seethiere. — Zeitschrift fur Wis- senschaftliche Zoologie 12:35-51. 1865. Beitrage zur anatomischen und sys- VOLUME 99, NUMBER 4 tematischen Kenntniss der Sipunculiden. — Zeitschrift fiir Wissenschaftliche Zoologie 15: 404-445. Koren, J., and D. C. Danielssen. 1875. Bidrag til de norske Gephyrea Naturhistorie.— Nytt Magasin for Naturvidenskapene 21:108-138. , and 1877. The natural history of the Norweigian gephyreae. In M. Sars, ed. Fauna Littioralis Norvegiae, Druck und Verlag von Jo- hann Dahl, Christiania, pp. 111-156. Lanchester, W. F. 1905. On a collection of sipun- culids made at Singapore and Malacca.—Pro- ceedings of the Zoological Society of London 1: 26-28. Leroy, P. 1936. Les sipunculiens du Muséum d’His- toire Naturelle de Paris.— Bulletin du Muséum National d’Histoire Naturelle, Paris 8(2):423- 426. Lindroth, A. 1941. Echiurida, Sipunculida und En- teropneusta aus dem Skagerack 1933.—Zoolo- giska Bidrag fran Uppsala 20:443-452. Murina, V. V. 1957. Sipunculids collected on the first trip of the complex Antarctic expedition on the ‘Ob’ in 1956.—Zoologicheskii Zhurnal 36(7): 992-998. 1958. On the systematics of two closely re- lated species of the deep-water sipunculids of the genus Golfingia according to the materials of the “Vitjaz” expeditions in 1950-1955.— Zoologicheskii Zhurnal 37:1624-1633. 1964a. New and rare species of sipunculids of genus Golfingia. —Studies of the Institute of Oceanology, Academy of Science USSR 69:216- 253. 1964b. Sipunculid fauna of the Mediterra- nean Sea.—Studies of the Sevastapol Biological Station 17:51-76. . 1967a. On the sipunculid fauna of the littoral of Cuba.—Zoologicheskii Zhurnal 46:35—47. . 1967b. Report on the sipunculid worms from the sublittoral zone of Cuba and the Gulf of Mexico.— Zoologicheskii Zhurnal 54(9):1329- 1339. 1967c. New facts about the distribution of marine sipunculid worms in the Red Sea and the Gulf of Aden. Jn ““Some results of Expedi- tion III to the Red Sea Inbuma.” Express in- formation, Academy of Sciences Ukrainian So- viet Socialist Republic, pp. 52-56. . 1968a. On the distribution of new sipunculid fauna from Cuba and Gulf of Mexico.— Revue Roumaine de Biologie-Zoologie 13(6):42 1-423. 1968b. New data concerning the occurrence of abyssal sipunculids in the Atlantic Ocean. — Zoologicheskii Zhurnal 47(2):195-199. . 1971a. On the species composition and ecol- ogy of sipunculids of the Red Sea. Jn A. O. Sal Kovalevski, Shelf benthos of the Red Sea, Academy of Science USSR, pp. 76-88. . 1971b. On the occurrence of deep-sea sipun- culids and priapulids in the Kurile-Kamchatka Trench.— Trudy Instituta Okeanologii Akade- mii Nauk SSSR 92:41-45. 1972. Contribution of the sipunculid fauna of the Southern Hemisphere.—Zoologii Insti- tuta Akademii Nauk SSSR 11(19):294-314. 1973a. New species of Pacific Ocean sipun- culids.—Zoologicheskii Zhurnal 52:942—945. 1973b. The fauna of sipunculids from the Peruvian-Chilean Trench.—Zoologicheskii Zhurnal 5:66-71. . 1974a. Contributions to the knowledge of the fauna of sipunculid worms from the South At- lantic based on data of the ““Akademik Kurcha- tov’? expedition in 1971.—Trudy Instituta Okenologii Akademii Nauk SSSR 98:228-239. 1974b. New subspecies of sipunculid worms. — Zoologicheskii Zhurnal 53:1713-1717. 1975. New taxa of the genus Golfingia. — Zoologicheskii Zhurnal 54:1085-1089. 1976. New and rare species of sipunculids from the East China Sea.— Vestnik Zoologii 2: 62-67. 1977. A new species of Sipuncula from the Atlantic Ocean. — Galathea Report 14:133-134. . 1978. On the sipunculid and priapulid fauna of the southern Pacific.—Trudy Instituta Okeanologii 113:120-131. Ostroumov, A. 1909. Sur les gephyriens du nord de la mer du Japan.—Ezhegodnik Zoologischeskii Muzei 14:319-321. Paul, G. 1909. Uber Petalostoma minutum Kefer- stein und verwandte Arten.—Jahrbucher Ab- theilung fiir Anatomie 29:1-50. Pergament, T. S. 1940. Ona new genus Nephasoma from the Arctic Ocean. Results of cruising ex- pedition on icebreaker G. Sedov 1937-1940.— Arctic Science Institute of Arctic Sea Route, Council of Ministry USSR:1-3. Quatrefages, A. 1865. Gephyrea Inermia.— Histoire naturelle des Annelés marins et d’eau douce 2: 599-632. Roule, L. 1896. Gephyriens. Résultats scientifiques de la campagne du “‘Caudan”’ dans le Golfe de Gascogne, 1895.—Annales de |’Université de Lyon 26:473-474. 1898. Sur les gephyriens des grans fonds de la mer.—Comptes Rendus de |’Académie des Sciences, Paris 127:197-199. 1906. Annelides et Gephyriens.—Expedi- tions Scientifiques du ““Travaileur” et du “Tal- isman” 8:1-102. Saiz-Salinas, J. I. 1980. Estudio de los sipunculidos (Sipuncula) de los fondos litorales y circalitor- 572 ales de las costas de la peninsula Iberica, Islas Baleares, Canarias y mares Adyacentes. [Un- published masters thesis.] Departamento de Biologia, Universidad del Pais Vasco, 120 pp. 1984. Gusanos sipunculidos (Sipuncula) de varias localidades de la costa de Portugal. Jn V. Monteiro-Marques, ed. ACTAS do 4th Sim- posio Iberico de Estudos do Benthos Marinho, Associacao des Estudantes da Faculdade de Cien- Cias de Lisboa, pp. 177-188. Sars, M. 1851. Beretning om en i Sommeren 1849 foretagen zoologisk Reise i Lofoten og Finmar- ken.—Nytt Magasin for Naturvidenskapene 6: 121-125, 190-211. Selenka, E. 1885. Report on the Gephyrea collected by H.M.S. Challenger during 1873—76.— Report of Scientific Results of Voyage of Challenger, Zoology 13(36):1-25. —., J. G. de Man, and C. Biilow. 1883. Die Si- punculiden, eine systematische Monographie. — Semper, Reisen in Archipelago Philippinen 2, 4:1-131. Shipley, A. E. 1899. A report on the Sipunculoidea collected by Dr. Willey at the Loyalty Islands and in New Britain. Jn A. Willey, ed. Zoological Research, pp. 151-160. Sluiter, C. P. 1881. Beitrage zu der Kenntnis der Gephyreen aus dem Malayischen Archipel. Zweite Mittheilung. — Natuurkundig Tijdschrift voor Nederlandisch-Indie 41:148-171. 1891. Die Evertebraten aus der Sammlung des Koniglichen naturwissenschaftlichen Ver- eins in Nederlandisch-Indien in Batavia.—Na- tuurkundig Tijdschrift voor Nederlandisch-In- die 50:102-123. 1900. Géphyriens (Sipunculides et Echiuri- des) provenant des campagnes de |’Hirondelle et de la Princesse Alice, 1886-1897, Fascicule 15:1-29. 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. 1912. Géphyriens (Sipunculides et Echiuri- des) provenant des campagnes de |’Hirondelle Alice, 1898-1910.—Résultants Campagnes Scientifique Prince Albert I 36:1—36. Southern, R. 1908. A new Irish gephyrean.—Irish Naturalist 17:83-86. 1913. Gephyrea of the coasts of Ireland.— Scientific Investigations of the Fisheries Branch, Department of Agriculture for Ireland 3:1—46. Stephen, A. C. 1934. The Echiuridae, Sipunculidae, and Priapulidae of Scottish and adjacent waters.— Proceedings of the Royal Society of Edinburgh 22:159-186. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1948. Sipunculids.—B.A.N.Z.A.R.E. Re- ports, Series B 5(4):213-—220. Stephens, A. C., and S. J. Edmonds. 1972. The phyla Sipuncula and Echiura. London, Trustees of the British Museum (Nat. Hist.), 528 pp. tenBroeke, A. 1925. Westinische Sipunculiden und Echiuriden. — Bijdragen tot de Dierkunde 24:8 1— 96. 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. 1905. Northern and Arctic invertebrates in the collection of the Swedish State Museum I. Sipunculids.—Kungliga Svenska Vetenskaps- Akademiens Handlingar 39(1):1—130. . 1911. Priapulids and Sipunculids dredged by the Swedish Antarctic Expedition 1901-1903 and the phenomenon of bipolarity.— Kungliga Svenska Vetenskaps-Akademiens Handlingar 47(1):3-36. Thompson, B. E. 1980. A new bathyal sipunculan from southern California, with ecological notes. — Deep-Sea Research 27a:95 1-957. Verrill, A. E. 1874. Explorations of Casco Bay by the U.S.F.C. in 1873.—Proceedings of the Ameri- can Association for the Advancement of Science 22:340-393. Wesenberg-Lund, E. 1925. Gephyreer.—Conspectus Faunae Groenlandicae Bianco Lunos Bogtryk- keri, Kobenhavn 23:81-91. 1930. Priapulidae and Sipunculidae.—The Danish “‘Ingolf’ Expedition 4(7):1-44. 1932. The Godthaab Expedition 1928 Ge- phyrea.— Meddelelser om Gronland 79(3):1-18. 1933. The collection of gephyreans in the Royal Museum of Natural History Belgium.— Bulletin du Musée Royal d’Histoire Naturelle de Belgique 9(6):1—16. . 1937a. Gephyrea.— Zoology of Iceland 2(23): 1-16. 1937b. The zoology of East Greenland Ge- phyreans.— Meddelelser om Grgnland 121(1): 1-25. 1939. Gephyreans from Swedish waters in the Museum of Natural History of Gothen- burg.—Goteborgs Kungliga Vetenskaps Vitter- hets-Samhalles Handlingar Femte Foljden 6(6): 3-35. : 1954. Priapuloidea, Sipunculoidea and Echiuroidea.— Bulletin Institut Royal des Sci- ences Naturelles de Belgique 30(16):1-18. 1955. Reports of Lund University Chile Ex- pedition, 1948-49. 19. Gephyrea from Chile.— Lunds Universitets Arsskrift 51(10):1-23. 1959. Sipunculoidea and Echiuroidea from VOLUME 99, NUMBER 4 573 tropical West Africa.—Atlantide Report 5:177- Appendix 1.—Certain morphological characters of 210. Nephasoma species. 1963. South African sipunculids and echiu- roids from coastal waters. — Videnskabelige ate pets Meddelelser fra Dansk Naturhistorisk Forening vert anterior Origin Muscles 125:101-146. Aronk length # of to anus of retrac- : ength as % gut by % of tors (% Fix- in mm trunk coils trunk oftrunk) Spindle ing (NJC) Biology Department, Hamilton Nephasoma flagriferum College, Clinton, New York 13323; (EBC) 120 108 70 8 23 i) Biology Department, Utica College of Syr- 97 15 &S 4 28 hah acuse University, Utica, New York 13502. 90 5170 1 22 sad EO 85 67 6 4 Di + 2 83 87 ~=—80 2 37 + 0 82 52 60 4 29 + 0 77 130 70 4 35 + 0 VW 56 ~=6.60 1 23 + 1 75 ? 60 1 24 0 0 75 85 60 4 47 + 0 UD ? 85 7 22 + 0 63 106 60 3 28 + 1 61 162 80 2 48 + 0 58 121 25 2 16 + 1 55 89 8650 4 33 + 0 54 111 2 4 20 + 1 40 165 6 4 35 ot 1 35 71 14 9 37 =f 0 32) 147 45 9 25 sF 0 DY 96 ? ? ? ? D 9 200 25 11 39 +f 0 7 457 20 7 57 + 1 6 DSO 2D 7 42 + 0 5 140 25 20 50 u v 5 100 20 0 40 ? ? 4 100 20 0 50 ry 0 35 143 15 14 43 ? ? Nephasoma constrictum (posterior) 30 33 = 40 3 53 0 0 29 62 40 5 55 0 0 29 45 ? 7 69 ar 1 25 88 30 4 64 0 0 23 104 25 7 57 ar 0 22 59 40 9 59 0 0 20 60 35 3 55 + 0 20 80 §@630 10 50 + 0 18 78 35 6 56 0 1 17 82 28 18 59 + 1 15 60 35 3 67 + 1 13 Ti 30 12 38 + 0 12 83 =. 20 8 67 + 0 10 80 30 10 50 + 0 8 113 +20 6 50 ? ? 7 157 15 11 71 0 0 7 214 20 14 57 + 0 PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 574-579 ON THE SPECIES AND POPULATIONS OF THE GENUS ACANTHOCEPHALUS (ACANTHOCEPHALA: ECHINORHYNCHIDAE) FROM NORTH AMERICAN FRESHWATER FISHES: A CLADISTIC ANALYSIS Omar M. Amin Abstract. —Cladistic analysis provided greater understanding and further sup- port of the evolutionary relationships among the species and populations of the genus Acanthocephalus from North American freshwater fishes outlined earlier by Amin (1985). It is proposed that A. dirus, A. tahlequahensis and A. alabamensis are monophyletic with A. dirus representing a probable successful and persistent general ancestor. The two southern species are more closely related to each other than either one is to 4. dirus, probably evolved allopat- rically, and now exhibit new restricted distributions. Within A. dirus, the Mis- sissippi River and Wisconsin-Lake Michigan populations are more closely related to each other than either one is to the New England population, which may have been evolving in isolation longer than the former two. The recent revision of the genus Acan- thocephalus from North American fresh- water fishes by Amin (1984) and Amin and Huffman (1984) established the presence of three species, A. dirus (Van Cleave 1931), A. tahlequahensis Oetinger and Buckner, 1976, and A. alabamensis Amin and Wil- liams, 1983. Acanthocephalus dirus has the widest host and geographical distribution and exhibits the greatest morphological variability. It is found in 65 species and 16 families of fish mostly in the Mississippi River drainage system (or waters previously connected to it in Ohio and Lake Erie) in 13 states in the U.S. (Amin 1985). Acan- thocephalus tahlequahensis is found in four species and two families of fish from Okla- homa and A. alabamensis in six species and four families from Alabama. Three distinct populations are recognized within A. dirus by Amin (1984): the Mis- sissippi River population; the Wisconsin- Lake Michigan population (=A. parksidei Amin, i975) and the New England popu- lation (=A. jacksoni Bullock, 1962). The Mississippi River population has the widest geographical (in 10 states) and host (46 fish species in 11 families) distribution and is more variable morphologically than the Wisconsin-Lake Michigan population (in two states and 22 species and 10 families of fish) or the New England population (in two states and 14 species and nine families). Meristogram patterns (unpublished) were distinctly different in each of the three species. Within A. dirus, the meristogram patterns were similar in all the populations studied by Amin (1984) except the Gleason population from Kentucky which showed some elements of uniqueness corresponding with certain peculiarities discussed by Amin (1984). Based on geological evidence as well as on variability, host and geographical distri- bution, Amin (1985:214) proposed that “early A. dirus ancestors were probably found in the Mississippi River basin before the Wisconsin glaciation.”’ The establish- ment of the geographically isolated Wiscon- sin-Lake Michigan population “‘must have taken place with Mississippi River elements from the Des Plaines-Illinois River system VOLUME 99, NUMBER 4 575 Table 1.—Character distribution among species of genus Acanthocephalus from North American freshwater fishes and outgroups. North American species of Acanthocephalus Outgroups (Europe) Ill X, X, I II A. tahlequa- Character distribution A. lucii' A. anguillae' A. dirus A. alabamensis hensis (1) Mean male body length (mm) Large Large Moderate Small Small (>5) (>5) (3-4) (<3) (<3) (2) Mean female body length (mm) Large Large Moderate Small Small (> 10) (> 10) (7-9) (<6) (<6) (3) Body shape Cylindrical Cylindrical Cylindrical Cylindrical Spindle (4) Mean anterior testis length (um) Large Large Moderate Small Small (> 800) (> 800) (450-800) (<450) (<450) (5) Mean anterior testis length/body Small Small Large Large Small (%) (<15) (<15) (> 19) (> 19) (<15) (6) Mean anterior testis width/body Large Moderate Large Large Small width (%) (> 50) (42-46) (>50) (>50) (<42) (7) Mean no. proboscis hooks/row Small Small Moderate Small Large (males) (<9) (<9) (9-10) (<9) (>11) (8) Mean no. proboscis hooks/row Small Small Moderate Moderate Large (females) (<9) (<9) (9-11) (9-11) (>11) (9) Mean length of largest proboscis Large Large Moderate Small Small hooks (males) (um) (> 60) (> 60) (50-60) (<50) (<50) (10) Mean length of largest proboscis Large Large Moderate Small Small hooks (females) (um) (>70) (>70) (60-70) (<60) (<60) (11) Meristogram patterns A? B? C D E ' From Petrochenko (1956). 2 Presumably. be) (tributaries of the Mississippi River... in post-glacial streams like the Pike River “*_.. after the withdrawal of the Lake Mich- igan lobe of the Wisconsinan ice sheet from the area between 15,000 and 12,000 years B.P.” (before the present). Based on mor- phological evidence alone, the geographi- cally isolated New England population ap- pears to have also originated from a Mississippi River A. dirus-like source (Amin 1985). Acanthocephalus tahlequahensis has a limited distribution in an Oklahoma trib- utary of the Mississippi River and 4. ala- bamensis is found in the Mobile Bay drain- age system which had continuous fauna with the Mississippi River before barriers to re- cent dispersal arose. The two southern species are found more typically in certain species of Etheostoma not parasitized by other species of Acanthocephalus. The above information provides a brief background of the inter- and intraspecific associations within the genus Acanthoceph- alus from North American freshwater fishes which lends itself to further analysis within a cladistic context. To date, cladistic anal- ysis has not been used as an aid to the un- derstanding of acanthocephalan evolution- ary biology. This method, however, was found amenable to this study as it provided greater understanding of and further sup- port for the evolutionary relationships brief- ly outlined above. Materials and Methods Methods in Brooks et al. (1985) and Wi- ley (1981) as well as suggestions by Drs. D. R. Brooks and J. N. Caira were instrumental in the understanding of the findings pre- 576 Table 2.—Character distribution among populations of Acanthocephalus dirus and outgroups. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Outgroups Acanthocephalus dirus populations me I Il xX, A. tahlequa- X; Mississippi Wisconsin- Ill Character distribution A, alabamensis hensis A, lucti River Lake Michigan New England (1) Mean male body Small Small Large Moderate Moderate Moderate length (mm) (<3) (<3) (>5) (3-4) (3-4) (3-4) (2) Mean female body Small Small Large Moderate Moderate Moderate length (mm) (<6) (<6) (> 10) (7-9) (7-9) (7-9) (3) Body shape Cylindrical Spindle Cylindrical Cylindrical Cylindrical Robust an- teriorly (4) Mean anterior testis Small Small Large Moderate Moderate Moderate length (um) (<450) (<450) (800) (450-800) (450-800) (450-800) (5) Mean anterior testis Large Small Small Large Large Moderate length/body length (>19) (<15) (<15) (> 19) (> 19) (17-18) (%) (6) Mean anterior testis Large Small Large Large Large Moderate width/body width (%) (>50) (<42) (> 50) (> 50) (> 50) (42-46) (7) Mean no. proboscis Small Large Small Moderate Moderate Small hooks/row (males) (<9) (> 11) (<9) (9-10) (9-10) (<9) (8) Mean no. proboscis Moderate Large Small Moderate Moderate Small hooks/row (females) (9-11) (>11) (<9) (9-11) (9-11) (<9) (9) Meanlengthoflargest Small Small Large Moderate Moderate Large proboscis hooks (<50) (<50) (> 60) (50-60) (50-60) (> 60) (males) (um) (10) Meanlength oflargest Small Small Large Moderate Moderate Large proboscis hooks (fe- (<60) (<60) (>70) (60-70) (60-70) (>70) males) (um) (11) Meristogram pattern A B 2 D D D ' From Petrochenko (1956). * Presumably. sented in a cladistic context and in the con- struction and interpretation of cladograms. Results and Discussion Cladograms were made with the assump- tion that the three North American species are each others’ closest relatives, and that none has closer relatives on other continents (see following parts). Adequate data were available for four species of Acanthocepha- lus: A. lucii (Muller, 1776), A. anguillae (Muller, 1780), A. clavula (Dujardin, 1845), and A. falcatus (Frolich, 1789) for use as outgroups. Resolving all characters used in the analysis (Table 1) was possible by using the first two species, i.e., their plesiomor- phic states satisfactorily polarized ingroup characters. These plesiomorphic states in- cluded large and cylindrical bodies, large testes, larger and fewer proboscis hooks per row. The choice of those characters was based on characteristics of body, testes, and proboscis hooks, the evolutionary signifi- cance of which was discussed by Amin (1984). Note that in the absence of other data (Tables 1, 2) a linear transformation series is to be preferred (Mickevich 1982). The data matrix is shown in Table 3. The resulting cladogram (Fig. 1) has a high over- all consistency index (minimum number of steps/actual number of steps; see Farris et al. 1970a, b) of 95% and synapomorphic consistency of 100% indicating a low degree of parallel evolution in the characters used VOLUME 99, NUMBER 4 and giving a high degree of confidence in the pattern hypothesized. Two other clado- grams with lower consistency values were excluded from consideration. The hypoth- esized pattern (Fig. 1) indicates that (1) the three North American species (ingroup) are monophyletic. X, and X, plus the three North American species are postulated to form a monophyletic group based on their shared possession of a suite of four derived traits (3, 5, 6, 7 in Fig. 1). Using the two ““X”? species as outgroups, I, II and III are postulated to be a monophyletic group based on traits 1, 2, 4, 8, 9, 10 in Fig. 1. The hypothesis of monophyly is supported by the very high goodness of fit statistics. (2) A. alabamensis and A. tahlequahensis are more closely related to each other than either one is to A. dirus. The two branching points on the cladogram indicate a north-south split (A. dirus and the ancestor of A. alabamensis and A. tahlequahensis). Prior to its differ- entitation into distinct populations, A. dirus was of the same age as the common ancestor of the two southern species. The most reasonable events are a wide- spread ancestor divided into a southern population giving rise to A. alabamensis and A. tahlequahensis and a more northern pop- ulation forming A. dirus. Under the above conditions of monophyly, speciation would be allopatric. Figure 1 and Wiley’s (1981) identical diagram (Fig. 2-10b) indicate al- lopatric speciation. While the identification of the persistent ancestral A. dirus popula- tion is possible, it is not easily detectable given available data above. It is proposed, however, that based on its broad geograph- ical and host distributions and its great mor- phological variability. Acanthocephalus di- rus appears to be a successful persistent general ancestor while A. tahlequahensis and A. alabamensis seem to exhibit new re- stricted distributions. Acanthocephalus di- rus may also be interpreted as a successful adaptable new species with the other two species exhibiting relictual distributions. Some combinations of the two interpreta- 577 Table 3.—Data matrix of six taxa and 11 characters based on Table 1. Character state * x % = = = = KB OwoO ONDA MN PWN Cc & © © © © OC CO So © wD ao oo = Eo eC eo OS © oe aa Bore ore BRNNRK CORK NON WN OMANNNRK KH ONK KN NY tions may also be theorized. The first in- terpretation is, however, preferred because of the ancestral position of A. dirus on the phylogenetic tree (Fig. 1). This is concor- dant with the contention that the anatom- ical diversity and the broad host and geo- graphical distributions of this species do represent generalized persistent ancestral traits and that distinguishing states of the restricted A. alabamensis and A. tahlequa- hensis, e.g., small size, are more recently derived. The evolutionary relationships among the three A. dirus populations are expressed in cladogram Fig. 2 based on data summarized in Tables 2 and 4. This cladogram has an overall and synapomorphic consistency in- dex of 100%. It suggests that (1) the Mis- sissippi River and the Wisconsin-Lake Michigan populations are more closely re- lated to each other than either one is to the New England populations, (2) the latter population may have been evolving in iso- lation longer than the former two if equal rates of evolution of all characters are as- sumed. Of the three extant populations, the New England population is the most highly differentiated, and (3) the present Missis- sippi River population and that of Wiscon- sin-Lake Michigan are equally derived. The above interpretation of cladogram 578 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1, 2. Acanthocephalus from freshwater fishes (Fig. 1, from Tables 1 and 3) and among the three populations of A. dirus (Fig. 2, from Tables 2 and 4). For asterisks see Tables 1 and 3. Circled numbers (A-E, Table 1 and A-D, Table 2) indicate specific meristogram patterns different in each species. Fig. 2 provides additional evidence to those based on geological and morphological grounds (Amin 1985) supporting the prop- osition that the Wisconsin-Lake Michigan population of A. dirus dispersed from an early Mississippi River-based source and then became geographically isolated less than 15,000 years ago. Given sufficient time and continued isolation, this situation would be a classical example of Van Cleave’s (1952) earlier proposition that in Palaeacantho- cephala, isolation might allow the normal extremes in a highly variable species to be- come segregated as distinct species. The in- terpretation also provides further insights into those based on morphological and clin- al variations suggesting a similar process for the New England population even though the origin of its source remains to be iden- tified. Cladogram Fig. 2 indicates that this highly differentiated population may have been evolving in isolation longer than the other two. At the present time, it is not known whether the dispersal of A. dirus led to the isolation of A. alabamensis and A. tahle- Cladograms showing the phylogenetic relationships among the three North American species of quahensis due to competitive exclusion, or that some geological or host related changes which isolated the latter two species also allowed the dispersal of A. dirus after the fact. It is, however, clear that certain geo- logical events affected other aspects of the distribution and evolution of these acan- thocephalans, e.g., the geographically iso- lated northern populations of A. dirus, mak- Table 4.—Data matrix of six taxa and 11 characters based on Table 2. Taxon Character state Pad * a4 — = = = = q KODOMAIDWAWHH— MoVototetoloiOnrolore NOOFKFKF KH OK OO Qeewoo HH ore BNNONDONONWN BNNONGTCONOON Db Bere NONNNNNW — pt VOLUME 99, NUMBER 4 ing the proposed geological scenario the best hypothesis presently possible. Acknowledgments I am grateful to Drs. D. R. Brooks, Uni- versity of British Columbia, Vancouver, and J. M. Caira, University of Maine, Orono, for help with the cladistic analysis. The sup- port and inspiration of Michele Meinert are gratefully noted. Literature Cited Amin, O. M. 1984. Variability and redescription of Acanthocephalus dirus (Acanthocephala: Echi- norhynchidae) from freshwater fishes in North America. — Proceedings of the Helminthological Society of Washington 51:225-—237. 1985. Hosts and geographic distribution of Acanthocephalus (Acanthocephalus: Echino- rhynchidae) from North American freshwater fishes, with a discussion of species relation- ships.— Proceedings of the Helminthological Society of Washington 52:210-—220. , and D. G. Huffman. 1984. Interspecific vari- ability in genus Acanthocephalus (Acanthoceph- 57/9) alus: Echinorhynchidae) from North American freshwater fish, with a key to species.—Pro- ceedings of the Helminthological Society of Washington 51:238-240. Brooks, D. R., J. N. Caira, T. R. Platt, and M. H. Pritchard. 1984. Principles and methods of phylogenetic systematics: A cladistics work- book. The University of Kansas Museum of Natural History Special Publication No. 12, v + 92 pp. Farris, J. S., A. G. Kluge, and M. J. Eckardt. 1970a. A numerical approach to phylogenetic system- atics.—Systematic Zoology 19:172-189. , , and 1970b. On predictivity and efficiency.—Systematic Zoology 19:363- 372. Mickevich, M. F. 1982. Transformation series anal- ysis. —Systematic Zoology 31:461—478. Petrochenko, V.I. 1956. Acanthocephala in domestic and wild animals. Akademia NAUK, Moscow, 435 pp. Wiley, E. O. 1981. Phylogenetics. The theory and practice of phylogenetic systematics. Wiley-In- terscence Publications, New York, 439 pp. Department of Biological Sciences, Uni- versity of Wisconsin-Parkside, Box 2000, Kenosha, Wisconsin 53141. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 580-583 A NEW SUBSPECIES OF TURDUS SWALEST (AVES: PASSERIFORMES: MUSCICAPIDAE) FROM THE DOMINICAN REPUBLIC Gary R. Graves and Storrs L. Olson Abstract.—A new subspecies, Turdus swalesi dodae, is described from the Sierra de Neiba and Cordillera Central of the Dominican Republic. The arid Cul-de-Sac/Valle de Neiba Depression forms a barrier between J. s. swalesi of the Massif de la Selle and Sierra de Bahoruco and T. s. dodae. The La Selle Thrush, 7urdus swalesi, one of the last species of birds to be described from the island of Hispaniola (Wetmore 1927, Wetmore and Swales 1931), was long believed to be endemic to the Massif de la Selle in southeastern Haiti. Within the past fifteen years, however, populations have been found in several localities in the Do- minican Republic (Fig. 1). One of these near the Haitian border in the Sierra de Baho- ruco was not unexpected, as this is only an eastward extension of the La Selle ridge (Bond 1977). Of greater interest was the subsequent discovery of two populations in the mountains north of the arid Cul-de-Sac/ Valle de Neiba Depression. Although 7. swalesi may be locally com- mon (Bond 1928, Bond 1978), it is evi- dently represented by fewer than a dozen specimens in museums. From the small se- ries at hand, there appears to be no geo- graphic variation among specimens of the nominate form from the mountains south of the Cul-de-Sac Depression. The two specimens collected north of the depression, in the Sierra de Neiba and the Cordillera Central, however, represent a distinctive new subspecies. ies le Distribution of collected specimens of Turdus s. swalesi (circles) and T. s. dodae (triangles) on Hispaniola. Hatched area represents the Cul-de-Sac/Valle de Neiba Depression, which forms a low arid barrier between the “north” and “‘south” montane islands. VOLUME 99, NUMBER 4 581 Fig. 2. Dorsal view (from left to right) of Turdus s. dodae (MNHN 647 2; USNM 536701 8) and T. s. swalesi (USNM 264705 2; USNM 264704 6; MNHN 197 @, subadult). Turdus swalesi dodae, new subspecies Fig. 2 Holotype. —National Museum of Natural History, USNM 536701; male, from near divide between Rio Baiguate and Rio Ji- menoa, about 15 miles (ca. 24 km) NE of Constanza, La Vega Province, Dominican Republic, elevation 5900 ft (1800 m); coll. 6 Mar 1976 by Francis M. Greenwell and William M. Perrygo. Diagnosis. — Turdus swalesi dodae differs from Turdus s. swalesi in having an oliva- ceous brown black (Saccardo’s Umber to Umber-Brown; Ridgway 1912) contrasting sharply with the black of the hindneck and wings, whereas the dorsum of 7. s. swalesi is uniformly black (Fig. 2). Range. — Montane forest in the Sierra de Neiba and Cordillera Central, Dominican Republic. Measurements. —See Table 1. Specimens examined.—Turdus s. swale- si: Haiti: Massif de la Selle (USNM 264704 6, 13 Apr 1927; 264705 9, 13 Apr 1927; 582 Table 1.—Measurements (in mm) of 7. s. swalesi and 7. s. dodae. Wing and tail measured to nearest mm. Culmen measured from anterior edge of nostril. Cul- Wing Tar- (chord) Tail sus men swalesi Massif de la Selle USNM 264707 6 (type) 12 105 42.2 17.4 USNM 264704 ¢ 131 105 44.7 15.4 USNM 264705 2 125 99 43.0 16.2 Sierra de Bahoruco MNHN 197 2 (subadult) 121 94 40.5 13.2 MNHN 879 ¢ 128 100 44.0 16.4 dodae Sierra de Neiba MNHN 647 2 125 99 43:5 15.7 Cordillera Central USNM 536701 4 129 108 42.0 16.3 264707 (type) 46, 15 Apr 1927). Dominican Republic: Loma de Toro, Sierra de Baho- ruco (Museo Nacional de Historia Natural, Santo Domingo [MNHN] 197 2 subadult, 1 Sep 1972; 879 2, 15 Apr 1976). Turdus s. dodae: Dominican Republic: Sierra de Nei- ba (MNHN 647 g, 15 May 1975); NE Con- stanza (USNM holotype). Etymology. —We take pleasure in naming this subspecies for Annabelle Dod, in rec- ognition of her contributions to ornithology in the Dominican Republic. Variation in Plumage The type specimen of 7. s. dodae was erroneously reported as an “immature spec- imen” (Bond 1977:12), possibly because of its olivaceous brown black. Unfortunately, the collectors did not include gonad or skull ossification data on the specimen tag. The unspotted, sleek glossy plumage and silvery auriculars indicate that the holotype is an adult in a definitive plumage; the only evi- dence that could be construed as indicating PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Dorsal plumage of subadult (top) and adult (sexes nearly identical) T. s. swalesi. Note the exposed apteria near the rictus and on the sides of the throat in the subadult. immaturity are the pointed rectrices and faint brownish edgings of the greater wing coverts. The juvenal plumage of 7. swa/lesi is un- known. One of the specimens from the Sier- ra de Bahoruco (MNHN 197) appears to be a subadult in prebasic molt—the rictus is enlarged and fleshy, the plumage is lax and fluffy, especially on the breast and flanks, and apteria are exposed along the sides of the throat (Fig. 3). Compared with an adult female (MNHN 879) from the same local- ity, the plumage of the immature is similar in pattern, but duller and less glossy. Oth- erwise, faint spotting across the pectoral re- gion of the immature is the only age-related difference. The presence of a “black back” in immature and definitive plumages of T. s. swalesi precludes the possibility that the olivaceous back, the diagnostic character of T. s. dodae, is just a feature of immature plumage in the species. It should be noted VOLUME 99, NUMBER 4 that the distal 14 to '2 of the dorsal feathers of 7. s. dodae are olivaceous, not merely the edges or tips. The two examples of T. s. dodae differ from one another in several details: (1) the posterior margin of the black hindneck is V-shaped in the holotype but is rounded in the Sierra de Neiba specimen; (2) the oli- vaceous brown portion of the dorsum in the holotype is more restricted in distribution and contrasts more with the adjacent black plumage than in the Neiba specimen. We are unsure as to whether these differences represent intra- or inter-populational dif- ferences or sexual dichromatism. Given a larger sample, the Sierra de Neiba and Cor- dillera Central populations may be taxo- nomically separable from one another as well as from T. s. swalesi. Discussion The pattern of distribution of the two sub- species of Turdus swalesi is similar to that of numerous other Hispaniolan vertebrates, with the low arid areas of the Cul-de-Sac/ Valle de Neiba presumably forming a bar- rier to gene flow between the southern pen- insula of Haiti and the remainder of His- paniola. During periods of marine transgression, this valley formed a channel separating the two land areas into discrete islands. The two-island theory has been used to explain the origin of a variety of species- pairs of birds, reptiles and amphibians (Pre- gill and Olson 1981). In some instances, members of these pairs have spread beyond their island of origin and co-occur in some regions (e.g., Todus subulatus and T. angus- tirostris). In other cases, allopatric taxa are still restricted either to the “north” or “south” island. Variance in the degree of 583 differentiation or sympatry among former isolates doubtless reflects differences in the time of splitting of ancestral populations, in dispersal ability, and in habitat preference. Acknowledgments We thank Steve Cardiff, Annabelle Dod, Frank Greenwell, George Reynard, and Mark Robbins for information. Sixto In- chaustegui, Antonio Domingo Siri, and Manuel Campora of the Museo Nacional de Historia Natural, Santo Domingo, were in- strumental in providing specimens. Pho- tographs were provided by Victor Krantz. Literature Cited Bond, J. 1928. The distribution and habits of the birds of the Republic of Haiti.— Proceedings of the Academy of Natural Sciences of Philadel- phia 80:483-521. . 1977. Twenty-first supplement to the check- list of birds of the West Indies (1956). Academy of Natural Sciences of Philadelphia. 16 pp. 1978. Twenty-second supplement to the check-list of birds of the West Indies (1956). Academy of Natural Sciences of Philadelphia. 20 pp. Pregill, G. K., and S. L. Olson. 1981. Zoogeography of West Indian vertebrates in relation to Pleis- tocene climatic cycles.—Annual Review of Ecology and Systematics 12:75—-98. Ridgway, R. 1912. Color standards and color no- menclature. Washington, D.C.: published by the author. Wetmore, A. 1927. A thrush new to science from Haiti.— Proceedings of the Biological Society of Washington 40:55-56. , and B. H. Swales. 1931. The birds of Haiti and the Dominican Republic.— Bulletin of the United States National Museum 155:1—483. Department of Vertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 584-601 A NEW ACTINOPTERYGIAN FISH (PALEONISCIFORMES) FROM THE UPPER MISSISSIPPIAN BLUESTONE FORMATION OF WEST VIRGINIA Robert E. Weems and John F. Windolph, Jr. Abstract.—A new genus and species of deep-bodied paleonisciform fish, Tanypterichthys pridensis, is described from near the base of the Pride Shale Member of the Bluestone Formation of the Upper Mississippian Series (Na- murian A equivalent) in West Virginia. Its unusually large size (roughly 0.5 m), diamond-shaped deep body, very large pectoral fins, and the details of its scale ornamentation demonstrate that this is a new and unique paleonisciform. The type specimen of Tanypterichthys, found immediately above a basal rubble zone representing a marine transgressive event, is associated both with goniatit- ic cephalopods and with plant and tree fragments. This association indicates that it was entombed in a shallow coastal marine or marginal marine deltaic depositional environment. The functional morphology of Tanypterichthys and its burial environment suggest that this fish may have inhabited quiet waters along the coastal reaches of rivers and brackish estuaries, where kelplike colonial algae or canelike aquatic plants grew in dense stands. A large fossil fish, missing most of the skull and the caudal and dorsal fins, was discovered in an ellipsoidal carbonate con- cretion in a roadcut near Princeton, West Virginia (Fig. 1), by John Windolph during geologic investigations in the U.S. Geolog- ical Survey Upper Mississippian/Pennsyl- vanian stratotype project (Englund et al. 1979). The concretion is one of many, most- ly unfossiliferous, in a discrete zone near the base of the Pride Shale Member of the Bluestone Formation (Upper Mississippian Series) (Fig. 2). The fish was discovered by splitting the concretion, whereupon an internal view of the scale pattern and left posterior skull ele- ments (Fig. 3) was revealed. Much of the scale pattern and the conformation of the anal fin was discernible from this internal view, but nothing could be ascertained about the pelvic fins and little about the pectoral fins or the external ornamentation of the scales. For this reason, we decided that the left side of the fish should be acid prepared to try to determine something about its ex- ternal appearance. The exposed parts of the left side first were photographed by Debo- rah Dwornik of the U.S. Geological Survey, then the left half of the fish was taken to the National Museum of Natural History where Arnold Lewis and Daniel Chaney impreg- nated the exposed part of the left side of the fish in plastic. After the plastic polymerized, they removed all but a 1-inch slab of the concretion with a diamond saw. The au- thors then immersed the block in a formic acid bath to remove most of the external concretionary covering. Unfortunately, some layers were rich in pyrite and were nearly unaffected by the formic acid. In those areas, material could not be removed with- out dislodging fragments of scales. This caused a certain amount of breakage to the specimen, but most of the outer nodular coating was successfully removed by the acid treatment to reveal a complex papillate to stoutly ridged pattern on the scales and the outlines of the pectoral and pelvic fins (Fig. VOLUME 99, NUMBER 4 VIRGINIA ~:~ Ve : fo ee Y cn? KENTUCKY *. ae! 4 Be VIRGINIA, 37° 29° 25° N Fig. 1. (marked by X) of the type specimen of Tanypterichthys pridensis. B\ = Bluefield, W. Va.; Ch = Charleston, W. Va.; Wa = Washington, D.C. Map showing the locality in West Virginia 4). The right half of the fish remains in its original concretionary coat. The pectoral fin, pelvic fin, most of the anal fin, posterior skull margin, and central body region are present and well preserved, but the caudal fin, dorsal fin, upper body outline, and much of the skull are missing. The nearly perfect articulation of the fossil fish suggests that the entire animal probably was once in the rock, but that the parts of the skeleton not enclosed by the concretion were destroyed by weathering and erosion. Geologic Setting Tanypterichthys was discovered in a large ellipsoidal limestone concretion collected from a zone of concretions that lies ap- proximately 6 inches above the base of the 585 1) GLADY FORK SANDSTONE MEMBER Formation PRIDE SHALE MEMBER SERIES Bluestone Zz < a a D op) 7p) ce) = UPPER Sandstone Princeton Fig. 2. Detailed stratigraphic column of the upper Princeton Sandstone and lower Bluestone Formation at the discovery locality for Tanypterichthys pridensis. The fish-bearing concretion zone occurs near the base of the Pride Shale Member of the Bluestone Formation. The unconformably underlying Princeton Sandstone and the conformably overlying Glady Fork Sandstone Member of the Bluestone Formation were formed in deltaic environments, as shown by the presence of coal and root casts, but at least the basal Pride Shale Mem- ber formed in a marginal-marine environment of de- position as shown by the successive presence of cone- shaped nautiloid and coiled goniatitic cephalopod shells in the 6-inch interval below the fish-bearing nodule zone, and the presence of inarticulate brachiopods and wood fragments within the fish-bearing nodule zone. 586 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Internal view of the left side of the type specimen of Tanypterichthys pridensis (USNM 391949) before it was mounted in plastic and exhumed from the matrix on the other side with formic acid. These and the following photographs were taken by Deborah Dwornik (U.S. Geological Survey, Reston). Pride Shale Member (Fig. 2). The Pride Shale Member is the basal member of the Blue- stone Formation (Upper Mississippian Se- ries) and is correlative with late Chesterian age rocks of the midcontinent and with Na- murian A strata of western and central Eu- rope. The Princeton Sandstone immediately underlies the Pride Shale Member and is approximately 60 feet thick at the fossil site. Outcrops of the Princeton extend for more than 100 miles along the southeastern edge of the Appalachian Basin, forming a belt of clastic wedges and deltaic sequences that thin to the northwest. The Princeton Sand- stone, which resulted from erosion after a widespread tectonic event, consists of lenses of medium-light-gray to medium-gray poly- mictic conglomerate, coarse- to fine-grained conglomeratic subgraywacke, sandstone, siltstone, shale, underclay, and coal. Basal conglomerate and sandstone beds grade coarse to fine upward, are massive to thin- bedded, and form impressive bluffs to the southeast along the Bluestone River. The clasts are diverse in size and source, con- sisting of well-rounded to angular fragments of quartz sandstone, siltstone, shale, lime- VOLUME 99, NUMBER 4 |e bat | ROC. ry ered =e (HE Hulud 587 Fig. 4. External view of the left side of the type specimen of Tanypterichthys pridensis (USNM 391949) after being treated with formic acid. stone, ironstone, chert, and coalified plant and tree-trunk fragments. Many of the lithic fragments were derived from limestone and clastic sediments immediately underlying the Princeton, but others came from more distant sources of older Paleozoic sedimen- tary and Proterozoic to Paleozoic meta- morphic and igneous rocks that occur to the southeast. Sandstone beds, located at distal parts of clastic wedges and locally at the upper part of deltaic sequences, are light- gray, moderately quartzose, and lenticular. This description suggests the formation of beach and barrier-bar deposits by winnow- ing and reworking of sediments through high-energy coastal and long-shore process- es. The Princeton Sandstone locally in- cludes thin-bedded to nonbedded medium- gray to greenish-gray siltstone, shale, lime- stone concretions, coal, and underclay. The several rooted underclays, overlain by thin coalbeds no more than 2 inches thick, in- dicate periods of protected and stable swamp-forming conditions. Thin roof shales above the coal beds contain partially abrad- ed plant fossils (identified by W. H. Gilles- pie in Englund et al. 1985) including Sphe- nophyllum tenerrimum, Stigmaria stellata, 588 Pecopteris aspera, Archaeocalamites sp., and Sphenopteris elegans. These floral elements are characteristic of the Upper Mississip- pian of North America and the Namurian A of western and central Europe. They occur in zone 3A, above the Upper Mississippian Fryopsis zone (zone 3 of Read and Mamay 1964) and below the Neuropteris pocahon- tas zone (zone 4 of Read and Mamay 1964), which marks the base of the Pennsylvanian System. Locally, the Princeton Sandstone is over- lain by an irregularly bedded rubble zone that ranges from an inch to more than a foot in thickness. It consists of a polymictic con- glomerate in a silty sandstone matrix and includes well rounded quartz pebbles and other diverse lithic fragments as much as 1 inch in diameter. This rubble zone forms an extensive resistant ledge at the base of the Pride Shale Member, and it has been traced for more than 5 miles to the southeast of the locality where the type specimen of Tanypterichthys was found. This zone marks a disconformable contact that was formed by a widespread marine-transgressive event at the beginning of Bluestone deposition. The Pride Shale Member is the basal unit of the Bluestone Formation and is approx- imately 100 feet thick at this locality. It is dark-gray, very fissile, carbonaceous, and silty. In places, it is bioturbated and in- cludes thin, silty lenticular beds and flaser bedding. A few thin grayish-red units are present at the base, beneath the more wide- spread zone of sparsely fossiliferous lime- stone concretions in which the fossil fish was found. Exposed in roadcuts northwest of the fossil site are several large slump, scower, and channel features that are over- lain and underlain by horizontal bedding. These unrooted features suggest the action of strong tidal influences and submarine currents. Associated with these beds are ma- rine and brackish invertebrate fossil assem- blages, which were collected by T. W. Henry of the U.S. Geological Survey and which are consistent with a shallow bayfill or la- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON goonal depositional environment for the Pride Shale Member. Above the Pride, coarse clastic rocks characterize the suc- ceeding Glady Fork Sandstone Member of the Bluestone Formation. These clastic rocks are similar to those previously described from the underlying Princeton Formation. Taxonomic Description The fish described here (see Figs. 5, 6, 7, 8) would seem to be readily assigned to the Paleonisciformes in view of the persistent development of branchiostegal rays in its cheek region and the unreduced number of rays in its fins. The order Paleonisciformes traditionally has been divided into two sub- orders (Moy-Thomas 1971), one (Paleonis- coidei) for normally shaped, fusiform types and the other (Platysomoidei) for deep-bod- ied forms reminiscent of our fish. Up to four families (Amphicentridae, Platysomidae, Bobasatranidae, and Dorypteridae) have been recognized, though variations on this theme have been proposed. The Amphi- centridae and Platysomidae were combined into a single family by Berg et al. (1964). Fowler (1958) suggested that the term Uropterygidae should be favored over the term Platysomidae because the latter name is pre-empted, but this practice has not been followed by any subsequent worker. Berg (1940b) suggested placing the Bobasatran- idae into a separate order, the Bobasatran- iformes. This practice generally has been followed. In recent years, this relatively simple tax- onomy has collapsed. Campbell and Phuoc (1983) have pointed out that the skull of the type species of Platysomus, the Permian P. gibbosus, shows derived characteristics which indicate that it, and by definition the genus Platysomus, belong in the Bobasa- traniformes. But it is unlikely that Missis- sippian and Pennsylvanian species which have been classified in Platysomus (or any of the other genera formerly classified in the Platysomidae and Platysomoidei) share these derived traits. Therefore, although VOLUME 99, NUMBER 4 589 Centimeters 1 eo ¥ sa we a, : ; Fig. 5. (Top) Detail of the left pectoral fin of the type specimen of Tanypterichthys pridensis (USNM 391949). Note that the dorsa! edge of the fin is lined by an enlarged row of scales, which are shown in more detail below. (Bottom) Detail of dorsal scale row on the pectoral fin of the left side of the type specimen of Tanypterichthys pridensis (USNM 391949). Scales are near distal end of fin. 590 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. (Top) Detail of anterior scale pattern on the anal fin of the type specimen of Tanypterichthys pridensis (USNM 391949), seen in internal view on the left side before acid preparation. (Lower left) Detail of internal flank scale pattern on the left side of the type specimen of Tanypterichthys pridensis (USNM 391949) before acid preparation. (Lower right) Detail of the left side of the type specimen of Tanypterichthys pridensis (USNM 391949) showing the vermiform sculpture on the posterior cranial elements, and the pustulose pattern on the anterior flank scales. Fig. 7. (Top) Detail of internal ventral skull element pattern on the left side of the type specimen of Tany- pterichthys pridensis (USNM 391949) before acid preparation. (Bottom) Detail of lower anterior flank region of the left side of the type specimen of Tanypterichthys pridensis (USNM 391949), showing the ventral pectoral fin rays (upper right diagonal) and the ridgelike texture on the ventral flank scales beneath the pectoral fin (lower left diagonal). 592 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. (Top) Detail of the ornamentation pattern on a single antero-medial flank scale from the left side of the type specimen of Tanypterichthys pridensis (USNM 391949). (Bottom) Detail of papillate ornamentation on a single postero-medial flank scale from the left side of the type specimen of Tanypterichthys pridensis (USNM 391949), VOLUME 99, NUMBER 4 these older forms can be properly retained within the Paleonisciformes, the type genus for the family Platysomidae and suborder Platysomoidei now is removed so that the taxonomy of the remaining deep-bodied pa- leonisciforms is near chaos. It is beyond the scope of this paper to sort through the var- ious phylogenetic pathways in this complex of genera and to classify them correctly, so for now our new form is simply placed in Paleonisciformes without familial assign- ment. Future work could even demonstrate that it deserves its own familial assignment. Because a meaningful classification for deep-bodied paleonisciform fishes has not been worked out, the genera which have been described from rocks of Early Missis- sippian to Middle Triassic age simply are considered in seriatim. These are Mesolepis, Paramesolepis, Platysomus (in part), Schaef- ferichthys, Soetendalichthys, Wardichthys, Cheirodopsis, Cheirodus (=Amphicentrum), Eurynotus, Paraeurynotus, Proteurynotus, Adroichthys, Globulodus (=Lekanichthys), Eurynotoides, and Caruichthys. Most of these fish are extremely deep bod- ied and narrow, but Mesolepis (Y oung 1866, Traquair 1879, Ward 1890, Traquair 1907, Pruvost 1919, Van der Heide 1943) and Eurynotus (Agassiz 1833-43, Koninck 1878, Traquair 1879) evolved into only slightly deep-bodied forms and even Proteurynotus (Moy-Thomas and Dyne 1938) is not nearly so deep bodied as Tanypterichthys. Adro- ichthys (Gardiner 1969), Paramesolepis (Traquair 1881, Moy-Thomas and Dyne 1938), Cheirodopsis (Traquair 1881, Moy- Thomas and Dyne 1938), and Wardichthys (Traquair 1874, 1879, 1907) all have a short- based and posteriorly located anal fin; this contrasts sharply with the elongate, antero- ventrally expanded anal fin in Tanypterich- thys. Soetendalichthys (Gardiner 1969) is somewhat similar to Tanypterichthys in body shape and possibly in anal fin elon- gation, but it contrasts with our form in that its flank scales are not nearly so vertically elongated. Additionally, it has a much more 593 tubercular scale ornamentation, and the pin and socket connections between scales in each column are at the center of each scale rather than along the front or back margin. Cheirodus (Newberry and Worthen 1870; Hancock and Atthey 1872; Traquair 1879; Ward 1890; Pruvost 1919, 1930; White 1937; Dyne 1939; Van der Heide 1943; Bar- dack 1979) is markedly distinct from Tany- pterichthys in its small size (5-15 cm), tu- bercular scale ornamentation, the extreme constriction between the tail and body, and the presence in at least some forms of body ““horns”’ drawn up before the dorsal and anal fins like keels. Platysomus has been used as a generic name for more species of deep-bodied pa- leonisciforms than any other (Agassiz 1833-— 43; Eichwald 1857; Hancock and Atthey 1872; Ward 1890; Cope 1891; Pruvost 1930; Moy-Thomas and Dyne 1938; Van der Heide 1943; Wilson 1950; Zidek 1972; Simpson 1979; Schaumberg 1976, 1980). As noted above, the type species of Platy- somus now is considered to be bobasatran- iform, so at least part of the Permian species assigned to that genus belong in that order. Mississippian and Pennsylvanian species assigned to Platysomus are grossly similar to Tanypterichthys, but they differ consis- tently from our fish in their possession of a finely vermiform scale-ornamentation pat- tern (aligned in parallel vertical rows), pos- session of a more constricted region be- tween the body and the tail, much smaller pectoral fins, and pelvic fins which are never drawn forward far enough to lie beneath the pectorals. Schaefferichthys (Dalquest 1966) is a poorly known form from the Permian of Texas. It is generally platysomid in form and has a finely vermiform scale-ornamen- tation pattern aligned in parallel vertical rows as in Platysomus. It may be possible that this species is a primitive bobasatran- iform, but the scale-ornamentation pattern debars it from any close relationship with Tanypterichthys. Paraeurynotus (Obruchev 1962) is poorly 594 known, but the scale proportions (only two times higher than wide) and ornamentation (nearly smooth) are markedly different from those of Tanypterichthys. Additionally, it is remote in time from our form (Early Permi- an). Eurynotoides (Berg 1940a) has very near- ly a normal fish shape and scale structure, and it is also remote in time (Late Permian) from Tanypterichthys; obviously it is not closely related to our specimen. Caruichthys (Broom 1913, Lehman 1966) of the Early Triassic is remote in time from Tanyperichthys, a smaller form (about 20 cm long), and covered with scales that are ornamented by irregular prominent trans- verse ridges. Globulodus (Munster 1842, King 1850, Young 1866, Traquair 1879, Woodward 1891, Brough 1934, Westoll 1941, Berg et al. 1964, Schaumberg 1980) shows some obvious similarities to Tanypterichthys. The scales are coarsely striated and their pin and socket arrangement is located along the edge of the scale, the size is large (a maximum of 40 cm), and the constriction between the body and the tail is persistently thick for a deep-bodied paleonisciform fish. Yet the anal fin is not nearly so elongated and the adjacent postero-ventral flank scale col- umns are not turned downward and forward as they are in Janypterichthys. In these characteristics, Globulodus is persistently the more primitive form, even though it is much younger in age (Late Permian). This tem- poral progression in specialization from de- rived to primitive seems to debar any direct lineage between these two forms. Probably the similarity in tail structure is a shared primitive trait retained in both forms and should not be considered to be of phylo- genetic signficance; the position of the pin and socket arrangement also is shared by many other forms in this family. Large size (40 cm) and a relatively thick tail constric- tion also are characteristics of Adroichthys, which otherwise is not especially close in its morphology to either Globulodus or Tany- pterichthys. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Some other forms formerly placed among the deep-bodied paleonisciforms (“Eury- notus’’ uspallatensis (Rusconi 1946a), ‘**Platysomus’’ pehuenchensis (Rusconi 1946b) and “Platysomus”’ cajonensis (Rus- coni 1948), from the Triassic and Jurassic of Argentina) probably do not belong there. What little that is known of them suggests that their scales are not so elongate as those of Tanypterichthys and not similarly orna- mented. The unnamed platysomid from the Triassic of Australia mentioned by Banks (1978) is, if correctly identified, probably the youngest representative of this family. Two other recognized families, also deep bodied and frequently considered to be pa- leonisciforms, are the Dorypteridae and the Dorsolepidae. The monogeneric Dorypter- idae (Westoll 1941, Schaumberg 1980) is known only from Late Permian marine beds. This family is characterized by a very un- usual body shape and the loss of nearly all trace of a scale coat, characteristics that are not remotely similar to those found in our fish. The monogeneric Dorsolepidae, known only from the Early Triassic, is represented only by the form Dorsolepis (Jorg 1969a, b; Gall et al. 1974). It has been associated with the platysomids, but only its narrow and deep body form strongly suggests any affin- ity. Its scales are greatly reduced and it is very small (5 cm), so it has no close simi- larity to Tanypterichthys. The marine-going Bobasatraniformes, which includes Bobasatrania canadensis (Schaeffer and Mangus 1976), B. groenlan- dica (Stensio 1932, Nielsen 1952, Lehman 1957), B. mahavavica (White 1932, Leh- man 1957), B. nathorsti (Stensio 1921), Ebenaqua ritchiei (Campbell and Phuoc 1983), Ecrinesomus dixoni (Woodward 1910, Lehman 1957), Lambeichthys cana- densis (Lambe 1914, Russell 1951, Lehman 1966), ““Platysomus”’’ brewsteri (Warren 1936, Lehman 1966), Tompoichthys abra- movi (Berg et al. 1964, Lehman 1966), and Sinoplatysomus meishanensis (Wei 1980), are all forms that are grossly similar in ap- pearance to Tanypterichthys. However, their VOLUME 99, NUMBER 4 scale ornamentation, where described, is finely vermiform in vertically parallel ridges, and the constriction between the body and tail is extreme, being only about four scale rows thick. Sinoplatysomus and Ebenaqua are from the Late Permian, and the other members of this family are from the Early Triassic. The pectoral fin of Tanypterich- thys is similar to that of Bobasatrania, but otherwise shows no special similarity to any member of this family except in its rela- tively large size. Moreover, the preserved portion of the cheek region in Tanypter- ichthys shows that it contains numerous branchiostegal rays. In contrast, bobasa- traniforms have the cheek region highly modified from the primitive paleonisciform condition and the branchiostegal rays have been largely or wholly removed from lateral view (Schaeffer and Mangus 1976). Its gross similarities to the bobasatraniform fishes are considered here to be the result of conver- gence rather than any intimate relationship. Thus Tanypterichthys, although it falls within the definition of the Paleonisci- formes and bears some resemblance to the Bobasatraniformes, cannot be shown to have a clear and close affinity to any other genus so far described within either of those two orders. Tanypterichthys therefore warrants recognition as a new and separate genus that can be defined as follows: Order Paleonisciformes Tanypterichthys, new genus Diagnosis. — Tanypterichthys differs from all other paleonisciforms by the following combination of characters. Scales vertically elongate (as much as 4 times longer than wide), ornamented with a complex pattern consisting of globular papillae and random- ly oriented, short, thick ridges reminiscent of the scale ornamentation in Wardichthys; scales arrayed in about 50 columns (from behind the head to the constricted portion of the tail) and in rows of 15 (near the tail) to no more than 26 (near the middle of the body). Posterior skull elements ornamented with a fine, vermiform, vertically aligned 595 pattern of parallel ridges very different from the body scale pattern. Pectoral fins greatly elongated and expanded as in Eurynotus, Caruichthys, and Bobasatrania, more than half as long as the body. Pelvic fins long, thin, subcylindrical, and inserting far for- ward on the body beneath the distal part of the pectoral fins, length about one-third that of the body. Anal fin greatly elongated, bor- dering along the entire length of the back half of the body and turning the adjacent postero-ventral scale columns forward in a manner similar to that seen in Platysomus striatus. Constriction between the body and the tail pronounced, but less so than in other deep-bodied members of this order except Adroichthys and Globulodus. Size very large, as much as 0.5 m. Type species.—Tanypterichthys priden- sis, new species. Tanypterichthys pridensis, new species Figs. 3-11 Type specimen. —USNM 391949, a large calcareous concretion containing the bulk of the body, the back edge of the skull, pec- toral fins, pelvic fins, and the anal fin. Type locality.—At the northwest end of the Camp Creek Interchange along the West Virginia Turnpike, Mercer County, West Virginia, at latitude 37°29'25”N and lon- gitude 81°06'25’W. Horizon. — About 6 inches above a rubble zone at the base of the Pride Shale Member of the Bluestone Formation (Upper Missis- sippian Series). Collector. —John F. Windolph, Jr., 20 Nov 1982. Diagnosis. — As for the genus. Etymology. —The pectoral fin is unusual in appearance, being very large and elon- gate, and it is this striking characteristic to which the generic name alludes (tany- = stretched out, pteryx = fin or wing, ich- thys = fish). The specific name reflects the occurrence of the specimen in the Pride Shale Member of the Bluestone Formation. Discussion. —Electron photomicrographs were made of a fragment of one bony scale 596 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 9. Electron photomicrographs of details ofa single scale from the postero-medial region of the left flank of the type specimen of Tanypterichthys pridensis (USNM 391949). Upper left, scale surface showing ridges (running from upper left to lower right) and nutrient foramina (top center); upper right, detail of the nutrient foramina in the top center of the upper left picture; lower left, more detailed view showing the cellular structure of the scale; lower right, more detailed view showing the central region of the lower left picture. Photos by Richard Larson (U.S. Geological Survey, Reston). VOLUME 99, NUMBER 4 a Wah - es Ly, UM aT iy Ai ‘ Da ane Mi i JME NA ! Zi ay) He HN My ae 597 o va 2a S aa SS WN) a Hr Pon Fig. 10. Sketch showing the scale pattern and anal fin shape as seen from the internal view of the left side - of the type specimen of Tanypterichthys pridensis (USNM 391949) before acid preparation. by Richard Larson (U.S. Geological Survey, Reston), and details of the well preserved bone structure and nutrient channels are shown at four different magnifications (Fig. 9). SEM spectral analysis of the fragment showed calcium and phosphorus as major elements, and silicon, aluminum, iron, and sulfur as minor elements. No trace of the appendicular skeleton was observed. Possibly the concretion split through the scales on one side of the body, so that the remains of a bony skeleton were not revealed. If so, there is no trace of the shapes of such bones revealed by warpage of the visible internal scale surfaces. There- fore, it seems more likely that the concretion split through the middle of the fish, and the appendicular skeleton had left no trace of its former presence. The proportions of the bones at the back of the skull are normal for a platysomid and require no special dis- cussion, though their parallel vermiform pattern (reminiscent of Platysomus) is strik- ingly different from the scale-ornament pat- tern. Parts of the front, top, and back of the body are missing, but general proportions can be estimated from the available re- mains. Deep-bodied paleonisciform fishes are known to have 15 (Paramesolepis tuber- culata) to 26 (Eurynotus, Soetendalichthys) rows of scales present in each vertical col- umn near the deepest part of the body. Be- cause about 22 scale rows are preserved in the type specimen of Tanypterichthys, the top of the body almost certainly did not extend any higher than 4 more short scale- 598 \\ vi vii Fig. 11. aN AA Ue PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0 SOQ ANS WN SS \) Sketch showing scale pattern, pectoral fin, and pelvic fin as seen from the external view of the left side of the type specimen of Tanypterichthys pridensis (USNM 391949) after acid preparation. Shape of the anal fin was derived from Fig. 10; the general body form is assumed to be typically platysomid. Inset shows the details of the construction and ornamentation of a single flank scale (shaded black on the fish body) and its relationships to surrounding scales. heights above the level of the preserved up- per margin. Likewise, when a large anal fin is found in other fishes in this family, it is matched by an equally large or larger dorsal fin (for example, Platysomus superbus or Ebenaqua ritchiei). The gross shape of the skull and the shape of the tail are not greatly variable in known members of this group; therefore, the rough outlines of the body reasonably can be surmised in addition to the observable detailed patterns of the pre- served parts (Figs. 10, 11). Functional Anatomy The outline of the flank scales is markedly different in internal and external aspect (see Figs. 3, 4). This is because the scales are hinged along their dorsal and ventral bor- ders in such a manner that the internal an- terior dorsal border forms a pinlike slip. This slip inserts beneath the internal anterior ventral border of the next scale above it where it is embayed from the internal side to form a socket (see Fig. 11). This general kind of structure is probably normal among deep-bodied paleonisciform fish, though in Eurynotus (Traquair 1879) and Soetendal- ichthys (Gardiner 1969) the pins and sockets are in the center of the scales, and in Ad- roichthys (Gardiner 1969) they are slightly off-center. In Globulodus (Agassiz 1833-43, King 1850), Platysomus (Traquair 1879), Cheirodus (Traquair 1879), and Tanypter- VOLUME 99, NUMBER 4 ichthys, the pin is formed by an upper corner of the scale. The function of this arrange- ment seems to have been to strenghten the ganoid scale coating and thus to form a chain-mail effect through a combination of antero-posterior overlapping and vertical interlocking of the scales. Such an inter- locking arrangement would be mobile only around a vertical axis, and the body move- ment of these fishes would be restricted to lateral flexure. Such restriction in body mo- tion is probably correlated with the very deep body form of the animal. The fact that the body armor did not evolve to a greater rigidity suggests that these fishes did need to preserve their ability to move in a lat- erally undulatory, normal piscine fashion. The dorsal fin of Tanypterichthys is un- known, but in better known members of the family that also have expanded anal fins (for example, Platysomus superbus (Moy- Thomas and Dyne 1938) and Ebenaqua rit- chiei (Campbell and Phuoc 1983)) great ex- pansion and elongation of the anal fin is matched by an equal or greater develop- ment of the dorsal fin. Such well developed sets of anal and dorsal fins in modern fishes usually function to propel their bearer at slow speeds by undulations of these two fins without marked undulation of the body as a whole. The well-developed anal fin, pres- ent in Tanypterichthys and in many of the other deep-bodied paleonisciforms and bobasatraniforms, suggests that a similar mode of propulsion probably was em- ployed. The pectoral and pelvic fins are strongly modified from the generalized actinopteryg- ian condition, which suggests that they were specialized in function. The large pectorals were braced along their dorsal border by an enlarged row of scales (see Fig. 5), which probably afforded them exceptional strength for an actinopterygian. These large fins could have poled the fish forward while it cruised near the bottom or functioned as oars for maneuvering the body fore, aft, and in tight circles. The pelvic fins are long but very 599 thin. They could have functioned in murky water as feelers to keep the fish slightly above the bottom or as claspers during mating as similarly shaped pelvic fins do in many modern actinopterygians. Literature Cited Agassiz, L. 1833-43. Recherches sur les poissons fos- siles. Vol. 2, Part 1, pp. 1-336. Banks, M.R. 1978. Correlation chart for the Triassic system of Australia.— Australia Bureau of Min- eral Resources, Geology, and Geophysics Bul- letin 156C:1-39. Bardack, D. 1979. Fishes of the Mazon Creek Fauna. InM. H. Nitecki, ed., Mazon Creek fossils. New York, Academic Press. Pp. 501-528. Berg, L.S. 1940a. Two new palaeoniscid genera, Eu- rynotoides and Amblypterina, from the Upper Permian of Kargala (Basin of the Ural River). — Akademiia Nauk SSSR, Izvestiia, Seriia Biol- ogicheskaia 3:414—419. 1940b. Classification of fishes, both recent and fossil.— Travaux de I’Institut d’Zoologique de l’Academie des Sciences de l’USSR (Trudy Zoologicheskogo Instituta Akademii Nauk) 5: 85-517. — A.A. Kazantseva, and D. V. Obruchev. 1964. (Superorder Palaeonisci).—Osnovy Paleonto- logii (Agnathes, Pisces) 11:336-370. Broom, R. 1913. On some fishes from the lower and middle Karroo beds.— Annals of the South Af- rican Museum 12:1-5. Brough, J. 1934. On Lekanichthys howsei—a new do- rypterid fish from the Permian.—Annals and Magazine of Natural History (10) 14(32):353- 366. Campbell, K. S. W., and L. D. Phuoc. 1983. A Late Permian Actinopterygian fish from Australia. — Palaeontology 6(1):33-70. Cope, E. D. 1891. On the characters of some Paleo- zoic fishes.— Proceedings of the U.S. National Museum 14:447-463. Dalquest, W. W. 1966. An unusual paleonisciform fish from the Permian of Texas. — Journal of Pa- leontology 40(3):759-762. Dyne, B. 1939. The skull of Amphicentrum granu- losum. — Proceedings of the Zoological Society of London 109(B):195—210. Eichwald, E. von. 1857. Beitrag zur geographischen Verbreitung der fossilen Theire Russlands. Alte Periode.— Bulletin de la Société Imperiale des Naturalistes de Moscou 30(2):305-354. Englund, K. J., H. H. Arndt, and T. W. Henry [eds.]. 1979. Proposed Pennsylvanian System strato- tpye, Virginia and West Virginia.—Field trip Number 1, Ninth International Congress of Car- 600 boniferous Stratigraphy and Geology, American Geological Institute Selected Guidebook Series 1:1-138. ——., W. H. Gillespie, C. B. Cecil, J. F. Windolph, Jr., and T. J. Crawford. 1985. Characteristics of the Mississippian-Pennsylvanian Boundary and associated coal-bearing rocks in the South- ern Appalachians.—Geological Society of America 1985 Annual Meeting, Field Trip Guidebook. Fowler, H. W. 1958. Some new taxonomic names of fishlike vertebrates.—Notula Naturae of the Academy of Natural Sciences of Philadelphia 310:1-16. Gall, J. C., L. Grauvogel, and J. P. Lehman. 1974. Faune du Buntsandstein: V. Les poissons fos- siles de la collection Grauvogel-Gall.— Annales de Paleontologie (Vertébrés) 60(2):129-147. Gardiner, B. G. 1969. New palaeoniscoid fish from the Witteberg series of South Africa.— Zoolog- ical Journal of the Linnean Society 48(4):423- 452. Hancock, A., and T. Atthey. 1872. Descriptive notes on a nearly entire specimen of Pleurodus Ran- kinii, on two new species of Platysomus and a new Amphicentrum, with remarks on a few oth- er fish remains found in the Coal Measures at Newsham.—Annals and Magazine of Natural History (4) 9(52):249-262. Jorg, E. 1969a. Eine Fischfauna aus dem Oberen Buntsandstein (Unter-Trias) von Karlsruhe- Durlach (Nordbaden).— Beitrage zur Naturkun- dlichen Forschung Siidwestdeutschland 28(2): 87-102. . 1969b. Fischfunde im Oberen Buntsandstein (Untertrais) von Karlsruhe-Durlach. —Zeit- schrift der Deutschen Geologischen Gesell- schaft 121:105-110. King, W. 1850. A monograph of the Permian fossils of England.— Memoirs of the Palaeontograph- ical Society 1:1—253. Koninck, L. G. de. 1878. Faune du calcaire Carboni- fére de la Belgique; (1) Poissons et genre nau- tile. —Annales du Musée Royal d’Histoire Na- turelle de Belgique 2:1—152. Lambe, L. M. 1914. Description of a new species of Platysomus from the neighborhood of Banff, Al- berta.—Transactions of the Royal Society of Canada 8(4):17—-23. Lehman, J.-P. 1957. Complements a l’étude des gen- res Ecrinesomus et Bobasatrania de \’Eotrias de Madagascar. — Annales de Paleontologie 42:65-— 94. 1966. Actinoptergyii, im Jean Piveteau, ed., Traite de paleontologie, Book 4, volume 3. Paris, Masson et Cie. 242 pp. Moy-Thomas, J. A. [revised by R. S. Milesj. TO ale PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Palaeozoic fishes. Philadelphia, W. B. Saunders Company. 259 pp. , and Bradley Dyne. 1938. The actinoptery- gian fishes from the Lower Carboniferous of Glencartholm, Eskdale, Dumfriesshire. — Transactions of the Royal Society of Edinburgh 59:437-480. Munster, G. von. 1842. Beschreibung einiger merk- wurdigen Fische den Kupferschiefern von Ri- chelsdorf und Eisleben. — Beitrage zur Petrefak- tenkunde 5:35-50. Newberry, J. S., and A. H. Worthen. 1870. Descrip- tions of fossil vertebrates. —Illinois Geological Survey 4(2,1):345-374. Nielsen, E. 1952. A preliminary note on Bobasa- trania groenlandica.—Dansk Geologisk Fore- ning, Meddelelser 12(2):197—204. Obruchev, D. V. 1962. (Permian fishes): (Paleozoic biostratigraphy of the Saian-Altai mountain re- gion, volume 3).—Trudy SNIGGIMS 21:440- 442. Pettigrew, T. H. 1984. Fossils from the Marl Slate of northeast England.— Fossils Quarterly 2(4):18- 23. Pruvost, P. 1919. Introduction a l'étude du Terrain Houiller du nord et du Pas-de-Calais: La fauna continentale du Terrain Houiller du Nord de la France; memoires pour servir a l’explication de la Carte Geologique Détaillee de la France. Paris, Ministere de Travaux Publics. 584 pp. 1930. La faune continentale du Terrain Houiller de la Belgique.—Memoires du Musée Royal d’Histoire Naturelle de Belgique 44:103- Die Read, C. B., and S. H. Mamay. 1964. Upper Paleo- zoic floral zones and floral provinces of the United States.— U.S. Geological Survey Profes- sional Paper 454:K1—-K35. Rusconi, C. 1946a. Nuevos peces Triasicos de Us- pallata.—Anales Sociedad de la Cientifica Ar- gentina 141:185-—190. . 1946b. Un pez Jurasico de Mendoza. —Divi- sao de Geologiae Mineralogia de Brasil, Notas Preliminares e Estudos 32:3-8. 1948. Nuevo plesiosaurio, pez y langostas del mar, Jurasico de Mendoza.—Revista del Museo de Historia Natura de Mendoza 2(1/2): 3-12. Russell, L. S. 1951. Bobasatrania? canadensis (Lambe), a giant chondrostean fish from the Rocky Mountains.— National Museum of Can- ada Bulletin 123:218-224. Schaeffer, B., and M. Mangus. 1976. An Early Trias- sic fish assemblage from British Columbia.— Bulletin of the American Museum of Natural History 156(5):517—563. Schaumberg, G. 1976. Uber ein gut erhaltenes der- VOLUME 99, NUMBER 4 males Schadeldach von Platysomus striatus Agassiz (Palaeonisciformes, Actinopterygii, Os- teichthyes) aus dem Kupferschiefer von Ri- chelsdorf (Perm, Hessen).—Geologisches Jahr- buch Hessen 104:39—42. . 1980. Der Richelsdorfer Kupferschiefer und seine Fossilien, III. Die tierischen Fossilien des Kupferschiefers; 2, Vertebraten.—Der Auf- schluss 28(8/9):297-357. Simpson, L. C. 1979. Upper Gearyan and lower Le- Onardian terrestrial vertebrate faunas of Okla- homa.— Oklahoma Geology Notes 39(1):3-19. Stensio, E.A. 1921. Triassic fishes from Spitzbergen, Part 1. Vienna, Adolf Holzhausen. 307 pp. 1932. Triassic fishes from East Greenland, collected by the Danish expeditions in 1929- 1931.— Meddelelser om Gronland 83(3):1—305. Traquair, R.H. 1874. On some fossil fishes from the neighbourhood of Edinburgh.— Annals and Magazine of Natural History (4) 15(34):258- 268. 1879. On the structure and affinities of the Platysomidae.— Transactions of the Royal So- ciety of Edinburgh 29:343-391. . 1881. Report on the fossil fishes collected by the Geological Survey of Scotland in Eskdale and Liddesdale; Part 1. Ganoidei.—Transac- tions of the Royal Society of Edinburgh 30:15- 71. . 1907. Report on fossil fishes collected by the Geological Survey from the shales exposed on the shore near Gullane, East Lothian. —Trans- actions of the Royal Society of Edinburgh 46: 103-117. Van der Heide, S. 1943. La faune ichthyologique du Carbonifére superieur des Pays-bas.—Me- dedeelingen Netherlands Geologische Stichting (Series C-IV-3) 2:1-66. Ward, J. 1890. The geological features of the North Staffordshire Coal-fields, their organic remains, their range and distribution, with a catalogue of 601 the fossils of the Carboniferous system of North Staffordshire.— Transactions of the North Staf- fordshire Institute of Mining and Mechanical Engineers 10:1—189. Warren, P. S. 1936. Two new fossil fish from the Canadian Rockies.— Transactions of the Royal Society of Canada (3) 30(4):55—58. Wei, F. 1980. The discovery of a fossil platysomid in the Changxing Limestone of Zhejiang Prov- ince.—Ku Sheng Wu Hsueh Pao [Acta Geolo- gica Sinica] 16(2):293-296. Westoll, T. S. 1941. The Permian fishes Dorypterus and Lekanichthys. — Proceedings of the Zoolog- ical Society of London 111(B):39-58. White, E.I. 1932. Ona new Triassic fish from north- east Madagascar.—Annals and Magazine of Natural History (10) 10:80-83. . 1937. The fishes of the “Crangopsis Bed” at Ardross, Fifeshire.—Geological Magazine 74: 411-428. Wilson, J. A. 1950. A platysomid from the Double Mountain group of Texas.—Journal of Paleon- tology 24(3):386-389. Woodward, A. S. 1891. Catalogue of the fossil fishes in the British Museum (Natural History 2:1- 567. 1910. On some Permocarboniferous fishes from Madagascar.—Annals and Magazine of Natural History (8) 5:1-6. Young, J. 1866. On the affinities of Platysomus and allied genera.— Quarterly Journal of the Geo- logical Society 22:301-317. Zidek, J. 1972. Oklahoma paleoichthyology: Part 1. A review and commentary.— Oklahoma Geol- ogy Notes 32(6):171-187. (REW) Mail Stop 928, U.S. Geological Survey, Reston, Virginia 22092; (JFW) Mail Stop 956, U.S. Geological Survey, Reston, Virginia 22092. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 602-603 LARVAE OF XJPHOPENAEUS KROYERI (HELLER, 1862) (CRUSTACEA: DECAPODA: PENAEIDAE) FROM OFFSHORE WATERS OF VIRGINIA, U.S.A. Robert C. Maris Abstract. — A range extension is provided for the shrimp Xiphopenaeus kroy- eri (Heller, 1862). Larvae of X. kroyeri were collected from offshore waters of Virginia, U.S.A. near the Chesapeake Light Tower (36°54'N, 75°43’W). Zoeal stages I-III, V—VII, and IX were obtained at depths ranging from neuston to epibenthic, during the day and night, with a maximum concentration of 6.8 per cubic meter. This report extends the known range of X. kroyeri about 190 km northward from Cape Hatteras, North Carolina. Zooplankton collections taken off the coast of Virginia were found to contain lar- val specimens of Xiphopenaeus kroyeri (Heller, 1862). This species was previously known to occur from capes Hatteras and Lookout, North Carolina, through the Gulf of Mexico and Caribbean Sea to Brazil (Wil- liams 1984). A station near the Chesapeake Light Tow- er (36°54'N, 75°43'W) was occupied for a continuous 72-hour period, 13-16 August 1985, (Maris 1986). Quantitative plankton samples were collected at three-hour inter- vals from the following depths: neuston (0.10-0.15 m), 1 m, 3 m, 6 m, epibenthic (12.8 m). A total of 125 samples were collected, of which 12 (9.6%) contained specimens of_X. kroyeri. Zoeal stages I, I, III, V, VI, VII, and IX were obtained and identified using descriptions provided by Kurata (1970). The larvae were found at all depths sampled, during both day and night collections, with a maximum concentration of 6.8 per cubic meter. A summary of the entire collection of larval X. kroyeri can be found in Table 1. The occurrence of larval X. kroyeri might indicate breeding populations in Virginia waters, but possible transport from the south cannot be eliminated. However, the pres- ence of early zoeae casts doubt on long-dis- Table 1.—Offshore Virginia collections of larvae of Xiphopenaeus kroyeri (Heller, 1862). Date Density (1985) Time (EDT) Depth (m) Stage (no./m?) 13 Aug 1500 6 Il 0.7 14 Aug 0000 1 Il DED; 15 Aug 0000 0 IX 0.8 15 Aug 0000 6 VI 2.0 15 Aug 0300 6 Il 1.1 15 Aug 0300 6 VII et 15 Aug 0900 6 I 1.1 15 Aug 1500 6 II 1.0 15 Aug 1500 6 Vv 1.0 16 Aug 0000 0 V 1.8 16 Aug 0300 13 I 0.7 16 Aug 0600 3 II 6.8 16 Aug 0900 6 I 1.1 16 Aug 0900 6 Vil eI 16 Aug 1200 6 II Mp) 16 Aug 1200 6 Il Dep) tance transport as the sole dispersal mech- anism. Also, the abundance and variety of later zoeal stages seems to imply that at least limited metamorphosis is occurring off Vir- ginia. Thus, this report extends the known range of X. kroyeri about 190 km northward from Cape Hatteras, North Carolina. Acknowledgments I am grateful for the reviews and critical comments by Drs. Anthony J. Provenzano, VOLUME 99, NUMBER 4 Jr. and John R. McConaugha. The crew of the R/V Linwood Holton, (Captains Robert N. Bray and James D. Padgett and Engineer Nelson Griffin) are given much appreciation for making the field collections possible. I thank the following volunteers for helping with preparation, transportation, and actual sampling: Emily Deaver, Dean Devereaux, Lore Hantske, Rebecca Lovingood, Yvonne Maris, Cathy McConaugha, Les Parker, Jean Pletl, Jim Pletl, Eric Stern, Lyle Varnell, and Ann Wry. I give special thanks to my wife, Yvonne Maris, for typing the manuscript, understanding, and encouragement. This work is a result of research supported by the Office of Sea Grant, NOAA, Department of Commerce, Grant NA81AA-D-0025. Literature Cited Kurata, H. 1970. Studies on the life histories of deca- pod Crustacea of Georgia: Part III. Larvae of 603 decapod Crustacea of Georgia. Final Report, University of Georgia Marine Institute, Sapelo Island, Georgia, 274 pp., 105 pls. Maris, R. C. 1986. Patterns of diurnal vertical dis- tribution and dispersal-recruitment mecha- nisms of decapod crustacean larvae and post- larvae in the Chesapeake Bay, Virginia and ad- jacent offshore waters. Ph.D. dissertation, Old Dominion University, Norfolk, Virginia, 222 pp. Williams, A. B. 1984. Shrimps, lobsters, and crabs of the Atlantic Coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C., 550 pp. Department of Oceanography, Old Do- minion University, Norfolk, Virginia 23508-8512. Present address: Box 204, Cheney University, Cheney, Pennsylvania 19319. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 604-611 TWO NEW CRABS, PARAPAGURISTES TUBERCULATUS AND PALAEOXANTHO LIBERTIENSIS, FROM THE PRAIRIE BLUFF FORMATION (MIDDLE MAASTRICHTIAN), UNION COUNTY, MISSISSIPPI, U.S.A. Gale A. Bishop Abstract. —Two new genera and species of fossil crabs, Parapaguristes tuber- culatus and Palaeoxantho libertiensis, occur in the Prairie Bluff Formation in Union County, Mississippi. Associated with these taxa are Prehepatus harrisi Bishop, 1985, Raninella? sp., Protocallianassa sp., an indeterminate claw of a portunid, and claw fragments of a large lobster. The genus Parapaguristes is erected to include P. tuberculatus n. sp. and P. whitteni (Bishop, 1983). The new genus Palaeoxantho is morphologically similar to the xanthid genera Xan- thosia, Xantho, and Syphax. During October 1982, I was guided to a fossil echinoderm locality by Mr. Ralph Harris, a paraprofessional collector working with me in studies of fossil Cretaceous Mis- sissippi decapod crustaceans. The exposure is situated just west of and below the Liberty School and Church in the SW '4, SW 14, Sec. 3, T 8S, R 3E, Union County, Mississippi. This locality, herein called The Liberty School Locality (Gale A. Bishop Locality 52 indicated herein GAB 52), is a scraped area just off the road to the northeast of a small ridge perpendicular to Sand Creek (Fig. 1). The rocks exposed in the scraped area consist of a few centimeters of marlstone overlain by three meters of sandy, fossilif- erous limestone assignable to the Prairie Bluff Formation (Fig. 2), which is laterally equivalent to the clastic Owl Creek For- mation. The Prairie Bluff Formation is un- conformably overlain by the Paleocene Clayton Formation and unconformably overlies the Ripley Formation (Russell et al. 1982:11). The age of the Prairie Bluff Formation is Middle Maastrichtian (Russell et al. 1982:24). The fauna from the Prairie Bluff appears to be diverse (Stephenson and Monroe 1940: 204). At the Liberty School Locality, the fauna (Fig. 3) is dominated by the heart ur- chin Linthia variabilis Slocum but includes numerous other taxa including Baculites sp., Scaphites sp., Turritella sp., Exogyra cos- tata, Pinna sp., Urceolabrum sp. and nu- merous other molluscs. A small but integral part of this fauna are the decapods Proto- callianassa sp., Parapaguristes tuberculatus n. gen., n. sp., Palaeoxantho libertiensis n. gen., n. sp., Prehepatus harrisi Bishop, 1985; Raninella? sp., claw fragments of Hoplo- paria sp., and an indeterminate claw of a portunid crab. Stephenson and Monroe (1940:208a) presented a faunal list for the Prairie Bluff Formation that included Cal- lianassa sp., Avitelmessus n. sp., Prehepatus n. sp., Raninoides ovalis Rathbun, a por- tunid, and crab claws. These specimens are in the collections of the United States Na- tional Museum of Natural History but could not be located during a visit in 1984. Order Decapoda Latreille, 1803 Infraorder Anomura H. Milne-Edwards, 1832 Superfamily Paguroidea Latreille, 1803 Family Paguridae Latreille, 1803 VOLUME 99, NUMBER 4 605 Ween cae ARS Union County Fig. 1. Subfamily Pagurinae Latreille, 1803 Parapaguristes, new genus Type species. —Paguristes whitteni Bish- op, 1983. Diagnosis.—Claws large, equal, longer than high, thick and biconvex; upper and lower margins convex, proximal margin strongly oblique, distal margin vertical, par- alleled by furrows on front and back; fixed finger short, blunt, and downturned; top Margin on inner face more or less over- turned into horizontal crest; surface orna- mented by coarse tubercles. Etymology. — Parapaguristes is derived from Para (Greek, near to) + Paguristes (Greek, a genus of (hermit) crab); gender masculine. Occurrence. —The two species that com- prise Parapaguristes are both from the east- central Mississippi Embayment occurring le Geographic setting of the Liberty School Locality. ere in the Coon Creek Formation (P. whitteni Bishop, 1983) and Prairie Bluff Chalk (P. tuberculatus n. sp.) and range from early to middle Maastrichtian in age. Parapaguristes whitteni (Bishop, 1983) Fig. 5B Synonymy. —Paguristes whitteni (Bish- op, 1983:420-—424, figs. 3F, 4. Diagnosis. —Chelipeds large, equal, sim- ilar, biconvex and inflated, upper margin overturned into horizontal crest; finger downturned; ornamented by longitudinal rows of tubercles on outerface. Remarks. — Parapaguristes whitteni is well known, being represented by nearly all ele- ments of the chelipeds. In my assignment to Paguristes, I pointed out (Bishop 1983) the relatively poor concurrence of this species with the genus Paguristes, with the 606 | Prairie Bluff 250 Chalk MAASTRICHTIA Bluffport Marl Member = _ © E N= ° wt ay PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | Demopolis Chalk STRATIGRAPHIC POSITION of the LIBERTY SCHOOL LOCALITY UPPER CRETACEOUS Tt ! nt UNION COUNTY, MISSISSIPPI CAMPANIAN HH AE | | HH Tombighee Sand Member Fig: 2. suggestion of Forest (Bishop 1983:419) and now myself that it is better to erect a new taxon for fossils such as Paguristes whitteni than to try to assign them to existing taxa in which they really do not fit. The descrip- tion of taxa based solely upon claws is nec- essary because certain taxa have differen- tially mineralized exoskeletons leading to preferential preservation, as in the hermit crabs. Parapaguristes whitteni was described from a sample of 30 specimens, all claws. Parapaguristes tuberculatus, new species Figs. 3D, 4, 5A Diagnosis. —Claws large, biconvex, equal, palm longer than high; overturned horizon- tal crest on top margin of inner face reduced to subtle ridge paralleling margin; entire claw es LONG are as EA Stratigraphic setting of the Liberty School Locality. ornamented by random pattern of large tu- bercles alternating with granules. Description. —Claws large, biconvex, longer than high (L/H = 1.4), highest at dis- tal third, upper and lower margins convex; distal margin vertical, paralleled by shallow furrows; proximal margin oblique, round- ing evenly onto upper margin. Entire sur- face covered by coarse tubercles inter- spersed with very small granules in random pattern except for relatively smooth area on inner face along proximal articulation and bottom of distal marginal grooves. Tuber- cles conical; those on upper and lower mar- gins tilted slightly distally; fixed finger ap- parently downturned. Comparison. — Parapaguristes tubercula- tus differs from Parapaguristes whitteni by being ornamented more richly and evenly, by its less well developed overturned crest VOLUME 99, NUMBER 4 607 Fig. 3. Important elements of the Liberty School fauna. A, Prehepatus harrisi Bishop, 1985 (GSCM 1687); B, Portunid(?) claw (GAB 52-14); C, Raninella? sp. (GAB 52-13); D, Parapaguristes tuberculatus n. sp. (GSCM 1690); E, Palaeoxantho libertiensis n. sp. (GSCM 1692); F—G, Occulusional surface of crushing claw of lobster or crab (GAB 52-5); H, Callianassa sp. (GAB 52-4); I, Linthia variabilis Slocum, 1909 (GAB 52-6); J, Urceo- labrum sp. (GAB 5207); K, Baculites sp. (GAB 52-8); L, Scaphites sp. (GAB 52-9); and M, Exogyra costata Say, 1820 (52-10); exhibiting healed predation breaks. (Bar scales—1 cm; Scale 1, Figs. A, D, F—G, I—M; Scale 2, Figs. B—C, E, H.) on the upper part of the inner face, and its smaller size (Fig. 5). Etymology. —Named for the exceedingly tuberculate claws. Types. —The holotype (GSCM 1690) and paratype (GSCM 1691a, b) of Parapagu- ristes tuberculatus are deposited in the col- lection of the Georgia Southern College Mu- seum, Georgia Southern College, Statesboro, Georgia 30460. Occurrence, sample size, and preserva- tion. —Four specimens were collected from the Liberty School Locality: the holotype, a single left claw; the paratypes, a pair of claws found in close proximity to one another and a single right claw. The exoskeleton is pre- served intact and the claws are filled with sediment. Section Brachyrhyncha Borradaile, 1907 Superfamily Xanthoidea MacLeay, 1838 Family Xanthidae MacLeay, 1838 Palaeoxantho, new genus Type species. —Palaeoxantho libertiensis Nn. sp. 608 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON F Fig. 4. Photographs of Parapaguristes tuberculatus. A-D, Outer, top, bottom, and inner views of holotype left claw (GSCM 1680); E-F, Outer view of paratype claw pair (GSCM 1691a, b). (Bar scale = 1 cm.) Diagnosis.—Carapace oval-pentago- nal, wider than long, widest across epibran- chial lobes; anterolateral margins upturned, front rounded; posterolateral margins rela- tively straight; converging posteriorly; pos- terior margin unknown; rostrum bilobed and sulcate; orbits small, bifissured above; fronto-orbital width 58% of carapace width. Carapace well differentiated into tumid lobes by well defined grooves. Etymology.—From the combination of Paleo (Greek, old) + Xantho (Greek, one of Cyrene’s attendant nymphs). Occurrence. —Palaeoxantho is known only from the middle Maastrichtian Prairie Bluff Formation at the Liberty School Lo- cality (GAB 52) in Union County, Missis- sIppi. Remarks. — Palaeoxantho is similar to the xanthid genera Syphax A. Milne-Edwards, 1863 (Eocene France), Xantho Leach, 1804 (Miocene-Pleistocene, Europe and Fiji; Re- cent, Indopacific, Mediterranean, Eastern Atlantic), and Xanthosia (Albian- ? Paleo- cene, Europe, North America, ? W. Africa). Palaeoxantho has the same general shape and groove morphology as Syphax but has better differentiated, more tumid regions, a trend carried even further by Xantho with its more differentiated but less tumid car- apace. Palaeoxantho differs from these by its upturned lateral margins, its general flat- ness, and the tumidity of its carapace. Pa- laeoxantho is relatively narrower and thick- er than Xanthosia but bears a resemblance to that genus (and many other xanthids). VOLUME 99, NUMBER 4 Palaeoxantho libertiensis, new species Figs. 3E, 6 Diagnosis. —Carapace ovate-pentagonal, wider than long, widest near middle. An- terolateral margin and frontal margins broadly rounded, anterolateral margins up- turned; front short, bilobed, medially sul- cate, 10% of carapace width; orbits small, bifissured above, upturned; frontal-orbital width 58% of carapace. Posterolateral mar- gins relatively straight, converging poste- riorly; hind margin unknown. Carapace well differentiated by grooves into tumid lobes. Description.—Carapace small, rounded pentagonal, wider than long (L/W = 0.80), widest across epibranchial lobes near mid- length. Frontal and anterolateral margins broadly rounded, upturned, posterolateral margins fairly straight, convergent poste- riorly, posterior margin unknown. Front short, medially grooved, bilobed, 10% of carapace width; frontal region between in- ner angles of orbits 33% of carapace width; orbits small, upturned, bifissured above, distance between outer edges of orbits 58% of carapace width. Four anterolateral lobes present (frontal, hepatic, and 2 epibranchi- al), hepatic placed at anterior of convergent posterolateral margin. Carapace well differ- entiated by grooves into tumid regions; scapular arch separated from cephalic arch by sinuous cervical furrow. Cephalic arch differentiated into mesogastric lobe, trian- gular behind with narrow anterior tongue; very large, tumid protogastric lobes; small longitudinally elongate epigastric lobes; slightly tumid hepatic lobes, and poorly de- fined frontal areas. Scapular arch well dif- ferentiated into tumid regions; cardiac lobe rounded-hexagonal, separated from meso- gastric lobe by broad transverse depression (continuing across entire dorsal shield), epi- branchial regions differentiated into 3 boss- es, 2 marginal, anterior at widest part of carapace and both produced into upturned wings, and one large boss forming most of region; epibranchial region separated from 609 Fig. 5. Comparison of outer views of left claws of (A) Parapaguristes tuberculatus (GSCM 1690) and (B) Paraguristes whitteni (Bishop, 1983) (GSCM 1683). (Bar scale = 1 cm.) mesogastric lobe by aforementioned broad, transverse depression; mesobranchial lobe developed into transverse ridge, in line with cardiac region, with slight tumid area along margin of dorsal shield; intestinal and meta- branchial regions unknown. Carapace finely and sparsely granulate over entire surface, especially on summits of bosses. Sides of carapace downturned, nearly vertical, slant- ing slightly under dorsal shield; sides gran- ulate, faint grooves present on sides in line with ends of cervical groove and groove sep- arating marginal epigastric bosses. Comparison. —Palaeoxantho libertiensis resembles Syphax crassus (A. Milne-Ed- wards, 1864) from the Lower Tertiary of southern France in carapace outline but dif- fers significantly from it by having straighter posterolateral margins, a better areolated carapace, and upturned anterolateral mar- gins. Palaeoxantho libertiensis is similar to Titanocarcinus serratifrons A. Milne-Ed- wards in carapace outline but differs in hav- ing a better areolated carapace, upturned anterolateral margins, and lacking the pro- nounced dentate anterolateral margins. Pa- laeoxantho libertiensis resembles Xantho impressus (Lamark) from the Recent in its general carapace shape and areolation but differs from it by being relatively narrower, having a better areolated carapace, and by its lack of differentiation of the protogastric and epibranchial lobes. Palaeoxantho lib- ertiensis differs from species of Xanthosia by being relatively more oval, thicker (i.e., not flattened), narrower, by having a more rounded front, a better areolated carapace, c D Fig. 6. Palaeoxantho libertiensis seen in: A, dorsal; B, anterior; C, right lateral; and D, oblique anterior views, holotype (GSCM 1692). (Bar scale = 1 cm.) and upturned anterolateral margins. In spite of these major differences, P. libertiensis re- sembles species of Xanthosia in a general way, and specifically is more similar to Xan- thosia granulosa (McCoy) and Xanthosia buchii (Reuss) but still very different from them as expressed above. Etymology. —The trivial name /ibertien- sis is derived from the type locality near the Liberty School and Church. Type.—The holotype (GSCM 1692) of Palaeoxantho libertiensis is deposited in the collection of the Georgia Southern College Museum, Georgia Southern College, States- boro, Georgia 30460. Occurrence.—One partial carapace has been collected from the Prairie Bluff For- mation near the Liberty School and Church in Union County, Mississippi. Conclusions The decapods of the Prairie Bluff For- mation represent the youngest Late Creta- ceous fauna known in North America. As more of this fauna is described and com- pared to the older and younger Gulf Coast crab faunas, it may be possible to document the presence or absence of a mass extinction event among decapods of the Gulf Coastal Plain at the Cretaceous-Tertiary boundary. The decapods of the Prairie Bluff occur as rare elements in the fossil assemblages dominated by echinoids and molluscs, not PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON as decapod-dominated assemblages. It is anticipated that few new additional taxa will be discovered; hence, the importance of de- scribing the known decapods becomes more significant. To attain this end, I have de- scribed a new Prehepatus from the Prairie Bluff (Bishop 1985) and in this paper have described two new species (Parapaguristes tuberculatus and Palaeoxantho libertiensis). Three additional Prairie Bluff decapod taxa are known and in hand. One of those is represented by large claw fragments of a shell-crushing decapod (Fig. 3F, G). This taxon is known from many localities rang- ing from the Demopolis Formation through the Prairie Bluffin Mississippi and Alabama and will soon be described. The fauna also includes a raninid crab assigned to Rani- nella? (Fig. 3C) which may be the same tax- on reported by Stephenson and Monroe (1940:208a) as Raninoides ovalis Rathbun. The collection also contains a claw of a brachyuran (Fig. 3B) which probably be- longs to a portunid crab. A similar claw, from the Coon Creek Formation, was named and assigned as a lobster, Eryma flecta Rathbun, 1926, and maintained in that tax- on in 1935 (Rathbun 1935:21). Both claws appear to belong to portunid crabs. I hesi- tate to name and make such reassignment until the portunid cited by Stephenson and Monroe (1940:208a) is examined. A calli- anassid claw (Fig. 3J) is likewise not named, because it is difficult to make even a generic assignment based on unique claws of the callianassids (Rathbun 1935:29). Prehepa- tus harrisi Bishop, 1985, may prove to be the same as Prehepatus n. sp. cited by Ste- phenson and Monroe (1940:208a). Iam also looking forward to seeing the specimen of Avitelmessus n. sp. cited by them. The decapods now known from the mid- dle Maastrichtian Prairie Bluff Formation of Northern Mississippi include: Fig. 3F—G Claw fragments indet. Fig. 3J Callianassa? sp. Fig. 3D Parapaguristes tuberculatus n. sp. Fig. 3C Raninella(?) sp. (?=Raninoides VOLUME 99, NUMBER 4 ovalis Rathbun: Stephenson and Monroe, 1940) Fig. 3A Prehepatus harrisi Bishop, 1985 (?=Prehepatus n. sp. Monroe) Fig. 3E Palaeoxantho libertiensis n. sp. Fig. 3B Portunid claws indet. (?=Eryma cf. flecta Rathbun and ?=portunid of Ste- phenson and Monroe 1940) No Fig. Avitelmessus n. sp. (Stephenson and Monroe, 1940) Acknowledgments The assistance of Mr. Ralph Harris in col- lecting decapods from the Cretaceous of Mississippi is gratefully acknowledged. Di- rect support of this research was received from the National Science Foundation (Grant DEB 8011570) and the National Geographic Society (Grant NGS 1629). In- direct support was received from the De- partment of Geology and Geography and the Faculty Research Committee, Georgia Southern College. The manuscript was typed by Phyllis Wiggins, Donna Cain, and Judith Underwood. The manuscript was strength- ened considerably by R. M. Feldmann and J. S. H. Collins, whose constructive criti- cism is especially appreciated. 611 Literature Cited Bachmayer, F., and R. Mundlos. 1968. Die tertiaren Krebse von Helmstedt bei Braunchweig Deutschland.—Annalen Naturhistorische Mu- seum Wein 72:649-692. Bishop, G. A. 1983. Fossil decapod Crustacea from the Late Cretaceous Coon Creek Formation, Union County, Mississippi.—Journal of Crus- tacean Biology 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 59:1028-1032. Rathbun, M. J. 1926. Decapoda. Jn B. Wade, ed., The fauna of the Ripley Formation on Coon Creek, Tenn.— U.S. Geological Survey Profes- sional Paper 137:184-191. 1935. Fossil Crustacea of the Atlantic and Gulf Coastal Plain.—Geological Society of America, Special Paper 2:1—160. Russell, E. E., D. M. Keady, E. A. Mancini, and C. E. Smith. 1982. Upper Cretaceous in the lower Mississippi Embayment of Tennessee and Mis- sissippi: Lithostratigraphy and biostratigraphy. Earth Enterprises, Inc., 50 pp. Stephenson, L. W., and W. H. Monroe. 1940. The Upper Cretaceous deposits.— Mississippi State Geological Survey, Bulletin 40:1-296. Institute of Arthropodology and Parasi- tology, Department of Geology and Geog- raphy, Georgia Southern College, States- boro, Georgia 30460-8149. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 612-615 A NEW SPECIES OF TUBIFICOIDES LASTOCHKIN (OLIGOCHAETA: TUBIFICIDAE) FROM BERMUDA AND THE BAHAMAS Berit Rasmark and Christer Erséus Abstract. — Tubificoides bermudae, new species, is characterized by (1) pap- illated body wall in postclitellar segments, (2) possession of only bifid setae, 2-3 per bundle anteriorly, 1-2 per bundle in postclitellar segments, (3) vasa deferentia that are about as long as atria, and (4) conical penis sheaths with smooth outline and slightly distended distal end. The species appears closely related to 7. brownae Brinkhurst and Baker, and 7. motei Brinkhurst, but is easily distinguished from these by the characteristic shape of its penis sheaths. While studying the marine oligochaete fauna of Bermuda, the second author en- countered a new species of Tubificoides La- stochkin at several subtidal stations of mud- dy sand. The same species was also found in material collected in Bermuda and the Bahamas by Dr. Meredith L. Jones, Na- tional Museum of Natural History (USNM), Washington, D.C. Material and Methods Specimens were sorted from sand col- lected at various sites in Bermuda in 1977 and were fixed in Bouin’s fluid. Seventeen specimens were sectioned and stained in Azan or Heidenhain’s hematoxylin and eo- sin. The bulk of the individuals were, how- ever, stained whole in paracarmine; 6 of these were dissected, the remaining 33 mounted whole in Canada balsam. Three additional whole-mounted specimens, one from Bermuda and two from the Bahamas, were kindly placed at our disposal from the USNM. The type-series of 7. bermudae is deposited in the USNM. Tubificoides bermudae, new species Figs. 1, 2 Holotype. —USNM 98945, whole- mounted specimen from Smith’s Sound (SE end of Smith’s Island), Bermuda, 32°22'08’N, 64°39'25”W, 5 m, muddy fine sand (coll. C. Erséus, 2 Dec 1977). Paratypes. -USNM 98946-98951, 1 sec- tioned, 1 dissected and 4 whole-mounted specimens from type locality. Other material. -USNM 98954, 1 whole- mounted specimen from Blue Hole area, base of causeway, Bermuda, sand (coll. M. L. Jones, Aug 1975). USNM 98952-98953, 2 whole-mounted specimens from Andros Island, Bahamas (coll. M. L. Jones, Mar 1966): 1 from S shore of Fever Cay, Middle Bight, lower intertidal, sand; and 1 from S side of Fresh Creek, “‘deeper flats with fil- amentous algae.’ Second author’s collec- tion: 12 sectioned, 5 dissected and 28 whole- mounted specimens collected in Bermuda by C. Erséus (Aug and Dec 1977); 9 worms from type locality, others from 4 different stations, 0.2-15 m depth, largely muddy sands (often with slight smell of H.S). Etymology. —The name bermudae refers to the geographic origin of most of the ma- terial studied. Description. —Body wide anteriorly, par- ticularly at sexual maturity, behind clitel- lum abruptly narrower; postclitellar part slender with elongated segments (Fig. 1A). Color greyish red. Length (8 complete, fixed specimens) 6.4—12.3 mm, about 50-60 seg- VOLUME 99, NUMBER 4 Yes Be ath SS X A x ae ~ NGNATE — 2 -— " \ 7 ‘eh af * ; \« 4 fe fe \\ *, ff \ é \ ge ee j i cee 1 Fig. 6. Photomicrographs of spicules: a, Cliona mazatlanensis, tylostyles and microscleres; b, C. mazatla- nensis, microspined oxeas and microrhabds; c, C. vastifica, tylostyles and microscleres; d, C. vastifica, microspined oxeas and microrhabds; e, C. canadensis, tylostyles and microscleres; f, C. canadensis, microspined oxeas and microrhabds; g, C. muscoides, microspined oxea and centrotylote microrhabds; h, C. cervina, tuberculated oxea and microrhabds bearing microspine clusters. Scales: a, c, e, 100 um; b, d, f—h, 25 um. \ SS PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ab otter ESS 2 e see. f eee a Fig. 7. Photomicrographs of spicules: a, Thoosa bulbosa, amphiasters and reduced oxyaster (“‘bird wing’’); b, T. cactoides, amphiasters; c, Cliona (=Aka) labyrinthica, oxeas; d, C. (=Aka) nodosa, oxea; e, C. (=Paracor- nulum) purpurea, tylotes and acanthotornotes; f, C. (=Paracornulum) purpurea, acanthotorotes and palmate isocheles. Scales: a, b, f, 25 wm; c, d, e, 100 um. final conclusions can be drawn without study of fresh and complete material, entire pop- ulations, and knowledge of environmental parameters. Topsent (1932:558) already had evidence that environmental conditions, such as salinity, can modify the spiculation of Cliona vastifica. Availability of dissolved silicic acid is another important factor known to influence spicule shape and size (Simpson 1981). On the other hand, pop- ulations of Cliona lampa (group (1), above) studied in Bermuda (Ritzler 1974) display very stable spicule characteristics but turn out to represent two distinct species distin- guishable by color, spicule size, and growth habit, as confirmed by long term observa- tions and field experiments (Ritzler, in prep.). Excavating sponges described by Han- cock but not belonging to the Clionidae are VOLUME 99, NUMBER 4 Cliona labyrinthica and C. pupurea. The former, transferred to Aka, is now viewed by us as a member of the Oceanapiidae (Haplosclerida), and the latter, transferred to Paracornulum, may belong to the Coe- losphaeridae (Poecilosclerida). Acknowledgments We are much indebted to A. Nimmo, for- mer curator at The Hancock Museum, whose interest and determination were in- strumental in recovering the Albany Han- cock microscope slides. We also wish to ex- tend our sincere thanks to A. M. Tynan, curator of The Hancock Museum, for his patience during the period of this study, and to P. Davis, now at The Hancock Museum, for his help in interpreting some of the cura- torial evidence. Kate Smith provided tech- nical assistance. Literature Cited Bowerbank, J. S. 1866. A monograph of the British Spongiadae, Vol. 2. Ray Society, London, 388 pp. Hallman, E. F. 1920. New genera of monaxonid sponges related to the genus Clathria. —Pro- ceedings of the Linnaean Society, New South Wales 44:767-792. Hancock, A. 1849. On the excavating powers of cer- tain sponges belonging to the genus Cliona; with descriptions of several new species, and an allied form.—Annals and Magazine of Natural His- tory (2) 3:321-347. 1867. Note on the excavating sponges; with descriptions of four new species.— Annals and Magazine of Natural History (3) 19:229-242. Hartman, W. D. 1958. Natural history of the marine sponges of southern New England.— Peabody Museum of Natural History, Yale University, Bulletin 12:1-155. Johnson, J. Y. 1899. Notes on some sponges be- longing to the Clionidae obtained at Madeira. — Journal of the Royal Microscopal Society, Transaction 9:461—463. Kirkpatrick, R. 1900. Description of sponges from Fanafuti.—Annals and Magazine of Natural History (7) 6:345-362. Laubenfels, M. W.de. 1936. A discussion of the sponge fauna of the Dry Tortugas in particular and the West Indies in general, with material for a re- vision of the families and orders of the Porif- era.— Carnegie Institution of Washington, Pa- pers from the Tortugas Laboratory 30:1—225. 675 1950. The Porifera of the Bermuda Archi- pelago.— Transactions of the Zoological Society of London 27:1-154. Lendenfeld, R. von. 1896. Die Clavulina der Ad- ria.—Nova Acta, Deutsche Akademie der Na- turforscher (Halle) 69:1—251. Old, M. C. 1941. The taxonomy and distribution of the boring sponges (Clionidae) along the Atlan- tic coast of North America. — Chesapeake Bio- logical Laboratory, Solomons Island, Maryland, Publication 44:1—30. Ritzler, K. 1971. Bredin-Archbold-Smithsonian Bi- ological Survey of Dominica: Burrowing sponges, genus Siphonodictyon Bergquist, from the Ca- ribbean.— Smithsonian Contributions to Zool- ogy 77:1-37. . 1974. The burrowing sponges of Bermuda. — Smithsonian Contributions to Zoology 165:1- 32. . [In prep.] Co-occurrence of Cliona carpenteri and C. /ampa (Porifera: Hadromerida: Clioni- dae) in Bermuda. Simpson, T. L. 1981. Effects of germanium on silica deposition in sponges. Pp. 527-550 in T. L. Simpson and B. E. Volcani, eds., Silicon and siliceous structures in biological systems. Springer-Verlag, New York. Topsent, E. 1888. Contribution a l’étude des clio- nides.— Archives de Zoologie Expérimentale et Générale, series 2, 5 (supplement): 1-165. 1891. Deuxiéme contribution 4 l’étude des clionides.—Archives de Zoologie Expérimen- tale et Générale, series 2, 9:555-592. 1900. Etude monographique des spongiaires de France, III. Monaxonida (Hadromerina). — Archives de Zoologie Expérimentale et Génér- ale, series 3, 8:1—328. 1907. Cliona purpurea Hck. n’est pas une clionide.— Archives de Zoologie Expérimentale et Générale, series 4, 7:X VI-XX. 1932. Notes sur des clionides.— Archives de Zoologie Expérimentale et Générale, 74 (Jubilee Volume):549-579. Volz, P. 1939. Die Bohrschwamme (Clioniden) der Adria.— Thalassia 3 (2):1-64. Vosmaer, G. C. J. 1933. The sponges of the Bay of Naples, Porifera Incalcaria, with analyses of genera and studies in the variations of species, Vol. I. Martinus Nijhoff, The Hague, 456 pp. (KR) Department of Invertebrate Zool- ogy, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (SMS) Department of Zoology, Brit- ish Museum (Natural History), Cromwell Road, London SW7 5BD, England. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 676-702 NEWLY ESTABLISHED FAMILIES OF THE ORDER BRANCHIOBDELLIDA (ANNELIDA: CLITELLATA) WITH A SYNOPSIS OF THE GENERA Perry C. Holt Abstract. — General remarks and a brief survey of the branchiobdellid liter- ature are followed by a review of the group’s taxonomic characters, methods of study and speculations as to its origins. A key to newly recognized families and eighteen genera is given. The order is divided into five families: Bran- chiobdellidae, Bdellodrilidae, Xironodrilidae, Caridinophilidae, Cambarincol- idae. A synopsis of the genera is included with the familial diagnoses. The annelid worms of the order Bran- chiobdellida are currently attracting a mod- est degree of attention. Since 1950, nine of the eighteen genera and more than eighty of their 132 included species have been diag- nosed. The opinion once held that the bran- chiobdellidans are a homogeneous group (Stephenson 1930:796) has long since been abandoned; their former association with either the Hirudinea or Oligochaeta has been dissolved (Holt 1965b). This proliferation of taxa and the recog- nition of the structural diversity expressed thereby now make it appropriate to segre- gate the genera of this previously monotypic order into families and to present a synopsis of the included genera. A brief summary of branchiobdellidan natural history will be followed by a review of selected literature devoted to them, a consideration of the methods and anatomical characters that have been used in taxonomic studies of them and a discussion of the group’s origin and distribution. This introduction is followed by keys to the families and genera and di- agnoses of these taxa with illustrations, lit- erature citations, number of included species and geographical range. It is to be empha- sized that these efforts should be considered as provisional and transitional in nature: probably much less than half of the world fauna of the worms is known. The branchiobdellidans are obligate sym- bionts of Holarctic (with limited excursions into the borders of the Neotropical and Ori- ental regions in Central America and China) freshwater crustaceans (crayfish, crabs, shrimps and isopods) that move in a leech- like fashion over the bodies of their hosts and feed on a variety of substances: the host’s blood, eggs and (?) young, other external symbionts and, most often, the bacterial and algal gloea that often covers the host’s body. Among those found on astacoidean cray- fishes, different species tend to occur on dif- ferent regions of the host animal and their feeding habits may reflect their occupancy of these microhabitats. Nearly all astacoideans carry these sym- bionts. Although there is no known species- to-species host specificity, some _ bran- chiobdellidans, e.g., the isopod egg-eating Cambarincola aliena Holt, 1963, may be restricted to a single species of host because of such feeding adaptations. Branchiobdel- lidans have been rather diffidently reported as living away from any host (Holt 1973a: 152-159), but there is no record of them depositing cocoons except on the host’s body. The first report of branchiobdellidans from other than astacoidean hosts (Hobbs and Villalobos 1958) was followed by the additions of several such instances: fresh- water crabs in Central America, Mexico and VOLUME 99, NUMBER 4 677 Fig. 1. Louisiana; troglobitic isopods from Ten- nessee and Mexico; pseudothelphusid crabs from Central America and Mexico; fresh- water shrimps from China. These instances seem to be from unusual habitats (caves) or from regions at the periphery, or beyond (Nicaragua and Costa Rica), of the range of the usual hosts. Little is known of other as- pects of the life history, ecology, and phys- iology of these worms. Some studies of their natural history are marred by improper or inadequate taxonomic determinations. The taxonomy of the group is incomplete and in some ways unsatisfactory simply be- cause a large proportion of the branchiob- dellidan fauna is undescribed: many new forms are known to await diagnosis in the collections of the National Museum of Nat- ural History of the Smithsonian Institution. A brief history of studies of the branchiob- dellidans. —Only the major taxonomic works are considered herein; other refer- ences may be found in them. In Europe, the number of nominal species of the genus Branchiobdella Odier, 1823, had reached forty-three when Pop (1965) reduced it to four species, one with three subspecies. This is almost surely an extreme example of “lumping,” but there are prob- ably only a few species of branchiobdelli- dans in Europe. Generalized branchiobdellid. Abbreviations: I-XI, trunk segments; b, bursa; br, brain; c, circulatory system; ed, ejaculatory duct; int, intestine; j, jaws; np, nephridiopore; ov, ovary; ps, penial sheath; sg, spermiducal gland; sp, spermatheca; vn, ventral nerve cord (from Holt 1969). The Japanese and Korean branchiobdel- lidan fauna was studied by Yamaguchi (1934). A few species have been recognized by Chinese workers (Liang 1963; Liu 1964, 1984; Liu and Chang 1964; Liu and Zhang 1983). The Asiatic fauna, as now known, consists of 32 species assigned to six genera (two of these genera are new ones to be pro- posed by Gelder and Liu (pers. comm.) and one is the North American genus Cambar- incola represented by the introduced C. okadai Yamaguchi, 1933). The principal homeland of the bran- chiobdellidans is North America. Impor- tant early contributions to a knowledge of these worms were made by Moore (1895b) in his anatomical study of Bdellodrilus il- luminatus, which has served as a model for all subsequent such studies, and the recog- nition of new forms (Moore 1894, 1895a). Ellis (1912, 1918, 1919) diagnosed the gen- era Cambarincola (1912), Xironodrilus (1918), and Xironogiton (1919) and recog- nized a total of seven new species in these and other genera. Hall (1914) raised the family Branchiobdellidae to the status of a superfamily and erected the genus Cerato- drilus. Goodnight (1940) assembled these works and others, divided the Branchiob- dellidae into two subfamilies and added some generic diagnoses and species descrip- 678 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON aL .[) Ellisodrilus Cambarincola Sh). Pterodri Ceratodrilus rilus Xironogiton ae Ankyrodrilus MD, () Bdellodrilus S47 ) Triannulata atl Cirrodrilus Me) Tettodrilus pu.) Branchiobdella Oedipodrilus | , = S S Xironodril BRANCHIOBDELLIDAE veneers Sathodrilus BDELLODRILIDAE See Caridinophila CAMBARINCOLIDAE XIRONODRILIDAE CARIDINOPHILIDAE PROTOBRANCHIOBDELLID Fig. 2. Suggested relationships of the genera and families of the order Branchiobdellida. VOLUME 99, NUMBER 4 tions. Hoffman (1963) treated 21 species, 12 of them new, in his monograph of the genus Cambarincola which also contains important reflections on taxonomic char- acters. Holt (1949) demonstrated the im- portance of the reproductive systems as a source of taxonomically important charac- ters. Since then, he has described 55 new species and erected eight new genera (Holt 1960b, 1965a, 1967a, b, 1968b, 1977b) and published monographs of the genera Pter- odrilus (Holt 1968c) and Xironogiton (Holt 1974b). The argument for raising the family Branchiobdellidae to ordinal rank (Holt 1965b) was followed by a review that is semi- popular in style (1968a) and zoogeographic treatments (Hobbs et al. 1967, Holt 1969), among other studies. Methods of collection, preservation and preparation for study. — An inhibiting factor in the study of the branchiobdellidans is the difficulty of doing anatomical studies of them. The worms range from about one to six millimeters in length (when contracted) and are opaque unless special methods are used. The preparation of serial sections is often hampered by the presence of diatoms in the gut, but it is essential that beginning students familiarize themselves with the ba- sic anatomy of the animals by studies of such sections and of dissections. The prep- aration of stained whole mounts of the an- imals requires live specimens to be relaxed (? with 1-5% magnesium chloride), fixed and stained. The more usual method is to dehydrate specimens in ethanol, clear in clove oil and mount unstained in Canada balsam. Microscopic study of specimens so treated requires the use of an objective for the microscope that is adjusted to an un- usually long working distance (approxi- mately 1.5 mm) that enables the student to see through the total thickness of the worm or to turn over the slide on which it is mounted in order to view it from both sides. Another nuisance occurs because much of the material presented to the student has 679 been taken from the sediment at the bottom of jars in which crayfish have been collected. The students of crayfish usually preserve their animals in 70% ethanol and jam all the crayfish they can find into a collecting jar. The result is rotten worms often totally useless for serious study. For the best re- sults, using the method described above, the branchiobdellidans should be preserved in 5% formalin in 70% ethanol and the col- lecting jars should not be overcrowded. Taxonomic characters.—The students of the branchiobdellidans have used a number of features as taxonomic characters. To evaluate these usages a brief description of a generalized branchiobdellidan (Fig. 1) is required. The body consists of 15 segments of which four constitute a head and the re- mainder a trunk with a terminal sucker. Only the post-cephalic body segments have tra- ditionally been numbered. The prostomium is absent and the first cephalic segment, the peristomium, is usu- ally divided into upper and lower lips which may be further subdivided into lobes; oc- casionally lateral lobes are present. The up- per lip in some forms may be furnished with digitate projections or tentacles. The body segments are indicated externally by inter- segmental furrows and are usually subdi- vided by a circular groove which produces a secondary annulus. The anterior annulus of some segments in some species is greater in diameter than the secondary annulus: there are “‘dorsal ridges.’ The dorsal ridges may carry fan-like or finger-like projections. The greater diameter of the anterior annuli in these cases is produced by the insertion of slips of the longitudinal segmental mus- cles into the outer body wall posterior to the intersegmental furrows and into the sec- ondary furrow that delimits the posterior, secondary annulus, or sometimes in front of this furrow (Holt 1960b:fig. 7). The body outline is said to be smooth if the dorsal ridges are absent. The anterior nephridia open on segment III through two dorsolat- 680 eral pores or by a median dorsal pore. The anus opens dorsally on what apparently is always segment X. The posterior sucker (the peristomium forms an anterior one) is formed from the eleventh segment with, ap- parently, in some cases portions of segment X included. Both suckers are aided in their function by glandular secretions (Weigl 1980). Three otherwise dissimilar genera are characterized by a dorsoventral flattening of some of the mid-body segments. The nervous and circulatory systems are annelidan, apparently invariant, and rarely mentioned in taxonomic works. The diges- tive system likewise presents few features of taxonomic interest: the gut is expanded much more noticeably in some segments of dorsoventrally flattened forms, the pharynx has one or more expansions (pharyngeal sul- ci); and all branchiobdellidans possess, dor- sally and ventrally, pharyngeal placoids (jaws) that normally bear teeth and that vary in shape, size and number of teeth. Oral papillae, thought to be sensory in function, surround the mouth opening in several forms. However, it is possible that these small structures are present in most bran- chiobdellidans and because of their size have been simply overlooked in the descriptions of many species. There are two pairs of nephridia. An an- terior pair lies asymmetrically alongside the gut in segments I-IV, and discharges to the exterior on segment III. Variation in the position of anterior nephridia may exist among members of a species or between species (Moore 1897:329-330), but this possibility has not been further investigat- ed. The posterior pair lie, one on each side, in segment VIII (Moore 1897:332) and open on the anterior lateral surface of segment IX (Freeman 1963). The female reproductive system consists of paired ovaries, placed laterally on the posterior face of septum 6/7 and a sper- matheca in segment V. There are no ovi- ducts; eggs are expelled through a pair of ventrolateral pores in the body wall of seg- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ment VII. The spermatheca opens by a me- dian pore on the venter of segment V and is a blindly ending sac that may be variously modified along its length. The epidermis of segments VI and VII is provided with glands forming a mucus-secreting clitellum. The male system is more complex. Two pairs of testes located on the posterior faces of septa 4/5 and 5/6 break up at maturity into morulae that release spermatozoa into the body cavity. Paired male funnels in the posteroventral quadrats of the testicular segments open into thin ducts, the vasa ef- ferentia, that unite to form vasa deferentia. The latter fuse to form a glandular organ, the spermiducal gland, which may have a diverticulum of greater or lesser distinc- tiveness, the prostate. In most forms a mus- cular tube joins the spermiducal gland to the penial sheath, the ental part of the bursa, which encloses the penis. The latter projects into a cavity, the bursal atrium, which opens mid-ventrally on segment VI. The penis takes several forms: in some genera it is eversible, consisting of a cuticular tube with or without hooks and with or without strands (presumably muscular) that attach it to the inner wall of the penial sheath. In other forms it is clearly muscular and is protruded by the eversion of the wall of the atrium as a cone-shaped mass. In some species this mass is narrowed and similar in composi- tion and apparent eversibility to those of species in which a clearly eversible penis has strands connecting it to the inner wall of the penial sheath (Holt 1982:254—255). There are numerous permutations of the elements of the reproductive systems, of the variations of the jaws and their teeth and of the features of the body surface. Most of the variations in structure shown by the branchiobdellidans are stable and discon- tinuous and no instance is now known with certainty of North American species that gradually vary in space. Consequently no subspecies of these have been recognized (but cf. Cambarincola osceolai Hoffman, 1963:331). VOLUME 99, NUMBER 4 Some variations in body form are ob- vious adaptations to a particular way of life, but nothing can be said about the signifi- cance of most of the differences in structures that are used in the taxonomy of the group. Members of a genus usually present a common facies, although interspecies dif- ferences are common in some external fea- tures (for example, the presence or absence of peristomial tentacles or dorsal ridges which occur within the confines of a single genus). Two of the three dorsoventrally flat- tened genera and several terete ones cannot be separated on the basis of external ap- pearance and recourse must be had to fea- tures of the male reproductive system to do so. The same is true of the jaws: although members of a genus share a basic plan of jaw structure, one type of jaw may be shared by several genera. The number of anterior nephridiopores (one or two) are of necessity shared by several genera. Supraspecific taxa must, perforce, be based on modifications of the male reproductive system. There have been other discussions of taxonomic characters used in studies of the branchiobdellidans (Hoffman 1963; Holt 1953, 1960a, 1965a, 1968a, b, 1973c; Holt and Hoffman 1959, inter alia) and suppo- sitions expressed as to “primitive” as op- posed to “‘advanced”’ characters (cf. Holt 1973c:3). These efforts, however, have been directed to features diagnostic of both gen- era and species. Herein attention will be di- rected to features that characterize genera and suprageneric taxa and many features used in the diagnosis of species ignored. Their usage may be derived from the taxo- nomic works cited above. Of the somatic (non-genitalic) characters, only one is important in the present context. (The shape of the jaws has not been carefully considered, though Ellis (1919:241—243) laid the basis for a system utilizing the jaws). The ancestral branchiobdellidans probably were provided with separate openings of the anterior nephridia: five genera are so char- acterized; the remaining genera have a com- 681 mon opening (one anterior nephridiopore). In all cases, these openings, whether one or two, are obscure pores on the dorsum, or latero-dorsum, of segment III. In two of these five genera the middle segments are dorsoventrally depressed, but there is no reason to believe that there is any phylo- genetic relationship between the presence of two anterior nephridiopores and a de- pressed body form (segment III of all bran- chiobdellidans is terete). There is no way of knowing, except by uncertain inferences, the anatomical shapes and relationships of the components of the reproductive systems of the ancestors of the branchiobdellidans. Among living forms, the female system is relatively invariant. The spermatheca may be absent, reduced to ves- tigialness or “bifid” (having a lateral branch). Otherwise the variations in the presence or absence of an ental bulb, the shape of the spermathecal bulb (the sperm-storing part of the organ) and of the ectal spermathecal duct and bursa are only uncertainly, and in not all cases, associated with groups of species forming genera. This is not true of the male system. The testes may consist of one or two pairs. The vasa deferentia may enter the spermiducal gland ectad to its inner end or entally and there may or may not be a prostate or ru- diment thereof associated with it. The penis is variable in its structure. The composition of its wall ranges from a cuticular tube, with or without hooks, to a dense cone-shaped mass of muscle: the former type is eversible; the latter protrusible. There are left, then, relatively few features whose states may serve to characterize su- prageneric groupings and presumptively “‘primitive”’ characters (Holt 1968a:83-84) are found in more than one of any possible groupings of genera. Whether there are one or two nephridiopores; the number of pairs of testes and their location; the nature of the junction of the vasa deferentia with the spermiducal gland; the presence or absence of a prostate associated with the spermi- 682 Table 1.—Data matrix for some major characters. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Caridino- philidae | Cambarincolidae Xironodrilidae Bdellodrilidae Branchiobdellidae Spermiducal gland: globose — elongate globose elongate elongate elongate elongate Vasa deferentia enter sper- miducal gland: terminally — subtermi- nally — terminally terminally subterminally subterminally Spermatheca: present — absent absent absent-present present present present Mid-body depression: present — absent absent absent present absent absent-present Anterior nephridiopores: 2-1 1 1 2 1 2 Prostate: absent — present absent absent-present absent absent absent Testes 2 pairs — | pair 2 2 2 2 1 or 2 Penis: eversible — protrusible h eversible- protrusible —eversible eversible protrusible ducal gland and its degree of development; and the composition of the penis and its eversibility or protrusibility seem to be the totality of such characters. One would need to know in detail the phylogenetic history of the members of the order, and there are no clear clues to this history offered by the study of the anatomy of the worms or their present distribution, in order to assert that any of these features are primitive (Table 1). Origins and distributions. —It is probably futile to seek the ancestral stock of the bran- chiobdellidans among the other clitellates. Stephenson (1930:704—705) regarded them as oligochaetes and derived them from the Lumbriculidae. Holt (1965b) showed that they are not oligochaetes, but said nothing as to their origins. Brinkhurst (in Brinkhurst and Jamieson 1971:169-172, fig. 4.2) ar- gued for the primitive position of the lum- briculids and in his schema of phylogenetic relationships (Brinkhurst and Jamieson 1971:191, fig. 4.4) places the branchiobdel- lidans next to the former, but otherwise does not mention them. The branchiobdellidans are derived from an ancient stock of the Ur- oligochaetes, possibly quite different from any living form. The problem cries for re- study with the use of modern techniques, including cyto-chemistry and electron mi- croscopy (see Weigl 1980). Nonetheless, some speculations as to the origins and historical wanderings of the or- der and its members deserve, if for no other reason than refutation, brief mention. The distribution of the families and gen- era of the branchiobdellidans is not con- gruent with that of the astacoideans (Holt 1968a:85-86). Holt’s speculations therein antedate the general acceptance of the geo- logical theories of plate tectonics and may now be regarded as an example of the efforts of earlier historical zoogeographers to ex- plain the distribution of animals. More credible is Hobbs’ (1974:2; 1981:52) belief that the unusual distribution of the families of the Astacoidea is the result of separate invasions of freshwaters by different ances- tral marine stocks. But the branchiobdellidans originated in freshwater: no saline-tolerant branchiob- dellidans are known. The uncertainties of the consequences of VOLUME 99, NUMBER 4 floating and colliding continental land masses (and fragments, “‘terranes,”’ thereof) and the utter lack of a fossil record preclude any conclusions as to the times of origin prior to the Cretaceous and the subsequent migrations of the branchiobdellidans. They arose in the nothern regions of Pangaea that today are incorporated into the two Hol- arctic lands. The branchiobdellidans are an indepen- dent group, evolving in response to a myr- iad of environmental factors, only a few of which are determined by their means of transport. They were most likely adapted to symbiosis with unknown freshwater ani- mals (? crustaceans) when the various stocks of the astacoideans came in out of the sea. They can, thus, hardly be expected to bear the burden of explaining the evolutionary vagaries of their hosts. This is not to deny that plausible theories correlating the dis- tribution of both worms and hosts that now occupy a common area cannot be devised (Hobbs et al. 1967; Holt 1968c, 1969, 1973c, 1974b, inter alia). Problems of classification. — Any attempt at grouping the genera of the branchiob- dellidans into families is not only faced with the usual difficulties of such efforts, but is complicated by an unusual number of ap- parent convergences. Two such cases are of particular relevance: the penis appears to have evolved from a cuticular eversible tube into a muscular organ that may be either eversible or protrusible in more than one lineage or the opposite; the prostate is of variable development or absent in what otherwise are clearly related groups. These inconsistencies defy all attempts at organi- zation. Two salient features of members of the order are used to derive the arrangement which follows: the number of anterior ne- phridiopores and the points of entry of the vasa deferentia into the spermiducal gland. The problem of the variations in the struc- ture of the penis and the apparent conver- gencies presented by these differences may well be solved by more careful studies of the nature of the muscular penes in several 683 genera. (The problem has received little at- tention and has only recently been broached by Holt (1978a:479; 1982:254).) And the muscular penes of such genera as Ankyrod- rilus Holt, 1965; Xironogiton Ellis, 1919; and Adenodrilus Holt, 1977, among others, are different from and may not be homol- ogous with those of the much better known ones of Cambarincola Ellis, 1912. If this should prove to be the case, the arrange- ment presented herein is basically sound (Fig. 2). One other unrelated difficulty exists: Liu and Zhang (1983) described a Chinese species that they assigned to Branchiobdella as B. heterorchis with only one pair of testes in segment VI. If there is only one pair of testes in this segment in this species it pos- sibly should be placed in a separate family. Stuart R. Gelder and Liu Si-Cheng (pers. comm.) are considering this in a forthcom- ing review of east Asian branchiobdelli- dans. The species will not be further referred to herein. But it is conceivable that work yet to come on the Eurasian fauna will jus- tify the creation of suborders on the basis of the number and location of pairs of testes. For now the characters discussed herein do not clearly require this, though neither do they negate such a classification. The following, then, is proposed as a su- prageneric classification of the branchiob- dellidans. Class Clitellata Order Branchiobdellida Family Branchiobdellidae Bdellodrilidae Xuironodrilidae Caridinophilidae Cambarincolidae Key to the Families and Genera of the Order Branchiobdellida Holt A key to the then known branchiobdel- lidan genera of North America was provid- ed by Holt (1978b:292-295) for Pennak’s (1978) Freshwater Invertebrates of the 684 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON United States, but no comprehensive one for all the genera of the order has been pub- lished since Goodnight’s (1940:28-30) now outdated one. The one presented below is followed by familial and generic synony- mies, diagnoses, location of types, ranges, number of species and literature references. K Vasa efferentia enter spermiducal gland separately; no vasa defer- entia; spermiducal gland globose. Family Caridinophilidae ....... Caridinophila Vasa efferentia unite to form vasa deferentia; spermiducal gland elongate . (1) Vasa deferentia enter ental end of spermiducal gland Vasa deferentia enter ectad to ental end of spermiducal gland ....... 12 (2) Some body segments flattened; two anterior nephridiopores. Fam- ily Xironodrilidae Xironodrilus Terete; one anterior nephridio- pore. Family Cambarincolidae .. 4 (3) Penis eversible cuticular tube Oedipodrilus Penis ectally a muscular epithelial tube, or connected by strands to inner wall of penial sheath, or densely muscular (4) Ectal end of penis cellular ep- ithelial tube enclosing eversible cuticular tube; prostate arises en- tad to junction of spermiducal gland and ejaculatory duct Penis variously muscular . (5) Penis eversible, connected to inner wall of penial sheath by strands Penis densely muscular . (6) Body ornamented with peristo- mial tentacles and dorsal projec- tions Ceratodrilus Both without tentacles or dorsal projections Sathodrilus . (6) Spermiducal gland without a prostate 1S: Prostate or prostatic rudiment as- sociated with spermiducal gland 10 . (8) Penis composed of muscular, ae wet alle eversible ejaculatory duct Bursa long, muscular; retracted penis short, eversible; spermiducal gland long, slender ..Magmatodrilus . (8) Spermatheca absent; penis composed of atrial fold. . Ellisodrilus Spermatheca present; penis pro- trusible muscular cone . (10) Prostate incompletely divided from spermiducal gland .. Pterodrilus Prostate arises at junction of sper- miducal gland and ejaculatory duct te JM: OS OO Be Cambarincola . (2) One anterior nephridiopore. Family Bdellodrilidae .......... 13 Two anterior nephridiopores. Family Branchiobdellidae ...... 15 . (12) Penis cuticular tube lying free in elongated penial sheath ...... Ee aS RA ee Cronodrilus Penis muscular... 425 s5eee eee 14 . (13) Penis and penial sheath ever- sible; ejaculatory duct short, dilat- ed; bursa with pair of glands; sper- matheca bifid .......... Bdellodrilus Penis densely muscular; ejacula- tory duct short, with three layers of muscle; no bursal glands; sper- matheca without lateral branches 2, Le See Adenodrilus (12) Penis cuticular tube; one pair of testes in segment V Penis membraneous or muscular 16 . (15) Penis membraneous, eversi- ble, attached to inner wall of penial sheath by thin strands ... Cirrodrilus Penis muscular’... 2.22) See iv . (16) Penial sheath eversible; ejacu- latory duct short, ectal tip forming protruded penis; ental end of sper- miducal giand bifurcated LD SRL A Ses F Meee Ankyrodrilus Bursa asymmetrically spherical enclosing protrusible muscular pe- CeOLt DeoernO VOLUME 99, NUMBER 4 685 Fig. 3. Branchiobdella astaci: A, Lateral view of animal; B, Male efferent apparatus; C, Spermatheca; D, Jaws; E, Tip of everted penis. Abbreviations: b, bursa; m, male funnels; p, penis; sb, spermathecal bulb; sd, ectal duct of spermatheca; se, ental process of spermatheca; sg, spermiducal gland; sm, median duct of sper- matheca; ve, vas efferens; vd, vas deferens. (A, B, after Pop 1965:22; C, D, E, after Dorner 1864: plate 37.) Order Branchiobdellida Clitellate annelids with constant number of segments (15); without setae or prosto- mium; peristomium forming anterior suck- er; with posterior sucker; dorsal and ventral pharyngeal dental placoids; body divided into head of 4 segments, trunk of 11; anus dorsal, subterminal; 2 pairs of nephridia, asymmetrical pair in trunk segments I-IV, symmetrical pair in trunk segments VIII- IX; muscles non-syncytial, composed of cells with outer contractile and inner undiffer- entiated cytoplasm; testes in trunk segments V and VI or one of these segments only; paired male funnels and ducts in each tes- ticular segment; male efferent apparatus Opening through single pore on venter of trunk segment VI; a pair of ovaries and ovi- pores without oviducts and funnels in trunk segment VII; unpaired spermatheca, if pres- ent, in trunk segment V; clitellum on trunk segments VI and VII; Holarctic symbionts of freshwater crustaceans (modified from Holt 1965b:30-31). Family Branchiobdellidae Branchiobdellinae Goodnight, 1940:27 (in part). Diagnosis. —Two anterior nephridio- pores; vasa deferentia enter spermiducal gland ectad to latter’s ental end; no prostate. Genus Branchiobdella Fig. 3 Branchiobdella Odier, 1823. [Type species, by subsequent designation (Goodnight 1940:28), Branchiobdella astaci Odier 1823:75. Gender; feminine. ] Deposition of type specimens. —Un- known, presumably never designated. 686 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Cirrodrilus cirratus: A, Dorsal view; B, Male efferent apparatus; C, Spermatheca; D, Longitudinal section through bursa and penis. Abbreviations: ed, ejaculatory duct; others as for Fig. 3 (after Yamaguchi 1934, figs. 4, 5). Diagnosis. —One pair of testes in segment V; penis eversible, cuticular tube. Range. —Europe; Asia: China, Korea, Ja- pan. Number of species.—Pop (1965) recog- nized four species, one with three subspe- cies; Karaman (1970), six species, two with a subspecies each in Europe; seven east Asian species are accepted as valid by Gelder and Liu (in ms.). References. —Pierantoni 1906a, 1912; Stephenson 1930; Yamaguchi 1934; Good- night 1940; Georgevitch 1955, 1957; Pop 1965; Karaman 1970; Gelder and Liu (ms.). Note.—The literature pertaining to the genus Branchiobdella is confusing. The species currently assigned to it in both Eu- rope and Asia may well include more than one genus in both regions. Genus Cirrodrilus Fig. 4 Cirrodrilus Pierantoni, 1905:2. [Type species, by subsequent designation (Goodnight 1940:63), Cirrodrilus cirratus Pierantoni, 1905:1. Gender: masculine.] Ceratodrilus Yamaguchi, 1932a:361, 366. Carcinodrilus Yamaguchi, 1932b:62. VOLUME 99, NUMBER 4 Fig. 5. Ankyrodrilus koronaeus: A, Dorsal view; B, Longitudinal section through bursa and penis; C, Ventral view, reproductive systems; D, Jaws, upper to left. Abbreviations: sp, spermatheca; others as above (after Holt 1965a, figs. 6, 9, 1, 4, 5). Stephanodrilus (Stephanodrilus) Yamagu- chi, 1934:191-192. Stephanodrilus (Ceratodrilus) Yamaguchi, 1934:191-192. Cirrodrilus Goodnight, 1940:63; Holt, 1960a:54—-57, 1967b:3. Disposition of type specimens. —Un- known; some of Pierantoni’s material is de- posited in the Zoologische Museum at Hamburg (Holt 1967b:5). Diagnosis.—Two pairs of testes in seg- ments V and VI; penis membraneous, ever- sible, attached by strands to inner wall of penial sheath. [This genus is heterogeneous and should be subdivided. See Yamaguchi 1934:199-200.] Range. — Asia: Japan, Korea, China. Number of species. —Twenty-one species are accepted by Gelder and Liu (23, with the removal of C. truncatus (Liang, 1963) to a new genus, are proposed in ms. by these authors); other names are available. References. —Pierantoni 1905, 1906b, 1912; Yamaguchi 1932a, b, 1934; Good- night 1940; Holt 1960a, 1967b. Genus Ankyrodilus Fig. 5 Ankyrodrilus Holt, 1965a:10. [Type species, by original designation, Ankyrodrilus ko- ronaeus Holt, 1965a:10. Gender: mas- culine. ] Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Holt 1965a:11). Diagnosis. —Two pairs of testes; ental end of spermiducal gland bifurcated; penis mus- cular, penial sheath eversible; ejaculatory duct absent; mid-portion of body de- pressed. Range. —Eastern North America: south- west Virginia to middle Tennessee. Number of species. —Two. References. —Holt 1965a, Hobbs et al. 1967. 688 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON te on La D E Fig. 6. Xironogiton instabilis: A, Dorsal view; B, Ventral view, reproductive systems; C, Longitudinal section through bursa and penis; D, Upper jaw; E, Lower jaw. Abbreviations: af, atrial fold; others as above (after Holt 1974b; fig. 2). Genus Xironogiton Fig. 6 Xironogiton Ellis, 1919:247. [Type species, by original designation, Xironogiton ore- gonensis Ellis, 1919:248. Gender: mas- culine.] Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Ellis 1919:249). Diagnosis.—Two pairs of testes; penis, muscular, protrusible; bursa large, spherical to asymmetrical; spermiducal gland not bi- furcated entally; spermatheca small; mid- portion of body depressed. Range.—North America: higher eleva- tions of Appalachian uplands; north portion of Pacific versant. Number of species. — Five. References. —Ellis 1919; Goodnight 1940; Holt 1949, 1974b. Family Bdellodrilidae Cambarincolinae Goodnight, 1940:53 (in part). Diagnosis. —One anterior nephridiopore; vasa deferentia enter spermiducal gland ec- tad to its ental end; no prostate. Genus Bdellodrilus Fig. 7 Bdellodrilus Moore, 1895b:498. [Type species, by subsequent designation (Goodnight 1940:53) Brachiobdella illu- minata Moore, 1894:421. Gender: mas- culine. ] Disposition of type specimens. —Uncer- tain; Moore’s collections are now in those of the National Museum of Natural History, Smithsonian Institution. Diagnosis. —Penis eversible, epithelial (Moore 1895b:591, figs. 19, 20); bursa pro- vided with 2 lateral glands enclosed in its muscular wall; spermatheca with lateral process; body wall thin, glandular. Range. —North America: eastern United States, Mexico. Number of species. —One. References. — Moore 1895b, Hobbs et al. 1967; Holt 1973c. VOLUME 99, NUMBER 4 689 Fig. 7. Bdellodrilus illuminatus: A, Lateral view; B, Lateral view, reproductive systems; C, Lateral view of part of male efferent apparatus; D, Jaws. Abbreviations: bg, bursal gland; others as above (B, C, after Moore 1885b, figs. 19, 20; A, D, after Hobbs et al. 1967, fig. 13). Genus Cronodrilus Fig. 8 Cronodrilus Holt, 1968a:308. [Type species, by original designation, Cronodrilus ogygius Holt, 1968a:308. Gender: mas- culine. } Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Holt 1968a:308). Diagnosis. —Penis eversible, cuticular; penial sheath very long; ejaculatory duct short. Range. —North America: Georgia. Fig. 8. miducal gland; D, Ectal end of penis; E, Ental end of penis; F, Oblique view, upper jaw; G, Oblique view, lower jaw. Abbreviations: as above (after Holt 1968b, figs. 7, 8). Number of species. —One. Reference. —Holt 1968a. Genus Adenodrilus Fig. 9 Adenodrilus Holt, 1977b:727. [Type species, by original designation, Adenodrilus hemophagus Holt, 1977b:727.] Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Holt 1977b:727). Diagnosis.—Penis muscular, (?) eversi- ble; ejaculatory duct composed of 3 layers G Cronodrilus ogygius: A, Lateral view; B, Lateral view, reproductive systems; C, Lateral view, sper- 690 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 9. Adenodrilus hemophagus: A, Lateral view; B, Lateral view, reproductive systems; C, Longitudinal section through bursa and ejaculatory duct; D, Jaws. Abbreviations: as above (after Holt 1977b, figs. 1, 3, Se): eo rSes Fig. 10. Xironodrilus formosus: A, Dorsal view; B, Longitudinal section through bursa and penis; C, Upper jaw; D, Lower jaw; E, Ventral view, reproductive systems. Abbreviations: as above (original). VOLUME 99, NUMBER 4 Fig. 11. F, I, C, D). of muscles; spermiducal gland constricted at junction with vasa deferentia; body wall thin, glandular. Range. —North America: Oregon. Number of species. —One. Reference. —Holt 1977b. Family Xironodrilidae Cambarincolinae Goodnight, 1940:48 (in part). Diagnosis. —Two anterior nephridio- pores; two pairs of testes; vasa deferentia enter ental end of spermiducal gland; no prostate; penis muscular, protrusible. 691 C Caridinophila unidens: A, Lateral view; B, Ventrolateral view, male efferent apparatus; C, Same, lateral view, diagrammatic; D, Upper Jaw; E, Lower jaw. Abbreviations: as above (after Liang 1963, fig. 3A > Genus Xironodrilus Fig. 10 Xironodrilus Ellis, 1918:49. [Type species, by subsequent designation (Ellis 1919: 244), Xironodrilus formosus. Gender: masculine. ] Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Ellis 1919:244). Diagnosis.— As for the family; mid-por- tion of body depressed. Range. —North America: Michigan southeast to the Piedmont of North Caro- lina. 692 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 12. Ocedipodrilus oedipus: A, Lateral view; B, Longitudinal section through bursa and penis; C, Lateral view, reproductive systems; D, Tip of partially everted penis; E, Jaws, upper to left. Abbreviations: pr, prostate; others as above (A, C, E, after Holt 1967a; figs. 1, 4, 2, 3; B, D, after Holt 1984a, fig. 3). Number of species. — Five. References.—Moore 1894; Ellis 1919; Goodnight 1940, 1943; Holt and Weigl 1979. Family Caridinophilidae Diagnosis. —One anterior nephridiopore; 2 pairs of testes; vasa efferentia enter sub- spherical spermiducal gland separately (no vasa deferentia); no prostate; no sperma- theca. Genus Caridinophila Fig. 11 Caridinophila Liang, 1963:565, 569. [Type species, by original designation, Caridi- nophila unidens Liang, 1963:565. Gen- der: feminine. ] Disposition of type specimens. —Un- known. Diagnosis. —As for the family. Range. — Asia: Yunnan Province, China. Number of species. —One. Reference. —Liang 1963. Family Cambarincolidae Cambarincolinae, Goodnight, 1940 (in part). Diagnosis. —One anterior nephridiopore; 2 pairs of testes; vasa deferentia enter ental end of spermiducal gland. VOLUME 99, NUMBER 4 693 Fig. 13. Magmatodrilus obscurus: A, Lateral view; B, Longitudinal section through bursa and penis; C, Lateral view, male efferent apparatus; D, Jaws, upper to left. Abbreviations: as above (A, C, D, after Holt 1967b, figs. 3, 4, 1, 2; B, after Holt 1974a, fig. 4). Genus Oedipodrilus Fig. 12 Oedipodrilus Holt, 1967a:58. [Type species, by original designation, Oedipodrilus oedipus Holt, 1967a:58. Gender: mascu- line. ] Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Holt 1967a:58). Diagnosis. —Penis eversible, cuticular, provided with many small recurved hooks; prostate rudimentary to weil developed lobe of ental portion of spermiducal gland. Range. —North America: Kentucky, Tennessee, Mexico. Number of species. —Three. References. —Holt 1967a, 1984a. Genus Magmatodrilus Fig. 13 Magmatodrilus Holt, 1967b:3. [Type species, by original designation (Holt 1967b:3), Stephanodrilus obscurus Good- night, 1940:55. Gender: masculine.] Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Holt 1967b:3). Diagnosis. —No prostate, penis short, eversible; bursa large, atrial region elongat- ed; spermiducal gland long. 694 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON C Fig. 14. Triannulata magna: A, Lateral view; B, Laterodorsal view, reproductive systems; C, Longitudinal section through bursa and penis; D, Ventral jaw. Abbreviations: as above (after Holt 1974a, figs. 1, 2, 3). Range. —North America: California. Number of species. —One. References. —Goodnight 1940; Holt 1967b, 1974a. Genus Triannulata Fig. 14 Triannulata Goodnight, 1940:56. [Type species, by original designation, 7rian- nulata magna Goodnight, 1940:56. Gen- der: feminine.] Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Goodnight 1940:57). Diagnosis. —No prostate; no ejaculatory duct; penis long, muscular, eversible; atrial portion of bursa spherical; spermiducal gland short, subspherical; spermatheca short, muscular. Range. —North America: Oregon, Wash- ington. Number of species. —One. References. —Goodnight 1940, Holt 1974a. Genus Ceratodrilus Fig. 15 Ceratodrilus Hall, 1914:191. [Type species, by original designation, Ceratodrilus thy- sanosomus Hall, 1914:191. Gender: mas- culine.] VOLUME 99, NUMBER 4 695 oles Fig. 15. Ceratodrilus thysanosomus: A, Lateral view; B, Lateral view, reproductive systems; C, Longitudinal section through penial sheath and penis; D, Everted penis; E, Jaws, upper to left. Abbreviations: pp, prostatic protuberance; others as above (after Holt 1960a, figs. 11, 12, 6, 7, 15, 16). Stephanodrilus (Ceratodrilus) Yamaguchi, (Goodnight 1940:63), Cirrodrilus cirratus 1934:192. [Type species, by subsequent Pierantoni, 1905:2.] designation (Goodnight 1940:55), Ste- Ceratodrilus (Holt, 1960a:53-73). phanodrilus sapporensis Pierantoni, Disposition of type specimens. — 1906b:3.] “U.S.N.M. (Bureau of Animal Industry Cirrodrilus Pierantoni, 1905:2. [Type Helminthological Collection” (Hall 1914: species, by subsequent designation 191). 696 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 16. Ellisodrilus clitellatus: A, Lateral view; B, Male efferent apparatus; C, Longitudinal section through bursa; D, Jaws (after Holt 1960b, figs. 1, 2, 3, 4). Diagnosis. — Digitate projections on dor- References.—Hall 1914; Yamaguchi sum of segments II—-VII; peristomium ten- 1934; Goodnight 1940; Holt 1960a. taculated; prostatic bulb near ental end of spermiducal gland; penis membraneous, Genus Ellisodrilus eversible, attached by strands to inner wall Fig. 16 of penial sheath; bursa elongate. Ellisodrilus Holt, 1960b:170. [Type species, Range.—North America: Utah, Idaho, by original designation, Ellisodrilus cli- Oregon, Wyoming. tellatus Holt, 1960a:170. Gender: mas- Number of species. —Two. culine. ] loa a A aos D B C Fig. 17. Sathodrilus carolinensis: A, Lateral view; B, Lateral view, reproductive systems; C, Longitudinal section through bursa and penis; D, Jaws (after Holt 1968b, figs. 2, 1). VOLUME 99, NUMBER 4 Fig. 18. Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Holt 1960b:170). Diagnosis.—Prostate arises from mid- portion of spermiducal gland; penis reduced to pore on ectal surface of penial sheath; bursa asymmetrically subspherical; no sper- matheca. Range.—North America: Kentucky, In- diana, Michigan. Number of species. —Two. Reference. —Holt 1960b. 697 Tettodrilus friaufi. A, Lateral view; B, Lateral view, reproductive systems; C, Longitudinal section through bursa and penis; D, Jaws (after Holt 1968b, fig. 9). Genus Sathodrilus Fig. 17 Sathodrilus Holt, 1968b:294. [Type species, by original designation, Sathodrilus car- olinensis Holt, 1968b:294. Gender: mas- culine.] Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Holt 1968b:294). Diagnosis. —Prostatic protuberance, or prostate arising along mid-portion of sper- 698 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ES) ei D Fig. 19. Pterodrilus alcicornus: A, Lateral view; B, Lateral view, reproductive systems; C, Longitudinal section through bursa and penis; D, Jaws (A, B, from Holt 1968c, fig. 7; C, D, original). miducal gland or absent; penis membra- neous, attached by strands to inner wall of penial sheath; bursa usually elongate; dorsal body surface without digitate projections. Range. —North America. Number of species. — Fifteen. References. —Holt 1968b, 1973b, 1977a, 1978a, 1981b. Genus Tettodrilus Fig. 18 Tettodrilus Holt, 1968b:312. [Type species, by original designation, Tettodrilus friaufi Holt, 1968b:314. Gender: masculine.] Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Holt 1968b:312). Diagnosis. —Penis eversible, cuticular; ejaculatory duct projects as epithelio-mus- cular tube into atrium of bursa, enclosing ectal (when retracted) portion of penis; pros- tate arises at mid-portion of slender sper- miducal gland. Range. —North America: Tennessee. Number of species. —One. Reference. —Holt 1968b. Genus Pterodrilus Fig. 19 Pterodrilus Moore, 1895a:449. [Type species, by subsequent designation (Goodnight 1940:58), Pterodrilus alcicor- nus Moore, 1895a:449. Gender: mascu- line.] Disposition of type specimens.—(?) Lost (Holt 1968c:6). Diagnosis. — Anterior annulus of segment VIII with elevated dorsal ridge, those of oth- er segments often so, some or all such ridges often with fan-like digitate projections; small worms; prostate arising at mid-portion of short spermiducal gland; penis muscular, protrusible; bursa ovoid to pyriform. Range. —North America: eastern United States. Number of species. — Eight. References.—Moore 1895a; Ellis 1919; Goodnight 1940; Holt 1968c, 1973c. VOLUME 99, NUMBER 4 D E 699 Fig. 20. Cambarincola macrodontus: A, Lateral view; B, Lateral view, reproductive systems; C, Jaws (after Holt and Hoffman 1959, figs. 1, 2, 4, 3). Cambarincola illinoisensis: D, Longitudinal section through bursa and penis; E, Everted bursa, protruded penis (after Holt 1982, fig. 1D, E). Genus Cambarincola Fig. 20 Cambarincola Ellis, 1912:481. [Type species, by original designation, Cam- barincola macrodontus Ellis, 1912:481. Gender: masculine.]} Disposition of type specimens. — National Museum of Natural History, Smithsonian Institution (Ellis 1912:451). Diagnosis. — Prostate arising at ectal end (Gunction with ejaculatory duct) of spermi- ducal gland; penis muscular, protrusible; body segments often with raised anterior annuli (dorsal ridges), never with fan-like or digitate projections. Range. —North America; introduced into Japan. Number of species. — Forty-seven. References. —Ellis 1912, 1919; Good- night 1940; Hoffman 1963; Holt 1949, 1954, 1963, 1964, 1973b, c, d, e, 1974a, 1978a, 198la, 1982, 1984a, b; Leidy 1851. Acknowledgments I am grateful to Drs. James E. Carico, Brent D. Opell, and Stuart R. Gelder for reading the manuscript. Their comments have been most helpful. Dr. Susan E. H. West assisted with the preparation of Fig. 2. This paper is based in part on a paper presented at Louisville, Kentucky, on 27 December 1982, as part of a symposium “Crayfish Distribution Patterns,’ spon- sored by the Division of Invertebrate Zo- ology of the American Society of Zoologists and the Crustacean Society, in recognition of the seventieth birthday of Horton H. Hobbs, Jr. This paper is dedicated to Dr. Hobbs. Literature Cited Brinkhurst, Ralph O., and B. G. M. Jamieson. 1971. Aquatic Oligochaeta of the world. University of Toronto Press, Toronto and Buffalo, 860 pp. 700 Dorner, Herman. 1864. Ueber die Gattung Bran- chiobdella Odier. — Zeitschrift fiir Wissenschaft- liche Zoologie 14(3):464—493, pls. 36, 37. Ellis, Max M. 1912. A new discodrilid from Colo- rado.—Proceedings of the United States Na- tional Museum 42(1912):481—486, figs. 1-8. . 1918. Branchiobdellid worms from Michigan crayfishes.— Transactions of the American Mi- croscopical Society 37(1):49-S1. 1919. The branchiobdellid worms in the col- lections of the United States National Museum, with descriptions of new genera and new species. — Proceedings of the United States Na- tional Museum 55 (2267):241-265, pls. 10-13. Freeman, Carl J. 1963. An anatomical study of the posterior nephridial systems of four species of the Branchiobdellidae. Master’s Thesis, Virgin- ia Polytechnic Institute, pp. 1-25, figs. 1-2. Georgevitch, Jivoin. 1955. Sur les branchiobdellides des écrevisses du Lac Dojran.— Acta Macedoni- ci Scientiarum Naturalium II (10/21):199-221, figs. 1-33. 1957. Les branchiobdellides de Yugosla- vie. — Bulletin de l’ Académie Serbe des Sciences 18(5):5—23, figs. 1-35. Goodnight, Clarence J. 1940. The Branchiobdellidae (Oligochaeta) of North American crayfishes. — Illinois Biological Monographs 17(3):1-75, pls. 1-3. 1943. Report on a collection of branchiob- dellids.— Journal of Parasitology 29(2):100—-102, figs. 1-5. Hall, Maurice C. 1914. Description of a new genus and species of the discodrilid worms.—Pro- ceedings of the United States National Museum 48(2071):187-193, figs. 1-3. Hobbs, Horton H., Jr. 1974. Synopsis of the families and genera of crayfishes (Crustacea: Decapo- da).—Smithsonian Contributions to Zoology 164:i-111, 1-32, figs. 1-27. . 1981. The crayfishes of Georgia.—Smithson- ian Contributions to Zoology 318:i—viii, 1-549, figs. 1-194. , Perry C. Holt, and Margaret Walton. 1967. The crayfishes and their epizoGtic ostracod and branchiobdellid associates of the Mountain Lake, Virginia, region.—Proceedings of the United States National Museum 123(3602):47-74, figs. 12-22. , and Alejandro Villalobos. 1958. The exo- skeleton of a freshwater crab as a microhabitat for several invertebrates.— Virginia Journal of Science 9, new series (4):395-396. 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 1|14(3470:271-371, figs. 1-79. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Holt, Perry C. 1949. A comparative study of the reproductive systems of Xironogiton instabilius instabilius Moore and Cambarincola philadel- phica Leidy (Annelida, Oligochaeta, Branchiob- dellidae).—Journal of Morphology 84(3):535- 572, pls. 1-4. 1953. Characters of systematic importance in the family Branchiobdellidae (Oligochae- ta).— Virginia Journal of Science 4, new series (2):57-61. 1954. A new branchiobdellid of the genus Cambarincola Ellis (Oligochaeta, Branchiob- dellidae) from Virginia.— Virginia Journal of Science 5, new series (3):168—172, figs. 1—S. 1960a. The genus Ceratodrilus Hall (Bran- chiobdellidae, Oligochaeta) with the description of a new species.— Virginia Journal of Science 11, new series (2):53-73, pls. 1-4. 1960b. On a new genus of the family Bran- chiobdellidae (Oligochaeta).—American Mid- land Naturalist 64(1):169-176, figs. 1-6. 1963. A new branchiobdellid (Branchiob- dellidae: Cambarincola).—Journal of the Ten- nessee Academy of Science 38(3):97—100, figs. 1-6. . 1964. A new branchiobdellid (Annelida) from Costa Rica.—Tulane Studies in Zoology 12(1): 1-4, figs. 1-4. . 1965a. On Ankyrodrilus, a new genus of bran- chiobdellid worms.— Virginia Journal of Sci- ence 16, new series (1):9-21, figs. 1-14. 1965b. The systematic position of the Bran- chiobdellidae (Annelida: Clitellata).—System- atic Zoology 4(1):25-32, figs. 1-3. . 1967a. Oedipodrilus oedipus, n.g., n. sp. (An- nelida: Clitellata: Branchiobdellida).—Trans- actions of the American Microscopical Society 86(1):58—60, figs. 1-4. 1967b. Status of the genera Branchiobdella and Stephanodrilus in North America with the description of a new genus (Clitellata: Bran- chiobdellida).— Proceedings of the United States National Museum 124(3631):1—10, figs. 1-4. 1968a. The Branchiobdellida: Epizootic an- nelids.— The Biologist 50(3—4):79—94, figs. 1—2. . 1968b. New genera and species of branchiob- dellid worms (Annelida: Clitellata).— Proceed- ings of the Biological Society of Washington 81: 291-318, figs. 1-9. 1968c. The genus Pterodrilus (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 I: Invertebrates.— Virginia Polytechnic Institute, VOLUME 99, NUMBER 4 Blacksburg Research Division Monograph 1, 191-219, figs. 1-10. 1973a. A free-living branchiobdellid (An- nelida: Clitellata)?— Transactions of the Amer- ican Microscopical Society 92(1):152-153. 1973b. Epigean branchiobdellids (Annelida: Clitellata) from Florida.— Proceedings of the Bi- ological Society of Washington 86(7):79-104, figs. 1-8. 1973c. A summary of the branchiobdellid (Annelida: Clitellata) fauna of Mesoamerica. — Smithsonian Contributions to Zoology 142:- iii, 1—40, figs. 1-19. . 1973d. An emended description of Cambar- incola meyeri Goodnight (Clitellata: Branchiob- dellida).— Transactions of the American Micro- scopical Society 92(4):677-682, figs. 1-5. 1973e. Branchiobdellids (Annelida: Clitel- lata) from some eastern North American caves, with descriptions of new species of the genus Cambarincola.—International Journal of Spe- leology 5:219-256. 1974a. An emendation of the genus Trian- nulata Goodnight, 1946, with the assignment of Triannulata montana to Cambarincola Ellis, 1912 (Clitellata: Branchiobdellida).— Proceed- ings of the Biological Society of Washington 87(8):57—72, figs. 1-5. 1974b. The genus Xironogiton Ellis, 1919 (Clitellata: Branchiobdellida).— Virginia Jour- nal of Science 25(1):5-19, figs. 1-6. 1977a. An emendation of the genus Sath- odrilus Holt, 1968 (Annelida: Branchiobdelli- da), with the description of four new species from the Pacific drainage of North America. — Proceedings of the Biological Society of Wash- ington 90(1):116-131, figs. 1-7. . 1977b. A gill-inhabiting new genus and species of the Branchiobdellida (Annelida: Clitellata). — Proceedings of the Biological Society of Wash- ington 90(3):726-734, figs. 1-5. 1978a. The reassignment of Cambarincola elevatus Goodnight, 1940 (Clitellata: Bran- chiobdellida) to the genus Sathodrilus Holt, 1968.— Proceedings of the Biological Society of Washington 91(2):472—482, figs. 1-4. 1978b. Key to genera of Branchiobdellida. In Robert W. Pennak, Freshwater invertebrates of the United States. John Wiley and Sons, New York, 803 pp. . 1981a. Aresumé of the members of the genus Cambarincola (Annelida: Branchiobdellida) from the Pacific drainage of the United States. — Proceedings of the Biological Society of Wash- ington 94(3):675—695, figs. 1-5. . 1981b. New species of Sathodrilus Holt, 1968 (Clitellata: Branchiobdellida) from the Pacific drainage of the United States, with the synon- 701 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. . 1984a. On some branchiobdellids (Annelida: Clitellata) from Mexico with the description of new species of the genera Cambarincola and Oedipodrilus.—Proceedings of the Biological Society of Washington 97(1):35—42, figs. 1-3. 1984b. A new species of the genus Cambar- incola (Clitellata: Branchiobdellida) from Cali- fornia.— Proceedings of the Biological Society of Washington 97(3):544—549, figs. 1-2. , and Richard L. Hoffman. 1959. An emended description of Cambarincola macrodonta Ellis, with remarks on the diagnostic characters of the genus (Oligochaeta: Branchiobdellidae).—Jour- nal of the Tennessee Academy of Science 34(2): 97-104, figs. 1-6. — ., and Ann M. Weigl. 1979. A new species of Xironodrilus Ellis, 1918 from North Carolina (Clitellata: Branchiobdellida).— Brimleyana 1(1): 23-29, figs. 1-2. Karaman, S. Mladen. 1970. Beitrag zur Kenntnis der europdischen Branchiobdelliden (Clitellata, Branchiobdelloidea).— Internationale Revue der Gasamten Hydrobiologie 55(3):325-333, figs. 1-18. Leidy, Joseph. 1851. Contributions to helmintholo- gy.—Proceedings of the Academy of Natural Science, Philadelphia 5:205-209. Liang Yan-lin. 1963. Studies on aquatic Oligochaeta of China. 1. Descriptions of new naids and bran- chiobdellids.— Acta Zoologica Sinica 15(4):560- 570, figs. 1-4. [Chinese with English summary] Liu Sze-cheng. 1964. A second report of Branchiob- dellidae in Liaoning Province with descriptions of three new species.—Acta Zoologica Sinica 16(4):602-610, figs. 1-7. [Chinese with English summary] . 1984. Descriptions of two new species of the genus Stephanodrilus from north east China and notes on St. truncatus Liang from Guangdong Province (Oligochaeta: Branchiobdellidae).—Acta Zootaxonomica Sinica 9(4):351-355, figs. 1-2. [Chinese with English summary] , and Chang Chen-an. 1964. On four new species of Branchiobdellidae from crayfish Cambaroides dauricus (Pallas).— Acta Zoologi- ca Sinica 16(1):33-38, figs. 1-6. [Chinese with English summary] , and Zhang De-cai. 1983. Three new species of the genus Branchiobdella (Oligochaeta: Bran- 702 chiobdellidae) from China.—Acta Zootaxono- mica Sinica 8(3):246—-251, figs. 1-3. [Chinese with English summary] Moore, J. Percy. 1894. On some leech-like parasites of American crayfishes.—Proceedings of the Academy of Natural Sciences of Philadelphia (for 1893):419-—428, pl. 12. 1895a. Pterodrilus, a remarkable discodri- lid.— Proceedings of the Academy of Natural Sciences of Philadelphia, 1894, 3:449-454. 1895b. The anatomy of Bdellodrilus illumi- natus, an American discodrilid.—Journal of Morphology 10(2):497—540, pls. 28-32. 1897. On the structure of the discodrilid ne- phridium.—Journal of Morphology 13(3):327- 372, pls. 20-23. Odier, Auguste. 1823. Mémoire sur le branchiob- delle. — Mémoires de la Société d’Histoire Na- turelle de Paris 1:71-78. Pennak, Robert W. 1978. Freshwater invertebrates of the United States. John Wiley and Sons, New York, 803 pp. Pierantoni, Umberto. 1905. Cirrodrilus cirratus, n.g. n.sp. parassita dell’ Astacus japonicus. —Annu- ario del Museo Zoologica della R. Universita di Napoli (Nuova serie) 1:1—3, pls. 1-6. 1906a. Observazioni sul genere Branchiob- della Odier.—Annuario del Museo Zoologico della R. Universita di Napoli (Nuova serie) 2(4): 1-9, pl. 2. . 1906b. Nuovi discodrilid del Giappone e del- la California.—Annuario del Museo Zoologico della R. Universita di Napoli (Nuova serie) 2(11):1-9, pl. 5. 1912. Monografia dei Discodrilidae.—An- nuario del Museo Zoologico della R. Universita di Napoli 3(24):1—27, pl. 5, figs. 1-14. Pop, Victor. 1965. Systematische Revision der eu- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ropaischen Branchiobdelliden (Oligochaeta). — Zoologischer Jahrbuch, Systematik 92:219-238, figs. 1-11. Stephenson, J. 1930. The Oligochaeta. Oxford, Clar- endon Press, i-xi + 978 pp., 242 figs. Weigl, Ann M. 1980. The structure of the epidermis of three species of American branchiobdellids (Annelida: Branchiobdellida): An ultrastruc- tural and histochemical study. Ph.D. disserta- tion, Wake Forest University, i-xvi + 116 pp., 66 figs. Yamaguchi, Hideji. 1932a. On the genus Cirrodrilus Pierantoni, 1905, with a description of a new branchiobdellid from Japan.—Annotationes Zoologicae Japonensis 13(4):36 1-367, figs. 1-3. . 1932b. Description ofa new branchiobdellid, Carcinodrilus nipponicus, n.g. et n.sp.—Journal of the Faculty of Science, Hokkaido Imperial University, Series VI (Zoology) 2(1):6 1-67, figs. 1-2, pl. 3. 1933. Description of a new branchiobdellid, Cambarincola okadai, n.sp., parasitic on Amer- ican crayfish transferred into a Japanese lake. — Proceedings of the Imperial Academy (of Japan) 9:191-193, figs. 1-2. . 1934. Studies on Japanese Branchiobdellidae with some revisions on the classification.— Journal of the Faculty of Science, Hokkaido Im- perial University, Series VI, 3(3):176—219, figs. 1-17, pls. 12-13. 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. 99(4), 1986, pp. 703-708 OCCURRENCE OF LARVAL JLLEX ILLECEBROSUS AND OTHER YOUNG CEPHALOPODS IN THE SLOPE WATER/GULF STREAM INTERFACE Michael Vecchione and Clyde F. E. Roper Abstract. —Specimens and data suggest that larval short-finned squid, J/lex illecebrosus, may concentrate in water where density (c,) is approximately 26.7, indicating that spawning occurs in the subsurface interface where Slope Water mixes with the Gulf Stream. Because these water masses occur year-round within the range of J. i/lecebrosus, spawning may be a prolonged or progressive process along the shoreward edge of the Gulf Stream. The distribution of other species indicates mesoscale spatial separation of young cephalopods in the vicinity of the Gulf Stream. Among many unanswered questions in cephalopod ecology, two currently are re- ceiving much attention: (1) what are the sea- son and site(s) of spawning for the com- mercially important species I//ex illecebrosus and (2) what factors in the early life history of the species are crucial to its survival and recruitment to the fishery? A multinational effort was organized by Canada, Japan, and the United States to attempt to answer these questions. Inten- sive directed sampling for the early stages of I. illecebrosus was conducted during Feb- ruary and March 1982 on cruise 8201 of the Japanese Research Vessel Kaiyo-Maru. Per- sonnel from all three sponsoring nations participated in the two-part cruise. The area sampled was centered on the Gulf Stream east of New England and south of Nova Scotia. During the second leg of the cruise, from which our specimens were obtained, the Gulf Stream was meandering through the sampling area. Thus, the 40 sta- tions for which we have data include water masses characteristic of Boreal Slope Water, the Gulf Stream, and the Sargasso Sea. Materials and Methods Sampling was done primarily with oblique tows. A model 1271 open bongo-net system with 61 cm diameter mouth opening and 0.5 mm mesh was used to sample between the surface and 200 m depth. Sampling be- tween the surface and 1000 m depth was done with an open midwater trawl (KMT; 100 m long, 18 mm mesh cod-end), and a rectangular midwater trawl (KYMT; 3 x 3 m mouth opening, 17 m long, 1.7 mm mesh cod-end). Data on hydrographic conditions were collected using expendable bathyther- mographs and multiple bottle casts for con- ductivity, temperature, and dissolved oxy- gen. The biological samples were sorted at sea and field identifications were attempted for larval and juvenile ///ex, sometimes under rather severe weather conditions. Preserved specimens then were divided among the representatives of the participating coun- tries: 50% to the Japanese, 25% to the Ca- nadian, and 25% to the U.S. participants. Results The material available for analysis con- sisted of 66 specimens that were identified to 18 taxa. Young J/lex (we believe J. ille- cebrosus; see Vecchione 1979) was the most abundant cephalopod collected (Table 1). The material also included several other taxa for which our current knowledge of early 704 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Ranking by abundance of cephalopod specimens from Leg II of Kaiyo-Maru cruise 8201.* Mantle length (mm) Taxon No. of specimens Rank Min Median Max. Illex sp. (cf. illecebrosus) 16 l 3.0 4.0 10.7 Octopodidae 6 3 3.7 4.0 6.1 Gonatus fabricti 6 3 Dyed] 7.0 13.3 Ommastrephes sp. 6 3 Dall 3.8 8.1 Leachia sp. 4 5.5 14.7 17.9 28.0 Pterygioteuthis sp. 4 5.5 3). 4.6 10.0 Abralia sp. 3 8 2.8 4.3 4.7 Octopoteuthis sp. 3 8 3.1 6.3 6.8 Teuthowenia megalops 3 8 4.3 5.0 5.7 Pyroteuthis margaretifera 2 12.5 Dol 3.3 3.9 Onychoteuthis banksii 2 12.5 3.8 4.8 5.8 Onykia carribaea 2 12.5 6.7 7.0 7.3 Ctenopteryx sicula 2 12.5 2.8 3.0 3.1 Brachioteuthis sp. 2) OES Del DS 2.8 unid. oegopsid 2 12.5 2.3 2 3.1 Abraliopsis sp. 1 16 8.2 8.2 8.2 Histioteuthis sp. 1 16 Dell D7] Ded Ornithoteuthis antillarum (?) 1 16 2.6 2.6 2.6 Total 66 specimens 18 taxa * Represents 25% of total collected; see text. life history is even more limited. These taxa include young Gonatus fabricii, as well as several other ommastrephids, onychoteu- thids, enoploteuthids, cranchiids, and oc- topodids. Station locations and isotherms at the surface and at 200 m depth are shown in Fig. 1. These depths were selected because they represent the sampling envelope of the oblique bongo tows. The temperature struc- ture shows the turbulent nature of the cur- rent in the sampling area, as is typical of the transition zone between the Gulf Stream and the North Atlantic Drift. It also shows the substantial variability in thermal structure within the sampled depth range. Distribu- tion of the five most abundant taxa is shown also in Fig. 1. Based on this limited subset of the complete set of data, we see indica- tions of mesoscale spatial segregation among these taxa. Whereas Gonatus fabricii was collected only along the Slope-Water edge of the system, J//ex illecebrosus and the un- identified octopods were concentrated along the central axis of the system, and Leachia sp. and Ommastrephes sp. appeared to be concentrated on the Sargasso Sea side of the system. The six stations at which J//ex were col- lected included a broad range of surface temperatures (Fig. 2A). Inferences on the distribution of young J//ex based only on surface conditions where oblique tows col- lected specimens would indicate eurytopic habitat requirements, from 8°C to nearly 20°C. However, based on our earlier work with sampling programs from shelf and slope waters of the western North Atlantic (Roper and Lu 1979, Vecchione 1979), we have reason to believe that larval J. i/lecebrosus are concentrated deeper in the water col- umn. At approximately 100 m depth all of the temperature sections begin to converge and by 150 m the temperatures concentrate around 13°-18°C. If we assume, based on earlier studies, that this is close to the depth at which most of the ///ex were collected, then we have reason to point out strong similarities among all of these stations. The temperatures and salinities of these VOLUME 99, NUMBER 4 705 KAIYO- MARU CRUISE 82-0OI - 3 MAR 82 l2 FEB Fig. 1. +39°N xX ILLEX ILLECEBROSUS O OMMASTREPHES SP. 4 GONATUS SP. O LEACHIA SP. + UNID. OCTOPOD ¢ PHYSICAL DATA ONLY — SURFACE —--—-- 200m ISOTHERM ISOTHERM Study area showing station locations, surface and 200 m isotherms, and occurrence of the five most abundant cephalopod taxa (Kaiyo-Maru cruise 8201, Feb—Mar 1982). stations at 150 and 200 m depths are shown in Fig. 2B. These waters are approximately isopycnal at o, of about 26.7, an indication of subsurface mixing. The isopycnal mixing extends between what Wright and Parker (1976) refer to as the “slope-water ther- mostad”’ and the Gulf Stream. Some cap- tures are in the discrete water types and some are in the intermediate water; the aggregate of the captures is what would be expected in subsurface mixing between two water types because it extends along isopycnal lines. Based on these indications, we believe that hatching probably occurs in subsurface waters at the interface between the Slope Water and the Gulf Stream. Since both of these water masses occur within the distri- butional range of J. i/lecebrosus throughout the year, these data may also indicate a pro- longed spawning season along the shore- ward edge of the Gulf Stream over a very extensive latitudinal range. Discussion and Recommendations Prior to O’Dor and Durward’s (1979) demonstration that J/lex illecebrosus spawns pelagic egg masses, it was assumed that I/lex probably spawned demersally on the con- tinental slope (Roper and Lu 1979, Vec- chione 1979). The concept that an obligate relationship exists between the Gulf Stream and the early life history of J//ex is now gaining support (Coelho 1985, O’Dor and 706 Illex illecebrosus STATIONS KAIYO- MARU CRUISE 820] TEMPERATURE (°C) O05) 10015) 20)es ) 100 E 200 x la a WwW Q 300 SALINITY (x 10) 200 37.0 ° 500 rd > & x WW a = ry) B = Fig. 2. A, Temperature profiles at Kaiyo-Maru sta- tions where I/lex illecebrosus larvae were captured (station numbers indicated); B, T-S diagram of Kaiyo- Maru stations where I/lex illecebrosus larvae were cap- tured with notation of water-mass type (cruise 8201, Feb—Mar 1982). Balch 1985, Rowell et al. 1985, Hatanaka etal. 1985). Additionally, our data from this cruise and unpublished observations from other cruises (Vecchione et al. 1986) indi- cate that similar relationships may exist for other cephalopod species. Hypotheses are now being proposed to explain this relationship for J. i/lecebrosus, but conclusive tests of these hypotheses are very difficult because of problems with sam- pling and taxonomy. Not only has it so far been impossible to collect [//ex eggs in the field (O’Dor and Balch 1985) and difficult to collect planktonic young, but specific identification of the young must be inferred from adult distributions. South of Cape Hatteras such identifications are confound- ed by the presence of a complex of three species (J. i/lecebrosus, I. coindetii, and I. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON oxygonius) which even as adults are very similar morphologically. Thus, distribu- tional patterns of larvae collected on cruises in or near the area of species overlap (e.g., Rowell et al. 1985) may result from spawn- ing by one or more of these species. Often, inferences on larval ecology have been based on surface conditions at stations where the larvae have been collected in oblique subsurface tows. This practice is particularly faulty for a species like J/lex illecebrosus for which data on vertical dis- tribution are extremely limited. For in- stance, assumptions like that of Rowell et al. (1985) that “‘it is likely that the majority [of I//ex larvae] were taken in the upper 50 m”’ of oblique tows between the surface and 200-300 m depth are not supported by data. Observations on J. illecebrosus egg masses by O’Dor and Balch (1985) indicate that the egg masses in nature are probably suspend- ed in midwater at about the depth of the pycnocline (about 150 m in our samples). Furthermore, O’Dor et al. (1986) have shown that laboratory-hatched J. illecebro- sus larvae avoid the surface but are capable of diel vertical migration from mesopelagic depths. This is consistent with the very lim- ited observations by Hatanaka et al. (1985) of a diel shift in the depth of maximum abundance of larval I/lex. An analysis of the distribution of J//ex illecebrosus collected off Nova Scotia during Kaiyo-Maru cruise 8201 has been compiled by Hatanaka et al. (1985). This report pre- sumably was based on field-log entries of identifications for all specimens, since sev- eral specimens remained in our possession as a result of subdividing the catch upon completion of the cruise. During the first leg of the cruise, discrete-depth bongo tows were attempted at four stations. These attempts were successful at only one station in col- lecting a complete set of samples (from sev- eral depth strata between the surface and 200 m depth). Subsequently, discrete-depth sampling was abandoned. Inferences by Ha- tanaka et al. (1985) on J//ex larval distri- VOLUME 99, NUMBER 4 bution were therefore based on physical conditions in the 0-100 m depth range at oblique-tow stations (however, their very limited discrete-depth data indicated that larval J//ex may be concentrated at 100 m or greater depth during the daytime). Infer- ences by Hatanaka et al. (1985) about the relationship between J//ex larval distribu- tion and the boundary between the Gulf Stream and Slope Water do not contradict our impressions which are based on reex- amination of a subset of the collections, but we do feel, however, that some deficiencies in the data must be pointed out so that sim- ilar problems can be avoided in the future. Although we are able to speculate about the larval occurrence and distribution of J. illecebrosus and other species, a more pre- cisely defined sampling program probably would have provided sufficient data to test our hypothesis. To achieve this, we rec- ommend several changes in future sampling programs. First, once the general area and season of larval abundance have been de- termined for the target species, discrete- depth sampling should be considered a re- quirement for the study of early life history. Such a sampling regime can be accom- plished either with bongo systems or with multiple opening/closing systems such as the MOCNESS Multiple Opening-Closing Net and Environmental Sensing System (Weibe et al. 1976). If time and funding constraints are imposed on the number of samples that can be collected and processed, then the number of stations occupied should be re- duced to ensure that discrete-depth sam- pling can be accomplished. Second, defini- tive sorting and verification of species-level identifications should not be attempted at sea. Our cross-checks of the field logs against subsequent laboratory identifications indi- cate that several of the field identifications were incorrect. Even under the best of con- ditions use ofa microscope at sea is difficult. Hatchlings of only about 2 mm or less in length are easily overlooked sloshing about in a petri dish of plankton; furthermore, the 707 minute taxonomic characters required for identification are difficult to distinguish, let alone confirm. Thus, we cannot be confident enough in the shared, complete set of field-log entries of identifications to infer distribution and abundance based on all specimens collected but subdivided prior to verification of identifications. A complete set of samples from this cruise, examined by a specialist in cephalopod sys- tematics, would have been of considerable value in fulfilling the goals of this study. An additional benefit of onshore examination of all specimens by a single specialist is that the distribution of non-target species, such as Gonatus fabricii, could confidently be as- sessed along with the distribution of larval Illex. We very strongly support international collaborative studies and coauthorship as an effective, efficient means of helping to answer many of the important questions we face in biological oceanography, fisheries bi- ology, and systematics. To help achieve these ends, we recommend that future collections be identified and analyzed at one institu- tion, prior to dividing the material among participating organizations. This will ensure consistency of analysis and interpretation without diminishing the collaborative ef- fort. Acknowledgments This paper is a modification of one pre- sented at the North Atlantic Fisheries Or- ganization Special Session on Squid during 5-7 September 1984, in Dartmouth, Nova Scotia. We wish most heartily to thank Mi- chael J. Sweeney, National Museum of Nat- ural History, Smithsonian Institution, who participated in Leg II of Kaiyo-Maru cruise 8201 and collected the specimens discussed here. We very much appreciate being in- vited by the U.S. National Marine Fisheries Service to participate in the cruise and to study the material; we especially thank Anne T. Lange. The officers, crew, and scientific party of the Kaiyo-Maru are gratefully ac- 708 knowledged for their hospitality and co- operation during the cruise. We further ap- preciate reviews of this paper by John Wormuth, Texas A&M University, T. E. Bowman, Smithsonian Institution, and two anonymous reviewers. Literature Cited Coelho, M.L. 1985. Review of the influence of ocean- ographic factors on cephalopod distribution and life cycles.—Northwest Atlantic Fisheries Or- ganization Scientific Council Studies 9:47—57. Hatanaka, H., A. T. Lange, and T. Amaratunga. 1985. Geographical and vertical distribution of short- finned squid (J//lex illecebrosus) larvae in the northwest Atlantic.— Northwest Atlantic Fish- eries Organization Scientific Council Studies 9: 93-99. O’Dor, R. K., and N. Balch. 1985. Properties of J//ex illecebrosus egg masses potentially influencing larval oceanographic distribution.— Northwest Atlantic Fisheries Organization Scientific Coun- cil Studies 9:69-76. , and R. D. Durward. 1979. A preliminary note on J/lex illecebrosus larvae hatched from eggs spawned in captivity.— Proceedings of the Biological Society of Washington 91(4):1076- 1078. , E. A. Foy, P. L. Helm, and N. Balch. 1986. Locomotion and energetics of hatchling squid, Illex illecebrosus.—American Malacological Bulletin 4:55-60. Rowell, T. W., R. W. Trites, and E. G. Dawe. 1985. Distribution of short-finned squid (I/lex illece- brosus) larvae and juveniles in relation to the Gulf Stream frontal zone between Florida and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cape Hatteras.—Northwest Atlantic Fisheries Organization Scientific Council Studies 9:77— 92. Roper, C. F. E.,andC.C. Lu. 1979. Rhynchoteuthion larvae of ommastrephid squids of the western North Atlantic, with the first description of lar- vae and juveniles of J/lex illecebrosus.—Pro- ceedings of the Biological Society of Washington 91(4):1039-1059. Vecchione, M. 1979. Larval development of J//ex Steenstrup, 1880 in the northwestern Atlantic with comments on ///ex larval distribution. — Proceedings of the Biological Society of Wash- ington 91(4):1060-1075. , C. F. E. Roper, C. C. Lu, and M. J. Sweeney. 1986. Distribution and relative abundance of planktonic cephalopods in the western North Atlantic.—American Malacological Bulletin 4: 101. Weibe, P. H., K. H. Burt, S. H. Boyd, and A. W. Morton. 1976. A multiple opening/closing net and environmental sensing system for sampling zooplankton. — Journal of Marine Research 34: 313-326. Wright, W. R., and C. F. Parker. 1976. A volumetric temperature/salinity census for the Middle At- lantic Bight.—Limnology and Oceanography 21(4):563-571. (MV) Department of Biological and En- vironmental Sciences, McNeese State Uni- versity, Lake Charles, Louisiana 70609, U.S.A.; (CFER) Department of Inverte- brate Zoology—Mollusks, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560, U.S.A. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 709-716 ASTYANAX SCOLOGASTER, A NEW CHARACID (PISCES: OSTARIOPHYSI) FROM THE RIO NEGRO, SOUTH AMERICA Marilyn J. Weitzman and Richard P. Vari Abstract. —Astyanax scologaster is described from a series of localities in the middle and upper Rio Negro of Venezuela and Brazil. The presence of exserted anterior spines of the pelvic bones serve to define a subunit of Astyanax con- sisting of the relatively small-sized species A. scologaster, A. essequibensis, A. mucronatus, and A. zonatus. The extensive development of the exserted pelvic spine, large eye diameter, branched anal-fin ray count, number of pored lateral- line scales, and pigmentation pattern of Astyanax scologaster delimit that species within this assemblage. The presence of exserted strong spine-like pelvic bones which project anteriorly from the anteroventral body wall was first noted for characids by Eigenmann (1911:180, and Fig. 2) in his description of Deuterodon acanthogaster. In that species account Ei- genmann noted a similar condition for As- tyanax mucronatus Eigenmann, although no mention of the modification had been made in the original description (Eigenmann 1909: 19-20) and subsequent treatments of that species until 1921. In that year, Eigenmann (p. 280) inserted “‘... innominate bones protruding as spines in front” in his rede- scription of Astyanax mucronatus. Recent collecting efforts in the Rio Negro basin of Venezuela and Brazil have revealed the presence of an undescribed Astyanax species with very well developed anteriorly exserted pelvic bones. Comparative studies associ- ated with the description of the new species have uncovered the presence of comparable spines in individuals of three additional As- tyanax species and in a second Deuterodon species. Materials and methods. —Counts and measurements in the description follow the methods outlined in Fink and Weitzman (1974). All measurements are given as per- centages of standard length (SL) except for subunits of the head which are presented as percentages of head length (HL). Vertebral counts were taken from radiographs and cleared and counterstained specimens. This number includes the four vertebrae incor- porated into the Weberian apparatus and considers the fused PU, + U, as a single element. In the meristic values detailed, the range of the holotype and measured para- types is presented first, with the value for the holotype indicated in brackets. The following abbreviations are used for institutions: AMNH, American Museum of Natural History, New York; ANSP, Acad- emy of Natural Sciences of Philadelphia; BMNH British Museum (Natural History), London; CAS, California Academy of Sci- ences, San Francisco; FMNH, Field Mu- seum of Natural History, Chicago; MBUCV, Museo de Biologia, Instituto de Zoologia Tropical, Universidad Central de Venezue- la, Caracas; MZUSP, Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo; and USNM, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. Astyanax scologaster, new species Figs. 1 and 3, Table 1 Holotype. —MBUCV V-15249, 37.5 mm SL, Venezuela, Territorio Federal Amazo- nas, Departamento Rio Negro, lower por- 710 Fig. 1. tion of Cano Manu, which drains into the Rio Casiquiare about 250 m upstream of Solano (approx. 02°00'N, 66°57’W); coll. R. P. Vari, C. J. Ferraris, Jr., O. Castillo, and J. Fernandez, 7 Dec 1984. Paratypes.—(All from Venezuela, Terri- torio Federal Amazonas, Departamento Rio Negro, unless noted otherwise): MBUCV V-15250, 5, 28.8-37.2 mm SL; USNM 272612, 8, 24.5—36.2 mm SL (4, cleared and counterstained for bone and cartilage); AMNH 56173, 3, 27.5—30.0 mm SL; ANSP 157596, 3, 26.7-28.6 mm SL; BMNH 1985.10.14:1-3, 3, 26.9-30.0 mm SL; CAS 57471, 3, 24.9-35.0 mm SL; FMNH 96616, 3, 26.5-29.6 mm SL; MZUSP 36288, 3, 24.7-29.9 mm SL; taken with the holo- type. —AMNH 56174, 5, 38.7-40.7 mm SL; USNM 276503, 5, 39.6-42.0 mm SL; MBUCV V-15251, 5, 38.0—-40.6 mm SL, Rio Negro at San Carlos de Rio Negro, 2 Feb 1984.—USNM 272609, 4, 35.3-36.0 mm SL, Cano Chola, where crossed by road from San Carlos de Rio Negro to Solano (01°58’N, 67°00'’W), 5 Dec 1984.—USNM 272611, 5, 29.6-33.0 mm SL, Rio Negro, one-half hour upstream of San Carlos de Rio Negro, 4 Dec 1984.—USNM 272610, 5, 32.2-36.0 mm SL, San Carlos de Rio PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Astyanax scologaster, new species, holotype, MBUCV V-15249, 37.5 mm SL, Venezuela, Cano Manu. Negro, Rio Negro margin upstream of town landing, 3 Dec 1984.—MZUSP 30252, 6, 31.8-38.9 mm SL; USNM 276505, 6, 37.0- 41.4 mm SL, Brazil, Amazonas, Rio Negro, Massarabi, sandy beach just above Barcelos (approx. 00°57'S, 62°56’W), 18 Oct 1979.— MZUSP 30247, 6, 32.7-37.3 mm SL; USNM 276504, 5, 35.7-39.8 mm SL, Bra- zil, Amazonas, Rio Negro, beach, Ilha Ta- maquaré just downstream from Tapuru- cuara (approx. 00°25’S, 64°55'’W), 10 Oct 1979. The following non-type specimens were also examined: USNM 276502, 14; MBUCV V-15253, 24; Rio Negro, one-half hour upstream of San Carlos de Rio Ne- gro.—USNM 276501, 15, San Carlos de Rio Negro, Rio Negro margin, upstream of town landing.—AMNH 56175, 33; MBUCV V-15252, 20, Rio Negro at San Carlos de Rio Negro. Diagnosis.—The presence of prominent anteriorly divergent pelvic bones that are exserted in most individuals of Astyanax scologaster distinguishes the species from other characids with the exception of A. mu- cronatus Eigenmann, A. essequibensis Ei- genmann, A. zonatus Eigenmann, Deuter- odon pinnatus Eigenmann and OD. VOLUME 99, NUMBER 4 Mu Fig. 2. Astyanax mucronatus Eigenmann, cotype, FMNH 53521, 54.6 mm SL, Guyana, Potaro River, Tukeit. acanthogaster Eigenmann. Astyanax scolo- gaster,as other Astyanax species, has strong central cusps on all teeth and an abrupt change in the size of the teeth on the dentary which distinguishes the new species from all Deuterodon species which have flattened teeth without strong central cusps, and den- tary teeth in a graduated size series. As- tyanax scologaster has 21 or 22 branched anal-fin rays compared with 24 or 25 in A. mucronatus, 19 or 20 in A. essequibensis, and 25 to 29 in A. zonatus. Astyanax scol- ogaster has 33 to 35 scales in the lateral series versus 40 to 45 for Astyanax zonatus. Although the pigmentation patterns of As- tyanax scologaster and A. mucronatus are similar, the anterior humeral spot of A. scol- ogaster 1S narrower and more vertically elongate than the horizontal wedge-shaped humeral spot of A. mucronatus (compare Figs. 1 and 2). The relatively large eye of Astyanax scologaster (42.2—48.8% of HL) distinguishes that species from A. zonatus (37.7—40.6% of HL) and A. essequibensis (38.5—44.0% of HL). Astyanax scologaster is not as deep bodied as A. mucronatus (34.9-42.1 vs. 36.9-45.4% of SL; compare also Figs. 1 and 2), but deeper bodied than A. essequibensis (30.7—36.8% of SL) or A. zonatus (31.8-35.9% of SL). Description. —Table 1 gives morphomet- rics of holotype and measured paratypes. No evidence of sexual dimorphism in me- ristics or morphometrics was found. Body moderately deep, somewhat compressed, broadest in region of exsertion of pelvic spines in both sexes. Greatest body depth between origins of dorsal and pelvic fins, except in population from Massarabi in which greatest depth is in region of pelvic- spine exsertion, about midway between origins of pectoral and pelvic fins. Origin of dorsal fin closer to tip of snout than to hy- pural joint and often anterior to vertical line through origin of pelvic fin. Distance from snout to pelvic-fin origin (x = 50.9% of SL) usually greater than distance from snout to dorsal-fin origin (x = 49.2% of SL). Dorsal profile of head and body slightly convex from dorsal to orbit to dorsal-fin origin. Dorsal profile of body along dorsal-fin base nearly straight, posteroventrally oriented, nearly straight to slightly convex between dorsal and adipose fins. Caudal peduncle slender, compressed, rather short. Ventral profile of body slightly convex from sym- Fig. 3. Astyanax scologaster, new species, USNM 276503, 42.0 mm SL. Ventral view of pelvic-fin in- sertion and anteriorly proximate portion of body show- ing anteriorly divergent, exserted pelvic bones. Area in stipple pattern is without scales. S, pelvic spine. P, pelvic fin. B, body profile. physis of lower jaw to origin of pelvic fin, sometimes with sharp angle at point of pel- vic-spine exsertion from body; body trans- versely rounded anterior to this fin. Profile nearly straight between pelvic and anal fins (or nearly straight between pelvic-spine exsertion and anal-fin origin as in Fig. 1); straight and posteroventrally oriented along base of anal fin. Ventral profile of caudal peduncle slightly concave. Head length moderate, slightly more than one-quarter of standard length, snout rounded in profile, moderate in length, jaws equal and mouth terminal, or lower jaw slightly shorter and mouth barely subter- minal. Mouth nearly horizontal or slightly posteroventrally slanted; gape moderate, wider than snout length. Eye large. Inter- orbital width moderate, flat to slightly rounded transversely, but narrower than upper jaw width. Maxilla long, slightly lobed posteriorly; reaching slightly posterior to vertical line through anterior border of eye, and ventral to horizontal line through ven- tral border of eye. Maxilla with 3 to 5 very small com- pressed teeth; usually with 4 cusps, but with 5 cusps in some populations, third cusp longest. Premaxillary teeth in two series. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON aD Fig. 4. Map of portion of northern South America showing collecting localities of specimens of Astyanax scologaster, new species, A. mucronatus Eigenmann and A. essequibensis Eigenmann reported on in this paper (dotted lines depict international boundaries). Locali- ties 1 to 6 are collection sites of A. scologaster: 1, Cano Manu; 2, Cafio Chola; 3, upstream of San Carlos de Rio Negro; 4, San Carlos de Rio Negro; 5, Massarabi; 6, Ilha Tamaquaré. See listing of holotype, paratypes, and non-type specimens examined for detailed locality and collection information. Localities 7 to 9 are col- lection sites of A. mucronatus and A. essequibensis: 7, Crab Falls; 8, Tumatumari; 9, Tukeit. See listing of ‘“Comparative Material Examined” for more detailed locality information. Outer series usually in rather uneven row of 4 teeth of differing forms; third tooth from symphysis narrower, rather round in cross section, and with 3 cusps. Remaining teeth of outer premaxillary series broader, more ovoid in cross section, with at least a very small additional pair of cusps for a total of 5. Inner series of premaxillary teeth con- sisting of 5 broad teeth extending across gape. Each tooth with prominent central cusp flanked by two or, more usually, three pairs of cusps of decreasing size arranged in semicircle progressing around anterior bor- der of tooth. Lateralmost premaxillary tooth smaller, semi-compressed, sometimes al- most indistinguishable in form from dor- salmost maxillary tooth. Dentary with 5 large teeth along anterodorsal margin of VOLUME 99, NUMBER 4 WAS) Table 1.—Morphometrics of Astyanax scologaster, new species. Standard length is expressed in mm; mea- surements | to 12 are percentages of standard length; 13 to 16 are percentages of head length. Paratypes (59) Holotype Range Average Standard length 37.5 28.8-42.0 1. Greatest body depth 38.9 34.9-42.1 38.2 2. Snout to pelvic-fin origin 50.1 48 .3-53.2 50.9 3. Snout to dorsal-fin origin 48.3 47.3-52.0 49.2 4. Dorsal-fin origin to caudal base 56.0 53.6-57.6 55.6 5. Snout to anal-fin origin 64.3 61.9-67.4 64.1 6. Caudal peduncle length 12.3 11.2-13.6 12.5 7. Caudal peduncle depth 9.9 8.8-10.6 9.7 8. Pelvic-fin length 16.8 15.1-18.3 17.0 9. Length pelvic bone 10.9 10.0-12.6 11.5 10. Distance between exserted tips of pelvic bones 8.3 5.3-14.2 9.4 11. Snout to pectoral-fin origin 28.5 26.5-30.3 28.6 12. Bony head length 28.8 27.7-31.2 29.6 13. Horizontal eye diameter 46.3 42.2-48.8 45.9 14. Snout length 29.6 23.6-31.1 27.4 15. Upper jaw length 42.6 38.4—44.0 41.4 16. Bony interorbital width 29.6 26.7—34.2 30.6 bone. Large dentary teeth similar in form to inner premaxillary teeth, fitting closely behind latter when mouth closed. Central cusp long and strong; usually flanked by two pairs of smaller cusps. Larger dentary teeth followed posteriorly by 6 to 8 very small teeth with | to 3 uneven cusps. Infraorbital series complete, all elements relatively small, ventral and posterior in- fraorbitals not contacting sensory tube of preopercle. Frontoparietal fontanel extend- ing into posterior portion of ethmoid; pa- rietals completely separated; frontals in contact only at epiphyseal bar. Fontanel gradually widening posteriorly, extending onto dorsomedial portion of supraoccipital. Supraoccipital spine moderately developed. Adipose eyelid (a thick band of clear con- nective tissue) extending posteriorly from nostrils over lateral surface of head and an- terior midlateral portion of body to anterior humeral spot; with large, almost circular Opening Over eye. Dorsal-fin rays 11,8,i or 11,9 [11,9]; second unbranched ray usually rather short, less than one-half length of longest fin-ray; lon- gest rays are first and/or second branched rays. Dorsal fin length moderate. Pectoral- fin rays 1,11 to 1,14 [i,13]. Pectoral fin length moderate, usually not extending posteriorly to vertical through pelvic-fin origin. Clei- thrum large, anterodorsal border ap- proaches posterior border of preopercle. Posterolateral tips of coracoid bones widely separated. Pelvic-fin rays 1,7 in all speci- mens. Fin moderate in length, tips usually extending to near anal-fin origin. No hooks present on fin rays. Overall shape of pelvic bone modified anteriorly into strong spine. Anterior tips of two pelvic bones diverging widely and usually exserted from body wall. Length of pelvic bone measured from in- sertion of fin rays to tip of exserted spine 10.0-12.6% of SL, x = 11.5 [10.9], up to nearly half of length of bone exserted; dis- tance between anterior tips of spines 5.3- 14.2% of SL, x = 9.4 [8.3]. Anal fin with 2 or 3 unbranched anterior rays (first ray very small in many specimens examined, suggesting that very small first 714 ray typically present but not always visible) [2], followed by 20 to 23 branched rays, X = 21.6 [23]. Last ray divided to base and counted as two rays in most individuals, but in many specimens from Massarabi and Ta- maquaré posterior portion of ray un- branched and not included in branched-ray count. Anal-fin margin concave with longer anterior rays forming small anterior lobe; following rays abruptly shorter; fin shape similar in both sexes. No hooks present on anal-fin rays, but absence of hooks on anal- and pelvic-fin rays may be seasonal. Basal sheath of scales on anal fin short, consisting of 5 to 7 scales along anterior portion of fin base. Caudal-fin rays i1,9/8,i in all specimens examined. Caudal fin moderate in length, rather narrow, deeply forked, ventral lobe usually slightly larger than dorsal. Adipose fin of moderate size. Scales in lateral series 33 to 36 [35]. Scale rows in vertical series between dorsal- and pelvic-fin origins usually 7/5 [7/5]. Vertebrae of holotype and 25 paratypes: 337°C) 234,20) 93516): Color in life. —Overall coloration of head and body bright silver. Color in alcohol. —Color description based on holotype (Fig. 1). Scales lacking guanine, ground color pale yellowish tan, slightly darker dorsally. Head with small to medium dark brown chromatophores dor- sally and anteriorly. Few large brownish chromatophores ventrally near maxilla and on first infraorbital bones; larger, darker chromatophores on opercle and preopercle. Dark blackish chromatophores extending along base of dorsal fin and posteriorly to dorsal caudal-fin rays. Dark chromato- phores present along base of anal fin. Scat- tered fine dark chromatophores along bor- ders of scales of median predorsal scale row, horizontal scale rows 1 to 3 on region an- terior to dorsal fin, and scales of most of body posterior to dorsal fin. Anteroventral portions of body pale except for few chro- matophores along scale borders and ventral portions of humeral spots. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Two humeral spots present. Anterior a dusky vertical bar darkest on third and fourth scales of sixth horizontal scale row. Bar extending dorsally into third horizontal scale row and ventrally into ninth scale row; somewhat paler ventral to lateral line. Pos- terior humeral spot about two scales behind anterior spot. On holotype two humeral spots about same length but posterior spot narrower dorsally and much lighter. Most other specimens with posterior humeral spot smaller. Light dusky midlateral band of widely scattered small dark chromato- phores extending from posterior border of posterior humeral spot to caudal-fin base, widening on caudal peduncle to form pale, indistinct caudal spot terminating poste- riorly at caudal-fin base. Few large brown chromatophores proximally on membranes of middle caudal-fin rays. Dusky patches on dorsal and ventral lobes of caudal fin aligned with dorsal and ventral corners of caudal peduncle spot. Dorsal-fin membranes with patch of small black chromatophores forming indistinct horizontal band across middle of fin. Anal fin with band of dusky chromatophores along middle of fin and basal dusky spot anteriorly. Median portion of pectoral fin dusky. Dorsal border of adipose fin with several black chromatophores. Pelvic and pectoral fins almost clear. Etymology. —The specific epithet, scolo- gaster from the Greek skolos, thorn or pointed object, and gaster, stomach, refers to the exserted spinous pelvic bones on the ventral surface of the body of this species. Relationships. —The common possession of an exserted pelvic bone and Astyanax- type dentition (see “‘Diagnosis”) unites Astyanax scologaster, A. mucronatus, A. es- sequibensis, and A. zonatus. The exact re- lationships of the species within this group- ing remain unresolved. Ecology. — Astyanax scologaster is an in- habitant of the black-water main river chan- nels and tributary streams over much of the Rio Negro basin. The species has been cap- VOLUME 99, NUMBER 4 tured both along the margins of the swiftly flowing main channel of the Rio Negro and in slowly flowing terra-firma tributary streams. Comparative daytime and night- time sampling with nets at two localities on the upper Rio Negro has shown that the species is active at night, but absent during the daylight hours (RPV, pers. obs.). Comparative material examined.—As- tyanax mucronatus Eigenmann: FMNH 53520, holotype; Guyana, Essequibo, Po- taro River, Tumatumari Cataract.— FMNH 53521, 5 cotypes; CAS 39347, 4 cotypes; MCZ 29963, 1 cotype; Guyana, Essequibo, Potaro River, sandbank at Tukeit. Astyanax essequibensis Eigenmann: FMNH 53519, holotype; CAS 39144, 24 paratypes; SU 21956, 5 paratypes; Guyana, Essequibo, Potaro River, Tumatumari Cat- aract.—FMNH 52962, 11 cotypes; MCZ 29958, 2 cotypes; CAS 39146, 23 cotypes; Guyana, Essequibo, Essequibo River, Crab Falls. Astyanax zonatus Eigenmann: MCZ 20768, 1 syntype; MCZ 20766a, 1 syntype; MCZ 20753, 2 syntypes; Brazil, Amazonas, Rio Solimoes, Tabatinga. Deuterodon pinnatus Eigenmann: FMANH 53525, holotype; FMNH 53526, 3 para- types; MCZ 29952, 2 paratypes; Guyana, Essequibo, Potaro River, Amatuk Cata- ract.—MCZ 29951, 2 cotypes; Guyana, Es- sequibo, Essequibo River, Warraputa Cat- aract. Deuterodon acanthogaster Eigenmann: FMNH 54748, holotype; Brazil, Mato Grosso do Sul, Corumba.—FMNH 54750, 5 cotypes; Brazil; Mato Grosso do Sul, Rio Jauru, SW of Sao Luis de Caceres. -USNM 232385, 3; Paraguay, Amambay, Rio Aqui- daban, Parque Nacional Cerro Cora. — MZUSP 30868, 5; Brazil, Para, Rio Itacai- nos, Serra dos Carajas.—MZUSP 30543, 5; Brazil, Para, Rio Tapajos between Jacare- a-Canga and Itaituba. Remarks.—Eigenmann (1911:180, Fig. 2) in his original description of Deuterodon acanthogaster noted that the species was 715 characterized by protruding innominate bones (= pelvic bones) but did not comment on the presence of the structures in any oth- er members of the genus. Comparative studies associated with this study have dis- closed comparable spines in Deuterodon pinnatus Eigenmann. Resumen. —Astyanax scologaster es des- crita provenientes de varias localidades del medio y alto Rio Negro de Venezuela y Bra- sil. Esta especie se caracteriza por poseer la seccion anterior de los huesos pélvicos modificados en unas espinas lateralmente divergentes, las cuales se extienden a través de la piel en la mayoria de los individuos. Una modificacion similar ha sido obser- vada en Astyanax essequibensis, A. mucro- natus, A. zonatus, Deuterodon pinnatus, y D. acanthogaster. Las tres especies de As- tyanax con los huesos pélvicos prominentes pueden ser distinguidas de las especies de Deuterodon, por una serie caracteres den- tales. Astyanax scologaster se diferencia de sus congéneres por poseer 21 6 22 radios anales bifurcados, comparado con 24 6 25 en A. mucronatus, 19 6 20 en A. essequi- bensis y 25-29 in A. zonatus. Los patrones de coloracion y diferencias morfométricas y meristicas, distinguen aun mas a A. scol- ogaster del resto de las especies. Acknowledgments The specimens that served as the basis for this study were collected by a joint MBUCV-USNM expedition to the upper Rio Negro, and as a corollary to AMNH collecting efforts associated with the expe- dition to Cerro de la Neblina National Park organized by La Fundacion para el desarro- llo de Las Ciencias Fisicas, Matematicas y Naturales. We thank Dr. Antonio Macha- do-Allison (MBUCV) for his efforts in or- ganizing the MBUCV-USNM expedition. He, along with Mr. Carl J. Ferraris, Jr., Lic. Justa M. Fernandez, Lic. Otto Castillo, and Sr. Ramiro Royero assisted in collecting the specimens. Specimens from the Brazilian portions of the Rio Negro were collected in 716 1979 by Dr. Michael Goulding, then of the Instituto Nacional de Pesquisas da Ama- zonia, Manaus, Brazil. That material, sub- sequently sorted and housed at the Museu Paraense Emilio Goeldi, Belém, Para, Bra- zil, and now deposited at MZUSP, was loaned for this study by Dr. Naércio A. Me- nezes. Mr. Andrew G. Gerberich, Mr. Kurt A. Bruwelheide and Ms. Ann M. Williams provided technical assistance at USNM. Figures | and 2 were prepared by Mr. Theo- philus Britt Griswald. The Spanish trans- lation of the ““Resumen” was provided by Dr. Antonio Machado-Allison. Participa- tion by one of us (RPV) in collecting efforts on the upper Rio Negro was made possible by a grant from the Scholarly Studies Pro- gram of the Smithsonian Institution. Re- search associated with this study was sup- ported in part by the I.E.S.P. Neotropical Lowland Research Program of the Smith- sonian Institution. This paper was im- proved by the comments and suggestions of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Dr. Stanley H. Weitzman and Dr. Wayne C. Starnes. Literature Cited Eigenmann, C.H. 1909. I. Reports on the Expedition to British Guiana of the Indiana University and the Carnegie Museum, 1908.—Annals of the Carnegie Museum 6(1):4—54. 1911. III. New characins in the collection of the Carnegie Museum.— Annals of the Carnegie Museum 7(1):164-181. . 1921. The American Characidae.— Memoirs of the Museum of Comparative Zoology 43(3): 209-310. Fink, W. L.,andS.H. Weitzman. 1974. The so-called Cheirodontin fishes of Central America with de- scriptions of two new species (Pisces: Charac- idae).—Smithsonian Contributions to Zoology 172:1-—46. Department of Vertebrate Zoology (Fish- es), National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 717-730 BARNACLES (CIRRIPEDIA: BALANIDAE) FROM THE LOWER PLEISTOCENE JAMES CITY FORMATION, NORTH CAROLINA COASTAL PLAIN, WITH THE DESCRIPTION OF A NEW SPECIES OF BALANUS DA COSTA Victor A. Zullo and William Miller, III Abstract. —Numerous specimens of Balanus neusensis, new species, and B. improvisus Darwin, and a few specimens of B. calidus Pilsbry occur in asso- ciation with a Crepidula fornicata (Linnaeus) biostrome in the lower Pleistocene James City Formation at Johnson Point, Neuse River, Craven County. Balanus improvisus and B. neusensis also are reported from an isolated James City outcrop at Dam Creek on the Neuse River and from a tidal inlet deposit of presumed equivalent age at Woodside, Pender County. The extant tropical American and Carolinian species B. calidus is common in inner shelf deposits of early Pleistocene age in the Carolinas, whereas the extant North Atlantic species B. improvisus is more typically found in younger Pleistocene marginal marine deposits of the region. Balanus neusensis appears to be the youngest surviving species of the B. pacificus Pilsbry group in the western Atlantic, and a descendent of the Pliocene species B. oppidieboraci Ross. Bulk sediment samples from two locali- ties in the lower Pleistocene James City For- mation on the Neuse River, Craven County, North Carolina yielded remains of three species of barnacles (Fig. 1). The samples were taken and processed by Miller as part of a paleoecologic study of the James City fauna (Miller and DuBar, in prep.). The ma- jority of barnacle specimens were associated with a Crepidula fornicata (Linnaeus) bio- strome at Johnson Point in the type area of the formation (UNCW locality Z-807). The second locality (UNCW locality Z-808) near Dam Creek, is an isolated outcrop of James City sediments that yielded relatively few barnacle remains. The abundance of bar- nacles at the Johnson Point locality is re- lated to the numerous attachment sites for sessile, epibenthic organisms provided by the large, tabular accumulation of the sed- entary epifaunal gastropod Crepidula for- nicata. Two of the Johnson Point barnacles, Balanus improvisus Darwin and B. calidus Pilsbry, are extant species with Pleistocene fossil records on the Atlantic coast. The third and most abundant barnacle is a new species that bears considerable superficial resem- blance to the extant estuarine Atlantic Coast species B. eburneus Gould, but is, rather, related to the Balanus pacificus Pilsbry group. Extant members of the B. pacificus group are known only from the Pacific ba- sin, but Neogene representatives are known from the Atlantic Coast Pliocene, and Bal- anus oppidieboraci Ross, a likely precursor of the new species, occurs in the Pliocene Yorktown Formation of Virginia. Only a few specimens of the new species and of B. improvisus were identified at the Dam Creek locality. As discussed in the sec- tion on paleoenvironments and paleoecol- ogy, differences in abundances of barnacle remains at the two sites are related to the location of the two sites within the James City embayment and to concomitant dif- ferences in depositional environment. The Johnson Point biostrome accumulated in a relatively high energy, shallow water envi- 718 Fig. 1. Location of James City Formation collect- ing sites on the Neuse River, Craven County, North Carolina. ronment where sedimentation rates were low. Barnacles, in addition to requiring the hard substrata for settlement provided by the Crepidula fornicata biostrome, are filter feeders that thrive in higher energy sites with little sedimentation. The Dam Creek site, by comparison, represents a deeper water, bay-center deposit where very fine-grained sediments were being deposited and few hard substrata were available. It is likely that the barnacle remains preserved at the Dam Creek locality are allochthonous. A few, worn, disarticulated specimens of the new species and some juvenile, but well preserved entire individuals of B. improvi- sus were obtained from a tidal inlet deposit of presumed equivalent age at Woodside, Pender County, North Carolina (UNCW lo- cality Z-750; Fig. 2). This is the only other known occurrence of the new species. Bal- anus improvisus, however, is common in later Pleistocene, marginal-marine deposits both in North and South Carolina and is presently found to range from the nearshore subtidal up through the mid-intertidal zone, primarily in marginal-marine environ- ments. To the north of Cape Hatteras, B. improvisus 1s essentially restricted to estu- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON NORTH CAROLINA O Fayetteville Georgetown fe Fig. 2. Location of North and South Carolina Pleis- tocene sites discussed in text. aries (oligohaline through polyhaline), but to the south this species can be abundant in polyhaline and coastal euhaline waters. Balanus calidus is probably the most commonly encountered barnacle in lower Pleistocene deposits of the Carolinas. Pleis- tocene examples are usually found in great numbers on shells of Ostrea sculpturata Conrad from localities that have been in- terpreted as representing open water envi- ronments or embayments with unrestricted circulation (e.g., Lee Creek Mine, Beaufort County, North Carolina). Extant popula- tions are found in marine waters at subtidal depths and are best developed on the inner and middle shelf. Stratigraphic Setting Three lithostratigraphic units crop out along the Neuse River east of New Bern (DuBar and Solliday 1963, Mixon and Pil- key 1976, Miller 1985). These are, in as- cending order: 1) lower Pleistocene James City Formation, exposed for the most part along the south shore from Black Beacon Point downstream to Johnson Point; 2) middle Pleistocene Flanner Beach Forma- tion, in bluffs along both sides of the river extending from Johnson Point to the Suffolk (or Grantsboro) Scarp; and 3) upper Pleis- VOLUME 99, NUMBER 4 tocene Core Creek sand, which is poorly exposed downstream from the Suffolk Scarp (Fig. 1). All three units were deposited dur- ing glacio-eustatic high stands that occurred at 1,000,000-800,000 yr, 200,000 yr, and 100,000 yr B.P., respectively (McCartan et al. 1982, Cronin et al. 1984). The oldest Pleistocene depositional cycle is represented by fine-grained, fossiliferous, nearshore marine beds of the James City Formation (DuBar and Solliday 1963, DuBar and Howard 1969, DuBar et al. 1974). The Johnson Point exposure consists of a bluish gray to olive gray, shelly mud- stone to siltstone, broadly divisible into a lower bed made up of discontinuous, thin layers of Crepidula fornicata shells in life position (stacks of adhering shells) and an upper bed characterized by varied, abun- dant fossil bivalves, especially Anadara aequicostata Conrad, Ostrea sculpturata Conrad and Noetia limula Conrad. At Dam Creek, slightly over one meter of the James City Formation is overlain disconformably by the Flanner Beach Formation. Here, the James City is a bluish-gray, moderately fos- siliferous mudstone to claystone containing valves of the small bivalve Nuculana acuta (Conrad). Near the base of the outcrop, heads of the ahermatypic coral Septastrea occur in apparent life position (Miller 1985:120). The James City beds, both at Johnson Point and Dam Creek, were deposited as muddy, more or less shelly sediments with- in a protected, subtropical marine embay- ment. Johnson Point appears to be part of a regressive sequence, with deeper subtidal deposits containing the Crepidula bank fau- na succeeded in the section by shallower subtidal deposits containing a great variety of mollusks and other organisms. Dam Creek seems to be a deeper, bay-center de- posit that accumulated basinward of the Johnson Point sequence. Presumably contemporaneous, sandy, Open marine beds are exposed 40 km north of the Neuse River in the Lee Creek Mine at Aurora (referred to as “Croatan Forma- 719 tion’”’ by many workers; see contributions in Ray 1983). The exact temporal and pa- leogeographic relationships between the James City Formation in the Neuse River Valley area and the Waccamaw Formation in southeastern North Carolina and north- eastern South Carolina, traditionally re- garded as coeval units, are far from being clear. A recent interpretation based on mul- tiple lines of evidence suggests that the James City Formation may be a little younger than most of the Waccamaw (Cronin et al. 1984). Paleoenvironments and Paleocommunities Table 1 lists the most abundant autoch- thonous invertebrate fossils in samples col- lected from Johnson Point and Dam Creek. This list gives a general sense of the benthic paleocommunity contexts for balanids liv- ing in James City depositional environ- ments. A part of this ecosystem-level con- text at Johnson Point included long-term community replacement resulting from sus- tained, unidirectional habitat alteration in subevolutionary time (Miller 1986). The faunal transition at Johnson Point involved the fairly abrupt decline of the biostrome and its replacement by a more diverse, level- bottom community with many infaunal, suspension-feeding bivalves, carnivorous and scavenging gastropods, and occasional clumps of oysters. Little is known about the conditions con- ducive to extensive buildups of C. fornicata into banks the size of the Johnson Point biostrome, which is estimated to have cov- ered up to 15,000 m2? of sea floor during maximum development (Miller and DuBar, in prep.). A long period of low turbidity, consistent currents carrying food supplies, and near-normal marine salinity probably were key factors (see Johnson 1972, DuBar et al. 1974:111, Hoagland 1979). Changes in currents, probable shallowing in the Johnson Point area, and a shift in local dis- turbance regime (seasonality, frequency, in- tensity, and geographic extent of physical disturbance; see Sousa 1985, Connell and 720 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Most important components of fossil as- sociations from the James City Formation in the Neuse valley. Only organisms with individuals making up =0.5% of the association are listed. Format includes taxon, relative abundance, followed by organism type/ substrate niche + feeding category. All abbreviations explained at foot of table; sample numbers refer to Fig. 1. I. Crepidula biostrome association (Johnson Point, samples | and 2) Table 1.—Continued. Anadara aequicosta- II. III. Crepidula fornicata* 43.2% Ga/EPSF Boonea seminuda 29.5 Ga/EPPA Balanus spp. 8.6 Ci/EPSF Nuculana acuta 4.8 Bi/INDF Ostrea sculpturata 4.3 Bi/EPSF Anadara aequi- costata 2.3 Bi/INSF Anachis lafresnayi var. 1.5 Ga/VAGC A. obesa 1E5 Ga/VAGC Nassarius albus 0.8 Ga/VAGS Ostrea clump association (Johnson Point, sample 3) Crepidula fornicata 30.2% Ga/EPSF Boonea seminuda 27.9 Ga/EPPA Balanus spp. 10.8 Ci/EPSF Nuculana acuta 7.4 Bi/INDF Ostrea sculpturata* 4 7/ Bi/EPSF Anadara aequi- costata Dell Bi/INSF Mulinia lateralis 1e2. Bi/INSF Turbonilla (Chem- nitzia) sp. Hol Ga/EPPA Sphenia sp. ili Bi/INSF? Boonea impressa ail Ga/EPPA Anachis obesa 1.0 Ga/VAGC A. lafresnayi var. 0.9 Ga/VAGC Mercenaria sp. cf. M. permagna 0.7 Bi/INSF Abra aequalis 0.7 Ga/VAGC Urosalpinx per- rugata 0.7 Ga/VAGS Busycon sp. 0.6 Ga/VAGS Nassarius albus 0.6 Ga/VAGS brachyurans 0.5 Ma/VAGS Anadara-Noetia association (Johnson Point, sample 4) Crepidula fornicata 17.8% Ga/EPSF Boonea seminuda 7/2 Ga/EPPA Ostrea sculpturata 11.9 Bi/EPSF Nuculana acuta 11.4 Bi/INDF Balanus spp. 8.7 Ci/EPSF ta* 8.0 Bi/INSF Abra aequalis 4.4 Bi/INDF Sphenia sp. aN Bi/INSF? brachyurans 5) Ma/VAGS Anachis lafresnayi 1.5 Ga/VAGS Boonea impressa 1.4 Ga/EPPA Cumingia tellinoides itil Bi/INSF Mercenaria sp. cf. M. permagna 1.1 Bi/INSF Mulinia lateralis 0.9 Bi/INSF Turbonilla (Pyrgis- CUS) Sp. 0.9 Ga/EPPA Anachis obesa 0.8 Ga/VAGC Turbonilla (Chem- nitzia) sp. 0.7 Ga/EPPA Melanella conoidea 0.7 Ga/VAGU Urosalpinx per- rugata 0.7 Ga/VAGC Nucula proxima 0.6 Bi/INDF Nassarius albus 0.6 Ga/V AGS Polinices sp. 0.5 Ga/VAGC Prunum sp. 0.5 Ga/VAGU Noetia limula* 0.5 Bi/INSF Vermicularia sp. cf. V. knorri 0.5 Ga/EPSF? IV. Nuculana association (Dam Creek) Nuculana acuta* 61.3% Bi/INDF Mulinia lateralis 24.2 Bi/INSF Parvilucina multi- lineata 4.0 Bi/INSF Abra aequalis Dal Bi/INDF Anadara aequi- costata ill Bi/INSF Crassinella lunulata ili Bi/INSF Ensis sp. 1.0 Bi/INSF Balanus spp. 1.0 Ci/EPSF Prunum sp. 0.7 Ga/VAGU Vermicularis sp. cf. V. knorri 0.5 Ga/EPSF? Ostrea sp. 0.5 Bi/EPSF Gemma gemma 0.5 Bi/INSF Symbols used in table: *—dominant organism in terms of biovolume; lends name to association. Or- ganism type: Bi—bivalve mollusk, Ga—gastropod mollusk, Ci—cirriped crustacean, Ma—malacostracan crustacean. Substrate niche + feeding category: INSF— infaunal suspension-feeder, INDF—infaunal deposit- feeder, EPSF—epifaunal suspension-feeder, EPPA— epifaunal parasite, VAGC — vagrant carnivore, VAGS— vagrant scavenger, U—unknown. VOLUME 99, NUMBER 4 Keough 1985) led to disruption of the Cre- pidula-dominated community and estab- lishment of the community represented by the Ostrea clump and Anadara-WNoetia fossil associations (Table 1). In terms of changing spatial patterns, laterally extensive blankets of Crepidula stacks gave way to a much patchier and consequently more diverse community with only a few isolated Cre- pidula aggregations persisting in the area. Although barnacles are abundant through- out the transition sequence, they are more numerous in shelly beds within the bio- strome probably because of widespread availability of hard substrata in the form of exposed Crepidula shells. By comparison, balanids are uncommon at Dam Creek (Table 1) in a fossil asso- ciation dominated by the infaunal, depos- it-feeding bivalve Nuculana acuta. This association was derived from a deeper subtidal community inhabiting a very fine- grained, soft to possibly thixotropic bottom, offshore from the Crepidula biostrome and Anadara-Noetia-Ostrea paleocommunities (see Parker 1976). Locality Descriptions University of North Carolina at Wilming- ton (UNCW): Z-635 Cape Fear coquina, south side of Snows Cut, 650 m west of U.S. Highway 421 bridge over Atlantic Intracoastal Waterway, southern New Hanover Coun- ty, North Carolina. V. Zullo, coll., 5 Sep 1980. Z-642 Upper Pleistocene shell bed in bor- row pit (since reclaimed) for construction of Mark Clark Expressway, on north side of Ashley Hall Road, 1370 m east of in- tersection with State Highway 61 (Ashley River Road) and west of Ashley River, Charleston County, South Carolina. V. Zullo, coll., 7 Feb 1981. Z-750 Waccamaw Formation, quarry at seaward edge of +7.6 m terrace, south- 721 east of U.S. Highway 17 at Woodside, Pender County, North Carolina. V. Zullo, coll., 26 Jun 1985. Z-807 James City Formation, north side of Johnson Point in steep-sided bank, 500 m upstream (northwest) from tip of point and 400 m downstream (southeast) from pier at Veterans of Foreign Wars building, south shore of Neuse River, Craven County, North Carolina. W. Miller, coll., Aug 1982. Z-808 James City Formation, basal bed in low bank 3.8 km upstream (northwest) from mouth of Slocum Creek and 700 m downstream (southeast) from mouth of Dam Creek, south shore of Neuse River, Craven County, North Carolina. W. Mil- ler, coll., Aug 1982. Z-809 “‘Croatan Sand,” Lee Creek Mine, south shore of the Pamlico River, Rich- land Township, north of Aurora, Beau- fort County, North Carolina. Aura Baker, coll., date unknown. Z-812 Waccamaw Formation, Cedar Creek Village quarry, southeast side of U.S. Highway 17, Little River, Horry County, South Carolina. V. Zullo and W. B. Har- ris, coll., 1977. Systematic Paleontology Suborder Balanomorpha Pilsbry, 1916 Superfamily Balanoidea Leach, sensu Newman and Ross, 1976 Family Balanidae Leach, sensu Newman and Ross, 1976 Subfamily Balaninae Leach, sensu Newman, 1980 Genus Balanus Da Costa, 1778 Balanus improvisus Darwin, 1854 Fig. 3 Balanus improvisus Darwin, 1854:250, pl. 6, fig. la—c; Pilsbry, 1916:84, text fig. 16a, pl. 24, figs. 3-3b, 5-5d; Henry and McLaughlin, 1975:68, text fig. 16, pl. 5, figs. a-f, g, h-j (see for complete synon- ymy). 722 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 99, NUMBER 4 Material examined. —Johnson Point, Neuse River, UNCW locality Z-807: 16 complete shells, over 200 compartmental plates, over 50 opercular plates.—Dam Creek, Neuse River, UNCW locality Z-808: 18 compartmental plates, 1 scutum.— Woodside, Pender County, UNCW locality Z-750: 6 shells, some with opercular plates, 8 isolated scuta, 3 isolated terga.—Snows Cut, New Hanover County, UNCW locality Z-635: 7 shells, 3 scuta, 5 terga.—Charles- ton County, South Carolina, UNCW local- ity Z-642: 9 compartmental plates, 1 ter- gum. Disposition of specimens. —Figured hy- potypes USNM 405359 through 405365 are in the collection of the Department of Pa- leobiology, National Museum of Natural History, Washington, D.C. Remaining specimens are in the collection of the De- partment of Earth Sciences, University of North Carolina at Wilmington. Geologic and geographic range. — Recent: Scotland and Baltic Sea to West Africa; Mediterranean, Black, Caspian and Red seas; Nova Scotia to Patagonia; Oregon to Peru; Japan; Australia (Henry and Mc- Laughlin 1975). Documented fossil occur- rences: Pleistocene, North and South Car- olina. Discussion. — Balanus improvisus is a common element of estuarine faunas of the North Atlantic basin, and has been intro- duced to various localities throughout the world by ships. Fossils of this species have not been reported previously from the East Coast of North America. Kolosvary (1955, 1959, 196la, b) cited B. improvisus from the Miocene (Burdigalian and Tortonian) of Hungary and the USSR without accompa- —_— 723 nying descriptions or illustrations of the specimens. Because of the antiquity of this extant species suggested by these records, and because of the distance of these local- ities from the known natural range of B. improvisus, Kolosvary’s identifications are in doubt. The specimens from the James City For- mation and from Woodside are typical for the species, which is identified by: (1) its narrow radii with nonseptate sutural edges; (2) its scutum which lacks external radial striae and bears a well-developed adductor ridge; and (3) its tergum with its long, very narrow tergal spur and partially infolded spur furrow. Balanus improvisus is also known from late Pleistocene deposits in the Carolinas, including the Cape Fear “‘co- quina’”’ in southern New Hanover County, North Carolina (UNCW locality Z-635) and an upper Pleistocene shell bed northwest of Charleston, South Carolina (UNCW local- ity Z-642; Zullo 1986). Balanus neusensis, new species Figs. 4a-e, 5, 6 Holotype. —Shell with opercular plates, USNM 405366. Type locality.—Crepidula fornicata bio- strome, UNCW locality Z-807, James City Formation, Johnson Point, Neuse River, Craven County, North Carolina. Diagnosis.—Smooth to irregularly pli- cate, high conic shell with moderately- toothed, subtriangular orifice and broad ra- dii with oblique, crenate summits; parietal tubes open throughout; basis porous, pores not septate; scutum very narrow, with prominent radial striae, markedly denticu- Fig. 3. Balanus improvisus: a, Side view of shell, hypotype USNM 405359, x3; b, Interior of scutum, hypotype USNM 405360, x18; c, Interior of tergum, hypotype USNM 405361, x21; d, Exterior of tergum, hypotype USNM 405362, x21; e, Exterior of scutum, hypotype USNM 405363, x25; f, g, Interiors of scutum and tergum, hypotype USNM 405364, x37; h, Exterior of tergum, hypotype USNM 405365, x33. Figures a— e, UNCW locality Z-807, Johnson Point; f, g, UNCW locality Z-750, Woodside; h, UNCW locality Z-642, Charleston. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 724 VOLUME 99, NUMBER 4 late occludent margin, prominent, high ad- ductor ridge extending nearly from apex to basal margin and separated from erect ar- ticular ridge by deep, narrow cleft; tergum relatively narrow, with infolded, partially closed spur furrow and short, narrow spur placed its own width from basiscutal angle; basal margin of tergum not embayed on ca- rinal side. Description. —Shell (Figs. 4a—e, 5d—g) of moderate size (largest 2.4 cm in height and 2 cm in greatest basal diameter), high conic to subcylindric, with large, moderately- toothed, subtriangular orifice; parietes smooth or irregularly plicate, plications re- flecting irregularities of substratum; radii thick, solid, moderately broad, glossy, hor- izontally striate, with roughly crenulated summits sloping about 30° from horizontal; sutural edges of radii septate, with promi- nent denticulae on lower sides of septa; edges of parietes receiving sutural edges of radii also septate, with denticulae on upper sides of septa; alae thick, broad, short, with mod- erately convex summits and finely-septate sutural edges; sheath solid, short, confined to upper third of interior of shell; basal mar- gin of sheath thin, dependent, forming deep, narrow, non-vesiculate cavity between edge of sheath and interior of parietes; interior of parietes strongly and regularly ribbed near base, ribs fading towards base of sheath; internal ribs conforming to parietal septa; parietal tubes numerous, round to oval, moderately large, open from apex to base; occasional, single, fine, secondary septa ex- tending into parietal tubes from external lamella; basis porous, thin at center, thick- ening at margin; basal tubes small, non-sep- tate. — 2s) Scutum (Figs. 5c, d, g, 6a—c) nearly flat, very narrow, thick; narrow strip of tergal margin sharply reflexed inward at 90°; ex- terior ornamented by prominent growth ridges (best developed on lower half of plate) that are broken into nodes where they are crossed by deeply incised radial striae on central third of plate; striate central third of plate noticeably sulcate; reflexed tergal mar- gin and narrow strip of plate adjacent to occludent margin without radial striae; ma- jor growth ridges extending onto occludent margin to form large, obliquely-inclined oc- cludent teeth; apex and basioccludent angle acute, basitergal angle truncate at about 45°; occludent margin straight; tergal margin straight to slightly convex; basal margin sin- uous, with central third convex, reflecting central, external sulcus of plate; length of basal margin no more than one-half length of occludent margin; articular ridge prom- inent, erect, with broad, flat upper surface, and extending from just below apex of plate to just beyond midpoint of plate; articular furrow moderately broad, deep, crossed by 2 or 3, low, sharp, nearly vertical growth ridges; adductor ridge prominent, highest in center, inclined toward articular ridge, very long, extending from just below upper end of articular ridge nearly to basal margin; adductor ridge not confluent with articular ridge, leaving deep, narrow cleft between ridges extending nearly to apex of plate; ad- ductor muscle pit large, usually deep, oval, located near midpoint of plate between ad- ductor ridge and occludent margin; 2 short, low, thin, nearly equally-spaced, vertical ridges occurring on lower third of interior of scutum between adductor ridge and basi- tergal angle; ridge closest to truncate basi- Fig. 4. a-e, Balanus neusensis, UNCW locality Z-807, Johnson Point: a, Shell clump, including holotype USNM 405366 (lower center) and paratype lot USNM 405367, x2; b, Side view of shell, paratype USNM 405368, x 2.4; c, Rostrocarinally elongate shell growing on scutum, paratype USNM 405369, x 6.5; d, Oblique view of shell with convex basis, previously attached to interior of Crepidula shell, paratype USNM 405370, x 2.4; e, Oblique view of subcylindric shell, paratype USNM 405371, x<2.4; fj, Balanus calidus, hypotype USNM 405372, UNCW locality Z-807, Johnson Point: f, Oblique view of shell, x6.5; g, Interior of tergum, x 32; h, Exterior of broken tergum, x32; 1, Exterior of scutum, x33; j, Interior of scutum, x33. 726 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 99, NUMBER 4 727 Fig. 6. Balanus neusensis, UNCW locality Z-807, Johnson Point. a—c, Exteriors of scuta showing range in development of external radial striae: a, paratype USNM 405377, x6; b, paratype USNM 405378, x6; c, paratype 405379, x12; d, Interior of carina, paratype USNM 405380, x 2.5; e, Interior of rostral plate, paratype USNM 405381, 2.5; f, Interior of lateral plate, paratype USNM 405382, x2.5; g, Interior of lateral plate, paratype USNM 405383, x 2.5. tergal angle forming lateral border of large, deeply impressed, ovoid to triangular, lat- eral depressor muscle pit situated next to basitergal angle; rostral depressor muscle pit long, narrow; upper part of interior of plate between adductor ridge and occludent bor- der markedly ridged or papillate especially in larger specimens; basal margin of larger specimens nodose, nodes reflecting external radial striae. Tergum (Figs. 5a—c, e) relatively thin, nar- row, with acute apex, slightly concave scutal margin and gently convex carinal margin; — Fig. 5. basal margin broadly V-shaped; exterior or- namented by low growth ridges crossed by faint radial striae; tergal furrow partially in- folded, but open from apex to base of spur; tergal spur narrow, relatively short, straight to slightly arched toward scutum, with par- allel sides and gently rounded base; width of spur between one-fifth and one-fourth width of basal margin; distance of spur from basiscutal angle approximately equal to its width; length of spur about one-fifth length of tergum; articular ridge prominent, sharp, arcuate, nearly erect, highest in upper third Balanus neusensis, UNCW locality Z-807, Johnson Point: a, Exterior of scutum, paratype USNM 405373, x13; b, Interior of tergum, paratype USNM 405374, x13; c, Interior of scutum, paratype USNM 405375, x12; d, Interior of scutum, paratype USNM 405376, x14; e-g, Tergum exterior, scutum exterior, scutum interior, holotype USNM 405366, x18. 728 of plate, and extending from apex to, or nearly to, basiscutal angle; articular furrow broad, shallow, crossed by 2 or 3 low, sharp, steeply oblique growth ridges; depressor muscle crests well developed, short, increas- ing in length toward carinal margin, up to 6 in number; interior of tergum between articular ridge and carinal margin markedly rugose, rugosities fading toward basal mar- gin; interior surface of apex marked by sharp, closely-spaced growth ridges. Material examined. —Johnson Point, Neuse River, UNCW locality 807: 37 com- plete shells, one with opercular plates; over 1000 compartmental plates, over 500 oper- cular plates. Dam Creek, Neuse River, UNCW locality 808: 12 compartmental plates, 1 scutum. Woodside, Pender Coun- ty, UNCW locality Z-750: three worn scuta. Disposition of types.—Holotype USNM 405366, paratypes USNM 405368405371 and 405373-405383, and paratype lot 405367 are in the collection of the Depart- ment of Paleobiology, National Museum of Natural History, Washington, D.C. Geologic and geographic distribution. — Early Pleistocene, North Carolina. Etymology.—The specific name is taken from the Neuse River, which is adjacent to the type locality at Johnson Point, Craven County. Discussion.—Balanus neusensis is typi- cally a robust, thick-walled, conic barnacle. There are, however, some rather odd spec- imens from the Johnson Point locality that are small (up to 5 mm in greatest basal di- ameter), laterally asymmetric, and tear-drop shaped, being widest at the rostral, and nar- rowest at the carinal end (Fig. 4c). These specimens are attached to scuta of B. neu- sensis, and their shape is governed by the form of that plate. Because of the consis- tency in asymmetry and shell alignment, it is probable that these small barnacles were growing on scuta of living adults. The shell of B. neusensis, in external as- pect, bears considerable resemblance to that PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of the extant western Atlantic species B. eburneus Gould, but is readily distinguished by the lack of transverse septa in the parietal and basal tubes, and the solid, rather than vesicular sheath. The scutum of B. neusen- sis also resembles that of B. eburneus in bearing prominent external radial striae, but is otherwise quite distinct, being consider- ably narrower, and in having a much longer and more prominent adductor ridge which is not confluent with the base of the articular ridge. The deep, narrow cleft formed be- tween the adductor and articular ridges is a very distinctive feature of B. neusensis. The tergum is very unlike that of B. eburneus, having a well-defined, partially infolded spur furrow, and lacking any indication of an em- bayment on the carinal side of the basal margin. The shell and opercular plates of B. neu- sensis most closely resemble those of B. op- pidieboraci Ross from the Pliocene York- town Formation of Virginia and its equivalents in the Carolinas (see Ross 1964, Zullo 1986). The shells of the two species are virtually identical in form, but can be distinguished by the summits of the radii. Although Ross (1964:490) described the summits of the radii of B. oppidieboraci as varying from 45° to subhorizontal, exami- nation of nearly 100 specimens from the Yorktown Formation of Virginia indicates that oblique radii are very rare in this species and are confined to individual plates of specimens with otherwise subhorizontal summits. The summits of the radii of B. neusensis, on the other hand, are always oblique and are, additionally, crenate, whereas those of B. oppidieboraci are smooth. The scutum of B. oppidieboraci bears finer, more numerous radial striae over almost the entire external surface of the plate, is not sulcate longitudinally, and the growth ridges are not broken into promi- nent nodes. The articular ridge is reflexed over the articular furrow, and its upper sur- face is not flattened. The tergum is broader, VOLUME 99, NUMBER 4 the tergal spur is broader and longer, and the tergal furrow, although open throughout its length, is narrower as a result of infolding from the sides. Balanus neusensis and B. oppidieboraci appear to be related to species of the B. amphitrite Darwin complex and, especially, to the B. pacificus Pilsbry group, which is represented in the Virginia Pliocene fauna by B. pacificus prebrevicalcar Ross. The B. pacificus group is not known in the extant Atlantic fauna, surviving only in the eastern Pacific. It is possible that B. neusensis is a derivative of B. oppidieboraci and the last survivor of the B. pacificus group in the North Atlantic basin. Balanus calidus Pilsbry, 1916 Fig. 4f-j Balanus spongicola var. Darwin, 1854:225, pl. 4, fig. 1d. Balanus calidus Pilsbry, 1916:118, pl. 25, figs. 1-lc, text fig. 32a—f.—Zullo, 1966: 235.—Newman and Ross, 1976:65 (see for complete synonymy). Material examined. —Johnson Point, Neuse River, UNCW locality Z-807; 2 com- plete shells (one with opercular plates), 4 compartmental plates.—Lee Creek Mine, Beaufort County, UNCW locality Z-809: over 200 shells on Ostrea sculpturata, many with opercular plates.—Waccamaw Fm., Little River, South Carolina, UNCW lo- cality Z-812: over 500 shells on Ostrea sculpturata, many with opercular plates. Disposition of specimens.—Figured hy- potype USNM 405372 is in the collection of the Department of Paleobiology, Nation- al Museum of Natural History, Washing- ton, D.C. Remaining specimens are in the collection of the Department of Earth Sci- ences, University of North Carolina at Wil- mington. Geologic and geographic range. — Recent: North Carolina south to the West Indies and the Gulf of Mexico. Pleistocene: North and 729 South Carolina (Wells and Richards 1962; and reported herein); (?) northern Colombia (Nilsson-Cantell 1939). (?) Miocene (Tor- tonian), Bulgaria (Kolosvary 1962). Discussion.—Balanus calidus was the most common balanoid barnacle on the continental shelf of the Carolinas (Zullo 1966), but in recent years has been replaced by the related tropicopolitan species B. tri- gonus Darwin. There is some suggestion that B. trigonus is a recent introduction to the Atlantic coast of the United States (see Wells 1966:84), a suggestion that is supported by the Pleistocene fossil record. Balanus cali- dus is abundant in lower Pleistocene marine deposits of the Carolinas, whereas no spec- imen of B. trigonus has yet been identified positively from the region (see Ross 1965: 273). The Pleistocene shells from the James City Formation have the small orifice, rel- atively narrow radii and ribbed parietes characteristic of B. calidus. The externally striate scutum is rather thick and narrow, and bears a rather short and indistinct ad- ductor ridge. The tergum is distinguished by its broad, basally truncate spur. Literature Cited Connell, J. H., and M. J. Keough. 1985. Disturbance and patch dynamics of subtidal marine animals on hard substrata. In S. T. A. Pickett and P. S. White, eds., The ecology of natural disturbance and patch dynamics. Academic Press, Orlando, pp. 125-151. Cronin, T. M., L. M. Bybell, R. Z. Poore, B. W. Black- welder, J. C. Liddicoat, and J. E. Hazel. 1984. Age and correlation of emerged Pliocene and Pleistocene deposits, U.S. Atlantic Coastal Plain.—Palaeogeography, Palaeoclimatology, Palaeoecology 47:21-51. Darwin, C. 1854. A monograph on the sub-class Cir- ripedia, with figures of all the species. The Ba- lanidae, the Verrucidae, etc. Ray Society, Lon-~ don, 684 pp. DuBar, J. R., and J. F. Howard. 1969. Paleoecology of the James City Formation (Plio-Pleisto- cene?), Neuse River estuary, North Carolina. — Geological Society of America Abstracts with Programs, pt. 4:20. , and J. R. Solliday. 1963. Stratigraphy of the 730 Neogene deposits, lower Neuse estuary, North Carolina.— Southeastern Geology 4:213-233. ——., J. R. Solliday, and J. F. Howard. 1974. Stra- tigraphy and morphology of Neogene deposits, Neuse River estuary, North Carolina. Jn R. Q. Oaks and J. R. DuBar, eds., Post-Miocene stra- tigraphy, central and southern Atlantic Coastal Plain. Utah State University Press, Logan, pp. 102-122. Henry, D. P., and P. A. McLaughlin. 1975. The bar- nacles of the Balanus amphitrite complex (Cir- ripedia, Thoracica).— Zoologische Verhandelin- gen 141:1-254. Hoagland, K. D. 1979. The behavior of three sym- patric species of Crepidula (Gastropoda: Pro- sobranchia) from the Atlantic, with implications for evolutionary ecology.— The Nautilus 94:143— 149. Johnson, J. K. 1972. Effect of turbidity on the rate of filtration and growth of the slipper limpet, Crepidula fornicata Lamarck, 1799 [sic.].—The Veliger 14:315-—320. Kolosvary, G. 1955. Uber stratigraphischer Rolle der fossilen Balaniden.—Acta Universitatis Szege- densis, Acta Biologica, nova series I(1—4):183- 188. 1959. A statistical study of the Miocene bal- anids from Hungary.—Journal of Paleontology 33(1):196-198. . 1961la. Einige interessante Balanus-funde aus der Burdigalienstuff von Turkmenistan.— Acta Universitatis Szegedensis, Acta Biologica, nova series VII(1—2):99-102. . 1961b. Further fossile [sic] balanids from the USSR.—Acta Universitatis Szegedensis, Acta Biologica, nova series VII(3—4):149-154. 1962. Balanids from the Bulgarian Tertiary age.— Annuaire de |’Université de Sofia, Livre 2, Geologie 55:85-89. McCartan, L., J. P. Owens, B. W. Blackwelder, B. J. Szabo, D. F. Belknap, N. Kriausakul, R. M. Mitterer, and J. F. Wehmiller. 1982. Com- parison of amino acid racemization geochro- nometry with lithostratigraphy, biostratigraphy, uranium-series coral dating, and magnetostra- tigraphy in the Atlantic Coastal Plain of thé southeastern United States.—Quaternary Re- search 18:337-359. Miller, W., II]. 1985. The Flanner Beach Formation (middle Pleistocene) in eastern North Caroli- na.— Tulane Studies in Geology and Paleontol- ogy 18:93-122. 1986. Paleoecology of benthic community replacement.— Lethaia 19(3):225-231. , and J. R. DuBar. [in prep.] Benthic com- munity replacement in early Pleistocene James City Formation, North Carolina Coastal Plain. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Mixon, R. B., and Pilkey, O. H. 1976. Reconnais- sance geology of the submerged and emerged Coastal Plain province, Cape Lookout area, North Carolina.—U.S. Geological Survey Professional Paper 859:1-45. Newman, W. A., and A. Ross. 1976. Revision of the balanomorph barnacles; including a catalog of the species.—San Diego Society of Natural His- tory, Memoir 9:1—108. Nilsson-Cantell, C. A. 1939. Recent and fossil bal- anids from the north coast of South America. — Capita Zoologica 8(4):3-7. Parker, R. H. 1976. Classification of communities based on geomorphology and energy levels in the ecosystem. /n R. W. Scott and R. R. West, eds., Structure and classification of paleocom- munities. Dowden, Hutchinson and Ross, Stroudsburg, Pennsylvania, pp. 67-86. Pilsbry, H. A. 1916. The sessile barnacles (Cirripedia) contained in the collections of the United States National Museum; including a monograph of the American species.— Bulletin of the United States National Museum 93:1-366. Ray, C. E. [ed.] 1983. Geology and paleontology of the Lee Creek Mine, North Carolina, I.—Smith- sonian Contributions to Paleobiology 53:1—529. Ross, A. 1964. Cuirripedia from the Yorktown For- mation (Miocene) of Virginia.—Journal of Pa- leontology 38(3):483—491. 1965. A new barnacle from the Tamiami Miocene.— Quarterly Journal of the Florida Academy of Sciences 27(4):273-277. Sousa, W. P. 1985. Disturbance and patch dynamics on rocky intertidal shores. Jn S. T. A. Pickett and P. S. White, eds., The ecology of natural disturbance and patch dynamics. Academic Press, Orlando, pp. 101-124. Wells, H. W. 1966. Barnacles of the northeastern Gulf of Mexico.— Quarterly Journal of the Flor- ida Academy of Sciences 29(2):81—95. Wells, H. W., and H. G. Richards. 1962. Invertebrate fauna of coquina from the Cape Hatteras re- gion.—Journal of Paleontology 36(3):586—-591. Zullo, V. A. 1966. Thoracic Cirripedia from the con- tinental shelf off South Carolina, U.S.A. —Crus- taceana | 1(3):229-244. 1986. Pliocene barnacles (Cirripedia, Tho- racica) from South Carolina, with a key to Plio- cene balanoids of the eastern United States.— South Carolina Geology 29:1-18. (VAZ) Department of Earth Sciences, University of North Carolina at Wilming- ton, North Carolina 28403; (WM) Depart- ment of Geology, Humboldt State Univer- sity, Arcata, California 95521. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 731-734 THE SYNONYMY OF CIROLANA TUBERCULATA (RICHARDSON, 1910) GSOPODA: FLABELLIFERA: CIROLANIDAE) Paul M. Delaney Abstract.—The syntypes of Alcirona tuberculata Richardson, 1910 (Coral- lanidae) were examined and determined to be more appropriately placed in the genus Cirolana Leach, 1818 (Cirolanidae). The species is redescribed here as Cirolana tuberculata. Cirolana Leach, 1818 Diagnosis.—Rostral process minute or absent. Frontal lamina flat. Antennular pe- duncle 2- or 3-articulate; articles 1 and 2 of antennule not forming right angles to one another (as in Eurydice). Antenna peduncle 5-articulate. Mandible with broad triden- tate incisor; lacinia mobilis and molar pro- cess well-developed. Maxilliped usually with 1-3 coupling hooks (occasionally more) on endite. Pereopod dactyls with small sec- ondary ungues. Pleonite 1 often concealed by pereonite VII; pleonite 5 narrower than 1-4, with lateral margins covered by pleo- nite 4. Pleopods 1 and 2 similar to each other; endopod of pleopod 5 without setae. Appendix masculinum arising basally from endopod of pleopod 2. Inner angle of uro- pod peduncle strongly produced; uropod rami with or without plumose marginal se- tae. (Modified after Brusca and Iverson 1985.) Cirolana tuberculata (Richardson, 1910) Figs. 1, 2 Alcirona tuberculataRichardson, 1910:8, figs. 7a,b. Material examined. — Philippine Islands; U.S. Bureau of Fisheries Steamer Albatross, sta 5141, Jolo Light, 5.5 mi. S 17 E, (6°09’00"N, 120°58'00”E) 53 m, “in coral and sand” USNM No. 40910, syntypes 1 male, 7.5 mm 1 female, 8.0 mm. 15 Feb 1908. Description of male.—Cephalon without tubercles; eyes lateral (Fig. 1a). Antennular peduncle 2-articulate, flagellum 14-articu- late, with simple setae and aesthetascs (Fig. 2c); extending to midlength of pereonite I (Fig. 1a). Antennal peduncle 5-articulate, 4— 5 elongate; flagellum 20-articulate (Fig. 2a), extending to midlength of pereonite III (Fig. 1c). Frontal lamina flat, pentagonal; clypeus short, broad; labrum short, broad, quad- rangular (Fig. 2b). Mandible incisor broad; lacinia mobilis lobular, rounded, with sim- ple setae and 3 distal spines; molar process triangular, with 12 spines and many “imple setae; 3-articulate palp arising from middle of mandible, middle article with simple marginal setae, distal article with simple and comb-like setae (Fig. 2d). Maxillule, lateral lobe with 10 apical spines, medial lobe with 3 large plumose spines (Fig. 2g). Maxilla, lateral lobes with 4 and 9 simple setae; me- dial lobe with 6 simple and 6 plumose setae (Fig. 2e). Maxillipedal palp with simple, bi- fid and comb-like setae as figured; endite extending to second palp article, with 1 cou- pling hook, apex with 2 plumose and 1 sim- ple setae (Fig. 2f). Pereon length 5 times pleon length; pere- on widest at pereonite V; pereon without dorsal tubercles or setae. Coxal plates large, increasing in size and acuteness posteriorly; 732 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Cirolana tuberculata: a, Dorsal view, pleotelson and uropod setae and spines not shown; b, Pleotelson and uropods; c, Lateral view; d, Pereopod I; e, Pereopod III; f, Pereopod VII. coxae IV—VII extending beyond posterior no spines or setae (Fig. 1c). Pereopods I-III margins of their respective pereonites; cox- prehensile, dactyl with small secondary un- ae VII extending to midlength of pleonite guis, merus with 4—5 short, blunt spines on 2; coxae II-VII with single oblique carina, medial margin; simple, bifid and plumose VOLUME 99, NUMBER 4 733 Fig. 2. Cirolana tuberculata: a, Antennal peduncle; b, Frontal lamina, clypeus and labrum; c, Antennule; d, Mandible; e, Maxilla; f, Maxilliped; g, Maxillule; h, Pleopod 1; 1, Pleopod 2. 734 setae as figured (Fig. ld, e). Pereopods [V- VII with small secondary unguis, and short blunt spines on medial margins; simple, bi- fid and plumose setae as figured (Fig. 1f). Pleon widest at pleonites 3—4; pleonite 1 covered by pereonite VII; lateral margins of pleonite 5 overlapped by pleonite 4; pleo- nites 1-5 without dorsal setae or tubercles. Pleopods with plumose marginal setae (ex- cept endopod of pleopod 5), peduncles with 3 coupling spines and 2—4 plumose setae on medial margins (Fig. 2h, i). Male pleopod 2 with rodlike appendix masculinum arising from proximal medial margin of endopod (Fig. 21). Pleotelson triangular, lateral mar- gins slightly concave; apex with simple setae and 7 spines. Pleotelson dorsum with two parallel rows of 4 tubercles, decreasing in size posteriorly (Fig. 1b). Uropods extend- ing well beyond posterior margin of pleo- telson; exopod one-half width of endopod; apex acute, margins with setae and 7-8 spines; endopod broadly rounded, margins with setae and 11 spines. Uropodal pedun- cle, inner margin with 3-5 simple and 2 plumose marginal setae (Fig. 1b). Female.—Similar to male but with less distinct tuberculation on pleotelson. Remarks. — Richardson (1910) described this species as Alcirona tuberculata from two specimens collected in the Philippines, but figured only the posterior portion of the body and pereopod I of the male. The genus A/- cirona Hansen, 1890, is distinguished from other genera in the family Corallanidae pri- marily by mouthpart morphology, includ- ing characters of the maxillule (lateral lobe with 2 large recurved spines, and occasior+ ally 1-3 smaller spines between the 2 large ones), maxilla (short rounded lobe), and maxilliped (palp articles of approximately equal length, endite absent). For a key to the genera of Tridentellidae and Corallan- idae see Delaney and Brusca (1985). The mouthparts of this species (Fig. 2d—g) show that it clearly does not belong in Alcirona but to Cirolana (Cirolanidae). The linear pattern of pleotelson tuber- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON culation in Cirolana tuberculata closely re- sembles that of C. pleonastica Stebbing, (1900:629, pl. LX VIIa) from New Britain in the southwest Pacific. Cirolana tubercu- lata differs from that species in number of pleotelson tubercles (4 pairs in C. tubercu- lata, 5 pairs in C. pleonastica), lack of dorsal pleotelson setae, lack of pereonal and pleon- al tuberculation, number of coupling spines on peduncles of pleopods 1—2 (3 in C. tuber- culata, 4 in C. pleonastica) and shape of labrum and maxilliped. Acknowledgments This study was done during a 1984 Grad- uate Fellowship in the Division of Crusta- cea, National Museum of Natural History, Smithsonian Institution. Literature Cited Brusca, R. C., and E. W. Iverson. 1985. A guide to the marine isopod Crustacea of Pacific Costa Rica.—Revista de Biologia Tropical, Univer- sidad de Costa Rica 33, Supplement 1:1—77. Delaney, P. M., and R. C. Brusca. 1985. Two new species of Tridentella Richardson, 1905 (Isop- oda: Flabellifera: Tridentellidae) from Califor- nia, with a rediagnosis and comments on the family, and a key to the genera of Tridentellidae and Corallanidae.—Journal of Crustacean Bi- ology 5(4):728-742. Hansen, H. J. 1890. Cirolanidae et familiae nonnul- lae propinquae Musaei Hauniensis. — Kongelige Danske Videnskabernes Selskabs Skrifter, 6 Raekke, Naturvidenskabelig og Mathematisk Afdeling 3:237—426. Leach, W. E. 1818. Cymothoadées. Jn Levrault, F. G., Dictionnaire des Sciences Naturelles 12:338— 354. Paris and Strasbourg. Richardson, H. R. 1910. Marine isopods collected in the Philippines by the U.S. Fisheries steamer Albatross in 1907-1908. Bureau of Fisheries Document No. 736:1-44. Stebbing, T. R. R. 1900. On Crustacea brought by Dr. Willey from the South Seas. Zoological re- sults based on material from New Britain, New Guinea, Loyalty Islands and elsewhere collected during the years 1895, 1896, and 1897. Part 5, pp. 605-690. Cambridge University Press. Division of Life Sciences, Los Angeles County Museum of Natural History, Los Angeles, California 90007. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 735-738 NOTES ON THE CRAYFISH PROCAMBARUS (ORTMANNICUS) XILITLAE (DECAPODA: CAMBARIDAE) Horton H. Hobbs, Jr. and Andrew G. Grubbs Abstract. — With the acquisition of the exuviae of a first form male revealing the structure of the secondary sexual features (described and illustrated herein), the affinities of the troglobitic crayfish Procambarus (Ortmannicus) xilitlae are reassessed. Its tentative assignment to the subgenus Scapulicambarus by Hobbs and Grubbs (1982) was found to be in error when it was discovered that a shoulder was lacking on the cephalic surface of the first pleopod of the first form (“‘adult’’) male. Procambarus (Scapulicambarus) xilitlae Hobbs and Grubbs, 1982, was described from Hoya de las Guaguas, near the town of Xilitla, San Luis Potosi, Mexico. The au- thors based their description on a second form male and two females, and they ten- tatively assigned the crayfish to the subge- nus Scapulicambarus, making the state- ment that “The availability of a first form male of this new crayfish would greatly strengthen our confidence in the inferences that we have made in assessing its relation- ship to other crayfishes.’’ While they were confident that it is a member of the genus Procambarus, they were “‘. . . less certain as to which of the subgenera it should be as- signed, Ortmannicus or Scapulicambarus”’ (Hobbs and Grubbs 1982:49). The only feature that serves consistently to separate the members of these two sub- genera is the presence of a strong, usually angular shoulder on the cephalic surface of the left first pleopod of the male; the shoul- der on the right pleopod is folded cau- domesially and pressed against the mesial face of the appendage. Ignoring the shoul- der, the two subgenera seem clearly to merge through forms like Procambarus (O.) ever- manni (Faxon, 1890), P. (O.) xilitlae, P. (S.) paeninsulanus (Faxon, 1914), and P. (S.) strenthi Hobbs (1977). A shoulder is present in the second form male of all members of Scapulicambarus, but it is often much weaker than that of first form individuals (see figs. 280 to 284 in Hobbs 1974). The small bulge on the cephalic surface of the pleopod of the holotypic male, form II, of P. (O.) xilitlae (see fig. 1b in Hobbs and Grubbs 1982) appeared to represent such a shoulder, thus prompting the original as- signment of this crayfish to the subgenus Scapulicambarus. With the rearing of a second form male in the laboratory by one of us (AGG) and preserving its exuviae, one of which was shed when molting from form I to form II (see below), we discovered that the first pleopod lacks a shoulder. thus the species must be transferred to the subgenus Ort- mannicus. The absence of pigment from the body and eyes, the non-faceted cornea, the pres- ence on the first pleopod of the first form male of a well developed caudal process, and the absence of an adventitious process will serve to distinguish this crayfish from all others. A description and illustrations of the secondary sexual and certain other fea- tures of the first form exuviae of this topo- typic male are offered to complete the basic description of this crayfish. 736 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Procambarus (Ortmannicus) xilitlae (all from exuviae of first form male): a, Mesial view of first pleopod; b, Caudal view of first pleopods (proximal parts probably shrunken, but asymmetry real); c, Proximal podomeres of third, fourth, and fifth pereiopods; d, Lateral view of first pleopod; e, Distal podomeres of second pereiopod; f, Same of third pereiopod; g, Same of fifth pereiopod; h, Same of fourth pereiopod; i, Same of first pereiopod; j, Lateral view of distal part of first pleopod without subapical setae. VOLUME 99, NUMBER 4 Procambarus (Ortmannicus) xilitlae Fig. | Procambarus (Scapulicambarus) xilitlae Hobbs and Grubbs, 1982:45—50, 1 fig. Male, Form I.—Chela (Fig. 11) differing from that of holotype in following respects: mesial margin of palm with row of 7 small tubercles; opposable margin of fixed finger with row of 8 small corneous teeth along proximal third; larger, more ventrally sit- uated one as in holotype; fifth tubercle from base largest of 8 present on proximal third of opposable margin of dactyl. Ischia of third and fourth pereiopods with simple hooks (Fig. 1c), that on third overreaching bas- ioischial suture; neither opposed by tubercle on basis. First pleopods (Fig. la, b, d, j) reaching coxae of third pereiopods, distinctly asym- metrical but contiguous basally; cephalic surface lacking shoulder; distal extremity bearing large slender tapering mesial pro- cess directed distolaterally; cephalic process short, with corneous acute apex, curved caudodistally parallel to short, corneous, acute, beaklike central projection; caudal process corneous, short, wedgelike, its distal margin sloping cephalodistally to angle, apex of which directed toward tip of central pro- jection. Cephalic process and central pro- jection overreaching apex of caudal process but falling short of that of mesial process. Sockets of distally directed subapical setae forming arc on lateral surface just proximal to bases of terminal elements. Relationships. — As pointed out in the in- troductory remarks, this crayfish is allied to those species that appear to bridge the gap between the subgenera Ortmannicus and Scapulicambarus. Seemingly it is rather dis- tantly related to the other troglobitic mem- bers of Ortmannicus which populate much of the peninsula and eastern part of the pan- handle of Florida even though the annulus ventralis is much like those of P. (O.) horsti Hobbs and Means (1972) and P. (O.) orci- nus Hobbs and Means (1972). Among the 737 Mexican crayfishes, it shares more in com- mon with P. (O.) caballeroi Villalobos (1944), P. (O.) acutus cuevachicae (Hobbs, 1941), P. (S.) strenthi, and P. (Pennides) roberti Villalobos and Hobbs (1974) than with any others. Remarks.—The topotypic specimen from which the exuviae of the first form was ob- tained was collected on 26 November 1981, and, when brought into the laboratory, had a carapace length of 26.8 (postorbital car- apace length, 21.7) mm. It underwent three molts, remaining in the second form, with- out any increase in the total length of the carapace. The dates on which the first two exuviae were removed from the aquarium were not recorded, but the third was found on 5 November 1982. Another test was dis- covered in the aquarium on 23 February 1983 in which the corresponding lengths of the carapace were 27.8 and 22.0 mm. The next was removed on 29 July 1983, but it was so torn that accurate measurements could not be made. Presumably it was at this molt that the transformation to form I was accomplished, for the next exuviae, found on 12 December 1983, furnished the models for the illustrations depicted herein. The corresponding lengths of the carapace were 29.2 and 23.1 mm. The crayfish, found dead on 6 November 1984, had reached a stage of decomposition such that measure- ments of the carapace could not be made. All of these exuviae and the partly decom- posed remains, in form II, are in the col- lection of the Smithsonian Institution. To our knowledge, other than the type series and the specimen cited above, only five specimens of P. (O.) xiiitlae have been collected. These were obtained at the type locality by R. Rhorer and AGG on 4 August 1982 and deposited in the Smithsonian In- stitution (1 II, 1 js, 1 j2) and the Texas Memorial Museum (1 Q, | jé). Acknowledgments For their criticisms of the manuscript, we extend our thanks to Horton H. Hobbs III 738 of Wittenberg University, Brian Kensley of the Smithsonian Institution, and Douglas G. Smith of the University of Massachu- setts. Literature Cited Faxon, Walter. 1890. Notes on North American Crayfishes, Family Astacidae.— Proceedings of the United States National Museum 12(785): 619-634. 1914. Notes on the crayfishes in the United States National Museum and the Museum of Comparative Zoology, with descriptions of new species and subspecies.—Memoirs of the Mu- seum of Comparative Zoology at Harvard Col- lege 40(8):351—427, 13 plates. Hobbs, Horton H., Jr. 1941. A new crayfish from San Luis Potosi, Mexico (Decapoda, Asta- cidae).— Zoologica 26(1):1—4. 1 figure. 1974. A checklist of the North and Middle American crayfishes (Decapoda: Astacidae and Cambaridae).—Smithsonian Contributions to Zoology 166:i1i + 161 pages, 294 figures. 1977. A new crayfish (Decapoda: Cambari- dae) from San Luis Potosi, Mexico. —Proceed- ings of the Biological Society of Washington 90(2):412-419, 2 figures. —, and Horton H. Hobbs, Jr. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON , and Andrew G. Grubbs. 1982. Description of a new troglobitic crayfish from Mexico and a list of Mexican crayfishes reported since the publication of the Villalobos monograph (1955) (Decapoda, Cambaridae).— Association for Mexican Cave Studies Bulletin 8:45—SO0, 1 fig- ure/Texas Memorial Museum Bulletin 28:45-— 50, 1 figure. ———.,, and D. Bruce Means. 1972. Two new trog- lobitic crayfishes (Decapoda, Astacidae) from Florida.— Proceedings of the Biological Society of Washington 84(46):393—409, 2 figures. Villalobos Figueroa, Alejandro. 1944. Estudios de los cambarinos mexicanos, II: Dos especies nuevas del género Paracambarus. — Anales del Instituto de Biologia, Universidad Nacional Autonoma de México 15(1):161—174, 2 plates. 1974. Three new crustaceans from La Media Luna, San Luis Po- tosi, Mexico.—Smithsonian Contributions to Zoology 174:1-18, 8 figures. (HHH) Department of Invertebrate Zo- ology, Smithsonian Institution, Washing- ton, D.C. 20560; (AGG) Route 2, Box 279A, San Marcos, Texas 78666. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 739-747 A NEW SPECIES OF ELACHOCHARAX (TELEOSTEI: CHARACIDAE) FROM THE RIO NEGRO REGION OF VENEZUELA AND BRAZIL Stanley H. Weitzman Abstract.—Elachocharax mitopterus is described as a new species of the characid fish subfamily Characidiinae from the Rio Negro system of Venezuela and Brazil. The new species is distintuished from Elachocharax pulcher Myers, E. junki (Géry), and E. geryi Weitzman and Kanazawa by a variety of color pattern, morphometric, and meristic characters outlined in the key and text. Elachocharax pulcher is the sister group of the other three species which form an unresolved trichotomy. Although some of these species have seemingly allopatric distributions, it is probable that their geographic ranges broadly overlap. The three previously known species of Elachocharax Myers (1927:114) were treat- ed most recently by Weitzman and Kana- zawa (1978) and Weitzman and Géry (1981). The phylogeny of Elachocharax was dis- cussed and a cladogram of the relationships of the recognized species was presented by Weitzman and Géry (1981:898-910). The genus is not confined to the Amazon region as stated by Géry (1984:356) since two species, Elachocharax pulcher Myers (1927: 115) and Elachocharax geryi Weitzman and Kanazawa (1978:173), previously were re- ported from both the Orinoco and Amazon basins. Elachocharax junki (Géry, 1971: 154) and the new species, Elachocharax mi- topterus, are so far known only from Am- azon drainages, but both species occur in the Rio Negro (the new one in the upper portion of that system) and may be expected to be found at least in black water streams of the upper portions of the Rio Orinoco. Methods and Materials Counts and measurements are recorded as described by Fink and Weitzman (1974: 1-2). Body depth is measured vertically from dorsal-fin origin. All measurements other than standard length (SL) are expressed as a percentage of SL except subunits of head which, unless otherwise stated, are recorded as a percentage of head length. Total ver- tebral counts, taken from radiographs and from one cleared, Alizarin and Alcian blue stained specimen, include all vertebrae of Weberian apparatus and with fused PU + U of caudal skeleton counted as a single vertebra. In text and table, count or mor- phometric character given first is of holo- type unless otherwise noted; next series of figures is range of all specimens (given in parentheses in text). This is followed by mean (xX) of all specimens. Specimens ex- amined for this study are deposited in Mu- seo de Biologia, Instituto de Zoologia Trop- ical, Universidad Central de Venezuela (MBUCV-V), Museu de Zoologia da Uni- versidade de Sao Paulo (MZUSP), and Na- tional Museum of Natural History, Smith- sonian Institution (USNM). Artificial Key to the Species of Elachocharax Myers 1. Premaxillary teeth tricuspid ante- riorly followed by a few bicuspid and/or unicuspid teeth. Dentary with tricuspid teeth anteriorly fol- lowed by none, one, or a few bicus- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. partamento Rio Negro, Territorio Federal Amazonas, Venezuela, 5 Dec 1984. N pid teeth and then several unicuspid teeth lateroposteriorly. Adipose fin present Premaxillary teeth and dentary teeth all unicuspid. Adipose fin absent Total dorsal-fin rays 17 or 18, usu- ally 18. Scales in lateral series to hy- pural joint 24 or 25. Branchiostegal rays 5. Horizontal scale rows be- tween dorsal- and pelvic-fin origins 7. Total vertebrae 29 Ai ane Elachocharax pulcher As bea getteti Elachocharax mitopterus Total dorsal-fin rays 19 to 22, usu- ally 21 or 22. Scales in lateral series 27 to 33, usually 28 to 32. Bran- chiostegal rays 4. Horizontal scale rows between dorsal- and pelvic-fin origins 8. Total vertebrae 31 to 33, usually 31 or 32 . Caudal fin without obvious vertical dark bars. Teeth on ectopterygoid 8 to 13. Inner row dentary teeth 19 to 20. Upper limb gill rakers 5 to 6. Anterior anal-fin origin in advance ofa vertical line drawn through dor- sal-fin base. Snout length about 5.4 to 6.1% of SL. Caudal peduncle length about 19.0 to 22.0% of SL shh sageltcge siege anak. Elachocharax junki Elachocharax mitopterus, new species, holotype MBUCV-V-15270, SL 13.9 mm, Cano Chola, De- — Caudal fin with 13 or 14 narrow ver- tical dark bars. Teeth on ectopter- ygoid 4 or 5. Inner row dentary teeth 13 to 16. Upper limb gill rakers 3. Anal-fin origin at or slightly poste- rior to a vertical line drawn through posterior termination of dorsal-fin base. Snout length about 6.0 to 8.0% of SL. Caudal peduncle length about 16.0 to 19.0% SL. .Elachocharax geryi Elachocharax mitopterus, new species Figs. 1-3, Table 1 Holotype. —MBUCV-V-15270, SL 13.9 mm, Venezuela, Territorio Federal Ama- zonas, Departmento Rio Negro, Cano Cho- la, where crossed by road from San Carlos de Rio Negro to Solano, about 01°58'N, 67°00’W, 5 Dec 1984, R. P. Vari, A. Ma- chado-Allison, C. Ferraris, O. Castillo, J. M. Fernandez. Paratypes. —1, MBUCV-V-15271, SL 13.8 mm.—2, USNM 270147, SL 12.3-12.7 mm (jaws and opercle of right side of one specimen cleared and stained).—1, cleared and stained, USNM 274101, SL 12.6 mm. All preceding paratypes with same collec- tion data as holotype.—3, MZUSP 29871, VOLUME 99, NUMBER 4 741 Fig. 2. Elachocharax mitopterus, new species, from a paratype, USNM 274101, SL 12.6 mm, Cano Chola, Departamento Rio Negro, Territorio Federal Ama- zonas, Venezuela, 5 Dec 1984. Lateral view of pre- maxilla, maxilla, and dentary of right side; anterior to right. Outer dentary tooth row lies in a deep groove. Replacement teeth of this row develop within groove. Inner-row dentary teeth present posterior to or near visible dorsal border of ridge seen internal to outer- row dentary teeth. Note that third dentary tooth from midline partly eroded away and its replacement tooth (to its left) easily visible posterior and interior to it. Anterior to right. SL 11.7—13.2 mm, Brazil, Amazonas, igara- pe Tarumazinho, about 45 km north of Manaus on road between Manaus, Ama- zonas and Caracarai, Roraima, about 2°42’S, 60°02’W, 25 Sep i976, Ivanzier Vieira. —2, USNM 274100, SL 12.7—13.3 mm (jaws and opercle of right side of one specimen cleared and stained), with same collection data as MZUSP 29871. Diagnosis. — Distinguished from all other species of Elachocharax by having 29 in- stead of 30 to 33 vertebrae, 7 instead of 8 scale rows between anterior base of dorsal fin and pelvic fin, and 24 to 26 scales in lateral series instead of 27 to 33. This species, Fig. 3. Elachocharax mitopterus, new species, from a paratype, USNM 274101, SL 12.6 mm, Cano Chola, Departamento Rio Negro, Territorio Federal Ama- zonas, Venezuela, 5 Dec 1984. Medial view of right dentary. Seventeen fully-developed inner-row teeth visible on dentary ridge. Five replacement teeth present interior to inner-row teeth. These teeth point inwards and not fully or not at all attached to dentary ridge. Full complement of inner-row teeth about 19, counting in-place fully-developed teeth and their replacement teeth as one. Anterior to left. E. geryi, and E. junki are distinguished from E. pulcher by having only conic teeth in jaws (no tricuspid or bicuspid teeth). Elacho- charax mitopterus is similar to E. geryi and especially E. pulcher but unlike E. junki in having two strong black pigment bars on anal fin and one on pelvic fin. Elachocharax mitopterus has only five narrow dark bars on base of caudal fin rather than 13 to 14 of these bars as in E. geryi. As in E. geryi, anal fin of E. mitopterus originates at or near imaginary vertical line drawn through pos- terior basal termination of dorsal fin rather than anterior to such a line as in E. junki or slightly posterior to one as in E. pulcher. Description. —See Table 1 for morpho- metric values. Body relatively short, as in other species of Elachocharax; greatest depth at dorsal-fin origin. Profile of head and back from snout tip to dorsal-fin origin gently convex. Dorsal-fin origin in advance of imaginary vertical line drawn through pelvic-fin origin. Anal-fin origin at or near imaginary vertical line drawn through pos- terior termination of dorsal-fin base. Profile of ventral surface of head somewhat con- vex. Belly slightly convex or nearly flat from 742 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Morphometrics of two population samples of E/lachocharax mitopterus. Standard length expressed in mm. Other measurements are percentages of standard length except for eye diameter, snout length, and interorbital width which are percentages of head length. Venezuela, Cano Chola Brazil, igarapé Tarumazinho Holotype Range, n = 5 re Range, n = 5 x Standard length 13.9 12.3-13.9 11.7-13.3 Depth at dorsal-fin origin 30.2 29.7-30.9 30.4 29.5—30.8 30.4 Snout to dorsal-fin origin SB 52.2-54.3 53.1 52.0-53.1 D257) Snout to pelvic-fin origin 57.6 55.8-58.3 S72 56.3-59.0 fle! Snout to anal-fin origin 79.1 78.3-79.1 78.8 78.0-79.7 78.7 Caudal peduncle depth 14.1 13.8-15.4 14.7 15.2-16.5 16.0 Caudal peduncle length 15.1 15.1-17.1 16.2 14.2-16.7 15.0 Pectoral-fin length 32.4 32.4-37.8 34.7 32.3-36.7 34.9 Pelvic-fin length 26.6 23.2-26.8 25.3 21.2-23.6 22.8 Bony head length 31.7 31.2-33.9 37? 31.4-33.8 B22 Horizontal eye diameter 33.0 31.4-33.0 32.5 32.2-35.1 34.0 Snout length 18.2 15.2-18.2 16.8 17.9-20.0 18.8 Least width interorbital 34.1 27.9-34.1 31.2 30.5-33.8 30.5 region of isthmus to pelvic-fin origin. Profile of body between pelvic-fin origin and anal- fin origin gently convex to nearly straight. Caudal peduncle short, deep. Both dorsal and ventral profiles of caudal peduncle slightly concave. Head moderately long. Mouth terminal. Snout relatively blunt; eyes large, horizontal diameter exceeds snout length. Teeth on dentary in two rows (Figs. 2, 3). Outer row with 11 (12-13) conic teeth. In- ner row uncounted in holotype (18-19, 3 Alizarin preparations) conic teeth, Figs. 3, 4. Premaxilla with single row of 9 (12-12, 3 Alizarin preparations) conic teeth. Max- illa toothless. Ectopterygoid teeth uncount- ed in holotype (4—9, 3 Alizarin preparations) conic teeth in single row. Branchiostegal rays uncounted in holoe type (5 in 3 Alizarin preparations); two slen- der anterior rays attached to anterior part of ceratohyal followed by broad ray at- tached to posterior part of ceratohyal; fourth broad ray attached to epihyal. Gill rakers 4 (3-4) on dorsal limb and 8 (7-8) on ventral limb. Range of gill raker counts based on 3 Alizarin preparations. Frontal-parietal fo- ramen greatly reduced. A narrow antorbital bone present dorsal and posterolateral to posterior and dorsal portion of anterior (first) infraorbital. First infraorbital with latero- sensory canal and anterior process extend- ing dorsal and anterior to anterior dorsal slender process of maxillary bone. Second infraorbital bone primarily a laterosensory tube but some laminar bone present. Other infraorbital bones absent. Scales cycloid, usually 4-5 radii on ex- posed field. Lateral line with 5 (5-6, x = 5.1, n = 10) perforated scales. Scales in lat- eral series 26 (24-26, X = 25.2, n = 10). Horizontal scale rows between dorsal and pelvic fins 7 (7, n = 10). Horizontal scale rows around caudal peduncle 10 (10, n = 5 in specimens from type locality; 12-14, X = 12.8, n = 5 in specimens from igarapé Tarumazinho). (See section on population variation.) Predorsal scales 8 (8-9, x = 8.3, n = 10). Dorsal-fin rays ii, 16 (11,15, n = 2, 11,16, n = 8); total dorsal-fin rays 18 (17-18, x = 17.8, n = 10). Last dorsal-fin ray not split to its base. Adipose fin absent. Externally visible anal-fin rays ii,16 in all specimens. Extremely small, but not externally visible, third anterior unbranched ray revealed in 9 of 10 radiographed and cleared and stained specimens. Posterior anal-fin ray split to its VOLUME 99, NUMBER 4 base. Pectoral fin elongate, reaching to or beyond midlength of pelvic fin when both fins are laid against body. Pectoral-fin rays usually undivided, sometimes rays three and/or four with some division. Total rays 6 (6-7, X = 6.5, n = 10). Pelvic fin 1,7 in all specimens. Pelvic fins reach somewhat pos- terior to anal-fin origin, reaching level of second or third anal-fin rays. Caudal fin forked, principal ray count 10/9 in all spec- imens. Total number of vertebrae including We- berian apparatus 29, n = 10. Color in alcohol. — Background body col- or pale yellowish-brown. About ten irreg- ular, rather indefinite dark vertical bars present on body; see Fig. 1 of holotype. These nearly absent on somewhat faded paratype from igarapé Tarumazinho, Ama- zonas. Elachocharax mitopterus also shows variability in body pigment noted by Weitz- man and Géry (1981:898) for other three species of Elachocharax. Approximately ten dark vertical body bars of E. mitopterus similar to those of E. pulcher (8-10) and E. geryi (10-12) rather than those found in E. junki (16 to 18). Dark bars on body of E. mitopterus more distinct posteriorly than anteriorly and continuous with dark pig- ment of pelvic and anal fins. Belly with same color pattern as body sides. Top of head pale brown to very dark brown. A dark bar from dorsoposterior bor- der of eye to near nape present as in other species of Elachocharax. Snout pale to dark brown dorsally, a prominent dark stripe ex- tends from oral border of premaxilla and anterior part of maxilla to anterior border of eye. Stripe complete in holotype (Fig. 1) but in some specimens stripe does not ex- tend over premaxilla to snout tip although mottled dark pigment is present in this re- gion. Anterior tip of lower jaw dark brown. Dark brown chromatophores ventral to eye organized into one broad or two more or less distinct, short vertical bars. Opercle with scattered dark chromatophores which may be expanded and coalesced into variable 743 blotches. As on body, pigment of head var- les according to expansion or contraction of dark brown chromatophores. Ventral sur- face of head also pale to dark, depending on chromatophore expansion. Caudal fin with none (faded?) to as many as five narrow vertical bars. Bars occur on basal area of fin and are absent distally. Pigment of these bars occurs on rays or at least along ray borders. Posterior border of three large ter- minal scales over base of caudal fin usually broadly covered with dark pigment (Fig. 1), producing a heavy, irregular vertical bar in addition to five or so narrow vertical bars on fin rays. Intraradial membranes of cau- dal fin nearly without pigment except im- mediately adjacent to fin rays. Dorsal fin with two prominent horizontal pigment rows in unfaded specimens. As in other species of Elachocharax, a dark horizontal band is present along length of dorsal fin near its base. More distal band also well developed in unfaded specimens of E. mitopterus but dark chromatophores less concentrated. In both bands, dark chromatophores occur both on rays and membranes between them. Distal one-fourth of dorsal fin appears free of dark chromatophores. Anal and pelvic fins with two broad dark vertical bars (Fig. 1). Pectoral fins dusky in unfaded speci- mens, pigment concentrated along ray bor- ders. Dark band present at basal region of pectoral fin. Relationships. —In this cladistic analysis, outgroup information is the same as that derived in Weitzman and Géry (1981:898-— 905). Outgroup analysis for additional char- acters is presented here. The reasons for placing the new species in Elachocharax are quite clear. These are the high number of dorsal-fin rays, 17 or 18 in this species, and the position of the origin of the anal fin at Or near an imaginary vertical line drawn through the posterior termination of the dorsal fin. These features agree with char- acters one and two used as synapomorphies for Elachocharax by Weitzman and Géry (1981:898-901). Within Elachocharax, E. 744 mitopterus belongs to the group containing E. junki and E. geryi and consistently hav- ing unicuspid jaw teeth and no adipose fin. Weitzman and Géry (1981:902) also used a reduction (from 5 to 4) of branchiostegal rays to diagnose FE. junki and E. geryi as a group. Elachocharax mitopterus has five branchiostegal rays and would not fit a group diagnosed by four branchiostegal rays as a synapomorphy. Weitzman and Géry (1981: 902) note that the sample size of E. junki and EF. geryiis small and that in characidiine outgroups there is variation in this character within genera and species and occasionally between the two sides of the head ofa single specimen. If the character is at all useful as a synapomorphy within Elachocharax it would now allow recognition of EF. junki and E. geryi as a sister group of EF. mitopterus. This hypothesis is weakly supported by a synapomorphy consisting of an increased number of dorsal-fin rays, 19 to 21 in E. Junki and 20 to 22 in E. geryi. Elachocharax mitopterus has 17 or 18 dorsal-fin rays, a number more nearly approaching that of most outgroup species in the Characidiinae, 11 to 14 rays. However, E. pulcher, the nearest outgroup to E. mitopterus, E. junki, and E. geryi, has 17 to 20 dorsal-fin rays, variously overlapping in full, or in part, the ranges of the rays of the other species of Elachocharax. With this much overlap and such small samples at hand it seems pre- mature to suggest that a decreased number of branchiostegal rays and an increased number of dorsal-fin rays constitute synap- omorphous evidence of a group formed by E. junki and E. geryi. Regarding autapomorphies six and seven, the number and width of vertical body bars, in Weitzman and Géry (1981:903, 904), E. mitopterus has about 10 narrow, relatively uninterrupted vertical body bars. Thus the high number (16 or 17) and also the inter- rupted nature of each of the vertical body bars of E. junki appear to be autapomor- phies for that species. Elachocharax pulcher has about 8 to 10 wide vertical body bars. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Elachocharax geryi and E. mitopterus share 9 to 12 narrow body bars. The narrow na- ture and number of the bars in E. geryi and E. mitopterus appear to be plesiomorphic, more like the vertical bar pattern in out- group genera such as Characidium Rein- hardt and Klausewitzia Géry than the aut- apomorphous wide bars in E. pulcher or the relatively derived high number of inter- rupted autapomorphous bars in E. junki. Characters nine to twelve in Weitzman and Géry (1981:904) are autapomorphies for E. geryi. Two of these, numbers nine and eleven (number of inner-row dentary teeth and number of caudal-fin bars) remain clearly autapomorphous for that species. Elachocharax mitopterus has 18 or 19 teeth in the inner dentary row; E. pulcher has 20 to 24 and E. junki 19 to 20. Elachocharax geryi has a reduced number of 13 to 16. Similarly, E. mitopterus, E. pulcher, and E. Junki have O to 5 caudal-fin bars. Elacho- charax geryi has an increased number of 13 or 14. The remaining two characters, ten and twelve (number of ectopterygoid teeth and number of epibranchial gill rakers) are no longer clearly autapomorphous for E. geryl since E. mitopterus demonstrates character overlap. The number of ectopter- ygoid teeth in E. pulcher is 7 to 10, in E. Junki, 8 to 12, in E. mitopterus, 4 to 9, and in E. geryi itis 4 or 5. Similarly, the number of epibranchial gill rakers in E. pulcher is 4 or 5, in E. junki, 5 or 6, in E. mitopterus, 3 or 4, and in E. geryi it is 3. There might be a synapomorphous “trend” towards re- duction in these two characters shared by E. mitopterus and E. geryi. However, such “trends” are poor indicators of synapo- morphy because of overlap and, especially in this case, because they are based on so few cleared and Alizarin stained or partly ‘dissected samples, 3 in E. mitopterus, 2 in E. junki, 2 in E. geryi, and 10 in E. pulcher. Since the discovery of an additional taxon shows that these two characters are not as distinct as when utilized by Weitzman and Géry (1981:904), it now seems premature VOLUME 99, NUMBER 4 to use them as synapomorphies or autapo- morphies for these species. Elachocharax mitopterus has several unique characters appearing to be autapo- morphies, such as 29 vertebrae versus 30 to 33 in other species of Elachocharax, 24 to 26 scales in lateral series versus 26 to 33, 7 scale rows between the dorsal-fin origin and the pelvic-fin origin versus 8, and a tendency to have fewer scale rows around the caudal peduncle, 10 to 14 versus always 14 in the other species of Elachocharax. These unique reductions in meristic values in E. mitopterus make it unlikely that this species 1s a hybrid between other known species of Elachocharax, or that any one of them could be a hybrid between E. mitop- terus and one of the other species. Also, these counts are low compared to most species in other genera of the Characidiinae with at least two exceptions. Ammocrypto- charax elegans Weitzman and Kanazawa (1976:335) and Klausewitzia aphanes Weitzman and Kanazawa (1977:151) have 12 scale rows around the caudal peduncle. The most parsimonious cladogram of character distribution among characidiine taxa indicates that this similarity is homo- plaseous. Based on the above analysis, it seems that E. pulcher is the sister group of the other three species of Elachocharax, all charac- terized by two synapomorphies, unicuspid Jaw teeth and loss of an adipose fin. Except for a reduced number of branchiostegal rays and perhaps a reduced number of dorsal- fin rays in E. geryi and E. junki, the only apomorphies that so far seem clear and dis- tinct for E. junki, E. geryi, and E. mitop- terus are a series of autapomorphies for each. As discussed above, two putative synapo- morphies that might diagnose E. junki and E. geryi as a group seem somewhat dubious, especially the dorsal-fin ray count. There- fore E. junki, E. geryi, and E. mitopterus are currently considered to form an unre- solved trichotomy. Population variation. —Only small num- 745 bers, five each, of two population samples are available, making population variation study nearly impossible. However, one striking difference is present in the two sam- ples. Specimens from near San Carlos, Ven- ezuela always had 10 horizontal scale rows around the caudal peduncle, a very reduced number compared to the other species. The sample from igarapé Tarumazinho, Brazil had 12 to 14, x = 12.8. This appears to be an absolute difference in the two population samples of E. mitopterus. However, the samples are small and the possible magni- tude of the variation is unknown. That vari- ation of this sort occurs at all is surprising. Usually in Elachocharax the caudal pedun- cle scale row count is a constant 14. I would expect a count of 10 to be constant in E. mitopterus, as it appears to be in the San Carlos sample. The variation in the count in the igarapé Tarumazinho sample does not appear to be due to hybridization with other species of Elachocharax because this sample is like the San Carlos sample in hav- ing the other low meristic values character- istic for this species. An analysis of the cau- dal row scale count variation in E. mitopterus must await collection of larger samples. At this time it would seem pre- mature to recognize the two samples at hand as representing separate taxa. Ecological notes.—The specimens from Cano Chola were taken in a black water forest stream with a temperature of 28.0°C. The stream flow was slow to moderate and the depth of the water was to 1.5 meters. The bottom was sand and logs. Other forest detritus was present as well as filamentous algae and emergent aquatic vegetation. No field notes accompany the specimens from igarape Tarumazinho but when I visited the stream at this locality in January 1976 I found it to be a clear, slightly brown, terra firmae stream surrounded by forest. It was similar to the description here for Cano Chola except the water was less dark. Etymology.—The specific name mitop- terus is from the Greek mitos meaning thread 746 and pteron meaning wing, hence fin. The name is given in reference to the long thread like rays of the pectoral fin. Additional Locality Records for Other Species of Elachocharax Weitzman and Géry (1981:890) reported a few locality records for E. pulcher addi- tional to those recorded by Weitzman and Kanazawa (1978:162—164). Since then a few new locality records have come to hand. One specimen of E. pulcher, USNM 270142, SL 14.8 mm was found in the same collec- tion as the type locality of E. mitopterus, Cano Chola, between San Carlos de Rio Ne- gro and Solano, Departamento Rio Negro, Territorio Federal Amazonas, Venezuela. A short distance away, four specimens of E. geryi, USNM 270146, SL 10.7-13.8 mm, were taken from Cano Manu, a tributary of the Casiquiare Canal, about 250 meters up- stream of Solano, at about 02°00'N, 60°59'W. These were taken on 7 Dec 1984 by Richard P. Vari and Antonio Machado- Allison. This locality was a blackwater for- est stream. The water temperature was about 27.5°C and the water was moving very slow- ly. The bottom was sand and forest detritus, the water depth to about 0.5 meter at the capture site. It seems very likely that E. pul- cher, E. geryi, E. mitopterus, and probably E. junki, are sympatric in at least part of their range. All four species are known to live in black acid waters of terra firmae and all have locality records near the Rio Negro, Amazonas, see Weitzman and Kanazawa (1978:162-164, 173) and Weitzman and Géry (1981:890). In addition, nine specimens of Ela- chocharax pulcher, Naturhistoriska Riks- museet, Stockholm (NHM SOK) 1984312.3974, SL 9.7—-14.7 mm, were col- lected from Peru, Departamento Loreto, Rio Yavari (Javari) system, a quebrada tribu- tary to Rio Galvez, about 25 minutes up- stream from Colonia Angamos, about 05°16'S, 73°00'W, by Sven O. Kullander, A. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Urteaga C., N. Buendia Y., and A. Hoge- born-Kullander on 31 Jul 1984. This pop- ulation sample has some specimens with lateral tooth cusps on jaw teeth reduced in size, making them difficult to see except in stained examples. An adipose fin is present in all specimens. Acknowledgments The specimens from near San Carlos, Venezuela were collected by a joint MBUCV/USNM expedition made possible by funding from the Scholarly Studies Pro- gram of the Smithsonian Institution. I thank Dr. Antonio Machado-Allison (MBUCV) and Dr. Richard P. Vari (USNM) for mak- ing the specimens available. These individ- uals along with Mr. Carl J. Ferraris, Jr., American Museum of Natural History, Lic. Justa M. Fernandez, and Lic. Otto Castello (both MBUCV) are thanked for their col- lecting efforts. The specimens from igarapé Tarumazinho, Brazil were collected by Ivanzier Viera, Universidade Federal de Juiz de Fora, Minas Gerais, Brazil and were made available by Dr. Heraldo Britski and Dr. Naércio Menezes (MZUSP). Marilyn Weitzman, Lynn Norrod, Edgar N. Gram- blin, and Andrew G. Gerberich provided technical assistance. The drawings were pre- pared by Sara V. Fink. The study was sup- ported in part by the I.E.S.P. Neotropical Lowland Research Program of the Smith- sonian Institution. The manuscript benefit- ed from comments and suggestions by Rich- ard Vari, Wayne C. Starnes, and Marilyn Weitzman. Literature Cited Fink, W. L.,andS.H. Weitzman. 1974. Theso-called cheirodontin fishes of Central America with de- scriptions of two new species (Pisces: Charac- idae).—Smithsonian Contributions to Zoology 172:1—46. Géry, J. 1971. Une sous-famille nouvelle de poissons characoides sud-Americains: Les Geisleri- inae.— Vie et Milieu series C 12(1):153-166. . 1984. The fishes of Amazonia, pp. 353-370. VOLUME 99, NUMBER 4 In Monographie Biologicae, vol. 56, Harald Sioli, editor. The Amazon. Limnology and landscape ecology of a mighty tropical river and its basin. Dr. W. Junk Publishers, Dordrecht, Boston, Lancaster, 763 pp. Myers, G. S. 1927. Description of new South Amer- ican fresh-water fishes collected by Dr. Carl Ter- netz.— Bulletin of the Museum of Comparative Zoology 68(3):107-135. Weitzman, S. H., and J. Géry. 1981. The relation- ships of the South American pygmy characoid fishes of the genus Elachocharax, with a rede- scription of Elachocharax junki (Teleostei: Characidae). — Proceedings of the Biological So- ciety of Washington 93(4):887-913. , and R. H. Kanazawa. 1976. Ammocrypto- charax elegans, anew genus and species of riffle- inhabiting characoid fish (Teleostei: Charac- idae) from South America.— Proceedings of the 747 Biological Society of Washington 89(26):325- 346. ——., and 1977. A new species of pygmy characoid fish from the Rio Negro and Rio Amazonas, South America (Teleostei: Charac- idae).— Proceedings of the Biological Society of Washington 90(1):149-160. , and 1978. The South American fish genus Elachocharax Myers with a description of a new species (Teleostei: Characidae).—Pro- ceedings of the Biological Society of Washington 91(1):158-183. Department of Vertebrate Zoology (Fish- es), National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 748-752 NEW SPECIES OF CUMMINGSIA FERRIS (MALLOPHAGA: TRIMENOPONIDAE) FROM PERU AND VENEZUELA Roger D. Price and K. C. Emerson Abstract.— Descriptions and illustrations are given for two new species of Cummingsia: C. barkleyae off a probable new species of Thomasomys Coues from Peru and C. gardneri off Marmosa impavida (Tschudi) from Venezuela. In a recent review of the mallophagan ge- nus Cummingsia Ferris (Trimenoponidae), Timm and Price (1985) recognized a total of six species, with five of these in the nom- inate subgenus and the sixth in the subgenus Acanthomenopon Harrison. We have re- cently obtained two series of Cummingsia, each representing a new species; it is our intent here to describe and illustrate these new species. For brevity, the generic and subgeneric characters are as given by Timm and Price (1985) and will not be repeated here. All measurements are in millimeters. Cummingsia (Cummingsia) barkleyae, new species Figs. 1-3 Type host.—Thomasomys species, prob- ably new [Rodentia: Cricetidae]. Male.—As in Fig. 1. Dorsal head chae- totaxy as shown, without any heavier spi- niform setae. Both ventral spinous head processes on each side blunt, close together. With well-developed heavily pigmented ca- rina across posterior head margin and lack- ing medioposterior protrusion. Pronotum with median setae only slightly shorter than pair of lateral setae on each side near end of transverse thickening; metanotum with anterior setae all of essentially similar size. Prosternal plate with 7 long, 7 short stout setae; mesosternal plate with 5 long, 5 short stout setae; metasternal plate with 16-21 short to long setae. Tergal setae: I, 6 (with outermost very short, other 4 subequally long); II, 4-5; HI-—VII, 6; VIII, 4; [X, 1 very long, 1 medium on each side. Pleura II—VIII each with 3 marginal setae, with middle seta much longer than those adjacent to it. An- terior pleural setae on II, 2; III, 0-1; IV, 1- 2; V-VI, 1; VI, 0-1; VIII, 0. Marginal ster- nal setae: I, 9-11; II, 24—27; III, 21—25; IV, 15-21; V, 12-15; VI, 11-12; VII, 6. Ante- rior sternal setae: I, 4—6; II, 13-16; III, 5— 11; IV, 1-7; V, 0-1; VI-VII, 0. Subgenital plate (fused VIII-IX) with 10 setae. Di- mensions: preocular width (POW), 0.27- 0.30; temple width (TW), 0.34—0.38; head length (HL), 0.23-0.25; prothorax width (PW), 0.31-0.34; metathorax width (MW), 0.38-0.42; abdomen width at segment V (AW), 0.43-0.47; total length (TL), 1.27- 1.41. Genitalia as in Fig. 2, with genital plate broadly tapered, with sac sclerite thin, ““U”’- to “V”-shaped, and with tip of paramere with subapical seta well removed from end; genitalia width (GW), 0.07-0.08; genital plate width (GPW), 0.05-0.06; genital plate length (GPL), 0.04—0.05. Female.—Much as for male, except as follows. Prosternal plate with 7—9 short stout setae; mesosternal plate with 4—5 long setae. Anterior pleural setae: II, 2—3; III, 1-2; VIII, 0-1. Marginal sternal setae: I, 10—13; II, 27— 31; II, 22-27; IV, 19-23; V, 14-18; VI, 13- 15; VII (fused with VIII), 6-7. Anterior ster- nal setae: II, 15—19; III, 7-13; IV, 5—9; V, 1-4; VI, 1-2. Terminalia as in Fig. 3. Subgenital plate (fused VIII-IX) with 11- 12 setae, including 4 minute medioposterior 749 VOLUME 99, NUMBER 4 Wee TH ZS CIS Figs. 1-3. Cummingsia barkleyae: 1, Dorsal-ventral male; 2, Male genitalia; 3, Dorsal-ventral female ter- minalia. 750 setae. Anus oval, with 26-29 minute to me- dium setae. Dimensions: POW, 0.29-0.32; TW, 0.39-0.42; PW, 0.35-0.39; MW, 0.46-— 0.49; AW, 0.57-0.65; TL, 1.62-1.70. Remarks. — By having no abdominal ter- gites with more than six setae, C. barkleyae is allied with C. albujai Timm and Price and would be identified with the latter in couplet 2 of the key provided by Timm and Price (1985). However, C. barkleyae may be readily distinguished by its having (1) no dorsal head spiniform setae, (2) only four to five setae on tergite II, (3) consistently more marginal sternal setae on II-VI, with ranges well separated especially on II-IV, (4) more anterior sternal setae on II-IV, and (5) all abdominal pleurites with three mar- ginal setae, instead of only two on most pleurites of C. albujai. The chaetotaxy of the abdominal pleu- rites and dorsal thorax is similar for C. barkleyae, C. inopinata Mendez (the only other species known from Thomasomys), and C. perezi Timm and Price. Both C. in- opinata and C. perezi have more than six setae on at least five abdominal tergites; C. inopinata has distinctly fewer metasternal setae and marginal and anterior sternal se- tae on II-III; and C. perezi has distinctively different dorsal head and female subgenital plate chaetotaxy, as well as being larger in most widths and the male genitalia dimen- sions. Etymology. —This new species is named in honor of Linda J. Barkley, Los Angeles County Museum of Natural History, in rec- ognition of her interest in ectoparasites and for collecting the hosts bearing these lice. Holotype. —Adult 6, ex Thomasomys species, probably new, Peru, Dpto. Huanu- co, Unchog, NNW Acomayo, 3450 m, 3 Aug 1984, coll. L. J. Barkley (LJB field cat- alog number 2438); in collection of U.S. National Museum of Natural History. Paratypes. —Ex Thomasomys species, probably new: 34 44, 46 22, same locality and collector data as holotype, 18 Jul—5 Aug 1984 (off 17 host individuals with the fol- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lowing LJB field catalog numbers: 2340, 2341, 2342, 2366, 2376, 2389, 2390, 2391, 2412, 2426, 2432, 2434, 2438, 2439, 2440, 2443, 2449). Paratypes will be distributed among the University of Minnesota, Okla- homa State University, U.S. National Mu- seum of Natural History, and Field Mu- seum of Natural History. Note: Linda J. Barkley informs us that the host is most likely an undescribed species, which will ul- timately be described when her studies of the genus Thomasomys in Peru are com- pleted. Cummingsia (Acanthomenopon) gardneri, new species Figs. 4-6 Type host.—Marmosa impavida (Tschu- di) [Marsupialia: Didelphidae]. Male.—As in Fig. 4. Dorsal head chae- totaxy as shown. Slender pointed ventral inner spinous head process well separated from outer. With reduced carina across pos- terior head margin and with medioposterior protrusion. Pronotum with median setae very short and fine, with much longer pair of setae on each side near end of transverse thickening; metanotum with anterior setae all of similar length. Prosternal plate with 6 long, 12-15 short stout setae; mesosternal plate with 4 long, 13-15 short stout setae; metasternal plate with 21-22 short to long setae. Tergal setae: I, 6 (shortest outermost, longest innermost); II, 12-14; III, 16-17; IV, 15-19; V—VII, 17-19; VIII, 10; IX, 1 very long, | medium on each side. Pleura II and VI each with 4 marginal setae; IIJ- V with 4—5; VII-VIII with 3; all with 1 long among much shorter setae. Anterior pleural setae on II-III, 1-4; IV, 1-2; V—VI, 0-1; VII, 1; VUI, 0. Marginal sternal setae: I, 12- 13; II, 29-32; II-VI, 22-24; VII, 18-19. Anterior sternal setae: I, 3; II, 11-12; IJ- VII, 0. Subgenital plate with 20-21 setae. Dimensions: POW, 0.27-0.28; TW, 0.34— 0.37; HL, 0.19-0.22; PW, 0.29-0.32; MW, 0.33-0.34; AW, 0.46-0.49; TL, 1.14-1.17. 751 VOLUME 99, NUMBER 4 i: 4, Dorsal-ventral male; 5, Male genitalia; 6, Dorsal-ventral female ter- Figs. 4-6. Cummingsia gardner minalia. 752 Genitalia as in Fig. 5, with genital plate elongated, with small semicircular sac scler- ite, and with straight to slightly curved par- ameres each bearing minute seta near tip; GW, 0.07-0.08; GPW, 0.05; GPL, 0.07- 0.08. Female.—Much as for male, except as follows. Metasternal plate with 21-24 setae. Tergal setae: I, 8-9; II, 14-16; III, 16-18; IV, 19-20; V, 21-23; VI, 20-22; VII, 16- 21; VIII, 12. Pleura III-V each with 5 mar- ginal setae. Anterior pleural setae on II-III, 3-5; V-VI, 1. Marginal sternal setae: I, 13- 14; II, 31-34; IN-—VI, 22-26; VII, 20—22. Anterior sternal setae: I, 4—5; II, 12—14; III, Q-1;: IV—VI, 0; VII, 0-2. Terminalia as in Fig. 6. Anus with 25—29 minute to medium setae. Dimensions: POW, 0.28-0.29; PW, 0.31-0.33; MW, 0.37-0.40; AW, 0.52-0.54; TL, V-25=1.29: Remarks. —Since the subgenus Acantho- menopon contains only a single previously- described species, C. peramydis Ferris, C. gardneri resembles it and keys to it in cou- plet 1 in Timm and Price (1985), thereby differing grossly from all members of the subgenus Cummiungsia. The distinguishing features for C. gardneri are (1) the consis- tently smaller size of the female in all di- mensions and the male in POW, HL, AW, and TL, (2) the female with more tergal se- tae on I-III, and (3) the male genitalia small- er in all dimensions, with paramere tips not pronouncedly outwardly curved. Etymology. —This new species is named in honor of A. L. Gardner, U.S. National Museum of Natural History, in recognition PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of his interest in ectoparasites and his par- ticipation in the collection of the hosts bear- ing these lice. Holotype. — Adult 6, ex Marmosa impav- ida (Tschudi), Venezuela, T. F. Amazonia, Cerro de la Nebina, 4 Feb 1985, coll. A. L. Gardner and P. J. Spangler (ALG field cat- alog number 14418); in collection of the U.S. National Museum of Natural History. Paratypes.—Ex M. impavida: 6 46, 8 99, same locality and collector data as holotype, 3-4 Feb 1985 (off 2 hosts with the following ALG field catalog numbers: 14410, 14418). Paratypes will be distributed among the University of Minnesota, Oklahoma State University, and U.S. National Museum of Natural History. Acknowledgments This paper is published as Paper No. 14,911 of the Scientific Journal Series of the Minnesota Agricultural Experiment Station on research conducted under Minnesota Agricultural Experiment Station Projects No. Min-17-015 and Min-17-016. Literature Cited Timm, R. M., and R. D. Price. 1985. A review of Cummingsia Ferris (Mallophaga: Trimenopon- idae), with a description of two new species. — Proceedings of the Biological Society of Wash- ington 98:391—402. (RDP) Department of Entomology, Uni- versity of Minnesota, St. Paul, Minnesota 55108; (KCE) 560 Boulder Drive, Sanibel, Florida 33957. PROC. BIOL. SOC. WASH. 99(4), 1986, pp. 753-756 PARACYMOTHOA THOLOCEPS, A NEW FRESHWATER PARASITIC ISOPOD FROM SOUTHERN VENEZUELA (FLABELLIFERA: CY MOTHOIDAE) Thomas E. Bowman Abstract.— Paracymothoa tholoceps is described from a single specimen col- lected from Hoplias macrophthalmus in the Baria River, a tributary of the Orinoco River, in Amazonas State, Venezuela. It differs from the two known species, P. astyanactis and P. parva, in its larger size, convex anterior margin of the head, convex posterior margin of the telson, longer and more exposed pereonite 7, and less reduced dactyl of pereopod 7. The diagnosis of Paracy- mothoa is emended to accommodate the new species. The genus Paracymothoa was established by Lemos de Castro (1955) for a Brazilian isopod taken from the mouth of Astyanax bimaculatus. A second species, P. parva, de- scribed by Taberner (1976), was found par- asitizing Hyphessobrycon callistus in Argen- tina. A third species, from southern Venezuela, is described below. Paracymothoa tholoceps, new species Fig. 1 Material. —Venezuela, Amazonas terri- tory, Baria River, near base camp of expe- dition to Cerro de la Neblina, elev. 140 m (ca. 1°45'N, 66°W), ex Hoplias macroph- thalmus (Pellegrin) (Erythrinidae), leg. Ra- miro Ruyero, 26 Feb 1985; 1 2 without oos- tegites, 24.3 mm, holotype USNM 231092. Etymology.—From the Latin “‘tholus”’ (dome, rotunda) plus “‘-ceps” (head), refer- ring to the round anterior margin of the head. Diagnosis. — Paracymothoa tholoceps may be distinguished from its two congeners by its greater size, the shape of the head and pleotelson, and the well developed pereo- nite 7 and dactyl of pereopod 7. The prin- cipal differences between the three species are set forth in Table 1. Discussion As defined by Lemos de Castro (1955) and Taberner (1976), Paracymothoa differs from its presumed marine ancestor, Cy- mothoa Fabricius, 1793, by the short pereo- nite 7, almost completely covered by pereo- nite 6, the short dactyl of pereopod 7, much shorter than the dactyls of the other pere- opods, and the weak carinae on the bases of pereropods 4—6, lacking in pereopod 7. Paracymothoa tholoceps is closer to Cy- mothoa than P. astyanaxi and P. parva. Pe- reonite 7 and the dactyl of pereopod 7 are less reduced, and pereopods 5-6 have well developed carinae on the basis. The mandible was extremely difficult to remove, and appeared to be firmly attached to the labium. When finally extracted, it had an extraordinary form (Fig. If). Brusca (1981) has stated that some authors (in- cluding Bowman 1960: fig. 1h) have illus- trated a mandible with one lobe of the par- agnath attached to it and erroneously considered part of the mandible. Perhaps this explains the unusual form of the man- dible in Fig. 1f, yet when removed the la- bium appeared to be undamaged (Fig. 1g). The labium itself is unusual; the lateral lobes are rather slender and the anterior apices VOLUME 99, NUMBER 4 Table 1.—Characteristics of the species of Paracymothoa. SS) Character P. astyanactis! P. parva P. tholoceps Length = a — fo] sTi0W Inystonsban | aig Assad Din Fos fasta enw noise ait oe tw Li tis Aunuet Metin, & nigalsat hsrrlaedisg vith oaygeadl a { | witea tre wi ie Oc ad srwae a! Nee wy Ay * site toe ra : emer’ Stes 1900 bn ihe Waldo Seni See A.4 pin’ “agi'92) Pate! TWebit Our fener ete 7 ; = . ase Wan! AAV Ste Tats : fost See nent ; Pea wis 7 7 “ore re rhe? he j re, : j § a » ‘ s i ' v Bie! 5 wud te yeas WW 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-163, 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. Manuscripts are reviewed in order of receipt by a board of Associate Editors and appropriate referees. Proofs. —First proofs are submitted to authors for correction and approval. Reprint orders are taken with returned proofs. Publication charges. — Authors are asked to assume publication costs of page-charges, tabular material, and figures, at the lowest possible rates. Authors are required to pay the charges for figures, tables, changes at proof stage, and reprints. Payment of full costs will facilitate speedy publication. ‘Costs: Printed page @ $60.00, figures @ $10.00, tabular material $3.00 per printed inch. One ms. page = approx. 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. Manuscripts in dot-matrix will not be accepted. Submit two facsimiles (including illustrations) with the original, and retain an author’s copy. Pages must be numbered consec- utively. 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 A revision of the genus Nephasoma (Sipuncula: Golfingiidae) Norma J. Cutler and Edward B. Cutler On the species and populations of the genus Acanthocephalus (Acanthocephala: Echinorhyn- chidae) from North American freshwater fishes: a cladistic analysis Omar M. Amin A new subspecies of Jurdus swalesi (Aves: Passeriformes: Muscicapidae) from the Dominican Republic Gary R. Graves and Storrs L. Olson A new actinopterygian fish (Paleonisciformes) from the Upper Mississippian Bluestone For- mation of West Virginia Robert E. Weems and John F. Windolph, Jr. Larvae of Xiphopenaeus kroyeri (Heller, 1862) (Crustacea: Decapoda: Penaeidae) from offshore waters of Virginia, U.S.A. Robert C. Maris Two new crabs, Parapaguristes tuberculatus and Palaeoxantho libertiensis, from the Prairie Bluff Formation (Middle Maastrichtian), Union County, Mississippi, U.S.A. Gale A. Bishop A new species of Tubificoides Lastochkin (Oligochaeta: Tubificidae) from Bermuda and the Bahamas Berit Rasmark and Christer Erséus Four new genera of Dorvilleidae (Annelida: Polychaeta) from the Gulf of Mexico Paul S. Wolf Three new species of Dorvilleidae (Annelida: Polychaeta) from Puerto Rico and Florida and a new genus for dorvilleids from Scandinavia and North America Paul S. Wolf Redescription of Sheina orri Harding, 1966, a myodocopid ostracode collected on fishes off Queensland, Australia Louis S. Kornicker The first Anilocra and Pleopodias isopods (Crustacea: Cymothoidae) parasitic on Japanese fishes, with three new species Ernest H. Williams and Lucy Bunkley Williams Discovery and significance of Albany Hancock’s microscope preparations of excavating sponges (Porifera: Hadromerida: Clionidae) | Klaus Ritzler and Shirley M. Stone Newly established families of the Order Branchiobdellida (Annelida: Clitellata) with a synopsis of the genera Perry C. Holt Occurrence of larval J/lex illecebrosus and other young cephalopods in the slope water/Gulf Stream interface Michael Vecchione and Clyde F. E. Roper Astyanax scologaster, a new characid (Pisces: Ostariophysi) from the Rio Negro, South America Marilyn J. Weitzman and Richard P. Vari Barnacles (Cirripedia: Balanidae) from the Lower Pleistocene James City Formation, North Carolina coastal plain, with the description of a new species of Balanus Da Costa Victor A. Zullo and William Miller, III The synonymy of Cirolana tuberculata (Richardson, 1910) (Isopoda: Flabellifera: Cirolanidae) Paul M. Delaney Notes on the crayfish Procambarus (Ortmannicus) xilitlae (Decapoda; Cambaridae) Horton H. Hobbs, Jr. and Andrew G. Grubbs A new species of E/achocharax (Teleostei: Characidae) from the Rio Negro region of Venezuela and Brazil Stanley H. Weitzman New species of Cummingsia Ferris (Mallophaga: Trimenoponidae) from Peru and Venezuela Roger D. Price and K. C. Emerson Paracymothoa tholoceps, a new freshwater parasitic isopod from southern Venezuela (Flabel- lifera: Cymothoidae) Thomas E. Bowman International Commission on Zoological Nomenclature. Applications and Opinions Biological Society of Washington: Proceedings of the 113th Annual Meeting Table of contents, Volume 99 Index to new taxa, Volume 99 547 574 580 584 602 604 612 616 627 639 647 658 676 703 709 717 fal 735 139 748 SS Wem 760 763 770 ——-” f= — — 7 EE NOILNLILSNI NVINOSHLIAWS “OL THSONIAN INSTITUTION OShe SN N\ N\ NV S3!1YVYEIT LIBRARIES SMITHS H NOILNLILSNI NOILA.LILSNI LIBRARIES. Sh LIBRARIES Sh NVINOSHLII INSTITUTION NOILONLILSNI NOSHLINS S3JIYVYSIT LIBRARIES ~ SMITHSONIAN INSTITUTION NOILOLILSNI NVINOSHLIA Sy SJi4YVYdiT LIBRARIES SMITHSONIAI THSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWS 7) Ww fea < faa faa) sail JOSHLINS SAJIYVYSIT LIBRARIES > oMITHSONIAN INSTITUTION NOILNLILSNI bee z E = . XS ° ow o “oOo w a H ~ ~ - w 5 A Dw ~ . ’ \ ., = > Ey > > We = = - * i 2 SAX, 2 i Sn: : a . £ = wo &THSONIAN INSTITUTION NOILALILSNI_ NVINOSHLINS (S3 IyVvuad Hot IBRARI ES SMITHSON f Sey aes) ee < = = \ a. ee 5 s =z 5 = . 5 SN =z w no 2) 2) ” “ Tp) => re) =o o.% ba \. Oo FE Zz = = = y =< = ett ie = > = \ ae w ° Pt w as w) e Zz i (OSHLIWS $3 bYVesiT LIBRARI ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIV we = ad z ae NN WwW w WwW p ow lu : SS cc = x. (Wi yy = cc = S\N < < Pay, ti = Fa NVINOSHLIWS SMITHSONIAN if NVINOSHLIAWS SMITHSONIAN Ca > pe 4 Y o, 7 fe OSHLINS S3ZIYVYSIT LIBRARIES SMITHSONIAN NOILNLILSNI LIBRARIES NOILNLILSNI NOILNLILSNI INSTITUTION NOILOLILSNI NVINOSHLIAS Saluyvuygli INSTITUTION INSTITUTION INSTITUTION SIJIYVYSIT_LIBRARIES S S3IYVUYSIT LIBRARIES SMITHSONIAN -HSONIAN INSTITUTION NOILOLILSNI NVINOSHLIWS SMITHSONIA NVINOSHLINS S3IYVYUSIT LIBRARIES z NS 2 z = = Ne > WS Sf ‘Z 5 “Ss Qs See. S Ws a ro) a Oo WY 2 NR 2 2 By 2 Re fs » 2 i z = \. S > = > = . 7p Die ee w Foe w” : J JOSHLINS $3 byvVusit_ LIBRARI ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIW: Rc! “ = 7) ss SS SS ox = ee Yb fp = oc = SN SS. ae S z gs Y 3 = 2 WSS o aw oe Po . BR Gy ra Jy he = sy => «74 THCONIAN INGTITIITION NOLIQOTLICATL NIWIAINGCUTIMAIC CSI tuWwWUodIS hh hUDTL TR DOA WI Co)) 6 CRAITLIGCONIIA? — UTION NOILMLILSNI NVINOSHLINS S31YVYEIT_LIBRARIES SMITHSONIAN INSTITUTION <= “Rh. Sy = oy — a a — (ee) 2 2 ” a Ww z w JTION NOILALILSNI_ NVINOSHLIWS LIBRARIES SMITHSONIAN INSTITUTION | uw aa nid S Ww > ” us v = a uw a us oa — jag _ a ip eae a < a < = < nz < oc ar te ar a er « oa 3S = rs) = S ane a z is z = z ar Yai] LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3I1uVudiT | . = S ~ S 5 Es o — Bee! = = wo ~ % ~ = 0 ff Pe) AI = rf My s = F 2 <= te ¢ Vfy a ‘i E af uw My w ey) Oo z i aw = z i 2) = =a am wn oO = > = NVINOSHLINS S3I¥V¥slt 2 ey | Be z ae en a | Za = =z Z AYN: ~ ff ? &GM 2 aN > =a = =) = = a > GQ = > = x s * '. os w me Ww) “ rT) A ygit_ LIBRARI SSN A STITUTION Mer T TES Ny NWINOSHINS 22 byvVyesdIt > neh z — uw 2 Sy ul z 4 = “S z 2 NX = 4 a <3 c = a S\aee a = = = e = Sih ing = o o Ge re) am rs) 2 re) z , ey z ad eZ Seiya Uoah 2 UTION NOILNLILSNI NVINOSHLINS S3!YVUSIT LIBRARIES SMITHSONIAN 5 as SM aN | = z eyes 5 : : : : 5 $43 : - = at > = % > = - = E cae - ,. 2 E z z i z He Z LIBRARIES INSTITUTION NOILALILSNI NVINOSHLINS S31¥YVUSIT = Ww Ae w = w ; z = = < = iH. = a7 = < ar ee fo = z = z a ra yy. = g 2 g E 8 Hh ? g ~ = = ie =a Ale iia = 2 = = > = SS =) = > = 7) z a hee z2 7) z v) NVINOSHLINS S3IY¥VYRGIT LIBRARIES INSTITUTION ep) =e Ww S w = ; W uw 7) uw w = w = a = ec = fe a 2 < = . x c < ae < lad = < . = =f ~ = sg ro) ™' SS oe ro) ers Oo = z 2 ® a = ce = a Malt LIBRARIES SMITHSONIAN INSTITUTION NOJLNLILSNI NVINOSHLINS S3iy¥vudit = 5 ~ S L Bite 7 s = 5 7 = 2 = x 2 i s F Z = s His z j w z Oo _ =z v UTION NOILNLILSNI NVINOSHLINS SSid¥VuaIT_ LIBRARIES SMITHSONIAN INSTITUTION | w re, w =< os uv z= owe dp ne = < = «= = < Ko = were? 4108 1 “44 WSL NE N S = z i Z = z a z a 3 a 3 NOLLALILSNI NVINOSHLINS S3IuYvVuaIT s\i ISS SS N lyYvugil LIBRARIES SMITHSONIAN INSTITUTION LIBRARIES SMITHSONIAN __INSTITUTION NOILNLILSNI LIBRARIES LIBRARIES NOLIOLiILSNi LIBRARIES NOILNLILSNI NOILNLILSNI NOILOL i i SMITHSONIAN will LIBRARIES 3 9088 01205 2221 ill eee eet ee rer een Pee Pad ree ene era em eee eaten ate LLL OLA AL AD Mie fe AAD {URW Ns a NK Were nts we ine en ee or ewes